R e~. BioL Trop. 46. Supl. 6: 41-62, 1998 www .ucr.ac .cr

The Pilargidae (Annelida: Polychaeta) of the Pacific Coast of Costa Rica

Harlan K. Dean Department of In~ e.rtebmte Zoology, Museum of Com pamti~e Zoology, 26 Oxford St., C3mbridge. MA 02138 USA, and John Hazen White School of Arts and Sciences, Johnson and Wales Uni~ersi t y. 6 Abbott Parle Plllce, Pro~iden ce, Rhode Island 02903, USA. E-m ail: [email protected]

(Ree. 22-X-I997. R e~. 26-111-1998. Acep. 14.v1- 1998 j

Abstract: Thirteen species of Pilargidae (Annelida: Poly ehaeta) from the subtidal and inlerth.lal Pacific Coast of Costa Rica are reported. A new species, Sigambra vargasi n. sp., is described and an as yet undescribed species, Si­ gambra sp. A, is also reported. The occurrence of Sigambra 'en/aeu/ara (Treadwell). Loandalia riojai Salazar­ Vallejo, Pilargls berkeleyl Monro, ArrclJlrosyl//s nama/a H3rtman and AneWrosyllis jonesei Peltibone are range extensions for these species. The occurrence of Sigambra pellibaneae Hartmann-SchrOder, Pilrandillia Iricuspi! (MUlier), Cubira inartu Webster, Ancis/rosyllis papillosa Jones and Ancislrosyllis narlmanae Pettibone are new re cords of these species for the eastern Pacific Ocean. Synelmls cf. albini (Lan gcrhan s) is also reported. A taxono­ mic key to these species is included.

Key words: Polychaeta, Pilargidae, Pacific coast, Costa Rica.

The Pilargidae are a family of polychaetes mainly by the presence of emergent notopo­ reported in sediments from the intertidal to dial spines or hooks, two pairs of tentacular abyssal depths. Since these organisms are usu­ cirri and entirely simple neurosetae. More ally quite small and present in low densities recent results of cladistic analysis by Licher there is very little information regarding their and Westheide (1994) have indicated that the ecology (Faucha1d & Jumars 1979). They do Pilargidae might be more appropriately inclu­ not build a tube and have a muscular eversible ded within the family Hesionidae and, in their pharynx which is presumed to be used in the review of the genus Sigambra (Licher & capture of prey or perhaps intake of detritus Westheide 1997), regarded the Pilargidae as a and microa1gae (Day 1967). "subtaxon" of the Hesion idae. Further study The systematics of the Pilargidae is also may support this view, however, until such poorly known. Historically, species have been time the family Pilargidae is herein retained assigned to the Orbiniidae (MUlier 1858), the (see Fauchald & Rouse 1997). Syllidae (Mcintosh 1879. Webster 1879), and Relationships of the genera within the Pi­ quite often the Hesionidae (Ehlers 1908; largidae have always been difficult due to the Fauvel 1920. 1923, Augener 1927, Monro wide range of character states. Salazar.vallejo 1933, Berkeley & Berkeley 1941 . and Tread­ (1987(1986» divided the Pilargidae into two well 1941). Eventually Hartman (1947) and subfamilie s, the Synelminae with cyl indrical Pettibone (1966) united a disparate group of body and emergent notopodial spines, and the genera into the family Pilargidae characterized Pilarginae ("'Sigambrinae) with a dorsoven- 48 REVISTA DE ()fOLOG1A TROPICAL trally depressed body and either emergent MATERIALS AND METHODS nOlopodial hooks or emergent nOlopodial setae lacking. These subfamilies are nol supported Subtidal material from the Gu lf of Nicoya was by the results of cladistic analysis based on collected during a cooperative research effort gross brain morphology by Fitzhugh and Wolf between the University of Costa Rica and the (1990) or the analysis of Licher and Wes­ University of Delaware (USAXsee Maurer & theide (J 994) based on 28 morphological Vargas (\ 984), and Vargas (1995) for loca­ characters. Licher and Westheide (1994) iden­ tions and review). Intertidal material from the tified several clades or groups of genera cha­ Punta Morales region of the Gulf of Nicoya racterized by such characters as presence of a was collected by the author during a series of papillated integument or gross morphology of collecting trips supported by the University of the hind brain and indicated that the Pilargidae Costa Rica 's Centro de Investigaci6n en Cien­ may not be a monophyletic group. cias del Mar y Limnologia (CIMAR). Subtidal Despite the confusion regarding phyloge­ material from Golfo Dulce was collected by netic relationships, the species within the Pi­ the R. V. Victor Hensen Costa Rican ex­ largidae have been well characterized. espe­ pedition (1993 / 1994) and material from that cially in the eastern Pacific Ocean. Two spe­ cruise provided by J. Vargas and R. Le6n cies have been recorded from the region be­ (C IMAR). Specimens from the intertidal of tween Alaska and Washington State (USA) by Golfito Bay, Golfo Dulce were collected by J. Monro ( 1933) and Banse & Hobson (1974). Vargas (CIMAR). Voucher specimens of most The pilargids of the California coast (USA) of the listed species have been deposi ted in the have been well sampled and nine species re­ Museo de Zoologia, Universidad de Costa corded by Hartman ( 1947, 1968) and Emerson Rica. The remainder of the material is deposi­ & Fauchald (1971). Salazar-Vallejo (1987 ted in the Museum of Comparative Zoology, (1986), 1990) and Le6n-Gonzalez (1991) have Harvard University, Camhridge, MA , USA. sampled extensively along the Pacific coast of Mexico and published several keys and spe­ Key 10 the Pilargidae of Ih e Pacific cies li sts which include 18 species from this coast of Cosla Rica region. Hartmann-SchrOder (1959) recorded two species of Pilargidae from the mangroves I . Integument smooth...... 2 of EI Salvador. Synelmis albini (Langerhans) I a. Integument papillate ...... 8 was also included in collections by Westheide (1974) from the Galapagos Islands and 2. Antennae and tentacu lar cirri present ...... 3 Fauchald (1977) from the Pacific side of Pan­ 2a. Antennae and tentacu lar cirri absent ...... 7 ama. In prelimi nary lists of polychaetes from the Gulf of Nicoya and Golfo Dulce, Costa 3. Three weakly developed antennae, furcate Rica, Dean (1996a,b) reported 10 species be­ neuropodial spines present. (Fig. 1) ...... longing to the family Pilargidae...... Syne/mis ef a/bini (Langerhans) This paper reviews 13 species of Pilargidae 3a. Three well developed antennae present, collected from the Paci fic Coast of Costa Ri ca. dorsal parapodial cirri of setiger I longer Four species are added to the lists of Dean that those of other setigers ...... 4 (1996a, b) while specimens originally identi­ fied as Sigambra bassi are herein described as 4. Emergent notopodial hooks from setiger 4. a new species, S. vargasi n. sp. Another spe­ (Fig. 2) ... Sigambra lenlaeu/ala (Treadwell) cies included in the list of Pilargidae by Dean 4a. Emergent notopodial hooks begin poste· ( I 996a), Synelmis Ielalli Friedrich, has been rior to setiger 4 ...... 5 moved to the family Hesionidae and redes­ cribed by Licher ( 1994). Specimens identified 5. Emergent notopodial hooks from setigers as S. /dalli from Costa Rica (Dean I 996a) 8-9. (Fig. 3) ...... sigambra have been descri bed as a new species of petfiboneae Hartmann-SchrMer Glyphohesione in the fam ily Hesionidae Sa. Emergent notopodial hooks begin poste- (Dean 1998). rior to seliger 10 ...... 6 Aql,latie c ~OSySlc ll1$ of Costa R i~a 49

