Redescription of Hapalotrema mistroides (Monticelli, 1896) and Hapalotrema synorchis

Luhman, 1935 (Digenea: Spirorchidae), with Comments on Other Species in the Genus
Author(s): Thomas R. Platt and David Blair
Source: The Journal of Parasitology, Vol. 84, No. 3 (Jun., 1998), pp. 594-600
Published by: Allen Press on behalf of The American Society of Parasitologists
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 J. Parasitol., 84(3), 1998 p. 594-6(X)
? American Society of Parasitologists 1998

REDESCRIPTION OF HAPALOTREMA MISTROIDES (MONTICELLI, 1896) AND

HAPALOTREMA SYNORCHIS LUHMAN, 1935 (DIGENEA: SPIRORCHIDAE),
WITH COMMENTS ON OTHER SPECIES IN THE GENUS

Thomas R. Platt and David Blair*

Department of Biology, Saint Mary's College, Notre Dame, Indiana 46556

ABSTRACT: Hapalotrema mistroides (Monticelli, 1896) Stiles and Hassall, 1908 and Hapalotrema synorchis Luhman, 1935 are
redescribed using specimens from the loggerhead turtle, Caretta caretta (L.). The redescription of H. mistroides is based on
specimens collected by Looss and described as Hapalotrema constrictum Leared, 1862 and renamed Hapalotrema loossi Price,
1934, which is now considered a junior synonym of H. mistroides. Hapalotrema orientalis Takeuti, 1942 is considered a junior
synonym of Hapalotrema synorchis. The redescriptions are consistent with the originals but provide an unreported range of
variation while adding new information about the reproductive systems. Reexamination of Hapalotrema dorsopora Dailey, Fast,
and Balazs, 1993 confirms the presence of a reproductive system typical for the genus and the absence of a dorsal genital pore;
therefore, H. dorsopora is reduced to a junior synonym of Hapalotrema mehrai Rao, 1976. Four valid species of Hapalotrema
are recognized: H. mistroides, H. synorchis, H. postorchis, and H. mehrai, and a key to the species is provided.

As currently constituted, the genus Hapalotrema Looss, 1899 H. dorsopora Dailey, Fast, and Balazs, 1993, holotype (USNPC 82326),
paratype (USNPC 82327); Hapalotrema postorchis Rao, 1976, voucher
(Digenea: Spirorchidae) contains 8 species reported exclusively
(USNPC 82324, Dailey et al. [1993]); H. synorchis Luhman, 1935,
from the circulatory systems of marine turtles (Smith, 1997). holotype (USNPC 8909), and vouchers (USNPC 73323, Fischthal and
The discovery of specimens of Hapalotrema spp. from turtles Acholonu [1976]). Additional specimens from the junior author's col-
off the coast of Australia prompted a reassessment of Hapalo- lection were used, as was information from published descriptions.
trema mistroides (Monticelli, 1896) and Hapalotrema synorchis Numerous specimens of spirorchids have been collected from marine
turtles by the junior author during the past 2 decades. Many of the
Luhman, 1935.
turtles were examined opportunistically and some had been dead for
Price (1934) recognized significant nomenclatural problems undetermined lengths of time before necropsy. Methods of killing and
involving species assigned to both Hapalotrema and Learedius fixation varied with circumstances. Specimens were stained in Mayer's
and proposed several changes to rectify the situation. The hematoxylin, following standard procedures, cleared, and mounted in
names applied to species of both genera sensu Price (1934) Canada balsam as whole mounts for microscopic examination.
The following measurements are used in species descriptions: genital
have become the standard for the past 60 yr. We feel that Price
pore/TL = distance from the anterior end to the genital pore/total length
erred in several of his decisions (Platt and Blair, 1996), but that
and presented as a percentage; forebody/TL = distance from anterior
strict application of the rules of zoological nomenclature end to body constriction/total length and presented as a percentage.
(Anonymous, 1985) would not be in the best interests of no- Drawings were made with the aid of a drawing tube. Measurements are
menclatural stability. The International Commission for Zoo-in micrometers unless otherwise stated and presented as the range fol-
lowed by the mean in parentheses.
logical Nomenclature exercised its plenary power to recognize
Hapalotrema loossi Price, 1934 as the type species of the genus
and to retain the concept of Hapalotrema Looss, 1899 and DESCRIPTION
Learedius Price, 1934 as defined by Price (1934) (Anonymous,
Hapalotrema mistroides (Monticelli, 1896) Stiles and Hassall,
1997).
1908
Cribb and Gordon (1998) reexamined the status of several (Figs. 1, 2; Table I)
species of Hapalotrema and concluded that Hapalotrema dor-
sopora Dailey, Fast and Balazs, 1993 and Hapalotrema pam- Synonyms: Mesogonimus mistroides Monticelli, 1896; H. constrictum
banensis Gupta and Mehrotra, 1981 were junior synonyms of (Leared, 1862) Looss, 1899; H. loossi Price, 1934.
Hapalotrema mehrai Rao, 1976. It is our intention to review Diagnosis (based on 8 whole mounts and 1 sectioned specimen):
the remaining species of Hapalotrema, with particular attention
Body robust, maximum width in posterior /2 of body; division into
to the species commonly reported from the loggerhead turtle, forebody and hindbody indistinct, near acetabulum. Small tegumental
Caretta caretta. spines present anteriorly, diminishing posteriorly, absent posterior to
acetabulum. Oral sucker subterminal, wider than long; esophagus slight-
ly sinuous, expanded distally, surrounded by glandular cells at cecal
MATERIALS AND METHODS bifurcation. Intestinal ceca bifurcate at level of acetabulum, terminate

