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, México.

*To whom correspondence and reprint requests should be addressed.

E-mail: maria_ana_tovar@yahoo.com

Tel: 52-983-8350440 ext. 4329.

(Accepted May 3, 2005)

Abstract María Ana Tovar-Hernández and Sergio I. Salazar-Vallejo (2006) Sabellids (Polychaeta: Sabellidae) from the Grand Caribbean. Zoological Studies 45(1): xxx-xxx. 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.

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

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Introduction

The Sabellidae Latreille 1825 is a family of polychaete annelids (Polychaeta Grube 1850) commonly known as “fan worms”, “feather-duster worms”, or “sea flowers”. They are easily recognized because living specimens have a colorful crown frequently projecting from the mouth of their tubes. According to Giangrande (in Rouse and Pleijel 2001), the family consists of 490 species, of which 75 belong to the subfamily Fabriciinae (13 genera), and over 400 correspond to the Sabellinae (30 genera) (Fitzhugh and Rouse 1999).

The Grand Caribbean region comprises the Gulf of Mexico, Caribbean Sea, Bermuda, and the northern littoral of Brazil (Salazar-Vallejo 2000). In the entire area, a number of important studies on the polychaetes have been carried out: Schmarda (1861), McIntosh (1885), Ehlers (1887), Treadwell (1901 1917 1921 1924a b 1928 1936 1939), Augener (1906 1922 1927), Monro (1928 1933a b), Hartman (1942 1951), Rioja (1946), Carpenter (1956), Renaud (1956), Marsden (1960), Jones (1962), and Uebelacker and Johnson (1984), among others. However, none of these studies focused exclusively on sabellids. Salazar-Vallejo (1996) listed 50 species of sabellids, belonging to 22 genera, occurring in the Grand Caribbean region.

The state of knowledge of the species of sabellids occurring in the Grand Caribbean, however, is still very poor and doubtful due to several reasons: for many years, identifications of Caribbean material were made using literature and taxonomic keys from other regions of the world. Moreover, the identification of species rested mainly on drawings (which were seldom original), without comparison to the type material. Also, inadequate preservation of the material may sometimes have led to misinterpretations of body structures or proportions. Finally, many of the original descriptions used to identify local fauna are incomplete and not illustrated. All of the above reasons led to misidentifications and records of species from localities far away from the Grand Caribbean.

Thus, the main objectives of this research were to determine the composition of species of sabellids occurring in the Grand Caribbean region, describe 4 new species, and provide an updated checklist and taxonomic key for the species occurring in the area.

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MATERIALS AND METHODS

Type and non-type materials were loaned from the following institutions: BMNH, ECOSUR, FSBC, LACM-AHF, UMML, USNM, and ZMA (see explanation of acronyms below). Samples from different localities in the Mexican Caribbean were collected at different depths from the intertidal zone to 10 m, by skin or scuba diving. Polychaetes were obtanined from dead coral blocks, algae, sponges, sea grasses, or sediments. Worms were placed in plastic bags, transferred to fresh water for up to 15 min in the shade, causing an osmotic shock which enhanced the relaxation of organisms and avoided contraction at fixation; samples were then fixed with a 10% formaline solution, washed in the lab with tap water for 24 h, and transferred to 70% ethanol for long-term preservation.

Specimens were generally examined with the aid of a stereomicroscope; characteristics of the chaetae, eyes, internal structure of the dorsal lips, and number of thoracic segments were observed using a compound microscope. Every specimen was analyzed to obtain the following measurements: width of the posterior thorax, body length (from the peristomium to the pygidium), length of the radiolar crown, and number of thoracic segments. Drawings were made with a camera lucida, while some illustrations of diagnostic structures were obtained using scanning electron microscopy (SEM).

 In order to standardize the relative size of the manubria and facilitate comparison between species, the following clasification was used: a) manubria were considered short if they were shorter than the distance between the breast and crest; b) medium, if they were as long as the distance between the breast and crest; c) long, if they were 1.5 times longer than the distance between the breast and crest; and d) extra long, if they were 2 or more times longer than the distance between the breast and crest. Further, uncini were separated into 2 different types with respect to the relative development of the breast; thus, breasts were considered well developed if they had a rounded projected lobe, or breasts were reduced, if this lobe was missing, having a sligthly rounded lobe in lateral view.

The checklist for the sabellids from the Grand Caribbean region is arranged in alphabetic order, including information as follows: author(s) and date of publication, type locality (TL), abbreviations of museums where the type material is deposited; other letters in parentheses are HT for holotype, PT for paratype, ST for syntype, LT for lectotype, NT for neotype, SST for schizosyntype, and in some cases, remarks. In this contribution, the species of Branchiomma and fabriciin sabellids are only included in the checklist. The revisions of the tropical American species of Branchiomma and the Grand Caribbean species of Chone are going to be published elsewhere (Tovar-Hernández and Knight-Jones, in press; Tovar-Hernández, in press). Fabriciin sabellids are documented in a series of contributions by Fitzhugh (1983 1990 1996).

Abbreviations

The following abbreviations are used in the text or in the checklist:

AMNH, American Museum of Natural History, New York, NY, USA; BMNH, The Natural History Museum, London, UK; ECOSUR, El Colegio de la Frontera Sur, Chetumal, Mexico; FSBC, Florida Marine Research Institute, St. Petersburg, FL, USA; LACM-AHF, Los Angeles County Museum of Natural History, Allan Hancock Foundation, Los Angeles, CA, USA; MCZ, Museum of Comparative Zoology, Harvard Univ., Cambridge, MA, USA; NHMW, Naturhistorisches Museum Wien, Wien, Austria; NMI, National Museum of Ireland, Dublin, Ireland; NMW, National Museum and Galleries of Wales, Wales, UK; UMML, Marine Invertebrates Museum, Rosenstiel School of Marine and Atmospheric Science, Univ. of Miami, Miami, FL, USA; USNM, The Natural History Museum, Smithsonian Institution, Washington DC, USA; UZIU, Museum of Evolution, Zoology Section, Uppsala Univ., Uppsala, Sweden; ZMA, Zoological Museum, Univ. of Amsterdam, Amsterdam, The Netherlands; ZMH, Zoologisches Institut und Museum, Univ. Hamburg, Hamburg, Germany; and ZMUC, Zoologisk Museum, Univ. København, Copenhagen, Denmark.

Key for sabellids from the Grand Caribbean

1 Breast of thoracic uncini narrow, poorly developed, giving uncini an acicular appearance (Fig. 7L) 2

- Breast of thoracic uncini well developed, giving uncini a Z-shaped or avicular appearance (Fig. 1M) 10

2 (1) Abdominal uncini with an elongate manubrium below dentate region.................... 3

- Abdominal uncini without or with reduced manubrium (Fig. 7K)............................... 7

3 (2) Branchial crown with 2 pairs of radioles........................................... Manayunkia

- Branchial crown with 3 pairs of radioles................................................................... 4

4 (3) Entire anterior peristomial ring collar membranous, or reduced to a low ridge...... 5

- Anterior peristomial ring collar as a low ridge dorsally and laterally, ventrally as a well-developed lobe 6

5 (4) Anterior peristomial ring collar complete middorsally; ventral filamentous appendages, if present, vascularized............................................................................................... Pseudofabriciola

- Entire anterior peristomial ring collar reduced to a low ridge, except for conical to elongate lobes on either side of dorsal midline; inferior thoracic neurochaetae in chaetigers 3~8, pseudospatulate Fabricinuda

6 (4) Ventral filamentous appendages branched; dorsal lips well developed, erect….Augeneriella;

Peristomial eyes rounded, well developed...................... A. hummelincki

- Ventral filamentous appendages unbranched; dorsal lips reduced to low ridges......Novafabricia

Inferior thoracic pseudospatulate notochaetae limited to chaetigers 3~5; manubrium of abdominal uncini twice longer than dentate region...................................................... ...N. infratorquata

7 (2) Anterior peristomial ring with narrow, triangular, ventral lobe; abdominal uncini with equal-sized teeth above main fang..................................................................................................... Amphicorina

- Anterior peristomial ring without triangular ventral lobe; abdominal uncini with unequal-sized teeth above main fang 8

8 (7) Palmate membrane absent........Jasmineira;

Collar well developed; dorsal lobes triangular, ventral lobes rounded..

........................................................................................... .J. bilobata

- Palmate membrane present...................................................................................... 9

9 (8) Last several abdominal chaetigers modified as a ventral anal depression......Euchone;

Anal depression in last 3 chaetigers; collar ventrally entire; 8 abdominal chaetigers E. incolor

- Anal depression absent................................................................................... Chone

10 (1) Manubrium of thoracic uncini extra long............Potamethus;

Collar ventral lobes very prolonged; thorax with 8 chaetigers...... P. spathiferus

- Manubrium of thoracic uncini short, medium-sized, or absent.................................. 11

11 (10) Abdomen with neuropodial chaetae in a group............................................. 12

- Abdomen with neuropodial chaetae in a row.......................................................... 16

12 (11) Companion chaetae present (Figs. 1K, L, 2C, 5G, 11E, F, 13G, 21E)......... 13

- Companion chaetae absent.................................................................................... 14

13 (12) Abdominal neurochaetae arranged in an incomplete spiral or C-shaped pattern Bispira

- Abdominal neurochaetae arranged in a tight spiral pattern............................... Sabella

14 (12) Radioles with stylods................................................................................... 15

- Radioles without stylods.........Sabellastarte;

Base of the crown involuted, forming spirals, thorax with 7 or 8 chaetigers..............

. S. magnifica

15 (14) Stylodes poorly developed (Fig. 18B); compound radiolar eyes present or absent…Pseudobranchiomma ;

Radioles with serrations, radiolar tips long, digitiform........... ...P. emersoni

- Stylodes well developed; compound radiolar eyes present, set in pairs ....Branchiomma

16 (11) Abdomen with paleate neurochaetae (Fig. 1N, O)........................................ 17

- Abdomen without paleate neurochaetae................................................................. 19

17 (16) Crown with dorsal and ventral basal flanges (Figs. 1C, 3C)........... Anamobaea

- Basal flanges absent............................................................................................... 18

18 (17) Chaetiger 1 with chaetae arranged in a row (Fig. 12A)...................... Notaulax

- Chaetiger 1 with chaetae arranged in a bundle......................................... Hypsicomus

19 (16) Inferior thoracic notochaetae broadly hooded (Fig. 10J)............................... 20

- Inferior thoracic notochaetae paleate...................................................................... 21

20 (19) Radioles with subdistal compound eyes (Fig. 9C, D).................... Megalomma

- Radioles without compound eyes................................................................ Demonax

21 (19) Radioles with compound eyes, except the dorsalmost pair (Fig. 19A)......... ......

......................................................................................... Pseudopotamilla;

3 or 4 compound eyes per radiole; dorsolateral collar margins entire (Fig. 19C).. .Pseudopotamilla fitzhughi sp. nov.

- Radioles without compound eyes...Perkinsiana;

long ventral lappets (Fig. 17A); up to 20 thoracic chaetigers............... P. fonticula

Amphicorina

1 Posterior peristomial ring collar crenulate laterally, with 10~12 abdominal chaetigers A. anneae

- Posterior peristomial ring collar with smooth margin; with 5 abdominal chaetigers

......... A. androgyne

Anamobaea

1 Two dorsal kidney-shaped shields over anterior margin of base of crown; flanges smooth (without papillae) (Fig. 3C)........................................................................... Anamobaea phyllisae sp. nov.

- Dorsal kidney-shaped shields absent; flanges wrinkled (with papillae) (Fig. 1C)..........

.......................................................................................... Anamobaea orstedi

Bispira

1 Radioles with eyes arranged in pairs................................................ B. melanostigma

- Radioles without eyes.............................................................................................. 2

2 With dorsal spongy cushions (Fig. 6A); collar with short rounded ventral lobes (Fig. 6B) B. paraporifera sp. nov.

- Without spongy cushions; collar with long triangular ventral lobes (Fig. 4B).................

......... B. brunnea

Branchiomma

1 Macrostylodes strap-like, tongue-like at midlength of radiole; collar with rounded ventral lobes; uncini with 3 rows of teeth above main fang.................................................................................... B. bairdi

- Microstylodes at midlength of radiole twice as long as the others; uncini with 1 row of teeth above main fang B. nigromaculatum

Chone

1 Pygidial cirrus present; crown with 10 pairs of radioles; 43 abdominal chaetigers

......... C. americana

- Pygidial cirrus absent (Fig. 7A); crown with 5 pair of radioles; 17 abdominal chaetigers Chone sp. 1

Demonax

1 Ventral shield of collar segment longer than wider..................................... D. flecatus

- Ventral shield of collar segment at least twice as wide as long................................... 2

2 Radioles with numerous pairs of minute ocelli; branchial crown as long as 1/4 of body; radioles with 4~6 dark bands D. microphthalmus

- Radioles without ocelli; branchial crown as long as 1/2 of body................................. 3

3 Ventral shield of collar segment as long as wide; thoracic tori not reaching ventral shields D. lacunosus

- Ventral shield of collar segment twice wider than long; thoracic tori indenting ventral shields (Fig. 8B) D. jamaicensis

Fabricinuda

1 Branchial crown dorsally displaced; anterior peristomial ring dorsally entire.................

......... F. trilobata

- Branchial crown not dorsally displaced; anterior peristomial ring ventrally U-shaped F. pseudocollaris

Manayunkia

1 Dorsalmost radiole with unpaired pinnules; ventral filamentous appendage wrinkled M. speciosa

- Dorsalmost radiole with paired pinnules; ventral filamentous appendage smooth........ ..

. M. aestuarina

Megalomma

1 Dorsal margins of collar fused to fecal groove; caruncle on anterior margin of shelf between dorsal lips, above mouth (Fig. 10B)............................................................................................. M. lobiferum

- Dorsal margins of collar not fused to fecal groove; caruncle absent............................ 2

2 Eyes present on most radioles................................................................................. 3

- Eyes present only on dorsalmost pair of radioles, large, with visible ommatidia

......... M. bioculatum

3 Eyes projected; segment 1 with quadrangular ventral shield (Fig. 9B).........................

......... M. heterops

- Eyes flat; segment 1 with rectangular ventral shield (wider than long)...........................

......... M. perkinsi sp. nov.

