Systematic Parasitology24: 99-110, 1993.

© 1993 KluwerAcademic Publishers. Printedin the Netherlands.

Lepidapedon spp. (Digenea: Lepocreadiidae) from deep-sea gadiform fishes

of the NW Atlantic Ocean, including four new species

Ronald A. Campbell 1 and Rodney A. Bray2

1Department of Biology, University of Massachusetts, Dartmouth, North Dartmouth, MA 02747, USA
2Department of Zoology, The Natural History Museum, Cromwell Road, London SW 7 5BD, UK

Accepted for publication5th March, 1992

Abstract

Five species are described from fishes in deep waters of the northwestern Atlantic Ocean: Lepidapedon
gaevskayae n. sp. from Coryphaenoides (Nematonurus) armatus; L. merretti n. sp. from Phycis chesteri;
L. zubchenkoi n. sp. from Coryphaenoides (Chalinura) leptolepis; L. beveridgei n. sp. from C. (N.)
armatus; and Lepidapedon sp. innom, from Antimora rostrata. The genus Lepidapedon is discussed and
divided into groups of species distinguished by the vitelline distribution and the extent of excretory
vesicle. The new species are placed in their appropriate groups and differentiated from the other species
in the group.

Introduction related hosts or distant localities. For example,

The genus Lepidapedon Stafford, 1904 belongs to
the small lepocreadiid subfamily Lepidapedinae
which predominantly infests fishes of the order
Gadiformes and has many deep-sea species (Bray
& Gibson, 1991). In this paper we describe four
new species from deep-sea gadiformes (macrour-
ids and gadids) from the NW Atlantic. Multivari-
ate analyses have strongly indicated that the genus
Lepidapedon contains many superficially similar,
but distinguishable, species and that they are oiox-
enic, or at least stenoxenic (Bray & des Clers,
1992). In view of this finding, and in anticipation
of a full revision of the genus (Bray & Gibson, in
prep.) and of molecular (ribosomal RNA gene
sequence) results, we have recognised as being
distinct forms which differ from their congeners
in minor morphological features and proportions
or by host or geographical separation. We feel
that there would be a damaging potential loss of
information if the forms we describe were not
distinguished from related forms from distantly
one of the species we describe from a deep-sea
macrourid in the northern NW Atlantic is in prac-
tical terms morphologically indistinguishable from
two species in Antarctic nototheniids and zoar-
cids. We feel that these are unlikely to be con-
specific. The four new species described here,
however, can be readily distinguished from each
other by clear morphological features. The multiv-
ariate analyses (Bray & des Clers, 1992) on which
we base our findings was made on worms from
five gadiform host-species all fixed in similar ways.
Unfortunately, the fixation of the worms de-
scribed here is not of a similar type to that used
by Bray & des Clers and, therefore, these worms
are not entirely comparable with their results.
Materials and methods
Live worms were dissected in sea water from
freshly caught fish and immediately fixed in alcoh-
ol-formalin-acetic acid (A.F.A.) under coverslip
100 R.A. Campbell and R.A. Bray
pressure at room temperature for approximately
20 minutes. Fixed specimens were stored in vials
containing AFA for the duration of the cruise.
Fish hosts were always dead when obtained from
depths greater than 1,500 m regardless of season.
Live or moribund fish were obtained from shal-
lower depths, especially when the surface waters
were cold. Additional worms were obtained from
data-catalogued fishes fixed in 10% buffered for-
malin and stored in 70% ethanol. All digeneans
were transferred to 70% ethanol prior to further
processing. Whole-mounts were stained with
Mayer's paracarmine, paraffin sections were cut at
8-10/~m, stained with Harris' haematoxylin and
counterstained with eosin. All hosts were ident-
ified by Dr Richard L. Haedrich prior to examin-
ation for parasites.
Measurements are quoted as range (with mean
in parentheses) in micrometres throughout.
Family Lepocreadiidae Odhner, 1905
Subfamily Lepidapedinae Yamaguti, 1958
Genus Lepidapedon Stafford, 1904
The species of this genus may be divided into
(probably not natural) groups based, predomin-
antly, on the anterior extent of the vitelline fol-
licles. The specimens we describe belong to two
of these groups, which will be briefly discussed
below.
Group A. This group includes species where the
anterior extent of the vitellarium is distinctly
within the hindbody, i.e. posterior to the posterior
margin of the ventral sucker. This is the group
designated the 'Lepidapedon elongatum species-
complex' by Bray & des Clers (1992). The group
can be further subdivided by the anterior extent
of the excretory vesicle thus (with species listed):
i. Excretory vesicle reaches to about level of post-
erior margin of anterior testis:
L. elongatum (Lebour, 1908) (syn: L. microc-
otyleum Odhner in Dollfus, 1953)
L. coelorhynchi Yamaguti, 1938
L. gadi (Yamaguti, 1934)
ii. Excretory vesicle reaches to about level of
posterior margin of posterior testis:
L. antimorae McCauley, 1968
L. australis Manter, 1954
L. filiformis McCauley, 1968
L. lepidum Gaevskaya & Rodyuk, 1988
L. oregonensis McCauley, 1968
L. taeniaturn Gaevskaya & Rodyuk, 1988
L. yaquina McCauley, 1968
iii. Excretory vesicle reaches to about level of
ovary:
L. ostorhinchi Korotaeva, 1974
iv. Excretory vesicle not described:
L. clavatum Linton, 1940
L. gymnacanthi (Issaitschikov, 1928)
Group B. Those species where the vitelline fol-
licles reach well into the forebody, usually to
about the level of the intestinal bifurcation. This
group can also be subdivided on the length of the
excretory vesicle thus (with species listed):
i. Excretory vesicle reaches to the level of the
posterior margin of the anterior testes or else-
where well into the region between the testes:
L. notogeorgianus Zdzitowiecki, 1990
L. tertius Zdzitowiecki, 1990
ii. Excretory vesicle reaches to about the level of
the posterior margin of the posterior testis:
L. calli Acena, 1947
L. carcini Mordvinova, 1985
L. garrardi (Leiper & Atkinson, 1914) (syn:
L. antarcticum Byrd, 1963)
iii. Excretory vesicle reaching into forebody to
about level of intestinal bifurcation:
L. congeri Manter, 1954
iv. Excretory vesicle not described:
L. golphick Oshmarin, 1968
Cirrus-sac
The form of the cirrus-sac in Lepidapedon is
characteristic of the subfamily Lepidapedinae and
does not need repetitive description. It has been
described many times, but its features were
stressed by Shimazu & Shimura (1984) and by
Bray & Gibson (1989), who distinguished it (as
the 'Lepidapedon-type') from the other major le-

