Pectinariidae (Polychaeta) from Caribbean and Associated Waters

PECTINARIIDAE (POLYCHAETA) FROM CARIBBEAN AND ASSOCIATED WATERS

CHARLENE D. LONG Museum of Comparative Zoology, Harvard University

ABSTRACT Three species of Pectinaria, P. regalis Verrill, P. gouldii (Verrill), and Pectinaria meredethi, n. sp., from various localities of the eastern Gulf of Mexico and eastern Caribbean are described. Previous work on Caribbean species is reviewed.

INTRODUCTION This study was initiated during a visit in 1970 to the marine laboratory of the University of Puerto Rico on Maguayes Island, off La Parguera. All Caribbean specimens of Pectinariidae deposited in the Yale Peabody Mu- seum (YPM), United States National Museum (USNM), Rosenstiel School of Marine and Atmospheric Science of the University of Miami (UMML), the University of Puerto Rico (UPR), and the Natuurwetenschappelijke Studiekring voor Suriname en de Nederlandse Antillen (NSSNA), were examined. I would like to thank Professors H. Baraclough Fell and Charles Cutress, Mrs. Bertha Cutress, and Drs. Willard Hartman, Lowell Thomas and H. A. ten Hove for their cooperation and help. I am particularly indebted to Dr. Marian Pettibone for her patient and valuable guidance. This work was supported in part by NSFGB-3532.

FAMILY PECTINARllDAE QUATREFAGES, 1865 The family was studied in detail by Nilsson (1928) and has since been discussed by several workers, including Hartman (1941) and Day (1967). A comparison of the terminology used in these and other publications indicates that there is considerable variation in the names used for different structures. The location, identity, and terminology of the characteristic features used in this report are shown in Figure 1. The number of teeth of the uncinus, as seen from the side view, is usually included as a part of the description of the species. This mayor may not indicate the number of teeth actually present in uncini with two or more vertical rows; the distance between the teeth determines the number seen from the side. If the teeth are sufficiently staggered and the teeth in both vertical rows are showing, then part or all of each tooth will be counted as a separate tooth when viewed from the side. If, however, the teeth are close together, then only some of the total number show laterally. There- fore, the tooth count should be taken when the uncini are face on. In the species discussesd here, there are two or more small teeth on the crown of 858 Bulletin of Marine Science [23(4)

Anterior tentacular cirrus Palp Posterior tentacular cirrus Lateral margin of cephalic veil Mid-ventral lobe of first ventral ridge Posterior gill First notosetiger

ANTERIOR END OORSAL VIEW First unci niger

UNCINI

(' J~ FRAGMENT OF TUBE NOTOSETAE

.SCAPHAL SETAE ~ J"r3 t~·_·~ Last unc;n;ger-4J_-.'-' .. . Achaete Last notoset;ger_·~. ~. ', Scaphal setae Scaphe----· • . . Anal cirrus "'1<.ll..llJ.U\ - '~Anal flap WHOLE ANIMAL SCAPHE VENTRAL VIEW DORSAL VIEH

