SystematicParasitology 31: 201-213, 1995.

201
© 1995KluwerAcademicPublishers. Printedin theNetherlands.

Fellodistomidae and Zoogonidae (Digenea) of deep-sea fishes of the NW

Atlantic Ocean

R o d n e y A. B r a y 1 a n d R o n a l d A. C a m p b e l l 2
1Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK
2Department of Biology, University of Massachusetts Dartmouth, North Dartmouth, Massachusetts 02747, USA

Acceptedfor publication4th March, 1994

Abstract

The following species are described from fishes in deep-waters of the northwestern Atlantic Ocean: Fellodistomidae:
Steringophorus furciger from Antimora rostrata, Coryphaenoides (Lionurus) carapinus, Lycenchelys paxillus, L.
verrillii and Lycodes atlanticus; S. blackeri from Xenodermichthys copei, S. haedrichi n. sp. from Porogadus
milesi, characterised by its lobate testes, sucker-ratio and long vitelline fields Megenteron crassum from Dicrolene
intronigra; Zoogonidae: Brachyenteron rissoanum n. sp. from Polyacanthonotus rissoanus, characterised by the
ovary position, the saccular seminal vesicle, the marginal genital pore and egg-size; Koiea notacanthi n. g., n. sp.
from Notacanthus (?) chemnitzi, characterised by the Y-shaped excretory vesicle and the posteriorly placed testes.

