Some nemerteans (Nemertea) from Queensland and the Great Barrier Reef, Australia

15 Pages·2001·2.1 MB·

TThhee ZZoooloolgiocagliSoccaielty Society oofJfap anJapan
ZOOLOGICAL SCIENCE 18: 1259-1273 (2001) @ 2001 Zoologica lSociety of Japan
SomeNemerteans (Nemertefar)om Queensland
and

the GreatBarrierReef,Australia

Ray Gibson"' Sundberg2
andPer
`School oi Bioiogic aanld Eamb Sciences ,Liverpoo Jlohn Moores Llpivers iBtyy;rom Street,
LivetpooLl33,4EU.K.

2Departrnent of Zloolog yG;Oteboig Univetsi tPy.} O. Box 463,
SE-40530GditeboigS,vveden

ABSTRACT-Threespeciesof･marinenemerteansdescribedandillu$tratedfromQueenslandandtheGreat
Barrier Reef, Australia ,include ene new genus and two new species: these are the monostiliferous
hoplonemertean sThallasionemertes ieucocephal agen ,et sp, nov, and Correanemertes potyt?phthai mspa.
nov. A new colour variety of the heteronemertea nMicrura callima is also reported, this species previously
only being known from Rottnes tlslan dW,estern Australia .A key for the fiel didentificati oonf the marine
nemertean6 recorded from coastal Queensland and the Grea tBarrl eRreef is provided.
Key words:Nemertea, Australi aK,ey, new taxa
theGreatBarrierReefarea.SpecimenswereanaesthetizedinMgC12,
tNTRODUCTION examined fo rexternal featur easnd then tixe din a seawater Bouin's
G a imTaher dea' rsl( i1e8s tr3e p3bor)rti oe f afAcucsoutnrta loifa ntnewmeor tseapnescie si s oQtu moayr ianned sfdoielun pwtoaixosni. ta neSdd ei cns tttahiienoe nMdsu wse breeyu mt hsue b oMstea qTlurelnootplryiy tcra ilQ cucuhetre oanmts e7lm aeptnmhdo di(.n M5 TT8y0QpC)e Tm ,mo.apwt,en rpsiavalri ailsfl-e,
heteronemertean sB,orlasia vittata (no wLineus vittatus) from Australia.
Tasmania and Borlasia vin'dZ (ssubseque snytnolnyymised with
NotospermustTicuspidatusf)romNew SouthWales(TabI1e).
SYSTEMATICS
Excludin gterrestri aalnd freshwate traxa, a tota lof 61 species
of nemerteans has now been recorded from Australian Clas sAnopla
waters{Gibson,1995,1997,1999;SundbergandGibson,1995) SubclassHeteronemertea
(Tab l1e). Of these, 25 species {41% )have been recorded Genus Mictura Ehrenberg ,1828
from Queenslan dand the Grea tBarrie Rreef (Tab i1e) ,two- Diagnosis :The followin gdiagnosi sof the genus MiOrur ais
third sof these belongi ntgo the anoplan subclass Hetero- based upon informati ogniven by Sundberg and Gibson (1 995:
nemertea. 120) :heteronemertean swjth single pair of horizont aIlateral
DuringavisittoQueenslandin1995,severalturtherspe-cephalic furrows ,posterior elnylarged to form wide bays, from
cies of nemerteans were discovere dby PS. One of these which ciliated cerebral canals emerge from median or ventral
proved to be a new colour morph of a species, the hetero- walls; probosc iunsbranched, typicall ycontaining two muscle
nemertean Micrur acalLima Sundberg and Gibson, 1995, pre- layers (out ecirrcular, inner longitudinal )s,ome $pecies with
viously known only from Rottnes tlslan dW,estern Australia incomplet eouter longitudi nallayer ,and none, one or two
(Sundber agnd Gibson ,1995; Gibson ,1999). The remaining muscle crosses; rhynchocoel circular musculature not inter-
taxa were previous lunyknown and two of these, both woven with adjacent body wall inner bongitudi nmauslcle layer;
monostiliferous hoplonemerteans a,re named and described dorsa ltibrou scores ef cerebral gangli aforke donly at rear
for the fir sttime in the presen tpaper. int oupper and lower branches ;nervous system with neither
neurochords nor neurochord cells, neu roganglionic tissue osf
MATERIALS AND METHODS brai nlobes not usually separated from body wall muscles by
outer neurilemma; foregu twith or without somatic muscles, if
The nemerteans were collected during July and August 1995 presen vtariably composed of circular andlor longitudi nfail-
fromseveralsamplingsites,eitherontheQueenslandcoastorwithin bres ;dermis variable, mostly with distin cctonnective tissue
'Correspondingauthor:Tel,FAX.[phone]i Iaye rseparating glandula rzone from body wall musculature;
[phone]. caudal cirrus present ;toregu twithout subepithelia lgland cell
NNIII-IE-leEcltreoncitcronic Library Service
TThhee ZZoooloolgiocagliSoccaielty Society ooff JJaapapnan
1260 R. Gibso nand P. Sundberg
Table 1. Species of marine nemerteans previousl yrecorded from Australi aT.axa reported from Queensland and the Great Barrie rReef are
liste idn boid itaiic sH,igher taxonomic categories are used in accordance with Sundberg's (1991 p)roposals .E = possibl yendemic species
which thus far have only been found in Australia wnaters.
Taxon Reference fir sretcording species from Australia wnaters and
comments
ClassAnopta

