Some earthworms from the wet tropics and from Bunya Mountains, Queensland (Megascolecidae: Oligochaeta)

24 Pages·1994·7.8 MB·English

by  B G M Jamieson

SOME EARTHWORMS FROMTHE WETTROPICS AND FROM BUNYAMOUN-
TAINS, QUEENSLAND(MEGASCOLECIDAE: OLiGOCHAETA
>
B.G.MJAMIESON
Jamicson. B.G.M* 1994 12 01; Some earthworms from the Wet Tropics and from Bunva
Mountains, Queensland (Megascolccidae: Oligochaela). Memoir* of the Queensland
Muscum$7(\y 157-180. Brisbane. ISSN0079-883?
Asystematicstudy,supportedbyacladisticanalysis,indicatesthatNorthQueenslandspecies
of Diporochaeta warrant separate generic status, as Tenl$8WQlk6rlus gen. nov
Ftetcherodrilus gen. nov. represents an apomorphic clade of this assemblage which is,
nevertheless, retained as a separate genus. The Bunya Mountains isolate ofDiporochaeta
merits separate generic status, as Hiatidrilus bunya, and Cryptodrilus semicinctus, horn
Grafton, on the Clarence River, New South Wales, is congeneric with it. The residue of
Diporochaeta is a paraphylelic and polyphyletic entity requiring further resolution. The
cladisticanalysisalsoinvestigateswiderrelationshipsoftheabovegenerawiththeremainder
of the Perionychini. In the analysis the type-species of Diporochaeta, D. intermedia, b
separatedbyothergenerafromthoseDiporochaetaspecieshereassignedtoTerrisxwalkerius
and, indeed, from all other included Diporochaeta spccic9. The analysis suggests that
Perionychella, currently subsumed in Diporochaeta, is the sister-taxon ot the Ter-
risswalkerius-Fletcherodrilus assemblage. The phylogram is not inconsistent with the
generic status of the remaining perionychin genera Graliophilus, Heteroporodrilus,
Paraplutellus,Perionychella*Pinguidritus,Plutellus,Pseudoperichaeta*Stmsia,andWood-
ward'tellabutsignificantautapomorphiesrenmintobeestablishedforsomeofthese. Within
lerrisswalkeriusxhcmostplcsiomorphic species appears tobe 7*. biounti. 7*. atavius, is the
most derived species and forms the sister-laxon ofFletcherodrilus, Pairs ofsister species
are; T. grandis and T. terraereginae, T. mitiaamiltaa and T. mcdonaldi sp. nov., T. athcr-
tonensisandT. oculatus, T. kurandaand T. canaliculatus,and T. nashiand T. libersp. nov.
A monophyletic, though weakly defined, species group contains T. uthertonensix, T.
oculatus* T. canaliculatus, T kuranda* T,crateris* T. nasfuand T, liher. The new species7*.
covaccvichaeandFletcherodrilusmenurusare alsodescribedO'/Vrm.vvvr//A-/rr/w,T%en.nov.
Hiatidrilusgen.nov,, Fletcherodrilus, Diporoclwta, Perionychini, chidisrirs
M
B.G Jamicson, ZoologvDepartment, UniversityofQueensland, Brisbane Queensland
,
4072, Australia; 28August IV94.
Diporochaeta was erected by Bcddard (1890) ThreenewspeciesareaddedtoTerrisswalkerius.
with,asustype-species,theNewZeaJandspecies The status and phylogcnetic affinities ot
Diporochaeta i-Perichaeta) intermedia Bed- Fletcheroxirilus will also be investigated,
dard, 1889 Prior to the present work, Jamicson (1976b) observed lhat little other than
Diporochaeta contained 78 Australian species, the unpaired genital pores warranted separation
Thesespecies had been initially placed,oncrcc- of Fletcherodhtus from Diporochaeta, that in-
tion, in some eight genera in addition to elusioninthelattergenusasasubgenuswouldbe
Diporochaeta. Of these, five (Cryptodrilus, sustainable, and thai Fletcherodrilus may be
Megascolex* Megascolides, Perionyx. Pluielius) regarded as the apomorph sister-group of
are still recognised as distinct genera, two Diporochaeta, A new species is added lo
{Perionychella and Vesiculodrilus) have been Fletcherodrilus. Erection ofa distinct genus for
subsumed in Diporochaeta (see Jamieson, north Queensland species of Diporochaeta and
1976b) and one (Pertchaeta) in Me^asolex (see the apomorphic status of Fletcherodrilus have
Miehaelsen, 1900). Here evidence will be been independently suggested by Dyne & Wal-
prcsented, in a cladistic analysis, in support of lace (1994).
removal on 'intuitive' grounds, of 16 species The status of the predominantly Western
from North Queensland from Diporochaeta and Australian genus GraliophilusJamicson. 1971b,
their transfer lo a new genus, Terrisswalkerius, was rendered uncertain by addition of North
and for transfer of two species to a further new Queensland species to Dij>orockaeta by
genus, Hiatidrilus. The number of Australian Jamicson (1976b) which blurred the distinction
species of Diporochaeta is thus reduced to 60. between the two genera. Graliophilus was
15R MEMOIRS OFTHEQUEENSLANDMUSEUM
retained by Jamieson (1974a) on the grounds, female, male and spermathecal porcs absent. A
t
inter alia that in Western Australian species the pair of combined pores of v*sa deferentia and
number ofspcrmathccac was less (2 or 3 pairs) tubular or lubuloracemose prostates on XVIH.