Notaulax

1 Ventral margin of collar entire.................................................................................. 2

- Ventral margin of collar incised................................................................................ 3

2 Collar segment as long as wide, longer than 3 following segments (Fig. 12B); radioles completely flanged except for small part of ocular region (Fig. 12D); about 30 ocelli per group........... N. bahamensis

- 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. 14C); 30~70 ocelli per group N. nudicollis

3 Dorsal margin of collar with a deep incision (Fig. 16A); short incision in ventral margin of collar (Fig. 16B); radioles with 4 or 5 ocelli (Fig. 16C), in small groups......................................... N. paucoculata

- Dorsal margin of collar without deep incision; long incision in ventral margin of collar. 4

4 Thorax with 13 chaetigers; radioles with many ocelli per row (> 50)............................... N. circumspiciens

- Thorax with 8 chaetigers; radioles with fewer ocelli per row (up to 30)..................... 5

5 Numerous ocelli (30 per row)............................................................ N. occidentalis

- Few ocelli (7 per row)............................................................................ N. midoculi

Pseudofabriciola

1 Ventral filamentous appendages present, vascularized; dorsal lips well developed

......... P. quasiincisura

- Ventral filamentous appendages absent.................................................................... 2

2 Anterior margin of anterior peristomial ring with a large conical structure (dorsal to mouth, separated from collar); abdominal uncini with manubrium 1.5-times longer than dentate region..... P. sofla

- Anterior margin of anterior peristomial ring with no prominent structure; abdominal uncini with manubrium as long as dentate region....................................................................................... P. longa

Anamobaea Krøyer, 1856

Anamobaea is known from a single Caribbean species, A. orstedi, for 150 years, which is a very attractive species due to the colors of the branchial crown. In the phylogenetic analysis of Fitzhugh (1989), Anamobaea is defined by 1 reversal of the chaetal state: the posterior rows of the anterior abdominal neurochaetae are modified, elongate, narrowly hooded. In this revision, we redescribe A. orstedi and describe a new species from Guana Island (Anamobaea phyllisae).

Anamobaea orstedi Krøyer, 1856

(Figs. 1, 2)

Anamöbæea Orstedii Krøyer 1856: 32.

Anamobaea orstedi.-- San Martín et al. 1994: 556, fig. 1A-I.— Humann 1992: 135.

Material: [ECOSUR] Mexican Caribbean: I. Cozumel, SEDENA, Coll. A. Medina, 24 Mar. 2001 (9). I. Contoy, Punta Sur, Coll. S. I. Salazar, 2 Mar. 2001 (5); Campamento de pescadores, Coll. M. A. Tovar, 1 Mar. 2001 (7). Cancún, Punta Nizuc, Coll. S. I. Salazar and L. F. Carrera, 1 Apr. 1997 (1). Buenavista, E-24, Coll. S. I. Salazar and L. F. Carrera, 27 Sept. 1996 (3). Xahuayxol, CR8, RC1, Coll. S. I. Salazar and L. F. Carrera, 2 June 1998 (2). Veracruz, I. Verde, Coll. S. I. Salazar, 15 June 1985 (3). [LACM-AHF] British Virgin Is.: Vc 0564, AF00-59, Guana, Beef I., Trellis Bay, Coll. G. Hendler, J. Martin, K. Fitzhugh, and R. Wear, 12 July 2000, tubes taken from coral rubble (1). [LACM-AHF] BVI 2000, Sta. 59 (1). [LACM-AHF] Vc 0556, airport (15). [LACM-AHF] LH 668, Sta. 110, Coll. T. Zimmerman, 25 July 2000, between White Beach and Harris Ghut (1). [UMML] Jamaica: 22.754, Pedro Bank, S Dominican Republic, R/V Pillsbury, Sta. 1254, 17°11'N, 78°31'W, Cruise 7006, 14 July 1970, 20 m (1). [UMML] Lesser Antilles: 22.753, R/V Pillsbury, 852, 11°53'N, 61°53'W, 3 July 1969, 13 m (3). [UMML] Puerto Rico: 22. 763, 1423, 21°41'N, 71°23'W, 19 July 1971, 18 m (3).

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

Anamobaea phyllisae sp. nov.

(Fig. 3)

Material: Holotype [LACM-AHF POLY 2148] British Virgin Is.: Off Guana Head, ARM, Coll. G. Hendler and R. Wear, 10 July 2000, 21.6 m, sand bottom, fine sand mixed with silt. [LACM-AHF] AF0056, VC0519, White Bay, 18.2 m directly off beach house, Coll. K. Fitzhugh, 11 July 2000, 2 m, crevice on side of large brain coral with sabellid tubes extending out (1).

Etymology: This species is named in honor of Dr. Phyllis Knight-Jones (Univ. of Swansea, UK) in recognition of her many years of dedication to the taxonomy of sabellid polychaetes.

Description: Holotype complete, 37 mm long. Crown long (21 mm), rigid, with 26 pairs of radioles. Basal flanges and basal 1/3 of radioles pale brown, next 1/3 pale, distal 1/3 yellowish-brown (in preserved material). Two pairs of erect, rigid, and smooth flanges (not papillated) on basal lamina, 1 pair dorsal, the other ventral, terminating on posterior peristomial ring (Fig. 3C). Two dorsal kidney-shaped shields on anterior margin of dorsal pair of basal flanges (Fig. 3A, C). Palmate membrane present above basal lamina (1.5-times length of basal lamina). Simple radiolar eyespots, orange, in a small group above palmate membrane (Fig. 3D), in a narrow band slightly above radiolar midlength. Radiolar skeleton with 8 cells in cross-section (Fig. 3 I). Dorsal lips short, with dorsal pinnular appendages. Ventral lips kidney-shaped, with parallel lamellae. Collar with dorsal margin indented by fecal groove (Fig. 3 A). Ventral lappets rounded, overlapping, (Fig. 3B), brownish-red. Ventral shield of segment 1 divided into 2 triangles, longer than the following shields. Thorax 20 mm long, 5 mm wide, with 74 chaetigers. Inferior thoracic notochaetae paleate, symmetrical (Fig. 3F, G), arranged in 2 transverse rows of 5 or 6 chaetae each. Superior thoracic notochaetae spinelike, short, 5 or 6 per group (Fig. 3E). Thoracic tori long, diminishing in length towards posterior end, not indenting ventral shields. Thoracic uncini avicular, with 18 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), breast well developed, manubrium medium (Fig. 3H). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 3J-M). Abdominal chaetigers 70. Abdominal neurochaetal fascicles in 2 transverse rows; anterior and posterior rows on all chaetigers with asymmetric paleate chaetae, mucroate, with small teeth at base of mucro (Fig. 3N) and modified, elongate, narrowly hooded chaetae (Fig. 3O). Abdominal uncini avicular, with 27 equal-sized teeth above main fang in lateral view, covering 3/4 of extension of main fang (seen under light microscopy), breast well developed (Fig. 3P), manubrium long. Tube distally thinner, semi-transparent, covered by fine sand and algae; thicker at midlength, leathery, with incrusted calcareous algae; very hard basally.

Remarks: Anamobaea phyllisae sp. nov. is unique by the presence of smooth basal flanges, without rounded papillae and 2 dorsal kidney-shaped shields located on the anterior margin of the dorsal pair of the basal flanges, and a dorsal collar margin divided by a deep fecal groove. In the Caribbean, there is another undescribed species of Anamobaea (Anamobaea sp. 1: [LACM-AHF] BVI 99, Sta. 123, LH 1005, Guana: Bigelow Beach, Coll. T. Zimmerman, 15 Aug. 1999, 9~11 m (14). AF0042, VC0489, Off Guana Head, Coll. G. Hendler and R. Wear, 10 July 2000, 21.6 m, sand bottom, fine sand mixed with silt). It differs from A. phyllisae by having a different radiolar color pattern in preserved material, as follows: white-purple-transparent-orange-transparent with the transparent flanges, and for lacking the kidney-shaped shields (see Humann 1992: 131). Anamobaea orstedi and Anamobaea sp. 1 have different radiolar color patterns in live specimens (red-white in A. oerstedi and white-purple-transparent-orange-transparent in Anamobaea sp. 1), but both have bright white patches dorsally and ventrally on the basal flanges.

Bispira Krøyer, 1856

Krøyer (1956) defined Bispira without including any species. Knight-Jones and Perkins (1998) designated Amphitrite volutacornis Montagu, 1804, as the type species. Bispira is recognized by the independent adquisition of radiolar flanges and dorsal pinnular appendages (Fitzhugh 1989). Knight-Jones and Perkins (1998) recognized 19 valid species; only B. brunnea (Treadwell, 1917) and B. melanostigma (Schmarda, 1861) are found in the Grand Caribbean.

Bispira brunnea (Treadwell, 1917)

(Fig. 4)

Metalonome (sic) brunnea Treadwell 1917: 268, pl. 3, figs. 24-27.

Sabella bahamensis Augener 1922: 48 fide Knight-Jones and Perkins 1998.

Bispira brunnea.-- Knight-Jones and Perkins 1998: 433, figs. 19, 20.

Material: [ECOSUR] Mexican Caribbean: Punta Allen, Coll. P. Gómez, 23 Apr. 1992 (34). Puerto Morelos, Coll. M. S. Jiménez, 7 Sept. 1986 (22). Majahual, Coll. J. R. Bastida and P. Salazar, 21 Mar. 2000 (73). Buenavista, Coll. S. I. Salazar and L. F. Carrera, 27 Sept. 1996 (52). I. Cozumel, Chankanaab, Coll. S. I. Salazar, 2 Apr. 1992 (78); SEDENA, Coll. M. A. Tovar, 5 Mar. 2001 (89). Laguna de Términos, July 1984, 30.27a (3). [LACM-AHF] British Virgin Is.: Vc 0627, Sta. 59 Guana, Beef I., Trellis Bay, Coll. G. Hendler, J. Martin, K. Fitzhugh, and R. Wear, 12 July 2000 (30). [LACM-AHF] Grand Ghut, Coll. G. Hendler, 6 July 2000, algal clumps near Graham Forrester (15). [LACM-AHF] AF00-16A, Vc 0236, Grand Ghut, near Graham Forrester's transect, reef slope beyond spur and groove zone, algal clumps, 3~4 cm high, mainly Laurencia and Dictyota, 19.8 m, 7 June 2000. [LACM-AHF] Vc 0237. [LACM-AHF] Vc 0238. [ZMA] St. Martin: V. Pol. Great Bay, Pt. Blanche Bay, Coll. P. Wagenaar Hummelinck, Sta. 1125A, 26 Apr. 1949, conglomeratic rocks with balanids, some sand, 0.5~1.5 m (88).

Description: Gregarious species. Body long (5.5~36 mm). Crown brown at base with white radioles (in preserved materials). Crown longer than body, 6~38 mm (Fig. 4A), radioles arranged in 2 semicircles with slightly involuted ventral edges, with 13 or 14 pairs of radioles. Radioles with bare tips as long as equivalent space of 16 pinnules (Fig. 4C), without eyes. Radiolar skeleton with 8 cells in cross-section disposed in a reniform arrangement at base of radioles; narrow flanges along edges of outer surface. Dorsal lips with radiolar appendages extending beyond palmate membrane; pinnular appendages present, surrounding sheath tissue absent, radiolar appendage skeleton composed of a single row of cells, and connective tissue at dorsal lip lamellae. Ventral lips and parallel lamellae present. Collar with wide dorsal gap, lateral margins extending well beyond origin of crown. Long, triangular ventral lappets (Fig. 4B), with small lobular extensions overlapping at midline. Anterior margin of shield of segment 1 W-shaped, following thoracic shields rectangular with wide gap to adjacent tori (Fig. 4B). Spinelike chaetae on segment 1. Thoracic and abdominal inter-ramal eyespots present. Thorax 2~6.5 mm long, 1.5~2 mm wide, with 10 or 11 chaetigers. Superior and inferior thoracic notochaetae spinelike (Fig. 4F) arranged in bundles, with irregular, longitudinal rows of chaetae. Thoracic uncini avicular, with 4 or 5 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), breast well developed (Fig. 4D), manubrium medium. Companion chaetae with distinctly asymmetrical membranes. Abdomen with 22~84 chaetigers. Abdominal neuropodia as conical lobes. Abdominal neurochaetal fascicles partially spiraled or C-shaped. Anterior abdominal neurochaetae spinelike on anterior and posterior rows; posterior abdominal neurochaetae spinelike on anterior rows, and modified, elongate, narrowly hooded on posterior rows. Abdominal uncini avicular, with 5 or 6 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), breast well developed, manubrium short (Fig. 4E). Tube built with fine white sand.

Remarks: Bispira brunnea is similar to B. porifera (Grube, 1878), as both species lack radiolar eyes; however, the collar segment in that species is very elongate and has a dorsal spongy structure, both characters of which are absent from B. brunnea. This is a gregarious species, very common in the Caribbean, easily recognized alive by the white tubes and bicolored crown.

Bispira melanostigma (Schmarda, 1861)

(Fig. 5)

Sabella melanostigma Schmarda 1861: 36, pl. 22, fig. 190.-- Uebelacker 1984: 54-42, 54-43.

Sabella variegata Krøyer 1856: 29-30 fide Knight-Jones and Perkins 1998: 415.

Sabella thoracica Krøyer 1856: 31-32 fide Knight-Jones and Perkins 1998: 415.

Sabella bipunctata Baird 1865: 158-159 fide Knight-Jones and Perkins 1998: 415.

