J. Parasitol., 91(6). 2005. pp.

1379-1384
() American Society or Parasitologists 2005

CULUWIY A CICHLIDORUM N. SP. (DIGENEA: HAPLOPORIDAE) FROM THE BLACK-BEL T

CICHLlD VIEJA MACULICAUDA (PISCES: CICHLlDAE) FROM NICARAGUA
M. Leopoldina Aguirre-Macedo and Tom Scholz'
Laboratorio de Parasitologa, Centro de Investigacin y Estudios Avanzados del Instituto Politcnico Nacional (CINVESTAV-IPN),Unidad
Mrida, Carretera Antigua a Progreso Km 6, A.p. 73 "Cordemex", C,P. 97310. Mrida, Yucatn, Mexico. e-mail: leo@mda.cinvestav.mx
ABSTRACT:

A new haploporid trematode, Culuwiya cichlidorum n. sp., is described from the intestine of the black-belt cichlid

Vieja maculicauda (=Cichlasoma maculicauda (Regan)) from the Atlantic coast of Nicaragua. It differs from congeneric species
from freshwater and estuarine mugi1id, cichlid, and gobiid fishes, previously placed to Saccocoelioides Szidat, 1954, and Carassotrema Park, 1938, in possessing the intestinal ceca reaching up to the posterior half of the testis and a bipartite, swollen
extemal seminal vesic1e. The new species also is characterized by a small, pyriform body, a well-developed prepharynx, a large
pharynx, extensive vitelline follic1es, confluent posterior to the testis that is situated near the posterior extremity, a uterus limited
to the pretesticular and postacetabular region, and few, but relatively large. eggs (67-81 X 36-47 /Lm, Le., about 12-14% of the
body length) and miracidia without eye-spots. Culuwiya cichlidorum is also reported from Tomocichla tuba (Meek) in Nicaragua
and species of Cichlasoma, Herichthys, Oreochromis, Parachromis, Petenia, Theraps, and Vieja from Mexico.

Aguirre-Macedo et al. (2001a) reported 3 species of haploporid trematodes of SaccocoeUoides Szidat, 1954, from freshwater tishes in the drainages of the Autonomous Region of the
South Atlantic (RAAS) of Nicaragua, namely SaccocoeUoides
sogandaresi Lumsden, 1963, from the sail-tin moIly Poecilia
velifera (Regan, 1914), SaccocoeUoides sp. 1 from the blackbelt cichlid Cichlasoma macuUcauda (Regan, 1805) (= Vieja
macuUcauda (Regan, 1905-see Froese and Pauly, 2004), and
SaccocoeUodes sp. 2 from the banded astyanax Astyanax fasciatus (Cuvier, 1819). A subsequent study has revealed that
specimens designated as SaccocoeUoides sp. 1 represent a new,
hitherto unknown species of the newly erected genus, Culuwiya
Overstreet and Curran, 2005, that is described in this artic1e.
MATERIALS AND METHODS
Adult trematodes found in the intestine of Vieja maculicauda, designated as Saccocoelioides sp. I (referred by mistake as Saccocoelioides
sp. 2 in Table 2) by Aguirre-Maeedo et a\. (2001a), were fixed with hot
4% formaldehyde solution, stained with Mayer's carmine, and mounted
in Canada balsam as permanent preparations. A few specimens were
flattened and fixed with GAP (glycerin: ammonium picrate) according
to the method of Malmberg (1957) and Ergens (1969) used for monogeneans. In total, 32 specimens from V. maculicauda from the Atlantic
coast of Nicaragua were studied (8 specimens from the Torsuani River
and 24 from Caio Marain in Bluefields, both RAAS).
ln addition, trematodes collected by the present authors and their
coworkers (Vidal-Martnez et a\., 2001) in the intestine of Tomocichla
tuba (Meek, 1912) from the Atlantic coast of Nicaragua (Torsuani River) (Coleccin helmintologica de CINVESTA Merida, CHCM No. 468),
Theraps coeruleus Stawikowski and Werner, 1987, from Chiapas
(CHCM No. 20), Vieja synspilum (Hubbs, 1935) from Campeehe
(CHCM No. 104 and 492), and Oreochromis niloticus (Linnaeus, 1758)
from Yueatn (CHCM No. 491), all Mexico, were studied for eomparison. The amount and quality of specimens from these other fish hosts
were not suffieient to provide biometrieal data; different methods of
fixation have also eaused that these speeimens should not be used in
standard deseriptions of the new taxa. Therefore, only speeimens fixed
with hot formalin that are neither eontraeted or unnaturally flattened
were measured. The names of fishes follow.those in Froese and Pauly
(2004). Observations and drawings were made using an Olympus mieroseope with Nomarski interferenee eontrast. Measurements are given
in mierometers.