pocreadiid pattern, which they designated the
'Opechona-type' arrangement. It has a muscular
outer wall which is dwarfed by the very thick wall
of the internal male duct occupying much of the
lumen of the sac, and which is separated from the
outer wall by a narrow layer of fibrillar connective
tissue and gland-cells. The lumen of the male duct
is narrow proximally (the internal seminal vesicle)
and distally forms a globular to oval vesicular pars
prostatica. The proximal male duct enters the pars
prostatica from the side or proximally through a
small papilla. The thick wall of the male duct
forms a short, conical permanent cirrus distally,
which protrudes slightly into the genital atrium.
Lepidapedon gaevskayae n. sp. (Figs 1-2)
Synonym: Lepidapedon elongatum of Campbell,
Haedrich & Munroe (1980).
Material studied
Host: Corphaenoides (Nematonurus) armatus
(Hector), Macrouridae.
Site: Intestine.
Type-locality: Hudson Canyon, 39012' N,
71o47 ' W. Depth: 2,293 m. (No. RHB 206).
Other localities: 39011' N, 71021' W. Depth:
2,603 m. (No. RHB 16, 18); 39°35 ' N, 70031 ' W.
Depth: 2,295m. (No. RHB C86-36); 39°10'N,
71°35'W, Depth: 2,481m. (No. RHB 162);
39001 ' N, 71o10' W. Depth: 2,763 m. Upper Conti-
nental Rise (No. RHB 268); 39°12 ' N, 71047' W.
Depth 2,293 m. Hudson Canyon (No. RHB 201).
Registration numbers: BM(NH) holotype
1991.10.30.20, paratypes 1991.10.30.14-19,
USNM 82190 (2 specimens).
Description
Based on 9 specimens, one in serial section. Body
elongate; flattened; 2,400-3,700 (2,970)× 280-
600 (443); width 11.2-20.3 (15.37)% of body
length (Fig. 1). Body-surface spinose throughout,
spines most prominent in forebody. Oral sucker
subterminal; rounded; 136-210 x 144-280
(178 x 197). Ventral sucker rounded to oval; 128-
232 x 144-224 (149 x 152); in anterior third of
body. Sucker-ratio 1:0.79-0.95 (0.85). Forebody
Lepidapedon from the NW Atlantic 101
26-33% of body length. Prepharynx 30-160 (86).
Pharynx 96-130 x 56-96 (97 x 71). Oesophagus
156-258 (212). Intestinal bifurcation in posterior
forebody. Caeca terminate blindly near posterior
extremity.
Testes two; tandem; subspherical to rounded;
margins smooth; anterior 144-272 x 168-352
(223 x 263); posterior 184-330 x 200-376 (254-
x 299). Post-testicular region 10-20% of body
length. External seminal vesicle 156-281 x 30-65
(229 x 51); surrounded by delimited mass of
gland-cells; outline of cell-mass may be smooth or
indented. Cirrus-sac of typical Lepidapedon-type;
114-190 x 68-120 (151 x 88) (Fig. 2). Genital at-
rium small. Genital pore just antero-sinistral to
ventral sucker.
Ovary ovoid; pretesticular; 152-244 x 176-280
(202 x 236). Seminal receptacle postero-dorsal to
ovary. Laurer's canal not seen (sections damaged
in this region). Uterus passes anteriorly from
ovary. Metraterm short, restricted to forebody;
wall muscular; ensheathed in gland-cells. Enters
genital atrium from left. Eggs numerous; tanned;
operculate; 59-78 x 38-48 (69 x 41). Vitellarium
follicular; follicles extend to level of seminal ves-
icle, about 2 diameters of ventral sucker from
ventral sucker; distinct bands present between
gonads; gaps at level of testes and ovary may
occur, but are not consistent; lie ventrally and
encroach dorsally to caeca.
Excretory pore terminal; vesicle reaches into
region between testes almost to anterior testis.
Discussion
L. gaevskayae n. sp. fits into group A(i) (see
above), which includes three very similar species.
L. elongatum is a shallow-water form from the
north Atlantic predominantly found in cod
(Gadus spp.), L. gadi is the north Pacific vicariant
(and may be a synonym) of L. elongatum and L.
coelorhynchi is a north Pacific macrourid form
(Lebour, 1908; Dollfus, 1953; Yamaguti, 1934,
1938). A comparison of some important pro-
portions of L. gaevskayae and L. elongatum (using
the measurements taken from 25 specimens of the
latter species size [range 1,740-3,970 (2,882)/~m]
102 R.A. Campbell and R.A. Bray

