Digenetic Trematodes of Indian Marine Fishes

Rokkam
Madhavi · Rodney A. Bray
Digenetic
Trematodes of
Indian Marine
Fishes
Digenetic Trematodes of Indian Marine Fishes
Rokkam Madhavi Rodney A. Bray
•
Digenetic Trematodes
of Indian Marine Fishes
123
Rokkam Madhavi Rodney A. Bray
Department of Zoology Life Sciences
Andhra University Natural History Museum
Visakhapatnam, Andhra Pradesh London
India UK
ISBN 978-94-024-1533-9 ISBN 978-94-024-1535-3 (eBook)

https://doi.org/10.1007/978-94-024-1535-3
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Preface
The present century is witnessing a sharp decline in taxonomy research and an acute
dearth of taxonomic expertise all over the world, and this has been the cause of
much concern to scientists. The reasons for this crisis are many, a major one being
the difficulties in getting access to the literature which is making taxonomic
research strenuous and unattractive to the younger workers. In particular, the tax-
onomy of digeneans of marine fishes has received lot of attention and India has
produced many eminent taxonomists, not to speak of the large number of publi-
cations. A glance at the literature reveals, however, that it is widely scattered and
some of it published in remote journals is not readily accessible to the taxonomists.
The introduction of molecular methods in taxonomy has given a slight boost to
taxonomic research but molecular taxonomy cannot be taken as a replacement to
traditional taxonomy, it can only be an adjunct to it. Application of both basic and
modern tools together in taxonomy makes it a strong and attractive field.
To revive the research interest in taxonomy, emphasis should be placed on the
publication of monographs, checklists and reviews and databases. Concerted efforts
of scientists from public and private institutions are needed to ignite the spirit of
taxonomy in the young scientists. Keeping these facts in view, we have brought out
this monograph on digenetic trematodes of Indian marine fishes. Almost all the
species of digeneans so far recorded, numbering over 600, are covered in this
monograph. For each species information on hosts, localities, geographic distri-
bution, references and a brief description accompanied by an illustration are fur-
nished. A host-parasite list is also presented. The pertinent areas for future research
are indicated. We are hoping that taxonomists working in this field will find this
manual to be a useful guide.
We have taken help directly or indirectly from a number of scientists in the
context of preparing this monograph. The publications of Dr. Gibson, Dr. Cribb,
Dr. Overstreet, Dr. Justine have proved to be extremely useful. We are thankful to
them for providing the literature whenever it is needed. Madhavi extends her
grateful thanks to a number of Indian scientists without whose help this monograph
would not have been completed. She is particularly grateful to Prof. K. C. Pandey
and Nirupama Agrawal who supplied most of the Indian literature and to the
v
vi Preface
director of Zoological Survey of India for providing the holotype slides and the
literature. She is thankful to her former research students Dr. Shameem,
Dr. Umadevi, Dr. Janaki Ram, Dr. Jayasree and Dr. Sailaja for their strong support
and continuous help in bringing the manual to the final stage. She remains for ever
grateful to her teacher and guide late Professor K. Hanumantha Rao who introdued
her to this field and offered valuble guidane and support. Finally, the financial
assistance provided by funding agencies like UGC, CSIR, MOEF and DST is
gratefully acknowledged.
The textbook includes quite a good number of figures taken from different
journals. In this context, the authors express their grateful thanks to the publishers:
Allen Press, Cambridge University Press, Springer and De Gruyter and the directors
of Zoological Survey of India, Pakistan Journal of Zoology and Indian Society of
Parasitology for permitting reproduction of figures published in their journals.
Visakhapatnam, India Rokkam Madhavi

London, UK Rodney A. Bray
Contents
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
2 Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3 The Digenetic Trematodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
General Organization and Life Cycles (Fig. 3.1a–e) . . . . . . . . . . . . . . 9
Life Cycles of Digenetic Trematodes . . . . . . . . . . . . . . . . . . . . . . . . . 12
Classification of Digenea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Part I Systematic Account—Order Diplostomida Olson,

Cribb, Tkach, Bray & Littlewood, 2003
4 Superfamily Schistosomatoidea Hassall, 1898 . . . . . . . . . . . . . . . . . 21
Family Aporocotylidae Odhner, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . 21
Part II Systematic Account—Order Plagiorchiida La Rue, 1957

5 Superfamily Bivesiculoidea Yamaguti, 1934 . . . . . . . . . . . . . . . . . . 29
Family Bivesiculidae Yamaguti, 1939 . . . . . . . . . . . . . . . . . . . . . . . . 29
6 Superfamily Bucephaloidea Poche, 1907 . . . . . . . . . . . . . . . . . . . . . 35
Family Bucephalidae Poche, 1907 . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
7 Superfamily Gymnophalloidea Odhner, 1905 . . . . . . . . . . . . . . . . . 83
Family Fellodistomidae Nicoll, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . 83
Family Tandanicolidae Johnston, 1927 . . . . . . . . . . . . . . . . . . . . . . . . 101
8 Superfamily Haplosplanchnoidea Poche, 1926 . . . . . . . . . . . . . . . . 109
9 Superfamily Hemiuroidea Looss, 1899 . . . . . ....... . . . . . . . . . . 119
Family Hemiuridae Looss, 1899 . . . . . . . . . . . ....... . . . . . . . . . . 119
Family Accacoeliidae Odhner, 1911 . . . . . . . . . ....... . . . . . . . . . . 182
Family Dictysarcidae Skrjabin & Guschanskaja, 1955 . . . . . . . . . . . . . 186
vii
viii Contents
Family Didymozoidae Monticelli, 1888 . . . . . . . . . . . . . . . . . . . . . . . 188

Family Hirudinellidae Dollfus, 1932 . . . . . . . . . . . . . . . . . . . . . . . . . 236
Family Lecithasteridae Odhner, 1905 . . . . . . . . . . . . . . . . . . . . . . . . . 238
Family Sclerodistomidae Odhner, 1927 . . . . . . . . . . . . . . . . . . . . . . . 256
10 Superfamily Apocreadioidea Skrjabin, 1942 . . . . . . . . . . . . . . . . . . 263
Family Apocreadiidae Skrjabin, 1942 . . . . . . . . . . . . . . . . . . . . . . . . . 263
11 Superfamily Transversotrematoidea Witenberg, 1944 . . . . . . . . . . . 273
Family Transversotrematidae Witenberg, 1944 . . . . . . . . . . . . . . . . . . 273
12 Superfamily Haploporoidea Nicoll, 1914 . . . . . . . . . . . . . . . . . . . . . 275
Family Haploporidae Nicoll, 1914 . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
Family Atractotrematidae Yamaguti, 1939 . . . . . . . . . . . . . . . . . . . . . 295
13 Superfamily Lepocreadioidea Odhner, 1905 . . . . . . . . . . . . . . . . . . 299
Family Lepocreadiidae Nicoll, 1935 . . . . . . . . . . . . . . . . . . . . . . . . . . 300
Family Aephnidiogenidae Yamaguti, 1934 . . . . . . . . . . . . . . . . . . . . . 353
Family Enenteridae Yamaguti, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . 357
Family Gyliauchenidae Fukui, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . 363
14 Superfamily Brachycladioidea Odhner, 1905 . . . . . . . . . . . . . . . . . 367
Family Acanthocolpidae Lühe, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . 367
15 Superfamily Opecoeloidea Ozaki, 1925 . . . . . . . . . . . . . . . . . . . . . . 405
Family Opecoelidae Ozaki, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 405
Family Opistholebetidae Nicoll, 1915 . . . . . . . . . . . . . . . . . . . . . . . . 486
16 Superfamily Monorchioidea Odhner, 1911 . . . . . . . . . . . . . . . . . . . 493
Family Monorchiidae Odhner 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . 493
17 Superfamily Gorgoderoidea Looss, 1899 . . . . . . . . . . . . . . . . . . . . . 551
Family Gorgoderidae Looss, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . 551
18 Superfamily Microphalloidea Ward, 1910 . . . . . . . . . . . . . . . . . . . . 567
Family Zoogonidae Odhner, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . 567
Family Faustulidae Poche 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 578
19 Superfamily Opisthorchioidea LaRue, 1957 . . . . . . . . . . . . . . . . . . 597
Family Cryptogonimidae Ward, 1917 . . . . . . . . . . . . . . . . . . . . . . . . . 597
20 Miscellaneous Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 613
Family Cladorchiidae Fischoeder, 1901 . . . . . . . . . . . . . . . . . . . . . . . 613
Family Microscaphidiidae Looss, 1900 . . . . . . . . . . . . . . . . . . . . . . . 616
Family Cephalogonimidae Looss, 1899 . . . . . . . . . . . . . . . . . . . . . . . 618
Family Opisthorchiidae Looss, 1899 . . . . . . . . . . . . . . . . . . . . . . . . . 622
21 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 623
Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 631
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 649
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 685
Chapter 1
Introduction
Marine fishes by virtue of their inherent characteristics such as large body size, varied
diet, high vagility and wide geographic distribution serve as hosts for a wide range
of parasites belonging to diverse groups and occupying different niches as ecto-
and endoparasites. The role of different species of marine fish as hosts also varies
depending on a multitude of ecological and host factors. Based on the overwhelming
number of parasite species recorded from marine fishes, it is assumed that the number
of parasites far exceeds the number of species of marine fish. The ubiquitous nature
of these parasites also indicates that they will have some commercial importance in
fisheries and aquaculture. On the positive side, they can be used as biological tags
for stock discrimination (Mackenzie, 2002) and they are also receiving importance
as indicators in environmental monitoring. On the negative side, many parasites
are severe pathogens of commercially important fishes and cause mass mortalities
especially under culture conditions.
A vast literature now exists on marine fish parasites dealing with a variety of
aspects including taxonomy, life cycles, ecology and host–parasite relationships.
The main focus in research has been on taxonomic aspects devoted to morphological
descriptions, identification up to species level and analysis of phylogenetic relation-
ships of different taxa of marine parasites. In recent years, taxonomic research has
made rapid advances because of the application of molecular methods.
So far as the Indian scenario is concerned, marine fish parasites have not received
due recognition. Studies are yet to be undertaken on a large scale. India has a vast
coastline of about 7520 km spread along the margins of the Bay of Bengal on the
east coast and the Arabian Sea on the west coast. These two coasts join at the Gulf of
Mannar, situated between India and Sri Lanka. Additionally, there are a few island
territories such as the Andaman and Nicobar archipelagoes in the Bay of Bengal
and Lakshadweep in the Arabian Sea (Fig. 1.1). There are 9 coastal states, 5 along
the west coast namely Mumbai, Gujarat, Karnataka, Kerala and Goa and 4 along
the east coast namely West Bengal, Orissa, Andhra Pradesh and Chennai (Madras
coast). The coastal waters support a rich and diverse fish fauna. It is estimated that
there are about 2564 species of marine fish inhabiting Indian coastal waters. Many
© Crown 2018 1
R. Madhavi and R. Bray, Digenetic Trematodes of Indian Marine Fishes,
https://doi.org/10.1007/978-94-024-1535-3_1
2 1 Introduction
Fig. 1.1 Map of India showing the various coastal areas
institutions and universities situated in coastal districts are undertaking investiga-

tions on various aspects of marine biology on a large scale. One shortcoming in
these studies is the lack of contributions to Marine Parasitology, although it is well
recognized that knowledge on this aspect is very important and should form an inte-
gral part of Marine Biological studies. It is well known that a wide range of parasites
belonging to Protozoa, Monogenea, Digenea, Cestoda, Nematoda, Acanthocephala,
Myxozoa and Crustacea infect marine fishes. Studies undertaken so far are confined
to morphological aspects. Knowledge of the life cycles, pathogenicity and ecology
is meagre. The present knowledge of marine parasites is at a basic level.
1 Introduction 3
Among the various groups of parasites infecting marine fishes, the digeneans have
received more attention, because of their common occurrence in marine fishes and the
great diversity exhibited by them in their morphology and the life cycle patterns. They
occur in a variety of sites, exhibiting rather variable, but often strict host specificity
for each species or higher group of fishes. Approximately 18,000 nominal species
are reported worldwide including well over 5,000 species from marine fishes alone
(Cribb, 2005b), constituting perhaps the largest group of metazoan parasites (Cribb
et al. 2001). A huge literature exists now on digenean parasites of marine fishes.
The volumes on “Keys to the Trematoda” edited by Gibson, Bray and Jones served
as a boon to scientists working on Digenean taxonomy. The contributions made by
scientists of yesteryears, e.g. Yamaguti, Manter, Dawes, and the present century, e.g.
Gibson, Bray, Cribb, Overstreet, are substantial.
Historical account: Research on Digenea of marine fishes in India has a long
history. Investigations were initiated in 1906 but the major work commenced in the
1930s. Early workers like Lühe (1906) and Southwell (1913) identified a few dige-
neans collected from marine fish of Gulf of Mannar. The first detailed study was that
of Srivastava (1936–1941) who published a series of papers on digeneans of marine
fishes from the east and west coasts of India and reported several new species. Another
major contribution was by Chauhan (1943, 1945) who reported digeneans belong-
ing to Bucephalidae and Hemiuridae and published monographs on these groups
(Chauhan, 1954a, b). After a gap of nearly 20 years, a comprehensive study of dige-
nean parasites of marine fishes was undertaken by Hafeezullah (Hafeezullah and
Siddiqi, 1970; Hafeezullah, 1971–1990; Hafeezullah and Dutta, 1998) who not only
reported many species of digeneans but also discussed the taxonomic relations of the
species and the validity of various species. Many surveys were undertaken especially
during the 1960s, 1970s and early 1980s. Scientists from Lucknow University such
as Ahmad, V. Gupta, P. C. Gupta, R. C. Gupta, S. P. Gupta, from Punjab University,
e.g. N. K. Gupta and Andhra University, e.g. Hussain et al., reported several species.
The senior author of this monograph (Rokkam Madhavi) carried out investigations
on Digenea of marine fishes of the Visakhapatnam coast of the Bay of Bengal from
1970 to 1982 and published a series of papers. A comprehensive study on didymo-
zoid trematodes of tunas of Visakhapatnam was undertaken and several new species
were reported (Murugesh and Madhavi 1990–1993). Investigations were undertaken
on metazoan parasites of Rastrelliger kanagurta, and helminth parasites of mullets
(Madhavi and Trivenilakshmi, 2011; Rekharani and Madhavi, 1985).
Recently, there has been a general decline in studies on this aspect and just a
few publications dealing with description of one or a few parasites have appeared
(Gupta, 2007; Govind, 1985; Job, 1961–1965; Karyakarte, 1968–1969; Karyakarte
and Yadav, 1976a, 1976b, 1977a, 1977b; Khan and Karyakarte, 1984, 1987a, 1987b;
Kumar and Agrawal, 1984; Bijukumar, 1997; Dutta and Manna, 1993, 1998). At
present, it has come to a standstill. Madhavi (2011) published a checklist on Digenea
of Indian marine fishes by compiling all the published information. All the digeneans
reported till then together with their hosts, localities and references were covered in
this checklist.
4 1 Introduction
A perusal of the literature reveals that many more species are yet to be described.
The studies undertaken so far were confined to fish from just a few regions such as
the Puri coast, Visakhapatnam and the Mumbai coast. Vast areas along the coast still
remain to be explored. The validity of many of the species reported has been ques-
tioned and many species whose identifications are based on a single specimen have
been placed in the species/taxon inquirendae category. In spite of these controversies,
it is surprising that molecular studies have not been undertaken and not even a single
species of digeneans of marine fish from this region has been subjected to molecular
study, although this is receiving global attention. Further, practically no informa-
tion is available on aspects such as host specificity, biodiversity, pathogenicity and
geographic distribution.
The present monograph is an attempt to consolidate all the species of Digenea so
far recorded from marine fishes of the Indian coasts of the Bay of Bengal and the
Arabian Sea. The primary aim in preparing the monograph is to bring together all the
information scattered in different journals, most of which is not readily accessible.
It covers over 600 species of digeneans. Most species are illustrated and described
briefly, information on the hosts, locality, geographic distribution, and the validity
status is presented. Keys and definitions are given for families, subfamilies, genera
and species. A host–parasite list is also presented. Attention is drawn to some poorly
known or controversial species, but they are not necessarily described or illustrated.
In the preparation of the account, testing the validity of each and every species
reported by referring the diverse body of literature has proved a herculean task. In
this context, the data presented in World Register of Marine species (WoRMS) has
proved to be extremely useful. These data are a tremendous resource for trematodes
studies, created largely through the efforts of Dr. D. I. Gibson. The reviews published
by Bray and Cribb on different families of digeneans were of great help.
We have attempted to cover all the valid species and provide means for their
identification including diagnostic features for families, subfamilies and genera and
keys for their separation. The “Keys to the Trematoda” volumes 1, 2 and 3 served
as useful guides for preparing this document. The diagnostic characters of families,
subfamilies and genera were derived from them. The classification suggested in these
accounts is adopted except where superseded by recent molecular studies.
One problem encountered relates to the date of publication of some papers. In
some cases, the journal was actually published later than the nominal date on the
front page of the journal part. The actual date of publication may be listed, often in
small print, on the last page or inside the cover. We have not been able to check this
in every case, so there may be discrepancies between the dates given here and those
quoted in other publications and online sources.
The purpose of the monograph is to serve as a guide to Indian scientists for the
identification of digeneans. We hope it will stimulate studies on the taxonomy of fish
parasites adopting modern taxonomic tools, as well as inspire workers to undertake
studies on life cycles, ecology and allied aspects.
Chapter 2
Materials and Methods
Digeneans infect almost every species of marine fish. According to the estimates
of the Zoological Survey of India, Fisheries Division there are approximately 1,400
species of marine fish in the Indian Ocean. It is reckoned that on average each species
of marine fish harbours one species of digenean (Poulin & Morand, 2000). From the
records, it is evident that only 50% of digeneans have so far been recorded and there is
vast scope for future studies. For proper identification of digeneans it is necessary to
see that uniform methods are followed for collection, fixation, staining and mounting
of the parasite and that the slide prepared is of a high standard permitting detailed
study of its organization on which depends the identification of the parasite.
Morphological analysis requires that the material is in good condition. For this
purpose, it is essential that only live and actively moving individuals are used for the
study. This can be achieved by following certain precautionary measures. The ideal
method for collecting trematodes is to examine the host while still alive or freshly
caught. Flukes that are dead and degenerating should be avoided. Proper fixation
of parasites is also of crucial importance to the study. The most commonly used
method is to fix the flukes under slight cover glass pressure. In this method, the fluke
is placed on a slide, and flattened under a cover glass pressure, and fixed in AFA or
70% alcohol, or 10% formaldehyde; the pressure exerted depends on the size and
thickness of the parasite. Usually, the pressure that is just sufficient to keep the fluke
flat is applied. This method, although commonly followed, is criticized as disrupting
the internal organs and causing damage to the parasite.
Cribb and Bray (2010) suggested fixation by pipetting live parasites into nearly
boiling saline and immediately transferring them to the fixative. Cribb and Bray
(2010) described the procedure in sufficient detail with illustrations. Most of the
present-day scientists are adopting this method. Later, Justine et al. (2012) presented
a slightly modified version of this method.
Another method suggested is fixation in Berland’s fluid (95% glacial acetic acid
+ 5% formalin). The parasite is pipetted into Berland’s fluid and almost immediately
preserved in 80% ethanol. This method may be useful for the parasites having a thick
muscular tegument, but the material will not be suitable for molecular study.
© Crown 2018 5
https://doi.org/10.1007/978-94-024-1535-3_2
6 2 Materials and Methods
Fixation is followed by staining in carmalum or haematoxylin, dehydrating in

alcohol series, clearing in creosote or methyl salicylate and mounting in Canada
balsam.
In India the number of parasitologists trained in systematics is declining and this
is particularly the case with digeneans. Only traditional taxonomic methods have so
far been employed for identifying the digeneans. The present-day trend in parasite
taxonomy is to use both traditional and molecular methods; the latter techniques
are gaining special emphasis. In this method, DNA sequences are used to address a
variety of questions on the relationships and species boundaries of organisms. This
is found to be a very useful approach that complements traditional taxonomy based
on morphology. The protocol for molecular taxonomy consists of DNA extraction
from the parasite, amplification of the DNA extracted, sequencing and construction
of phylogenetic tree using proper software. The database containing sequences of
digenetic trematodes is quickly growing and being used for taxonomic identification
of many species. In India, however, molecular methods are yet to find their place and
not even a single species of digenean has been subjected to analysis of sequences. It
is essential to initiate these studies on a priority basis. Technical details for molecular
methods have been well established and a number of publications have appeared (e.g.
Nolan & Cribb, 2005; Blasco-Costa et al. 2016).
Preparation of the document: The vast literature published in various journals
on digenetic trematodes from marine fishes from different coastal regions of India
is the main source for the preparation of this document. In this context, the data
presented in world register of marine species (WoRMS) compiled mainly by David
Gibson (2004–2016) has proved to be extremely useful in getting details of species
of digeneans recorded under each digenean genus, their validity, taxonomic status
and the relevant literature.
The major part of the document deals with systematic account on digenetic trema-
todes collected from marine fishes of the Indian region. Only adult digeneans are
treated in this manual. The metacercarial stages and the cercariae are not covered.
Almost all the species recorded are covered. Keys are provided for the differentiation
of subfamilies, genera and species. Keys are designed to include only species occur-
ring in Indian region. Diagnoses are provided for all the taxa down to generic level.
Brief descriptions for each species including the key characters have been furnished.
Illustrations have been copied from published literature after obtaining necessary
permission from the publishers. Information on the hosts and geographical distribu-
tion is also provided for each species. The host scientific names given in the original
description have been revised following ‘Fish Base’. All the measurements are given
in microns. Information on the number of specimens collected by the author for each
species is also provided. This information is felt desirable because for some species
the descriptions are based on only one specimen and these are to be accepted with
reservation.
2 Materials and Methods 7
Abbreviations are used for indicating the localities along east and west coast
of India. These are: AS, Arabian Sea; BOB, Bay of Bengal; PR, Puri Coast; VSK
Visakhapatnam coast; BOM, Bombay coast; GOM, Gulf of Mannar; MS, Madras
coast.
Chapter 3
The Digenetic Trematodes
General Organization and Life Cycles (Fig. 3.1a–e)
The vast majority of digeneans are endoparasitic, except for a few such as Transver-
sotrema spp. which are ectoparasitic. The preferred site of infection is the gut; there
are also reports from the urinary bladder (gorgoderids), air bladder (Elongoparorchis)
and blood vessels (aporocotylids). The didymozoid trematodes are exceptional in that
they occur encysted in pairs in almost all the organs of the fish body where they appear
as prominent yellowish cysts. Wide variations also exist in the life cycle patterns.
As an adaptation to the site of infection, the digenean body organization varies in
different species, while at the same time reflecting similarities to the basic structure
of the group.
The body is leaf-like, oval or elongate, filamentous, transversely oval in some
(Transversotrema spp.). Didymozoids are peculiar in that the body is often divisible
into a filiform forebody and a cylindrical hindbody. In some hemiurids, the posterior
part of the body is modified into retractable tail-like structure known as ecsoma. The
body size also varies from less than a millimetre (bivesiculids) to more than 5 cm
(Hirudinella).
The body typically bears two suckers, the oral and ventral suckers. The oral
sucker surrounds the mouth at the anterior end, except in bucephalids where the
mouth is located at the ventral midbody (gasterostomate condition) and the sucker
at the anterior end is then known as the anterior sucker which may be simple or
provided with a solid, muscular rhynchus with or without tentacles. The oral sucker
may be simple or provided with a ring of spines (Stephanostomum) or a collar-like
structure (Orientodiploproctodaeum). In the genus Karyakartia, the rim of the oral
sucker is provided with anteriorly directed tentacles. The ventral sucker is usually
located ventrally in the body proper (distome), or at the posterior end of the body
(amphistome), or may be totally absent (monostomate). It is usually sessile or may
be borne on a peduncle as is the case in the genus Pseudopecoeloides Yamaguti,
1940. In the genus Opegaster Ozaki, 1942, the anterior and posterior lips of the
© Crown 2018 9
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10 3 The Digenetic Trematodes
ventral sucker are provided with digital papillae. Usually, the size relationships of
the two suckers are indicated by the sucker ratio, which is the ratio of the transverse
diameters of the two suckers, with the diameter of oral sucker taken as one. Similarly,
the region of the body between anterior end and anterior margin of ventral sucker
is known as the forebody and that between posterior margin of ventral sucker and
posterior end is known as the hindbody.
The outer layer of the body wall is known as the tegument, a dynamic syncytial
cytoplasmic layer connected to underlying subtegumental cells through cytoplasmic
extensions. The tegument may be thin or thick, smooth, spined or annulated.
Almost all the digeneans possess a rudimentary digestive system. Mouth is usually
at the anterior end, opening terminally or subterminally. In bucephalids, the mouth is
located at about the ventral midbody (gasterostomatous condition). The mouth leads
into prepharynx, pharynx and oesophagus, the relative sizes of which vary from
species to species. The pharynx is a muscular, bulbous structure. The oesophagus
usually bifurcates into two caeca, usually in the forebody, rarely in the hindbody. The
caeca terminate blindly close to the posterior end of the body or open through two ani
(e.g. Bianium) or open into the excretory bladder forming an uroproct (Stephanos-
tomum). The caeca may also unite forming a cyclocoel (e.g. Coitocaecum) which
may in turn open through a single anus (Opegaster). In the haplosplanchnids, the
caecum is single and lined by prominent cells. In some hemiurids, the oesophagus
is provided with Drüsenmargen, a glandular structure.
Most digeneans are hermaphroditic and protandrous. Only a few digeneans such
as didymozoids and schistosomes are gonochoristic with separate male and female
individuals either totally free or partially or fully fused. The genital pore is single
and is usually located in the forebody, rarely in the hindbody, and may be median,
submedian or lateral. The genital pore opens into a genital atrium into which open
the male and female terminal ducts. Typically, there are two testes, oval, lobed or
follicular, located in the hindbody or forebody, disposed symmetrically, obliquely or
in tandem. The testis is single in some monorchiids, and multiple in Pleorchis spp.
The vasa differentia lead from the testes into a seminal vesicle which is saccular, or
bipartite or sinuous and either lies free in the parenchyma or is enclosed in a cirrus-
sac. Sometimes, a part of the seminal vesicle lies outside the cirrus-sac, dividing it
into two parts, the internal and external seminal vesicles (lepocreadiids). In some
opecoelids, the cirrus-sac is absent and the seminal vesicle lies free in parenchyma.
The seminal vesicle continues as the pars prostatica and then as an ejaculatory duct,
which may extrude to form cirrus, which leads into genital atrium. The ejaculatory
duct may be lined with spines (monorchiids). The pars prostatica is surrounded
by prostatic cells that often fill the cirrus-sac. In some forms such as hemiurids,
the cirrus-sac is absent and the metraterm joins the ejaculatory duct and forms a
hermaphroditic duct which may be enclosed in a sac, the sinus-sac also called a
hermaphroditic sac. The genital atrium maybe thin-walled and inconspicuous, long
and tubular as in some acanthocolpids, and in some monorchiids its lumen as well
as the cirrus bears prominent spines. In bucephalids, where the genital pore is at the
posterior end of body, the genital atrium is spacious, and the cirrus-sac is extended
into it and bears blunt papillae. In tandanicolids and Monodhelmis spp., the genital
General Organization and Life Cycles (Fig. 3.1a–e) 11
atrium is provided with an accessory copulatory organ consisting of a solid anteriorly

directed digitiform process and associated glandular tissue.
The female reproductive system is complex, consisting of several organs and a
system of ducts connecting the various organs. Typically, there is a single spherical or
lobed ovary, from which arises the oviduct. Near the origin of the oviduct, there is a
valve-like structure known as the ovicapt. The oviduct feeds into a seminal receptacle
and also Laurer’s canal. The latter usually opens through a dorsal pore; but in some
worms, such as Elongoparorchis pneumatis, it opens into a blind sac known as Juel’s
organ which contains a thick fibrous wall and is filled with degenerating cellular
components like sperm, ova and vitelline cells (Madhavi & Rao, 1974). In some
digeneans, the seminal receptacle is absent and the sperm is stored in the proximal
part of the uterus and is known as the uterine seminal receptacle. The oviduct joins
the vitelline duct and forms the ootype surrounded by Mehlis’ gland, consisting of a
cluster of gland cells. The functions of Mehlis’ gland are still not clearly established,
but it is believed that the secretion helps in the release of shell globules from the
vitelline cells, that ultimately fuse and form the eggshell in the ootype. The ootype
continues as the uterus where the eggs constructed in the ootype accumulate. The
uterus may be short, straight or long and coiled, may be confined to a small region
of the body or occupy practically the entire body. The distal part of the uterus is
known as the metraterm, a muscular tube which opens into the genital atrium or
joins the ejaculatory duct to form the hermaphroditic duct. In monorchiids, there is a
terminal organ associated with the genital atrium, which may be simple or bipartite
and spined or not, and the metraterm opens into the terminal organ at its base or its
middle part. The uterus contains few or numerous eggs. The eggs are oval, operculate
or non-operculate, and the capsule may rarely extend to form a filament. Another
prominent structure in the female reproductive system is the vitellarium, which may
be follicular occupying a varying extent of the lateral fields of the body or may be
reduced to one or two compact masses situated near the ovary.
The excretory system is of the protonephridial type and performs two functions:
osmoregulation and removal of waste. Its main components are the excretory bladder,
numerous flame cells and the connecting tubules. The excretory bladder may be I-
shaped, V-shaped or Y-shaped and opens through a posterior pore situated terminally
or subterminally. The shape of the excretory bladder and its anterior extent is of
taxonomic importance. In Prosogonotrema, there occurs a long and wide accessory
vesicle in the hindbody dorsal to the normal excretory vesicle, which is known as
Manter’s organ. Some digeneans also possess a lymphatic system (Apocreadiidae).
The nervous system is of a primitive type consisting of a pair of ganglia (brain)
located in the pharyngeal region, three pairs of anterior and three pairs of posterior
longitudinal trunks and several commissures. Some digeneans also retain eyespots
from the cercarial stage.
Life Cycles of Digenetic Trematodes
The life cycles of digeneans involve an alternation of generations with the sex-
ual generation represented by the adult fluke occurring in the definitive host and the
asexual generation represented by the ‘larval’ stages (sporocyst and redia—known as
parthenitae) occurring in the first intermediate host, usually a mollusc and occasion-
ally an annelid. The adult stage with high fecundity produces large numbers of eggs,
and the asexual multiplication in the first intermediate host results in the generation
of numerous infective stages. Both of these processes ensure successful invasion of
the definitive host and a built up of heavy infections.
A wide range of patterns exist in the life cycles of digeneans depending on the
behaviour, the food and feeding habits of the definitive host, the nature of the inter-
mediate host and the surrounding environment. These patterns are discussed in detail
by Cribb et al. (2003) and Niewiadomska & Pojmanska (2011). These are discussed
briefly here, with examples from digeneans infecting marine fishes. Usually, the
Fig. 3.1 a–e Organisation of digenetic trematodes. a Adult stage b, c, e Male terminal genitalia: b
typical digenean c hemiurid e haploporid. d Female reproductive system. Abbreviations c, cirrus;
cs, cirrus-sac; ed, ejaculatory duct; ga, genital atrium, gp, genital pore; hd, hermaphroditic duct; hs,
hermaphroditic sac; ic, intestinal caecum; lc, Laurer’s canal; mg, Mehlis’ gland; mt, metraterm; oc,
ovicapt; od, oviduct; oe, oesophagus; oot, ootype; os, oral sucker; ph, pharynx; pp, pars prostatica;
pph, prepharynx; rs, seminal receptacle; ss, sinus sac; sv, seminal vesicle; t, testis; ut, uterus; vd,
vitelline duct, vr, vitelline reservoir; vt, vitellarium
Life Cycles of Digenetic Trematodes 13
eggs of the parasite are released into the surrounding water medium, and they either
hatch into a tiny motile stage known as the miracidium which actively penetrates the
molluscan intermediate host or the eggs with the miracidium are swallowed by the
molluscan host in the tissues of which they are stimulated to hatch. The miracidium
develops into a mother sporocyst in the tissues, which in turn produces daughter
sporocysts or rediae. The germ cells inside these stages multiply rapidly by asexual
means and produce large numbers of cercariae which are released into the external
environment. They infect the definitive host either through the active skin penetration
(aporocotylids) or follow a passive mode of penetration by encysting either in the
open environment or algae (haplosplanchnids, gyliauchenids) or in the tissues of a
second intermediate host to develop into metacercariae which are consumed with
the second intermediate host by the definitive host. The metacercariae excyst in the
gut of the definitive host and the immature flukes migrate into the preferred habitat
to develop into egg-producing stages. Details are given in Fig. 3.1.
The typical life cycle involves three hosts, two intermediate hosts and a definitive
host, which pattern is followed in most digeneans, such as bucephalids, opecoelids,
lepocreadiids, acanthocolpids, cryptogonimids and monorchids to mention a few.
In some digenean families, abbreviation of the life cycle occurs usually through
the elimination of the second intermediate host. The simplest type of life cycle is
reported in bivesiculids such as Paucivitellosus fragilis and P. hanumanthai, which
do not have a second intermediate host, but infect the fish host directly when the
cercaria is ingested by the fish. In some digeneans such as hemiurids, the life cycle
Fig. 3.1 (continued)

Fig. 3.2 Diversity in the life cycle pattern of Digenea of marine fishes
is extended by the addition of a paratenic (transport) host which helps in the transfer
of the parasite from the second intermediate host to the definitive host.
Information available from India on the life cycles of marine digeneans is scanty.
The life cycles have so far been described for three species; Paucivitellosus hanum-
nathai by Gnanamani (1989), Stephanostomum cloacum by Madhavi & Shameem
(1993) and Helicometra gibsoni by Murugesh et al. (1993). Investigations on life
cycles of these digeneans have to be undertaken on a large scale employing both
experimental studies and molecular methods (Fig. 3.2).
Classification of Digenea 15
Classification of Digenea
The Digenea is recognized as a subclass in the Class Trematoda along with the sub-
class Aspidogastrea. Aspidogastrea is a small group comprising about 80 species
occurring as adults in molluscs, fishes and reptiles. On the other hand, the Digenea,
comprising about 18,000 nominal species, represent the largest group of internal
metazoan parasites (Cribb et al, 2001), capable of infecting members of all groups of
vertebrates. They are especially common in marine fishes. In view of the great diver-
sity exhibited by them in their morphology, life cycle pattern and the extraordinary
variation in the host selection, classifying them in a form reflecting their phylogeny
has proved to be a challenging issue. Cribb et al (2001) provided a historical review of
provisional classifications suggested for Digenea. Early classifications relied mostly
on adult morphological features such as the number and arrangement of suckers
which are now considered as of limited taxonomic value. As knowledge on cercarial
characters and the life cycles of digeneans has accumulated, this information along
with a stress on the excretory system was used in later classifications. The system
proposed by LaRue (1957) covers these various aspects and it was the most widely
accepted classification for many years.
With the progress of knowledge on ultrastructure and DNA sequences of differ-
ent species of digeneans, many changes have been suggested in the phylogenetic
relationships of different taxa of digeneans and the overall classification. The best
classification covering all families of digeneans and utilizing both morphological
and molecular data is that followed in ‘Keys to the Trematoda’ volumes 1 to 3,
proposed by Gibson, Bray and Jones (2002–2008). Although higher taxa were not
covered, this classification dealing up to superfamilies is found to be most useful and
comprehensive and served as the basis for other classifications proposed later. The
important conclusion from this study is that both the morphological and molecular
matrices are complimentary and both kinds of data are of value in inferring relation-
ships among Digenea. Prior to this work, Cribb et al. (2001) attempted a combined
approach using morphological characters for all stages of the digenean life cycle and
complete18sRDNA sequences for 75 species of digeneans. The conclusions drawn
from these two studies are similar. Another comprehensive study in this direction is
that of Olson et al (2003) who, employing all the available molecular data on different
taxa of digeneans, created a classification that is most acceptable and accurate. The
notable feature in this classification is that only two orders were recognized instead
of three as in the earlier classifications: order Diplostomida consisting of 3 suborders
and Plagiorchiida with 13 suborders. Recently a number of families or species of
digeneans have been subjected to molecular phylogenetic analysis employed mainly
to determine the monophyly of the groups and the validity of different taxa and
their interrelationships. This type of information is now available on digeneans of
marine fishes belonging to the Lepocreadioidea by Bray et al. (2009), Opecoelidae
by Bray et al. (2016), Bunocotylinae, by Perez-Ponce-de-Leon& Brooks (2016),
Hemiuroidea by Blair et al (1998), Gyliauchenidae by Blair& Barker (1993), and
many significant contributions were made on their phylogenetic relationships. This
information is presented here in the accounts dealing with individual families of dige-
neans. Kostadinova & Perez-del-Olmo (2014) in the article entitled ‘The Systematics
of the Trematoda’ provided a comprehensive review on the various classifications
and other contributions on the molecular analysis of digeneans and compared the
Table 3.1 Classification of Digenea Adopted for Digenea of Indian marine fish
Superfamily Family
Order Diplostomida Olson, Cribb, Tkach &
Littlewood, 2003
Schistosomatoidea Stiles & Hassall, 1898 Aporocotylidae Odhner, 1912
Order Plagiorchiida La Rue, 1957
Apocreadioidea Skrjabin, 1942 Apocreadiidae Skrjabin, 1942
BivesiculoideaYamaguti, 1934 BivesiculidaeYamaguti, 1934
Bucephaloidea Poche, 1907 Bucephalidae Poche, 1907
Gymnophalloidea Odhner, 1905 Fellodistomidae Nicoll, 1909
Tandanicolidae Johnston, 1927
Haplosplanchnoidea Poche, 1926 Haplosplanchnidae Poche, 1926
Hemiuroidea Looss, 1899 Hemiuridae Looss, 1899; Accacoelidae
Odhner, 1911; Derogenidae Nicoll,
1910,DictysarcidaeSkrjabin&Guschanskaja,
1955; Didymozoidae Monticelli, 1888;
Hirudinellidae Dollfus, 1932; Lecithasteridae
Odhner, 1905; Ptychogonimidae Dollfus,
1937; Sclerodistomidae Odhner, 1927;
Sclerodistmoididae Gibson &Bray, 1979;
Syncoeliidae Looss, 1899.
Transversotrematoidea Witenberg, 1944 Transversotrematidae Witenberg, 1944
Haploporoidea Nicoll, 1914 Haploporidae Nicoll, 1914,
AtractotrematidaeYamaguti, 1939
Lepocreadioidea Odhner, 1905 Lepocreadiidae Odhner, 1905;
AephnidiogenidaeYamaguti, 1934;
Enenteridae Yamaguti, 1958; Gyliauchenidae
Fukui, 1929; LepidapedidaeYamaguti, 1958.
Brachycladioidea Odhner, 1905 Acanthocolpidae Luhe, 1906
Opecoeloidea Ozaki, 1925 Opecoelidae Ozaki, 1925; Opistholebetidae
Fukui, 1929
Monorchioidea Odhner, 1911 Monorchiidae Odhner, 1911
Gorgoderoidea Looss, 1899 Gorgoderidae Looss, 1899
Microphalloidea Ward, 1910 Zoogonidae Odhner, 1902; Faustulidae Poche,
1926
Opisthorchioidea La Rue, 1957 Cryptogonimidae Ward, 1917; Cladorchiidae
Fischoeder, 1901; MicroscaphidiidaeLooss,
1900; Cephalogonimidae Looss, 1899;
OpisthorchiidaeLooss, 1899
Classification of Digenea 17
two classifications namely the one followed in the Keys to the Trematoda and the
other proposed by Olson et al (2001).
The relationships at the superfamily level appear to be well resolved, but several
issues remain to be solved at the higher digenean taxa. In the present account, the
classification of Digenea proposed in ‘Keys to the Trematoda’ is modified to include
only the families relevant to the present work. Some other changes have also been
made. All the superfamilies are distributed under two orders: the Diplostomida Olson,
Cribb, Tkach & Littlewood, 2003, and Plagiorchiida La Rue, 1957. No attempt was
made to include suborders (Table 3.1).
Part I
Systematic Account—Order
Diplostomida Olson, Cribb, Tkach,
Bray & Littlewood, 2003
Chapter 4
Superfamily Schistosomatoidea Hassall,
1898
Family Aporocotylidae Odhner, 1912
[Syn. Sanguinicolidae von Graff, 1907]
Two different family names have been in use for the blood flukes of fishes: the
Aporocotylidae Odhner, 1912, and the Sanguinicolidae von Graff, 1907. The San-
guinicolidae was widely accepted and is followed by Yamaguti (1958, 71) and by
Smith (2002) in the Keys to the Trematoda. Overstreet & Køie (1989) considered
Aporocotylidae to be a junior synonym of Sanguinicolidae. However, Bullard et al.
(2009) from a deep and critical analysis of the literature recommended the family
name Aporocotylidae for these flukes, and subsequently, this family name is used in
all the publications.
The family comprises trematodes that principally parasitise the vascular system
of fishes (Smith, 2002). The body organization shows several unique features such as
a flat leaf-like body and the usual absence of suckers. The eggs are thin, transparent
and the eggshell is made of elastin (Madhavi & Rao, 1971a, b).
Diagnosis: Body flat, leaf-like. Tegument with transverse rows of spines. Oral
sucker absent or rudimentary. Ventral sucker absent. Pharynx absent. Oesophagus
surrounded by gland cells. Intestinal caeca H- or X- or inverted U-shaped. Testis
single, oval or follicular or transversely coiled, in middle third of body. Cirrus-sac
present or absent. Ovary slightly lobed, post-testicular. Seminal receptacle present or
absent. Uterus postovarian. Eggs thin-shelled. Genital pore dorsal, postovarian, near
posterior end of body. Vitellarium follicular, mostly anterior to ovary. Type genus:
Aporocotyle Odhner, 1900.
Only two species of aporocotylids have been reported from India: Orchispirium
heterovitellatum Madhavi & Rao, 1970 from an elasmobranch fish and Paradeonta-
cylix megalaspium Trivenilakshmi & Madhavi, 2007 from a teleost fish.
© Crown 2018 21
R. Madhavi and R. A. Bray, Digenetic Trematodes of Indian Marine Fishes,
https://doi.org/10.1007/978-94-024-1535-3_4
22 4 Superfamily Schistosomatoidea Stiles & Hassal, 1898
Fig. 4.1 Orchispirium

heterovitellatum (after
Madhavi & Rao, 1970, J.
Parasitol, 56, p 42 Courtesy
Allen Press)
Genus Orchispirium Madhavi & Rao, 1970
Diagnosis: Body lanceolate. Oesophagus long, surrounded by gland cells, enlarged

posteriorly to form vesicle. Intestine with posterior caeca only, appearing as inverted
U-shaped. Testis single, transversely coiled tube, between and posterior to caeca.
Cirrus-sac poorly developed, encloses seminal vesicle and short, eversible cirrus.
Ovary lobed, post-testicular. Uterus postovarian. Metraterm short. Eggs thin-shelled.
Vitellarium follicular, with follicles much more extensive on right side than on left
side. Type and only species: O. heterovitellatum Madhavi & Rao, 1970.
Orchispirium heterovitellatum Madhavi & Rao, 1970 (Fig. 4.1)
Host: Dasyatidae: Brevitrygon imbricata (Bloch & Schneider)
Location: Mesenteric veins
Locality: VSK, BOB
Reference: Madhavi & Rao (1970a).
Description: Body spatulate, 3230–4480 long 990–1280 wide, tapering gradually

towards ends. Tegument tuberculated, spines absent. Mouth small, surrounded by
small sucker, visible only under high magnification. Oesophagus long, surrounded
Family Aporocotylidae Odhner, 1912 23
by gland cells, with vesicle at posterior end. Caeca narrow, terminate preovarian.
Testis large, transversely coiled, with 9–11 loops, anterior surface of loop smooth,
posterior surface crenulated, situated in intercaecal space in front of ovary. Genital
pore single, submedian, postovarian. Cirrus-sac thin-walled, slender, extends from
posterior margin of ovary to genital pore, encloses long seminal vesicle and short
cirrus. Ovary multi-lobed, immediately post-testicular, to right of midline. Uterus
post-testicular. Metraterm short. Eggs small, thin-shelled, transparent, 23 by 16 in
size. Vitelline follicles small, asymmetrical in distribution, extending from nerve
commissure to level of ovary on right side, to level of intestinal bifurcation on left
side.
Remarks: The important characteristics of O. heterovitellatum are considered to be:

the flat leaf-like body, the lack of obvious suckers, and the presence of a vesicle at
the posterior region of the oesophagus, the U-shaped intestinal caeca, the testis with
transverse loops, the lobed ovary and asymmetrically distributed vitelline follicles.
The original description of the species was revised by Bullard & Jensen (2008) based
on a re-examination of holotype and paratype specimens. In the revised description,
a rudimentary oral sucker is stated to be present. However, it is visible in only a
few specimens, it is very small and almost indistinguishable. So far, this is the only
species in the genus.
Genus Paradeontacylix McIntosh 1934
Diagnosis: Body small, flat. Tegument with transverse rows of spines. Suckers and
pharynx absent. Intestinal caeca X or H-shaped. Testes 19–71, arranged in two or
three rows between caeca. Cirrus-sac encloses seminal vesicle and pars prostat-
ica. Male genital pore opens dorsally near left body margin. Ovary shield or heart
shaped lying posterior to testis. Seminal receptacle absent. Uterus largely preovarian.
Metraterm present. Female genital pore opens dorsally in midline, anterior to male
pore. Eggs thin-shelled, ovoid. Vitellarium follicular, extensive, from near anterior
end of body to near ovary.
Type species: O. sanguinicoloides McIntosh, 1934
Paradeontacylix megalaspium Trivenilakshmi & Madhavi, 2007 (Figs. 4.2 and 4.3)
Host: Carangidae: Megalaspis cordyla (Linnaeus)