Introduction were transferred to 70% ethanol prior to further pro-

Members of the families Fellodistomidae Nicoll, 1909
and Zoogonidae Odhner, 1902 are common parasites
of marine fishes at all depths, and are two of the major
families of digeneans found in deep-water fishes. In
this paper we describe six species, four fellodistomids
and two zoogonids. One fellodistomid and both zoogo-
nids are considered new. Special justification is neces-
sary for the description of the two new zoogonids, as
both are based on single specimens. We believe that
both forms are so distinct morphologically that there
can be no doubt as to their status. In conjunction with
the difficulty in acquiring deep-sea material, we believe
that this makes a good case for their description.
Materials and methods
The trematodes were dissected from freshly caught
fish and immediately fixed in alcohol-formol-acetic
acid (AFA) under minimal coverslip pressure at room
temperature for approximately 20 minutes. Fixed spec-
imens were stored in vials containing AFA for the dura-
tion of the cruise. Additional worms were obtained
from data-catalogued fishes fixed in 10% buffered
formalin and stored in 70% ethanol. All digeneans
cessing. Whole-mounts were stained with Mayer's
paracarmine, and mounted in Canada balsam. Paraf-
fin sections were cut at 8-10 #m, stained with Harris'
haematoxylin and counterstained with eosin. Figures
were drawn from mounted material with the aid of a
drawing tube. Measurements are given as the range
(with mean in parentheses) in micrometres through-
out the descriptions. All hosts were identified by Dr
Richard L. Haedrich prior to examination for para-
sites. Distribution lists are numbered in accordance
with the FAO Major Fishing Areas. The abbreviation
RHB refers to numbers assigned to host specimens.
Specimens have been deposited in the BM(NH)
collection at The Natural History Museum, London
and in the United States National Collection (USNM),
Beltsville, Maryland.
Family Fellodistomidae Nicoll, 1909
Subfamily Fellodistominae Nicoll, 1909
Genus Steringophorus Odhner, 1905
Bray, Soto & Rollinson (1994) have re-assessed the
status of the genus Steringophorus using partial small
sub-unit (18S) rRNA sequences and a retrospective
consideration of isoenzyme information from Bray &
Rollinson (1985). Their results suggest that the genus
202 R . A . Bray and R . A . Campbell
as currently conceived (e.g. Bray & Gibson, 1980)
is paraphyletic, and that those species with the bulk
of their vitelline distribution in the forebody [i.e.S.
agnotus (Nicoll, 1909) and S. sebastodis (Yamaguti &
Matumura, 1942)] should be transferred back to the
genus Fellodistomum Stafford, 1904.
Steringophorus furciger (Olsson, 1868) Odhner,
1905
Syns: See Bray & Gibson (1980), also Steringophorus
sp. of Campbell et al. (1980) in part.
Material studied
Antimora rostrata GUnther, Gadiformes: Moridae.
Site: Intestine. Localities: 39°13 ' N, 71055 ' W,
depth 1,828 m (No. RHB-C35-16); 39°11' N, 70012'
W, depth 2,730 m (No. RHB-C86-7); 39012 ' N,
71°47 ' W, depth 1,945 m (No. RHB-337); 39°12 ' N,
71°47 ' W, depth 1,945 m (No. RHB-338); 39018 ' N,
71056 ' W, depth 1,700 m (No. RHB-345). BM(NH)
1993.11.11.1-12. USNM 83384.
Coryphaenoides (Lionurus) carapinus (Goode &
Bean), Gadiformes: Macrouridae. Site: Intestine.
Localities: 39009 ' N, 71o22 ' W, depth 2,609 m (No.
RHB-561); 39012 ' N, 71047 ' W, depth 2,293 m
(No. RHB-604); 39°31 ~ N, 70020 ' W, depth 2,400 m
(No. RHB-615). BM(NH) 1993.11.11.13-16. USNM
83385.
Lycenchelys paxillus (Goode & Bean), Perciformes:
Zoarcidae. Site: Intestine. Localities: 39o45 ' N, 70°43 ~
W, depth 1,926 m (No. 521), 39 ° 52' N, 70056 ~W, depth
918 m (No. 106). BM(NH) 1993.11.11.24-25.
Lycenchelys verrillii (Goode & Bean), Zoarcidae. Site:
Intestine. Locality: 39°55 ~ N, 70049 ' W, depth 550 m
(No. 103). BM(NH) 1993.11.11.17-18.
Lycodes atlanticus Jensen, Perciformes: Zoarcidae.
Site: Intestine. Localities: 39°43 ' N, 71 ° 12' W, depth
1,691 m (No. RHB-C35-28); 39o18 ' N, 71°56 ' W,
depth 1,700 m (No. RHB-718); 39017 ' N, 71°59 ~ W,
depth 1,730 m (No. RHB-729); 39°25 ~ N, 72007 '
W, depth 1,275 m (No. RHB 1306). BM(NH)
1993.11.11.19-23. USNM 83386.
Description
The measurements of these worms are given in Table
I. The species has recently been described in detail by
Bray & Gibson (1980) and Machida (1988).
Type-host and locality: Limanda limanda [first host
listed], Varberg, Sweden.
Records: Including this paper, we have traced 67 arti-
cles mentioning records of S.furciger. Fifteen of these
records include some descriptive material. The para-
site is reported from 67 fish species, of 41 genera, 17
families and five orders. It is reported from the Arctic
Sea, the northern Atlantic Ocean, the northern Pacific
Ocean and the Mediterranean Sea.
Discussion
Some of the specimens in this collection are contracted
such that the hindbody is shorter and broader than is
usual. This, of course, has caused the forebody to be
relatively large. The hindbody in uncontracted speci-
mens differs considerably in length, depending on the
number of eggs present (see e.g. figures in Bray &
Gibson, 1980).
The large variation in egg-size was first described
by Odhner (1905), and was discussed later by Yam-
aguti (1934), Bray & Gibson (1980), Machida (1988)
and Machida & Araki (1992). Yamaguti (1934) noted
that the lightly tanned eggs could reach to 75 ~zm in
length and that the egg-size changes during the passage
of the egg through the uterus. Machida (1988) agreed
that the degree of development of the egg was a fac-
tor, but other factors such as 'different hosts, different
localities' also played a part. It may be that more than
one form is at present to be found under the name S.
furciger, and the variety of depths from which it is
recorded reinforces this view, but at present it is not
practical to distinguish these forms. Molecular studies
underway at the moment may give indications of the
status of this species.
Zoarcid fishes are widely recorded as hosts of S.
furciger, so our reports from Lycenchelys spp. and
Lycodes atlanticus are not surprising. Zoarcids consti-
tute 9.1% of host-records in the literature. The reports
from Antimora and Coryphaenoides were unexpected,
as neither morids nor macrourids have been report-
ed as hosts before. S. thulini Bray & Gibson, 1980
has been reported from the macrourids Trachyrincus
scabrus [as T. trachyrhynchus] (by Bray & Gibson,
1980) and Macrourus berglax (by Zubchenko, 1975,
see Bray, 1983). It differs from S. furciger, including
our specimens from C. (L.) carapinus, in its sucker-
ratio and deeply lobed testes. The record from Antimo-
ra is the first for this genus from a morid. Both morids
and macrourids belong to the order Gadiformes, which
constitute 5.5% of host-records of S. furciger.
The majority of records (63%) of S. furciger are
from pleuronectiform (flatfishes), which may well
 Fellodistomidae and Zoogonidae from deep-sea fishes 203

Table 1. The measurements of Steringophorusfurciger from various hosts.