SubclassPalaeonemertea
darinoma patricia eGibson ,1 979 E Gib$on ,1979a
Cephatotrich eallola aGibso nand Sundberg, 1992 Sundberg and Gibson, 1995
Hubrechtetl amalabarensis Gibson ,1979 EE Gibson ,1979b
Hubrechtetla queenslandica Gibson, 1979 Gibson, 1979a
Procephalothn' axrenarius Gibson ,1990 Sundber gand Gibson ,1995
SubclassHeteronemertea
Aetheothynchus actites Gibson, 1981 EE Gibson, 1981a
Austratineu salbidecus Gibson, 1990 Gibson, 1990a
Baseodiscus deiineatus (Del lCheiaje ,1825) Punnett ,1900, as Eupolia defineata
Saseodiscus hemprichii (Ehrenbe r1 8g3,1 ) Gibson, 1979c
Baseodiscus quinquelineat (uQsuo yand Gaimard, 1833) BOrger, 1895, as Etpolia septemlineata
BennettieSl ainsulari sCGibso 1n9,81 ) EE Gibson, 19Bla, as Bennetti ainsularis
Cerebratulu saustralis (Stimpso n18,57) Stimpson, 1857, as Meckeiia australis; inadequately
described
CerebratuluhsaddoniPunnett1,900 EEEEEPEunnett1,900;inadequatelydescribed

CerebratuljuoshnstonWhieeler, 1940 Wheeler,1940;inadequatelydescribed

Cerebratul umsagneticus Gibson ,1981 Gibson ,19Bla
Cerebratulus queenslandicus Punnett ,1900 Punnett ,1900; jnadequatel ydescribed
CerebratulutsonesianusPunnett,1900 Punnett1,900;inadequatedleyscribed

Colemanietla aloulus (Gibso tn9,81) Gibson, 1981a, as Coiemania ahulus
Goigonothynchus repens Dakin and Fordham, 1 931 Dakin and Fordham, 1931
Kitsteuer iaabocellus Gibson, 1978 EEEEEGibson ,1978
Kbhnia rottnestensis Sundberg and Gibson, 1 995 Sundberg and Gibson, 1995
Lineus biocutatu sSundberg and Gibson, 1995 Sundberg and Gibson, 1995
LineusgitviceSpusndbergand Gibson,1995 Sundbergand Gibson,1995

Lineus vittat u{sQuo yand Gaimard, 1833) Quoy and Gaimard, 1833, as Bodasia vittata; inadequately
described
Micrura cailima Sundberg and Gibson, 1995 E Sundberg and Gibson, 1995
Notospermus geniculatu(sDellCheiaje,1828) Gibson,1981a,as Micturatriclacnaree;described byRiser,

1991Quoy
Notospermus tricuspidatu s(Quo yand Gaimard ,1833) and Gaimard ,1833, as Bottasi aviri(fiS
ltarborlasi hautehingsae Gibson ,1978 EEEEEGibson ,1978
Quasiline luuscidocula Gtiubsson ,1990 Gibson ,1990a
Quasilineu psuicherrimus Gibson, 1981 Gibson,1981a
Un'chotemma nigricans Sundberg and Gibsen, 1995 Sundberg and Gibson, 1995
Vlaiencinin aalbula Gibson, 1981 Gibson, 1981b
ClassEnopla

Subclass Hoplonemertea
SuperorderMonostilifera
Aegiatonernerte cshlorophthalma Gibson, 1990 EEEE Gibson, 1990a
Aenigmanemertes norenbuigi Sundberg and Gibson, 1995 Sundberg and Gibson, 1995
Ammonemertes erseusiGibson, 1990 Gibson,199ea
Amphiporu sthomboi(t a(tSitsimps o18n5,5) . Stimpson 1,855, as PoEa thomboidali si;nadequately
described ,identifi easd a nomen dubium by Gibson &
Crandall19,89

Carcinonemertes austratiensis Campbell, Gibson and Evans, 1989 E Campbel let aL 1989
Carcinonemertes ct, carcinophila (Kblli 1k8e4r5), Bell and Hickman, 1 985; identif aise sdimilar to
variety imminuta Humes, 1942
Catcinonemertes humesiGibson and Jones, 1990 EEEEEGEiEbEsEon and Jones, 1990
Ctybelonemertesarenicolus Sundbergand Gibson,1995 Sundbergand Gibson,1995

Digononemertes austrafiensis Gibson, 1990 Gibson,1990a
Eonemerte semmyakos Gibson ,1 990 Gjbson,1990a
Gononemerte saustrafiensis Gibson, 1974 Gjbson, 1974
Haimanemertes slacksmithae Gibson ,1990 Gibson,1990a
lschyronemertes aloanyensis Gibson, 1990 Gbson, 1990a
tschyronemerte setythrophleps Gibson ,1990 Gibson,1990a
tschyronemertestetrophthalma Gibson,1990 Gibson,1990a