6. Emergent notopodial hooks from setiger 12. leaving on ly 2 species S. albini (Langemans) (Fig 4) ...... sigambra sp A and S. simplex Chamberlin in the genus with 6a. Emergent notopodial hooks from setigers the synapomorphy of two furcate neurosetae. 14·I S. ( Fig. S·I2) .. .sigambra vargasi n. sp. Synelmis cf albini ( Langemans 1881 ) 7. Ventrolateral branchiae present. (Fig. 13) ... (F;g. I) ...... Loandali a riojai Salazar· Vallejo 7a. Ventrolateral branchiae absent. ( Fig. 14) ... Synelm is albini (Langemans). Pettibone 1996: ...... Parondolio Iricwpis (MUller) 191 . Fauchald 1977:17, Westheide 1974. Sala­ zarNallejo 1987 ( 1986): 20S. Maurer & Var­ 8. Parapodia weakly developed. proboscis gas 1984:10 1 (in part), Maurer et a/. 1988:48 with dentic1es. (Fig. IS & 16) ...... (in part), Dean 1996a:74...... Cabiro incerla Webster 8a. Pharynx not cuticularized. parapodia well Materillli Examined.G ulf of Nicoya, Sla. J, developed ...... 9 9"S7')0"N, 84°S3'OO"W, 46 m, mud. Jul y 1980,(IS) and (I XUSNM 079952). Sla. 2, 9° 9. Emergenl nOlopodial spines or hooks absent 55'28"N, 84°S2'OS"W. 18 m, muddy sand, Ju ly (Fig. 17) ...... Pi/argis berkeleyi Monro 1980,(2XUSNM 079973). Sla. 3, 9"S2'00"N, 9a. Emergent dorsal hooks present. Genus 84°48'10"W, 33 m, sandy mud, July 1980, ( I) Ancislrosyllis ...... 10 (USNM 0799S3). Sio. 4, 9°S3'40"N, 84°46' 10"W, 40 m. mUd. July 1980,(1 ) ( USN M 10. Two antennae present ...... 11 0799S4). Sta. 13. 9"S2'30"N. 84°43'SO"W, 26 103. Three antennae present ...... 12 m, July 1980.(3)(USNM 07995S & 0799S6). Sra . 14, 9"57'05"N, 84 °4 S ' 3 0 ~ W , 9 m, sandy I I. Emergenl notopodial hooks from seti· mud, Jul y 1980,(2S)( USNM 079966 & 07996 gers 4·7, dorsal parapodial cirri with 7). Sta. 15. 9°S7'40"N, 84°47'OO"W, IS m, small papillae. (Fig. 18) ...... sandy mud, July 1980, (7I XUSNM 079974 & ...... Ancistrosyllis hamata Hartman 0799S5). Sta. 13, 9"48'3S"N, 84°43'SO"W, 35 11 3. Emergenl notopodial hooks from setiger m, mud, July 1980,(1) (USNM 079959). Slo. 3. dorsal parapodial cirri with prominent 24, 9°49'2S"N. 84°4I'20"W. 11 m. sand, Oct elongated papillae. (Fig. 19) ...... 1980,(27); Jan 1981 ,( I); Apr 1981 ,( 11 ); Aug ...... Ancistrosyllis papillosa Jones 1981,(7). Sta. 25, 9"SO'OS"N, 84°S2'OO"W, 20 m, mud/sand, July 1980,(J XUSNM 079960). 12. Emergent notopodial hooks from setiger 6. Sta. 26, 9°Sl'SO"N, 84°S3'20"W, 17 m, muddy (Fig. 20) ...... Ancistrosyllisjonesi Pettibone sand, Jul y 1980,(I)(USNM 079961 ). Sla. 27, 12a. Emergent notopodial hooks from seti· 9"S I'57"N, 84°50'SO"W, 12 m, muddy sand, ger 3, fle xed anterior region of the di· July 1980,(2) (USNM 079963 & 079963). Sra . gestive tract. (Fig. 2 1) ...... 28, 9°S2' 16"N, 84°4S'30"W, 26 m, mud, June ...... Ancistrosyllis harlmonae Pettibone 1981,(1). Sta. 29, 9"S4'S5"N, 84°45' 15"W, 18 m, muddy sand, Oct 1980,( 18); Jan 1981 ,(32); Apr 198 1, (4); June 1981 , (4); Aug 198 1. (20); Species Descriptions Apr 1982,(22). Sta. 30, 9"S4'40"N, 84°4S' SO"W, 18 m, muddy sand, Oct 1980,(11); Jan Family Pilargidae 1981 ,(6); Apr 198 1,(SI ); June 1981,(4). Sla. 31, 9°44'OO "N, 84"59'2S"W. 20 m, mud/sand, Genus Synelmis Chamberlin 1919 July 1980,(5 1XUSNM 079947, 079948, 0799 50 & 07996S). Sla. 32, 9"S3'47"N, 84°49' Fitzhugh & Wolf (1990) separated this genus 3S"W, 24 m. mud/sand, July 1980,(2).(USNM into "complex A" species with two furcate 079971). S IO . 34, 9"SS'30"N, 84°S0'OS"W, 24 neurosetae and "complex B" species lacking m, sand, Ju ly 1980,( 12)(USNM 079949 & furcate neurosetae. Based upon their cladistic 0799S I). Sta. 35, 9"SS'42"N, 84°47'40"W, 13 analysis Licher & Westheide (1994) united the m, sand, July 1980,(32)(USNM 079976 & 07· "complex B" species with the genus Litacorsa 9977). Sla. 37. 9"S7'38"N, 84°48'20"W, 14 m, so REV1STA DE 1lI0LOGIA TROPICAL muddy sand, July 1980, (2 1) (USNM 079978 With two pi gmented cyespots and a cyl indri­ & 079979). Punta Morales, IO"N, 8S oW, in­ cal pharynx (Fig. 1). Emergent nOlopodial tenidal, coarse sand, Aug 1996;(12, UCR spines beginning at seliger 11 accom pan ied by 109). one or two thickened notacicula. Capi llary ncurosetac include one or IWO longer and two Remarks. Com plete specimen 136 seligers, or three shoner setae as we ll as one or two approximately 50 mm long and 0.7 mm wide. furcate setae in a single fascicle .