Material: [BMNH] St. Vincent: [1839.12.27./27]. 16~20 m, Rev. Landstone Guilding Coll. (3 specimens labeled as syntypes of Dasychone nigromaculata, correspond to Bispira melanostigma). [ECOSUR] Mexican Caribbean: Akumal, 12 Apr. 1986 (2). Banco Chinchorro, 28 July 1990, 2.5 m (1). Canal Zaragoza, Coll. J. Espinosa, 9 Oct. 1997, in mangrove roots (1). I. Cozumel, Chankanaab, 2 Apr. 1992 (3). Desembocadura de Bacalar Chico, Coll. J. Espinosa, 9 Oct. 1997, in Rhizophora mangle (15). DIF Puerto Aventuras, Coll. S. I. Salazar, 21 Mar. 1992 (5). Hualalpich, 17 Apr. 1986 (1), 19 Apr. 1986 (5). I. Contoy, Punta Sur, Coll. M. A. Tovar and S. Frontana, 28 Feb. 2001 (3). Majahual Norte, Coll. M. A. Tovar, 19 Jan. 2001 (21); muelle, Coll. H. ten Hove, 18 Mar. 2001 (various dozens). Cancún, Laguna Nichupté, 20 Apr. 1988, E7M1+1 (2); Punta Nizuc, Coll. S. I. Salazar and L. F. Carrera, 1 Sept. 1997 (3). Punta Gavilán, 9 Apr. 1992, Coll. S. I. Salazar (5); Coll. M. Londoño, 10 Feb. 2001, 4 m (7). Punta Río Indio, Coll. M. A Tovar, 17 Mar. 2001 (3). Río Lagartos, Coll. J. R. Bastida and S. I. Salazar, 18 Feb. 1999 (1). Santa Cecilia, bolsa 101, 4 Nov. 1990 (1). Xahuayxol, Coll. S. I. Salazar and L. F. Carrera, 27 Sept. 1996 (1). Xamach, Coll. E. Donath, 28 Feb. 1986 (2). Xcacel, Coll. S. I. Salazar, 3 Apr. 1992 (3). Xcalay, B 115, 4 Nov. 1990 (10). Yalahau, 8 Sept. 1993 (1). Panama: Fuerte Sherman, Colón, Coll. S. I. Salazar, 2 June 2002 (3). [LACM-AHF] British Virgin Is.: BVI-99-12F, LH 0143, Guana, North Bay, Coll. L. Harris, 25 July 1999, west rocky intertidal, clumps of zoanthids Zoanthus pulchellus (3). [LACM-AHF] BVI-99-25F, LH 142, Tortola, Toll bridge, Coll. L. Harris, 29 July 1999, mangroves (6). [LACM-AHF] BVI-99-53, LH 0361, Pelican Ghut, Coll. T. Zimmerman, 3 Aug. 1999 (1). [LACM-AHF] BVI-99-81, LH 0754, Tortola, Coll. L. Harris, 7 Aug. 1999 (8). [LACM-AHF] AF00-04, Vc 0050, White Bay, Coll. K. Fitzhugh, 4 July 2000, 0.5 m, tubes attached to coral boulder (1). [LACM-AHF] AF00-04, Vc 0051 (1). [LACM-AHF] AF00-04, Vc 0052F (6). [LACM-AHF] AF00-04, Vc 0054 (1). [LACM-AHF] AF00-54, Vc 0555, Muskmelon Bay, Long Point, near ARM site, Coll. G. Hendler and R. Wear, 11 July 2000, 9.1~15.2 m, gorgonians and sponges, sediment extremely silty, covered by red, green, and brown algae (2). [LACM-AHF] AF00-57, Vc 0480, White Bay, Coll. K. Fitzhugh, 11 July 2000, 2 m, coral rock (1). [LACM-AHF] AF00-59, Beef I., Trellis Bay, Coll. G. Hendler, J. Martin, K. Fitzhugh, and R. Wear, 12 July 2000 (15). [LACM-AHF] AF00-59, Vc 0628 (1). [LACM-AHF] AF00-60, Vc 0673, Muskmelon Bay off Crab Cove, Coll. G. Hendler, T. Zimmerman, J. Martin, and R. Wear, 13 July 2000, 21.3 m (1). [LACM-AHF] Bahamas: LH03-29, Sta. 15; LH03-30, Sta. 15; LH03-31, Sta. 15; LH03-34, Sta. 010 (5); LH03-72, Sta. 21; LH03-203, Sta. 045 (2); LH03-204, Sta. 045 (1); LH03-215, Sta. 55; LH03-225, Sta. 55. [UMML] Mexican Caribbean: 22. 764, Majahual, muelle, Coll. H. ten Hove, 18 Mar. 2001 (20); 22.765, Xcalay, B115, Nov. 4, 1990 (6). [ZMA] Jamaica: V. Pol. Kingston Harbour, Small Boat Channel, E of Port Royal, Coll. P. Wagenaar Hummelinck, Sta. 1678, 7 May 1973, Rhizophora timber, 0~1 m (50). V. Pol. flood gate of The Falsees Great Salt pond, Coll. P. Wagenaar Hummelinck, Sta. 024, 8 May 1975 (10). [ZMA] Antigua: V. Pol. Dickinson Bay N, Coll. P. Wagenaar Hummelinck, Sta. 1540B, 26 July 1967, eroded Thalassia flat, 0.5~1 m (7). [ZMA] St. Martin: V. Pol. Great Bay, Pt. Blanche Bay, Coll. P. Wagenaar Hummelinck, Sta. 1125A, 26 June 1949, rocks, sand, low tide and lower zone (20).

Description: Medium-sized specimens, 21~68 mm long. Thorax inflated (Fig. 5B), with anterior purple spots. Crown purple (in live and preserved material), 13~30 mm long, with 16~21 pairs of radioles in semicircular arrangement. Radioles with 2 or 3 pairs of dark-brown compound eyes (Fig. 5C). Radiolar skeleton with 8~10 cells in cross-section, obtuse epithelial corners on outer surface (Fig. 5D). Dorsal lips with radiolar appendages extending beyond palmate membrane; pinnular appendages present. Ventral lips and parallel lamellae present. Dorsal margins of collar prominent and separated by fairly wide gap, lateral margins just covering origin of crown (Fig. 5A). Ventral lappets prominent and medially involuted, forming small pockets flanking midline cleft (Fig. 5B). Ventral thoracic shields trapezoidal on segment 1. Spinelike chaetae on segment 1. Purple patches on ventral sacs and dorsally on thorax, dark patches close to dorsal sides of all parapodia, and small marks at ventral end of each thoracic torus and ventral to each abdominal fascicle. Thorax with 8~14 chaetigers, 4.5~9 mm long, 4.5~7 mm wide. Thoracic ventral shields trapezoidal, not indented by tori. Inferior thoracic notochaetal arranged in bundles, with irregular, longitudinal rows of spinelike chaetae. Thoracic uncini avicular, with 3 or 4 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main faing (seen under light microscopy), breast well developed, manubrium medium (Fig. 5E). Companion chaetae with distinctly symmetrical distal membranes (Fig. 5G). Abdomen with 64~72 chaetigers. Abdominal neurochaetal fascicles in partially spiraled or C-shaped arrangement formed by posterior chaetal row, posterior abdominal neurochaeta spinelike on anterior rows, and modified, elongate, narrowly hooded on posterior rows. Abdominal uncini avicular, with 3 or 4 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main faing (seen under light microscopy), breast well developed, manubrium short (Fig. 5F). Tubes made of fine white sand.

Remarks: Bispira melanostigma is a common species in the Caribbean, being very attractive because of the purple color of the branchial crown. Knight-Jones and Perkins (1998) considered Sabella thoracica Krøyer, 1856, B. variegata (Krøyer, 1856), and Bispira melanostigma (Schmarda, 1861) as synonyms. Although S. thoracica and B. variegata have priority over B. melanostigma, those names have not been used since 1856, except B. variegata by Perkins (1984). Conversely the name melanostigma has been in frequent use since its inception and is the name used by the curators of most museums. Therefore, to conserve the name melanostigma, the precedence of S. thoracica and B. variegata should be waived (Art. 23, Sect. 9, International Code of Zoological Nomenclature, 2000).

Bispira paraporifera sp. nov.

(Fig. 6)

Material: Holotype [LACM-AHF POLY 2149] Mexican Caribbean: Holotype, DIF Puerto Aventuras, Coll. S. I. Salazar, 22 Mar. 1992, on dead coral.

Etymology: Specific name refers to the similarity between this species and B. porifera.

Description: Holotype complete, 16 mm long. Crown pale (in preserved material), 8 mm long with unflanged bases and 14 pairs of radioles, arranged in 2 semicircles with slightly involuted ventral edges. Radioles with 4 pairs of eyes, long bare tips, as long as equivalent space of 8 pinnules, radiolar flanges present in area between eyes and radiolar tip. Radiolar skeleton with 8 cells in cross-section. Dorsal lips purple, with radiolar appendages extending beyond palmate membrane; pinnular appendages present. Ventral lips and parallel lamellae present. Collar dorsal margins with orange spongy, cushion-like mass extending to 6th thoracic chaetiger, superior border heart-shaped, inferior border V-shaped (Fig. 6A); ventral lappets small, rounded with purple spots, overlapping (Fig. 6B). Thorax 5 mm long, 3 mm wide with 8 chaetigers. Ventral shield of segment 1 W-shaped (Fig. 6B); following shields rectangular, not indented by tori. Thoracic and abdominal inter-ramal eyespots present (Fig. 6B). Inferior thoracic notochaetal fascicles arranged in bundles, with irregular, longitudinal rows of spinelike chaetae (Fig. 6F, G). Thoracic uncini avicular, with 5 equal-sized teeth in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), breast well developed, manubrium medium (Fig. 6C). Companion chaetae with distinctly asymmetrical distal membranes (Fig. 6E). Abdomen with 38 chaetigers. Abdominal neuropodia as conical lobes, in partially spiraled or C-shaped arrangement. Abdominal neurochaetae spinelike on anterior and posterior rows (Fig. 6H), and modified, elongate, narrowly hooded on posterior rows (Fig. 6I, J). Abdominal uncini avicular, with 3 equal-sized teeth in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), breast well developed, manubrium short (Fig. 6D). Gametes visible on 4th abdominal chaetiger.

Remarks: The spongy cushion of Bispira paraporifera sp. nov. is similar to that of B. porifera (Grube, 1878), described from the Philippines. However, B. paraporifera has the spongy cushion-like mass restricted to the dorsal side of the thoracic segments, its inferior border is V-shaped, and it extends to the 6th thoracic chaetiger, while in B. porifera, the spongy cushion-like mass extends around the peristomium to the ventral sacs into smaller cushions; its inferior border is rounded and less developed, and it extends to the 8th thoracic chaetiger. Further, B. porifera has radioles without eyes, with incipient paired flanges along the outer surface becoming more distinct distally, radiolar tips very short, and ventral lappets prominent (see Knight-Jones and Perkins 1998, fig. 16A-P). Willey (1905) suggested that the spongy cushions of B. porifera are glandular, while Knight-Jones and Perkins (1998) suggested that variability in texture and degree of coverage in spongy cushions in B. porifera may be due to a developmental series and may be involved in the incubation of embryos.

Chone Krøyer, 1856

Banse (1972) emended Chone, stressing the girdle of glands on the 2nd chaetiger, the ventral shields, the “bayonet-like chaetae”, and the “subespatulate chaetae”. He included Megachone Johnson, 1901, redescribed or improved the knowledge of some species, described 3 new species, and recognized that the taxonomy of this genus was confusing because some species present variations in the morphology of the abdominal uncini.

Fitzhugh (1989) placed Chone within the subfamily Sabellinae, giving emphasis to the presence of the branchial skeleton and to the origin of the collar. He pointed out that Chone is not defined by any synapomorphy; also, he emended the genus because some specimens previously identified as C. infundibuliformis have dorsal lips with dorsal radiolar appendages, whereas other specimens do not have such structures. Fitzhugh (1989) named those specimens Chone1 and Chone2, respectively. In addition, Fitzhugh (1989) suggested that Chone possibly be divided into several monophyletic genera based on the retention or loss of dorsal radiolar appendages and the morphology of the abdominal uncini.

Chone sp. 1

(Fig. 7)

Material: [ECOSUR] Panama: Club de Yates, Colón, Coll. S. I. Salazar, 1 June 2002 (1). [ZMA] Grenada: V. Pol. Hog I. near Point Salines, Thalassia in muddy sand near Rhizophora, 0.5~1 m, 8 July 1967, Coll. P. Wagenaar Hummelinck, Sta. 1551A (1).

Description: Specimen from Panama complete, small, cylindrical body 5.5 mm long (Fig. 7A). Crown 1.5 mm long, palmate membrane along 3/4 of radiolar length, 5 pairs of radioles with broad radiolar flanges (Fig. 7C) and short bare tips, as long as equivalent space of 6 pinnules (Fig. 7B). Pinnules of same length along radioles. Dorsal margins of collar fused to fecal groove; lateral and ventral margins of collar entire. Radiolar skeleton with 2 rows of cells in cross-section. Three pairs of ventral radiolar appendages. Dorsal lips erect, ventral lips rounded. Anterior margin of anterior peristomial ring with narrow-ventral lobe. In lateral view, ventral margin slightly elevated (Fig. 7A). Segment 1 with broad, dark ventral shield, with rounded anterior margin. Thorax with 8 chaetigers, each with 2 pigmented bands (Fig. 7C). Glandular girdle present on chaetiger 2. Inferior thoracic notochaetae in 2 transverse groups: anterior rows with 3 or 4 bayonet chaetae (Fig. 7J), posterior rows with 3 or 4 paleate chaetae with small mucro (Fig. 7D-G). Nine acicular uncini per tori, with 5 equal-sized teeth above main fang in lateral view (seen under light microscopy) (Fig. 7L). Abdomen with 17 chaetigers. Abdominal neurochaetae arranged in 2 transverse rows per fascicle; anterior row on anterior chaetigers with 4 or 5 elongate, narrowly hooded chaetae; posterior row with modified, elongate, narrowly hooded chaetae; posterior abdominal fascicles with modified, elongate, narrowly hooded chaetae in both rows (Fig. 7H, I). Abdominal uncini with main fang surmounted by 3 or 4 small-sized teeth in lateral view (seen under light microscopy), breast well developed, manubrium absent (Fig. 7K). Pygidium rounded, cirrus and eyespots absent. Sperm with rectangular heads, and 2 small mitochondrial lobes.

Remarks: In the Grand Caribbean Region no species of Chone has been described. Chone americana Day, 1973 (described from Beaufort, NC) has a pygidial cirrus, while our specimens have 5 pairs of radioles, broad flanges, short radiolar tips, 17 abdominal segments, all paleate chaeta with well-developed mucro, sperm with rectangular heads, and no pygidial cirrus. Material of Chone sp. 1 is available at ECOSUR and ZMA; we included this species in the present paper in order to facilitate and encourage further research.

Demonax Kinberg, 1867

Generic diagnosis was provided by Knight-Jones (1983), Knight-Jones and Walker (1985), Perkins (1984), and Fitzhugh (1989). The primary character distinguishing Demonax is that companion chaetae are distally dentate (Fitzhugh 1989). Only 3 species of Demonax have been described from the Grand Caribbean: D. flecatus (Hoagland, 1919) from Puerto Rico, D. jamaicencis (Augener, 1922) from Jamaica, and D. lacunosus Perkins, 1984 from Hutchinson Island, FL.

Demonax jamaicensis (Augener, 1922)

(Fig. 8)

Parasabella jamaicensis Augener 1922: 48.

Demonax jamaicensis.-- Johansson 1927: 136.-- Perkins 1984: 315-317, fig. 17.

Material: [ZMA] St. Thomas: V. Pol. Benner Bay lagoon, 0~1 m, 30 Apr. 1973, Coll. P. Wagenaar Hummelinck, Sta. 1674, Rhizophora (3).

Description: Medium-sized body, 13~23 mm long, 1.5 mm wide. Branchial crown 5 mm long, with 10 pairs of radioles, with 5 yellow crossbands on pinnules (in preserved material). Radioles in semicircles, without flanges or stylodes. Dorsal margin of dorsal lip extending up along inner margin of dorsalmost radiole and fused with 1~3 of the more-proximal pinnules. Ventral lips triangular, well developed, beginning between ventral lappets extending dorsally to near dorsal lips. Collar short, widely separated dorsally by fecal groove, beginning slightly anterior and medial to chaetae from segment 1 (Fig. 8A). Thorax with 8 chaetigers. Ventral lappets rounded (Fig. 8B). Ventral shield of segment 1 rectangular, about twice as wide as long, broader than following segments, anteriorly indented at midlength; following shields laterally indented by tori (Fig. 8B). Spinelike chaetae on segment 1 in 2 short rows. Inferior thoracic notochaetae broadly hooded (Fig. 8E), arranged in several transverse rows. Thoracic uncini avicular, with equal-sized teeth above main fang, breast well developed, manubrium medium (Fig. 8C). Companion chaetae with very stout shaft and long mucro. Abdomen with 55 chaetigers. Abdominal neurochaetae in 2 transverse rows of elongate, narrowly hooded chaetae (Fig. 8F). Abdominal uncini avicular, with main fang surmounted by 7 equal-sized teeth in lateral view, covering 1/2 of main fang length (seen under light microscopy), breast well developed, manubrium short (Fig. 8D).