FIGURE 1. Diagnostic characteristics of the Pectinariidae. 1973] Long: Pectinariidae (Polychaeta) 859 the uncinus and a variable number of much smaller teeth between the last large tooth and the lower scoop; the smaller teeth are usually not counted. Another diagnostic characteristic subject to some confusion is the total number of setigers. On the specimens examined for this study, the first 15 setigers were always distinct. On the last, or 16th, setiger, it was sometimes difficult to locate more than a few notosetae protruding from a reduced parapodiallobe; often, it appeared achaetous. The setigerous counts of the Caribbean specimens of Pectinaria gouldii indicate that the difference between 15 and 16 total setigers is not accompanied by specific, much less generic, differences. The use of total setigerous counts for generic defini- tions may need to be reconsidered. The number of papillae on the cephalic veil is often given as part of a species diagnosis. On the specimens of P. regalis, there was a general correlation between total body length and number of papillae, with a total difference of ] 1 (24-35) papillae from the smallest to the largest specimens. On the other hand, Car.ibbean specimens of P. gouldii did not show such a correlation and, in fact, had a range of 12 to 38 papillae not correlated with size. Obviously, the number of cephalic-veil papillae is not diagnostic for the Caribbean pectinariids thus far studied. As used in this study, the first ventral ridge extends transversely between the posterior tentacular cirri. In the Caribbean pectinariids, it was found that examination of the midventral lobe of this ridge was very helpful in making preliminary identifications of well-preserved specimens. Thus, the three species discussed here are quite easily separated as follows: P. rega/is has a smooth, single midventrallobe; P. gouldii has a group of small papillae in this area; P. meredithi has a striated or ridged lobe. As with all soft parts, however, this character must be used with care on contracted, inflated, or otherwise distorted specimens.

GENERA The number and definition of the genera and subgenera of the Pectinari- idae have differed depending upon the author. In her catalogue, Hartman (1959) recognized five genera: Amphictene Savigny, 1818a; Cistenides Malmgren, ] 866; Lagis Malmgren, 1866; Pectinaria Savigny, 1818b; and Petta Malmgren, ] 866. The distinction between Pectinaria and Cistenides has undergone much discussion. Hartman (1941) separated these two genera on the basis of the number of vertical rows of teeth in the uncini, with Pectinaria having two or more rows, and Cistenides having only one. Following this, the three species reported here are members of the genus Peetinaria.

HISTORICAL REVIEW OF CARIBBEAN SPECIES There have been several reports of species of Pectinariidae from Carib- bean and associated waters. The earliest record was that of Donovan 860 Bulletin of Marine Science [23(4) ( 1826), who described and illustrated Peetinaria guildingii from St. Vin- cents (sic), West Indies. The description is not sufficient to determine its present standing. The species has not been referred to subsequently and, since the whereabouts of the original specimen is unknown (Dr. M. H. Pettibone, pers. commun.), it is here declared nomen dubium. The next published account of this family in North America was by Gould (1841), who reported Pectin aria belgiea (Pallas) from Massachu- setts. Later, P. groenlandiea, (?) Grube was reported from Grand Manan Island, New Brunswick, Canada, by Stimpson (1854), and P. aurieoma (Muller) was briefly described from Rhode Island and New Jersey by Leidy (1855). These false extensions of range of European species were corrected in part by Verrill (1873), who described Cistenides gouldii from New England and included the above records of Gould and Leidy as synonyms. Verrill (1881: 290) subsequently referred gouldii to Peetinaria. Another North American species, Peetinaria (Lagis) dubia from Virginia, was described by Webster (1879). Nilsson (1928) supplemented the description of P. gouldii, based on a study of specimens from Massachusetts, New York, and Jamaica, and placed in synonymy Stimpson's record of P. groenlandiea and Webster's P. dubia. Another report of Pectinariidae from the Caribbean is found in the work of Ehlers (1887), who described Peetinaria (Petta) pellucida, based on a single specimen from Santarem Channel in 486 meters, apparently referring to the Bahamian Santaren Channel. This genus has not since been reported from the Caribbean. Verrill (1901) described Peetinaria regalis from Bermuda. The description was subsequently supplemented and referred to Cistenides by Hartman (1941, 1942). Treadwell (1901, 1939) reported P. gouldii from Puerto Rico; these specimens have been reexamined and are referred herein to P. regalis. In 1941, Treadwell reported P. gouldii from Bermuda; this specimen is apparently no longer in existence (Dr. Ernst Kirsteucr, pers. commun.). Marsden (1960) reported the Pacific P. ehilensis (Nilsson) from Barbados, West Indies. I have been unable to locate these specimens.