1 2

.. " ,

4

Figs. 1-4. Fig. I. Synelm;J cf albmi. dorsal view of anterior. Fig. 2.S'ga",brn Itnlaf;fl/U/(l. dorsal view of anterior. Fig. 3. Sigamhrapelliboneat, dorsal vicw ofanterior. Fig . 4.sigambra sp . A. dorsal view ofantcrior. Scale bar" ] mrn . AQLlalic ecosystems ofCOSla Rica " Distribution. One of the most common poly­ mm long, maximum width 0.8 mm (0.6 not chaetes from the Punta Morales intertidal from including parapodia). NOlOpodial emergent a wide range of sediments. What has been hooks from setiger 4, ventral cirrus lacking on identified as Synelmis afbini (Langerhans) has setiger 2, median antenna longer than lateral been reported from the Galapagos Islands antennae, tips of neurosetae entire, notosetae (Westheide 1974) and the Pac ific side of Pan­ absent. These speci mens lack the row of ama (Fauchald 1977). It is beli eved, however, peristomial papillae described by Katzmann er that S. a/bini may be a complex of seve ral a/. (1974) for material iden tified as S. lenla­ species and awaits further systematic analysis culala from the Mediterranean. (Wolf 1984). Distribution. I)reviously reported from New Genus Sigambra MO ller 1858 York, USA, to Argentina in the At lantic Ocean and from California, USA, to Manzani- Members of this gen us possess well devel­ 110, Col., Mex ico, in the Pacific. The material oped lateral and median antennae, tentacular identified as S. lenlaculala by Katzmann el 01. cirri, and dorsal parapodial cirri . Lieher & 1974 from the Mediterranean may be a differ­ Westheide ( 1994) regard all these features as ent species. This extends the range of thi s spe­ pleisio'morphic indicating a hi gh similarity be­ cies in the Eastern Pacific to Golfo Dulce, tween Ih is genus and juveni le hesionids. Costa Rica. Sigambra lenlaclliara has been reported from the intertidal to abyssal depths. Sigambra lelllacli/afa (Treadwell 194 I) (Fig. 2) Sigambra pellibolleae Hartmann-SchrOder 1979 ( Fig. 3) Sigambra fen/aclilata Treadwel l 1941 : I. Pet­ tibone 1966: 182. Sigambra pellibolleae Hanmann-SchrMer Maurer & Vargas 1984 : 101 (in part). Wolf 1979:84. 1984:29.8. Material Exa mined, Golfo Dulce: Golfito Salazar-Vallejo 1987(1986):202. Maurer el al. Bay, g038'N, 83°10'W, intertidal, muddy sand 1988:48 (in part). Salazar-Vallejo et al. 2 Km west of main dock, Sep 1986; ( 109 1990:2 16. Le6n-Gonzalez 199 1:3 14 . Sa lazar­ +7(UCR 108). Vallejo & Orensanz 1991 :275. Blake 1997: 275. Dean 1996a:74; 1996b:83. Remarks. Complete specimen 59 setigers, 5. 8 mm long, 0.7 mm max imum width (0.5 mm Material Examined. G ulf of Nicoya, Sla. 6, wi thout parapodia). Emergent notopodial 9"4 S'20"N, 84°46'25"W, 42 m, mud, Jul y hooks beginn ing at seligers 8-9, ventral cirri 1980,( IXUSNM 079936). Sla. 22, 9"48'25"N, absent on setiger 2, and the median antenna 84°S2'40"W, 22 m, muddy sand, July extends to setigers 4-6. The neurosetae include 1980,( IXUSNM 079937). Slo. 24, 9°49'25"N. short coarsely toothed and longer capill ary 84"4 I'20"W, II m, sand, Oct 1980,( 1); Apr setae. 198 1.(1); Jun 198 1,( 1); Aug 198 1,( 1). Sta. 29, 9°54'55"N, 84°45' 15"W, 18 01 , muddy sand, Distribution. Type locality is intenidal sand Jan 198 1,(2). Sla. 30, 9"54'40"N. 84°45'50"W, with organic detritus from Broome in north­ 18 m, muddy sand, Jan 1981,(3). Sla. 31, western Australia. This is the first record of 9°53'47"N, 84°49'35"W, 24 m, mud/sand, July this species from the eastern Pacific and the 1980,( IXUSNM 079944). S/a. 33, 9"53'40"N, first report of this species outside of the type 84°53'20"W, I I m, sandy mud, Ju ly 1980, locality. (IXUSNM 079942). Punta Morales, 10"N, 85°W, intertidal, Aug 1996:(11 ), mud; (8), Sigambra sp. A sandy mud; (II + 3(UCR 1100I »,coarse sand. (Fig. 4) Golfo Dulce, Sla. CDI2, 8"2 I'N, 83" 14'W, 200 m, mud, Dec 1993, ( I). Material Exa mined, Golfo Dulce, Colfito Bay, 8°38'N, 83°IO'W, intertidal, muddy sand Remarks. Complete specimen 57 setigers, 7.3 2 Km west of main dock, Sep 1986;( 1). " REVISTA DE BI OLOGIA TROPICAL Remarks. Complete specimen with 48 seti­ disregard by Licher & Westheide (1997) for gers, 3.9 mm in length and 0.5 mm in width the significance of the first appearance of no­ anteriorly (0.35 mm without parapodia). Body topodial hooks in their review of the genus. wide anteriorly, tapering posteriorly; dorso­ ventrally flattened with deeply incised Sigambra vargasi, n. sp. parapodia. Color in alcohol light tan. (F;g.5·12)

