Memoirs of Museum Victoria 58(1): 39–75 (2000)
A REVIEW OF THE TUBIFICIDAE (ANNELIDA: OLIGOCHAETA) FROM
AUSTRALIAN INLAND WATERS
ADRIAN M. PINDER1 AND RALPH O. BRINKHURST2
Department of Conservation and Land Management, Wildlife Research Centre,
PO Box 51, Wanneroo, Western Australia 6946, Australia (adrianp@calm.wa.gov.au)
1
205 Cameron Court, Hermitage, Tennessee 37076, USA (Oligo1@aol.com)
2
Abstract
Pinder, A.M. and Brinkhurst, R.O., 2000. A review of the Tubificidae (Annelida: Oligochaeta)
from Australian inland waters. Memoirs of Museum Victoria 58(1): 39–75.
The Australian limnic representatives of the cosmopolitan oligochaete family Tubificidae
are reviewed. The four Australian species of Telmatodrilus Eisen, 1879 are placed in new
genera (Breviatria and Biprostatus) and these are considered to belong to Rhyacodrilinae
rather than Telmatodrilinae. Four new species, Rhyacodrilus megaprostatus, Ainudrilus
nharna, Breviatria arvensis and Biprostatus duplex are described and Rhyacodrilus fultoni
Brinkhurst, 1982 is transferred to Ainudrilus Finogenova, 1982. Limnodrilus claparedeianus
Ratzel, 1868 and Aulodrilus limnobius Bretscher, 1899 are recorded in Australia for the first
time. The Australian fauna now consists of 29 described species, including 11 of the most
widespread species of cosmopolitan genera (Tubifex Lamark, 1816, Potamothrix Vejdovský
and Mrazek, 1902, Limnodrilus Claparède, 1862, Aulodrilus Bretscher, 1899, Rhyacodrilus
Bretscher, 1901, Bothrioneurum Stolc, 1886 and Branchiura Beddard, 1892). Antipodrilus
Brinkhurst, 1971, with five species, and the two new rhyacodriline genera, with six species,
are essentially Australian genera, except that one species of Antipodrilus and possibly one of
Biprostatus also occur in New Zealand. Seven endemic species belong to more widespread
genera (Rhizodrilus Smith, 1900, Rhyacodrilus and Ainudrilus). The Australian tubificid fauna
appears to have biogeographic affinities with northern hemisphere and marine tubificids.
provide a foundation for future work, complementing the recent review of the Australian
Phreodrilidae (Pinder and Brinkhurst, 1997). Our
definition of the Tubificidae is conservative,
excluding the Naididae which are generally
acknowledged to be part of the same taxon
(Brinkhurst, 1994; Erséus, 1990b), although a
formal proposal to merge the families has yet to
be made.
Introduction
The first tubificid to be described from Australia
was Branchiura pleurotheca Benham, 1907 from
Blue Lake, Mt Kosciusko. Further tubificid
records and descriptions from Australia were not
published until Brinkhurst (1971a, b) reviewed
the aquatic oligochaete faunas of the world
and Australia. In these publications, Branchiura
pleurotheca was synonymised with Rhyacodrilus
coccineus (Vejdovský, 1875), several other new
species and a new genus (Antipodrilus) were
described and the presence of numerous cosmopolitan species was recorded. Subsequent
records and descriptions of new species were provided by Brinkhurst and Fulton (1979) and
Brinkhurst (1982, 1984).
In this paper we list previous records and add
many new ones for most species, improve some
descriptions, describe four new species and
address problems relating to the taxonomic
position of Australian species of Telmatodrilinae by moving them to two new rhyacodriline genera. This contribution is intended to
Materials and methods. Most specimens were
stored in ethanol, often after fixation in formalin.
They were then whole-mounted, dissected or
serial sectioned. Dissected or whole-mounted
specimens were usually first stained in
Grenacher’s borax carmine. Sectioned specimens were cut at 6 mm and stained in Mayer’s
haematoxylin and eosin.
Descriptions are limited to brief diagnoses,
with important distinguishing characteristics,
except for new species, or where an original
description can be substantially amended.
The term chaetae is used in preference to setae,
following a determination at the International
39
40
A. M. PINDER AND R. O. BRINKHURST
Symposium on Aquatic Oligochaete Biology,
Presque Isle, Maine, USA, 1997. Roman numerals refer to segment numbers and two arabic
numerals separated by a slash refer to the septum
(or intersegmental furrow) separating two adjacent segments. Body lengths are for preserved
specimens and are indicative of the length of
mature specimens only.
Where details of the holotype are not given, as
is the case for some of the earliest described cosmopolitan species, this information is unknown
and not listed in Reynolds and Cook (1976, 1981,
1989, 1993).
Institutions and collections are abbreviated as
follows: AMS, Australian Museum, Sydney,
NSW; AMP, Adrian Pinder collection; AWT,
Australian Water Technologies, Sydney, NSW;
BGMJ, Barrie Jamieson collection, Brisbane,
Qld; CALM, Wildlife Research Centre, Department of Conservation and Land Management,
Perth, WA; ERISS, Environmental Research
Institute of the Supervising Scientist, Jabiru, NT;
IFC, Inland Fisheries Commission, Hobart, Tas..
(all collections now in QVM); MDFRC, Murray
Darling Freshwater Research Centre, Albury,
NSW (J. Hawking); MNHN, Museum National
d’Histoire Naturelle, Paris, France; MON, Ecology and Evolutionary Biology, Monash University, Melbourne, Vic.. (P.S. Lake); MUR, Aquatic
Ecosystems Research, Environmental Science,
Murdoch University, Perth, WA (J. Davis);
NMV, Museum Victoria, Melbourne, Vic.; NTM,
Museum and Art Gallery of the Northern Territory, Darwin, NT; QVM, Queen Victoria
Museum, Launceston, Tas.; ROB, Ralph
Brinkhurst collection; SAM, South Australian
Museum, Adelaide, SA; TAS., Tasmanian
Museum, Hobart, Tas.; UCA, University of
Canberra, Canberra, ACT (R. Norris);
USNM, National Museum of Natural History,
Washington, USA; WAM, Western Australian
Museum, Perth, WA; ZMUH, Zoological
Museum, University of Hamburg, Germany.
Abbreviations used in the figures are as
follows: a, atrium; c, copulatory chamber; e, ejaculatory duct; f, female funnel; g, gut; gd, gut
diverticulum; gt, glandular tissue; mf, male funnel; n, nerve cord; o, ovary; p, prostate; pa, papillae; pc, penial chaetae; pe, peritoneal tissue;
s, spermatheca; sc, spermathecal chaetae; sd,
spermathecal duct; t, testis; v, vas deferens.
Where possible, localities are given with latitude and longitude, Australian map grid (AMG)
1:100 000 coordinates (four digit map number
followed by three eastings and three northings) or
Tasmap 1: 500 000 coordinates (three eastings
and three northings). Australian states and territories are abbreviated as follows: Australian
Capital Territory (ACT), New South Wales
(NSW), Northern Territory (NT), South Australia (SA), Queensland (Qld), Victoria (Vic.),
Tasmania (Tas.) and Western Australia (WA).
Tubificidae
Diagnosis. Genitalia bilaterally paired. One pair
of testes and 1 pair of ovaries, usually in X and XI
respectively. Spermathecae mostly in the testicular segment. Male funnels on posterior septum of
testicular segment leading to vasa deferentia, atria
and male pores in the ovarian segment. Female
funnels leading to pores on posterior furrow of
ovarian segment. Chaetae from 1 to many per
bundle, hair chaetae usually restricted to dorsal
bundles or absent. Reproduction normally sexual.
Tubificinae
Diagnosis. Spermathecal pores in the middle of
the testes segment in most species. In all but a few
genera, sperm organised as spermatozeugmata of
a specific type, with non-fertile sperm encapsulating fertile sperm. Atria each with a single stalked
prostate gland (prostate rarely absent). Pendant
penes usually present, often with cuticle of penis
sac thickened to form a penis sheath. Spermathecal chaetae, when present, usually single,
enlarged, grooved distally and with large chaetal
glands. Penial chaetae normally absent, but when
present then usually of similar form to the spermathecal chaetae. Coelomocytes not large and
abundant.
Remarks. The spermatozeugmata found in tubificines are of a distinct form, with fertile sperm
cells enclosed by a cortex of bound infertile
sperm cells (Braidotti and Ferraguti, 1982;
Ferraguti et al., 1994; Jamieson, 1992), distinct
from the looser sperm agglomerations found in
the rhyacodrilinae.
Tubifex Lamarck
Type species. Lumbricus tubifex Müller, 1774.
Diagnosis. Hair chaetae usually present dorsally,
accompanied by pectinate chaetae, but polymorphic forms occur. Vasa deferentia long, often
of two widths. Atria comma-shaped, with vasa
deferentia opening close to the apex anteriorly,
each with a large stalked prostate gland. Pendant
penes present, each with a thin tub-shaped sheath.
Genital chaetae not modified. Spermatozeugmata
present.
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
Tubifex tubifex (Müller)
Lumbricus tubifex Müller, 1774: 27 (in part).
Tubifex rivulorum Lamark, 1816: 225.
Tubifex
tubifex.—Michaelsen,
1900:
48.—
Brinkhurst, 1971a: 453, figs 8.1A–D, 8.3G–J.—
Brinkhurst, 1971b: 10, fig. 2A.—Timms, 1973: 11.—
Timms, 1977: 170.—Timms, 1978: 435.—Timms,
1981: 188.—Timms, 1982: 126.—Marshall and
Winterbourn, 1979: 205.
Material examined. NSW: Wingecarribee Creek,
Berrima, 34º29´S 150º20´E, 7 Nov 1991, 4 specimens
(AWT). Tas.: stream 4.9 km above Barrow Creek, 26
Nov 1978, 1 on slide (ROB); creek E of The Needles,
13 Nov 1978, 2 on slides (ROB); Arthur R., 31 Oct
1978, 1 on slide (ROB). Vic.: Beechworth Trout Hatchery, May 1994, 1 on slide (AMP) and Sep 1994, 2 on
slides (NMV F77644–F77645); Thomson R. at Thompson Valley Rd, 24 Nov 1976, 2 on slides and 3 in
alcohol (NMV F77646–F77648); Phytotelmata in fig
tree, South Yarra, Melbourne, several on slide (MON).
WA: Soak NW of Corrigin, 32º10´38´´S 117º49´08´´E,
21 Oct 1997, several on slides and in alcohol (CALM);
Paperbark Swamp, E of Corrigin, 32º24´58´´S
118º05´52´´E, 8 Oct 1997, 1 on slide (CALM); Farm
dam near Corrigin, 32º19´44´´S 118º02´24´´E, 7 Oct
1997, 2 on slides (CALM).
Other Australian records. Qld: Downfall Creek
(Brinkhurst, 1971b). SA: A creek near Adelaide, no
date, 1 on slide (AMP). Vic.: Lakes Bong Bong,
Bridgewater, Purrumbete and Elusive (Timms, 1973,
1981, 1977). WA: Lake Monger, Perth (Brinkhurst,
1971b). Tas.: Lake St Clair (Timms, 1978).
Distribution and habitat. Cosmopolitan, including New Zealand (Brinkhurst, 1971b; Marshall
and Winterbourn, 1979; Timms, 1982). Although
widespread in Australia, T. tubifex is not commonly encountered. In the northern hemisphere
T. tubifex is commonly found in marginal sites,
such as those with heavy organic pollution or
intense sedimentation or in highly oligotrophic
waters (Brinkhurst, 1996). In sites with heavy
organic pollution this species is usually associated
with Limnodrilus hoffmeisteri Claparède, 1862
and these are the dominant oligochaetes or even
the dominant benthic invertebrates (Brinkhurst,
1996). Studies of gross organic pollution are less
common in Australia (as are ecological studies in
which oligochaetes are speciated) and the Australian records of T. tubifex span a wide variety of
habitats. This species can tolerate salinities up to
10 ppt (Chapman et al., 1982) and mature worms
can encyst to survive short periods of drought
(Kaster and Bushnell, 1981).
Diagnosis. Length 20–200 mm. Anterior dorsal
bundles with 1–6 hair chaetae and 3–5 pectinate
chaetae with irregular intermediate teeth. Hair
chaetae and pectinate chaetae reduced posteriorly.
41
Ventral chaetae bifid, 6–10 per bundle anteriorly,
with the upper teeth thinner than but not much
longer than the lower, posteriorly with the upper
teeth relatively shorter and fewer chaetae. Polymorphic forms with hairs short and bifid chaetae
dorsally, or bifid chaetae both dorsally and
ventrally, may be found, especially where conductivity is high. Vasa deferentia elongate,
divided into thin and thick sections, entering the
comma-shaped atria on the concave side, near
the where the prostate glands join. Penis sheaths
tub-shaped but often indistinct.
Limnodrilus Claparède
Limnodrilus Claparède, 1862: 221, pl I figs 1–3, pl.
III fig. 12, pl. IV fig. 6.—Michaelsen, 1900: 42.—
Stephenson, 1930: 747.—Brinkhurst, 1971a: 463.
Type
species.
Claparède, 1862.
Limnodrilus
hoffmeisteri
Diagnosis. Dorsal and ventral chaetae bifid, hairs
absent. Vasa deferentia long, entering short beanshaped atria apically. Prostates large, entering
atria by short stalks. Atria connect, via long ejaculatory ducts, to pendant penes lying free within
thick penis sheaths (formed by the thickened cuticle of the penis sac wall) of varying lengths, some
longer than the body width. No modified genital
chaetae. Spermatozeugmata present.
Remarks. Mature specimens of the three species
present in Australia are readily separable by
examining the penial sheaths. These are usually
visible through the body wall of cleared and slide
mounted specimens.
Limnodrilus hoffmeisteri Claparède
Limnodrilus hoffmeisteri Claparède, 1862: 248, pl. I,
figs 1–3, pl. III, fig. 12, pl. IV, fig. 6.—Brinkhurst,
1971a: 464, figs 8.3M, O, 8.4C, H, I, 8.5E.—
Brinkhurst, 1971b: 112, fig. 2B.—Brinkhurst and
Fulton, 1979: 1.—Fulton, 1983a: 778.—Fulton, 1983b:
792.—Timms, 1978: 428.—Timms, 1982: 126.—
Marshall and Winterbourn, 1979: 205.
Clitellio (Limnodrilus) hoffmeisteri.—Valliant, 1890:
424.
Tubifex hoffmeisteri.—Ditlevsen, 1904: 422.
Material examined. ACT: Murrumbidgee R.,
1987–1988, returned to R. Norris (UCA). NSW:
Georges R. at Liverpool Weir, 1995, 1 on slide (AMS
W7876); Meggaritys Creek at Warradale Rd, 33º54´S
150º36´E, 9 Feb 1993, 1 on slide (AWT); Rushy Billabong, 36º02´S 146º42´E, 5 Sep 1984, 1 on slide
(MDFRC); Ryans Billabong, 36º06´S 146º58´E, 30 Jun
1981, 1 on slide and 1 in alcohol (MDFRC); Orphan
School Creek, Kings Park, 3 May 1996, 1 on slide
(AWT). Qld: Bulimba Creek, Brisbane, 4.8 km from
mouth, 4 on slides (AMS W18081). SA: Valley Lake,
42
A. M. PINDER AND R. O. BRINKHURST
Mt Gambier, 37º50´S 140º46´E, 26 Feb 1978, 1 on slide
(AMS W18084). Vic.: La Trobe R. at Moe—Willow
Grove Bridge Rd and 100 m downstream of Moe Drain,
AMG 8121–346773 and 8121–359769 respectively, Jul
1980, 2 on slides (NMV F77626–7); Mitta Mitta R., 2
km downstream of Dartmouth Dam, 31 Jan 1975 and
Nov 1975, 3 on slides (NMV F77628, F77629, F77631)
and 4 km N of Eskdale, 1 Feb 1975, 1 on slide (NMV
F776230). WA: Munday Swamp, Perth, 22 Nov 1989,
1 on slide (WAM 41–95); Fire dam on Seven Day Rd,
Warren R. catchment, 34º19´S 115º58´E, 4 in alcohol,
1 on slide (WAM 39–95 and 40–95); stream across
Bussell Highway, 34º08´32´´S 115º06´41´´E, 15 Sep
1996, 1 on slide (AMP).
