Systematic Parasitology 50: 219–229, 2001.

© 2001 Kluwer Academic Publishers. Printed in the Netherlands.

219

Two new species of Acanthobothrium Beneden, 1849 (Tetraphyllidea:

Onchobothriidae) from horn sharks in the Gulf of California, Mexico

J. N. Caira & Shawn D. Zahner

Department of Ecology & Evolutionary Biology, University of Connecticut, 75 N. Eagleville Rd, Storrs,
Connecticut 06269-3043, USA

Accepted for publication 20th March, 2001

Abstract
Between 1993 and 1996, 26 individuals of two species of horn sharks were collected from the upper Gulf of
California and their spiral intestines examined for onchobothriid tapeworms. Heterodontus francisci was found to
host Acanthobothrium puertecitense n. sp. Based on the criteria of Ghoshroy & Caira (2001), this is a Category
4 species (with >15mm total length, >50 proglottids, >80 testes and a symmetrical ovary). It differs from the
four Category 4 species previously reported from either the western Atlantic or eastern Pacific Oceans, including
A. bajaense (emend.) which was described from H. francisci, in hook-shape, total length, number of proglottids,
number of testes and number of columns of vitelline follicles. H. mexicanus was found to host Acanthobothrium
santarosaliense n. sp., which, based on the criteria of Ghoshroy & Caira (2001) is a Category 3 species (with
>15mm total length, >50 proglottids, >80 testes and an asymmetrical ovary). A. santarosaliense differs from the
five Category 3 species previously reported from either the western Atlantic or eastern Pacific Oceans in its size,
euapolytic nature, lack of dark matrix from the bases of the hooks, hook-shape, and distribution and number of
testes. In addition to the difference in ovarian symmetry, A. puertecitense possesses fewer testes, fewer proglottids,
is a shorter worm, possesses a more anterior genital pore and vitelline follicles which extend further forward
anteriorly than in A. santarosaliense. This represents the first report of onchobothriids from H. mexicanus and the
first report of onchobothriids from H. francisci in the Gulf of California. The apparent host-specificity of both new
species should be viewed with caution until a greater sample of host individuals of both species can be examined.
The fact that both new tapeworm species are known only from the Gulf is also considered to be a preliminary result,
given the small sample of host individuals examined from these regions to date. This brings the total number of
species of Heterodontus known to host Acanthobothrium to four and the total number of Acanthobothrium species
described from heterodontiform sharks to five.