Prostomium with indistinctly separated biar­ Material Examined. Gulf of Nicoya, Sta. I, ticulated palps; palpophores large, anteroven­ 9°57'30"N, 84°53'OO"W, 46 m, mud, July !ral palpostyles small, digitate. Median an­ .l980,(2XUSNM 079935). Sta. IS, 9"57'40"N, tenna arising from posterior of prostomium, 84°47'00"W, 15 m, sandy mud, July 1980, extending to fifth seliger; lateral antennae (lXUSNM 079946). Sta. 23, 9°48'35"N, 84° ari se slightly anterior to median antenna, ex­ 43'50"W, 35 m, mud, July 1980,(IXUSNM tend to the first seliger. Proboscis with 14 079938). Sta. 24, 9°49'25"N, 84°4 1'20"W, II conical distal papillae. Two pair subequaJ m, sand, Oct 1980,(1) ; Jan 1981,(5) ; Apr tentacular cirri from peristomium, cirri slightly 1981, (5); June 1981,(10); Aug 1981,(16). Sta. longer than lateral antennae. Dorsal cirri of 25, 9°50'05"N, 84°52'00"W, 25m, mud/sand, seliger 1 elongate, extending to the fifth seti­ July 1980, (4)(USNM 079939). Sta. 26, 9°51' ger. Ventral cirri absent from setiger 2. 50"N, 84°53'20"W, 17 m, sandy mud, July 1980,(IXUSNM 079940). Sta. 27, 9°51'57"N, Notopodia with a single acicula, accompa­ 84°50'50"W, 12 m, muddy sand, July 1980, nied by a stout emergent notopodial hook be­ (I) (USNM 079941). Sla. 28, 9°52'16"N, 84° ginning at setiger 12, a capillary seta may also 45'30"W, 26 m, mud, July 1980,(18); Oct occur in posterior notopodia. Neuropodial 1980, (18) (MCZ 20019); Jan 1981,(4) ; Apr lobe conical with numerous capillaries, shorter 1981 , (1 + 5(UCR 1602» ; June 1981 ,(12); capillaries with fine denticulations, and short Aug 1981,(17). Sta. 29, 9°54'55"N, 84°45' pectinate setae. Dorsal cirri extend beyond 5"W, 18 m, muddy sand, Jan 1981,(1); Aug setal lobe; ventral cirri subequal in length to 1981,(21); Apr 1982,(6). Sla. 30, 9°54'40"N, setal lobe. Pygidium with pair of long anal 84°45'50"W, 18 m, muddy sand, July 1980, cirri. (1)(USNM 079943); Oct 1980,(2); Jan 1981, (6); Apr 1981,(6); June 1981,(4); Aug 1981, This spec ies most closely resembles a re­ (I+I(HOLOTYPE MCZ 20017)+2 (PARA­ cently described species Sigambra phulcetensis TYPES MCl 20018). Sta. 32, 9°53'47"N, Licher & Westheide 1997 in that they both 84°49'35"W, 24 m, mud/sand, July 1980,(6) have ventral cirri on the second setiger, 14 (USNM 079944). Sta. 36, 9°55'32"N, 84°45' distal papillae on the pharynx lack emergent 20"W. 18 m, sandy mud, July 1980,(2) (US spines, have a longer median than lateral an­ NM 079945). tennae and possess pectinate neurosetae. While S. phulcetensis is described as having Species description. Holotype (MCZ 20017) emergent notopodial hooks beginning at seli­ 5.2 mm long, 1.0 mm wide (0.5 mm not in­ ger 3-23, S. sp A has notopodial hooks from cluding parapodia) with 60 setigers. Paratype setiger 12. The median antenna of S. sp A is (UCR 160-2) 9.7 mm long, 0.8 mm wide (0.5 much longer than the lateral antennae extend­ mm not including parapodia), with 54 setigers. ing posteriorly to the 5th setiger (4-5 times the Paratype MCZ (20019) 18.4 mm long, 0.5 mm length of the laterals) while the lateral anten­ wide (0.3 mm not including parapodia), with nae are shorter relative to the body width than 136 setigers. Body white to light brown in those reported for S. phuketensis. Addition­ ethanol. First four segments cylindrical, 3 ally, the dorsal cirri of S. puketensis are ap­ times as wide as long, middle and posterior proximately 2/3 the body width in median and setigers dorsoventrally flattened, deeply in­ posterior setigers while those of S. sp A are cised, 4 times as wide as long (Fig. 5). much shorter than that. It would be premature Prostomium two rounded lobes; two biar­ to formally describe this species based upon a ticulate palps indistinctly separate from the single specimen especially in the light of the prostomium, palpophores large with small Aquatic ecosystems of Costa Rica " anteroventral palpostyles (Fig. 5). Median that these are the only two species in the genus antenna extending to seventh setiger (x = 6.9, with an terior setigers possessing a capillary SD = 1.29, n = 10); paired lateral antennae notoseta unaccompanied by emergent hooks. attached slightly anterior to and approximately S. vargas; n. sp. differs from S. qingdaoensis 1/3 the length of median antenna. Eversible in the first occurrence of notopodial hooks at proboscis with 8 distal papillae. Peristomium seliger 15-1 7 rather than setigers 3-8, the with 2 pair tentacular cirri, dorsal slightly much greater length of the median antenna longer than ventral, 113 length of lateral an­ and the greater length of the anal cirri. tennae. Distribution. From II to 46 m in muddy to First setiger with elongate dorsal cirrus 4-5 sandy sed im ents. setigers in length (Fig. 9). Setiger 2 with short dorsal cinus, ventral cirrus lacking (Fig. 10). Etimology. This species is named for Prof. Setigers 3-5 with subequal dorsal and ventral Jose A. Vargas (C IMAR) for his many years cirri extending beyond the parapodial lobe of work furthering benthic studies in Costa (Fig. II). Remain ing setigers with dorsal cir­ Rica. rus approximately twice as long as ventral cinus, ventral cinus subequal in length to Loandalia, Ptuandalia .{ TalellSapia Group parapodiallobes (Fig. 12). This group of pilargids, identified by the Notopodia reduced, notacicula present at base phylogenetic analysis of Licher & Westheide of dorsal cirrus, accompanied by a short capi­ (1994), is characterized by the absence of an­ llary notoseta beginning at setiger 4. Emergent tennae, tentacular cirri, and dorsal parapodial notopodial hooks from setiger 15-1 7 (x = 15 .2, cirri, the possession of capillary notosetae in SD = 0.48, n = 31), accompanied by a single addition to the emergent spine, and a lack of notacicula and occasionally one or two short notacicula. Members of this group are inclu­ capillary setae. In median and posterior seti­ ded within the Pilargidae based mainly on the gers notopodial hooks may be replaced by a presence of emergent notopodial spines. These short capillary seta. genera may be considered high ly derived or possibly be unrelated to the other pilargids Neuropodia with fascicle of maximally 17-18 (Licher& Westheide 1994). simple neurosetae. Neurosetae include longer serrate bladelike setae (Fig. 6), shorter pecti­ Genus Loandalia Monro 1936 nate setae with smooth distal tip (Fig. 7), and short pectinate setae with fine teeth extending This is the only genus in the Pilargidae with to distal tip (Fig. 8); neuropodia of posterior branchiae. setigers with one or two smooth capillary setae (F;g. 12). Loandalia rioja; Salazar· Val lejo 1987( 1986) (Fig. 13) Pygidium with two long, thin, cirriform anal cirri. Loandalia rioja; Salazar-Vallejo 1987(1986): 202. Salazar-Vallejo et al. 1990:2 16. Le6n­ Remarks. Sigambra vargasi n. sp. is similar Gonzalez, 1991 :3 14 to S. oce/Iata (Hartmann-Schroder 1959), S. bidentata Britaev & Saphronova 1981, and S. Material Examined. Gulf of Nicoya, Sta. q;ngdaoensis Licher & Westheide 1997 in the 29,9°54'55'N, 84°45'15'W, 18 m, muddy sand, possession of 8 distal papillae on the pahrynx Apr 1982 ( I, UCR 2003). and the absence ofa ventral cirris on seliger 2. It is unlike S. ocellata in the absence of eyes­ Remarks. Complete specimen 65 setigers, 40 pots and the greater length of the dorsal relati­ mm long, maximum width 1.6 mm ( 1.1 mm ve to the ventral cirris. S. vargas; n. sp. lacks without parapodia). First occurrence of emer­ the bidentate neurosetae seen in S. bidentata. gent notopodial spines is on seti ger 7, dorso­ S. vargasi n. sp. is similar to S. qindaoensis in lateral fields of proboscoid organs present. REVISTA DE BIOLOGIA TROPICAL

