New Fanworm Species (Polychaeta: Sabellidae: Fabriciinae) from Phuket, Thailand, with Comments on Fabriciola flammula Rouse and Fabriciola cri Rouse Kirk Fitzhugh1 ABSTRACT.TwonewfanwormspeciesinthegeneraFabriciolaFriedrichandFseudoaugeneriellaFitzhugh are described from Panwa Bay, Phuket, Thailand. Fabriciola pbuketensis n. sp. is similartoFabriciolasp. cf. F. berkeleyi Banse, F. mediaseta Fitzhugh, F. rubra Fitzhugh, F. flammula Rouse, and F. cri Rouse in havinga lowanteriorperistomialringcollar,redperistomialandpygidialeyes,andabdominalneuropodial pinhead setae. The new species is distinguished from F. mediaseta and F. rubra bythe absenceofthoracic broadly hooded flagellate setae and differs from Fabriciola sp. cf. F. berkeleyi, F. flammula, and F. cri in that it lacks crown and trunk pigmentation; it differs in collar construction as well. Fabriciolaflammula and F. cri were originally described as having broadly hooded flagellate setae. Examination of paratype material reveals that all inferior thoracic notosetae are narrowly hooded; descriptions of the species are appropriately emended. The total number ofdescribed Fabriciola species is increasedto 16. Fseudoauge- neriella brevirama n. sp. is the second species describedinthegenus anddiffersfromF. uniramaFitzhugh inhavingvascularizedventralfilamentousappendagesthatarenomorethanhalfthelengthofthebranchial crown. Appendages in P. unirama are almost the same length asthecrown. ThemonophylyofPseudoau- generiella is discussed, and cladisticrelationships among Fabriciola species are presented. INTRODUCTION branched to some extent. Fitzhugh (1998; see also Fitzhugh, 1990b) pointed out, however, that fila- The present account describes a new species ofFa- ments in Augeneriella are initially unbranched, briciolaFriedrich, 1939, andanewspeciesofFseu- with branching developing as animals grow. Based doaugeneriella Fitzhugh, 1998, from the intertidal on a cladistic analysis ofFabriciinaetaxa, Fitzhugh zone atPhuketIsland,Thailand. Recognitionofthe (1998) found the presence ofunbranchedvascular- diversity ofFabriciola has especially increased dur- ized filaments to be plesiomorphic for the subfam- ing the past several years. Including the species de- ily, such thatitisnotpossible to includeP. unirama scribed here, there have been seven Fabriciola spe- inAugeneriella. Justification for the placementofa cies described since 1990 (Fitzhugh, 1990a, 1992a, second species in Fseudoaugeneriella as well as 1998; Rouse, 1993, 1996), givinga total of 16 spe- monophyly ofthe genus are provided by an update cies. In the process ofcomparing the Fabriciolade- ofthe cladistic analysis offabriciingenera andspe- scribed here to similar species, I found that the de- cies performed by Fitzhugh (1998). scriptions of F. flammula Rouse, 1993, and F. cri Rouse, 1996, must be emended with respect to the SYSTEMATICS type of thoracic notosetae in each. Fseudoaugeneriella was originally described Family Sabellidae Latrielle, 1825 ffrroomm OaksiinngalweaspIescliaensd,,JP.apuanni.rFaimtazhFuigthzh(u1g9h9,8)19n9o8t,- Subfamily Fabriciinae, Rioja, 1923 ed that the genus resembles Augeneriella Banse, Fabriciola Friedrich, 1939 1957,inthatbothhaveananteriorperistomialring collaronlydevelopedventrallyasawidelobe,well- Fabriciola pbuketensis n. sp. developed triangular dorsal lips, and vascularized Figures 1-2 ventral filamentous appendages. The difference be- tween the two genera is that the filamentous ap- MATERIAL EXAMINED. Indian Ocean, Andaman pendages in Fseudoaugeneriella are unbranched, Sea, Thailand, Phuket Island. Holotype: LACM-AHF whereas fully formed filaments in Augeneriella are 1896, Panwa Bay, just north (about 100 m) of Phuket Marine Biological