Syst Parasitol (2007) 67:25–42
DOI 10.1007/s11230-006-9068-0
ORIGINAL PAPER
The taxonomic status of Opegaster Ozaki, 1928
and the description of four new species of Opecoelus Ozaki,
1925 (Digenea: Opecoelidae) from marine teleosts
in Australian waters
Thelma O. Aken’Ova
Received: 17 August 2005 / Accepted: 17 April 2006 / Published online: 2 December 2006

Springer Science+Business Media B.V. 2006
Abstract The similarities between Opecoelus
Ozaki, 1925, Coitocaecum Nicoll, 1915, Opegaster
Ozaki, 1928 and Paropecoelus Pritchard, 1966 and the
difficulty of separating Opecoelus and Opegaster are
discussed. It is proposed that Opegaster be reduced to
synonymy with Opecoelus and the diagnosis of the
latter amended to accommodate both forms. Four
new species of Opecoelus are described from marine
teleosts in Australian waters. These are Opecoelus
woolcockae n. sp. from Acanthopagrus butcheri and
A. australis from off South Australia, New South
Wales and southern Queensland, O. pomatomi n. sp.
from Pomatomus saltatrix off New South Wales,
O. crowcrofti n. sp. from Atherinomorus ogilbyi off
southern Queensland and O. queenslandicus n. sp.
from Apogon fasciatus off southern Queensland. The
following new combinations are formed: Opecoelus
gonorhynchi (Gavrilyuk, 1979) n. comb., O. elongatus
(Yamaguti, 1959) n. comb., O. pentadactylus (Manter,
1940) n. comb., O. apogonichthydis (Yamaguti, 1938)
n. comb., O. cameroni (Caballero & Caballero, 1969)
T. O. Aken’Ova (&)
Department of Microbiology and Parasitology,
The University of Queensland, Brisbane, Queensland
4072, Australia
e-mail: akenovat@yahoo.com
Present Address:
T. O. Aken’Ova
Department of Biological Sciences, Ahmadu Bello
University, Zaria, Kaduna State, Nigeria
n. comb., O. dendrochiri (Yamaguti, 1970) n. comb.,
O. hawaiiensis (Yamaguti, 1970) n. comb., O. jamu-
nicus (Srivastava, 1968) n. comb., O. longivesiculus
(Yamaguti, 1952) n. comb., O. mastacembalii (Har-
shey, 1937) n. comb., O. mehrii (Harshey, 1937) n.
comb., O. synodi (Manter, 1947) n. comb., O. tamori
(Yamaguti, 1938) n. comb., O. bothi (Yamaguti, 1970)
n. comb., O. caulopsettae (Manter, 1954) n. comb.,
O. beliyai (Pande, 1937) n. comb., O. brevifistulus
(Ozaki, 1928) n. comb., O. cryptocentri (Yamaguti,
1958) n. comb., O. dactylopteri (Yamaguti, 1970) n.
comb., O. dermatogenyos (Yamaguti, 1970) n. comb.,
O. ditrematis (Yamaguti, 1942) n. comb., O. gobii
(Yamaguti, 1952) n. comb., O. hippocampi (Shen,
1982) n. comb., O. iniistii (Yamaguti, 1970) n. comb.,
O. lobulus (Wang, 1977) n. comb., O. macrorchis
(Yamaguti, 1938) n. comb., O. parapristipomatis
(Yamaguti, 1934) n. comb., O. pritchardae (Over-
street, 1969) n. comb., O. syngnathi (Yamaguti, 1934)
n. comb., O. lutiani (Bravo-Hollis & Manter, 1957) n.
comb., O. ovatus (Ozaki, 1928) n. comb., O. plotosi
(Yamaguti, 1940) n. comb. and O. rectus (Ozaki, 1928)
n. comb.; all the new combinations were previously
species of Opegaster.
Introduction
In this paper, four new species of Opecoelus
Ozaki, 1925 are described from Australian marine
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fishes, using a new concept of the genus. The four
species were recovered from two species of
Acanthopagrus and one each of Pomatomus,
Atherinomorus and Apogon. All but one of the
new species were recovered from a single host
species.
Materials and methods
Fish were caught in Moreton Bay off the coast of
south-east Queensland, off New Brighton, New
South Wales and off the South Australian coast.
They were dissected fresh and any worms recov-
ered were washed in saline, fixed in hot saline and
preserved in 5% formalin. Whole-mounts were
prepared by staining worms in Mayer’s acid hae-
matoxylin. Sagittal sections (7lm) were prepared
and stained in haematoxylin and eosin. The
stained whole-mounted worms and sections were
dehydrated in ethanol, cleared in methyl salicy-
late and mounted in Canada balsam. The whole-
mounted worms were measured using a calibrated
ocular micrometer. Drawings were prepared with
the aid of a drawing tube mounted on an Olym-
pus BH2 microscope. Measurements in micro-
metres are indicated as a range followed by the
mean in parentheses.
Abbreviations: QM, Queensland Museum,
Brisbane, Australia; BMNH, Natural History
Museum, London.
Family Opecoelidae Ozaki, 1925
Subfamily Opecoelinae Ozaki, 1925
Genus Opecoelus Ozaki, 1925
Syn. Opegaster Ozaki, 1928
Opecoelus Ozaki, 1925 was erected for
O. sphaericus Ozaki, 1925 by Ozaki (1925). He
also included a second new species in the genus.
In his account, Ozaki implied that Opecoelus was
the first digenean to be reported with what he
called a true anus. Prior to the discovery of
Opecoelus, most known digeneans had caeca that
terminated blindly and, in one instance, caeca
that opened into the excretory vesicle to form a
uroproct. The unique features of Opecoelus
formed the basis for the erection of the family
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Syst Parasitol (2007) 67:25–42
Opecoelidae and the subfamily Opecoelinae.
Ozaki (1925) commented on the close similarity
between Opecoelus and another opecoeline
genus, Coitocaecum Nicoll, 1915. The similarities
have been discussed under the latter genus by
Aken’Ova & Cribb (1996).
Opecoelus was first defined properly by Ozaki
(1928). As well as defining the genus, Ozaki de-
scribed two more new species of Opecoelus and
erected a new genus, Opegaster Ozaki, 1928,
which is very similar to Opecoelus. Although
Ozaki did not comment on the similarity between
Opecoelus and Opegaster, this has been the sub-
ject of wide comment. Manter (1940) separated
them on the basis of the anterior extent of the