Pigmented eyespots were not observed for this Distribution. Previously reported from BahIa specimen. Salazar.vaJlejo (1987(1986» re­ de Manzanillo, Mexico, to the west coast of ported branchiae beginning on setigers 21 to Baja California. This record extends the range 24 and branchiae first appeared on setiger 24 of L. rioja; to the Gulf of Nicoya, Costa Rica. in this specimen_ Le6n-GonzAlez (1991) re­ Subtidal from 18 to 140 meters. ported branchiae from setiger 31-39 in two specimens of L. rioja; from Baja California Genus Parandalia Emerson & Fauchald 1971 Sur.

6 I , 7 8

Figs. "j..,. Fig. 5. Sigam~ WlTgaJi , n. sp. (Ho]otype MCZ 20011). Dorsal view of anterior. Figs. 6-8_Slgonrbra vargrui (p~pe MCZ 2OO1~) . Fig. 6. Long se!""'te ~uroseta. Fig: 1. Shon pectinate neuroseta with smooth distal tip. FigS. Shon pectinate nc:urosetawlth finely tootheddLStal tip. Scale bar Fig. 5 " 100 /1m. Figs. 6-S " 20/lm.. Aquatic rcruystcms of Costa Rica " This genus resembles Loandolio but lacks The presence of dentides on the pharynx are branchiae. The other genus in this subgroup, synapomorphic for this genus. Talehsopia, differs from Porondalio in the possession of jaw like structures. Cabira incerta Webster 1879 Flg.(16& 17). Parondalio tricuspis (MUller 1858) (Fig. 14) Cabil'a incerta Webster 1879:267. Pettibone 1966:178. Wolf 1984:29.5. Salazar-Vallejo & Parandalia trjcuspi.~ (MUller) Salazar-Vallejo Orensanz 1991 :272. 1990:5 12, Salazar-Vallejo & Orensanz 1991 : AncistrosylJis sp. (in pan) Maurer et al. 268, Dean 1996a:74. 1988A8. Cabiro sp. A. Dean 1996a:74. Porondalia sp. A. Wolf 1984:29.30. Material Examined. Gulf of Nicoya, Sla. 24, Parondalio sp. Maurer et al. 1988:48. 9°49'25"N, 84°41'20"W, II m, sand, Apr 1981 ,(2); Aug 1981 ,(1 + I , UCR 6802)). Slo. Material Examined.Gulf of Nicoya, S10. 23, 28. 9°52'J6"N, 84°45'30"W, 26 m, mud, Ju ly 9°48'35"N,. 84°43'50"W, 35 m, mud, July 1980, ( 1)(USNM 079930); Aug 1981,(1). Sto. 1980,(1) (USNM 079934). Sta. 24, 9°49'25"N, 30, 9"54'40"N, 84°45'50"W, 18 m, muddy 84°41 '20"W, II m, sand. Jan 1980,(1XUSNM sand, Apr 1981,(3). Punta Morales, JO"N , 080033); Oct 1980, (3 + I , VCR 2302); June 85"W, intertidal, coarse sand, Aug 1996; (3), 1981,( I )(USNM 080032); Aug 198 1,( I). Remarks. Complete specimen 35 setigers, 6.9 Remarks. Complete specimen 48 seligers, 6.4 mm long, 0.6 mm maximum width. Variable mm \ong .. anterior 7 setigers 0.8 mm wide, number (6-1 7) of swollen anterior segments, remaining setigers maximum width 0.7 mm emergent dorsal hooks frol11 setiger 7, and 24 (0.5 without parapodia). These specimens smooth capillary neurosetae. Wolf ( 1984) re­ agree with the new combination by Salazar­ ported neurosetae with a minutely bidentate Vallejo (\990). Ventral parapodial cirri first tip but this was not true of these specimens. appear on setiger four and emergent notopo­ Salaur-Vallejo & Orensanz (1991) reported dial spines lacking a hood on seliger 7. Eye­ ventral parapodial cirri on all setigers for spots were weakly pigmented, one spec imen specimens from Rio La Plata, Uraguay-Ar­ with only a single eyespot on the right side. gent ina, but ventral parapodial cirri begin on Posterior neurosetae in two series with ap­ setiger 3 in the Costa Rican specimens as was proximately 10 neurosetae per parapodia. All previously described for C. incerto by Petti ­ parapodia biramous. bone ( 1966) and Wolf(1984).

Distribution. Previously known from the up­ Distribution. Previously reported from Chesa­ per Gulf of Mexico to Argentina in the Atlan­ peake Bay, USA to Rio La Plata, Uruguay· tic Ocean. This is the first record of this spe­ Argentina in the Atlantic Ocean. This is the cies in the eastern Pacific. P. Iricuspis has first record of this species in the Eastem Paci­ been collected from the intertidal to 55 meters fic. Known from the intertidal to 80 meters depth. depth.

Anclstrosyllls group Genus Pilargis Saint-Joseph 1899

The genera Arn::istrosyllis, Pilorgis, Cabira, This genus is characterized by the absence of and Paracobiro are considered a monophy­ emergent hooks or spines. letic group by Licher & Westheide (1994) due to the papillated integument. Pilargis berkeleyae Monro 1933 (Fig. 17).