Other Australian records. Tas.: Arthurs Lake and Lake
Sorrel (Fulton, 1983ab); Great Lake (Brinkhurst and
Fulton, 1979); Lakes St Clair, Dove, Sorell, Crescent,
Leake and Toombs (Timms, 1978).
Distribution and habitat. Cosmopolitan, including New Zealand (Brinkhurst, 1971b; Timms,
1982; Marshall and Winterbourn, 1979). This is
the most commonly encountered Limnodrilus and
is found in many habitat types, reaching very high
abundance in organically enriched sites, often
with T. tubifex (Brinkhurst, 1965b). It can tolerate
salinities up to 10 ppt (Chapman et al., 1982).
Diagnosis. Length 20–35 mm. Anterior bundles
with 3–10 chaetae with upper teeth varying in
length from slightly shorter than, to longer than,
the lower and usually thinner than the lower.
Fewer chaetae posteriorly. Penis sheaths up to 20
times longer than broad, ental ends slightly flared,
ectal ends either with elaborate hood opening at
right-angles to shaft or sometimes a plate with
scalloped edges.
Limnodrilus udekemianus Claparède
Limnodrilus udekemianus Claparède, 1862: 243, pl.
I, figs 4–5, pl. III, figs 13, 13a, pl. VII, fig. 1.—
Brinkhurst, 1971a: 467, figs 8.4A–B.—Brinkhurst,
1971b: 113, fig. 2E.—Timms, 1978: 428.—Marshall
and Winterbourn, 1979: 205.
Clitellio (Limnodrilus) udekemianus.—Vaillant
1890: 425.
Tubifex udekemianus.—Ditlevsen, 1904: 422.
Material examined. NSW: Dairy drains, Taree, 1 Apr
1987 (AMS). Qld: Bulimba Creek, Brisbane, 4.8 km
from mouth, 2 on slides (immature, but with characteristic chaetae) (AMS W18082). SA: Valley Lake, Mt
Gambier, 37º50´S 140º46´E, 26 Feb 1978, 1 (immature,
but with characteristic chaetae) on slide (AMS
W18083). WA: Katanning Creek at Katanning sale
yards, 33º42´S 117º34´E, 16 Sep 1997 1 on slide
(CALM).
Other Australian records. Qld: Moggill Creek, Brisbane; Downfall Creek, Chermside. Tas.: Lake Pedder
(all previous from Brinkhurst, 1971b), Lake St Clair
and Dove Lake (Timms, 1978). WA: Lake Monger,
Perth, uncertain identification (Brinkhurst, 1971b).
Distribution and habitat. Cosmopolitan, including New Zealand (Brinkhurst, 1971b; Marshall and Winterbourn, 1979). Often, though
not exclusively, found in organically polluted
situations.
Diagnosis. Length 20–90 mm. Anterior chaetal
bundles with 3–8 bifid chaetae with the upper
teeth much thicker and longer than the lower,
fewer chaetae posteriorly with teeth more nearly
equal. Penis sheath usually only up to 4 times as
long as wide with a simple plate-like hood usually
reflected back over the shaft.
Limnodrilus claparedeianus Ratzel
Limnodrilus claparedeianus Ratzel, 1868: 590, pl.
XLII, fig. 24.—Michaelsen, 1900: 45.—Brinkhurst,
1971a: 468, figs 8.3Q, R.—Brinkhurst, 1971b: 113,
fig. 2F.
Tubifex claparedeianus.—Ditlevsen, 1904: 422.
Material examined. Vic.: Lake Thirlemere, 8 Sep 1981,
2 on 3 slides (NMV F81432).
Distribution
and
habitat.
Cosmopolitan
(Brinkhurst, 1971b), occurring in a wide range
of habitat types.
Diagnosis. Length 30–60 mm. Anterior bundles
with 4–9 bifid chaetae, some with the upper teeth
much longer than and as thick or somewhat
thicker than the lower, fewer chaetae posteriorly.
Penis sheaths up to 50 or even 80 times longer
than broad with small triangular plates set at an
angle to the shaft distally.
Remarks. This is the first and only record of this
species from Australia. The shape of the penis
sheath is normally diagnostic except in North
America, where variants between this species and
Limnodrilus cervix Brinkhurst, 1963 can be
observed.
Potamothrix Vejdovský and Mrazek
Potamothrix Vejdovský and Mrazek, 1902: 24.—
Brinkhurst, 1971a: 486.—Holmquist, 1985: 311.—
Finogenova and Poddubnaja, 1990: 55.
Euilyodrilus Brinkhurst, 1962: 329.
Type
species.
Potamothrix
Vejdovský and Mrazek, 1902.
moldaviensis
Diagnosis. Hair chaetae present in dorsal bundles
of most species. Vasa deferentia short, joining
elongate tubular atria apically, prostate glands
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
rudimentary or absent. Pendant penes small, without cuticular sheaths. Large grooved genital
chaetae associated with spermathecal pores.
Spermatozeugmata present.
Remarks. While the genus is largely holarctic, and
mostly palearctic, some of the many PontoCaspian species appear to have been introduced to
North America, Africa and South America (Peru)
and others are spreading through Europe (G. Milbrink, pers. comm.). The single species occurring
in Australia, Potamothrix bavaricus (Öschmann,
1913), is a cosmopolitan species, although it may
have been introduced to some parts of its current
range, e.g., South America (Brinkhurst and
Marchese, 1989).
Potamothrix bavaricus (Öschmann)
Tubifex (Ilyodrilus) bavaricus Öschmann, 1913: 559,
figs 1–5.
Ilyodrilus bavaricus.—Stephenson, 1930: 746.
Euilyodrilus bavaricus.—Brinkhurst, 1962: 321.
Potamothrix bavaricus.—Brinkhurst, 1971a: 492,
figs 8.12H,I.—Brinkhurst, 1971b: 113, fig. 2G.—
Timms, 1981: 188.—Marshall and Winterbourn, 1979:
205.
Holotype. Type locality unknown (ZMUH 7704).
Material examined. NSW: Gowrie Creek, 54 km downstream of sewage treatment plant, 27º20´S 151º27´E,
28 Sep 1995, few in alcohol (AWT). Tas.: Big Waterhouse Lake, 40º53´20´´S 147º36´50´´E, 1 Mar 1997, 2
on slide (AMP). Vic.: Curdies R., 38º20´S 143º08´E, 7
and 29 Feb 1992, 2 on slides (NMV F77641–2). WA:
Malaga Wetlands, Perth, 2 Nov 1989, 2 on slides
(WAM 76–95, 77–95); Herdsman Lake, Perth, 2 and 13
Nov 1989, 13 on slides, 8 in alcohol (slides WAM
55–95, 67–95, alcohol 54–95, 78–95); Lakes Nowergup, 31º38´16´´S 115º44´42´´E, Yonderup, 31º34´S
115º41´E, and Carabooda, 31º37´S 115º43´E, 8 on
slides (WAM 68–95 to 75–95); Coogee Springs, Perth,
9–14 Nov 1989, several on slides (MUR); Shark Lake,
N of Esperance, 33º46´S 121º51´E, 5 Sep 1996, many
specimens in alcohol (AMP); Congelin Dam,
32º49´07´´S 116º53´05´´E, 26 Oct 1997, 3 on slides
(CALM).
Other Australian records. Vic.: Lake Purrumbete,
Camperdown (AMS W4275) (Timms, 1981). WA:
Corio Pool, Rottnest Island (Brinkhurst, 1971b).
Distribution and habitat. Eurasia, North, Central
and South America, Australia and New Zealand
(Brinkhurst, 1971a; Brinkhurst and Marchese,
1987; Coates and Stacey, 1994; Marshall and
Winterbourn, 1979). Potamothrix bavaricus is
quite widespread in southern Australia, occurring
in both lentic and lotic sites and often very
abundant when present.
43
Diagnosis. Length 15–35 mm. Dorsal anterior
bundles with 1–5 hair chaetae and 2–5 pectinate
chaetae. Ventral chaetae with teeth equally long
but the upper thinner than the lower. Spermathecal chaetae relatively stout, about equal in
length to the somatic ventral chaetae, with triangular blade-like ectal half. Vasa deferentia short
joining tubular atria apically. Prostate glands
absent. Penis sheaths absent.
Antipodrilus Brinkhurst
Antipodrilus Brinkhurst, 1971a: 523.—Brinkhurst
and Wetzel, 1984: 50.
Type species. Tubifex davidis Benham, 1907.
Diagnosis. Dorsal chaetal bundles with or without
hairs. Vasa deferentia long and narrow, joining
atria apically or subapically. Atria globular, narrowing to form ejaculatory ducts that terminate in
pendant penes (or protrusible pseudopenes?).
Apparent penis sheaths (formed by thickened
cuticular lining of the ectal part of the male duct)
present in some species. Single stalked prostate
gland attached apically each atria. Spermathecae
with elongate spermatozeugmata. Spermathecal
chaetae thin with grooved elongate tips.
Remarks. Antipodrilus is known only from Australia (all five species) and New Zealand (one
species, A. davidis). In a cladistic analysis of the
subfamily by Brinkhurst (1991), Antipodrilus
appeared to group with two genera that are widely
distributed in the northern hemisphere: Psammoryctides Hrabe, 1964 and Haber Holmquist,
1978. These northern genera have protrusible
pseudopenes, with a crumpled cuticular lining to
the inverted ectal part of the male duct, which
then forms a penis sheath when the penis is
everted. Antipodrilus magelensis Brinkhurst,
1984 and possibly Antipodrilus timmsi Brinkhurst, 1971, appear to have similar crumpled
sheaths within the ectal part of the male duct and
so the genus was coded as having protrusible
pseudopenes in Brinkhurst (1991). Unfortunately
further Antipodrilus specimens were not available
for this study. Antipodrilus davidis (Benham,
1907) does appear to have pendant penes (though
these may be protruded pseudopenes) and the
nature of the penes is not clear for the other two
species (Antipodrilus multiseta Brinkhurst and
Fulton, 1979 and Antipodrilus plectilus Brinkhurst and Fulton, 1979). Further anatomical study
is required to confirm the presence of pseudopenes and to generally improve descriptions of
Antipodrilus species.
44
A. M. PINDER AND R. O. BRINKHURST
Antipodrilus davidis (Benham)
Tubifex davidis Benham, 1907: 252, figs 1–6.—
Hrabe, 1962: 307.—Hrabe, 1966: 74.
Euilyodrilus heuscheri (Bretscher).—Brinkhurst,
1963: 49 (in part).
Antipodrilus davidis.—Brinkhurst, 1971a: 523.—
Brinkhurst, 1971b: 112, fig. 2b.—Brinkhurst, 1982:
1.—Timms, 1979: 121.—Timms, 1982: 126.—
Marshall and Winterbourn, 1979: 205.
Holotype. Blue Lake, Mt Kosciusko, NSW, 1906,
sectioned on 4 slides (AMS G5610–11).
Material examined. NSW: Holotype, as above. SA: The
Bubbler Spring, near Lake Eyre, 3 Dec 1974, 2 on
slides, 1 in alcohol (SAM E2823–E2825); Coopers
Creek, southwest branch, 27 Mar 1987, 1 on slide
(SAM E2817); Freeling Springs, 1987 (AMS
W198716); Davenport Spring Bore, near Hermit Hill,
Lake Eyre South, 21 Sep 1989, 5 on slides (SAM
E2818–E2821). WA: Spectacles Lake (North Eye),
Brownmans Swamp, Lake Mariginiup and Thomson
Lake, Perth, 30 Oct and 3–9 Nov 1989, 11 on slides, 8
in alcohol (WAM 42–95 to 47–95, 49–95 to 53–95 and
81–95).
Other records. NSW: Mt Kosciusko lakes (Timms,
1979). Following records all from Brinkhurst (1971b).
SA: Lakes Edward and Leake, Glencoe. Vic.: Lakes
Surprise and Mumblin, western Victoria. Tas.:
Cambridge, 30 Aug 1966 (TAS. K221). Also recorded
from New Zealand (Brinkhurst, 1971b; Marshall and
Winterbourn, 1979; Timms, 1982).
Distribution and habitat. Southern Australia and
New Zealand (Brinkhurst, 1971b; Marshall and
Winterbourn, 1979; Timms, 1982). Favours lentic
habitats.
Diagnosis. Length 25–40 mm. Anterior dorsal
bundles with 2–5 non-hispid hair chaetae and 2–4
pectinate chaetae with teeth approximately equal.
Ventral chaetal bundles with 3–5 bifid chaetae
with the upper teeth longer and thinner than the
lower anteriorly and teeth equal or the upper
shorter posteriorly. Spermathecal chaetae single,
grooved or with short lower tooth (probably when
partially developed), about equal in length to
somatic ventral chaetae, rarely absent. Vasa deferentia long, atria small, narrowing abruptly to
form long ejaculatory ducts leading to (pendant?)
penes.
Antipodrilus timmsi Brinkhurst
Antipodrilus timmsi Brinkhurst, 1971b: 112, fig.
12c.—Brinkhurst and Wetzel, 1984: 50.—Timms,
1981: 187.—Timms, 1983: 168.
Holotype. Lake Bullenmeri, Vic., 13 Aug 1969, 5 on
slides (AMS W4276–W4278, typus amissus).
Material examined. NSW: Lake Jillamatong, near
Cooma, 3 Apr 1986, many in alcohol (NMV F77614).
Vic.: Type locality and date, 5 on slides (ROB); Lake
Colangulac, 2 on slides (NMV 77612–F77613).
Other Australian records. Vic.: several lakes in western
Victoria (Timms, 1981, 1983).
Distribution and habitat. Lakes in southeastern
mainland Australia. Has been collected in sites
with up to 25 ppt salinity, in both sandy and
muddy sediments and at depths of up to 65 m
(Timms, 1981, 1983).
Diagnosis. Length at least 14.5 mm (length of
largest fragment). Anterior dorsal bundles with
2–5 serrate hairs and 3–5 pectinate chaetae with
the upper teeth slightly longer than the lower,
fewer posteriorly with upper teeth becoming
longer and pectinations less apparent or absent.
Ventral bundles with 3–5 chaetae anteriorly with
the upper teeth markedly longer and somewhat
thinner than the lower, fewer chaetae posteriorly
with upper teeth less markedly longer than the
lower. Spermathecal chaetae single, grooved and
narrowly blade-shaped distally, slightly longer
than somatic ventral chaetae. Vasa deferentia
long, atria small and almost spherical, narrowing
abruptly to form ejaculatory ducts. Penes present,
possibly with thickened inner cuticle.
Remarks. The crumpled cuticle within the penes
is indicative of a protrusible penis, but the structure of this requires closer examination from fresh
specimens.
Antipodrilus magelensis Brinkhurst
Antipodrilus magelensis Brinkhurst, 1984: 142, figs
1–2.
Holotype. Mudginberri Billabong, Magela Creek, NT, 4
May 1981, 1 on slide (USNM 80692).
Material examined. NT: Holotype, as above, and
paratypes from type locality as well as Georgetown and
Bowerbird Billabongs, Magela Creek, 11 on slides, 19
in alcohol (USNM 80693–80697, 80700) and several
on slides (ROB); Nankeen Billabong, Magela Creek, 1
immature on slide (ROB); South Alligator R., Oct 1988
and Dec 1989, 1 on slide (NMV F77611) and 2 on
slides (ERISS). SA: Charles Angus Bore, 15 km N of
Alberri Creek railway siding, Lake Eyre South, 2 Dec
1974, 1 on slide (SAM E2815); Coopers Creek, northwest branch, 27 Mar 1987, 1 on slide (SAM E2816).
Distribution and habitat. Northern Territory
(coastal riverine sites) and South Australia (few
arid inland sites). The new South Australian specimens are the first to be recorded outside of the
Northern Territory.