vitelline follicles, the posterior extent of the
seminal vesicle and the form of the ventral sucker.
He noted the existence of some intermediate
forms and the fact that the presence or absence of
papillae on the ventral sucker was not considered
a generic character. In the definition of Opegaster,
Ozaki (1928) described the ventral sucker as
having ‘rudimentary marginal finger-like papillae
or none’. Manter (1954) remarked that, although
Opegaster and Opecoelus are very close, it was
convenient to retain the two genera, because of
the large number of species involved. He sug-
gested an arbitrary separation. Manter (1954)
recognised 15 species of Opecoelus and presented
a key for their identification. Crowcroft (1947)
was in favour of considering Opecoelus and
Opegaster as synonyms. He remarked that using
the distribution of the vitelline follicles and
the size of the seminal vesicle to distinguish
Opecoelus from Opegaster was unsatisfactory.
Other workers who have commented on the
similarity of the two genera include Banerjee
(1965), Mehra (1966), Prudhoe & Bray (1973),
Cribb (1985), Bray (1987), Shimazu (1988) and
Bray & Cribb (1989). Banerjee (1965) separated
the two genera using the distribution of the
vitelline follicles, whereas Mehra (1966) used the
degree of protrusion of the ventral sucker. Prud-
hoe & Bray (1973) noted that, as more species of
each of the two genera were described, their
distinctness became less apparent because their
features graded into one another. They also felt
that, of all the features used by different authors
to separate the genera, the most satisfactory was
Syst Parasitol (2007) 67:25–42
the distribution of the vitelline follicles. Although
several workers have commented to the effect
that Opecoelus and Opegaster were almost indis-
tinguishable morphologically, none of them ex-
cept Cribb (1985) made a move to formally merge
them. Both genera were retained most probably
for convenience, since they each contain large
numbers of species, as observed by Manter
(1954). Cribb (1985) suggested that Opegaster be
reduced to synonymy with Opecoelus and pro-
posed a comprehensive definition to include both
forms. In his definition of Opecoelus, Cribb (1985)
addressed the inconsistence in the anterior extent
of the vitelline follicles, the presence or absence
of papillae on the ventral sucker and the variation
in the number of papillae. Shimazu (1988) con-
curred with Cribb (1985) after he re-examined
Yamaguti’s (1940) type-series. He stated that it
was practically impossible to distinguish Opegas-
ter from Opecoelus. Bray & Cribb (1989) did not
follow the actions of Cribb (1985) and Shimazu
(1988) because the vitelline follicles of the species
of Opegaster they examined did not show the type
of distribution characteristic of species of Ope-
coelus. Cribb (2005), in his key to the Opecoeli-
dae, retained Opecoelus and Opegaster as
separate genera, because this situation ‘is so en-
trenched in the literature’, but also considered
that it was ‘not likely to reflect a phylogenetic
distinction’.
Opecoelus is also very similar to Paropecoelus
Pritchard, 1966, another opecoeline erected by
Pritchard (1966) for Opecoelus-like species with
either peripheral or peripheral and apertural
ventral sucker papillae and with a pronounced
predilection for hosts of the family Mullidae.
Opecoelus is here recognised in the sense of
Cribb (1985) and Shimazu (1988) and considered
to include ‘opecoelids with caeca uniting and
opening through one anus, an antero-sinistral
genital pore, with vitelline follicles restricted to
the hindbody or extending into the forebody and
a ventral sucker with three or more pairs of
apertural papillae’. Apertural papillae are absent
in a few species assigned to the genus, following
the definition for Opegaster that was given by
Ozaki (1928). Yamaguti (1971) listed 27 species
of Opecoelus and 30 of Opegaster.
27
Four forms of Opecoelus, each represented by
more than one specimen, were recovered from
the marine fish examined and are described
below.
Opecoelus woolcockae n. sp.
Type-host: Acanthopagrus butcheri (Munro)
(Sparidae).
Other host: Acanthopagrus australis (Günther)
(Sparidae).
Type-locality: Harriet River, South Australia,
3556¢S, 13739¢E.
Other localities: Off S.E. Qld: Hope Island,
2753¢S, 15323¢E; Wynnum 2727¢S, 15310¢E;
Amity Point 2724¢S, 15326¢E and Point Lookout
2726¢S, 15332¢E, Stradbroke Island; off NSW:
Iluka 2924¢S, 15321¢E; New Brighton 2831¢S,
15333¢E.
Site: Intestine.
Material studied: Ex Acanthopagrus butcheri: 17
from 8 hosts from the Harriet River, S.A., 8
December, 1995. Ex A. australis: From Moreton
Bay, Qld.: 3 from 2 hosts from off Hope Island,
January & February, 1993; 1 from off Amity
Point, January, 1994; 1 from off Point Lookout,
November, 1994; 1 from off Wynnum, February,
1995. NSW: 4 from 3 hosts, 4 from 2 hosts from
off New Brighton, April, June 1993; 2 from 1 host,
April 1994.
Type-material: Holotype: QM G225326, paratypes
(from A. butcheri): QM G225327-36, BMNH
2006.2.1.16-17; vouchers (from A. australis) QM
G225337-56, BMNH 2006.2.1.18-28.
Etymology: This species is named for V. Woolcock
in recognition of her contribution to the taxonomy
of the Digenea.
Description (Figs. 1–6)
[Based on 16 gravid unflattened, whole-mounted
specimens and 1 set of serial sagittal sections from
Acanthopagrus butcheri. Some measurements and
ratios from 10 specimens from Acanthopagrus
australis are given in brackets. Measurements
used in the following description are of 10
specimens from A. butcheri.]
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1 2