Centerpier, scrapingoflowmatofde- tritus and algae from shale rock, midintertidal zone, col- 1. Invertebrate Zoology Section, Research and Collec- lectionmadeatlowtide,depth0m, 18August1997,coll. tions Branch, Natural History Museum of Los Angeles K. Fitzhugh. Paratypes: LACM-AHF 1897, 11 specimens County, 900 Exposition Boulevard, Los Angeles, Califor- (10complete, 1 missingcrown),samelocalityasholotype. nia 90007. Paratypes: LACM-AHF 1898, 11 specimens (2 complete, Contributionsin Science,Number477, pp. 1-17 Natural HistoryMuseum ofLos Angeles County, 1999 9 lackingcrown), same locality as holotype,scrapingsoff 1C); uncini in setigers 9-11 number 13, 12, and dead, branching, Acropora-like coral, midintertidalzone, 10, respectively. Anus midventral, along anterior collection made at low tide, depth 0 m, 18 August 1997, margin of pygidium. Males with spermiogenesis coll. K. Fitzhugh. occurring in setigers 4-8, oocytes in females in se- ETYMOLOGY. The specific epithetrefers to the tiger 4. Preserved specimens white, no pigmenta- occurrence of the species at Phuket Island. tion on crown or body wall. Tubes composed of DESCRIPTION. Holotype a complete male fine, flocculent detrital material, total tube width with 8 thoracic and 3 abdominal setigers; bran- about 2 times greater than body width. Brooding chial crown length, 0.45 mm; remainder of body, of young not observed. 1.70 mm long; maximum width, 0.12 mm (Fig. REMARKS. This species falls within the Fabri- 1A). Body slender, nearly uniform in width except ciola species complex defined by the presence of for slight tapering posteriorly. Branchial crown abdominal neuropodial pin-head setae (sensu Ben- with 3 pairs of radioles, distal ends filamentous, Eliahu, 1975), which includes F. mediaseta Fitz- same width as pinnules. Branchial or “radiolar” hugh, 1990 (Aldabra Atoll, westernIndianOcean), skeleton (sensu Rouse 1993) not observed. Radi- Fabriciola sp. cf. F. berkeleyi Banse, 1956 (sensu oles each with 3 pairs ofpinnules, all terminating Fitzhugh, 1992a; California), F. flammula Rouse, at about same height as ends of radioles. Dorsal 1993 (Belize),F. criRouse, 1996 (PapuaNewGuin- lips erect, triangular, rounded distally, and distinct ea), and F. rubra Fitzhugh, 1998 (OkinawaIsland). from dorsal-most radioles; low, distally rounded, Fabriciola mediaseta and F. rubra have broadly ventral liplikeprocesspresentatbase ofproximal- hooded, flagellate setae in thoracic notopodia, most pinnule of ventral-most radioles (Fig. IB). whereas inferior notosetae in F. phuketensis, Fabri- Nonvascularized ventral filamentous appendages ciola sp. cf. F. berkeleyi, F. flammula, andF. crihave present, slightly shorter than total length of radi- elongate narrowly hooded setae. The body of F. oles, surfaces smooth, about same width as pin- phuketensis lackspigmentation,whereasFabriciola nules (Figs. IB, 2B-C). Dorsal margins of bran- sp. cf. F. berkeleyi and F. cri have peristomial and chial lobes not fused to one another. Branchial branchial crown pigmentation, respectively. The hearts present. Anterior margin of anterior peri- posterior margin of the middorsal collar gap in F. stomial ring as low membranous collar (Fig. 2), flammula (Rouse, 1993: figs. 1-2) andF. cri(Rouse, with distal margin smooth all around; middorsum 1996: figs. 1-2) is noticeably expanded, whereas completely separated by narrowgap; collarheight the gap in F. phuketensis is uniformly narrow (Fig. uniform. Collar of even thickness throughout, 2A). about same length as posterior peristomial ring. Rouse (1993, 1996) allied F. flammula and F. cri Annulation between collar and posterior peristo- with F. mediaseta on the basis of the fact that the mial ring only