Table Of ContentNew 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