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Figs 1-6. Lepidapedon spp. from deep sea fishes. 1. Lepidapedon gaevskayae n. sp. from Coryphaenoides (Nematonurus) armatus,
ventral view. 2. L. gaevskayae n. sp., terminal genitalia. 3. Lepidapedon merretti n. sp. from Phycis chesteri, dorsal view. 4. L.
merreni n. sp., excretory vesicle in immature worm. 5. Lepidapedon sp. innom, from Antimora rostrata, dorsal view. 6. Lepidapedon
sp. i n n o m from A. rostrata, terminal genitalia, dorsal view.
 Lepidapedon from the NW Atlantic 103

Table 1. Lepidapedon gaevskayae n. sp., L. merretti n. sp. and L. elongatum: comparison of some morphometric proportions (as
percentage of body-lengthunless otherwise stated).
Feature L. gaevskayae L. merretti L. elongatum
Width 11.2-20.3 (15.4) 14.1-27.2 (20.7) 5.5-14.1 (7.75)
Oral sucker 5.4-7.6 (6.4) 4.7-7.8 (6.7) 2.2-4.2 (2.8)
Pharynx 2.5-3.7 (3.0) 3.3-5.4 (4.7) 1.6-2.6 (1.9)
Oesophagus 5.4-9.7 (7.0) 1.8-2.7 (2.5) 4.1-12.8 (6.3)
Ventral sucker 4.4-7.5 (5.7) 5.8-8.4 (6.8) 2.4-3.9 (3.0)
Intestinal bifurcation as % of forebody 20.6-23.2 (21.9) 22.8-36.1 (28.9) 25.2-50.5 (43.3)
Ventral sucker to ovary 24.7-30.6 (27.6) 14.7-23.4 (18.8) 16.3-34.2 (28.4)
Anterior testis 6.1-10.0 (8.4) 7.6-12.2 (10.1) 4.1-9.0 (5.9)
Posterior testis 7.2-11.9 (9.6) 8.3-13.9 (11.5) 4.3-9.3 (6.2)
Post-testicular region 16.4-16.9 (16.7) 10.3-21.7 (17.0) 10.3-18.0 (13.7)