KEY TO Peetinaria FROM THE EASTERN CARIBBEAN 1A. Scaphal setae 0-4, straight, tapered, not curved (Fig. 3, D). Oper- cular rim incised (Fig. 3, A). First ventral ridge with one smooth midventrallobe (Fig. 2, A). Notosetae without boss or incision (Fig. 3, B). Tube formed of single layer of large, variously sized, opaque shell fragments and debris (Fig. 2, B) P. regalis lB. Scaphal setae 14-26, straight and abruptly tapered to off-center tip, when viewed from above, and curved, when viewed laterally (Fig. 4,D). Opercular rim not incised. First ventral ridge with 3 or more papillae midventrally (Fig. 4, A). Some notosetae with ] 973] Long: Pectinariidae (Polychaeta) 861 bosses separated from shafts by incision (Fig. 4, E). Tube usually formed of single layer of small, evenly sized, transparent sand grains (Fig. 4, C) P. gouldii 1C. Scaphal setae 7-9, straight to slightly curved and tapered, when viewed from above, and curved when viewed laterally (Fig. 5,D). Opercular rim not incised. First ventral ridge with single striated or ridged midventrallobe (Fig. 5, A). Some notosetae with bosses directly adhering to shafts and blades of varying lengths or absent (Fig. 5, E, G). Tube formed of two layers of variously sized, overlapping, opaque shell fragments and debris (Fig. 5, C) P. meredithi

