06 dean pilargidae (1)

8/13/2019 06 Dean Pilargidae (1)

1/16

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

The Pilargidae (Annelida: Polychaeta) of thePacific Coast of Costa Rica

Harlan K. DeanDepartment of e . r t e b m t e Zoology, Museum of o m p a m t i ~ e Zoology, 26 Oxford St., C3mbridge. MA02138 USA, and John Hazen White School of Arts and Sciences, Johnson and Wales U n i ~ e r s i 6 AbbottParle Plllce, r o ~ i d e n Rhode Island 02903, USA. E-m ail: hdc3n@ici .met

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

Abstract: Thirteen species of Pilargidae (Annelida: Polyehaeta) from the subtidal and inlerth.lal Pacific Coast ofCosta 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 Salaza rVallejo Pilargls berkeleyl Monro, rrclJlrosyl//s nama/a H3rtman and AneWrosyllis jonesei Peltibone are rangeextensions for these species. The occurrence of Sigambra pellibaneae Hartmann-SchrOder, Pilrandillia Iricuspi(MUlier), Cubira in rtu Webster, Ancis/rosyllis papillosa Jones and Ancislrosyllis narlmanae Pettibone are newre cords of these species for the eastern Pacific Ocean. Synelmls cf. albini (Langcrhans) is also reported. A taxonomic key to these species is included.Key words: Polychaeta, Pilargidae, Pacific coast, Costa Rica.

The Pilargidae are a family of polychaetesreported in sediments from the intertidal toabyssal depths. Since these organisms are usually quite small and present in low densitiesthere is very little information regarding theirecology (Faucha1d Jumars 1979). They donot build a tube and have a muscular eversiblepharynx which is presumed to be used in thecapture of prey or perhaps intake of detritusand microa1gae (Day 1967).The systematics of the Pilargidae is alsopoorly known. Historically, species have beenassigned to the Orbiniidae (MUlier 1858), theSyllidae (Mcintosh 1879. Webster 1879), andquite often the Hesionidae (Ehlers 1908;Fauvel 1920. 1923, Augener 1927, Monro1933, Berkeley Berkeley 1941 . and Treadwell 1941). Eventually Hartman (1947) andPettibone (1966) united a disparate group ofgenera into the family Pilargidae characterized

mainly by the presence of emergent notopodial spines or hooks, two pairs of tentacularcirri and entirely simple neurose tae. Morerecent results of cladistic analysis by Licherand Westheide (1994) have indicated that thePilargidae might be more appropriately included within the family He sionidae and, in theirreview of the genus Sigambra (LicherWestheide 1997), regarded the Pilargidae as asubtaxon of the Hesionidae. Further studymay support this view, however, until suchtime the family Pilargidae is herein retained(see Fauchald Rou se 1997).Relationships of the genera within the Pilargidae have always been difficult due to thewide range of character states. Salazar.vallejo(1987(1986 divided the Pilargidae into tw osubfamilies, the Synelminae with cyl indricalbody and emergent notopodial spines, and thePilarginae ( 'Sigambrinae) with a dorsoven-


2/16

48 REVISTA DE ()fOLOG1A TROPICAL

trally depressed body and either emergentnOlopodial hooks or emergent nOlopodial setaelacking. These subfamilies are nol supportedby the results of cladistic analysis based ongross brain morphology by Fitzhugh and Wolf1990) or the analysis of Licher and Westheide J 994) based on 28 morphologicalcharacters. Licher and Westheide 1994) identified several clades or groups of genera characterized by such characters as presence of apapillated integument or gross morphology ofthe hind brain and indicated that the Pilargidaemay not be a monophyletic group.Despite the confusion regarding phylogenetic relationships, the species within the Pilargidae have been well characterized. espec iall y in the easte rn Pacific Ocean. Two species have been recorded from the region between Alaska and Washington State USA) byMonro 1933) and Banse Hobson 1974).The pilargids of the California coast USA)have been well sampled and nine species recorded by Hartman 1947, 1968) and EmersonFauchald 1971). Salazar-Vallejo 19871986), 1990) and Le6n-Gonzalez 1991) havesampled extensively along the Pacific coast ofMexico and published several keys and species lists which include 18 species from thisregion. Hartmann-SchrOder 1959) recordedtwo species of Pilargidae from the mangrovesof EI Salvador. Synelmis albini Langerhans)was also included in collections by Westheide1974) from the Galapagos Islands andFauchald 1977) from the Pacific side of Panama. In prelimi nary lists of polychaetes fromthe Gulf of Nicoya and Golfo Dulce, CostaRica, Dean 1996a,b) reported 10 species belonging to the family Pilargidae.This paper rev iews 13 species of Pilargidaecollected from the Paci fic Coast of Costa Ri ca.Four species are added to the lists of Dean1996a, b) while specimens originally identified as Sigambra bassi are herein described asa new species, S. vargasi n. sp. Another species included in the list of Pilargidae by Dean( I 996a), Synelmis Ielalli Friedrich, has beenmoved to the fami ly Hesionidae and redescribed by Licher 1994). Specimens identifiedas S. /dalli from Costa Rica Dean I 996a)have been descri bed as a new species ofGlyphohesione in the fam ily HesionidaeDean 1998).

MATERIALS AND METHODSSubtidal material from the Gu lf of Nicoya wascollected during a cooperative research effortbetween the University of Costa Rica and theUniversity of Delaware USAXsee MaurerVargas (\ 984), and Vargas 1995) for locations and review). Intertidal material from thePunta Morales region of the Gulf of Nicoyawas collected by the author during a series ofcollecting trips supported by the University ofCosta Rica s Centro de Investigaci6n en Ciencias del Mar y Limnologia CIMAR). Subtidalmaterial from Golfo Dulce was collected bythe R V Victor Hensen Costa Rican expedition 1993 / 1994) and material from thatcruise provided by J. Vargas and R. Le6nC IMAR). Specimens from the intertidal ofGolfito Ba y, Golfo Dulce were collected by J.Vargas CIMAR). Voucher spec imens of mostof the listed species have been deposi ted in theMuseo de Zoologia, Universidad de CostaRica. The remainder of the material is deposited in the Museum of Comparative Zoology,Harvard University, Camhridge, MA , USA.