Whenever possible type material was examined. Specimens near wereposterior

ob- end of body. Acetabulum slightly pedunculate, ovoid,
tained from the United States National Parasite Collection (USNPC), with minute spines on margin. Testes in preovarian and postovarian
Beltsville and the Swedish Museum of Natural History (SMNH), Stock- groups. External seminal vesicle elongated transversely, located be-
holm. Specimens examined were: Hapalotrema constrictum (Leared) tween preovarian testes and ovary. Cirrus sac well developed; internal
(SMNH 1570, 1571, 1840, 1843, 1855, 2663, 2664, 2665, 2666, plus seminal vesicle coiled; cirrus spined. Common genital pore sinistral,
1 slide containing an entire worm in serial section with no number,near inside margin of left cecum. Ovary large; slightly lobed, between
from Looss [1899]); vouchers (USNPC 9634 and 51669, Looss [1899]); testicular groups. Oviductal seminal receptacle present, small; Laurer's
canal present, opening dorsally. Vitelline follicles diffuse, granular, be-
ginning slightly posterior to acetabulum, confluent medially anterior to
Received 26 November 1997; revised 3 February 1998; accepted 3 anterior testes group, terminating near cecal ends. Vitelline reservoir
February 1998. small, immediately posterior to ovary. Uterus small, passes left of ovary
to common genital pore at midovarian level. Excretory vesicle Y-
* Department of Zoology, James Cook University, Townsville, Queens-
land 4811, Australia. shaped.

594

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 PLATT AND BLAIR-REDESCRIPTION OF HAPALOTREMA SPP. 595

FIGURE 1-3. 1. Hapalotrema mistroides adult, ventral (scale bar = 1 mm); lectotype of Hapalotrema loossi. 2. Hapa
genitalia, ventral (scale bar = 250 am); lectotype of Hapalotrema loossi. 3. Hapalotrema synorchis adult, ventral (scale bar = 1 mm).