Host Antimora Coryphaenoides Lycenchelys Lycenchelys Lycodes

rostrata (Lionurus) verrilli paxillus atlanticus
carapinus

n 10 5 6 2 5

Body L 1,656-5,104 1,472-3,632 1,168-1,546 1,760-1,840 1,224-2,704

W 918-2512 960-2144 618-739 670-824 685-1534
M 2515 × 1266 2406 x 1537 1377 X 681 1934 × 1151
BW % BL 36--69 (54) 55-75 (66) 43-57 (50) 38-44 52-76 (61)
Forebody L 483-1392 592-952 258-380 406-415 470)944
M 691 754 314 729
FB % BL 22-4 1 (29) 25-44 (34) 22-25 (23) 23 35--44 (38)
Oral sucker L 225-419 206-354 144-189 193-219 225-328
W 247-431 246-303 190-217 201-203 231-411
M 278 × 288 275 x 280 179 × 204 275 × 324
Ventral sucker L 328-654 264-579 278-328 335-386 316-567
W 502-832 457-898 290-399 425-444 421-664
M 420 × 575 437 x 665 307 x 364 419 x 550
Sucker-ratio 1:1.88-2.11 1:1.86-2.68 1:1.41-2.00 1:2.11-2.19 1:1.27-1.89
M 1:2.01 1:2.14 1:1.79 1:1.73
Pharynx L 119-232 109-200 90-109 104-129 129-174
W 108-222 116-142 90-109 97-116 104-148
M 141 X 142 170 × 125 102 x 101 147 × 127
Oesophagus L 39-218 13-135 32-52 77-90 45-200
M 104 63 45 102
PCR L 190-1139 152-383 ? ? 122-379
M 696 268 285
Testes L 126-560 151-386 109-180 138-225 103--415
W 142-689 193-412 97-174 135-219 109-486
M 251 x 264 271 x 285 137 × 147 252 x 282
PTR L 451-2112 332-1629 396-590 644q579 264-960
M 1014 902 512 546
trl'R % BL 26--45 (39) 23-45 (34) 34-42 (37) 37 19-36 (27)
Cirrus-sac L 213-773 245-476 200-254 225-232 213-509
W 158-547 190-361 126-155 135-148 117-439
M 324 x 246 354 × 294 223 x 145 335 x 277
Ovary L 180-631 232-386 161-232 335-341 135-380
W 187-496 174-348 132-290 207-274 161-444
M 314 × 284 280 × 257 199 x 180 271 x 263
Vitelline field L 251-1096 213-599 174-277 384--425 188-599
W 174-880 264-683 135-245 174-283 168-509
M 480 x 357 419 x 464 239 x 188 372 x 359
Eggs L 38-64 48-73 41-45 42-46 51-67
W 19-35 25-30 19-24 20-22 24-29
M 47 x 24 60×27 45 x 21 6 0 x 27

Abbreviations: BL, body-length; BW, body-width; FB, forebody; L, length; M, mean; PCR, post-caecal region;
PTR, post-testicular region; W, width.
204 R.A. Bray and R.A. Campbell
reflect a shallow-water sampling bias. In deeper waters
it appears that zoarcids and, possibly gadiforms, are the
major hosts.
Steringophorus blackeri Bray, 1973
Syn.: Abyssotrema sp. of Markle & Wenner (1979).
Material studied
Xenodermichthys copei Gill, Alepocephalidae. Site:
Intestine. Locality: Environs of Norfolk Marine
Canyon, off Virginia. BM(NH) 1993.11.11.26-28.
USNM 83387.
Description
Measurements of three worms are as follows. Body
5,072-6,112 x 2,096-2,256. Oral sucker 361-399 ×
438-464. Ventral sucker 451-470 x 486-535. Sucker-
ratio 1:1.05-1.20. Forebody 17-18% of body length.
Pharynx 155-182 x 167-200. Oesophagus 213-393.
Cirrus-sac 316--412 x 180-257. Ovary and testes
obscured by eggs. Eggs 48-53 x 25-29.
Type-host and locality: Xenodermichthys copei [syn:
X. socialis], Bay of Biscay (44023-38 ~ N, 02012 ~ W,
depth 575-665 m).
Records: 1. Bray (1973); 2. Markle & Wenner (1979 -
also Campbell et al., 1980; Campbell, 1983); 3. Bray
& Gibson (1980); 4. Aleshkina & Gaevskaya (1985);
5. Present study.
