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ON SOME P:OLYCHABTOUS FROM JAMAICA, .THE WEST INDIES

MEREDITH L. JONES

BULLETIN OF THE AMERCANMUSEUM OF NATURAL HISTORY VOLUME 124,: ARTICLE 5 NWYOK 1962

ON SOME POLYCHAETOUS ANNELIDS FROM JAMAICA, THE WEST INDIES

ON SOME POLYCHAETOUS ANNELIDS FROM JAMAICA, THE WEST INDIES

MEREDITH L. JONES Assistant Curator, Department of Living Invertebrates The American Museum of Natural History

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 124 : ARTICLE 5 NEW YORK: 1962 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 124, article 5, pp. 169-212, text figures 1-146, plate 52, table 1 Issued October 15, 1962

Price: $1.00 a copy INTRODUCTION

ALTHOUGH MANY COLLECTIONS Of pOly- It will be noted that in some cases, i.e., the chaetes have been made in the Gulf of cosmopolitan , the synonymies given Mexico and Caribbean area, most of our below are not complete. It was felt that, for knowledge concerning these annelids derives the present, it would be best to restrict the from work done by Schmarda (1861), Tread- synonymies and locality records to those of well (1901, 1917, 1921, 1924a, 1924b, 1928, the Gulf of Mexico and Caribbean region. In 1936, and 1939), Augener (1906, 1922a, and addition, it should be noted that certain of 1927b), Monro (1928, 1933a, 1933b, and the synonymy entries are enclosed in quota- 1933c), Hartman (1942b and 1951b), and tion marks; these signify that the citation in Rioja (1946). More recently, single papers question refers, not to a new locality record, have been contributed by Carpenter (1956), but to generalized localities that are based on on the of the northern Gulf, previous records in the literature. Renaud (1956), on those of Miami, Florida, The following determinations have been and Bimini, Bahamas, and Marsden (1960), made for this collection: on the polychaetous annelids of Jamaica and Barbados. Polyodontes oculea (Treadwell), 1901 In spite of the number of faunistic and Pareulepis sulcatisetis, new species distributional studies of this area, the present Chloeia viridis Schmarda, 1861 as well as the examination Hesione picta Muller, 1858 work, preliminary Typosyllis corallicola (Verrill), 1900 of other collections from northwest Florida, Glycera abranchiata Treadwell, 1901 indicates that there is still much to be done ?Eunice tridentata Ehlers, 1905 in the compiling of a definitive species list of Lysidice ninetta Audouin and Milne-Edwards, 1833 polychaetes for the Gulf of Mexico and the Polydora ancistrata, new species Caribbean. Nerinides goodbodyi, new species The 21 species reported upon in the present Armandia nonpapillata, new species paper were collected and kindly sent to me Dasybranchus sp. by Dr. Ivan M. Goodbody, Department of Nicomache antillensis Augener, 1922 Zoology, University College of the West Sabella melanostigma Schmarda, 1861 Indies, Jamaica. Duplicate specimens are in Branchiomma nigromaculata (Baird), 1865 Branchiomma arenosa (Treadwell), 1924 the reference collection at the University Pseudobranchiomma emersoni, new genus, new College, and all type material has been de- species posited in the collections of the American Sabellastarte magnifica (Shaw), 1800 Museum of Natural History. One additional Olga elegantissima, new genus, new species species (Eunice, new species) will be con- Salmacina amphidentata, new species sidered later in a separate paper. Eupomatus alatalateralis, new species

173 SYSTEMATIC TREATMENT

FAMILY POLYODONTIDAE PFLUGFELDER, 1934 bases of the palpi, there are two short conical GENUS POLYODONTES RENIERI IN AUDOUIN AND tentacles. An unpaired tentacle, similar in MILNE-EDWARDS, 1832 shape to the anterior pair, arises on the mid- Polyodontes oculea (Treadwell), 1901 line in the anterior third of the prostomium. Panthalis oculea TREADWELL, 1901, pp. 18S- About midway along the length of the 189; Puerto Rico. prostomium there is a pair of eye spots, and Polyodontes oculea, MONRO, 1928, pp. 572-575; near the posterior prostomial margin there Trinidad, Col6n (Caribbean end of the Panama are two more pairs of eye spots. The sides Canal); Panama [City], Taboga, Balboa (Pacific of the prostomium are nearly straight, the end of the Panama Canal). posterior margin is broadly rounded, and the Panthalis oculea, TREADWELL, 1939, pp. 192- anterior margin, in the vicinity of the palpi, 193; "Puerto Rico." is very slightly attenuated. The posterior U.C.W.I. COLLECTION: One ovigerous speci- two-thirds of the prostomium is hidden by men, in two fragments, measures 38 mm. in medial extensions of the first elytrophores. total length and has 68 setigerous segments. These are fused to the dorsal mid-line of the The specimen, 3 mm. in width, excluding prostomium and the point of fusion extends parapodia, and 6 mm. in width, including anteriorly, to the level of the anterior pair of parapodia, was dredged near Pickinng Beacon eye spots. on January 29, 1960. The first three parapodia are directed DISTRIBUTION: Tropical: throughout the anteriorly, and the fourth is at right angles to Greater and Lesser Antilles, from Jamaica to the anteroposterior axis. In the holotype, the Trinidad; on both sides of Panama. prostomium of which is not everted, all these anterior parapodia are in the same FAMILY PAREULEPIDAE HARTMAN, 1939 frontal plane. GENUS PAREULEPIS DARBOUX, 1899 In the paratype (figs. 2 and 3), the everted Pareulepis sulcatisetis, new species proboscis has caused a shifting of the relative position of certain structures. The bases of Figures 1-27 the pair of palpi are separated and are lateral DIFFERENTIAL DIAGNOSIS: Pareulepis with to the pair of prostomial tentacles. The first 12 fringed elytra which bear no dorsal tuber- parapodia come to lie directly above the cles; with tripartite marginal processes on the second, almost in line with the first elytro- twelfth elytra; with three pairs of prostomial phore. The proboscis of Pareulepis sulcatisetis eyes; with heavy, bent notosetae, the tips of has a smooth exterior, and its opening is which may be spatulate or gouge-like; and bordered with a single row of papillae, ap- with 32 setigerous segments. proximately 15 above and 18 below. The U.C.W.I. COLLECTION: Two specimens of mouth opening of the proboscis of the para- Pareulepis sulcatisetis were collected in mud, type is sufficiently wide so that it can be Green Bay, Jamaica, on February 11, 1960. seen that there are no chitinous teeth or The holotype is 9.5 mm. long and approxi- jaws associated with this structure. mately 2.5 mm. broad, including parapodia. The first parapodium (fig. 4) is more The paratype, which is fragmented into two elongate than the remaining parapodia. Both pieces, has a total length of 10.0 mm. and is the ventral cirrus and the notopodial lobe also approximately 2.5 mm. wide. Both spec- (not a dorsal cirrus, because it is supported imens are comprised of 32 setigerous seg- by the notoaciculum and a number of setae ments. are inserted on its dorsomedial surface) ap- DESCRIPTION: The prostomium (fig. 1), pear to insert on the parapodium by means which is slightly longer than wide, bears a of an articulation. The ventral cirrus is pair of tapering palpi on its anterior margin. longer, and it tapers to a somewhat capitate The bases of these are but barely separated tip, while the dorsal lobe tapers to a rounded at the point of insertion. Just dorsal to the tip. The parapodium is supported by a pair 174 1962 JONES: POLYCHAETOUS ANNELIDS 175

FIGS. 1-6. Pareulepis sulcatisetis, new species. 1. Dorsal view of prostomium, tips of third and fourth parapodia omitted. Central dashed line in posterior part of prostomium indicates point of fusion with base of first elytrophore; dotted line, anterior overhang of base of first elytrophore. 2. Dorsal view of anterior region of paratype, with proboscis extruded. Setae omitted from left side, and right first elytrum is missing. 3. Lateral and slightly dorsal view of paratype. 4. Anterior view of right first parapodium. 5. Posterior view of left second parapodium. 6. Posterior view of right sixth parapodium. 1-3, scale A; 46, scale B.

of acicula, with the notoaciculum extending Ventral cirri are present from the first into the base of the dorsal lobe and the neuro- parapodium (fig. 4) and in the anterior region aciculum recurving dorsally near the tip of (fig. 5) are relatively long and narrow and the neuropodial lobe. have Icapitate tips. By the sixth parapodium The second parapodium (fig. 5) has the (fig. 6) they are quite short, less than one- general shape of succeeding parapodia, i.e., fourth of the length of the ventral cirrus of the notopodial lobe is rather rounded and the the second parapod, and are composed of two neuropodial lobe is somewhat flattened in a parts, a round, portion and a narrow, transverse plane. In the second parapodium apical piece. The ventral cirri tend to be in- the neuropodial lobe shows a rounded lateral serted on the posterior face of the parapodia. margin, and it is supported by a heavy acic- Dorsal cirri first appear on the third para- ulum which is inserted upon the anterior podium as pyriform structures (fig. 3) which face of a flattened chitinous plate. In more are laterally situated, near the fascicle of posterior parapodia the acicular plate be- heavy notosetae. After two setigers with comes elongated, and the aciculum comes to elytra, they are next present on the sixth be inserted in the mid-region of the plate. parapodia and are more medially placed, in The anterior notoacicula have straight tips, line with the elytrophores of the fifth and but the tips of those more posterior are seventh parapodia (fig. 7). In addition to the ventrally recurved. dorsal cirrus the sixth parapod also bears a 176 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124

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0 AIMM 0 B 1.0MM. I..I I 1 I I I I FIG. 7. Pareulepis sulcatisetis, new species, dorsal views of right dorsal cirri. A. Of sixth setiger. B. Of tenth setiger. C. Of fourteenth setiger. D. Of twentieth setiger. E. Of twenty-fifth setiger. F. Of twenty-eighth setiger. G. Of thirtieth setiger. H. Of thirty-second setiger. 7A-F, scale A; 7G, H, scale B. pyriform structure on the posterior face of one anterior and one posterior to the lateral the notopodial lobe. Dorsal cirri posterior to series. The lateral elytral processes are gen- the sixth parapodium become larger, so that erally clavate, but those of the second and by the tenth, the dorsal cirrus is as long as third elytra (fig. 9) show a constriction mid- the parapodium. It is here a broad lobe, the way along their length, and some of those of sides of which are extended into wide margins the last (twelfth) elytrum (fig. 11) are actu- and the generally pyriform tips of which ally tripartite and articulated. No micro- appear to be articulated to the base. The structures were seen on the dorsal surfaces of dorsal surfaces of these cirri are smooth, but the elytra, and they are not pigmented. their ventral surfaces bear two or three With the exception of the first, the para- ridges which appear to be ciliated. By the podia of Pareulepis sulcatisetis bear the same twenty-fifth dorsal cirri (fig. 7), the lateral types of setae, with some minor structural limbations are less obvious, and those more variations along the length of the . posterior show only flattened, articulated, The first parapodium bears three types of pyriform tips of decreasing size (fig. 7). capillary setae. Two of the three types origi- Pareulepis sulcatisetis has 12 elytra which nate medial to the cirrus-like notopodial lobe are inserted on parapodia 2, 4, 5, 7, 9,... 19, on the posterior face of the first parapod. 21, and 24. The first elytrum (fig. 8) is sub- The setae of the more proximal fascicle circular, with relatively straight medial and (fig. 12) are abruptly bent at approximately posterior margins. Several papillar processes the mid-point of their length, and there is a are inserted on the dorsal surface along the short series of serrations just proximal to the anterior edge. The second to fourth elytra bend. Those of the second type (fig. 13), (fig. 9) increase both in length and width. inserted just distal to the first, are gently From the fifth to twelfth (figs. 10 and 11), curved and show a longer series of serrations the width remains approximately the same, in the mid-region. The remaining setae of but there is a gradual lengthening of elytra this parapodium are all smooth, thin capil- to the twelfth, which is the longest. Pairs of laries, and arise from three areas of the elytra nearly meet at the mid-line, and the neuropodial lobe: from the anterior face, posterior edge of the last pair extends poster- dorsal to the recurved neuroaciculum; from iorly to cover the twenty-eighth parapodia; the posterior face, ventral to the neuroacicu- the last four pair of parapodia and the lum; and from the anterior face, near the pygidial structures are visible from above. base of the ventral cirrus. All elytra but the first have a series of from On all of the remaining parapodia, a three to 10 processes inserted dorsally along fascicle of many capillaries is found on a lobe their lateral margins (figs. 9, 10, and 11). In just ventral to the notoaciculum, on the addition to these lobular structures, the bases posterior parapodial face. Two types occur of which are constricted at their point of here, thin smooth capillaries and serrate attachment, the free margin of the elytron is capillaries (fig. 14) which are similar to one drawn out into an additional pair of processes, of the capillary types of the first parapod 0-