Location: Gill blood vessels
Locality: VSK, BOB
Reference: Trivenilakshmi & Madhavi (2007).
Description: Body long, slender, worm-like, 1920–3520 long and 96–192 wide.
Tegument thin, transparent, with spines arranged in 400–450 regular rows, each
row with 8–10 spines. Suckers absent. Oesophagus long, sinuous. Caeca H-shaped,
anterior branches short, posterior ones long. Testis follicular, approximately 70–86
follicles, arranged in two irregular rows in intercaecal region. Cirrus-sac encloses
seminal vesicle and short cirrus. Male genital pore submedian on dorsal side, at
some distance from posterior end. Ovary post-testicular appears as large irregular
24 4 Superfamily Schistosomatoidea Stiles & Hassal, 1898
Fig. 4.2 Paradeontacylix

megalaspium (after
Trivenilakshmi & Madhavi,
2007, Zootaxa, 1512, p 67)
Fig. 4.3 P. megalaspium

Enlarged view of posterior
part of body (after
Trivenilakshmi & Madhavi,
2007)
mass. Seminal receptacle absent. Uterus fills post-testicular area. Female genital pore
submedian, just anterior to male pore. Eggs small, oval, 20–24 long 12 wide. Vitelline
follicles small, occupy most of area between nerve commissure and genital pore.
Remarks: By possessing characters such as H-shaped caeca, a follicular testis, sep-
arate male and female genital pores and a postovarian distribution of uterine coils,
Family Aporocotylidae Odhner, 1912 25
the present form fits into the genus Paradeontacylix. This species differs from all the
other species of the genus by possessing the following combination of characters:
the long, slender worm-like body, with 400–450 rows of marginal tegumental spines,
the absence of enlarged posterior tegumental spines and the presence of 70–86 testis
follicles occupying the major part of the body.
Part II
Systematic Account—Order
Plagiorchiida La Rue, 1957
Chapter 5
Superfamily Bivesiculoidea
Yamaguti, 1934
Family Bivesiculidae Yamaguti, 1939
[Syn. Paucivitellosidae Yamaguti, 1971, Treptodemidae Yamaguti, 1971]

The family Bivesiculidae is comprised of a small group of digeneans that are par-
asitic in the digestive tract of marine fishes. Members of this family are characterized
by the small body size, the lack of suckers, the fairly large eggs and the V-shaped
excretory bladder. Molecular studies by Blair et al. (1998) revealed no close relation-
ship between this family and any other digenean family and the molecular studies of
Olson et al. (2003) and Littlewood et al. (2015) indicated that it is the most basally
derived family in the Plagiorchiida. The simple organization and the two-host life
cycle have also led to the conclusion that the family retains many plesiomorphic
characters, although a three-host life has occasionally been reported (Cribb et al.
1998).
Diagnosis: Body plump, elongate or transversely oval. Tegument spined. Eyespot
pigment present. Suckers absent. Prepharynx short. Pharynx oval. Oesophagus dis-
tinct. Caeca long, terminate blindly. Testis single, at midbody or just posteriorly
located. Cirrus-sac present. External seminal vesicle present or absent. Genital pore
on ventral midline. Ovary pretesticular. Uterine seminal receptacle present. Uterus
of varying length. Eggs large. Vitellarium composed of widely distributed follicles
or two small lobed masses. Excretory vesicle V-shaped. Type genus Bivesicula Yam-
aguti, 1934.
Three species of bivesiculids from three genera Bivesicula Yamaguti, 1934,
Bivesiculoides Yamaguti, 1938, and Paucivitellosus Coil, Reid & Kuntz, 1965, have
been recorded from Indian marine fishes.
Key to genera:
1. Vitellarium two small lobed masses, eggs thin-shelled …………… Paucivitel-
losus Coil, Reid & Kuntz, 1965
© Crown 2018 29
https://doi.org/10.1007/978-94-024-1535-3_5
30 5 Superfamily Bivesiculoidea Yamaguti, 1934
Vitellarium follicular, follicles distributed widely, eggshell thick …................. 2

2. Uterus extends posterior to testis ………………................................……
Bivesicula Yamaguti, 1934
Uterus not passing posterior to testis ……................. ................….…. Bivesicu-
loides Yamaguti, 1938
Genus Bivesicula Yamaguti, 1934
Diagnosis: Body oval. Caeca long or short. Suckers absent. Testis single, at midbody.
Ovary pretesticular. Uterus intercaecal, passing anterior to testis. Eggs thick-walled,
unembryonated. Vitelline follicles extensive, inter- and extracaecal. Excretory arms
extend to level of caeca. Type species: B. claviforme Yamaguti, 1934.
Bivesicula australis Crowcroft, 1947 (Fig. 5.1)

[Syn. B. congeri Yamaguti, 1970]
Host: Unidentified marine fish
Locality: Andamans
No. 8
Distribution: Australia, Hawaii, Andamans
Reference: Hafeezullah and Dutta (1980)
Fig. 5.1 Bivesicula australis

(after Hafeezullah and Dutta
1980, Rec. Zool. Soc. India,
45, p 81)
Family Bivesiculidae Yamaguti, 1939 31
Description: Body 990–1072 long, 385–453 wide, uniform. Eyespot pigment

present, dissociated. Ventral sucker absent. Pharynx 35–42 in diameter, squarish.
Oesophagus of medium length, muscular. Caeca simple, extend slightly beyond
testis. Testis single, postequatorial. External seminal vesicle attached to anterior end
of cirrus-sac. Cirrus-sac directed anteriorly, encloses internal seminal vesicle, thick-
walled pars prostatica and highly muscular cirrus, and numerous prostate cells. Geni-
tal atrium shallow. Genital pore equatorial or postequatorial. Ovary almost equatorial,
submedian. Vitellarium consists of vitelline follicles extending between oesophageal
level and level of anterior margin of testis, tending to coalesce medially. Uterus
descending well posterior to testis, then ascending. Eggs collapsed 70–87 by 42–45.
Remarks: B. australis was first reported from Australia by Crowcroft (1947).
Hafeezullah and Dutta (1980) considered B. congeri Yamaguti, 1970, from Hawaii
a synonym of B. australis. The Indian forms differ from the original description of
the species in the squarish shape, the size of the pharynx and the posterior extent
of the vitelline follicles reaching the level of the testis. Crowcroft (1947), Yamaguti
(1970) and Hafeezullah & Dutta (1980) all refer to the muscular part of the alimen-
tary system as an ‘oral sucker’. Cribb (2002) stated that the ‘digestive system has
just one muscular structure, which is now usually (but not universally) interpreted as
a pharynx’. Le Zotte (1954) gave histological evidence for this interpretation, which
has been followed by most recent workers (e.g. Sey et al., 2003; Nahhas et al., 2004;
Trieu et al., 2015; Atopkin et al. 2017).
Genus Bivesiculoides Yamaguti, 1938
Diagnosis: Body small, oval. Caeca terminate in posterior third of body. Testis pos-
terior to midbody. Ovary pretesticular. Uterus not extending posterior to testis. Eggs
thick-walled unembryonated. Vitelline follicles extensive, inter- and extracaecal, co-
extensive with caeca. Type species B. atherinae Yamaguti, 1938.
Bivesiculoides scari Hafeezullah, 1971 (Fig. 5.2)
Host: Scaridae: Scarus ghobban Forsskal
Locality: Trivandrum, GOM
Reference: Hafeezullah (1971b)
Description: Body 1682–2434 long 670–824 wide, provided with cap-like fold at
anterior end. Mouth terminal. Prepharynx present. Pharynx globular. Caeca obscured
by vitellarium. Testis oval, in middle of posterior half of body. Cirrus-sac in middle
of body, contents not clear. Vitellarium follicular, between anterior cap-like fold and
posterior end of body. Uterus pretesticular. Eggs 59–77 × 44–59. Excretory vesicle
V-shaped.
Remarks: There are too many details missing from the description given for the
species. More specimens need to be examined. Nevertheless, it can be seen that it
differs from the three species of Bivesiculoides in the shape of the body and the
pharynx and the extent of the vitellarium.
32 5 Superfamily Bivesiculoidea Yamaguti, 1934
Fig. 5.2 Bivesiculoides

scari (after Hafeezullah,
1971, J. Helminthol., 45, p
81, Courtesy Oxford Press)
Genus Paucivitellosus Coil, Reid & Kuntz, 1965.
Diagnosis: Body small, squat. Eyespot pigment present. Testis single. Ovary ante-
rior to testis. Uterus extends posterior to testis. Eggs thin-walled and embryonated.
Vitellarium two, small compact lobed masses. Type species: P. fragilis Coil, Reid &
Kuntz, 1965.
Paucivitellosus hanumanthai Gnanamani, 1989 (Fig. 5.3)
Host: Mugilidae Mugil cephalus L.
Locality: VSK, BOB
Reference: Gnanamani (1989)
Description: Body pear shaped 480–592×20–384. Tegument with minute spines.

Mouth subterminal. Pharynx muscular. Oesophagus long. Intestinal bifurcation at
junction of anterior and middle third of body. Caeca broad, extend to level of anterior
margin of testis. Eyespots prominent. Testis single, large, median, in posterior third of
body. Cirrus-sac median, ellipsoidal, immediately posterior to intestinal bifurcation.
Family Bivesiculidae Yamaguti, 1939 33
Fig. 5.3 Paucivitellosus

hanumanthai (after
Gnanamani, 1989)
Ovary dextral, between cirrus-sac and testis. Vitellarium formed of two compact bi-
lobed masses on either side of cirrus-sac. Eggs large, amber-coloured, thin-shelled,
56–68 × 24–32.
Remarks: Gnanamani (1989) differentiated P. hanumanthai from P. fragilis Coil,

Reid & Kuntz, 1965, the type species of the genus, on the basis of its being larger in
size and having a larger ovary and larger testes and eggs. However, Cribb et al.
(1994), who examined large numbers of P. fragilis collected from different hosts
and localities, found wide variations in body size, egg size and vitellarium shape
and considered P. hanumanthai as a synonym of P. fragilis. Atopkin et al. (2017)
described P. vietnamensis Atopkin, Besprozvannykh, Ngo, Van Ha, Van Tang,
Ermolenko & Beloded, 2017, from the mullet Liza subviridis in waters off Vietnam
and compared the partial 18S and 28S rDNA sequences with those of the species P.
fragilis (Australian form reported by Cribb et al. (1994) and sequences reported by
Cribb et al. (2001)) and two unnamed host-specific forms from other mullet species
from the same waters. P. fragilis was originally described from the mullet Planiliza
macrolepis as [C. troscheli] off Taiwan (Coil et al. 1965), and no molecular data
are available from this source, so it is distinctly possible that the Australian worms
are not conspecific to this. Atopkin et al. (2017) gave the opinion that their own
‘observations do not preclude P. fragilis sensu Cribb et al. (1994) from representing
a set of cryptic species’. Considering these findings and pending additional evidence,
P. hanumanthai is tentatively retained here as a valid species.
Gnanamani (1989) showed that the snail Clypeomorus clypeomorus Jusseaume

serves as the intermediate host for P. hanumanthai and gave details of the structure
of the cercaria and pattern of the life cycle.
Chapter 6
Superfamily Bucephaloidea Poche, 1907
Family Bucephalidae Poche, 1907
Bucephalidae is a large family of digenean parasites occurring in the gut of marine,

brackish and freshwater piscivorous fishes. Members of this family are easily recog-
nised by the following combination of characters: the position of the mouth at mid-
body on the ventral surface, the presence of an anterior rhynchus, the absence of a
ventral sucker, the position of the genital pore near the posterior extremity of the
body and the presence of a genital lobe within the genital atrium. The structure of the
rhynchus is of considerable importance in the separation of various bucephalid gen-
era. The exact systematic placement of Bucephalidae within the Digenea has been
much debated and the higher classification proposed by Olson et al. (2003) placing
it in the order Plagiorchiida LaRue, 1957, and suborder Bucephalata Olson, Cribb,
Tkach, Bray & Littlewood, 2003, is followed here.
So far about 56 species of bucephalids spread over 8 genera and 2 subfamilies
have been recorded from Indian marine fish. The maximum numbers of species have
been recorded in the genus Prosorhynchus Odhner, 1905, followed by Bucephalus
Baer, 1827. The validity of some of bucephalid species remains equivocal since the
descriptions provided are inadequate and the figures lack details.
Diagnosis: Body spinose, anterior end with solid or sucker-like rhynchus, may or
may not bear tentacles. Oral and ventral suckers absent. Mouth on ventral surface,
at about midbody. Pharynx and oesophagus associated with mouth. Intestinal cae-
cum simple, sac-like or elongate. Testes two. Genital pore terminal, near posterior
end of body. Cirrus-sac conspicuous, located posteriorly, contains seminal vesicle,
pars prostatica, ejaculatory duct leading into a sac-like genital atrium enclosing a
muscular genital lobe. Ovary single, spherical, pre or intertesticular. Uterus occupy-
ing much of body, distal portion opening into genital atrium. Eggs oval. Vitellarium
follicular. Excretory vesicle I-shaped, pore terminal, anterior extent variable. Type
genus: Bucephalus Baer, 1827.
© Crown 2018 35
https://doi.org/10.1007/978-94-024-1535-3_6
36 6 Superfamily Bucephaloidea Poche, 1907
Key to subfamilies of Indian genera of bucephalidae

1a. Anterior end usually with sucker-like rhynchus, unmodified or modified, with
lobes, tentacles or hood ……………………………..……………………….
Bucephalinae Poche, 1971
1b. Anterior end with solid rhynchus, a muscular cone topped by simple mus-
cular disc or with lobes or circles of long or short spines …………
Prosorhynchinae Nicoll, 1914
Subfamily Bucephalinae Poche, 1907
Diagnosis: Body cylindrical, spined. Rhynchus consists of simple sucker with or

without dorsal hood, lobes or retractable tentacles. Caecum saccular, at midbody.
Pharynx and oesphagus present. Testes 2, oval to subglobular. Cirrus-sac well devel-
oped, encloses seminal vesicle, straight pars prostatica and thin-walled ejaculatory
duct, opening into genital atrium. Genital atrium a thin-walled sac enclosing geni-
tal lobe with terminal genital pore. Uterus coils throughout body. Vitelline follicles
restricted to lateral fields of body. Excretory vesicle I-shaped.
Key to Indian genera of Bucephalinae

1. Rhynchus with tentacles…………………………………………………….… 2
Rhynchus without tentacles…………………………………………….……… 3
2. Rhynchus funnel-shaped, bears 7 or more tentacles ....………...... Alcicornis
MacCallum, 1917
Rhynchus sucker-like, usually with 7 tentacles……………….…. Bucephalus
Baer, 1827
3. Rhynchus a simple sucker without hood ….…………...… Prosorhynchoides
Dollfus, 1929
Rhynchus a simple sucker covered by a muscular hood, simple or with
fleshy lobes ……………………………………………………... Rhipidocotyle
Diesing, 1858
Genus Alcicornis MacCallum 1917
Genus Alcicornis was erected by MacCallum (1917) with A. carangis MacCallum,
1917, from Caranx ruber from Florida as the type species. It differs from the closely
related genus Bucephalus Baer, 1827, in the shape of anterior rhynchus, which is
funnel-shaped in Alcicornis in contrast to the rounded sucker in Bucephalus.
Diagnosis: Body elongate. Rhynchus either funnel-shaped or short U-shaped struc-

ture, crowned by ring of 7–21 long fleshy retractable tentacles. Mouth near midbody.
Gut sac-like directed anteriorly or posteriorly from pharynx. Testes oblique. Cirrus-
sac well developed. Seminal vesicle spherical or ovoid. Pars prostatica long. Ovary
pretesticular. Uterus extensive. Vitelline follicles in anterior lateral rows. Excretory
vesicle long. Type species: A. carangis MacCallum, 1917.
Subfamily Bucephalinae Poche, 1907 37
Nine species belonging to this genus have been recorded from marine fishes of India.
Of these, A. jairajpuri Gupta & Gupta, 1989, in the shape of the rhynchus and the
general organization of the body fits more appropriately into the genus Bucephalus.
However, more specimens of this species need to be examined to determine its exact
status. It is here regarded as a taxon inquirendum. Two more species A. inglisi Gupta
& Jain, 1993 and A. rauschi Gupta & Jain, 1993 are also considered here as species
inquirendae since the description of these species based on macerated specimens are
inadequate.
Alcicornis carangis MacCallum, 1917 (Fig. 6.1)
Hosts: Carangidae: Carangoides malabaricus (Bl. & Schn.); C. chrysophrys (Cuvier)
Locality: VSK (BOB)
Geographic distribution: Florida, Cuba, Curacao, Bermuda.
Reference: Madhavi (1974b)
Description: Body elongate 1580–2100 long, 300–400 wide. Rhynchus wedge-

shaped, prominent, 566–633 in width, tentacles 7, vary in size and shape, each with
2 lateral prongs and terminal filament, proximal prong longer than distal one. Mouth
at level of ovary. Caecum mostly anterior to testis. Testes tandem, contiguous. Cirrus-
sac extends to mid-level of posterior testis, encloses oval seminal vesicle, long pars
prostatica and prostatic cells. Ovary entire, anterior to testis, uterus voluminous,
extends from rhynchus to posterior end of body. Genital atrium wide, genital pore at
some distance from posterior end. Vitellarium in two lateral groups of 12–17 follicles
each, mostly in middle third of body commence at level of testes. Eggs 18–22 by
13–15.
Fig. 6.1 Alcicornis carangis

(after Madhavi 1974)
Remarks: This species was first described by MacCallum (1917) as the type of
the genus Alcicornis from Caranx ruber of Florida. Perez-Vigueras (1955) gave a
more detailed description of the species on the basis of specimens collected from
Cuba. Nahhas & Cable (1964) collected the species from carangids off Curacao and
Jamaica, and later, Rees (1970) recorded the species from the coast of Bermuda. There
were also reports of A. carangis from carangids off Puerto Rico (Dyer et al., 1997).
From Bay of Bengal, it was recorded from the carangids Carangoides malabaricus
and C. chrysophrys (Madhavi, 1974b). Bray & Justine (2010) examined some of the
specimens from C. malabaricus from New Caledonia and concluded that they had
some features distinguishing them from the Atlantic form described by MacCallum
and others.
Alcicornis hemiramphi Gupta & Tandon, 1983 (Fig. 6.2)

Hosts: Hemiramphidae: Hemiramphus brachynotopterus Bleeker
Locality: PR (BOB)
Number of specimens: One
Reference: Gupta, S.P. & Tandon (1983a)
Description: Body elongate, slender, 1380 long 125 wide. Rhynchus oval, 170 by
169, with seven short tentacles, each with two lateral processes. Mouth ventral, poste-
quatorial, close to ovary, 1000 from anterior extremity. Pharynx spherical. Oesopha-
gus short. Caecum saccular, anteriorly directed, extends up to just posterior to middle
of body. Excretory bladder tubular extends up to posterior end of rhynchus. Genital
pore subterminal. Testes oval, tandem, subequal, close together, in posterior half of
body. Cirrus-sac elongated, extends up to posterior end of anterior testis. Genital
atrium spacious, encloses genital lobes. Ovary ovoid, pretesticular, partially overlap-
ping anterior testis. Uterus extends anteriorly to anterior level of vitellarium and then
Fig. 6.2 Alcicornis

hemiramphi (after Gupta &
Tandon, 1983)
extends posteriorly to open into genital atrium. Eggs 20–21 by 18–20. Vitellarium
follicular, in two lateral rows, 16 on right side and 17 on left side extends from just
anterior to midbody to middle of anterior testis.
Remarks: This species was differentiated from all the known members of the genus
in the extent of the vitellarium from anterior to the middle of the body to the middle
of the anterior testis. However, more specimens of this species collected from the
type host and type locality have to be examined to determine its validity.
Alcicornis indicus Gupta & Sharma, 1972 (Fig. 6.3)
Host: Haemulidae: Pomadasys maculatus Gunther
Locality: RT (AS)
Number: 62
Reference: Gupta, A.N. & Sharma (1972a)
Description: Body elongate, narrow, 3150–5020 long, 330–370 wide at testicular

level. Rhynchus wedge-shaped, 120–165 by 90–150, with seven tentacles, each tenta-
cle possesses two prongs and terminal filament. Pharynx spherical, situated between
ovary and posterior extent of vitelline fields. Intestine tube-like extends to vitelline
level. Testes spherical, postovarian, tandem, separated by uterine coils. Cirrus-sac
long, tubular, thick-walled reaches to about posterior testis, encloses ovoid semi-
nal vesicle, long pars prostatica, prostatic cells, weakly developed ejaculatory duct.
Genital atrium spacious. Ovary entire, spherical, pretesticular, close to anterior testis.
Uterine coils extend anteriorly as far as halfway between anterior extent of vitelline
follicles and rhynchus. Vitelline follicles scanty, arranged in two groups laterally,
in linear fashion, 11–17 follicles on left, 9–16 on right side, anterior extension just
equatorial, posterior extension near to ovarian level. Eggs 15–21 by 7.
Remarks: This species was differentiated from A. carangis in the larger body size
and the smaller size of the rhynchus and the eggs.
Fig. 6.3 Alcicornis indicus

(after Gupta & Sharma,
1972)
Alicornis inglisi Gupta & Jain, 1993

Host: Carangidae: Alepes djedaba (Forskal) (= Caranx kalla)
Locality: PR (BOB)
No. 1
Reference: Gupta, V & Jain (1993)
Remarks: The description of the species is based on a single specimen. From the
figure provided for the species, it is evident that it is in a decomposed state. Hence,
it is considered here as a species inquirendum.
Alcicornis jairajpuri Gupta & Gupta, 1989
Host: Trichiuridae: Eupleurogrammus muticus (Gray)
Locality: PR (BOB)
Number: 2
Reference: Gupta, P.C. & Gupta, V.C. (1989)
Remarks: This species is here considered as taxon inquirendum.

Alcicornis multidactylus Madhavi, 1974 (Figs. 6.4 and 6.5)
Host: Caesionidae: Caesio caerulaurea Lacepede
Locality: VSK (BOB)
Fig. 6.4 Alcicornis

multidactylus entire (after
Madhavi, 1974)
Fig. 6.5 Alcicornis

multidactylus, anterior
extremity
Description: Body long, slender, 2240–2640 long by 192–288 wide. Tegument

entirely spined. Rhynchus pad-like, expanded anteriorly into fan-like structure,
148–175 by 94–175, muscles in anterior part of rhynchus arranged in regular rows
giving ribbed appearance. Tentacles 15, usually retracted, each tentacle short, with-
out lateral processes, those at corners elongated and horn-like. Mouth opening to left
of anterior testis or ovary. Pharynx spherical. Oesophagus long. Caecum long and
sac-like, directed anteriorly, median, preovarian. Testes smooth, elliptical, tandem,
contiguous, situated in posterior third of body. Cirrus-sac extends to level of anterior
testis, encloses crescent-shaped seminal vesicle, long pars prostatica joins seminal
vesicle laterally. Prostatic cells numerous. Genital atrium spacious. Ovary spherical,
pretesticular, separated from anterior testis by uterine coils, situated at junction of
middle and posterior thirds of body. Uterus with narrow coils, extends anteriorly
slightly beyond level of vitellarium. Eggs 27–30 by 16–17. Vitelline follicles large,
discrete, arranged in two lateral rows extend from 384 to 992 posterior to rhynchus
to level of anterior end of caecum on right side and to level of ovary on left side,
consist of 34 follicles. Excretory bladder I-shaped extends to level of rhynchus.
Remarks: This species is characterized by the presence of 15 tentacles on the

rhynchus, the mouth at the level of the ovary or testes, an anteriorly directed gut, the
elliptical testes in the posterior third of the body, a cirrus-sac extending to the level of
the anterior testis, asymmetrically distributed vitelline follicles and the uterine coils
extending anteriorly beyond the vitellarium.
Alcicornis rauschi Gupta & Jain, 1993

Host: Carangidae: Caranx sexfasciatus Quoy & Gaimard
Locality: PR (BOB)
Reference: Gupta, V & Jain (1993).
No. 5
Remarks: The description of the species is based on five macerated specimens.

Hence, it is considered here as a species inquirendum.
Alcicornis scomberi Gupta & Ahmad, 1976 (Fig. 6.6)

Host: Scombridae: Rastrelliger kanagurta (Cuvier)
Locality: PR (BOB)
No. One
Reference: Gupta V. & Ahmad (1976a)
Description: Body elongate, slender, slightly tapering anteriorly, rounded poste-

riorly, 3520 long, 420 wide. Rhynchus oval, wedge-shaped, 170 long, 175 wide,
with six short tentacles, without lateral processes. Mouth ventral, post-equatorial,
overlapping ovary. Pharynx spherical. Oesophagus short. Caecum saccular, directed
anteriorly as far as middle of body. Genital pore subterminal just anterior to posterior
end of body. Testes oval, tandem, subequal, close together, in middle of posterior
one-third of body. Cirrus-sac elongate, extends to posterior third of anterior testis.
Genital atrium spacious, encloses genital lobes. Ovary spherical, pretesticular, sepa-
rated from anterior testis by uterine coils. Uterus runs anteriorly up to some distance
anterior to level of vitellarium and then runs posteriorly to open into genital atrium.
Eggs oval, operculated, 15–24 long. 13–19 wide. Vitellarium follicular, in two lateral
rows, 15 on right side and 14 on left side, extend posteriorly up to middle of ovary.
Fig. 6.6 Alcicornis

scomberi (after Gupta &
Ahmad, 1976)
Remarks: The distinguishing features of this species are the occurrence of six tenta-
cles without lateral processes on the rhynchus, the extension of the vitellarium to the
middle of the ovary and the extension of the uterus to a significant distance anterior
to the level of the vitellarium.
Alcicornis thapari Hafeezullah & Siddiqi, 1970

Host: Carangidae: Caranx sexfasciatus Quoy & Gaimard (1); Alepis djedaba Forskal
(= Caranx kalla) (2)
Locality: Tuticorin (AS) (1); PR (BOB) (2)
Number: 4
Reference: 1. Hafeezullah & Siddiqi (1970a), 2. Gupta, V. & Jain (1993)
Description: Body elongate, slightly tapering anteriorly, rounded posteriorly,

2822–3670 wide. Rhynchus 105–122 long 108–116 wide, wedge-shaped, provided
with 7 short tentacles, Mouth postequatorial, at about junction of middle and poste-
rior one third of body. Pharynx 78–102 in diameter. Caecum saccular, short, directed
anteriorly. Testes tandem, situated in middle of posterior one third of body, to right of
midline. Cirrus-sac cylindrical, in posterior part of body, not quite reaching posterior
testis, contains saccular seminal vesicle and long pars prostatica surrounded by pro-
static cells. Genital atriumquite wide. Genital pore ventral, just anterior to posterior
end. Ovary globular, pretesticular, contiguous with anterior testis. Uterine seminal
receptacle present. Vitellarium follicular, in two lateral groups in middle one third of
body. Uterus reaches posteriorly behind genital atrium and anteriorly to anterior level
of vitellarium. Eggs 12–22 × 7–15. Excretory vesicle reaches anteiorly to middle of
body.
Remarks: This species is distinguished from all the species of Alcicornis by the fol-
lowing combination of characters: the wedge-shaped rhynchus with seven tentacles,
the distribution of vitelline follicles along lateral fields in the middle third of the
body, and the extent of the cirrus-sac with respect to the position of the testes.
Key to Indian species of Alcicornis

1. Tentacles fifteen…………………………………………….. A. multidactylus
Madhavi, 1974
Tentacles fewer than fifteen…………………………………………..…………2
2. Tentacles six…………………………………………………A. scomberi Gupta
& Ahmad, 1976
Tentacles seven …………………………………………………….…………. 3
3. Uterus extends to anterior level of vitellarium ……………………….………. 4
Uterus extends beyond anterior level of vitellarium ……………………………5
4. Cirrus-sac not reaching posterior testis ……….…….. A. thapari Hafeezullah &
Siddiqi, 1970
Cirrus-sac overlapping posterior testis ……………… A. hemiramphi Gupta &
Tandon, 1983
5. Rhynchus large, prominent, triangular…………….……………A. carangi

MacCallum, 1917
Rhynchus smaller, wedge-shaped ………………..……….. A. indicus Gupta &
Sharma, 1972
Genus Bucephalus Baer, 1827
[Syn: Eubucephalus Diesing, 1855; Gasterostomum von Siebold, 1848; Labratrema
Maillard, 1975]
Diagnosis: Body fusiform or elongate. Rhynchus sucker-like, usually with seven
retractable tentacles. Mouth near midbody, leads into pharynx. Caecum sac-
like, directed anteriorlyor posteriorly. Testes two, oblique. Cirrus-sac thin-walled,
encloses spherical or ovoid seminal vesicle, long straight pars prostatica and numer-
ous prostatic cells. Ejaculatory duct opens into genital atrium in close association
with protrusible genital lobe. Ovary pretesticular. Uterine coils extensive may or may
not extend anterior to vitelline zone. Vitelline follicles in two lateral fields anterior
to ovary. Excretory vesicle I-shaped, variable in length.
From the Indian region, eleven species of Bucephalus have been recorded from
marine fishes. Literature on morphology and relationships could be obtained for all
the species excepting B. arabiana Varma, 1982.
Bucephalus arabiana Varma, 1982
Host: Carangidae: Carangoides armatus (Ruppell)
Locality: Arabian Sea
Reference: Varma (1982a)
Bucephalus arabiansis Dwivedi, 2007 (Fig. 6.7)

Host: Carangidae: Alepes melanoptera (Swainson, 1839) Caranx nigripinnis Day
Locality: Cochin coast, Kerala, AS
Number: 4
Reference: Dwivedi (2007)
Fig. 6.7 Bucephalus

arabiansis (after Dwivedi,
2007)
Description: Body elongated 1020–1530 long, 170–540 wide. Tegument aspinose.

Rhynchus sucker-like, 160–280 in diameter, with five tentacles, tentacles without lat-
eral processes. Mouth at middle of body at level of anterior testis or at anterior margin
of posterior testis. Pharynx spherical. Oesophagus short. Caecum saccular, lateral to
ovary. Testes subequal, tandem, postequatorial, contiguous. Cirrus-sac elongated,
extends to posterior margin of anterior testis. Seminal vesicle oval. Pars prostatica
long. Ejaculatory duct narrow. Genital pore subterminal. Ovary spherical or sub-
spherical, entire, pretesticular, overlapping anterior testis. Uterus extends anteriorly
beyond anterior level of vitelline zone. Eggs 20–30 by 10–30 (?). Vitellarium two
lateral rows of rounded follicles, preovarian, equatorial, 10–15 on right side and
10–11 on left side.
Remarks: This species is characterized by the aspinose body, the presence of five
tentacles on the rhynchus, overlapping testes and the ovary situated laterally to the gut.
In having five tentacles on the rhynchus, this species resembles B. barina Srivastava,
1938 and B. yamagutii Gupta & Singh, 1985, but differs from B. barina in having
an aspinose body, tentacles lacking processes and in the overlapping testes and from
B. yamagutii in the extension of the uterus anteriorly beyond the vitelline follicles.
It is likely that the lack of tegumental spines is the result of loss during fixation or
processing.
Bucephalus barina Srivastava, 1938 (Fig. 6.8)

Hosts: Scatophagidae: Scatophagus argus (Linnaeus); (1) Sciaenidae: Johnius dus-
sumieri (Cuvier) (= J. sina), J. carutta Bloch, J. aneus (Bloch), J. belengeri (Cuvier),
Nibea soldado (Lacepede) (2)
Number: Numerous
Fig. 6.8 Bucephalus barina

(after Srivastava, 1938)
Locality: Puri (1), VSK (2), (BOB)

Distribution: India, China
Reference: 1. Srivastava (1938a); 2. Madhavi (1974b)
Description: Body 1520–2800 long, 50–68 wide. Tegument covered with fine back-
wardly directed spines. Rhynchus 140–200 by 120–220, anterodorsal surface bears
five tentacles, each tentacle armed with rose thorn-like processes, arranged radially
around basal half. Mouth at level of testes. Pharynx small transversely oval. Oesoph-
agus narrow. Caecum small, sac-like, directed anteriorly, lies at level of anterior
testis. Testes two, transversely oval, lie in contact, in tandem, closely posterior to
anterior half of body. Cirrus-sac extends anteriorly as far as level of anterior margin
of posterior testis. Seminal vesicle small, oval. Pars prostatica long, prostatic cells
present. Ejaculatory duct short, opens at base of well developed genital lobe, extends
into genital atrium. Genital pore subterminal. Ovary pretesticular, at middle of body,
separated from testis by Mehlis’ gland. Uterus well developed, extends just ante-
riorly to vitellarium. Eggs 15–19 by 9.5–11. Vitellarium consists of small follicles
arranged roughly in pairs along lateral sides of body, extend from level of ovary to
first sixth of body. Excretory bladder extends to anterior limit of vitellarium.
Remarks: Srivastava (1938a) reported B. barina from Scatophagus argus from off
the Puri coast. From Visakhapatnam coast, it was recorded from several species of
sciaenid fishes (Madhavi, 1974b). This species is characterized by the presence of
five tentacles, each tentacle provided with a number of hooks arranged radially at
the base.
Bucephalus elacatus Yadav, 1977 (Fig. 6.9)
Host: Rachycentridae: Rachycentron canadum (Linnaeus) (= Elacatanigra (Bloch))
Locality: Ratnagiri coast, Arabian Sea
No. 4
Reference: Yadav (1977)
Description: Body elongate 4160–5120 × 620–680. Rhynchus sucker-like, 130 by

180, with four tentacles. Mouth and gut located in posterior half of body. Caecum
situated just anterior to ovary. Testes diagonal, situated in posterior third of body.
Cirrus-sac extends to level of anterior margin of posterior testis. Ovary pretesticular,
postequatorial. Uterus passes between testes and extends to anterior third of body.
Eggs 24–25 b 9–11. Vitellarium distributed in lateral fields in middle third of body.
Remarks: According to Yadav (1977), this species closely resembles B. gangeticus

Srivastava, 1938, but differs from it in the position of the pharynx and testes, the
extent of the cirrus-sac and the larger eggs. The figure given for the species indi-
cates that it may not belong to the genus Bucephalus, since the rhynchus lacks the
tentacles characteristic of Bucephalus. The structure of the rhynchus on the other
hand indicates that it may be a species of Rhipidocotyle. We consider it as a taxon
inquirendum.
Fig. 6.9 Bucephalus

elacatus (after Yadav, 1977)
Bucephalus fischthali Gupta & Tewari, 1983 (Fig. 6.10)

Host: Terapontidae: Terapon theraps Cuvier
Locality: Madras coast, Bay of Bengal
No: 2
Reference: Gupta, V. & Tewari (1983a)
Description: Body elongated, aspinose, 2520–4000 long, 670–740 wide. Rhynchus
spherical, 210–240 in diameter, with 6 tentacles, without lateral processes. Mouth
opening at posterior third of body. Pharynx ovoid, post-equatorial. Oesophagus tubu-
lar. Caecum simple, ovoid, lateral to anterior testis. Genital pore at posterior end
of body. Testes subspherical, tandem, subequal, post-equatorial. Cirrus-sac tubular
extends from posterior end to level of caecum. Seminal vesicle elongated. Ejaculatory
duct long opens into spherical genital sinus. Prostate glands numerous, surround pars
prostatica. Ovary entire, spherical, overlaps anterior testis. Uterus extends anteriorly
as far as vitellarium, and then turns towards posterior extremity to open at genital
atrium. Eggs spherical, 32–35 in diameter. Vitelline follicles extend from anterior
end of ovary to anterior half of body.
Remarks: In having six tentacles and in the anterior extent of cirrus-sac up to the
anterior limit of the posterior testis, this species resembles B. jagannathai Verma,
1938, but differs from it in the position of the caecum at the level of the anterior testis
and in the absence of short processes at the anterior end of the tentacles.
Fig. 6.10 Bucephalus

fischthali (after Gupta &
Tewari, 1983)
Bucephalus jagannathai Verma, 1936 (Fig. 6.11)

Host: Scombridae: Scomberomorus guttatus (Bl. & Schn.) (= Cybium guttatum)
No. 12
Locality: PR, BOB
Reference: Verma (1936)
Description: Body 1100–1700 long, 420–540 wide in region of ovary. Rhynchus

shallow, ventroterminal, scoop-shaped, 106–193 by 110–170, with six tentacles,
each with short lateral process in middle. Pharynx equatorial, overlapped by ovary.
Caecum sac-like, short. Gonads packed together, overlapping one another, mostly
in third-fourth of body. Testes spherical, posterior testis smaller or nearly equal to
anterior. Cirrus-sac sickle-shaped, one third as long as body, reaches to level of

jagannathai (after Verma,
1936)
anterior margin of anterior testis. Ovary lateral to caecum, near commencement of

third quarter of body. Uterine coils extend from near genital atrium to anterior level
of vitellarium. Vitelline fields reach to different levels in second quarter of body,
follicles large, rounded, 14–15 on the ovarian side 16–17 on the other. Genital atrium
large. Genital lobe well developed, muscular, protrusible beyond genital pore. Gen-
ital pore ventro-terminal leads into atrium by short sinus. Excretory bladder tubular
extends to midway between vitellarium and rhynchus. Eggs 18.6–20 by 11.6–13.3.
Remarks: Distinguishing features of this species are the presence of six tentacles,
each provided with a short lateral process, the long cirrus-sac reaching to the level
of the anterior margin of the anterior testis and the uterus not extending anterior to
the vitelline zone
Bucephalus kanagurtai Gupta & Tewari, 1983

Locality: MS (BOB)
Number: 2
Reference: Gupta, V. & Tewari (1983a)
Description: Body elongated, 3200–3570 long, 820–950 wide. Tegument aspinose.