Taxonomic summary Remarks

Type host: Caretta caretta, the loggerhead turtle.
Other hosts: Chelonia mydas, the green turtle.
Type locality: Mediterranean Sea near Naples, Italy (Monticelli,
1896).
Other localities: Mediterranean Sea near Abukir, Egypt; Shark Bay,
Western Australia, Australia (present study; voucher specimens QM
G214179-81 from C. caretta).
Monticelli's (1896) description of H. mistroides was not complete by
current standards and our attempts to locate his specimens have been
unsuccessful. Price (1934) noted striking similarities between Monti-
celli's description and H. loossi. He preferred, however, to recognize 2
species based on several differences in the illustrations of the 2 forms:
number of testes in the postovarial group, 16 in H. mistroides and 10
in H. loossi; vitelline fields confluent anterior to the preovarian testes

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596 THE JOURNAL OF PARASITOLOGY, VOL. 84, NO. 3, JUNE 1998

TABLE I. Measurements of Hapalotrema mistroides (Monticelli, 1896) from the loggerhead turtle, Caretta caretta.

Lectotype Type series

Looss (1899) Looss (1899) Current (liver)

Locale Mediterranean Sea Mediterranean Sea Shark Bay,

Abukir, Egypt Abukir, Egypt Western Australia
(n = 8) (n = 5)

Character

Length 7,125 3.47-7.13 mm (5.13) 2.55-4.33 mm (3.84)

Width 1,400 1.05-1.25 mm (1.20) 0.66-1.02 mm (0.89)
Oral sucker: length 225 130-225 (193) 195-235 (212)
Width 297 225-346 (273) 183-250 (229)
Acetabulum: length 346 277-396 (343)
Width 416 287-416 (358) 267-327 (307)
Ovary: length 703 455-713 (570) 317-535 (448)
Width 614 495-693 (620) 376-713 (537)

Testis no.

Anterior group 8 8-9 8-9

Posterior group 11 10-12 10-15

Seminal vesicle: length 535 257-564 (447) 277-465 (383) [3]*

Width 240 88-240 (186) 163-243 (204) [3]
Cirrus sac: length 346 288-366 (324) 225-270 (257) [3]
Width 143 85-158 (129) 123-133 (127) [3]
Vitellaria to:

Cecal bifurcation 356 188-356 (276) 18-178 (117)

Posterior end 248 18-248 (94) 13-50 (38)

Anterior to cecal bifurcation 911 663-1,060 (873) 634-851 (770)

Genital pore to posterior end 4.15 mm 1.5-4.15 mm (2.68) 1.28-2.1 mm (1.79)
Genital pore/TL (%) 58.2 42.0-60.3 (51.2) 41.4-50.2 (46.8)
Forebody/TL (%) 18.3 18.3-36.7 (26) 24.9-37.6 (30)

* Numbers in brackets represent the sample size for that character.