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26 FIGS. 8-27. Pareulepis sulcatisetis, new species. 8. Dorsal view of right first elytrum. 9. Dorsal view of right third elytrum. 10. Dorsal view of right eighth elytrum. 11. Dorsal view of right twelfth elytrum. 12. Proximal notoseta from first parapodium. 13. Distal notoseta from first parapodium. 14. Serrate capillary notoseta. 15. Pointed, bent notoseta. 16. Spatulate, bent notoseta. 17. Surface view of spatulate tip of figure 16. 18. Gouge-tipped, bent notoseta. 19. Dorsal neuroseta from an anterior setiger. 20. Dorsal neuroseta from a posterior setiger. 21. Dorsal limbate neurosetae. 22. Ventral limbate neuroseta. 23. Scoop-tipped, bent neuroseta. 24. Posterior limbate neuroseta. 25. Far posterior, scoop-tipped, bent neuroseta. 26. Proximal region of anal cirrus. 27. Distal region of anal cirrus. 8-11, scale A; 12-25, scale B; 26, 27, scale C. 178 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 (fig. 13). Another type of seta occurs on the with the same distribution of fine teeth as on anterior face of the notopodial lobe. In those setae of the mid-region, and the ventral anterior parapodia the latter change from unilimbate setae remain unchanged. bilimbate pointed setae in the second para- The only remaining group of setae is to be podia, to heavy to stout, sharply bent setae, found on the posterior face of the parapodium, the tips of which are either pointed (fig. 15) lateral to the ventral cirrus. These thin, or have a spatulate tip (figs. 16 and 17). The smooth capillaries are present on all para- latter, which are found from the third para- podia from the second to the next to last. podium to at least the twenty-fifth para- The single anal cirrus (figs. 26 and 27) podium, have smooth shafts. By the twenty- arises from a small, knob-like cirrophore on ninth parapodium these setae have become the right side. The left cirrophore is also a further modified, and in the last four para- knob-like structure, but it shows no evidence podia of Pareulepis sulcatisetis, they are of bearing, or having borne, a cirrus. The heavy and sharply bent, with fine teeth on anal cirrus is ornamented along its length by the convex surface of the bend (fig. 18). a series of small papillar structures, some of The shaft, distal to the bend, has the appear- which are similar in shape to the posterior ance of a gouge chisel, the margin of which ventral cirri, i.e., there is a bulbous basal bears more fine teeth. portion and a more slender terminal portion. The most dorsal of the neurosetae is a These appear to be distributed in a single row heavy, pointed seta which carries a sub- along the cirrus; on the basal one-tenth of terminal, pectinate row of 12-14 teeth. The its length, these structures are relatively more proximal teeth are longitudinally di- small and numerous (fig. 26). More distally, rected, and the more distal are nearly at they are less numerous and are cirriform (fig. right angles to the setal axis. In anterior 27). parapodia, this seta is straight (fig. 19), but The trivial name, sulcatisetis, refers to the from the twenty-first parapodia, this seta is peculiar gouge-like setae of the posterior bent (fig. 20). Usually there is but one of region. these pectinate setae per parapodium, but TYPE DISPOSITION: The holotype (A.M.- occasionally there may be two. They are N.H. No. 3605) and the paratype (A.M.N.H. found on all parapodia from the second to No. 3606) are both deposited in the collec- the next to last. In anterior parapodia the tions of the American Museum of Natural remaining heavy neurosetae are all pointed History. and appear to be of the same type, with a DISCUSSION: Previous to the present writ- grading of thickness of shaft from the heavier, ing, eight species of Pareulepis had been pro- more dorsal setae (fig. 21) to somewhat more posed: P. hamifera (Grube), 1875, the type slender ventral setae (fig. 22). There also species, from the Philippines; P. challengeriae seems to be a tendency for the more dorsal (McIntosh), 1885, from a "Challenger" sta- of these to be bilimbate and the more ventral tion near Sombrero Island, the West Indies; to be unilimbate. Through the mid-region the P. wyvillei (McIntosh), 1885, from another difference in limbation becomes obscure, and "Challenger" station near Bermuda; P. it is only at approximately the twenty-fifth fimbriata (Treadwell), 1901, and P. splendida parapodium that two setal types finally (Treadwell), 1901, both from "Fish Hawk" emerge. The dorsalmost of these setae are stations near Puerto Rico; P. malayana here rather sharply bent (fig. 23), and their (Horst), 1913 (and Horst, 1917), from a tips have become rather scooped (similar to "Siboga" station near Great Kei Island; the gouge-chisel type of the posterior noto- P. geayi (Fauvel), 1918, from Madagascar; podia, fig. 18). There are fine teeth on the and P. weberi (Horst), 1922, from Curagao. convex surface of the bend, but the terminal However, Augener (1906, p. 129; 1918, p. portion is without denticulations of any kind. 156) has suggested that P. splendida is The more ventral setae in this region are synonymous with P. wyvillei (McIntosh), gently recurved and are unilimbate (fig. 24). 1885, and the suggestion has been confirmed Posterior to the twenty-ninth parapodia, the by Hartman (1959); thus, there were seven scooped neurosetae become heavier (fig. 25), recognized species of Pareulepis. 1962 JONES: POLYCHAETOUS ANNELIDS 179 The most obvious initial separation of Pareulepis sulcatisetis can be separated species in this genus is concerned with the from P. fimbriata and P. geayi on the basis number and the marginal ornamentation of of the following criteria: (1) the tripartite the elytra. Pareulepis weberi is the only species marginal processes of the twelfth elytra (fig. with 36 elytra rather than 12; P. challengeriae 11); (2) the presence of two pairs of posterior is the only species of which the elytral mar- and one pair of central eyes (fig. 1); (3) gins are entire; and the elytral margins of P. the free ends of anterior, heavy, bent noto- hamifera (Grube, 1878, pp. 52-53), P. wyvil- setae (fig. 17) are provided with spatulate lei, and P. malayana are merely notched, and tips (note, however, that Hartman [1939, are not provided with dependent processes. p. 144] states, in a figure legend, that the free Pareulepis fimbriata, P. geayi, and P. sul- end of these setae in P. fimbriata are flat- catisetis all exhibit such dependent processes. tened and depressed; she makes no further As Augener (1918, p. 153), and Fauvel mention of them, nor does she figure them), (1919b) in his second description of P. geayi, and those of the posterior region are gouge- have pointed out, Treadwell's description of like (fig. 18); and (4) there are only 32 P. fimbriata is rather brief. Fauvel (1919b) setigerous segments (rather than 33 to 39, and Hartman (1939) have presented further as in P. fimbriata, or 36 to 38, as in P. geayi). comments and observations on P. fimbriata; FAMILY AMPHINOMIDAE SAVIGNY, 1818 the latter suggested that P. geayi is synony- mous with P. fimbriata (Treadwell) 1901. GENUS CHLOEIA SAVIGNY, 1818 Recently, Day (1951 and 1957) and Tebble Chloeia viridis Schmarda, 1861 (1955) have maintained that P. geayi is a Chloeia viridis SCHMARDA, 1861, pp. 144-146; valid species. They base this contention on Jamaica; Caribbean region (= Antillenmeer?). the posterior, or at least central, placement Chloeia euglochis EHLERS, 1887, pp. 18-24; of the unpaired, median, prostomial antenna [Dry] Tortugas, Florida; Cape Florida [?]. (as contrasted with the extreme anterior Chloeia modesta EHLERS, 1887, pp. 21-23; location in P. fimbriata) and the lack of eyes [Dry] Tortugas, Florida; Cape Florida [?]. P. are eyes, two Chloeia euglochis, TREADWELL, 1901, p. 194; (in fimbriata, there three Puerto Rico. posterior and lateral, and a single one medial Chloeia euglochis, AUGENER, 1906, p. 96; and anterior). The absence of eyes may well Dominica; Barbados. be of questionable validity, for Fauvel, in Chloeia candida KINBERG, 1910, p. 33; St. his subsequent description (1919b, p. 338), Thomas. mentions that a smaller specimen of P. geayi Chloeia euglochis, AUGENER, 1922a, p. 51; carries, on the lateral posterior borders of the "West Indies." prostomium, very small black eyes, one on Chloeia viridis, MONRO, 1933a, pp. 1-10; each side, which are not visible dorsally. He Taboga Island, Panama (Pacific side). makes no mention of an anterior medial eye. Chloeia euglochis, TREADWELL, 1939, pp. 176- 177; "Tortugas, Cape Florida [?], and Puerto It is also well to note that Fauvel has ap- Rico." parently contradicted himself as regards the Chloeia viridis, HARTMAN, 1942b, p. 96; Bahia number of anal cirri of P. geayi. In his orig- de Cochinos, Cuba. inal description (1918, p. 504), he states Chloeia viridis, HARTMAN, 1951b, p. 29; south- that there are two long filamentous anal ern Florida. cirri, but in the subsequent treatment (1919b, Chloeia viridis, ANDREW AND ANDREW, 1953, p. 338, paragraph 2), he comments that there pp. 3-4; Bimini, Bahamas. is a single filiform cirrus (its counterpart on Chloeia viridis, RENAUD, 1956, pp. 7-8; Fort the other side is degenerate) and later (p. Lauderdale and Biscayne Bay, Florida (Atlantic 338, paragraph 3), that the anal cirrus is side). clearly unpaired. U.C.W.I. COLLECTION: A single specimen Apparently the latter-day discussions con- of 30 setigers (length, 45 mm.; width, 8 mm.) cerning P. geayi (Fauvel) have convinced was collected at Port Royal, Jamaica, on Hartman of its validity, for she has recently November 18, 1959. It was swimming free listed it as a valid species of Pareulepis in the surface waters. (Hartman, 1959, p. 123). DISTRIBUTION: Subtropical to tropical: 180 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 from south Florida throughout the Greater Syllis (Typosyllis) catenula VERRILL, 1900, p. and Lesser Antilles; Pacific side of Panama. 604; Bermuda. Syllis jugularis VERRILL, 1900, p. 606; Ber- FAMILY HESIONIDAE MALMGREN, 1867 muda. GENUS HESIONE SAVIGNY, 1818 Typanosyllis [sic] fertilis VERRILL, 1900, pp. 616-617; Bermuda. Hesione picta Muller, 1858 Typosyllis corallicola, TREADWELL, 1924b, p. Hesione picta MtLLER, 1858, p. 213; St. 10; Pelican Island, Antigua. Catherine Island, Brazil [approximately latitude Syllis corallicola, MONRO, 1933c, pp. 249-250; 270 S.]. Dry Tortugas, Florida. Hesione proctochona SCHMARDA, 1861, pp. 79-80; Typosyllis corallicola, TREADWELL, 1936, p. 50; Jamaica. Bermuda. Fallacia proctochona, WEBSTER, 1884, p. 31 1; Syllis (Typosyllis) corallicola, TREADWELL, Bermuda; Sarasota Bay to Key West, Florida. 1939, pp. 212-213; near Culebra Island, Puerto Hesione vittigera EHLERS, 1887, pp. 143-147; Rico (latitude 180 19' N., longitude 65° 19' W.). Key West, Florida. Typosyllis corallicola, HARTMAN, 1942a, pp. Hesione proctochona, TREADWELL, 1901, p. 187; 47-48; Bermuda. Puerto Rico. A single ovigerous Fallacia protochona [sic], VERRILL, 1901, p. 39; U.C.W.I. COLLECTION: Bermuda. specimen of 198 setigers, 64 mm. in length Hesione proctochona, HOAGLAND, 1919, p. 571; and from 1.5 to 2.0 mm. in width, was col- Bermuda. lected from sponges at Kingston Harbor, Hesione proctochona, AUGENER, 1922a, p. 52; Jamaica, on January 29, 1960. "West Indies." DISCUSSION: The original description of Hesione proctochona, HORST, 1922, pp. 200-201; this species was not accompanied by figures, Curagao. and it remained for Monro (1933c) and Hart- Hesione proctochona, AUGENER, 1927b, p. 49; of certain Curagao. man (1942a) to present figures Hesione proctochona, TREADWELL, 1928, p. 473; structures of this species. Certain other fea- near Saba Island. tures have been noted in the present study Hesione proctochona, AUGENER, 1933a, p. 224; which make it desirable to present further Santa Marto, Colombia; Isla de Margareta, figures of Typosyllis corallicola. Venezuela. The prostomium is basically oval, with an Hesione proctochona, TREADWELL, 1939, pp. extensive, wide cleft in the posterior margin 217-218; "Key West; Bermuda; west Florida; (fig. 28). Of the two pairs of eyes, the anterior near Saba Island; Puerto Rico." pair are the larger and are somewhat more Hesione picta, HARTMAN, 1951b, p. 35; south- laterally placed than the posterior pair. western Florida. Both pairs have lenses. The paired prostomial U.C.W.I. COLLECTION: Five specimens, all palps are separate along their medial length of 17 setigers (lengths are 28, 27, 23, 16, and and have somewhat swollen bases. The un- 8 mm.), come from Kingston Harbor, paired median prostomial antenna is nearly Jamaica. The collection was made on Janu- four times the length of the palps and is ary 29, 1960. moniliform, with approximately 50 articles. DISTRIBUTION: Subtropical to tropical: The two lateral prostomial tentacles are from Florida throughout the Greater and nearly two-thirds of the length of the medial Lesser Antilles; northern coast of South and consist of approximately 35 articles. America. Dorsal and ventral peristomial cirri are com- FAMILY GRUBE, 1850 prised of approximately 50 and 24 articles, respectively. GENUS TYPOSYLLIS LANGERHANS, 1879 Dorsal cirri are also moniliform and alter- Typosyllis corallicola (Verrill), 1900 nate regularly in the anterior and posterior Figures 28-40 regions, both in length and in number of Syllis (Typosyllis) corallicola VERRILL, 1900, articles (figs. 29, 30, 33, and 34). In the mid- p. 603; Bermuda. region, consecutive dorsal cirri are of ap- Syllis (Typosyllis) corallicola var. lineolata proximately the same length and have ap- VERRILL, 1900, p. 604; Bermuda. proximately the same number of articles 1962 JONES: POLYCHAETOUS ANNELIDS 181

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FIGS. 28-39. Typosyllis corallicola (Verrill). 28. Dorsal view of anterior region, some dorsal cirri on right side lacking. 29. Posterior view of right fourteenth setiger. 30. Anterior view of right fifteenth setiger. 31. Posterior view of left seventieth setiger. 32. Anterior view of left seventy-first setiger. 33. Anterior view of left 110th setiger. 34. Anterior view of left llth setiger. 35. Posterior view of right 170th setiger. 36. Anterior view of right 171st setiger. 37-39. Three compound setae from left seventy-first setiger. 28, scale A; 29-36, scale B; 37-39, scale C. (figs. 31 and 32). The ratio of number of from three to four acicula in those of the articles in dorsal cirri of anterior setigers is anterior and middle regions and only a single approximately 1:3, but in more posterior aciculum in the more posterior parapodia. In setigers it is closer to 1:2. the posterior parapodia (figs. 35 and 36), Parapodia are uniramous, and there are dorsal cirri are shorter, and there are second- 182 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124