Rhynchus spherical, 240–260 long 260–280 wide, with 7 tentacles, without lateral
processes. Mouth opening at posterior third of body. Pharynx ovoid, post-equatorial.
Oesophagus tubular. Caecum simple, ovoid. Genital pore at posterior end of body.
Testes subspherical, tandem, subequal, postequatorial. Posterior testis smaller than
anterior. Cirrus-sac tubular extends from midlevel of hind testis to posterior end of
body. Seminal vesicle elongated. Pars prostatica elongated. Ejaculatory duct small,
opens into a spherical genital sinus. Prostate glands numerous, surround pars pro-
statica. Ovary entire, spherical, overlaps anterior testis. Uterus extends anteriorly up
to vitellarium. Vitelline follicles extend from anterior end of body to level of anterior
margin of ovary. Eggs spherical, 40–70 in diameter.
Remarks: This species is characterized by the extent of the cirrus-sac to the midlevel
of the posterior testis and the presence of seven tentacles without lateral processes.
The egg size given for the species is unusual and needs verification.
Bucephalus margaritae Ozaki & Ishibashi, 1934 (Fig. 6.12a)

[Syn: Bucephalus carangoides Yamaguti, 1970; Bucephalus pseudovaricus
Velasquez, 1959;’ Bucephalus retractilis Yamaguti, 1952; Bucephalus ulua Yam-
aguti, 1970. Bucephalus varicus Manter, 1940. B. polymorphus of Gupta & Mehrotra
(1970b)].
Hosts: Terapontidae: Terapon jarbua (Forskål); (1, 2), Haemulidae: Pomadasys mac-
ulatus Bloch (1); Carangidae: Caranx sexfasciatus Quoy & Gaimard; Decapterus
russelli (Ruppell); Carangoides malabaricus Bloch; C. chrysophrys (Cuv.) (2);
Polynemidae: Polynemus plebeius (Broussonet)
Locality: Ernakulam, AS (1); VSK (BOB) (2)
Number: Numerous
Fig. 6.12 a Bucephalus (a) (b)

margaritae b Bucephalus
thapari (after Gupta &
Tewari, 1983)
Geographic distribution: Widely distributed in tropical and subtropical waters:

Caribbean Sea, Pacific coast of Panama, Mexico, Brazil, Venezuela, Guinea-Bissau,
China, Mediterranean coast of Israel, Kuwait, Red Sea, Arabian Sea, Hawaii, India,
Santa Catarina coast (Amato, 1982; Takemoto et. al., 1995, 1996; Pereira Junior et al.
(1996); Luque et al. (2000); Nahhas et al., (2006) Luque et Alves (2001), Alves et al.,
(2004); Marchier et al. (2010) Luque et al. (2001).
Reference: 1. Gupta, N.K. & Mehrotra (1970b); 2. Madhavi (1974b)
Description: Body small, elongate with spined tegument, 618–2288 long. Rhynchus
truncate, 68–168 in diameter, bears 7 tentacles arranged around dorsal side of ante-
rior edge; tentacles broad at base, each with two processes, larger and tongue-shaped
in middle, and smaller and knob-like at base. Caecum sac-like, directed anteriorly
or posteriorly. Testes spherical, tandem, contiguous, subequal situated in posterior
half of body towards right side. Cirrus-sac elongate encloses ovoid seminal vesicle,
long pars prostatica surrounded by numerous prostatic cells, opens posteriorly into
genital atrium. Ovary spherical, smooth, smaller than testes, pretesticular, contigu-
ous with anterior testis. Uterine coils numerous, extend anteriorly to anterior limit
of vitelline follicles, open posteriorly into genital atrium. Eggs 21–27 by 13–23.
Vitelline follicles arranged in two longitudinal lateral groups one on each side situ-
ated in pretesticular region, number of follicles on each side varies from 19 to 24.
Excretory bladder tubular extends almost to level of anterior sucker.
Remarks: According to Nahhas et al. (2006), the most important diagnostic feature
for B. margaritae is the presence of seven tentacles, each bearing two projections, one
big and basal and the other small and distal. Bray (1984) provided a historical review
of B. margaritae. B. margaritae is now known from a wide range of geographical
areas and hosts. Carangids are the most common hosts. It has also been recorded
from many non-carangid hosts. From the Indian region, Madhavi (1974b) recorded
it from both carangid and non-carangid hosts and considered B. polymorphus of
Gupta & Mehrotra (1970) from Terapon jarbua as a synonym of this species. The
known hosts for B. margaritae including carangid hosts from different geographical
localities are listed by Marchiori et al. (2010). They also demonstrated the life cycle
of the species from the coast of Santa Catarina State, Brazil, with the mussel Perna
and the blennid Hypopleurochilus fissiconis as intermediate hosts. It is likely that

this is a complex of cryptic species.
Bucephalus thapari Gupta & Tewari, 1983 (Fig. 6.12b)

Host: Trichiuridae: Eupleurogrammus muticus (= Trichiurus muticus) (Griffith)
Locality: MS (BOB)
Number: 3
Reference: Gupta, V & Tewari (1983a)
Description: Body elongate, 3470–3850 long, 900–950 wide in region of ovary.

Tegument aspinose. Rhynchus spherical, 250–300 in diameter, with 4 tentacles,
without lateral processes. Mouth opening at posterior third of body. Pharynx ovoid,
post-equatorial. Oesophagus tubular. Caecum simple, ovoid. Genital pore at poste-
rior end of body. Testes subspherical, tandem, subequal, postequatorial. Cirrus-sac
tubular extends from midlevel of anterior testis to posterior end of body. Seminal
vesicle elongated. Pars prostatica elongate. Ejaculatory duct opens into a spherical
genital sinus. Ovary entire, spherical, equatorial or postequatorial, separated from
anterior testis. Uterus extends anteriorly as far as anterior extent of vitellarium. Eggs
spherical, 16–18 in diameter. Vitelline follicles extend from about midway between
rhynchus and mouth to level of ovary.
Remarks: Characterised by the presence of 4 tentacles, the extent of the cirrus-sac to

the midlevel of the anterior testis and by the separation of the ovary from the anterior
testis.
Bucephalus uranoscopi Yamaguti, 1934 (Fig. 6.13)
Host: Uranoscopidae: Uranoscopus guttatus Cuvier
Location: Intestine
Locality: VSK (BOB)

uranoscopi (after Yamaguti,
1934)
Number: 15
Geographic distribution: Pacific coast off Japan, Bay of Bengal
Reference: Madhavi (1974b); Yamaguti (1934)
Description: Body long, slender, 4860 long, 450 broad. Tegument spined through-
out body. Rhynchus scoop-shaped, 150 by 125, bears 7 tentacles, each with small
distal and larger proximal protuberance. Mouth opening in posterior part of middle
third of body. Pharynx globular. Oesophagus as long as pharynx, directed anterodor-
sally. Caecum arcuate. Testes oval, separated, at beginning of posterior third of body.
Cirrus-sac in posterior third of body. Seminal vesicle saccular. Pars prostatic and pro-
static cells well developed. Genital atrium opening ventroterminally. Ovary situated
near caecum. Uterus extends as far anteriorly as anterior limit of vitellarium. Eggs
18 by 12. Vitelline follicles large, 70–90 in diameter, distributed in lateral fields in
anterior third of body. Excretory vesicle extends further forwards than anterior extent
of vitellarium.
Remarks: This species is characterized by the long slender body bearing seven ten-
tacles anteriorly, each tentacle provided with two processes: the widely separated
gonads and the short cirrus-sac. The original description given by Yamaguti (1934)
is based on a single specimen collected from Uranoscopus japonicas from Japan.
Madhavi (1974b) collected 15 specimens of the species from Uranoscopus guttatus
from Bay of Bengal.
Bucephalus yamagutii Gupta & Singh, 1985 (Fig. 6.14)

Host: Carangidae: Carangoides malabaricus (Bl. & Schn.) (= Caranx malabaricus)
Locality: PR (BOB)
Number: 3
Reference: Gupta, P.C. & Singh (1985a)

yamagutii (after Gupta &
Singh, 1985)
Description: Body elongate, 1460–1570 by 230–250 in size. Tegument spinose.

Rhynchus terminal, subspherical, 130–140 by 100–130, with five tentacles, provided
with lateral processes, having broad base and tapering terminal parts. Mouth poste-
quatorial, at level of testes. Pharynx ovoid, lateral to midlevel of testes. Oesophagus
small. Caecum elongate, saccular, extends anterior to ovary. Testes entire, subspher-
ical, tandem, close together, in posterior half of body. Cirrus-sac elongate, slender,
extends anteriorly as far as midlevel of posterior testis. Seminal vesicle oval. Pars
prostatica tubular. Ejaculatory duct narrow, tubular, thick-walled. Genital pore sub-
terminal. Ovary pretesticular overlaps anterior border of testis. Uterus ascends as far
as anterior limit of vitelline follicles. Eggs 15–18 by 9–11. Vitellarium follicular,
oval or rounded, in two lateral rows, 15–16 on right side, 13–16 on left side, extend
from about anterior third of body length to slightly anterior to ovary
Remarks: This species resembles B. barina in having five tentacles but differs from
it in having uterine coils extending anteriorly to the level of the vitelline follicles
instead of anterior to it, the ovary overlapping the anterior border of the anterior
testis, and the caecum sac longer and extending beyond the anterior margin of the
ovary.
Key to species of Bucephalus from Indian marine fishes*
1. Tentacles four………………………………………………………………….. 2
Tentacles more than four ……………………………………………………… 3
2. Caecum at level of anterior testis ……………..……………. B. thapari Gupta &
Tewari, 1983
Gut anterior to ovary ………………………………………..……… B. elacatus
Yadav, 1977
3. Tentacles five…..…………………………………….….…………..………….4
Tentacles more than five ……………………………………………………….6
4. Tentacles with rose thorn like structures at base……………… B. barina
Srivastava, 1938
Tentacles without rose thorn like structure…………………………....…………5
5. Uterus extends anterior to vitellarium, testes overlapping ……. B. arabiansis
Dwivedi, 2007
Uterus extends anteriorly as far as level of vitellarium, testes not overlapping
……………………….………………… B. yamagutii Gupta & Singh, 1985
6. Tentacles six ................................................................................................…. 7
Tentacles seven ………………………………………………………………..8
7. Tentacles with short lateral processes................................................……
B. jagannathai Verma, 1936
Tentacles without lateral processes ………………..……B. fischthali Gupta &
Tewari, 1983
8. Body elongate ……………………………………..……… B. uranoscopi
Yamaguti, 1934
Body elliptical or oval ………………………………………………………… 9
9. Tentacles without lateral processes, cirrus-sac short ….. B. kanagurtai Gupta &
Tewari, 1983
Tentacles with lateral processes, cirrus-sac fairly long….. B. margaritae Ozaki
& Ishibashi, 1934
*Information not available for B. arabiana
Genus Prosorhynchoides Dollfus, 1928

[Syn: Bucephalopsis Diesing 1855; Bucephaloides Hopkins, 1954; Neobucephalop-
sis Dayal, 1948]
Members of this genus have been referred to under the name Bucephaloides Hop-
kins, 1954, for a long time, but Srivastava & Chauhan (1973) suggested that the name
Prosorhynchoides is appropriate to the genus. Overstreet & Curran (2002) and Bott &
Cribb (2005) provided extensive reviews on the genus. Three genera Bucephalopsis
Diesing, 1855, Bucephaloides Hopkins, 1954, and Neobucephalopsis Dayal, 1948,
were regarded as synonyms of Prosorhynchoides by Overstreet & Curran (2002). All
the species with a simple sucker-like rhynchus and a pretesticular ovary are trans-
ferred to the genus Prosorhynchoides which now contains 63 nominal species (Bott
& Cribb, 2005).
Diagnosis: Anterior attachment organ a simple sucker without tentacles. Mouth
usually in middle third of body. Caecum saccular to tubular. Testes tandem or
oblique, near midbody or in posterior half of body. Cirrus-sac well developed. Ovary
pretesticular or at level of testis. Uterus normally reaches anterior to mouth. Genital
atrium contains fleshy lobes. Vitelline follicles in symmetrical lateral fields, mainly
or entirely preovarian. Type species: P. ovatus (Linton, 1900)
In India the genus is represented by six valid species in marine fishes and several
species in freshwater fishes.
Prosorhynchoides belonea (Srivastava, 1938) Srivastava & Chauhan, 1973
[Syn. Bucephalopsis belonea Srivastava, 1938]
Host: Belonidae: Strongylura strongylura V. Hasselt.
Locality: Allahabad (India)
Reference: Srivastava (1938a)
Description: Body pyriform, 1680–2800 long, 820–1200 wide. Anterior sucker
large, 300–400 in diameter. Pharynx circular, situated in median line at junction of
middle and anterior third of body. Oesophagus straight narrow tube. Caecum sac-
shaped, situated in median line at level of anterior testis, directed anteriorly. Testes
large, oblique, in middle third of body. Cirrus-sac well developed, extends as far
as level of anterior end of oesophagus, encloses oval seminal vesicle, long pars
prostatica and narrow ejaculatory duct. Genital tongue extends into funnel-shaped
genital sinus. Genital pore just anterior to posterior end. Ovary pyriform, pretesticular.
Uterus extends anteriorly as far as anterior limit of vitellarium. Eggs 34–36 × 11–13.
Vitelline follicles large, rounded, arranged longitudinally in pairs on lateral sides of
body, extending from level of middle of ovary to level of anterior sucker.
Remarks: This species closely resembles P. karvei Bhalerao, 1937, in the shape of
the body, the position of the genital pore and the extent of the vitellarium but differs
from it in being twice its size and the anterior extent of the cirrus-sac which reaches
to the posterior level of the anterior testis.
Prosorhynchoides brayi (Ahmad, 1987) Madhavi, 2011 (Fig. 6.15)

[Syn: Bucephaloides brayi Ahmad, 1987]
No: 19
Locality: Goa (AS)
Reference: Ahmad (1987b)
Description: Body elongate, narrow, 5319–7730 long, 510–830 wide. Tegument

spined. Anterior sucker spherical, 150–270 in diameter. Mouth ventral, median,
situated in posterior part of anterior third of body. Pharynx 100–160 in diameter.
Oesophagus long. Caecum saccular, directed posteriorly. Testes two, smooth,
tandem, equal in size, separated from one another by a distance of 150–380, anterior
testis lies in posterior part of middle third of body, posterior testis lies at junction of
middle and posterior thirds of body, testes distinctly separated. Cirrus-sac thick-
walled, small. Seminal vesicle oval. Pars prostatica long. Ejaculatory duct short.
Genital lobes unequal, project into genital atrium. Genital atrium triangular. Genital
Fig. 6.15 Prosorhynchoides

brayi (after Ahmad, 1987,
Pakistan J. Zool. 19, p. 328))
pore subterminal. Ovary round. Uterus extends from posterior margin of ovary to
anterior level of genital atrium. Vitelline follicles in narrow fields on either side
of gut, follicles numbering 8–10 on each side. Eggs oval, 11–14 by 10–12 in size.
Excretory vesicle extends just beyond anterior testis.
Remarks: This species differs from all the known Indian species of the genus in
having a long and slender body, the uterus restricted to the postovarian region and
the mouth lying some distance anterior to the anterior limit of the vitelline fields.
Prosorhynchoides chorinemi (Gupta & Ahmad, 1976) Madhavi, 2011
[Syn: Bucephalopsis chorinemi Gupta & Ahmad, 1976]
Host: Carangidae: Scomberoides lysan (Forsskal) (= Chorinemus moadetta)
Locality: Puri, BOB
Number: 12
Reference: Gupta, V. & Ahmad (1976a)
Description: Body elongate, slender, 3057–4750 long. 295–460 wide. Tegument
spinose. Anterior sucker subspherical, subterminal, 80–120 long. 90–124 wide.
Mouth ventral, equatorial. Pharynx oval. Oesophagus long. Caecum saccular,
directed posteriorly, extends to posterior margin of ovary. Genital pore subtermi-
nal. Testes oval, tandem, subequal, separated from each other by uterine coils, in
posterior part or middle one-third of body. Cirrus-sac elongate extends either to mid-
dle of posterior testis or just posterior to it. Seminal vesicle oval. Pars prostatica long.
Ejaculatory duct short. Genital atrium oval encloses genital lobes. Ovary oval, sub-
median, post-pharyngeal, pretesticular. Uterus coiled, extends from posterior edge of
ovary to genital atrium. Eggs oval, operculated, 19–29 long. 12–17 wide. Vitellarium
follicular, in two lateral rows, 14–18 follicles on right side, 14–19 on left side, extend
from middle of anterior testis to distinctly anterior to pharynx.
Remarks: In the body shape, in the extension of the vitelline fields distinctly anterior
to the pharynx and in having testes separated from each other by uterine coils this
species resembles P. microcirrus but differs from it in the arrangement and number
of vitelline follicles, in the extension of the cirrus-sac to the middle of the posterior
testis.
Prosorhynchoides karvei (Bhalerao, 1937) Srivastava & Chauhan, 1973 (Fig. 6.16)
[Syn. Bucephaloides karvei Bhalerao, 1937]
Host: Belonidae: Xenentodon cancila (Hamilton) Belone cancila
Locality: Indian Ocean
Reference: Bhalerao (1937)
Description: Body pyriform, wider anteriorly, length 500–965, width 270–570.
Tegument spined. Anterior sucker subterminal. Pharynx large, muscular, at about
posterior third of body. Oesophagus long, slender. Caecum rounded, with thick walls.
Testes almost tandem, dextral, close to intestine, anterior usually larger. Cirrus-sac
sinistral, longer than half length of body. Ovary dextral, anterolateral to intestine.
Uterine coils on left side of body. Eggs 18–21 × 9–13. Vitelline fields anterolateral,
follicles in compact groups, rounded or elongate.

karvei (after Bhalerao, 1937)
Remarks: B. karvei is characterised mainly by its small body size, the relatively
long cirrus-sac reaching anterior to the midbody, and the vitellarium arranged in two
widely separated groups of follicles.
Prosorhynchoides microcirrus (Chauhan, 1943) Srivastava & Chauhan, 1973

(Fig. 6.17)
[Syn: Bucephaloides microcirrus Chauhan, 1943]
Host: Sciaenidae: Johnius belengeri (Cuvier) (1), Scombridae: Scomberomorus gut-
tatus (Bl. & Schn.) (= Indocybium guttatum) (2)
Locality: 1. Mumbai, AS; 2. VSK, BOB
Reference: 1. Chauhan (1943a); 2. Madhavi (1974b)
Description: Body long, slender, 1710 long, 120 in width. Anterior sucker oval, sub-
terminal 16 by 4. Mouth situated at midbody. Pharynx globular. Oesophagus straight.
Caecum narrow thin-walled, posteriorly directed. Testes two, globular, postovarian,
median, tandem and separate, anterior slightly larger than posterior. Cirrus-sac small,

microcirrus (original)
ovoid, not reaching testes, encloses ovoid seminal vesicle elongate pars prostatica,
ejaculatory duct elongate. Genital atrium large and globular. Genital pore just ante-
rior to posterior extremity. Ovary globular, pretesticular, dorsal to gut. Uterus short.
Eggs 19 × 16. Vitelline follicles few, large and rounded, arranged in two lateral rows
15–17 on right and left side, extend from half distance between ovary and anterior
testis to well anterior to mouth. Excretory vesicle tubular.
Remarks: This species is characterized by the very long slender body, the compar-
atively small cirrus-sac, the extension of vitellarium from anterior to the caecum to
the ovary, the position of ovary at the posterior end of the elongated caecum, the
separation between the ovary and the anterior testis, between the testes and between
the testes and the cirrus-sac. Chauhan (1943) reported this species from Johnius
belengeri from the Arabian Sea. Madhavi (1974b) recorded it from Scomberomorus
guttatus from Bay of Bengal.
Prosorhynchoides pritchardae (Govind, 1985) Bott & Cribb, 2005 (Fig. 6.18)
[Syn. Bucephalopsis pritchardae Govind, 1985; B. thapari Gupta & Singh, 1985;
Prosorhynchoides guptai (Gupta & Singh, 1985) Bott & Cribb, 2005] n. syn.]
Host: Carangidae: Scomberoides lysan Forsskal (= Chorinemus moadetta (Cuv.
&Val))
Locality: PR (BOB)
Reference: Govind (1985); Gupta, P.C. & Singh (1985b)
Description: Body elongate, slender, 2620–3030 long, 410–430 wide. Tegument

spinose. Anterior sucker subterminal, 50–60 by 60–70. Mouth pre-equatorial. Phar-
ynx subglobular. Oesophagus small. Caecum saccular, extends to posterior mar-
gin of ovary. Genital pore subterminal at posterior end of body. Testes entire,
subspherical, tandem, separated by uterine coils, situated in middle third of body.
Cirrus-sac elongate, curved, not reaching to posterior testis, encloses ovoid seminal

pritchardae (after Govind,
1985)
vesicle, long tubular pars prostatica, tubular ejaculatory duct. Distinct genital lobe
in genital atrium. Ovary subspherical, pre-equatorial, pretesticular. Uterus occupies
space between ovary and genital atrium. Eggs 17–19 long 12–13 wide. Vitellar-
ium follicular, follicles rounded, in two lateral rows, 16 on either side extends from
anterior to mouth to distinctly anterior to anterior testis.
Remarks: Gupta & Singh (1985b) described Bucephaloides thapari Gupta & Singh,
1985, from the same host and locality as that of P. pritchardae. Bott & Cribb (2005)
renamed it as P. guptai since P. thapari was preoccupied by Prosorhynchoides tha-
pari (Dayal, 1948) Bott & Cribb, 2005. P. guptai appears to be a synonym of P.
pritchardae. Both the species were collected from the same host and locality, and
morphologically, they were similar. The year of publication of P. pritchardae and B.
thapari is the same, but as B. thapari would become a junior homonym if transferred
to Prosorhynchoides; it is clear that P. pritchardae is the best name for this taxon and
is used here, where we are considering ourselves the first revisers. The renaming of
B. thapari as P. guptai occurred in 2005, so clearly this does not have priority.
Key to species of Prosorhynchoides recorded from Indian marine fish.
1. Body small, pear shaped …………………………………………………….. 2
Body elongate or long and slender……………………………….………….. 3
2. Cirrus-sac extends to posterior level of anterior testis …….. P. belonea Srivastava,
1938
Cirrus-sac extends to level of ovary ………………………………P. karvei
Bhalerao, 1937
3. Mouth and pharynx in front of vitelline field ……………… P. brayi (Ahmad,
1987)
Mouth and pharynx in middle of vitelline field..………………………….…..4
4. Cirrus-sac very small, extends halfway between testes and posterior
extremity …………………………………………………………. P. microcir-
rus (Chauhan, 1943)
Cirrus-sac longer, reaching to level of testis ………………………………… 5
5. Caecumat level of anterior testis ………………….….. P. chorinemi (Gupta &
Ahmad, 1976)
Caecum distinctly anterior to anterior testi ……….……..…… P. pritchardae
(Govind, 1985)
Genus Rhipidocotyle Diesing 1858
Diagnosis: Body cylindrical, spined. Rhynchus a simple sucker with pentagonal
cap or hood like expansion with or without papillae. Mouth opening usually in
middle third of body. Caecum short. Testes tandem or oblique, in posterior half of
body. Cirrus-sac long, cylindrical. Seminal vesicle spherical or ovoid. Pars prostatica
curved, never straight. Ovary pretesticular. Vitellarium divided into two preovarian
groups. Excretory vesicle variable in length. Type species R. galeata (Rudolphi,
1819). Nine species of this genus have been recorded from marine fishes of India.
Of these, R. karthai Hafeezullah & Siddiqi, 1970, was regarded as a synonym of R.
ghanensis Fischthal & Thomas, 1968, by Madhavi (1974b)
Fig. 6.19 Rhipidocotyle

apapillosa (after Chauhan,
1943)
Rhipidocotyle apapillosa Chauhan, 1943 (Fig. 6.19)

Host: Clupeidae: Clupea sp.
Locality: Mumbai, AS
Reference: Chauhan (1943a)
Description: Body small, elongate, 1510 × 180, anterior third of body slender.
Rhynchus subterminal, weakly muscular, 95 by 85. Hood or cephalic disc with
dorsal pad-like structure, semicircular anterior edge and posterior edge prominently
notched midventrally, disc measures 90 by 60. Mouth post-equatorial. Caecum
extends posteriorly as far as middle of anterior testis. Gonads contiguous, crowded
together almost at same level. Testes obliquely tandem, contiguous. Cirrus-sac
extends to anterior end of posterior testis. Ovary small, subspherical, sinistral to cae-
cum, anterior to anterior testis, slightly overlapping it. Uterus does not extend beyond
anterior limit of vitellarium. Eggs minute, 12.5 by 8. Vitelline fields in midbody.
Remarks: This species is characterised by the absence of papillae on the hood, the
contiguous testes, the ovary in contact with the anterior testis and the cirrus-sac
reaching to the level of the posterior testis.
Rhipidocotyle ghanensis Fischthal & Thomas, 1968 (Fig. 6.20)
[Syn: R. karthai Hafeezullah & Siddiqi, 1970]
Host: Psettodidae: Psettodes erumei (Bl. & Schn.)
Locality: VSK (BOB)
Distribution: Ghana, Bay of Bengal
Description: Body 1770–2510 long, 290–450 wide at testicular level. Rhynchus
190–205 by 143–175, subterminal with seven-lobed polygonal hood. Mouth dis-
tinctly post-equatorial. Caecum terminates anterior to or at pharyngeal level. Gonads
smooth, tandem, usually overlapping, dextral, posterior to midbody length. Cirrus-
sac extends to level of testes. Ovary pretesticular. Uterus extensive extends anteriorly

ghanensis (original)
beyond vitellarium. Eggs 23–27 by 16–18 (15–21 by 15–18). Vitelline fields extend
from about midway between ovary and anterior extremity to level of ovary.
Remarks: This species was first reported from Psettodes belcheri from Ghana (Fis-
chthal & Thomas, 1968) and later from P. erumei from the Bay of Bengal (Madhavi,
1974b). R. karthai Hafeezullah & Siddiqi, 1970, from the same host and locality was
synonymized with this species (Madhavi, 1974b). It is characterised by the rhynchus
with a broad and hexgonal cap-like hood, with one median, two pairs of anterolateral
and one pair of postero-lateral papillae. R. ghanenis is closely related to R. laruei
Velasquez, 1959, reported from Psettodes erumei from the Philippines (Velasquez,
1959) in having the mouth near the midbody, the caecum anteriorly directed and
the excretory bladder extending anteriorly to the level of the Caecum Vitellarium
with 8–15 follicles on the right, 14–18 on the left.
Rhipidocotyle indicus Gupta & Ahmad, 1976 (Fig. 6.21)
Host: Cynoglossidae: Cynoglossus lida (Bleeker)

indicus (after Gupta &
Ahmad, 1976)
Locality: PR, BOB

Number: 5
Description: Body flattened, tapering towards ends, 1500–2490 long, 420–680

wide. Rhynchus cup-like, 150–190 long, 190–210 wide; hood polygonal or fan-
shaped, with seven processes; a median dorsal, two dorsolateral, two ventrolateral
and two ventral papillae. Mouth equatorial, close to anterior testis. Caecum short,
directed posteriorly. Testes in middle third of body. Cirrus-sac extends either to
level of pharynx or to middle of anterior testis. Ovary pretesticular. Uterus anterior
extent just posterior to anterior extent of vitelline fields. Eggs oval, 18–25 by 11–17
wide. Vitelline fields extend to distinctly anterior to ovary to about midway between
ovary and anterior extremity.
Remarks: This species very closely resembles R. ghanensis in having a polygonal

fan-shaped hood with seven processes but differs from it in having a short caecum
and in the extension of cirrus-sac to the level of the pharynx.
Rhipidocotyle khalili Nagaty, 1937 (Fig. 6.22)

Host: Sphyraenidae: Sphyraena obtusata Cuvier
Locality: VSK, BOB
Geographic distribution: Red Sea, BOB
Description: Body long, slender, 2558–3300 long, 314–363 wide. Rhynchus

rounded posteriorly, truncate anteriorly, 122 in diameter, surrounded by crown-
shaped structure provided with three small papillae. Mouth equatorial. Caecum

khalili (after Nagaty, 1937)
reaches to near anterior border of ovary. Testes in posterior part of third quarter of
body. Cirrus-sac reaches to testicular level. Ovary spherical, pretesticular, in anterior
half of third quarter of body. Uterus extends anteriorly as far as junction of anterior
and second quarters of body, does not extend anterior to vitelline fields. Eggs 19–23
by 13–15. Vitelline fields in third quarter of body.
Remarks: R. khalili Nagaty, 1937, was originally described from the milkfish
Chanos chanos in the Red Sea off Koseira. Subsequently, it was reported from
Sphyraena spp. off Makassar, Sulawesi (Yamaguti, 1953), from S. obtusata from
the Bay of Bengal off the Waltair coast (Madhavi, 1974b), from S. japonica from
the Indian Ocean off Mozambique (Reimer, 1985), and recently from S. putnamae
in a lagoon off New Caledonia (Bray & Justine, 2011).
Rhipidocotyle ligulum Chauhan, 1943 (Fig. 6.23)

Host: Ariidae: Arius arius (Richardson)
Locality: BOM, AS
Description: Body narrow, elongate, 2680 long 210 wide. Rhynchus with promi-
nent but feebly developed muscular, crescent-shaped hood without papillae. Mouth
equatorial. Caecum reaching posteriorly as far as ovarian level. Testes in anterior
part of posterior third of body, separated. Cirrus-sac not reaching to posterior testis.
Ovary pretesticular. Vitelline fields in middle third of body.
The description of this species is based on a single specimen collected from Arius
falcarius from the Mumbai coast. It is characterised by the long slender body, the
rhynchus with a crescent-shaped hood but lacking papillae, the separated oblique
testes, the cirrus-sac not extending to the level of the testes and the vitelline fields
extending slightly anterior to the pharynx.

ligulum (after Chauhan,
1943)
Rhipidocotyle pentagonum (Ozaki, 1924) Eckmann, 1932 (Fig. 6.24)

[Syn: Nannoenterum pentagonum Ozaki, 1924]
Hosts: Scombridae: Auxis thazard Lacepede, Euthynnus affinis (Cantor)
Locality; VSK, BOB
Distribution: Japan, Mediterranean, BOB, Rio de Janeiro
Description: (based on English version in Ozaki, 1928). Body elongate, 2220–3240

long, 350–530 wide. Rhynchus with pentagonal hood. Mouth equatorial. Caecum
saccular, passing posteriorly, reaching to ovary. Testes in anterior part of posterior
third of body slightly separated. Cirrus-sac reaching almost to posterior testis. Ovary
pretesticular, contiguous with anterior testis. Uterus extends to about level of mouth.
Eggs 20–22 by 13–15. Vitelline fields reach from level of mouth anteriorly, to about
midway between mouth and anterior extremity.
Remarks: This species was originally described from Scomberomorus niphonius

from Japan and has since been reported in different hosts and localities. It has been
reported in different species of scombrid fishes including: Thunnus thynnus from
the Mediterranean by Eckmann (1932) and from the Pacific Ocean by Yamaguti
(1938a); from Auxis thazard and Euthynnus affinis from the Bay of Bengal by Mad-
havi (1974b), and E. lineatus from Mexico by Castillo-Sánches et al. (1997) and A.
thazard and Katsuwonus pelamis from Rio de Janeiro (Fernandes et al. 2002).

pentagonum (after Ozaki,
1924)

septapapillata (after Krull,
1934)
Rhipidocotyle septapapillata Krull, 1934 (Fig. 6.25)

Host: Sparidae: Acanthopagrus berda (Forsskal) (= Chrysophrys berda)
Locality: Bombay, AS
Distribution: Japan
Reference: Chauhan (1943a).
Remarks: According to Chauhan (1943a), the specimens collected by him were
much larger than those collected by Krull (1934) in Virginia from the freshwater fish
Eupomotus gibbosus. It is unlikely that this is a correct identification considering its
marine source and geographical separation.
Rhipidocotyle sphyraenae Yamaguti, 1958 (Fig. 6.26)
Host: Sphyraenidae: Sphyraena obtusata (Cuvier)
Locality: VSK, BOB
Reference: Yamaguti (1958), Madhavi (1974b)
Geographic distribution: Japan, BOB
Description: Body subcylindrical 900–1700 × 160–260. Rhynchus bowl-shaped,
70–110 long 96–140 wide with semicircular apical hood with seven double papillae.
Mouth pre-equatorial. Caecum elongated, saccular, terminates at level of anterior part
of ovary. Testes tandem, post-equatorial. Cirrus-sac extends to posterior testis. Ovary
subglobular, post-equatorial, immediately pretesticular. Uterus reaches to rhynchus.
Eggs 18–21 by 13–14. Vitelline fields reach between anterior testis and midway
between mouth and anterior extremity.
Remarks: This species differs from the most closely related R. barracudae Manter,
1940, from Sphyraena barracuda from the coast of Japan, in the apical hood of the
rhynchus being provided with seven marginal double papillae. In R. barracudae,
the apical hood is pentagonal and devoid of papillae. Yamaguti (1959) reported the
species from S. pinguis from Sagami Bay, Japan.

sphyraenae (after Yamaguti,
1938)
Rhipidocotyle theraponi Gupta & Tandon, 1983 (Fig. 6.27)

Host: Terapontidae: Terapon theraps (Cuv. & Val.)
Locality: PR, BOB
Number of specimens: 3
Reference: Gupta, S.P. & Tandon (1983)
Description: Body squat, tapers towards both ends, 1080–2110 long 450–680 wide.
Rhynchus cup-like, 160–230 long 210–330 wide, hood polygonal or fan-shaped
with seven processes and seven papillae. Mouth equatorial. Caecum short, reaches

theraponi (after Gupta
& Tandon, 1983)
anteriorly to mouth. Testes tandem, overlapping, in third quarter of body. Cirrus-

sac extends to midbody or just anterior. Ovary pretesticular. Uterus reaches to level
of anterior vitelline extent. Eggs 20–24 by 10–15. Vitelline fields extend from just
posterior to rhynchus to level of pharynx or anterior end of cirrus-sac.
Remarks: This species resembles R. indicus Gupta & Ahmad, 1976, R. septpapillata
Krull, 1934, R. laruei Velaquez, 1959, and R. ghanensis Fischthal & Thomas, 1968,
in having a polygonal hood with seven processes but differs from all these species
in possessing a long cirrus-sac extending to midbody or just anterior to midbody.
Key to Indian species of Rhipidocotyle

1. Body elongate, flat; rhynchus with a pentagonal hood-like structure and seven
papillae…………………………………………………….…………………...2
Body long, slender, rhynchus hood without papillae ………………………… 6
2. Cirrus-sac long extends to level of mouth…………… R. theraponi Gupta &
Tandon, 1983
Cirrus-sac extends to level of testes …………………………………………….3
3. Caecum very short …………………………………….. R. indicus Gupta &
Ahmad, 1976
Caecum well developed………………………………………………………..4
4. Uterus not reaching anterior to mouth………………..……R. pentagonum
(Ozaki, 1924)
Uterus reaching distinctly anterior to vitelline fields ………………..…………5
5. Mouth post-equatorial, gonads contiguous..…………….. R. ghanensis Fischthal
& Thomas, 1968
Mouth equatorial, gonads slightly separated……………..…… R. sphyraenae
Yamaguti, 1959
6. Uterus extending anterior to vitelline zone ……………………..……… R. khalili
Nagaty, 1937
Uterus not extending anterior to vitelline zone …………….. 7
7. Rhynchus with a crescent shaped hood, testes not contiguous, cirrus-sac not
reaching level of posterior testis ………….………………………………..….
R. ligulum Chauhan, 1943
Rhynchus with a small pad-like hood, testes contiguous. Cirrus-sac reaches level
of posterior testis …………………………R. apapillosum Chauhan, 1943
Subfamily: Prosorhynchinae Nicoll, 1914
Diagnosis: Body oval or elongate. Tegument spined. Rhynchus muscular, glandular,

infundibuliform, capped with muscular disc, disc with or without long or short spines
or lobes. Caecum simple, sac-like. Testes two, at midbody. Cirrus-sac thin- to thick-
walled containing seminal vesicle, looped pars prostatica, ejaculatory duct opening
into genital atrium. Genital atrium thin-walled, encloses genital lobe. Ovary usually
subglobular. Vitellarium follicular, follicles arranged in two lateral groups or one
curved group near or anterior to midbody. Uterus tubular, coiling throughout body.
Excretory vesicle tubular.
Key to genera of Prosorhynchinae
1. Rhynchus small, funnel-shaped bearing disc……………….………………….2

Rhynchus shallow funnel-shaped, surrounded by muscular disc encircled by
three to four rows of spines…………………………………………………….
Dollfustrema Eckmann, 1934
2. Ovary pretesticular………………………………………… Prosorhynchus
Odhner, 1905
Ovary post-testicular………………………………….….. Neoprosorhynchus
Dayal, 1948
Ovary intertesticular …………………………………..…………. Neidhartia
Nagaty, 1837
Genus Dollfustrema Eckmann, 1934

[Syn: Dollfusina Eckmann, 1932 (Preoccupied); Neodollfustrema Long & Lee, 1964]
Diagnosis: Body elliptical. Rhynchus a shallow funnel topped by muscular mass

having three to four circlets of short spines. Mouth in posterior half of body. Caecum
elongate. Testes oblique. Ovary smooth, between testes or at level of anterior testis.
Vitellarium in two clumps of follicles. Type species: D. vaneyi (Tseng, 1930)
Dollfustrema bengalense Madhavi, 1974 (Figs. 6.28 and 6.29)

Host: Muraenidae: Gymnothorax undulatus (Lacepede)
Locality: VSK, BOB
Fig. 6.28 Dollfustrema

bengalense (after Madhavi,
1974)
Fig. 6.29 Dollfustrema

bengalense terminal genitalia
(after Madhavi, 1974)
Description: Body pyriform, 1920–2430 long 830–990 wide. Tegument entirely

spined. Rhynchus broad anteriorly, rounded posteriorly filled with muscular and
glandular tissue, 230–350 by 310–410, anterior edge with five rows of obliquely
set spines, spines long, slender, anterior most spines smallest. Mouth postequatorial,
1200–1540 from anterior end. Pharynx large, globular, surrounded by gland cells.
Caecum sac-like, in middle third of body. Testes ovoid, tandem or oblique, in middle
third of body. Genital pore subterminal. Cirrus-sac stout, thick-walled, anterior end
near mouth. Seminal vesicle long, bent at tip, about half length of cirrus-sac. Pars
prostatica long recurved, extends to base of cirrus-sac before looping back and widen-
ing to form terminal portion lined with microvilli. Genital atrium spacious encloses
three finger-shaped lobes, produced anteriorly into fine tube. Ovary intertesticular,
spherical, at about midbody. Uterus fills most of body space posterior to vitellarium,
terminal part of uterus modified into narrow thick-walled metraterm. Eggs 23 by
16. Vitelline follicles in two lateral groups in testicular zone, confluent anteriorly
forming arch in front of testes.
Remarks: This species differs from all the other species of Dollfustrema except D.
gymnothoracis Nahhas & Cable, 1964, in the uterus not extending to the level of
the rhynchus and the vitellarium arranged in an arcuate manner. It differs from D.
gymnothoracis in the much larger size of the spines on the rhynchus, the more anterior
gonads and the smaller eggs.
Genus Prosorhynchus Odhner, 1905