Genus Cobiro Webster 1879 Pi/argis berkeleyoe Monro 1933:673. Petti­ bone 1966:161. " REVISTA DE BIOLOOIA TROPICAL

10

12

Figs. 9·\2. SifO".b1'tH'u~i n. sp. (Parltype MCZ 20018). Fig. 9. Seliger 1, anterior view. Fig. 10. Setiger2, Interior vi(;w. Fig. II . Setiger), If1lcriorview. Fig. 12. Seliger 19,anteriorview. Scale b. Fig. 9-40 )1m. Fig.s 10.12 -)1m. Aquatic cCOS),51Cms ofCosUt Ri c. "

n

14

15 16

FilS. 13·16. Fia. 13. Lomd:JJia riC!/oi, dorsJIvicw ofillterior. Fig. 14.Parafldolla t,icu.JpIJ, dorsal view ofJlllerior. Fil. 15. Cobini' irIc¥,Ia,llknIview of..mor. fi&- 16.Cabil"tl illc",o , dorSlI view or prouomiunund pcrislomiunl.. SelIc bIr • I n.n. l8 REvrSTA DE BIOLOGIA TROPICAL

17 18

19

20

Figs. 17-21. Fig. 17. Pifarg~ beruleyae, dorsal view of anterior. Fig. 18.Ancis/fW)1/is hamata , dorsal view of ante rior. Fig. 19. AncutrosyJlis papillQ$Q, dorsal view of anterior. Fig. 20.Ancislrosyllisjcmesi , dorsal view ofanle rior. Fig. 2 l. AncislrosyiJis lIar/manat, dorsal view of anterior. Scale bar " I mm. Aquatic ecosystems of Costa Ri ca " Im ajima 1987: 162. Salazar.Vallejo & Oren­ 16"N , 84°45'30"W. 26 m. mud, July 1980. sanz 199 1:274. Dean 1996a:74. Blake (I)(USNM 079929); Aug 1981 .(1). Sta. 29, 9" 1997:270. 54'55"N. 84°45'1 5"W. 18 m. muddy sand. July 1980,(1); Oct 1980,(1 , UCR III); Apr 1981. Material Examined. Gulf of Nicoya, SIO. 29, (I); June 1981 ,( 1); Aug 1981 ,( 1). Sta. 30. 9°54'55 n N, 84°45' 15 "W, 18 m, muddy sand, 9°54'40"N. 84°45'50"W, 18 m, muddy sand. July 1980,( I XUSNM 079933). Oct 1980,(1); Jan 1981,( 1); Apr 1981 , (I ). Aug 1981 .( 1). Golfo Duke. Sta. COI2. Rema rks. Neurosetae with fine teeth and 08"2I'N, 83°14'W, 200 m, mud. Dec 1993,( 1). bidcntate tip, dorsal cirri in anterior setigcrs inflaled. Remarks. Complete specimen 39 set igers. 6.2 mm long. 0.55 mm maxi mum width (0.33 mm Distribution. Previously known from the without parapod ia). Vent ral cirri begin on northern Gulf of Mexico to Argentina in the setiger 3 and emergent nOl opodial begin at Atlantic and from Canada to California in the seti gers 6·7. Pacific Ocean. This report extends its range to the Gulf of Nicoya, Costa Rica. From inter­ Distribution. Prev io usly known from Califor­ tidal to abyssal depths. nia to the Gulf of Los Angeles. Baja California as well as the French Mediterranean Genus Ancistrosyllis Mcintosh 1879 (Katzmann et al. 1974). This extends its range in the eastern Pacific 10 Golfo Du lce. Costa The synapomorphy of this genus is possession Rica. Reponed from eleven meters 10 abyssal of a five lobed hind brain (Fitzhugh & Wo lf depths. 1990). The genus Allcistargis is sim il ar to All· cistrosyllis differing only in its lack. of a me· Ancislros),lIis IJQpi/losa (Jones 196 1) dian antenna. Licher & Westheide (1994) (Fig. 19). synonomize these two genera due to a lack of Ancislargis papi/loslls Jones 1961:3 a "synapomorphic character for Ancistrosyllis­ Allcistrosyllis IUlpi//osa Pettibone 1966: 170. species with three antennae" if considered Allcislrosyllis sp. Maurer et al. 1988:48 (in separate taxa. The median antenna of A IIcis­ part) largis species is usua ll y weakly developed Allcistargis hamata (1·lanman). Dean 1996a: (Fig. 11- 13) and the presence or absence of 74 (in part) such an inconspicuous structure does not war· rant separation at the generic level. Material Ex amined. Gulf of Nicoya, Sta. I . 9°57'30"N, 84°53'00"W. 46 Ill. mud, Ju ly Aneisfrosyllis hamata (Hartman 1960) 1980,( I)(USNM 079928). Sta. 14, 9"49'25"N. (F;g. 18). 84°4I'20"W, I I m, sand, June 1981.( 1 + I (UCR 1304» ; Aug 1981 ,(2): Sta. 28. 9°52' Pi/argis hamatus Hartman 1960:88. Reish 16"N, 84°45'30"W, 26 m, mud. July 1980, 1968:76. (I)(USNM 079931). Sta. 30. 9"54'40"N, 84° Ancislrosyllis hamata. Pettibone 1966: 168. 45'50"W. 18 m, muddy sand, Apr 1981.( I). Hartman 1968:377. Blake 1997:269. Remarks. Complete specimen 44 setigers. 9.0 Ancistargis sp. Maurer et al. 1988 :48. mm long, 0. 5 mm maximum width (0.3 mm without parapodia). Emergent notopodial Aneistargis hamata (Hartman). Dean 1996a: hooks begin on set igcr 3. paired ventral eye· 74, I 996b:83. spots weak ly pigmented and often inconspi­ cuous. a weakly developed median antenna on Material Examined.G ulf of Nicoya, Sta. 12, the posterior of the prostomium. Ventral cirri 9°48'25"N, 84°52'40"W, 22 m, muddy sand, begin on setiger 3 and the dorsal cirri possess July 1980,( 1). Sla. 24, 9"49'25 n N, 84°41' numerous elongate papillae. Setae are long 20"W, I m, sand, Jan 1981 ,(1 ); Apr 198 1,(1); capillaries and shoner pectinate setae with June 1981,(2); Aug 1981 ,(3). Sta. 28, 9"52' sligh tl y falca te ends, lacking Ihe ~ indist i nct se· 60 REVISTA DE BIOLOGIA TROPICAL condary loolh" ofWolf(1984, Fig. 29, 20 G· not including parapodia). Ventral cirri and H). emergent notopodial hooks begin o n setiger 3 (third on the right and fourth on the right in Distribution. Previously re poned from the Fig. 13), an inconspicuous median antenna. intert idal to 46 meters from Florida and Texas The disti nc t fl exed anterior region of the di­ in the Gulf of Mexico. This is the fi rst record gestive trac t described by Pettibone ( 1960) is of this species in the eastern Pacific. easily seen through the integument.