0 0.2 MM. § 4 4 4 45 4 4

4 4~~~~~~~~~ Ef H

Ii 4' K IN LJ M\J 0I FIG. 40. Typosyllis corallicola (Verrill), blades of compound setae. A. From setiger 10, dorsal. B. From setiger 10, ventral. C. From setiger 10, ventral. D. From setiger 15, dorsal. E. From setiger 15, ventral. F. From setiger 61, ventral. G. From setiger 71, dorsal. H. From setiger 71, dorsal. I. From setiger 71, ventral. J. From setiger 111, ventral. K. From setiger 111, ventral. L. From setiger 170, dorsal. M. From setiger 170, ventral. N. From setiger 170, ventral. 0. From setiger 171, dorsal. ary acicula embedded in the parapodia be- pigment streaks are the shorter; those more tween the dorsal cirri and the setal bundle. posterior extend from the cirrophore of one This has been noted in setigers posterior to side to that of the other (fig. 28). number 167 (of 198 setigers). In all cases Monro (1933c) has found that T. corallicola ventral cirri are smooth and tapered, and the from Dry Tortugas exhibits dorsal cirri which single pair of anal cirri are similar to the alternate in length (approximately 65 and 35 dorsal cirri of the mid-region. articles) and that setae from the anterior Setae are all composite; the basal shafts region have longer blades than those from are all similar, and the general structure of the middle and posterior regions. Hartman the blades is constant throughout, i.e., they (1942a), on the other hand, has found, in are bidentate falcigers with fine serrations viewing Verrill's types, that the alternating along the cutting edge (figs. 37 to 39). dorsal cirri have somewhat fewer articles However, the length of the blade varies (approximately 40 and 25) and that superior among setae of the same segment and from setae, apparently regardless of position along parapodium to parapodium. Generally, the the length of the body, have longer blades blades of those setae that are more dorsally than those more ventrally placed. Present placed (fig. 40A, D, G, H, L, and 0) tend to observations agree with those of Hartman, be longer than those that are more ventral except that longer and shorter blades may be (fig. 40B, C, E, F, I, J, K, M, and N). There intermixed to some extent, although the is also variation in the size of the apical longer blades do tend to be more superior. teeth, as well as in the distance that separates Hartman (1942a, p. 48), in her account of them (fig. 40). No rigid pattern of distribu- Verrill's types, also points out that Trypano- tion is apparent in regard to this latter varia- syllis fertilis might be a specimen of Typo- tion. syllis corallicola, but questions this on the The dorsum of the anterior quarter is basis of a larger number of acicula (three to flecked with minute reddish brown spots. four) present in parapodia of the former. On These usually are found in two rows on the the basis of my observations on the Jamaican dorsum of a given segment. The anterior specimen of T. corallicola, I confirm her sug- 1962 JONES: POLYCHAETOUS ANNELIDS 183 gestion of synonymy, for the number of from either of the two types present in G. acicula here varies from three to four in abranchiata. anterior setigers, to one in posterior setigers. As a further extension of Treadwell's DISTRIBUTION: Tropical: Bermuda; Key description of this species, figures of a para- West; throughout the Antillean chain from podium (fig. 47) and of a composite seta Jamacia to Antigua. (fig. 48) are presented. The parapodia are supported by two acicula, the tips of which FAMILY GLYCERIDAE GRUBE, 1850 extend barely beyond the parapodial lobes. A dorsal cirrus is inserted at the base of the GENUS GLYCERA SAVIGNY, 1818 parapodium, and the triangular ventral Glycera abranchiata Treadwell, 1901 cirrus is broadly inserted on the posterior Figures 41-48 surface of the ventral parapodial margin. Glycera abranchiata TREADWELL, 1901, pp. There are two presetal lobes, which are 200-201; Puerto Rico. somewhat longer than those shown for G. Glycera abranchiata, TREADWELL, 1924b, p. 14; tesselata by either Ehlers (1864-1868, pl. 24, Antigua. fig. 33) or Fauvel (1923, fig. 152c). Notosetae Glycera abranchiata, TREADWELL, 1928, p. 473; are simple, and neurosetae are composite. near Saba Island (latitude 170 39' N., longitude The blades of the composite setae (fig. 48) 630 17' W.). have a gentle sigmoid curvature and are ser- Glycera abranchiata, TREADWELL, 1939, pp. rate along one margin. 260-261; "Puerto Rico and near Saba Island." DISTRIBUTION: Tropical: Jamaica, Puerto U.C.W.I. COLLECTION: One incomplete Rico, and the Lesser Antilles. specimen of 109 setigers, 31 mm. long and 1.5 mm. in diameter, was collected at Ocho FAMILY EUNICIDAE SAVIGNY, 1818 Rios, Jamaica, on August 11, 1957. GENUS EUNICE CUVIER, 1817 DISCUSSION: This specimen agrees with the description given by Treadwell (1901) ?Eunice tridentata Ehlers, 1905 and with specimens in the collections of the Figures 49-51 American Museum of Natural History which Eunice tridentata, MONRO, 1933a, pp. 63-65; were identified by Treadwell (A.M.N.H. Coiba Island, Panama (Pacific end of the Panama Nos. 561, 907, 1280, and 2633). Treadwell, Canal). in his description of this species, did not Eunice tridentala, MARSDEN, 1960, p. 996; comment on the structure of the proboscidial Barbados. organs, and this omission led Augener U.C.W.I. COLLECTION: A single posterior (1922b, p. 205), Horst (1922, p. 201), and fragment of 125 setigerous segments (95 mm. Hartman (1950, pp. 77-78) to synonymize long and 3 mm. wide at the anterior end) this species with Glycera tesselata Grube. was collected at Lime Cay, Jamaica, on Observations of the proboscidial organs of August 7, 1960. this specimen show that two types are pres- DISCUSSION: The anterior 55 setigers (ap- ent. The first, and more common (figs. 41 and proximately 65 mm.) comprise a greenish 42), is rather slender, with a number of ovigerous section; the remainder of the speci- oblique ridges and a constriction at its base. men is reddish and is terminated by a single The second is larger and more oval in "frontal" pair of anal cirri. Ventral cirri are approxi- view (fig. 43) and from the "side" shows one mately one-half of the length of the single surface to be flat and the other to be convex dorsal cirri. No branchiae were seen on this (fig. 44). This latter type has no ridges. posterior fragment. Subacicular hooks are Proboscidial organs of a museum specimen brownish yellow and have bidentate tips (fig. (A.M.N.H. No. 907) are nearly identical 49). Composite setae have short falcigerous (figs. 45 and 46). Hartman (1950, pl. 10, blades which have bidentate tips (fig. 50). fig. 11) shows the proboscidial organ of G. The subapical tooth is often somewhat ob- tesselata to be a relatively long, narrow scure, and the cutting edge of blades is oc- structure with a non-constricted base and casionally eroded. Pectinate setae (fig. 51) longitudinal ridges. This is quite different are symmetrical and are provided with ap- 184 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124

41 42 44 43 01 45

48

B

O.S MM.- FIGS. 41-48. Glycera abranchiata Treadwell. 41. "Frontal" view of smaller proboscidial organ. 42. Profile of figure 41. 43. "Frontal" view of larger proboscidial organ. 44. Profile of figure 43. 45. "Frontal" view of smaller proboscidial organ of A.M.N.H. No. 907. 46. "Frontal" view of larger probo- scidial organ of A.M.N.H. No. 907. 47. Posterior view of right twentieth parapodium. 48. Compound seta from twentieth parapodium. FIGS. 49-51. ?Eunice tridentata Ehlers. 49. Posterior subacicular hook. 50. Posterior compound seta. 51. Posterior pectinate seta. FIGS. 52-54. Lysidice ninetta Audoin and Milne-Edwards. 52. Subacicular hook from 100th setiger. 53. Compound seta from second setiger. 54. Pectinate seta from fiftieth setiger. 41-46, 48-54, scale A; 47, scale B. proximately 16 subequal terminal teeth and off Tortugas; latitude 230 03' N., longitude 830 10' a lateral pair of teeth which are somewhat 05' W. (off the north coast of Cuba, west of longer. The subacicular hooks and composite Havana). setae Hart- Lysidice bilobata VERRILL, 1900, p. 645; agree with figures presented by Bermuda. man (1944, pl. 7, figs. 146, 148, and 149), Lysidice notata, TREADWELL, 1921, pp. 86-88; but the pectinate setae differ, in that there Key West; Bermuda; Dry Tortugas. are two longer teeth at each end of the row of Lysidice notata, AUGENER, 1922a, p. 52; "West teeth rather than a single longer tooth at one Indies." end. Lysidice ninetta, AUGENER, 1927b, p. 62; DISTRIBUTION: Tropical: scattered through- Curagao. out the Caribbean (Jamaica, Barbados, and Lysidice notata, TREADWELL, 1928, p. 477; Pacific side of Panama). near Saba Island. Lysidice ninetta, AUGENER, 1933b. pp. 143-144; GENUS LYSIDICE SAVIGNY, 1818 Jamaica; Curagao; Santa Marta, Colombia. Lysidice ninetta Audouin and Lysidice ninetta, MONRO, 1933a, pp. 70-71; Milne-Edwards, 1833 Taboga Island; Coiba Island; Balboa; Gorgona Island (all on Pacific side of Panama). Figures 52-54 Lysidice ninetta, RIOJA, 1946, p. 194; Veracruz, Lysidice brachycera SCHMARDA, 1861, p. 121; Mexico. Jamaica. Lysidice ninetta, HARTMAN, 195 1b, p. 58; "Vera- Lysidice notata EHLERS, 1887, pp. 100-102; cruz, Mexico." 1962 JONES: POLYCHAETOUS ANNELIDS 185 Lysidice ninetta, ANDREWS AND ANDREWS, are continued posteriorly as strap-like struc- 1953, p. 10; Bimini, Bahamas. tures with rounded tips. U.C.W.I. COLLECTION: An anterior por- The most common setal type throughout is tion of a single individual, in two fragments a unilimbate capillary (fig. 57) which is (total length, 58 mm. for 103 setigers), comes found in nearly all notopodia and in anterior from the lagoon at Port Royal, Jamaica. neuropodia, including the first setiger, which The specimen was collected on January 29, bears neurosetae only. The modified setae of 1960. the fifth setiger (fig. 58) are bidentate and DISCUSSION: Subacicular hooks are dark bear a subterminal collar which extends and bidentate, with a terminal sheath (fig. halfway around the circumference (fig. 59). 52). Composite setae (fig. 53) are provided These three to four special setae are accom- with falcigerous blades which are bidentate panied by several short, plain, pointed capil- and sheathed. Pectinate setae (fig. 54) are laries and a few flattened, pointed, trans- asymmetrical and have about 25 subequal parent, lanceolate setae (figs. 58 and 60). terminal teeth and a longer tooth on one side. The sixth setiger bears only neuropodial and DISTRIBUTION: Tropical: throughout the notopodial fascicles of unilimbate capillaries Caribbean area from Bermuda and Dry (fig. 57) and the seventh, as well as all re- Tortugas, through the Greater and Lesser maining setigers, bears neuropodial hooks in Antilles, to the northern coast of South addition to the unilimbate capillaries. The America; Mexico on the Gulf of Mexico; on hooks (fig. 61) are hooded and have one the Pacific end of the Panama Canal. large tooth, with a somewhat smaller ter- minal tooth on the most distal part of the FAMILY SPIONIDAE GRUBE, 1850 seta. In the pygidial region still another GENUS POLYDORA Bosc, 1802 setal type is to be found. On the last two to Polydora ancistrata, new species four setigers, neuropodial fascicles are pro- vided with special simple falcate hooks (figs. Figures 55-65 62-64), and the unilimbate capillary setae of DIFFERENTIAL DIAGNOSIS: Polydora with the more anterior setigers here have only a posterior notosetae modified as slightly re- narrow limbation (fig. 65). The special hooks curved falcate hooks; with special setae of are oriented with their tips pointing from the fifth setiger provided with a subterminal dorsal to posterior. collar and bifid tips; and with no notosetae The anus is dorsal, and the pygidium is a on the first setiger. flattened, subcordate structure posterior to U.C.W.I. COLLECTION: Numerous indi- the anal opening (fig. 62). viduals come from Port Royal, Jamaica. The trivial name, ancistrata, refers to the They were collected in May, 1960, and were shape of the notopodial falcate setae found associated with an unidentified sponge. Adult in the posterior region of this species. specimens are 6 mm. in length, approximately TYPE DISPOSITION: The holotype (A.M.- 0.25 mm. wide, and have about 45 setigerous N.H. No. 3607) and paratypes (A.M.N.H. segments. No. 3608) of Polydora ancistrata are deposited DESCRIPTION: The prostomium (fig. 55) in the collections of the American Museum is subtriangular from above and is slightly of Natural History. wider than long. The anterior tip is blunt, DISCUSSION: Of the approximately 50 and the medial region is slightly raised in a species of Polydora, the majority are pro- caruncle and extends back, with nearly vided with unmodified posterior notosetae. straight sides, to between the prostomial Eight species have posterior notosetae which palpi. The palpi in preserved specimens are are somewhat modified in that they are rela- about as long as the first 12 setigerous seg- tively heavy and awl-shaped or needle-like ments (fig. 56); however, in life they may be (P. armata Langerhans, 1880 [see also Fauvel, longer and their appearence here may be a 1927, and Hartman, 1941]; P. caeca (Oersted), fixation artifact. There are no eye spots 1843 [also Fauvel, 1927 and 1953];P. cardalia associated with the prostomium. Berkeley, 1927; P. caulleryi Mesnil, 1897 Branchiae begin on the seventh setiger and [also Fauvel, 19271; P. magna (Berkeley), 186 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124

-o

6f

-A

-o

-1.0MM. C

63

-0.I MM. FIGS. 55-65. Polydora ancistrata, new species. 55. Dorsal view of anterior region, palpi missing. 56. Dorsal view of anterior region of intact specimen. 57. Unilimbate capillary seta from setiger 6. 58. Setal complement of setiger 5. 59. Modified seta from setiger 5. 60. Transparent lanceolate seta from setiger 5. 61. Neuropodial hooded hook from setiger 20. 62. Dorsal view of posterior region. 63. Posterior, special falcate hook. 64. Second, posterior, special falcate hook. 65. Posterior unilimbate capillary seta. 55, 56, 62, scale A; 57, 61, scale B; 58-60, 63-65, scale C.