Pectinaria regalis Verrill, 1901 Figs. 2, 3 Pectin aria regalis Verrill, 1901: 38, pI. 8: figs. 6, 7.-Not Hartman, 1941: 332, pI. 50: figs. 9,10, pI. 51: fig. 18. Cistenides regalis, Hartman, 1942: 71-4, figs. 129, 132, 133, 136, 137, 140. Pectinaria gouldii, Treadwell, 1901: 206; 1939: 288, fig. 103.-Not Verrill, 1873. Material Examined.-BERMUDA: Coney Island, A. E. Verrill and class, collectors, 1901: 3 syntypes (YPM 2739).-Hungry Bay, W. G. Van Name, collector, 1901: 1 specimen (YPM 2804). PUERTORICO: Ensenada Honda, Culebra, FISH HAWK, 9 February 1899: 1 specimen (USNM 16121).-OfI Vieques Island, 25 meters, FISH HAWK Sta. 6085, 8 February 1899: 2 specimens (USNM 16122).- Majimo Reef, M. E. Rice, collector, 15 April 1967: 1 specimen (USNM 43438).-Puerto Rico, C. Mangum, collector: 2 specimens (USNM 43448). -Cayo Enrique, leeward side, soft sand just below low water, C. E. Cutress, collector, 30 May 1968: 4 specimens (UPR).-West end of Cayo Cara- coles, La Parguera, in sand near Porites and sea grass, P. Castro and C. D. Long, collectors, 11 March 1970: 1 specimen (USNM 49217). VIRGIN ISLANDS: Lesser Lameshur Bay, St. John, G. Voss and H. Kumpf, collectors, 24 January 1959: 1 specimen (UMML 22.100). BARBADOS:Near Lord's Castle, Barbados, Barbados-Antigua Expedition 1918: 1 specimen (USNM 20295). BONAIRE:Lac, Awa Blancu, Bao di Dam, 700 meters ESE of Sorobon Pt.,1;2 meter, sand, Thalassia, NSSNA 1563, 25 August 1967: 1 specimen (USNM 49218). FLORIDA:Sewage Beach, Virginia Key, A. Bitler, collector, June 1960: I specimen (UMML 22.189). GEORGIA: Sapelo Whistle, about 20 meters, M. Gray, collector, 28 February 1962: 1 specimen (USNM 43437). Description.-Tubes 35-110 mm by 7-20 mm, tapered, moderately curved; composed of one layer of cream or white (rarely colored) large coral and 862 Bulletin of Marine Science [23(4) shell fragments, Foraminifera and small shells poorly fitted together but seldom overlapping (Fig. 2, B). Specimens 26-90 mm by 6-17 mm. Opercular plate without papillae; rim incised, slits or fusion marks reaching to opercular plate (Fig. 3, A). Paleae 8-14 pairs, tapering to fine, slightly curved tips; usually tips broken or worn (Fig. 3, A). Cephalic veil free and extending posteriorly into oral area; margin with 21-35 slender, well-spaced papillae (Fig. 2,A). Number of cephalic-veil papillae varying with size of specimen, greatest number (35) found in largest specimen (90 mm), with more or less regular reduc- tion in number in smaller specimens; smallest specimen (26 mm) with fewest papillae (24). First ventral ridge with four lobes on either side of smooth, midventral lobe (Fig. 2, A). Anterior pair of gills larger than posterior pair. Setigers 16 (1-3, notosetae only; 4-15 both uncini and notosetae; 16, notosetae only), usually additional posterior achaetous parapodium. Scaphe glandular to smooth dorsally, laterally with reduced parapodial lobes, posteriorly with two lobes separated by a median space and fringed with crowded papillae (Figs. 2, A; 3, A). Scaphal setae 0-4, with one or two still in flesh; straight and tapered from all views (Fig. 3,A,D). Anal flap smooth along margin and ventrally (Figs. 2, A; 3, A); middorsal cirrus present (not shown in figure). Notosetae all tapered, narrowly winged capillaries without boss or incision; some with wings wider on one side (Fig. 3, B). Uncini with larger teeth in two (occasionally three) vertical rows and 6-10 horizontal rows; crowned with 1-5 smaller teeth (Fig. 3,e). Remarks.-The specimens from Puerto Rico, identified by Treadwell (1901, 1939) as P. gauidU, were examined (USNM 16121-2) and are referred herein to P. regalis. The specimen from Socorro Island, Mexico, identified by Hartman (1941) as Cistenides regalis, is being described as a new species, according to Dr. Kristian Fauchald (pers. commun.). Study of the specimens of P. regalis indicates that the use of the number of cephalic-veil papillae as a taxonomic characteristic should be correlated with size measurements. The midventral portion of the first ventral ridge of the specimen from Bonaire was deeply sunken into the body and, there- fore, covered by the midventral portion of the second ventral ridge. When exposed, the midventrallobe of the first ventral ridge was an atypical, low striated mound similar to that seen in P. meredithi. Distributian.-Bermuda, West Indian Antilles (Puerto Rico, Virgin Islands, Barbados, Bonaire), Florida Keys, Georgia. Littoral to 25 meters.

~ FIGURE 2. Pectinaria regalis Verrill, 1901 (USNM 49217): A, entire worm, ventral view; B, fragment of tube. (a, midventrallobe of first ventral ridge; b, achaetous parapodium; c, scaphe.) 1973] Long: Pectinariidae (Polychaeta) 863

o ~mm I A, B

A 864 Bulletin of Marine Science [23( 4)