Reeeived 22 Oetober 2004; revised II April 2005; aeeepted II April

2005.
* To whom eorrespondence should be addressed.

DESCRIPTION
Culuw/ya c/chlldorum n. sp.
(Figs. 1-3)
Synonyms: Saccocoelioides sp. of Guajardo-Martnez (1984); Saccocoelioides nanii Szidat, 1954 of Andrade-Salas (1987); Saccocoelioides sp. of Seholz et a\. (1995); Saccocoelioides beauforti (Hunter and
Thomas, 1961) of Salgado-Maldonado et a\. (1997); Saccocoelioides
sp. 1 of Aguirre-Maeedo et a\. (200Ia); Saccocoelioides sp. of VidalMartnez et a\. (2001).
Diagnosis: Deseription based on 15 hot formalin-fixed, unflattened
specimens; measurements of the holotype in parentheses. Body pyriform (fusiform in immature speeimens), with maximum width between
seeond and last thirds of body, 470-720 (720) long and 150-335 (290)
wide. Tegument entirely spined; tegumental spines small, pointed. Two
pairs of remnants of eye-spots at prepharyngeal and pharyngeal level;
some granules dispersed between oral opening and genital pore. Unieellular gland eells situated dorsally, between pharynx and aeetabulum;
gland duets narrow, opening near anterior extremity.
Oral sueker subterminal, spherieal, 67-105 (100) long and 75-125
(120) wide. Aeetabulum spherieal, pre-equatorial, 57-120 (112) long
and 50-125 (115) wide, slightly larger or of same size as oral sueker;
sueker ratio 1: 1.00-1.17. Prepharynx present, 25-72 (25) long; pharynx oval, strongly muscular, large, situated obliquely in lateral view,
only slightly shorter than oral sueker length, 50-82 (75) long and 4775 (68) wide; esophagus long, extending to posterior half of aeetabulum, 100-137 (125) long. Ceea saceate, thiek-walled, terminating at half
to two thirds of testis length.
Testis oval to spherical, in posterior third of body, 72-175 (175) long
and 58-145 (143) wide. Extemal seminal vesic1e saceular, bipartite,
with proximal (posterior) part swollen, entering dorsally hermaphroditie
sae. Hermaphroditie sae oval to c1ub shaped, obliquely arehed around
ventral sueker dorsally, 77-162 (150) long and 62-135 (120) wide.
Internal seminal vesic1e oval to elongate, filling posterior third to half
of hermaphroditie sae length, 36-105 (85) long and 20-62 (50) wide;
ejaeulatory duct"short, entering long hermaphroditic duet; prostatie eells
in peripheral region of hermaphroditic sae. Genital pore preaeetabular,
prominent, thick walled.
Ovary spherieal, smooth, slightly submedian, postequatorial, just pretesticular, 37-87 (63) long and 35-87 (65) wide. Seminal reeeptac1e
absent; Laurer's eanal present. Uterus pretesticular and postaeetabular,
entering in its distal part to hermaphroditie sae; metraterm thick walled,
opening to hermaphroditie duet. Vitelline follic1es large, usually longitudinally elongate, distributed from level of posterior half of aeetabulum
to posterior extremity, eonfluent in posttesticular region. Eggs opereulate, 67-81 (eggs eollapsed in holotype) long and 36-47 (-), without
eye-spotted miracidia; at maximum 15 (4) eggs in uterus. Exeretory
vesic1e Y shaped, bifureating at anterior level of testis, without exeretory eoneretions; exeretory pore terrninaJ.
Taxonomlc summary
Type host: Blaek-belt ciehlid Vieja maculicauda (Regan, 1905) (Perciformes: Ciehlidae).

1379

1380

THE JOURNAL OF PARASITOLOGY.VOL. 91. NO. 6. DECEMBER 2005

FIGURES1-3.

Culuwiya

cichlidorum

n. sp. from Vieja maculicauda,

Nicaragua.

1. Hermaphroditic

sac and esophagus; dorsal view oC paratype

(CNHE, 5298). 2. Dorsal view oCgravid specimen; paratype (CNHE, 5299). 3. Dorsal view oCnongravid specimen; paratype (CNHE, 5299).