Key 10 the Pilarg id ae o f Ih e Pac ificcoast of Cosla RicaI . Integument smooth............. .. ... 2I a. In tegument papillate .........................82. Antennae and tentacu lar cirri present ... ... .32a. Antennae and tentacu lar cirri absent .. . .. . .73. Three weakly developed antennae, furcateneuropodial spines present. Fig. 1 ......... .. .. ........ Syne/mis f a/bini Langerhans)3a. Three well developed antennae present,dorsal parapodial cirri of setiger I longerthat those ofother setigers................... .44. Emergent notopodial hooks from setiger 4.Fig. 2) ... Sigambra lenlaeu/ala Treadwell)4a. Emergent notopodial hooks begin posterior to setiger 4 ................................. 55. Emergent notopodial hooks from setigers8-9. Fig. 3) ...... ..... sigambrapetfiboneae Ha rtmann-SchrMerSa. Emergent notopodial hooks begin poste-rior to seliger 10 6


3/16

Aql,latie ~ O S y S l c l l 1 of Costa i ~ a 49

6. Emergent notopodial hooks from set iger 12 .(Fi g 4). . . . . . . . . . . . . . . . ... .. ...... sigambra sp A6a . Emergent notopodial hoo ks from setige rs14I S. ( Fig. SI2 ..sigambra vargasi n. sp.7. Ventro lateral branchiae present. (Fig. 13) ...

...... ... .... .Loandalia riojai Salazar Vallejo7a. Ventrolateral branchiae absent. ( Fig. 14) ...

.................Parondolio Iricwpis (MUller)8. Parapodia weakly developed. probosciswith dentic1es. (Fig. IS 16) .............. .

..................... . ..Cabiro incerla Webster8a . Pharynx not cuticularized. parapodia well

deve loped ............... . ........... . ...... .. ...99. Emergenl nOlopodial spines or hooks absent(Fi g. 17) . . .. ..... .. Pi/argis berkeleyi Monro9a. Emergent dorsal hooks present. GenusAncislrosyllis ................................. 1010. Two antennae present ...................... 11103. Three antennae present ................... 12I I. Emergenl notopodial hooks from seti

gers 47, dorsal parapodial cirri withsma ll papillae. (Fig. 18).. .... ... .... .. . .... .... . . ..... .. . ...Ancistrosyllis hamata Hartman11 3. Eme rgenl notopodial hooks from seti ger3. dorsal parapodial cirri with prominentelonga ted papillae. (Fig. 19) .................. .

....... ... .Ancistrosyllis papillosa Jones12. Emergent notopodial hooks from set iger 6.

(F ig. 20) ..... Ancistrosyllisjonesi Pettibone12a. Emergent notopodial hooks from set iger 3, flexed anterior region of the digestive tract. (Fig. 2 1) ... . ........ ..... ......... ....... .. Ancistrosyllis harlmonae Pettibone

Species DescriptionsFami ly Pilarg idaeGenu s Synelmis Chamberlin 1919Fitzhugh Wolf (1990) separated this genusin to complex A species with two furcateneurosetae and complex B species lackingfurcate neurosetae. Based upon their cladisticanalysis Licher Westheide (1994) united thecomplex B species with th e genus Litacorsa

leaving on ly 2 species S. albini (Langemans)and S. simplex Chamberlin in the genus withthe synapomorphy of two furcate neurosetae.Synelmis cf albini ( Langemans 1881 )(F;g. I)Synelm is albini (Langemans). Pett ibone 1996:191 . Fauchald 1977 :17, Westheide 1974. Sala-zarNallejo 1987 ( 1986): 20S. Mau rer Var-gas 1984: 10 1 ( in part), Maurer et a/. 1988 :48(in part), Dean 1996a:74 .Materillli Exa mined.G ulf of Nicoya, Sla. J9 S7')0 N , 84 S3'OO W , 46 m, mud. Jul y1980,(IS ) and (I XUSNM 079952). Sla . 2, 955'28 N, 84S2'OSW. 18 m, muddy sand, Ju ly1980,(2XUSNM 079973). Sla. 3, 9 S2'00 N,8448'10 W, 33 m, sandy mud, July 1980, ( I)(USNM 0799S3). Sio. 4, 9S 3'40N, 8446'10 W, 40 m. mUd . Jul y 1980,(1 ) ( USN M0799 S4) . Sta. 13. 9 S2'30 N. 84 43'SO W , 26m, July 1980.(3)(USNM 0799 5S 0799S6) .Sra . 14 , 9 57'05 N, 9 m, sandymud , Jul y 1980,(2S)( USNM 079966 079967). Sta. 15. 9S7'40 N, 84 47'OOW, IS m,sandy mud, July 1980, (7I XUSNM 0799740799S5). Sta. 13 , 9 48'3S N, 8443'SO W, 35m, mud, Jul y 1980,(1) (USNM 079959). Slo.24 , 949'2S N. 84 4I'20W. 11 m. sand, Oct1980,(27); Jan 1981 ,( I) ; Apr 1981 ,( 11 ); Aug1981,7). Sta. 25, 9 S OOSN, 84S2 'OO W , 20m, mud/sand, July 1980,(J XUSNM 079960).Sta. 26, 9Sl'SON, 84 S3'20 W, 17 m, muddysand, Jul y 1980,(I)(USNM 079961 ). Sla. 279 S I'57 N, 84 50'S OW, 12 m, muddy sand,July 1980,(2) (USNM 079963 079963). Sra.28, 9S2' 16N, 84 4S'30W, 26 m, mud, June1981, 1). Sta. 29, 9 S4'S5 N , 84 45' 15W, 18m, muddy sand, Oct 1980,( 18); Jan 1981 ,(32);Apr 198 1, (4) ; June 1981 , (4); Aug 198 1. (20) ;Apr 1982,(22). Sta. 30, 9 S4'40 N, 844S'SO W , 18 m, muddy sand, Oct 1980,(11); Jan1981 ,(6); Apr 198 1,(SI ); June 1981,(4). Sla.31, 944' N, 84 59'2SW. 20 m, mud/sand,July 1980,(5 1XUSNM 079947, 079948, 079950 07996S). Sla. 32, 9 S 3'47 N, 84 49'3S W, 24 m. mud/sand, July 1980,(2).(USNM079971). S . 34, 9 SS'30N, 84S 0'OS W , 24m, sand, Ju ly 1980,( 12)(USNM 0799490799S I). Sta. 35, 9 SS'42 N, 8447'40 W, 13m, sand, July 1980,(32)(USNM 079976 079977). Sla . 3 7. 9 S7'38N, 8448'20 W, 14 m,


4/16

so REV1STA DE 1lI0LOGIA TROPIC L

muddy sand , July 1980, (2 1) (USN M 079978079979). Punta Morales, IO N, 8S oW, in-tenidal, coarse sand, Au g 1996;(12, UC R109).Rem rks. Complete specimen 136 seligers,approx imately 50 mm long and 0.7 mm wide.