Introduction quently reported this species, noting slight differences

The Order Heterodontiformes, or horn sharks, is a
relatively small group of sharks, consisting of the
single genus Heterodontus Blaineville with a total of
only eight species world-wide (Compagno, 1984). To
date, onchobothriid tapeworms have been reported
from three species of Heterodontus. Yamaguti (1934)
described Acanthobothrium cestraciontis Yamaguti,
1934 from H. japonicus (Maclay & Macleay) (as Ces-
tracion japonicus) in Japan. Drummond (1937) de-
scribed A. heterodonti Drummond, 1937 from H. por-
tusjacksoni (Meyer) (as H. philippi) off Lady Julia
Percy Island in Tasmania; Prudhoe (1969) subse-
in morphology, from the same host species in the Der-
went River in Tasmania. In North America, Appy &
Dailey (1973) described A. bajaense Appy & Dailey,
1973 (emend.) from H. francisci (Girard) from San
Quintin Bay on the west coast of the Baja Penninsula,
Mexico.
A recent survey of the metazoan parasites of elas-
mobranchs of the Gulf of California provided the
opportunity for investigation of the tapeworms of Het-
erodontus mexicanus Taylor & Castro-Aguirre, for
the first time, and of the tapeworms of H. francisci
from a new locality, the east coast of the Baja Penin-
sula in the waters of the Gulf of California. Two
220
new species of Acanthobothrium Beneden, 1849 re-
sulting from these collections are described below.
Host-specificity exhibited by Acanthobothrium in the
sympatric congeners H. francisci and H. mexicanus is
also addressed.
Materials and methods
All sharks were collected by local fishermen with
gill-nets at four localities in the Gulf of California,
Mexico, in 1993 and 1996. Nineteen Heterodontus
mexicanus were examined for tapeworms. These con-
sisted of one male and two females from Puertecitos,
nine males and five females from Santa Rosalia and
one male and one female from Loreto. Seven indi-
viduals of H. francisci were examined for tapeworms.
These consisted of two males and two females from
Puertecitos, one male from Bahia de Los Angeles and
one male and one female from Santa Rosalia. The spi-
ral intestine of each shark was removed, opened with
a longitudinal incision along the primary mesenteric
vessel and fixed in 10% buffered formalin in the field.
Worms were subsequently removed and transferred to
70% ethanol for storage.
Specimens prepared for light microscopy were
hydrated in a graded ethanol series, transferred to
distilled water, stained in Delafield’s hematoxylin,
blued in tap-water, dehydrated in a graded ethanol se-
ries, cleared in xylene and mounted on glass slides
in Canada balsam. Sufficient material of one of the
new species was available to allow examination with
scanning electron microscopy (SEM). This specimen
was hydrated in a graded ethanol series, transferred
to distilled water, postfixed in 1% osmium tetrox-
ide overnight, dehydrated in a graded ethanol se-
ries, transferred to hexamethyldisilazane (Ted Pella
Inc., Redding, CA) for approximately 15 minutes
and, following removal of the bulk of the hexam-
ethyldisilazane, was allowed to air-dry. The dried
specimen was mounted on an aluminum stub with
carbon tape and grounded with carbon paint before
being sputter coated with ca. 10 nm of gold/palladium.
Scanning electron microscopy was performed using a
LEO/Zeiss DSM 982 Gemini Field Emission Scan-
ning Electron Microscope.
Illustrations were prepared with the aid of a draw-
ing tube. Measurements are given in micrometres
unless specified otherwise. The measurements for both
new species are given in the text as the range, fol-
lowed in parentheses by the mean, standard deviation,
number of observations made when more than one
structure per worm was measured and number of
worms measured. Hook measurements were taken and
are presented following the formula of Euzet (1959);
hook terminology follows that of Caira (1985). Total
length of the scolex was measured from the anterior
margin of the apical sucker of the bothridium to the
posterior margin of the cephalic peduncle. Testis num-
ber is presented, following Caira et al. (1999), as the
total number of testes as well as the number in the
primary testicular field and the post-vaginal field.
Museum abbreviations used are as follows: CNHE,
Coleccion Nacional de Helmintos, Instituto de Bi-
ologia, Universidad Nacional Autónoma de Méx-
ico, Mexico City, Mexico; LRP, Lawrence R. Pen-
ner Parasitology Collection, Department of Ecology
& Evolutionary Biology, University of Connecticut,
Storrs, Connecticut, USA; MPM, Meguro Parasitol-
ogy Musum, Tokyo, Japan; USNPC, US. National
Parasite Collection, Beltsville, Maryland, USA.
Acanthobothrium is currently the most speciose of
the onchobothriid genera (Schmidt, 1986), with well
over 100 nominal species. This genus was comprehen-
sively treated by both Goldstein (1967) and Williams
(1969). However, more than 50 new species have
been assigned to the genus since 1969. The strat-
egy described and followed by Ghoshroy & Caira
(2001), and also utilised by Caira & Burge (2001),
has been adopted here to facilitate comparisons be-
tween the new species and the numerous previously
described species of Acanthobothrium. Thus, both
new species are compared in detail to the 58 species
of Acanthobothrium previously reported from the east-
ern Pacific or western Atlantic Oceans, as listed by
Ghoshroy & Caira (2001) and augmented by Caira &
Burge (2001). This was accomplished as follows: the
new species are characterized according to their pos-
session of one of two states of each of the following
four characters: total length (≤ 15 vs > 15 mm), num-
ber of proglottids (≤ 50 vs > 50), number of testes (≤
80 vs > 80) and ovarian symmetry (poral and aporal
lobes symmetrical or asymmetrical with one another
in terms of their anterior extent). Articulation of dif-
ferentiating features is provided for all congeners ex-
hibiting the same coding, and thus the same Category
designation, as either of the new species. For con-
venience, the Categories recognised by Ghoshroy &
Caira (2001) for coding of these features are repeated
here: 1=SFFS, 2=SFFA, 3=LMMA, 4=LMMS,
5=LMFS, 6=LMFA, 7=LFFA, 8=SMFS, 9=LFFS,
10=SMMS, where each category represents the condi-