thanthefivepairsusual in Diporochaetaandthat Last hearts in XII orXIII. Gizzard in V or VI (or
the avesiculate nephndia and totally lumbriein VII?)(welldeveloped);intestinalcaecaandtyph-
conditionofthesciae were constant whereas the losole absent. Extramural calciferous glandsab-
combination of these characters was known in sent Nephridia stomale exonephric
.
only one species of Diporochaeta from eastern holonephndia with or without bladders; their
Australia (Tasmania). It was concluded, how- pores in straight or sinuous lines but never with
ever, thaithe low pheneiieresemblance between regular alternation. Spcnnatheeac 1 to 5 pairs.
DiporochaetaandGratiophilusdemonstrated by rarely unpaired midventral, pre-testicular, divcr-
Wallace(1972)andtheverydistinctmorphology ticulate;diverticulumusually single, unilorulate;
ofsome Gratiophilus species suggested that this rarely double, rarely multiloculate.
genus should be retained. The present study ex-
aminescladistic relationshipsofGraiiophilus to
DESCRIPTION
Dthiepotryopceh-asepteacifecslato.fanaldl innculmueddesginentehreaanoaflystihse Terrestrial worms. Fonm circularin cross se. -
Perionyehini, vifc, ihc 13 genera Diporochucia* slioovncnotrrallelsys;caonmumsontlerymisnlailg.htlWyitdehproerssweidthdoourt-
Fleteherodriltis,Graiiophilus.Hcteroporodrilus*
pigmentation. Prosiomium proepilobous to
HtaJiiinhiA, Paraplutellus Perionychtlla* Pin*
y tanylobous. Body usually with a dorsal groove
guidrilus. Pluteilus. Pseudoperichaeta. Simsia*
Tcrrisswalkerius, and Woodwardiella. (canalicula) in part or the whole of its length.
Dorsal pores present; in 3/4 - 6/7, usually 5/6.
Retention ofPerionychetla, for forms lacking
Setae pcrichactinc throughout. Nephridia in
ncphridial bladders (and possibly with racemose
single lateral series throughout, or in irregularly
prostates, as inthetype-species) as a subgenusof
Diptfrothaetadeservesconsiderationbutwasnot sinuous series, varying from far dorsally to fai
ventrally but never showing regular alternation,
upheld (Jamieson. 1976b) as the distinction be-
tween an avesiculate and a vesiculate group of Posterior limit ofthe clitellum shortly anteriorto
species had become increasingly blurred, Thus the male pores or including these. A pairofmale
poreson XVUl,eaeh pore thecombinedopening
Tasmanian Difwrochaeta monorti has avesicu-
of a prostate gland and the corresponding vosa
late and, typically, vesiculate individuals
D defcrentia. Female pores paired on XIV. Sper-
(Jamicson, 1974a). grandishasvesiculate and
mathecal pores to 5 pairs, rarely unpaired mid-
avesiculate populations, oreven segments in the I
sameindividualandinDt terracregimie{botharc vreanrterlayl,shboerttlwyeebnehiinntdertsheegimnetntearlsefgumrernotws4/5-8/9,
Queensland species) ncphridial ducts are transi-
tional between the twoconditions. Perhaps more Dorsal bloodvesselsingle,continuousontoihc
significantly, in an unpublished taxonometrie pharynx; last hearts in XII or. rarely, XIII; hearts
(numerical) study vesiculate and iivesiculalc in X posteriorly latero-ocsophagcal, each receiv-
species formed mixed groups, especially where ingaconnective fromthesupra-oesophageal ves-
all ncphridial characters are omitted It is sel and a lesser connective (sometimes absent'/)
noteworthy that Michaelscn. who ignored the fromthedorsal vessel. Supra-ocsophagcal vessel
presence or absence of bladders, considered the restricted to the post-gi/.zard oesophagus,
avesiculate D. atavia to be merely a form ofthe moderately to well developed. Subneural blood
vesicularD athertonensis,adecision indicating vessel nbseni Gizzard in V or, more frequently,
the closeness of the entities with the antithetic VI (or VII9) (well developed). Extramural cal-
conditions, notwithstanding the view (Jamieson, ciferous glands not developed but oesophagus
1976b)that they are distinct species. vascularized and often internally rugose in some
Unless otherwise stated material studied is .cuts from VIII to XVI. Intestine commenc-
Imi ivjj in ihcQueensland Museum (QM). ing in XVI to XIX, most frequently XVII; intes-
tinal caeca and ryphlosole absent. Nephndia
simple, exonephric holoncphridia throughout;
TerrLss*walkeriusgcn. nov.
tufted nephridia absent; bladders usually present,
with or without diverticula; exceptionally (D.
DlACWNJS gt&todiS, pail.) avesiculate in the forchody and
Setae numerous (>I4> per segment. Genital vesiculate inthehindbody. Holandric(testesin X
markings other than porophores bearing the and XI). seminal vesiclestwo to fourpairs, in IX
.