[Syn: Gotonius Ozaki, 1924; Paraprosorhynchus Kohn, 1967]
The important diagnostic characters of the genus are: a conical or pyriform
rhynchus, the ovary is anterior to the testes and the pars prostatica is curved.
Diagnosis: Body elongate or ellipsoidal. Rhynchus funnel-shaped, capped by mus-

cular disc. Mouth position variable. Testes oblique, near midbody. Pars prostatica
curved. Genital lobe prominent. Ovary pretesticular. Vitellarium in two lateral groups
or in one confluent group of follicles. Excretory vesicle of variable length. Type
species: P. squamatus (Odhner, 1905).
Prosorhynchus Odhner, 1905, is a large genus of bucephalids with over 70 nominal
species, about 60 of which are marine and about 10 are from freshwater (Bray &
Justine, 2006). Fifteen species of Prosorhynchus are known from Indian marine
fishes. Of these, the descriptions of three species P. bengalensis Gupta & Gupta,
1987; P. caballeroi Gupta & Ahmad, 1976, and P. orientalis Gupta & Ahmad, 1986,
Fig. 6.30 Prosorhynchus

atlanticus (after Hafeezullah
& Siddiqi, 1970)
each based on a single specimen obtained from their respective hosts, lack details.
Hence, these are not included in this account. P. vinodae Gupta & Gupta, 1987, is
considered here as a synonym of P. manteri Srivastava, 1938. Similarly, P. tewariae
Gupta & Gupta, 1990, is regarded here as a synonym of P. chorinemi Yamaguti,
1952. Therefore, the descriptions are provided for the remaining ten species.
Prosorhynchus atlanticus Manter, 1940 (Fig. 6.30)
Host: Serranidae: Epinephelus malabaricus (Bl. & Schn.)
Locality: Karwar, AS
Reference: Hafeezullah & Siddiqi (1970a)
Geographic distribution: Caribbean
Remarks: Hanson (1950) considered P. atlanticus as a synonym of P. pacificus, and
Winter (1960), Overstreet (1969), Madhavi (1974b) and Overstreet et al. (2009)
agreed, whereas Nahhas and Cable (1964) and Bray & Justine (2006) did not. Bray
& Justine (2006), from a comparison of the measurements of P. atlanticus and P.
pacificus, concluded that in the former the vitellarium reaches to or close to the
rhynchus, the rhynchus is relatively longer, the cirrus-sac is relatively shorter and
eggs are larger. P. atlanticus is essentially an Atlantic species, reported originally in
serranids. Hafeezullah and Siddiqi (1970a) reported it from the serranid Epinephelus
malabaricus from the Arabian Sea.
Prosorhynchus arabianum Srivastava, 1938 (Fig. 6.31)
Host: Soleidae: Brachirus orientalis (Bl. & Schn.) (= Synaptura orientalis)
Locality: Karachi (AS)
Reference: Srivastava (1938a); Chauhan (1954)
Description: Body elongate, 3300–4500 × 400–600. Rhynchus conical, 220–300
by 160. Mouth about three fifths of body length from anterior end. Caecum bulb-
shaped. Testes tandem. Cirrus-sac small, extends to middle of posterior testis. Genital

arabianum (after Srivastava,
1938)
tongue small, hook-shaped. Ovary small, ovoid. Uterus extend to just pre-equatorial
level. Eggs 23 × 12. Vitelline fields lateral, between mouth and ovary, not confluent
anteriorly.
Remarks: This species possesses the following combination of characters: an elon-
gate body, with contiguous gonads in the posterior third of the body, the caecum lying
anteriorly to the ovary, the vitelline fields lateral between the level of the mouth and
the ovary and the uterus extending anteriorly beyond the vitelline zone.
Prosorhynchus bengalensis Gupta & Gupta, 1987
Host: Sphyraenidae: Sphyraena obtusata (Cuv. & Va)
Locality: PR, BOB
Number: one
Reference: Gupta, P.C. & Gupta, V.C. (1987a)
Remarks: The description of the species based on a single specimen lacks many
details. The validity of the species is to be verified.
Prosorhynchus caballeroi Gupta & Ahmad, 1976
[Syn. of P. indicus Madhavi, 1974 (New synonymy)]
Host: Carangidae: Alepes djedaba Cuvier (= Caranx kalla)
Locality: PR, BOB
Number: one
Remarks: P caballeroi is very similar to P. indicus Madhavi, 1974, reported from
Scomberoides tala from Bay of Bengal, except for the smaller body size (2,980 ×
110 as against 3,360–4,480 × 400–500 in P. indicus). The rhynchus structure of P.
caballero was not described in detail. From the figure, it is apparent that it was
damaged. Pending examination of more specimens, P. caballeroi is here regarded as
a synonym of P. indicus.

chorinemi (after Madhavi,
1974)
Prosorhynchus chorinemi Yamaguti, 1952 (Fig. 6.32)

Host: Carangidae: Scomberoides tala (Cuvier)
Locality: Madras, VSK, BOB
Distribution: Celebes, BOB
Reference: Hafeezullah & Siddiqi (1970a); Madhavi (1974b); Yamaguti (1952)
Description: Body elongate. Rhynchus with anterior disc and posterior cone. Mouth
just post-equatorial. Caecum reaches posteriorly towards ovary. Testes dextral, about
midway between pharynx and posterior extremity, separated by loops of uterus.
Cirrus-sac reaches to about inter-testicular level. Ovary oval, between intestine and
anterior testis. Uterus reaches just posterior to anterior extent of vitelline zone. Eggs
18 × 11–12 Vitelline fields anterior extent about midway between anterior extremity
and ovary.
Remarks: The original description of the species was based on a single specimen col-
lected from Chorinemus moadetta (Cuv. & Val.) off Macassar, Sulawesi (Yamaguti,
1952). Hafeezullah & Siddiqi (1970a) obtained a single specimen of this species from
Scomberoides tala from the Madras coast, BOB. Madhavi (1974b) collected several
specimens of the species from Scomberoides tala from Visakhapatnam coast, BOB
and redescribed the rhynchus. According to the redescription, the rhynchus comprises
two parts, an anterior disc-like part and a posterior conical part.
Prosorhynchus epinepheli Yamaguti, 1939 (Fig. 6.33)
Hosts: Serranidae: Epinephelus undulosus Quoy & Gaimard; E. chlorostigma(Val.);
E. diacanthus(Val.); Serranus wandersii Day
Locality: Tuticorin, Karwar, Veeraval, AS
Distribution: Inland Sea of Japan, Arabin Sea, Arabian Gulf
Reference: Yamaguti (1939), Hafeezullah & Siddiqi (1970a), Nahhas et al. (2006)

epinepheli (after Yamaguti,
1939)
Description: Body 875–1400 × 325–800. Rhynchus length 125–350, width

190–200. Pharynx near junction of testes. Gonads near midbody, anterior to cirrus-
sac. Testes slightly diagonal Cirrus-sac reaches to posterior testis. Ovary at level of
anterior testis, occasionally intertesticular. Uterus extends anteriorly beyond level of
caecum, but not to anterior extent of vitellarium. Eggs 25–33 by 13–18. Vitellarium
in two lateral groups of follicles, extends from level of testicular junction, forming
an arch in region between caecum and rhynchus. Excretory vesicle tubular, extends
to level of gonads.
Remarks: P epinepheli was originally described from the grouper, Epinephelus

akaara from the Inland Sea of Japan (Yamaguti, 1939). The Japanese and Indian
specimens agree in most details but Indian forms are different in the following char-
acters: The testes may be arranged symmetrically or the left one anterior to the right.
The pharynx may be entirely posterior to the testis. The caecum extends far anterior
to the ovary. The number of vitelline follicles varies between 22–26 (Hafeezullah &
Siddiqi, 1970a).
Prosorhynchus indicus Madhavi, 1974 (Fig. 6.34)

[Syn. P. caballeroi Gupta & Ahmad, 1976 (new synonym)]
Host: Carangidae: Scomberoides tala (Cuvier)
Locality: Visakhapatnam coast, Bay of Bengal
Description: Body elongate, slender, 3360–4480 long by 400–500 wide, with long
neck. Rhynchus small, triangular, 171–234 by 154–195, anterior part disc-like
with lateral papillae, posterior part conical. Mouth slightly post-equatorial. Caecum
extends posteriorly to reach level of ovary. Gonads in posterior third of body. Testes
tandem, separated by uterine coil, in posterior quarter of body length. Cirrus-sac
small, extends to level of posterior testis. Ovary pretesticular. Eggs 17–19 × 8–11.
Vitelline fields confined to posterior half of body, between ovary and about equator.

indicus (after Madhavi,
1974)
Remarks: In having an elongated body and tandem gonads, P. indicus closely resem-
bles P. facilis (Ozaki, 1924) and P. tsengi Tsin, 1933, but differs from both these
species in having a long neck not occupied by uterine coils, the caecum extending
posteriorly and the much smaller eggs.
Prosorhynchus madhaviae Gupta & Gupta, 1990 (Fig. 6.35)

Host: Stromateidae: Parastromateus Niger (Bloch)
Locality: PR, BOB
Number: 5
Reference: Gupta, P.C. & Gupta, R.C. (1990)
Description: Body elongate, slender, with long neck, 1550–1810 by 410–490.

Rhynchus oval, 40–60 by 30–48. Mouth median, just postequatorial. Caecum extends
anteriorly to just anterior to ovary or anterior testis. Testes tandem, overlapping, in
middle third of body. Cirrus-sac extends to midlevel of posterior or anterior testis.
Ovary subspherical, pre-equatorial, pretesticular, overlapping anterior testis. Uterus
ascends beyond anterior extent of vitelline fields. Eggs 14–16 by 12–14. Vitellaria
fields extend from ovary to about midway between ovary and anterior extremity.

madhaviae (after Gupta &
Gupta, 1990)
Remarks: The important features of the species are the elongate body, the post-
equatorial location of the mouth, the anteriorly directed caecum, the post-equatorial
gonads and the uterus extending anteriorly beyond the vitelline fields.
Prosorhynchus manteri Srivastava, 1938 (Fig. 6.36)

Hosts: Tetraodontidae: Takifugu oblongus (Bloch); Trichiuridae: Trichiurus lepturus
Bloch; Eupleurogrammus muticus (Griffith)
Location: Intestine
Locality: PR, VSK, BOB,
Distribution: Arabian Gulf off the coast of Kuwait (Nahhas et al., 2006), Bay of
Bengal
Reference: Srivastava (1938a), Madhavi (1974b), Gupta & Ahmad (1976a), Gopal
et al. (1985)
Description: Body elongate, 1460–1600 long, 330–450 wide. Rhynchus small,

plug-shaped, 110–190 long, 80–90 wide. Mouth pre-equatorial, in anterior fifth of
body. Caecum saccular directed posteriorly to level of ovary. Testes oval, tandem,
contiguous or separated by uterine coils, in middle third of body. Cirrus-sac extends
to just posterior to posterior testis. Ovary oval, pretesticular, distinctly separated from
anterior testis by uterine coils. Uterus extends anteriorly just pre-mouth but not to
anterior vitelline extent. Eggs 18–24 long, 12–15 wide. Vitelline fields lateral extend
between anterior margin of ovary or just anterior to it to mid pre-mouth region.
Remarks: This species is characterized by the small rhynchus, the anteriorly situated
mouth and the posteriorly directed caecum. The ovary is distinctly separated from
the testes by the uterus. The caecum is preovarian.

manteri (original)
Prosorhynchus orientalis Gupta & Ahmad, 1976

Hosts: Scombridae: Rastrelliger kanagurta (Cuvier)
Location: Intestine
Locality: PR, BOB
Number: one
Reference: Gupta, V & Ahmad (1976a)
Remarks: The description of the species, based on a single specimen, lacks many
details. The validity of the species is to be verified.
Prosorhynchus pacificus Manter, 1940 (Fig. 6.37)
Host: Serranidae: Epinephelus tauvina (Forskal)
Locality: VSK, BOB; Karwar, AS
References: Manter (1940); Hafeezullah & Siddiqi (1970a); Madhavi (1974b)
Description: Body elongate, blunt at anterior end. Length 1206–1444; width

300–397. Rhynchus large, conical, longer than wide, width 172–232. Mouth between
1/3 and 1/2 body length from anterior end. Caecum sac-shaped, extends anteriorly.
Testes large, subspherical, near midbody, diagonal or tandem, slightly separated.
Cirrus-sac large, overlapping posterior testis. Ovary spherical, at pharynx level, over-
laps anterior testis. Uterus entirely postovarian. Eggs 24–27 by 12–17. Vitelline fields
extend from level of ovary to just anterior to caecum. Excretory pore terminal, excre-
tory vesicle extends to anterior testis.
Remarks: P. pacificus belongs to a group of Prosorhynchus spp. with the uterus

restricted to the postovarian region. P. atlanticus Manter, 1940, and P. luzonicus
Velasquez, 1959, have been synonymized with this species but Bray & Justine (2006)
did not support this synonymy. Originally, P. pacificus was recorded by Manter

pacificus (after Manter,
1940)
(1940) from the Galapagos Islands. There are several reports of P. pacificus from
serranids. These records are by Winter (1960) from Pacific Ocean off Sinaloa, Mex-
ico, Hafeezullah & Siddiqi (1970a) from Karwar, India, Leong & Wong (1988, 1990)
from Malaysia and Nahhas et al. (2006) from Arabian Gulf off the coast of Kuwait.
Leong & Wong (1989) reported P. pacificus in a lutjanid, Lutjanus johni off Malaysia.
Most of these records were made assuming P. pacificus and P. atlanticus are synony-
mous. In the light of the views expressed by Bray and Justine (2006) which indicate
that the two species are distinct, the various records of P. pacificus and P. atlanticus
need confirmation.
Prosorhynchus platycephali (Yamaguti, 1934) Srivastava, 1938 (Fig. 6.38)
[Syns. Gotonius platycephali Yamaguti, 1934; Prosorhynchus tsengi of Hafeezullah
& Siddiqi (1970)]
Host: Platycephalidae: Grammoplites scaber (Linnaeus) (= Platycephalus scaber
(L.))
Locality: Cochin, AS
Distribution: Japan, China
Reference: Hafeezullah & Siddiqi (1970a)
Remarks: P. platycephali is a parasite of flatheads throughout the Indo-West Pacific
region. Bray & Palm (2009) suggested that the illustration of P. tsengi by Tsin (1933)
shows a lobed rhynchus with an aperture and straight pars prostatica, which indicate
that the species may belong to Rhipidocotyle. Hafeezullah & Siddiqi (1970a) reported
the species from Grammoplites scaber from Cochin and provided an illustration but
no description. Bray & Palm (2009) pointed out that this illustration did not agree with
the original description of P. tsengi, but fitted that of P. platycephali. This elongate
worm is characterized by the widely separated testes, with Mehlis’ gland posterior
to the anterior testis and the ovary contiguous with the anterior testis.

platycephali (after
Hafeezullah & Siddiqi, 1970)
Prosorhynchus tewariae Gupta & Gupta, 1990

[Syn: P. chorinemi Gupta &Tewari, 1985 nec Yamaguti, 1952]
Host: Carangidae: Scomberoides lysan (Forsskål, 1775) (= Chorinemus moadetta
(Cuv. & Val.))
Locality: Madras, BOB
No: 2
Reference: Gupta, P.C & Gupta, V.C. (1990)
Remarks: This species was originally named as P. chorinemi which is preoccupied

by P. chorinemi Yamaguti, 1952. Gupta & Gupta (1987a) proposed the name P.
tewarii for the species. The name was emended as P. tewariae by Gupta & Gupta
(1990). P. tewariae is distinguished from other species of the genus in the elongated
body, the occurrence of only four vitelline follicles, two on each side. This latter
character needs verification since the illustration given for it indicates that it is based
on a gravid specimen and there is a possibility that the vitellarium is overlapped by
the uterus. In other respects, it is similar to P. chorinemi Yamaguti, 1952, and may
be synonymous with it.
Prosorhynchus truncatus Verma, 1936 (Fig. 6.39)

Host: Ariidae: Cephalocassis jatia (Hamilton) (= Arius jatius Hamilton)
Locality: PR, BOB
Reference: Verma (1936)

truncatus (after Verma,
1936)
Description: Body elongate, 1760 × 420. Rhynchus triangular, 250 in diameter.

Pharynx situated at one third of body length from posterior end, 150 in diameter.
Caecum saccular. Testes oval, obliquely tandem. Cirrus-sac short, extends to level of
posterior margin of posterior testis. Ovary at level of mouth, contiguous with anterior
testis. Eggs 35–40 × 18–20. Vitelline fields confluent anteriorly, forming arc around
anterior edge of caecum.
Remarks: P truncatus is based on two specimens, one ovigerous and lost and the
other without eggs, from the intestine of the catfish, C. jatia off Puri, Bay of Bengal.
It is characterized by a posteriorly located mouth and a short cirrus-sac.
Prosorhynchus vinodae Gupta & Gupta, 1987

Host: Trichiuridae: Eupleurogrammus muticus (Griffith).
Locality: PR, BOB
Reference: Gupta, P.C.& Gupta, V.C. (1986a)
Remarks: This species is synonymized here with P. manteri Srivastava, 1938,

reported from trichiurids from Visakhapatnam and Puri Coasts by Srivastava (1938a),
Madhavi (1974b), Gupta & Ahmad (1976a), Gopal et al. (1985).
Genus Neidhartia Nagaty, 1937

This genus differs from all other prosorhynchine genera by the intertesticular position
of the ovary. The rhynchus is bulbous and has a glandular base in the form of a cone
that extends into the body proper. Two species belonging to this genus have been
recorded from marine fish in India. N. neidharti Nagaty, 1937, from Belone sp. and
N. microrhyncha Chauhan, 1943, from the alimentary canal of Psettodes erumei (Bl.
& Schn.) both from the coastof Mumbai (AS) (Chauhan, 1943a).
Diagnosis: Body elongate. Rhynchus conical, provided with lobes or circlet of small
spines. Mouth opening in posterior half of body. Intestinal caecum simple and sac-
like. Testes tandem. Ovary intertesticular. Vitelline follicles in two indistinct groups.
Excretory vesicle tubular, reaching to level of rhynchus. Type species: N. neidharti
Nagaty, 1937.
Neidhartia microrhyncha Chauhan, 1943 (Fig. 6.40)

Locality: BOM, AS
Description: Body elongate, 1390–2930 × 230–400. Rhynchus 144–230 × 70–106.

Mouth just post-equatorial. Caecum saccular, oriented anteriorly. Gonads just post
equatorial. Testes tandem. Cirrus-sac reaching to about half of post-testicular dis-
tance. Ovary smaller than testes. Uterus restricted to postovarian region, but no
mature eggs described. Vitelline fields reaching from about level of ovary to about
mid pre-mouth distance.
Fig. 6.40 Neidhartia

microrhyncha (after
Chauhan, 1943)
Fig. 6.41 Neidhartia

neidharti (after Nagaty,
1937)
Neidhartia neidharti Nagaty, 1937 (Fig. 6.41)

Host: Belonidae: Belone sp.
Locality: BOM, AS
No.: 5
References: Nagaty (1937); Chauhan (1943a)
Description: Body wedge-shaped, wider anteriorly, 842–2112 × 198–347.
Rhynchus relatively massive, 118–209 × 114–190. Mouth in posterior third of
body. Caecum elongate, saccular, oriented anteriorly almost reaching to rhynchus.
Gonads relatively massive, in middle third of body. Testes tandem. Cirrus-sac
reaching to ovary. Ovary of similar size to testes. Uterus reaches to rhynchus. Eggs
23–29 × 15–19. Vitelline fields reaching from about level of ovary to rhynchus.
Remarks: In the description by Nagaty (1937), the size of the worms is quoted as
561–908 × 149–215 × 230–400 and the eggs as 30 × 15. In the table of measurements
(Nagaty, 1937, pp. 166–167), the measurements given above are quoted. Unfortu-
nately, no scale bar was given in the original figure and the worm has not been
redescribed. Bray & Justine (2013) were doubtful if this worm is correctly identified
in Indian waters.
Key to Indian species of Neidhartia Nagaty, 1937 (Chauhan, 1943a)

Body wedge-shaped, rhynchus large, cirrus-sac long, reaching to ovary, vitelline
fields reach or overlap rhynchus .………………………………….……..….….....
N. neidharti Nagaty, 1937
Body elongate, rhynchus small, cirrus-sac short, not reaching testes, vitelline fields
terminate well short of rhynchus…………………………………………….….….
N. microrhyncha Chauhan, 1943
Genus Neoprosorhynchus Dayal, 1948
Dayal (1948a) erected the genus Neoprosorhynchus with N. purius as the type
species, collected from Epinephelus lanceolatum from the Bay of Bengal, off the
Puri Coast. This genus is characterized by the post-testicular position of ovary and
the Y-shaped excretory vesicle. Apart from the type, one more species N. dayali
Ahmad, 1984, has been added to the genus from Eupleurogrammus muticus (Gray,
1831) from Bombay Coast. The type species and N. dayali need to be evaluated
carefully (Overstreet & Curran, 2002).
Diagnosis: Body elongate, rhynchus conical, surrounded by plain muscular disc.

Mouth in anterior half of body. Testes tandem, ovary post-testicular. Type species:
N. purius Dayal, 1948.
Neoprosorhynchus dayali Ahmad, 1984 (Fig. 6.42)

Host: Trichiuridae: Eupleurogrammus muticus (Gray, 1831)
Locality: Bombay, AS
No. 17
Reference: Ahmad (1984a)
Description: Body elongate, 1820–2240 long, 340–520 wide. Rhynchus well devel-
oped, conical, with flattened top, 183–315 long, 155–245 wide at anterior end, ante-
rior edge broadly indented ventrally. Mouth equatorial, between testes. Caecum sac-
cular, directed anterio-sinistally, reaches anterior limit of vitellarium or just beyond.
Testes symmetrical, equatorial, well separated. Cirrus-sac extends to posterior mar-
gin of ovary. One ventral and two smaller atrial lobes. Ovary spherical, submedian,
dextral at junction of middle and posterior thirds of body. Uterus extends anteriorly
almost to rhynchus. Eggs 30–34 by 22–25. Vitelline fields occupying space of middle
third of body, not forming arch anteriorly.
Remarks: N. dayali was differentiated from the type species N. purius Dayal, 1940,
in having symmetrical rather than tandem testes and in the larger eggs. The excretory
bladder in N. dubius is reported to be Y-shaped, whereas in N. dayali it is tubular.
Fig. 6.42 Neoprosorhynchus

dayali (after Ahmad, 1984,
Pakistan J. Zool,16, 47)
Neoprosorhynchus purius Dayal, 1948

Host: Serranidae: Epinephelus lanceolatus (Bl.)
Locality: PR, BOB
Reference: Dayal (1948a)
Description: Body subcylindrical, wider posteriorly, 2400 × 370. Rhynchus 150,

conical. Mouth just pre-equatorial. Caecum broadly saccular, reaches anteriorly,
beyond gonads, uterus and vitellarium. Testes oval, tandem, sinistral, just pre-
equatorial. Cirrus-sac reaches to about mid postovarian region. Genital lobe large,
muscular. Ovary just postero-dextral to mouth. Uterine coils fill up entire postovar-
ian space (according to description, in illustration it reaches to anterior testis). Eggs
25–27 × 18–21. Vitelline fields at level of caecum. Excretory bladder Y-shaped.
Chapter 7
Superfamily Gymnophalloidea
Odhner, 1905
Family Fellodistomidae Nicoll, 1909
Fellodistomidae are common parasites of a wide range of marine and freshwater

fishes and some marine invertebrates. They are characterized by a smooth tegument,
a V- or Y-shaped excretory vesicle, post-gonadal uterus, restricted vitelline fields,
uterine seminal receptacle and an anterior opening of Laurer’s canal (Bray, 2002).
Brooks et al. (1985) in their phylogenetic analysis of Digenea have not included
Fellodistomidae which they have considered ‘insertae sedis’ and polyphyletic. Sub-
sequently, Hall et al. (1999) supported the polyphyletic nature of Fellodistomidae
and based on molecular evidence, distributed the members of the family in three
families: Fellodistomidae Nicoll, 1909, Faustulidae Poche, 1926 (Syn. Baccigerinae
Yamaguti, 1954) and Tandanicolidae Johnson, 1927.
Fellodistomidae sensu stricto includes forms with the following characteristics:
Diagnosis: Body oval or elongate. Tegument smooth. Oral and ventral suckers small
to large. Pharynx large. Caeca usually two, blind, narrow to wide, reach to testis or
beyond, occasionally single. Testes two, entire or lobed, symmetrical or oblique or
tandem, in anterior or posterior hindbody, postovarian. Cirrus-sac well developed
contains seminal vesicle, pars prostatica and ejaculatory duct. Genital pore usually
sinistral in middle or posterior forebody. Ovary entire to multilobate, in hindbody.
Uterine seminal receptacle present. Uterus extends posterior to testes. Eggs numer-
ous, small, operculate. Vitelline follicles in two lateral fields. Excretory pore terminal,
vesicle V or Y-shaped. Type genus: Fellodistomum Stafford, 1904.
The family includes three subfamilies: Fellodistominae Nicoll, 1909, Piriforminae

Skrjabin and Koval, 1957, and Tergestinae Skrjabin and Koval, 1957. The twenty
species of fellodistomids belonging to six genera, recorded from Indian marine fish
fall into two subfamilies differentiated by the following key:
© Crown 2018 83
https://doi.org/10.1007/978-94-024-1535-3_7
84 7 Superfamily Gymnophalloidea Odhner, 1905
Key to subfamilies of Fellodistomidae:

Cirrus-sac unipartite; oral sucker lacks ring of muscular papillae, muscular flanges
on margins of anterior forebody lacking …………… Fellodistominae Nicoll, 1909
Cirrus-sac usually bipartite, oral sucker usually with ring of muscular papillae; mus-
cular flanges on margin of anterior forebody present ………… Tergestiinae Skrjabin
and Koval, 1957.
Subfamily Fellodistominae Nicoll, 1909
Diagnosis: Body small to large, oval or elongate. Oral and ventral suckers large
or small. Pharynx well developed. Caeca usually two, occasionally single, reach to
testes or beyond. Testes two, oval or lobed, symmetrical, oblique or tandem, in hind-
body. Cirrus-sac well developed, oval to claviform, contains seminal vesicle, pars
prostatica, prostatic cells and ejaculatory duct. Seminal vesicle usually bipartite,
occasionally oval or sinuous. Genital pore usually sinistral, in middle or posterior
half of forebody. Ovary entire to multilobate, pretesticular. Uterus fills entire mid-
body. Vitellarium follicular, in two lateral fields. Excretory vesicle Y or V-shaped
with anterior arms reaching about level of pharynx. Parasitic in gut of marine and
freshwater fishes. Type genus: Fellodistomum Stafford, 1904.
According to Bray (2002), there are 23 genera in this subfamily. Subsequently,
two more genera were added to the subfamily viz Steringovermes Bray, 2004 and
Thryssatrema Sailaja and Madhavi, 2012. Four genera and 17 species of digeneans
belonging to this subfamily have been recorded from marine fish of India. The various
genera are differentiated by features such as the shape of the excretory vesicle (V-
versus Y-shaped), the shape of the seminal vesicle (uni- or bipartite or sinuous), the
orientation of gonads and the distribution of vitelline follicles.
Key to genera
1. Caecum single ………………………………………… Monascus Looss, 1907
Caeca two …………………………………………..………………………… 2
2. Excretory vesicle Y-shaped, vitelline fields far posterior to ventral sucker
…………..…………………………. Thryssatrema Sailaja and Madhavi, 2013
Excretory vesicle V-shaped, vitelline fields in the area between gonads and ventral
sucker………………………………………………………………………… 3
3. Body plump, testes symmetrical, in anterior hindbody ….. Lintonium Stunkard
et Nigrelli 1930
Body elongate, testes oblique to symmetrical, in posterior part of hind-
body.…………………………………… Elopsium Fischthal and Thomas, 1972.
Genus Elopsium Fischthal and Thomas, 1972
[Syn: Protomonascus Thulin 1973]
Diagnosis: Body elongate. Ventral sucker bigger, smaller or subequal to oral sucker.
Caeca long. Testes rounded or lobed, symmetrical or oblique in posterior half of
hindbody. Cirrus-sac saccular, at level of ventral sucker. Seminal vesicle bipartite or
coiled, pars prostatica long, surrounded by thick zone of gland cells, cirrus muscular.
Genital pore median or submedian, in forebody. Ovary spherical or lobed, pretestic-
ular, separated from testes by uterine coils. Uterus from ventral sucker to posterior
Family Fellodistomidae Nicoll, 1909 85
extremity. Eggs oval, thick-shelled. Vitelline follicles extend from ventral sucker to
testes or reach post-testicular zone. Excretory vesicle V-shaped. Type species: E.
ghanensis Fischthal and Thomas, 1972.
The genus includes six species. As many as five species of the genus Elopsium,
4 of them from clupeoid fishes, were reported by Ahmad (1982) from India, but all
these may not belong to this genus. E. puriensis Ahmad, 1982, E. secundus Ahmad,
1982 and E.stunkardi Ahmad, 1982 possess a coiled seminal vesicle and fit into the
genus Elopsium sensu stricto. E. manteri Ahmad, 1982 and E. indicum Ahmad, 1982
possess a bipartite seminal vesicle and may not fit into this genus. According to Bray
(2002), these two species bear some resemblance to Steringophorus Odhner, 1905.
A revision of the genus Elopsium is necessary in order to determine the validity of
various Indian species. Tentatively all the five species are retained in Elopsium.
Elopsium indicum Ahmad, 1982 (Fig. 7.1)
Host: Clupeidae: Dussumieria acuta (Valenciennes)
Locality: PR, BOB
No. of specimens: 9
Reference: Ahmad (1982)
Fig. 7.1 Elopsium indicum

(after Ahmad, 1982)
Description: Body elongate, 3728–4302 long, 779–832 wide. Oral sucker 242–262
in diameter. Ventral sucker 490–516 in diameter, pre-equatorial. Sucker ratio 1:2.
Caeca simple with prominent shoulders at bifurcation, terminate just anterior to
posterior extremity. Testes oblique, in anterior part of posterior third of body. Cirrus-
sac saccular, overlaps right margin of ventral sucker. Seminal vesicle bipartite. Ovary
4–5 lobed, pretesticular. Uterine coils fill entire hindbody. Eggs 47–56 by 26–37.
Vitelline follicles small, in lateral fields, extend from posterior border of ventral
sucker to posterior border of testis.
Remarks: Important features of the species are: The oral sucker is smaller than the
ventral sucker, the cirrus-sac extends to the middle of the ventral sucker, the seminal
vesicle is bipartite and the caeca have prominent shoulders at the level of the intestinal
bifurcation.
Elopsium manteri Ahmad, 1982 (Fig. 7.2)
Host: Megalopidae: Megalops cyprinoides (Broussonet)
Locality: PR, BOB
Fig. 7.2 Elopsium manteri

(after Ahmad, 1982)
Description: Body elongate, 4230–4780 long by 784–896 wide. Oral sucker

286–312 in diameter. Ventral sucker 580–626 in diameter, pre-equatorial. Sucker
ratio 1:1.84–1.96. Caeca long with prominent shoulders, terminate near posterior
end of body. Testes two, symmetrical, multilobed, post-equatorial, close together.
Cirrus-sac saccate, overlaps left border of ventral sucker, contains bipartite seminal
vesicle, long pars prostatica and muscular cirrus. Genital pore immediately anterior
to ventral sucker. Ovary 5-lobed, pretesticular. Uterus fills all space in hindbody. Eggs
82−91 by 42−48. Vitelline fields extend from halfway between posterior border of
ventral sucker and ovary to caecal ends.
Remarks: Differs from all the other species in the genus in having lobed gonads.
Elopsium puriense Ahmad, 1982 (Fig. 7.3)
Host: Elopidae: Elops inermis (Mitchill)
Locality: BOM, AS
No. of specimens: 31
Fig. 7.3 Elopsium puriense

(after Ahmad, 1982)
Description: Body elongated, 4409–4960 long by 925–1109. Oral sucker spheri-

cal, 433–446 in diameter. Ventral sucker of same size as oral sucker, pre-equatorial.
Sucker ratio 1:1. Caeca long, terminate near posterior end. Testes two, symmetri-
cal, smooth, post equatorial, close together. Cirrus-sac saccular, entirely in forebody,
lies horizontally, encloses coiled seminal vesicle, long pars prostatica and muscular
cirrus. Genital pore immediately anterior to ventral sucker. Ovary spherical, subme-
dian, pretesticular, in posterior part of middle third of body. Uterus extensive between
ventral sucker and posterior extremity. Metraterm short, thick-walled. Eggs 35–40
by 23–26. Vitelline fields extend from posterior border of ventral sucker to anterior
border of testes.
Remarks: Characterized by the subequal suckers, spherical ovary and symmetrical

testes. The specific name ‘puriense’ is not appropriate since the species was collected
from fish off Bombay coast, but named after Puri coast.
Elopsium secundum Ahmad, 1982 (Fig. 7.4)
Host: Elopidae: Elops saurus (L.)
Locality: PR, BOB
Number: 15
Fig. 7.4 Elopsium

secundum (after Ahmad,
1982)
Description: Body elongated, 3892–4784 long by 1100–1352 wide at level of ovary.

Oral sucker spherical, 443–470 in diameter. Ventral sucker 308–340 in diameter,
smaller than oral sucker, pre-equatorial. Sucker ratio 1:0.695–0.723. Caeca long,
slender. Testes two, obliquely tandem, post-equatorial, close together but separated
by uterine coils. Cirrus-sac saccular. Seminal vesicle coiled. Genital pore immedi-
ately anterior to ventral sucker. Ovary trilobed. Uterine coils extensive between ven-
tral sucker and caecal ends. Metraterm short, thick-walled. Eggs 20–40 by 18–21.
Vitelline fields extend from anterior border of ventral sucker to caecal ends.
Remarks: This species differs from E. ghanensis Fischthal and Thomas, 1972 the
type species, in the smaller egg size, the extent of the vitellarium and in the sucker
ratio.
Elopsium stunkardi Ahmad, 1982 (Fig. 7.5)
Host: Clupeidae: Dussumieria elopsoides (Bleeker)
Locality: BOM, AS
Fig. 7.5 Elopsium stunkardi

(after Ahmad, 1982)
Description: Body elongated, 3965–4793 long by 890–1107 wide. Oral sucker

365–423 in diameter. Ventral sucker of same size as oral sucker, in anterior part
of middle third of body. Pharynx larger than oral sucker. Oesophagus absent. Intesti-
nal bifurcation dorsal, nearer to oral sucker than to ventral sucker. Caeca simple,
terminate at hind end of body. Testes entire, oblique, subequal, postovarian, close
together, in anterior part of posterior third of body. Cirrus-sac saccular, immediately
anterior to ventral sucker. Seminal vesicle coiled. Genital pore in posterior forebody.
Ovary spherical, in posterior part of middle third of body. Uterine coils extensive,
intercaecal between ventral sucker and posterior extremity. Eggs 67–72 by 32–37.
Vitelline follicles small, in lateral fields, extend from anterior border of ventral sucker
to caecal ends.
Remarks: This species differs from all the other species in the genus by the very
large pharynx. It closely resembles E. puriense but differs from it in having oblique
testes, instead of symmetrical, larger eggs and vitelline fields extending from the
anterior border of the ventral sucker to the caecal ends.
Key to Indian species of the genus Elopsium
1. Ventral sucker smaller or subequal to oral sucker……………………………2

Ventral sucker bigger than oral sucker………………………………….……4
2. Vitelline follicles not extending posterior to testis …….….… E. puriense Ahmad,
1982
Vitelline follicles extending posterior to testis ……………………….……..… 3
3. Ventral sucker smaller than oral sucker, pharynx of medium size, ovary
trilobed…….E. secundum Ahmad, 1982
Ventral sucker of same size as oral sucker, pharynx very large, ovary entire
……… E. stunkardi Ahmad, 1982
4. Testes lobed, symmetrical……. ………………….………………….. E. manteri
Ahmad, 1982
Testes entire, oblique ……………………………………..…………. E. indicum
Ahmad, 1982.
Genus Lintonium Stunkard & Nigrelli, 1930

[Syn: Paradiplangus Gupta, 1968]
Diagnosis: Body plump, with bulbous hindbody and narrow anterior part. Tegu-
ment thick, smooth. Suckers well developed, ventral sucker large, transversely ovate.
Prepharynx absent. Pharynx oval. Oesophagus short or absent. Caeca terminate dis-
tinctly separated from posterior extremity. Testes symmetrical, in anterior hindbody.
Cirrus-sac small, oval, internal seminal vesicle bipartite, pars prostatica surrounded
by numerous prostatic cells. Genital pore median, bifurcal or at level of posterior
part of pharynx. Ovary lobed or oval, pretesticular or intertesticular. Uterus mainly
post-testicular, intercaecal, reaches post-caecally. Vitelline follicles in two lateral
fields in hindbody, mainly post-testicular. Type species: L. vibex (Linton, 1900).
Five species belonging to the genus Lintonium have been recorded from Indian
marine fish, including L. pulchrum (Johnston, 1913); L. puriense Gupta & Ahmad,
1977 and L. pseudovibex Madhavi, 1975. Madhavi (1975b) considered the genus
Paradiplangus Gupta, 1975 as a synonym of Lintonium and transferred the two
species into the genus as L. indicum (Gupta, 1968) and L. tetradontis (Gupta, 1968).
The differences between the various species were not well marked and ultimately all
these may prove to belong to one species.
Lintonium indicum (Gupta, 1968)
[Syn. Paradiplangus indicus Gupta, 1968]
Host: Tetraodontidae: Tetraodon viridipunctatus (Gunther)
Locality: MS, BOB
Reference: Gupta, A. N. (1968a)
Remarks: The description of the species given by Gupta (1968) is brief and lacks
details. Tentatively it is considered here as species inquirendum. Madhavi (1975b)
thought this may well be a synonym of L. pulchrum.
Lintonium pseudovibex Madhavi, 1975 (Fig. 7.6)

Host: Monacanthidae: Paramonacanthus choirocephalus (Bleeker)
Locality: VSK, BOB
Fig. 7.6 Lintonium

pseudovibex (after Madhavi,
1975)
Description: Body pyriform, broad posteriorly narrowed anteriorly, 3040–3584

long, 1392–1456 wide. Oral sucker subterminal, 320–416 by 384–416. Ventral sucker
large, broader than long, 560–640 by 752–800 wide, with transverse aperture, slightly
pre-equatorial. Sucker ratio 1:1.9. Oesophagus absent. Intestinal bifurcation dis-
tinctly anterior to ventral sucker. Testes subglobular, symmetrical, separated by uter-
ine coils, situated just posterior to ventral sucker, Cirrus-sac small, elliptical, median
in mid forebody, not reaching ventral sucker, seminal vesicle bipartite with very
small posterior part and globular anterior part. Genital pore median, bifurcal. Ovary
irregularly lobed, intertesticular, or slightly pretesticular, almost touching posterior
border of ventral sucker. Seminal receptacle absent. Uterus fills entire hindbody, eggs
small, brownish, 31 by 20. Vitelline follicles grouped into 6–8 bunches on each side,
extend from posterior border of ventral sucker to ends of caeca or slightly beyond.
Remarks: According to Madhavi (1975b), L. pseudovibex shows characters inter-

mediate between L. vibex (Linton, 1900) and L.laymani (Layman, 1930) Skrjabin &
Koval, 1958 which according to Manter & Pritchard (1962) differ in the sucker ratio
(1:1.9–2.0 in L. vibex 1:2.3–3.4 in L. laymani) and egg size (45–59 by 23–29 in L.
vibex 26–40 by 13–24 in L. laymani). In L. pseudovibex the sucker ratio approaches
that of L. vibex while its egg size falls within the limits given for L. laymani. Madhavi
further regarded L. vibex reported by Parukhin and Chikunova (1964) from Abalistes
stellaris from South China Sea as synonymous to L. pseudovibex.
Lintonium pulchrum (Johnson, 1913) Yamaguti, 1954
[Syn. Steringotrema pulchrum Johnston, 1913].
Host: Tetraodontidae: Gastrophysus lunaris (Bloch)
Locality: VSK, BOB
Distribution: Australia
References: Madhavi (1975b); Kishore et al. (1989); Johnston (1913)
Description: Body plump, highly muscular, 4080–5760 long, 1280–1760 wide.

Sucker ratio 1:2.3–2.7. Testes subglobular, intercaecal, symmetrical. Genital pore
median, at base of pharynx. Ovary pretesticular. Uterus fills up intercaecal area.
Vitelline fields from testicular level, terminating short of caecal extremities. Eggs
43–48 × 32–35.
Remarks: Crowcroft (1950) regarded L. pulchrum as a synonym of L. consors (Luhe,

1906) while Machida (1971) synonymized it with L. vibex (Linton, 1900). Manter
and Pritchard (1962) characterized it by the shorter caeca, the intercaecal testes and
the vitellarium not covering the ends of the caeca. According to Madhavi (1975b),
the best criteria for the separation of L. pulchrum from the other species of Lintonium
are the more anterior genital pore located at the base of pharynx and the somewhat
rounded eggs (43–48 × 32–35).
Lintonium puriense Gupta & Ahmad, 1977
Host: Tetraodontidae: Gastrophysus lunaris (Bloch)
Locality: PR, BOB
Number: 3
Reference: Gupta, V & Ahmad (1977b)
Description: Body fusiform, 2630–6270 long, 1020–1700 wide. Oral sucker subter-
minal, 280–500 in diameter. Ventral sucker pre–equatorial, 530–800 long, 610–850
wide. Oesophagus short. Testes entire, symmetrical or diagonal, in anterior hindbody.
Cirrus-sac extends either to anterior margin of ventral sucker or just anterior to it.
Genital pore bifurcal. Ovary trilobed, one large dorsal lobe and two smaller ventral
lobes, submedian, pretesticular or overlaps anterior margin of right testis. Uterus
fills up intercaecal area behind ovary. Eggs 32–49 long, 23–34 wide. Vitelline fields
extend either from anterior margin of ovary or testes to posterior extremity of caeca.
Remarks: According to Gupta & Ahmad (1977b), this species closely resembles L.
pulchrum but differs from it in the extension of the cirrus-sac either up to the anterior
margin of the ventral sucker or a little anterior to it, in the vitellarium extending from
anterior level of ovary or testes to the posterior end of the caeca and in having a
bipartite seminal vesicle.
Lintonium tetradontis (Gupta, 1968)
[Syn. Paradiplangus tetradontis Gupta, 1968]
Locality: MS, BOB
References: Gupta, A. N. (1968a); Madhavi (1975b)
Remarks: The description of the species as well as the figure lack details. It is
here regarded as species inquirendum. Madhavi (1975b) thought this may well be a
synonym of L. pulchrum.
Genus Monascus Looss, 1907
[Syn. Haplocladus Odhner, 1911; Karachitrema Bilqees, 1973]
Diagnosis: Body elongate. Oral sucker elongate, larger than ventral sucker. Ventral
sucker in anterior third of body. Pharynx elongate. Oesophagus short. Caecum single,
long. Testes two, tandem, separated by uterine coils, in posterior half of hindbody.
Cirrus-sac oval, anterosinistral to ventral sucker. Seminal vesicle bipartite. Pars pro-
statica tubular, curved. Ejaculatory duct wide, muscular. Genital atrium large. Genital
pore sinistral, in posterior half of forebody. Ovary entire to 3-lobed, pretesticular,
separated from testis by uterine coils. Uterine coils fill much of hindbody. Vitellarium
in two symmetrical lateral fields between levels of ventral sucker and testes. Type
species: M. filiformis (Rudolphi, 1819).
The most significant feature of the genus Monascus is the occurrence of a single
caecum, which has been used to validate subfamily and even family rank for this
genus. But the phylogenetic studies of Hall et al. (1999) indicate that this morpho-
logical character has no significance at the subfamily or family level. Six species
belonging to this genus have been recorded from India: M. chauhani Kumari, 1975;
M. elongatus Karyakarte and Yadav, 1976; M. filiformis (Rudolphi, 1819); M. ori-
entalis (Srivastava, 1941); M. ovatus Kumari, 1975 and M. typicus (Odhner, 1911)
Looss, 1912. Bray and Gibson (1980) regarded only M. filiformis as valid and syn-
onymized the remaining species with this.
Monascus filiformis (Rudolphi, 1819) Looss, 1907 (Fig. 7.7)

[Syns: Monascus chauhani Kumari, 1975; M. elongatus Karyakarte & Yadav, 1976;
M. orientalis (Srivastava, 1941); M. ovilobatus Kumari, 1975 and M. typicus (Odhner,
1911) Looss, 1912]
Hosts: Stromateidae: Pampus argenteus (Euphrasen) (1, 8); P. sinensis (Euphrasen)
(1); Parastromateus niger (Bloch) (2); Scombridae: Rastrelliger kanagurta (Cuvier)
(3,4); Soleidae: Brachirus orientalis (Bl. & Schn.); Ariidae: Arius jella (Day) (5,6);
Carangidae: Decapterus russelli (Ruppell) (7); Caranx sp. (9)
Locality: Coasts of West Bengal, Andhra Pradesh, Tamilnadu, Kerala and Maha-
rashtra (BOB, AS) Distribution: Southwest Atlantic, South America, Mudany coast,
Turkey, Arabian Gulf, Indian ocean
References: 1. Kumari (1975); 2. Karyakarte & Yadav (1976c); 3. Kumari & Sri-
vastava (1976); 4. Hafeezulah & Dutta (1998); 5. Srivastava (1941a); 6. Kumari
& Srivastava (1976); 7. Hafeezullah & Siddiqi (1970b); 8. Madhavi (1975b); 9.
Hafeezullah & Dutta (1998).
Description: Body elongate, narrow, 1618–2700 × 490–675. Tegument smooth.