tions of the above features exhibited by a species: total
length (Short or Long), number of proglottids (Few
or Many), number of testes (Few or Many) and ovary
symmetry (Symmetrical or Asymmetrical).
In addition, each new species is explicitly dis-
tinguished from species from other regions of the
world with which it is similar in all three of the
following conspicuous and easy to assess morpholog-
ical features: total length, number of proglottids, and
number of testes per proglottid. The ranges of these
three features considered to be similar enough to jus-
tify explicit comparison are provided at the beginning
of the ‘Remarks’ section for each new species. If a
new species is not explicitly compared with another
species, it should be assumed that it differs from that
species in at least one of these three characters. Fi-
nally, the new species are also explicitly compared
to the three species of Acanthobothrium previously
described from horn sharks.
In addition to the material of Acanthobothrium
listed in Ghoshroy & Caira (2001) and Caira & Burge
(2001), one paratype (USNPC No. 72568) and two
voucher specimens (from the personal collection of
Murray Dailey) of A. bajaense Appy & Dailey, 1973
and the type of A. cestraciontis Yamaguti, 1934 (MPM
No. SY3090) were examined. We were unable to lo-
cate the type-material of A. heterodonti Drummond,
1937; however, several voucher specimens of what
appear to be A. heterodonti, collected from H. portus-
jacksoni in Australia, were examined.
Acanthobothrium puertecitense n.sp.
Description (Figures 1-9)
(Based on 6 whole-mounts; scolex of one prepared for
SEM). Worms euapolytic, 15.2-24.6 (18.7 ± 4.3; 4)
mm long, greatest width at scolex; 110-165 (133 ±
21.9; 5) proglottids per worm. Scolex 1,040-1,380
(1, 182 ± 130.3; 5) × 550-624 (583.5 ± 34.6; 4),
consisting of cephalic peduncle and 4 muscular ac-
etabula in form of bothridia. Bothridia free anteriorly
and posteriorly, 608-768 (694.4 ± 50.1; 10; 5) × 264-
344 (301.8 ± 27.5; 8; 5); each bothridium with 3
loculi and specialised anterior region in form of mus-
cular pad. Anterior loculus 200-320 (271 ± 38.4; 10;
5) long; middle loculus 110-170 (134.6 ± 20; 10; 5)
long; posterior loculus 90-144 (116.4 ± 18.9; 10; 5)
long; ratio of loculus lengths (anterior: middle: pos-
terior) 2.1-2.6 (2.4 ± 0.2; 10; 5): 1-1.3 (1.2 ± 0.1;
221
10; 5): 1; greatest width of scolex at level of first
costa. Velum present between medial margins of adja-
cent bothridia approximately at level of second costa.
Muscular pad 50-80 (70 ± 9.8; 7; 5) × 153-180
(166.2 ± 9.9; 7; 5), with conspicuously rounded pos-
terolateral margins, bearing single apical sucker and
one pair of hooks; apical sucker oval, 30-50 (43.2 ± 5;
10; 5) × 45-80 (58.1 ± 13.0; 9; 5), with inconspic-
uously thickened anterior margin. Hooks bipronged,
hollow; internal channels of axial and abaxial prongs
continuous, smooth; hooks each with single knob-like
tubercle on proximal surface of axial prong; medial
and lateral hooks approximately equal in size. Ax-
ial prongs of medial and lateral hooks conspicuously
longer than abaxial prongs. Lateral hook formula:
75 − 85(80.8 ± 3.4; 6; 5)145 − 177.5(160.8 ± 10.5; 6; 5)95 − 118(103.4 ± 0.1; 6; 5)
202.5 − 247.5(220.9 ± 15.5; 7; 5)
Medial hook formula:
80 − 90(83.8 ± 3.5; 6; 5)150 − 167.5(156.1 ± 6.6; 7; 5)95 − 113(101.9 ± 5.9; 7; 5)
215 − 247.5(220.8 ± 12.8; 8; 5)
Bases of medial and lateral hooks approximately equal
in length, dark grey in colour; base of lateral hook
resting upon base of medial hook. Bases and anterior
portions of both prongs of medial and lateral hooks
embedded in musculature of scolex. Cephalic pedun-
cle 540-770 (634 ± 85.6; 5) × 256-320 (291.2 ± 28.6;
5).
Scolex proper and velum covered with short fili-
form microtriches only. Proximal bothridial surfaces
(Figure 8) and cephalic peduncle (Figure 9) densely
covered with blade-like spiniform microtriches inter-
spersed with long filiform microtriches. Distal both-
ridial surfaces (Figure 7) densely covered with short
filiform microtriches and occasional blade-like spini-
form microtriches. Surfaces of strobila covered with
long filiform microtriches only.
Proglottids slightly craspedote. Immature proglot-
tids 108-163 (131.4 ± 22.2; 5) in number, initially
wider than long, becoming longer than wide with
maturity. Mature proglottids 1-2 (1.7 ± 0.6; 3) in num-
ber, 968-1,780 (1, 461.6 ± 300.4; 5; 3) × 410-550
(474 ±62.3; 5; 3). Gravid proglottids not seen. Genital
pores lateral, irregularly alternating, opening anterior
to ovary, 41-60% (50.5±5.8; 8; 5) of proglottid length
from posterior of proglottid, slightly more anterior
in terminal proglottid than in subterminal proglottids.
Testes 74-111 (96.8 ± 10.9; 12, 6) in total number, 62-
102 (85.3 ± 10.7; 12; 6) in primary field, 8-16 (11.4 ±
1.9; 12; 6) in post-vaginal field, generally arranged in
4-6 irregular columns in primary field and 1-2 irregular
columns in post-vaginal field, extending from ovarian
222