Descriptions: 1,3.
Definitive hosts: Alepocephalidae: Xenodermichthys
copei ( 1,2,3,4,5).
Distribution: 21 Atlantic, NW [off Virginia (2,5)], 27
Atlantic, NE [Bay of Biscay (1,3), offW Ireland (3), off
NW Scotland (3)], 34 Atlantic, Central [off Morocco
(1,3), off Western Sahara (4), off Guinea Bissau (4)],
47 Atlantic, SE [Mid-Atlantic Ridge NE of Ascension
(4)].
Discussion
This species is apparently oioxenic to and common in
X. copei in the Atlantic Ocean. Our specimens are not
distinguishable from those described by Bray (1973)
and Bray & Gibson (1980).
Steringophorus haedrichi n. sp. (Figs 1-3)
Material studied
Porogadus milesi Goode & Bean, Gadiformes: Ophidi-
idae. Site: Intestine. Locality: 38°28 ~ N, 70°52 ~ W,
depth 2,949 m (No. RHB 495). BM(NH) holotype
1994.2.28.1, paratypes 1994.2.28.2-3.
Description
Based on 3 specimens (2 mature), including one set
of sections. Body large, robust; linguiform; 3,600-
4,096 x 1,344-1,776 (Fig. 1). Tegument unarmed.
Oral sucker large, 420-464 x 412-502; subglobular;
subterminal. Prepharynx not seen. Pharynx subglobu-
lar; 174-187 x 155-187. Oesophagus 113-180 long.
Intestinal bifurcation in posterior half of forebody. Cae-
ca narrow; reach posterior to testes, but not into poste-
rior half of hindbody; extremities obscured by eggs in
gravid whole-mounts. Ventral sucker large, 522-528
x 563-580; rounded. Sucker-width ratio 1:1.12-1.41.
Forebody 956-979 long, 24-27% of body length.
Testes symmetrical or subsymmetrical; in anterior
hindbody; lobate; 386-522 × 335-431. Post-testicular
region 1,549-1,936 long, 43-47% of body length.
Cirrus-sac broadly oval; 406-470 x 264-309; over-
lapping ventral sucker. Internal seminal vesicle bipar-
tite. Pars prostatica wide; lined with filaments (Fig. 2).
Ejaculatory duct convoluted, complex. Genital atrium
large; wall pocketed and complex. Genital pore about
midway between suckers, median or slightly submedi-
an.
Ovary pretesticular, close to or overlapping ven-
tral sucker, slightly dextral to mid-line; multilobate;
367-390 x 277-354. Mehlis' gland large; dorsal to
ovary. Laurer's canal short; straight; ensheathed in
thick layer of gland cells; opens dorsally to ovary. Uter-
ine seminal receptacle present. Uterus coils posterior
to testes, reaching close to lateral margins and posteri-
or extremity; with more weakly tanned younger eggs
mainly on left side passing posteriorly; dextral coils
pass anteriorly with eggs more deeply tanned; nar-
row distal sling passes between testes, to left of ovary
and dorso-sinistral to ventral sucker, and enters genital
atrium from left. Metraterm short, about 33% length of
cirrus-sac; ensheathed in gland-cells. Eggs numerous;
operculate; 35-41 x 19-22; shells without ornamenta-
tion. Vitellarium follicular; 2 more or less symmetrical
lateral fields between levels of mid-ventral sucker and
lateral or postero-lateral to testes; overall measure-
ments of fields 715-992 x 335-708.
Excretory pore terminal. Vesicle Y-shaped; stem
reaches into anterior half of hindbody; arms reach to
 Fellodistomidae and Zoogonidae from deep-sea fishes 205