Diagnosis. Length up to 9 mm. Anterior bundles
with 4–6 bifid chaetae with teeth equal in length
and breadth, diminishing in number posteriad to 2
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
per bundle with upper teeth becoming shorter and
thinner than the lower. Spermathecal chaetae
single with grooved tips, about equal in size to
somatic ventral chaetae. Vasa deferentia moderately long, joining atria subapically. Short atria
terminate in coiled ejaculatory ducts, which lead
to (protrusible?) penes with crumpled cuticular
lining forming apparent penis sheaths.
Antipodrilus multiseta Brinkhurst and Fulton
Antipodrilus multiseta Brinkhurst and Fulton, 1979:
2, figs 6–8.—Brinkhurst and Wetzel, 1984: 50.—
Fulton, 1983a: 778.—Fulton, 1983b: 792.
Holotype. Great Lake, Tas., 1975 (QVM:14:316).
Material examined. Tas.: Holotype and paratypes from
type
locality
and
date,
four
on
slides
(QVM:14:316–319); Lake Sorell, 3 on slides, uncertain
identification (ROB).
Other Australian Records. Tas.: Great Lake, Arthurs
Lake and Lake Sorell (Fulton, 1983ab).
Distribution and habitat. Known only from large
lakes of Tasmania’s central plateau. Collected at
depths of up to 17.5 m and in both silt/sand and
clay sediments (Fulton, 1983b).
45
Diagnosis. Length at least 20 mm. Thin, red
worms that may be found in clusters of up to 50
specimens. Dorsal and ventral bundles with 7–9
chaetae anteriorly, reduced in number posteriorly
to 3 or fewer per bundle. Chaetae of II with upper
teeth shorter and thinner than the lower, chaetae
of other preclitellar bundles with teeth equally
long and with teeth equal in width or the upper
teeth thinner than the lower. Posterior chaetae
with upper teeth shorter and thinner than the
lower. Spermathecal chaetae single, thin and
groove tipped, twice as long as somatic ventral
chaetae. Vasa deferentia moderately long, atria
elongate, distended where vasa deferentia and
prostates attached. Ejaculatory ducts moderately
long.
Remarks. The type specimens have all chaetae
bifid, but those from Lake Sorell have pectinate
chaetae posteriorly. Further investigation may be
required to substantiate the unity of this taxon,
although chaetal polymorphism is known for
other oligochaetes (Chapman and Brinkhurst,
1986, 1987).
Aulodrilus Bretscher
Diagnosis. Length up to 50 mm. Chaetae 12–15
per bundle in II, gradually diminishing in number
to 3 or 4 posteriorly. Anterior chaetae with blunt
teeth of equal thickness, the upper slightly longer,
posterior chaetae with upper teeth shorter than the
lower. Spermathecal chaetae single, thin with
grooved-tips, about equal in size to somatic
ventral chaetae. Vasa deferentia moderately long,
atria and prostate glands small, ejaculatory ducts
long and thin, leading to small penes. Posterior
end of body tapering to a coiled tail. Living within tubes of fine sand grains (at least in the type
locality).
Aulodrilus Bretscher, 1899: 388.—Michaelsen,
1900: 55.—Stephenson, 1930: 752.
Antipodrilus plectilus Brinkhurst and Fulton
Remarks. Asexual reproduction is common in
Aulodrilus (Brinkhurst, 1984) and most Australian records are of immature worms. However,
the three species found in Australia have distinctive chaetae that permit identification of immature
specimens. Finogenova and Arkhipova (1994)
provide scanning electron micrographs of the
distinctive chaetae of the following species.
Antipodrilus plectilus Brinkhurst and Fulton,
1979: 2, figs 1–5. —Brinkhurst and Wetzel, 1984:
50.—Fulton, 1983a: 778.—Fulton, 1983b: 792.
Holotype. Great Lake, Tas., 1975 (QVM:14:309).
Material examined. Tas.: Holotype and paratypes from
type locality and date (QVM:14:309–315); Lake Sorell,
2 Feb 1981, 7 on slides (ROB).
Other Australian records. Tas.: Great Lake and Arthurs
Lake (Fulton, 1983a, b).
Distribution and habitat. Known only from large
lakes of Tasmania’s central plateau. Collected at
depths of up to 17.5 m and in both silt/sand and
clay sediments (Fulton, 1983b).
Type species. Aulodrilus limnobius Bretscher,
1899.
Diagnosis. Dorsal bundles with or without hair
chaetae. Often with gonads replicated and/or genital apparatus shifted forwards due to asexual
reproduction. Vasa deferentia short, atria globular, bean-shaped or cylindrical with solid prostate
glands. Penes pendant without cuticular sheaths.
Penial chaetae (but not spermathecal chaetae)
may be modified. Sperm stored as loose bundles,
spermatozeugmata absent.
Aulodrilus limnobius Bretscher
Aulodrilus limnobius Bretscher, 1899: 388, figs
8.23G–H.—Brinkhurst, 1971a: 524, figs 8.24G, H. —
Finogenova and Arkhipova, 1994: 8, figs 1–2, 4–5, 19.
Material examined. NSW: Murray R. at Heyward’s
Gauge, 36º06´S 147º01´E, 6 Jan 1988, 1 on slide
(MDFRC). Vic.: Wentworth R., 17.7 km upstream of
46
A. M. PINDER AND R. O. BRINKHURST
Tabberabbera, 37º31´S 147º23´E, 2 Oct 1975, 1 in
alcohol (NMV F77618); Mitta Mitta R., 2 km downstream of Dartmouth Dam wall, 5 Nov 1976, 1 in alcohol (NMV F77615); Branjee Creek, Goulburn Valley,
36º42´S 148º20´E, 23 Apr 1992, 1 on slide, 3 in alcohol
(NMV F77616, F77617).
Distribution and habitat. Recorded from Africa,
Europe, Asia and North and South America
(Brinkhurst, 1971a) and now southeastern
Australia.
Diagnosis. Length 12–15 mm. Anterior dorsal
and ventral chaetae similar with up to 10 bifid
chaetae with upper teeth shorter and thinner than
lower, the chaetae of first bundles often shorter
and thicker than the rest. Chaetae of mid and posterior segments with lateral wings which appear
as narrow oar-shaped blades when viewed
frontally, with the upper teeth visible as a small
projection beyond the blade. The wings can be
discerned as a ridge on the lateral aspect of the
chaetae. Vasa deferentia long, entering long
cylindrical atria apically. Pendant penes large.
Genitalia may be shifted forwards.
Remarks. These are the first Australian records of
this otherwise widespread species.
Aulodrilus pluriseta (Piguet)
Naidium pluriseta Piguet, 1906: 218.
Aulodrilus pluriseta.—Piguet and Bretscher, 1913:
118, figs 1–2.—Brinkhurst, 1971a: 524, figs
8.23J–N.—Brinkhurst, 1971b: 114, fig. 2I.—
Brinkhurst, 1984: 147.—Timms, 1982: 126.—Marshall
and Winterbourn, 1979: 205. —Finogenova and
Arkhipova, 1994: 11, figs 11–13, 20–22.
Aulodrilus japonicus Yamaguchi, 1953: 298, fig. 12,
pl. VII, figs 5–7.—Finogenova and Arkhipova, 1994: 8,
figs 3, 6, 9, 10.
Material examined. No new specimens examined.
Other Australian records. NT: Georgetown and Mudginberri Billabongs, Magela Creek, Kakadu National
Park (Brinkhurst, 1984). Qld: Yule Creek, Yanda Yarra
(Brinkhurst, 1971b).
Distribution
and
habitat.
Cosmopolitan
(Brinkhurst, 1971b) but in Australia only known
from the north so far.
Diagnosis. Length 10–25 mm. Anterior dorsal
bundles with up to 8 short hair chaetae and up to
10 bifid chaetae with the upper tooth shorter and
thinner than the lower and frequently replicated.
Anterior ventral chaetae up to 16 per bundle with
very thin, short upper teeth. Vasa deferentia long,
entering ovoid atria apically. Atria with large
lobate prostate gland, terminating in pendant
penes. Genitalia may be shifted forwards.
Remarks. All Australian specimens observed so
far have had replicate upper teeth on the dorsal
chaetae. Specimens with replicate teeth were
originally described as a separate species, Aulodrilus japonicus Yamaguchi, 1953, but this
species was synonymised with A. pluriseta by
Brinkhurst (1971a). Some Russian specimens,
identified as A. japonicus by Finogenova and
Arkhipova (1994), had replicate upper teeth, male
pores in X and there were other minor differences
in the form of the male ducts. Replicate teeth on
these specimens were shown to be lateral, occurring either side of the main teeth. These may be a
separate species, but forward shifting of the
reproductive organs is commonly associated with
asexual reproduction and other A. pluriseta
(including other specimens described by Finogenova and Arkhipova (1994) have testes as far anterior as V (Brinkhurst, 1971a). Furthermore, the
replicate teeth are essentially a form of pectination which has been shown to vary in other tubificids, including as a response to environmental
conditions (Chapman and Brinkhurst, 1986,
1987).
Aulodrilus pigueti Kowalewski
Aulodrilus pigueti Kowalewski, 1914: 625, fig. 12.—
Brinkhurst, 1971a: 526, fig. 8.23I.—Brinkhurst, 1971b:
114, fig. 2J.—Brinkhurst, 1984: 147. —Finogenova
and Arkhipova, 1994: 13, figs 14–17.
Material examined. NSW: Ryans Billabong (station
61), 36º06´S 147º58´E, 30 Aug 1978, 1 on slide
(MDFRC); Manning R. at Wingham Bridge, 31º52´S
152º23´E, 29 May 1996, 1 immature (AWT). NT: Main
stream in Compartment L of CSIRO Kapalga Research
Station, 20 Aug 1992, 10 in alcohol (NMV F77619).
Vic.: La Trobe R. below Moe Drain, AMG 8121
359769, 3 Jul 1979 and Jul 1980, 7 immature in alcohol
(NMV F77621–F77624); Steavenson R., 19 Feb 1991,
1 immature in alcohol (NMV F77625); Branjee Creek,
Goulburn Valley, 23 Apr 1992, 1 immature in alcohol
(NMV F77620). WA: Geegully Creek, Kimberley
region, 18º13´34´´S 123º44´36´´E, 5 May 1995, few
immature in alcohol (CALM); Meda R. near Lennard
R. Gorge, 17º20´58´´S 124º30´27´´E, 20 May 1996, 2
mature in alcohol (CALM).
Other Australian records. NT: Georgetown, Mudginberri and Nankeen Billabongs, Magela Creek, Kakadu
National Park (Brinkhurst, 1984). WA: Pine Hill Rock,
near Balladonia, locality uncertain (Brinkhurst, 1971b).
Qld: Pond near Tully (Brinkhurst, 1971b).
Distribution
and
habitat.
Cosmopolitan
(Brinkhurst, 1971b) and widespread in Australia,
occurring in a variety of habitats.
Diagnosis. Length 2–28 mm. Dorsal anterior bundles with 2–5 hairs and 4–5 (rarely to 10) other
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
chaetae which are either simple-pointed or bifid
with the upper teeth shorter and thinner than the
lower. Beyond VII the bifid dorsal chaetae
become oar-shaped when viewed laterally (i.e. in
the same plane as the teeth), often with rudimentary teeth visible on the end of the oar blade. Dorsal chaetae or just hair chaetae may be absent
from some or all of II–VII. Ventral bundles with
3–6 bifid chaetae, with upper teeth shorter and
thinner than the lower, not becoming oar-shaped.
Large spoon-shaped penial chaetae beside each
male pore on XI in mature specimens. Vasa deferentia short, atria bean-shaped. Pendant penes
open via a median inversion of the body wall.
Remarks. The chaetae of A. limnobius are also
oar-shaped, but more narrowly so than in A.
pigueti and the blade-like wings of the former are
in a plane perpendicular to the orientation of the
teeth rather than in the same plane.
Rhyacodrilinae Hrabe
Diagnosis. Spermathecal pores close to anterior
border of their segment (usually the testes bearing
segment, normally X). Spermathecal ampullae
containing loose or loosely bundled sperm. Atria
with prostate glands diffuse or formed into distinct glands, rarely absent. Pendant penes generally absent; protrusible or eversible pseudopenes
often present. Penial chaetae often modified,
often numerous and typically arranged fanwise or
in a row, with the blunt or bifid ectal ends close
together. Spermathecal chaetae usually not modified. Coelomocytes usually large and abundant,
rarely absent.
Remarks. Without doubt, the most problematic
Australian freshwater tubificids are the four
species placed in the Telmatodrilinae by
Brinkhurst (1971a) and Brinkhurst and Fulton
(1979) and two similar new species to be
described below. The Telmatodrilinae is diagnosed by the presence of multiple prostate glands,
each attached to the atria by short stalks. Telmatodrilus multiprostatus Brinkhurst, 1971 and Telmatodrilus pectinatus Brinkhurst, 1971 were
placed within the Telmatodrilinae because the
prostate cells are grouped into several glands,
each of which appear to have a discrete connection to the atrial lumen. Later, two other Australian species, Telmatodrilus (Alexandrovia)
papillatus Brinkhurst and Fulton, 1979 and Telmatodrilus? (Telmatodrilus?) bifidus Brinkhurst
and Fulton, 1979, were tentatively placed here for
the same reasons, although T. bifidus has only two
prostates per atrium and these are not stalked. Of
47
the two new species described below, one has
prostate tissue similar to that of T. multiprostatus
and the other has a pair of unstalked prostate
glands, like that of T. bifidus.
Prior to the description of the Australian forms,
the subfamily Telmatodrilinae consisted of three
or four northern hemisphere species in two genera
(or subgenera): Telmatodrilus Eisen, 1879 from
western USA and Canada, and Alexandrovia
Hrabe, 1962 from Alaska and Russia. This subfamily is generally considered to be closely
related to the Tubificinae, based on cladistic analyses using the few characters available (Erséus,
1990b; Brinkhurst, 1994).
The Australian species differ from the holarctic
forms in many respects. Their atria are ovoid to
pear-shaped with apical prostates, rather than
tubular with prostates along their length. In two of
the Australian species the prostates lack stalks,
instead being attached rather broadly to the anterior and posterior walls of the atria (Figs 9E, 11A,
B). The Australian species have simple male
pores, usually on the inner aspects of ventrolateral
folds of the body wall, whereas the northern
forms have pendant penes. Grooved spermathecal
chaetae are found in two of the northern species
and most of the Australian species, but all except
one of the Australian forms also have modified
penial chaetae which are absent in the northern
forms. Coelomocytes are present in the two new
species but have not been observed in the other
Australian forms and appear to be absent in the
holarctic species.
Before discussing the subfamilial relationships
of these Australian species we should indicate
that we intend to assign them to two new genera,
described below. Breviatria gen. nov. will include
B. multiprostatus, B. pectinatus, B. papillatus and
B. arvensis sp. nov. and Biprostatus gen. nov. will
include B. bifidus and B. duplex sp. nov. These
names are mentioned here to facilitate the
following discussion.
Erséus (1990b), in a cladistic analysis of the
Naididae and subfamilies of Tubificidae, showed
the two subfamilies Tubificinae and Telmatodrilinae (based on the northern species) as sharing
plesiomorphic character states for modified spermathecal chaetae, absence of penial chaetae,
unmodified oesophagus, absence of multiple
large granular coelomocytes, lack of atrial ciliation, and absence of posterior prostates. Tubificinae and Telmatodrilinae share the apomorphic
states for penes and spermatozeugmata present.
While these subfamilies share the presence of
stalked prostates, the former has one per atrium,
the latter many, and the analysis suggested that
48
A. M. PINDER AND R. O. BRINKHURST
the stalked prostates of these two groups had
‘evolved independently from the diffuse prostate
condition in the tubificid ancestor’. These subfamilies are also separated in that analysis by the
apomorphic state of the spermathecal pore position in midsegment in Tubificinae (anterior in the
Telmatodrilinae). Erséus (1990b) found that the
telmatodrilines and tubificines formed a sister
clade to another containing the phallodrilines,
marine and freshwater rhyacodrilines and the
Naididae, which share the presence of numerous coelomocytes and penial chaetae. The codings in Erséus’ analysis depend on the coding of
the outgroup (Phreodrilidae) adopted and are not
necessarily accepted here.