Ancislrosyllisjonesi Pettibone 1966 Distributio n. Reported subtidally in 10-135 (F;g. 20). meters frOIll Chesapeake Bay. USA and the Gulf of Mexico. This ;s the first record of this Ancistrosylis jones; Salazar- Val lejo 1987 species in the eastern Pacific and its firs t re­ (1986),200. corded intertidal occurrence. Ancysfrosy/lis jonesi Pettibone 1966: 173. Le6n-Gonzalez 1991: 3 11. Dean 1996a:74. ACKNOWLEDGEMENTS Anc)'slrosyllis sp. (in part) Maurer ef o/. 1988048. The numerous collection trips to Costa Rica have been made possible through the aid and Material Examines. Gulf of Nicoya, Sla. ]4, encouragement of the University of Costa 9049'2S"N. 84"41'20"W, 11 m, sand, Oct Rica's Centro de Investigacion en Ciencias del 1980,{1 + I(UCR 11201». Sta. 28. 9052' 16"N, Mar y Limnologia (CiMAR) and its director 84"45'30"W, 26 m, mud, Ju ly 1980,( J)(USNM Jose A. Vargas. This manuscript was im­ 079932). SIO. 29. 9"54'55"N. 84"4S' I S"W, 18 proved by the review of an earlier draft by m, muddy sand, Jan 1981,(2): June 198 1.( I). Edward CUller, Department of Invertebrate Remarks. Complete specimen with 66 seli­ Zoology, Museum of Comparative Zoolvgy. gers, 15.7 mm long, 0.8 mm maximum width Harvard University and other errors corrected (0.4 mm withoUi parapodia). Body dorsoven­ by two anonymous reviewers. trally fl attened, dorsum with minute papi ll ae, notopodial ~m~rge nt hooks begin on setiger 6, ventral c irri begin on setiger 3, median an­ RESUMEN tenna inconspicuous Se infonna sobre 13 especies de Distribution. Previously descri bed from poliquetos de la familia Pilargidae (Annelida: Chesapeake Bay to Campeche, Mex ico in the Polychaeta) de ambientes submareales e in­ At lantic Ocean and from Baja California Sur, termareales en la costa Pacifica de Costa Rica. Mexico. This extends its range in the eastern Se describe la especie nueva Sigambra vargasi Pacific to the GulF of Nicoya, Costa Rica. Pre­ y se informa sobre Sigambra sp. A, au n no viously collected from the in te rt idal to 60 descrita. La ocurrencia en Costa Rica de Si­ meters. gambra lentaculata (Treadwe ll ), Loanda/ia riojai Salazar-Vallejo, Pi/argis berkeley; Ancystrosyllis hartmanae Pettibone 1966 Monro, Ancislrosyllis hamata Hartman and (F;g. 21). Ancistrosyl/is jonese; Penibone. representa ampliaciones de ambito para esas especics. La Ancystrosyllis hartmanae Pettibone 1966:172. ocurrencia de Sigambra pelliboneae Hamann­ Wolf 1984:29.24. Schroder. Parandalia Iricuspis (Muller), Cabira incerta Webster. Ancistrosyllis papil­ Material Enmined.Golro Dulce, GoJfito losa Jones and Ancistrosyllis hartmanae Petti­ Bay, 8OJ8'N. 83°10'W, intertidal 2 Km west of bone son nuevos hallazgos de estas especies main dock, muddy sand, Sept 1986; ( I + I en el Oceano Pacifico O ri ental costero de esta (UCR ). re gi6n. Synelmis cf albini (Langerh ans) tam­ bi~ n se menciona en este trabajo. Se ineluye Rem .... ks. Complete specimen 36 setigers, 8.0 una clave dicotomica para la identificaci6n de mm long, 0.7 mm maxim um width (0.5 mm las especies de Pilargidae de Costa Rica. AquRtic ecosystems o f Costa Rica 61

REFERENCES Fauvel, P. 1923. Polychtles Erranles. Fau. fran. 5: I· 488. Augencr. II. 1927. Polycllaeten von Silost und SOd­ f itzhugh. K. &. S. Wolf. 1990. Gross morphology of Australien. Videns. Medd. Dan. Natur. For. Kob. 83: 1' . 71-27S, the brain of pilargid polychaetes; Taxono mic and systcmatic implications. Am. Mus. Nov. 2992: 1-16. Banse, R. &. R. D. Bobson 1974. Bentllic errantiate poly_ chaetes of British Columbia and Washington. Bu ll. Hartman. O. 1947. Polychaetous annelids. g. Pilargiidac Fi sh. Res. Board Can. 185: I-I I I. [sic]. Allan Hancock Pacific Expedition 10: 483.523.