cs

pp

SV

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 Syst Parasitol (2007) 67:25–42
b Figs. 1–3 Opecoelus woolcockae n. sp. ex Acanthopagrus
butcheri. 1. Whole-mount, ventral view. 2. Ventral view of
terminal genitalia. 3. Ventral view of ventral sucker.
Abbreviations: cs, cirrus-sac; pp, pars prostatica; sv,
seminal vesicle. Scale-bars: 1, 250 lm; 2,3, 100 lm
Body elongate, robust, subcylindrical, tapering
to narrowly rounded anterior tip, broadly rounded
posteriorly; maximum width in region of ventral
sucker and gonads, 988–1,543 (1,334) · 232–352
(301) [1,161–1,425 (1,757) · 252–504 (379)]; width
to length ratio 1:4.1–4.8 (4.4) [3.8–5.4 (4.7)]. Oral
sucker ventrally subterminal, spherical to sub-
spherical, 81–112 (102) · 92–123 (111) [108–171
(136) · 108–162 (133)]. Ventral sucker on short
peduncle, spherical to subspherical, bearing 3
papillae each on anterior and posterior lips; papil-
lae interlock when ventral sucker aperture is closed
(Fig. 3), 110–143 (132) · 110–154 (134) [126–207
(174) · 135–225 (179)]; sucker-width ratio 1: 1.1–
1.3 (1.2) [1:1.1–1.6 (1.3)]. Forebody with 2 promi-
nent clusters of large gland-cells, 189–377 (287)
long, 17.8–25.7 (21.1) [19.4–30.8 (24.2)] % of body
length. Prepharynx distinct, short, usually entirely
dorsal to oral sucker, sometimes partly posterior to
oral sucker. Pharynx large, 59–72 (66) · 64–83 (76)
[68–108 (84) · 63–99 (83)], well developed,
spherical to sub-spherical; pharynx to oral sucker
width ratio 1:1.2–1.6 (1.5) [1:1.3–2.0 (1.6)].
Oesophagus distinct. Intestinal bifurcation 16–76
(36) anterior to anterior margin of ventral sucker.
Caeca long, unite near posterior extremity to open
through single ventrally subterminal anus.
Testes 2, in middle third of body, tandem,
usually entire, occasionally deeply incised at
posterior margin (n = 1), usually contiguous,
sometimes separate (n = 4), may be spherical,
sub-spherical or irregular; anterior 81–145
(118) · 88–160 (133); posterior 97–167
(137) · 101–157 (133). Post-testicular area 264–
471 (378), 23.3–31.0 (28.2) [26.8–36.4 (31.9)]%
body length. Cirrus-sac distinct, small, encloses
anterior portion of male terminal genitalia, ante-
ro-sinistral to intestinal bifurcation, 52–70
(59) · 20–33 (27). External seminal vesicle large,
saccular, 85–159 (118) · 43–67 (53), reaches var-
ious distances into hindbody. Pars prostatica
long, surrounded by gland-cells; distal extremity
enclosed within cirrus-sac. Ejaculatory duct short,
29
not clearly demarcated from pars prostatica.
Genital atrium distinct. Genital pore antero-
sinistral to intestinal bifurcation, about half way
between posterior margin of pharynx and pos-
terior limit of oesophagus, 168–240 (205) from
anterior end, 13.7–17.5 (15.4) [13.3–19.1 (16.1)]%
of body length.
Ovary pre-testicular, entire, indented at ante-
rior margin or not, usually contiguous with ante-
rior testis, sometimes separated (n = 2), 42–70
(61) · 51–118 (93). Mehlis’ gland distinct, occu-
pies areas anterior and antero-dorsal to ovary.
Uterine seminal receptacle present, occupies
proximal loops of uterus. Laurer’s canal present.
Uterine coils usually occupy area between ante-
rior margin of ovary and posterior margin of
seminal vesicle, sometimes reach to posterior
margin of ventral sucker. Metraterm distinct,
feebly developed, overlaps left caecum, sur-
rounded by gland-cells. Eggs operculate, 55–86
(67) · 21–42 (32) [54–77 (69) · 27–45 (38)].
Vitelline follicles usually restricted to hind-body,
extend from 318–534 (424) from anterior
extremity, 21.0–38.8 (32.4)% of body length, to 7–
22 (14) from posterior extremity; fields confluent
dorsally and ventrally in post-testicular area; fol-
licles lie mostly lateral and ventral to caeca, with
very few to none present dorsally; anterior extent
usually just anterior to, level with or just posterior
to posterior margin of seminal vesicle, sometimes
extends more anteriorly to anterior margin of
ventral sucker on one or both sides (n = 2),
occasionally to level of ventral sucker aperture
(n = 1).
Excretory pore more or less terminal, posterior
to anus. Excretory vesicle I-shaped, long, tubular;
anterior limit at point antero-dorsal to posterior
margin of ovary.
Comments
The known species of Opecoelus can be separated
into morphological groups A to J, based on the
anterior extent of the vitelline follicles, the pos-
terior extent of the seminal vesicle and the nature
of the testes and the ovary. It should be noted
that, although these features have been used in
morphological classification, some of them have
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4 5