I::?......
1
l ~l ,!~. ! .!!j~.i:-i:~).':.. l x::.:,~
' : ;:.'i"
: > .i;.i,"
: ' L~i:- ,.-

Figs 1-3. Steringophorus haedrichi n. sp. from Porogadus milesi. 1. Holotype, whole-mount, ventral view. 2. Paratype, sagittal section of
cirrus-sac. 3. Immature specimen showing excretory vesicle.
206 R.A. Bray and R.A. Campbell
level between oesophagus and anterior pharynx (Fig.
3).
Discussion
This species is similar to several members of the genus
Steringophorus, but differs as follows:
S. furciger (Olsson, 1868) has a greater sucker-ratio
(1:1.41-2.68), an almost totally pretesticular vitelline
distribution and unlobed testes (Bray & Gibson, 1980;
Machida, 1988; see also above).
S. blackeri Bray, 1973 has larger eggs (48-63 x 25-
34), unlobed testes and an inverted Y-shaped uterine
configuration in the post-testicular zone (Bray, 1973;
Bray & Gibson, 1980; see also above).
S. brevis (Ching, 1960) has a short post-testicular zone
and a sucker-ratio of 1:1.81 (known from one specimen
only) (Ching, 1960).
S. congeri Shen, 1987 is a large worm (7,871-10,268
long), the oesophagus is long (663-1,105) and the
vitellarium reaches into the forebody and not poste-
riorly to the testes (Shen, 1987).
S. foliatus (Yamaguti, 1970) has a sucker-ratio of
1:1.4-2.6, the caeca reach almost to the posterior
extremity and the vitellarium is restricted to two small,
pretesticular fields of follicles (Yamaguti, 1970).
S. magnus Manter, 1934 has a sucker-ratio of 1:1.93-
2.18 and an egg-size of 30-32 x 14-17 (Manter,
1934).
S. melanostigma (Noble & Orias, 1975) has an egg-size
of 63 x 31, unlobed testes, a short oesophagus and a
slightly lobed ovary (Noble & Orias, 1975) (see also
Benthotrema melanostigmi Parukhin & Lyadov, 1979
with its rounded ovary, a possible synonym (Parukhin
& Lyadov, 1979)).
S. pritchardae (Campbell, 1975) is an elongate worm
with unlobed, tandem testes, elongate vitelline fields, a
great distance between the testes and the ventral sucker
and a sucker-ratio of 1:0.85-0.94 (Campbell, 1975;
Bray & Gibson, 1980). Lomasoma kergeleni Parukhin
& Lyadov, 1979 is very similar to S. pritchardae and
differs from S. haedrichi in the same ways (Parukhin
& Lyadov, 1979).
S. profundus Manter, 1934 has a sucker-ratio of
1:1.62-1.70, a mainly pretesticular vitelline distribu-
tion, smooth or slightly lobed testes and an egg-size of
30-34 x 16 (Manter, 1934).
S. thulini Bray & Gibson, 1980 has a pretesticular
vitelline distribution and a smaller cirrus-sac (160-320
x 110-150) (Bray & Gibson, 1980).
The diagnostic characters of S. haedrichi are the
lobate testes, the sucker-ratio and the length of the
vitelline fields.
The new species is named for Dr Richard Haedrich
in gratitude for his contribution to this project.
Genus Megenteron Mantel', 1934
Yamaguti (1953) considered Megenteron a synonym
of Steringotrema, but Bray & Gibson (1980) consid-
ered it distinct, based on its long caeca, multilobed
ovary and relatively small ventral sucker. Megenteron
is distinguished from Steringophorus by its V-shaped
excretory vesicle and long caeca.
Megenteron crassum Manter, 1934 (Figs 4-6)
Syn.: Steringotrema crassum (Manter, 1934) Yam-
aguti, 1953
Material studied
Dicrolene intronigra Goode & Bean, Gadiformes:
Ophidiidae. Site: Intestine. Localities: 390171 N,
71059' W, depth 1,730 m (RHB-53, RHB-54, RHB-57,
RHB-722, RHB-727, RHB-728, RHB-104), 39025 I
N, 72007 ' W, depth 871 m (RHB-1309), 390091 N,
720111 W, depth 1,280 m (RHB- 108, RHB-114, RHB-
121), 34035 ' N, 75034 ' W, depth 1,015 m (RHB-722).
BM(NH) 1993.11.11.29-31. USNM 83388.
Description
Based on 94 specimens, 15 measured gravid specimens
and one set of serial sections. Body pyriform, taper-
ing slightly anteriorly; anterior end rounded; posterior
end abruptly truncated in mature specimens; 1,760-
4,120 x 940-2,160 (3,060 x 1,490) (Fig. 4). Tegu-
ment aspinous. Gland-cells form large, conspicuous
masses lateral to pharynx and oesophagus and around
proximal end of oesophagus. Oral sucker subterminal;
large, 464-632 (499) diameter. Ventral sucker 456-
744 (559) diameter. Sucker-ratio 1:0.98-1.18 (1.11).
Forebody 700-1,280 (926) long, 29-37 (33)% of body
length. Prepharynx short, within posterior concavity
of oral sucker. Pharynx ovoid; large, 160-380 × 184-
368 (284 x 263). Oesophagus longer than pharynx;
160-368 (252) × 64-96 (82). Caeca large, becom-
ing voluminous posteriorly; terminate blindly 40-400
(162), [ 1.3-12.6 (5.4)% of body-length] from posterior
extremity.
 FeUodistomidae and Zoogonidae from deep-sea fishes 207