c B

o .028mm I I c 1mm

4mm I A 1973] Long: Pectinariidae (Polychaeta) 865 Pectinaria gouldii (Verrill, 1873) Fig. 4 Cistenides gouldii Verrill, 1873: 612, 323, pI. 17: figs. 87, 87a.-Hartman, 1944: 342, pI. 52[20], fig. 9, 9a. Pectinariagouldii, Verrill, 1881: 290.-NotTreadwell, 1901: 206; 1939: 288, fig. 103. Pectin aria [Cistenides] gouldii, Nilsson, 1928: 24-28, fig. 7A-H. [?] Pectin aria gouldii, Treadwell, 1941: 29. Material Examined.-MAsSACHUSETTS: Barnstable Public Beach, Cape Cod, low tide, on sand flat, C. D. Long and J. Spiel, collectors, 17 June 1967: 1 specimen (USNM 49216).-Woods Hole, 12-15 meters, May- October 1971: 3 specimens; February 1972: 1 specimen. LONGISLANDSOUND: 8-14 meters, A. E. Verrill and class, collectors, 1866: 21 syntypes (YPM 169). FLORIDA(Atlantic): Upper Biscayne Bay, benthos, J. McNulty, collector, 8 June 1956: 2 specimens (UMML 22.117).-Matheson Ham- mock, Silver Bluff Shore, S. of Coconut Grove, Dade County, L. P. Thomas, D. R. Moore, and R. Work, collectors, 10 November 1960: 1 specimen (UMML 22.197). FLORIDA(Gulf of Mexico): Punta Rassa, 1.8 meters, H. Hemphill, collector, February 1884: 1 specimen (USNM 43439) .-Marco, in shell with hermit crab, H. Hemphill, collector, 1884: 1 specimen (USNM 839). -Coot Bay, Everglades National Park, D. C. Tabb, collector, 27 June 1958: 31 specimens (UMML 22.141 ).-Wiggan's Pass, Naples, inlet beach side, C. Long, collector, 28 December 1958: 1 specimen (USNM 43453) .-Sarasota, near Cape Haze Marine Laboratory, R. Brown, R. Cressey, J. Levi, collectors, 27 January 1966: 1 specimen (USNM 43443). -Wilson's Beach, St. Theresa, 0.3 meter, D. W. Kirtley, collector, 10 April 1966: 1 specimen (USNM 43440). CUBA: Sabanilla, New Granada, ALBATROSS,16-22 March 1884: 1 specimen (USNM 1702). PUERTORICO: Off Fajardo, 3 meters, N. Hulings and D. Feray, collectors, 1963: 1 specimen (USNM 42786). Description.-Tubes 12-32 mm by 1.5-5 mm, tapered, slightly curved; composed of one layer of small, evenly sized, transparent (occasionally orange-hued) sand grains, neatly fitted together and not overlapping (Fig. 4, C); few bits of black or grey stones or opaque shell fragments in most tubes.

~ FIGURE3. Pectin aria regalis Verrill, 1901 (USNM 49217): A, entire worm, dorsal view; B, notosetae; C, uncini; D, scaphal setae. (a, opercular rim incision; b, scaphal setae; c, anal flap, reflected; d, worn palea; e, scaphe.) 866 Bulletin of Marine Science [23(4)

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A 1973] Long: Pectinariidae {Polychaeta} 867 Specimens 5-45 mm by 1-9 mm. Opercular plate and rim both smooth. Paleae 9-15 pairs, tapering to fine, slightly curved tips when not worn or broken. Cephalic veil free and extending posteriorly into oral area; margins with 12-38 slender papillae (Fig. 4, A). Number of cephalic-veil papillae not varying with size of specimen. First ventral ridge with four lobes on either side of a midventral group of three to many papillae (Fig. 4, A); smaller specimens usually having fewer papillae. Anterior pair of gills larger than posterior pair. Setigers 15-16 (1-3, notosetae only; 4-15 both uncini and notosetae; 16 with or without notosetae), with 1-2 achaetous parapodia posteriorly (Fig. 4, A). Scaphe smooth dorsally; laterally with reduced parapodial lobes, posteriorly with row of crowded marginal papillae (Fig. 4, A, B). Scaphal setae 8-26, with some still in flesh; smaller specimens with fewer scaphal setae (Fig. 4, B). When viewed from above, scaphal setae straight, distal end (just proximal to off-center tip) slightly wider than base; when viewed laterally, appearing tapered and slightly curved (Fig. 4, D). Anal flap with papillose margin and smooth ventral surface; middorsal anal cirrus present (Fig. 4,B). Notosetae of two kinds: (1) tapered capillaries with narrow wings and (2) capillaries with boss separated from shaft by incision and with tapered, narrowly winged blade (Fig. 4, E). Uncini with larger teeth in 2-4 vertical rows and 6-8 horizontal rows; crown of 1-3 smaller teeth (Fig. 4,F). Remarks.- Treadwell (1901, 1939) reported P. gouldii from Ensenada Honda, Puerto Rico; as noted above, the specimens (USNM 16121-2) were examined and referred to P. regalis. Nilsson (1928) described a small (12 mm long) specimen of P. gouldii from Jamaica. Treadwell (1941) listed this species from Castle Harbor, Bermuda. This specimen is apparently no longer in existence (Dr. Ernst Kirsteuer, pers. commun.). Specimens of P. gouldii from southern waters have been consistently smaller than those from more northern waters: Cape Cod, 5-45 mm; Long Island Sound, 20-27 mm; North Carolina, 10 mm; SE Florida, 17-19 mm; NW Florida, 11 mm; SW Florida, 8-20 mm; Cuba, 15 mm; and Puerto Rico, 11 mm. The syntypes of C. gouldii (YPM 169) are extremely flaccid, with the skin partly inflated, making determinations of soft parts impossible. Tubes of specimens of P. gouldii from Coot Bay, Everglades National