cs

pp

SV

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 Syst Parasitol (2007) 67:25–42
bFigs. 4–6 Opecoelus woolcockae n. sp. ex Acanthopagrus
australis. 4. Whole-mount, ventral view. 5. Ventral view of
terminal genitalia. 6. Ventral view of ventral sucker.
Abbreviations: cs, cirrus-sac; pp, pars prostatica; sv,
seminal vesicle. Scale-bars: 4,6, 250 lm; 5, 100 lm
been known to show variation; some of the spe-
cies might therefore fall into more than one of the
groups indicated below. Species below listed as
new combinations have been transferred from
Opegaster.
Group A. Species with vitelline follicles posterior
to the anterior margin of the ventral sucker, a
seminal vesicle extending into the hindbody, en-
tire testes and a lobed ovary: O. bohaiensis Li,
Qiu & Zhang, 1988; O. goniistii Yamaguti, 1938;
O. lateolabracis Yamaguti, 1958; O. nipponicus
Yamaguti, 1951; O. ozakii Layman, 1930;
O. pacificus Caballero y C. & Caballero R.,
1976; O. rhadinotus Manter, 1963; O. sebastodis
Yamaguti, 1934 and O. sphaericus Ozaki, 1925.
Group B. Species with vitelline follicles posterior
to the anterior margin of the ventral sucker,
seminal vesicle extending into the hindbody, en-
tire testes and an entire ovary: O. adsphaericus
Manter & Van Cleave, 1951; O. arii Wang, 1982;
O. himezi Yamaguti, 1951; O. inimici Yamaguti,
1934; O. lotellae Manter, 1954; O. mexicanus
Manter, 1940; O. minor Yamaguti, 1934;
O. mulloidichthydis Yamaguti, 1970; O. noblei
Banerjee, 1965; O. sebastici Yamaguti, 1958; O.
variabilis Cribb, 19851. and O. zhifuensis Qui & Li
in Shen & Qiu, 1995.
Group C. Species with vitelline follicles posterior
to the anterior margin of the ventral sucker, a
seminal vesicle extending into the hindbody, lo-
bed to incised/irregular testes and a lobed ovary:
O. lobatus Ozaki, 1925; O. mutu Yamaguti, 19402
and O. platycephali Yamaguti, 1970.
Group D. Species with vitelline follicles posterior
to the anterior margin of the ventral sucker,
seminal vesicle not extending into hindbody,
lobed testes and a lobed ovary: O. atlanticus
1
The anterior extent of the vitelline follicles and the
posterior extent of the seminal vesicle vary in O. variabilis
2 (Yamaguti, 1952) n. comb.; O. hippocampi (Shen,
Manter (1954) and Pritchard (1966) considered O. mutu
a synonym of O. lobatus but the former was recognised by
Yamaguti (1971).
31
Zhukov, 1983; O. gonorhynchi (Gavrilyuk, 1979)
n. comb.; O. pagrosomi Yamaguti, 1958; O.
scorpaenidicola Prudhoe & Bray, 1973 and O.
xenistii Manter, 1940.
Group E. Species with vitelline follicles posterior
to the anterior margin of the ventral sucker, a
seminal vesicle not extending into hindbody, en-
tire testes and an entire ovary: O. acutus Manter,
1954; O. elongatus (Yamaguti, 1959) n. comb.; O.
kuhliae Yamaguti, 1970; O. minimus Tubangui,
1928; O. pentadactylus (Manter, 1940) n. comb.
and O. pteroisi Shen, 1986.
Group F. Species with vitelline follicles whose
anterior limit is posterior to the anterior margin
of the ventral sucker, a seminal vesicle not
extending into the hindbody, lobed testes and an
entire ovary: O. tasmanicus Crowcroft, 1947.
Group G. Species with vitelline follicles extending
anterior to the anterior margin of the ventral
sucker, a seminal vesicle extending into the hind-
body, entire to irregular testes and an entire ovary:
O. apogonichthydis (Yamaguti, 1938) n. comb.; O.
cameroni (Caballero & Caballero, 1969) n. comb.;
O. dendrochiri (Yamaguti, 1970) n. comb.; O. ha-
waiiensis (Yamaguti, 1970) n. comb.; O. jamunicus
(Srivastava, 1968) n. comb.; O. longivesiculus
(Yamaguti, 1952) n. comb.; O. mastacembalii
(Harshey, 1937) n. comb.; O. mehrii (Harshey,
1937) n. comb.; O. synodi (Manter, 1947) n. comb.
and O. tamori (Yamaguti, 1938) n. comb.
Group H. Species with vitelline follicles extend-
ing anteriorly beyond the anterior margin of the
ventral sucker: O. bothi (Yamaguti, 1970) n.
comb. and O. caulopsettae (Manter, 1954) n.
comb.
Group I. Species with vitelline follicles extend-
ing anteriorly beyond the anterior margin of the
ventral sucker, a seminal vesicle not extending into
the hindbody or just into the hindbody, entire to
irregular testes and an entire to irregular ovary:
O. beliyai (Pande, 1937) n. comb.; O. brevifistulus
(Ozaki, 1928) n. comb.; O. caballeroi Caballero,
1977; O. cryptocentri (Yamaguti, 1958) n. comb.;
O. dactylopteri (Yamaguti, 1970) n. comb.;
O. dermatogenyos (Yamaguti, 1970) n. comb.;
O. ditrematis (Yamaguti, 1942) n. comb.; O. gobii
1982) n. comb.; O. iniistii (Yamaguti, 1970) n. comb.;
O. lobulus (Wang, 1977) n. comb.; O. macrorchis
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32
(Yamaguti, 1938) n. comb.; O. parapristipomatis
(Yamaguti, 1934) n. comb.; O. piriformis Yamaguti,
1952; O. pritchardae (Overstreet, 1969) n. comb.
(this species has no ventral sucker papillae);
O. syngnathi (Yamaguti, 1934) n. comb. and
O. ukigori Shimazu, 1988. The freshwater species
Opegaster anguillii Harshey, 1933 is variable and
could be accommodated in groups H, I or J.
Group J. Species with vitelline follicles extending
anterior to the anterior margin of the ventral
sucker, a seminal vesicle not extending into the
hindbody, entire to irregular testes and a lobed
ovary: O. lutiani (Bravo-Hollis & Manter, 1957)
n. comb.; O. ovatus (1928) n. comb.; O. plotosi