NEWQUEENSLANDEARTHWORMS IS9
to XII, exceptionally (7". terraereginae) in VIII Spermathecalporesmedian toa lines. Spermathc-
(in addition to IX and XII), and often with pseu- cal ductlongand slenderthough shorterthan the
dovesiclesinXIII. Metagynous(ovariesandfun- ampulla. Lastheartsin XH 3
ncls in XIII). Prostates tubular or 3. Spermathecal pores of8/9c,0.02bodycircum-
tubuloracemose,neverracemose.Penialsetaeab- ferenceapart. Male poresc.0.03 bodycircum-
sent. Spermathecae diverticulate; diverticulum ferenceapart.Seminal vesicles in IXandXII.
usually single, uniloculate; rarelydoubleormul- Nephridialbladderspresent T. arm*as
tiloculate. Spermathecalporesof8/9c. 0.06bodycircum
ferenceapart. Male poresc.0.1 bodycircum-
ferenceapart. Seminal vesiclesinXIandXII (Or
Distribution absent) Nephridial bladdersabsent
_ ;
TheWetTropicsofeasternNorthQueensland, T.phalavrwt
from the Paluma Range, 19S., near Townsville, 4. Spermathecal poresin4intersegments,5/6-8/9or
north to the Upper Endeavour River shortly 4/5-7/8,pairedorsingle S
below 15°S. Spermathecal pores 3 pairsorfewer 9
TypeSpecies Spermathecal pores in 4 intersegments
Terrisswalkerius (=-Perichaeta) canaliculars
(Fletcher, 1887a); here selected because its 5.Spermalheeal poresin 5/6-8/9, unpaired, mid-
anatomy is well known and it is the most widely ventral T. mcdonaldisp. mo
distributed speciesofthegenus andthereforethe Spermathecal pores4pairs, in4/5-7/8or5/6-8rV
most likely to be available for molecular and 6
otherstudies.
6. Spermathecal pores4 pairs,in4/5-7/8
T. terrareginar
ETYMOLOGY Spermathecalpores4pairs,in5/6-8/9 7
Named in memory of Terry Walker Gender 7. Male pores mediantoa lines, theirpapillaecon-
masculine. tiguousin the midline. Nephridialbladders ab-
sent T. millaamiiiaa
IncludedSpecies Male poresbetween selai linesa andJ, their
papillae well separated in the midline.
T. atavius(Michaelsen, 1916); 7". athertonensis
Nephridialbladderspresent 8
(Michaelsen, 1916); T. barronensis (Fletcher,
c1a8nKa6lbi)c;ulaTt.usM(Foluetnctheir,(1J8a8m7iae)s;onT.,co1v9a7c6ebv)i;chaLe 8. PrNoespthormOipOurmetaaninylaobSotruasi,ghwtitsheriweisdoendoerascahlstiodnegu.e..
met
sp. nov.; T. cratcris (Jamieson, 1976b); T. eric't T.
Prostomiumcpilobousorprocpilobous; with a
(Michaelsen, 1916); T. grandis (Spencer, 1900); deepmiddorsalgroovewhichcontinues to the
T. kuranda (Jamieson, 1976b); T. liber sp. nov.; hindmarginoftheperistomium. Nephropores in
T. nicdonaldisp. nov.;T.millaamillaa(Jamieson, an irregularly sinuous series, varying from far
1976b); T. montislewisi (Jamieson. 1976b); 7". dorsaliy to far ventrally,oneach side
nashi(Jamieson, 1976b); T. oculatus(Jamieson, T. kuratuin
1976b); T. phalacrus (Michaelsen, 1916); 7". 9. Spermathecal pores3 pairs, in4/5-6/7 orin or
raveni (Jamieson, 1976b); T. terrareginae shortly behind6/7-8/9 10
(Fletcher, 1890). Spermathecal pores I or 2pairs, in someof5/(*
K/9 16
KEYTO SPECIES OF TERRISSWALKERIUS
Spermathecal pores 3 pairs
1. Spermathecal pores5 pairs, in orshortlybehind
intersegmental furrows4/5-8/9 ,2 10. Spermathecal pores3 pairs, in4/5-6/7 . . . . 1J
Spermathecal pores 1 to4pairs (rarelyunpaired, Spermathecal pores 3pairs, inorshortlybehind
midventral), inorshortly behind someorall of 6/7-8/9 13
intersegmental furrows5/6-8/9 4
1 1. Spermathecal diverticulum longand tortuous.
Spermathecal poresin thevicinity ofJ to elines
Spermathecal pores5 pairs 12
Spermathecal diverticulum shortandclavalc.
2.Spermathecal poresin alines. Spermalheealduct Spermathecalporesbetweena andb lines . .
veryshort. Lasthearts in XIII T grandis T. montislewisi (part.)
"
160 MEMOIRSOFTHEQUEENSLANDMUSEUM
12. Spermathecaldiverticulumlongerthanampulla
plusduct (Spermathecalporesindlines) . . .
T. barronensis
Spermathecaldiverticulumshorterthanampulla
plusduct. (Spermathecalporesind-elines) .