Megalomma Johansson, 1927

Megalomma was studied by Perkins (1984), Fitzhugh (1989), and Knight-Jones (1997). The latter author divided 23 species of Megalomma into 5 groups based on (1) whether the middorsal collar margins are fused to the fecal groove or not; (2) whether the dorsolateral margins of the collar form pockets or not; and (3) the extent to which eyes occur in the radioles. Nishi (1998) described a new species from Thailand, and added a new group (2D) to the description proposed by Knight-Jones. Fitzhugh (2002) described a new species from Thailand included in group 1A, and later (Fitzhugh 2003) described a new species from Taiwan (group 1B) and emended the diagnosis of the genus to point out that the radiolar appendages of the dorsal lips do not have an internal skeleton, and that dorsal pinnular appendages can be either present or absent. Furthermore, Fitzhugh emphasized the implications of those findings for the determination of cladistic relationships among Sabellinae genera.

Three species of Megalomma have been described from the Grand Caribbean: M. bioculatum (Ehlers, 1887), M. lobiferum (Ehlers, 1887), and M. heterops Perkins, 1984, all of them from Florida.

Megalomma heterops Perkins, 1984

(Fig. 9)

Megalomma heterops Perkins 1984: 359-363, figs. 42-43.

Material: [FSBC] Florida: Paratypes, 27696, St. Lucie Co., Hutchinson I., 27°20'24”N, 83°13’04”W, Sta. 4, 12.1 m, Coll. R. Gallagher and M. Hollinger, 15 Sept. 1971, Id. T. H. Perkins (1). [FSBC] I 27698, Hutchinson I., St. Lucie Co., 27°21’23”N, 80°13’24”W, 11.2 m, Coll. R. Gallagher and M. Hollinger, 15 Sept. 1971, Id. T. H. Perkins (1). [USNM] Florida: Paratypes, 067953, Sta. J, 20 miles W of Sanibel I., 26°24’N, 82°28’W, R/V Hernán Cortés, Coll. Presley, 11 May 1966, 18 m, Id. T. H. Perkins (5). [ECOSUR] Mexican Caribbean: I. Cozumel, Coll. S. I. Salazar, 24 Apr. 2001, coral (2). Cayo Valencia, Coll. M. S. Jiménez, 29 Apr. 1983 (1). DIF Puerto Aventuras, Coll. S. I. Salazar, 21 Mar. 1992 (1). Xcacel, Coll. S. I. Salazar, 3 Apr. 1992 (1). Ana y José, Coll. S. I. Salazar, J. R. Bastida and J. Ruíz, 11 Feb. 2001 (1). San Felipe, Coll. J. R. Bastida, 19 Feb. 1999 (3). Xahuayxol, Coll. S. I. Salazar, L. F. Carrera, 1 June 1997 (3). [LACM-AHF] British Virgin Is.: Vc 0566, Guana (1). [UMML] Florida: 22.250, Pennekamp Site 5, Florida Reef Tract, Grecian Rocks, July 1980 (1). [USNM] 067955, Off W coast, middle ground Florida, Sta. 151, 28°32’02”N, 84°18’36”W, 05 Oct. 1978, 25~27 m, on Madracis decactis, Id. T. H. Perkins (3). [USNM] Puerto Rico: 42791, J2 1963, 7.6 m, Coll. Hulings and Feray, muddy sand, Id. N. Foster. [USNM] 53233, Guayanilla Bay, Coll. T. Morgan, 28 Oct. 1971, Thalassia bed, 1.2 m, Id. T. H. Perkins.

Description (variations observed in paratypes given in parentheses): Body pale (in preserved material), 4~47 (4.5~27) mm long, anterior margin of segment 1 with paired purple spots dorsally below collar, paired spots on chaetigers 1~3 dorsal to notopodia. Branchial crown long, with a row of pigmented spots on palmate membrane, narrow purple bands on outer and lateral surfaces of radioles, not extending to pinnules, and 13~15 pairs of radioles alternating 5 or 6 purplish-yellow bands (in preserved material). Radioles with subterminal compound eyes, 1st pair on dorsalmost radioles enlarged with distinct ommatidia; other eyes much smaller, gradually decreasing in size ventrally, with smooth surface and ommatidia visible only under high magnification of light microscope; dorsalmost radioles subdistally enlarged, with short tips extending beyond eyes (Fig. 9C-E). Radiolar skeleton with 8 cells in cross-section surrounded by moderately thick sheath and columnar epithelium (Fig. 9F). Dorsal lips with dorsal radiolar appendages and dorsal pinnular appendages, pigmented on upper surface. Ventral lips with parallel lamellae, beginning between ventral lappets of collar on anterior margin of collar, extending anteriorly and dorsally to dorsal lips. Collar not fused to fecal groove (Fig. 9A), pockets present. Collar with 1 pair of distally rounded, overlapping, ventral lappets, which extend beyond beginning of radioles (Fig. 9B). Thorax with 8 chaetigers, 1~5 (3.5~9) mm long, and 0.5~2.5 (0.5~6.5) mm wide. Ventral shield of segment 1 entire, with rounded anterolateral corners and slight anteromedial indentation, twice as long as other shields and about the same width, ventral shields rectangular, not indented by tori (Fig. 9B). Tori gradually shortening posteriorly. Superior thoracic notochaetae, elongate narrowly hooded. Inferior thoracic notochaetae broadly hooded (Fig. 9G, H), arranged in 2 or more transverse rows. Thoracic uncini avicular, with 16 equal-sized teeth in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), breast well developed, manubrium long (Fig. 9I). Companion chaetae with roughly symmetrical tips as teardrop-shaped membranes. Abdomen with 36~98 (26~92) chaetigers. Abdominal neurochaetae in 2 transverse rows: anterior and posterior rows on anterior chaetigers with elongate, narrowly hooded and modified, elongate, narrowly hooded chaetae, respectively. Abdominal uncini avicular, with 13 equal-sized teeth in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), breast well developed, manubrium short (Fig. 9J). Pygidium rounded with irregularly arranged eyespots (Fig. 9K). Tubes with pieces of shells and coral fragments of different sizes, posterior end translucent amber.

Remarks: Perkins (1984) described M. heterops from Florida, and it corresponds to the group 2A (Table 1) designated by Knight-Jones (1997). Perkins indicated that there are 2 similar forms of M. heterops from North Carolina, Florida, and Venezuela (sp. A), and from Puerto Rico (sp. C), both with reduced flat eyes. Specimens of Megalomma sp. C., are in poor condition [USNM 42791 (1 incomplete) and USNM 53233 (1 complete)]; they have some rounded pigmented areas in the 3rd dorsal pair of the radioles, and distinct ommatidia in the eyes of the 2nd and 3rd dorsalmost pairs. Megalomma sp. A is formally described as M. perkinsi sp. nov. (see below).

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, pockets present	on most radioles	present	M. lobiferum	Florida
2A	Not fused to fecal groove, pockets present	on most radioles	absent absent absent	M. heterops M. neapolitanum M. perkinsi sp. n.	Florida Italy Florida
2B	Not fused to fecal groove, pockets absent	dorsalmost pair of radioles	absent present present absent	M. bioculatum M. pigmentum Megalomma sp. 1 Megalomma sp. 2	Florida Baja California Venezuela Florida

Megalomma lobiferum (Ehlers, 1887)

(Fig. 10)

Branchiomma lobiferum Ehlers 1887: 254-259, pl. 53, figs. 10-15.-- Hoagland 1919: 577.-- Treadwell 1924a: 18; 1939: 291-292, fig. 105.-- Fauvel 1953: 17.

Megalomma lobiferum.-- Johansson 1927: 132.-- Perkins 1984: 354-357, figs. 39, 40.

Material: [ECOSUR] Mexican Caribbean: I. Contoy, Punta Sur, Coll. S. I. Salazar, 2 Mar. 2001 (1). Veracruz, I. Verde, Coll. S. I. Salazar, 5 Aug. 1985 (1). Panama: Fuerte Sherman, Colón, Coll. S. I. Salazar, 2 June 2002, T=30, S=30/00 (1). [LACM-AHF] British Virgin Is.: AF00-59, Vc 0634, Guana, Beef I., Trellis Bay, Coll. K. Fitzhugh, 12 July 2000, tubes taken from coral rubble (2). [LACM-AHF] AF00-59, Vc 0565, Beef I., Trellis Bay, directly off airport, Coll. K. Fitzhugh, 12 July 2000 (1). [USNM] Panama: 073019, Galeta reef, Coll. A. A. Reimer, Id. T. H. Perkins, Apr. 1982 (6).

Description: Body 15~36 mm long. Branchial crown purple, 4~14 mm long with 15~27 pairs of radioles, cross-banded with yellowish-orange pigment extending to pinnules (in preserved material). Subterminal compound eyes on all radioles, moderately large on dorsalmost pair, gradually decreasing in size laterally and ventrally. Except for size, all eyes similar, well defined, spherical, with distinct ommatidia. Radiolar skeleton with 4 or more rows of cells in cross-section. Branchial crown with long fused region. Dorsal lips long, with dorsal radiolar appendages, with margin of upper lamellae fused proximally to base of basal pinnules of dorsalmost pair of radioles. Ventral lips short, rounded, with parallel lamellae. Caruncle above mouth, anterior to and in between dorsal lips (Fig. 10B). Dorsal margin of collar fused to fecal groove (Fig. 10A) forming deep pockets. Ventral lappets short, rounded, overlapping (Fig. 10B). Ventral shield of segment 1 up to twice as broad as long, convex laterally, straight to convex anteriorly (Fig. 10C); following shields concave, rounded laterally around tori. Thorax 3~6 mm wide, with 8 chaetigers. Collar with long, narrowly hooded chaetae (Fig. 10G-I). Superior thoracic notochaetae, elongate narrowly hooded. Inferior thoracic notochaetae broadly hooded, arranged in 2 or more transverse rows (Fig. 10J). Thoracic uncini avicular with several rows of equal-sized teeth in lateral view, covering 1/2 of extension of main fang ([not sure of the meaning?]a number of teeth not visible under light microscopy), breast well developed, manubrium long (Fig. 10N). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 10D-F). Abdomen with 43~84 chaetigers. Abdominal neurochaetae in 2 transverse rows: anterior and posterior rows on anterior chaetigers with elongate, narrowly hooded and modified, elongate, narrowly hooded chaetae, respectively (Fig. 10K, L). Abdominal uncini avicular, with 13 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (visible under light microscopy), breast well developed, manubrium short (Fig. 10M).

Remarks: The most-distinctive character in M. lobiferum is the “caruncle”; this organ also occurs in M. pigmentum and Megalomma sp. 1 (Table 1). Megalomma lobiferum differs of M. pigmentum and Megalomma sp. 1 in the occurrence of subterminal eyes on most radioles, dorsal collar margins fused to the fecal groove, and the presence of pockets.

Megalomma perkinsi sp. nov.

(Fig. 11)

Megalomma sp. A Perkins 1984.

Etymology: This species is named in honor of Thomas Perkins (Florida Marine Research Institute, St. Petersburg, FL) in recognition of his important contribution to the knowledge of polychaetes in general, and especially of sabellids.

Material: [USNM] Florida: Holotype 53976, Cape Lookout, NC, Coll. S. L. Gardiner, 16 Apr. 1976, intertidal sand mixed with gravel and shell fragments 129, Id. S. L. Gardiner. [USNM] Paratype 53977, Cape Lookout, NC, 28 July 1976, Coll. C. Jenner, intertidal sand mixed with gravel and shell fragments 146, Id. S. L. Gardiner as M. lobiferum.

Description (variations observed in paratype given in parentheses): Specimen large, 107 (92) mm long. Branchial crown 12 mm long, with 22 (23) pairs of radioles with 3 purplish-brown bands at 1st 1/2 of crown, dorsalmost pair longer and pale (in preserved material). Radioles with subterminal compound eyes on most radioles, flat, those from dorsalmost pair bigger than others (Fig. 11H), surrounding almost covering tip dorsally, small ventrally. Radiolar skeleton with 4 or more rows of cells in cross-section (Fig. 11G). Dorsal lips long with broad base (Fig. 11C), with orange pigment laterally and radiolar pinnular appendages. Ventral lips with parallel lamellae. Dorsal margins of collar not fused to fecal groove, with 2 concave lobes forming deep pockets (Fig. 11B); collar with a small lateral indentation (Fig. 11A). Ventral shield of segment 1 as wide as long (Fig. 11C), following shields rectangular. Thorax 11 (10) mm long, 4.5 (5) mm wide, with 8 chaetigers. Superior thoracic notochaetae, elongate narrowly hooded. Inferior thoracic notochaetae broadly hooded (Fig. 11K), arranged in 2 or more transverse rows. Thoracic uncini avicular, with 22 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang, breast well developed, manubrium long (Fig. 11D). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 11E-F). Abdomen with 66 (104) chaetigers. Abdominal neurochaetae in 2 transverse rows: anterior and posterior rows on anterior chaetigers with elongate, narrowly hooded and modified, elongate, narrowly hooded chaetae, respectively (Fig. 11J). Abdominal uncini avicular, with 18 equal-sized teeth in lateral view, covering 1/2 of extension of main fang, breast well developed, manubrium short (Fig. 11I). Maturating oocytes in abdomen (chaetiger 11). Tube with large shells and stone fragments.

Remarks: Megalomma perkinsi sp. nov. belongs to group 2A proposed by Knight-Jones (1997), which is defined by the dorsal margins of collar not being fused to the fecal groove, the presence of dorsolateral collar pockets, and the occurrence of eyes on most radioles. The other species belonging to this group include M. heterops Perkins 1984 from Florida and M. neapolitanum (Claparède, 1868) from Italy (Table 1).

Based on the description and illustration provided by Claparède (1868: pl. XXII, fig. 5), the anterior dorsal margin of the collar in M. neapolitanum is of even height, forming 2 small pockets, compound radiolar eyes are hemispherical with big ommatidia (more than in M. heterops), covering all radiolar tip. In M. perkinsi, the anterior dorsal margin of the collar is high; forming 2 big, deep pockets, compound radiolar eyes are rounded with small ommatidia, not covering the radiolar tip. Megalomma heterops differs from M. perkinsi because in the latter species, the anterior margin of the collar has a lateral depression, reduced and flattened radiolar compound eyes, with distinct ommatidia only in the 2nd and 3rd dorsalmost pairs, and rounded pigmented areas in 3rd dorsal pair of radioles.