(Yamaguti, 1940) n. comb. and O. rectus (1928) n.
comb.
Remarks
The material from Acanthopagrus australis did
not exhibit any significant morphological
differences from those from A. butcheri. The
minor differences observed were in the form of
the testes and the distribution of the vitelline
follicles. The testes in one of the specimens from
A. butcheri were incised deeply at their posterior
margins, whereas this feature was not observed in
the material from A. australis (Figs. 1, 4). The
vitelline follicles in the material from A. butcheri
extend further anteriorly more often than in the
material from A. australis. These variations were
reported to occur in the related freshwater form,
O. variabilis, by Cribb (1985) and are therefore
not considered to be taxonomically important in
the present material.
Opecoelus woolcockae n. sp. fits the concept of
Group B and can be distinguished from the other
species as follows: O. minor is very distinct from
this, and the other species in the group, due to its
most elongate form, its short forebody relative to
the length of the body, and by its long finger-like
papillae. O. woolcockae n. sp. is less elongate, has
a larger oral sucker relative to the ventral sucker
and a relatively larger pharynx than O. inimici.
The latter, like O. minor, also has long digitiform
ventral sucker papillae. O. arii differs from
O. woolcockae in its distinctly oblique testes and
vitelline fields reaching only about halfway
between the ovary and the ventral sucker. The
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Syst Parasitol (2007) 67:25–42
new species is more elongate than O. mexicanus,
has a slightly smaller ventral sucker, larger phar-
ynx relative to the size of the oral sucker and a
shorter post-testicular region relative to the
length of the body. O. noblei, unlike O. wool-
cockae, has vitelline fields reaching only to the
level of the ovary. O. adsphaericus, O. sebastici
and O. mulloidichthydis are more elongate and
the latter two have shorter forebodies relative to
the length of the body than the new species.
O. woolcockae has a relatively smaller ventral
sucker than O. lotellae and O. sebastici, as re-
flected in their sucker-width ratios. The new
species can further be distinguished from
O. adsphaericus by the consistently patchy dis-
tribution of the vitelline follicles with distinct bi-
lateral interruptions in the area of the gonads in
the latter; such interruptions were not featured in
the present material (Figs. 1, 4). O. zhifuensis has
narrow, elongate ventral sucker papillae and
vitelline fields reaching about halfway between
the ovary and ventral sucker. O. variabilis is a
freshwater species, which is generally smaller and
less elongate with a larger ventral sucker relative
to the size of the ventral sucker. The new species
is closest to O. himezi in its general appearance
and in most of its metrical features. However, it
can be distinguished from the latter by its larger
pharynx relative to the oral sucker and by its
more posteriorly situated genital pore relative to
the length of the body (see Table 1).
Opecoelus pomatomi n. sp.
Type-host: Pomatomus saltatrix (Linnaeus)
(Pomatomidae).
Type-locality: Iluka, New South Wales, 2924¢S,
15321¢E.
Site: Intestine.
Material studied: 18 from 4 of 10 hosts from Iluka,
N.S.W., April 1993.
Type-material: Holotype: QM G225357; para-
types: QM G225358-66 and G225392-4, BMNH
2006.2.1.13-15.
Description (Figs. 7–9)
[Based on 16 gravid, unflattened, whole-mounts
and 2 sets of serial sections; measurements on 11
 Table 1 Opecoelus species that are closest to the new species in Group B – with vitelline follicles whose anterior limit lies posterior to the anterior margin
of the ventral sucker, seminal vesicle extending into the hindbody, entire testes and an entire ovary
Species O. inimici O. adsphaericus O. himezi O. variabilis O. woolcockae n. sp. O. pomatomi O. crowcrofti
(ex Acanthopagrus butcheri) n. sp. n. sp.
Syst Parasitol (2007) 67:25–42

No. examined Holotype 7 2 28 10 11 10

Length 2,1260 2,500–3,100 732–1,463 988–1,543 999–2,480 1,503–1,847

Width 320 500–530 196–347 232–352 198–297 219–413
Mean 280 · 520 1,005 · 262 1,334 · 301 1,618 · 236 1,496 · 311
Length/Width 7.1 8.6* 5.4 3.8 4.1–4.8 5.0–8.6 4.3–5.3
Mean (4.4) (6.7) (4.8)
O.S. length 125–135 69–130 81–112 81–117 81–123
O.S. width 150 150 69–124 92–123 81–126 78–126
Mean 130 · 150 102 · 96 102 · 111 99 · 97 106 · 104
V.S. length 116–182 110–143 117–167 104–162
V.S. width 220–230 198–201 110–154 122–180 107–162
Mean 148 · 148 132 · 134 142 · 142 136 · 137
V.S./O.S. width 1.7* 1.3–1.7 1.5 1.2–1.9 (1.5) 1.2–1.3 (1.2) 1.3–1.6 (1.5) 1.2–1.4 (1.3)
Pharynx L. 110 65–80 59–72 41–72 55–84
Pharynx W. 74 72–75 64–83 45–77 55–91
Mean (72.5 · 73.5) 66 · 76 53 · 58 66 · 75
O.S./pharynx W. 2.0 1.6 2.0 1.6* 1.2–1.6 (1.5) 1.5–1.8 (1.7) 1.2–1.5 (1.4)
1
F.body% body L. 17.8* /8–1/5; 16.6* 20* 20–39 (29) 17.8–25.7 (21.1) 13.6–23.0 (17.9) 20.7–25.8 (22.7)
Post.T % body L. 32.6* 32.3* 30* 29.4* 23.3–31.0 (28.2) 25.4–39.6 (32.9) 18.4–26.2 (22.8)
Egg length 66 49–59 54–60 67–77 55–86 54–72 65–91
Egg width 35 32–38 33–36 43–48 25–42 27–45 32–45
Mean (71 · 45) (67 · 32) (62 · 38) (78 · 38)
G.pore % body L. 12.2* 10.1* 11.7* 14.5* 13.7–17.5 (15.4) 8.9–16.2 (12.2) 14.4–20.3 (16.6)