from the type-host Gadus morhua from various
localities [see Bray & des Clers, 1992]), mostly
expressed as a percentage of body-length (sucker,
pharynx and testes dimensions taken as
length + width/2) shows that the species differ in
the following characters: oral sucker, ventral
sucker, anterior testis, posterior testis, post-tes-
ticular region, and the distance from the intestinal
bifurcation to the ventral sucker as a percentage
of the forebody length (Table I). The species de-
scribed below, L. merretti, differs in width,
oesophagus length, the distance between ventral
sucker and ovary, and the distance from the intes-
tinal bifurcation to the ventral sucker as a per-
centage of the forebody length (Table I). The
north Pacific form from a macrourid host, L. co-
elorhynchi, differs in sucker-ratio and egg-size.
This species is named for Professor Albina V.
Gaevskaya of the Institute of Biology of the Sou-
thern Seas, Sevastopol, in recognition of her con-
tributions to the biology of deep sea fish hel-
minths.
Lepidapedon merretti n. sp. (Figs 3-4)
Material studied
Host: Phycis chesteri Goode & Bean, Gadidae.
Site: Intestine, pyloric caeca, stomach.
Type-locality: Hudson Canyon 39°57'N, 70 °
58' W. Depth 457 m. (No. RHB 526).
Other localities: 39032 ' N, 72024 ' W. Depth 498 m.
(No. RHB 44, 438); 39o50 ' N, 70o32 , W. Depth
768 m. (No. RHB 92, 93, 94); 39032 ' N, 72024 ' W.
Depth 497m. (No. RHB 441); 39o31' N ,
72023 ' w . Depth 550m. (No. RHB 504);
40000 ' N, 70047' W. Depth 283m. (No. RHB
510); 39°57'N, 70°58'W. Depth 457m. (No.
RHB 521,527, 535,536).
Registration numbers: BM(NH) holotype
1991.10.30.21, paratypes 1991.10.30.23-35,
USNM 82191 (2 specimens).
Description
Based on 10 mature specimens. Body elongate;
flattened; 1,835-2,780 x 391-518 (2,201 x 445)
(Fig. 3). Body-surface spinose throughout, spines
most prominent in forebody. Oral sucker subter-
minal; rounded; 113-190 x 109-170 (150 x 139).
Ventral sucker in anterior third of body; rounded;
120-164 x 136-170 (145 x 152). Sucker ratio
1:0.94-1.39 (1.11). Forebody 18-31 (24)% of
body length. Prepharynx 5-90 (53). Pharynx
muscular; 101-160 x 55-110 (128 x 78). Oesoph-
agus 39-92 (54); pseudoesophagus absent. Intesti-
nal bifurcation in posterior forebody. Caeca term-
inate blindly near posterior extremity.
Testes two; tandem; subspherical to rounded;
margins smooth or slightly indented; anterior 160-
262 x 190-268 (200 x 240); posterior 190-
418 x 195-300 (259 x 254). Post-testicular region
9-22 (17)% of body length. External seminal ves-
icle surrounded by delimited mass of gland-cells.
Cirrus-sac of typical Lepidapedon-type; 168-
224 x 80-156 (197 x 110). Genital atrium small.
Genital pore just antero-sinistral to ventral
sucker.
Ovary ovoid; smooth to slightly indented; pre-
104 R.A. Campbell and R.A. Bray
testicular; 80-189 x 176-232 (143 x 213). Semi-
nal receptacle postero-dorsal to ovary. Laurer's
canal not seen. Uterus passes anteriorly from
ovary. Metraterm short, just overlapping ventral
sucker; wall muscular; ensheathed in gland-cells;
enters genital atrium from left. Eggs numerous;
tanned; operculate; 64-79 x 32-40 (69 x 37). Vit-
ellarium follicular; follicles round to subspherical;
follicles extend to about level of posterior margin
of ventral sucker or distinctly posterior; distinct
bands present between gonads; gaps at level of
testes and ovary may occur, may be unilateral at
testis level, but are not consistent; follicles lie
ventrally and encroach dorsally to caeca.
Excretory pore terminal; 1-shaped vesicle re-
aches to anterior testis (Fig. 4).
Discussion
This species also fits into Group A(i) (see above).
A comparison of some body-proportions of L.
merretti and L. elongatum are given in Table I.
The features listed in the table which serve to
distinguish these species are the width, the oral-
sucker, the pharynx, the oesophagus, the ventral
sucker and the distance between the ventral
sucker and the ovary.
L. elongatum has been reported twice from
Phycis chesteri. Manter (1934) listed the associ-
ation in 366 m (200 fms) depth off Tortugas, Flor-
ida, with a prevalence of one in six. Scott (1987)
reported the association at a prevalence of 5%
from the central and northeastern Scotian Shelf
in water deeper than 100 fms (183 m). We have
seen five specimens (three whole-mounts and two
sets of sections) mentioned by Scott (BM(NH)
1983.1.24.1-2, 1984.3.22.2). These are large
worms (5,130-6,580/~m long) from frozen hosts
with many proportions different to those de-
scribed here. They differ from L. merretti parti-
cularly in the prepharynx length (as a percentage
of body-length: 8.2-9.6 (8.8) vs 1.1-4.0 (3.0), and
many other proportions are closer to those of L.
elongatum, which is much smaller. The status of
this large form is problematical. It may well be
that certain organs (sizes expressed here as per-
centages of body-length), such as oral sucker (2.4-
3.8 (3.1)), ventral sucker (3.2-4.4 (3.9)) or phar-
ynx (2.0-2.6 (2.2)) exhibit allometry, not growing
as fast as the body and, therefore, decreasing as
a proportion of body-length. Intermediate sized
forms would be needed to test this hypothesis. It
should be stressed again, however, that the most
reliable comparison of proportions is of worms
fixed in similar ways.
This species is named for Mr Nigel R. Merrett
of the Natural History Museum, London, in ac-
knowledgement of his kindness and help to one
of us (R.A.B.) and of his major contributions to
deep-sea ichthyology.
Lepidapedon sp. innom. (Figs 5-6)
Material studied
Host: Antimora rostrata Gtinther, Moridae.
Site: Intestine.
Localities: Grand Bank, Newfoundland, 45034 ' N,
48o07 ' W . Depth 1,497m. (No. GA 155);
45o38 ' N, 48°16 ' W. Depth 717 m. (No. GA 158,
161); 42040' N, 69034' W. Depth 2116m. (No.
RHB 39).
Registration numbers: BM(NH) 1991.10.30.44-
47.
Description
Based on 2 measured, 5 distorted specimens and
one large fragment. Body elongate; flattened;
5,495-5,825 x 705-720 (Fig. 5). Body surface spi-
nose anteriorly; spines absent or lost posteriorly.
Oral sucker subterminal; subglobular; 185-
215 x 192-208. Ventral sucker in anterior quarter
of body; rounded; 216-220 × 219-220. Sucker-
ratio 1:1.05-1.15. Forebody 24% of body length.
Prepharynx long 285-392. Pharynx large; elong-
ate oval; 178-190 x 120-132. Oesophagus shorter
than prepharynx; 128-159. Intestinal bifurcation
just posterior to mid-forebody. Caeca terminate
blindly near posterior extremity.
Testes two; oval; margins irregular; widely sep-
arated by 302-406; anterior 485-620 × 305-405;
posterior 472-620 x 345-399. Post-testicular re-
gion 14-22% of body length. External seminal
vesicle surrounded by delimited mass of gland-