Figures 1–5. Acanthobothrium puertecitense n. sp. 1. Scolex. 2. Hooks. 3. Terminal mature proglottid. 4. Detail of terminal genitalia. 5.
Reconstruction of whole worm. Arrows indicate boundaries between fragments of actual worm.
 223

Figures 6–9. Scanning electron micrographs of scolex of Acanthobothrium puertecitense n. sp. 6. Scolex. 7. Enlarged view of distal bothridial
surface. 8. Enlarged view of proximal bothridial surface. 9. Enlarged view of cephalic peduncle. Scale-bars: 6, 100 µm; 7-9, 2 µm.

isthmus between ovarian lobes to near anterior mar-
gin of proglottid, one layer deep, 25-50 (34.6 ± 6.5;
21; 3) × 40-95 (59.3 ± 16.1; 21; 3). Vas deferens
extensive, coiled, looping anteriorly along medial line
of proglottid into anterior sixth of proglottid. Cirrus-
sac pyriform, bent anteriorly, 104-176 (144 ± 30; 5;
3) × 184-264 (219.2 ± 32.3; 5; 3), containing coiled
cirrus; cirrus densely covered with slender spiniform
microtriches. Vagina thick-walled, slightly sinuous,
extending along anterior margin of cirrus-sac from
genital pore to median line of proglottid, slightly
overlapping proximal portion of cirrus-sac, extending
posteriorly along median line of proglottid to oötype;
base of vagina not conspicuously expanded; vagi-
nal sphincter absent. Ovary H-shaped in dorsoventral
view, tetralobed in cross-section, follicular, located at
posterior end of proglottid, 216-610 (408.8 ± 163.5;
5; 3) × 230-304 (262.8 ± 32.4; 5; 3); ovarian arms
symmetrical, both stopping approximately 2-4 testes
posterior to cirrus-sac, ovarian bridge in posterior half
of ovary. Mehlis’ gland present posterior to ovarian
bridge between posterior lobes of ovary. Vitellarium
224
consisting of 2 lateral bands; each band consisting of
one dorsal and one ventral column of follicles, extend-
ing from near posterior margin of ovary to anterior
margin of testicular field, interrupted by cirrus-sac and
vagina on poral side, not interrupted by ovary. Uterus
sacciform, extending anteriorly from ovarian bridge
along median line of proglottid stopping just short of
anterior margin of testicular field. Uterine pore absent.
Eggs not seen. Excretory ducts lateral.
Type-host: Heterodontus francisci (Girard), Horn
shark.
Site of infection: Spiral intestine.
Type-locality: Puertecitos, Gulf of California, Mexico
(30◦21 N, 114◦39 W).
Additional localities: None.
Specimens deposited: Holotype (CNHE No. 4175); 2
paratypes (CNHE No. 4176); 1 paratype (USNPC No.
90843); 2 paratypes (LRP Nos. 2105-2106).
Etymology: This species is named for its type-locality.
Remarks
Acanthobothrium puertecitense n. sp. is greater than
15 mm in total length, possesses more than 50 proglot-
tids and more than 80 testes and exhibits a symmet-
rical ovary. Thus, it is a Category 4 species based
on the criteria of Ghoshroy & Caira (2001). It can
be distinguished from the three Category 4 species
reported previously from the eastern Pacific Ocean (ta-
ble II of Ghoshroy & Caira, 2001) as follows. It is a
much smaller worm (<25 mm vs 22.4 cm), has fewer
proglottids (
165 vs 500) and fewer testes (74-111
vs 147-186) than A. bajaense, which is also reported
to parasitise H. francisci. Unlike A. microcephalum
Alexander, 1953 from Myliobatis californica Gill,
A. puertecitense has medial and lateral hooks in which
the abaxial prongs are conspicuously shorter than the
axial prongs (rather than axial and abaxial prongs that
are approximately equal in length), and has only two
(rather than three to four) columns of vitelline follicles
on either side of the proglottid. A. puertecitense can be
distinguished from A. robustum Alexander, 1953 from
Rhinobatos productus Ayres in having larger hooks
with abaxial prongs that extend well beyond the abax-
ial prong tubercle (rather than stopping short of the
tubercle) and in that its hooks lack the small process
associated with the base of the abaxial prong found in
A. robustum.
A. puertecitense is distinguished from A. terezae
Rego & Dias, 1976 from Potamotrygon motoro Nater-
rer, the only Category 4 species reported previ-
ously from the western Atlantic Ocean (table III of
Ghoshroy & Caira, 2001) in that it is a smaller worm
(15.2-24.6 vs 88-110 mm total length) with fewer
proglottids (110-165 vs 200-260) and fewer testes
(74-111 vs 120-148).
With respect to the two other species of Acan-
thobothrium previously reported from heterodontif-
orm sharks, A. puertecitense is a much smaller worm
(up to 15.2-24.6 vs
282 mm in total length) and has
relatively longer abaxial hook prongs (conspicuously
longer than the axial prong vs only slightly longer)
than A. cestraciontis. In addition, the bothridial mus-
cular pads in A. puertecitense bear an apical sucker,
whereas this feature appears to be lacking from the
muscular pads of A. cestraciontis. A. puertecitense is
a much smaller worm (15.2-24.6 vs
340 mm total
length) with smaller mature proglottids (up to 1.8 ×
0.55 vs up to 2.7 × 2 mm) than A. heterodonti.
Finally, A. puertecitense can be distinguished from
all described species of Acanthobothrium from other
regions of the world in that it is > 10 and < 30 mm in
total length, possesses > 100 and < 180 proglottids
and possesses > 70 and < 120 testes. It should be
noted that A. puertecitense is ‘Acanthobothrium n. sp.
1’ of Caira, Jensen & Healy (2001).
Acanthobothrium santarosaliense n. sp.
Description (Figures 10-14)
(Based on 4 whole-mounts consisting of 3 complete
specimens and one strobila without scolex). Worms
euapolytic, 37.7-44.9 (40.8±3.8; 3) mm long; greatest
width at scolex, 174-215 (188 ± 23.4; 3) proglottids
per worm. Scolex 1,440-1,520 (1,486.7 × 41.6; 3)
× 704-840 (772 ± 96.2; 2), consisting of cephalic
peduncle and 4 muscular acetabula in form of both-
ridia. Bothridia free anteriorly and posteriorly, 704-
888 (818.7 ± 59.9; 9; 3) × 340-432 (378 ± 35.1; 5; 3);
each bothridium with 3 loculi and specialized anterior
region in form of muscular pad. Anterior loculus 280-
360 (318.4 ± 29.6; 5; 3) long; middle loculus 130-184
(165.6 ± 20.9; 5; 3) long; posterior loculus 120-168
(139.6 ± 18.1; 10; 5) long; ratio of loculus lengths
(anterior: middle: posterior) 1.9-3 (2.3 ± 0.4; 5; 3):
1-1.4 (1.2 ± 0.2; 5; 3): 1; greatest width of scolex
at level of first costa. Velum present between medial
margins of adjacent bothridia slightly posterior to level
of second costa. Muscular pad 70-120 (92.5 ± 23.6;
 225