71 raveni
13. Maleporesveryclosetogether,almostcon-
tiguous midventrally,onacommonpapilla
. ,
T.athertonensis
Maleporeswell separated, betweensetallinesb
and/,on a pairofpapillae 14
14.Spermathecaldiverticulumsimple, shortly
clavate 15
Spermathecaldiverticulumcomposite,consist-
ingofasmany as4 parallelconjoinedtubes
with terminal, knoblikeseminalchambers;ap-
proximatelyaslongas thespermathec ....
T. oculatus
15. Spermathecal pores varying fromsetal lines3to
5. Maleporesbetween setal lines4to7. Prostate
glandsrestricted toXVU1 . . . T. canaliculatus
Spermathecalporesinblines. Maleporesinab.
XVW-
Prostateglandsconcealing thegut, in r^phroport-
XXI T. covacevichaesp.nov. XUI
temalepors
Spermathecal pores 2 pairs
16.Spermathecalpores2pairs, in5/6and6/7 ,. . -4J
T. montisiemsi(part) <vii — —
Spermathecalpores 1 pair, in 6/7 or7/8or8/9 XVll|
17 1 m0*r-*~'.
« _^-~"~
':'III
-—
Spermathecal pores 1 pair
mm
1
17.Spermathecalpores 1 pair, in8/9. Spermathecal
diverticulumlongand tortuous 18
FIG. 1. Terrisswalkerius athertonensis (Michaelsen,
Spermathecal pores I pair, in6/7 or7/8 . . . 19 1916). Mt Lewis, QMG9210. Forebody and clitellar
18. Spermathecalporesinsetal linesc tod. .... region, latcroventral view.
T. nashi
TypeLocality
Spermathecal poreswellmedianofsetal linesa 17B16,S.145B29,E., Atherton. 15P39*-
T. libersp.nov.
27°52,S.145°15,E.-15312,E., Cedar Creek. All
19. Spermathecal pores I pairin6/7 . . . T. crateris Queensland
Spermathecal poresin7/8 T. blounti
MaterialExamined
Terrisswalkeriusathertonensis Syntypes: Zoology Museum Hamburg HMV8481
(Michaelsen, 1916)comb. nov. (re-examined), HMV8482 (now represented by soil
(Figsl;2A,B) debrisonly). Stockholm MuseumNHRS1245.
NewRecord: tt^M'S. 145°17'E., Mt Lewis, in rain-
Perionyx (Diporochaeta) athertonensis Michaelsen, forest,F.LittleandQueenslandMuseumParty,7Nov
1975, a single, previously dissected, clitellate
1916: 7-9,pLl,fig.7. specimen, QMG9210. (This material was correctly
Diporochaeta athertonensis; Jamieson, 1971c: 83; identifiedbyR. RavenasDiporochaetaathertonensis,
Jamieson, 1976b:14, figs 1, 9a, table 1. thefirstsightingsinceMichaelsen*s 1916description);
Perionychella (Perionychella) athertonensis; 2clitellatespecimens, rainforest, 6kms fromMalanda
Jamieson, 1974a: 221. FallsEnvironmentalPark,onroadtoAtherton,B.G.M.
.
NKWQUEENSLANDEARTHWORMS
161
Jamieson. 12. Dec 1994. QMG211454. All in
Queensland. prostate,gland
Description
(Thefollowingaccountcombinesdescriptions
ofall knownmaterial, The single specimen from
Ml Lewisis a posterior amputee).
Length 36-68mm, width 3-5mm. segments
A
98-108. Pigmented violet brown, little colour —spermathecal
remaining in Mi Lewis specimen, Tanylobous. jnh" duct
Body canaliculate throughout First dorsal pore
5/6. Setal circlet regularly interrupted ventrally, apermathecal *& \^spermathecMl
dorsallyirregularly;alinesregularthroughout, h diverticulum %^-J ampulla
lines regular throughout or for much of body mm
B
length; c lines less regular; in mid- and hind- i
body, aa = 1.5-2 ah; dd 1.3-2.3 yz, in anterior
segmentsmuchlarger.Intersetalspacingdorsally
irregular and in general wider than vcntrally, FI1G9.16)2.. Mttl-rLrteswiiwsa,lkQerMiGus92at1h0e.rtoAn,enrsiigsht(pMriocshtaateeK;enB.,
except in forebody. Seial numbers (MiehaelsenI; righi posteriorspennatheca.