We reran the analysis with the Australian forms
as a separate group (AUS) using PAUP 3.1 and
the original options. AUS was coded as lacking
spermatozeugmata, atrial cilia and penes, but possessing coelomocytes, penial chaetae and multiple prostates (the coding was thus 10100 01101
000). Three equally parsimonious trees of 24
steps (CI 0.708) were obtained (unpublished
data). In all of these, the Australian species appear
within a clade which also included Phallodrilinae
and marine and freshwater Rhyacodrilinae, separate from a Tubificinae-Telmatodrilinae-Limnodriloidinae clade. This was predictable because of
coding for the presence of coelomocytes and
penial chaetae, and implies that the multiple
prostates of AUS are a convergent development
to those of the (northern) Telmatodrilinae. Within
this group, AUS was associated with the Phallodrilinae and marine Rhyacodrilinae, rather than
the freshwater Rhyacodrilinae and associated
Naididae, because of the lack of hair chaetae and
the apical vasa deferentia.
Brinkhurst’s (1994) cladistic analysis had the
broader aim of investigating the evolutionary
relationships within the whole of the Clitellata
and the characters and choice of states used
reflected this. Again, Telmatodrilinae was coded
based on the northern species, on the assumption
that these were plesiomorphic within the subfamily. A clade including Tubificinae, Telmatodrilinae and Limnodriloidinae, recognised by possession of spermathecal chaetae (not unique) and
stalked prostates, loss of penial chaetae (presence
of which was considered plesiomorphic for the
family), and organised bundles of sperm in the
spermathecae, separated from a clade containing
the Rhyacodrilinae and Naididae. As discussed
below, the Phallodrilinae was associated with
the tubificine clade rather than that including
Rhyacodrilinae.
This analysis was also rerun using PAUP 3.1
and the original options and weightings and with
all characters unordered. AUS was coded as lacking spermatozeugmata but having coelomocytes,
penial chaetae and stalked prostates (the coding
was thus 10110 11111 50010 01000 012). Twelve
trees of equal length (88, CI 0.568) were produced (unpublished data). In this rerun, AUS
appeared within a clade with the Tubificinae,
northern Telmatodrilinae, Phallodrilinae and
Limnodriloidinae, which formed a sister group to
the clade including Rhyacodrilinae and Naididae.
The appearance of AUS and Phallodrilinae with
the northern Telmatodrilinae, Tubificinae and
Limnodriloidinae, rather than with the Rhyacodrilinae (as in Erséus’ study), appears to result
from the fact that this analysis saw penial chaetae
as plesiomorphic for the whole family and that the
prostates were coded differently by Erséus.
The lack of consistency in the above analyses
requires that we consider some of the character
states in detail.
Prostate. Traditionally, the prostate tissue of
Rhyacodrilinae has been described as diffuse
(e.g., Brinkhurst, 1971a), meaning that the
prostate cells form a continuous layer over the
atrium, with each cell communicating with the
atrial lumen independently. More recently, this
view (which focused on the freshwater forms) has
been modified following the description of
numerous marine forms with prostate tissue that
is more clumped. Thus, Baker and Brinkhurst
(1981) describe the Rhyacodrilinae as “having
prostate glands forming a diffuse glandular covering; the cells communicating with the atrial lumen
independently or in small clusters”. Erséus (1984)
describes the subfamily as having “Prostates diffuse or irregularly scattered in two or more clusters of cells broadly attached to the atrial surface,
or absent”. Some rhyacodrilines, including the
marine Heronidrilus Erséus and Jamieson, 1981
and Heterodrilus Pierantoni, 1902, the freshwater
monotypic Macquaridrilus Jamieson, 1968 and
the Lake Baikal endemic Rhyacodriloides
Chekanovskaya, 1975, have some or all of the
prostate cells formed into two or more glands
(Erséus, 1981; Erséus and Jamieson, 1981;
Jamieson, 1968; Martin and Brinkhurst, 1998).
There are therefore precedents for including Breviatria and Biprostatus in Rhyacodrilinae in
respect to the form of the prostates. Although
most of the rhyacodrilines mentioned above have
the prostate clumps broadly attached to the
atrium, rather than narrowly stalked, the same is
true for the two Biprostatus species. Furthermore,
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
some Heterodrilus (H. inermis (Erséus, 1981) and
H. rapidensis Erséus, 1997) do have prostate
glands that narrow into stalks like Breviatria. The
Phallodrilinae is the only other subfamily to have
more than one prostate gland per atrium and so
should be mentioned here. Like Biprostatus, phallodrilines have a pair of prostates on each atrium,
but these are almost always stalked and typically
placed ental and ectal in position on the atria
(Erséus, 1992), unlike the symmetrically placed
prostate tissue of Biprostatus. A few phallodrilines (Pacifidrilus Erséus, 1992 and Pirodrilus
Erséus, 1992) do have the prostates placed symmetrically on the atria but this appears to be a
consequence of the small size of the atria. There
is a need for more knowledge on the histology
and morphogenesis of prostate tissue in various
tubificids to more adequately use prostate
characters in phylogenetic work, as highlighted
by Gustavsson and Erséus (1997).
Coelomocytes. An abundance of coelomocytes
(a rhyacodriline trait) has been found in both of
the new species described below but their presence in the older Australian taxa cannot be established because of the condition of the scarce
material. They have recently been observed in
rhyacodrilines such as Torodrilus Cook, 1970 and
Rhizodrilus Smith, 1900, in which they were previously thought to be absent (Erséus, 1994).
Coelomocytes have not been observed in Rhyacodriloides, but that genus is nonetheless now associated with the Rhyacodrilinae (Baker and
Brinkhurst, 1981; Martin and Brinkhurst, 1998).
Coelomocytes have not been reported for any of
the northern telmatodrilines (Holmquist, 1974)
and, with few exceptions, are not abundant in the
Phallodrilinae (Erséus, 1992).
Genital chaetae. Groove-tipped genital chaetae
are common in the Tubificinae and Limnodriloidinae and some northern Telmatodrilinae. These
are generally few in number (usually 1 or 2 per
bundle) and lie in enlarged chaetal glands, usually
on the spermathecal segment. A few rhyacodrilines, some Rhizodrilus and Rhyacodriloides, do
have similar chaetae (Baker and Brinkhurst,
1981; Erséus, 1990a; Martin and Brinkhurst,
1998), although these are on the male pore segment in the latter. The superficially similar
chaetae of the rhyacodriline Torodrilus, appear to
be a separate apomorphic development (Erséus,
1994). More typical in the Rhyacodrilinae and
Phallodrilinae is modification of the penial
chaetae associated with the male pores. These
have bifid or simple (often club-shaped) tips,
arranged in a row or fanned out with the tips close
49
together, always on the male pore segment.
Chaetae of the male pore segment are not modified in the northern Telmatodrilinae and only
rarely modified in the Tubificinae (two Varichaetadrilus Brinkhurst and Kathman, 1983 and
the few species of Krenedrilus Dumnicka, 1983).
Biprostatus and Breviatria are unusual within the
family in possessing both grooved chaetae on the
spermathecal segment (absent in two Breviatria)
and modified penial chaetae on the male pore segment (absent in one Breviatria). Krenedrilus
(Tubificinae) and some Rhizodrilus (Rhyacodrilinae) are the only other tubificids to have both
types of genital chaetae. This means the genital
chaetae may be of little help in classifying the
new genera, unless grooved spermathecal chaetae
are seen as plesiomorphic for the family (almost
identical chaetae do occur in the Phreodrilidae)
and modified penial chaetae are seen as an apomorphic development in the ancestral rhyacodriline, as tentatively suggested by Erséus (1990b).
In that case, the presence of penial chaetae in the
new genera might suggest their inclusion within
the Rhyacodrilinae.
Somatic chaetae. Somatic chaetae are generally
difficult to use in higher oligochaete taxonomy
because of a lack of consistency among and
within taxa. Both the northern telmatodrilines and
Breviatria and Biprostatus have some unusual
chaetae, but there are no chaetal features that
would indicate a close relationship between the
Australian and northern species. Neither do the
chaetae provide evidence of a relationship to any
other particular group of tubificids.
Sperm. Some Breviatria and Biprostatus
appear to have sperm in the spermathecae
arranged into numerous bundles with the nuclei
aligned at one end, others have the sperm cells in
loose masses, both arrangements typical of the
Rhyacodrilinae and Phallodrilinae. The spermatozeugmata more typical of the Tubificinae, with
fertile sperm encapsulated by infertile sperm
(Braidotti and Ferraguti, 1982; Ferraguti et al.,
1989; Ferraguti et al., 1994), are absent. Of the
northern telmatodrilines, Alexandrovia species
have some kind of sperm bundles (Holmquist,
1974), which require ultrastructural analysis to
investigate homology with tubificine spermatozeugmata, and Telmatodrilus have loose sperm
in the spermathecae. This character is thus
ambiguous in terms of relationships.
Spermathecal pores. In general, tubificines
have spermathecal pores near the middle of the
segment (Brinkhurst, 1991) while rhyacodrilines
and phallodrilines have these pores at or just
50
A. M. PINDER AND R. O. BRINKHURST
behind the anterior septa (Baker and Brinkhurst,
1981; Erséus, 1992). All Breviatria, at least one
of the two Biprostatus, and the northern telmatodrilines have anterior spermathecal pores but,
since this may be the plesiomorphic state for the
family (Erséus, 1990b; Brinkhurst, 1991), it indicates little about the relationship between these
worms.
Penes. Pendant penes may constitute an apomorphy for tubificines and northern telmatodrilines (Erséus, 1990b; Brinkhurst, 1994), although
they are also present in some Phreodrilidae.
Absence of penes in Biprostatus and Breviatria
may be a plesiomorphic state shared by rhyacodrilines and phallodrilines.
In summary, the argument that Breviatria and
Biprostatus belong within the Telmatodrilinae
simply because they have prostate cells formed
into multiple glands is not strong, particularly
considering the lack of knowledge about prostate
homology and the number of rhyacodrilines that
are now known to have clumped prostate tissue.
Breviatria and Biprostatus have several other features, including lack of pendant penes and spermatozeugmata and presence of penial chaetae and
coelomocytes (in at least some species) which
indicate a closer relationship with the Rhyacodrilinae. Therefore, as anticipated by the discussion of these and other problem taxa by Martin
and Brinkhurst (1998), we now propose to
consider the Australian former telmatodrilines to
be properly associated with Rhyacodrilinae,
described below as two new genera.
While this is undoubtedly not the last word on
the position of these worms, we believe that this
move represents a more parsimonious classification, pending further ontogenetic and ultrastructural analyses of some characters (especially
prostate glands and organised sperm) and further
phylogenetic work on the rhyacodrilines. It does
seem that the Rhyacodrilinae is paraphyletic
(Erséus, 1990b; Brinkhurst, 1994) and we may be
trying to force the Australian ex-telmatodrilines
into a group that has no phylogenetic reality. A
thorough study of rhyacodriline phylogenetics is
yet to be undertaken but Erséus (1990b) canvasses the possibility that the Rhyacodrilinae will
have to be split into a number of smaller groups to
achieve monophyly, perhaps with each raised to
family level. If this were to be given effect then
the Australian ex-telmatodrilines may form one
(or more) such groups.
Rhyacodrilus Bretscher
Rhyacodrilus Bretscher, 1901: 205.—Stephenson,
1930: 742.—Brinkhurst, 1971a: 544.
Diagnosis. Dorsal chaetal bundles with or without
hair chaetae. Vasa deferentia moderately long,
entering atria subapically, atria usually bulbous or
rounded, usually covered with a diffuse layer of
prostate cells. Protrusible pseudopenes present or
absent. Penial chaetae usually modified,
numerous with blunt tips held close together.
Coelomocytes usually large and abundant.
Rhyacodrilus coccineus (Vejdovský)
Tubifex coccineus Vejdovský, 1875: 193.—
Michaelsen, 1909: 32, figs 55–58.
Ilyodrilus coccineus.—Stolc, 1886: 656.
Rhyacodrilus coccineus.—Michaelsen, 1909: 31.—
Brinkhurst, 1971a: 537, figs 8.31A–D.—Brinkhurst,
1971b: 115, fig. 2K.
Branchiura pleurotheca Benham, 1907: 256, pl.
XLVI, figs 7–12.
Holotype: Type locality unknown (ZMUH 6948).
Material examined. NSW: Blue Lake, Mt Kosciusko,
36º24´24´´S 148º18´50´´E, Jun 1906, several on slides
(2 sectioned) (AMS W5612–W5613). WA: Power
Rock, Balladonia, 1 on slide, uncertain id. (ROB).
Distribution and habitat. Holarctic, Australia and
South America (Brinkhurst, 1971a; Brinkhurst
and Marchese, 1989). This species is not
commonly collected in Australia.
Diagnosis. Length 10–35 mm. Anterior dorsal
bundles with 3–5 hair chaetae and up to 5 pectinate chaetae, hairs missing in many posterior segments. Ventral chaetae 2–5 per bundle with the
upper teeth thinner and not much longer than the
lower (but longer in Australian specimens
described as B. pleurotheca). Ventral chaetae of
XI with 3–5 club-headed penial chaetae per
bundle, equal in size to somatic ventral chaetae.
Spermathecal pores ventrolateral (lateral in the B.
pleurotheca specimens). Vasa deferentia not
coiled around atria, which are globular and
covered with a diffuse layer of prostate cells.
Coelomocytes large and abundant.
Remarks. Benham (1907) described Branchiura
pleurotheca from Blue Lake, Mt Kosciusko, but
Brinkhurst (1971ab) considered this to be synonymous with R. coccineus. The long upper teeth
of the anterior ventral chaetae and the lateral position of the spermathecal pores of Benham’s material may prove to be diagnostic of a distinct
species once intra-specific variation in R. coccineus is better documented. Further specimens
from the type locality of B. pleurotheca (Blue
Lake, Mt Kosciusko) are required to substantiate
the synonymy. The Power Rock specimen is dissected but not stained and little can now be seen
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
of the genital anatomy. The most recent accounts
of species with hair chaetae are Ohtaka (1995)
and Timm (1990). The Tasmanian specimens
from Southwell R., mistaken as R. coccineus by
Pinder and Brinkhurst (1994), are attributable to
Ainudrilus fultoni Brinkhurst, 1982. Other worms
attributed to this species by Pinder and Brinkhurst
(1994) are described below as a new taxon.
Rhyacodrilus megaprostatus sp. nov.
Figure 1
Non Rhyacodrilus coccineus (Vejdovský, 1875).—
Pinder and Brinkhurst, 1994: 94.
51
Material examined. Holotype: Curdies R., Vic.,
(38º20´S 143º08´E), 7 Feb 1992, G. Quinn (MON), 1
mature on slide (NMV F81869).
Paratypes. Vic: Grassy Creek, Otway Ranges,
38º28´S 144º00´E, 6 Jun 1989, 1 mature on slide (NMV
F81868); Lake Purrumbete, 38º17´S 143º14´E, 31 Jan
1992, G. Quinn (MON), 1 mature on slide (NMV
F81866); Lake Purrumbete, no date, A. Glaister
(MON), 1 mature on slide (NMV F81867); Branjee
Creek, Goulburn Valley, 36º42´S 148º20´E, 23 Apr
1992, A. Pinder, 1 mature on slide (NMV F81870).
Distribution and habitat. All of the above localities are in central southern Victoria. Collected
from lentic and lotic sites and from a variety of
Figure 1, Rhyacodrilus megaprostatus sp. nov.: A, tips of anterior dorsal chaetae with section of hair; B, tip of ventral chaeta of II; C, tip of posterior ventral chaeta; D, atrium with prostate tissue and penial chaetae of holotype; E,
spermatheca of NMV F81868. Scales: A–C, 0.01 mm; D, 0.04 mm; E, 0.03 mm.