1927; P. quadrilobata Jacobi, 1883 [also P. colonia has a slightly emarginate prosto- Fauvel, 1927]; and P. saint josephi Eliason, mium (the prostomium of P. ancistrata is 1920). bluntly rounded) and bears notosetae on the The last group, into which P. ancistrata first setigerous segment (P. ancistrata has falls, has posterior notosetae which are only non-setigerous notopodial lobes on the modified as falcate hooks, the tips of which first setiger). In addition, there are differ- may be more or less recurved. Polydora ences in the morphology of the hooded hooks ancistrata can be separated, on the structure and in the relative length of the apical teeth of the special setae of the fifth setigerous seg- of the special setae of the fifth setiger. Poly- ment, from P. hamata Webster, 1879, and dora ancistrata and P. hoplura inhaca are P. hamata Langerhans, 1880 (the special more similar morphologically, and the pri- setae are simple and pointed in both), and mary difference lies in the shape of the modi- from P. hoplura Claparede, 1869 (the special fied posterior notosetae. In P. hoplura inhaca setae are bifid, but lack the subterminal col- these are strongly recurved, like a fish-hook, lar). and they lack an enlargement in the mid- Polydora colonia Moore, 1907, P. hoplura region, while in P. ancistrata they are but inhaca Day, 1957, and P. ancistrata all have barely recurved and show an enlargement in the same type of subterminal collar on the the mid-region. special setae of the fifth setiger. However, It will be noted above that a homonymy 1962 JONES: POLYCHAETOUS ANNELIDS 187 exists in the case of Polydora hamata Webster, setiger (fig. 68) the branchiae are long, gently 1879, and Polydora hamata Langerhans, 1880. tapered, and are basally fused with the For some time this has been recognized, for dorsal lamellae. The dorsal lamellae are asym- Mesnil (1896, p. 235) mentioned it and metrically cordate, and the ventral lamellae pointed out that the two species appeared to are semicircular lobes. The twentieth setiger be different. Fauvel (1927, p. 50) listed them (fig. 69) bears branchiae and dorsal lamellae as "? P. hamata Webster, Langerhans 1880" similar to those on the fifth, but at this level, in his synonymy of Polydora hoplura, and ventral lamellae are two-parted. The more Hartman (1959, p. 384) listed P. hamata dorsal of these is the smaller, and both are Langerhans as a homonym of P. hamata rounded. The thirtieth setiger shows elon- Webster. Inspection of the original descrip- gate, tapered branchiae fused to the bases of tions and figures (Webster, 1879; Langerhans, the dorsal lamellae (fig. 70), which are here 1880) confirms Mesnil's comments and re- subtriangular in shape. The two parts of the veals that there are major differences in the ventral lamellae are to be found on each side structure of the pygidium, of the special of the fascicle of hooded hooks; the ventral setae of the fifth setiger, and of the modified part is rather strap-like, with a rounded tip, posterior notosetae. Therefore, I propose a and the dorsal part is broadly attached be- new name, Polydora posthamata, for Polydora tween the notopodial and neuropodial setae, hamata Langerhans, 1880. and there is the suggestion of a dorsal at- GENUS NERINIDES MESNIL, 1896 tenuation. The dorsal lamellae of the forty- fifth setiger (fig. 71) are still basically tri- Nerinides goodbodyi, new species angular but here are nearly bifid. The dorsal Figures 66-82 portion of the ventral lamellae is broadly DIFFERENTIAL DIAGNOSIS: Nerinides with attached and rounded; the ventral portion is no occipital prostomial tentacle; with a smaller than its more anterior counterpart, pointed prostomium and no obvious pattern and its tip is more pointed. The ventral lamel- of pigmentation; with palpi which lack a basal lae of the fiftieth setiger are represented only sheath; with bidentate hooded hooks which by the counterpart of the dorsal portion of are recurved; and with strap-like ventral cirri. the ventral lamellae of previous setigers and U.C.W.I. COLLECTION: Twelve specimens this is smaller, with a rounded ventral margin; were collected at Green Bay, Port Hender- its dorsal margin still shows a slight attenua- son, Jamaica, on August 11, 1960. Adults are tion. from 16 to 17 mm. in length (approximately It is of interest to note that, although all 60 setigers) and are nearly 0.5 mm. in diam- branchiae have essentially the same shape, eter. Several are ovigerous. there is a group of what appear to be glandu- DESCRIPTION: The prostomium (fig. 66) lar cells on the distolateral margin of the tapers to an acute point and overlies the branchiae of the mid-region (from the peristomium which is approximately one- fifteenth setiger to the thirtieth; figs. 69 and half of the length of the prostomium. The 70). posterior portion of the prostomium is set off All setae of the anterior region are pointed from the peristomium by nearly straight capillaries which bear extremely fine spine- lateral margins. The occipital portion is lets (figs. 73 to 76). Both neurosetae and slightly raised and contains four embedded notosetae are disposed in anterior (longer) eye spots. The lateral pair are the larger, and posterior (shorter) series. The anterior and all four form a straight line. A pair of notosetae (fig. 73) are non-limbate and palpi are inserted on the peristomium near gently sinuous; the posterior notosetae (fig. the base of the prostomium, just anterior to 74) are unilimbate and curved. The anterior the first setiger. The posterior portion of the neurosetae (fig. 75) are bilimbate and nearly prostomium appears to extend back onto the straight; the posterior neurosetae (fig. 76) first setiger and is not tapered. are unilimbate and abruptly curved so that The first setiger (fig. 67) bears dorsal and their apical regions are nearly parallel to the ventral post-setal lamellae only; branchiae surface of the body. begin on the second setiger. By the fifth Hooded hooks begin at about the twenty- 66

-C

-c

KI.1MM.

FIGS. 66-82. Nerinides goodbodyi, new species. 66. Dorsal view of prostomial region, right palp missing. 67. Anterior view of left first setiger, dotted line indicating contour of prostomium and black spots showing position and shape of prostomial eye spots. 68-72. Anterior views. 68. Right fifth setiger. 69. Right twentieth setiger. 70. Right thirtieth setiger. 71. Left forty-fifth setiger. 72. Right fiftieth setiger. 73-76. Fifteenth setiger. 73. Anterior notoseta. 74. Posterior notoseta. 75. Anterior neuroseta. 76. Posterior neuroseta. 77, 78. Thirty-fifth setiger. 77. Anterior notoseta. 78. Posterior notoseta. 79. Capillary neuroseta accompanying hooded hooks of fiftieth setiger. 80-82. Hooded hooks from fiftieth setiger. 80. Bidentate. 81. Tridentate. 82. Tridentate, showing details of hood structure. 66, scale A; 67-72, scale B; 73-78, scale C; 79-82, scale D. 1962 JONES: POLYCHAETOUS ANNELIDS 189 eighth setiger and, throughout the length of more proximally on the dentate side than the Nerinides goodbodyi, are confined to the other. neuropodia. The notopodial capillary setae The pygidium is a rather thick, rounded are devoid of spinelets and are provided with structure, subcordate in shape. very narrow limbations. As in the anterior It is with great plesaure that I name this segments, there are two series of setae in the spionid in honor of Dr. Ivan M. Goodbody, notopodial fascicles. The setae of both are University College of the West Indies, who, quite slender, and those of the anterior series through care and patience in the relaxation (fig. 77) are longer than those of the posterior and preservation of the present collection, series (fig. 78). has made their study all the more enjoyable. The hooded hooks are accompanied by TYPE DISPOSITION: The holotype (A.M.- non-limbate, slender, pointed capillaries (fig. N.H. No. 3609) and paratypes (A.M.N.H. 79) only in the most posterior setigers. The No. 3610) of Nerinides goodbodyi are de- hooks themselves may bear either one (fig. posited in the collections of the American 80) or two (fig. 81) apical teeth. There is no Museum of Natural History. apparent to the distribution of these DISCUSSION: Of the several species of types (table 1); however, there is a general Nerinides, four are readily separable from increase in the number of hooks per fascicle Nerinides goodbodyi on the basis of the pres- in the more posterior setigers. ence of an occipital prostomial tentacle (N. The hoods of the hooks of Nerinides good- tridentata Southern, 1914; N. cantabra Rioja, bodyi (figs. 80 and 81) nearly envelop their 1918; N. papillosus Okuda, 1937; and N. toothed tips, and the apertures of the hoods yamaguchii Imajima, 1959). In addition, two are occasionally eccentrically placed. The recently described species, N. pigmentata hoods are quite transparent, and observa- (Reish), 1959, and N. maculata Hartman, tions of hooks that are not properly orientat- 1961, have blunt and subcircular prostomia, ed suggest that the hoods are entire and have respectively, and both exhibit distinctive no cleft. However, observations with oil-im- pigmentation patterns. mersion optics show that there is actually a The remaining species of Nerinides are cleft in the hood and, further, that there is a sufficiently close to N. goodbodyi to require second structure, a transparent cylinder, be- the listing of more subtle differences. The tween the hood proper and the hook (fig. 82). original description of N. acuta (Treadwell), The distal margin of the inner "cylinder" ap- 1914, although unclear on certain points, e.g., pears to be in contact with the distal margin details of the teeth of hooded hooks (" ... of the hood and closes the gap of the cleft. obscure terminal teeth . . . "), does show that The inner cylinder is in contact with the hook there is a single ventral lamella on the an-

TABLE 1 DISTRIBUTION OF BIDENTATE (B) AND TRIDENTATE HOODED HOOKS (T) ON VARIOUS SETIGERS, LISTED FROM DORSAL TO VENTRAL IN A GIVEN FASCICLE Setiger 30 Setiger 35 Setiger 40 Setiger 45 Setiger 50 Right Left Right Left Right Left Right Left Right Left B B B T B B T T B T B T B T T T T T T T T B T B T B T T T T B B T T T B T T B T T T T T B B B T B B T B B B T T B T B B B B No. of hooks per fascicle 3 4 5 6 7 7 7 8 8 7 190 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 terior parapodia and that the dorsal lamellae sufficient to place it in this genus, as both in this region are shaped quite differently Mesnil (1896, p. 152) and Southern (1914, p. from those of N. goodbodyi (cf. fig. 69). Sub- 97) have pointed out. Quatrefages (1843, pp. sequent observations by Berkeley and Berke- 12-14) makes no mention of hooks, dorsal or ley (1941) have shown that N. acuta possesses ventral, in the first 50 setigers (in other bidentate hooded hooks which are not api- species of Nerinides, hooded hooks first ap- cally recurved, as well as a basal sheath on the pear on from the eleventh to the thirtieth palpi. Neither of these features agrees with setiger). The same point has been made in the the present findings in the case of N. good- case of N. lamellata McIntosh, 1909 (South- bodyi. In addition, the posterior parapodia of ern, 1914, p. 97), for no mention is made of N. acuta have more reduced ventral lamellae hooded hooks in McIntosh's description than the present species (cf. figs. 70-72). (1909, pp. 175-176), and his illustration of Nerinides agilis (Verrill), 1873, is quite the fiftieth parapodium of N. lamellata (pl. 6, similar to N. goodbodyi with respect to the fig. 5) shows no hooks. In addition, N. lamel- shape of the prostomium, the distribution of lata has frontal prostomial horns (McIntosh, eye spots, and distribution of hooded hooks 1909, pl. 5, fig. 7), which is in conflict with (Verrill, 1873), as well as the structure of the Mesnil's (1896, p. 119) diagnosis for Nerini- hooks (Hartman, 1942a). However, Verrill des. Mesnil stated (p. 152) that he erected the (p. 600) mentions the presence of a "... new genus, Nerinides, for the species described small papilliform ventral cirrus . . . " which, by St. Joseph (1894, p. 74) as Nerine longiros- in the posterior region, " . . . becomes more tris Quatrefages and suggested that Malaco- prominent and separate from the setigerous ceros longirostris Quatrefages was perhaps lobe." The so-called ventral cirri of N. good- synonymous with Nerine cirratulus delle bodyi are not traceable to small papilliform Chiaje. As a point of information, Mesnil structures but are homologues of the lower (1896, pp. 164-165) also suggested that portion of the ventral lamellae present on all Nerinides agilis (Verrill) was also identical anterior setigers. The illustrations presented with Nerine cirratulus. by Hartman (1942a, figs. 126, 127) are com- Before we leave the present discussion of parable to figures 70 and 72 of the present Nerinides, one last item must be considered. paper. Her figures show that the branchiae of Hartman (1959, p. 381) listed "Nerinides the median parapodia of N. agilis are some- longirostris (Quatrefages, 1843)" as "geno- what longer than those of N. goodbodyi, and type" of Nerinides Mesnil, 1896. However, a the ventral cirri are triangular, not strap-like. personal examination of pertinent literature In more posterior setigers (Hartman's fig. and a consideration of current nomenclatural 127, twelfth last parapodium of N. agilis, and procedure indicate that such cannot be the fig. 72 of the present paper, fiftieth right para- case. podium of N. goodbodyi, are comparable), it is As mentioned above, the description of seen that there is no "ventral cirrus" in N. Nerinides longirostris (Quatrefages), 1843, is goodbodyi, that the dorsal portion of the ven- inadequate, insofar as its being placed in the tral lamella is truncated, and that the genus Nerinides; thus, the taxon is unrecog- branchia is approximately twice as long as nizable on the basis of present information. the dorsal lamella to which it is fused. In erecting the genus Nerinides, Mesnil Nerinides agilis differs in that there is a tri- (1896, p. 152) stated that "Je cree ce nouveau angular "ventral cirrus," the dorsal portion genre pour l'espece que de St. Joseph . . . decrit of the ventral lamella is narrow and rounded, sous le nom de Nerine longirostris Qfg." ("I and the branchiae are not fused to the dorsal create this new genus for the species which de lamellae and are only half again as long as the St. Joseph ... described under the name of lamellae. Nerine longirostris Quatrefages"), and, later, There now remain only some comments "L'espece de de St. Joseph serait nouvelle" concerning the last two species which have ("The species of de St. Joseph is new"). If N. been designated as belonging to Nerinides longirostris (Quatrefages) is assumed to be a (Hartman 1959, p. 381). The description of nomen dubium, and St. Joseph's taxon repre- N. longirostris (Quatrefages), 1843, is not sents a new species, then a new name must be 1962 JONES: POLYCHAETOUS ANNELIDS 191 assigned to the latter. Therefore, I propose a third parapodium. They are all of about the new name, Nerinides st. josephi for the species same length, although those more posterior described by St. Joseph (1894, pp. 74-77) as tend to be somewhat shorter. Nerine longirostris Quatrefages. Further, I The setae are all unornamented capillaries, point out that this species is, ipso facto, the the shafts of which are quite heavy. In the type species of the genus Nerinides, by Mes- first seven parapodia, these setae are disposed nil's original designation (Mesnil, 1896, p. in rather widespread, fan-shaped fascicles 152). transversely oriented; from the eighth para- podium on, they are in tighter bundles which FAMILY OPHELIIDAE MALMGREN, 1867 are nearly in a frontal plane. GENUS ARMANDIA FILIppi, 1861 Eleven pairs of lateral eye spots are found beginning between the sixth and seventh Armandia nonpapillata, new species parapodia (fig. 88) and extending to between Figures 83-90 the sixteenth and seventeenth. The first pair DIFFERENTIAL DIAGNOSIS: Armandia with are smaller than the rest and are circular; the 29 setigers; with 11 pairs of lateral eye spots; remaining eye spots are elliptical. and with neither papillae on the margin of The anal funnel of A. nonpapillata is prob- the anal funnel nor anal cirri. ably approximately as long as the last three to U.C.W.I. COLLECTION: A single specimen four setigers (figs. 89 and 90). In the specimen of Armandia nonpapillata comes from Kings- the posterior margin of the anal funnel is ton Harbor, Jamaica. It was collected on rolled inward, which probably represents a January 29, 1960, and is comprised of 29 fixation artifact. There are approximately 12 setigerous segments (length, 19 mm.). It is to 13 narrow vertical bands on the funnel, approximately 1 to 1.5 mm. in diameter. and its posterior margin is entire; there are no DESCRIPTION: The prostomium is some- anal papillae and no indications that they what abruptly tapered, and its tip is swollen might have been present and lost in fixation (figs. 83 and 84). The paired nuchal organs or handling. Ventrally, it is seen that the anal are everted and form irregularly shaped funnel is cleft (fig. 90) and that there is no structures, the bases of which are generally anal cirrus. circular, anterior to, and slightly medial to The trivial name, nonpapillata, refers to the first parapodia. No prostomial eye spots the lack of marginal papillae around the open- are present. ing of the anal funnel of this species. The first parapodium bears two fan-shaped TYPE DISPOSITION: The holotype (A.M.- fascicles of capillary setae and a ventral cir- N.H. No. 3611) is deposited in the collections rus; there are neither branchiae nor dorsal of the American Museum of Natural History. cirri. The ventral cirri of the first and second DISCUSSION: It has been established that parapodia are inserted on the posterior face of Armandia nonpapillata possesses an anal the parapod. On the third setiger, and all fol- funnel which is devoid of anal cirri and anal lowing, it is more ventrally placed and is papillae. While most other species of Ar- visible from the front (figs. 85-87). The ven- mandia have both anal cirri and papillae, this tral cirri in all cases are rounded subtriangu- condition is not without precedent in the lar processes, while the dorsal cirri (figs. 85- genus, for A. exigua Kiikenthal, 1887, ex- 87), where they occur, are small globular hibits a pygidial region similar to that of A. structures which are inserted just above the nonpapillata. notopodial setal fascicle on the posterior sur- Armandia nonpapillatacan bedifferentiated face of the parapod. Dorsal cirri are found on from Armandia exigua by its 29 setigerous all parapodia from the second to the next to parapodia, its plainly visible lateral eyes, and last. Branchiae are also found, beginning on its truncated anal funnel (A. exigua has 38 the second parapodium. These are ciliated setigerous parapodia, its lateral eyes are and are relatively long and gently tapered deeply embedded, and its anal funnel is distally to their rounded tips. Many of the attenuated). branchiae have been lost from the specimen, Although Moore (1906) and Hartman but they are present to at least the twenty- (1938) have stated that A. brevis possesses an 0