10/11, 15/V, 22/VHI, 24/X, 36/XIX, 35/XXVI,
36/LXX, 38/LXXI; 23 in XII,40 in LXXVI (Mt t\ midlength,two limbs parallel, ectal limb wick
Lewis). Nephropores (see also internal descrip- andoften berU nearpore (V-shaped bladders). In
tion) visible as a straight series on each side, anterior intestinal region ental limb has become
immediatelyventralofsctallines 10onclitellum. so narrow that it can scarcelybe considered part
Clitellum annular, XIIM/2XVITL XVin; inter ofbladderandissocloselyadherenttoectallimb
segmental furrows, setae, ncphridial and dorsal that bend has spurious appearance of a diver*
pores clearly visible or (Mt Lewis) dorsal pores tieulurn. This form persists to caudal extremity
occluded. Male pores scarcely visible, very near andalthoughenlallimbwidensslightly ilremains
ventral midline. 0.1 1 - 0.2mm apart (Mt Lewis) much narrower than ectal limb. In Mt Lewis
inatransversely oval midventralglandularfield. specimen ncphridial postscptal portions com-
FemaleporesfMt Lewis)apairofminutepoints, mence in II and a large preseptal funnel is
shortly median ofsetae a, neareranteriorborder demonstrable forihoseof111 and lesscertainly 11
than they arc to setal arc of XIV. Spermathccal ductsofthe first three pairs are dilated but ques-
pores, hardly externally recognizable, 3 pairs, in tionably to be considered bladders; tbereaiLJi
6/7-8/9- very nearventral midline,0.05-0.07mm hey form definite bladders; those of V are V
I
apart, or(Mt Lewis) 0.08mm ami about one half shaped and those of anterior intestinal region
aaapart. Genital markingsabsent. similarly conform with syntypes. Two pairs of
Septa from 6/7 well developed, 6/7-8/9 thin, iridescent sperm funnels in X and XI; racemose
9/10-12/13 slightly thickened,theremainderthin seminal vesiclesin IXand XII (MtLewis). Pros
(type material). Dorsal blood vessel single; last talestubular,almostconfined toXVIII (syntypes
hearts in XII; large and laiero-oesophagea! in and Mt Lewis); central lumen (Miehaelsen)
X-XIL Gizzard large, in VI or (Mt Lewis) ap- simple, about one eighth widih ofgland; wall of
parently V. Oesophagus inXI1-XIV with folded, gland with two layersofglandularcells overlain
vascularized walls or (Mt Lewis) X-XVI vas- by peritoneum, cells of inner layer in groups
cularized; but lacking extramural calciferous around apparently lumenle^s axes. Duct (all
glands and with no development of largepaired specimens) well defined from gland; about one
blood vessels. Intestinecommencing abruptly in fourth as long, much thinner and muscular. The
XVI or (Mt Lewis) XV11; lyphlosole and caeca two vasa deferentia of a side running close
absent. Ncphridial bladders (not mentioned by togetherandjoiningprostateductatitsmidlength
Miehaelsen),inthe syntypes,widelubes, thosein before (Miehaelsen)passingdistally to fuse with
IIconvolutedanddischarging in:. lines,succeed- its lumen shortly before distal end. Penial setae
ingbladdersprogressivelymoreventralsothatby absent. Ovarieswideandbushy,oviducalfunnels
XVIIi,he8yonarceliitnelsletuamlalnidne1s27c.auNdeaplhlryo.pBoyreVsviensi9cloens 3larpgaier;so:vaimdpuuctlslashsomrotoatnhd, sstarcaliigkhct.orSpoevromiad,thdeucealcs
takeformofa tube with an abrupt lateralbend at sharply demarcated, twothirds widlh ofampulla
162 MEMOIRS OFTHEQUEENSLANDMUSEUM
and somewhat shorter or (Mt Lewis) ap-
proximately as long; a slender clavate to
digitiform diverticulum, with simple lumen,
entering duct inbody wall or(Mt Lewis)nearits
base and shorterthan ampulla plus duct.
Remarks
The new material, from Mt Lewis, agrees
closely with the type material in most respects,
indicating conspecificity. The difference of one
segment recorded in the position of the gizzard
and intestinal origin between the type and Mt
Lewis material is not, however, usual in-
traspecifically and requires confirmation from
betterpreserved material ifavailable.
Terrisswalkeriuscovacevkhae sp. nov.
(Figs3;4A.B)
TypeLocality
17°33,S.145°34,E., Mt Fisher. N.E.
Queensland, J. Covacevich, V.E. Davies, R.
Raven, 23.iv. 1978.
t=f7l,3<r..nr-
MaterialExamined dile«lum
Holotype: QMG211447exQMGH1991.
Paratype:QMG21 1448 ex QMGH1991.
Description
Length (holotype, H, is a posterior amputee), iial&poio
125mm (paratype, P), width 5.0mm (P), 6.2mm
(H); segments 222 (P). Body notdepressed. Pig-
mentless,exceptingpalebrownishgreyclitellum,
in alcohol. Prostomium tanylobous; its anterior
half, and ventral surface bisected but body only
weaklv canaliculate; peristomium short. First
dorsafpore 5/6 (P), 6/7 (H).Setae 28 in XU, 47 FIhoGl.oty3.pe.TerQrMiGss2w1al1k4e4ri7u.s cFoovraecbeovdtcyhuaendsp.clintcolvl.a,r
caudally;aaand, mostly, zzconspicuousbreaks; region, ventral view.
alines regular, zlines irregular, in hindbody and
with some irregularity in first few segments; in present between porophoresand behind setal arc
XII, aa:ab;bc:cd:yz:zz= 2.2:1.0:1.2:1.0:1.0:6.6; ofXVII. Accessory genital markings absent (H,
setae a and b absent from XV111. Nephropores P). Female pores small transverse slits immedi-
clearly visible a short distance behind clitellum ately anteromedian to setae a of XIV (P). Sper-
near anterior borders oftheir segments, laterally mathecal pores 3 pairs of simple transverse slits
situated, in straight longitudinal lines in setal atanteriorborders ofVI, VII and VIII. inblines,
lines 1 1 or near this; not observed (though sometimeshidden in anteriorintersegmental fur-
present) in forebody. Clitellum annular, XIV- row (H, P).