Megalomma sp. 1

Megalomma pigmentum.-- Perkins 1984: 357-359, fig. 41C (non Reish).

Material: [USNM] Venezuela: 57945, Bahía de Mochima, Cumaná (7), Coll. Edwards, 22 June 1971, 6~10 m, in calcareous sand, Id. T. H. Perkins, 1981.

Description: Five complete, pale (in preserved material), small specimens, 4~23 mm long. Crown 3~7 mm long, with 12 pairs of radioles, base and interradiolar spots orange on preserved material. Dorsal lips with radiolar and pinnular appendages. Ventral lips with parallel lamellae. Ventral lappets lanceolated. Dorsal margins of collar not fused to fecal groove, pockets absent, eyes only present on dorsalmost pair of radioles. Caruncle thin, above the mouth, anterior to and in between dorsal lips. First ventral shield divided into 2 parts, basal part larger, remaining thoracic shields rectangular. Thorax 2.5~5.5 mm long, 1~1.5 mm wide, with 8 chaetigers. Inferior thoracic notochaetae broadly hooded, arranged in transverse rows. Companion chaetae with roughly symmetrical tips. Abdomen with 14~43 chaetigers. Abdominal neurochaetae with elongate, narrowly hooded chaetae.

Remarks: Perkins (1984) recorded M. pigmentum Reish from Florida and Venezuela; he assumed that the main difference between Atlantic specimens and the type material was the color. Nevertheless, the material from the Grand Caribbean region (USNM), identified by Perkins as M. pigmentum (non Reish), corresponds to 2 undescribed species, one from Venezuela (Megalomma sp. 1), the other from Florida (Megalomma sp. 2 see below). The 3 species correspond to group 2B according to the classifictaion proposed by Knight-Jones (1997) (Table 1).

In Megalomma sp. 1 the 1st shields are similar to those of M. pigmentum Reish; however, Megalomma sp. 1 has a caruncle, a structure previously recorded only for M. lobiferum, and in this study for M. pigmentum Reish. Megalomma bioculatum has a collar with triangular ventral lappets, and a quadrangular ventral shield on segment 1 with an anterior M-shaped margin, and lacks the caruncle, while M. pigmentum Reish has a collar with lanceolated ventral lappets, the ventral shield of segment 1 is divided into 2 asymmetrical parts, and there is a caruncle, as corroborated by topotypical specimens [ECOSUR: Bahía San Quintín, Coll. L. E. Calderón, 8 Dec. 1981].

We included this species in the present paper in order to facilitate and encourage further research, because the presence of a caruncle in species of Megalomma should be more frequently included in taxonomic studies since it provides additional information for investigation of phylogenetic relationships.

Megalomma sp. 2

Megalomma pigmentum Perkins 1984: 358, fig. 41A, B (non Reish).

Material: [USNM] Florida: 45691 (7), Tampa Bay, Coll. J. L. Taylor, 1963, Id. T. H. Perkins, 1981.

Description (data of best specimen given in parentheses): Two complete specimens and 7 fragments. Body pink (in preserved material), 3~19 (24) mm long. Crown 4~6 (5.5) mm long, with 10 or 11 (10) pairs of radioles. Dorsal lips with dorsal radiolar appendages and dorsal pinnular appendages. Ventral lips with parallel lamellae. Ventral lappets lanceolated. Dorsal margins of collar not fused to fecal groove, pockets absent, eyes only present on dorsalmost pair of radioles. Caruncle absent. First ventral shield divided into 2 parts, basal part larger, remaining thoracic shields rectangular. Thorax 2~5.5 (3.5) mm long, 1~2 mm wide, with 7~9 (8) chaetigers. Inferior thoracic notochaetae broadly hooded, arranged in transverse rows. Companion chaetae with roughly symmetrical tips. Abdomen with 3~21 (60) chaetigers. Tube covered with transparent crystals.

Remarks: Megalomma sp. 2 is close to Megalomma sp. 1, the 1st ventral shield of both is divided into 2 pairs; nevertheless, they only differ because Megalomma sp. 2 lacks a caruncle (Table 1). Drawings of Perkins (1984: 358, fig. 41A, B) illustrate the anterior region of Megalomma sp. 2 very clearly. Descriptions of Megalomma sp. 1 and Megalomma sp. 2 are brief and without drawings; however, the material is available at the USNM.

Notaulax Tauber, 1879

Perkins (1984) made a preliminary revision of Hypsicomus and Notaulax, in which he redescribed some species, moved most species of Hypsicomus to Notaulax, described 2 new species, and provided a key for the identification of species of Notaulax. The genus is characterized by the arrangement of collar chaetae, the presence of superior thoracic spinelike notochaetae, and the long branchial base flanges (Fitzhugh, 1989). Six species have been described from the Grand Caribbean: N. bahamensis Perkins, 1984 and N. paucoculata Perkins, 1984 from the Bahamas; N. circumspiciens (Ehlers, 1887), and N. midoculi (Hoagland, 1919) from Florida; N. nudicollis (Kroyer, 1856) from St. Thomas, and N. occidentalis (Baird, 1856) from St. Vincent.

Notaulax bahamensis Perkins, 1984

(Fig. 12)

Notaulax bahamensis Perkins 1984: 348, figs. 35, 36.

Material: [ECOSUR] Mexican Caribbean: I. Cozumel, Chankanaab, Coll. S. I. Salazar, 2 Apr. 1992 (1). [USNM] Holotype 062082, Lucaya, Grand Bahama I., Hydrolab, off Bell Channel, 26°33’00’’N, 78°34’00’’W, Coll. B. A. Vittor and T. S. Hopkins, 28 Jan. 1974, on coral.

Description: Body pale in preserved materials, 34 mm long. Crown with 4 brown bands (in preserved material), 2 in palmate membrane, another immediately above it, and the last at midlength of free parts of radioles. Crown 15 mm long, with 11 or 12 pairs of radioles united proximally by palmate membrane for about 40% of their length above basal lamina, flanged from palmate membrane to tips (Fig. 12C) except for a small part of region of simple eyespots (Fig. 12D). Basal lamina very long, as long as palmate membrane, eyes in oval groups (30), separated from palmate membrane for about 1/3 of its length; dorsal and ventral margins of basal lamina flanged, from bases of lobes to origins of dorsalmost and ventralmost radioles. Radiolar skeleton with 4 or more rows of cells in cross-section. Simple radiolar eyes present above palmate membrane (Fig. 12D). Dorsal lips with radiolar appendages; pinnular appendages absent. Ventral lips and parallel lamellae present. Collar as a single lobe, beginning near dorsal midline at margin of posterior peristomial ring and extending anteriorly, almost straight laterally, with convex ventral margin (Fig. 12A, B); segment 1 much longer than following segments (Fig. 12A, B), ventral shield subtriangular. Chaetae on segment 1, as single, elongate row of spinelike chaetae (Fig. 12E, F) on posterior 1/3 of collar, fascicles longitudinal to oblique (Fig. 12A). Thorax with 8 chaetigers, 5 mm long, 2.8 mm wide. Thoracic superior notochaetae spinelike (Fig. 12K, L); inferior notochaetae paleate, arranged in 2 transverse rows, symmetrical (Fig. 12I, J). Thoracic uncini avicular, with 16 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), hood absent, breast well developed, manubrium medium (Fig. 12G). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 12M). Abdomen with 108 chaetigers. Abdominal neurochaetal fascicles with 2 transverse rows of chaetae: anterior row on anterior and posterior abdominal chaetigers paleate, mucroate with small teeth at base of mucro (Fig. 12O, P); posterior row on anterior abdominal chaetigers elongate, narrowly hooded; posterior row in posterior abdominal chaetigers modified, elongate, narrowly hooded (Fig. 12N). Abdominal uncini avicular, with 14 equal-sized teeth above main fang in lateral view, covering 3/4 of extension of main fang (seen under light microscopy), breast well developed, manubrium short (Fig. 12H). Pygidial eyespots present.

Remarks: Notaulax bahamensis and N. nudicollis share a roughly similar arrangement of eyespots, but in the 1st species, the radioles are flanged except in a small portion on the eye band, while in the latter species, the radioles are completely flanged. Further, N. bahamensis has a very long collar (longer than the next 3 segments), while the collar of N. nudicollis is as long as the next 2 segments. The holotype is damaged, the crown is almost separated from the body, and some radioles were already removed.

Notaulax midoculi (Hoagland, 1919)

(Fig. 13)

Parasabella midoculi Hoagland 1919: 579, pl. 31, figs. 10-14, pl. 32, figs. 1, 2.

Parasabella sulfurea.-- Treadwell 1924a: 18 fide Perkins 1984: 343.

Hypsicomus midoculi.-- Johansson 1927: 141.

Material: [ECOSUR] Mexican Caribbean: I. Contoy, Boca Norte, Pueblo Viejo, Coll. M. A. Tovar, 1 Mar. 2001, in sponge on dead coral (1); Punta Sur, Coll. M. A. Tovar, 2 Mar. 2001 (4). I. Cozumel, SEDENA, Coll. A. Medina, 24 Mar. 2001 (4); Playa Azul, Coll. L. E. González, 25 Mar. 2001 (1); Chankanaab, Coll. S. I. Salazar, 2 Apr. 1992, QR-7 (1); Muelle Puerta Maya, Coll. A. Medina, 26 Mar. 2001 (2). Majahual Norte, Coll. S. I. Salazar, 18 Mar. 2001, on dead coral (1). [UMML] Florida: 22.166, Reef Tract, Margot Fish Shoal, 16 Feb. 1962, MFS (18 l), Coll. Jones, Work, Bayer, and Ebbs (1). [USNM] Barbados: 020304, Bathsheba, Coll. Univ. of Iowa, Summer, 1918, Barbados-Antigua Expedition, Id. T. H. Perkins, 1980 (1).

Description: Body 18~28 mm long. Crown 8~14 mm long, with 10~16 pairs of radioles united above basal lamina by a palmate membrane, extending for 1/6 of their length. Radioles with purple (in preserved material) very long basal lamina (Fig. 13A, B), followed by a white band (10 pinnules), at midlegth an orange band (13 pinnules), another white band (5 pinnules), and a pale-orange band (6 pinnules), the latter with 2 groups of 9~14 small simple eyespots at both sides of radiole (Fig. 13C). Radioles slightly flanged near tips. Radiolar skeleton with 6 rows of cells in cross-section (Fig. 13D). Dorsal lips with radiolar appendages but without dorsal pinnular appendages. Ventral lips and parallel lamellae present. Collar with rounded ventral lappets (Fig. 13B). Shield of segment 1 not well defined; following segments with trapezoidal ventral thoracic shields, lateral sides strongly indented by tori (Fig. 13B). Thorax 3~4 mm long, 2~3 mm wide, with 8 chaetigers. Single, elongate, longitudinal to oblique row of spinelike chaetae on segment 1. Superior thoracic notochaetae spinelike (Fig. 13M), inferior notochaeta paleate, arranged in 2 transverse rows, with symmetrical and asymmetrical chaetae (Fig. 13K, L). Thoracic uncini avicular, with 22 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), hood absent, breast well developed, manubrium medium (Fig. 13E). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 13G-J). Abdomen with 22~41 chaetigers. Abdominal neurochaetae fascicles in 2 transverse rows of chaetae: anterior row on anterior and posterior abdominal chaetigers paleate, with long mucro (Fig. 13N, O) with small teeth at base of mucro; posterior row on anterior abdominal chaetigers elongate, narrowly hooded; posterior row on posterior abdominal chaetigers modified, elongate, narrowly hooded (Fig. 13P). Abdominal uncini avicular, with 19 equal-sized teeth above main fang in lateral view, covering 3/4 of extension of main fang (seen under light microscopy); breast well developed, manubrium short (Fig. 13F).

Remarks: Perkins (1984) reviewed a specimen from Barbados in which the ventral margin of the collar had a slight incision, and the ventral lobes were not very well defined; in his figures, the shield of segment 1 is distinct and rectangular, and the other shields are rectangular with concave lateral sides. Specimens identified in this study as N. midoculi have a collar that is well incised ventrally, forming 2 rounded lobes, and the ventral shield of segment 1 is indistinct, as originally illustrated by Hoagland (1919 fig. 10) and Perkins (1984 fig. 32H) from the holotype. The USNM specimen was collected in 1918, and has been reviewed by Treadwell (as Parasabella sulfurea), Hartman (as Hypsicomus circumspiciens), and Perkins. It is broken into 3 parts, has a transverse incision showing the coelom full of oocytes, and another dorsal incision from chaetiger 5 (thorax) towards the midbody.

Notaulax nudicollis (Krøyer, 1856)

(Fig. 14)

Sabella nudicollis Krøyer 1856: 30-31.

Protulides elegans Webster 1884: 325-326, pl. 11, figs. 63-74 fide Perkins 1984: 332.

Notaulax nudicollis.-- Perkins 1984: 331, figs. 25-28.

Material: [ECOSUR] Mexican Caribbean: DIF Puerto Aventuras, QR-4, Coll. S. I. Salazar, 21 Mar. 1992 (3). Contoy, Coll. L. F. Carrera and S. I. Salazar, 10 June 1999 in Ircinia (3); Faro, Coll. P. Salazar, 1 Mar. 2001 (1); Camping, 22 Feb. 1999 (1); Coll. S. I. Salazar, 1 Mar. 2001, in Strombus gigas (3); Punta Sur, 28 Feb. 2001 (2). Cancún, Punta Nizuc, Coll. S. I. Salazar and L. F. Carrera, 30 Aug. 1997 (2). I. Cozumel, Paraíso, Coll. S. I. Salazar, 5 June 1996 (1). Buenavista, E-25, Coll. S. I. Salazar and L. F. Carrera, 27 Sept. 1996 (3); R-5 (1); R-4 (1); E-24 (1). Majahual, Coll. M. A. Tovar, 10 Jan. 2001 (1). Río Indio, Coll. S. Frontana, 17 Mar. 2001 (1). San Felipe, Coll. J. R. Bastida and S. I. Salazar, 19 Nov. 1999 (1). Sol y Mar (2). Xahuayxol, Coll. S. I. Salazar and L. F. Carrera, 27 Sept. 1996, E-4 (2), 3 June 1998, C28 RC1 (2); 1 June 1997, A-46, in Strombus gigas (3); 27 June 1997, E2 (1); 19 Oct. 1997, B-45, Coll. S. I. Salazar and L. F. Carrera (1). [LACM-AHF] British Virgin Is.: BVI-99, LH0995, Guana, Sta. 035 (3). [UMML] Florida: 22.87, Straits off Florida, R/V Gerda, Sta. 1033, 24°36’N, 81°06’W, 42 m, 26 Feb. 1969, Coll. G-1033, Id. T. H. Perkins (6). [USNM] 52042, Big Pine Key, FL, Coll. D. Maynahan, 14 Aug. 1973, from rock wall of canal, 0.6~1.8 m, Id. T. H. Perkins (3), lot contains a brown crown, 1 thorax, 2 fragments and rest of tranlucid tubes. [USNM] 57933, Safe Harbor, Stock I., near Key West, FL, Sta. 7A, 2.4 m, 7 July 1970, Coll. R. Chester, Id. T. H. Perkins (2), lot contains 1 complete specimen, 1 incomplete (missing posterior segments), and 2 crowns. [ZMA] North Carolina: V. Pol. Bogue Sound, wooden piling at Fisheries Institute, Coll. P. Wagenaar Hummelinck, 28 Aug. 1963, 0~1 m (7).