FIGURE4. Pectinaria gouldii (Verrill, 1873) (A-F, USNM 49216; G, UMML 22.141): A, entire worm, ventral view; B, posterior end, dorsal view; C, fragment of tube; D, scaphal setae; E, notosetae; F, uncini; G, fragment of tube. (a, midventrallobe of first ventral ridge; b, achaetous parapodium; c, scaphe; d, scaphal setae; e, anal flap.) 868 Bulletin of Marine Science [23(4) Park, Florida, differ from those of other specimens (Fig. 4, G). They are composed of Foraminifera and overlapping rounded shell fragments. While many of the tubes have a fine covering of sandy silt, the distinct quartz sand grains found in tubes of typical P. gouldii are not present. The number of cephalic-veil papillae does not appear to be correlated with the size of the animal, as in P. regalis. Thus, the smallest specimen of P. gouldii (5 mm) had the fewest number of papillae (12); a 15-mm specimen had the greatest number (38), and the largest specimen (45 mm) had only 29 papillae. The variability in the number of papillae indicates that this character cannot be used as a taxonomic feature for Caribbean gouldii. Of the specimens carefully checked for the number and arrangement of setigers, half had 15 setigers and half had 16 setigers. Specimens from Cape Cod and western Florida had 15 or 16 setigers; specimens from Long Island Sound, North Carolina, and Cuba had 16 setigers; specimens from Puerto Rico, Miami, and northern Florida had 15 setigers. The notosetae of the 16th setiger are sometimes so inconspicuous as to be difficult to locate. While Nilsson (1928) noted the possibility of larger specimens of P. gouldii having a greater number of setigers, he reported 15 setigers in the specimens he examined. On the specimens examined for this study, there was no correlation between length and number of setigers; thus, specimens 8 mm and 15 mm in length both had 16 setigers, while several specimens 5 to 20 mm in length had 15 setigers. Obviously, there is not a fixed number of notosetigers in P. gould ii, and its use as a generic character is also questionable. Two small (5 and 20 mm long) specimens from Woods Hole had three to four vertical rows of larger teeth on the uncini. Nilsson (1928) also examined uncini from specimens with more than two vertical rows of larger teeth. Examination of one of the three syntypes of Pectinaria (Pectinaria) hartmanae Reish (1968), from Bahia de los Angeles, Lower California (USNM 38399), showed that it resembles P. gouldii in the following characters: size, the opercular plate and rim, the cephalic veil and associated papillae, the paleae, the gills, the scaphe, the anal flap with lateral papillae, the shape of the scaphal setae, and the notosetae. As with most specimens of gouldii, the syntype of P. hartmanae is too flaccid to permit observations on the first ventral ridge. P. hartmanae differs from P. gouldii in regard to the tube, uncini, and number of scaphal setae. The tube, while composed of transparent sand grains, is much more uneven in appearance, due to the inclusion of some sand grains with a rough, exposed edge. This overall rough appearance was noted in one collection of P. gouldii from the Everglades (see above); the materials used, however, are quite different. For the present, it is impossible to determine the significance of tube differences in isolated 1973J Long: Pectinariidae (Polychaeta) 869 collections of P. gouldii. The uncini of P. hartmanae have at least three vertical and seven horizontal rows of evenly sized teeth and are constructed so as to make the face-on view reminiscent of a pineapple. The scaphal setae resemble those of P. gouldii, but are slightly fewer in number. In specimens of similar size (15-20 mm long), there would be 14-26 emergent scaphal setae in P. gouldii, compared to 8-10 in P. hartmanae. Distribution.-East coast of North America (Cape Cod to Florida), Gulf of Mexico (Florida), West Indies (Cuba, Jamaica, Puerto Rico). Littoral to 15 meters.