1

.. ,

With two pi gmented cyespots and a cyl indrica l pharynx (Fig. 1. Emergent nOlopodialspines beginning at seliger 11 accom panied byone or two thickened notacicu la. Capi llaryncurosetac include one or IWO longer and twoor three shoner setae as we ll as one or twofurcate setae in a sing le fascicle .

2

4

Figs. 1-4. Fig. I. Synelm;J f albmi. dorsal view of anterior Fig. 2.S'ga ,brn Itnlaf;fl/U/(l. dorsal view of anterior. Fig. 3.Sigamhrapelliboneat , dorsal vicw of nterior. Fig . 4.sigambra sp . A. dorsal view of ntcrior. Scale bar ] mrn .


5/16

AQLlalic ecosystems of C la Rica

Dis tr i butio n. One of the most common po lychaetes from the Punta Morales intertidal froma wide range of sediments. What has beenident ified as Synelmis afbini (Langerhans) hasbeen reported from the Galapagos IslandsWe stheide 1974) and th e Pac ific side of Pan

ama (Fauchal d 1977). It is be lieved, however,that S. a/bini may be a complex of seve ralspecies and awaits further system atic analys isWo lf 1984).

Genus Sigambra MO ller 1858Members of this gen us possess we ll devel

oped lateral and median antennae, tentacularcirr i, and dorsal parapodial cirri . Lieher &Westheide ( 1994) regard all th ese features aspleisio'morphi c indica tin g a high similarity betwee n Ih is genus and ju veni le hesionids.Sigambra lelllacli/afa Treadwe ll 194 I)(Fig. 2Sigambra fen/aclilata Treadwel l 1941 : I. Pe ttibone 1966: 182.Maurer & Vargas 19 84 : 101 (in part). Wolf1984 :29.8 .Sa lazar-Vallejo 1987(1986):202. Maurer el al.1988:48 (in part). Salazar-Vallejo et al.1990 :2 16. Le6n-Gonzalez 199 1:3 14 . Sa lazarVallejo & Orensanz 1991 :275. Bl ake 1997:275. Dean 1996a:74; 1996b:83.Material Exam in ed. G ulf of Nicoya , Sla. 69 4S'20N, 8446'25 W, 42 m, mud, Jul y1980,( IXUSN M 079936). Sla. 22, 948'25 N,84 S2'40 W, 22 m, muddy sand, July1980,( IXUSN M 079937). Slo. 24 , 949'25 N.844 I'20 W, II m, sand, Oct 1980,( 1) ; Apr198 1.( 1) ; Jun 198 1,( 1); Aug 198 1,( 1) . Sta. 29954'55 N, 8445' 15 W, 18 01 , muddy sand,Jan 198 1,(2). S la. 30 9 54'40 N. 8445'50 W,18 m, muddy sand, Jan 198 1,(3). Sla. 31953'47 N, 8449'35 W, 24 m, mud/sand, July1980,( IXUSNM 079944). S/a. 33, 9 53'40 N,84 53'20 W, m, sandy mud , Ju ly 1980,IXUSNM 079942). Punta Morales 10 N,85W, intertidal, Aug 1996:( 11 ), mud; (8),sandy mud ; II + 3(UCR 1100I ,coarse sand.

Go lfo Dulce, Sla. I2 8 2I'N, 83 14'W,200 m, mud, Dec 1993 , ( I).Remarks. Complete spec im en 57 setigers, 7.3

mm long, maximum width 0.8 mm (0.6 notincluding parapodia). NOlOpodial emergenthooks from setiger 4, ventral cirrus lacking onseti ger 2, median antenna longer than latera lantennae, tip s of neurosetae entire, notosetaeabsent. These speci mens lack the row ofperistomial papillae described by Katzmann era/. (1974) for materia l iden tified as S. lenla-culala from th e Mediterranean.Dis tribution. I)reviously reported from NewYork , USA, to Argentina in th e At lanticOcean and from California, USA, to Man zani- 0, Col. , Mex ico, in the Pacific. The materialid entified as S. lenlaculala by Katzmann el 01.1974 from the Mediterranean may be a different species. This extends the range of thi s spec ies in th e Eastern Pacific to Golfo Dulce,Costa Rica. Sigambra lenlac lliara has beenreported from the intertidal to abyssal depths.Sigambra pellibolleae Hartmann- SchrOde r1979 ( Fig. 3)Sigambra pellibolleae Hanmann-SchrMer1979:84.Mate rial Exa min ed, Golfo Dulce: GolfitoBay g038 'N, 83 10'W, in tertida l, muddy sand2 Km west of main dock, Sep 1986 ; ( 109+7(UC R 108).Rema rks. Complete spec im en 59 se tige rs, 5. 8mm long, 0.7 mm max imum width (0 .5 mmwi th out parapodia). Emergent notopod ialhooks beginning at seligers 8-9, ventral cirriabsent on set iger 2, and th e median antennaex tends to setigers 4-6. The neurosetae includeshort coarsely toothed and longer capi llaryseta e.Distribution. Type locality is intenidal sandwith organic detritus from Broo me in northwestern Australia. This is th e first record ofthi s species from th e eastern Pacific and th efirst report of this spec ies ou tside of the typelocality.Sigambra sp. A(Fig. 4)Material Exa min ed, Go lfo Dulce, ColfitoBay 838'N, 83 IO'W, intertidal, muddy sand2 Km west of main dock, Sep 1986;( 1).


6/16

REVISTA DE BI OLOGIA TROPICALRemarks. Complete specimen with 48 setigers, 3.9 mm in length and 0.5 mm in widthanteriorly (0.35 mm without parapodia). Bodywide anteriorly, tapering posteriorly; dorsoventrally flattened with deeply incisedparapodia. Color in alcohol light tan.Prostomium with indistinctly separated biarticulated palps; palpophores large, anterovenral palpostyles small, digitate. Median antenna arising from posterior of prostomium,extending to fifth seliger; lateral antennaeari se slightly anterior to median antenna, extend to the first seliger. Proboscis with 14conical distal papillae. Two pair subequaJtentacular cirri from peristomium, cirri slightlylonger than lateral antennae. Dorsal cirri ofseliger 1 elongate, extending to the fifth setiger. Ventral cirri absent from setiger 2.