present ventrally. Anterior peristo- 3 species have inferior, notopodial, broadly hood- mial ring, including collar, about same length as ed, flagellate setae {sensu Fitzhugh, 1990a: fig. 6C; posterior ring. Middorsal medial lobe just dorsal see also Fitzhugh, 1998: fig. IB) in some thoracic to mouth almost same height as collar. Pair of setigers. Rouse (1993: fig. 3; 1996: fig. 3) described round, red eyes in posterior peristomial ring. Se- these species as having these setae in setigers 2-8 tiger 1 about same length as posteriorperistomial and 2-6, respectively. After examiningparatypesof ring, distinctly wider than long; setigers 2-6 each F. flammula and F. cri, I find that these species do successively longer, with setigers 4-8 longer than not have broadly hooded, flagellate setae. I com- they are wide. Setiger 9 about same length as 1, pletely concur with Rouse (1996) that in F. cri, with setigers 10-11 each slightly shorter than se- there are qualitative differences in hood shape in tiger 9. Pygidium about same length as setiger 11, some setigers, and, as much as possible, these dif- posterior margin slightly tapered, rounded. Some ferences should be reported. Comparatively, how- paratype specimens exhibit greater longitudinal ever, I regard the setae in F. cri to come closest to contraction, with most thoracic setigers about as the condition of elongate narrowly hooded as op- long as they are wide. Pair of round, bright red posed to flagellate. In the latter condition, there is pygidial eyes. Superior thoracic notosetae elon- a marked disjunction between the hood and distal gthahotooer,daecnidacrbruntootwsolhsyoertthaeoreo,di1endp,seert3if-gae4srcispcel2re-.8fAabsacdlioscolmei.nnaIarnlfreonrweiluoy-r cFb.oentctrwii,eneuanantdhiooF.nodpohfuatknhedetedsnihssatfiatsl.sTshhhaoefwt.saetEsamemeoniondtaF.htiftolrnaamsnsmiuttoliaoF,n. ropodia of setigers 9-11 with very elongate, nar- rowly hooded setae, 1-2 per fascicle, and 2 pin- flammula and F. cri are provided in order to reflect these changes. head setae per fascicle. Thoracic uncini acicular, main fang slender; teeth above main fang slender and slightly decreased in size away from fang; Fabriciola flammula Rouse, 1993, Emended hood not observed; 4-5 uncini per fascicle in sin- Fabriciola flammula Rouse, 1993: 250-253, figs. gle rows (6-7 in some paratypes). Abdominal un- 1-10, 48. cini with 5-6 rows of teeth in profile, 3-5 teeth perrow;manubriumover2 times longerthanden- MATERIAL EXAMINED. Caribbean Sea, Belize, Car- tate region, slightly expanded proximally (Fig. rie Bow Cay. Paratypes: LACM-AHF 1630, 3 specimens, 2 Contributionsin Science, Number 477 Fitzhugh: Thailand Fanworms Figure 1. Fabriciola phuketensis n. sp. A, entire animal, lateral view (holotype, LACM-AHF 1896); B, right half of branchialcrown, innerview (paratype, LACM-AHF 1897);C,abdominal uncinusfromsetiger9 (paratype,LACM-AHF 1897). Abbreviations: bh, branchial heart; dl, dorsal lip; vfa, ventral filamentous appendage; vl, ventral liplikeprocess. Contributionsin Science,Number477 Fitzhugh: Thailand Fanworms 3 Figure 2. Fabriciola phuketensis n. sp. A-C, dorsal, lateral (right side), and ventral views of anterior end, respectively (holotype,LACM-AHF 1896).Abbreviations:apr,anteriorperistomialring;bh,branchialheart;ppr,posteriorperistomial ring; vfa, ventral filamentous appendage. south side of leeward jetty, algal turf on boulders and Fabriciola cri Rouse, 1996, Emended Strombus shells, sta. F400, 22 May 1991,coll. G. Rouse. Fabriciola cri Rouse, 1996: 1765-1768, figs. 1-12, REMARKS. Rouse (1993: 252) described inferi- 39, 42-43. or thoracic notosetae in setigers 2-8 as broadly hooded, flagellate. All inferior thoracic notosetae MATERIAL EXAMINED. Pacific Ocean, Papua New are elongate, with narrow hoods, as seen in most Guinea, Madang Province. Paratypes: LACM-AHF 1798, 10 specimens, fromloglyinginmangrovesoppositeRiwo other Fabriciola species. It should be noted, how- village,sta.F645,23August1993,coll. G.Rouse,K.Fau- ever, that the hoods ofinferior notosetae in setiger chald, L.A. Ward, and P. Scott. 