BerteLey, E. &. C, l3en.;eley 194 1. On a colI« tion of Bartman. O. ]968. Atl as of tile emmtiate polyehsclous I'olychaeta from southern California. Dull. South. Cal. annelids from California. Allan Hancock Foundation. Ac. Sc. 40: 16-60. University o f Southern California. Los Angeles. 828 p. Blake. J. A. 1997. Family Pilargidae Saint Joseph, 1899. In J.A. Blake, O. Hilbig &. 1'.11 . SemI (cds.). Tuo­ Hartmann-SchrOder. G. 1959. lur Okologie der Pol)'. nomic atlas of the benthic fauna o f the SMta Maria chaeten des Mangrove Estero Gebe ites von EJ Salva. dor. De it. Neo t. Fau. I: 69-183. ba~ln and western Santa Barbara channel, Volume 4 The Annelida Part I. 26 1-284. Sant. Darb. Mus. Nat. Hi st. Sama Darbara, California. Hartmann·SchrOder, G. 1979. lur Kenntnis des Eulitorals der australisehcn KOsten unter besonder OcrUcksi­ ChamtJerli n. R. V. 1919. The Annelida Polychacta. Mem. chtigung de r Polychaeten und Ostracoden, 2. Die Mus. Compo lool. Harv. Coil. 49: 1-5 14. Poly chaeten der tropishen Nordwestkllste (zwishen Derby im Norden und Pt. Hedland im SlIden). Mit. Hamb. Zool. Mus. lnst. 76: 77-218. Day. J. II. 1967. A mooograph on the Polychaetes o f Southern Africa. Part I: Errantia; Part 2: Sedentaril. Bri t. Mus. Publ. 656: 1·878. Imajima. M. 1987. Pilargidae (Annelida, I'olychaeta) from Japan (Part 1). Bull. Nat. Sci. Mus. SC I. De:lIl . II. K. 1996a. Subtidal benthic polychaetes (Zoology) 13: 151- 164. (Annelida) of the Gul f of Nicoya. Costa Rica. Rev. Iliol. Trop. 44 (suppl. 3): 69,80. Ka~man. W.. L. Laubier. &. J. Ramos 1974. Pil argidae (Annclides polychctes ~ rrante s) de MediterranCe. Denn. II. K. 1996b. Polychaete worms (Annelida) col­ Bulletin d'lnsitute Occanographique Monaco 71 : 140. lected in Golfo Dulce during the Victor lIensen Co~ta Ri ca expedition (1993f l 994). Rev . Ri ol. Trop. 44 Langerhans. P. 1881. Ober elnige eanarische Anneliden . (suppl. 3): 81-86. Deutsche Akademie der Naturforscher Nova Acta 42: 93-124. Dean. H. K. 1995 A new species o f Hesionidae. GJJ'ph~llotlf! lIico}'e'lJiJ (A nnelida: I'olychaeta) Le6n-GonzAlez, J. A. 1991 . Poliquetos de fondos blandos from the Gulf of Nicoya. Costa Rica. Dull. Bioi. Soc. de la costa occidental de Baja California Sur, M6cico. Wash. I I I; 257-262. I. Pilargidae. Cab. Bioi. Mar. 32: 3 11-321_ Ehlers, E. 1908. Die bodensllssigen Anneliden aus den Sammlungen der Deutschen Tiefsee· Expcdition. Licher. F. 1994. Resurrection of G!yp"~sione Fr ie. Wi ss. Ergeb. Deut. Tief Exp. 1897-1899 16: 1. 168. drich. 1950. wilh redescription of G. Haui Friedrich, 1950 and description of G. iongocirrala (Polychpeta: Emerson, R. R. &. K. Fauchald. 197 1. A revision of the Hesionidae). Proc. BioI. Soc. Wash. 107: 600-608. genus Loanda/ia Monro with description of a new ge­ nus and species of pilagiid polychaete. Bull. Sout. Licher, F. &. W. Westhe ide. 1994. The phylogeneti c Cnl. Acad. Sci. 70: 18-22. position o f the Pilargidae with a cladistic analysis of the taxon-faets and ideas. In J. C. Dauvin. L. Laubier, Fauchal d. F. 1977. Polychaetes from intertidal lll"Cas in & D. J. Re ish. (eds). Actes de la 4 me Conftn:ncc in· Pan.ama. with a ~view of p~vioU1 shallo w water reI;,­ ternational des Polyehaetes. Mem. Mus. Nat. Uist. ords. Smith. Contrib. lool. 221: 1-85. Nat. 162: 223·236.

Fauchald, K. a: P. Jumars 1979. The d iet o f worms: A Licher. F. &. w. Weslheide. 1997. Review o f the genus stud), o f polychaete f~ding guilds. Oceanog. Mar. Sigamb,a (PolychllC tl: lic:sionidae). redescription o f S. BioI. Ann. Rev. 17: 193-284. haul (Hartman 1947), and descriptions of two new species from Thailand and China. Steenstrupi a 23 : I· Faucllald, K. &. G. Rouse. 1997. Polychaete systematics: 20. Past and present. Zool. Scrip. 26: 71-138. Maurer, D. &. J. A. Vargas. 1984. Diversity of soft bonom Fauve l, P. 1920. Les genres Ancis/,osylfis et Pifa'giJ benthos in a tropical estuary: Gulf of Nicoya. Costa (Hesionidac) Bull. Soc. Zool. Fran. 45: 205-213. Rica. Mar. Bioi. 81 : 97·106. 62 REVIST A DE BIOl.OGIA TROPICAL

Maurer, D .. J. A. Vargas & H. Dean. 1988. Polychactous 1858 from Brazil and designation of neotypes (Poly­ annelids from the Gulf of Nicoya, Costa Rica. Inter. chaeta: Pi[argidae). Jour. Nat. Hi st. 24: 507-5 I 7. rev. ges. Hydrobiologie 73: 43-59. Salazar-Vallejo, S. I., 1. A. de Le6n-Gonzillel & J. C. Mclntosh, W. C. 1879. On the Annelida obtained during ChAvez-Companin. 1990. Poliquetos (Annelida: Poly­ the cruise of H.M,S. 'Valorous' to Davis Strait in chaeta) de La Bahfa de Manlanillo, con una clave 1875. Tran. Linn. Soc. 2: 499-511. ilustrada para las especies de Colima, Mexico. Rev. BioI. Trop. 38: 21 1-229. Monro, CA. 1933. On a new species of Polychac:ta of the genus Pi largis from Friday Harbour, Washington. Salazar-Vallejo, S. I. & J. M. Qrensanl 199 1. Pih\rgidos Ann. Mag. Nat. lIisl. 10: 673-675. (Annelida: Polychaeta) de Uruguay y Argentina. Cah. BioI. Mar. 32: 267-279. MOiler, F. IS58. Einiges Ubcr die Anne li den Fauna der In sel SI. Catherina an der Brazilianischen KlIste. Treadwell. A. L. 1941. Polychactous annelids from the Arch. Natur. (Berli n) 24: 211-220. New England region. Puerto Rico and Bralil. Am. Mus. Nov. 1138: 1-4. Pettibone, M. H. 1966. Revision of the Pilargidae (Annelida: Polychaeta), including descriptions of new Vargas, J. A. 1995. The Gu[f of Nicoya estuary. Costa species, and n:descriptions of the pe lagic Podarmus Rica: past, present, and future cooperative research. ploa Chamberlin (Polynoidac). Proc. Nat Mus. 118: Helgol. wiss. Meeres. 49: 821-828. 15S-20~ . Webster, H. E. 1879 . Annelida chaetopoda of the Vir­ ginian W bSt. Trans. Alban. [nst. 9: 202-269. Rei sh, D, J. 1968. A biological survey of Bahia de Los Angeles, Gu[f ofCa[ifornia, Me.xico. Il. Benthic poly­ Westheide, W, l!j , ' Interstitielle Fauna von Galapagos chaelous annelids. Tran. San Diego Soc. Nat. Hist. IS: Xl. Pisionidae. Ii< Jionidae, Pi largidac:, Syllidae 67-106. (Polych3eta). Mikrof Mem:sh. 44: 1-146.

Salazar-Vallejo, S. I. \987(1986). Pi largidae (Annelida: Wo lf, P. S. 1984. Chapter 29. F~mily Pilargidae Saint­ Po[ychaeta) de Mexico: Lista de especies. nueva es­ Joseph 1899. p. 29-1 to 29-41. In J. M. Uebclacker & pecies y biogralia. Cah. BioI. Mar. 27: 193-209. p, G. Johnson (cds). T..:..onomic guide to the poly­ chHetes of the northern Gu lf of Mexico, 7(4). Final re ­ Salazar-Vallejo, S. I. [990. Redescriptions of Sigambra port to the Minerals Management Service, contract grubii MOller 1858 and Hermundura Iricuspis MUller 14-\2-001-29091. Barry A. Vittor, Mobile. Alabama.