Notopodia with a single acicula, accompanied by a stout emergent notopodial hook beginning at setiger 12, a capillary seta may alsooccur in posterior notopodia. Neuropodiallobe conical with numerous capillaries, shortercapillaries with fine denticulations, and shortpectinate setae. Dorsal cirri extend beyondsetal lobe; ventral cirri subequal in length tosetal lobe. Pygidium with pair of long analcirri.This spec ies most closely resembles a recently described species Sigambra phulcetensisLicher & Westheide 1997 in that they bothhave ventral cirri on the second setiger, 14distal papillae on the pharynx lack emergent

spines, have a longer median than lateral antennae and possess pectinate neurosetae.While S. phulcetensis is described as havingemergent notopodial hooks beginning at seliger 3-23, S sp A has notopodial hooks fromsetiger 12. The median antenna of S. sp A ismuch longer than the lateral antennae extending posteriorly to the 5th setiger (4-5 times thelength of the laterals) while the lateral antennae are shorter relative to the body width thanthose reported for S. phuketensis. Additionally, the dorsal cirri of S puketensis are approximately 2/3 the body width in median andposterior setigers while those of S sp A aremuch shorter than that. It would be prematureto formally describe this species based upon asingle specimen especially in the light of the

disregard by Licher & Westheide (1997) forthe significance of the first appearance of notopodial hooks in their review of the genus.Sigambra vargasi n. sp.(F;g.5 12)

aterial Examined. ulf of Nicoya, Sta. I957'30"N, 8453'OO W, 46 m, mud, July.l980,(2XUSNM 079935). Sta. IS, 9"57'40"N,8447'00"W, 15 m, sandy mud, July 1980,(lXUSNM 079946). Sta. 23, 948'35"N, 8443'50"W, 35 m, mud, July 1980,(IXUSNM079938 . Sta. 24, 949'25"N, 844 1'20"W, IIm, sand, Oct 1980,(1) ; Jan 1981,(5) ; Apr1981, (5); June 1981,(10); Aug 1981,(16). Sta.25, 950'05"N, 8452'00"W, 25m, mud/sand,July 1980, (4)(USNM 079939). Sta. 26, 951'50"N, 8453'20"W, 17 m, sandy mud, July1980,(IXUSNM 079940). Sta. 27, 951'57"N,845 0'50"W, 12 m, muddy sand, July 1980,I) (USNM 079941). Sla. 28, 952'16"N, 8445'30"W, 26 m, mud, July 1980,(18); Oct1980, (18) (MCZ 20019); Jan 1981,(4) ; Apr1981 , (1 + 5(UCR 1602 ; June 1981 ,(12);Aug 1981,(17). Sta. 29, 954'55"N, 8445'5"W, 18 m, muddy sand, Jan 1981,(1); Aug1981,(21); Apr 1982,(6). Sla. 30 954'40"N,8445'50"W, 18 m, muddy sand, July 1980,(1)(USNM 079943); Oct 1980,(2); Jan 1981,(6); Apr 1981,(6); June 1981,(4); Aug 1981,(I+I(HOLOTYPE MCZ 20017)+2 (PARATYPES M l 20018). Sta. 32, 953'47"N ,8449'35"W, 24 m, mud/sand, July 1980,(6)(USNM 079944). Sta. 36, 955'32"N, 8445'20"W. 18 m, sandy mud, July 1980,(2) (US

NM 079945).Species description. Holotype (MCZ 20017)5.2 mm long, 1.0 mm wide (0.5 mm not including parapodia) with 60 setigers. Paratype(UCR 160-2) 9.7 mm long, 0.8 mm wide (0.5mm not including parapodia), with 54 setigers.Paratype MCZ (20019) 18.4 mm long, 0.5 mmwide (0.3 mm not including parapodia), with136 setigers. Body white to light brown inethanol. First four segments cylindrical, 3times as wide as long, middle and posteriorsetigers dorsoventrally flattened, deeply incised, 4 times as wide as long (Fig. 5).Prostomium two rounded lobes; two biarticulate palps indistinctly separate from theprostomium, palpophores large with small


7/16

Aquatic ecosystems of Costa Rica

anteroventral palpostyles (Fig. 5). Medianantenna extending to seven th setiger (x = 6.9,SD = 1.29, n = 10; paired lateral antennaeattached slightly anterior to and approximately1/3 the length of median antenna. Eversibleproboscis with 8 distal papillae. Peristomiumwith 2 pair tentacular cirri, dorsal slightlylonger than ventral, 113 length of lateral antennae.First setiger with elongate dorsal cirrus 4-5set igers in length (F ig. 9). Setiger 2 with shortdorsal cinus, ventral cirrus lacking (Fig. 10 .Setigers 3-5 with subequal dorsal and ventralcirri extending beyond the parapodial lobe(Fig. II . Remain ing setigers with dorsal cirrus approximately twice as long as ventralcinus, ventral cinus subequal in length toparapodiallobes (Fig. 12).Notopodia reduced, notacicula present at baseof dorsal cirrus, accompanied by a short capillary notoseta beginning at setiger 4 . Emergentnotopodial hooks from setiger 15-1 7 (x = 15 .2,SD = 0.48, n = 31), accompanied by a singlenotacicula and occasionally one or two shortcapillary setae. In median and posterior setigers notopodial hooks may be replaced by ashort capillary seta.Neuropodia with fascicle of maximally 17-18simple neurosetae. Neurosetae include longerserrate bladelike setae (Fig. 6), shorter pectinate se tae with smooth dista l tip (Fig. 7), andshort pectinate setae with fine teeth extendingto distal tip (Fig. 8); neuropodia of posteriorsetigers with one or two smooth capillary setae(F;g. 12).Pygidium with two long, thin, cirriform analcirri.Remarks. Sigambra vargasi n. sp. is similarto S. oce/Iata (Hartmann-Sc hroder 1959), S.bidentata Britaev Saphronova 1981, and S.q;ngdaoensis Li cher Westheide 1997 in thepossession of 8 distal papillae on the pahrynxand the absence ofa ventral c irris on seliger 2.It is unlike S. ocellata in the absence of eyespots and the greater length of the dorsal relative to the ventral cirris. S. vargas; n. sp. lacksthe bidentate neurosetae seen in S. bidentata.S. vargasi n. sp. is similar to S. qindaoensis in