D-

0 E 0.1MM.

9! ~~~~~II .2M

95 94

0 C 2 MM. 92 93

FIGS. 83-90. Armandia nonpapillata, new species. 83. Lateral view of anterior region. 84. Dorsal view of anterior region. 85. Anterior view of right fifth setiger. 86. Anterior view of left twelfth setiger, setae missing. 87. Anterior view of left twenty-third setiger. 88. Lateral view of body, from sixth to tenth setigers (anterior to right). 89. Lateral view of posterior region. 90. Ventral view of posterior region. FIGS. 91-95. Nicomache antillensis Augener. 91. Dorsal view of anterior region. 92. Notoseta from fifth setiger. 93. Notoseta from seventeenth setiger. 94. Neuropodial hook from second setiger. 95. Neuropodial hook from seventeenth setiger. 83, 84, 88-90, scale A; 85-87, scale B; 91, scale C; 92, 93, scale D; 94, 95, scale E. 1962 JONES: POLYCHAETOUS ANNELIDS 193 anal funnel which lacks anal papillae, Tread- FAMILY MALDANIDAE MALMGREN, 1867 well (1922) and Berkeley and Berkeley (1941) GENUS NICOMACHE MALMGREN, 1865 believe that the type of A. brevis was an im- Nicomache antillensis Augener, 1922 perfect specimen, and they have reported specimens which do bear an anal cirrus and Figures 91-95 papillae. Nicomache antillensis AUGENER, 1922a, p. 46; Barbados. Nicomache antiguensis TREADWELL, 1924b, pp. FAMILY CAPITELLIDAE GRUBE, 1862 16-17; Pelican Island, Antigua. Nicomache antillensis, AUGENER, 1927b, p. 70; GENUS DASYBRANCHUS GRUBE, 1850 Curagao. Dasybranchus sp. U.C.W.I. COLLECTION: This species is U.C.W.I. COLLECTION: Three specimens of represented by two posteriorly incomplete Dasybranchus sp. were taken from a mud sub- specimens. The first, comprised of the an- strate at Ocho Rios, Jamaica, on August 11, terior 10 setigerous segments, has a length of 1954. All are represented by anterior frag- 50 mm. and a maximum width of 4 mm. at ments, the longest of which is comprised of parapodia and 3 mm. between parapods. The 176 setigerous segments (length, 61 mm.; second, the anterior 17 setigers, is 155 mm. maximum width in the thoracic region, 1.0 long and 5 mm. wide at parapodia and 4 mm. mm.). The remaining specimens have 119 wide between parapods. The former was col- setigers (length, 32 mm.; maximum width at lected at Drunkenman Cay, Port Royal, the twenty-fifth setiger, 1.5 mm.) and 86 Jamaica, in January, 1959; the latter was setigers (length, 30 mm.; width, 1.5 mm.). collected from sand, "Port Royal Cays," in DISCUSSION: These specimens caused some November, 1959. confusion at the outset of their identification DISCUSSION: Although the anal funnel of on two accounts. First, the two largest carried this specimen, which would give an unequiv- capillary setae on the first 13 setigers, but the ocal determination of the genus (Nicomache smallest had capillary setae on only the first or Petaloproctus) is missing, the number and 12. Second, the apparent demarcation be- distribution of anterior neuropodial acicular tween the thorax and abdomen was between setae, the morphology of the head region, and the eleventh and twelfth setigers in the two the shape of the nuchal organs all confirm largest, and between the tenth and eleventh this identification. in the smallest. It was finally decided that the In one specimen the neuropodia of the first smallest specimen represented a variant and setiger bear five acicular setae on one side and that the three specimens were, indeed, three on the other; the second bears four on Dasybranchus sp. Further identification was each side; and the third has five on each side. not possible because no dorsal gills were None of the described species of Petaloproctus visible. Indeed, under the closest scrutiny, not bears more than one acicular seta on either even slit-like passages for the extrusion of side of the most anterior neuropodia, and this retractable gills were to be seen. Thus a more number and distribution agree with descrip- definitive identification of these specimens tions presented by both Augener (1922a) and must await further collections to show the Treadwell (1924b). shape of gills, if, indeed, they prove to be The nuchal organs of this specimen from present. If this species of Dasybranchus Jamaica are a pair of semicircular structures, actually has no gills, it may well be new. on each side of a low median ridge (fig. 91). The observation of abdominal hooded These are also a confirmatory feature. hooks added to the confusion concerning the During the course of observations, other identity of these specimens, for, under oil items of interest were noted, namely, the fine immersion, there appear to be at least four structure and distribution of setae. smaller teeth immediately above the large The notosetae of the most anterior setigers fang rather than the row of three small teeth are of two types. The first are smooth, un- reported by Hartman (1947, pp. 430-437, pls. ornamented, pointed capillaries. Those of the 55-57). second type, also pointed, at first glanceap- 194 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 pear to be bilimbate, but under oil immersion FAMILY MALMGREN, 1867 the "limbations" prove to be longitudinal GENUS SABELLA LINNAEUS, 1767 splits on each side of the axis of the seta. Sabella melanostigma Schmarda, 1861 These latter setae are found along the length of N. antillensis. Sabella melanostigma SCHMARDA, 1861, p. 36; Jamaica. By the fifth setiger unornamented noto- Sabella bipunctata BAIRD, 1865, PP. 158-159; podial capillaries are replaced by a setal type, St. Thomas. which in the past has been called " . . . soies Sabella melanostigma, WEBSTER, 1884, p. 327; . . . garnies d'epines tres dMlicates" (Fauvel, Bermuda. 1927, p. 168), setae with "deutliche Seiten- Sabella bipunctata, MCINTOSH, 1885, pp. 489- zahnchen" (Arwidsson, 1906, pp. 103, 107, et 490; St. Thomas. al.), "spined" (Chamberlin, 1919b, p. 407), or Sabella melanostigma (Schmarda) [sic], EHLERS, setae "with paired lateral plates whose free 1887, pp. 263-266; [Dry] Tortugas, Florida. margins are toothed" (Treadwell, 1924, p. Sabella melanostigma, TREADWELL, 1901, P. 17). After their first appearence these (fig. 92) 208; Puerto Rico. Sabella melanostigma var., HOAGLAND, 1919, are to be found on all remaining notopodial p. 577; Puerto Rico. fascicles (at least to the seventeenth setiger). Sabella melanostigma, AUGENER, 1922a, p. 52; In the more anterior setigers these setae tend "West Indies." to be much shorter than those more posterior. Sabella melanostigma, MULLIN, 1923, pp. 49-50; In some of the larger setae of this type the Barbados. true setal structure becomes apparent, for Sabella melanostigma, TREADWELL, 1924, p. 20; their tips may become unraveled (fig. 93), re- Barbados and Antigua. vealing that the so-called "spines," "teeth," Sabella melanostigma, AUGENER, 1927b, pp. 73- and "lateral plates" are double spirals, the 74; Curagao. distal edges of which are minutely serrate. Sabella melanostigma, TREADWELL, 1928, p. 482; near Saba Island. The long-handled hooks, which are present Sabella bipunctata, MONRO, 1933b, p. 1078; in all neuropodia posterior to the third (at Trinidad. least to the seventeenth), are of the usual Sabella melanostigma, MONRO, 1933c, p. 267; maldanid type. Each bears a subrostral tuft of Dry Tortugas. from five to six distally curled filaments. Sabella melanostigma, TREADWELL, 1939, p. Those of the anterior region (fig. 94) bear, in 295; "Barbados; Antigua; Puerto Rico; and near addition to a large primary tooth, a single Saba Island." series of five somewhat smaller teeth sur- Sabella melanostigma, HARTMAN, 1951b, pp. mounted by a sixth much smaller tooth. 116-117; Franklin and Sarasota counties, Florida. These more obvious features have been noted Sabella melanostigma, CARPENTER, 1956, p. 107; 27 and Franklin County, Florida. previously by Treadwell (1924, figs. Sabella melanostigma, RENAUD, 1956, p. 34; 29). However, present observations, under Bimini, Bahamas. rigorous optical and illumination conditions, Sabella melanostigma, MARSDEN, 1960, p. 1014; reveal a row of about five very fine teeth on Barbados and Jamaica. each side of the larger medial row of teeth. Hooks from more posterior setigers are simi- U.C.W.I. COLLECTION: Two specimens lar (fig. 95), but their large tooth is sur- were collected at Port Royal, Jamaica, in mounted by a single row of seven progres- November, 1959. The larger (124 setigerous sively smaller teeth and an eighth still smaller segments) has an over-all length of 62 mm. tooth. These, too, are flanked by a series of and a crown length of approximately 15 mm. very fine teeth on each side. A further differ- The smaller (103 setigers) has an over-all ence, which may be of little importance, is length of 40 mm. and a crown length of ap- that there is much less space between the proximately 12 mm. large tooth and the next one above it, in those DISTRIBUTION: Temperate, subtropical, hooks of the posterior region. tropical: northwest Florida; Bermuda; Ba- DISTRIBUTION: Tropical: Jamaica; Cura- hamas; Florida Keys; Jamaica; Lesser Antil- gao; Lesser Antilles. les; and the northern coast of South America. 1962 JONES: POLYCHAETOUS ANNELIDS 195 GENUS BRANCHIOMMA KOLLICKER, 1858 Dasychone ponce TREADWELL, 1901 [in part], Branchiomma arenosa (Treadwell), 1924 p. 209; Puerto Rico. Figures 96-106 Dasychone nigro-maculata, AUGENER, 1922a, Dasychone nigro-maculata ("Baird"), McIN- p. 52; "West Indies." TOSH, 1885 [not Branchiomma nigromaculata Dasychonopsis arenosa TREADWELL, 1924a, pp. (Baird) 1865], pp. 503-504; St. Thomas. 1-2; Puerto Rico.