Oral sucker subterminal, 153–230 × 95–170 with a longitudinal aperture. Ventral
sucker 78–150 × 88–125. Sucker ratio 1:0.60. Prepharynx absent. Pharynx oval. Cae-
cum single, extends to near posterior end of body. Testes elongate, tandem, separated
Fig. 7.7 Monascus filiformis

(after Odhner 1911)
by uterus. Cirrus-sac ovoid, immediately anterior to and overlapping ventral sucker,

encloses bipartite seminal vesicle, small pars prostatica with gland cells and small
ejaculatory duct. Genital pore in forebody. Ovary entire, small, pretesticular, sepa-
rated from anterior testis by uterine coils. Seminal receptacle absent. Vitelline folli-
cles extend from level of posterior margin of ventral sucker to anterior or posterior
testis. Uterus voluminous, fills all available space from posterior margin of ventral
sucker to posterior and of body. Metraterm present. Eggs 34–38 × 19–23. Excretory
vesicle Y-shaped.
Remarks: Køie (1979) studied the life history of M. filiformis and with the aid of
SEM found that the caecum is not unbranched, rather the right caecum is complete
and the left caecum is short and reduced.
Genus Thryssatrema Sailaja & Madhavi, 2012
Diagnosis: Body elliptical. Tegument smooth. Oral sucker somewhat funnel-
shaped. Ventral sucker small, rudimentary, in anterior quarter of body. Prepharynx
absent. Pharynx oval. Oesophagus long. Intestinal bifurcation anterior to ventral
sucker. Caeca slender, reach to posterior extremity. Testes two, oval, symmetrical,
in posterior half of hindbody. Cirrus-sac claviform extends well posterior to ven-
tral sucker, encloses bipartite seminal vesicle, prostatic vesicle, narrow ejaculatory
duct. Genital pore median, immediately anterior to ventral sucker. Ovary deeply
3-lobed, median, immediately pretesticular. Mehlis’ gland small, ventral to ovary.
Laurer’s canal opens dorsally at level of ovary. Uterine seminal receptacle not clear.
Uterus fills most of post-testicular space. Metraterm weekly developed. Eggs numer-
ous, small, thick-shelled, operculate. Vitelline follicles elliptical, in two small lateral
zones in front of ovary, far posterior to ventral sucker. Excretory vesicle V-shaped,
broad arms reach level of bifurcation. Type-species: T. hanumantharaoi Sailaja &
Madhavi, 2012.
Thryssatrema hanumantharaoi Sailaja & Madhavi, 2012 (Fig. 7.8)
Host: Engraulidae: Thryssa setirostris (Broussonet)
Locality: VSK, BOB
Reference: Sailaja & Madhavi (2012)
Description: Body elongate, with broad posterior region, narrow extensile anterior
region, 2400–3312 (2878) long, 330–624 (506) wide. Suckers poorly developed. Oral
sucker somewhat funnel-shaped, almost squarish, 88–172 (126) long, 98–156 (130)
wide. Ventral sucker small, circular, rudimentary, 48–80 (60) in diameter. Sucker
ratio 1:0.34–0.57 (0.48). Prepharynx absent. Pharynx large, spherical, major part
lies dorsal to oral sucker. Caeca long, narrow, terminate blindly close to posterior
extremity. Testes 2, oval, fairly large, subequal in size, symmetrically arranged in
posterior half of hindbody, separated by uterine coils. Cirrus-sac small, claviform,
pointed anteriorly, extends posterior to ventral sucker. Seminal vesicle bipartite, both
chambers of same size. Genital pore median, immediately anterior to ventral sucker.
Ovary distinctly 3-lobed, pretesticular, overlapping right testis. Uterine coils fill
entire post-testicular space. Eggs 24–28 long 12–16 wide. Vitelline follicles elliptical,
forming wide and compact lateral groups anterolateral to testes.
Fig. 7.8 Thryssatrema

hanumantharaoi (after
Sailaja & Madhavi, 2012);
Remarks: The V-shaped excretory vesicle, symmetrical testes, bipartite seminal

vesicle, pretesticular lobed ovary and distribution of vitelline follicles in hindbody
of Thryssatrema suggest an affinity to the genus Prudhoeus Bray & Gibson, 1980
erected by Bray and Gibson (1980) with P. nicholsi Bray & Gibson, 1980 as the
type species. However, Thryssatrema differs from Prudhoeus in many respects: the
shape of body, the sucker proportions viz the oral sucker much larger than the ventral
sucker, the position of the gonads far posterior to the ventral sucker, the extension of
the cirrus-sac posterior to the ventral sucker, the position of the genital pore immedi-
ately anterior to the ventral sucker rather than at the base of the intestinal bifurcation,
in the distribution of the vitelline follicles limited to the middle third of the body and
in the much smaller eggs. So far T. hanumantharaoi is the only species in the genus.
Subfamily Tergestiinae Skrjabin & Koval, 1957
Diagnosis: Body elongate. Oral sucker surrounded by ring of about 13–20 muscu-
lar cephalic lobes and neck region with six muscular flanges. Oral sucker smaller or
larger than ventral sucker, Ventral sucker in anterior half of body. Intestinal bifur-
cation in fore or hindbody. Caeca long, blind. Testes two, oval or irregularly lobed,
tandem to oblique, in mid to posterior hindbody. Cirrus-sac bipartite, extends well

posterior to ventral sucker. Genital pore median or submedian, in posterior forebody.
Ovary reniform to oval or irregularly lobed, pretesticular. Uterine coils extend to pos-
terior extremity or stop at level of testes. Vitellarium in two lateral fields of follicles
in hindbody. Excretory vesicle I-shaped. Type genus: Tergestia Stossich, 1899.
Key to Genera
1. Intestinal bifurcation in anterior hindbody, uterus extends into post-testicular
field ……………………………………………………………………………
Tergestia Stossich, 1899
Intestinal bifurcation in posterior forebody, uterus not extending into post-
testicular space ………………………………………………………….………
Theledera Linton, 1910.
Genus Tergestia Stossich, 1899
Diagnosis: Body elongate. Oral sucker surrounded by ring of 10–20 cephalic lobes.
Six muscular flanges occur on lateral surface at level of pharynx. Ventral sucker
in anterior third of body. Caeca long. Testes tandem in posterior half of hindbody.
Cirrus-sac bipartite, posterior part elongate, anterior part globular. Seminal vesicle in
posterior part, pars prostatica small, ejaculatory duct large, muscular, diverticulate,
eversible. Genital pore in posterior forebody. Ovary oval or lobed, in middle of
hindbody. Uterus fills much of hindbody, intercaecal, pregonadal. Vitelline follicles
in symmetrical lateral fields.
Type species: T. laticollis (Rudolphi, 1819)
Tergestia bengalensis Gupta & Singh, 1985 (Fig. 7.9)
Host: Carangidae: Caranx hippos (Bloch)
Locality: PR, BOB
Number of specimens: one
Reference: Gupta, P.C. & Singh (1985a)
Description: Body elongate, slender, 3220 × 470. Oral sucker ovoid, terminal,
220 × 190. Prepharynx very short. Pharynx elongate, cylindrical. Oral lobes mus-
cular, 11 in number. Forebody with 6 pairs of lateral muscular flanges. Oesophagus
long. Intestinal bifurcation just posterior to ventral sucker. Ventral sucker subspher-
ical, almost in middle of body, 390 × 360. Sucker ratio 1:1.9. Testes entire, ovoid,
oblique, close together, in extreme hind end of body. Cirrus-sac curved, bulbous ante-
riorly, extends to just posterior to posterior margin of ventral sucker. Seminal vesicle
curves around right side of ventral sucker. Pars prostatica bulbous, surrounded by
prostate gland cells. Ejaculatory duct 150 long. Genital pore submedian, anterior to
ventral sucker.
Remarks: According to Gupta and Singh (1985a), this species differs from all the
known species of the genus except T. clonacantha Manter, 1963 in having 11 oral
lobes. It differs from T. clonacantha in having a larger sucker ratio, a lobed ovary,
the extension of the cirrus-sac only up to the posterior margin of ventral sucker and
in the larger eggs.
Fig. 7.9 Tergestia

bengalensis (after Gupta &
Singh, 1985)
Tergestia clonacantha Manter, 1963 (Fig. 7.10)

Host: Hemiramphidae: Hemiramphus far (Forsskal)
Locality: Mandapam, GOM
Reference: Hafeezullah & Siddiqi (1970b)
Description: Length 1843–2527; width 341–455. Oral sucker 140 × 207. Ventral
sucker 160–297. Sucker ratio 1:1–1.2. Oral lobes 11 in number, round or bluntly
pointed, ventral lobe of each side with median branch, two median papilla like lobes,
ventral one on each side of mouth separated from other lobes. Forebody less than
one quarter body length with six pairs of lateral flanges. Pharynx oval. Intestinal
bifurcation at posterior edge of ventral sucker. Caeca not quite reaching posterior
extremity. Genital pore to left, about halfway between ventral sucker and posterior
end of pharynx. Testes ovoid, oblique, at about middle of hindbody, separated by
uterus. Cirrus-sac curves around left side of ventral sucker, reaches halfway or more
along distance between ventral sucker and ovary. Ovary ovoid, reniform, at midbody,
separated from anterior testis by one or more coils of uterus. Uterine coils extensive.
Eggs thin-shelled 15–18 by 9–12. Vitelline follicles scanty, lateral, from posterior
edge of ventral sucker to near posterior end of body.
Fig. 7.10 Tergestia

clonacantha (after
Hafeezullah & Siddiqi, 1970.
Courtesy Allen Press, J.
Parasitol, 56, p. 939)
Remarks: This species is characterized by the presence of 11 oral lobes. Wee et al.
(2017) mentioned that ‘Manter (1963) indicated that Tergestia clonacantha had 11
cephalic lobes with ‘two rudimentary, papilla like lobes, one on each side of the
mouth, separated from the other lobes”. They also said that their ‘examination of his
figures leads us to conclude that the ‘two papilla like lobes’ are regular cephalic lobes’.
They counted 13 cephalic lobes in worms they considered T. clonacantha. They also
pointed out that other species of Tergestia are reported with variable numbers of
cephalic lobes, indicating that these are not reliable distinguishing features. Manter
(1963) described the species from Hemiramphus sp. from Fiji. The only report of
the species from India is that of Hafeezullah & Siddiqi (1970b) from Hemiramphus
far from Mandapam coast.
Genus Theledera Linton, 1910
[Syn. Cithara MacCallum, 1917; Tergestina Nagaty & Abdel, 1964]
Diagnosis: Body elongate. Oral sucker surrounded by ring of 10–20 cephalic lobes.
Six muscular flanges on lateral sides at level of pharynx. Ventral sucker protuberant,
in anterior third of body. Intestinal bifurcation in posterior forebody. Caeca long.
Testes oval, entire or lobed, tandem or oblique close to posterior extremity. Cirrus-
sac bipartite, posterior part elongate, anterior part globular. Internal seminal vesicle
saccular. Pars prostatica small, ejaculatory duct large, muscular, deeply diverticulate,
may form cirrus. Genital pore submedian in posterior forebody. Ovary oval, entire
or lobed, pretesticular. Uterus fills space between anterior testis and ventral sucker.
Vitellarium in lateral fields in mid-hindbody. Type species: T. pectinata (Linton,
1905)
Theledera synganathusi (Gupta & Tandon, 1983) (Fig. 7.11)
[Syn. Tergestina synganathusi Gupta & Tandon, 1983]
Host: Syngnathidae: Trachyrhamphus serratus (Temm. & Schl.)
Locality: Quilon, Kerala, AS
Reference: Gupta, S. P. & Tandon (1983)
Description: Body elongate, 1845 long, 305 wide. Oral sucker terminal, oval, with
a crown of 9 muscular lobes rounded or sharply pointed, 165 long, 145 wide. Phar-
ynx elongate, 138 long, 85 mm wide. Oesophagus long, bifurcates well anterior to
ventral sucker. Intestinal caeca terminate near posterior extremity. Ventral sucker
Fig. 7.11 Theledera

synganathusi (after Gupta &
Tandon, 1983)
larger than oral sucker, 230 long, 235 wide. Sucker ratio 1:1.8. Neck with six pairs of
muscular flanges. Genital pore, median anterior to ventral sucker. Testes subequal,
post equatorial, in posterior part of body. Cirrus sac lateral to ventral sucker, between
intestinal bifurcation and ventral sucker. Seminal vesicle straight, lateral to ventral
sucker. Ovary spherical, post equatorial, distant from anterior testis. Seminal recep-
tacle round, pre-ovarian, just anterior to ovary, Vitelline follicles small, scattered in
lateral fields from just posterior to cirrus-sac to middle of posterior testis. Uterus with
descending coils to posterior end of anterior testis. Eggs small, 20–40 long, 15–26
wide.
Remarks: This species was originally included in the genus Tergestina Nagaty &
Abdel Aal, 1964 which was synonymized with Theletrum Linton, 1910 by Bray
& Gibson (1980). Tergestina synganathusi Gupta & Tandon, 1983 thus became
Theletrum synganathusi (Gupta & Tandon, 1985). The specific name given after
the host Syngnathus is misspelled as synganathusi. It is here retained as such. The
important characters of the species are: the reported occurrence of only 9 oral lobes,
and the testes and ovary not being contiguous. Gupta & Tandon (1983) interpreted
the posterior part of the bipartite cirrus-sac as an external seminal vesicle. This needs
confirmation and would be unique in the subfamily if correct.
Family Tandanicolidae Johnston, 1927
[Syn. Monodhelminthidae Dollfus, 1937, Monodhelminthinae Dollfus, 1937, Mehra-

trematinae Srivastava, 1939, Prosogonariinae Mehra, 1963 and Buckleytrematinae
Yamaguti, 1971]
The family Tandanicolidae is mainly characterized by the presence of a distinct
accessory copulatory organ associated with the genital atrium. It was included as a
subfamily Tandanicolinae under the family Fellodistomidae with Monodhelminthi-
nae Dollfus 1937 as its synonym (see Cribb & Bray, 1994). However, Hall et al.
(1999) presented a molecular and morphological phylogeny which indicated that
the Tandanicolinae and Fellodistominae do not form a clade, and they considered a
separate status as Tandanicolidae appropriate for this taxon. Bray (2002) provided
a review on the family and keys for the various genera included under it. Members
of this family occur in the intestine or swim bladder of siluriform fishes. They are
reported mainly from the Indo-Pacific region.
Diagnosis: Body elongate to oval, Tegument with or without spines. Oral sucker
simple. Ventral sucker smaller or larger than oral sucker, in midbody. Prepharynx
short. Pharynx present. Oesophagus distinct. Caeca extend into hindbody, terminate
blindly. Testes symmetrical or tandem usually in forebody. Cirrus-sac present. Genital
atrium large with thick muscular wall. Accessory copulatory organ situated in genital
atrium, consists of solid digitiform process and associated glandular material. Genital
pore midway between oral and ventral suckers. Ovary multilobed, lobes contiguous
or well separated. Uterus usually in hindbody. Vitellarium follicular, in two lateral

bunches. Excretory vesicle V-shaped. Type genus: Tandanicola Johnston, 1927
Key to genera:
1. Ventral sucker with semicircular laminar apparatus embedded in antero-ventral

part ……………. ……………………………. Prosogonarium Yamaguti, 1952
Ventral sucker not specialized ………………………………….………….… 2
2. Testes tandem …………………………..………….. Buckleytrema Gupta, 1956
Testes symmetrical ……………………………… Monodhelmis Dollfus, 1937.
Genus Buckleytrema Gupta, 1956
[Syn. Paramonodhelmis Oshmarin & Mamaev, 1963]
Diagnosis: Body elongate to oval. Tegument unarmed. Oral sucker simple. Ventral
sucker smaller or larger than oral sucker, in midbody. Prepharynx absent. Phar-
ynx globular. Oesophagus small. Caeca narrow, extend to mid-hindbody. Testes tan-
dem, contiguous, situated just posterior to ventral sucker. Cirrus-sac present, situated
antero-dorsal to ventral sucker. Genital atrium large with thick irregular muscular
wall. Accessory copulatory organ situated posteriorly in genital atrium consists of
solid anteriorly directed digitiform process and associated glandular tissue. Geni-
tal pore midway between oral and ventral suckers. Ovary trilobed. Laurer’s canal
present. Uterus largely post-testicular, extends to posterior end of body. Vitelline
fields lateral, from genital atrium to posterior testis. Type species: B. indica Gupta,
1956
Buckleytrema indica Gupta, 1956 (Fig. 7.12)
[Syns. Paramonodhelmis postacetabulorchis Oshmarin & Mamaev, 1963; Buck-
leytrema postacetabulorchis (Oshmarin & Mamaev, 1963) Yamaguti, 1971]
Host: Ariidae: Nemipterys nenga (Ham.)
Locality: Krusadai Isle, GOM (1); BOM, AS (2)
Distribution: AS, GOM, Queensland, Karachi Gulf of Tonking
References: 1. Gupta, N. K. (1956); 2. Hafeezullah (1971b).
Description: Body elongate oval, 1635–3576 × 356–691 long. Oral sucker

199–324 × 199–340. Ventral sucker at about midbody 98–142 × 94–158. Sucker
ratio 1:0.42–0.47. Prepharynx short, usually entirely dorsal to oral sucker. Pharynx
oval. Oesophagus distinct. Intestinal bifurcation in forebody. Caeca narrow, extend
to mid-hindbody. Testes oval, tandem, contiguous, anterior immediately dorsal to
ventral sucker, posterior immediately posterior to posterior testis. Cirrus-sac con-
tains oval seminal vesicle, prominent pars prostatica, short ejaculatory duct. Genital
atrium large, muscular. Accessory copulatory organ situated posteriorly in genital
atrium, consists of solid anteriorly directed digitiform process and associated glan-
dular tissue. Genital pore midway between oral and ventral suckers. Ovary trilobed,
immediately anterior to anterior testis. Laurer’s canal present. Uterus largely post-
testicular, extends to posterior end of body. Eggs 39–49 × 18–20. Vitelline follicles
in lateral fields of fore and hindbody.
Family Tandanicolidae Johnston, 1927 103
Fig. 7.12 Buckleytrema

indica (after Gupta, 1956)
Remarks: Gupta (1956) erected the genus Buckleytrema with B. indica from ‘a
marine cat-fish’ as the type species. Cribb & Bray (1994) redescribed Buckleytrema
indica from Arius graeffei, from southern Queensland and synonymized B. postac-
etabulorchis (Oshmarin & Mamaev, 1963) from Arius graeffei from Vietnam with
B. indica. B. indica shows specificity towards marine catfishes and has a wide geo-
graphic distribution.
Genus Monodhelmis Dollfus 1937
[Syn. Burnellus Angel, 1871; Mehratrema Srivastava, 1939; Parheterorchis Gupta
& Jain, 1991]
Diagnosis: Body elongate oval, spined. Ventral sucker smaller than oral sucker, in
midbody. Caeca extend to near posterior end of body. Testes symmetrical, situated
in forebody, posterior half of each overlapping anterior part of ventral sucker. Ovary
lobed, pretesticular. Uterus largely in hindbody, between caeca. Vitelline fields lat-
eral, from level of testes to hindbody. Type-species: M. torpedinis Dollfus, 1937.
The number of Indian species of the genus Monodhelmis has increased consider-
ably up to eight species following the synonymy of the genera Mehratrema Srivas-
tava, 1939 and Parheterorchis Gupta & Jain, 1991 with this genus. However, three
out of these eight species namely M. dollfusi Srivastava, 1939, M. polynemusinis
Chauhan, 1943 and M. skrjabini Karyakarte, 1969 were synonymized with the type
species (Cribb & Bray, 1994). This leaves only 5 valid species: M. chilkai Gupta &
Singh, 1999; M. militaris Dutta, 1995; M. orissai (Gupta & Jain, 1991b) M. thapari
(Ahmad, 1981) and M. torpedinis Dollfus, 1937. The descriptions of some of these
species are not adequate and the characters used for distinguishing them from M.
torpedinis Dollfus, 1937 are not clearly defined. Some of these are likely to fall
synonyms of the type species.
Monodhelmis chilkai Gupta & Singh, 1999
Host: Labridae: Halichoeres scapularis (Bennett) (=Platyglossus scapularis (Ben-
nett))
Loclity: Chilka Lagoon
Number: 6
Reference: Gupta, P. C. & Singh (1999)
Description: Body elongate, 1190–2740 long, 390–670 wide. Tegument spinose.

Oral sucker subterminal, 150–270 long, 230–1150 wide. Prepharynx present or
absent. Pharynx elongate. Oesophagus long. Intestinal caeca simple, extend almost
to posterior end of body. Ventral sucker subspherical, post equatorial, post-testicular,
smaller than oral sucker, 110–230 long, 100–210 wide. Testes smooth, ovoid, sym-
metrical, intercaecal, antero-dorsal to ventral sucker. Cirrus-sac described as absent.
Seminal vesicle elongated, pars prostatica long, opens directly into genital atrium,
surrounded by prostate gland cells. Accessory copulatory organ posterior to gen-
ital atrium. Genital pore median, post-bifurcal, distant from ventral sucker. Ovary
trilobed, submedian, pretesticular. Vitellarium follicular, extracaecal, extend either
from just posterior to anterior margin of ovary or anterior level of ventral sucker,
to just anterior to midlevel of hindbody. Uterine coils occupy entire post-testicular
space. Eggs numerous, 52–61 × 24–32. Excretory bladder V-shaped, with two longi-
tudinal arms extending anteriorly to just posterior to intestinal bifurcation; excretory
pore subterminal.
Remarks: M. chilkai was differentiated from other species of the genus in having
a trilobed ovary instead of an entire ovary and in having the vitellarium extending
anterior to the ventral sucker.
Monodhelmis militaris Dutta, 1995
Host: Ariidae: Osteogeniosus militaris (L.)
Locality: Krusadai Isle, GOM
Reference: Dutta (1995)
Remarks: This form is practically similar to M. torpedinis Dollfus, 1937 and may
be a synonym of that species.
Monodhelmis orissai (Gupta & Jain, 1991) Madhavi, 2011
[Syn. Parheterorchis orissai Gupta & Jain, 1991]
Host: Scatophagidae: Scatophagus argus (L.)

Locality: PR, BOB
Reference: Gupta, V. & Jain (1991b)
Remarks: This species was included in a new genus Parheterorchis Gupta & Jain,
1991. Cribb and Bray (1994) expressed the view that the authors mistook the genital
atrium for a ventral sucker and have overlooked the actual ventral sucker. Here we
consider M. orissai as a species inquirendum.
Monodhelmis thapari (Ahmad, 1981) Madhavi, 2011 (Fig. 7.13)
[Syn. Mehratrema thapari Ahmad, 1981]
Host: Ariidae: Plicofollis platystomus (Day, 1877) (=Arius platystomus Day)
Locality: PR, BOB
Number: 38
Description: Body fusiform, 1940–2480 long by 750–960 wide. Tegument thin,

unspined. Oral sucker 190–283 in diameter. Ventral sucker 130–240 in diameter,
aperture oval, triangular or slit-like. Sucker ratio 1:0.68–0.84. Prepharynx absent.
Pharynx oval. Intestinal bifurcation usually midway between oral sucker and genital
atrium. Caeca terminate close to posterior extremity. Testes two, oval, smooth, equal,
Fig. 7.13 Monodhelmis

thapari (after Ahmad, 1981)
symmetrical, intercaecal, immediately post-equatorial. Cirrus-sac arcuate, to left or

right of genital atrium, extends to anterior margin of testes, encloses tubular seminal
vesicle, prostate complex, ejaculatory duct. Genital pore median, usually midway
between intestinal bifurcation and ovary. Ovary entire, median or submedian, imme-
diately pretesticular, contiguous with testes, situated anterior to ventral sucker. Uterus
occupies intercaecal field of hindbody. Accessory copulatory organ pyriform, highly
muscular, with deeply stained gland cells. Eggs oval, yellowish, 32–35 long by 19–20
wide. Vitellarium follicular, lateral, extend from anterior level of genital sucker to
caecal ends. Excretory arms reach to the level of intestinal bifurcation; pore terminal.
Remarks: Originally included in the genus Mehratrema Srivastava, 1939, it differs

from M. torpedinis in the more extensive vitelline fields, distributed between the
intestinal bifurcation and close to the posterior extremity.
Monodhelmis torpedinis Dollfus, 1937 (Fig. 7.14)
[Syns. Mehratrema aria GU & Shen, 1979; M. dollfusi Srivastava, 1939; M. polyne-
musinis Chauhan, 1943; M. skrjabini Karyakarte, 1969; Monodhelmis aria Yam-
aguti, 1952; Monodhelmis dollfusi (Srivastava, 1939); Monodhelmis philippinenen-
sis Velasquez, 1961; Monodhelmis dollfusi (Srivastava, 1939); Ref.: Cribb & Bray,
1994, 2004]
Hosts: Scatophagidae: Scatophagus argus L. (1); Polynemidae: Polynemus indicus
Shaw; Muraenesocidae: Congresox talabonoides (Bleeker); Sciaenidae: Sciaena sp.
Fig. 7.14 Monodhelmis

torpedinis (after Chauhan,
1943)
(2); Terapontidae: Terapon puta (Cuv. & Val.) (3); Ariidae: Arius platystomus Day
(4); Arius jella Day (5)
Locality: AS(1), BOM, AS (2); Pamban, MS, BOB (3); Karwar, AS (4); DC, BOB
(5)
Distribution: Atlantic (Mauritania), Ghana, Indian Ocean, Pacific (south-east
Atlantic)
References: 1. Srivastava (1939f); 2. Chauhan (1943b); 3. Karyakarte (1969); 4.
Hafeezullah (1971b); 5. Kumari & Srivastava (1976)
Description: Body fusiform, 1920–2400 long by 980–1000 wide. Oral sucker spher-
ical, subterminal, 160–204 in diameter. Ventral sucker 172–192 in diameter, smaller
than the oral sucker, median. Prepharynx absent. Pharynx oval. Intestinal bifurca-
tion usually midway between oral sucker and genital sucker. Caeca terminate close to
posterior extremity. Testes two, oval, smooth, equal, symmetrical, intercaecal, imme-
diately post-equatorial. Cirrus-sac arcuate, thin and delicate, not prominent, to left or
right of accessory copulatory organ, extends up to anterior margin of testes encloses
tubular seminal vesicle, prostate complex, ejaculatory duct. Genital sucker median,
usually midway between intestinal bifurcation and ovary. Ovary 3-lobed, median
or submedian, immediately pretesticular, contiguous with testes, situated anterior to
ventral sucker. Vitellarium follicular, lateral, extend from anterior level of ventral
sucker to caecal ends. Uterus intercaecal, descends posteriorly but not quite to pos-
terior extremity. Accessory copulatory organ pyriform, 160–240 long by 110–170
wide, highly muscular, with deeply stained gland cells. Eggs oval, yellowish, 39–48
long by 19–27 wide. Excretory arms reach to the level of caecal bifurcation; pore
terminal.
Remarks: Cribb & Bray (1994) provided a review on the genus Monodhelmis and
considered several species namely Monodhelmis arii Yamaguti, 1952, Mehratrema
arii Gu & Shen, 1979, Mehratrema dollfusi Srivastava, 1939, Monodhelmis
philippinensis Velasquez, 1961, Mehratrema polynemusinis Chauhan, 1943 and
Mehratrema skrjabini Karyakarte, 1969 as synonyms of Monodhelmis torpedinis.
Monodhelmis elongatus Bilqees, 1970 is considered a species inquirendum.
Genus Prosogonarium Yamaguti, 1952
Diagnosis: Body oval, plump. Tegument spined. Ventral sucker in midbody, special-
ized with semicircular laminar muscular apparatus. Caeca extend to midbody. Testes
symmetrical, situated in posterior part of forebody, overlapping ventral sucker pos-
teriorly. Cirrus-sac present, situated in forebody between testes and intestinal bifur-
cation. Ovary trilobed, pretesticular. Seminal receptacle uterine. Uterus occupies
hindbody. Vitelline fields lateral, from ventral sucker to pharynx. Excretory vesicle
V-shaped. Type species: P. arii Yamaguti, 1952
Prosogonarium plotosi Madhavi, 1975 (Fig. 7.15)
Host: Plotosidae: Plotosus lineatus (Thunberg)
Locality: VSK, BOB
Distribution: BOB, Moreton Bay, Queensland [by Cribb & Bray (1994)]
Fig. 7.15 Prosogonarium

plotosi (after Madhavi, 1975)
Description: Body elliptical, 1600–4040 long, 800–1120 wide. Oral sucker

148–195 by 144–210. Ventral sucker large, 400–608 by 352–597 wide, discoidal,
mostly non-muscular, filled with deeply staining nuclei dispersed over muscles, con-
tains ring of circular muscles near opening. Aperture reduced owing to presence
of several muscular papillae which present ribbed appearance. Prepharynx absent.
Pharynx globular. Caeca slender, terminate at level of posterior margin of ventral
sucker. Testes oval, symmetrical, separated by uterine coils, intercaecal, in posterior
part of forebody. Cirrus-sac arcuate, lies close to right caecum, encloses oval seminal
vesicle, long pars prostatica surrounded by numerous prostatic cells. Genital atrium
large, spacious lies medially at level of ovary, Genital pore situated slightly posterior
to intestinal bifurcation, surrounded by strong circular muscles. Accessory copula-
tory organ well developed, consists of thick muscular wall enclosing atrial papilla,
surrounded by numerous gland cells forming compact mass near base of accessory
organ. Ovary with three distinct lobes, situated slightly anterior to testis in space
enclosed by two caeca. Uterine coils fill post-testicular region. Eggs small, elliptical,
35–39 by 20–27. Vitelline fields extend from midlevel of ovary to middle of testes.
Remarks: Yamaguti (1952) erected the genus Prosogonarium with P. arii Yamaguti,
1952 as the type species. Two more species are known in the genus: P. plotosus
Madhavi, 1975 and P. angelae Cribb & Bray, 1994. Cribb & Bray (1994) provided
a review on the family Tandanicolidae. P. plotosus was differentiated from P. arii
Yamaguti, 1952 by the smaller and less muscular ventral sucker; absence of uterine
coils in the forebody, more numerous and more extensive vitelline follicles and
smaller eggs (66–81 by 27–33 in P. arii).
Chapter 8
Superfamily Haplosplanchnoidea Poche,
1926
Family: Haplosplanchnidae Poche, 1926
Haplosplanchnidae Poche, 1925 is a small family of digeneans with representatives

in the gastro-intestinal tract of marine, estuarine and freshwater fishes, mullets being
the preferred hosts. The haplosplanchnids have been considered as closely related to
the Haploporidae Nicoll, 1914 and were included along with this family in the super-
family Haploporoidea Nicoll, 1914. However, phylogenetic analysis of the digeneans
employing molecular tools revealed no relationship whatsoever between haploporids
and haplosplanchnids. In accordance with this, Olson et al. (2003) referred it to the
Haplosplanchnoidea Poche, 1926 and to a new suborder Haplosplanchnata Olson,
Cribb, Tkach, Bray & Littlewood, 2003. Reviews on the family were provided by
Skrjabin & Guschanskaja (1955), Nahhas et al. (1997) and Madhavi (2005).
The most reliable distinguishing feature of the family is the presence of a single
caecum lined by prominent cells. Other important characters are the smooth tegu-
ment, the presence of eyespot pigment, a single testis, the absence of a cirrus-sac, and
a pretesticular ovary.
Diagnosis: Body small to medium, with smooth tegument, often with eyespot pig-
ment granules and cervical gland cells in forebody. Oral sucker normal or replaced
by muscular disc with or without lobes. Ventral sucker pre-equatorial, equatorial or
near posterior extremity. Prepharynx short or absent. Pharynx present. Caecum sin-
gle, short or long, terminates blindly, lined by prominent cells. Testis usually single,
rarely double, in fore or hindbody. Cirrus-sac usually absent, rarely present. Seminal
vesicle tubular, prostatic complex well developed. Genital pore in forebody, median.
Ovary pretesticular. Uterine coils between testis and genital pore, may extend to
post-testicular area. Eggs large, embryonated or not. Vitellarium follicular or tubu-
lar, of variable extent. Seminal receptacle present. Excretory vesicle Y-shaped, pore
terminal. Type genus: Haplosplanchnus Poche, 1926.
© Crown 2018 109

https://doi.org/10.1007/978-94-024-1535-3_8
110 8 Superfamily Haplosplanchnoidea Poche, 1926
Twelve species belonging to 3 genera: Haplosplanchnus Poche, 1926 (Syn.

Laruea Srivastava, 1937), Schikhobalotrema Skrjabin & Guschanskaja, 1955 and
Hymenocotta Manter, 1961 and three subfamilies Haplosplanchninae Poche, 1926,
Schikhobalotrematinae Skrjabin & Guschanskaja, 1955 and HymenocottinaeYam-
aguti, 1971 have been recorded from Indian marine fish with the majority in the genus
Haplosplanchnus. Some of these species have since been synonymized retaining only
a few valid species.
Key to subfamilies:
1. Oral sucker modified into muscular flat or lobed disc .......... HymenocottinaeYa-
maguti, 1971
Oral sucker unmodified ............................................................. 2
2. Vitelline follicles few, restricted in distribution ……… Haplosplanchninae
Poche, 1926
Vitelline follicles numerous ........ Schikhobalotrematinae Skrjabin & Guschan-
skaja, 1955
Subfamily Haplosplanchninae Poche, 1926
Diagnosis: Body fusiform or triangular. Suckers and pharynx well developed. Ven-
tral sucker circular or discoidal with or without protrusible lips. Caecum single.
Testis usually single, rarely two. Cirrus-sac usually absent, rarely present. Genital
pore midway between suckers. Ovary pretesticular. Vitelline follicles few or numer-
ous. Uterus extensive. Type genus Haplosplanchnus Looss, 1902.
Genus Haplosplanchnus Looss, 1902
[Syn: Laruea Srivastava, 1937]
Diagnosis: Body elongate or Y-shaped. Ventral sucker usually larger than oral
sucker. Pharynx well developed. Caecum single, extent variable. Testis single, in
hindbody. Cirrus-sac absent. Seminal vesicle tubular, extends into hindbody. Genital
pore median in between pharynx and anterior margin of ventral sucker. Ovary pretes-
ticular. Seminal receptacle canalicular. Uterus pretesticular. Eggs oval. Vitelline fol-
licles in hindbody. Excretory bladder Y-shaped.
Type species: H. pachysomus (Eysenhardt, 1902) Looss, 1902
Ten species belonging to this genus have been described from India. Some of the
species have been invalidated. H. indicus Gupta & Ahmad, 1979 was synonymized
with H. purii Srivastava, 1937; H. orientalis Gupta & Ahmad, 1979, H. stunkardi
Gupta & Ahmad, 1879 and L. straightum Jehan, 1973 were synonymized with H.
caudatus Srivastava, 1937(See Bray, 1984). Nahhas et al. (1997) synonymized H.
otolithi Gupta & Ahmad, 1979 with H. pachysomus (Eysenhardt, 1902). This leaves
only five valid species: H. bengalensis Gupta & Ahmad, 1985, H. caudatus Srivas-
tava, 1937, H. guptai Ahmad, 1982, H. purii Srivastava, 1937 and H. vinodae Ahmad,
1985. Nahhas et al. (1997) considered H. caudatus, H. straightum, H. orientalis, H.
stunkardi and H. guptai as conspecific with H. pachysomus and H. otolithi with
Family: Haplosplanchnidae Poche, 1926 111
H. purii. Madhavi (1979a) and Bray (1984) observed extensive intraspecific varia-
tions in respect of certain characters in H. caudatus. Evidently the genus requires a
thorough revision employing molecular taxonomic tools.
Haplosplanchnus caudatus (Srivastava, 1937) Skrjabin & Guschanskaja, 1954
(Fig. 8.1)
[Syn. Laruea caudatum Srivastava, 1937; L. straightum Jahan, 1973; H. orientalis
Gupta & Ahmad, 1979; H. stunkardi Gupta & Ahmad, 1979; H. straightum (Jahan,
1973) Ahmad, 1985]
Hosts: Mugilidae: Liza vaigiensis (Quoy & Gaimard) (1); Mugil cephalus L, Liza
macrolepis (Smith), Valamugil cunnesius (2); Pristigasteridae: Ilisha elongata (Day);
Acanthuridae: Acanthurus triostegus (Linnaeus) (3)
Locality: PR (1,3); VSK (2)
References: 1. Srivastava (1937); 2. Madhavi (1979a); 3. Gupta & Ahmad (1979b)
Description: Body Y-shaped, with unequal arms and broad stem with rounded pos-
terior end, 1240–1700 long, 640–690 wide. Oral sucker 140–148 long, 190–210
wide. Ventral sucker deep, tubular extended as long arm of body. Prepharynx short.
Pharynx ovoid. Caecum simple, straight, extends to level of ovary. Genital pore in
angle of two arms, between bifurcation of Y. Testis single, oval, in mid to posterior
hindbody. Seminal vesicle tubular, extends from anterior end of testis to oesopha-
gus level. Pars prostatica oval, surrounded by large number of prostate gland cells.
Genital atrium tubular. Ovary oval, immediately anterior to testis. Seminal receptacle
oval. Uterus pretesticular. Eggs elongated, contain developing miracidia, 40–42 long,
18–20 wide. Vitellarium follicular, poorly developed, follicles few, extend between
middle of testis and blind end of intestinal caecum. Excretory bladder Y-shaped.
Excretory pore subterminal.
Fig. 8.1 Haplosplanchnus

caudatus (after Srivastava,
1937)
Remarks: Srivastava (1937) erected the genus Laruea with L. caudatus Srivastava,
1937 as the type species. Skrjabin & Guschanskaja (1955) synonymized the genus
Laruea with Haplosplanchnus. As a result, L caudatus became H. caudatus (Srivas-
tava, 1937) Skrjabin & Guschanskaja, 1955. Yamaguti (1971) retained Laruea as a
valid genus but Pritchard & Manter (1961) and Nahhas & Cable (1964) agreed with
Skrjabin & Guschanskaja in synonymizing the genus Laruea with Haplosplanch-
nus. Most of the subsequent authors also agreed with this synonymy (Madhavi,
1979; Bray, 1984). Nahhas et al. (1997) in a review of the family Haplosplanch-
nidae, regarded H. caudatus as a synonym of H. pachysomus (Eysenhardt, 1902).
H. caudatus is here retained as a distinct species. Reports of H. caudatus are mainly
from the Indian Ocean (Srivastava, 1937; Madhavi, 1979a; Gupta & Ahmad, 1979b;
Ahmad, 1985) and South African waters (Bray, 1984). The hosts are different species
of mullets.
Haplosplanchnus guptai Ahmad, 1985 (Fig. 8.2)
Host: Mugilidae: Mugil cephalus (L.).
Habitat: Small intestine.
Locality: Panjim coast, Goa. AS
Prevalence: 9 specimens from 2 hosts out of 76 examined.
Description: Body Y-shaped, with unequal arms, posterior end semi-spirally rolled
1553–1750 long, 780–880 wide. Tegument thin, aspinose. Eyespot pigment granules
absent. Oral sucker 120–150 long, 190–250 wide. Ventral sucker massive, elongate,
pedunculate (no measurements given). Prepharynx absent. Pharynx longer than wide.
Oesophagus, long. Intestinal caecum single, small. Testis single, elongate, postequa-
torial. Seminal vesicle tubular, pretesticular, not reaching testis. Prostatic vesicle sac-
cular, unipartite, surrounded by prostate gland cells. Genital atrium tubular. Genital
pore between the angle of the two arms. Ovary immediately pretesticular. Seminal

guptai (after Ahmad, 1985)
receptacle longitudinally elongate, dextral to ovary. Uterus descending to poste-

rior end of body. Eggs elongate, operculate, containing developing miracidia with
prominent eyespots, 50–65 long, 26–37 wide. Vitellarium consists of 7–10 follicles,
postovarian, grouped together.
Remarks: This species is characterized by possessing the following combination

of characters: the contiguous gonads, the uterus extending into the post-testicular
region, the vitellarium consisting of 7–10 follicles grouped together and the small
seminal vesicle stopping well anterior to testis.
Haplosplanchnus purii Srivastava, 1937 (Fig. 8.3)
[Syn. H. indicus Gupta & Ahmad, 1979; H. otolithi Gupta & Ahmad, 1979)
Host: Mugilidae: Liza vaigiensis (Quoy & Gaimard) (1); Liza macrolepis (Smith)
(2, 3)
Locality: PR (1); VSK (2, 3), BOB.
Distribution: China (TWS, SCS) [Wu et al. (2001); Ku & Shen (1964); Shen (1990)]
References: 1. Srivastava (1937); 2. Madhavi (1979a); 3. Rekharani & Madhavi
(1985)
Description: Body roughly triangular, 1500−2300 × 540−1200. Oral sucker cup-

shaped, 180−220 × 250−340. Ventral sucker at junction of first and second thirds of
body. Pharynx dome-shaped. Prepharynx, oesophagus short. Caecum tubular. Testis
single, ovoid, situated close to posterior extremity. Seminal vesicle narrow, tubular,
sinuous. Pars prostatica surrounded by prostatic cells. Genital atrium deep, narrow.
Genital pore on prominent conical papilla, halfway between suckers. Ovary closely
anterior to testis. Seminal receptacle small, spherical. Uterus well developed, occu-
pies all space between ovary and ventral sucker. Eggs large, numerous, 40−68 by
23−34, enclose miracidia with eyespots. Vitelline follicles few, scattered in region
between ovary and ventral sucker. Excretory bladder Y-shaped.