Description: Body 12~24 mm long. Branchial crown 5~5.5 mm long with 5 brown to purple cross bands (in preserved material) and 10~14 pairs of radioles; radiolar eyes in elongate-oval groups of 30~70 eyes of each side, diminishing in number distally (Fig. 14C). Radiolar flanges beginning above eyes and extending to tips. Radiolar skeleton with 4 rows of cells in cross-section (Fig. 14D). Dorsal lips extending laterally in almost straight line, with radiolar appendages but without pinnular appendages. Ventral lips and parallel lamellae present. Dorsal and ventral margins with flanges. Dorsal margin of collar incised (Fig. 14A), ventral margin entire (Fig. 14B). Thorax with 8 chaetigers, 2.5 mm long and 1~1.5 mm wide. Ventral shield of segment 1 rectangular to trapezoidal (Fig. 14B). Chaetae of segment 1 in a longitudinal to oblique single elongate row of spinelike chaetae. Superior thoracic notochaetae spinelike. Inferior thoracic notochaeta paleate, nearly symmetrical (Fig. 14H-J), arranged in 2 rows. Thoracic uncini avicular with 17 or 18 equal-sized teeth above main fang in lateral view (seen under light microscopy), covering 1/2 of extension of main fang, hood absent, breast well developed, manubrium medium (Fig. 14E). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 14G). Abdomen with 29~115 chaetigers. Abdominal neurochaetae in 2 transverse rows of chaetae: anterior row paleate with short mucro on anterior and posterior abdominal chaetigers (Fig. 14L); posterior row on anterior abdominal chaetigers elongate, narrowly hooded; posterior row on posterior abdominal chaetigers modified, elongate, narrowly hooded (Fig. 14K). Abdominal uncini avicular, with 13 equal-sized teeth above main fang in lateral view (seen under light microscopy), covering 3/4 of extension of main fang, breast well developed (Fig. 14F), manubrium short.

Remarks: Perkins (1984) recorded N. nudicollis from North Carolina, Florida, Puerto Rico, the Virgin Islands, Brazil, Western Africa, and the Mexican Pacific; he stated: “Minor differences between specimens from the Caribbean Sea and adjacent areas and those from West Africa, Brazil, and Western Mexico are attributed to geographic separation but are not considered to be sufficient to indicate speciation”. Nevertheless, the morphology of the collar and the distribution of the eyes on the radioles were variable in his material; further, Perkins (1984: 332) synonymized Sabella torquata Grube, 1877, described from Western Africa, with N. nudicollis (Krøyer, 1856) described from St. Thomas. However, these species have remarkable differences: the ventral, anterior collar margin of the syntypes of S. torquata is triangular and medially indented (Perkins, Fig. 25F), and there are groups of more than 40 eyes, separated from the palmate membrane by a distance shorter than that in N. nudicollis (Perkins 1984, fig. 25D). Perkins mentioned that these characters were similar in 1 specimen from the Mexican Pacific (Zihuatanejo), but with up to 30 eyes. However, this record is questionable, since in Perkins' drawings (Perkins 1984, fig. 26C, D), the ventral collar is complete, of the same width all along its extension, and the dorsal margin of collar is more incised than in the holotype of N. nudicollis. Further, the material of Zihuatanejo (USNM 41528, Coll. Klawe, 6 Sept. 1968, on dead coral, Id. T. H. Perkins) consists only of a trunk fragment in a very poor condiction missing the crown, collar, and pygidium.

Two specimens from Bahía de los Ángeles, Baja California (ECOSUR: 23 Aug. 1987, Coll. SISV) are identical to Perkins’ drawings for the Zihuatanejo specimen; but we considered them as belonging to a different species. The number of eyes is a character which changes with age, but the pattern of radiolar bands is constant and useful to separate species. Thus, some species have bands near the basal membrane, others at radiolar midlength, and still others near the distal part of the radioles. Perkins’ (1984) drawings of N. nudicollis from Florida (Perkins 1984, fig. 27A-G), show at least 2 different forms, differentiated by the ventral morphology of the collar: one with a wide, entire, distally rounded anterior margin (Perkins 1984, fig. 27D), and the other with the collar segment wide, incised, with a heart-shaped anterior margin (Perkins 1984, fig. 27F). Thus, they may be different species, and some additional differences can be found by studying their chaetae.

Notaulax occidentalis (Baird, 1865)

(Fig. 15)

Sabella occidentalis Baird 1865: 159, pl. 5, figs. 7, 8.

Sabella alba Treadwell 1917: 266-267, pl. 3, figs. 10-15; 1939: 294, fig. 108 fide Perkins 1984: 339.

Parasabella sulfurea Treadwell 1917: 267, pl. 3, figs. 16-23 fide Perkins 1984: 339.

Hypsicomus purpureus Treadwell 1924: 20-21, pl. 2, figs. 30-33 fide Perkins 1984: 339.

Notaulax occidentalis.-- Perkins 1984: 339-342, figs. 29, 30.

Material: [ECOSUR] Mexican Caribbean: I. Cozumel, Paraíso, Coll. S. I. Salazar, 5 June 1995 (1); Chankanaab, Coll. S. I. Salazar, 2 Apr. 1992 (2). I. Contoy, Punta Sur, Barlovento, Coll. S. I. Salazar, 2 Mar. 2001 (3). DIF Puerto Aventuras, QR-4, Coll. S. I. Salazar, 21 Mar. 1992 (2). Majahual Norte, Coll. M. A. Tovar, 19 Jan. 2001 (2). Cancún, Punta Nizuc, Coll. S. I. Salazar and L. F. Carrera, 1 Sept. 1997 (2); 31 Aug. 1997, rocks 4 (2). Buenavista, R4, Coll. S. I. Salazar and L. F. Carrera, 27 Sept. 1996 (1). Venezuela: Turpialito, Cumaná, Coll. M. Liñero, 22 Feb. 2002, in Millepora (3). [LACM-AHF] British Virgin Is.: Vc 0675, Guana (1); AF00-49, Vc 0517 (1); AF00-65, Vc 0791 (1); AF01-67, Vc 0592 (1). [UMML] Florida: 22.761, Reef Tract, Margot Fish Shoal, Red Head, 16 Feb. 1962 (1), Coll. Jones, Work, Bayer, and Ebbs (1). [UMML] British Virgin Is.: 22.98, St. John, Great Cruz Bay, Virgin Is., Coll. Thomas, 15 Dec. 1958 (2). [USNM] Puerto Rico: 16218, Reef at Ponce, 1898-1899, Coll. Str. Fish Hawk, Donor US Fish. Comm., Id. T. H. Perkins (2). [USNM] Bahamas: 062083, Lucaya, Grand Bahama I., Hydrolab, off Bell Channel, 26°33’00”N, 78°34’00’’W, Coll. B. A. Vittor and T. S. Hopkins, 28 Jan. 1974, 15 m, Id. T. H. Perkins (1). [USNM] Panama: 073020, Galeta Reef, Coll. A. A. Reimer, 5 Oct. 1970, Id. T. H. Perkins (2).

Description: Body brown with a brown band along dorsum, dark brown ventrally, 22~57 mm long. Base of crown purple to brown (in preserved material). Palmate membrane darker than body, with 4~8 cross bands of purple to brown pigmentation, pigment extending into pinnules. Branchial crown 9~18 mm long with 17~30 pairs of radioles; palmate membrane extending for about 1/3 of total length of branchial crown, with basal lamina about 2/3 as long as palmate membrane; radiolar flanges extending until radiolar tips (Fig. 15C). Flanges beginning at about midlength of free parts, progressively widening distal; bare tips, flat, long, about the length of 2 or 3 pinnules, tongue-shaped (Fig. 15C); eyes in single rows, 10~30, beginning just above palmate membrane (Fig. 15J). Radiolar skeleton with 4 cells in cross-section. Dorsal lips longer than palmate membrane (1~2 mm), with dorsal radiolar appendages but without pinnular appendages. Ventral lips and parallel lamellae present. Branchial lobes very long, dorsal and ventral margins with flanges extending from base of lobes to origin of dorsal and ventralmost radioles. Collar bilobed (Fig. 15A), with triangular and distally rounded ventral lobes. Thorax 3.5~5.5 mm long with 8 chaetigers, 2.5~4 mm wide. Ventral shield of segment 1 rectangular, divided into 2 asymmetrical parts (Fig. 15B), following ventral shields trapezoidal, with broader anterior margin and lateral margins indented by tori. Segment 1 with fascicles modified as a single, elongate, longitudinal to oblique row of spinelike chaetae (Fig. 15K). Superior thoracic notochaetae spinelike (Fig. 15N). Inferior thoracic notochaetae paleate, asymmetrical (Fig. 15L, M), arranged in 2 transverse rows. Thoracic uncini avicular, with 18 equal-sized teeth above main fang in lateral view (seen under light microscopy), covering 1/2 of extension of main fang (Fig. 15F), hood absent, breast well developed, manubrium medium (Fig. 15D). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 15G-I). Abdomen with 78~162 chaetigers. Abdominal neurochaetae in 2 transverse rows of chaetae: anterior row on anterior and posterior abdominal chaetigers paleate, with long mucro (Fig. 15O-P); posterior row on anterior abdominal chaetigers elongate, narrowly hooded; posterior row on posterior abdominal chaetigers modified, elongate, narrowly hooded (Fig. 15Q). Abdominal uncini avicular, with 24 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (seen under light microscopy); breast well developed, manubrium short (Fig. 15E). Posterior abdominal segments containing coelomic-madurating oocytes. Pygidial eyes present.

Remarks: Both N. occidentalis and N. circumspiciens (Ehlers, 1887) have 2 rows of eyes per radiole; the 1st species has up to 30 eyes per radiole, while the 2nd has up to 50. In N. circumspiciens (described from Carysfort Reef, FL), the collar is divided into 4 lobes, and the ventral lobes are rounded and longer than those in N. occidentalis; in lateral view; the collar of N. circumspiciens has a shorter indentation than that in N. occidentalis.

Notaulax paucoculata Perkins, 1984

(Fig. 16)

Notaulax paucoculata Perkins 1984: 345, figs. 33, 34.

Material: [ECOSUR] Mexican Caribbean: I. Cozumel, Ixchen, Coll. S. I. Salazar, 4 June 1995 (1); SEDENA, Coll. L. F. Carrera, 24 Mar. 2001, in Ircinia strobilina (1); Coll. A. Medina (3). DIF Puerto Aventuras, Coll. J. R. Bastida and S. I. Salazar, 28 Nov. 1999 (1). Punta Herradura, Coll. L. F. Carrera and S. I. Salazar, 28 Oct. 1997, Com B90 (1). San Felipe, Coll. J. R. Bastida and S. I. Salazar (1). [UMML] Mexican Caribbean: 22.766, I. Cozumel, SEDENA, Coll. A. Medina, March 24, 2001 (3). [USNM] Holotype 062077, Lucaya, Grand Bahama I., Hydrolab, off Bell Channel, 26°33’00’’N, 78°34’00’’W, Coll. S. A. Earle and A. Hurley 8 July 1974, Sta. 8-1, 41.1 m.

Description: Body medium-sized, 12 mm long. Branchial crown very long at 11 mm with 8 pairs of radioles, 3 cross-banded with purplish-brown pigment (in preserved material). Radioles united proximally above basal lamina by palmate membrane for 2/5 of their length, flanged above palmate membrane for their its length. Simple radiolar eyes present above palmate membrane, 4 or 5 eyes in a small group on lateral borders of radioles (Fig. 16C), at distal end of middle color band. Radiolar skeleton with 4 rows of cells in cross-section. Dorsal lips with radiolar appendages, but without pinnular appendages. Ventral lips and parallel lamellae present. Collar long, covering origin of branchial crown, deeply incised dorsomedially (Fig. 16A). Thorax with 8 chaetigers, 3 mm long, 1.5 mm wide. Chaetae from segment 1 in a single, elongate, longitudinal to oblique row of spinelike chaetae (Fig. 16D, E). Ventral shield of segment 1 rectangular, divided into 2 asymmetrical regions, basal one well defined (Fig. 16B), following shields trapezoidal, indented by tori. Superior thoracic notochaetae spinelike (Fig. 16K). Inferior thoracic notochaeta paleate, asymmetrical and symmetrical, arranged in 2 transverse rows (Fig. 16I, J). Thoracic uncini avicular, with 18 equal-sized teeth above main fang in lateral view (seen under light microscopy), covering 1/2 of extension of main fang, hood absent, breast well developed, manubrium medium (Fig. 16G). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 16F). Abdomen with 31~42 chaetigers. Abdominal neurochaetal fascicles in 2 transverse rows of chaetae: anterior row throughout abdomen with symmetrical paleate chaetae with long denticulate mucro (Fig. 16L, M); posterior row on anterior abdominal chaetigers with elongate, narrowly hooded chaetae; posterior row on posterior abdominal chaetigers with modified, elongate, narrowly hooded chaetae (Fig. 16N). Abdominal uncini avicular, with 17 equal-sized teeth above main fang in lateral view, covering 3/4 of extension of main fang (seen under light microscopy), breast well developed, manubrium short (Fig. 16H). Pygidium with eyespots.

Remarks: Notaulax paucoculata has few radiolar eyes, and the dorsal margin of the collar is deeply incised. The holotype is incomplete; the crown and posterior end of the body are in poor condition.

Perkinsiana Knight-Jones, 1983

Perkinsiana was erected to accommodate 11 species which did not reasonably fit into Potamilla Malmgren,1866, Potamethus Chamberlin, 1919, or Demonax Kinberg, 1867 (Knight-Jones 1983). Fitzhugh (1989) found that Perkinsiana could not be diagnosed in terms of apomorphy. Only 1 species of Perkinsiana has been described from the Grand Caribbean: P. fonticula from Puerto Rico, recorded below.