XVII but dorsolateral^ from anterior 1/3X111; Thickest septa 10/11-12/13, strongly thick-
dorsal pores occluded in 14/15 to 16/17 where ened.Dorsalbloodvesselsingle,continuousonto
theseintersegmentsareweaker. Maleporesapair pharynx. Lasthearts in XII; those in X-XII large
ofminute pores, in ab, each with a whitish rim, and arising from supra-ocsophageal without
on large porophores which expand XVIII recognizableconnectiontodorsal vessel;thosein
anteriorly and posteriorly and are somewhat VTI-IX slender, and dorsoventral only. Gizzard
medianly inclined so that male pores seemto be large, barrel-shaped with anterior rim, and
near their median borders; a deep depression strongly muscular, in VI; septa 6/7 and 7/8
,
NEWQUEENSLANDEARTHWORMS
163
regionofforebodyductisbentonitselfanddistal
te duct limb is dilated as an elongate bladder which ex-
tends at bend as a very short diverticulum; in
intestinal region duct joins bladder near its
equator and an elongate diverticulum extends
laterallyfrombladder;egressofbladdersatbody
wall is more lateral in intestinal region than fur-
ther anteriorly (H, P). Holandric; testes (?) and
fairly small scarcely iridescent funnels in X and
rostate gland XI; large racemose seminal vesicles with several
large loculi which are themselves further sub-
mm divided, in IX and XII (H. P); those in DC with a
1 median wing-like or longue-like portion which
extends onto gizzard (H) or this extension not
evident, although investing gizzard (P). Ovaics,
spermathecal
eachalarge plateofoocytes, andsmall funnel, in
diverticulu spermathecal X1IL Prostates with massive glands enwrapping
duel
and concealing alimentary canal, in XVIII-XXI;
deeply incised by septa but otherwise smooth
spermathecal
surfaced and not notably subdivided. Sper-
ampulla
malhecae3pairs,eachwithalarge,ovoidampul-
laandawelldevelopedonlyslightlyshorterduct;
duct bears, at its ectal fourth, a subspheroidah
almost sessile, inseminated diverticulum. By in-
ternalseptationtherearenospermathecaeinVIII,
segmentanteriortothatcontaininganteriorsemi-
nal vesicles, last pair of spermathecae being in
VII.However,externalexaminationconfirmsthe
posteriorpairofspermathecal pores anteriorly in
B i mm VIII. The internal septation is therefore
anomalous, disagreeing withexternal segmenta-
tion (H, P).
FIhGolAo.tTyeprer,isQsMwGa2l1ke1r4i4u7s.coAv,acReivgihcthaperostsatpe., vnenotrva.l
Etymology
view;B, right anteriorspermatheca, ventral view.
Named for Ms Jeanette Covacevich.
diaphanous and applied to its posterior surface;
gizzard deflecting septa posteriorly so that its Remarks
posterior end is invested by a pair of lung,
anteriorlyprojecting lobesofseminal vesiclesof Although in the key T. covacevichae forms a
IX. Oesophagus with circumferential vascular couplet with T. canaliculatus, the latter differs
striaein 1X-XVI, virtuallyobliterated by gizzard substantially in the following features: zz is
inVIIandVIII;greatlyexpandedinXVandXVI wider, nephropore rows are irregular, male and
where itattains the width ofthe intestine; dilata- spermathecal pores are more lateral, intestinal
tionnotconstricted offfrom oesophageal lumen origin is more anterior (XVII), nephridial blad-
butslightly depressed middorsally, giving a sug- ders are adiverticulate, and seminal vesicles arc
gestion ofapaired condition and internally with in XI and XII.
circularly arranged, moderate vascular rugae.
Oesophagus narrow and whitish in XVII and Terrisswalkeriusmcdonaldi sp. nov.
XVIII. Intestinal origin XIX, although bulging (Figs5;6A,B)
forward so as to appear to commence in XVIII;
typhlosole and caeca absent (H, P). Intestinal
contents a fine reddish 'silt\ of unknown com- TypeLocality
position, and vegetablematter(including,in H. a 17°38'S. 145°32'E., Majors Mtn. N.E,
recurvedthorn fillingmuchoflumen). Nephridia Queensland,coll.J.Covacevich, K.R. McDonald
stomate, vesiculate holonephridia; in anterior andR. Raven, 14-19.iv. 1978.
MEMOIRSOFTHEQUEENSLAND MUSEUM
164
paired, midventral, in 5/6-8/9, not recognisable
unless intersegmental furrows are held open.
Dorsal blood vessel single. Last hearts in XII.
Gizzard large, firmly muscular, in VI (?), with
spermaUwcal conspicuousanteriorrim;precededbyalarge,but
pore1 not dilated, soft-walled proventriculus.