purii (after Srivastava, 1937)
Remarks: Srivastava (1937) reported the species from Mugil ‘waigiensis’ from
the Puri coast, Bay of Bengal. Later Madhavi (1979a) reported it from the mullets
Mugil cephalus and M. ‘waigiensis’ from the coast of Visakhapatnam. Subsequently
it has been reported from Liza macrolepis of Visakhapatnam (Rekharani & Madhavi,
1985), Liza tricuspidens from South Africa (Bray, 1984), from Liza vaigiensis and
M. cephalus from Suva coast, Fiji (Nahhas et al., 1997) and from M. cephalus from
the Yuedong area in Guangdong Province, China (Lu et al., 2001). Bray (1984) syn-
onymized H. bivitellosus Zhukov, 1971, H. cuneatus Tang & Lin, 1978, H. elongatus
Tang & Lin, 1978 and H. indica Gupta & Ahmad, 1979 with H. purii.
Haplosplanchnus vinodae Ahmad, 1985 (Fig. 8.4)
Host: Mugil cephalus (L.).
Locality: Panjim coast, Goa, AS
Prevalence: 7 specimens from 2 hosts out of 76 examined.
Description: Body plump, 1050–1230 long, 700–815 wide. Oral sucker cup-shaped,
120–160 long, 180–240 wide. Ventral sucker cup-shaped, slightly protuberant,
210–270 long, 215–260 wide. Sucker ratio 1: 1.08–1.19. Intestinal caecum single,
extends posteriorly but ending distinctly anterior to posterior end of body. Testis
single, oval, in posterior part of middle third of body. Seminal vesicle tubular. Pro-
static vesicle bipartite, both chambers of equal size, surrounded by prostate gland
cells. Genital atrium tubular. Genital pore between suckers. Ovary oval, ventro-lateral
to testis. Seminal receptacle oval, postovarian. Vitellarium poorly developed, with
8–15 pretesticular follicles. Uterus fills most of hindbody. Eggs enclose developing
miracidia with prominent eyespot, 68–82 × 36–46 µm wide.
Remarks: This species was characterized by the bipartite nature of the prostatic
vesicle and the limited number of vitelline follicles.

vinodae (after Ahmad, 1985)
Other species
Haplosplanchnus bengalensis Gupta & Puri, 1985
Host: Acanthuridae: Ctenochaetus strigosus (Bennett)
Locality: PR, BOB
No. One
Reference: Gupta & Puri (1985a)
Remarks: The validity of this species remains doubtful since the description of the
species is based on a single specimen and the characters used for separation of the
species from others are variable in nature and have little taxonomic significance.
Haplosplanchnus indica Gupta & Ahmad, 1979
[Syn. of H. purii Srivastava, 1937]
Host: Liza vaigiensis (Quoy & Gaim.).
Locality: Puri, Orissa.
Prevalence: One specimen from one host out of 41 examined.
Reference: Gupta & Ahmad (1979b).
Haplosplanchus orientalis Gupta & Ahmad 1979
[Synonym of H. caudatus Srivastava, 1937]
Host: Pristigasteridae: Ilisha elongata (Bennet).
Locality: PR, BOB
No. One
Reference: Gupta & Ahmad (1979b)
Haplosplanchnus otolithi Gupta & Ahmad, 1979
(Synonym of H. purii)
Host: Sciaenidae: Otolithus ruber (BI. Schn.).
Locality: PR, BOB.
No. Two
Haplosplanchnus stunkardi Gupta & Ahmad, 1979
[Syn. of H. caudatus]
Host: Acanthuridae: Acanthurus sandvicensis (Streets).
Location: Intestine.
Locality: PR, BOB.
Prevalence: Two specimens from one host out of 41 examined.
Subfamily Hymenocottinae Yamaguti, 1971
Diagnosis: Oral sucker replaced by muscular disc provided with lobes or not. Ventral
sucker in anterior half of body. Caecum single, long. Testis single. Cirrus-sac absent.
Ovary pretesticular. Vitellarium follicular, confined to hindbody. Uterus occupies
space between testis and ventral sucker. Eggs numerous, unembryonated. Type genus:
Hymenocotta Manter, 1961.
Genus Hymenocotta Manter, 1961
Diagnosis: Oral disc with six lobes. Pharynx large. Ventral sucker large, in anterior
quarter of body. Caecum long. Testis single, close to posterior extremity. Seminal
vesicle tubular. Genital pore ventral to pharynx. Ovary rounded, anterolateral to testis.
Vitellarium tubular, distributed in lateral and median fields between ventral sucker
and ovary. Metraterm well developed. Eggs large, thin-shelled, unembryonated. Type
species: H. mulli Manter, 1961.
Hymenocotta mulli Manter, 1961 (Fig. 8.5)
Host: Chirocentridae: Chirocentrus dorab (Forskål)
Locality: VSK, BOB
Distribution: Great Barrier Reef, Australia (Durio & Manter, 1968; Cribb et al., 1994)
Suva, Fiji (Nahhas et al., 1997); Japan (Machida, 1993); New Caledonia (Justine,
2010)
Description: Body elongate, 3040–4503 long. Oral sucker forms broad membra-
nous muscular disc with six short, rounded lobes, one pair anterior, two pairs latero-
posterior, posterior pair smaller and more or less pointed, with mouth as transverse
slit, oral disc 603–684 in diameter. Prepharynx covered by oral disc. Pharynx spher-
ical. Caecum single, reaches about 2/3 body length. Testis in posterior fourth of
body, elongate oval. Seminal vesicle tubular, long, almost straight, extends to almost
halfway from ventral sucker to testis. Pars prostatica short. Genital atrium very short.
Genital pore at level of posterior half of pharynx. Ovary rounded, overlapping ante-
rior end of testis. Seminal receptacle as large as ovary. Vitellarium forms broad tubes,
two on each side of body and two or three shorter median preovarian tubes, anterior
Fig. 8.5 Hymenocotta mulli

(after Hafeezullah, 1971,
courtesy Cambridge
University Press, J.
Helminthol, 45, p. 81)
extent about midway between ovary and ventral sucker, posterior extent midway
between testis and posterior end of body. Uterus between ovary and ventral sucker.
Eggs large, thin-shelled, 92–128 by 80–88.
Remarks: The genus Hymenocotta Manter, 1961 is very different from other hap-
losplanchnid genera in having a lobed muscular disc in place of an oral sucker. The
genus contains three species: H. mulli Manter, 1961, H. manteri Overstreet. 1969 and
H. mugili Wang & Wang, 1993. Hafeezullah (1971b) obtained only one specimen
identified as H. mulli Manter, 1961 from the gut of Chirocentrus dorab (Forskal)
from Visakhapatnam coast, Bay of Bengal. The single specimen measured 6,528
long and 1,020 wide. It has a ventral sucker well separated from the pharynx and
a seminal receptacle larger than the ovary. This constitutes the only report of the
species from India.
Hymenocotta mulli was originally described with a cirrus-sac (Manter, 1961),
but Cribb et al. (1994) examined the types and new material and could not detect a
cirrus-sac. In a recent molecular phylogeny, Huston et al. (2017) found Hymenocotta
to be sister to the other haplosplanchnids sequenced.
Subfamily Schikhobalotrematinae Skrjabin & Guschankaja, 1955
Diagnosis: Oral sucker simple. Ventral sucker in anterior half of body, with or
without paired appendages of various shapes. Caecum short. Testis single in middle
of hindbody or at level of ventral sucker. Cirrus-sac absent. Ovary close to testis.
Vitelline follicles profusely developed, distributed in fore and hindbody. Uterine
coils preovarian. Eggs few large, unembryonated. Type genus: Schikhobalotrema
Skrjabin & Guschanskaja, 1955.
Genus Schikhobalotrema Skrjabin & Guschankaja, 1955
Diagnosis: Forebody with eyespot pigment and gland cells. Ventral sucker may
be provided with one pair of digitiform appendages. Caecum of variable length.
Testis single, in posterior half of body. Seminal vesicle tubular. Prostatic complex
not well developed. Genital pore median, in forebody. Ovary pretesticular. Uterine
coils preovarian. Eggs not embryonated. Vitelline follicles large, numerous, extend
along each side of body from level of pharynx or ventral sucker to posterior end.
Type species; S. acutum.
Schikhobalotrema acutum (Linton, 1910) Skrjabin & Guschanskaja, 1955 (Fig. 8.6)
[Syn Deradena acuta Linton, 1910; Haplosplanchnus acutus (Linton, 1910)]
Host: Belonidae: Tylosurus crocodilus Péron & Lesueur; Hemiramphidae: Hemi-
ramphus marginatus (Forskål); Rhynchorhamphus georgei (Cuv. & Val.)
Locality: VSK, BOB
Distribution: Suva, Fiji (Nahhas et al., 1997); Atlantic coast of Panama (Sogandares
& Sogandares,1961); Republic of Panama (Caballero et al., 1953): Cubana (Perez
Vigueras 1956); Curacao (Nahhas & Cable, 1964); Galapagos (Manter, 1940); South
China Sea (Shen, 1990); Florida (Manter, 1937, 47; Overstreet, 1969): Brazil (Kohn
& Fernandes, 1982); Mediterranean (Fayek et al., 1990)
Reference: Madhavi (1979a)
Fig. 8.6 Schikhobalotrema

acutum (after Madhavi,
1979)
Description: Body elongate, widest at level of ventral sucker with smooth tegu-
ment and gland cells on either side of pharynx. Oral sucker subterminal. Ventral
sucker in anterior third of body, protrusible, with vertical aperture and one pair of
lateral lobes. Pharynx, oesophagus present. Caecum single, long, slender, termi-
nates at level of posterior margin of testis. Testis single, elliptical, in posterior half
of hindbody. Seminal vesicle tubular extends to posterior level of ventral sucker
or slightly beyond. Pars prostatica inconspicuous. Genital pore prominent, sinistral
to oesophagus. Ovary pretesticular, Uterus preovarian. Eggs fairly large, unembry-
onated. Vitellarium somewhat acinous, extends from level of oesophagus to posterior
end.
Remarks: Madhavi (1979a) collected flukes identified as S. acutum from Tylosurus

crocodilus, Hemiramphus marginatus and Rhynchorhamphus georgei from the Bay
of Bengal. She found her specimens to be morphologically similar to those reported
from other geographic regions in all respects, but differed in that they lacked sper-
matozoa both in the seminal vesicle and in the seminal receptacle and both these
structures are filled with a granular material. However, the eggs underwent nor-
mal development. She concluded that these forms may represent a parthenogenetic
species.
Chapter 9
Superfamily Hemiuroidea Looss, 1899
The superfamily Hemiuroidea comprises several families. Gibson & Bray (1979)
in their classification of the Hemiuroidea based on the functional morphology of
the adults, recognized 14 families. The first molecular phylogenetic study of the
Hemiuroidea, based on V4 variable domain of the 18SrRNA gene for 33 species
of Hemiuroidea, supported the monophyly of Hemiuroidea and revealed two main
groups, one group containing all members of Hemiuridae and Lecithasteridae and the
other one comprising the members of Derogenidae, Didymozoidae, Hirudinellidae,
Sclerodistomidae, Syncoelidae, and Accacoelidae (Blair et al., 1998). This classifi-
cation of Hemiuroidea adapted in ‘Keys to the Trematoda’ Volumes 1–3 is followed
here (Gibson, Bray, Jones 2002–2008).
Family Hemiuridae Looss, 1899
[Syn. Lecithochiriidae Luhe, 1901; Dinuridae Looss, 1907; Bunocotylidae Dollfus,

1950; Elytrophallidae Skrjabin & Guschanskaja, 1954; Biporidae Malhotra, Nanda,
Mukherjee, Ghosh, Sukul & Capoor, 1989]
Hemiurids are one of the most common and diverse groups of digeneans infecting
marine fishes, occurring usually in the stomach and occasionally in the bile duct. The
hemiurids can be easily identified by the presence of an ecsoma in most cases, the
thick smooth or plicated body wall, the prominent suckers, the compact or tubular
vitellarium and the extensive uterine coils. A cirrus-sac is absent but in its place there
is usually a sinus-sac enclosing the hermaphroditic duct or sinus organ. The organi-
zation of the genital complex as well as the presence or absence of an ecsoma are
important characters on which further classification of the family and differentiation
of genera is based. Useful reviews on hemiurids were provided by Gibson & Bray
(1979), Brooks (1985) and Gibson (2002).
© Crown 2018 119

https://doi.org/10.1007/978-94-024-1535-3_9
120 9 Superfamily Hemiuroidea Looss, 1899
Diagnosis: (Adapted from Gibson, 2002): Body fusiform to elongate. Ecsoma

present, reduced or absent. Body surface smooth or with annular plications. Preso-
matic pit occasionally present. Oral and ventral suckers well developed. Oesophagus
short. Caeca terminate blindly, rarely form cyclocoel. Testes two, tandem, oblique or
symmetrical, preovarian, in hindbody. Seminal vesicle tubular, saccular or constricted
into portions, muscular or thin-walled, in fore or hindbody. Pars prostatica of variable
length, tubular or vesicular, ejaculatory duct short. Sinus-sac usually well developed,
occasionally reduced or absent. Hermaphroditic duct usually enclosed within sinus-
sac. Sinus organ and genital atrium well developed or reduced or absent. Genital pore
mid ventral, at level of oral sucker or pharynx. Ovary oval, post-testicular. Mehlis’
gland postovarian. Laurer’s canal and seminal receptacle usually absent. Uterine sem-
inal receptacle and Juel’s organ present. Uterus coiled mainly in pre and postovarian
region of hindbody. Eggs numerous, small. Vitellarium with seven tubular branches
or paired oval or lobed masses mainly postovarian. Excretory vesicle Y-shaped, arms
united in forebody or not.
Subfamilies: The classification of hemiurids has remained unstable for many years.
Gibson (2002) discussed the various views proposed for the division of the family
Hemiuridae into subfamilies and proposed a system that is followed here. Altogether,
11 subfamilies are recognized. After excluding those that have no representatives in
Indian marine fish, the following key serves to differentiate the remaining eight
subfamilies:
1. Ecsoma present …………………………………………………………………2
Ecsoma absent …………………………………………………………………6
2. Prostatic vesicle present within sinus-sac ……………Lecithochiriinae Luhe,
1901
Prostatic vesicle absent …………………………………………………………3
3. Sinus-sac absent or poorly developed, ecsoma poorly developed …. Plerurinae
Looss, 1907
Sinus-sac present, well developed, ecsoma well developed ……………………4
4. Vitellarium entire or two lobed masses………..…………….……. Hemiurinae
Looss, 1899
Vitellarium seven distinct oval to tubular lobes………………………………….5
5. Seminal vesicle oval, with thick muscular wall, sinus organ delicate and amuscu-
lar ………………………………… Elytrophallinae Skrjabin & Guschanskaja,
1954
Seminal vesicle thin-walled, oval or constricted into partitions, sinus organ large
and muscular ………………………………………………………………. Din-
urinae Looss, 1907
6. Uterine seminal receptacle present, vitellarium one or two masses …………. 7
Blind seminal receptacle present …………………………Opisthadeninae Yam-
aguti, 1970
7. Body surface smooth, ridges present over body at level of oral and ventral suckers,
major part of uterus preovarian, vitellarium single…………………Bunocotyli-
nae Dollfus, 1950
Family Hemiuridae Looss, 1899 121
Body surface plicated, no ridges over body at level of oral and ven-
tral suckers, major part of uterus postovarian, vitellarium single or double
……..…Aphanurinae Looss, 1899
Subfamily Aphanurinae Looss, 1899
[Syn. Ahemiurinae Chauhan, 1954]
Diagnosis: Body small, ecsoma absent or extremely rudimentary. Body surface usu-
ally with plications. Ventral sucker in anterior half of body. Caeca end bindly. Testes
tandem to symmetrical, well posterior to ventral sucker. Seminal vesicle tubular in
forebody or saccular in hindbody. Pars prostatica tubular or vesicular, short or long.
Ejaculatory duct long, short or absent. Ovary oval, immediately post-testicular. Sem-
inal receptacle absent. Uterine seminal receptacle and Juel’s organ present. Major
part of uterus postovarian. Vitellarium one or two compact masses, immediately
posterior or lateral to ovary. Excretory arms united in forebody.
Type genus: Aphanurus Looss, 1907

Key to genera
Vitellarium composed of one distinct mass……………………………………….
Aphanurus Looss, 1907
Vitellarium composed of two masses……………………………..…………..
Ahemiurus Chauhan, 1954
Genus Ahemiurus Chauhan, 1954
Diagnosis: Body elongate, without ecsoma. Ventral sucker larger than oral sucker.
Caeca long. Testes symmetrical or oblique, seminal vesicle oval, in hindbody. Par
prostatica tubular, long. Ejaculatory duct long. Sinus-sac present. Genital pore at
base of oral suker. Ovary post-testicular, separated from testis by uterine oils. Vitel-
larium two oval compact masses, symmetrical, postovarian. Type species: A. karachi
(Srivastava, 1937).
Ahemiurus karachii (Srivastava, 1937) Chauhan, 1954 (Fig. 9.1)
[Syn. Sterrhurus karachii Srivastava, 1941]
Host: Clupeidae: Sardinella longiceps Valenciennes
Locality: Arabian Sea, Karachi (Pakistan).
Reference: Srivastava (1941b)
Description: Body narrow with prominent transverse annulation all over, 1400 ×
200. Ecsoma extremely rudimentary. Oral sucker 80 in diameter. Intestinal caeca
extend to posterior end, never extend into tail. Ventral sucker situated at end of first
quarter of body, 150 (diameter). Testes small, situated symmetrically in anterior part
of posterior half of body. Seminal vesicle pyriform, thin-walled, situated medially,
just anterior to testes. Pars prostatica long, narrow, surrounded by large number of
prostate gland cells. Hermaphroditic duct spindle-shaped, enclosed in a sinus-sac.
Genital pore at level of anterior margin of pharynx. Ovary situated just anterior to
Fig. 9.1 Ahemiurus karachii

(after Srivastava, 1937)
vitellarium. Vitellarium a pair of elongated, oval, compact bodies, situated symmet-

rically at junction of last two-thirds of body. Uterus occupies whole of intercaecal
space. Excretory bladder Y-shaped, main stem bifurcating behind ventral sucker; two
arms uniting dorsally to oral sucker.
Remarks: Srivastava included this species in the genus Sterrhurus as S. karachii

Srivastava, 1941. However, Manter (1947) suggested its inclusion in the genus Dero-
genes. According to him the tail is too rudimentary to be recognized as an ecsoma.
Chauhan (1954b) erected the genus Ahemiurus with A. karachii (Srivastava, 1941b)
as the type species. The genus Ahemiurus Chauhan, 1954, was differentiated from the
closely related Aphanurus Looss, 1909, in the relative positions of the genital pore
and seminal vesicle and in possessing two compact, elongated, oval, symmetrically
placed vitelline masses. A. karachii is the only species in the genus so far.
Genus Aphanurus Looss, 1907

[Syn. Chauhanurus Skrjabin & Guschanskaja, 1954; Minutocauda Srivastava &
Sahai, 1977; Helaphanurus Slusarski, 1957; Reference: Gibson (2002)]
Diagnosis: Body small. Tegument plicated. Testes oblique, symmetrical or tandem.

Seminal vesicle oval, in hindbody. Pars prostatica long, tubular. Ejaculatory duct
long or short. Sinus-sac tubular. Vitellarium a single large mass, postovarian.
Type species: A. stossichi (Monticelli, 1891)

Five species belonging to this genus have been recorded from clupeoid fishes of India,
of which A. microrchis Chauhan, 1954, is considered as a synonym of A. stossichii
(Monticelli, 1891) (Zhukov, 1977).
Aphanurus dussumierii Hussain, Rao & Shyamasundari, 1984 (Fig. 9.2)
Host: Clupeidae: Dussumieria hasseltii Bleeker
Fig. 9.2 Aphanurus

dussumierii (after Hussain,
Rao & Shyamasundari,
1984)
Locality: VSK, BOB

Number: 4
Reference: Hussain et al. (1984b)
Description: Body elongated, 880–940 long, 210–240 wide. Oral sucker cup-
shaped, 40–50 × 60–64 in size. Ventral sucker large, 170–180 by 140–144, situated
160–180 from anterior end. Sucker ratio 1:3.5. Caeca long. Testes 2, oval, tandem,
contiguous situated in middle third of body. Seminal vesicle long, situated obliquely
between ventral sucker and testes. Pars prostatica long, tubular, surrounded by pro-
static cells, continues as ejaculatory duct, joins metraterm close to anterior border
of ventral sucker. Hermaphroditic duct long, enclosed in elongated sinus-sac. Gen-
ital pore median, posterior to oral sucker. Vitellarium single, compact, postovarian.
Uterus extends to near posterior extremity. Eggs oval, 4–7 × 2–4 (?)
Remarks: This species was differentiated from A. harengulae Yamaguti, 1938, by

the larger ovary compared with the testes, by the length of the seminal vesicle and
the absence of body plications and from A. stossichii in the larger sucker ratio. But
Nahhas & Sey (2002) suggested that the relative sizes of the vitellarium, the ovary
and the testes are not reliable species characteristics. A. dussumierii differs from the
other species only in its larger sucker ratio. The eggs are unusually small.
Aphanurus harengulae Yamaguti, 1938 (Fig. 9.3)

Hosts: Mugilidae: Mugil cephalus L.; Valamugil cunnesius (Val.)
Locality: VSK, BOB
Distribution: Lake Hamana, Japan (Yamaguti, 1938); Celebes (Yamaguti, 1953)
Reference: Rekharani & Madhavi (1985)
Fig. 9.3 Aphanurus

harengulae (after
Yamaguti,1938)
Description: Body cylindrical, 700–880 long 140–180 wide, Tegumental annula-

tions conspicuous. Oral sucker 42–51 × 51–56. Caeca simple, terminate near pos-
terior extremity. Ventral sucker 110–126 in diameter. Sucker ratio 1:2.3–3.8. Testes
rounded, diagonal, in middle third of body. Seminal vesicle fusiform, with thick
muscular wall, extends obliquely from immediately behind ventral sucker to one
side of anterior testis. Pars prostatica runs forward curves dorsally to ventral sucker,
unites with uterus immediately ventral to intestinal bifurcation. Hermaphroditic duct
80–100 long. Genital pore median, ventral at level of pharynx. Uterus fills entire
hindbody. Eggs 16–19 by 7–8 in size. Vitelline mass entire or slightly indented,
contiguous with ovary.
Remarks: This species differs from A. stossichii in the egg size being much smaller
(26–28 × 13–15 in A. stossichii) and in the much longer seminal vesicle with a mus-
cular wall. Nahhas & Sey (2002) regarded the species as a synonym of A. stossichii.
Aphanurus microrchis Chauhan, 1945
Hosts: Mugilidae: Liza parsia (Hamilton)
Locality: BOM, AS
Distribution: Palawan island (Philippines) NE Atlantic region (off Spain), the
Mediterranean (off Spain and Turkey) and the Black Sea (off Bulgaria)
Reference: Chauhan (1945b)
Remarks: Conisdered synonym of A. stossichii (Reference: Zhukov, 1977).
Aphanurus stossichii (Monticelli, 1891) Looss, 1907 (Fig. 9.4)
[Syns. Sterrhurus monolecithus Srivastava, 1941; Aphanurus monolecithus (Srivas-
tava, 1941) Manter, 1947; Aphanurus microrchis Chauhan, 1945]
Fig. 9.4 Aphanurus

stossichii (after Looss, 1908)
Hosts: Clupeidae: Tenualosa ilisha (Hamilton); Clupea fimbriata (Val); Hilsa toli
(Val.); Ilisha elongata (Anonymous [Bennett], 1830)
Locality: PR, DC, BOB
Distribution: Mediterranean sea (Monticelli, 1891), Adriatic sea (Looss, 1907),
Atlantic ocean at the coast of Spain, Pacific coast of Japan (Yamaguti, 1938); Dni-
ester Estuary (Butskaya, 1952), Sea of Asov (Naidenova 1970); Palawan Island,
Philippines (Fischthal & Kuntz, 1964), Lake Hamana, Japan; Kuwait coast, Arabian
Sea (Nahhas & Sey, 2002), Off Spain, North Atlantic, Mediterranean off Turkey, and
Black sea off Bulgaria (Kostadinova et al., 2004).
Reference: Srivastava (1941b); Zhukov (1977), Hafeezullah & Dutta (1998)
Description: Body small, elongate, 704–928 long, 208–240 wide. Tegument with
annular plications all over. Oral sucker small, 23–27 by 23–36. Ventral sucker large,
66–72 near anterior end of body. Testes symmetrical, equatorial. Seminal vesicle
large, pyriform, in hindbody. Pars prostatica long, tubular, surrounded by prostatic
cells. Ejaculatory duct long. Sinus-sac present anterior to ventral sucker, encloses
hermaphroditic duct. Genital pore near bifurcation. Ovary small, post-testicular. Sem-
inal receptacle uterine. Vitellarium single, large ovoid mass, posterior to ovary. Uterus
reaches to caecal ends. Eggs 26–28 by 13–15.
Remarks: Zhukov (1977) reported the species from Ilisha elongata, Engraulis tol-
era and Coilia ramcarati (Hamilton) from the Hughli Estuary. He synonymized A.
microrchis Chauhan, 1945 with A. stossichii. Hafeezullah and Dutta (1998) consid-
ered A. microrchis as distinct from A. stossichii in the following characters: Position
of ventral sucker close to oral sucker, position of ovary immediately post-equatorial,
extracaecal position of sinus-sac, its posterior extent up to posterior margin of phar-
ynx and genital pore ventral to oral sucker. Manter (1947) transferred Sterrhurus
monolecithus Srivastava, 1941, from Hilsa ilisha from Puri, Bay of Bengal to the
genus Aphanurus as A. monolecithus. Chauhan (1954) regarded this species as a syn-

onym of A. stossichii, and expressed the view that the very rudimentary tail described
by Srivastava for Sterrhurus monolecithus is probably not an ecsoma but a temporary
fold of body wall. The species has recently been described in detail by Kostadinova
et al. (2004).
Aphanurus tuberculatus Hafeezullah, 1981 (Fig. 9.5)

Host: Clupeidae: Hilsa sinensis (L.); Sardinella fimbriata (Val.)
Locality: BOM, AS
Number: 18
Reference: Hafeezullah (1981)
Description: Body 728–854 long; 172–189 wide. Tegument moderately thick, with
plications more conspicuous in anterior region, extending to vitelline zone on ventral
side. Ventral sucker 112–122 in diameter, spherical, prominent. Oral sucker 52 in
diameter. Sucker ratio 1:2.25. Intestinal bifurcation anterior to ventral sucker. Caeca
simple reaching posterior end of body. Testes globular, almost equatorial. Seminal
vesicle elongated, fusiform with thick muscular wall, posterior to ventral sucker,
extends to anterior margin of posterior testis. Pars prostatica dorsal to ventral sucker,
surrounded by prostatic cells at proximal and distal ends, naked in middle. Sinus-
sac tubular, narrow, reaching from near anterior border of ventral sucker to posterior
border of oral sucker, encloses straight hermaphroditic duct and knob-like sinus organ
beset with fine tubercles, projecting through genital pore. Genital pore ventral to
oral sucker. Ovary entire, median, post-testicular. Uterus reaches posteriorly beyond
caeca. Eggs 14–18 by 8. Vitellarium globular, indented anteriorly.
Fig. 9.5 Aphanurus

tuberculatus (after
Hafeezullah, 1981, Bull.
Zool. Soc. India, 4, 114)
Remarks: A. tuberculatus was differentiated from all the other members of Apha-
nurus in the presence of a sinus-organ bearing fine tubercles (Hafeezullah, 1981).
Key to Indian species of the genus Aphanurus Looss, 1907
1. Sinus organ with tubercles ……………….……………A. tuberculatus Hafeezul-

lah, 1981
Sinus organ without tubercles ………………………………………………… 2
2. Ovary larger than testes ……..…….. A. dussumierii Hussain, Rao, Shyamasun-
dari, 1984
Ovary smaller than testes …………………………………………………….. 3
3. Seminal vesicle of medium size, oval, eggs large 26–28 × 13 …………………
A. stossichii (Monticelli, 1891)
Seminal vesicle large, elliptical, eggs 16–19 × 7–8 ……. A. harengulae Yam-
aguti, 1938
Subfamily Bunocotylinae Dollfus, 1950
Diagnosis: Body small, elongate, body surface smooth. Ridges present around body
at level of oral sucker and posterior margin of ventral sucker. Transverse septate
partitions of body may occur. Ventral sucker in anterior half of body. Caeca end
blindly or form cyclocoel. Testes tandem or oblique. Seminal vesicle saccular, oval,
in forebody or dorsal to ventral sucker. Pars prostatica short, tubular, or vesicular.
Sinus-sac absent or small and tubular. Ovary oval, uterine seminal receptacle present.
Vitellarium a single unlobed mass, immediately postovarian.
Type genus: Bunocotyle Odhner, 1928
Genus Saturnius Manter, 1969
Diagnosis: Body small, with two circular flanges, at level of oral sucker and just
posterior to ventral sucker. Caeca end blindly. Hindbody divided internally into four
segment-like parts by partitions formed of compact fibres. Testes tandem, in separate
compartments. Seminal vesicle saccular. Pars prostatica short. Sinus-sac pyriform.
Ovary single. Vitellarium in fourth or last segment. Uterus largely divided into seg-
ments by partitions. Excretory vesicle with short arms uniting dorsal to pharynx.
Type species: S. segmentatus Manter, 1969
Saturnius segmentatus Manter, 1969 (Fig. 9.6)

Hosts: Mugilidae: Mugil cephalus L.
Locality: VSK, BOB
Distribution: Queensland, Australia, Russian coast of Sea of Japan (Manter, 1969;
Blasco-Coasta et al. (2008)
Fig. 9.6 Saturnius

segmentatus (after Rekharani
& Madhavi 1985)
Description: Body elongate, 661–866, width 148–159. Ecsoma absent. Preoral lobe
well developed. Oral sucker 62–65 wide. Ventral sucker 93–101 wide. Sucker ratio
1:1.44–1.6. Two flanges or ridges encircle body, one at posterior edge of mouth,
another just posterior to ventral sucker. Lateral edge of body at level of posterior edge
of ventral sucker elevated to a nipple shape and provided with concentric muscles.
Caeca extend to near posterior end of body, end blindly. Hindbody divided internally
by four transverse, fibrous septa-like partitions resulting in four tandem segments of
increasing size. Partitions consist of numerous thick fibres. Testes smooth, tandem in
separate segments, anterior testis between second and third partitions, posterior testis
between third and fourth partitions. Seminal vesicle at anterior edge of ventral sucker,
saccular, recurved or bent once, prostatic cells few. Sinus-sac pyriform with small
spherical basal sac containing sperm cells and thick-walled genital sinus about half
length of sac. Genital pore a transverse slit near level of intestinal bifurcation. Ovary
globular, immediately posterior to fourth partition. Uterus divided by partitions into
five sacs. Eggs large 26–28 by 12–13. Vitellarium single, immediately postovarian,
larger than ovary.
Remarks: This bunocotyline hemiurid possesses many unique features like the
presence of partitions dividing the body into compartments and the uterus sepa-
rated into sac-like compartments. The species shows specificity towards mullets. It
was recorded from Mugil cephalus L from Queensland, Australia (Manter, 1969a);
Visakhapatnam coast, Bay of Bengal (Rekharani & Madhavi, 1985) and the Russian
coast of the Sea of Japan (Blasco-Costa et al. 2008).
Saturnius valamugilis Rekharani & Madhavi, 1985]

Host: Mugilidae: Valamugil cunnesius (Val.)
Locality: VSK, BOB
Remarks: Blasco-Costa et al. (2008) regarded this as species inquirendum.
Subfamily Dinurinae Looss, 1907
Diagnosis: Ecsoma well developed. Body surface plicated or smooth. Testes sym-
metrical to tandem. Seminal vesicle thin-walled, oval, tubular or constricted into two
or four portions, in forebody or in hindbody. Pars prostatica tubular to vesicular, short
or long. Sinus-sac present, small or large, not enclosing prostatic vesicle. Permanent
sinus organ large or small or apparently absent. Genital atrium well developed. Ovary
oval or reniform, or lobed. Vitellarium with seven tubular lobes, three on one side
and four on the other. Excretory arm united or not in forebody. Type genus: Dinurus
Looss, 1907.
Key to Genera
1. Seminal vesicle constricted into portions; permanent sinus organ present

………….2
Seminal vesicle not constricted into portions, sinus organ absent .....…..….... 4
2. Body surface with plications ……………………………………………Dinurus
Looss, 1907
Body surface without plications…………………….………………………….3
3. Seminal vesicle dorsal or postero-dorsal to ventral sucker…… Erilepturus Wool-
cock, 1935
Seminal vesicle in forebody………………………………………Qadriana
Bilqees, 1971
4. Ecsoma large, seminal vesicle oval or elongate
oval……….…………………….……..5.
Ecsoma normal, seminal vesicle tubular and sinuous…….. Tubulovesicula Yam-
aguti, 1934
5. Seminal vesicle in forebody……………………………. Allostomachicola Yam-
aguti, 1958
Seminal vesicle in hindbody……………………………….. Stomachicola Yam-
aguti, 1934
Genus Allostomachicola Yamaguti, 1958
Diagnosis: Soma long. Ecsoma enormous. Body surface smooth. Seminal vesicle
elongate oval, partly in forebody. Pars prostatica short, vesicular. Sinus-sac present,
small, oval. Permanent sinus organ absent. Ovary reniform or indistinctly lobed.
Uterus extends well into ecsoma. Excretory bladder Y-shaped, arms uniting near
middle of oral sucker. Type species: Allostomachicola secundus (Srivastava, 1937).
Allostomachicola lepturusi (Gupta & Gupta, 1976) Gupta &Singh, 1981 (Fig. 9.7)
[Syn. Stomachicola lepturusi Gupta & Gupta, 1978]
Host: Congridae: Uroconger lepturus (Richardson)
Locality: Kerala Coast
Number: One
Reference: Gupta, R.C. & Gupta, S.P. (1976); Gupta, P.C. & Singh (1981)
Description: Body elongate, with ecsoma nearly three times longer than body,
47400 long, 6000 wide. Oral sucker 1520 long 1600 wide. Caeca prominently shoul-
dered at pharyngeal level, terminate at posterior extremity. Ventral sucker 2,820 long,
2,760 wide. Genital pore post-bifurcal. Testes oval, diagonal, subequal, just poste-
rior to ventral sucker Seminal vesicle partly anterior to ventral sucker. Pars prostatica
bulbous, straight, surrounded by large number of gland cells. Hermaphroditic duct
enclosed in saccular sinus-sac. Ovary bean-shaped. Uterus runs posteriorly up to
one-third length of ecsoma. Eggs 28–30 × 20–27. Vitellarium composed of two long
coiled tubes overlapping caeca, from level of posterior testis to posterior margin of
ovary.
Remarks: This species closely resembles A. secundus (Srivastava, 1939), but differs
from it in having an entire ovary instead of lobed, in the extension of the vitelline
tubes up to the posterior margin of the ovary instead of to the testes and in the
extension of the uterine coils and seminal vesicle. Additional specimens need to be
examined to confirm the validity of the species.
Fig. 9.7 Allostomachicola

lepturusi (after Gupta &
Gupta, 1976)
Allostomachicola rauschi (Gupta & Ahmad, 1978) Gupta &Singh, 1981 (Fig. 9.8)
[Syn. Stomachicola rauschi Gupta & Ahmad, 1978]
Host: Chirocentridae: Chirocentrus dorab (Forsskål)
Locality: PR, BOB
Number: One
Reference: Gupta, V. & Ahmad (1978b); Gupta, P.C. & Singh (1981)
Description: Body elongate, with ecsoma more than twice length of soma, 15240
long, 1560 wide. Oral sucker 420 long, 510 wide. Intestinal caeca prominently shoul-
dered at pharyngeal level, terminate at posterior extremity. Ventral sucker larger than
oral sucker. 970 long, 1030 wide. Genital pore posterior to oral sucker. Testes entire,
oval, diagonal, subequal. Seminal vesicle elongate, saccular extends to middle of
ventral sucker. Pars prostatica bulbous, surrounded by many prostate gland cells.
Hermaphroditic duct in elongate muscular sinus-sac, continued into tubular genital
atrium. Ovary tetra-lobed. Uterus forming intercaecal and extracaecal loops extends
into half of ecsoma. Eggs small, 18–21 long, 9–12 wide. Vitellarium composed of
two long tubes, extend from level of anterior margin of right testis up to end of soma.
Remarks: This species differs from all the species within the genus except A. secun-
dus (Sivastava, 1939) in having two vitelline tubes and in the extension of the seminal
vesicle to the middle of the ventral sucker. It differs from A. secundus in having a tetra-

rauschi (after Gupta &
Ahmad, 1978)
lobed ovary rather than a three-lobed, in having the genital pore posterior to the oral
sucker rather than posterior to the pharynx.
Allostomachicola secundus (Srivastava, 1939) Yamaguti, 1958 (Fig. 9.9)
[Syns Stomachicola secundus Srivastava, 1937; Pseudostomachicola secundus (Sri-
vastava, 1937) Skrjabin & Guschanskaja, 1954].
Host: Hemiramphidae: Hemiramphus limbatus (Day) (1); Chirocentridae: Chirocen-
trus dorab (Forsskål) (2, 3)
Locality: BOB
Reference: 1. Srivastava (1939d); 2. Gupta, P.C. & Singh (1981); 3. Gopal et al.
(1984)
Description: Body elongate, with ecsoma longer than soma, 16580 long (including
ecsoma), 1520 wide; ecsoma 9900 long, 1150 wide. Oral sucker subspherical, 620
long, 640 wide. Intestinal caeca simple, prominently shouldered at pharyngeal level,
extend to posterior extremity of ecsoma. Ventral sucker larger than oral sucker, 1060
long, 1030 wide. Sucker ratio 1:1.6. Genital pore, just post-bifurcal. Testes entire,
unequal, asymmetrical, in anterior hindbody. Seminal vesicle elongate, saccular,
extends posteriorly to midlevel of ventral sucker. Pars prostatica bulbous, surrounded
by many prostate gland cells. Hermaphroditic duct in elongate, muscular sinus-
sac, continued into tubular thick-walled genital atrium. Ovary trilobed, median, in
posterior half of soma. Uterus long, coiled, extends into anterior three-fifth of ecsoma.
Eggs small, oval, numerous, 17–21 long, 7–9 wide. Vitellarium tubular, consisting
of two long coiled tubes, extends laterally from level just anterior to posterior margin
of left testis to some distance in ecsoma.

secundus (after Srivastava,
1939d)
Remarks: This species was initially included in the genus Stomachicola by Srivas-
tava (1939d) as S. secundus Srivastava, 1939 collected from Hemiramphus limbatus
but Yamaguti (1958) erected a new genus Allostomachicola for this species and
differentiated it from Stomachicola by the location of the seminal vesicle in the fore-
body, the bulbous pars prostatica and the lobed ovary. Subsequently, two more Indian
species have been added to the genus: A. lepturusi (Gupta & Gupta, 1974) and A.
rauschi (Gupta & Ahmad, 1974).
Key to species of the genus Allostomachicola:
1. Seminal vesicle reaches to middle of ventral sucker ………….….………… 2