Figures 10–14. Acanthobothrium santarosaliense n. sp. 10. Scolex. Note that view of scolex drawn is of lower pair of bothridia on mounted
specimen. 11. Hooks. 12. Mature proglottid. 13. Detail of terminal genitalia. 14. Whole worm.
226
4; 3) × 194-202 (199.2 ± 3.0; 4; 3), with conspic-
uously rounded posterolateral margins, bearing single
apical sucker and one pair of hooks; apical sucker oval,
60-90 (73 ± 15.4; 4; 3) × 90-109 (102.3 ± 10.7; 3;
2), with inconspicuously thickened anterior margin.
Hooks bipronged, hollow; internal channels of axial
and abaxial prongs continuous, smooth; hooks each
with single knob-like tubercle on proximal surface of
axial prong; medial and lateral hooks approximately
equal in size. Axial prongs of medial and lateral hooks
conspicuously longer than abaxial prongs.
Lateral hook formula:
88 − 96(92.5 ± 4.4; 5; 3)146 − 170(160 ± 10; 4; 3)88 − 110(99.3 ± 8.6; 5; 3)
228 − 234(230 ± 2.9; 4; 3)
Medial hook formula:
82 − 95(90 ± 6; 5; 3)158 − 165(160 ± 3.3; 4; 3)105 − 122.5(112.6 ± 7.4; 3; 5)
224 − 232(227.9 ± 3.3; 4; 3)
Bases of medial and lateral hooks approximately equal
in length, base of lateral hook resting upon base of me-
dial hook. Bases and anterior portions of both prongs
of medial and lateral hooks embedded in musculature
of scolex. Cephalic peduncle 740-860 (783.3 ± 66.6;
3) × 360-512 (429.3 ± 76.9; 3).
Microtriches observed with light microscopy only.
Spiniform microtriches present on proximal bothridial
and peduncular surfaces.
Proglottids slightly craspedote. Immature proglot-
tids 164-212 (181.7 ± 26.4; 3) in number, initially
wider than long, becoming longer than wide with
maturity. Mature proglottids 3-9 (5.6 ± 3.1; 3) in num-
ber, 900-1,620 (1, 239.4 ± 263.3; 16; 4) × 480-720
(638.1 ± 41.8; 16; 4). Gravid proglottids not seen.
Genital pores lateral, irregularly alternating, open-
ing anterior to ovary, 38-49% (43 ± 0.04; 15; 4) of
proglottid length from posterior of proglottid. Testes
114-172 (134.8 ± 20.5; 14; 4) in total number, 98-
149 (117.9 ± 17.8; 14; 4) in primary field, 12-25
(16.8 ± 3.9;14; 4) in post-vaginal field, generally
arranged in 5-7 irregular columns in primary field and
2-3 irregular columns in post-vaginal field, extend-
ing from ovarian isthmus between ovarian lobes to
near anterior margin of proglottid, one layer deep,
20-57.5 (39.3 ± 8.3; 32; 4) × 28-82 (55.8 ± 13.7;
32; 4). Vas deferens extensive, coiled, looping an-
teriorly along medial line of proglottid into anterior
fifth of proglottid. Cirrus-sac conspicuously bent an-
teriorly, 72-152 (106.5 ± 24.3; 19; 4) × 208-280
(247.2 ± 22.5;19; 4), containing coiled cirrus; prox-
imal portion of cirrus densely covered with slender
spiniform microtriches. Vagina thick-walled, slightly
sinuous, extending along anterior margin of cirrus-
sac from genital pore to median line of proglottid, not
overlapping proximal portion of cirrus-sac, extending
posteriorly along median line of proglottid to oötype;
base of vagina not conspicuously expanded; vagi-
nal sphincter absent. Ovary H-shaped in dorsoventral
view, tetralobed in cross-section, conspicuously fol-
licular, located at posterior end of proglottid, 240-430
(341.9 ± 71.9; 15; 4) × 320-432 (366 ± 35.5;15;
4); ovarian arms asymmetrical; aporal ovarian arm
extending almost to level of posterior margin of cirrus-
sac; poral arm stopping approximately 3 testes short of
posterior margin of cirrus-sac; ovarian bridge in pos-
terior half of ovary. Mehlis’ gland present posterior
to ovarian bridge between posterior lobes of ovary.
Vitellarium consisting of 2 lateral bands; each band
consisting of one dorsal and one ventral column of fol-
licles, extending from near posterior margin of ovary
to approximately 3 testes short of anterior margin of
testicular field, interrupted by cirrus-sac and vagina on
poral side, not interrupted by ovary. Uterus sacciform,
extending anteriorly from ovarian bridge along median
line of proglottid, stopping 3-4 testes short of anterior
margin of testicular field. Uterine pore absent. Eggs
not seen. Excretory ducts lateral.
Type-host: Heterodontus mexicanus Taylor & Castro-
Aguirre, Mexican hornshark.
Site of infection: Spiral intestine.
Type-locality: Santa Rosalia, Gulf of California, Mex-
ico, 27◦ 19 N, 112◦17 W.
Additional localities: None.
Specimens deposited: Holotype (CNHE No. 4177);
one paratype (CNHE No. 4178); one paratype (US-
NPC No. 90844); one paratype (LRP No. 2107).
Etymology: This species is named for its type-locality.
Remarks
Acanthobothrium santarosaliense n. sp. is greater than
15 mm in total length, possesses more than 50 proglot-
tids and more than 80 testes, and exhibits an ovary
that is asymmetrical. Thus, based on the criteria of
Ghoshroy & Caira (2001), it is a Category 3 species. It
can be distinguished from the four other potential Cat-
egory 3 species reported previously from the eastern
Pacific Ocean (table II of Ghoshroy & Caira, 2001) as
follows. A. santarosaliense differs from A. cleofanus
Monks, Brooks & Ponce de Leon, 1996 from Dasyatis
longus (Garman) in that it is a much smaller worm
(37.7-44.9 vs 370-670 mm in total length), is euapoly-
tic (vs anapolytic) and has hooks that lack the dark
matrix seen on the bases of the hooks of A. cleofanus.