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Figs 4-6. Megenteron crassum Manter from Dicrolene intronigra. 4. Whole-mount, gravid specimen, ventral view. 5. Detail of terminal
genitalia. 6. Immature specimen showing excretory vesicle.
208 R.A. Bray and R.A. Campbell
Testes 2, irregular in shape; oblique, with left testis
anterior; in midbody, widely separated laterally, part-
ly overlapping caeca ventrally; 280-420 x 152-320
(364 x 251). Post-testicular region 560-1,520 (968)
long, 27-38 (33)% of body length. Cirrus-sac clav-
iform; 280-360 × 88-152 (339 x 134); extending
medially and posteriorly from genital pore, overlap-
ping ventral sucker (Fig. 5). Seminal vesicle internal;
bipartite. Pars prostatica vesicular; lined with anuclear
filaments, surrounded by gland-cells. Ejaculatory duct
short; rectilinear. Genital atrium small. Genital pore
slightly to antero-sinistral to ventral sucker.
Ovary multilobate; dextral, slightly overlapping
right caecum ventrally, just anterior to right testis; 184-
360 x 152-220 (244 x 187). Mehlis' gland antero-
dorsal to ovary. Laurer's canal convoluted; surrounded
by gland-cells; opens dorsally to ovary. Small amount
of spermatozoa in initial coils of uterus indicating uter-
ine seminal receptacle. Uterus initially passes anterior
for short distance prior to abruptly turning posterior-
ly and passing between testes to form loose coils in
hindbody that traverse caeca, then coils pass anteriorly
in mid-line towards genital atrium. Metraterm short,
weakly differentiated; enters genital atrium sinistrally
to cirrus-sac. Eggs amber; operculate, lack terminal
process; 23-30 x 15-19 (25 x 17). Vitellarium follic-
ular; 2 lateral fields of small follicles extend between
levels of mid-ventral sucker and anterior testis; ventral
to caeca.
Excretory vesicle large, V-shaped; arms volumi-
nous posteriorly in intercaecal space, crossing caeca
in hindbody, tapering until they terminate laterally at
pharyngeal level (Fig. 6).
Type-host and locality: Diplacanthopoma brachyso-
ma, Tortugas, Florida, USA.
Records: 1. Manter (1934); 2. Campbell et al. (1980);
3. Present study.
Descriptions: 1,3.
Definitive hosts: Gadiformes: Bythitidae: Dipla-
canthopoma brachysoma (1); Ophidiidae: Dicrolene
intronigra (2,3).
Distribution: 21 Atlantic, NW [New York Bight (2,3)],
31 Atlantic, W Central [Florida (1)].
Discussion
Manter (1934) described short polar processes on the
egg of this species; but, although we could not detect
this in our specimens, we could find no other differ-
ence between our specimens and Manter's description.
Polar processes are not evident on the of all speci-
mens in Manter's paratype series. Examination of these
paratypes showed that the oesophagus is equal to or
slightly longer than the pharynx. Manter's illustration
does not indicate the dorsal loop in the oesophagus
that gives it a shortened appearance in some speci-
mens. This condition was observed in several of his
specimens as well as our own.
The two host species belong to closely related gad-
iform families.
Family Zoogonidae Odhner, 1902
Bray (1987a,b) revised the family Zoogonidae. Brooks
(1990) re-interpreted Bray's data and, although he
refrained from making name changes, he reckoned
that fewer genera should be recognised. Subsequent-
ly, Brooks & McLennan (1993) changed many con-
tentious generic designations.
Subfamily Lepidophyllinae Stossich, 1903
Genus Brachyenteron Manter, 1934
Brachyenteron rissoanum n. sp. (Figs 7-9)
Material studied
Polyacanthonotus rissoanus (De Filippi & Verany),
Notacanthiformes: Notacanthidae. Site: Stomach.
Locality: 39012' N, 71047 ' depth 2,293 m (No. RHB
C124-1). BM(NH) holotype 1994.2.28.4.
Description
Based on one whole-mount specimen. Body fusiform;
extremities rounded; 1,322 x 368 (Fig. 7). Large
spines embedded in tegument over entire body surface
(Fig. 8). Numerous gland-cells embedded in parenchy-
ma. Oral sucker subglobular; subterminal, occupy-
ing almost entire width of anterior extremity; 232 x
256; aperture wide. Prepharynx short; within poste-
rior concavity of oral sucker. Pharynx large, 147 x
142; oval. Oesophagus 64 long. Intestinal bifurcation
midway between suckers. Caeca very short, 138-160
long; clavate; terminating in forebody, 52 from ventral
sucker, anterior to vitelline follicles. Ventral sucker
rounded; 155 x 180. Sucker-width ratio 1:0.70. Fore-
body 618 long, 46% of body-length.
Testes oval, entire; oblique, with right testis ante-
rior, overlapping posterior margin of ventral sucker;
 Fellodistomidae and Zoogonidae from deep-sea fishes