96 97

104 103\

FIGS. 96-106. Branchiomma arenosa (Treadwell). 96. Hook from right fourth thoracic setiger of holotype (A.M.N.H. No. 1616). 97. Hook from right fifteenth abdominal setiger of holotype. 98. External appendages of a radiole of holotype. 99. Shorter limbate seta from left fourth thoracic setiger of specimen from Jamaica. 100. Longer limbate seta from left fourth thoracic setiger (Jamaica). 101. Hook from left fourth thoracic setiger (Jamaica). 102. Shorter limbate seta from left fifteenth abdominal setiger (Jamaica). 103. Longer limbate seta from left fifteenth abdominal setiger (Jamaica). 104. Hook from left fifteenth abdominal setiger. 105. External appendages of a radiole (Jamaica). 106. Distal tip of a radiole (Jamaica). 96, 97, 99-104, scale A; 98, 105, 106, scale B. 196 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 Dasychone nigromaculata, AUGENER, 1927b [in well's (1924a) original description, is accept- part], p. 76; Curagao. able for Branchiomma arenosa. In addition, [?] Dasychone nigromaculata, AUGENER, 1934, figures of critical points of comparison are p. 113; Isla de Margareta, Venezuela. herein Dasychonopsis arenosa, TREADWELL, 1939, pp. included (figs. 99-106), as well as 296-297; "Puerto Rico." figures of comparable structures in Branchi- Branchiomma nigromaculata, HARTMAN, 1951b, omma nigromaculata (figs. 107-114). It is seen pp. 114-115; St. Lucie County, Florida (Atlantic); that there are but slight differences in the Sarasota County, Florida (Gulf of Mexico). limbate setae of the two species; however, Branchiomma nigromaculata, CARPENTER, 1956, there are somewhat more substantial differ- p. 107; Wakulla County, Florida (northern Gulf of ences in the structure of the thoracic hooks Mexico). (figs. 101 and 109) as well as those of the Brianchiomma [sic] nigromaculata, RENAUD, abdomen (figs. 104 and 112). The main 1956, p. 34; Key West. morphological differentiation of B. arenosa Branchiomma nigromaculata, MARSDEN, 1960, from B. nigromaculata lies in the structure of pp. 1014-1015; Jamaica. the radioles of the branchial plume, especially U.C.W.I. COLLECTION: Two specimens, those of the dorsal and lateral areas. The ex- both of 62 setigers, were collected at Port ternal appendages of the radioles of B. arenosa Royal, Jamaica, in November, 1959. The are all of approximately the same length (fig. over-all length of one was 30 mm. (the bran- 105), while at least two pairs of those of B. chial plume was 11 mm. long, and the speci- nigromaculata are twice the length of the re- men was 1.5 mm. at its widest); of the other, maining appendages (fig. 113). There also ap- 38 mm. (the branchial plume was 9 mm. long, pears to be a difference in the structure of the and the specimen was 2.0 mm. at its widest). tips of the radioles, for in B. arenosa the tips DISCUSSION: As a result of critical com- bear filaments to their apices (fig. 106), while parisons of the texts and figures presented by those of B. nigromaculata are bare (fig. 114). Baird (1865) for his Sabella nigro-maculata DISTRIBUTION: Subtropical to tropical: and by McIntosh (1885) for his Dasychone from the northern Gulf coast of Florida bairdi and D. nigro-maculata, it was concluded through the Greater and Lesser Antilles to that Branchiomma bairdi is a junior synonym the Venezuelan coast. of B. nigromaculata (see discussion of B. nigromaculata below). Branchiomma nigromaculata (Baird), 1865 In the establishing of the valid name for the Figures 107-114 species referred by McIntosh to the taxon that has heretofore been accepted as Sabella nigro-maculata BAIRD, 1865, p. 159; St. Branchiomma nigromaculata, it is seen that Vincent. Dasychone bairdi MCINTOSH, 1885, pp. 495-4971 the next available name, B. ponce, is actually Bermuda. unavailable, since there are ambiguities in the Dasychone conspersa EHLERS, 1887, pp. 266- description, as mentioned below. 270; Key West, Florida. Comparative observations have been made Dasychone ponce TREADWELL, 1901 [in part], on the holotype of Dasychonopsis arenosa p. 209; Puerto Rico. Treadwell (A.M.N.H. No. 1616), and, on the Dasychone conspersa, HOAGLAND, 1919, p. 577; basis of the structure of thoracic (fig. 96) and Puerto Rico. abdominal (fig. 97) hooks and the form of the Dasychone bairdi, AUGENER, 1922a, p. 49; Dry external appendages of the radioles (fig. 98), Tortugas; St. Thomas; Jamaica; Haiti; Veracruz, Dasychonopsis arenosa is considered a syno- Mexico. nym of B. nigromaculata (sensu McIntosh, Dasychone conspersa, MULLIN, 1923, pp. 50-51; 1885). Antigua. It is, then, Treadwell's (1924a) taxon, Dasychonopsis conspersa, TREADWELL, 1924b, p. 18; Barbados and Antigua. Dasychonopsis arenosa, which is the earliest Dasychone nigromaculata, AUGENER, 1927b valid name available for this species. [in part], p. 76; Curagao. For the most part, McIntosh's descriptive Branchiomma nirgomaculata [sic], JOHANSSON, account (1885, pp. 503-504, pl. 31A, figs. 4-6, 1927, pp. 162-164; Curagao; Barthelmy (St. pl. 39A, fig. 6, pl. 53, fig. 5), as well as Tread- Bartholomew); and Bermuda. 1962 JONES: POLYCHAETOUS ANNELIDS 197

fit

FIGS. 107-114. Branchiomma nigromaculata (Baird). 107. Shorter limbate seta from right fourth thoracic setiger. 108. Adjacent longer (left) and shorter limbate setae from right fourth thoracic setiger. 109. Hook from right fourth thoracic setiger. 110. Shorter limbate seta from right fifteenth abdominal setiger. 111. Longer limbate seta from right fifteenth abdominal setiger. 112. Hook from right fifteenth abdominal setiger. 113. External appendages of a radiole. 114. Distal tip of a radiole. 107-112, scale A; 113, 114, scale B.

Dasychone bairdi, MONRO, 1933c, p. 267; Dry 299; "Key West and Puerto Rico." Tortugas. Branchiomma bairdi, HARTMAN, 1951b, p. 115; [?] Dasychone nigromaculata, AUGENER, 1934, "Dry Tortugas." p. 113; Isla de Margareta, Venezuela. Dasychone bairdi, RIOJA, 1952, pp. 513-516; Dasychonopsis ponce, TREADWELL, 1939, p. Veracruz, Mexico. 298; "Puerto Rico." Branchiomma bairdi, MARSDEN, 1960, p. 1015; Dasychonopsis conspersa, TREADWELL, 1939, p. Barbados and Jamaica. 198 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 U.C.W.I. COLLECTION: Four specimens viduals have two pairs of these dorsal appen- were collected at Port Royal, Jamaica, three dices much larger than the rest . . . this does in November, 1959 (one of which was oviger- not appear in all specimens." This would in- ous), and one on August 8, 1961. The over- dicate to me that Treadwell was probably all lengths of the three specimens collected in dealing with the two West Indian species of November ranged from 30 to 34 mm. (the Branchiomma recognized by McIntosh as length of the branchial plumes, from 11 to 12 Dasychone nigromaculata Baird and D. bairdi mm.), and they were all 2.0 mm. wide at the McIntosh. thoracic region. The number of setigerous seg- Similarly, Augener (1927b, p. 76) ap- ments varied from 58 to 70. The single com- parently found these two species in the collec- plete individual taken in August was 33 mm. tions from Curagao. He noted the differences in length, for 70 setigers (branchial plume, 12 in length of the appendages of the radioles, mm.), and was 3.0 mm. wide. but minimized these and was moved to sy- DISCUSSION: In the original description of nonymize Dasychone nigromaculata and D. Branchiomma nigromaculata (Baird, 1865, p. bairdi, retaining the former name as the 159), it is stated that on the outer margin of senior synonym. the "rachis" of the branchial radioles, there It remained for Johansson (1927, pp. 162- are "I . .. very short filaments, set in pairs. 164) to assign Dasychone and Dasychonopsis Near the base of the filament [radiole] spring to K1llicker's genus Branchiomma and to a pair longer and broader, and near the middle synonymize Dasychone conspersa Ehlers, D. of its length another pair of the same kind." ponce Treadwell, and D. arenosa Treadwell In addition, Baird's figure 6 confirms this with B. nigromaculata (Baird). It should be statement. noted, in passing, that Johansson's identifica- Later, McIntosh (1885, pp. 503-504, pl. tion of this species (B. nigromaculata) was 31A, figs. 4-6, pl. 39A, fig. 6, pl. 53, fig. 5) pre- based on McIntosh's (1885) description, in sented figures and the description of speci- spite of his (Johansson) observing that some mens which he identified as Baird's species. of the external appendages of the branchial However, McIntosh (p. 503) observed that radioles were longer than others. Further, on the external surface of the radioles " . . . a Johansson indicated (p. 167) that, although series of rather short appendages occur in it was not present in his collections, he recog- pairs ... which are numerous and slender." nized B. bairdi as a valid species. An accompanying figure (McIntosh, 1885, pl. The final item in the synonymy that re- 39A, fig. 6) shows that the appendages in quires comment is Augener's (1934) record of question are all of approximately the same B. nigromaculata from the Venezuelan coast. size. Strangely, in the same Challenger re- Unfortunately, he makes no mention of the port, and some pages earlier (pp. 495-497), morphology of the branchial plume, so it only McIntosh described Dasychone bairdi as a can be assumed that he was dealing with the "new" species of this genus. Of the branchial present species. radioles, McIntosh writes (1885, p. 496), DISTRIBUTION: Subtropical to tropical: "Each has externally, at intervals, a pair of from the Florida Keys through the Greater ligulate hypodermic processes .., a shorter and Lesser Antilles to the Venezuelan coast; pair often alternating with a longer pair." Bermuda; and Veracruz, Mexico. This observation is also confirmed by a figure (pl. 39A, fig. 2). GENUS PSEUDOBRANCHIOMMA, NEW GENUS It should be pointed out that Day (1955, p. TYPE: Pseudobranchiomma emersoni, new 445) has also suggested that these two species species. are synonymous. Pseudobranchiomma emersoni, new Ehler's species, Dasychone conspersa Eh- species lers, 1887, also exhibits both long and short Figures 115-124 external radiolar appendages. DIFFERENTIAL DIAGNOSIS: Sabellid with The status of Dasychone ponce Treadwell, four thoracic setigers in the adult; with re- 1901, is somewhat obscure, for Treadwell duced external appendages on the branchial mentions (p. 209) that although "some indi- radioles; with no eye spots on the radioles; 1962 JONES: POLYCHAETOUS ANNELIDS 199