AGUIRRE-MACEDO AND SCHOLZ-CULUWIYA

Other hosts (hosts and localities from which voucher specimens were
available to the present authors are marked with an asterisk): Cichlasoma urophthalmus (Gtinther, 1862), Herichthys minckleyi (Kornfield
and Taylor, 1983), Oreochromis aureus (Steindachner, 1864), O. mossambicus (Peters, 1852), *0. niloticus, Parachromis managuensis (Gtinther, 1867), Petenia splendida Gtinther, 1862,* Theraps coeruleus,*
Tomocichla tuba, * Vieja synspilum.
Site: lntestine.
Type locality: Torsuani River, Regin Autnoma del Atlntico del
Sur (RAAS), Nicaragua (1147'06"N, 8352'38"W) (see Aguirre-Macedo et aJ., 200lb for map of localities in Nicaragua).
Other localities: Mexico: Campeche: *Rancho II, *Santa Gertrudis;
Chiapas: *Ro Cedros; Coahuila: Cuatro Cinegas; Colima: Laguna
Amela; Quintana Roo: *Cenote Azul (Bacalar); Tabasco: EI Espino, EI
Guanal, Las Ilusiones, Santa Anita, Thcta; Nicaragua: *Bluefields (Caio
Marain), RAAS (Guajardo-Martnez,
1984; Andrade-Salas, 1987;
Prez-Ponce de Len et aJ., 1996; Salgado-Maldonado et aJ., 1997;
Aguirre-Macedo et aJ., 2001a; Vidal-Martnez et a!., 2001).
Deposition oj types: Holotype and 5 paratypes in the Coleccin Nacional de Helmintos, lnstituto de Biologa, Universidad Nacional Autnoma de Mxico, Mexico City, Mexico (CNHE Nos. 5298 and 5298),
3 paratypes in the U.S. National Parasite Collection (USNPC Nos.
095844.00 and 095845.00), Beltsvi11e, Maryland, USA; 3 paratypes in
The Natural History Museum, London, U.K. (BMNH Nos. 2005.3.30.13), 4 paratypes at the lnstitute of Parasitology, AS CR, Cesk Budejovice, Czech Republic (IPCAS 0-511), and 3 paratypes at the Laboratory
of Parasitology CINVESTAV-IPN Unidad Mrida, Mexico (CHCM No.
490).
Etymology: The species name refers to the family name of the fish
host.
Remarks: The specimens from Nicaragua are placed to Culuwiya, a
newly proposed genus of the haploporid subfamily Waretrematinae Srivastava, 1937 sensu Overstreet and Curran (2005), on the basis of the
following characteristics: body pyriform, less or equal to I mm in
length; prepharynx present (vs. absent or very short in species of Saccocoelioides sensu stricto that now belongs to the Chalcinotrematinae
Overstreet and Curran, 2005-see Overstreet and Curran, 2005); esophagus long; ceca saccate (whereas often longer, cylindrical caeca in Saccocoelioides spp.); testis near posterior extremity but separated by vitelline follic1es elongated in longitudinal axis (vs. testis often far anterior from posterior end and posttesticular vitelline follic1es lacking in
several species now retained in Saccocoelioides); external seminal vesicle bipartite with swollen posterior (proximal) part; uterus confined
longitudinally between hermaphroditic sac and ovary (vs. preacetabular
and post-testicular in Saccocoelioides s. str.); eggs (few in number) relatively large; miracidia in uterine eggs without eye-spots (vs. eye-spotted in most Saccocoelioides spp.).
The genus Culuwiya, as proposed by Overstreet and Curran (2005),
inc1udes 3 species from Mugil cephalus L. (Mugilidae), previously
placed in Saccocoelioides, namely Culuwiya beauJorti (Hunter and
Thomas, 1961) (type species) in southeastern United States, inc1uding
the Gulf of Mexico, Culuwiya overstreeti (Bargiela, 1988), and Culuwiya papernai (Bargiela, 1988), both from Chile, Culuwiya agonostomus (Dyer, Bunckley-Wi11iams, and Williams, 1999), described also as
a member of Saccocoelioides from the freshwater mullet Agonostomus
monticola (Bancroft) and the goby Sicydium plumieri (Bloch) in Puerto
Rico, and Culuwiya tilapiae (Nasir and Gmez, 1976) (syn. Carassotrema tilapiae Nasir and Gmez, 1976) from the cichlid Tilapia mossambicus (= Oreochromis mossambicus) in Venezuela.
Culuwiya cichlidorum differs from all of these taxa in possessing
longer intestinal ceca that reach to the midlength of the testis or up to
its posterior half (vs. pretesticular or ending near the anterior margin of
the testis in other species of Culuwiya), and a bipartite, proximally
swollen external seminal vesic1e (vs. saccular, spherical, or c1ub shaped
in other taxa).
The new species also can be distinguished from the type species, C.
beauJorti, by the absence of prominent excretions in the excretory bladder (I or 2 prominent excretions always present in C. beauJorti-Hunter
and Thomas, 1961; Overstreet, 1971), a larger pharynx (much shorter
than the length of the oral sucker in C. beauJorti), and more numerous
and extensive posttesticular vitelline follic1es (vs. fewer follic1es limited
in their distribution to a smaller posttesticular space-Hunter
and
Thomas, 1961; Overstreet, 1971).