Pantinonemertes daguilarens iGsibson and Sundberg, 1 992 Sundberg and Gibson, 1995
Pantinonemertes enalios Moore and Gibson, 1981 E Moore and Gibson ,1981
NII-Electronic Library Service
TThhee ZZoooloolgiocaglical SSoociceityety ooff JJaapapnan
NemerteansfromAustralia 1261
Ranthionemertes mooteae Gibson, 1982 Gibson, 1982
Flantinonemertes -rinsori Moore and Gibson, 1981(stri acntEEly Moore and Gibson, 1981
estuarinespecies)
Flaranemertopsi vsvelsi Gibson, 1990 EEEEEGEibson, 1 990a
Pheroneonemertes dianae Gibson, 1990 Gibson, 1990a
Poseidonemertes bothwetlae Gibson, 1982 Gibson, 1982
fetranemertes hennaphtvditicus <Gibso n1,982) Gibson, 1982, as Nemerte sherrnaphroditicus
fetrastemma tristloru Snuandber gand Gibson ,A995 Sundberg and Gibson, 1995
Z)(gonemertes Mxactemupensis Gibson ,1999 Gibson ,1999
SuperorderPolystilifera
Drepanophorell atasmaniWheeler, 1940 E Wheeler, 1940; inadequatel ydescribed
Drepanophorus senatiCo"is (Hubrec h18t7,4) Hubrecht ,1887; no longer a valid species (Gibso 1n9,95),
the identificatio nof this taxon remains unknown {Gibson,
1997)Gibson,
Urichonemerte psilottrynchu sGibson, 1 983 EE 1983
Xenonemertes rhamphocephaius Gibson, 1983 Gibson ,1983
layer c;ephalic gland snormally well developed, occasionally
weakly formed or absent; apical organ usually present ;eyes
presen tor absent; sexes separate.
Micrura caMma Sundberg and Gibson ,1 995
Materia lexaminect 1 specimen collected 12 July 1995 from
rock, covered with coralline algae, exposed at low tide ,Tu rtle
Beach,CapeFerguson(19e16.21'4S70,3.4'E)1;specimen

collected 5 August 1995 from coral rubble on the reef flat,
LizardlslandW,atson Bay (14040.0'1S4,5026.7'E).

ExtemalieaturesT:he generaalppearance resembled that

describe dtor Micrur acaMma but the colour patte rdinffere din
being black with a white head tinge dorange towards its tip,
the ventral surface was completely black ,and the dorsa iside
banded and $triped in various shades of grey-blac (kFi g1.}.
The two small eyes we re distinctl ryed, the two dorsa llongitu-
dina lstripe$ had a fain torange tin tand the transverse bands
a tinge of sulphurous yellow T.he specimens were up to about
20 mm long and 1-2 mm wide.
Interna lanatomy and remarks: The anatomy of Micrura
callima, a species previousl yknown only from Rottnest
lslan dW,estem Austral i(aSundbe arndg Gibson 1995; Gibson
1999) ,has already been full dyescribed S.tudie son sections
of the presen tmaterial closely conform with the original de-
scri ptio ngiven by Sundberg and Gibson ( 1995: 1 16-1 20, figs
15-17), the only differenc ebseing tound relating to dimen-
sions of tissues and organs which can vary quit ceonside rably
in nemerteans depending upon their degree of contraction or
extension at the time of preservatio nI .tis theretore on mor-
phologic aglrounds not possibl eto distingui sbhetween the
vividly coloured species (dorsa al lgeynera lpurplish-brown
colour marked with magenta-pink and orange-brown longitu- Fig ,1. Micrura ca"ima, Drawing of complete individu ailn dorsa lview
dina lstripes, ventrally a brigh mtagenta) known from Rottnest basedonaoolourphotographofthelivingspecimen.Scalebar=5mm.
lslan dand the presen tspecimens. However, in all the
Queensland examples the colour patter nt,hough resembling
that of the We$tern Australi a$npecimens in terms of it scom- evident between the two colour morphs, the presen tspeci-
plex patter onf longitudi naandl transvers beands, white head mens merely represent a colour variety of Micrur acaMma
patch and do;solater sapelckle$, i$ pr miarily in shades of black, which has not previousl byeen described O,ur conclusion is
grey and white, with only fain ttraces of other colours. We supported by the fac tthat a few of the specimens previously
conclude that, with no significant morphological differences found at Rottnes tlslan dwere distinc tdlayrker than others,
NII-Electronic Library Service
TThhee ZZoooloolgiocagliSoccaielty Society ooff JJaapapnan
1262 R. Gib$on and P. Sundberg
though stil lshowing the overall brilli acnoltour patter norigi-
nally described for thi staxon (P,S p.e,rs .obs,).
Class Enopla
SubclassHoplonemertea

SuperorderMonostilifera

Genus TT;allassionemertes
gen. nov.
Diagnosis :Monostilifero umsarine hoplonemerteans ;rhy-
nchocoel extending to posteri toir pof body, with wall contain-
ing two distin cmtuscle layers ;anterior region of proboscis
with outer circular and inne rlongitudi nmaulscle layers p;ro-
bosci sarmature consisting ot single central stylet and two
accessory stylet pouches; body wall musculature without
diagona llayer ,longitudina lmuscles not divide adnteriorly; pre-
cerebral septum ventrally incomplete; front aolrgan present,
front agllands arranged as short dorsa land paire dventrolat-
eral groups ;cephaiic glands absent; cerebral sensory organs
massive, posterior lryeaching beiow fron tof brain lobes;
cerebrai gangli alarge w,ith neither neurochord cells nor inner ff
neurilemma; later anlerve cords without acce$sory nerves;
foregu tdivisib liento oesophagus, stomach and pylorus,
intestin caalecum dorsa lsh,ort, without later adliverticu bluat
with pai rof long anterior pouches; blood system consisting of
simple vascular Ioop in head and three post-cerebr avelssels
not transversely linke dby pseudometameric connectives;
parenchyma extreme;y sparse; excretory system confined to
foregu tregion of body; eyes numerous, irregularl dyistr biuted;
sexes probabl yseparate.
Etymoiog) tT:he generi cname is formed by prefixin gthe name
nemertes with the Greek thallassio s{=marin e()masculine}. /1/
7)pespecies:thaMassionemertesloucocephalasp.nov.