Oesophagus with a pairofcircumferential blood
vessels, and dilated, in XIV, XV and XVI, with
many discrete internal vascular folds: these
permathecal dilatations not constricted off from oesophageal
pore-! lumen, but probably to be considered true cal-
ciferous glands; oesophagus in XVII short and
narrow and invaginated into posterior face of
calciferous glands of XVI. Intestinal origin
XVIII;typhlosolcabsent;contentsfinely particu-
femalepore late vegetable matter Nephridia stomate,
avcsieulate holonephridia; each discharging at
body wall through a long duct which, although
lacking a bladder, has a clearly visible lumen;
egressofducts is in vicinity ofsetal rows 10and
malepore 1 1 in forebody and in anterior intestinal region,
the row on each side not notably irregular.
Holandric; large, iridescentfree funnels in X and
XI; seminal vesicles small but elongate in XI,
very small in Xll, not subdivided into loculi.
Ovaries, with several delicate strings ofoocytes,
which are terminally large, in XIII; ovisacs not
recognizable. Prostates tubuloracemose, exter-
FIG.5. Terrisswulkeriusmcdonaldisp. nov.,hololypc,
QveMntGra2l1vi14e5w0..Regionofmaleandspermathecalpores, wnailtlhystohmiceklmyintuubtuelasrurafnadceaplampousltatsimoon,otrheswtrailclteedd
toXVIII inwhichthey extendlaterally fromtheir
MaterialExamined ducts; the distal third recurved on remainder of
Holotype: QMG211450m QMGH1994 gland; each muscular duct describing a
posterolateral loop; conjoined vasa defcrentia o\'
Description its side joining duct at about one seventh of its
length from gland. Penial setae absent. Four un-
Length 31mm, width (midclitellar) 2.4mm,
segments 106. Body not depressed. A faint trace pairedspermathecacpresent,eachwithatortuous
of purplish pigmentation dorsally in alcohol vasadeterentla
Prostomium tanylobous (?); body not canalicu-
late. First dorsal pore 4/5. Setae 36 in XII; 36 prostateduct
caudally; ventral and dorsal interruptions small
butvisiblethroughoutinXllaa:ab:be:cd:vz.:z=
1.8:1.0:1.0:0.7:1.1:1.8. Nephropores not'exter-
nally recognizable. Clitellum annular, yellow
brown, 1/2 XNI-XVI1; intersegmental furrows
obliterated; dorsal pores occluded- Male pores a spermathecaldiverticulum spermathecal
pairin setal arc ofXVUI,on very small papillae spermathecalduct ampulla
separated medianly by a minute papilla, situated
inanelongateoval areawhichextendsfromsctal
arc ofXVII toposterior XIX andhas atumidrim mm
B 1
surrounding a depressed and apparently glan-
dularareaanteriorandposteriortomalepapillae.
Femaleporespaired(?)inatransverseslitshortly FIG.6. Terrisswulkeriusmcdonaldiup, nov.,holotype,
anteriortosetal arcofXIV, inatransversely oval QMG21 1450. A, rightprostate; B, rightspermatheca
slightly tumid field. Spermathecal pores 4, un- ofVffl.
NEWQUEENSLANDEARTHWORMS
165
digitiform or narrowly saccular ampulla, con-
tinuous with and not demarcated from is duct; a
subspheroidal diverticulum almost sessile on y\\\—j
spermatheca ai a point which arbitrarily may be
considered the enlal endofduct
spermalhetal
pore
ETYMOLOGY
For Keith R. Mcdonald.
Remarks
Terrisswalkcrrus medonatdi appears closest
morphologically and, presumably phylogeneti-
cally, to T. (=Diporochaeta) miUaamillaa
(Jamieson, 1976b),fromnearbyMillaaMillaa.T.
miUaamillaa is also a very small species resem-
bling D. tncdonaldi in locationofthe first dorsal
pore in 4/5; nephropores in straight or nearly
straightrows;clitellumonXTV-XVII;malepores
median to a lines in an oval field; absence of
accessory genital markings (as in all Ter-
risswallcerius); spermaihccal pores in 4interseg-
ments, 5/6-8/9; last hearts in XII; prostates
restricted to XVIII, recurved entally. and sper- —
mathecaeelongatesacciform,withpoorlydemar- .litHli'm
catcd duct and clavate, almost sessile
diverticulum. It differs from T. miUaamillaa in
the unpaired condition ofthespermathecalpores
which isunknownelsewhereinTerrisswalkerius in;,I- |:,
although invariable for Fletcherodrilus. Dilata-
tion and internal folding ofthe oesophagus in 7.
miUaamillaa (in XV and XVI) is less developed
than in T. mcdotialdi.
Terrisswalkeriuslibersp. nov. FTG. 7. Terrisswalkerius liber sp. nov., holotype,
(Figs7;8A.B) QMG211445.Regionofmaleandspermathecalpores,
ventral view.