right testis 190 x 161; left testis 216 × 171. Post-
tesicular region 290, 21% of body-length. Cirrus-sac
elongate-claviform; lies diagonally across forebody,
reaches almost to level of ventral sucker; 354 × 47
(Fig. 9). Seminal vesicle in proximal half of cirrus-sac;
elongate-saccular; 191 x 46; walls delicate. Pars pro-
statica short; slightly vesicular; surrounded by gland-
cells. Ejaculatory duct long, narrow, surrounded by
scattered gland-cells. Genital atrium small. Genital
pore dextral, lateral, at mid-pharyngeal level.
Ovary subglobular; antero-dextral to right testis,
dorso-sinistral to ventral sucker; 122 x 117. Semi-
nal receptacle and Laurer's canal not seen. Mehlis'
gland and vitelline reservoir postero-medial to ovary.
Uterus fills most of hindbody and ascends between
testes and dorsally to ventral sucker; terminal part of
uterus runs straight, dorsal to cirrus-sac; distal por-
tion about 102 long, slightly more muscular, forming
metraterm; opens into genital atrium dorsal to male
system. Eggs tanned; operculate; 38-41 x 14-20.
Vitellarium follicular; 2 lateral clusters of 8 (poral) and
9 (aporal) rounded follicles at antero-lateral margin of
ventral sucker, immediately posterior to caeca.
Excretory pore terminal; vesicle obscured by
uterus.
Discussion
Although only one specimen is available, it is distinct
enough for us to describe as new. Some features were
not seen, but it was felt that these, for example the prox-
imal female system and the excretory vesicle, were not
likely to be diagnostic enough to warrant the sacrifice
of this worm for sectioning.
This form keys to Brachyenteron in Bray (1987b),
although unusual in that the genital pore is in the ante-
rior forebody. It cannot be keyed down to any of the
species in Bray's (1987b) key to the genus, as it has a
combination of characters which separate in the key.
At couplet 2, it has a mixture of characters: ovary
overlapping ventral sucker; seminal vesicle elongate
saccular; genital pore marginal; eggs greater than 33
/zm.
The phylogenetic trees given for this genus by
Bray (1987b), Brooks (1990) and Brooks & McLen-
nan (1993) are identical. B. rissoanus appears to form
another unresolved branch at the basal polytomy. The
apomorphy of the species is the anteriorly placed gen-
ital pore.
B. campbelli Bray & Gibson, 1986 was reported
from the same host in the NE Atlantic (Bray & Gibson,
209
1986). It differs in the position of the genital pore (fur-
ther posterior), the size of the pharynx, the sucker-ratio,
the bipartite seminal vesicle, the infundibuliform oral
sucker, the narrow and parallel caeca, the pre-gonadal
position of much of the uterus, and the vitellarium
reaches more posteriorly.
Genus K o i e a n. g.
Zoogonidae, Lepidophyllinae. Body fusiform, bear-
ing spines. Oral sucker subterminal. Prepharynx wide,
short. Pharynx large. Oesophagus long. Intestinal
bifurcation in posterior forebody. Caeca elongate,
pa(allel; terminating blindly close to mid-hindbody.
Ventral sucker rounded; small; in mid-body. Testes
2; oblique; close to posterior extremity. Cirrus-sac
elongate-oval; lies across forebody. Seminal vesi-
cle internal; coiled, tubular. Pars prostatica vesicu-
lar. Ejaculatory duct muscular. Genital atrium distinct.
Genital pore very close to ventro-sinistral margin in
mid-forebody. Ovary median, pretesticular, separated
from testes by uterus, just posterior to level of caecal
extremities. Uterus extensive, from level of middle of
posterior testis to level of cirrus-sac; field narrower
between caeca, metraterm short, muscular, with dis-
tinct sphincter distally. Eggs numerous, tanned. Vitel-
larium 2 small lateral fields of follicles along inner
margin of caeca. Excretory pore dorsally subterminal;
vesicle Y-shaped; arms reaching into forebody.
Type-species: Koiea notacanthi n. g., n. sp.
Koiea notacanthi n. sp. (Figs 10--12)
Material studied
Notacanthus (?) chemnitzi Bloch, Notacanthiformes:
Notacanthidae. Site: Intestine. Locality: 46018 ~ N,
45018 ~ W (Flemish Cap), depth 632 m (No. GA 185).
BM(NH) holotype 1994.2.28.4.
Description
Single whole-mounted worm studied. Body fusiform,
tapering posteriorly; 2,080 × 580 (Fig. 10). Tegu-
ment spinous throughout; spines acuminate (Fig. 11),
more tightly packed anteriorly. Oral sucker subtermi-
nal; subglobular, slightly wider posteriorly; 306 × 325.
Prepharynx very wide, short; not measurable as phar-
ynx is forced up into it; walls folded. Pharynx large,
161 × 211; wider aperture anteriorly. Oesophagus
210 R.A. Bray and R.A. Campbell