Fig. 2. Thallassionemerte lseucocepha glean .et sp, nov, Drawing
TTrallassionemene sleucocephala sp. nov, of holotype ,viewed dorsall yb,ased on a colour photograp ohf the
Tyt) especjmen: Holotype immature female ,complete set of IMng specjmen, Scal ebar=5 mm,
mixed transverse and longitudi nsaelction$, 5 slides ,MTQ
G20023, Proximall cyonnective tissue dermis i s2-4 pm thick ,it souter
7)tp elocalit Tyu;rtle Beach {19016.2' 1S4,703.4'E), Cape surtace formin gcup-iike developments describe dfor many
Ferguson ,Queenslan dA,ustralia ,on rock covered with coral- other hoplonemertea nspecies.
line algae, exposed at low tide. Body wall musculature con$ists of outer circular and inner
Etymoiogy :The specific epithet, referring to the colour of the longitudina llayers (Fi g3.) ,respectively 3-5 pm and 15-20
head, i sa composite between the Greek words teukos (=white)pm thick .Both muscle layers extend to tip of head. Longitudi-
and kephale (= ahead). nal laye rnot anteriorly divide dand no trace of diagona lIayer
E>(tema tieature s:Body $lender, about 1 mm maximum distinguishe dT.here are also neither dorsoventr amluscle
width, 25 mm long ,tapering posterior ltyo end in sharply bundle$ in any part ot body, nor somatic muscles associated
pointe dtai l{Fi g2.) ,General colour dark greyish-brow np,aler with foregut,
posterior lByl,untl ryounded head white, median dorsa lregion Parenchymatous connective tissue$ moderately devel-
speckled with large number of irregularl dyj$tribut gerdey and oped, particula rarloyund alimentary tract.
black fleck sW.hite cephalic area sharply marked off from re- Probosci sapparatus: Probosci spore opens ventrally ,almost
mainder of body. at tip of head, from short, median ciliate dfurrow ,l tleads into
Body varal lm,usculature andparencbyma: Epidermis thickest thin-walledrhynchodaeumwhoseepitheliumisneitherciliated
in cerebral and cephalic regjons of body, where i tvaries from norglandular.
20-40 LLm in height b,ut becomes progressive lthyinner pos- Rhynchocoel extends to posterio trip of body. It swall
terior laynd in intestin arelgions is at most only 12-15 pm contains separate outer ci rcular and inner longitudin mauslcle
tal lT.hroughout it slengt hacidophjlic epidermal gland sare layer s(Fi g3.) ;their delicat selender appearance, in places
the most obvious, but larg egoble tcells also evident and re- only a single fibr ethick i,s most probabl ya consequence of
semble mucus-secreting type describe dby Norenburg < 1985), massive probosci sbein gin ful lreytracted positio nP.roboscis
NNIII-IE-leEcltreoncitcronic Library Service
TThhee ZZoooloolgiocaglical SSoociceityety ooff JJaapapnan
NemerteansfromAustralia 1263

RC
ax
-
Fig. 3. 7hallassionemette lseucocephal agen. et sp. nov. Transverse section of holotyp eto show the arrangement of the various body struc-
tures in the foreg urtegion, Scale bar;1O Opm, AP, anterior pouch of intestin aclaecum; CM, body wall circular muscle layer ;DE, dermis; EP,
epidermis; EX, excretory tubule; LM, body wall longrtudin amulscle layer ;LN, later anlerve cord; LV, Iater abllood vessel; PR, probosci$ ;RC,
rhynchocoel; ST, stomach.
inserti elnocate dimmediatel yin tron tef brain f,ormed by lon-
gitudin amulscle fibre sleadin ginwards from body wall longi-
tudinal muscle layer,
Extreme anterior porti ootn probosci asb,out 80-90 pm
long,consistsofepitheliumwhich containsno glandcells in-

ternal lbyounded by distin ccotnnective tissu elining ,a longi-
tudina lmuscle laye rin which rudimentary probosci snerves
can be distinguish eadn,d inner connective tissu eepithlium,
Epitheliu mof thi sregion increase sin thicknes sposteriorly,
commensurate with increas ein overall diameter ot proboscjs, s
Main anterior chamber of probosci sgraduall yincrease sin
diameter posterior tloy maximum of about 250 um, eventually
occupying some 90% of body diameter ,i,e ,r,eiative to size ot
specimen, itismassive (Figs3,5a},Epithelium,60 um or

more thick ,not developed into distinc tpapill aase, found in
many hoplonemertea nspecies, nor is i trichly provid ewdith
gland cells ,only irregularl yscattered glands being evident
throughou tit slength .Musculatur eof thi sproboscis region
cgoimtpruidseisn all aoyutee rr25 c-ir3c0ul aurm l tahyiecr 3k,(-F7 ip g4m). .aEcraocssh, o fa ntdi$ isnun elear lyoenr-s -
Fig.4. 77ialassionemertesteucocephalagen.etsp.nov.Transverse
separated from those adjacent by thi nbut distin ccotnnective
tissue coat, radial strands from inner connective tissue Iayer soef ctthioe nh tool sohtowype t.Shec astlr uecbtaurre=1 0of0 tphme. a ntPeCri,o rp rpoobrotisoc n oifcsi trhceu lparr omubsocslceis
and that separating the two probosci smuscle layer sextend- Iayer ;PE, proboscis epithelium, Pl, probosci sinner linin lgayer ;PL,
ing between longitudina lmuscle fibre sT,he 12 large probos- probosci slongitudin amluscle layer ;PN, probosc inserve; PS, pro-
cis nerves {Fi g3s, 4, 5a), linked by distinc tcircumferential boscis neural sheath.
neural layer ,situated in proxim ahlalf of probosc ilsongitudi-
nal muscle layer,
NII-Electronic Library Service
TThhee ZZoooloolgiocagliSoccalelty Society oofJfap anJapan
1264 R. Gibso nand P. Sundberg
Fig.5. Thallassionemertesieucocephalagen.etsp,