1 of most Fabriciola are narrower than those seen in subsequent setigers. REMARKS. Rouse (1996: 1767,fig. 3) described Rouse (1993: 251) stated thatventral lips areab- inferior thoracic notosetae insetigers2-6 as broad- sent. At the base of each proximalmost pinnule of ly hooded, flagellate. I consider these setae to be the ventral radioles, there is a distinct, distally elongate, narrowly hooded, as seen in most other rounded swelling, similar to that described here in Fabriciola species. As I noted in the remarks on F. F. phuketensis (Fig. IB) andseveralotherFabriciola phuketensis, however, I agree with Rouse’s assess- (e.g., Fitzhugh, 1990a: fig. IE for F. baltica Fried- mentthatthe hoods ofinferiorsetae in setigers2-6 rich, 1939; Fitzhugh, 1998: fig. 2B for F. rubra) as are somewhat broader than those seen in setigers 1 a “ventral lip-like process.” and 7-8. 4 Contributionsin Science, Number 477 Fitzhugh: Thailand Fanworms Rouse (1993: 251) stated that ventral lips are ab- Pseudoaugeneriella Fitzhugh, 1998 sent. The bases oftheproximal-mostpinnulesofthe ventral radioles do show some slight swelling, but Pseudoaugeneriella brevirama n. sp. this is not pronounced and does not impart the ap- Figures 3-4 pearance of the “ventral lip-like process” described MATERIAL EXAMINED. Indian Ocean, Andaman above in F. phuketensis (Fig. IB) or several other Sea, Thailand, Phuket Island. Holotype: LACM-AHF Fabriciola species (e.g., Fitzhugh, 1990a: fig. IE for 1899, detrital scrapings off dead, branching, Acropora- F. baltica; Fitzhugh, 1998: fig. 2B for F. rubra). like coral, midhintertidal zone, collection made at low tide, depth 0 m, 18 August 1997, coll. K. Fitzhugh. Par- REVISED KEY TO FABRICIOLA SPECIES atypes: LACM-AHF 1900, 17 specimens (1 complete, 16 missingposteriorend),samelocalityasholotype,scraping The following key is modified from Fitzhugh (1998). As oflow mat ofdetritus and algae from shale rock, midin- with that key, this one does not include F. spongicola tertidal zone, collection made at low tide, depth 0 m, 18 (Southern, 1921) orF.pacifica(Annenkova, 1934),asma- August 1997, coll. K. Fitzhugh. tceormipalleitse.unaFvaabirliacbiloleaantodneorreilglianaBladnessec,ri1p9ti5o9n,siasreintcoloudiend- ETYMOLOGY. The specific epithetrefers tothe based on the original description. short, vascularized, ventral filamentousappendages of the crown. la. Two abdominal setigers . . . F. minuta Rouse DESCRIPTION. Holotype complete with 8 tho- b. Three abdominal setigers 2 racic and 3 abdominal setigers; branchial crown 2a. Pygidial eyes present 3 length, 0.4 mm; remainder of body, 1.3 mm long; b. Pygidial eyes absent F. parvus Rouse maximumwidth, 0.15 mm. Bodyslender,slightlyta- 3a. Abdominal neuropodial pin-head setae pres- peringanteriorlyandposteriorly(Fig. 3A).Branchial ent, peristomial and pygidial eyes range from crown with 3 pairs of radioles, distal ends filamen- red to brown 4 tous, same width as pinnules. Radioles eachwith4- b. Pin-head setae absent, eyes black to light 5 pairs ofpinnules, terminating at or slightly below brown 9 distal ends of radioles. Dorsal lips erect but low, 4a. Thoracic notopodia withinferiorflagellatese- broadlyrounded distally, distinctfromradioles;low, tae in setigers 3-7 5 distally rounded, ventral liplike processes presentat b. Flagellate setae absent 6 bases of proximalmost pinnules of ventral radioles 5a. Peristomial and pygidial eyes bright red . . (Fig. 3B). Vascularized ventral filamentous append- F. rubra Fitzhugh ages present, about one-half the total length of ra- b. Peristomial and pygidial eyes faint red dioles, surfaces slightly to verywrinkled, aboutone- F. mediaseta Fitzhugh third wider than pinnules; interior of each filament 6a. Peristomial and pygidial eyes bright red . . 