7 8

i
9

Figs 7-9. Brachyenteron rissoanum n. sp. from Polyacanthonotus rissoanus. 7. Holotype, whole-mount, ventral view. 8. Detail of tegumental
spines. 9. Terminal genitalia, holotype, dorsal view.
 FeUodistomidae and Z o o g o n i d a e f r o m deep-sea fishes
thick-walled, muscular; 335 long. Intestinal bifurca-
tion close to (58 anterior to) ventral sucker. Caeca
narrow; diverge almost at right angles then run paral-
lel posteriorly; end blindly at level of ovary, 335-361
posterior to ventral sucker. Ventral sucker small, 224
× 219; rounded. Sucker-width ratio 1:0.67. Forebody
876 long, 42% of body length.
Testes 2; irregularly shaped but with smooth out-
lines; oblique, close to posterior extremity; posterior
testis sinistral, reniform, 261 × 135; anterior testis
dextral, oval but with truncated oblique anterior mar-
gin, 265 x 206. Post-testicular region 126 long, 6% of
body- length. Cirrus-sac claviform; 380 x 145; runs
almost at fight angles to longitudinal axis; wall thin
(Fig. 12). Seminal vesicle internal; coiled, tubular.
Pars prostatica vesicular; lined with anuclear filaments,
ensheathed in thick layer of gland-cells which also
overlap seminal vesicle and ejaculatory duct. Ejacu-
latory duct distinct; of similar length to pars prostati-
ca; thick-walled. Genital atrium large; surrounded by
gland-cells. Genital pore close to left margin, ventral;
at about halfway between suckers.
Ovary oval; 148 × 192, transverse diameter great-
est; median, pretesticular, separated from ventral suck-
er by 328, from anterior testis by gap of 155 filled by
uterus, just posterior to ends of caeca. Other details of
proximal female system not detected. Uterus extends
anteriorly from mid-ventral region of posterior testis,
reaches close to lateral margins in post-caecal zone,
runs dorsally to ovary; field narrows in caecal region
to become mostly inter-vitelline; slings pass dorsally
to ventral sucker, some in posterior forebody poste-
rior to cirrus-sac. Metraterm short, overlaps extreme
distal end of cirrus-sac; strongly muscular; forms very
prominent, large sphincter before debouching into gen-
ital atrium from dorsal side. Eggs numerous; tanned;
32-35 × 16-17. Vitellarium consists of 2 lateral fields
of small, subglobular follicles lying along median mar-
gins of caeca; 13 follicles on poral side, reaching from
mid-ventral sucker to close to caecal extremity, 277
from ventral sucker; 11 on aporal side, reaching from
mid-ventral sucker to just anterior to caecal extremity,
206 from ventral sucker.
Excretory pore dorsally subterminal. Vesicle Y-
shaped, initially narrow and surrounded by gland cells,
divides dorsally to testes; narrow arms can be traced
into forebody close to proximal end of cirrus-sac on
aporal side and about midway between cirrus-sac and
ventral sucker on poral side.
211
Discussion
This is a fascinating and puzzling specimen. Most char-
acteristics suggest that it is a zoogonid, but the excre-
tory system appears to be Y-shaped, an unusual feature
for a zoogonid, although the lepidophylline zoogonid
Urinatrema Yamaguti, 1934 has been described on
having such (Yamaguti, 1934; Schiller, 1956). Yam-
aguti (1934) stated that in Urinatrema the stem is rel-
atively short, but in Koiea n. g. the stem and arms are
long and distinct. While such features as the lateral gen-
ital pore, spined tegument, cirrus-sac contents, short
caeca and few vitelline follicles indicate the placement
of this worm in the subfamily Lepidophyllinae, it is
distinguished from other genera by several features,
notably the posteriorly situated testes. In the key giv-
en by Bray (1987b), this form keys (more or less)
to the genus Brachyenteron, but differs from it in the
posteriorly placed testes, pre-testicular uterus, and cae-
ca and vitellarium in the hindbody. As far as we are
aware, the only new lepidophylline genus described
since Bray (1987b) is Parasteganoderma Machida &
Araki, 1990. This genus differs from other members
of the subfamily, including Koiea, in its longitudinally
divided ventral sucker and lobed oral sucker (Machida
& Araki, 1990).
Koiea appears to form an additional branch leading
from the basal polytomy in Bray's (1987b) phylogenet-
ic tree to the genera of the subfamily Lepidophyllinae.
Brooks (1990) and Brooks & McLennan (1993) give
a rather different tree to the genera of this subfam-
ily, but Koiea again appears to form a branch from
a polytomy, in this case the one which leads also to
the genera Steganoderma Stafford, 1904, Lepidophyl-
lum Stossich, 1903 and Overstreetia Bray, 1985. The
new genus does not, then, help resolve either tree. The
same could be said of the other new genus Parastegan-
oderma, which may be related to the worms on the
branch of Bray's (1987b) tree leading to Steganoder-
matoides Parukhin & Lyadov, 1979, Neosteganoderma
Byrd, 1964 and Proctophantastes Odhner, 1911, gen-
era which were considered synonymous by Brooks &
McLennan (1993). It shares a divided ventral suck-
er with these genera, but the division in this case is
longitudinal, not transverse.
The genus is named for Dr Marianne Kgfie, of
the Marine Biology Laboratory, Helsing¢r, Denmark,
in recognition of her major contributions to digenean
biology.
212 R.A. Bray and R.A. Campbell

10
11
I

0.1ram
I I

Figs 10-12. Koiea notacanthi n.g., n. sp. from Notacanthus chemnitzi (?). 10. Holotype, whole-mount, ventral view. 11. Detail of tegumental
spines. 12. Terminal genitalia, holotype, dorsal view. Abbreviations." ga, genital atrium; mt, metraterm.
 Fellodistomidae and Zoogonidae from deep-sea fishes 213
Acknowledgements
We wish to thanks Richard L. Haedrich for the collec-
tion and identification of hosts and for making research
space available for RAC aboard research vessels at the
Woods Hole Oceanographic Institution and the Memo-
riai University of Newfoundland. Thanks are also due
to Dr D.I. Gibson for reading the manuscript. This
work was supported by NSF Grant DEB76-20103 to
RAC and in part by NSF 22339 and National Sciences
and Engineering Research Council Grant A-7230 to
R.L. Haedrich.
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