nov. (a >Transverse section through the anterior por-
tionoftheproboscis,showing the12distinncertves;

one of the nerves is indicate dby the arrow. (b )Ob-
lique section through the mtd-brain region, showing
theoesophagus(arrowed)belowtheventralcerebral
commissure. (c )Tran$verse section to show the ex-
treme posterio rpylonc portio nof the foregut lying
ventrally belov vthe intestin aeplithelium {d >Trans-
verse section through a later anlerve cord, anterior
pouch ot the intestin acalecum <indjca btye tdhe as-
terisk )and part of the excretory system; the arrows
indicat meyofibrillae running at the margin of the neu-
repil.(e)Transversesectionthroughthetipofthehead
to show the fronta lorgan. (f )Transverse section
throughtheventralpartoftheheadtoshow their-

regularly-shaped, acidophilic submuscular glands
'
swr (arrowe da)nd part of a cerebral sensory organ with
the dorsa land ventral regions flankin ga ciliate dca-
nal (asterisk e(dg) }T.ransverse section close to the
tip of the head tQ show one of the cephalic furrows;
note the differen icn ethe appearance of the furrow
epithelium (whi taseterisk) compared with the epider-
mis Black asterisks indicat tehe th ree groups of fron-
tal glands ,Sca[e bars :a=150 pm; b, f ,g=100 pm; c,
d=50 pm, e=25 pm.BR, brain lobe ;CO, cerebral sen-
soryorgan; EP,epidermis; EX,excretorytubuie;FO,

frontalorgan;IE,intestinalepithelium;LV,lateralblood
vessel, PE, probosci sepithelium; PR, proboscis ;PY,
pyloricportionofforegut;RD,rhynchodaeum.AlIpho-
tomicrograph sof sections of holotyp setained with the
Mallor ytnchrome method,
Style btulb region exhibits typica lmonostiliferous appear- Posteriorchamberofprobosciscompriseslowepithelium
ance, with no unusual features, Two accessory styletpouches dominated by basophilicglandcells, outer circular muscle

each contain two incompletetydeveloped reserve stylets,the layer,thinlongitudinalmuscle coat and innerlining.Posterior