in H. mistroides but not in H. loossi; differences in the sucker ratios H. loossi Price, 1934 and the remaining specimens listed in the Mate-
and the relative distances between the suckers in the 2 species. Neither rials and Methods from SMNH as paralectotypes. Considering the
of the original authors provided information on the range of variation strong similarities in the original descriptions of these taxa, the fact that
found in their specimens. both were taken from the same location in the same host species, and
Reexamination of Looss's specimens revealed 10-12 postovarian tes- that both turtles were taken in the Mediterranean Sea, we feel H. mis-
tes (Table I). Specimens of this morphotype from the type host collectedtroides should be considered a senior synonym of H. loossi, and H.
at Shark Bay, Western Australia, have 10-15 postovarian testes. Half mistroides becomes the recognized name for this species. The interested
of Looss' specimens show the anterior vitelline fields confluent anterior reader is directed to Platt and Blair (1996) for the nomenclatural details
to the preovarian testes, and the remainder probably would have, had and Anonymous (1997) for the ICZN ruling.
the specimens not been so mercilessly flattened.
Monticelli's (1896) figures 2 and 3 present distinctly different con-
DESCRIPTION
cepts of relative sucker size and sucker position. Figure 2 of Monticelli
(1896) shows the acetabulum larger than the oral sucker and the dis- Hapalotrema synorchis Luhman, 1935
tance between the suckers consistent with Looss' material. Monticelli's (Fig. 3; Table II)
(1896) figure 3 appears to be a caricature with the posterior portion of
Synonym: Hapalotrema orientalis Takeuti, 1942.
the worm greatly enlarged in order to present adequately the internal
organs, and the acetabulum and oral sucker are shown nearly equal in Diagnosis: Body robust, maximum width attained in anterior testic-
size. These discrepancies are curious and difficult to explain. There is ular field. Small tegumental spines anteriorly, diminishing posteriorly,
a great deal of overlap in meristic values between Looss' specimens absent posterior to acetabulum. Oral sucker subterminal, ovoid. Esoph-
and material collected from C. caretta in Shark Bay (Table I). Looss' agus sinuous, surrounded by glandular cells at cecal bifurcation. Intes-
specimens, as mentioned previously, appear to have been flattened se-tinal ceca bifurcate anterior to acetabulum, terminate near posterior end
verely during preparation. The range of measurements provided for H.of body. Acetabulum slightly pedunculate, ovoid, with minute spines
mistroides from the current study are somewhat smaller as they were on margin. Testes in preovarian, postovarian groups; individual testes
killed and fixed without mechanical flattening. difficult to differentiate. External seminal vesicle transversely elongate,
We petitioned the International Commission of Zoological Nomen- between preovarian testes and ovary. Cirrus sac well developed; cirrus
clature to exercise its plenary power to recognize H. loossi Price, 1934 spined, opening into a common genital atrium. Genital pore submedian,
as the type species of the genus (Platt and Blair, 1996; Case 2932) based sinistral; surrounded by sucker-like musculature without delimiting
on Art. 70b (misidentification of type species). Whereas Looss (1899) membrane forming true sucker. Ovary large, slightly lobed, between
did not describe a new species when erecting Hapalotrema, and his testicular groups. Oviductal seminal receptacle present, small; Laurer's
designation of H. constrictum was based on misidentification, no type canal not observed. Mehlis' gland not seen. Uterus small, passes anter-
specimen is currently recognized for this taxon. We, therefore, desig- iad and medially to enter genital atrium. Vitellarium granular, filling
nated specimen 2663 i 12 (left) (Alexandria vii.01) as the lectotype for extracecal space from posterior acetabular margin to cecal end, contin-

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 PLATT AND BLAIR-REDESCRIPTION OF HAPALOTREMA SPP. 597

4 5

:...: ..... ,

FIGURE 4-5. 4. Ha
cirrus sac; e., egg
posterior group; v.
(scale bar = 250 pJ
ovary; t., anteriorm