Abbreviations: O.S., oral sucker; V.S., ventral sucker; L, length; W, width; F.body, forebody; Post.T., post-testicular area;
G.pore, genital pore.
* Data calculated from the published figures.
33

123
34 Syst Parasitol (2007) 67:25–42

Figs. 7–9 Opecoelus

pomatomi n. sp. ex
Pomatomus saltatrix. 7.
8
Whole-mount, ventral 7
view. 8. Ventro-lateral
view of terminal genitalia.
9. Postero-ventral view of
ventral sucker. cs
Abbreviations: cs, cirrus- pp
sac; pp, pars prostatica; sv,
seminal vesicle. Scale-
bars: 7, 500 lm; 8,
100 lm; 9, 250 lm

sv

123
Syst Parasitol (2007) 67:25–42
gravid, unflattened, dorso-ventrally mounted
worms.]
Body elongate, slender, subcylindrical; maxi-
mum width in region of ventral sucker, 999–2,480
(1,618) · 198–297 (236); width to length ratio
1:5.0–8.6 (6.7). Oral sucker ventrally subterminal,
almost always spherical, 81–117 (99) · 81–126
(97). Ventral sucker close to anterior end,
spherical to subspherical, 117–167 (142) · 122–
180 (142); sucker-width ratio 1:1.3–1.6 (1.5).
Forebody short, containing several large and
small gland-cells, 216–351 (277) long, 13.6–23.0
(17.9)% body length. Prepharynx distinct, short,
entirely or partly dorsal to oral sucker. Pharynx
spherical to subspherical, 41–72 (53) · 45–77 (58);
pharynx to oral sucker width ratio 1:1.5–1.8 (1.7).
Oesophagus distinct, moderately long, usually
looped or sigmoid, flanked by large gland-cells.
Intestinal bifurcation close to anterior extremity,
13–32 (22) anterior to anterior margin of ventral
sucker. Caeca long, unite close to posterior
extremity to form cyclocoel, open through ven-
trally subterminal anus.
Testes 2, usually entire, sometimes incised
around dorso-ventral muscle (n = 2), usually
separate, occasionally contiguous (n = 2), in
middle third of body; anterior 95–171 (141) · 72–
135 (110); posterior 90–198 (154) · 72–144 (115).
Post-testicular area moderately long, 279–981
(547) in length, 25.4–39.6 (32.9)% body length.
Cirrus-sac small, encloses anterior portion of
male terminal genitalia, 42–58 (50) · 19–26 (23).
External seminal vesicle long, sinuous to convo-
luted anteriorly, large, saccular posteriorly, ex-
tends from close to origin of left caecum to point
just posterior to posterior margin of ventral
sucker; posterior portion 84–192 (142) · 42–80
(61). Pars prostatica enclosed by cirrus-sac, sur-
rounded by gland-cells. Ejaculatory duct short.
Genital atrium small. Genital pore sinistrally
submedian to sinistral, antero-sinistral to intesti-
nal bifurcation, 138–237 (190) from anterior end,
at 8.9–16.2 (12.2)% of body length.
Ovary pre-testicular, contiguous with or sepa-
rated from anterior testis, entire, usually spherical
to subspherical, sometimes notched on anterior
margin, 45–117 (79) · 63–117 (89). Mehlis’ gland
distinct, well developed, anterior or antero-dorsal
to ovary. Uterine seminal receptacle present,
35
occupies proximal loops of uterus. Laurer’s canal
present. Uterine coils occupy area between
anterior margin of ovary and posterior margin of
ventral sucker or level of posterior margin
of seminal vesicle, just overlapping inside margins
of caeca. Metraterm indistinct, surrounded by
gland-cells anteriorly. Eggs operculate, 54–72
(62) · 27–45 (38). Vitelline follicles, usually re-
stricted to hindbody, extend from 207–621 (423)
from anterior extremity, 14.2–36.0 (27.3)% of
body length, to 14–27 (20) from posterior
extremity; fields confluent dorsally and ventrally
in post-testicular area; most follicles lie lateral
and ventral to caeca, and dorsal or not; anterior
extent may be level with or just posterior to
posterior margin of ventral sucker, occasionally
level with posterior margin of seminal vesicle
(n = 1); 3 isolated follicles found just anterior to
anterior margin of ventral sucker in 1 specimen.
Excretory pore terminal. Excretory vesicle
I-shaped, long, tubular, reaches anteriorly, to
overlap posterior margin of ovary, terminates just
posterior to anterior third of body.
Remarks
Opecoelus pomatomi n. sp. can also be accom-
modated in Group B (see above). It can be dis-
tinguished from the other species in the group
using morphological characteristics, as follows:
O. pomatomi is more elongate than O. mexicanus
and O. himezi and has a larger pharynx relative to
the oral sucker than the latter. O. lotellae has a
larger ventral sucker, as reflected in its sucker
width ratio, and the anterior limit of its
vitelline follicles is just anterior to the ovary,
whereas it is just posterior to the ventral sucker in
O. pomatomi. O. himezi, O. sebastici and
O. mulloidichthydis are larger species. O. sebastici
has a shorter forebody relative to the length of
the body and larger eggs than the new species;
this is despite the fact that Yamaguti (1958)
measured the eggs of O. sebastici from live spec-
imens. The post-testicular area of O. mulloidich-
thydis is relatively shorter than that of the new
species, and the anterior limit of its vitelline fields
is at the level of the posterior third of the uterus,
whereas it is just posterior to the posterior margin
of the ventral sucker in O. pomatomi . O. arii has
123
36
distinctly oblique testes and vitelline fields
reaching only about halfway between the ovary
and the ventral sucker. O. noblei has vitelline
fields reaching only to the level of the ovary.
O. zhifuensis has a narrow, elongate ventral
sucker papillae and vitelline fields reaching only
about halfway between the ovary and ventral
sucker. O. variabilis has a less elongate and
more robust body form which is reflected in its
width to length ratio, and it has a relatively
longer forebody and larger eggs. The anterior
extent of the vitelline follicles and the posterior
extent of the seminal vesicle in O. variabilis are
variable, whereas they are consistent in the new
species. O. pomatomi is a more elongate worm
with a smaller oral sucker relative to the ventral
sucker than O. woolcockae n. sp. O. pomatomi is
very close to O. inimici and O. adsphaericus
(Table 1), but it can be distinguished from
O. inimici by its shorter ventral sucker papillae
versus long digitiform papillae and by its saccu-
lar seminal vesicle compared with the convo-
luted one in O. inimici. The new species can be
distinguished from O. adsphaericus mainly by its
slightly larger eggs, its slightly less elongate body
and the distribution of vitelline follicles which
extend further anteriorly in the new species
without forming distinct gaps in the area of the
gonads as featured in O. adsphaericus.
Opecoelus crowcrofti n. sp.
Type-host: Atherinomorus ogilbyi (Whitley)
(Atherinidae).
Type-locality: Pumicestone Passage, Bribie
Island, Moreton Bay S.E. Qld, 2700¢S, 15305¢E.
Other locality: Off Dunwich, Stradbroke Island,
2730¢S, 15324¢E.
Site: Intestine, gut.
Material studied: From Moreton Bay, Qld: 20 (16
from a pooled gut wash) from Pumicestone Passage,
off Bribie Is., June 1995; 2 from a single host from off
Dunwich, Stradbroke Is., August 1995.
Type-material: Holotype: QM G225367, para-
types: QM G225368-86, BMNH 2006.2.1.1-12.
Etymology: This species is named for P. W. Crowcroft
in recognition of his contribution to the systematics
of the opecoelids.
123
Syst Parasitol (2007) 67:25–42
Figs. 10–13 Opecoelus crowcrofti n. sp. ex Atherinomorusc
ogilbyi. 10. Whole-mount, ventro-lateral view with ovary
in the usual position, directly anterior to anterior testis. 11.
Whole-mount, ventro-lateral view with ovary almost
posterior to anterior testis. 12. Lateral view of terminal
genitalia. 13. Ventral view of ventral sucker. Abbrevia-
tions: cs, cirrus-sac; pp, pars prostatica; sv, seminal vesicle.
Scale-bars: 10,11,13, 250 lm; 12, 100 lm
Description (Figs. 10–13)
[Based on 20 gravid, unflattened, whole-mount
specimens and 2 sets of serial sagittal sections;
measurements on 10 gravid, unflattened whole-
mount specimens.]
Body elongate-oval, robust, sub-cylindrical;
1,053–1,847 (1,496) · 219–413 (311); anterior end
tapered; posterior end rounded; hindbody of al-
most uniform width; maximum width in area of
gonads, width to length ratio 1:4.3–5.3 (4.8). Oral
sucker spherical to sub-spherical; 81–123
(106) · 78–126 (104); aperture nearly terminal.
Ventral sucker on very short peduncle, spherical
to sub-spherical, with 5 papillae on both anterior
and posterior lips, 104–162 (136) · 107–162 (137);
sucker-width ratio 1:1.2–1.4 (1.3). Forebody
272–415 (339) long, 20.7–25.8 (22.7)% body
length. Prepharynx distinct, narrow, always pos-
terior to oral sucker. Pharynx large, spherical to
sub-spherical, 55–84 (66) · 55–91 (75); pharynx to
oral sucker width ratio 1:1.2–1.5 (1.4). Oesopha-
gus moderately long. Intestinal bifurcation always
dorsal to ventral sucker, 7–26 (16) posterior to
anterior margin of ventral sucker. Caeca unite
close to posterior extremity, open through single
ventrally subterminal anus.
Testes 2, entire, spherical to subspherical, tan-
dem, usually contiguous, sometimes separate
(n = 4), in posterior half of body; anterior
130–188 (152) · 123–194 (149); posterior 136–233
(187) · 130–201 (154). Post-testicular area
194–467 (345) long, 18.4–26.2 (22.8)% body
length. Cirrus-sac distinct, well developed, fairly
long, overlaps outer margin of left caecum,
97–214 (145) · 26–32 (29). Seminal vesicle
external, large, saccular, extends into hindbody,
117–220 (174) · 39–84 (60). Pars prostatica
narrow, surrounded by gland-cells, enclosed in
cirrus-sac. Ejaculatory duct short. Genital
atrium distinct, large, cylindrical. Genital pore
Syst Parasitol (2007) 67:25–42 37