central style tbasi sis 30 pm maximum diamete arnd about probosci sregion much smaller than anterior ,with maximum
50-60 pm long .Main stylet not evtdent in sections. overall diameter of about 65-70 pm.
NNIII-IE-leEcltreoncitcronic Library Service
TThhee ZZoooloolgiocagliSoccaielty Society ooff JJaapapnan
NemerteansfromAustralia 1265
Atimentat ycanafi Oesophagus opens from ventral margin of mid-dor$al ve$sel so compressed that i tis almost or even
rhynchodaeum jus itn fron tof probosci sinsertio Int. sepithe- completely indistinguish Oarbilgei .nof mid-dorsal blood ves-
lium not ciliated but contains scattered acidophilic gland cell$ sel, and whether or not i tforms vascular plug ,could not be
and is enclosed by thin but distin cctonnective tissue base- determined.
ment layer A,s j tpasses below ventral cerebral commissure it Throughout post-cerebr alregions of body the three
form$ slender tubula rcanal about 15 pm diameter (Fi g5b.). longitudj nballood vessels do not communicate with each other
Immediatel ybehind ventral commissure oesophagus until they meet at posterio srub-intestinal connective, Blood
enlarges and it sepithelium begins to develop cili aas i tmerges system of present species thus represents simple mono-
with anterior par tof stomach. Extreme anterior po rtio onf stom- stiliferous hoplonemertea npattern.
ach short, about 35-40 pm Iong ,and comprises unfolded Nervous s)tste mBr:ain lobes large relative to size of body
epithelium 20 pm or more thick which has no acidoph"ic (Fi g5.b) ,ventral lobes being somewhat larger than dorsal,
glands ,but for most of lengt hstomach typicall yhop[one- Thin but distin cctonnective tissue outer neurilemma invests
mertean with deeply folded epithelium (Fi 3g), which contains brain Iobes as a whole, but there is no inner neurilemma sepa-
both acidophilic and basophili gclan dcells. Stomach epithe- rating fibrou sand ganglioni nceural components. Dorsa lce-
Iium ,8-30 pm thic kdepending upon degree of foldin ge,n- rebralcommissure,1O-12pmthick,situatedanteriortothicker
closed by distin cctonnective tissu ebasement laye rbut has (30-3 5pm) ventral commissure. No evidence of neurochord
no somatic muscles. Overal llengt hof stomach, excluding cells in any part of brain L,atera nlerve cords throughou ttheir
pylor ircegion, about 230-250 pm. Towards rear of stomach lengt hcontain only single fibrou sneuropil (Fig 3s, 5d) ,i.e.,
it swall becomes progressive llyess folded t,hinne rand dors- there is no accessory later anelrve, Myofibril leaxteend through
oventrall ycompressed as i tleads int oshort pylori rcegion of inner latera lmargins of neuropil (Fi g5d.).
toregut w,hich is only about 70 pm long ,Pylori cepithelium Peripheralnervesupplywel1developed.Thicknervetrunk
comptetely lacks gland cells. Unu$uaHeature of pyloriclintes-leads from outer surface ot each dorsaliobe j,us btehind ven-
tina jlunct iios tnhat pyloru sleads directl yinto anterior intes- tra lcommissure, and curves anteroventrall yand outwards to
tine but with intesti ncaalecum that extends anteriorly above lead to cerebral senso ry organ. Severa lthic knerves also lead
pyloru s(Fi g5.c), not below as typica lof most hoplone- forward sfrom fron tof brain lobes int ohead, but ultimate fate
merteans. of most of these nerves not traced ;one of nerves from tron tof
Intestinalcaecumatfirstformswide,dorsoventrallycoema-ch dorsa llobe enters probosci sinserti otno form origin of
pressedtube,extendinganteriorlytoraboutonlyabout30pmprobosci sneural supply.
befor ebranchin gto form pairo flong ,later aalnterior pouches Flrontaloig afni,ont galtands and submuscular glandS :Single
(Fi g3). which reach forward sto end immediatel ybehind or front aolrgan, appearing as ciliate dpi tabout 35 gem in diam-
above brain .One of anterior pouches throughout it slength eter, opens neartjp of head (Fi g5e,). I tlead$ int oshort, thick-
runs immediatel aybove a later naelrve cord and ends at rear walled and ciliated chamberfrom which three groups of lightly
of brain o,ther for most of its length extends below nerve cord basophil ifcront agllands extend back (Fi g5g.), one median
but passes around it sinner margin close behind brain and dorsa lcap reaching for about 80-90 pm and two ventrolat-
ends dorsolatera lalboyve dorsa lbrai nlobe. Shorter of the eral bundles reaching somewhat furthe brack on either side
two anterior pouches some 220 pm long ,longe rabout 280 of rhynchodaeum. Cells of tront aolrgan chamber have dis-
pm, Neithe rintestin caalecum nor anterior pouche spossess tinctl yvacuolate appearance.
lateraldiverticula. SmaH,irregularly-shapedacidophilicsubmuscularglands
Intesti nfeorms thin-wall eddo,rsoventral lcoympressed occur only on ventral side of head (Fi g5.f) ,thei rdistribution
canal extending to posteri oernd of body below rhynchocoel. extending from anterior margin of cerebral sensory organs
I tdoes not possess later adliverticu bluat ,in it sapproximately back to leve lof probosci sinsertion.
posterior half ,does have shallow and irregula rdliystributed Typical ,lobular ,neutrophilic cephalic glands ,as found in
later aptouches extending between gonads, many hoplonemerteans, completely missing from present
Blood systern: Just behind probosci spore simple supra- specles.
rhynchodaeal vascular loop crosses head. Thick-walled, Sense oigans: Cerebra lsensory organs enormou$, almost
paired ,cephalic vessels run posterior loyne, close on either as larg eas brai nlobes .They fi lmluch of precerebr arlegion,
side of rhynchodaeum. A$ they pass cerebral sensory organs opening from short oblique ventrolateral ciliate fdurrow s(Fig.
vessels run in an inner median `groeve' along organs, effec- 5g) which, on each side of head, run from jus btehin dleve lof
tivel yflanke dabove and below by neural and glandular probosci spore obliquely backwards to later ablody margins,
components of cerebral organs, Behind probosci sinsertion about 70 pm behind ti pot head. Ciliat ecderebral canals, 20-
cephalic vessets enter cerebral ring, becoming small and 25 pm in diameter ,thick-wall etdh;ey lead obliquely inwards
inconspicuou sas they do so. Withi nbrai nregion course of and backuards to meet anterior glandula rregions of cerebral
vessels could not be traced ,but by beginnin gof stomach organs. These consist of large ,lobula rdorsa land ventral
region three longitudin ballood vessels evident, one running accumulations of vacuolar cells containing dark brown gran-
below each latera lnerve cord (Fi g3,} and one medially ules and orange-staining globule tsh,e two glandula rregions