Pectinaria meredithi, n. sp. Fig. 5 Material Examined.-BAHAMAS: Fresh Creek, south side of Andros Island, 24° 43.2' N, 77° 47.5' W, bushy alga (?) and Thalassia, M. L. Jones, collector, 1 March 1966: holotype (USNM 43442).-Northwest coast of Andros Island, 24° 53.3' N, 78° 12.1' W, lower intertidal, M. L. Jones, collector, 17 March 1966: para type (USNM 43441). FLORIDA: Lower Matacumbe Key, among grass below low tide, Henry Hemphill, collector, before 1888: 1 specimen (USNM 41l0).--South side of Tavernier Key, M. L. Jones, collector, July 1960: 1 specimen (USNM 43447). Type-Material.-The holotype is 26 mm by 4 mm; the tube is 32 mm by 6 mm. The paratype is 9 mm by 2 mm; the tube is 10 mm by 3 mm. Description.-Tubes 10-36 mm by 3-7 mm, tapered, slightly curved; composed of two layers of variously sized, overlapping, opaque, cream and white shell fragments with occasional bits of colored debris (Fig. 5, C). Tube of holotype and paratype from Bahamas with flat fragments of shells; tube of specimen from Tavernier Key (USNM 43447) with rounded fragments of shells; tube of specimen from Lower Matacumbe Key (USNM 4110) with mixture of flat and rounded fragments. Specimens 9-26 mm by 2-4 mm. Opercular plate and rim both smooth. Paleae 8-11 pairs (holotype: 10), tapering to fine, slightly curved tips when not worn or broken. Cephalic veil free and extending posteriorly into oral area; margins with 16-21 (holotype: 21) slender papillae (Fig. 5, A). Number of cephalic-veil papillae not varying with size of specimen. First ventral ridge of holotype with four lobes on either side of ridged midventral lobe; lobe slightly incised medially (Fig. 5, A). Other specimens with three to four lobes on either side of striated to ridged midventral lobe; lobe incised to varying degrees. Anterior pair of gills larger than posterior pair. Setigers 16 (1-3, notosetae only; 4-15, both uncini and notosetae; 16, notosetae only) with 1 posterior achaetous parapodium. Scaphe smooth 870 Bulletin of Marine Science [23(4)

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KEY 9 Pcctinaria goutdii~ Long 1972 (g) Poctinaria goutdii~ Nilsson 1928 or Treadwell 1941 Pcctinaria rogaZis) Long 1972 Pcctinaria mercdithi Long 1972 Pcctinaria chilcnais~ Marsden 1960 Pcctinaria guildi1:gi Donovan 1826