The hooks of A. santarosaliense differ in shape from
those of A. holorhini Alexander, 1953 and A. mac-
ulatum Riser, 1955 from Myliobatis californica and
A. chilense Rego, Vincente & Herrera, 1968 (emend.)
from Sarda chilensis; whereas the hooks of A. san-
tarosaliense bear axial prongs that are conspicuously
longer than the abaxial prongs, the axial and abax-
ial prongs of the hooks in all three of the former
species are almost equal in length. In addition, both
A. holorhini and A. maculatum possess fewer testes
than A. santarosaliense (60-77 and 75-85 vs 114-172,
respectively). A. chilense is also a larger worm (up to
170 vs 37.7-44.9 mm in total length) with more testes
(up to 300 vs 114-172) than A. santarosaliense.
A. santarosaliense differs from A. tortum (Linton,
1916) Baer & Euzet, 1962 from Aetobatus narinari
(Euphrasen), the only Category 3 species reported pre-
viously from the western Atlantic Ocean (table II of
Ghoshroy & Caira, 2001) in that it is smaller (37.7-
44.9 vs 220 mm in total length) but has more testes
(114-172 vs c. 90). A. santarosaliense also possesses
post-vaginal testes, which are apparently lacking in
A. tortum.
With respect to the four species of Acan-
thobothrium previously reported from heterodontif-
orm sharks, A. santarosaliense is smaller (37.7-44.9
vs up to 282 mm) than A. cestraciontis and has hooks
that differ in shape (the abaxial prong is conspicuously
longer than the axial prong vs only slightly longer). In
addition, whereas the muscular pads on the bothridia
of the former species each clearly exhibit an apical
sucker, this feature appears to be lacking from the
muscular pads of A. cestraciontis. A. heterodonti from
Heterodontus portusjacksoni possesses bothridia and
loculi that are approximately twice as long as those of
A. santarosaliense (0.6, 0.3, 0.3 vs 0.32, 0.16, 0.14
for anterior, middle and posterior loculi, respectively).
In addition, the abaxial prongs of the hooks in A. het-
erodonti are only slightly shorter than the axial prongs,
whereas they are conspicuously shorter in A. santaros-
aliense. A. santarosaliense is smaller than A. bajaense
from H. francisci (37.7-44.9 vs up to 224 mm in total
length) and possesses fewer proglottids (174-215 vs