cells. Cirrus-sac of typical Lepidapedon-type;
253-260 x 145-152 (Fig. 6). Genital atrium small.
Genital pore just antero-sinistral to ventral
sucker.
Ovary oval; with irregular margins; 145-190
posterior to posterior testis; size 297-345 x 310-
328. Seminal receptacle large when full; postero-
dorsal to ovary. Laurer's canal opens dorsally at
level just posterior to ovary, laterally, close to
left caecum. Uterus passes anteriorly from ovary;
intercaecal. Metraterm muscular; about same
length as, and running parallel to cirrus-sac; en-
sheathed in gland-cells distally; enters genital at-
rium from left through distinct sphincter. Eggs
numerous; tanned; operculate; 67-72 x 32-38
(69 × 33.5). Vitellarium follicular; follicles reach
to about level of posterior margin of ventral
sucker, may be just posterior or very slightly an-
terior; bands present between gonads; gaps occur
irregularly at level of ovary and testes; follicles lie
ventrally and encroach dorsally to caeca.
Excretory pore terminal; vesicle reaches to or
almost to posterior margin of anterior testis.
Discussion
This worm clearly belongs in group A(i) (see
above) by virtue of its relatively long excretory
vesicle. It differs from the Atlantic members of
this group, L. elongatum, L. gaevskayae and L.
merretti, by several proportions. Comparison of
this form with the features enumerated for distin-
guishing L. elongatum in the multivariate analyses
undertaken by Bray & des Clers (1992) shows that
this form overlaps the range of many features but
is not close to the mean, although with only two
specimens, the value of this comparison is limited.
The important features compared are (given as
percentage of body length) in Table II.
We are not giving a new name to this form as
we have only two measured worms, because it
appears closest to the unnamed form from Ono-
gadus argentatus in Bray & des Clers (1992) for
which a description based on 26 specimens is in
progress, and because the worms were not fixed
in a similar way to those used for the multivariate
analyses.
Lepidapedon from the NW Atlantic 105
This form differs from L. antimorae from the
same host in the Pacific Ocean off Oregon in the
length of the excretory vesicle and prepharynx,
the distinct separation of the gonads and the broad
bands of vitelline follicles between the gonads.
Lepidapedon zubchenkoi n. sp. (Figs 7-9)
Synonyms: Lepidapedon luteum abyssensis and
Lepidapedon sp. n. of Campbell et al. (1980).
Material studied
Host: Coryphaenoides ( Chalinura) leptolepis
Giinther, Macrouridae.
Site: Intestine.
Type-locality: New York Bight, slope, 38°29 ' N,
70010 ' W. Depth: 3,250 m. (No. RHB C86-19).
Other localities: 39033' N, 70044' W. Depth:
2,397m. (No. RHB 541); 38028'N, 70052' W.
Depth: 2,949m. (No. RHB 544); 38°20'N,
70013 ' W. Depth: 3,300m. (No. RHB 546, 547,
548); 38°11 ' N, 69050' W. Depth 3,623 m. (No.
RHB 787); 39033 ' N, 70044 ' W. Depth 3,623 m.
(No. RHB 788, 790).
Registration numbers: BM(NH) holotype
1991.10.30.22, paratypes 1991.10.30.4-13, USNM
82192 (2 specimens).
Description
Based on 5 specimens. Body large; elongate oval;
5,470-7,380 x 1,190-1,970 (6,662 x 1,582) (Fig.
7). Tegument spined throughout. Oral sucker
subglobular; subterminal; 310-390 x 170-400
(350 x 334). Ventral sucker in anterior third of
body; rounded; 300-410 x 360-450 (374 × 414).
Sucker-ratio 1:1.05-1.15 (1.08). Prepharynx 180-
230 (192); wall in posterior part may be wrinkled
giving impression of anterior processes on phar-
ynx. Pharynx ovoid; 180-250 x 117-170
(216 x 155). Oesophagus 430-500 (484) long. In-
testinal bifurcation in about mid-forebody. Caeca
reach close to posterior extremity and terminate
blindly.
Testes two; tandem; irregular, subspherical,
slightly lobed; anterior 385-870 x 460-850
(613 x 642); posterior 490-800 x 610-770 (616
x 702). External seminal vesicle surrounded by
106 R.A. Campbell and R.A. Bray