11
10

12

cs

pp

sv

13

123
38
antero-sinistral to intestinal bifurcation, at point
slightly posterior to posterior margin of pharynx,
207–298 (246) from anterior end, 14.4–20.3
(16.6)% of body length.
Ovary entire, may be sub-spherical to roughly
triangular or irregular, usually pre-testicular,
antero-dextral or anterior to anterior testis,
occasionally dextral to testes (n = 1), usually
contiguous with anterior testis, flattened at con-
tiguity, sometimes separate, 65–130 (102) · 65–
156 (111). Mehlis’ gland distinct, anterior to
ovary. Uterine seminal receptacle present, in
proximal loops of uterus. Laurer’s canal present,
opens at point dorsally antero-sinistral to ovary,
in area of uterus. Uterine coils usually extend
from level of ovary to level of posterior portion of
seminal vesicle or ventral sucker, sometimes ex-
tend more posteriorly to level of testes when
ovary is antero-dextral to anterior testis (n = 3).
Metraterm distinct, joins ejaculatory duct at point
slightly posterior to genital pore. Eggs relatively
large, operculate, 65–91 (78) · 32–45 (38). Vitel-
line follicles extend from 250–499 (371) from
anterior extremity, 20.0–32.4 (24.9)% of body
length, to 19–78 (49) from posterior extremity;
fields confluent ventrally and more or less dorsally
in post-testicular area; most follicles lie lateral
and ventral to caeca, with few to none dorsally;
anterior limit usually level with or just posterior
to anterior margin of ventral sucker; sometimes
follicles reach more anteriorly to point just ante-
rior to anterior margin of ventral sucker (n = 5).
Excretory pore more or less terminal. Excre-
tory vesicle I-shaped, tubular, narrow at posterior
end; broader anterior portion reaches past testes
to terminate just anterior to posterior margin of
ovary.
Remarks
Opecoelus crowcrofti n. sp. has the morphological
features of the worms in Group B (see above); it
can be distinguished from O. inimici by its less
elongate and more robust body, its larger oral
sucker relative to the ventral sucker, a relatively
larger pharynx, a longer forebody and shorter
post-testicular area relative to the length of the
body, and larger eggs. O. mexicanus is less elon-
gate than O. crowcrofti, and it has a smaller
123
Syst Parasitol (2007) 67:25–42
pharynx relative to the oral sucker, a longer post-
testicular area relative to the body length and
smaller eggs. O. adsphaericus is a more elongate
species with a slightly larger ventral sucker and
pharynx relative to the oral sucker, a shorter
forebody and longer post-testicular area relative
to the length of the body. O. arii has distinctly
oblique testes and vitelline fields reaching only
about halfway between the ovary and the ventral
sucker. O. noblei has vitelline fields reaching only
to the level of the ovary. O. zhifuensis has narrow,
elongate ventral sucker papillae and vitelline
fields reaching only about halfway between the
ovary and ventral sucker. O. crowcrofti has a
larger oral sucker relative to the ventral sucker, a
larger pharynx relative to the oral sucker and
larger eggs than O. himezi and O. lotellae. It is a
less elongate worm with a relatively longer fore-
body, a shorter post-testicular area and larger
eggs than O. sebastici. The new species has a less
elongate body, a relatively longer forebody and
shorter post-testicular area than either O. mul-
loidichthydis or O. woolcockae n. sp.; further-
more, it has a smaller ventral sucker and a smaller
pharynx, both relative to the oral sucker, and
shorter eggs than O. woolcockae. O. crowcrofti is
closest to O. variabilis, but can be distinguished
from the latter by its slightly more elongate form,
its smaller ventral sucker as reflected by their
sucker width ratios, a longer post-testicular area
relative to the body length (Table 1), and the fact
that the latter is a freshwater species.
Opecoelus queenslandicus n. sp.
Type-host: Apogon fasciatus White (Apogoni-
dae).
Type-locality: Off Mud Island, Moreton Bay, S.E.
Qld, 2721¢S, 15314¢E.
Other localities: Central Moreton Bay, 2720¢S,
15307¢E; off Tangalooma, Moreton Bay, 2714¢S,
15325¢E.
Site: Intestine, pyloric caeca.
Material studied: Four from central Moreton Bay,
Qld, February 1968; 1 from off Mud Island, Feb-
ruary 1993; 1 from off Tangalooma, December
1993.
Type-material: Holotype: QM G225387, Para-
types: QM G225388-91, BMNH 2006.2.1.7-9.
Syst Parasitol (2007) 67:25–42
Description (Figs. 14–16)
[Description and measurements based on 6
gravid, unflattened, whole-mounted specimens.]
Body elongate-oval, 950–1,950 (1,476) · 243–
450 (352); lateral margins nearly parallel, tapering
at anterior end, broadly rounded at posterior end;
maximum width in region of gonads; width to
length ratio 1:3.9–4.6 (4.2). Oral sucker ventrally
subterminal, spherical to subspherical, 81–126
(99) · 77–117 (99). Ventral sucker spherical to
subspherical, bearing 5 papillae each on anterior
and posterior lips, 153–252 (199) · 162–252 (202);
sucker-width ratio 1:1.6–2.3 (2.0). Forebody 243–
378 (317) long, 19.4–25.6 (21.8)% body length.
Prepharynx distinct, dorsal or posterior to oral
sucker. Pharynx well developed, spherical to
subspherical, 45–72 (58) · 41–81 (61); pharynx to
oral sucker width ratio 1:1.4–1.9 (1.7). Oesopha-
gus moderately long, thick-walled. Intestinal
bifurcation usually 7–45 (24) anterior to ventral
sucker, occasionally level with anterior margin of
ventral sucker (n = 1). Caeca long, thick-walled,
unite close to posterior extremity to form cyclo-
coel, open through single anus.
Testes 2, usually entire, sometimes incised at
anterior and posterior margins around dorso-
ventral muscle fibres, spherical to subspherical or
irregular, usually contiguous, occasionally sepa-
rate (n = 1), in about middle third of body; ante-
rior 81–180 (126) · 90–171 (126); posterior 90–198
(139) · 72–180 (122). Post-testicular area 288–666
(466) long, 29.3–34.2 (31.4)% body length. Cirrus-
sac distinct, small, encloses anterior portion of
male terminal genitalia, antero-sinistral to intes-
tinal bifurcation, 48–75 (59) · 18–30 (23). Exter-
nal seminal vesicle long and sinuous anteriorly,
large and saccular posteriorly, extends from point
antero-sinistral to intestinal bifurcation to point
variously anterior to posterior margin of ventral
sucker; posterior portion 83–194 (131) · 38–71
(54). Pars prostatica distinct, small, surrounded by
gland-cells, enclosed within cirrus-sac. Ejaculatory
duct short, thick-walled. Genital atrium small.
Genital pore at level of oesophagus, antero-sinis-
tral to intestinal bifurcation, 162–233 (194) from
anterior end, 10.3–17.1 (13.6)% of body length.
Ovary pre-testicular, irregular to entire,
always contiguous with anterior testis, 90–140
39
(116) · 108–189 (140). Mehlis’ gland distinct,
well developed, dorsally antero-dextral to ovary.
Uterine seminal receptacle present, in posterior
loops of uterus. Uterus coils between anterior
margin of ovary and point level with or anterior to
posterior margin of ventral sucker, only just over-
lapping inner margin of caeca, reaches past seminal
vesicle to genital pore without coiling. Metraterm
distinct, extends from level of genital atrium to
level of left caecum. Eggs oval, operculate, 45–63
(55) · 27–45 (36). Vitelline follicles extend from
216–284 (260) from anterior extremity, 14.3–22.7
(18.1)% of body length, to 18–63 (28) from pos-
terior extremity; lateral fields confluent dorsally
and ventrally in post-testicular area, separate in
forebody; follicles usually lie lateral, ventral and
dorsal to caeca; no follicles dorsal to caeca in single
specimen; anterior extent always anterior to ante-
rior margin of ventral sucker, may be level with,
just anterior to or just posterior to posterior limit of
oesophagus.
Excretory pore terminal. Excretory vesicle
I-shaped, extends anteriorly to overlap ovary
dorsally, terminates just posterior to its anterior
margin.
Remarks
Opecoelus queenslandicus n. sp. can be accom-
modated in Group I as defined above. The fol-
lowing species can be readily distinguished
from all the species in the group as follows:
O. parapristipomatis n. comb. is the most elongate
species, as reflected in its length/width ratio and it
has the longest post-testicular area relative to
body length; O. piriformis n. comb. is flask-shaped
and the vitelline follicles are arranged in small
clusters; O. cryptocentri n. comb. has the broadest
pharynx relative to the oral sucker and it has the
longest forebody and the shortest post-testicular
area relative to the body length; O. pritchardae n.
comb. and, apparently, O. lobulus n. comb. (based
on the illustration in Zhang et al., 1999, original
description not seen) are the only species in the
group without ventral sucker papillae and
O. hippocampi n. comb. has the smallest eggs.
O. piriformis n. comb. has a shape that is unusual
for species of Opecoelus, but the many anomalous
eggs in the single specimen from which it was
123
40 Syst Parasitol (2007) 67:25–42