124

0 122

120 121 Il 19

-0 -o

II, )I -A II II -B

-0.1MM. 118

116 117 115 - 1.0 MM. II FIGS. 115-124. Pseudobranchiomma emersoni, new genus, new species. 115. Longest limbate seta from right second thoracic setiger. 116. Medium limbate seta from right second thoracic setiger. 117. Shortest limbate seta from right second thoracic setiger. 118. Hook from right second thoracic setiger. 119. Longer limbate seta from right fifteenth abdominal setiger. 120. Shorter limbate seta from right fifteenth abdominal setiger. 121. Different view of shorter limbate seta from right fifteenth abdominal setiger. 122. Hook from right fifteenth abdominal setiger. 123. External appendages of a radiole. 124. Distal tip of radiole. 115-122, scale A; 123, 124, scale B. and with the tips of the branchial radioles ated with an unidentified colonial ascidian. devoid of pinnules. DESCRIPTION: The general body color is a U.C.W.I. COLLECTION: Many specimens brownish to pinkish white, and the branchial were collected on July 1, 1961, from settling plumes bear purple splotches which give the panels in shallow water (4 feet) at Port Royal, impression of narrow irregular bands. The Jamaica. Various sizes, from very small indi- pigmentation of the branchial radioles is con- viduals to adults, are represented, and the fined to their sides and is carried onto those larger ovigerous specimens are approximately internal pinnules that are immediately adja- 25 to 30 mm. in length for about 90 setigers cent (fig. 124). The base of the branchial (branchial plumes comprise 6 to 9 mm.); they plume bears a series of longitudinal purple are from 1.0 to 1.5 mm. wide in the thoracic streaks. The anterior region of Pseudo- region. All specimens collected were associ- branchiomma emersoni is provided with ir- 200 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 regular purple splotches, especially on the col- a recurved tip; the longer are straighter and lar and the ventral body surface. In each seg- have a somewhat more narrow limbation ment there are rather large, circular, purple (fig. 117). spots (not eyes) between the neurosetae and Thoracic hooks (fig. 118) have approxi- the notosetae, and, dorsal to the notosetae, mately five teeth (or rows of teeth) above the there is a much lighter, more irregular spot. main fang, which is sharply bent back on the There is a large purple spot at each of the rounded basal portion. There is a distinct dorsal ends of the collar and a distinct spot on broad extension of the basal region which either side of the ventral notch. may represent a short manubrium. These are There are no eyes on the radioles, only ir- not accompanied by so-called "pickax" setae. regular patches of purple. The external ap- Neurosetae of the abdomen are also of two pendages of the radioles, which are so promi- types: thin, attenuated, unilimbate capillaries nent in the genus Branchiomma, are here re- (fig. 119), and shorter, heavier, geniculate duced to stubby processes, where they are unilimbate capillaries (fig. 120). Some of the best developed (figs. 123 and 124). Approxi- latter, which are properly oriented (fig. 121), mately 10 pairs of such reduced appendages show that these are actually bilimbate setae are to be found on the external margin of the which appear unilimbate owing to their branchial radioles, and the members of each orientation. This condition probably holds pair are separated by a rather pronounced for the apparently unilimbate setae of the longitudinal groove which runs the length of thorax. the radiole. The tips of the radioles are fili- Abdominal hooks (fig. 122) are similar to form and lack pinnules (fig. 124). The ra- those of the thorax. dioles are fused basally for approximately Tubes are flexible and are covered with a one-eighth of their total length. fine gray sediment. Dorsally, the collar is separated, and it ex- The generic name refers to the superficial tends only to the notosetal fascicles on each resemblence of this genus to the genus side. Ventrally, the collar is deeply incised, Branchiomma. The trivial name was chosen and the lobes so formed do not overlap. An- to honor Dr. William K. Emerson, the Ameri- terior to the ventral notch of the collar there can Museum of Natural History, without is a pair of foliaceous flaps which appear to whose encouragement and criticism the form a connection between the collar and the present paper might not have been written. base of the branchial plume. The pair of oral TYPE DISPOSITION: The holotype (A.M.- tentacles are approximately one-fourth of the N.H. No. 3612) and paratypes (A.M.N.H. length of the plume. Dorsally, between the No. 3613) of Pseudobranchiomma emersoni base of the branchial plume and the dorsal are deposited in the collections of the Ameri- ends of the collar, there is a longitudinal can Museum of Natural History. ridge on each side of the mid-line. DISCUSSION: The separation of Pseudo- In ovigerous specimens, the thoracic region branchiomma from Branchiomma, the most is comprised of four segments which are closely related genus of the Sabellidae, is represented by four notopodial fascicles of based mainly on the number of thoracic seti- limbate setae and three neuropodial tori of gers. A comparison of the 22 established spe- avicular hooks. In the case of smaller indi- cies of Branchiomma shows that all but B. viduals, the thorax may consist of five or of bombyx, B. curta, and B. natalensis have eight six segments; one other smaller specimen was thoracic setigers. According to Fauvel (1927, noted which bore four notopodial fascicles of p. 319), B. bombyx may have from five to setae on one side and five on the other. eight setigerous segments in the thoracic re- Thoracic notosetae are of two basic types: gion. Ehlers (1901), in his original description a long, rather straight, unilimbate capillary of B. curta, reports from four to six setigers in (fig. 115), and shorter capillaries, the limba- the thorax of this species (which may, ac- tions of which are restricted to their subapical cording to Augener [1922b, pp. 211-217], region (figs. 116 and 117). The shorter of the represent regenerative stages), and Kinberg latter setae (fig. 116) are slightly geniculate (1910) indicates that B. natalensis possesses and are provided with a broad limbation and seven thoracic setigers. In addition, Mc- .1962 JONES: POLYCHAETOUS ANNELIDS 201 Intosh (1885, p. 497) states that the number Indies; Veracruz, Mexico; Puerto Cabello, Vene- of "anterior segments" (=thoracic setigers) zuela; St. Croix; Surinam. of B. picta could not be determined, and later ? [Sic] Bispira melania, MULLIN, 1923, pp. 52- (1885, p. 499), that the number of thoracic 56; Antigua. setigers of B. orientalis "appears to be eight, Bispira melania, TREADWELL, 1924b, pp. 18-19; of is not attain- Antigua. but ... a clear view these Sabellastarte magnifica, AUGENER, 1925, pp. able." In ovigerous Pseudobranchiomma emer- 10-11; St. Thomas and St. Croix. soni, the number of thoracic setigers is always Sabellastarte magnifica, AUGENER, 1927b, p. 73; four. Curagao. On the basis of the structure of the external ? Sabellastarte indica, MONRO, 1933b, p. 1079; appendages of the radioles, a closer relation- Margarita Island; Col6n, Panama (Caribbean end ship may be postulated between Pseudo- of the Panama Canal). branchiomma and Branchiomma picta (Mc- ? Sabellastarte indica, RIOJA, 1946, pp. 198-199; Intosh), 1885, B. orientalis (McIntosh), 1885, Veracruz, Mexico. and B. kumari (Aziz), 1938. All of these have Sabellastarte magnifica, HARTMAN, 195 1b, p. external appendages which are poorly de- 116; "West Indies and Veracruz, Mexico." veloped, and, further, none bears eyes on the Sabellastarte magnifica, ANDREWS AND AN- radioles-only patches of pigment. Yet DREWS, 1953, pp. 12-13; Bimini, Bahamas. another sabellid has the reduced dorsal ap- Sabellastarte magnifica, MARSDEN, 1960, p. pendages that these species possess. Recently, 1012; Barabados and Jamaica. Hartman (1959, p. 541) has indicated that U.C.W.I. COLLECTION: Two complete spec- Fauvel's (1921) Dasychone odhneri is a imens and the branchial crown of a third synonym of Sabellastarte longa (Kinberg). were collected on piers at Port Royal, Ja- Earlier, Augener (1926, pp. 257-258) con- maica, in October, 1959. The larger complete sidered it to be a synonym of Branchiomma specimen (155 mm. long, 12 mm. wide) has a serratibranchis (Ehlers), 1907, and Day (1934, branchial crown 65 mm. in length; the smaller p. 75) suggested the possibility of its being a (143 mm. long, 10 mm. wide), a crown 60 mm. synonym of Sabellastarte indica (Savigny). On in length. The unattached crown is 55 mm. the basis of the reduced external appendages long. that it possesses ("Appendices branchiaux DISCUSSION: Comparison of these speci- reduits d de simple crates lateral," Fauvel, mens with the holotype of Laonome sanjuan- 1921, p. 24), I suggest that it is a Branchi- ensis (A.M.N.H. No. 2892) and its descrip- omma and that it has affinities with B. picta, tion (Treadwell, 1941) indicates that L. san- orientalis, and B. kumari, insofar as external juanensis should not be synonymized with appendages are concerned, and with B. Sabellastarte magnifica (cf. Hartman, 1956, p. natalensis, with reference to its spirally ar- 299). Generally, my observations on the ranged branchial plumes. holotype of Laonome sanjuanensis agree with Finally, I would suggest that if Branchi- Treadwell's description; however, there are omma picta and B. orientalis prove to have several features, one of which is of prime im- four thoracic setigers, then it will become portance, which were overlooked both by necessary to transfer them to the genus Treadwell (1941), and by Hartman (1956) in Pseudobranchiomma. her inspection of this specimen. GENUS SABELLASTARTE SAVIGNY, 1818 Treadwell states (p. 3) that "the thorax is Sabellastarte magnifica (Shaw), 1800 composed of six somites. . . ," and (p. 4) that "uncinigerous tori begin on somite 2, Tubularia magnifica SHAW, 1800, pp. 227-229; each carrying a single row of uncini." The Jamaica. out the Sabella lingva KR0YER, 1856, p. 27; West Indian first statement has been borne by seas. present observations, but it is uncertain Sabella melania SCHMARDA, 1861, p. 35; Jamaica. whether or not the second is wholly true. The Sabella splendida KINBERG, 1867, p. 353; uncertainty arises from the fact that the first Guadeloupe. two left thoracic setigers have been removed, Sabellastarte magnifica, AUGENER, 1922a, p. in toto, and the first two right thoracic setigers 48; St. Thomas; Jamaica; Haiti; Barbados; West bear only notopodial setal fascicles; there are 202 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 no corresponding rows of neuropodial uncini reports that "thoracic neurosetae are entirely associated with either. The remaining four avicular." However, close inspection of the thoracic setigers have both neuropodial and sixth left thoracic neurosetae shows that, in notopodial fascicles. This condition may well addition to uncini, there are small, nearly represent a morphological abnormality, for transparent, "pennoned setae" (cf. Fauvel, there is further evidence that the orderly ar- 1927, fig. 103, k and 1 ["soies en pelle"]; and rangement of abdominal setigers has been Fauvel, 1953, fig. 232, k and 1 ["shovel pick- upset; the right and left twenty-sixth and the axe setae"]). On this basis alone, this species right twenty-seventh setigers of the abdomen can be neither a Metalaonome nor a Sabella- are inverted, i.e., there are notopodial capil- starte. In my opinion this is a species of lary setae and neuropodial uncini. Sabella. Proper specific designation must The two types of limbate capillary setae await a thorough study of the literature and described by Treadwell have been noted, further observations. rather than the one type reported by Hart- DISTRIBUTION: Tropical: Bahamas; Great- man (1956, p. 299). er and Lesser Antilles; northern coast of The last, and most important, point at South America; and perhaps the Caribbean variance with Treadwell's description is the coast of Panama and western Mexico. fact that there are pairs of so-called dorsal appendages on the outer surfaces of the FAMILY SERPULIDAE SAVIGNY, 1818 branchial filaments. To be sure, these are GENUS OLGA, NEW GENUS rather delicate and nearly transparent, but TYPE: Olga elegantissima, new species. they are present all along the length of the Olga elegantissima, new species filament. Further, they are of varying lengths, similar to those of Branchiomma nigromacu- Plate 52, figures 1 and 2; text figures 125-128 lata (Baird) [ =B. bairdi McIntosh], and longer DIFFERENTIAL DIAGNOSIS: Serpulid with appendages are followed by two pairs of six thoracic setigers; with an operculum shorter appendages (cf. McIntosh, 1885, pl. which possesses horns on its surface and wing- 39A, fig. 2). Distally, all dorsal appendages like processes on its peduncle; without collar are short. I concur with Treadwell in his ob- setae; and with spirally arranged branchial servation that there are no eye spots on the plumes. gill filaments. In addition, there are no pig- U.C.W.I. COLLECTION: A single specimen mented spots between the neuropodial and was collected from "Port Royal Cays," Ja- notopodial areas. maica, in 1957. It is approximately 44 mm. in In my opinion Laonome sanjuanensis length (crown length is 17 mm.) and repre- Treadwell is not Sabellastarte magnifica sents an anterior fragment (to the seven- (Shaw), but, rather, is a member of the genus teenth abdominal setiger); it is about 7 mm. Branchiomma Kollicker (non Claparede), and in diameter in the abdominal region. is herein designated Branchiomma sanjuanen- DESCRIPTION: The branchial crown of sis (Treadwell). Olga elegantissima (pl. 52, fig. 1) consists of a In addition, a comparison of the holotype pair of whorls of branchial filaments which (A.M.N.H. No. 982) and paratypes (A.M.- are disposed in decreasing spirals. There are N.H. No. 981) of Metalaonome brunnea and eight whorls in each part which decrease from its description (Treadwell, 1917) with these a basal diameter of approximately 1 cm. to specimens suggest that this, too, should not the apical tip. The bases of the branchial fila- be synonymized with Sabellastarte magnifica ments are united for one-third of their length. (cf. Augener, 1927b, p. 40; Hartman, 1956, Dorsally, an operculum arises from between p. 299). In general, my observations of the the two parts of the branchial crown; the tip holotype and paratypes confirm Treadwell's of the operculum is a subcircular plate which description of Metalaonome brunnea and bears a central, low tooth and a pair of hooked Hartman's comments on this species. Tread- horns. Just below the opercular plate there well states (p. 268) that he " . . . was unable are paired wings on each side of the opercular to discover any pennoned setae in the peduncle (pl. 52, fig. 2). thorax of these forms," and Hartman (p. 299) The thorax is comprised of six setigers, the BULLETIN AMER. Mus. NAT. HIST. VOL. 124, PLATE 52