Table H. Lepidapedonsp. innom, and L. elongaturn: comparisonof some morphometricproportions (as percentageof body-length
unless otherwisestated).
Lepidapedon sp. L. elongatum
Width 12.4-12.8 5.5-14.1 (7.75)
Forebody 24.0-24.3 20.8-32.9 (27.1)
Intestinal bifurcationto ventral sucker (% of forebody) 36.6-36.7 25.2-50.5 (43.3)
Ventral sucker to ovary 18.0-21.4 16.3-34.2 (28.4)
Oral sucker 3.5-3.6 2.2-4.2 (2.8)
Ventral sucker 3.7-4.0 2.4-3.9 (3.0)
Post-testicular region 14-22 10.3-18.0 (13.7)

delimited gland-cell mass; 350-600x270-430 or slightly anterior, and, therefore, the vitelline
(495 × 365). Cirrus-sac of typical Lepidapedon- follicles do not reach far into the forebody
like construction; 320-460 x 180-290 (400 x 250) (Acena, 1947; Ching, 1961).
(Fig. 9). Genital atrium small. Genital pore adjac- L. carcini from the crab Carcinus mediterraneus
ent and antero-sinistral to ventral sucker. in the Black Sea, is small (1,218-1,400/zm), with
Ovary pre-testicular; smooth, rounded or sub- its ventral sucker in the mid-body, and with the
rectangular; 310-530 x 440-540 (404 x 486). gonads crowded into the posterior quarter of the
Mehlis' gland sinistral to ovary. Seminal recepta- body. It is not clear from the description (Mordvi-
cle postero-dorsal to ovary. Laurer's canal convol- nova, 1985) that the terminal genitalia are of the
uted; ensheathed in gland-cells; opens dorsally to Lepidapedon-type.
left caecum at level of ovary. Uterus lies anterior This species is named for Dr A.V. Zubchenko
to ovary. Metraterm commences just in hindbody; of the Polar Research Institute of Marine Fisher-
enters genital atrium sinistrally; with muscular ies and Oceanography, Murmansk, in recognition
wall ensheathed in gland-cells. Eggs numerous; of his contributions to the marine parasitology of
tanned; operculate; 80-90 x 50-60 (84 x 56). Vit- the north Atlantic.
ellarium follicular; follicles reach anteriorly to
about level of intestinal bifurcation; confluent in
forebody; do not intrude between gonads; ventral,
but not dorsal to caeca.
Lepidapedon beveridgei n. sp. (Figs 10-14)
Excretory pore terminal; vesicle short, reaches
Synonyms: Lepidapedon sp. n. and Neolepidape-
to or almost to posterior testis (Fig. 8).
don sp. n. of Campbell et al. (1980).

Discussion

This species belongs in Group B(ii) (see above).

It differs from the three species in this group in Material studied
the following characters: Host: Coryphaenoides (Nematonurus) armatus
L. garrardi, an Antarctic form from noto- (Hector), Macrouridae.
theniid, channichthyid and bathydraconid fishes, Site: Intestine.
is smaller (450-2,450/~m), the intestinal bifur- Type-locality: New York Bight, slope, 38005' N,
cation is in the posterior forebody, the external 69o49 ' W. Depth: 3,740 m. (No. RHB C86-14).
seminal vesicle reaches almost to the ovary and Other localities: 38044' N, 70o23' W. Depth:
the eggs tend to be huge (e.g. 83-156 x 43- 2,833 m. (No. RHB OC5-1); 38°11 ' N, 71031 ' W.
75/zm) (Byrd, 1963; Prudhoe & Bray, 1973; Zdzi- Depth: 2,603m. (No. RHB 33); 35039'N,
towiecki, 1979, 1987). 71035 ' W. Depth: 2,481 m. (No. RHB 155).
L. calli from pleuronectids in the northeastern Registration numbers: BM(NH) holotype
Pacific Ocean is tiny (940-1,246/zm), the intesti- 1991.10.30.36, paratypes 1991.10.30.37-43.
nal bifurcation is at the level of the ventral sucker USNM 82193 (2 specimens).
 Lepidapedon from the NW Atlantic 107

Figs 7-9. Lepidapedon zubchenkoi n. sp. from Coryphaenoides (Chalinura) leptolepis. 7. rlolotype,whole-mount,ventral view. 8.
Immature specimenshowingexcretoryvesicle. 9. Holotype,terminal genitalia.