7 occupied by large blood vessel (Figs. 3A-B, 4B-C). b. Peristomial and pygidial eyes reddish-brown Dorsal margins of branchial lobes not fused to one Fabriciola sp. cf. F. berkeleyi Banse another. Branchialheartspresent. Anteriormarginof 7a. Body without pigmentation 8 anterior peristomial ring is a low ridge dorsally and b. Base of branchial crown with black pigment laterally (Figs. 3A,4A-B). Collardevelopedventrally F. cri Rouse aslow,triangularlobe;widebasally,taperingdistally 8a. Posterior margin of middorsal gap in peristo- to broadlyroundedmargin (Fig.4C).Annulationbe- mial collar distinctly expanded tweenanteriorandposteriorperistomialringsvisible F. flammula Rouse ventrally. Middorsal medial lobe just dorsal to b. Middorsal collar gap narrow along entire mouthlow, triangular. Pairofreniform(dorsalview) length F. phuketensis n. sp. or rounded (lateral) black eyes in anterior half of 9a. Anteriorperistomialringcollarrelativelyeven posterior peristomial ring. Posteriorperistomialring in height all around 10 is fourto five timeslongerthan anteriorring. Setiger b. Collar higher ventrally 11 1 slightlyshorterthanposteriorperistomialring,dis- 10a. Middorsal gap in collar very wide tinctlywiderthanitislong;remainingsetigersslight- F. ghardaqa Banse ly longer but all wider than they are long. Setiger 9 b. Collar gap narrow F. berkeleyi Banse slightly shorter than 8; setigers 10-11 each about 11a. Thoracic uncini few in number, 2-3 per fas- one-half the length of 9. Pygidium about the same cicle 12 length as setiger 11, posterior margin slightly ta- b. Thoracic uncini more numerous, 5-8 per fas- pered, rounded. Pair of round, black pygidial eyes. cicle 13 Superior thoracic notosetae elongate, narrowly 12a. Dorsal andlateralmarginsofanteriorperisto- hooded, 3 per fascicle. Inferiorthoracicnotosetaein mial ring collar relatively high setigers2 and 7-8 alsonarrowlyhoodedbutshorter, F. baltica Friedrich 1-2 per fascicle; setigers 3-6 each with 2 pseudos- b. Dorsal and lateral margins ofcollar very low patulate setae (Fig. 3C). Abdominal neuropodia of F. liguronis Rouse setigers 9-11 with very elongate, narrowly hooded 13a. Branchial crown comprises 1/5 to 1/8 oftotal setae,2-3perfascicle.Thoracicunciniacicular,main body length ... F. brevibranchiata Fitzhugh fang slender; single large tooth slightly offset over b. Branchial crown longer . . F. tonerella Banse main fang, followed by a series of smaller teeth; Contributions in Science,Number477 Fitzhugh: ThailandFanwormsI5 Figure 3. Pseudoaugeneriella brevirama n. sp. A, entire animal, lateral view (holotype, LACM-AHF 1899); B,righthalf of branchial crown, inner view, entire crown length is 0.62 mm, ventral filamentous appendage length is 0.27 mm (paratype, LACM-AHF 1900); C, inferior thoracic notopodial pseudospatulate seta from setiger 3 (paratype, LACM- AHF 1900); D, thoracic uncinus from setiger 3 (paratype, LACM-AHF 1900). Abbreviations: dl, dorsal lip; vfa,ventral filamentous appendage; vl, ventral liplikeprocess. hood present (Fig. 3D); 4-7 uncini per fascicle in posedofdetritusandmucus. No broodingofyoung straight or irregular single rows. Abdominal uncini observed. with 7-8 rowsofteethinprofile, 3-4teethperrow; REMARKS. Pseudoaugeneriella brevirama is manubrium about 1.5 times longerthan dentatere- nearly