14
15

pp

cs

sv

16

123
 Syst Parasitol (2007) 67:25–42
bFigs. 14–16 Opecoelus queenslandicus n. sp. ex Apogon
fasciatus. 14. Whole-mount, ventral view. 15. Ventral view
of terminal genitalia. 16. Ventral view of ventral sucker.
Abbreviations: cs, cirrus-sac; pp, pars prostatica; sv,
seminal vesicle. Scale-bars: 14,16, 250 lm; 15, 200 lm
described may be indicative of some abnormality.
There appear to be no other reports of the species
since it was first reported by Yamaguti (1952).
O. queenslandicus can be distinguished from
the remaining species in Group I as follows:
O. brevifistulus n. comb. is a slightly less elongate
species, it has a relatively larger oral sucker and
smaller ventral sucker, as indicated by the sucker
width ratio and a longer forebody relative to the
length of the body; in addition, it has an indistinct
oesophagus compared with a distinct oesophagus
in the new species. O. queenslandicus has a larger
ventral sucker relative to the oral sucker, smaller
eggs and it is less elongate than O. syngnathi n.
comb. The new species is a more elongate species
with a longer post-testicular area relative to the
body length than O. macrorchis and O. ditrematis
n. comb., and has a relatively shorter forebody
than the latter. The seminal vesicle in the new
species extends to just beyond the aperture of the
ventral sucker posteriorly, whereas it extends
more posteriorly to a point just posterior to
the posterior margin of the ventral sucker in
O. macrorchis n. comb. The distribution of the
vitelline follicles in O. queenslandicus is different
from that of O. ditrematis n. comb., being more
profuse and reaching further anteriorly to the le-
vel of the pharynx in the latter, whereas they
consistently extend to the level of the intestinal
bifurcation in the new species. The genital pore in
the present material is at the level of the oesoph-
agus, whereas it is at the level of the pharynx in
O. ditrematis n. comb. O. queenslandicus has a
longer post-testicular area in relation to the body
length than O. gobii n. comb., O. inistii n. comb.,
O. caballeroi n. comb., O. dermatogenyos n. comb.
and O. ukigori n. comb., and it is more elongate
than the first three species, as reflected by its width
to length ratio. The new species has a relatively
smaller ventral sucker and a more anteriorly sit-
uated genital pore than O. gobii n. comb.; it has a
relatively larger ventral sucker than O. ukigori
and a larger pharynx relative to the size of the oral
41
sucker than O. caballeroi. The genital pore in
O. queenslandicus is at the level of the oesophagus
and its seminal vesicle does not reach the posterior
margin of the ventral sucker, whereas the genital
pore in O. dermatogenyos is at the level of the
pharynx and the seminal vesicle reaches posteri-
orly beyond the posterior margin of the ventral
sucker. O. beliyai n. comb. is a stouter species than
O. queenslandicus with six papillae on the ventral
sucker and the vitelline fields reach interiorly to
the middle of the oesophagus.
The new species is very close to O. dactylopteri
n. comb. in most of its metrical features but can
be distinguished mainly by its longer post-testic-
ular area relative to the body-length (31.4% ver-
sus 25.3%) and by the seminal vesicle which does
not extend beyond the posterior margin of the
ventral sucker whereas it extends into the hind-
body in O. dactylopteri n. comb.
Manter & Pritchard (1960) recovered Opegaster
ditrematis from eels and noted that the varia-
tions they observed in their material agreed with
Yamaguti’s (1942) worm, which he described from a
single specimen. These authors concluded that the
variation in the anterior extent of the vitelline folli-
cles, the shape of the testes and in the sucker-width
ratio were insufficient to erect a new species. Bray &
Cribb (1989) also recovered O. ditrematis from
Apogon sp. off Heron Island. Their material is very
similar to Opecoelus queenslandicus in its body form,
the position of the genital pore and in most of its
metrical features. The ventral sucker and the pharynx
in the present material are, however, larger than of
Bray & Cribb’s material, as reflected in the sucker-
width ratio of 1:1.6–2.3 (2.0) versus 1:1.5–1.8 (1.6) and
the pharynx to oral sucker width ratio of 1:1.4–1.9
(1.7) versus 1:2.2 in the latter. These differences are
here considered to reflect species distinctness,
although the occurrence of both species in fish of the
same genus from the same general geographical area
seems to indicate otherwise. On the difference in the
position of the genital pore in Yamaguti’s (1942)
material and their material, Bray & Cribb (1989)
remarked that they had not found this feature to vary
in unflattened opecoelids and that its more anterior
position in Yamaguti’s specimen may have resulted
from flattening. The position of the genital pore
was used to distinguish the new species from
O. ditrematis, as described by Yamaguti (1942).
123
42
Bray & Cribb (1989) suggested that O. ditrematis
may represent a complex of species, which might
eventually be recognised by life-cycle, host-specific-
ity and biochemical studies.
In view of the morphological variations
observed in species of Opecoelus by workers such
as Manter & Pritchard (1960) and Cribb (1985), it
is important to have a large number of specimens
so that variation can be assessed properly.
Acknowledgements I thank the Australian and Nigerian
governments and Ahmadu Bello University, Zaria, Nigeria
for financial support. I also thank the ABRS (Australian
Biological Resources Study) for financial support through
a grant to Dr T.H. Cribb, who supervised the study. I thank
Drs T.H. Cribb and R.A. Bray, and Alistair Dove, Trudy
Wright, Scott Burge and Sylvie Pichelin for assistance in
the field. Dr R.A. Bray kindly read the earlier version of
this manuscript and made useful suggestions.
References
Aken’Ova, T.O., & Cribb, T. H. (1996). Two species of
Coitocaecum Nicoll, 1915 (Digenea: Opecoelidae)
from Moreton Bay, Queensland, Australia. Systematic
Parasitology, 33, 217–224.
Banerjee, A. C. (1965). Some opecoelid trematodes from
California with a description of Opecoelus noblei, sp.
nov. American Midland Naturalist, 74, 451–456.
Bray, R. A. (1987). Some helminth parasites of marine
fishes of South Africa: Family Opecoelidae (Digene-
a). Journal of Natural History, 21, 1049–1075.
Bray, R. A., & Cribb, T. H. (1989). Digeneans of the
family Opecoelidae Ozaki, 1925 from the southern
Great Barrier Reef, including a new genus and three
new species. Journal of Natural History, 23, 429–473.
Cribb, T. H. (1985). The life cycle and biology of Ope-
coelus variabilis, sp. nov. (Digenea: Opecoelidae).
Australian Journal of Zoology, 33, 715–728.
123
Syst Parasitol (2007) 67:25–42
Cribb, T. H. (2005). Family Opecoelidae Ozaki, 1925. In
A. Jones, R. A. Bray, & D. I. Gibson (Eds.), Keys to
the Trematoda (Vol 2, pp. 443–531). Wallingford:
CABI Publishing and the Natural History Museum.
Crowcroft, P. W. (1947). Some digenetic trematodes from
fishes of shallow Tasmanian waters. Papers and Pro-
ceedings of the Royal Society of Tasmania, 81, 5–25.
Manter, H. W. (1940). Digenetic trematodes of fishes from
the Galapagos Islands and the neighboring Pacific.
Allan Hancock Pacific Expeditions, 2, 364–473.
Manter, H. W., & Pritchard, M. H. (1960) Some digenetic
trematodes of eels of Hawaii. Journal of Parasitology,
46, 651–658.
Manter, H. W. (1954) Some digenetic trematodes from
fishes of New Zealand. Transactions of the Royal
Society of New Zealand, 82, 475–568.
Mehra, H. R. (1966). Revision of Allocreadioidea Nicoll,
1934. Allahabad: Published by the author, 58 p.
Ozaki, Y. (1925). Preliminary notes on a trematode with
anus. Journal of Parasitology, 12, 51–53.
Ozaki, Y. (1928). On some trematodes with anus. Japanese
Journal of Zoology, 2, 5–33.
Pritchard, M. H. (1966). Studies on digenetic trematodes
of Hawaiian fishes: family Opecoelidae Ozaki, 1925.
Zoologische Jahrbücher. Abteilung für Systematik,
Ökologie und Geographie der Tiere, 93, 173–202.
Prudhoe, S., & Bray, R. A. (1973). Digenetic trematodes
from fishes. B.A.N.Z. Antarctic Research Expedition
Reports, Series B (Zoology and Botany), 8, 199–225.
Shimazu, T. (1988). Trematodes of the genera Coitocaecum,
Dimerosaccus and Opecoelus (Opecoelidae: Opeco-
elinae) from freshwater fishes of Japan. Proceedings of
the Japanese Society of Systematic Zoology, 37, 1–19.
Yamaguti, S. (1942). Studies on the helminth fauna of
Japan. Part 39. Trematodes of fishes mainly from
Naha. Transactions of the Biogeographical Society of
Japan, 3, 329–398.
Yamaguti, S. (1971). Synopsis of digenetic trematodes of
vertebrates (Vol. 1, 1074 p.; Vol. 2, 349 p). Tokyo:
Keigaku Publishing Company.
Zhang, J.-y., Qiu, Z.-z., & Ding, X.-j., et al. (1999) Para-
sites and parasitic diseases of fishes. Beijing: Science
Press, 735 p. (In Chinese).