52
A. M. PINDER AND R. O. BRINKHURST
habitats including sandy sediment and rocks and
bricks used for colonisation studies.
Etymology: Named for the size of the prostate
gland in relation to the small atrium.
Description. Length 5–9 mm, diameter when
slide mounted about 0.3 mm. Number of
segments 40–70. Coelomocytes abundant.
Hair chaetae in preclitellar dorsal bundles normally 3–4 per bundle (1 long and 1 short in one
Lake Purrumbete specimen), 270–300 µm long.
Hairs accompanied by 3–6 pectinate chaetae,
100–110 µm long, with upper teeth slightly
shorter than lower and few pectinations (Fig. 1A).
Fewer dorsal chaetae posteriorly. Anterior ventral
bundles with 3–5 chaetae (those from Curdies R.
with as many as 7), 90–140 µm long, with upper
teeth thinner than lower, teeth about equal in
length on II (Fig. 1B), upper teeth slightly shorter
than lower posteriorly (Fig. 1C). Penial chaetae in
XI bluntly simple-pointed, 90–100 µm long, 1–3
per bundle, projecting medial to the line of the
somatic ventral chaetae (Fig. D).
Paired male and spermathecal pores in line with
ventral chaetae.
Atria small, spherical (Fig. 1D), opening into
smaller spherical chamber with a lobed pore.
Atria enclosed by a voluminous prostate gland
(Fig. 1A), concealing the vas deferens-atrial
union. Prostate not closely applied to the sides of
the atrium. Ovaries large and obscuring the vasa
deferentia. Male funnels observed in some specimens, often closely associated with the spermathecae and a part of vasa deferentia folded and
adhered to funnels. Spermathecae ovoid (Fig. 1E)
containing sperm arranged in distinct masses.
Remarks. The apparent attachment of the prostate
at the top of the atrium needs to be confirmed
from sectioned material as this may indicates that
the prostate is not as diffuse as is normal in
Rhyacodrilus. Within the genus, the large volume
of prostate tissue compared to the atria is unique
to this new species.
According to the key to Rhyacodrilus in
Brinkhurst (1971a) and notes in the update by
Brinkhurst and Wetzel (1984), R. megaprostatus
is closest to R. coccineus, from which it differs
mainly in the volume of the prostate. Several
other species with hair and pectinate chaetae have
been described more recently. In Rhyacodrilus
okamikae Giani and Rodriguez, 1988 the pectinate chaetae have short upper teeth, the ventral
chaetae have long upper teeth and there is only a
single median spermatheca. Rhyacodrilus suputensis Timm, 1990 has lyre-shaped pectinate
chaetae and tubular atria. Rhyacodrilus hiemalis
Ohtaka, 1995 has voluminous spermathecae with
narrow ducts, more numerous chaetae and a less
voluminous prostate gland.
Rhyacodrilus bifidus Brinkhurst
Rhyacodrilus bifidus Brinkhurst, 1982: 3, pl. 6.
Holotype: Blue Lake, Mt Kosciusko plateau, NSW
(QVM:14:232).
Material examined: NSW: Holotype and paratype from
type locality, 36º24´24´´S 148º18´50´´E, 7 Feb 1977, 5
on slides (QVM:14:232–233) and 7 on slides (ROB);
Lake Cootapatamba, Mt Kosciusko plateau,
37º27´58´´S 148º15´48´´E, 7 Feb 1977, 1 on slide
(ROB). Vic.: Thomson R. at Thomson Portal Rd, 3 on
slides (NMV F81671–81873); Wentworth R., 17.7 km
upstream of Tabberabbera, 37º31´S 147º23´E, 2 Oct
1975, 1 on slide, tentative identification (NMV
F81874). New Zealand: Lake Matheson, Canterbury
Highlands, 2 Nov 1978, 2 on slides (ROB); Lake Gault,
Canterbury Highlands, 5 Nov 1978, 2 on slides (ROB).
Distribution and habitat. Known only from the
above lakes and rivers of the far southeast of
mainland Australia and the lakes on the South
Island of New Zealand (Brinkhurst, 1982),
although the identification of the latter could not
be confirmed as the genitalia are no longer visible
in the specimens examined. Australian specimens
have been collected from coarse sediments in
rivers: sand and gravel, often between cobbles
and pebbles (Ahern and Blyth, 1979; Malipatil
and Blyth, 1982) but possibly in finer sediments
from the Mt Kosciusko sites (Timms, 1979).
Diagnosis. Length up to 7 mm. Ventral and dorsal bundles with 3–8 chaetae anteriorly, bifid with
upper teeth 2–3 times as long as, but thinner
than, the lower. Penial chaetae 6–10 bifid per
bundle in XI, about 2 times longer than somatic
chaetae and slightly curved ectally. Vasa deferentia as long as atria. Atria 4 times longer than
broad, ending in wide bulbs with narrow pores on
the anterior face of penis sacs which open to the
exterior via narrow pores. Abundant prostate
tissue attached to atria subapically. Coelomocytes
not abundant.
Remarks. The reproductive structures of the
Wentworth R. specimen are not visible and so the
identification cannot be confirmed. There are
only three penial chaetae per bundle in XI, but
these are large and bifid and the somatic chaetae
are mostly four or five per bundle with the upper
teeth much longer than the lower. Similarly, the
identification of the whole-mounted New Zealand
specimens could not be confirmed.
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
Ainudrilus Finogenova
Ainudrilus Finogenova, 1982: 1255.—Erséus, 1990a.
Vadicola Baker, 1982: 3232.
Type species. Ainudrilus oceanicus Finogenova,
1982.
Diagnosis. Hair chaetae present or absent.
Chaetae of penial segment modified, chaetae of
spermathecal segment slightly modified in one
species. Vasa deferentia usually wide and at least
partially granulated, entering atria subapically.
Atria variably shaped but more or less erect or
directed posteriad, usually consisting of an
ampullae, often with spacious lumen containing sperm, usually leading to thick, and often
equally large and well developed, ejaculatory
ducts, often giving the appearance of a bi-partite
atria. Prostate absent. Penes absent. Spermathecae
with distinct, and often complex ducts. Sperm
loose in ampullae. Coelomocytes large and
abundant.
Remarks. The lack of prostate on the atria distinguishes Ainudrilus from almost all other rhyacodrilines, bar some other highly unusual forms
such as Jolydrilus Marcus, 1965 and Epirodrilus
Hrabe, 1930 (see Brinkhurst, 1971a; Baker and
Brinkhurst, 1981), and one freshwater Rhyacodrilus. The latter, Rhyacodrilus simplex (Benham,
1903), lacks prostate tissue but also differs from
other Rhyacodrilus in having vasa deferentia that
coil around the atria and which join the atria apically. Further study of this species may lead to it
being moved back to a resurrected Taupodrilus,
which was established by Benham (1903) for its
reception.
Ainudrilus billabongus (Brinkhurst, 1984),
Ainudrilus stagnalis (Erséus, 1997) and the following new species and new combination are the
only freshwater representatives described so far.
There are eight marine species, from Hong Kong,
Northern Australia, the Pacific Ocean and the
Caribbean (Erséus, 1990ab, 1997; Finogenova,
1982).
Ainudrilus billabongus (Brinkhurst)
Rhyacodrilus billabongus Brinkhurst, 1984: 144,
fig. 3.
Ainudrilus billabongus.—Erséus, 1990a: 265.
Holotype. Bowerbird Billabong, Magela Creek, NT
(USNM 80698).
Material examined: NT: Paratype, from type locality, 1
on slide, 21 Aug 1981 (ROB); Magela Creek, 24 Feb
1988, 1 on slide (ERISS).
Distribution and habitat. Recorded only from
53
lentic and lotic sites of the Magela Creek system,
Northern Territory.
Diagnosis. Length at least 7.5 mm, all specimens
incomplete. Coelomocytes large and abundant.
Anterior chaetae bifid with teeth of equal length,
4–6 per bundle anteriorly, reduced in number posteriorly. Up to 6 simple-pointed penial chaetae in
each ventral bundle of XI, strongly recurved at the
tip and about 2 times longer than somatic chaetae.
Vasa deferentia wide and glandular, about twice
the length of the muscular upright atria, entering
the atria basally. Atria leading directly to small
circular pores. Spermathecal ducts long and
narrow, sharply set of from the globular ampullae.
Remarks. The original description of A. billabongus made mention of a bundle of sperm lying
outside of the atrium. Most fully mature Ainudrilus seem to maintain sperm in the atrium or
expanded ejaculatory ducts so the sperm bundle
of the holotype illustrated in Brinkhurst (1982)
was most likely extruded from the atrium during
dissection.
Ainudrilus stagnalis Erséus, 1997
Ainudrilus stagnalis Erséus, 1997: 9, figs 2E–F.
Holotype. Billabong associated with Stephens Creek,
Port Darwin West Arm, NT (NTM W00020).
Material examined. No new specimens examined.
Distribution and habitat. Known only from the
freshwater type locality, in fine sand (Erséus,
1997).
Diagnosis. Length at least 3.2 mm (only specimen
incomplete). Coelomocytes abundant. Hair
chaetae absent. Anterior chaetae bifid with upper
tooth much longer than lower, 4–6 per bundle
anteriorly, 3–4 posteriorly. A single bifid chaeta,
not larger than other ventral chaetae, near each
male pore. Vasa deferentia not described. Atria
with non-muscular ampullae and thick muscular
ejaculatory ducts terminating in simple pores
medial to the ventral chaetal line. Spermathecae
absent in the only known specimen.
Remarks. Erséus (1997) suggested that the chaeta
near each male pore is not a modified genital
chaeta, but that such might be developed at a later
stage.
Ainudrilus fultoni (Brinkhurst) comb. nov.
Figure 2
Rhyacodrilus fultoni Brinkhurst, 1982: 2, pl. 1–5.
Holotype. Lake Sorell, Tas. (QVM:14:230).
54
A. M. PINDER AND R. O. BRINKHURST
Figure 2, Ainudrilus fultoni (Brinkhurst) from Lake Sorell: A, somatic chaetae; B, penial chaetae; C, genitalia of
QVM:14:4202. Scales: A–B, 0.01 mm; C, 0.05 mm.
Material examined. Tas.: Holotype and paratypes,
type locality, several sectioned specimens and others in
alcohol, 2 Feb 1981 (QVM:14:230–231 and ROB);
Southwell R., 41º34´50´´S 145º44´04´´E, 20 Dec 1988,
2 on slides (QVM:14:4201–4202); Ouse R. at Marlborough Highway, 41º59´22´´S 146º38´43´´E, 16 Mar
1997, 1 on slide (AMP); Lake Perry, 43º13´10´´S
146º45´17´´E, 11 Mar 1997, 2 on slides, 3 in alcohol
(AMP).
deferentia enter atria subapically (Fig. 2C). Atria
consisting of a thick walled ental portion with a
narrow lumen, which leads into a thinner walled
ectal portion with a wide lumen which usually
contains sperm in mature specimens (Fig. 2C).
Spermathecae with ampullae full of loose masses
of sperm, ducts short and broad opening through
large pores (Fig. 2C).
Distribution and habitat. Known only from lakes
and rivers in Tasmania. The Ouse R. and Lake
Perry specimens were collected from sand
between cobbles in shallow water.
Remarks. Specimens from Southwell R. were
attributed to R. coccineus by Pinder and Brinkhurst (1994) but conform to the description of A.
fultoni. As these specimens are dissected, rather
than sectioned like the type material, the reproductive structures are illustrated here to give a different perspective (Fig. 2C). The atrium consists
of a thick-walled portion in which the lumen
can be seen in only one dissected specimen and a
thinner walled ectal chamber (essentially an
ejaculatory duct), with a sperm bundle within the
latter in examined specimens.
Diagnosis. Length up to 15 mm. Coelomocytes
large and fairly abundant. Chaetae all bifid with
teeth equally long, 10–13 per bundle anteriorly,
fewer posteriorly (Fig. 2A). Ventral bundles of XI
with 7–9 straight penial chaetae with blunt,
slightly curved tips, about equal in length to
somatic ventral chaetae (Fig. 2B). Short vasa
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
It seems preferable to transfer R. fultoni from
Rhyacodrilus to the expanding Ainudrilus, which
includes most species without prostate glands,
whereas all Rhyacodrilus (except R. simplex,
discussed above) have diffuse prostate tissue.
Ainudrilus fultoni can be distinguished from its
freshwater congeners by the more numerous
somatic and penial chaetae.
Ainudrilus nharna sp. nov.
Figure 3
Material examined. Holotype: From Frankland R. at
Roe Rd ford, Western Australia, 34º41´02´´S
116º51´13´´E, 9 Sep 1996, A. Pinder and R. Brinkhurst,
serially sectioned specimen on three slides (WAM
1–99).
55
Paratypes: Collection details as for holotype, 2 specimens serially sectioned on slides and others in alcohol
(WAM 2–99 to 4–99).
Other material: WA: Collection details as for types, 7
on slides, including 1 serially sectioned (AMP); Lake
Wilson at track off Scott Rd, 34º25´23´´S
115º07´52´´E, 14 Sep 1996, 2 on slides (AMP);
Thomas Spring, Augusta, 34º21´00´´S 115º09´35´´E,
17 Sep 1996, A. Pinder and R. Brinkhurst, 5 on slides
(AMP); stream crossing Thomson Rd, approximately 5
km N of Granite Peak, 34º40´48´´S 116º42´13´´E, 12
Sep 1996, A. Pinder and R. Brinkhurst, 1 on slide
(AMP); northern tributary of Collier Creek on
Cemetary Rd, 34º58´30´´S 116º45´12´´E, 11 Sep 1996,
A. Pinder and R. Brinkhurst, 1 on slide (AMP); Boyndiminup Rd, Lake Muir wetland system, WA, 34º30´S
116º35´E, Jul and Oct 1993, P. Horwitz (ECU), 3 on
slides and others in alcohol (WAM 90–97, 91–97,
Figure 3, Ainudrilus nharna sp. nov.: A, tip of dorsal chaeta; B, tip of ventral chaeta; C, tip of penial chaeta; D, genitalia, based mostly on holotype. Scales: A–C, 0.01 mm; D, 0.1 mm.
56
A. M. PINDER AND R. O. BRINKHURST
92–97, 98–97 and 99–97); Peaty seepage in logging
coup, Johnsons Rd, upper tributary of Deep R., 34º41´S
116º36´E, Jan 1993, P. Horwitz (ECU), 4 on slides
(WAM 95–97 to 97–97, 93–97); creek in sandy shrub
land at Bevan Rd, Kent R. catchment, 34º39´S
116º58´E, P. Horwitz (ECU), Oct 1993, 1 on slide
(WAM 94–97); Beedelup Brook South, 34º22´39´´S
115º56´E, Sep 1993, K. Trayler (MUR), 1 on slide
(WAM 36–98); Beedelup Brook South, 34º22´50´´S
115º55´E, Oct 1994, K. Trayler (MUR), 1 on slide
(WAM 37–98); Carey Brook at Pile Rd, 34º21´29´´S
115º54´29´´E, 16 Sep 1996, A. Pinder and R.
Brinkhurst, 1 on slide and 3 in alcohol (AMP); Lake
Walbyring, 32º50´19´´S 117º35´27´´E, 18 Dec 1996,
M. Smith (CALM), 1 on slide (WAM 38–98) and 2 on
slides (AMP).
Distribution and habitat. Ainudrilus nharna
appears to be widespread in the southwest of WA,
occurring in springs, lakes, and both flowing and
still reaches of streams and rivers. Sediments at
collecting sites ranged from peat to mud to coarse
sand. Specimens from Lake Walbyring, a brackish to saline lake, were collected in Dec 1996
when salinity in the lake was 2.8 ppt, but not in
Oct 1997 when salinity was 20 ppt (unpublished
data).
Etymology: From nharna, worm in Nhungar
aboriginal language.
Description. Length about 9 mm. Number of
segments 75–90. Width at clitellum (slidemounted worms) 0.3–0.5 mm. Coelomocytes
present.
Clitellum inconspicuous, half-X–half-XII.