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FIGS. 125-128. Olga elegantissima, new genus, new species. 125, 126. Left second thoracic setiger. 125 Limbate seta. 126. Uncinus. 127, 128. Left twenty-fifth abdominal setiger. 127. Uncinus. 128. Seta. FIGS. 129-138. Salmacina amphidentata, new species. 129-131. Collar setae. 129. Largest type. 130. Second type. 131. Third type. 132. "Salmacina type" of seta from thoracic notopodium. 133. Thoracic uncinus. 134. Face view of thoracic uncinus. 135. Abdominal uncinus. 136. Face view of abdominal uncinus. 137. Larger abdominal seta. 140. Smaller abdominal seta. 125, scale A; 126-128; scale B; 129-138, scale C. setae and uncini of which are embedded in the basal "gouge-shaped" tooth. The neuropodial thoracic collar. There are no notopodial collar setae of the abdomen (fig. 128) are expanded fascicles. Thoracic notopodial setae are uni- at their tips and are asymmetrically attenu- limbate pointed capillaries (fig. 125), and ated. Their distal margin is finely denticulate. notopodia are provided with elongate uncini Even after four years in preservative which bear approximately 20 teeth and a (formalin), this specimen still retains its basal "gouge-shaped" tooth (fig. 126). color. The general body, the opercular pe- The thoracic region of 0. elegantissima is duncle, and the branchial filaments are white; separated from the first abdominal setae by the anterior portion of the thoracic collar and about the length of the collar. The abdominal the central axis of the branchial plumes are a uncini (fig. 127) are smaller than those of the deep blue; and the rim of the operculum and thorax. They are subtriangular in shape and its horns are red. This is truly a beautiful bear approximately 12 curved teeth and a animal. 204 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124 The generic name was chosen to honor Dr. Jamaica, on December 8, 1959. Ovigerous in- Olga Hartman, Allan Hancock Foundation, dividuals are from 2.5 to 3.5 mm. in length University of Southern California, whose bib- for 23 to 25 setigers, and are approximately liography (Hartman, 1951a) and catalogue 0.25 mm. wide, in the thoracic region. (Hartman, 1959), as well as her many mono- DESCRIPTION: The branchial crown of Sal- graphic works, have contributed so much to macina amphidentata is comprised of eight the study of the Polychaeta and have made pinnate branchiae, the tips of which are but the way easier for those of us who follow her. slightly inflated. The dorsal collar extends The trivial name refers to the esthetic ap- from the base of the crown to the posterior pearance of the animal. end of the thorax, which is composed of seven TYPE DISPOSITION: The holotype (A.M.- (rarely, six) setigerous segments. The first N.H. No. 3614) of Olga elegantissima is de- setiger is represented only by a pair of noto- posited in the collections of the American podial setal fascicles, the so-called collar Museum of Natural History. setae, and subsequent thoracic segments DISCUSSION: According to Bush (1904, pp. have both notopodial capillaries and neuro- 221-227), Chamberlin (1919b, pp. 473-476), podial uncini. The abdomen is comprised of and Fauvel (1927, pp. 348-350; 1953, pp. 16 to 17 segments, which are provided with 453-454), of the five serpulid genera that lack notopodial uncini and neuropodial capillaries. collar setae (Placostegus Philippi, Bonhourella The setae of the collar fascicle are of three Gravier, Pomatoleios Pixell, Rhodopsis Bush, types. The first, and largest (fig. 129), have a and Ditrupa M. Berkeley), three are char- distal wing which is finely denticulate on one acterized by flat opercula (Placostegus, Bon- margin. This portion is separated by a wide, hourella, and Pomatoleios), one by a chitinous shallow notch from a toothed protuberance, operculum with a rosette of chitinous spines the distal teeth of which are larger than those (Rhodopsis), and the last by an operculum that are more proximal. The teeth of this re- bearing an inverted cone (Ditrupa). Further, gion are not confined to the profile edge, but the gill filaments of four of these genera are are distributed over the surface of this en- disposed not in spirals, but as semicircles; the larged structure. The second type of collar disposition of gill filaments of Rhodopsis is not seta (fig. 130) is long and narrow and is pro- given by Bush (1904, pp. 298-290). vided with a pair of narrow limbations, one of On the basis of these criteria (lack of collar which is finely denticulate, basally. The third setae, opercular ornamnentation, and the dis- type (fig. 131) is also narrow but is non- position of gill filaments), Olga is proposed as limbate; rather, it is provided with a basal a new genus of the Serpulidae. denticulate wing. The lack of collar setae would indicate an The remaining thoracic notopodial setae affinity to the five genera listed above. How- are of two types: one is similar to the seta last ever, it should be noted that Olga must also described (fig. 131), and the other, which usu- be closely related to Spirobranchus Blainville. ally occurs singly in a given fascicle, is the so- Such a relationship is indicated by the horn- called "Salmacina type" of seta (fig. 132). shaped processes of the operculum, the wing- These are pointed and are gently double- like structures on the peduncle of the opercu- curved. The inner margin of the distal curve lum, the general shape of the uncini, and the bears blunt teeth, and the blade bears fine spirally arranged branchiae. arcuate striations. The thoracic uncini (fig. GENUS SALMACINA CLAPARADE, 1870 133) are provided with what appear to be eight curved teeth and a large basal tooth. Salmacina amphidentata, new species Actually the "eight" teeth represent approxi- Figures 129-138 mately four rows of teeth (fig. 134); the large DIFFERENTIAL DIAGNOSIS: Salmacina with basal tooth is but a single tooth. collar setae, the basal toothed area of which Abdominal uncini (fig. 135) are smaller bears spines of various sizes and is separated than those of the thorax but appear to bear 10 from the apical blade by a wide shallow notch. curved teeth (approximately five rows in U.C.W.I. COLLECTION: An aggregation of "frontal" view; fig. 136), with a longer, from 35 to 40 individuals, many of which are stouter, single basal tooth. Abdominal neuro- ovigerous, were collected at Port Royal, podial pointed capillaries are of two types: 1962 JONES: POLYCHAETOUS ANNELIDS 205 larger gently curved setae (fig. 137) which are spine which bears a small knob at the summit dentate on the outer margin of the curve, and of its curve. smaller setae (fig. 138) which are provided U.C.W.I. COLLECTION: Four specimens of with a single denticulate limbation. Eupomatus alatalateralis were collected at The trivial name refers to the gradation of Port Royal, Jamaica, in May, 1960. All speci- the size of the spines on the basal toothed mens are approximately 30-35 mm. in total prominence of the collar setae. length, of which the branchial plumes com- TYPE DISPOSITION: The holotype (A.M.- prise approximately 4 mnm. Diameters are N.H. No. 3615) and paratypes (A.M.N.H. about 2 mm. No. 3616) of Salmacina amphidentata are de- DESCRIPTION: The branchial plume is made posited in the collections of the American up of about 22 pinnate filaments. The oper- Museum of Natural History. cula (and pseudo-opercula, where they occur) DIscusSION: Salmacina amphidentata can arise from the base of the plume on the dorsal be differentiated most readily from other spe- side. The opercular peduncle is smooth and cies of Salmacina by the structure of the terminates apically in a crown composed of a largest of the collar setae. It is pointed out proximal circlet of short teeth and a distal above that the large collar setae of S. amphi- circlet of spines (fig. 139). There are from 36 dentata are characterized by a denticulate to 43 teeth in the proximal circlet, and these prominence which is set apart from the finely are centrifugally oriented. The 15 or 16 spines toothed apical wing by a wide notch, and by of the distal circlet each have a recurved tip the gradation in size of the teeth of the basal which is centripetally oriented. There are prominence, the more proximal teeth being limbations on each side of the upright portion finer than the more distal. In S. australis of each spine (figs. 139, 140). At the base of Haswell, 1884, the basal prominence is closely each spine (fig. 140) there is a shorter, in- associated with the distal blade, and there is wardly directed spine which curves down- no notch separating the two regions. The col- ward. At the summit of the curve of these lar setae of S. setosa Langerhans, 1884, are basal spines, there is a small protuberance. provided with a V-shaped notch between the Except for the lateral limbations, there are no proximal area and the distal wing. Salmacina spines or any other ornamentation on these dysteri (Huxley), 1855, and S. incrustans spines of the distal circlet. Claparede, 1870, show a wide shallow notch The thoracic membrane of Eupomatus ala- (similar to that of S. amphidentata), but the talateralis extends to the second or third ab- basal area of S. incrustans is provided with dominal setiger. In the anterior region of the four to six large teeth, while that of S. dysteri thoracic membrane, there is a notopodial col- has numerous fine teeth. lar fascicle of setae which is unaccompanied Although Hartman (1959, p. 591) includes by a neuropodial torus of uncini. There are an operculate serpulid, Salmacina tribranchi- two types of setae in the collar fascicle. The ata (Moore), 1923, in this genus, I feel that first is a heavy, pointed seta which bears two this species cannot belong to Salmacina, for short, heavy teeth in the mid-region (fig. by definition (Claparede, 1869, p. 176; 1870, 141). The other collar seta (fig. 142) is a pp. 518-519), the genus includes only non- longer, more slender, pointed seta which may operculate forms. Therefore, I suggest that be finely limbate. The remaining notosetae of Salmacina tribranchiata be returned to the the thorax are unilimbate and pointed (fig. genus Filograna. 143); their margins are smooth and unorna- mented. Thoracic uncini are provided with a GENUS EUPOMATUS PHILIPPI, 1844 single row of from six to seven heavy, curved Eupomatus alatalateralis, new species teeth (fig. 144). The abdominal region of Eupomatus alata- Figures 139-146 lateralis is provided with notopodial uncini DIFFERENTIAL DIAGNOSIS: Eupomatus which are somewhat smaller than those of the with an operculum that is provided with a thorax and which have a single row of six distal calyx, the spines of which point in- curved teeth (fig. 145). Uncinigerous tori of ward; and with distal opercular spines pro- the abdominal region nearly meet at the vided with lateral limbations and a basal dorsal mid-line. Abdominal neurosetae (fig. 206 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 124

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FIGS. 139-146. Eupomatus alatalateralis, new species. 139. Operculum. 140. Spine from distal circlet of operculum. 141, 142. Collar setae. 141. Larger type. 142. Second type. 143. Thoracic notoseta. 144. Thoracic uncinus. 145. Abdominal uncinus. 146. Abdominal neuroseta. 139, scale A; 140, scale B; 141, scale C; 142-146, scale D. 1962 JONES: POLYCHAETOUS ANNELIDS 207 146) have slender shafts which are apically man (1951b, p. 119), "The shorter [spines of expanded. These are asymmetrical and ap- the distal calyx] . .. have a short boss or pear to be slightly flared like a trumpet. The spine, about half way down, on the side facing margin of the elongate oval opening of these the center and the outer side is geniculate setae is provided with fine teeth of varying near the same place, on its outer side." How- sizes. ever, Morch (1863, p. 378) has stated, con- Eupomatus alatalateralis was collected in cerning the spines of the distal calyx, that, on smooth, white, calcareous tubes, the only these spines, there is a " . . . spina parva in ornamentation of which was fine, transverse, latere externo paululum infra medium sita." growth lines. Further, Fauvel (1919a, pp. 478-479) ob- DISCUSSION: The various species of Eupo- serves that the distal spines of E. sanctae cru- matus are separable into general groups on the cis may be curved in or out, or both, but, re- basis of the structure of the operculum. The gardless of the direction of curvature, each is groups, based for the most part on original geniculate and there is an outer spine. Fauvel descriptions or figures presented by subse- also presents illustrations of both types of quent reputable workers, are as follows: distal spine (Fauvel, 1919a, fig. 2f-h). 1. Species with opercula that are provided Previous to the present description, only with a distal calyx, the spines of which are not two species of Eupomatus [E. intereans Cham- all the same size, as regards their thickness or berlin, 1919, and E. spongicola (Benedict), length, or both: E. albiceps (Grube), 1870 1887] had been described as possessing a distal [also Pixell, 1913]; E. dianthus (Verrill), 1873 calyx comprised of thin attenuated spines [also Hartman, 1945]; E. exaltatus Marenzel- which are inwardly curved and symmetric- ler, 1884 [also Pixell, 1913]; E. helmatus Iroso, ally arranged (as in the case of E. alatala- 1921 [also Fauvel, 1927, and Rioja, 1947]; E. teralis). The separation of Eupomatus alata- inermis (Monro), 1933; and E. ralumianus lateralis from E. intereans is based on Cham- (Augener), 1927. berlin's comment that the incurving distal 2. Species with opercula that are provided spines of the operculum of E. intereans are not with a distal calyx, the spines of which spread provided with basal teeth (Chamberlin, centrifugally or are erect, not incurving: E. 1919a, p. 23). Eupomatus alatalateralis is so elegantulus Bush, 1910; E. fusicola (M6rch), provided with internally oriented basal teeth 1863; E. gracilis Bush, 1904; E. humilis Bush, (figs. 139, 140). The separation of E. alata- 1904; E. lunulifer Claparede, 1870; E. lateralis from E. spongicola can be made on novae-pommeraniae (Augener), 1923; E. pro- the basis of several structural differences. The tulicola (Benedict), 1887; and E. uncinatus distal opercular spines of E. alatalateralis Philippi, 1844. have a more sharply recurved basal spine 3. Species with opercula that are pro- which bears a boss at its summit (fig. 140), vided with a distal calyx, the spines of which and the main spine is provided with lateral are asymmetrically arranged: E. dipoma limbations (figs. 139, 140). There is no boss on Schmarda, 1861; and E. similis Treadwell, the basal spine of E. spongicola, nor are limba- 1929. tions present on the main spine (Benedict, 4. Species with opercula that are provided 1887, pl. 20, fig. 12). Moreover, E. alatala- with a proximal calyx, the spines of which teralis has approximately 22 branchial fila- have T-shaped apices: E. dirampha (M6rch), ments and 36 to 43 teeth in the basal opercu- 1863; and E. gairacensis (Augener), 1934. lar calyx, while E. spongicola has 30 branchial The remaining species require some com- filaments and about 65 teeth in the basal ment and discussion. The descriptions of E. opercular calyx. blumenbachii (Morch) (1863, p. 373), E. The trivial name refers to the limbations plateni Kinberg (1867, pp. 351-352), and, as that are to be found on the sides of the spines given by M6rch (1863, p. 372), E. euplaeana of the distal opercular circlet. (delle Chiaje), 1828, are not sufficiently de- TYPE DISPOSITION: The holotype (A.M.- tailed to be determined. N.H. No. 3617) and paratypes (A.M.N.H. Some confusion has arisen concerning the No. 3618) of Eupomatus alatalateralis have spines of the distal calyx of E. sanctae crucis been deposited in the collection of the Ameri- (Kroyer) in M6rch, 1863. According to Hart- can Museum of Natural History. LITERATURE CITED

ANDREW, WARREN, AND NANCY V. ANDREW seen von Leiden und Amsterdam. II. 1953. Some and sipunculid worms of Ibid., vol. 16, pp. 107-128. the Bimini region. Amer. Mus. Novi- 1934. Polychaeten aus den zoologischen tates, no. 1617, pp. 1-16. Museen von Leiden und Amsterdam. ARWIDSSON, IVAR IV. (Schluss). Ibid., vol. 17, pp. 67-160. 1907. Studien uber die skandinavischen und Aziz, NAJMUD-DIN arktischen Maldaniden nebst Zusam- 1938. Fauna of Karachi. 2. Polychaetes of menstellung der ubrigen bisher be- Karachi. Mem. Dept. Zool. Panjab kannten Arten dieser Familie. Zool. Univ., vol. 1, pp. 19-52. Jahrb., Abt. fur Syst., vol. 25, pp. BAIRD, WILLIAM 1-308. 1865. On new tubicolous annelids, in the col- AUGENER, HERMANN lection of the British Museum. Part 2. 1906. Reports on the results of dredging, un- Jour. Linnean Soc. London, vol. 8, pp. der the supervision of Alexander Agas- 157-160. siz, in the Gulf of Mexico and the BENEDICT, JAMES E. Caribbean Sea, and on the east coast of 1887. 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Selsk. Forhandl., Co- Occas. Paper, no. 23, pp. 1-628. penhagen, pp. 1-36. 1961. Polychaetous annelids from California. KtKENTHAL, WILLY Allan Hancock Found. Publ., Pacific 1887. Die Opheliaceen der Expedition der Expeds., vol. 25, pp. 1-226. "Vettore Pisani." Jenaische Zeitschr. HASWELL, WILLIAM A. Naturwiss., vol. 21, pp. 361-373. 1884. The marine annelides of the order LANGERHANS, PAUL Serpulea. Some observations on their 1880. Die Wurmfauna von Madeira. III. anatomy, with the characteristics of the Zeitschr. Wiss. Zool., vol. 34, pp. 87-143. Australian species. Proc. Linnean Soc. 1884. Die Wurmfauna von Madera. IV. Ibid., New South Wales, vol. 9, pp. 649-675. vol. 40, pp. 247-285. HOAGLAND, RUTH MCINTOSH, WILLIAM C. 1919. Polychaetous annelids from Porto Rico, 1885. Report on the Annelida Polychaeta col- the Florida Keys, and Bermuda. Bull. lected by H.M.S. Challenger during the Amer. Mus. Nat. Hist., vol. 41, pp. years 1873-76. 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