Description rounded; smaller than oral sucker; in anterior

Based on 2 size forms, 10 specimens of larger
(Figs 10,12) and 5 of smaller (Fig. 11); measure-
ments in Table III. Body elongate-oval. Tegument
spined throughout, more prominent in forebody.
Oral sucker rounded; subterminai. Ventral sucker
third of body. Prepharynx length varies with state
of contraction, may be deeply wrinkled. Pharynx
oval. Intestinal bifurcation in mid-forebody or just
posterior. Caeca terminate near posterior ex-
tremity.
Testes 2; tandem; entire; in posterior half of
108 R.A. Campbell and R.A. Bray

12

Figs 10--14. Lepidapedon beveridgei n. sp. from Coryphaenoides (Nematonurus) armatus. 10. Holotype, whole-mount, large form
with deeply lobed gland-cell mass around seminal vesicle. 11. Paratype, whole-mount, small form with deeply lobed gland-cell mass
around seminal vesicle. 12. Paratype, whole-mount, large form with weakly lobed gland-cell mass around seminal vesicle. 13.
Holotype, terminal genitalia, with deep lobation of gland-cell mass around seminal vesicle. 14. Paratype, terminal genitalia, with
shallow lobation of gland-cell mass around seminal vesicle.
 Lepidapedon from the NW Atlantic 109

Table III. Dimensions of large and small forms of Lepidapedon beveridgei n. sp. from Coryphaenoides (Nematonurus) armatus) in
micrometres.
Code number RHB-OC5-1 RHB-33
Quantity studied 10 5
Body (1") 5,600-7,800 (6,490) 1,330-3,700 (2,794)
Body (w*) 820-1,060 (950) 460-1,280 (915)
Oral sucker (1) 240-376 (330) 250-370 (292)
Oral sucker (w) 288-400 (340) 270-370 (308)
Ventral sucker (1) 256-360 (297) 200-310 (246)
Ventral sucker (w) 248-344 (294) 220-340 (262)
Sucker ratio 1:0.78-0.94 (0.86) 1:0.74-0.91 (0.83)
Prepharynx (1) 320-408 (406) 80-270 (128)
Pharynx (1) 160-324(219) 160-260 (194)
Pharynx (w) 130-232 (185) 100-130 (114)
Oesophagus (1) 208-440 (328) 120-200 (160)
Anterior testis (1) 504-712 (594) 250-340 (307)
Anterior testis (w) 340-496 (414) 350-420 (390)
Posterior testis (l) 460-936 (683) 300-470 (376)
Posterior testis (w) 416-624 (460) 390-480 (414)
Seminal vesicle (1) 552-840 (742) 350-440 (392)
Seminal vesicle (w) 112-460 (314) 210-360 (262)
Cirrus-sac (1) 320-560 (440) 173-230 (210)
Cirrus-sac (w) 96-176 (120) 86-106 (94)
Eggs (1) 72-80 (76) 65-100 (77)
Eggs (w) 40-52 (45) 48-60 (52)
* 1, length; w, width.