Type Locality
late. Firstdorsal pore 5/6 (H, P). Setae24 inXII;
J5°45'S. 145°17*E.,HomeRule,nearSlatyCk, aa:ab:bc:cd:yz:zz = 2.3:1.0:1.3:1.4:1.9:25.5;
QN.uEe.enQsuleaenndsMlaunsde.uUmnpdaerrty,fa2l5l.exn.1s9i7a4g.hom fern, vweindterailnbhrienadkbnoadryr,odwoirnsaflorberbeoadkyebxuttremmoedleyrawtiedley
throughoutforebody,sothatsetaearevisibleonly
MaterialExamined
HOLOTYPE: QMG211445exQMG9010 laterally; narrowerbutvery irregularposteriorly;
Paratyphi QMG21446exQMG9010 several of the more dorsal rows not visible on
clitellumbutvisiblebeforeandbehindit; setaein
Description forebody very strongly protruding in a posterior
Length (holotype.H, isposterioramputee) 170 direction. Nephropores, each with a dark encir-
mm (previously dissected paratype, P), width 7 cling field,conspicuousinanalmoststraightline
mm,segments 150.Bodynotdepressed.Purplish on each side on clitellum, almost exactly lateral,
greydorsaJly;intersegmentalfurrowsandventral in setal lines 1 1 relative to XIU. Clitellum an-
surface pale flesh coloured, in alcohol. Pros- nular, not strongly developed; pale reddish
tomium prolobous (H) orvery slightly indenting brown; from setal annulus of XIII to presctal
peristomium (P), with open lateral margins ex- annulus ofXVIII, inclusive; intersegmental fur-
tending to halfperistomium; deeplycanaliculate rowsweakerandnotpaler; dorsal poresretained;
dorsally and bisected ventrally; body canalicu- canalicula weaker. Maleporescruciform slits,in
MEMOIRSOFTHEQUEENSLANDMUSEUM
!<fe
origin 1/2XVII(H,P).Nephridiastomaic,vesicu-
prostategland late holonephridia with large pyriform bladders;
slenderductjoiningbladderslightlysubterminal-
ly in pharyngealregion and increasingly ectal of
apex of bladder posteriad but recognition of a
diverticulum unwarranted. Holandric; very large
iridescent seminal funnels in X and XI; seminal
orosfateriurt vesiclesapairoflargesimplealmostsmooth-sur-
mm faced sacs on posteriorseptum ofIX (H, but not
1
P); and anterior septa of XII and, remarkably,
XIII (H, P). Ovaries moderate sized laminae of
,;i^iMi.-.lhu> i manyoocytesventral toputativeseminalvesicles
dud of XIII (H, P). A pair of small sacs on anterior
ijpefmalheral face of XIV may be ovisacs. Prostates slender,
-J- l|'lI.I.I tubular and much coiled, in XVII! and XIX (P),
or extending into XXI (H); slender muscular
sinuous ducts discharging near midventral in
XVIIl, each receiving parallel vasa deferentia of
its side at midlength (H, P). Spermathecae a
single pair, each with a very large saccular am-
pullawith, in itsectal half, a papillated surface, a
well demarcated annulated duct; and a terminal.
tubular,sinuous,inseminateddiverticulumwhich
islongerthancombined ampullaand duct(H,P).
FJG. 8. Tetrivtwalkertvi Ubtr sp. nov., paraiypc.
QMG211446. A, right prostate; B. leftspermatheca.
dorsal view. Etymology
setal arc of XVIII, in a lines, on a pair of small From the Latin liber = free, reflectingthename
rounded papillae which are poorly defined ofthe type-locality.
laterally and fill middle third ofsegmentand are
almost in contactmidventrally(H, Pi Accessory Remarks
genital markings absent. Female pores immedi-
ately anteromedian to setae a, on a low whitish Terrisswalkeriui liber is very similar mor-
presetal elevation in XIV (H, P). Spermathecal phologically to T. nashi (Jamieson, 1976b; see
pores 1 pair, shortly posterior to intersegmental below)and isundoubtedly itssisteMaxon. Ofthe
furrow 8/9, each a clearly visible minuieporeon few species of the genus with a single pair of
a very small but well defined wart-like papilla in spermalheca! pores, T, liber and 71 nashi are the
acommon shaMrply defineddepression. only ones in which they are at intersegment X/9.
Septa 10/1 2/13 greatly thickened(H). Dor- 7*. liber differs in the close apposition of the
spermathecal and male pores which are sig-
sal blood vessel single; dorsoventral commis-
nificantly widerapart in T. nashi. Sympatry of 7*.
surals in V-XIJ; those in X-XII large, heart-like nashi and T. liber at Home Rule strengthens
and latero-oesophageal, originating by a large
rather than weakens the grounds for separating
branch from the supra-oesophageal vessel and a
the two entities and the difference in separation
thin, scarcely recognizable branch from the dor- of the genital pores suggests that interbreeding
sal vessel(H,P).Gizzardverylarge,elongateand would not occur between the two forms. Never-
firmly muscular, in V, ensheathed by the ex- theless, reproductive isolation remains to be es-
tremelythinsepta5/6and6/7; itsposteriorendat tablished.
level of VIII relative to external segmentation;
obliterating oesophagus in VI and VII (H, P); a
large muscular proventriculus in IV giving an Terrisswalkeriusnashi
almost digiceriate appearance (H). Oesophagus (Jamieson. 1976b) comb, nov
segmentally dilated and vascular, witha circum-
ferential blood vessel on each side, in XIII-XV1; Diporachaeta nashi Jamieson, 1976b: 41-42. figs I.
but dilatations not cut off from gut. Intestinal Ilb»14j,15m. I6n-p table2.
7