500) than A. bajaense. A. santarosaliense is larger
(37.7-44.9 vs 15.2-24.6 mm total length), has more
proglottids (174-215 vs 110-165), more testes (114-
172 vs 74-111) and a more posterior genital pore (38-
49% vs 41-60% from posterior) than A. puertecitense
n. sp. from H. francisci. In addition, whereas the
vitelline follocles in A. santarosaliense stop short of
227
the anterior margin of the testicular field, they extend
to the anterior margin of this field in A. puertecitense.
A. santarosaliense can be distinguished from all
species of Acanthobothrium from other geographical
regions, except A. ijimai Yoshida, 1917 from Dasyatis
akajei (Muller & Henle) and A. microcephalum from
Myliobatis californica, in that it is > 20 and < 50 mm
in total length, possesses > 150 and < 250 proglottids
and possesses > 100 and < 200 testes. A. santaros-
aliense differs from both A. ijimai and A. micro-
cephalum in the shape of its hooks, which possess
abaxial prongs that are substantially longer than the
axial prongs. It is further distingued from A. ijimai in
the form of its bothridia which are free only slightly
anteriorly and posteriorly, rather than free extensively
posteriorly, giving the scolex the appearance of a
four-leaved clover, and from A. microcephalum in the
shape and size of its mature proglottids which are
conspicuously longer than wide in the former species
(900-1,620 × 480-720) but only slightly longer than
wide in the latter species (490-630 × 430-470).
Discussion
Based on their results for the stingray Dasyatis brevis
(Garman), Ghoshroy & Caira (2001) briefly addressed
whether the Gulf of California hosts a unique fauna of
elasmobranch tapeworms. However, with only a single
record of tapeworms from D. brevis outside the Gulf,
and that record coming from a locality well to the
south, their conclusions were limited. Data available
for Heterodontus francisci are somewhat more appro-
priate for addressing this question because Appy &
Dailey (1973) examined this species from a locality
on the western coast of the Baja Penninsula. Whereas
individuals of H. francisci in the Gulf hosted only
A. puertecitense n. sp., Appy & Daily (1973) found
that individuals in the eastern Pacific Ocean hosted
only A. bajaense. These data are consistent with the
hypothesis that differences exist between the elasmo-
branch tapeworm fauna within and outside of the Gulf
of California. It is of note, however, that both stud-
ies involved examination of only a very small number
of sharks (seven in this study; three by Appy & Dai-
ley, 1973). Unless prevalences of both Acanthoboth-
rium species are normally rather high, these results
may not reflect all species of tapeworms found in H.
fancisci in either geographical region. As this repre-
sents the first record of onchobothriid tapeworms from
228
Table 1. Acanthobothrium species known from Heterodontus species.

Heterodontus species Acanthobothrium species Locality

H. francisci A. bajaense Appy & Dailey, 1973 San Quintin Bay, Mexico
A. puertecitense n. sp. Puertecitos, Mexico
H. japonicus A. cestraciontis Yamaguti, 1934 Japan
(as Cestracion japonicus)
H. mexicanus A. santarosaliense n. sp. Santa Rosalia, Mexico
H. portusjacksoni A. heterodonti Drummond, 1937 Perey Island, Tasmania
(as H. philippi)