Gonopores in line with the ventral chaetae, spermathecal pores anterior on X. Male pores open
into a transverse groove, lateral to the penial
chaetae in the anterior half of XI.
Anterior dorsal bundles with up to 4 hair
chaetae (up to 420 µm long) and up to 5 pectinate
chaetae (up to 130 µm long), latter with upper
teeth thinner than and equal in length to lower,
with indistinct pectinations (Fig. 3A). Ventral
bundles with bifid chaetae of similar size and
form to the dorsals (but without pectinations), 3–5
per bundle anteriorly, fewer posteriorly (Fig. 3B).
Penial chaetae 4–7 per bundle in XI, up to 120 µm
long and slightly curved ectally (Fig. 3C). Penial
chaetae protrude close to the mid-line, on either
side of the ventral nerve cord medial to the male
pores.
Testes anteroventral in X (Fig. 3D). Atria in XI,
bulbous with thick lining tissue and ciliated
lumen, leading to folded protrusible pseudopenes,
which are protruded on one of the paratypes.
Prostate tissue absent. Vasa deferentia thickened,
appearing glandular and often twisted or folded
once or twice between male funnel and atria,
attaching anteromedially to the atria (Fig 3D).
Spermathecal ampullae large and ovoid with
sperm formed into loose masses, clearly separated
from the ducts. Spermathecal ducts constricted at
one or more points, leading to pores which are
often on a protrusion within small vestibulae,
located anteriorly on X (Fig 3D). Ovaries lying
near the vasa deferentia in XI, female pores not
observed.
Remarks. The absence of prostate tissue and the
thickened vasa deferentia suggest that this new
species belongs in the genus Ainudrilus. Ainudrilus taitamensis Erséus, 1990a, of Hong Kong,
which was only tentatively included in the genus,
is the only other species to have ciliated atria. The
atria and pseudopenes resembles what was called
a two-part atria in A. fultoni and the ectal part of
the male duct is formed into pseudopenes or elaborate ejaculatory ducts in some other species
(Erséus, 1990a). The thick vasa deferentia are difficult to follow in both sectioned and dissected
worms as they are tightly folded with an indistinct
lumen and are obscured by the ovaries. Ainudrilus
taitamensis and Ainudrilus brendae Erséus, 1997,
both marine/brackish, are the only other Ainudrilus to have hair chaetae. These differ from the
new species in that the former has long tubular
atria and the latter has only two small straight
penial chaetae per bundle and ventral chaetae
with upper teeth much longer than the lower.
Rhizodrilus Smith
Rhizodrilus Smith, 1900: 44.—Baker and Brinkhurst,
1981: 952.
Monopylephorus Levinson (in part).—Brinkhurst,
1963: 62.—Brinkhurst, 1965a: 148.—Brinkhurst,
1971a: 555.
Type species. Rhizodrilus lacteus Smith, 1900.
Diagnosis. Chaetae all bifid, hairs absent. Vasa
deferentia moderately long, entering atria subapically. Atria normally tubular with diffuse prostate
tissue, terminating in ejaculatory ducts which exit
either on the inner aspect of large ventrolateral
folds in the body wall or into a median invagination of the ventral body wall (copulatory bursa),
sometimes via protrusible pseudopenes. Spermathecal chaetae and/or penial chaetae modified
in mature specimens, the former associated with
large glands. Some or all of the spermathecal genitalia in IX. Sperm in spermathecae in loose
masses or in oriented bundles. Coelomocytes
abundant.
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
Remarks. Rhizodrilus consists of a few marine
and freshwater species with apparently disparate
distributions (Baker and Brinkhurst, 1981;
Erséus, 1990a). These are united by the presence
in IX of at least some of the spermathecal genitalia. The following species was tentatively
included in Rhizodrilus by Baker and Brinkhurst
(1981).
Rhizodrilus arthingtonae (Jamieson)
Rhyacodrilus arthingtonae Jamieson, 1978.
Rhizodrilus arthingtonae.—Baker and Brinkhurst,
1981: 956.
Holotype. Brown Lake, North Stradbroke Island, Qld
(QM G8881).
Material examined: Qld: Holotype and paratype, locality as above, 31 Jul 1975, dissected on slides (QM
G8881–2).
Distribution and habitat. Known only from the
type locality, a perched acidic lake. Specimens
collected in organic material in sandy sediment
(Jamieson, 1978).
Diagnosis. Length 66 mm (paratype). Coelomocytes numerous. Anterior chaetal bundles with
3–5 bifid chaetae with the upper teeth slightly
longer than the lower, posteriorly teeth about
equal. Hair chaetae absent. The chaetae are
smaller anteriorly and posteriorly, largest in the
clitellar region. Several clearly bifid penial
chaetae per bundle in XI, 1.5–2 times longer than
somatic chaetae and slightly curved towards the
tip. Vasa deferentia narrow, coiled anteriorly,
passing through the prostate before entering atria
subapically. Atria fusiform to slenderly bulbous,
prostate tissue compact but lobulated, enclosing
the atrium and communicating with the lumen at
numerous points. Slender ejaculatory ducts lead
to male pores on the inner wall of a ventrolateral
fold in the body wall. Accessory genital markings
present, consisting of a pair of conspicuous elliptical papillae posteromedian to ventral bundles of
IX, each with a short diagonal groove passing
posterolaterally from its centre to 9/10. Spermathecae in X, with indistinct ventrolateral pores
at 9/10. Worm squarish in cross-section.
Remarks. Erséus (1994) rightly questions whether
R. arthingtonae belongs within this genus, preferring to include only those species with large tubular atria and a clear involvement of spermathecal
genitalia in IX, rather than X as is normal in the
family. R arthingtonae has only minor involvement of spermathecal genitalia in IX (just
papillae on IX with grooves leading back to the
57
spermathecal pores on 9/10). However, Rhizodrilus lacteus Smith, 1900, which has just the
spermathecal chaetae and associated glands in IX
but retains the spermathecae themselves in X is
intermediate between R. arthingtonae and other
Rhizodrilus in this respect and so R. arthingtonae
may represent one extreme of a continuum. Arguing against this proposition is the fact that the
atria are more upright and less tubular than those
of other Rhizodrilus and Ferraguti et al. (1994)
note the more plesiomorphic spermatozoal ultrastructure of R. arthingtonae compared to Rhizodrilus russus Erséus, 1990a and the related
Monopylephorus Levinson, 1884. If R. arthingtonae is excluded from Rhizodrilus, then, short of
creating a new genus, it is difficult to see a more
suitable place for it. Erséus (1994) summarises
the possible relationships between Rhizodrilus
and other relevant rhyacodriline genera and, since
we suggest no change to the status quo, these need
not be repeated here. A more satisfactory placement of R. arthingtonae may have to await further
phylogenetic studies of this and related rhyacodrilines.
Breviatria gen. nov.
Type species. Telmatodrilus multiprostatus
Brinkhurst, 1971a, as the first of two species
described in the original account.
Diagnosis. Atria ovoid to erect pear-shaped, with
unciliated lumen, bearing numerous glands of
prostate tissue apically, communicating independently with atrial lumen. Male pores simple on
XI, on inner aspect of protruding ventrolateral
folds of the body wall surrounding a broad
median copulatory chamber. Spermathecal
chaetae large and grooved, penial chaetae numerous with blunt or bifid tips. Spermathecae normally with ventrolateral to dorso-lateral pores,
located anteriorly on X. Sperm in spermathecae in
organised bundles (not spermatozeugmata) or
loose. Coelomocytes abundant for at least one
species, unknown in the others.
Etymology. Breviatria referring to the short atria,
in contrast to the tubular atria of other genera with
prostate formed into multiple clumps.
Remarks. Of the four species assigned to this new
rhyacodriline genus, three were previously placed
within the Telmatodrilinae and one is new. The
reasons for re-assigning the earlier species were
discussed above (in the Rhyacodrilinae remarks
section). Breviatria is distinguished from other
Rhyacodrilinae by the ovoid to pear-shaped atria
58
A. M. PINDER AND R. O. BRINKHURST
with prostate tissue formed into discrete glands
with separate (usually stalked) connections to the
ectal part of the atria. Other rhyacodriline genera
with clumped prostate tissue, mostly marine
forms such as Heronidrilus Erséus and Jamieson,
1981 and Heterodrilus Pierantoni, 1902, usually
have the clumps more broadly attached along the
length of elongate tubular atria, although
prostates are more restricted in position in a few
species. The other new genus, Biprostatus,
described below, also has small atria but these
have only two large prostate glands which are
broadly attached to the anterior and posterior
sides of the atria. Other rhyacodrilines with short
atria either have evenly distributed diffuse
prostate tissue (such as most Rhyacodrilus) or
lack prostate tissue (Ainudrilus and Rhyacodrilus
simplex). Rhizodrilus arthingtonae (discussed
above) also has ovoid atria but the lobes of
prostate tissue are tightly packed over the surface
of the atria and broadly attached rather than
stalked. The new species described below has
numerous coelomocytes but this feature needs to
be re-examined when fresh material of the other
species becomes available.
Breviatria multiprostatus (Brinkhurst)
comb. nov.
Figures 4, 5
Telmatodrilus multiprostatus Brinkhurst, 1971a: 535,
figs 8.27A–F.—Brinkhurst, 1971b: 117, fig. 3C.—
Brinkhurst and Fulton, 1979: 3.—Brinkhurst, 1982: 3,
pl. 7.—Holmquist, 1974: 256, figs 10–11.—Bayly,
1973: 305.—Timms, 1978: 430.—Marshall, 1975: 22,
fig. 3.—Marshall and Winterbourn, 1979: 205. The
species description was duplicated in Brinkhurst
(1971b) but the 1971a publication should be used as the
original description.
Holotype. Lake Pedder, Tas. (AMS W4178).
Material examined. Tas.: Holotype and paratypes,
locality as above, 2–4 Mar 1966 (AMS
W4178–W4182) and 1 on slide (ROB); Cuvier R., 10
Oct 1978, 1 on slides (ROB); Collingwood R., 4 Nov
1978, 1 on slides (ROB); trickle under the Needles, 11
Nov 1978, 1 on slides (ROB); creek in Frodshams Pass,
12 Nov 1978, 1 on slides (ROB); Lake Sorell, 2 Feb
1981, 1 sectioned (ROB); Crossing R., south of Lake
Pedder, southwest Tasmania, 10 Feb 1987, 1 on slide
(QVM:14:3002) and 3 in alcohol (AMP): Lake Surprise, AMG 8112 441869, 13 Nov 1976, 1 on slide
(QVM:14:3001) and 1 in alcohol (AMP). Tentatively
identified immature specimens from various sites in
Tasmania listed by Brinkhurst (1982).
Other records. Tas.: Lake Pedder (Bayly, 1973); South
Esk R. (Brinkhurst and Fulton, 1979); Lakes Sorell and
Crescent (Timms, 1978). New Zealand: Leeston Drain,
near Killinchy, 56 km S of Christchurch (Marshall,
1975; Marshall and Winterbourn, 1979); Freshwater R.,
Stewart Island (Cowie et al., 1978).
Distribution and habitat. Recorded from lakes,
streams and rivers in Tasmania and possibly from
New Zealand. The Lake Crescent specimens were
collected from a variety of sediment types,
including sand, detritus and mud (Timms, 1978).
Specimens from Great Lake were collected at
depths of up to 16.5 m (Fulton, 1983b).
Description. Length up to 16 mm. Pharyngeal
glands present on either side of septa 2/3 to 4/5,
gut expanded in VII to form a diverticulum which
completely encircles the gut in VII and protrudes
forward into VI (Fig. 4A).
Somatic chaetae all simple-pointed (Fig. 5A),
up to 8 per bundle anteriorly, fewer posteriorly.
Ventral chaetae missing on penial segment of
mature specimens, or if present then not modified.
Spermathecal chaetae (Fig. 5B) larger than
somatic chaetae, with extensive musculature,
projecting into an enlarged chaetal sac posteriorly on X, into which also empties a separate
elongate gland (Fig. 4C). In one paratype, the
spermathecal chaetal sac opens anteriorly in X,
with the chaetal shaft protruding well anteriad
into IX (Fig. 4B).
Testes anteroventral in X, with sperm funnels
on 10/11 feeding broad vasa deferentia which
wind upwards to enter atria apically (Figs 4B, C).
Atria elongate pear-shaped, may be folded over
dorsally, with unciliated lumen and thick lining
tissue. Atria narrow basally, opening on inner
aspect of folds in the body wall lateral to a median
ventral copulatory chamber. Prostate cells
grouped into numerous glands which narrow
basally and connect to apical third of atria (Figs
4B, C, Fig. 5C). Spermathecae large in X with
short ducts that lead to ventral pores just behind
septa 9/10 (or medially in X in the Lake Sorell
specimen). Sperm loose in ampulla. Ovaries
attached to anterior septa of XI near sperm
funnel. Female funnels large, leading to pores at
intersegmental furrow 11/12 (Figs 4B, C).
Remarks. This is the only Australian tubificid to
have all chaetae simple-pointed. However, many
aquatic Enchytraeidae also have multiple simplepointed chaetae in all bundles and immature
forms could be confused with B. multiprostatus.
Of the Tasmanian records, the only identifications
that can be confirmed by re-examination of specimens are those from Lake Pedder and Lake
Sorell. Specimens from the other sites listed
above are either immature (and so are tentatively
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
59
Figure 4, Breviatria multiprostatus (Brinkhurst): A, anterior of specimen from Lake Sorell; B, genitalia of paratype
AMS W4182; C, genitalia of Lake Sorell specimen. Scales: A–C, 0.15 mm.
60
A. M. PINDER AND R. O. BRINKHURST
Figure 5, Breviatria multiprostatus (Brinkhurst): A, typical somatic chaetae; B, bent spermathecal chaeta of holotype; C, atria dissected from holotype. Scales: A, 0.02 mm; B, 0.05 mm; C, 0.2 mm.
identified from the chaetae only) or their identification can no longer be verified because of their
poor condition. It is not known whether identifications of specimens from New Zealand were
based on the genitalia of mature specimens or just
the chaetae of immature worms so the identification must be considered tentative. One of the New
Zealand records (Leeston Drain) is unusual in that
the site was enriched with septic tank leachate and
dairy effluent (Marshall and Winterbourn, 1979)
whereas the Australian records are all from nonenriched sites, including minimally disturbed
alpine streams.
The position of the spermathecal pores and
spermathecal chaetae appears to vary. Paratypes
W4180 and W4182 (Fig. 4B) both have spermathecal pores anterior in X, but spermathecal
chaetae posterior and anterior (projecting forward
into IX) in X respectively. The Lake Sorell specimen has spermathecal chaetae posterior in X but
the spermathecal pore medial on X (Fig. 4C). All
other features of these worms seem similar and
we consider the variation in spermathecal characters to be intraspecific for now. The holotype is
dissected rather than sectioned and the original
position of the spermathecae is unclear.
Breviatria pectinatus (Brinkhurst) comb. nov.
Figure 6
Telmatodrilus pectinatus Brinkhurst, 1971a: 535, figs
8.25D–H.—Brinkhurst, 1971b: 117, fig. 3D.—Bayly,
1973: 305.—Holmquist, 1974: 257, fig. 12.—Timms,
1974: 276.—Timms, 1978: 435. The species description was duplicated in Brinkhurst (1971b) but the 1971a
publication should be used as the original description.
Holotype. Lake Pedder, Tas. (AMS W4176).
Material examined. Tas.: Holotype and paratypes, Lake
Pedder, 2 Mar 1966, 2 on slides (AMS W4176–7) and
5 on slides (ROB).
Other Australian records. Tas.: Lake Pedder (Bayly,
1973); Lake St Clair (Timms, 1978). Vic.: Lake Tali
Karng (Timms, 1974).
Distribution and habitat. Recorded only from the
above lakes in Victoria and Tasmania. Bayly
(1973) collected this species from Lake Pedder
(in 1972) in sandy sediment but the Lake Tali
Karng specimens were recovered from mud at a
depth of 12 m.
Description. Length unknown, but greater than 10
mm.