Perkinsiana fonticula (Hoagland, 1919)

(Fig. 17)

Parasabella fonticula Hoagland 1919: 579, pl. 31, figs. 3-9.

Potamilla fonticula.-- Fauchald 1977: 62, fig. 13c-k.

Perkinsiana fonticula.-- Knight-Jones 1983: 282, fig. 17A-L.

Material: [ECOSUR] Mexican Caribbean: Majahual, dead coral, Coll. S. I. Salazar and B. Trujillo, 7 June 2003 (1). DIF Puerto Aventuras, Coll. S. I. Salazar, 21 Mar. 1992 (5). Tulum, Punta Piedra, Coll. LEGE, 11 Feb. 2001 (3). I. Contoy, Punta Sur, Coll. S. I. Salazar, 2 Mar. 2001 (1). I. Cozumel, Puerta Maya, Coll. A. Medina, 26 Mar. 2001 (1).

Description: Body medium-sized, 22~30 mm long, 1 mm wide. Branchial crown supported by a deep cartilaginous base that joins 2 halves dorsally (Fig. 17B), 5 mm long, with 6~8 pairs of radioles fused at bases. Anterior margin of anterior peristomial ring low, posterior peristomial ring collar present (Fig. 17A, B). Dorsal lips tapered with outer lamella fused to slightly enlarged pinnules. Elongated triangular collar lappets (Fig. 17C), parallel lamellae present. Anterior margin of shield of segment 1 rounded, M-shaped (Fig. 17C). Thorax with 20 chaetigers. Thoracic fascicles very small with only 1 or 2 slender, narrowly hooded superior chaetae (Fig. 17F, G), and about 5 inferior paleate notochaetae with short mucro (Fig. 17D, E), arranged in 2 or more transverse rows. Thoracic tori with 6~9 avicular uncini, with 16 equal-sized teeth above main fang in lateral view, covering 1/2 of extension of main fang (seen under light microscopy), hood absent, breast well developed, manubrium long. Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Fig. 17I, J). Abdominal neurochaetae in 2 transverse rows of chaetae: anterior and posterior rows with elongate, broadly hooded chaetae throughout. Abdominal uncini avicular, with 19 equal-sized teeth above main fang in lateral view, covering 1/3 of extension of main fang (seen under light microscopy), breast well developed, manubrium long.

Remarks: The holotype of P. fonticula as specimens recorded in this study has 20 thoracic chaetigers. Fauchald (1977) recorded specimens from Panama with 14~22 chaetigers, but probably this variation is due to regeneration of the anterior end of the body or may depend on the age of the organisms.

Pseudobranchiomma Jones, 1962

Fitzhugh (1989) used 3 characters to separate Pseudobranchiomma from Branchiomma: the presence of radiolar flanges, dorsal lips without pinnular support, and abdominal fascicles with long chaetae. Knight-Jones (1994) indicated, however, that the last character is also present in Branchiomma; the presence or absence of the pinnular support in the dorsal lip is not very robust, since the holotype of P. emersoni has 1 lip with this appendage, while the other has a free end. In the same work, she indicated that P. odhneri and P. longa (Kinberg, 1867) have radiolar eyes (this feature was sufficient to emend the genus), and that radiolar serrations extend through a variable range, since they can run along the radiole, are restricted to the distal region, or are missing completely.

Nogueira and Knight-Jones (2002) suggested that a thorax with few chaetigers can be attributed to an imperfect regeneration after scissiparity; some records of Pseudobranchiomma species show variations of 4, 6, and 8 chaetigers. Also, they indicated that the external radiolar appendages of Pseudobranchiomma are only regular serrations of the radiolar flange, which differs from the paired stylodes of Branchiomma.

Knight-Jones and Giangrande (2003) divided 13 species of Pseudobranchiomma into 3 groups: (A) those with paired serrated flanges along all or most of the extensions of radioles, as in the type species P. emersoni; (B) those with such flanges only on the distal parts of the radioles; and (C) those with reduced or absent flanges (i.e., without serrations). Only 2 species of Pseudobranchiomma have been described from the Grand Caribbean: P. emersoni Jones, 1962 (Jamaica) and P. perkinsi Knight-Jones and Giangrande, 2003 from Cape Canaveral.

Pseudobranchiomma emersoni Jones, 1962

(Fig. 18)

Pseudobranchiomma emersoni Jones 1962: 198-201, figs. 115-124.

Material: [FSBC] Florida: I 23705, North Biscayne Bay, Sta. 28040349, 25°49’34”N, 80°08’20”W, 1 m, 22 Feb. 1979, Id. T. H. Perkins, Coll. Biosystems Research, Miami (1). [FSBC] I 43728, Sta. 32, 25°44’03”N, 80°12’05”W, 11 Oct. 1981, Id. H. D. Rudolph (3). [FSBC] I 43729, Sta. 4a, 25°49’34”N, 80°08’20”W, 1.2 m, 10 Mar. 1982, Id. H. D. Rudolph (1). [FSBC] I 43730, near Miami Beach shore, 25°48’49”N, 80°08'48”W (2). [FSBC] I 43731, Sta. 40, 25°47’47”N, 80°11’06”W, 1.7 m, 11 Mar. 1982, Id. H. D. Rudolph (2). [FSBC] I 43732, Sta. 38, 25°46’40”N, 80°11’57”W, 2.7 m, 11 Mar. 1982, Id. H. D. Rudolph (1). [FSBC] I 43733, South Biscayne Bay, 25°32’01”N, 80°10’17”W, 2.4 m, 21 Nov. 1982, Id. H. D. Rudolph (2). [UMML] 22.762, Broad River Delta, Chatham River Delta, Coot Bay Delta, Coll. W. L. Rouse, 29 Dec. 1965.

Description: Body pale or pink (in preserved material), small, 1~14 mm long. Base of crown involuted midventrally, each side forming a partial circle with dark inter-radiolar spots (Fig. 18C). Radioles without eyes, with paired serrated stylods for most their length (5~9 serrations) (Fig. 18K). Palmate membrane extending for 1/8 length of branchial crown. Radioles with bare tips, as long as equivalent space of 9 pinnules (Fig. 18K). Dorsal lips long (1/2 lengths of radioles) with dorsal radiolar appendages. Ventral lips rounded. Midline fecal groove deep on 1st segment forming mounds at each side, collar well separated dorsally. Ventral lappets rounded, overlapping (Fig. 18B). Anterior margin of 1st ventral shield M-shaped (Fig. 18A). Interramal eyespots present (Fig. 18C). Thorax, 0.5~2 mm long and 0.3~1.5 mm wide with 4~7 chaetigers. Spinelike chaetae on segment 1 arranged in compact fascicles. Thoracic notochaetae arranged in irregular, oblique rows of superior and inferior chaetae; superior chaetae spinelike with knee slightly broader than shaft (Fig. 18H); inferior chaetae spinelike with knee up to twice as broad as shaft (Fig. 18I). Thoracic uncini avicular, with 4 rows of equal-sized teeth above main fang in frontal view (Fig. 18E), occupying 1/2 of extension of main fang (seen under light microscopy), breast well developed, manubrium short (Fig. 18F). Abdomen with 18~23 chaetigers. Abdominal neuropodia as conical lobes; anterior abdominal row with short, spinelike chaetae, posterior row with modified, elongate, narrowly hooded chaetae (Fig. 18J). Abdominal uncini avicular, with 3 equal-sized teeth above main fang in lateral view (Fig. 18G), covering 1/2 of extension of main fang (seen under light microscopy), breast well developed, manubria short (Fig. 18G). Pygidium bilobed (Fig. 18D). Tubes flexible with fine sand.

Pseudopotamilla Bruguière, 1789

Müller (1771: 194) described “Die nierenförmige Amphitrite” from Iceland. Bruguière (1789: 57) gave a binomial name to Müller´s species: Amphitrite reniformis, and considered it to be the type species for Pseudopotamilla. Unfortunatelly, the type material is missing; Knight-Jones et al. are working in a revision of species from the North Sea, and describing the neotype of P. reniformis.

Pseudopotamilla reniformis (Müller, 1771) has numerous questionable records in many areas far apart from its type locality (Iceland): Fauchald and Reimer (1975) and Fauchald (1977) recorded it from Panama; Perkins and Savage (1975) listed it from Florida, the Gulf of Mexico, and the Caribbean; Laverde-Castillo and Rojas-García (1983) from Bahía de Cartagena (Colombia); and Uebelacker (1984) from Florida. We herein state that the specimens studied by Uebelacker (1984), from the northern Gulf of Mexico and identified as P. cf. reniformis correspond to a new species described below as P. fitzhughi sp. nov.

Pseudopotamilla fitzhughi sp. nov.

(Figs. 19-21)

Potamilla cf. reniformis Uebelacker 1984: 53-36, 54-38, figs. 54-29.

Etymology: This species is named in honor of Dr. Kirk Fitzhugh (Los Angeles County Museum of Natural History) for his significant contribution to the knowledge and taxonomy of the Sabellidae, and for his sustained advice and recommendations throughout this study.

Material: Holotype [LACM-AHF POLY 2150] Mexican Caribbean: I. Contoy, camping, on dead coral, 1.5 m, Coll. S. I. Salazar, 1 Mar. 2001. [LACM-AHF POLY 2151] One paratype. [ECOSUR 0050] Three paratypes (one mounted in gold for SEM). [UMML 22.767] One paratype. [ECOSUR] I. Contoy, Coll. L. F. Carrera and S. I. Salazar, June 1999 (5). San Felipe, Coll. J. R. Bastida and S. I. Salazar, 19 Feb. 1999 (5). Cancún, Punta Nizuc, Coll. S. I. Salazar and L. F. Carrera, 30 Mar. 1997, rock 2 (2). I. Cozumel, SEDENA, Coll. S. I. Salazar, 24 Mar. 2001 (1). Majahual, Coll. J. R. Bastida and P. Salazar, 22 Mar. 2000 (1). [FSBC] Florida: I 23695, east edge of Jeff’s Reef, east coast of Florida, 27°32.8’N, 79°58.8’W, Sta. JSL I-438 B, 23 Aug. 1977, Id. T. H. Perkins as P. reniformis, Coll. L. E. Edmiston, 81.4 m (20). [FSBC] Curaçao: I 60311, Netherlands Antilles, 9.1 m, Coll. Y. Steward-Van Es, Oct. 1977, Id. T. Perkins as P. reniformis (3). [UMML] Mexican Caribbean: 22.768, San Felipe, Coll. J. R. Bastida and S. I. Salazar, 19 Feb. 1999 (4). [USNM] Texas: 090424, Hospital rock, 27°32’0.5"N, 96°28’19’’W, Coll. BLM, Sta. HR1, 1976, 75 m. Id. J. M. Uebelacker as P. cf reniformis (1). [USNM] 090425, Southern Bank, 27°26’06’’N, 96°31’47”W, BLM-OCS, Spring 1976, Sta. SB3, 82 m, Id. J. M. Uebelacker (1). [USNM] Alabama: 090426, 29°40’30”N, 87°37’00”W, BLM-OCS, May 1974, Sta. 16 F, 36 m, Id. J. M. Uebelacker (1). [USNM] 090429, off Mobile Bay, 29°35’00’’N, 87°20’02”W, Coll. BLM-OCS, Feb. 1978, Sta. VI-2645, 106 m, Id. J. M. Uebelacker (3).

Description (variations observed in paratypes given in parentheses): Holotype complete, with medium-sized body, 8 (1.5~11.5) mm long, 1 (0.75~1.5 mm) mm wide with purple crown and thorax (in live and preserved materials). Radiolar crown long, measuring about 1/2 of body length, 4 (1.5~5.5) mm long. Base of crown with erect rigid dorsal, basal flanges. Crown with unpaired compound eyes only in dorsal radioles, except on dorsalmost pair, limited to proximal 1/2 of radioles, 3 or 4 teardrop-shaped eyes per radiole (Figs. 19D, 21A). Nine pairs of radioles, diminishing in length ventrally. Radioles with bare tips, as long as the space equivalent to 8 pinnules. Outer radiolar margins quadrangular in cross-section. Radiolar skeleton with 4 rows of cells in cross-section. Dorsal lips long, measuring about 1/2 of length of radioles, triangular and with distinct longitudinal ridge (mid-rib), with radiolar and pinnular appendages (Fig. 19E). Ventral lips and parallel lamellae present. Anterior margin of anterior peristomial ring low, of even height in lateral view (Fig. 19C), forming 2 deep pockets dorsally (Fig. 19A); posterior peristomial ring collar present. Ventrally, collar with a pair of triangular lappets, not overlapping, nearly reaching proximal margin of 1st ventral shield, when reflexed (Fig. 19B). Dorsal margin of collar fused to fecal groove (Fig. 19A). Ventral shield of segment 1 divided into 2 asymmetrical regions, anterior one with distal margin rounded (Fig. 19B). Thorax 1.5 mm long, 0.3~1 mm wide, with 9 chaetigers. Each ventral thoracic shield divided into 2 regions by a dark band. Segment 1 with spinelike chaetae. Following thoracic chaetigers with 3 spinelike superior notochaeta (Fig. 19F), and 5~7 paleate inferior notochaetae (Fig. 19F), arranged in 2 transverse rows, upper row with short and medium length mucro (Fig. 19A), lower row with short mucro (Fig. 19B, C). Thoracic tori longer than abdominal tori, not reaching ventral shields. Thoracic avicular uncini18 or 19 per torus, with main fang surmounted by 19~22 rows of equal-sized teeth in frontal view (seen under SEM), occupying 1/2 of extension of main fang, breast well developed, manubrium long (Figs. 19G, 20D). Thoracic uncini reaching 3/4 of length of manubria of companion chaetae (Fig. 19D). Companion chaetae with roughly symmetrical tips, as teardrop-shaped membranes (Figs. 20D, 21E). Abdomen with 11~32 chaetigers. Tori smaller than those on thorax, with 10~13 uncini. Abdominal neurochaetae in 2 transverse rows: anterior and posterior rows with 3 or 4 elongate, broadly hooded chaetae on all chaetigers (Fig. 20E), those of anterior rows shorter (Fig. 21F). Abdominal uncini avicular, with 22-24 rows of equal-sized teeth above main fang in frontal view (seen under SEM), occupying 3/4 of extension of main fang, breast well developed, manubrium short (Fig. 19H). Tubes made of mucus and fine sand.

Remarks: Pseudopotamilla fitzhughi sp. nov. resembles P. oligophthalmus (Grube, 1878), described from Singapore and later synonymized by Johansson (1927) with P. lacioniosa Ehlers, 1904 from New Zealand. Pseudopotamilla oligophthalmus and P. fitzhughi sp. nov. have entire dorsolateral collar margins, with rounded dorsal margins of the collar near the fecal groove (Table 2). They differ because (A) P. oligophthalmus has rounded ventral lappets, while in P. fitzhughi sp. nov., they are triangular.