between gut and rhynchocoel. In many parts of foregu tregion being medially separated by neural tissues of cerebral organs
NNIII-IE-leEcltreoncitcronic Library Service
TThhee ZZoooloolgiocagliSoccaielty Society ooff JJaapapnan
1266 R. Gibso nand P. Sundberg
(Fi g5.f) .Ciliat ecdanal then turns posterior atnyd continues anterior region of body ,outer portion reaches togethe rwith
back through middle of cerebral organs, it souter margin circularmusclelayerintotipofhead;precerebralseptumlack-
becoming enclosed by neuroganglionic tissues. Ciliated ing ,probosci sinserti ofnormed by fiber fsrom inne rportio onf
canals end in small, thin-walled chamberat about leve lof pro- longitudi nmaulsculature; head retractors related to outerand
bosci sinsertio Cne.rebra lorgans themselves about 90 pm in inner portio nof longitudin amulsculature; cerebral organs
diameter ,140-150 pm long, extending from short distance anterior to brain ;later anlerve cords with one fibrou score;
behind probosci spore back to below ventrolateral border sof musculature of probosci ssheath in separate layers ,rhy-
bral nlobes, nchocoel without dive rticula a,nd extending into posteri hoarlf
ln histologic aselctions indistin ceytes, 10-12 pm in of body; foregut opens into rhynchodaeum; intestina claecum
diameter ,each has appearance of a spherical accumulation present;blood-vascularsystemwithoutcephaliclacunaeand
of fin ebrown particl esusrrounding small central lumen, but extrace rebral vessels; excreto ry system present (;sexe psrob-
they lack typi cpiaglment cup ocellus construction of mo$t other ably separate)."
hoplonemerteans T,here are about 8 eyes each side of head, 7)pe species: CorTeanemerte sbioculatu (sCorre 1a9,58)
irregula rslcayttered from close to anterior ti pback to above
anterior regions of cerebral organs, Coneanemertes polyophthaima sp. nov,
E)(creto r$yystem: Excretor ysystem extends for most of fo- 7}tp especimens Holotype ,sex undetermined, series of trans-
regut length, anteriorly terminating alongside outer margins verse sections through the anterior body region, t3 slides,
of brai nlobes. Mostly consists ot single tubule ,about 1O pm MTQ G20024; paratype ,immature, transverse sectionS
in diameter ,extending along dorsolater ablody margin, close through anterior body and anterior intestina lregion, 5 slides,
above later anlerve cords and outside anterior pouches of MTQ G20025,
intestin caalecum <Fi g3). .As i tapproaches rear of brai nlobes DNA-sequence: Sequence for the 18S rDNA gene from
tubul ebecomes thjckeF walled and somewhat convoluted, a another specimen of thi sspecies collected irom the type lo-
slender efferent canal leadin gfrom this region to open at lat- cality at the same occasion as the holotyp eis deposite dwith
eral body surface via smail nephridiopore, Genbank (access inuombner AY 062924). A small pa rt ot the
Reproductiv esystem: Singl sepecimen sectioned an imma- specimen was placed in 70% ethanol for late rDNA extraction
ture female .Ovarie sscattered between intestin aplouches using the Dneasy ki t(QIAge nln,c. )A.n approximately 1900
in intestina blody regions. Whether sexes separate or not re- bp region of the 5' end of the 18S rDNA gene was amplified
mains unknown, by the Polymerase Chain Reactio (nPCR )based on eukary-
Systema tdiicscussio nT:he Hoplonemertea is a monophyl- otic specitic primer s(Medl ietn aL, 1988) using a PTC-1OOT",
etic group according to the phylogeneti acnalysis based on MJ Research lnc ,Amplification swere pertormed in 50 pl vol-
1 8S rDNA gene sequences in Sundberg etal. (i npress) T.heir ume of a solution containing 5-1 OO ng templat eDNA, MgC12
analysis inciude donly monostiliferous hoplonemerteans, 2.0 pM, each prime r{PCRA and PCRB) at 2.0 pM, each dNTP
of which there are at presen t97 recognised genera (Gibson, at 200 pM, 1 x reaction buffe r( 1O mM Tris-HCI pH 8.3, 50 mM
1995; Sundberg and Gibson ,1995; Rogers et al., 1996; KCI) )and 2 units of Taq polymerase (Perk Eilnmer) .The PCR
Chernyshev ,1998; Cranda lalnd Gibson, 1998; Kajihar aet cycling parameter sfor double stranded amplification were 2
aL, in press) in eight famili eTsh.ese familie sh,owever, are min. 30 sec. at 95eC for initi daelnaturatio nf,ollowe dby 60
poorl dyefine dand thei rdeflnitio inn sgenera lare not based cycles of 30 sec, at 950C, 30 sec, at 45 -580C, and 2 min, at
on explici tphylogene taniaclyses. We are therefor ereluctant 720C. The cycling was ended with 7 min. sequence extension
to place thi snew species in any taxon more inclusi vtehan the at 720C, Amplifi epdroduc twsere purifi beyd QIAquick PCR
genu$. We erect a new genus for this species because of the Purificatio nKit (QIAge lnnc.).
unusual arrangement of the alimentary canal where although Sequencing was pertorme udsing Cy5-labelle pdrimers
the pyloru sIeads direct liynt othe anterior intestine ,there is on an ALF-Express Automatic Sequencer (Pharmacja).
an intestin acalecum with long anterior pouches extending ThermoSequenase sequencing ki t(Amersha mw)as used
forvvar dasbove the pyloru sb,etween i tand the rhynchocoel for the sequencing reactions applying a two-step cycle 2
wail (se eAlimentar ycanal above), No comparable arrange- min, denaturati oant 96"C, foltowe dby 20 cycles of 30 sec, at
ment has been reported tor any previou smonostiliferous 950C and 40 sec. at the annealingtextensien temperatur eB.oth
hoplonemertea nand we regard this as a synapomorphy which strands were sequenced twice for the specimen.
allows 7haLLassionemert teos be identifie ads a monophyletic T)pe loaalj A4rnnong atgae T,unde Beach (1ge16. 124S7℃,.4'E>,
genus for which only the species thahassionemertes leuco- Cape Fenguson Q,ueensLand A,ustrel tona ,roc* exposed et bw tide.
cephala sp. nov. is so farknown. EtymotQgy :The specific epithet is a composite of the Greek
words poly (=many a)nd ophthaimos (=th eeye) and refers to
Genus Correanemertes Kirsteue r1,967 the numerous eyes which occur in this species,
Diagnosis: Kir$teue (r1 974: 1 59) gave the followin egmended AdditiOnalmateriatOnespecimenamongalgaecollectedfrom
diagnosi fsor the genus Correanemertes ":Body shape resem- the fla otf Rib Reef ( 18028.8'S ,146052.4]E) north-east of the
blin gthat of Amphiporus species; body wall musculature with- Palm lsland sd,ept habout 2 m,
out diagonaHiber layer ,longitudi nmaulscle laye rdivide idn E)rtema lteature sU:p to 80 mm long ,1-2 mm in maxjmum
NNIII-IE-leEcltreoncitcronic Library Service
TThhee ZZoooloolgiocaglical SSoociceityety ooff JJaapapnan
NemerteansfromAustralia 1267
OL
oc
5t t:,. axED
t:t: ' r-:- PRDE
dt-t
--J-J
:-