uous across the vitellaria began at the interce

posterior edge of the acetabulum, rather than
cle Y-shaped. the anterior margin. They reported 20-34 testes anterior and 12-29
testes posterior to the ovary (Fischthal and Acholonu, 1976); however,
Taxonomic summary we could not determine the number of testes in their material. In our
material the anterior and posterior testicular masses are highly lobed
Type host: Caretta caretta, loggerhead turtle. and show evidence that they may represent the overlapping of numer-
Other hosts: Eretmochelys imbricata, C. mydas. ous, individual testes. Measurements of 10 individuals (Tables II, III)
Type locality: Tortugas, Florida. overlap the measurements taken from Luhman's type specimen.
Other localities: Caribbean Sea near Cabo Rojo, Puerto Rico (Fisch- Fischthal and Acholonu (1976) also noted that the genital sucker did
thal and Acholonu, 1976); Gulf of Mexico near Tampico, Mexico (Cab- not possess an "outer, limiting membrane"; a condition we can confirm
allero y C., 1962). Current study (all from C. caretta): Indian Ocean, from the type material, vouchers, and our specimens. Also, the genital
Shark Bay, Western Australia, Australia (QM G214174-5); Pacific sucker may be faint and difficult to see in some individuals. They also
Ocean; Heron Island, Queensland, Australia (QM G214176-7), and Mon reported a "definite sucker surrounds the opening of Laurer's canal."
Repos, Queensland (QM G214178). The presence of the same structure in our material suggests that, al-
though we were unable to observe Laurer's canal directly, it is present.
Remarks
Takeuti (1942) described Hapalotrema orientalis from Eretmochelys
The original description of this species (Luhman, 1935), based japonica
on a (=E. imbricata) on the basis of 5 specimens (3 whole mounts,
single specimen from C. caretta, includes measurements of length 2 sectioned)
(6 and differentiated it from H. synorchis based on the ab-
mm), maximum width (1.5 mm), and diameters of the oral sucker sence of a genital sucker and failure of the anterior vitellaria to unite
(383)
and acetabulum (583). The testes were described as "numerous, in two in the median field anterior to the preovarian testes in H. orientalis,
large, compactly massed groups, one anterior, the other posterior to the whereas this does occur in H. synorchis. The medial union of the an-
ovary." Individual testes could not be distinguished, and hence counted, terior vitellaria appears to be variable, dependent on the degree of flat-
in the type specimen. The specimen appears to have been flattened tening of the specimens, and not suitable for differentiating species of
firmly. The diagnostic feature was the presence of a sucker surrounding Hapalotrema. The genital sucker is often difficult to discern, perhaps
the common genital pore. more so when subjected to mechanical flattening, as Takeuti (1942)
Fischthal and Acholonu (1976) reported H. synorchis from E. imbri- indicated his had been. The few measurements provided in the original
cata (hawksbill turtle); the specimens were smaller than the type, and description (length X width: 8.2 mm x 2.08 mm, 8.45 mm X 1.85 mm,

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598 THE JOURNAL OF PARASITOLOGY, VOL. 84, NO. 3, JUNE 1998

TABLE II. Measurements of Hapalotrema synorchis Luhman, 1935 from the loggerhead turtle, Caretta caretta.

Luhman (1935) type Current (heart)

Character (n = 1) (n = 10)

Tortugas, Florida Shark Bay, Western Australia;

Heron Isl. and Mon Repos, Queensland
Length 5.68 mm* 5.25-9.18 mm (6.95 mm)
Width 1.33 mm 0.95-2.13 mm (1.6 mm)
Oral sucker: length 317 277-445 (376)
Width 346 330-445 (416)
Acetabulum: length 545 510-790 (660) [8]t
Width 584 510-790 (661)
Ovary: length 378 340-802 (586)
Width 663 550-1,010 (739)
Seminal vesicle: length 703 440-1,025 (719) [8]
Width 213 203-310 (257) [8]
Cirrus sac: length 225 235
Width 53 55

Vitellaria to:

Cecal bifurcation 0 1.55-2.39 mm (1.78 mm)

Posterior end 110 20-297 (103)

Anterior to cecal bifurcation 1.25-1.83 mm (1.49 mm)

Genital pore to posterior end 1.73 mm 1.43-3.80 mm (2.19 mm)
Genital pore/TL (%) 69.5* 58-73.2 (68.9)
Forebody/TL (%) 24.3-36.4 (31.2)

* Due to the condition of the specimen, this is an estimate.

t Numbers in brackets represent the sample size for that character.