Seminal vesicle entirely in forebody…………………. A. lepturusi Gupta &
Gupta, 1974
2. Ovary tetra-lobed………………………..…………. A. rauschi (Gupta &
Ahmad, 1978)
Ovary 3-lobed……………………………………………A. secundus (Srivas-
tava, 1939)
Genus Dinurus Looss, 1907
Diagnosis: Characterized by thin-walled 3–4 partitioned seminal vesicle, vitellar-

ium with seven digitiform lobes, sinus-sac and sinus organ present, long pars pro-
statica surrounded by dense or sparcely distributed gland cells.
Seven species of this genus have been recorded from India: D. barbatus Cohn, 1902,
D. breviductus Looss, 1907, D. hippuri Nadakal, Kappikarayil & Jacob, 1991, D.
ivanosi Rekha & John, 2004, D. longisinus Looss, 1907, D. thapari Gupta & Gupta,
1980 and D. tornatus Rudolphi, 1819. Except for D. thapari, all the species have been
recorded from Coryphaena hippurus L. For D. thapari the host is Thryssa malabari-
cus (Bloch) (= Engraulis malabaricus). The various species have been distinguished
by differences in sucker proportions, number of chambers in the seminal vesicle,
the size of the sinus organ, the number of vitelline lobes and extent of the uterus
into the ecsoma. Opinions differ as to the validity of various species of the genus.
Dawes (1946, 47) considered D. barbatus, D. breviductus and D. longisinus as syn-
onyms of D. tornatus Rudolphi, 1819. Manter (1947) considered all the species as
distinct. Gupta & Gupta (1980) considered D. breviductus as a synonym of D. barba-
tus, Bray (1990) regarded D. hippuri as a synonym of D. longisinus although Rekha
& John (2002) claim it to be a distinct species.
Dinurus barbatus Cohn, 1902 (Fig. 9.10)

Hosts: Coryphaenidae: Coryphaena hippurus L.
Locality: Kerala, AS
Distribution: Tortugas, Florida (Manter, 1947) Puerto Rico (Siddiqi & Cable, 1960;
Dyer et al. 1997)
Reference: Rekha & John (2003)
Fig. 9.10 Dinurus barbatus

(after Looss, 1908)
Description: Body stout and straight. Soma 4160 long, ecsoma 3800 long. Tegu-
mental plications prominent. Tuft of 18–24 tubular in tegumentary outgrowths
present ventrally behind oral sucker. Oral sucker circular, 385 × 372. Ventral sucker
877 × 811. Sucker ratio 1:2.2. Preoral lobe present. Papillae prominent around oral
sucker. Testes globular, tandem, located close together. Seminal vesicle thin-walled,
trilobed, lobes equal in size. Pars prostatica long weakly coiled, sparsely covered
with small prostatic cells except at ends. Sinus-sac very short. Genital atrium short.
Ovary ovoid. Uterus short, descending into part of ecsoma. Eggs 15–19 × 19–22.
Vitellarium seven digitiform lobes, two of them directed forward.
Remarks: This species is characterized by the tuft of tegumentary papillae occurring

behind the oral sucker on the ventral side, three-lobed seminal vesicle and the two
anteriorly directed vitelline lobes.
Dinurus breviductus Looss, 1907 (Fig. 9.11)

Host: Coryphaenidae: Coryphaena hippurus L.
Distribution: Tortugas, Florida (Manter, 1947) Puerto Rico (Siddiqi & Cable, 1960,
Dyer et al., 1997) and records from elsewhere in the Atlantic, Indian and Pacific
Oceans.
Description: Body long, slender, 1233 by 832. Soma longer than ecsoma. Sucker
ratio 1:2. Caeca extend to posterior end of ecsoma. Testes of equal size and shape.
Seminal vesicle trilobed, middle lobe smallest, pars prostatica more or less straight,
Fig. 9.11 Dinurus

breviductus (after Looss,
1908)
prostatic cells numerous closely packed leaving the ends free. Sinus-sac situated
anterior to ventral sucker. Uterus passes into ecsoma. Eggs 25 × 20. Vitelline tubes
seven, two of them directed anteriorly.
Remarks: This species differs from other species of Dinurus by possessing the fol-
lowing combination of characters: Soma longer than ecsoma, sucker ratio 1:2, sinus-
sac situated anterior to the ventral sucker and the trilobed seminal vesicle with a
smaller middle lobe.
Dinurus hippuri Nadakal, Kappikarayil & Jacob, 1991

Reference: Nadakal et al. (1991)
Remarks: This species is synonymized with D. longisinus by Bray et al. (1993)
Dinurus ivanosi Rekha & John, 2004 (Fig. 9.12)

Description: Body curved. Tegument with plications, 3040–3470 long. Ecsoma

5080–5187 long. Oral sucker terminal, preoral lobe absent. Sucker ratio 1:1.54.
Caeca long, terminate in ecsoma. Testes oblique. Seminal vesicle well developed,
adjacent to anterior testis, thick-walled, trilobed, distal lobe smaller than middle lobe.
Fig. 9.12 Dinurus ivanosi

(after Rekha & John, 2004)
Pars prostatica more or less straight, prostatic cells ensheath middle of duct leaving
the ends free. Sinus-sac very small, far anterior to ventral sucker. Ovary 172–159 ×
202–266. Uterus descends to almost three fourth length of ecsoma. Eggs 15 × 9.
Vitellarium seven, tubular, uncoiled, two directed anteriorly.
Remarks: The most important features of the species are the general shape, and
the proportions of the body, the terminal position of the oral sucker, the elongated
pharynx, the small distal lobe of the seminal vesicle, the larger anterior testis, the
smaller eggs and the shorter and slender vitelline lobes.
Dinurus longisinus Looss, 1907 (Fig. 9.13)
[Syn. D. hippuri Nadakal, Kappikarayil & Jacob, 1991]
Distribution: Indian Ocean, Red Sea, Gulf of Mannar, Arabian Sea, Bay of Bengal,
Galapagos (Manter, 1940); Western Australia (Bray, 1990c)
Reference: Rekha & John (2003); Parukhin (1976, 89); Zhukov (1977)
Description: Soma longer than ecsoma. Plications crenulate, reaching to near poste-
rior end. Preoral lobe absent. Sucker ratio 1:1.75–2.00. Seminal vesicle thin-walled,
three-lobed, with large middle lobe. Sinus-sac long, extends to midlevel of ventral
sucker. Vitellarium seven-lobed, three directed anteriorly. Eggs 17 × 11.
Remarks: Bray et al. (1993) gave a detailed account of its diagnostic features, hosts
and geographic distribution. It is a stenoxenic species with 60% of its records from
C. hippurus. Distribution mostly Indo-Pacific with few records from Atlantic.
Fig. 9.13 Dinurus
longisinus (after Looss,
1908)
Dinurus thapari Gupta & Gupta, 1980

Host: Engraulidae Thryssa malabaricus (Bloch)
Reference: Gupta, S.P. & Gupta, R.C. (1980b)
Description: Body 4980–5000 long, 940–780 wide. Ecsoma longer or shorter than
soma, 3600–6540 long, 780 wide. Oral sucker 44–50 long, 56–60 wide. Ventral
sucker 43–66 long, 51–57 wide. Genital pore submedian at mid-level of oral sucker or
posterior to it. Testes tandem, contiguous or nearly contiguous. Seminal vesicle pre-
testicular, constricted into three portions, middle lobe smaller. Pars prostatica tubular,
winding, surrounded by large number of prostate gland cells. Genital atrium short.
Ovary ovoid. Vitellarium seven long, narrow winding tubules extend from posterior
testis to some distance into posterior half of soma. Uterus extends posteriorly some
distance into ecsoma. Eggs oval, 13–20 long, 8–11 wide.
Remarks: Details of the sinus organ and sinus-sac were not provided in the descrip-
tion. Without these details, it is difficult to assign it to the genus Dinurus. Moreover,
the occurrence of a member of this genus in an engraulid fish is unusual.
Dinurus tornatus Rudolphi, 1819 (Fig. 9.14)
Distribution: Tortugas, Florida (Manter, 1947) Puerto Rico (Siddiqi & Cable, 1960,
Dyer et al., 1997).
Fig. 9.14 Dinurus tornatus
(after Looss, 1908)
Remarks: Resembles D. longisinus in having a long sinus-sac, but differs in the

sucker ratio 1:1.14–1.27 as against 1:1.75–2.00. Further differs from D. longisinus
in that the body annulations begin dorsally at about the level of the posterior border
of ventral sucker and the hermaphroditic duct extends past the posterior border of
the ventral sucker. Most of the records are from the Atlantic Ocean.
*Key to Indian species of the genus Dinurus

1. Sinus organ long, extends to midlevel or posterior margin of ventral sucker
……….. 2
Sinus organ short, not reaching ventral sucker…………………………….3
2. Seminal vesicle trilobed with large middle lobe, sucker ratio 1:2 or
more…………………………….……………………………….……………D.
longisinus Looss, 1907
Seminal vesicle trilobed with small midde lobe, sucker ratio less than
1:2……………………………………………………………..….…D. tornatus
Rudolphi, 1819
3. Large integumental outgrowths present ventral to oral sucker.… D. barbatus
Cohn, 1902
No integumental outgrowths ventral to oral sucker ………………….………. 4
4. Soma longer than ecsoma, sucker ratio 1:2 …………….… D. breviductus Looss.
1907
Soma shorter than ecsoma, sucker ratio less than 1:2 … D. ivanosi Rekha &
John, 2004
*D. thapari is not included in the key.
Genus Erilepturus Woolcock, 1935

[Syns Uterovesiculurus Skrjabin & Guschanskaja, 1954; Exoticotyle Sahai & Sri-
vastava, 1978; Quasimmetra Sahai & Srivastava, 1978; Transversolecithus Sahai &
Srivastava, 1978;
Chrysophrytrema Gupta & Jain, 1992; Reference:: Bray et al. (1993a), Gibson
(2002)]
Diagnosis: Body surface smooth. Preoral lobe prominent. Testes oblique or tandem.
Seminal vesicle oval, tubular to trilocular, dorsal or postero-dorsal to ventral sucker.
Pars prostatica short, tubular, connected to seminal vesicle by short, aglandular duct.
Sinus-sac present, sinus organ present. Ovary oval. Distal extremity of uterus may
be vesicular. Excretory arms united in forebody.
Type species: E. tiegsi Woolcock, 1935.
Erilepturus hamati (Yamaguti, 1934) Manter, 1947 (Fig. 9.15)
[Syn. E. berdae Gupta & Sayal, 1979; E. caranxi Gupta & Gupta, 1988; E. chilkai
Gupta & Gupta, 1986; E. indicus Sahai & Srivastava, 1998; E. orientalis Sahai &
Srivastava, 1998; E. paralichthydis (Yamaguti, 1968); E. thrissocli Ahmad, 1980; E.
trichiurusi Gupta & Jain, 1992; E. yamaguti Ahmad, 1980]
The synonymies given above follow Bray et al. (1993a) who provided a detailed
review on the genus.
Hosts: Wide range of marine fish belonging to different taxa. Individual host species
are given below under each synonymized species (Table 9.1)
Location: Stomach
Distribution: Worldwide
References: Bray et al. (1993); Nahhas & Sey (2002).
Fig. 9.15 Erilepturus

hamati (after Bray, Cribb &
Barker, 1993)
Table 9.1 Species of Erilepturus (Syn. Uterovesiculurus) recorded from marine fish of India, their
hosts, localities and valid status
Species Synonym Host Locality References
E. bengalensis (Ahmad, U. bengalensis Pristipoma furcatum PR, BOB Ahmad
1980) Madhavi, 2011 Ahmad, 1980 (Bl. & Schn.) (1980a)
E. berdae (Gupta & E. hamati Acanthopagrus Laccadive Gupta,
Sayal, 1979) Madhavi, (Yamaguti, 1934) berda Forsskål Isles, N.K. &
2011 GOM Sayal
(1979)
E. caranxi (Gupta & E. hamati Caranx gallus Chilka Gupta,
Gupta, 1986) Madhavi, (Yamaguti, 1934) (Kluzinger) Lagoon P.C. &
2011 Gupta,
V.C.
(1986b)
E. chilkai (Gupta & E. hamati Etroplus suratensis Chilka Gupta,
Gupta, 1986) Madhavi, (Yamaguti, 1934) (Bloch) Lagoon P.C. &
2011 Gupta,
V.C.
(1986b)
E. gazzi (Ahmad, 1980) E. hamati Gazza minuta (Bl.) PR, BOB Ahmad
Madhavi, 2011 (Yamaguti, 1934) (1980a)
E. guptajaini Uterovesiculurus Eupleurogrammus PR, BOB Gupta, V.
Madhavi, 2011 indica Gupta & Jain, muticus Gray & Jain
1991 nec U. indica (1991a)
Sahai & Srivastava,
1978
E. hamati Yamaguti, Sillagolutea, Sillago VSK, Sujatha et
1934 soringa, BOB Madhavi
Sillaginopodys (1997)
chondropus
E. indicus (Sahai & E. hamati Rastrelliger South Sahai &
Srivastava, 1978) kanagurta (Cuvier) Indian Srivastava
Ocean (1978a)
E. neoorientalis Uterovesiculurus Abudefduf PR, BOB Ahmad
Madhavi, 1911 orientalis Ahmad, bengalensis (1980a)
1980, E. hamati (Gunther)
E. orientalis (Sahai & E. hamati Acanthopagrus South Sahai &
Srivastava, 1978) Bray, berda (Forsskål) Indian Srivastava
1993 Ocean (1978a)
E. paralichthydis E. hamati Eleutheronema BOM; AS Gupta,
(Yamaguti, 1934) tetradactylum N.K. &
Manter, 1947 (Shaw) Sehgal
(1970a)
E. puriensis (Gupta & Sphyraena obtusata PR, BOB Gupta, V.
Jain, 1991) (Cuv. & Val.) & Jain
(1991a)
(continued)
Table 9.1 (continued)

Species Synonym Host Locality References
E. sarbai (Gupta & Jain, Chrysophitrema Rhabdosargus sarba PR, BOB Gupta, V.
1992) Gibson, 2002 sarbai Gupta & Jain, (Forsska°l) & Jain
1992 (1992b)
E. thrissocli (Ahmad, E. hamati Thryssa mystax (Bl. PR, BOB Ahmad
1980) & Schn.) (1); Sillago (1980a),
Madhavi, 2011 lutea McKay; Sujatha &
Sillago soringa Dutt Madhavi
& (1997)
Sujatha;
Sillaginopodys
chondropus
(Bleeker) (2)
E. trichiurusi (Gupta & Eupleurogrammus PR, BOB Gupta, V.
Jain, 1991) Madhavi, muticus Gray & Jain
2011 (1991a)
E. yamagutii (Ahmad, E. hamati Trachinocephalus PR, BOB Ahmad
1980) Madhavi, 2011 myops (Forster) (1980a)
Description: Body elongate. Soma 1200–4375 × 380–1170, forebody 500–1850,

hindbody 450–150; forebody to hindbody ratio 1:0.8–1.3. Tegument smooth. Oral
sucker subterminal, 120–280 × 100–310, preoral lip present. Ventral sucker near
midbody, 250–725 in diameter. Sucker ratio 1:1.9–2.54. Intestinal caeca extend into
ecsoma. Testes intercaecal, diagonal in most, slightly posterior to ventral sucker.
Seminal vesicle thick- or thin-walled, Postero-dorsal to ventral sucker. Pars prostatica
long, narrow, surrounded by prostate cells at its distal portion. Hermaphroditic duct
enclosed in sinus-sac, opening into tubular genital atrium. Genital pore at level of
posterior margin of oral sucker. Ovary unlobed, midway between testis and posterior
end of soma. Uterine coils extensive, not intruding into ecsoma. Uterine seminal
receptacle present or absent. Vitelline tubes digitiform, in two lateral groups of 4 and
3, not reaching end of soma. Eggs 17–24 × 10–15 (Table 9.1).
Genus Qadriana Bilqees, 1971
[Syns Atheria Hafeezullah, 1975; Quasimmetra Sahai & Srivastava, 1978; Refer-
ence:: Gibson (2002)]
Diagnosis: Body stout. Body surface smooth. Preoral lobe large. Testes postero-
lateral to ventral sucker, symmetrical. Seminal vesicle saccular, in forebody. Pars pro-
statica short, tubular, connected to seminal vesicle by short duct. Sinus-sac present.
Sinus organ long, muscular. Ovary oval. Genital pore at level of pharynx. Uterus
forming muscular dilatation just before joining male duct.
Qadriana zakiae (Hafeezullah, 1975) Gibson, 2002 (Figs. 9.16 and 9.17)
[Syn. Atheria zakiae Hafeezullah, 1975]
Fig. 9.16 Qadriana zakiae

(after Hafeezullah, 1975)
Fig. 9.17 Qadriana zakiae

Enlarged anterior part
Hosts: Haemulidae: Pomadasys hasta (Bloch); Sciaenidae: Johnius carulta Bloch;

Sillaginidae: Sillago sihama (Forsskål); Terapontidae: Terapon jarbua (Forsskål)
Locality: VSK, MS, Gopalpur, BOB
Reference: Hafeezullah (1975a)
Description: Body 2832–5335 long including ecsoma, 1072–2008 wide. Body sur-
face thick, smooth. Oral sucker 192–453 in diameter. Ventral sucker 550–1100 in
diameter, at about middle of body. Testes globular, symmetrical, postero-lateral to
ventral sucker. Seminal vesicle sac-like, situated between ventral sucker and intesti-
nal bifurcation. Pars prostatica reduced. Prostatic cells around pars prostatica and
distal part of sinus-sac. Sinus-sac barrel-shaped. Genital atrium short. Genital pore
ventral to pharynx. Ovary globular, pretesticular. Vitellarium with seven long wind-
ing tubules, three right and four left sometimes descending into ecsoma. Uterus may
or may not extend into ecsoma, forms a muscular bulb just before entering sinus-sac.
Eggs 17–18 by 10–12.
Remarks: Hafeezullah (1975) erected the genus Atheria to include the single species
A. zakiae Hafeezullah, 1975. Gibson (2002) considered the genus Atheria as a
synonym of Qadriana Bilqees, 1971 finding no major morphological differences
between the two. A. zakiae became Q. zakiae. Genus Qadriana Bilqees, 1971, closely
resembles Erilepturus Woolcock, 1935, but differs from it in the seminal vesicle sit-
uated in forebody and in the greatly reduced pars prostatica.
Genus Stomachicola Yamaguti, 1934

[Indostomachicola Gupta & Sharma, 1973; Linguastomachicola Srivastava & Sahai,
1978; Pseudostomachicola Skrjabin & Guschanskaja, 1954; Acerointestinecola
Jahan, 1970; Cameronia Bilqees, 1971; Segmentatum Bilqees, 1971; Cestodera
Bilqees, 1971.
Reference: Hafeezullah (1998) Gibson (2002)]
Diagnosis: Body long, ecsoma enormous. Body surface smooth. Seminal vesicle
oval, in hindbody. Pars prostatica tubular, long, sinuous, external gland cells not
evenly distributed throughout length. Sinus-sac present. Ovary oval to reniform.
Majority of uterine coils within ecsoma, usually fill half of ecsoma.
Yamaguti (1934) erected the genus Stomachicola with S. muraenesocis from the
stomach of Muranesox cinereus in the Inland Sea of Japan as the type species. The
genus is characterized by the extremely long ecsoma, the long filiform vitelline
lobes and a sinuous pars prostatica. Two genera Indostomachicola Gupta & Sharma,
1973, and Linguastomachicola Srivastava& Sahai, 1978 have been synonymised
with Stomachicola. Several species belonging to this genus have been recorded from
India and Pakistan from marine fishes mostly eels. However, most of these were
later considered junior synonyms of the type species (Hafeezullah, 1980; Gupta
& Gupta, 1991). Bilqees (1971) established three genera with eight species from
marine fish in Pakistan: Cameronia (C. octovitellaria, C. pakistani), Segmentatum
(S. karachiensis, S. gadrii, S. cinderensis and S. magnaesophagustum) and Cestodera
(C. gastrocecus, C. unicecus). Gibson (2002) considered these three genera together
with Linguastomachicola Srivastava & Sahai, 1978 as junior synonyms of Stomachi-
ola. According to Gibson (2002) Indostomachicola Gupta & Sharma 1973 is also a
synonym of this genus. Hafeezullah (1980) expressed the view that the eight species
described by Bilqees (1971) conformed to the original description of S. muraene-
socis and all of them should be considered as junior synonyms of S. muraenesocis.
Altogether eleven species of Stomachicola have been described from marine fish of
Indian region. Of these, S. singhi Gupta & Ahmad, 1974; S. chauhani Pandey &
Tewari, 1984, S. chauhani Gupta & Singh, 1982, S. kinnei (Gupta & Sharma, 1973)
were synonymized with S. muraenesocis (Reference: Gupta & Gupta, 1991). Three
species S. lepturusi Gupta & Gupta, 1976, S. rauschi Gupta & Ahmad, 1978 and
S. secundus Srivastava, 1937 were transferred to the genus Allostomachicola Yam-
aguti 1958 which differs from Stomachicola in the location of the seminal vesicle
in the forebody and in the small vesicle-like pars prostatica. This leaves only five
valid species in the genus: S. muraenesocis Yamaguti, 1934, S. guptai Ahmad, 1978;
S. mastacembeli Verma, 1973; S. pelamysi Gupta & Gupta, 1976 and S. serpentine
(Srivastava & Sahai, 1978). Recently, a new species S. bengalensis Mishra, Chandra,
Johri, Bajpai & Saxena, 2014 (published in a conference proceedings) is reported
under the genus. The description of the species lacks details and it may not belong to
the genus Stomachicola. Here it is considered as species inquirendum. In the present
account detailed, description is provided for Stomachicola muraenesocis and only
information on the hosts and distribution is provided for the remaining species which
in due course of time may fall as synonyms of S. muraenesocis. A table is created giv-
ing details of the hosts, localities of the species synonymized with S. muraenesocis
and also the species tentatively considered as valid.
Stomachicola muraenesocis Yamaguti, 1934 (Fig. 9.18)
[Syn. Pseudostomachicola rubea (Linton, 1910), P. magna (Manter, 1931),
Lecithocladium longicaudatum Tseng, 1935, S. longicaudatum (Tseng, 1935)
Chauhan, 1945, Acerointestinecola karachiensis Jahan, 1970, Cameronia octovitel-
larii Bilqees, 1971, C. pakistani Bilqees, 1971, Segmentatum karachiensis Bilqees,
1971, S. qadrii Bilqees, 1971, S. cinerensis Bilqees, 1971, S. magnaesophagus-
tum Bilqees, 1971, Cestodera gastrocecus Bilqees, 1971, C. unicecus Bilqees,
1971, Indostomachicola kinnei Gupta & Sharma, 1973, Stomachicola mastacembeli
Verma, 1973, Stomachicola pelamysi Gupta & Gupta, 1976, Stomachicola chauhani
Pandey & Tiwari, 1984, n. syn. Linguastomachicola serpentina Srivastava & Sahai,
1978]
Fig. 9.18 Stomachicola

muraenesocis (after
Yamaguti, 1934)
Table 9.2 Nominal Indian species of Stomachicola, their hosts and localities
Name of the Status Host Locality References
species
S. Valid Muraenesox cinereus BOB, AS 1. Bhalerao
muraenesocis (Forsskål) (1), M. (1945); 2.
Yamaguti, talabanoides (Bleeker) (2), Chauhan
1934 Sarda chiliensis (Cuvier) (3) (1945b);
Monopterus cuchia Gupta, N.K.
(Hamilton); Ophisternon & Sehgal
bengalense (McClelland); (1970a);
Ariosoma mauritianum Gupta, Gupta,
Pappenheim) (4, 5) A.N. &
Sharma
(1973)
3. Gupta, R.C.
& Gupta, S.P.
(1974); 4.
Gupta, P.C. &
Gupta, V.C.
(1991); 5.
Gupta, P.C. &
Singh (1981)
S. chauhani Syn of S. Congridae: Uroconger BOB, AS Pandey &
Pandey & muraenesocis lepturus Richardson Tewari (1984)
Tewari, 1984 Gupta, P.C. &
Gupta, V.C.
(1991)
S. chauhani Syn of S. Congridae: Uroconger PR, BOB Gupta &
Gupta & muraenesocis lepturus Richardson Singh (1981)
Singh, 1981 Gupta, P.C. &
Gupta, V.C.
(1991)
S. pelamysi Syn of S. Sarda chiliensis (Cuvier) PR, BOB Gupta &
Gupta & muraenesocis Singh (1981)
Gupta, 1974
S. singhi Syn of S. Acanthurus triostegus (L.) PR, BOB Gupta, V. &
Gupta & muraenesocis Ahmad
Ahmad, 1978 (1978b)
S. kinnei Syn of S. Leptocephalus conger (Linn.) Ratnagiri, AS Gupta &
(Gupta & muraenesocis Sharma,
Sharma, Syn. Indos- (1973)
1973) tomachicola
kinnei Gupta
& Sharma,
1973
(continued)

Name of the Status Host Locality References
species
S. secundus Included Hemiramphus limbatus (Day) BOB Srivastava
Srivatava, 19 under (1); Chirocentrus dorab (1939d);
Allostom- (Forsskål) (2, 3) Gupta, P.C. &
achicola Singh (1981);
Gopal et al.
(1984)
S. rauschi Included Chirocentridae: Chirocentrus PR, BOB Gupta, V. &
Gupta & under dorab (Forsskål) Ahmad
Ahmad, Allostom- (1978b);
1978 achicola Gupta, P.C. &
Singh (1981)
S. singhi Syn. of S. Acanthuridae: Acanthurus PR, BOB Gupta, V. &
Gupta & muraenesocis triostegus (L.) Ahmad
Ahmad, 1978 Reference: (1978b)
Hafeezullah
(1985)
S. guptai valid
Ahmad, 1978
S. Syn. of S. Mastacembelus armatus
mastacembeli muraenesocis
Verma, 1973 Reference:
Hafeezullah
(1985)
S. serpentinea Syn. of S. Muraenesocidae: Muraenesox AS Srivastava &
(Srivastava & muraenesocis talabanoides (Forsskål) Sahai (1978)
Sahai, 1978). Reference:
Hafeezullah
(1985)
Hosts and localities: See table 9.2

Distribution: Indo-Pacific
Reference: Hafeezullah (1985)]
Description: Soma very long, 5.8–6.6 mm or longer, broadest at level of genital

organs. Ecsoma very long, several times length of soma, 14.7–17.28 mm long. Tegu-
ment thick, smooth. Oral sucker subterminal, ventrally flexed, 156–183 by 183–214
in size. Ventral sucker 640–840 × 672–800. Pharynx postero-dorsal to oral sucker.
Oesophagus short. Caeca sinuous, often filled with black pigment, extend to poste-
rior end of ecsoma. Testes oval, symmetrical, posterior to ventral sucker. Seminal
vesicle oval, lies immediately posterior to ventral sucker. Pars prostatica long, sin-
uous, unites with uterus forming hermaphroditic duct enclosed in sinus-sac which
opens into wide genital atrium. Genital pore just posterior to oral sucker. Ovary reni-
form, post-testicular. Uterus extends into tail, for about one-fifth of its length. Eggs
oval, 17–22 × 13–14. Vitellarium consists of seven long, slender tubes, extending
into ecsoma.
Remarks: Characterized by the large body size, the well-developed pars prostatica,
the voluminous seminal receptacle and long, filiform vitelline tubes. S. muraenesocis
has been reported from Muraenesox cinereus from several localities: Japan (Yam-
aguti, 1934), China (Tseng, 1935), Indian Ocean (Bhalerao, 1944, Bilqees, 1971);
and Formosa (Reid et al. 1966). Chauhan (1945b) and Gupta & Sehgal (1970) also
reported this species from Muraenesox talabanoides from India. Chauhan (1945b)
transferred Lecithocladium longicaudatum Tseng, 1935 to Stomachicola as S. long-
icaudatum (Tseng, 1935). Yamaguti (1954) has synonymized this species with S.
muraenesocis (Table 9.2).
Genus Tubulovesicula Yamaguti, 1934

[Syn. Transversolecithus Sahai & Srivastava, 1978; Reference: Gibson (2002)]
Diagnosis: Soma spindle-shaped, with ecsoma. Oral sucker subterminal. Caeca ter-
minate near posterior end. Testes in hindbody, obliquely juxtaposed, separated by
uterus. Seminal vesicle tubular, sinuous, anterior to testes. Pars prostatica long, sur-
rounded by numerous prostatic cells. Hermaphroditic duct enclosed by sinus-sac,
expanded at base. Genital pore at level of pharynx. Ovary close to posterior testis.
Uterus descends into ecsoma. Vitellarium with seven digitiform lobes. Excretory
vesicle Y-shaped, arms uniting dorsal to pharynx. Type species: T. spari Yamaguti
1934.
Two species of the genus have been recorded from India; T. angusticaudatum
(Nicoll, 915) and T. lindbergi (Layman, 1930) Yamaguti, 1934.
Tubulovesicula angusticauda (Nicoll, 1915) Yamaguti, 1934
[Syn. Transversolecithus scateri Sahai & Srivastava, 1978; Tubulovesicula scateri
(Sahai & Srivastava, 1978; T. anguillae Yamaguti, 1934; T. australica Lebedev, 1968;
T. longicaudata Lebedev, 1968; T. magna Lebedev & Nighat, 1981; T. muraensocis
Yamaguti, 1934; T. pseudorhombi Yamaguti, 1934, T. sauridia Gu & Shen, 1978; T.
serrani Nagaty, 1956 (Reference: Bray, 1990)
Host: Synodontidae: Saurida tumbil (Bloch) (1) Platycephalidae: Platycephalus
scaber (Linn.)
Reference: 1. Radhakrishnan & Nair (1986); Sahai & Srivastava, 1978; 2. Hafeezul-
lah (1990)
Remarks: T. angusticauda has been recorded from a wide range of marine fishes
from geographic localities such as Australia, New Zealand, Celebes, Hawaii, Red
Sea. From India, there are two reports: Radhakrishnan & Nair (1986) reported it
from Saurida tumbil and later Hafeezullah (1990) recorded it from Platycephalus
scaber and synonymized Transversolecitha scateri Sahai & Srivastava, 1978 with
this species. Bray (1990) while recording the occurrence of this species in Trachichth-
odes gerradi from South Australia synonymized eight species of the genus with T.
angusticauata.
Tubulovesicula lindbergi (Layman, 1930) Yamaguti, 1934 (Fig. 9.19)
[Syns. Lecithaster lindbergi Layman, 1930, Tubulovesicula spari Yamaguti, 1934, T.

muraenosocis Yamaguti, 1934; T. californica Park, 1938; T. pseudorhombi Yamaguti,
1934; T. madurensis Nigrelli, 1940; T. nanaimoensis (McFarlane, 1936) Manter,
1947]
Host: Synodontidae: Saurida tumbil (Bloch)
Locality: Mandapam, GOM
No. 1
Reference: Gupta, N.K. & Sehgal (1970a)
Description: Body fusiform, 8.34 mm long, 1290 wide. Tegument smooth. Oral
sucker 238 × 231, suberminal. Ventral sucker in anterior part of body, 612 × 605.
Caeca smooth, terminate near posterior end of body. Testes spherical, diagonally
symmetrical. Seminal vesicle tubular, extends from posterior margin of ventral sucker
to middle level of anterior testis. Pars prostatica elongated, surrounded by prostatic
cells. Hermaphroditic vesicle within sinus-sac, ovoidal, situated posterior to intestinal
bifurcation. Ovary spherical, post-testicular, larger than testis. Uterine coils extend to
middle of ecsoma, metraterm well developed. Eggs numerous 27 × 13. Vitellarium
7 slender clavate tubes, extending anterior as well as posterior to ovary.
Remarks: This species has been recorded from the Pacific coast of Japan (Yamaguti,
1934) and Gulf of Mannar (Gupta & Sehgal, 1970a). It differs from T. angusticauda
in the posterior extent of the seminal vesicle and in the larger egg size.
Fig. 9.19 Tubulovesicula

lindbergi (after Yamaguti,
1934)
Subfamily Elytrophallinae Skrjabin & Guschanskaja, 1954

Diagnosis: Ecsoma well developed. Body surface smooth or plicated. Presomatic
pit absent. Testes tandem to symmetrical, usually oblique. Seminal vesicle with
thick muscular wall, oval, not constricted into portions, in forebody or in hindbody.
Pars prostatica long or short. Sinus-sac present, tubular, not enclosing ejaculatory
or prostatic vesicle. Sinus organ well developed. Genital atrium deep. Ovary oval.
Vitellarium seven tubular to tear-shaped lobes, three on one side and four on the other
which may form rosette. Excretory arms united in forebody.
Type genus: Elytrophallus Manter, 1940
Key to genera
1. Sinus-sac long and narrow, reaching to level of seminal vesicle, vitelline lobes
tear-shaped………………………………….. Elytrophalloides Szidat, 1955
Sinus-sac relatively long, but not reaching to level of seminalvesicle, vitelline
lobes tubular………………………………….. Lecithocladium Luhe, 1901
Genus Elytrophalloides Szidat, 1955
Diagnosis: Body surface with plications. Seminal vesicle large, reaching back to
level of testes. Pars prostatica sinuous, in hindbody. Sinus-sac long, narrow reaching
back to level of seminal vesicle. Vitelline lobes tear-shaped.
Type species: E. oatesi Szidat, 1955
Elytrophalloides parukhini Gupta & Gupta, 1997
Host: Lutjanidae: Lutjanus monostigma (Cuvier)
Locality: PR, BOB
No. 1
Description: Body elongate, 2776–2887 long (excluding ecsoma) by 617–648 wide.
Ecsoma 1286–2887 long by 348–410 wide. Body surface with plications. Oral sucker
terminal, 182–211 by 248–256. Intestinal caeca simple, extend to posterior tip of
ecsoma. Ventral sucker spherical, 331–338 in diameter. Sucker ratio 1:1.3–1.4. Testes
ovoid, obliquely tandem, entire, pre-equatorial. Seminal vesicle elongated extends
obliquely posteriorly to mid level of anterior testis. Pars prostatica tubular, surrounded
by numerous prostrate gland cells, runs posteriorly and then anteriorly to join base
of sinus-sac. Sinus-sac muscular. Hermaphroditic duct convoluted within sinus-sac.
Genital atrium tubular. Genital pore median or submedian, ventral to median level of
pharynx or more posteriorly. Ovary transversely elongated. Vitellarium consisting
of 7 long, winding tubular lobes, sometimes entering into ecsoma. Uterine coils
descending into ecsoma. Eggs oval, operculate, 15–16 long by 5–7 wide.
Remarks: E. parukhini differs from the type species E. oatesi Szidat, 1955 in having
the genital pore at the mid level of the pharynx instead of at the mid level of oral
sucker, the intestinal caeca extending to thde posterior extremity instead of confined
to the soma, the uterus extending into the ecsoma instead of confined to the soma
and larger body size.
Genus Lecithocladium Luhe, 1901

[Syns Clupenurus Srivastava, 1935; Colletostomum Sahai & Srivastava, 1978; Cleft-
icolletta Sahai & Srivastava, 1978; Bengalotrema Malhotra, Nanda, Mukherjee,
Ghosh, Sukul & Capoor, 1989; Reference: Gibson (2002)]
The genus Lecithocladium is characterized by the possession of a funnel-shaped
oral sucker, a long and cylindrical pharynx, and an unpartitioned seminal vesicle,
long tubular vitelline tubes usually seven in number.
Diagnosis: Body surface with plications. Oral sucker usually infundibuliform and
larger than ventral sucker. Pharynx cylindrical. Seminal vesicle large, thick-walled,
fusiform, in hindbody. Pars prostatica long, sinuous, mainly or entirely in hindbody.
Sinus-sac tubular, narrow, not reaching level of seminal vesicle, mainly in forebody.
Vitelline lobes long and tubular.
Type species: L. excisum (Rudolphi, 1819)
Lecithocladium Luhe, 1901 is one of the largest genera of digeneans comprising over
83 nominal species (Bray & Cribb, 2004), now nearly 100 (Gibson, 2013). From the
Indian region alone as many as 38 species have been recorded.
The descriptions provided for many of these species are inadequate and the valid-
ity of many of the species remained questionable. Lack of host specificity amongst
the species of the genus, extensive intraspecific variations in respect of certain char-
acters and fixation artifacts have magnified the problem of species differentiation in
the genus. Gibson & Bray (1986) cleared the confusion by reducing the number of
species of the genus in the Indo-Malaysian region from 40 nominal species to six,
pointing out that previously ‘little account has been taken of intraspecific variations,
fixation artifacts or functional variation of the genus. Possibility exists that many
more species of the genus also fall synonyms. According to Bray & Cribb (2004) the
taxonomy of the genus is difficult and the existing taxonomy needs to be tested with
molecular techniques. Table 9.3 lists the species recorded from India, their validity
status and also provides details of hosts and localities for each species.
Species descriptions: It is evident from the table that a total of 45 species were
recorded from India of which 30 species were synonymized leaving only 15 valid
species. Descriptions are given below for some of the valid species.
Lecithocladium annulatum Chauhan, 1945
Body 9.87 mm long and 1.10 mm broad across posterior region of the soma.
Oral sucker cup-shaped, 425 × 325 in size. Intestinal caeca extend into ecsoma and
end in front of its posterior end. Ecsoma 3.61 mm in length. Ventral sucker 350 in
diameter, situated in anterior sixth of body. Testes obliquely placed. Seminal vesicle
pear-shaped, with muscular wall, touches anterior border of posterior testis. Pars-
prostatica quite long, tubular, extends up just anterior to ventral sucker, surrounded
by prostate gland cells. Genital pore at antero-ventral border of oral sucker. Ovary
kidney-shaped, situated just at about middle region of body. Vitellarium consists
of seven long, filiform tubules in two groups of four and three. Uterine coils not
extending intoecsoma. Metraterm well developed. Eggs elliptical, 12 × 4.8.
Table 9.3 Species of Lecithocladium recorded from Indian marine fish

Species Validity status Host Locality References
L. Valid Stromateidae: Pampus BOM (1), 1. Chauhan
annulatum argenteus (Euphrasen) Calicut (2), (1945); 2.
Chauhan, (1); Carangidae: AS Gupta, N.K.
1945 Apolectus niger (Bloch) & Sehgal
(2) (1970a)
L. Syn. L. indicum Gupta Scombridae: Rastrelliger VSK, BOB Madhavi &
angustiovum & Gupta, 1978; L. kanagurta (Cuvier) Trivenilak-
Yamaguti, manteri Gupta & Puri, shmi
1953 1981; L. seriolellae (2011)
Gupta & Gupta, 1983;
L. microlepidotusi
Gupta & Govind, 1981;
L. siddiquii Gupta &
Gupta, 1983; L. tewarii
Gupta & Gupta, 1983;
L. apolecti Syn. L. excisiforme of Carangidae: Formio Veeraval, AS Hafeezullah
Velasquez, Gupta & Sehgal, 1971; niger (Bloch) (1975)
1962 L. excisiforme of
Radhakrishnan & Nair,
1979
L. Valid Carangidae: Decapterus PR, BOB Reference:
bengalense russelli (Ruppel) Ahmad
Ahmad, (1981a)
1981
L. Valid Leiognathidae: Gazza Saxena &
bengalense minuta (Bl.) Prakash
Saxena & (2006)
Prakash,
2006
L. Syn. of L. harpodontis Carangidae: BOB Srivastava
brevicaudum Srivastava, 1942 Chrysophrys bifasciata (1942a)
Srivastava, (Forsska°l)
1942
L. caranxi Valid Carangidae: PR, BOB Gupta, P.C.
Gupta & Carangoides praestus & Govind
Govind, Bennett 1981)
1981
L. carultum Syn. of L. glandulum Sciaenidae: Johnius BOM, AS Chauhan
Chauhan, Chauhan, 1945 carultum (Day); (1945)
1945 Synodontidae Harpodon Gibson &
neherius (Hamilton) Bray (1986)
L. chauhani Syn. of L. harpodontis Clupeidae: Hilsa ilisha Hafeezullah
Hafeezullah, Srivastava, 1942 Hamilton (1975a)
1975
L. cybiumi Syn. of Scombridae: PR, BOB Gupta &
Gupta & L.angustiovumYamaguti, Scomberomorus guttatus Jain (1992b)
Jain, 1992 1953 (Bl. & Schn.)
(continued)

L. elongatus Syn of L. parviovum Carangidae: Alepis PR, BOB Gupta, V. &
Gupta & Yamaguti, 1953 djedaba (Forsska°l) Puri (1981)
Puri, 1982 Gibson &
Bray (1986)
L. Valid Stromateidae: BOM, AS Gupta N.K.
excisiforme Stromateus niger, Trigon & Sehgal
Cohn, 1903 sp. (1970a)
L. excisum Valid Carangidae: Apolectus Hugli Zhukov
(Rudolphi) niger (Bloch); Estuary (1977)
Luhe Stromateidae: Pampus Hafeezullah
argenteus (Euphrasen) & Dutta
(1998)
L. fotedari Syn. of L. parviovum Carangidae: PR, BOB Gupta, P.C.
Gupta & Yamaguti, 1953 Carangoides praeustus & Singh
Singh, 1982 (Bennett) (1982)
Gibson &
Bray (1986)
L. Valid Leiognathidae: Gazza PR, BOB Gupta &
gazzi Gupta minuta (Bleeker) Jain (1992b)
& Jain, 1992
L. Valid Lutjanidae: Lutianus BOB, AS Reference:
glandulum johnii (Bloch); 1. Chauhan
Chauhan, Mugilidae: Valamugil (1945); 2.
1945 speigleri (Bleeker) (1); Gupta, N.K.
Synanceiidae Minous & Sehgal
monodactylus (Bl. & (1970a);
Schn.); Sciaenidae: 3.
Kathala Hafeezullah
axillaris (Cuvier); (1975a)
Stromateidae: Pampus 4. Kumari &
argenteus Euphrasen; Srivastava
Carangidae: Caranx (1976)
sansum (Forsskål) (2);
C. carangus (Bloch) (3)
Polynemidae: Polynemus
paradiseus (L.) (4)
L. guptai Synodontidae: Saurida PR, BOB Gupta, P.C.
Gupta & tumbil (Bloch) & Govind
Govind, (1981)
1981
L. hanuman- Syn. of L. megalaspis Carangidae: Alepis PR, BOB Gupta, V. &
thai Gupta & Yamaguti, 1953 djedaba (Forsskål) Jain (1991b)
Jain, 1991 Bry & Cribb
(2004)
(continued)