H. mexicanus in any locality, no such comparative Acknowledgements

geographical data are available for A. santarosaliense.
With respect to host-specificity of Acanthoboth-
rium species in the sympatric congeners H. francisci
and H. mexicanus, the data presented here support
strict host-specificity; A. puertecitense is described
solely from H. francisci and A. santarosaliense is
described solely from H. mexicanus. However, we en-
countered a number of immature specimens of Acan-
thobothrium that could not be identified to species.
This, combined with the small sample sizes on which
the results of this study are based, suggest that addi-
tional sampling is required to confirm these prelimi-
nary observations.
This brings the total number of Acanthobothrium
species reported from heterodontiform sharks to five.
These species are summarised in Table 1 along with
their hosts. That all four of the eight species of
Heterodontus examined have been found to be par-
asitised by one or more species of Acanthobothrium
suggests that the other four species of Heterodontus
likely host additional species of Acanthobothrium. In-
vestigation of the four species of Heterodontus not
previously examined for tapeworms, i.e. H. galea-
tus (Günther), H. quoyi (Fréminville), H. ramalheira
(Smith) and H. zebra (Gray), is highly recommended.
Current records indicate that heterodontiform sharks
are generally parasitised by relatively large species
of Acanthobothrium; all five known species are over
1.5 cm in total length and the largest, A. heterodonti,
obtains a length of up to 34 cm. Thus, all are Cate-
gories 3 or 4 species sensu Ghoshroy & Caira (2001).
Based on these data we predict that the above four Het-
erodontus species will also be found to host relatively
large species of Acanthobothrium.
We are grateful to P. Cislo, S. Curran, K. Jensen, P.
Olson and G. Tyler for their assistance with dissections
of sharks in Mexico. This research was supported by
National Foundation grants No. DEB 9300796 (with
an REU supplement to SDZ) and No. DEB 9532943
to JNC (and G. W. Benz and C. Henry, respectively).
All collections in Mexico were made under the condi-
tions set forth in collecting permit No. 120496-213-03
issued to JNC by the Secretaria de Medio Ambiente
Recursos Naturales y Pesca, México.
References
Appy, R.G. & Dailey, M.D. (1973) Two new species of Acan-
thobothrium (Cestoda: Tetraphyllidea) from elasmobranchs of
the Eastern Pacific. Journal of Parasitology, 59, 817-820.
Caira, J.N. (1985) Calliobothrium evani sp. n. (Tetraphyllidea: On-
chobothriidae) from the Gulf of California, with a redescription
of the hooks of C. lintoni and a proposal for onchobothriid
hook terminology. Proceedings of the Helminthological Society
of Washington, 52, 166-174.
Caira, J.N. & Burge, A.L. (2001) Three new species of Acan-
thobothrium (Cestoda: Tetraphyllidea) from the ocellated electric
ray, Diplobatis ommata, in the Gulf of California, México.
Comparative Parasitology, 68, 52-65.
Caira, J.N., Jensen, K. & Healy, C.J. (2001) Interrelationships
among tetraphyllidean and lecanicephalidean cestodes. In: Lit-
tlewood, D.T.J. & Bray, R.A. (Eds) Interrelationships of the
Platyhelminthes. London: Taylor & Francis, pp. 135-158.
Compagno, L. (1984) FAO species catalogue. VI. 4, Parts 1 and
2: Sharks of the world. An annotated and illustrated catalogue
of shark species known to date. FAO Fisheries Synopsis, 125,
655 pp.
Drummond, F.H. (1937) Cestoda. Lady Julia Percy Island. Reports
of the expedition of the McCoy Society for Fish Investigations
and Research. Proceedings of the Royal Society of Victoria, 49,
401-404.
Euzet, L. (1959) Recherches sur les cestodes tétraphyllides des
sélaciens des côtes de France. Docteur és Sciences Naturelles
Thèse, Université de Montpellier, Montpellier, France, 263 pp.

Ghoshroy, S. & Caira, J.N. (2001) Four new species of Acan-
thobothrium (Cestoda: Tetraphyllidea) from the whiptail stingray
Dasyatis brevis in the Gulf of California. Journal of Parasitology,
87, 354-372.
Goldstein, R.J. (1967). The genus Acanthobothrium van Beneden,
1849 (Cestoda: Tetraphyllidea). Journal of Parasitology, 53,
455-483.
Prudhoe, S. (1969) Cestodes from fish, birds and whales. Reports of
B.A.N.Z. Antarctic Research Expedition, 1929-1930, Series B, 8,
171-193.
229
Schmidt, G.D. (1986). Handbook of tapeworm identification. Boca
Raton: CRC Press, 675 pp.
Williams, H.H. (1969). The genus Acanthobothrium Beneden, 1849
(Cestoda: Tetraphyllidea), Nytt Magasin for Zoologi, 17, 1-56.
Yamaguti, S. (1934) Studies on the helminth fauna of Japan. Part 3.
Cestodes of fishes. Japanese Journal of Zoology, 6, 1-112.