and 9 mm X 2.1 mm; oral sucker and acetabulum 0.46 mm and 0.62
ovary slightly left of the midline of the body. The genital open-
mm in diameter, respectively; Takeuti [1942]) fall within the range of
ing shows muscular striations with a limiting membrane (Cribb
our specimens from C. caretta (Table II). The remaining features, sub-
median genital pore, genital pore approximately 30% of total length and Gordon, 1998). This is distinct from the genital musculature
from the posterior end of the body, and relative size of the oral sucker seen in H. synorchis. Laurer's canal arises from the oviduct near
and acetabulum, also fall within the range of H. synorchis. We consider the midline of the body, prior to the junction of the oviduct and
H. orientalis to be a junior synonym of H. synorchis.
vitelline duct, courses almost directly dorsad and is surrounded
by a raised muscular structure (see fig. 7 of Dailey et al.
DISCUSSION
[1993]). The meristic comparison of the 2 forms provided by
Cribb and Gordon (1998) presented new information on H. Cribb and Gordon (1998) reveals no significant differences. We
mehrai Rao, 1976 and considered H. dorsopora Dailey, Fast agree, therefore, that H. dorsopora should be regarded as a
and Balazs, 1993 a junior synonym. Dailey et al. (1993) de- junior synonym of H. mehrai.
scribed H. dorsopora from green turtles off the coast of Hawaii.The seminal receptacle is reported as either absent (Rao,
The new species was considered most similar to H. mehrai but 1976) or a separate saccate (Dailey et al., 1993) or canalicular
differed in its larger size, different disposition of the anterior (Cribb and Gordon, 1998) structure. We interpret the seminal
testes in the preovarian groups, anterior extent of vitellarium, receptacle to be an expansion of the oviduct from our exami-
and most importantly, the presence of separate male and female nation of the types of H. dorsopora, voucher material, and spec-
genital pores, with the female pore opening on the mid-dorsal imens in the junior author's collection. The same condition is
surface between the vitelline reservoir and the anteriormost tes- found in the 2 species of Hapalotrema redescribed above.
tes of the postovarian group. The female genital pore was sur- Mehrotra (1973) first proposed H. pambanensis as a new spe-
rounded by a raised sucker-like structure (Dailey et al., 1993). cies in an abstract that did not meet conditions for availability
Reexamination of the type material of H. dorsopora reveals a (Art. 13(a)(i); Anonymous, 1985) and is regarded as a nomen
typical arrangement of the reproductive system with the open- nudum. Gupta and Mehrotra (1981) subsequently described H.
ing of Laurer's canal misidentified as the female genital pore. pambanensis from the green turtle, hence the correct name and
We were unable to trace the oviduct from the ovary to the point authorship is H. pambanensis Gupta and Mehrotra, 1981, but
where it expands to form an oviductal seminal receptacle. From they failed to compare this species to H. mehrai. Specimens of
there the oviduct courses sinistrally and is joined by the duct both were collected from the same location in the same host
from the vitelline reservoir. The uterus turns sharply anteriad species from the same locale. Meristic comparison with H. meh-
and there is a partially collapsed egg in the uterus just under- rai (see Cribb and Gordon [1998]) revealed overlap in every
neath the junction of the left vitelline duct and the transverse character, and we agree that H. pambanensis should be consid-
vitelline duct. The uterus continues on to open, with the cirrus ered a junior synonym of H. mehrai.
sac, in the genital atrium located at the posterior edge of the Rao (1976) described H. postorchis as a new species in green

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 PLATT AND BLAIR-REDESCRIPTION OF HAPALOTREMA SPP. 599

TABLE III. Reports of Hapalotrema spp. (Spirochidae) from marine turtles (Cheloniidae) from the coasts of Queensland (Qld.), Wester
(WA), and the Torres Straits, 1976-1993.

No. of
Host Parasite species Coll. no. Locality Year Location slides Accession no.*

Chelonia mydas Hapalotrema postorchis 1143 Badu Isl., Torres Straitt 1979 Artery near heart 1 QM G214162
1144 Badu Isl., Torres Strait 1979 Artery near heart 1 QM G214163
1145 Badu Isl., Torres Strait 1979 Artery near heart 1 QM G214164
1887 Townsville Harbor, Qld. 1993 Heart/liver 2 QM G214165
1890 Townsville Harbor, Qld. 1992 Various 10 QM G214166
Townsville Harbor, Qld.
Hapalotrema mehrai 1138 Magnetic. Isl., Qld. 1978 Heart 8 QM G214167
DB 1138
1139t Magnetic Isl., Qld. 1979 Heart 0 DB 1139
1140 Townsville Harbor, Qld. 1978 Heart 4 QM G214168
1141 Dalrymple Island, Qld. 1979 Heart 3 QM G214169
Dalrymple Island, Qld. DB 1141
1142 Townsville Harbor, Qld. 1979 Heart 1 QM G214170
DB 1142
1890 Townsville Harbor, Qld. 1992 Various 9 QM G214171
Hapalotrema sp. 1873 Townsville Harbor, Qld. 1992 Liver wash 3 QM G214172
1886 Townsville Harbor, Qld. 1992 Liver/lung wash 2 QM G214173
Caretta caretta Hapalotrema synorchis 1146t Shark Bay, WA 1979 Heart 0 DB 1146
1148 Shark Bay, WA 1979 Heart 2 QM G214174
DB 1148