CICHLlDORUM N. SP. FROM CICHLlDS

1381

Culuwiya cichlidorum differs from C. tilapiae in possessing a round,

entire ovary instead of a triangle-shaped ovary in C. tilapiae, the absence of a receptaculum seminis (present in C. tilapiae), relative size
of the pharynx (much smaller in C. tilapiae), and the number of eggs
(at maximum 4 in C. tilapiae) (Nasir and Gmz, 1976).
The new species can be distinguished from C. agonostomus by the
presence of a prepharynx (absent in C. agonostomus) and the shape of
an external seminal vesic1e (saccular in the latter species) (see Dyer et
aJ., 1999).
The only haploporid species reported from cichlids in the Neotropical
Region is Saccocoelioides carolae Lunaschi, 1984, described from
Cichlasoma Jacetum (Jenyns) from the Buenos Aires Province, Argentina by Lunaschi (1984) and redescribed by Martorelli (1986). The new
species differs markedly from S. carolae in the postacetabular uterus
(preacetabular in S. carolae), the extent of intestinal ceca (pretesticular
in S. carolae), the position of the testis (just near the posterior extremity
in S. carolae), the shape of the body (elongate in S. carolae), length of
the preparynx (absent or very short in S. carolae), and relative size of
the pharynx (much smaller than the length of the oral sucker in the
latter taxon) (see Lunaschi, 1984; Martorelli, 1986).
Saccocoelioides sogandaresi Lumsden, 1964, was described from the
sail-fin molly Mollienisia latipinna Lesueur (now Poecilia latipinna) in
Texas and then reported from guppies (Poeciliidae) in Mexico and Nicaragua (Scholz et al., 1995; Aguirre-Macedo et a!., 2001a), but also
from species of other fish families (Characidae, Gobiidae, Goodeidae,
Mugilidae), inc1uding 2 cichlids (Nandopsis istlanum and Oreochromis
aureus) from Mexico (Salgado-Maldonado et aJ., 2004). However, records of S. sogandaresi from other fish than poeciliids should be revised
because they may inc1ude other species of haploporids, inc1uding species of Culuwiya.
Saccocoelioides sogandaresi, as a member of Saccocoelioides sensu
Overstreet and Curran (2005), differs from C. cichlidorum in the posttesticular uterus, more anterior position of the testis, a shorter prepharynx (only 1-22 II-m long in S. sogandaresi) and hermaphroditic duct,
the shape of the external seminal vesic1e, the presence of eye-spotted
miracidia within uterine eggs, and much shorter hermaphroditic sac
(Lumsden, 1963).
DISCUSSION

Haploporid trematodes are cosmopolitan parasites of the alimentary tract of fishes and are characterized by the presence
of a hermaphroditic sac (Overstreet and Curran, 2005). Controversial opinions have been presented as to the classification of
the family, and the validity of its genera and species also has
been questioned (e.g., Yamaguti, 1971; Nasir and Gmez, 1976;
Kohn, 1986; Daz and Gonzlez, 1990; Lunaschi, 1996,2002).
ln addition, limited information exists about life cycles of haploporids (Szidat, 1970, 1973; Martin, 1973; Martorelli, 1986;
Daz and Gonzlez, 1990; Ditrich et al., 1997).
Recently, the systematics of the Haploporidae has been revised by Overstreet and Curran (2005), who proposed a new
classification based on evaluation of type and voucher spec imens as well as critical evaluation of existing literature, and
erected 2 new haploporid genera. In this article, their conclusions are followed and the newly described species, previously
considered by the present authors (Aguirre-Macedo et al.,
2001a) to be a member of Saccocoelioides, is here considered
to be a species of Culuwiya, a genus established by Overstreet
and Curran (2005) to accommodate 5 other taxa from North
and South American freshwater and estuarine fish (Mugilidae,
Gobiidae, and Cichlidae), previously placed to Saccocoelioides
and Carassotrema.
Based on field experience, it is assumed that haploporid species of Culuwiya and Saccocoelioides from cichlid, poeciliid,
and characid fishes in Central America and Mexico may exhibit
a narrow host specificity (at least limited to 1 family of fish