Anterior bundles with 9–14 chaetae, upper
teeth longer than the lower and slightly hooked.
Posterior bundles with 5–8 pectinate chaetae with
upper teeth not so long (Fig. 6A). Spermathecal
chaetae single, thin with grooved tips (Fig. 6B),
up to twice as long as somatic ventral chaetae.
Penial chaetae (Fig. 6C) modified with rough,
blunt to bifid tips (absent on paratype W4176),
about 1.5 times longer than somatic ventral
chaetae.
Vasa deferentia broad, coiled once or twice
before entering atria apically (Fig. 6D). Atria
pear-shaped with numerous prostates with short
stalks distributed around the upper half. Stalks
enveloped by the atrial muscle tissue. Individual
peritoneal cells form diffuse cover over basal half
of atria and some present apically (Fig. 6D). Atria
narrowing to pores on the inner aspect of lateral
folds in the body wall around a shallow median
ventral copulatory chamber. Details of the
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
spermathecae difficult to decipher but possibly
quite folded, with lateral or dorsolateral pores and
containing loose sperm.
Remarks. Pectinate chaetae are more common in
61
the Tubificinae than the Rhyacodrilinae and the
former usually has pectinations more developed anteriorly, rather than posteriorly as in
B. pectinatus.
Figure 6, Breviatria pectinatus (Brinkhurst): A, tips of pectinate dorsal chaetae of holotype; B, spermathecal chaeta
of holotype; C, penial chaetae of holotype; D, male ducts and spermathecal chaetae of paratype AMS W4177.
Scales: A, 0.01 mm; B–C, 0.015 mm; D, 0.05 mm.
62
A. M. PINDER AND R. O. BRINKHURST
Breviatria papillatus (Brinkhurst and Fulton)
comb. nov.
Figure 7
Telmatodrilus (Alexandrovia) papillatus Brinkhurst
and Fulton, 1979: 2, figs 9–14.—Fulton, 1983a: 778.—
Fulton, 1983b: 792.
Holotype. Great Lake, Tasmania (QVM:14:320 and
321).
Material examined. Tas.: Holotype and paratypes, type
locality, 26 Sep 1975, 3 on slides (QVM:14:320, 321,
326 and 327) and 1 on slide (ROB); Arthurs Lake, 4 Jul
1977, 8 on slide, 2 mature and several immature in alco-
hol (NMV F88870, F88871 and ROB); Lake Sorrel, no
date, 4 on slides (QVM:14:322–325).
Distribution and habitat. Recorded only from the
above lakes of the central plateau of Tasmania.
Collected in both sandy and clayey sediments, at
depths up to 17.5m (Fulton, 1983b).
Description. Length 10–20 mm. Coelomocytes
not observed. Worms encrusted with fine foreign
material, body wall papillate with several rings of
papillae per segment (Fig. 7A). Prostomium and
segment I (and sometimes II) may be retracted
into the body. Posterior segments narrow and
Figure 7, Breviatria papillatus (Brinkhurst and Fulton): A, view of body showing copulatory bursa on XI; B, tip of
anterior chaeta; C–D, tips of transitional chaetae; E, tip of posterior hair chaeta; F, male ducts of NMV F88870; G,
spermatheca of NMV F88870. Scales: A, 0.2 mm; B–E, 0.01 mm; F, 0.05 mm; G, 0.025 mm.
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
appearing highly contracted in the preserved
specimens.
Anterior chaetae bifid, 5–7 per bundle, with
each tooth broad and flattened (Fig. 7B). Chaetae
changing to a hair-like form, some with vestiges
of teeth (Figs. 7C, D) between VII and X so that
postclitellar bundles have 5–9 curved hair chaetae
(Fig. 7E). Penial chaetae slightly shorter than preclitellar somatic chaetae, 3–5 per bundle with
bifid tips (Fig. 7F). Spermathecal chaetae not
modified.
Long vasa deferentia connect to short pearshaped atria subapically (Fig. 7F). Atria narrow
gradually to enter small median copulatory chamber (which is everted in some preserved worms)
near the penial chaetae which project from the
protruding lateral walls of the chamber. Three or
more lobes of prostate tissue on atria apically
(Fig. 7F). Spermathecae small and globular
(Fig. 7G) with short ducts leading to lateral
pores anteriorly on X. Sperm not observed in
ampullae.
Remarks. While the chaetae and papillae of B.
papillatus are unusual, the genitalia suggest that
this species is simply an apomorphic member of
Breviatria and to exclude it would leave Breviatria paraphyletic. Specimens from Arthurs Lake,
some of which were dissected and illustrated for
this study (Fig. 7), clearly have spermathecae
with lateral pores whereas spermathecae were not
seen in the type specimens from Great Lake. The
spermathecae of these specimens are very small
(100–150 µm) and do not contain sperm, so may
not be fully developed. Unfortunately, the type
specimens have become too cleared for this feature to be checked. The holotype (the only type in
which the genitalia can be seen) has ‘at least 2 or
3 prostate glands’ (Brinkhurst and Fulton, 1979),
but the size of these and their attachment to the
atria is uncertain due to clearing and distortion
during slide mounting. The prostate on one atrium
of the holotype appears to be formed into two or
three large, stalked lobes but on the other atrium
the prostate appears to be a more cohesive mass
with a number of short connections. The prostate
tissue of the Arthur R. specimens is clearly
formed into numerous small glands but connections to the atria are obscured. Fresh specimens
from the type locality are required to determine
the unity of this taxon.
63
Marchant (NMV) and A. Pinder, dissected on slide
(QVM:14:3878).
Paratypes: Collection details as for holotype, 1
mature and 1 immature sagitally halved on slide, with
posterior portion of 1 of these in alcohol, and 1 immature whole-mounted on a slide (QVM:14:3879–3881).
Distribution and habitat. Known only from the
type locality, collected from sand/silt under
cobble at a depth of 10–20 cm.
Etymology: Named arvensis for the type locality.
Description. Length at least 20 mm (none complete), width of slide mounted worms 0.25–0.3
mm. Prostomium squarish in preserved type
material. Coelomocytes abundant but without
consistent shape, entirely filling coelom in most
of body, including the prostomium, making the
worms appear quite solid when handled.
All chaetae bifid, 55–85 µm long, with upper
teeth longer than lower and nodulus slightly ectal
(Fig. 8A). Up to 8 chaetae per bundle anteriorly,
reduced in number posteriorly. Chaetae of spermathecal segment (X) present but not modified,
penial chaetae of XI 4 per bundle, straight, with
bifid ectal ends and no nodulus (Fig. 8B),
150–165 µm long, projecting somewhat diagonally towards the midline to open into median
copulatory chamber (Fig. 8C).
Testes anteroventral in X. Sperm funnels large,
vasa deferentia wide and slightly coiled near funnel. Point of union of atria and vasa deferentia not
established. Atria ovoid, thin walled, with short
ejaculatory ducts leading to pores on inner aspect
of folds in body wall lateral to a median ventral
copulatory chamber (Fig. 8C). Prostate tissue present as numerous discreet glands, which presumably connect with atria individually, although this
could not be confirmed. Spermathecae bulbous
with short ducts leading to dorso–lateral pores
anteriorly on X. Sperm in ampullae formed into
numerous bundles. Ovaries anteroventral in XI,
female pores not observed.
Remarks. Although the connections between the
atria and prostate tissue are obscured in the available specimens, the numerous separate glands are
similar to those of other Breviatria. The long
straight penial chaetae with bifid tips also resemble those of other Breviatria. The bifid chaetae of
B. arvensis, with their long upper teeth, differ
from those of all other Breviatria but resemble
those of Biprostatus duplex, described below.
Breviatria arvensis sp. nov.
Figure 8
Biprostatus gen. nov.
Material examined. Holotype: Arve R. at Arve Rd,
43º11´20´´S 146º46´05´´E, Tasmania, 12 Mar 1997, R.
Diagnosis. Atria small and spherical, bearing
broadly attached prostate tissue formed into
64
A. M. PINDER AND R. O. BRINKHURST
Figure 8, Breviatria arvensis sp. nov.: A, tip of ventral chaeta; B, tips of penial chaetae; C, male genitalia of holotype. Scales: A–B, 0.01 mm; C, 0.06 mm.
two large glands, one anterior and one posterior. Atria leading to simple pores on inner
aspects of folds in body wall, lateral to a
median ventral copulatory chamber. Spermathecal chaetae large and grooved, penial
chaetae multiple and bluntly bifid. Spermathecal pores lateral, sperm in ampullae in loose
bundles. Coelomocytes abundant in anterior segments in at least one species, to be confirmed for
the rest.
Type species: Telmatodrilus bifidus Brinkhurst
and Fulton, 1979.
Remarks. The type species of Biprostatus was
originally placed within the telmatodriline genus
Telmatodrilus, but is transferred to this new rhyacodriline genus for reasons already discussed
above (see remarks for Rhyacodrilinae). This
genus is separable from most others in the Rhyacodrilinae by virtue of the small atria with large
bilobate prostate glands. Although some Heterodrilus, such as H. inermis and H. rapidensis, also
have short atria with large paired prostate glands,
these glands appear to be stalked and the chaetae
of Heterodrilus (1 or 2 per bundle, at least some
of which are trifid) are diagnostic (Erséus, 1981).
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
Biprostatus bifidus (Brinkhurst and Fulton)
comb. nov.
Figure 9
Telmatodrilus ? (Telmatodrilus ?) bifidus Brinkhurst
and Fulton, 1979: 3, figs 15–19.
Holotype. Great Lake, Tas. (QVM:14:328).
Material examined. Tas.: Holotype and paratypes,
Great Lake, 1975, 10 on slides (QVM:14:328–339);
Arthurs Lake, 4 Apr and 22 May 1977, 4 on slides
(ROB).
Distribution and habitat. Recorded only from the
above lakes of the central plateau of Tasmania.
Collected in both sandy and clayey sediments, at
depths up to 17.5m (Fulton, 1983b).
Diagnosis. Length up to 25 mm. Anterior bundles
with up to 13 bifid chaetae with upper teeth
longer than the broad lower. (Fig. 9A) Number
of chaetae diminishing from VIII–X with
65
upper teeth becoming shorter than the lower after
the clitellar region (Fig. 9B). Chaetae of some
species have groove-like markings (ornamentations) ectally. Spermathecal chaetae
straight, single, with grooved tips (Fig. 9C),
varying in width and length, from shorter than to
3 times longer than somatic chaetae. Penial
chaetae bifid (Fig. 9D), up to 7 per bundle, twice
as long and thick as somatic ventral chaetae.
Atria small, elongate ovoid with short ejaculatory
ducts (Fig. 9E). Each atrium with a pair of
broadly attached prostate glands, one anterior,
which envelops the vas deferens, the other
posterior, occupying much of the coelom (Fig.
9E). Male pores and penial chaetae on the lateral
walls of a large median copulatory chamber.
Spermathecae ovoid, with short duct-like extension entally and lateral pores. Sperm in ampullae
formed into loose bundles. Coelomocytes not
recorded.
Figure 9, Biprostatus bifidus (Brinkhurst and Fulton): A, tip of anterior chaeta; B, tips of posterior chaetae; C, tips
of spermathecal chaetae of Arthurs Lake specimen; D, tips of penial chaetae of holotype; E, genitalia of
QVM:14:338. Scales: A–B, 0.015 mm; C, 0.015 mm; D, 0.01 mm; E, 0.12 mm.
66
A. M. PINDER AND R. O. BRINKHURST
Biprostatus duplex sp. nov.
Figures 10, 11
Telmatodrilus sp. 2.—Pinder and Brinkhurst, 1994.
Material examined. Holotype: Que R., just below Que
R. Mine dam, Tasmania, AMG 8014 914956, 27 Apr
1987, Wayne Fulton and Brett Mawbey (IFC), transverse serially sectioned specimen (with the posterior
whole-mounted) on 15 slides (QVM:14:4207).
Paratypes: Collection details as for holotype, 5 serially sectioned and 3 mounted whole or dissected on
slides, (QVM:14:4208, 4210–4215, 4218); Que R. at
Murchison Highway, Tasmania (AMG 8014 901963),
27 Apr 1987, W. Fulton and B. Mawbey (IFC), 2 serially sectioned and 2 whole-mounted on slides
(QVM:14:4209, 4216, 4217, 4223).
Other material. Tas: Southwell R., above Hellyer
mine, AMG 8014 945960, 20 Dec 1988, S. Chilcott
(IFC), 1 on slide, 5 in alcohol (AMP); Douglas Creek,
AMG 8114 206692, 16–17 Feb 1990, S. Chilcott and L.
Mellefont (IFC), 1 on slide (AMP); Creek at Charlies
Hill, AMG 8111 297938, 19 Feb 1990, B. Mawbey
(IFC), 2 in alcohol (AMP).
Distribution and habitat. The above sites are all
rivers in western Tasmania.
Etymology: Named duplex for the double prostate
gland.
Description. Length up to 20 mm. Prostomium
short, bluntly conical. Coelomocytes large,
abundant in a few preclitellar segments.
Anterior chaetae 3–7 per bundle (Figs 10A, C),
fewer posteriorly, nodulus slightly distal, upper
teeth much longer than lower. Posterior chaetae
2–4 per bundle, nodulus clearly distal, upper teeth
also much longer than lower. Chaetae shortest in
II (75 µm) and posteriorly, otherwise about 110
µm. Spermathecal chaetae single in X and/or IX,
140–160 µm long and grooved ectally (Figs 10D,
E), associated with large glands (Fig. 10G), protruding from the body wall in line with somatic
ventral chaetae. Penial chaetae 4–7 per bundle,
straight with notched ectal ends (Fig. 10F), no
nodulus, 120–160 µm long. Penial chaetae lie
medial to atria and project somewhat diagonally
toward midline to open into median depression of
the body wall.
Sperm funnels large, vasa deferentia thin,
slightly coiled at first, then running posteriad to
enter the small globular atria subapically. Atria
with prostate tissue formed into large anterior and
posterior lobes (Fig. 11B). Short ejaculatory ducts
lead to pores on inner aspect of folds in body wall
lateral to a median copulatory chamber (Figs
11A, B). Thick glandular tissue present ventrally
on either side of 10/11, more so in X (Figs 10G,
11B). Spermathecae oval (Fig. 10G), with short
thick-walled ducts and pores antero-laterally in X.
Sperm in ampullae formed into bundles.
Remarks. This species is similar to B. bifidus in
many respects. The genitalia, bifid or notched
penial chaetae and anterior somatic chaetae are
similar in the two species, although the latter are
less numerous in B. duplex. They differ in that the
upper teeth of the posterior somatic chaetae of B.
duplex are longer than the lower (like Breviatria
arvensis) whereas the upper teeth are shorter than
the lower on the posterior chaetae of B. bifidus.
The tips of the spermathecal chaetae also appears
to differ.
Bothrioneurum Stolc
Bothrioneurum Stolc, 1886: 647.—Michaelsen,
1900: 54.—Stephenson, 1930: 752.—Brinkhurst,
1971a: 539.
Type species. Bothrioneurum vejdovskyanum
Stolc, 1886.
Diagnosis. All chaetae bifid, hair chaetae absent.
Prostomium with a middorsal sensory pit. Vasa
deferentia short to long. Atria tubular and covered
with diffuse prostate gland cells except for ectal
portion, leading to protrusible pseudopenes bearing diverticula (paratria) with accessory glands.
Sperm stored within external spermatophores,
attached to body wall. Spermathecae absent.
Penial chaetae may be modified. Coelomocytes
large and abundant.
Bothrioneurum vejdovskyanum Stolc
Bothrioneurum vejdovskyanum Stolc, 1886: 647.—
Michaelsen, 1900: 54.—Brinkhurst, 1971a: 540, figs
8.29A–D.—Brinkhurst, 1982: 1.
Material examined. NSW: North Katoomba sewage
treatment plant constructed wetlands, 10 Aug 1994,
tentative identification of immature worms (AWT).
Other Australian records. NSW: Barrington R.,
Gloucester (Brinkhurst, 1982). Tas.: Arve R.