that these are the only two species in the genusw ith an terior setigers possessing a capillarynotoseta unaccompanied by emergent hooks.S. vargas; n. sp. differs from S. qingdaoensisin the first occurrence of notopod ial hooks atseliger 15-1 7 rather than setigers 3-8, themuch greater length of the median antennaand the greater length of the ana l cirri.Distribution . From to 46 m in muddy tosa ndy sed im en ts.Etimology. This species is named for Prof.Jose A. Vargas (CIMAR) for h is many yearsof work furthering benthic studies in CostaRica.Loandalia Ptuandalia .{ TalellSapia GroupThis group of pilargids, identified by thephylogenetic analysis of Licher Westheide(1994), is characterized by the absence of antennae, tentacular cirri, and dorsal parapod ia lcirri, the possession of cap illary notosetae inaddition to the em ergent spine, and a lack ofnotacicula. Members of this group are included within the Pilargidae based mainly on thepresence of emergent notopodial spines. Thesegenera may be considered highly derived orpossibly be unrelated to the other pilargids(Licher Westheide 1994).Ge nus Loandalia Monro 1936This is the on ly genus in the Pilargidae withbranchiae.Loandalia rioja; SalazarVal lejo 1987( 1986)(Fig. 13)Loandalia rioja; Salazar-Vallejo 1987(1986):202. Salazar-Vallejo et . 1990:216. Le6nGonzalez, 1991 :3 14Material Examined. Gulf of Nicoya, Sta.29,954'55'N, 8445'15'W, 18 m, muddy sand,Apr 1982 ( I, UCR 2003).Remarks. Complete specimen 65 setigers, 40mm long, maximum width 1.6 mm ( 1.1 mmwithout parapodia). First occurrence of emergent notopodial spines is on se tiger 7, dorsolateral fields of proboscoid organs present.


8/16

REVISTA DE BIOLOGIA TROPICAL

Pigmented eyespots were not observed for thisspecimen. Salazar.vaJlejo (1987(1986 reported branchiae beginning on setigers 21 to24 and branchiae first appeared on setiger 24in this specimen_ Le6n-GonzAlez (1991) reported branchiae from setiger 31-39 in twospecimens of L rioja; from Baja CaliforniaSur.

Distribution. Previously reported from BahIade Manzanillo, Mexico, to the west coast ofBaja California. This record extends the rangeof L rioja; to the Gulf of Nicoya, Costa Rica.Subtidal from 18 to 140 meters.Genus Parandalia Emerson Fauchald 1971

6

,,I

7 8

Figs. "j.. . Fig. 5. S i g a m ~ WlTgaJi n sp. (Ho]otype MCZ 20011). Dorsal view of anterior. Figs. 6 8_Slgonrbra vargruip ~ p e MCZ 2 O O ~ Fig. 6. Long se ""'te ~ u r o s e t a Fig: 1 Shon pectinate neuroseta with smooth distal tip. FigS. Shonpectinate nc:urosetawlth finely tootheddLStal tip. Scale bar Fig. 5" 100 /1m Figs.6-S" 20/lm..


9/16

Aquatic rcruystcms of Costa Rica

This genus resembles Loandolio but lacksbranchiae. The other genus in this subgroup,Talehsopia, differs from Porondalio in thepossession of aw ike structures.Parondalio tricuspis (MUller 185 8)(Fig. 14)Parandalia t r j c u s p i ~ (MUller) Salazar-Vallejo1990:5 12, Salazar-Vallejo & Orensanz 1991 :268, ean 1996a:74.Porondalia sp. A. Wolf 1984:29.30.Parondalio sp. Maurer et al. 1988:48Material Examined .Gulf of Nicoya, S10 23,948'35 N,. 8443'50 W, 35 m, mud, July1980,(1) (USNM 079934). Sta. 24, 949'25 N,8441 '20 W, m, sand. Jan 1980,(1XUSNM080033); Oct 1980, (3 + I , VCR 2302); June1981,( I)(USNM 080032); Aug 198 1,( I).Remarks . Complete specimen 48 seligers, 6.4mm \ong . anterior 7 setigers 0.8 mm wide,remaining setigers maximum width 0.7 mm(0.5 without parapodia). These specimensagree with the new combination by SalazarVallejo (\990). Ventral parapodial cirri firstappear on setiger four and emergent notopodial spines lacking a hood on seliger 7. Eyespots were weak ly pigmented, one spec imenwith only a single eyespot on the right side.Posterior neurosetae in two series with approximately 10 neurosetae per parapodia. A llparapodia biramous.Distribution. Previously known from the upper Gulf of Mexico to Argentina in the Atlantic Ocean. This is the first record of this species in the eastern Pacific. P Iricuspis hasbeen collected from the intertidal to 55 metersdepth.Anclstrosyllls groupThe genera Arn::istrosyllis, Pilorgis, Cabira,and Paracobiro are considered a monophyletic group by Licher & Westheide (1994) dueto the papillated integument.Genus Cobiro Webster 1879

The presence of dentides on the pharynx aresynapomorphic for this genus.Cabira incerta Webster 1879Flg.(16 17).Cabil a incerta Webster 1879:267. Pettibone1966:178. Wolf 1984:29.5. Salazar-Vallejo &Orensanz 1991 :272.AncistrosylJis sp. (in pan) Maurer et al.198 8A8Cabiro sp. A. Dean 1996a:74.Materia l Examined. Gu lf of Nicoya, Sla. 24,949'25 N , 8441'20 W, m, sand, Apr1981 ,(2); Aug 1981 ,(1 + I , UCR 6802)). Slo.28. 952'J6 N, 8445'30 W, 26 m, mud, Ju ly1980, ( 1)(USNM 079930); Aug 1981,(1). Sto.30, 9 54'40 N, 8445'50 W, 18 m, muddysand, Apr 1981,(3). Punta Morales, JO N ,85W, intertidal, coarse sand, Aug 1996; (3) ,Remarks. Complete spec im en 35 setigers, 6.9mm long, 0.6 mm maximum width . Variablenumber (6-1 7) of swollen anterior segments,emergent dorsal hooks frol11 setiger 7, and 24smooth capillary neurosetae. Wolf ( 1984) reported neurosetae with a minutely bidentatetip but this was not true of these specimens.Salaur-Vallejo & Orensanz (1991) reportedventral parapodial cirri on all setigers forspecimens from Rio La Pl ata, Uraguay-Argent ina, but ventral parapodial cirri beg in onsetiger 3 in the Costa Rican specimens as waspreviously described for C. incerto by Petti bone ( 1966) and Wo lf( 1984).Distribution. Previously reported from Chesapeake Bay, USA to Rio La Plata, UruguayArgentina in the Atlantic Ocean. This is thefirst record of this species in the Eastem Pacific. Known from the intertidal to 80 metersdepth.Genus Pilargis Saint-Joseph 1899This genus is characterized by the absence ofemergent hooks or spines.Pilargis berkeleyae Monro 1933 (Fig. 17).Pi/argis berkeleyoe Monro 1933:673. Pettibone 1966:161.


10/16

REVISTA DE BIOLOOIA TROPICAL

10

2

Figs. 9\2. i f O . b 1 t H u ~ n.sp . Parltype MCZ 20018) .Fig. 9. Seliger 1,anteriorview .Fig. 10.Setiger2, Interior vi ;w. Fig .II .Setiger), If1lcriorview. Fig. 12.Se liger 19,anteriorview.Scaleb Fig. 9 40 )1m. Fig.s 10.12 -)1m.