tL
BR
IC
B
-
Fig.7. CorTeanemertespolyophthalmasp.nov.Transversesection

through the posteri coerrebral region of holotype to show the divided
body wall longitudi mnuasclulature and arrangement of various body
structures, Scaie bar=250 pm. BR, brain lobe; DE, dermis ;ED, effer-
ent canal of excretory system; EP, epidermis; EX, excretory tubule;
IC, body wall inner circular muscle layer ;IL ,body vvall inner longitudi-
nai muscle Iayer ;OC, body wall outercircular muscle layer ;OE, oe-
sophagus; OL, body wall outer longitudi nmauslcle layer P:R, proboscis.
cerebral septum as define dby Kirsteue r(1974 )Bo.th inner
and outer portion ost the longitudi nmauslcie laye rcontinue in
tron tot the brain to torm cephalic retractor muscle bundles .1n
posteri foorregut region of body longitudina lmuscle layer up
to 60 pm or more thick o,uter circula r1O-12 pm.
In stomach region an incomplet eiayer of inner circular
musclefibresextendstromclosetothemid-dorsal1ine,around
Flg.6. Coneanemertespotyophthatmasp.nov.(A)Drawingofcom- the rhynchocoel and later aalnd ventrolaterat foregu tborders
piet $epecimen in dorsal aspect based on a colour photograp ohf the almost to the mid-ventral line T.hese inner circular muscles
leyievsi,n aSgnciamlale.b a(rB= )1EOnlmamrg(ermeenft eortt sohAeaodn ltyo) .show the distribu toif othne are separated from the rhynchocoel wall musculature by a thin
but distin ccotnnective tissu emembrane, Effectiv etlhye$e in-
nerciroularmusclesappeartheequivalentofdorsoventralmuscles.
width, body long and slender with a pointe pdosterio tri p(Fig. Cephati rcetractor muscles derive dfrom outer portio nof
6A), with a well demarcated ,tlatten eadnd rather pointe dhead divide dbody wall Iongitudina lmusculature, in fron tof brain
(Fi g6,B), The overali colour is a uniform chocolate brown. separating off to extend forwards as discret feibr ebundles
Just at the rear of the head a slight reddish tinge indicate tshe enclosed by connective tissu emembranes, i,e, f,asciculated.
positio onf the brai nlobes .On either side of the head there Parenchyrnatousconnectivetissuesfairlywelldeveloped
are about 15-20 variably sized eyes arranged in a single wide around the various body structures,
and scattered row. Proboscis apparatus: Probosci spore ventral, $ubterminal.
Body ivaza musculatute and parenchyma: CIose behind brain Rhynchodaeal walt thin ,without it sown musculature but
epidermis 45 pm or more thick ,internal bloyrdered by di$tinct flanke ddorsall aynd lateral blyy longitudi nmauslcle bundles
connective tissu edermis up to 8-10 pm across. Body wall which appear to represent cephalic retractors derive dfrom
musculature consisting of outer circular layer m,ostly 6-7 pm the inner portio nof the divide dlongitudin amulscles. Inner
thick a,nd inner longitudi nlaalye r30-35 pm deep. No diago- and outer retractor sare separated by parenchymatous con-
nal muscle laye revident. Just behind brain the Iongitudinal nective tissue behind the cephalic glands.
layer divide sint oinner and outer portion swhich are sepa- Rhynchocoel wall with separate circular and longitudinal
rated by connective tissue (Fi g7s, 8a), The inner fibre asione muscle layers. Extends almost to rear of body, Large num-
contribute to the probosci sinsertio in.,e ,t,here is thus no pre- bers of gregarinifo rpmarasit etasirl yabundant in the rhy-
NNIII-IE-leEcltreoncitcronic Library Service
TThhee ZZoooloolgiocagliSoccaielty Society ooff JJaapapnan
1268 R.Gibsonand P,Sundberg

/. ,÷.
pt,xaj,ur
.tt
1?
t'
':i'E':･'#/･'#'r
･'l

iltt.
/.t'v-1
-
fffs
..ptf
g
- -.
Fig. S. Correanemertes potyophthatma sp, nov. (a )Transverse section through part of the body wall in the pQsterio rbrain region to show
bundles of the inner longitudin amulscle layer (indica btye adrrows), an excretory tubule and the disti nocutter neurilemma enclosing the brain
lobes ,(b )Transverse section through the head, showing the size of the probosci sand mid-dorsal bundle of cephalic glands; asterisks indicate
the cerebral senso ry organs. (c )Transverse section through the cerebral region jus itn fron tof the ventral ce rebral commissure to show the inner
and outer portion sof the divide dbody wall longitudin amulscle layer and the oesophagus (arrowe (dd) ).Part Qf the dorsolater aclephalic region
intransversesectiontoshow threeoftheindependentducts(astensktehdro)ughwhich thecephalJcglandsdischargetothebodysurface.(e}

Transversesectionthroughtheanteriorheadtoshowoneofthepigment-cupocelli(arrowed}.{f)Transversesectionthroughacerebralsensory
organ, Scale bars: a, d, e=1 OO -m; b, c=250 um;f=50 pm. BR, brain lobe; CG, cephaiic gland sC;O, cerebral sensory organ; EP, epjdermis; EX,
excretory tubule ;PR, proboscis A.l lphotomicrograph sof sections of holotyp estained with the Mallor ytrichrom emethod.
nchocoel of one additional specimen from type locali ttyh,ese nantly basophilic ,epithelium up to 25-30 pm thick ,then thick
parasite sbeing up to about 25 pm long and 7-8 pm wide, outer connective tissue zone which extends peripherally
with a single nucleus about mid-body level. to form core of epithelial papill aoeut,er circula rmuscle layer
Main, anterior, chamber of probosci scomprises richly about 15 pm maximum thicknes sm,iddle connective tissue
glandularepitheliumdevelopedintopapillae,glandspredomi- laye rwhich is moderately thic kand distin citn,nerlongitudinal
NNIII-IE-leEcltreoncitcronic Library Service