>
CI)
'"

TABLEI. Comparative measurements and morphological characteristics of Culuwiya species most similar to C. cichlidoum and Saccocoelioides carolae from cichlids and S. sogandaresi
from Centra I and North America.
Species
C. cichlidorum n. sp.

Saccocoelioides sp.
(= C. cichlidorum)

C. beauforti (Hunter
and Thomas, 1961)

C. agonostomus
(Dyer et aJ., 1999)

S. carolae
Lunaschi, 1984

S. sogandaresi
Lumsden, 1963

Host
Distribution

Vieja maculicauda
Nicaragua, Mexico

Vieja synspilum
Mexico

Prochilodus lineatus
United States

Agnostomus monticola
Puerto Rico

Cichlasoma facetum
Argentina

Reference
Body shape
Body length
Body width
Oral sucker

Present study
Pyriform
470-680
150-335
67-105
75-125
57-120
50-125
1 : 1.0-1.2

Andrade-Salas (1987)
Pyriform
400-690
157-243
58-81
70-98
64-104
70-104
Acetetabulum slightly
larger
Short
43-74
49-80
1/3 of testis length

Overstreet (1971)
Fusiform
458-968
152-337
59-109
64-102
63-102
70-123
1 : 0.8-1.2

Dyer et aJ. (1999)

Pyriform
451-647
167-310
81-101
68-114
72-99
77-112
Oral sucker slightly
larger
Absent
68-77
68-80
Anterior margin of
testis
121-176
85-100
Saccular

Lunaschi (1984)
Elongate
548-825
208-349
92-132
80-132
92-138
96-140
1 : 1.0-1.2

Poecilia latipinna
United States, Mexico,
Nicaragua
Lumsden (1963)
Pyriform
341-512
165-307
55-87
60-87
62-105
75-103
1 : 1.2

Short
43-73
42-43
Posterior border of
ovary
116-170
63-97
Spherical to saccular

1-12
45-73
37-70
Anterior margin of
testis
45-98
50-84
Saccular

86-114
67-136
40-43
35-46
Pretesticular
66-77
35-46
Absent
Posttesticular

32-189
54-135
41-120
43-128
Posttesticular
72-123
41-68
Present
Pretesticular

75
112
41-62 in diameter

Acetabulum
Sucker ratio
Prepharynx
Pharynx
Extent of intestinal ceca
Hermaphroditic sac
External seminal vesic1e
Testis
Ovary
Uterus
Eggs
Eye-spotted miracidia
Vitellarium

36-72
50-82
47-75
1/2 to 2/3 of testis
length
77-162
62-135
Bipartite, swollen
72-175
58-145
37-87
35-85
Pretesticular
67-81
36-47
Absent
Posttesticular

86-150
75-93
Spherical
78-142
59-114
30-76
29-60
Pretesticular
68-89
42-53
Absent
Posttesticular

12-54
37-63
37-65
Anterior margin of
testis
72-182
54-138
Spherical to c1ub
shaped
105-266
61-170
37-84
30-72
Pretesticular
70-107
40-56
Absent
Posttesticular

Posttesticular
78-87
37-55
Present
Posttesticular

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AGUIRRE-MACEDO AND SCHOLZ-CULUWIYA