1150 Shark Bay, WA 1979 Heart 1 QM G214175

DB 1150

1151:t Shark Bay, WA 1979 Heart 0 DB 1151

1152 Heron Isl., Qld. 1979 Heart 3 QM G214176
1153 Heron Isl., Qld. 1979 Blood clot in bile duct 1 QM G214177
1154 Mon Repos, Qld. 1976 Heart 3 QM G214178
Hapalotrema mistroides 1148 Shark Bay, WA 1979 Heart 1 QM G214179
DB 1148

1149 Shark Bay, WA 1979 Liver wash 3 QM G214180

1150 Shark Bay, WA 1979 Heart/liver 4 QM G214181
DB 1150

Eretmochelys imbricata Hapalotrema mehrai? 1156 Magnetic Isl., Qld. 1978 Heart 1 QM G214182
1157 Magnetic Isl., Qld. 1978 Heart 4 QM G214183

* QM = Queensland Museum; DB = private collection of David Blair.

t All reports represent new locality records.
: Tentative identification based on unmounted specimens.
? New host record.

turtles from the Gulf of Manar. While Rao (1976) reported find- We recognize 4 valid species of Hapalotrema Looss, 1899
ing numerous specimens of this fluke, he found only a single,that can be differentiated using the following key.
intact specimen to provide measurements for the description.
Dailey et al. (1993) provided a thorough redescription of H. Key to the species of Hapalotrema Looss, 1899
postorchis from Hawaii based on a wider range of material. Wela. Testes in anterior and posterior groups forming a compact mass,
have found this species in green turtles from various locations individual testes difficult to discern; genital sucker lacking
in the Coral Sea. limiting membrane ........... H. synorchis Luhman, 1935
An abbreviated list of marine turtles examined and Hapalo- (syn. H. orientalis)
lb. Testes discrete, not forming a compact mass ............. 2
trema spp. collected by the junior author during the past 22a. de-Total testes number >50 ............. H. mehrai Rao, 1976
cades is provided, along with information on the deposition of (syns. H. dorsopora, H. pambanensis)
voucher specimens in the Queensland Museum (Table III). Due 2b. Total testes number <30 ........................... 3

to the opportunistic nature surrounding the collection of much
of the material, meaningful estimates of prevalence and inten-
sity are not possible. All reports represent new locality records,
and H. mehrai from the hawksbill turtle (E. imbricata) is a new
host record. We feel that the information and specimens may
prove useful to other workers. Additional information on the
hosts and localities may be obtained from either of the authors T.R.P. was supported by a Lilly Open Fellowship during a
of the current work.
3a. Genital pore located in anterior 1/2 of body ...... H. mistroides
(Monticelli, 1896) (syn. H. loossi)
3b. Genital pore located in posterior /2 of body; genital sucker pres-
ent, with limiting membrane ...... H. postorchis Rao, 1976
ACKNOWLEDGMENTS
sabbatical leave and thanks T Cribb (University of Queensland)

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600 THE JOURNAL OF PARASITOLOGY, VOL. 84, NO. 3, JUNE 1998

for a stimulating discussion of spirorchid anatomy. D.B. was
supported by internal grants from James Cook University. J.
Glazebrook and R. Lester assisted in the collection of blood
flukes, and C. Limpus provided specimens from turtles exam-
ined as part of other studies.
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