(Brinkhurst, 1982).
Distribution
and
habitat.
Cosmopolitan
(Brinkhurst, 1971a) but uncommon in Australia.
Diagnosis. Length 28–35 mm. Anterior bundles
with 4–6 bifid chaetae with upper teeth longer
than lower, posteriorly fewer with teeth equally
long. Ventral bundles of XI with 4 slightly
hooked, club-headed penial chaetae arranged in a
fan. Vasa deferentia long, joining tubular atria
apically. Atria covered with prostate gland,
except for terminal portion, and leading to protrusible pseudopenes which bear paratria with
accessory glands. Spermathecae absent. Body
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
67
Figure 10, Biprostatus duplex sp. nov.: A, ventral chaeta of II; B, ventral chaeta of other anterior segments; C, two
posterior chaetae; D, enlarged tip of spermathecal chaeta of QVM:14:4214; E, whole spermathecal chaeta of
QVM:14:4214; F, tips of penial chaetae of QVM:14:4214; G, transverse section through segment X, composite
from several sections of holotype. Scales: A–C, E and F, 0.01 mm; G, 0.05 mm.
68
A. M. PINDER AND R. O. BRINKHURST
Figure 11, Biprostatus duplex sp. nov.: A, transverse section through segment XI, composite from several sections
of paratype QVM:14:4223; B, reconstruction of genitalia from several specimens. Scales: A–B, 0.05 mm.
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
wall may appear spotted or papillate, but smooth.
Prostomium with dorsal sensory pit, often difficult to detect but visible as a small indentation
when a slide mounted specimen viewed laterally.
Remarks. The only mature specimens examined
are from Barrington R. and these matched the
description of the species, except that the prostomial pit was not observed. All other identifications are of immature specimens based on
the presence of the prostomial pit and the form of
the somatic chaetae. An electron micrograph
of the prostomial pit is provided by Brinkhurst
and Gelder (1991).
Branchiura Beddard
Branchiura Beddard, 1892.—Michaelsen, 1900:
40.—Stephenson, 1930: 749.—Brinkhurst, 1971a: 562.
Type species. Branchiura sowerbyi Beddard,
1892.
Diagnosis. Hair chaetae present. Vasa deferentia
short, joining atria medially. Atria cylindrical,
covered in prostate cells. A glandular diverticula
(paratria) attached to atria where the latter join
large eversible pseudopenes. Coelomocytes not
conspicuous. Dorsal and ventral gill filaments
present posteriorly.
Branchiura sowerbyi Beddard
Branchiura sowerbyi Beddard, 1892: 325, pl. XIX,
figs 1–15.—Michaelsen, 1900: 40.—Stephenson, 1930:
750.—Brinkhurst, 1971a: 563, figs 8.36D–F.—
Brinkhurst, 1971b: 114, fig. 2H.—Brinkhurst, 1982:
1.—Hogg and Norris, 1991:515.—Timms, 1981: 188.
Kawamuria japonica Stephenson, 1917: 89, figs 1–5.
Holotype. Royal Botanical Gardens, London (Typus
amissus).
Material examined. NSW: Parramatta R., 17 Oct 1981
(AMS W197365); Wingecarribee R. at Greenstead, 14
Nov 1991, 2 in alcohol (AWT); Murray R. at Dora
Dora, 24 Mar 1994 (MDFRC); Megarritys Creek,
Fernsworth Avenue, 3 Feb 1993, 1 in alcohol (AWT);
Rushy Billabong, 16 Jun 1981, 1 on slide (MDFRC);
Ryans Billabong, 27 Jul and 6 Dec 1982, 1 on slide, 1
in alcohol (MDFRC). Qld: Leichhardt R., Coolullah
H.S., 115 km NW Cloncurry, 25 Apr 1988, 1 on slide
(SAM E2814). Vic.: junction of Thomson and MacAlister R.s, 31 Jun 1979, 1 in alcohol (NMV F77640);
Mitta Mitta R., 1 Feb 1975, 2 Dec 1980 and 3 Mar
1982, 5 on slides, 3 in alcohol (NMV F77633–8). WA:
Herdsman Lake, 2 Nov 1989, 1 on slide (MUR); Wungong Brook, Jan 1998, 1 two-tailed worm in alcohol
(AMP).
Other Australian records. ACT: Murrumbidgee R.
(Hogg and Norris, 1991). Qld: Brookfield, Goldcreek
(Brinkhurst, 1971a). Vic.: Thomson R. (Brinkhurst,
69
1982); Lake Purrumbete (Timms, 1981). WA: Lake
Claremont and Lake Monger, Perth (Brinkhurst,
1971b).
Distribution
and
habitat.
Cosmopolitan
(Brinkhurst, 1971a), widespread in Australia,
occurring in a wide range of environments.
Diagnosis. Length 30–185 mm. Dorsal anterior
chaetal bundles with 1–4 (or more) short hair
chaetae and 5–12 (or more) chaetae that vary
from simple-pointed to bifid with short upper
teeth (which may be replicated). Posteriorly hairs
fewer and shorter and non-hair chaetae with less
replication of upper teeth. Ventral bundles with
6–11 bifid chaetae with upper teeth shorter than
lower, even simple-pointed anteriorly. Gills and
genital anatomy as for genus.
Other tubificid taxa
Timms (1978) records the tubificine Peloscolex
sp. (and ‘Peloscoiex’, undoubtedly a misprint)
from Lake St Clair, based on identifications by K.
V. Naidu. These refer to some form of papillate
tubificid, though presumably differing in chaetal
morphology from Breviatria papillatus. These
should not be considered definitive generic identifications since body wall papillae are characteristic of, but not diagnostic for, Peloscolex Leidy,
1850, now revised as Spirosperma Eisen, 1879,
and other genera (Brinkhurst, 1979; Holmquist,
1978, 1979). We have seen several papillate
immature worms from Victoria (including specimens from the Plenty, Glenelg, Curdies and La
Trobe Rivers) and New South Wales (Georges
R.) that have chaetae very similar to some Spirosperma, but mature specimens are required for
identification.
The species coded Telmatodrilus sp. 1 in Pinder and Brinkhurst (1994) cannot yet be described
from the few poorly preserved specimens. This
species, which may be a Biprostatus, has anterior
chaetae bifid (like those of B. duplex), posterior
chaetae simple pointed, and has modified penial
and spermathecal chaetae. Specimens have been
collected from Que and Southwell Rivers in
northwest Tasmania.
Numerous other undescribed tubificids, mostly
from Tasmania and Western Australia, have been
accumulated by the senior author since the preparation of this manuscript and no doubt others
await discovery.
Zoogeography
Discussion of tubificid biogeography in the
region is limited by the low number of specimens
70
A. M. PINDER AND R. O. BRINKHURST
that have been examined from Australia, New
Zealand and Asia. No area of Australia has been
thoroughly surveyed, although projects are underway to improve coverage of particular areas such
as the southwest of Western Australia, Tasmania
and South Australia. Nonetheless, some patterns
can be discerned.
Twenty-nine species of Tubificidae, in 12 tubificine and rhyacodriline genera, have been
recorded from Australian inland waters so far
(Table 1). Of these, 16 species are restricted to
Australia and two more are known only from
Australia and New Zealand.
The nonendemic component consists of 11
common species within widespread genera (i.e.
the tubificine genera Tubifex, Limnodrilus, Aulodrilus and Potamothrix and the rhyacodrilines
Rhyacodrilus, Branchiura and Bothrioneurum).
Several of these species also occur in New
Zealand (Brinkhurst and Fulton, 1979, 1982;
Marshall, 1975; Timms, 1982). Of the cosmopolitan species, only Aulodrilus pigueti, Aulodrilus
pluriseta and Branchiura sowerbyi have been
recorded from northern Australia (the Queensland
records of Limnodrilus and Tubifex are from the
southeast of the state), but this may reflect the low
number of worms examined from this region. So
far, records of Aulodrilus pigueti have been
restricted to northern Australia, but other cosmopolitan species are widespread in southern
Australia and seem to occur in a wide range of
habitats. The endemic species appear to have
more restricted distributions.
One of the three endemic genera, Antipodrilus,
is widespread, although most of the species are
not. Antipodrilus davidis, with the new range
extensions into Western Australia and South Australia, is the most widely distributed and is now
known from across southern Australia and New
Zealand. By contrast, A. magelensis appears to be
Table 1. Species list of Tubificidae known from Australia. * = known only from Australia, # = known
only from Australia and New Zealand, #? = New Zealand identification dubious, others cosmopolitan.
NSW includes Australian Capital Territory, n = north, sw = southwest.
Subfamily
Genus
Species
NT
Tubificinae
Tubifex
Limnodrilus
Limnodrilus
Limnodrilus
Potamothrix
Antipodrilus
Antipodrilus
Antipodrilus
Antipodrilus
Antipodrilus
Aulodrilus
Aulodrilus
Aulodrilus
Rhyacodrilinae Rhyacodrilus
Rhyacodrilus
Rhyacodrilus
Ainudrilus
Ainudrilus
Ainudrilus
Ainudrilus
Rhizodrilus
Breviatrius
Breviatrius
Breviatrius
Breviatrius
Biprostatus
Biprostatus
Bothrioneurum
Branchiura
tubifex
hoffmeisteri
udekemianus
clapareianus
bavaricus
magelensis*
davidis#
timmsi*
multiseta*
plectilus*
pluriseta
pigueti
limnobius
coccineus
megaprostatus*
bifidus#
billabongus*
stagnalis*
nharnus*
fultoni*
arthingtonae*
multiprostatus#?
pectinatus*
papillatus*
arvensis*
bifidus*
duplex*
vejdovskyanum
sowerbyi
Qld
Australian state
WA(n) WA (sw) SA
●
●
●
●
●
●
●
●
●
●
●
●
●
●
NSW
Vic.
Tas.
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●?
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
TUBIFICID OLIGOCHAETES FROM AUSTRALIA
a species of the north and interior, A. timmsi is
known only from the south-east mainland while
A. plectilus and A. multiseta have not been
recorded outside a few sites in Tasmania. All
species show a preference for lentic habitats.
Both species of the new genus Biprostatus are
known only from Tasmania. The other new
genus, Breviatria, has four species, three of which
are apparently restricted to Tasmania. One of
these, Breviatria multiprostatus, may also occur
in New Zealand (Cowie et al. 1978; Marshall and
Winterbourn, 1979) but the identifications are
uncertain. The fourth, Breviatria pectinatus, is
known from a few lakes in Tasmania and
Victoria.
Other endemic rhyacodrilines belong to genera
with disjunct global distributions. Rhizodrilus
mostly consists of marine species with restricted
distributions, often limited to single records from
the type localities (Baker and Brinkhurst, 1981;
Erséus, 1990a). The only Australian species, Rhizodrilus arthingtonae, is known only from one
lake on North Stradbroke Island, Queensland
(Jamieson, 1978). Ainudrilus mostly consists of
coastal Pacific or Caribbean marine species
(Erséus, 1990ab; 1997), except for the four freshwater Australian species. Of these, A. stagnalis
and A. billabongus have been recorded only from
the Northern Territory, A. nharna only from
southwest Western Australia and A. fultoni only
from Tasmania.
Of the 18 species restricted to Australia and
New Zealand, ten occur in Tasmania and eight are
known only from that state (including most
species of the new rhyacodriline genera). Similarly, about half of the 26 species of Australian
Phreodrilidae are restricted to Tasmania (Pinder
and Brinkhurst, 1997). By contrast, only three
tubificids (Rhyacodrilus megaprostatus, Rhyacodrilus bifidus and Antipodrilus timmsi) and one
phreodrilid (Phreodriloides notabilis Benham,
1907) are known only from southeast mainland
Australia. This is despite the examination of
worms from twice as many localities (and
locality-dates) from Victoria and New SouthWales as from Tasmania.
Of the tubificids known only from Tasmania,
three (A. plectilus, A. multiseta and B. bifidus) are
known only from Great Lake and Arthurs Lake on
the eastern edge of the central plateau, and one (B.
papillatus) is known only from these lakes and
nearby Lake Sorell. While not wishing to place
too much emphasis on this, because oligochaete
records from other lakes are limited, it is interesting to note that two phreodrilids are also known
only from these lakes (Pinder and Brinkhurst,
71
1997). In addition, numerous other invertebrates
and fish are apparently restricted to these lakes,
though mostly to Great Lake (Frankenberg, 1974;
Fulton, 1983ab; Invertebrate Advisory Committee, 1994; Timms, 1985; Williams, 1974). These
lakes pre-date the late pleistocene glaciations that
led to the creation of most other lakes in Tasmania and lie outside earlier glacial influence
(Davies, 1974). Sutherland et al. (1973) and
Sutherland (1980) suggest that lacustrine environments may have existed in the Great Lake basin
since the late Oligocene when basalt flows
blocked the upper Ouse R.. The age of Great Lake
has been seen as a factor contributing to local
endemicity in its fauna (Timms, 1985). Other
ancient lakes, such as Lake Baikal (Russia), Lake
Ohrid (Europe) and Lake Tanganyikae (Africa)
also have numerous endemic oligochaetes,
whereas this is rare in more recent lakes (Martin,
1996). The few Tasmanian lakes of recent (Pleistocene/Holocene) origin from which oligochaetes
have been identified appear to have only species
that are widespread in the state. These include
pre- and post-impoundment Lake Pedder as
reported by Lake (1998), a few glacial lakes
(Dobson, Dove and St. Clair) surveyed by Timms
(1978) and a dune barrage lake in the north-east
(Blackman’s Lagoon, sampled by the first
author). However, records from many other Tasmanian lakes are required to confirm the apparent
endemism of the oligochaeta fauna of Great Lake,
Arthurs Lake and Lake Sorell.
Insufficient localities have been examined from
other parts of Australia to generalise about
regional diversity. The Phreodrilidae appear to be
quite diverse in the southwest of Western Australia (Pinder and Brinkhurst, 1997 and unpublished data) but only one endemic tubificid is
known from there so far. Three tubificids (Ainudrilus stagnalis, Ainudrilus billabongus and
Antipodrilus magelensis) are known only from
inland and northern Australia so far, but so few
localities have been examined in these regions
that it is difficult to speculate what further work
will reveal.
The Australian Tubificidae seem to have a different biogeographic history to the apparently
Gondwanan Phreodrilidae (Pinder and Brinkhurst, 1997). Antipodrilus appears most similar to
genera that are widespread in the northern hemisphere but absent in Africa and South America
(Brinkhurst, 1991). The new rhyacodriline genera, Breviatria and Biprostatus, are of uncertain
affinity as the phylogenetic relationships are
unclear in this possibly paraphyletic subfamily.
Other endemic rhyacodrilines are members of a
72
A. M. PINDER AND R. O. BRINKHURST
cosmopolitan freshwater genus (Rhyacodrilus
spp.) or appear to be allied to marine forms
(Ainudrilus spp.) or are only tentatively assigned
to a widespread genus whose monophyly is
uncertain (Rhizodrilus arthingtonae). As almost
nothing is known of the oligochaete fauna of
southeast Asia and New Zealand, little comment
can be made about affinities within the
Australasian region.
Acknowledgements
We express our gratitude to all those who have
provided specimens for this study. The work has
been funded mostly by grants from the Australian
Biological Resources Study. Richard Marchant of
Museum Victoria made available additional funding and has been extremely generous with all
aspects of support needed to complete this study.
Histological work was funded by the Queen Victoria Museum and Art Gallery’s Plomley Foundation and serial sections were prepared by Gordon
Thomson of Murdoch University. Dr M. Ferraguti
provided access to an unpublished manuscript.
Brenton Knott and Peter Davies kindly shared
their knowledge of Tasmanian lakes. Andrew
Boulton, Diana Jones and Stuart Halse have
allowed the first author to use their laboratories
and facilities during the last few years.
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Note in proof:
Recent survey work by Paul McEvoy (Australian
Water Quality Centre) has revealed the presence
of the cosmopolitan tubificids Potamothrix
bavaricus, Aulodrilus limnobius, Aulodrilus pigueti and Branchiura sowerbyi in South Australia.