hosts). In Nicaragua, infections of cichlids with haploporid species other than C. cichlidorum were not recorded. Similarly,
sail-fin moIly (P. velifera) harbored only S. sogandaresi and the
banded astyanax (A. fasciatus) was infected with another, not
yet identified species of Saccocoelioides (designated as Saccocoelioides sp. 2 in Aguirre-Macedo et al., 2001a), but the
latter fish never hosted species typical of guppies and cichlids,
i.e., S. sogandaresi and the new species (c. cichlidorum), respectively (Aguirre-Macedo et al., 2001a). The same observation has been made during long-term studies on the helminth
parasites of freshwater fish in southeastern Mexico (Scholz et
aI., 1995; Salgado-Maldonado et al., 1997; Vidal-Martnez et
al., 2001). On the other hand, S. sogandaresi has been reported
from a number of fish of several families (Salgado-Maldonado
et al., 2004).
These records require verification because species identification of haploporid trematodes is very difficult (Overstreet and
Curran, 2005). It is caused, among others, by a low number of
diagnostic characteristics based on morphology, by difficulties
in observing internal structures due to the presence of dark
pigments scattered throughout the body, and eggs and vitelline
follicles overlapping internal organs, and, in particular, by inadequate quality of many voucher specimens. The necessity of
adequate fixation (with hot fixative, without pressure) and careful pcocessing of specimens has also been emphasized by Overstreet and Curran (2005).
ln Mexico, haploporid trematodes found in cichlids have
been considered to belong to Saccocoelioides and have been
reported under different names (see the list of synonyms
above). A comparative study of voucher specimens from several cichlids from Mexican states of Campeche, Chiapas, Quintana Roo, and Tabasco has shown their conspecificity with C.
cichlidorum because no morphological and biometrical differences were found (see Fig. of Saccocoelioides sp. in VidalMartnez et al., 2001). Therefore, the records of Saccocoelioides fcom other cichlids are also tentatively identified as C. cichlidorum, although their voucher specimens were not available
to the present authors (Table I).
It should be noted that Overstreet and Curran (2005) mentioned C. beauforti to be reported by Scholz et al. (1995) fcom
several cichlids in cenotes in Mexico. In fact, the latter authors
(and Vidal-Martnez et al., 2001, as well) reported haploporid
trematodes fcom Cichlasoma and Petenia in Mexico under the
name Saccocoelioides sp., not S. beauforti. Actually, haploporids now considered to be conspecific with C. cichlidorum were
listed as S. beauforti in a checklist of the helminth parasites of
freshwater fish and cichlids of Mexico, respectively (PrezPonce de Len, 1996; Salgado-Maldonado et al., 1997). This
implies that C. beauforti has never been found in other hosts
than mugilids, similar to C. overstreeti (Bargiela, 1988) and S.
papernai (Bargiela, 1988). Thus, the genus Culuwiya now includes 2 species parasitic in cichlids, namely C. tilapiae and C.
cichlidorum.
The data available indicate that C. cichlidorum is a relatively
frequent parasite of cichlid fish occurring throughout Mexico
and in Central America. Further studies, especially those performed in Central America, will show us how widely this trematode is distributed in the Neotropical region and how many
species of cichlid fish it actually infects.

CICHLlDORUM N. SP. FROM CICHLlDS

1383

ACKNOWLEDGMENTS
The authors express their gratitude to Victor M. Vidal-Martnez and
Gregory Arjona-Torres, Laboratory of Parasitology, CINVESTAV-IPN
Unidad Mrida; and Petr Posel, Svetlana Dumailo, and Eduardo Siu,
Bluefields Indian and Caribbean University, Bluefields, Nicaragua, for
help with dissecting fish in Nicaragua, and to Clara Vivas-Rodrguez,
CINVESTAV-IPN Unidad Mrida, for technical assistance. Thanks are
also due to Agustn Jimnez-Ruz, University of Nebraska, Lincoln,
Nebraska, for providing the literature on haploporid trematodes from
South America; La Lunaschi, Universidad de La Plata, Argentina, for
sending us her papers on Saccocoelioides; Robin M. Overstreet and
Steve S. Curran, University of Southern Mississippi, Ocean Springs,
Mississippi, for providing galley proofs of their chapter on the Haploporidae for the Keys to the Trematoda, Volume 2, and Rodney A. Bray,
The Natural History Museum, London; and Anindo Choudhury, St.
Norbett College, De Pere, Wisconsin, for valuable advice. This study
was partly supported by the Grant Agency of the Academy of Sciences
of the Czech Republic (project A6022404 to T.S.) and the Institute of
Parasitology, AS CR, Cesk Budejovice (project Z6 022 909). Visits of
T.S. to Mexico were made possible due to financial support of the Academy of Sciences of the Czech Republic and the National Research
Council of Mexico (CONACyT) (interacademic exchange program).

LlTERATURE CITED
AGUIRRE-MACEDO, M. L., T. SCHOLZ, D. GONZLEZ-SOLS, V. M. VIDALMARTNEZ, P. POSEL, G. ARJONA-ToRRES, E. SIU-EsTRADA, AND S.
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