11/16

Aquatic cCOS ,51Cms of CosUt Ri c

n14

5 6

FilS. 1316. Fia. 3 . Lomd JJia riC oi, dorsJIvicw ofillterior. Fig. 14 .Parafldolla t icu.JpIJ dorsal view ofJlllerior. Fil . 5 .Cobini irIc,Ial lknIviewof mor.fi 16.Cabil tl i l lc ,o , dorSlI view orprouomiunund pcrislomiunl.. SelIc bIr I n n


12/16

l R v rSTA DE BIOLOGIA TROPICAL

17 18

9

20

Figs. 17-21. Fig. 17. i f a r g ~ beruleyae, dorsal view of anterior Fig. 18.Ancis/fW 1/is m t , 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 .


13/16

Aquat ic ecosystems ofCosta Ri ca

Im ajima 1987: 162. Salazar.Vallejo & Oren-sanz 199 1:274. Dean 1996a:74. Blake1997:270.Ma teria l Examined. Gu lf of Nicoya , SIO. 29 954'55 n N, 8445 ' 15 W, 18 m, muddy sand ,July 1980,( IXUSNM 079933).Rema rks. Neurosetae with fine teeth andbidcntate tip, dorsal c irri in anterior set igcrsinflaled.Di stribution. Prev ious ly known from thenorthern Gulf of Mexico to Argentina in th eAtlantic and from Canada to California in thePacific Ocean. This report extends its range tothe Gulf of Nicoya, Costa Rica. From inter-tidal to abyssal depths.Genus ncistrosyllis Mcintosh 1879The synapomorphy of this genus is possess ionof a five lobed hind brain (Fitzhugh & Wo lf1990). The genu s Allcistargis is sim ilar to Allcistrosyllis differing on ly in its lack. of a median antenna. Licher & Westheide (1994)synonomize these two genera due to a lack ofa synapomorphic character for ncistrosyl lis -species with three antennae if consideredseparate taxa. The median antenna of A cis-largis species is usually weakly developed(Fig. 11- 13) and the presence or absence ofsuch an inconspicuous struc ture does not warrant separation at the generic level.Aneis frosyllis hamata (Hartman 1960)(F;g. 18).Pi/argis hamatus Hartman 1960:88. Reish1968:76.Ancislrosyllis hamata. Pettibone 1966:168 .Hartman 1968:377 . Blake 1997:269.ncistargis sp. Maurer et al. 1988 :48 .

Aneistargis hamata (Ha rtman). Dean 1996a:74, I996b:83.Material Examined.Gulf of Nicoya, Sta. 12,948'25 N, 84 52'40W, 22 m, muddy sand,July 1980,( 1). Sla. 24, 9 49'25 n N, 84 41'20 W, I m, sand, Jan 1981 ,(1 ); Apr 198 1,(1) ;June 1981,(2); Aug 1981 ,(3). Sta. 28, 9 52'

16 N, 84 45'30 W. 26 m. mud, July 1980.(I)(USNM 079929); Aug 1981 .(1). ta . 29, 954'55 N. 8445'1 5 W. 18 m. muddy sand. July1980,(1); Oc t 1980,(1 , UCR III); Apr 1981.I ; June 1981 ,( 1); Aug 1981 ,( 1). Sta. 30.954'40 N. 84 45'50 W, 18 m, muddy sand.Oct 1980,(1); Jan 1981,( 1); Apr 1981 , I .Au g 1981 .( 1). Golfo Duke. Sta. C 208 2I'N , 8314'W, 200 m, mud . Dec 1993,( 1).

Rema rks . Complete specimen 39 set igers. 6 .2mm long. 0.55 mm maximum width (0.33 mmwithout parapod ia). Vent ral c irri beg in onsetiger 3 and emergent nOlopodial beg in atse tigers 67.Distribution. Prev iously known from Califor-nia to th e Gulf of Lo s An geles. Baja Californiaas well as the French Mediterranean(Katzmann et al. 1974 . This ex tends its rangein the eastern Pacific 10 Golfo Du lce. CostaRica. Reponed from el even meters 10 abyssa ldepths.Ancislros ,lIis Qpi/losa (Jones 196 1)(Fig. 19).Ancislargis papi/loslls Jones 1961:3Allc istrosyllis IUlpi//osa Pettibone 1966:170.Allcislrosyllis sp. Maurer et al. 1988:48 ( inpart)Allcistargis hamata (1lanman). Dean 1996a:74 (in part)Material Ex amin ed. Gulf of Nicoya , Sta. I .957'30N, 84 53'00 W. 46 . mud, Ju ly1980,( I)(USNM 079928). Sta. 14, 9 49'25 N.844I'20 W, I I m, sand, June 1981.( 1 + I(UCR 1304 ; Aug 1981 ,(2): Sta. 28. 952'16 N, 84 45'30 W, 26 m, mud. July 1980,(I)(USNM 079931). Sta. 30. 9 54'40N, 8445'50 W. 18 m, muddy sand, Apr 1981.( I).Remarks. Complete specimen 44 setigers. 9.0mm long, 0.5 mm maximum width (0.3 mmwithout parapodia). Emergent notopodialhooks begin on set igcr 3. paired ventral eyespots weak ly pigmented and often inconspi-cuous. a weakly developed median antenn a onthe posterior of th e prostomium. Ventral c irribegin on setiger 3 and the dorsal cirri possessnumerous e longate papillae. Setae are longcapillaries and shoner pectinate setae withsligh tly falca te ends, lacking Ihe i n d i s t se


14/16

60 REV ISTA DE BIOLOGIA TROPICAL

condary loolh ofWolf(1984, Fig. 29, 20 GH.Distribution . Previously re poned from theintert idal to 46 meters from Florida and Texasin th e Gu lf of Mex ico. This is the fi rs t recordof this species in the eastern Pacific.Ancislrosyllisjonesi Pettibone 1966(F;g. 20).Ancistrosylis jones; Salazar-Val lejo 1987(1986),200.Ancysfrosy/lis jonesi Pettibone 1966: 173.Le6n-Gonza lez 1991: 3 11. Dean 1996a:74.Anc)'s osyllis sp. (in part) Maurer ef o/.1988048.Material Examines. Gulf of Nicoya, Sla. ]49049'2S N. 8441'20 W, m, sand, Oc t1980 ,{1 I(UCR 11201. Sta. 28. 9052' 16 N,8445'30 W, 26 m, mud, Ju ly 1980,( J)(USNM079932). SIO. 29. 9 54'55 N. 844S IS W, 18m, muddy sand, Jan 1981,(2): June 198 1.( I).Remarks . Complete specimen with 66 sel igers, 15.7 mm long, 0.8 mm maximum width(0.4 mm withoUi parapodia. Body dorsoventrally flattened, dorsum with minute papi llae,notopod ial ~ m ~ r hooks beg in on setiger 6,ventral c irri begin on setiger 3, med ian antenna inconspicuousDis tribution. Previously desc ribed fromChesapeake Bay to Campeche, Mex ico in theAt lantic Ocean and from Baja Cal ifornia Sur,Mexico. This extends its range in th e easternPacific to the Gu lF of Nicoya, Costa Rica. Previously co llec ted from the in te rt ida l to 60meters.ncystrosyllis hartmanae Pettibone 1966(F;g. 21).