hindbody. External seminal vesicle tubular, sinu-
ous; extending to about mid-way between ventral
sucker and ovary; surrounded by large gland-cell
mass, depending on state of contraction this mass
can be deeply lobed, but delimiting membrane
can usually be seen (Figs 13,14). Cirrus-sac of
typical Lepidapedon-type; dorsal to ventral sucker
and overlapping ventral sucker anteriorly and sin-
istrally. Genital atrium small. Genital pore sinis-
tral, slightly anterior to ventral sucker, usually
ventral to left caecum.
Ovary rounded; pre-testicular, median, im-
mediately anterior to anterior testis. Mehlis' gland
anterior to ovary. Seminal receptacle posterior
and dorsal to ovary and vitelline reservoir. Laur-
er's canal convoluted; ensheathed in gland-cells;
opens dorsally to left caecum at about level of
anterior margin of anterior testis or level with
posterior margin of ovary. Uterus pre-ovarian,
intercaecal; relatively short. Metraterm about as
long as cirrus-sac; muscular, with glandular
sheath. Vitellarium follicular; follicles large, irreg-
ular; in dense lateral bands extending from about
level of intestinal bifurcation to near posterior
extremity; usually confluent in forebody; sur-
rounding gonads, filling post-testicular and inter-
testicular spaces; follicles lie ventral, but not dor-
sal to caeca.
Excretory pore terminal; vesicle I-shaped,
passes over dorsal surface of posterior testis and
just reaches to anterior testis.
Discussion
This species is represented here by specimens of
two distinct sizes. Some proportions (e.g. body-
length to width) also appear to distinguish these
size-forms, but we believe that the forms are so
similar morphologically, that these differences
must be related to age, allometry, variability and
fixation effects.
This species fits into Group B(i) (see above). It
is morphologically very similar to the two known
species, i.e. the Antarctic forms L. notogeorg-
ianus, from nototheniids, and L. tertius, from the
zoarcid Ophthalmolycus concolor, which have
closely adjacent testes with no intervening vitel-
line follicles (Zdzitowiecki, 1990).
110 R.A. Campbell and R.A. Bray
We have named this species for Dr I. Beveridge
of the School of Veterinary Science, University of
Melbourne, in recognition of his major contribu-
tions to helminth taxonomy and in thanks for his
kindness and help to both authors.
Acknowledgements
Thanks are due to Richard L. Haedrich for collec-
tion and identification of hosts and for making
research space available aboard research vessels
at Woods Hole Oceanographic Institution and the
Memorial 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 R.C. and in part by NSF
22339 and National Sciences and Engineering Re-
search Council Grant A-7230 to R.L. Haedrich.
References
Acena, S.P. (1947) New trematodes from Puget Sound fishes.
Transactions of the American Microscopical Society, 66, 127-
139.
Bray, R.A. & des Clers, S.A. (1992) Multivariate analyses of
metrical features in the Lepidapedon elongatum (Lebour,
1908) species-complex (Digenea, Lepocreadiidae) in deep
and shallow water gadiform fishes of the NE Atlantic. Sys-
tematic Parasitology, 21,223-232.
Bray, R.A. & Gibson, D.I. (1989) The Lepocreadiidae (Di-
genea) of fishes from the north-east Atlantic: review of the
genus Neolepidapedon Manter, 1954, with a description of
N. smithi n. sp. Systematic Parasitology, 13, 11-23.
Bray, R.A. & Gibson, D.I. (1991) The Lepocreadiidae (Di-
genea) of fishes from the north-east Atlantic: Profundiv-
ermis intercalarius n. g., n. sp. from the marine fish Cory-
phaenoides (Nematonurus) armatus (Hector) (Macrouridae)
from the Porcupine Abyssal Plain. Systematic Parasitology,
18, 121-125.
Byrd, M.A. (1963) Helminth parasites of Antarctic verteb-
rates. Part I. Digenetic trematodes of marine fishes. Pro-
ceedings of the Helminthological Society of Washington, 30,
129-148.
Campbell, R.A., Haedrich, R.L. & Munroe, T.A. (1980)
Parasitism and ecological relationships among deep-sea ben-
thic fishes. Marine Biology, 57, 301-313.
Ching, H.L. (1961) Redescription of the digenetic trematodes
Lepidapedon calli and L. pugetensis Acena, and new host
records for L. calli and Eurycreadium vitellosum Manter
from fishes of Washington State. Canadian Journal of Zo-
ology, 39, 615-621.
Dollfus, R.P. (1953) Aper~u g6n6ral sur l'histoire naturelle
des parasites animaux de la morue Atlanto-Arctique Gadus
callarius L. (= morhua L.). Encyclopgdie Biologique, 43,
428 pp.
Lebour, M.V. (1908) Fish trematodes of the Northumberland
coast. Report of the Investigations of the Northumberland
Sea Fish Commission, (1907), 11-57.
Manter, H.W. (1934) Some digenetic trematodes from deep-
water fish of Tortugas, Florida. Publications of the Carnegie
Institute, 28, 257-345.
Mordvinova, T.N. (1985) Helminth fauna of certain species of
the Black Sea highest Crustacea class. Ekologiya Morya,
20, 50-58 (In Russian).
Prudhoe, S. & Bray, R.A. (1973) Digenetic trematodes from
fishes. Reports. B.A.N.Z. Antarctic Research Expedition,
Series B (Zoology and Botany), 8, 195-225.
Scott, J.S. (1987) Helminth parasites of the alimentary tract
of the hakes (Merluccius, Urophycis, Phycis: Teleostei) of
the Scotian Shelf. Canadian Journal of Zoology, 65, 304-
311.
Shimazu, T. & Shimura, S. (1984) Paralepidapedon g. n. (Tre-
matoda: Lepocreadiidae), with descriptions of metacercar-
iae of Paralepidapedon hoplognathi (Yamaguti, 1938) comb.
n. and other species from sea urchins. Zoological Science,
1,809-817.
Yamaguti, S. (1934) Studies on the helminth fauna of Japan.
Part 2. Trematodes of fishes, I. Japanese Journal of Zoology,
5,249-541.
Yamaguti, S. (1938) Studies on the helminth fauna of Japan.
Part 21. Trematodes of fishes, IV. Ky6to: S. Yamaguti, 139
PP.
Zdzitowiecki, K. (1979) Digenetic trematodes in alimentary
tracts of fishes of South Georgia and South Shetlands (Ant-
arctica). Acta lchthyologia et Piscatoria, 9, 15-31.
Zdzitowiecki, K. (1987) Digenetic trematodes from the ali-
mentary tract of fishes off South Shetlands (Antarctic). Acta
Parasitologica Polonica, 32, 219-232.
Zdzitowiecki, K. (1990) Little known and new Antarctic Di-
genea of the genera Neolepidapedon and Lepidapedon (Le-
pocreadiidae). Acta Parasitologica Polonica, 35, 19-30.