identical to P. unirama, known only from gion, slightly expanded proximally; uncini in setig- Okinawa Island. Bothspecieshaveinferiorthoracic ers 9-11 number 17, 16, and 11, respectively. Anus pseudospatulate setae in setigers 3-6 and abdomi- midventral,alonganteriormarginofpygidium. Oc- nal uncini with manubria that are about 1.5 times currence of oocytes or sperm could not be deter- longer than the dentate region. The only difference mined. Branchial crown unpigmented in all speci- between the two species lies in the length of the mens. Darkto light brownpigmentpresentinmost ventral filamentous appendages. The appendages specimens (absent in holotype) in dorsum of pos- extend to nearly the distal end of the crown in P. terior peristomial ring and present dorsally, later- unirama (Fitzhugh, 1998: fig. 10B), whereas in P. ally, and ventrally in setigers 2 or 3; remainder of brevirama, appendages are one-half the length of bodycreamcolored. Specimens in loosetubescom- the crown. 6 Contributionsin Science, Number 477 Fitzhugh: ThailandFanworms mm 0.1 Figure 4. Pseudoaugeneriella brevirama n. sp. A-C, dorsal, lateral (left side), and ventral views ofanteriorend,respec- tively (paratype, LACM-AHF 1900). Abbreviations: apr, anterior peristomial ring; bh, branchial heart; ppr, posterior peristomialring; vc, ventral lobe extensionofanterior peristomialring; vfa, ventral filamentous appendage. The holotype ofP. brevirama is one ofthe small- ducted two separate analyses, taking into consider- er specimens in the type series. One of the largest ation different outgroup conditions forthe dentition specimens, which is complete except for the abdo- in thoracic uncini (character 11), and the following mm men, has a crown length of0.53 and a thorax statesweretreatedasplesiomorphicintherespective length of 1.20 mm. analyses: state 1, teeth gradually decrease in size away from the main fang; and state 2, large tooth THE STATUS OF PSEUDOAUGENERIELLA slightly offset from midline, followed by a series of smallerteeth. Separate analyseswere alsoperformed The description of the monotypic Pseudoauge- in the present study. Taxa included 58 fabriciin spe- neriella byFitzhugh (1998) wasbasicallydeveloped cies among 13 genera. The recognitionofFabriciola out of necessity as a means of accommodating a berkeleyi and Fabriciola sp. cf. berkeleyias different species that could not be placed in Augeneriella. species and thus their inclusion as separate entities With the discovery of a species nearly identical to intheanalysisherewasjustifiedbyFitzhugh(1992a: P. unirama, the question ofthe monophylyofPseu- 71). Character state assignments forspecies (Appen- doaugeneriellamust beaddressed. Thisisespecially dix II) are the same as those used by Fitzhugh critical since there are no features unique to Pseu- (1998), exceptin the cases ofP. flammula andP. cri, doaugeneriella that are not also found in at least which have been recoded with only narrowlyhood- some other Fabriciinae taxa. The most recent cla- ed inferior thoracic notosetae [state 13(0)] in accor- distic analysis of relationships among Fabriciinae dance with the emendations discussed above. Clad- taxa was that of Fitzhugh (1998), which provided ograms were constructed using the program Hen- the basis for recognizing Pseudoaugeneriella. The nig86 (Farris, 1988), with the heuristic command data from that study were used to perform a cla- options “mhennig*” and “bb*.” Character statedis- distic analysis here, with the inclusion ofP. brevir- tributionsamongtreeswereexaminedusingtheTree ama and Fabriciola pbuketensis. Gardener program (Ramos, 1997). A total of 21 characters were used (Appendix I), Both analyses produced over 1,071 trees (maxi- comprising a total of 40 apomorphic states, these mum held in computer memory), each with a length being the same ones used by Fitzhugh (1998) in an of 65 steps, a consistency index (ci) of 0.63, and a analysis of relationships among Fabriciinae genera retention index (ri) of 0.89. The consensus trees for and species. In that analysis, Fitzhugh (1998) con- bothanalyses (Figs. 5-6) aresimilartothosefoundby Contributionsin Science, Number477 Fitzhugh: Thailand Fanworms 7 GenusA Manayunkia+ Monroika Pseudofabriciola baltica liguronis minuta parvus berkeleyi ghardaqa brevibranchiata Fabhciola tonerella F. sp. cf. F. berkeleyi flammula cri phuketensis, n.sp. mediaseta rubra N \\\^ Pseudofabricia Fabricia \\ Parafabricia ^ Brifacia Augeneriella umrama Pseudoaugeneriella brevirama n.sp. | , Novafabricia Fabricinuda Figure 5. Strict consensus cladogram based on 1,071 cladograms from analysis with character 11 (thoracic unciniden- tition) coded as state 1 (teeth gradually decrease in size away from main fang) for the outgroup. Monotypicgenera are indicated by dashed branches; species are shown for Fabriciola and Pseudoaugeneriella nonmonophyletic genera are ; indicated by white bars. 8 Contributionsin Science, Number 477 Fitzhugh: ThailandFanworms GenusA Manayunkia+ Monroika Pseudofabriciola baltica liguronis minuta parvus berkeleyi ghardaqa brevibranchiata Fabriciola tonerella F. sp. cf. F. berkeleyi flammula cri phuketensis, n.sp. mediaseta rubra Pseudofabhcia Fabricia Parafabricia Brifacia Augeneriella unirama Pseudoaugeneriella brevirama, n.sp. | Novafabricia Fabricinuda Figure 6. Strict consensus cladogram based on 1,071 cladograms from analysis with character 11 (thoracic unciniden- tition) coded as state 2 (large tooth above main fang followed by series of smaller teeth) for the outgroup. Monotypic genera are indicated by dashed branches; species are shown for Fabriciola and Pseudoaugeneriella nonmonophyletic ; genera are indicated bywhite bars. Contributions in Science,Number477 Fitzhugh: ThailandFanworms 9 01 10 13 Monotypic genera are indicated by dashed branches; species-level relationships are shown for Novafabricia and Auge- neriella. The distribution of states for characters 10 (thoracic pseudospatulate setae) and 13 (manubrium length in abdominal uncini) is shown for terminal taxa. Note that the presence ofpseudospatulate setae in setigers 3-6 [10(3)] is a synapomorphyfor Pseudoaugeneriella (cf. Fig. 8). Fitzhugh (1998: figs. 18, 28), indicating the presence clade that contains at least Augeneriella Novafa- ; oftopologiesinwhichAugeneriellaandNovafabricia bricia Farafabricia Fitzhugh, 1992; Brifacia Fitz- ; Fitzhugh, 1990, are not monophyletic. As well, the hugh, 1998; Fabricia Blainville, 1828; and Fabri- consensus tree for the analysis with state 11(2) pie- cinuda Fitzhugh, 1990. The lack of resolution in siomorphic (Fig. 6) also allows forthepossibilitythat the consensus tree is the result of movements of Fabriciola may not be monophyletic (see also Fitz- Pseudofabricia Cantone, 1972, within and outside hugh, 1998: figs. 28, 31). Relationships among Fa- this clade (see Fitzhugh, 1998). briciola species will be addressed in the next section. The Fseudoaugeneriella clade is defined byeither In ail trees produced in both analyses,Fseudoau- state 10(3)-(distribution of inferior thoracic pseu- generiella is monophyletic, with P. unirama and P. dospatulate setae; Fig. 7) or 13(2)-(length of the brevirama as sister taxa (e.g., Figs. 7-8). In all to- manubrium of abdominal uncini; Fig. 8), depend- pologies, Fseudoaugeneriella is sister group to a ing on the topology. The two species ofPseudoau- 10 Contributionsin Science, Number477 Fitzhugh: Thailand Fanworms