Ancystrosyllis hartmanae Pett ibone 1966:172.Wo lf 1984:29.24.

aterial Enmined .Golro Dulce, GoJfitoBay 8OJ8'N. 8310'W, intertidal 2 Km west ofmain dock , muddy sand, Sept 1986; ( I I(UCR .Rem ....ks . Complete specimen 36 setigers, 8.0mm long, 0.7 mm maxim um width (0.5 mm

not including parapodia). Ventral cirri andemergent notopodial hooks begin o n setiger 3(th ird on the right and fourt h on th e right inFig. 13), an inconspicuous med ian antenna.The di sti nc t flexed anterior region of the digestive trac t desc ribed by Pettibone ( 1960) iseasily seen through the integument.Di st ributio n. Reported subtidally in 10-135meters frOIll Chesapeake Bay. USA and th eGu lf of Mexico . This ;s the first record of thisspecies in the eastern Pacific and its firs t recorded intertidal occurrence.

ACKNOWLEDGEMENTSThe numerous co llection trips to Costa Ricahave been made possible through the aid andencouragement of the University of CostaRica' s Centro de Investigacion en Ciencias delMar y Limnologia (CiMAR) and its d irectorJose A. Vargas. This manuscript was im proved by the review of an earl ier draft byEdward CUller, Department of InvertebrateZoology, Museum of Comparative Zoolvgy.Harvard University and other errors correctedby two anonymous reviewers.

RESUMENSe in fonna sobre 13 especies depoliquetos de la familia Pilargidae (Annelida:Polychaeta) de ambientes submareales e intermareales en la costa Pacifica de Costa Rica.Se describe la especie nueva Sigambra vargasiy se informa sobre Sigambra sp. A, au n no

descrita. La ocurrencia en Cos ta Rica de Si-gambra lentaculata (Treadwe ll), Loanda/iariojai Salazar-Vallejo, Pi/argis berkeley;Monro, ncislrosyllis hamata Hartman andncistrosyl/is jonese; Penibone. representa

amp liaciones de ambito para esas especics. Laocurre ncia de Sigambra pelliboneae HamannSchroder. Parandalia Iricuspis (Mu ller),Cabira incerta Webster. Ancistrosyllis papil-losa Jones and Ancistrosyllis hartmanae Pettibone son nu evos hallazgos de estas especiesen el Oceano Pacifico O riental costero de estaregi6n. Synelmis cf albini (Lange rh ans) tamse menciona en este trabajo. Se ineluyeuna clave dicotomica para la identificaci6 n delas especies de Pilarg idae de Costa Rica.


15/16

AquRtic ecosystems o f Costa Rica 61

REFERENCESAuge ncr . II . 1927. Polycllaeten von Silost und SOd

Austral ien . Videns . Medd. Dan. Natur. For. Kob. 83:7 27SBanse, R. . R. D. Bobson 1974. Bentllic errantiate poly_chaetes of Brit ish Columb ia and Washington. Bu ll.Fi sh. Res. Boa rd Can. 185: I-I I I.Bert eLey, E. . C, l3en.;eley 194 1. On a colI tion ofIolychaeta from so uthern California. Dull. South. Cal.Ac. Sc . 40: 16-60.Blake. J. A. 1997. Fam ily Pi largidae Sa int Joseph, 1899.In J .A. Blake, O. Hilbig . 1 .11 . SemI (cds.). Tuonomic atlas of the benthic fauna o f the SMta Maria

b a ~ l n and western Santa Barbara channel , Volume 4The Anne lida Part I. 26 1-284. Sant. Darb. Mus. Nat.Hi s t. Sama Darbara, California.ChamtJerlin. R. V. 1919. The Annelida Polychacta. Mem.Mus. Compo ool. Harv. Coil. 49 : 1-514 .Day. J. II. 1967 . A mooograph on the Polychaetes o f

Southern Africa. Part I: Errantia; Part 2: Sedentaril .Bri t. Mus. Publ. 656: 18 78.De:lIl . II. K. 1996a. Subtidal ben thic polychaetes(Annelid a) of the Gul f of Nicoya. Cos ta Rica. Rev.

Ilio l. Trop. 44 (suppl. 3): 69,80 .Denn. II. K. 1996b. Polychaete worms (Annelida) collected in Golfo Dulce during the Victor lIensen C o ~ t a

Ri ca expedition (1993f l 994). Rev . Ri ol. Trop. 44(sup pl. 3): 81-86.Dean. H. K. 1995 A new species o f Hesionidae .

G J J ' p h ~ l l o t l f lIico} e lJiJ (Annelida : I olychaeta)from the Gulf of Nicoya. Costa Rica. Dull. Bioi. Soc .Wash. I I I; 257-262.Ehlers, E. 1908. Die bodensllssigen Anneliden aus den

Samm lungen der Deutschen Tiefsee Expcdition.Wi ss . Ergeb. Deut. Tief Exp. 1897-1899 16: 1. 168.Emerson, R. R. . K. Fauchald. 197 1. A revi sion of thegenus o nda/ia Monro with description of a new genus and species of pilagiid polychae te. Bull. Sout.Cn l. Acad. Sci. 70: 18-22.Fauchald. F. 1977. Polychaetes from intertidal lll Cas inPan.ama . with a ~ v i e w of ~ v i o U 1 shallo w water reI; ords. Smith. Contrib. loo l. 22 1: 1-85 .Fauchald, K. a: P. Jumars 1979. The d ie t o f worms: A

stud), o f polychaete ~ d i n g gui lds. Oceanog. Mar.BioI. Ann. Rev. 17: 193-284.Faucllald, K. . G. Rouse. 1997 . Polychaete systematics:Past and present. Zool. Scrip. 26: 71-138 .Fauve l, P. 1920. Les genres Ancis/,osylfis et Pifa giJ(Hesionidac) Bull. Soc . Zool. Fran. 45: 205-213.