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

	Collar margin		Ventral lappets	Thoracic uncini length vs. shaft of companion chaeta	Substrate	Type locality
	Dorsally, near fecal groove	Dorsolaterally
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, among encrusting fauna on surfaces of sheltered, hard substrata	Iceland
P. saxicava	rounded	entire	distinct, triangular	1	in rocks	SW France

Other similar species are P. aspersa (Krøyer, 1856), P. reniformis (Bruguière, 1879), and P. saxicava (de Quatrefages, 1866); P. fitzhughi sp. nov. differs from these species because (A) in P. reniformis, the dorsal margins of the collar are quadrangular near the fecal groove (rounded in P. fitzhughi sp. nov.); (B) P. aspersa and P. reniformis have indistinct ventral lappets (distinct and triangular in P. fitzhughi sp. nov.); (C) the manubrium of the thoracic uncini in P. saxicava are as long as the shaft of the companion chaetae (3/4 in P. fitzhughi sp. nov.); and (D) P. aspersa and P. reniformis are not found in rocks (P. fitzhughi sp. nov. and P. saxicava live in rocks) (Table 2).

Sabellastarte Krøyer, 1856

Fitzhugh (1989) provided a complete diagnosis for the genus; he noted that Sabellastarte does not appear to be an apomorphic definable group. Knight-Jones and Mackie (2003) recognized 8 valid species in the genus, and included typical but not unique characters in their generic diagnosis. Only 1 species has been described from the Grand Caribbean: S. magnifica (Shaw, 1800) from Jamaica.

Sabellastarte magnifica (Shaw, 1800)

(Fig. 22)

Tubularia magnifica Shaw 1800: 228, pl. IX, figs. 1-6.

Sabella magnifica.-- Savigny 1822: 78.-- de Quatrefages 1866: 443.-- Fitzsimons 1965: 643, figs. 1-13.

Sabella lingva Krøyer 1856: 27 fide Knight-Jones and Mackie 2003.

Sabella melania Schmarda 1861: 35, fig. 188 fide Knight-Jones and Mackie 2003.

Sabella splendida Kinberg 1867: 353 fide Knight-Jones and Mackie 2003.

Sabellastarte magnifica.-- Augener 1922: 48.-- Rioja 1958: 287.-- Knight-Jones and Mackie 2003: 2278-2280, fig. 3A-P.

Material: [ECOSUR] Mexican Caribbean: Veracruz, La Galleguilla, Coll. M. A. Tovar and S. Frontana, 12 May 2001 (4); I. Verde, Coll. SISV, 15 June 1983 (1). Panama: Fuerte Sherman, Colón (1), Coll. S. I. Salazar, 2 June 2002 (1); Coll. L. Harris (8). [LACM-AHF] British Virgin Is.: BVI-00, Sta. 28 Guana, Coll. K. Fitzhugh (1); BVI-00, Sta. 59 (1); BVI-99, Sta. 13, LH128, 27 Aug. (1); BVI-99, Sta. 42, LH-316, Coll. T. Zimmerman, 1 Aug. 1999 (1). [UMML] Virgin Is.: 22.121, St. John, Greater Lameshur Bay, Coll. Voss, Kumpf, and Adams, 22 Jan. 1959 (2); [UMML] 22.752, Coll. Manning and Moore, 5 Jan. 1961 (1); [UMML] 22.127, Coll. and Id. Thomas, 14 Dec. 1958 (1); [UMML] 22.120, St. John, Kiddle Bay, Coll. G. Voss, Id. J. Evald, 23 Jan. 1959 (1). Venezuela: I. Larga, Coll. S. Tovar, 10 Nov. 1962 (1).

Description: Large-bodied species, 70~142 mm long. Branchial crown 53~69 mm long, each side of crown base involuted ventrally to almost form a circle on each side, outer surface of radioles dark with pale longitudinal lines. Radioles 69~82 pairs; radiolar tips short and blunt. Radiolar skeleton with 8~10 rows of cells in cross-section (Fig. 22D). Dorsal lips long (3/4 of length of crown), with radiolar appendages, but without pinnular appendages. Ventral lips with parallel lamellae. Lateral margins of collar transverse to axis of body, well above junction between crown and thorax (Fig. 22C); dorsal margins equally high, with notches above collar pockets, defining well-developed dorsal lappets, with rounded margins (Fig. 22A). Ventral lappets triangular, overlapping at midline (Fig. 22B). Thorax twice as wide as long, 5~12 mm long, 7~17 mm wide with 8~10 chaetigers. Thoracic inter-ramal eyespots present. First segment somewhat longer than following segments. Thoracic tori long, extending towards ventral shields. Inferior thoracic notochaetae arranged in bundles with irregular, longitudinal rows of spinelike chaetae (Fig. 22H, I). Thoracic uncini avicular, with 13 equal-sized teeth above main fang in lateral view (seen under light microscopy), covering 1/3 of length of main fang (Fig. 22E), hood absent, breast well developed, manubrium medium (Fig. 22F). Abdomen with 188~204 chaetigers. Abdominal neurochaetae in partially spiraled arrangement, formed by anterior chaetal row; original posterior chaetal row as a small bundle partially enclosed by anterior row. Anterior abdominal neurochaetae spinelike on anterior rows (Fig. 22K); elongate, narrowly hooded on posterior rows; posterior abdominal neurochaetae spinelike in anterior rows; modified, elongate, narrowly hooded on posterior rows (Fig. 22J). Abdominal uncini avicular, with 13 equal-sized teeth above main fang, covering 1/3 of main fang length (seen under light microscopy), breast well developed, manubrium short (Fig. 22G). Pygidium rounded.

Checklist of sabellids from the Grand Caribbean

Species with an asterisk (*) are not included in the key until any revision sustains their distribution, because those records from the Grand Caribbean area are questionable, either because material reviewed by some authors was unavailable, or their identifications have not been corroborated with the type material. In Salazar-Vallejo (1996), references of all these records are included.

Amphicorina androgyne (Rouse, 1994)

TL: Carrie Bow, Belize. USNM 157629 (HT).

Amphicorina anneae (Rouse, 1994)

TL: Florida. USNM 157620 (HT).

* Amphicorina rivularis (Annenkova, 1929)

TL: Schantar Is., Sea of Okhotsk, Russia. Type material not found.

* Amphiglena mediterranea (Leydig, 1851)

TL: Nice, France. USNM 5095 (HT).

Anamobaea orstedi (Krøyer, 1856)

TL: West Indies. ZMUC 18.9.1845 (HT).

Anamobaea phyllisae sp. nov.

TL: Guana I., British Virgin Is. LACM-AHF (HT, PT).

Augeneriella hummelincki Banse, 1957

TL: Salinja Plenchi, Bonaire. ZMH V-11917a (HT).

Bispira brunnea (Treadwell, 1917)

TL: Nassau Harbour, Bahamas. AMHN 982 (HT). Synonyms: Sabella bahamensis Augener, 1922 fide Knight-Jones and Perkins (1998: 433).

Bispira melanostigma (Schmarda, 1861)

TL: Jamaica. NHMW (ST). Synonyms: Sabella thoracica Krøyer, 1856 and Sabella variegata Krøyer, 1856 fide Knight-Jones and Perkins (1998: 415).

Bispira paraporifera sp. nov.

TL: Puerto Aventuras, Mexico. LACM-AHF (HT).

Branchiomma bairdi (McIntosh, 1885)

TL: Bermuda. BMNH 1885.12.1.391 (LT).

Branchiomma conspersum (Ehlers, 1887)

TL: Key West, FL. MCZ 848 (HT).

Branchiomma nigromaculatum (Baird, 1865)

TL: St. Vincent, West Indies. BMNH 1839.12.27.16.20 (LT, SST). Synonyms: Sabella crispa Krøyer, 1856, Dasychone ponce Treadwell, 1901. Syntypes of Dasychonopsis arenosa Treadwell, 1924b correspond in part to young B. nigromaculatum and B. conspersum.

* Branchiomma wyvillei (McIntosh, 1885)

TL: St. Thomas, West Indies. Type material is missing, but the original drawings of thoracic uncini do not have the typical shape of Branchiomma. It probably corresponds to Sabellastarte magnifica (Shaw, 1800).

Chone americana Day, 1973

TL: Beaufort, NC. USNM 43134 (HT).

* Chone duneri Malmgren, 1867

TL: Spitsbergen, Norway. Type material not found.

Demonax flecatus (Hoagland, 1919)

TL: Puerto Rico. AMHN (HT).

Demonax jamaicensis (Augener, 1924)

TL: Jamaica. ZMH V-6791 (HT).

Demonax lacunosus Perkins, 1984

TL: Hutchinson Island, FL. USNM 54725 (HT).

* Demonax leucaspis Kinberg, 1867

TL: San Lorenzo, Peru. NRS 575 (HT).

Demonax microphthalmus (Verrill, 1873)

TL: Vineyard Sound, MA. USNM 13079 (ST).

* Euchone incolor Hartman, 1965

TL: Continental shelf off New England. LAMNH-261 (HT), -262 (PT).

* Euchone southerni Banse, 1970

TL: Ballynakill Harbor, Ireland. NMI 77.1908 (HT).

Fabricinuda pseudocollaris Fitzhugh, 1990

TL: Florida. USNM 122023 (HT).

Fabricinuda trilobata (Fitzhugh, 1983)

TL: Belize. USNM 74679 (HT, PT).

Jasmineira bilobata (Day, 1973)

TL: Beaufort, NC. USNM 43134 (HT).

* Jasmineira pacifica Annenkova, 1937

TL: Peter the Great Bay, Asia. Type material not found.

Manayunkia aestuarina (Bourne, 1883)

TL: Western Europe. Species broadly distributed in European waters and in the North American coasts: Bell (1982) recorded it from South Carolina, Bishop (1984) from the Gulf of Mexico, and Rouse (1995) from Chesapeake Bay (Maryland).

Manayunkia speciosa Leidy, 1855

TL: Schuylkill River, Philadelphia, PA. Type material not found.

Megalomma bioculatum (Ehlers, 1887)

TL: Straits off Florida. MCZ 669 (PT), MCZ 824 (PT).

Megalomma heterops Perkins, 1984

TL: Florida. USNM 54721 (HT), FSBC I 27696, 27698 (PT).

Megalomma lobiferum (Ehlers, 1887)

TL: Florida. MCZ (HT).

* Megalomma pacifica Johansson, 1927

TL: Gilbert Islands, Melanesia. UZIU (HT).

Megalomma perkinsi sp. nov.

TL: Florida. USNM 53976 (HT).

* Megalomma pigmentum (Reish, 1963)

TL: San Quintín, Mexico. USNM 57945 (HT). Records of M. pigmentum from Perkins (1984) correspond to 2 undescribed species (informally described herein as Megalomma sp. 1 and Megalomma sp. 2 from Venezuela and Florida, respectively).

* Megalomma vesiculosum (Montagu, 1815)

TL: Devonshire, UK. NMW.1995.024 (NT).

Notaulax bahamensis Perkins, 1984

TL: Bahamas. USNM 62082 (HT).

Notaulax circumspiciens (Ehlers, 1887)

TL: Florida. MCZ (HT).

Notaulax midoculi (Hoagland, 1919)

TL: Dry Tortugas, FL. AMNH 1186 (HT).

Notaulax nudicollis (Krøyer, 1856)

TL: St. Thomas, West Indies. USNM 62084 (HT). Synonyms: Sabella brevicollaris Grube, 1859, S. torquata Grube, 1877, and Protulides elegans Webster, 1884 fide Perkins (1984: 331).

Notaulax occidentalis (Baird, 1865)

TL: St. Vincent, West Indies. BMNH (HT).

Notaulax paucoculata Perkins, 1984

TL: Bahamas. USNM 62077 (HT).

* Notaulax phaeotenia (Schmarda, 1861)

TL: Sri Lanka. NHMW (HT).

Novafabricia infratorquata (Fitzhugh, 1983)

TL: Twin Cays, Belize. USNM 74644 (HT).

Perkinsiana fonticula (Hoagland, 1919)

TL: Guanica, Puerto Rico. AMNH (HT).

Potamilla floridana Augener, 1922

TL: Tortugas, SW Channel, Bird Key Reef, FL. MCZ (HT).

Potamethus spathiferus (Ehlers, 1887)

TL: Florida. MCZ (HT).

Pseudobranchiomma emersoni Jones, 1962

TL: Jamaica. AMNH 3612 (HT).

Pseudobranchiomma perkinsi Knight-Jones and Giangrande, 2003

TL: Indian River at Cape Canaveral, FL. NMW CP94003 (HT), LACM-AHF, USNM, FSBC (PT).

Pseudofabriciola longa Fitzhugh, 1990

TL: Highland Beach, FL. USNM 122056 (HT).

Pseudofabriciola quassiincisura Fitzhugh, 1996

TL: Coco Plum I., Belize. USNM 169974 (HT).

Pseudofabriciola sofla Fitzhugh, 1996

TL: Florida. USNM 169980 (HT). Synonym: Fabricia sp. A Uebelacker, 1984.

* Pseudopotamilla reniformis (Müller, 1771)

TL: Iceland, Finland, Norway. Type material missing. Knight-Jones et al. are currently describing a neotype (Knight-Jones pers. comm.).

Pseudopotamilla fitzhughi sp. nov.

TL: I. Contoy, Mexico. LACM-AHF (HT), ECOSUR (PT).

Sabellastarte magnifica (Shaw, 1800)

TL: Jamaica. BMNH 1998.932 (NT).

Acknowledgments: This work was possible thanks to the support provided by several colleagues. Kirk Fitzhugh and Eduardo Suárez provided much useful advice throughout this study. João Miguel de Matos Nogueira revised the manuscript and provided valuable suggestions. Leslie Harris and Andy Cohen made possible the inclusion of one of the authors (SISV) in the Pew Exotic Species Expedition to Panama. Todd L. Zimmerman and J. W. Martin made possible a series of expeditions to Guana Island (British Virgin Islands) under a grant from the Biotic Surveys and Inventories Program of the US National Science Foundation (DEB 9972100), during which L. Harris collected important material. Harry ten Hove (ZMA) made an intense search of sabellids in his collections from the Antilles and provided some useful material, and also made important suggestions to the manuscript. Andrew Cabrinovic (BMNH), Sandra Farrington (FSBC), Nancy Voss (UMML), and Kristian Fauchald (USNM) loaned material revised in this study. Leslie Harris provided motivation and hospitality, both extremely important to culminate this work. We would like to thank the colleagues in the Grupo de Bentos (ECOSUR) for their help in collecting samples from the Mexican Caribbean coasts, and Guadalupe Nieto (ECOSUR) for helping make the SEM pictures. CONACyT provided a scholarship to one of us (MAT-H) in El Colegio de la Frontera Sur, Chetumal (Mexico).

----------------------------------------------------------------------------------------------------------------

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