r,lg! BERMUDA

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BONAI RE .~

r ".I?!s~~Re~~g~NT D DISTRIBUTION OF PECTINARIIOAE IN CARIBBEAN AND ASSOCIATEO WATERS

FIGURE 6. Distribution of Pectinariidae in Caribbean and associated waters. dorsally; laterally with reduced parapodial lobes, posteriorly with row of few, irregularly spaced marginal papillae (Fig. 5, A, B). Scaphal setae 7-9 (holotype: 8) with some still in flesh; tapered, straight to slightly curved when viewed from above, curved when viewed laterally (Fig. 5,D). Anal flap with smooth margin and papillate (4-6 papillae) ventral surface (Fig. 5, A, B); middorsal cirrus present (Fig. 5, B). Notosetae of two kinds; (1) tapered capillaries with narrow wings and (2) capillaries with boss directly adhering to shaft and with or without tapered, narrowly winged blade of varying length (Fig. 5, E, G). Uncini with larger teeth in 2-3 vertical rows and 6-8 horizontal rows; crown of 3-5 smaller teeth (Fig. 5,F).

~ FIGURE 5. Pectinaria meredithi, n. sp. (A-F, holotype, USNM 43442; G, USNM 43441): A, entire worm, ventral view; B, posterior end, dorsal view; C, fragment of tube; D, scaphal setae; E, notosetae; F, uncini; G, notosetae. (a, midventral lobe of first ventral ridge; b, achaetous parapodium; c, scaphe; d, scaphal setae; e, anal flap.) 872 Bulletin of Marine Science [23(4) Etymology.-It gives me great pleasure to name this species after Dr. Meredith Jones, who collected three of the four specimens studied. Remarks.-The four specimens of P. meredithi, one of which was collected nearly a hundred years ago, are quite distinct in all characters now con- sidered diagnostic for the two other species thus far studied from the Caribbean. Of particular note is the double-layered tube, which is so distinct as to immediately separate these specimens from others collected in the Caribbean. The holotype has a narrow midventral slit in the first ventral ridge, while the slit in the para type is wide; a third specimen is badly inflated, but midventral striations and more lobes are clearly visible. While the poor condition of the specimen from Lower Matacumbe Key (USNM 411 0) prevents verification of these structures, the tube, the scaphal setae, the notosetae, and the ventral papillae on the anal flap are unquestionably the same as in the other specimens. In all specimens, some of the bossed notosetae, at first glance, seem to have an incision separating the boss from the shaft; closer examination reveals a tear at this point (see Fig. 5, E, G). This is the point of breakage of most incomplete setae. The presence of the wing surrounding the boss distinguishes bossed setae that normally lack a blade from those that have subsequently lost the blades. The report of P. chilensis Nilsson from the Caribbean by Marsden (1960) necessitates a comparison of P. meredithi with Nilsson's description. Nilsson (1928) noted, among other things, that his single, 60-mm specimen from Chile had the anterior pair of gills inserted so closely to the midventralline that there was only a single lobe present between them, compared with three large lobes between the anterior gills in P. meredithi. Furthermore, he described and illustrated the anal flap with a laterally papillate margin, compared with an anal flap with a smooth margin in P. meredithi. Distribution.-Bahamas and the Florida Keys.

SUMMARY The three species of Pectinariidae from various parts of the eastern Gulf of Mexico and the eastern Caribbean are congeneric and sympatric (sec Fig. 6). The specimens of P. meredithi and gouldii are always smaller than the specimens of P. regalis from the same area. Additional collections should help distinguish ecological differences among the three species.

SUMARIO

PECTINARIIDAE (POLYCHAETA) DEL CARIBE Y AGUAS ASOCIADAS Las tres especies de Pectinariidae de varias partes del este del Golfo de Mexico y del este del Caribe son congenericas y simpatricas (vease mapa). 1973] Long: Pectinariidae (Polychaeta) 873 Los ejemplares de Pectinaria meredithi, n. sp., y P. gouldii son siempre mas pequeiios que los ejemplares de P. regalis de la misma area. Colecciones adicionales deben ayudar a distinguir las diferencias ecol6gicas entre las tres especies.

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