Fauvel, P. 1923. Polychtles Erranles. Fau. fran . 5: I488.f itzhugh. K. . 1' . S. Wolf. 1990. Gross morphology ofthe brain of pilargid polychaetes; Taxono mic and

systcmatic implications. Am. Mus. Nov . 2992 : 1-16.Hartman. O. 1947 . Polychaetous annelids. g. Pilargiidac[sic] . Allan Hancock Pacific Expedition 10: 483.523.Bartman. O. ]968 . Atlas of tile emm tiate polyehsc lousannelids from California. Allan Hancock Foundation .University o f Southern California. Los Angeles. 828p.Hartmann-Sch rOder. G. 1959. lur Okologie de r Pol) .chaeten des Mangrove Estero Gebe ites von EJ Salva.dor. De it . Neo t. Fau. I : 69-183.Hartmann SchrOder, G. 1979. lur Kenntnis des Eulitoralsder australi sehcn KOsten unter besonder OcrUcksichtigung de r Polychaeten und Ostracoden, 2. DiePoly chaeten de r tropis hen Nordwestkllste (zwishenDerby im Norden und Pt. Hedland im SlIden) . Mit.Hamb. Zoo l. Mus. lns t. 76: 77-218.Imajima. M. 1987. Pilargidae (Annelida, I olychaeta)from Japan (Part 1). Bull . Na t. Sc i. Mus. SC I.(Zoo logy) 13 : 151- 164.

a ~ m a n . W . L. Laubier. . J. Ramos 1974. Pil argidae(Annclides polychctes r r a n t e de MediterranCe.Bulletin d lnsitute Occanographique Monaco 71 : 140.Langerhans . P. 1881. Ober elnige eanarische Anneliden .Deutsche Akademie der Naturforscher Nova Acta 42:93 -124.Le6n-GonzAlez, J. A. 1991 . Poliquetos de fon dos blandosde la costa occidenta l de Baja California Sur, M6cico.I. Pilargidae . Cab. Bioi. Mar. 32 : 3 11-32 1_Licher. F. 1994 . Resurrection of G y p " ~ s i o n e Fr ie.drich. 1950. wilh redescription of G. au i Friedrich,1950 and description of G. iongocirrala (Polychpeta:Hesionidae). Proc. BioI. Soc. Wash. 107 : 600 -608.Licher, F. . W. Westhe ide. 1994. The phylogeneti cposition o f the Pilargidae with a cladi st ic analysis ofthe taxon-faets and ideas. In J. C. Dauvin. L. Laubier,D. J. Reish. (eds) . Actes de la 4 me Conftn:ncc internat ional des Polyehaetes. Mem. Mus . Nat . Uist.Na t. 162: 223236.Licher. F. . w. Weslheide. 1997. Review o f the genus

Sigamba (PolychllC tl : lic:sionidae). redesc ription o f S.haul (Hartman 1947), and descriptions of two newspecies from Thailand and China. Steenstrupi a 23: I20.Maurer, D. . J. A. Vargas. 1984. Diversity of softbonombenthos in a tropical estuary: Gulf of Nicoya. Cos taRica. Mar. Bioi. 81 : 97106.


16/16

62 REVIST A DE BIOl.OGIA TROPICAL

Maurer, D . J. A. Vargas H. Dean. 1988. Polychactousannelids from the Gulf of Nicoya, Costa Rica. Inter.rev. ges. Hydrobiologie 73: 43-59.Mclntosh, W. C. 1879. On the Annelida obtained duringthe cruise of H.M,S. Valorous to Davis Strait in1875. Tran. Linn. Soc. 2: 499-511.Monro, CA. 1933. On a new species of Polychac:ta of thegenus Pi larg is from Friday Harbour, Washington.

Ann. Mag. Nat. lI isl. 10: 673-675.MOiler, F. IS58. Einiges Ubcr die Anne liden Fauna derIn sel SI. Catherina an der Brazilianischen KlIste.Ar ch. Natur. (Be rli n 24: 211-220.Pettibone, M. H. 1966. Revision of the Pilargidae(Annelida: Polychaeta), including descriptions of newspec ies, and n:descriptions of the pe lagic Podarmusploa Chamberl n (Polynoidac). Proc. Nat Mus. 118:

1 5 S 2 0 ~

Rei sh, D, J. 1968. A biological survey of Bahia de LosAngeles, Gu[f ofCa [ifornia, Me.xico. Il. Benthic polychaelous annelids. Tran. San Diego Soc. Nat. Hist. IS:67-106.Salazar-Vallejo, S. I. \987 (1986) . Pi larg idae (Annelida:Po[ychaeta) de Mexico: Lista de especies. nueva especies y biogralia. Cah . BioI. Mar. 27: 193-209.Salazar-Vallejo, S. I. [990. Redescriptions of Sigambragrubii MOller 1858 and ermundura Iricuspis MUller

1858 from Brazil and designation of neotypes (Polychaeta: Pi[argidae). Jour. Nat. Hi st. 24: 507-5 I7.Salazar-Vallejo, S. I., 1. A. de Le6n-Gonzillel J. C.ChAvez-Companin . 1990. Poliquetos (Annelida: Polychaeta) de a Bahfa de Manlanillo, con una claveilustrada para las espec ies de Colima, Mexico. Rev.BioI. Trop. 38: 21 1-229.Salazar-Vall ejo, S. I J. M. Qrensanl 199 1. Pih\rgidos(Annelida: Polychaeta) de Uruguay y Argentina. Cah.BioI. Mar. 32: 267-279.Treadwell. A. L 1941. Polychactous annelids from theNew England region. Puerto Rico and Bralil. Am.Mus. Nov. 1138: 1-4.Vargas, J. A. 1995. The Gu[f of Nicoya estuary. Costa

Rica: past, present, and future cooperative research.Helgol. wiss. Meeres. 49: 821-828.Webster, H. E. 1879 . Annelida chaetopoda of the Virginian W bSt. Trans. Alban. [nst. 9: 202-269.Westheide, W, l j , ' Interstitielle Fauna von Ga lapagosXl. Pisionidae. Ii< Jion idae, Pi largidac:, Syllidae(Polych3eta) . Mikrof Mem:sh . 44: 1-146.Wo lf, P. S. 1984. Chapter 29. F ~ m i l y Pilargidae SaintJoseph 1899. p. 29-1 to 29-41. n J. M. Uebclacker

p, G. Johnson (cds). T..:..onomic guide to the polychHe tes of the northern Gulf of Mexico, 7(4). Final re port to the Minerals Management Service, contract14-\2-001-29091. Barry A. Vittor, Mobile. Alabama.

06 dean pilargidae (1)

Documents