L. Valid Synodontidae: BOM, AS; Srivastava
harpodontis Harpodon neherius DC, BOB (1942a),
Srivastava, (Hamilton); Kumari &
1942 Stromateidae: Pampus Srivastava
argenteus (Euphrasen) (1976)
Gibson &
Bray (1986)
L. indicum Syn. of L. angustiovum Scombridae: Rastrelliger Kerala, AS Gupta, R.C.
Gupta & Yamaguti, 1953 kanagurta (Cuvier) & Gupta,
Gupta, 1978 S.P. (1978)
Bray &
Cribb (2004)
L. inglisi Syn. of L. megalaspis Carangidae: Alepes PR, BOB Gupta, V. &
Gupta & djedaba (Forsskål) Ahmad
Ahmad, (1977a)
1977 Bray &
Cribb (2004)
L. Valid Rachycentridae: PR, BOB Ahmad
jagannathi Rachycentron canadum (1981a)
Ahmad, (L.)
1981
L. keralense Syn. of L. parviovum Psettodidae: Psettodes Kerala, AS Gupta, R.C.
Gupta Yamaguti, erumei (Bl. & Schn.) & Gupta,
&Gupta, 1953 S.P. (1978)
1978 Gibson &
Bray (1986)
L. magnum Syn. Cleftocolleta Carangidae: Madras, Sahai &
(Sahai & magnum Sahai & Scomberoides tala BOB Srivastava,
Srivastava, Srivastava, 1978 (Cuvier) (1978)
1978)
L. manteri Syn. of L. angustiovum Scombridae: Rastrelliger PR, BOB Gupta, V. &
Gupta & Yamaguti, 1953 kanagurta (Cuvier) Puri (1981);
Puri, 1981 Gibson &
Bray (1986);
Bray &
Cribb (2004)
L. Syn. L. hanumanthai Carangidae: Megalaspis BOM, AS Hafeezullah
megalaspis Gupta & Jain 1992; L. cordyla (L.) (1975a)
Yamaguti, inglisi Gupta & Ahmad,
1958 1977; L. purense Gupta
& Gupta, 1978’ L.
singhi Gupta & Singh,
1982
L. microlepi- Syn. of L. angustiovum Scombridae: Rastrelliger PR, BOB Gupta &
dotusi Gupta Yamaguti, kanagurta (Cuvier) Govind,
& Govind, 1953 1981, Bray
1981 (1990c)
(continued)

L. muthiahi Syn. Sciaenidae: BOM, AS Sahai &
(Sahai Colletostomummuthiahi Pseudosciaena diacanthi Srivastava
&Srivastava, Sahai & Srivastava, (Lacepede) (1942)
1978 1978 Gibson &
Syn. of L. harpodontis Bray (1986)
(Srivastava, 1942)
L. piscicola Syn. Clupenurus Clupeidae: Hilsa ilisha Srivastava
(Srivastava, piscicola Srivastava, (Hamilton) (1935)
1935) 1935 Hafeezullah
L. chauhani (1975a)
Hafeezullah, 1975
L. psettodi Syn. of L. glandulum Psettodidae: Psettodes PR, BOB Gupta, V. &
Gupta & Chauhan, 1945 erumei (Bl. & Schn.) Puri (1981)
Puri, 1981 Gibson &
Bray (1986)
L. purense Syn. of L. megalaspis Leiognathidae: Gazza PR, BOB Gupta, R.C.
Gupta & minuta (Bleeker) & Gupta,
Gupta, 1978 S.P. (1978),
Gibson &
Bray (1986);
Bray &
Cribb (2004)
L. puriense Valid Stromatidae: Pampus PR, BOB Ahmad
Ahmad, chinensis (Euphr.) (1981a)
1981
L. saurida Valid Synodontidae: Saurida PR, BOB Gupta, P.C.
Gupta & tumbil (Bloch) & Govind
Govind, (1981)
1981
L. sciani Syn. of L. parviovum Sciaenidae: Kathala PR, BOB Gupta, V. &
Gupta & Yamaguti, 1953 axillaris (Cuvier) Puri (1981)
Puri, 1981 Gibson &
Bray (1986)
L. seriolellae Syn. of L. angustiovum Scombridae: Rastrelliger PR, BOB Gupta, P.C.
Gupta & Yamaguti, 1953 kanagurta (Cuvier) & Gupta,
Gupta, 1983 V.C. (1983)
Gibson &
Bray (1986);
Bray &
Cribb (2004)
L. Pobably belongs to the Carangidae: Caranx Ernakulam, Gupta, N.K.
sexfasciatum genus Ectenurus Looss, sexfasciatus (Quoy & AS & Mehrotra
Gupta & 1907 Gaimard) (1970a);
Mehrotra Hafeezullah
1970 (1975a)
L. siddiquii Syn. of L. angustiovum Scombridae: Rastrelliger PR, BOB Gupta &
Gupta & Yamaguti, 1953 kanagurta (Cuvier) Gupta,
Gupta, 1983 (1983); Bray
(1990c)
(continued)

L. simhai Valid Synodontidae: Saurida PR, BOB Pandey &
Pandey & tumbil (Bloch) Govind
Govind, (2007)
2007
L. singhi Syn. of L. megalaspis Carangidae: Caranx ire PR, BOB Gupta &
Gupta Yamaguti, 1953 (Cuv. & Val.) Singh,
&Singh, (1982)
1982 Gibson &
Bray (1986)
Bray &
Cribb (2004)
L. Syn. of L. parviovum Leiognathidae: Gazza PR, BOB Gupta, V. &
srivastavai Yamaguti, 1953 minuta (Bleeker) Puri (1981)
Gupta & Gibson &
Puri, 1981 Bray (1986);
Bray et
Cribb (2004)
L. striatum Syn. Bengalotrema Clupeidae: Hilsa ilisha West Bengal, Malhotra
(Malhotra & striatum Malhotra & Hamilton River Ganga et al. (1989)
Sunita Sunita Nanda, 1989
Nanda,
1989)
L. Syn. of L. parviovum Stromateidae: Pampus PR, BOB Gupta, V. &
stromateusi Yamaguti, 1953 argenteus Euphrasen Puri (1981)
Gupta &
Puri, 1982
L. sulphurusi Species inquirendae Mullidae: Upeneus PR, BOB Gupta &
Gupta & sulphureus (Bleeker) Govind,
Govind, (1985)
1985
L. tewarii Syn. of L. angustiovum Rastrelliger kanagurta PR, BOB Gupta &
Gupta & Yamaguti, 1953 (Cuvier) Gupta,
Gupta, 1987 (1987)
L. thapari Syn. of L. glandulum Sphyraenidae: PR, BOB Gupta &
Gupta & Chauhan, 1945 Sphyraena obstusata Gupta,
Gupta, 1978 (Giinther) (1978)
L. triacanthi Valid Triacanthidae: Quilon, Gupta,
Gupta & Triacanthus strigilifer Keral, AS R.C.&
Gupta, 1978 (Day) Gupta, S.P.
(1978)
Remarks: Chauhan (1945) reported the species from Pampus argenteus off Bombay
coast, Arabian Sea. Later Gupta & Sehgal (1970 reported the species from Parastro-
mateus niger (=Apolectus niger) from the coast of Calicut.
Lecithocladium angustiovum Yamaguti, 1953 (Fig. 9.20)
[Syn. L. indicum Gupta & Gupta, 1978; L. cybiumi Gupta & Jain, 1992; L. manteri
Gupta & Puri, 1981; L. seriolellae Gupta & Gupta, 1983; L. siddiquii Gupta & Gupta,
1983; L. tewarii Gupta & Gupta, 1987; L. microlepidotusi Gupta & Govind, 1981]
Description: Body 3800–6528 long (excluding ecsoma) 750–1080 wide; ecsoma
2830–5740 long. Body surface plications extend from posterior end of oral sucker
to posterior end of soma. Oral sucker funnel-shaped. Caeca terminate at posterior
end of ecsoma. Ventral sucker smaller than oral sucker, in anterior third of body.
Testes ovoid, diagonal, subequal, contiguous. Seminal vesicle spindle-shaped, with
thick muscular wall, extends to anterior border of anterior testis or just posterior
to it. Pars prostatica curved, surrounded by large number of prostate gland cells.
Hermaphroditic duct elongate, thin-walled, genital pore near middle of oral sucker,
ovary ovoid, near posterior end of soma, separated by uterus from posterior testis or
contiguous to it. Vitellarium consists of seven long, winding narrow tubules, confined
to soma. Uterus descends up to 1/2 of ecsoma. Eggs 15–19 long, 5–8 wide.
Fig. 9.20 Lecithocladium

angustiovum (original)
Remarks: This species is characterized by the funnel-shaped oral sucker, cylindrical

pharynx and ventral sucker smaller than oral sucker. The many species of Lecitho-
cladium recorded from marine fishes of India in particular the carangid and scombrid
fishes, have been synonymized with L. angustiovum (Bray, 1990; Gibson & Bray,
1986; Bray & Cribb, 2004). It has been reported from a wide range of geographical
localities in the Indo-west Pacific Ocean (Bray, 1990) and the preferred hosts are
carangids and scombrids.
Lecithocladium apolecti Velasquez, 1962 (Fig. 9.21)
Description: Body elongate, 10.09–15.37 mm long, 730–1040 wide, with distinct

body annular plications. Ecsoma 3820–6910 long 730–1040 wide. Ventral sucker
in first quarter of body 364–418 long, 378–459 wide. Oral sucker infundibuliform,
397–451 wide. Sucker ratio 1:0.91–0.99. Pharynx cylindrical. Caeca extend to poste-
rior end of body. Testes tandem, in middle of soma. Seminal vesicle large, fusiform
or elliptical, thick-walled, pretesticular, giving rise anteriorly to short and curved
glandular duct. Pars prostatica long, sinuous, surrounded by well-developed pro-
static gland cells to middle of ventral sucker, joining metraterm dorsal to ante-
rior part of ventral sucker to form hermaphroditic duct. Sinus-sac tubular, encloses
hermaphroditic duct. Genital pore ventral to anterior part of oral sucker. Ovary in
posterior half of soma. Juel’s organ present posterior to ovary. Vitellarium consists
of seven long tubular lobes, posterior to ovary. Uterine coils descend to middle of
ecsoma. Eggs 18–20 × 11–12.
Remarks: This species was initially reported by Velaquez (1962) from the Philip-
pines. Hafeezullah (1990) reported the same species from the black pomfret Formio
niger (Bloch) from Veeraval and Okha coasts and regarded L. excisiforme of Gupta

apolecti (original)
& Sehgal (1971) and L. excisiforme of Radhakrishnan et al. (1979) to be identical

to this species. All the reports have been from the black pomfret Apolectus (Formio)
niger. The chief characteristics of the species are: tegument feebly plicate, gonads
are situated in the middle third of body, few vitelline tubules extend anterior to ovary,
a few of them entering ecosma, pars prostatica is not very sinuous and is surrounded
by feebly developed prostatic cells.
Lecithocladium bengalense Ahmad, 1981
Body 2730–3200 long by 580–660 wide; ecsoma 1510–1680 long by 460–570

wide. Tegumental plications throughout soma. Oral sucker inverted eye–drop’
shaped, subterminal, 341–400 long by 300–340 wide; Ventral sucker spherical,
294–320 in diameter. Caeca simple, terminate at posterior end of body. Testes
two, oval tandem, contiguous, pre-equatorial. Seminal vesicle spindle-shaped, thin-
walled, undivided, pre-testicular. Pars prostatica tubular, winding, surrounded by
numerous prostate gland cells. Sinus-sac tubular, thin–walled, starting from genital
pore extends up to anterior border of ventral sucker. Genital pore ventral to anterior
part of oral sucker. Ovary smaller than testes. Vitellarium in form of seven long,
winding tubules. Uterus descends posteriorly into ecsoma to half of its length. Eggs
20–23 long by 8–11 wide.
Remarks: This species closely resembles L. harpodontis Srivastava, 1942 but differs
from it in having ovary smaller than testes, oesophagus saccular, uterus entering to
half the length of ecsoma, anterior testis larger than posterior testis and in the smaller
sinus-sac extending up to anterior margin of ventral sucker.
Lecithocladium glandulum Chauhan, 1945 (Fig. 9.22)
[Syn. L. psettodi Gupta & Puri, 1981]
Description: Body elongated, 2800–4590 long, 420–880 wide, beset with strong
annulations in anterior part. Ecsoma short, 1020–1530 × 300–680. Lump-like skin
spur termed by Rudolphi (1819) ‘Nackenbuckel’ is present on skin in region of
pharynx on right side. Oral sucker subterminal, funnel-shaped 180–320 × 180–320.
Oral lip present. Caeca extend nearly to posterior end of ecsoma. Ventral sucker
180–280 × 180–300. Testes tandem. Seminal vesicle spindle-shaped, thick-walled,
situated in middle of soma posterior to ventral sucker. Pars prostatica, long, sinu-
ous, tubular, does not coil back over seminal vesicle, surrounded by prostate gland
cells. Sinus-sac in forebody. Genital pore at level of posterior margin of oral sucker.
Ovary dome-shaped, in posterior quarter of soma. Vitellarium consists of seven,
thick, convoluted tubules in two groups of three and four each. Uterine coils extend
into ecsoma. Eggs 24 × 10.
Remarks: This species has been recorded from Lutianus johnii and Mugil speigleri
from the Bombay coast (Chauhan, 1945), from Polynemus paradiseus from Digha
coast, BOB (Kumari & Srivastava 1976) and from Caranx carangus from Madras
coast, BOB (Hafeezullah, 1990). According to Gibson & Bray (1986), L. psettodi

glandulum (after Chauhan,
1945)
Gupta & Puri, 1981 from Psettodes erumei (Bl. & Schn.) from PR, BOB, is a possible
synonym of L. glandulum. This species is characterized by the spindle-shaped thick-
walled seminal vesicle, the ovary and the vitellarium situated in the posterior third
of body, the convoluted vitelline tubes and the uterus extending into ecsoma.
Lecithocladium guptai Gupta & Govind, 1981 (Fig. 9.23)
Description: Body elongate, with long ecsoma, 5.1–5.66 mm long, 720–770 wide;
ecsoma 2470–2750 long, 610–660 wide. Body surface plicated. Oral sucker ter-
minal, cup-shaped, 340 long, 310–330 wide. Caeca extend into ecsoma. Ventral
sucker spherical, slightly larger than oral sucker, 340–350 in diameter. Sucker ratio
1:1.06–1.1. Genital pore median, ventral to anterior part of oral sucker, Testes diago-
nal, contiguous, pre-equatorial. Seminal vesicle spindle-shaped, thick-walled, undi-
vided, extends to anterior testis. Pars prostatica curved, surrounded by many prostate
gland cells, anterior third naked. Hermaphroditic duct tubular, in thick-walled sinus-
sac. Ovary kidney-shaped, pre- or post-equatorial. Uterine coils descending to two-
thirds of ecsoma. Eggs oval, numerous, 23–24 long, 11–14 wide. Vitellarium consists
of seven long slender winding tubules, entering into ecsoma, four on right side, three
on left side.
Remarks: L. guptai differs from most species of Lecithocladium in having the oral
sucker smaller or subequal to the ventral sucker, the smaller seminal vesicle, the
absence of prostatic cells in the anterior part of the pars prostatica and larger eggs.

guptai (after Gupta &
Govind, 1981)
Lecithocladium harpodontis Srivastava, 1942 (Fig. 9.24)

[Syns: Colletostomum muthahi Sahai & Srivivastava, 19778; L. brevicaudum Sri-
vastava, 1942; L. chauhani Hafeezullah, 1975; L. ilishae Bashirullah & D’ Silvain
Basirullah, 1973 (Gibson & Bray 1981; 1986)]
Host: Synodontidae: Harpodon nehereus (Hamilton); Stromateidae: Pampus argen-
teus (Euphrasen)
Locality: BOM, AS; DC, BOB
Reference: Srivastava (1942a), Kumari & Srivastava (1976)
Description: Body elongate, about 5.26 mm long, 0.7 mm wide, with faint plica-
tions. Ecsoma shorter than soma. Oral sucker infundibuliform. Ventral sucker pre-
quatorial, smaller than oral sucker. Intestinal caeca with shoulders, extending into
ecsoma. Testes prequatorial. Seminal vesicle saccular, extending to testicular level.
Pars prostatics tubular, sinuous, surrounded by prostate gland cells. Sinus-sac tubular,
long, enclosing tubular hermaphroditic duct. Genital pore ventral to oral sucker near
anterior margin. Ovary spherical, slightly larger than testes, distant from junction of
soma and ecsoma. Uterine seminal receptacle formed. Vitellarium consists of eight
long-coiled tubules. Uterine coils largely postovarian, extending just into ecsoma.
Eggs 10 × 10.

harpodontis (after
Srivastava, 1942)
Remarks: Originally this species was described from the fish Chrysophrys datnia
Hamilton. It has been recorded in Pampus argenteus from Digha coast by Kumari
& C. B. Srivastava (1976a). It is characterized by its infundibuliform oral sucker, an
elongate pharynx, a small ventral sucker, a saccular seminal vesicle, a pars prostatica
which does not coil over the seminal vesicle, the vitellarium with eight tubules and
small eggs.
Lecithocladium jagannathi Ahmad, 1981 (Fig. 9.25)
Host: Rachycentridae: Rachycentron canadum (L.)
Locality PR, BOB
Number: 12
Description: Body elongate, with long ecsoma; soma 1900–2200 long by 500–540,
ecsoma 1140–1480 long by 420–470 wide. Tegumental plications confined to soma.
Oral sucker oval, subterminal, 260–290 long by 230–250 wide. Ventral sucker
240–270 in diameter, Pharynx elongate. Intestinal bifurcation midway between suck-
ers. Caeca terminate at posterior extremity. Testes contiguous, tandem to slightly
diagonal, anterior testis immediately pre-equatorial and posterior testis immedi-
ately postequatorial. Seminal vesicle spindle-shaped, thick-walled, pretesticular. Pars
prostatica tubular, winding, surrounded by numerous prostate gland cells. Sinus-
sac tubular, thin-walled, reaches to middle of ventral sucker, encloses thin-walled
hermaphroditic duct. Genital pore ventral to posterior part of oral sucker. Ovary near
posterior end of soma, smaller than testes and separated from posterior testis by
uterine coils. Vitellarium consists of seven long winding tubules just extending into
ecsoma. Uterus descends posteriorly into ecsoma to two-thirds of its length. Eggs
14–16 long by 9–11 wide.

jagannathi (after Ahmad,
1981)
Remarks: This species resembles L. angustiovum very closely but differs in having
an oval oral sucker, a thick-walled seminal vesicle, a sinus-sac extending to the
middle of the ventral sucker and the uterus extending into two-thirds of the ecsoma.
Lecithocladium megalaspis Yamaguti, 1953 (Fig. 9.26)
[Syns. L. dawesi Bashirullah e D’Silva, 1973; L. hanumanthai Gupta& Jain, 1992;
L. inglisi Gupta & Ahmad, 1977; L. purense Gupta & Gupta, 1978; L. singhi Gupta
& Singh, 1982]
Host: Carangidae: Megalaspis cordyla (L.)
Locality: BOM, AS
Reference: Hafeezullah (1990); Gibson & Bray (1986); Bray & Cribb (2004)
Description: Body rather slender, up to 3400 long, maximum width 580 at vitelline
level. Tegumental plications prominent on soma. Ecsoma 1.3 mm long Preoral lobe
distinct. Oral sucker approximately funnel-shaped 200 × 210, with membranous,
medianly notched flap on anterior ventral border. Ventral to this notch is another
small transverse flap. Ventral sucker 225 in diameter. Testes subglobular, tandem,
slightly pre-equatorial, with uterine loop between. Seminal vesicle elliptical, with
very thick wall of somewhat spiral muscle fibres, extending obliquely from left side
of anterior testis. Pars prostatica recurved, surrounded by prostate cells throughout
its length. Hermaphroditic duct somewhat enlarged posteriorly, with comparatively
thick wall of longitudinal muscle fibres, rectilinear, reaches just anteriorly to ventral
sucker. Genital pore at posterior end of oral sucker. Ovary subglobular, just post-
equatorial. Uterus descending to middle of ecsoma. Eggs elliptical, 18–20 × 10–12.

megalaspis (after
Hafeezullah, 1990)
Vitellarium consisting of three right and four left long tubular lobes, some of which
are subdivided near tip.
Remarks: This species differs from the most closely related Lecithocladium glan-
dulum Chauhan, 1945, in lacking the ‘Nackenbuckel’ of Rudolphi, and in the length
of the eggs.
Lecithocladium parviovum Yamaguti, 1953 (Fig. 9.27)
[Syn: L. elongatus Gupta & Puri, 1981; L. fotedari Gupta & Singh, 1982; L. keralense
Gupta & Gupta, 1978, L. sciani Gupta & Puri, 1981; L. srivastavai Gupta & Puri,
1981; L. stromateusi Gupta & Puri, 1981]
Description: Body elongate, 4400–8500 long (excluding ecsoma), 1200–1750
wide; ecsoma 3440–7340 long. Tegumental plications extend from anterior end of
body to posterior third of soma. Oral sucker funnel-shaped, 780–1100 long, 700–1100
wide, with preoral lip. Caeca have prominent shoulders and extend to posterior end of
ecsoma. Ventral sucker smaller than oral sucker, spherical, lying in anterior third of
body, 520–950 long, 550–900 wide. Testes ovoid, tandem or symmetrical, contigu-
ous, pre-equatorial, subequal. Seminal visicle spindle-shaped, undivided, surrounded
by muscular wall, extends to anterior border of anterior testis. Pars prostatica curved,
tubular, surrounded by many prostate gland cells. Sinus-sac elongate, thin-walled.
Genital pore near anterior ventral margin of oral sucker. Ovary transversely ovoid,
near posterior end of soma, close or separated from testes. Vitellarium postovarian,
consist of seven long, winding narrow tubules, confined to soma or extend into ante-
rior part of ecsoma. Uterus descends to half of ecsoma. Eggs numerous, small, 13–20
long, 8–11 wide.

parviovum (after Yamaguti,
1953)
Leithocladium puriense Ahmad, 1981

Desription: Body elongate, slender, with long ecsoma. Soma 4100–4860 long
by 800–910 wide. Ecsoma 3880–4140 long by 740–770 wide. Tegumental plica-
tions absent. Oral sucker cup–like, 350–380 long by 370–400 wide; Ventral sucker
380–400 in diameter, nearly same size as oral sucker. Pharynx barrel-shaped. Oesoph-
agus saccular. Caeca long, simple, unequal, terminate short of posterior extremity.
Testes two, oval, tandem. Seminal vesicle spindle-shaped, undivided, thin–walled,
pretesticular. Pars prostatica tubular, curved, surrounded by numerous prostate glands
to half its length. Sinus-sac tubular, thin–walled, reaches to middle of ventral sucker
Ovary larger than testes, well separated from posterior testis. Vitellarium consists of
eight winding tubules, extend anteriorly to posterior testis and posteriorly just into
ecsoma. Uterus entering ecsoma to half of its length. Eggs 13–15 long by 10–11
wide.
Remarks: This species was reported by Ahmad (1981) from Chondroplites chinensis
(Euphr.), (Stromateidae.) from Puri coast, Bay of Bengal. It is characterised by the
long ecsoma, the saccular oesophagus and the presence of eight tubules in the vitel-
larium, some of them extending anteriorly up to ovarian level. The lack of plications
may be a fixation artefact.
Subfamily Lecithochiriinae Lühe, 1901
[Sterrhurinae Looss, 1907; Umatrematinae Srivastava & Sahai, 1978
Diagnosis: Body with well-developed ecosma. Tegument smooth. Suckers well-

developed, presomatic pit present or absent. Caeca may or may not extend into
ecsoma. Testes two, tandem, symmetrical or oblique. Seminal vesicle globular or
elongate, constricted into portions or tubular and convoluted, usually in forebody,
may extend into hindbody. Pars prostatica short, vesicular or tubular. Sinus-sac
encloses distinct ejaculatory duct or prostatic vesicle and metraterm. Permanent sinus
organ absent. Genital atrium usually small. Ovary oval uterus distributed in pre- and
postovarian fields. Vitellarium seven digitiform or oval lobes, in lateral fields of three
and four. Excretory arms united in forebody.
Type genus: Lecithochirium Luhe, 1901

Genus Lecithochirium Lühe, 1901
[Syns. Sterrhurus Looss, 1907; Neohysterolecitha Ahmad, 1977; Anaplerurus Sri-
vastava & Sahai, 1978; Umatrema Srivastava & Sahai, 1978; Reference: Gibson
(2002)]
Generic Diagnosis: with characters of the subfamily as mentioned above.
The genus was discussed in detail by Gibson & Bray (1986) and later by Bray
(1991). It is a large genus with about 100 nominal species. Morphologically they are
complex and several differences occur that have a taxonomic significance including
modifications of the oral sucker, presence or absence and glandular nature of the
presomatic pit, differences in sinus-sac, prostatic vesicle, pars prostatica and seminal
vesicle. Bray (1991) devised a key for the differentiation of species. He has also
identified species groups, each group identified by a common feature. Molecular
characterization was carried out for two species: L. grandiporus (Rudolphi, 1819)
by Abdel-Ghaffar et al. (2013) and L. priacanthi Yamaguti, 1953 by Abdel-Ghaffar
et al. (2015).
The genus is represented by 23 species in Indian marine fish. Some of these species
were transferred from the genus Sterrhurus Looss, 1907, which is synonymized
with Lecithochirium. This synonymy has resulted in many homonyms which need
renaming. Apart from this, many of the species are not well characterized and the
descriptions lack details. The genus needs a thorough revision. Critical analysis of
the genus at this stage is difficult. Only species that are well defined are included in
this account. The description of the details of the terminal genitalia is essential for
accurate identification of members of this genus. In many cases this is not available.
Species of Lecithochirium so far recorded from marine fish of India, their valid
status, the hosts and the localities, authors are listed in Table 9.4. Descriptions are
provided for a few selected species.
Table 9.4 Species of Lecithochirium recorded from marine fishes of India, their hosts, localities
and valid status
Species of Lecithochirium Valid status Host Locality References
L. acutum Chauhan, 1945 Valid Ariidae: Arius BOM, Chauhan
falcarius Ernakulam, (1945, 54);
Trichiuridae: Calicut, (AS) Gupta, N.K.
Trichiurus sp. & Sehgal
Scombridae: (1970a) Gupta
Acanthocy- N.K. &
bium solandri Mehrotra
(Cuvier) (1970b);
Gupta, N.K. &
Puri (1981)
L. agarwali (Gupta & Gupta, To be assessed Trichiuridae: PR, BOB Gupta, R.C. &
1989 Eupleuro- Gupta,V.C. (1989)
grammus
muticus Gray
Lecithochirium anisotestes Not known Synodontidae: VSK, BOB Surekha &
Surekha & Vijayalakshmi, Saurida tumbil Vijayalakshmi
2007 Bloch (2007)
L. bilqeesae (Gupta & To be assessed Trichiuridae: PR (BOB) Gupta, P.C. &
Gupta, 1989) Eupleuro- Gupta,
grammus V.C. (1989)
muticus Gray
L. cynoglossi To be assessed Cynoglossidae: PR, BOB Gupta, V &
Gupta & Jain, 1991 Cynoglossus Jain (1991c)
sp.
L. durdanae (Gupta & Valid. Synodidae: PR, BOB Gupta, P.C. &
Gupta, 1989) Synodus Gupta,
(Saurus) V.C. (1989)
indicus (Day)
L. ernakulamense Dwivedi To be assessed Priacanthidae: Ernakulum, Dwivedi &
& Gupta, 2007 Priacanthus AS Gupta (2007)
blochi
(Bl. & Schn.)
L. fotedari Gupta & Gupta, To be assessed Ophithyidae: PR, BOB Gupta &
1990 Ophichthys Gupta (1990)
boro
(Hamilton)
L. indicum (Gupta & Clupeidae: Gupta &
Govind, 1984) Sar- Govind (1984)
dinella brachysoma
Bleeker
L. leiperi Gupta & Singh, Valid Carangidae: PR, BOB Gupta &
1981 Apolectus Singh (1981)
niger (Bloch)
L. mugili Gupta & Jain, 1991 To be assessed Mugilidae: PR, BOB Gupta, V &
Mugil sp. Jain (1991)
L. pampi Gupta & Puri, 1981 To be assessed Carangidae: Calicut, AS Gupta & Puri
Apolectus (1981)
niger (Bloch)
(continued)

Species of Lecithochirium Valid status Host Locality References
L. polynemi Chauhan, 1945 Valid Polynemidae: BOM, Chauhan,
Leptome- Ernakulam, (1945); Gupta
lanosoma AS N.K. &
indica Shaw Mehrotra
Mugilidae: (1970b).
Liza parsia Gupta, N.K. &
Trichiuridae: Sehgal
Trichiurus lep- (1970a);
turus Gupta, N.K. &
(Linnaeus) Puri (1981)
Scombridae:
Acanthocy-
bium
sp.
L. pristopomai Gupta & Jain, To be assessed Haemulidae: PR (BOB) Gupta, V &
1991) Pristipoma sp. Jain (1991)
L. pritchardae (Singh & To be Elopidae: PR (BOB) Singh &
Gupta, 1984) Madhavi, 2011 assessed Elops saurus Gupta (1984)
(Linn)
L. tanimae Gupta & Jain, To be assessed Marine fish PR (BOB) Gupta, V &
1991 Jain (1991)
L. testelobatus Surekha & Saurida VSK (BOB) Surekha &
Vijayalakshmi undosquamis Vijayalakshmi
(2005a)
L. tumbilense Surekha & Valid Synodontidae: VSK (BOB) Surekha &
Vijayalakshmi, 2005 Saurida tumbil Vijayalakshmi
a (Richardson) (2005b)
Lecithochirium acutum Chauhan, 1945 (Fig. 9.28)

Hosts: Ariidae: Arius fulcarius (1, 2); Trichiuridae: Trichiurus sp. (Forskål) (3, 4);
Scombridae: Acanthocybium solandri (Cuvier) (5)
Localities: BOM, AS (1, 2); Ernakulam, (3,4), Calicut (5), AS
Distribution: Indian ocean, Kuwait coast (6)
References: 1. Chauhan (1945); 2. Chauhan (1954); 3. Gupta, N.K. & Sehgal (1970a);
4. Gupta, N.K. & Mehrotra (1970b); 5. Gupta, N.K. & Puri (1981); 6. Nahhas & Sey
(2002)
Description: Body elongate 3232–4400 in length, 560–800 in maximum breadth.

Ecsoma 464–656 × 352–416 long. Preoral lobe well developed. Oral sucker sub-
terminal, 96–120 × 96–14.4. Ventral sucker large, 352–384 × 204–400. Sucker ratio
1:3.06–3.83. Pre-somatic pit well developed, glandular with oval opening. Caeca
extend to near posterior end of soma. Testes rounded or transversely oval, tandem or
slightly diagonal, lie posterior to ventral sucker. Seminal vesicle anterodorsal to ven-
tral sucker, tripartite, anterior portion thin-walled. Sinus-sac encloses hermaphroditic
Fig. 9.28 Lecithochirium

acutum
duct and small prostatic vesicle. Genital pore at level of oesophagus. Vitellarium two
indented masses just posterior to ovary, one consisting of 3 and the other of 4 lobes.
Uterus heavily coiled. Eggs 12 × 6.8–8.0.
Remarks: This species was first described by Chauhan (1945) from Arius fulcarius
and later from Trichiurus sp. by Chauhan (1954) and from T. haumela by Gupta &
Sehgal (1971). Nahhas & Sey (2002) reported it from T. lepturus from Kuwait.
Nama (1982) considered it a synonym of L. rufoviridae (Rud, 1819) Luhe, 1901 but
L. acutum is retained as a valid species (Nahhas & Sey, 2002).
Lecithochirium durdanae (Gupta & Gupta, 1986) (Fig. 9.29)

Host: Synodontidae: Saurua indicus (Day)
Locality: PR, BOB
No. 5
Body 3900–5880 long 920–1110 wide. Ecsoma partially extended. Oral sucker
160–190 by 180–250. Ventral sucker 460–560 × 420–570. Sucker ratio 1:2.28–2.33.
Caeca not extending into ecsoma. Testes spherical, preequtorial. Seminal vesicle
tripartite, thin-walled, extends to anterior margin of ventral sucker. Sinus-sac well
developed. Ovary entire. Vitellarium 7-lobed. Uterus confined to soma. Eggs 14–16 ×
8–9.
Remarks: Characterized by the thin-walled tripartite seminal vesicle and the uterus
not extending into ecsoma.

durdanae (after Gupta &
Gupta, 1986)
Family Hemiuridae Looss, 1899

Lecithochirium fotedari Gupta & Gupta, 1990
Host: Ophichthidae: Ophichthys boro (Hamilton)
Locality: PR, BOB
Number: 6
Reference: Gupta, P.C. & Gupta, V. C. (1990)
Description: Body 3050–3710 (including ecsoma) × 470–750. Ecsoma 190–490 ×

370–460. Oral sucker terminal, 120–150 in diameter. Caeca simple, entering into
ecsoma or not. Pit-like structure present in forebody. Testes pre-equatorial, over-
lapping each other or separate, obliquely tandem or tandem. Seminal vesicle thin-
walled, tripartite, extends dorsally to ventral sucker. Pars-prostatica surrounded
by many prostate gland cells, opening jointly with metraterm into hermaphroditic
duct. Hermaphroditic duct small, enclosed in sinus-sac. Genital pore lying posterior
to pharynx. Ovary entire, oval, median or submedian, postequatorial. Vitellarium
compact, consisting of seven lobes. Uterus coiled, not entering ecsoma, descending
to just posterior to vitellarium. Eggs oval, small, 17–19 × 7–8.
Remarks: L. fotedari is closely related to L. leiperi Gupta & Singh, 1981 but differs
from it in having obliquely tandem testes instead of symmetrical and the uterus not
entering the ecsoma. The genital pore is described as ‘marginal.’ This is not reported
in the Hemiuridae and the illustration indicates that the worm figured was lying on
its side, and the genital pore is in the usual, median position.
Lecithochirium indicum (Gupta & Govind, 1984) Madhavi, 2011 (Fig. 9.30)
[Syn: Sterrhurus indicum Gupta & Govind, 1984]
Host: Clupeidae: Sardinella brachysoma (Bleeker)
Locality: PR, BOB
Reference: Gupta, P.C. & Govind (1984a)

indicum (after Gupta &
Govind, 1984)
Description: Body elongate, 3170–4960 (including ecsoma) × 860–1060, ecsoma

910–1990 × 660–1020. Oral sucker subterminal, 170–200 × 160–200. Caeca extend
into ecsoma for more than half of its length. Forebody without pit. Ventral sucker
larger than oral sucker, 420–490 × 400–470. Sucker ratio 1:2.4. Genital pore subme-
dian, post-bifurcal. Testes symmetrical, pre-equatorial. Seminal vesicle thin-walled,
tripartite, reaches to and just overlaps ventral sucker. Pars prostatica short, surrounded
many prostate gland cells, opens jointly with metraterm into hermaphroditic duct,
enclosed in sinus-sac. Vitellarium with seven short digitiform lobes, lying postero-
lateral to ovary. Uterus coiled, not extending into ecsoma. Eggs 13–15 × 7–9.
Remarks: This species is characterized by the following combination of characters:

large ventral sucker, thin-walled tripartite seminal vesicle, digitiform vitelline lobes,
caeca extending into ecsoma, uterus confined to soma and small egg size.
Lecithochirium jairajpurii (Gupta & Govind, 1984) Madhavi, 2011

[Syn: Sterrhurus jairajpuri Gupta & Govind, 1984]
Locality: PR, BOB
Reference: Gupta, P.C.& Govind (1984a)
Description: Body elongate, smooth, 3.66–4.05 (including ecsoma). 802–870;

ecsoma 1800–1930 × 760–870. Oral sucker subspherical, 160–180 × 220–240.
Intestinal caeca extend into ecsoma for more than half of its length. No pit in fore-
body. Ventral sucker pre-equatorial, larger than oral sucker, 340–370 × 350–360.
Sucker ratio 1:1.5. Genital pore median or submedian, post-bifurcal. Testes sym-
metrical, close to posterior end of ventral sucker or slightly overlapping its posterior
margin. Seminal vesicle thin-walled, tripartite, anterior or anterolateral to ventral
sucker, extends up to just posterior to anterior margin of ventral sucker or up to

its level. Pars prostatica short, surrounded by large number of prostate gland cells;
opens jointly with metraterm into ductus hermaphroditicus. Hermaphroditic duct in
oval sinus-sac. Ovary post-testicular, Uterus extends up to middle of ecsoma. Eggs
17–19 × 8–9. Vitellarium compact, consist of seven digitiform lobes.
Remarks: This species was differentiated from S. indicum described above by the
smaller sucker proportions (1:2.1 as against 1:2.5), the digitiform vitelline lobes, the
extension of the uterus into ecsoma and in the location of ovary and the vitelline com-
plex near the posterior end of the soma.
Lecithochirium leiperi Gupta & Singh, 1981 (Fig. 9.31)
Host: Carangidae: Paratromateus niger (Bloch)
Locality: PR, BOB
Number: 2
Reference: Gutpa, P.C.& Singh (1981)
Description: Body elongate, 2510–3190 long, 570–750 wide. Ecsoma 620–730

long. Oral sucker subspherical, 150 long 140–200 wide Caeca extend into ecsoma
up to half of its length. Pit in front of ventral sucker surrounded by muscular wall,
present. Ventral sucker 350–460, 410–510 wide. Sucker ratio 1:2.5–3.0. Genital
pore submedian at posterior border of pharynx. Testes pre-equatorial, symmetrical.
Seminal vesicle thin-walled, tripartite extends anterolateral to ventral sucker up to
its anterior third level. Pars prostatica short, vesicular, surrounded by a large number
of gland cells. Ovary post-equatorial. Uterus extends into ecsoma. Eggs 17–21 by
9–11. Vitellarium consists of seven vitelline lobes, four sinistral and three dextral.
Remarks: This species closely resembles L. polynemi Chauhan, 1945, and L. acutum
Chauhan, 1945, but differs from L. polynemi, in the extent of seminal vesicle up

leiperi (after Gupta & Singh,
1981)
to anterior third level of ventral sucker, in the shape of pars prostatica, and from
L. acutum in having testes symmetrical, in the extent of caeca and uterus into ecsoma.
Lecithochirium polynemi Chauhan, 1945 (Fig. 9.32)
Hosts: Polynemidae: Polynemus indicus Shaw; Mugilidae: Liza parsia Hamilton (1);
Trichiuridae: Trichiurus lepturus (L) (2); Scombridae: Acanthocybium sp. (3)
Habitat: Digestive tract
Locality: BOM, Ernakulam (South India), AS
Reference: Chauhan, (1945); 2. Gupta N.K. & Mehrotra (1970b); 3. Gupta, N.K. &
Sehgal (1970a); 4. Gupta, N.K. & Puri (1981)
Distribution: New Caledonia (Manter, 1969)
Description: Body 3632–4288 mm in length and 696–712 in maximum breadth.
Ecsoma 704–1232 × 336–656 mm. Oral sucker 128–136 × 112–160, with well-
developed pre-oral lobe. Ventral sucker, 416–512 × 288–336 mm. Sucker ratio
1:3.31–3.76. Pre-somatic pit present, appears as small sucker. Caeca extend into
ecsoma, terminate at about half length of ecsoma. Testes oval or rounded in shape,
oblique, or tandem in anterior half of the soma. Seminal vesicle anterior to ventral
sucker or extends to middle of ventral sucker, tripartite, posterior part largest of all.
Pars prostatica short, leads into hermaphroditic duct. Genital pore at level of pharynx
or slightly behind it. Ovary post-testicular. Vitellarium compact in form of two dig-
itiform masses lying just behind ovary, one mass consisting of four lobes and other
three. Uterus extends into ecsoma. Eggs l4–16 × 8.
Remarks: According to Chauhan (1945) this species differs from all the species of
the genus then known in the posterior extent of caeca and uterus into the ecsoma,
in the disposition of uterine coils, relative position of the gonads, the nature of male

polynemi (after Chauhan,
1945)
terminal ducts and the extent of the seminal vesicle. Apart from the Indian Ocean, L.
polynemi has also been recorded from New Caledonia from a lutjanid fish (Manter,
1969).
Lecithochirium pristopomai (Gupta & Jain, 1991)
Host: Haemulidae: Pristipoma sp.
Locality: PR, BOB
Reference: Gupta & Jain (1991)
Remarks: Information about this species not available.
Lecithochirium pritchardae (Singh & Gupta,1984) Madhavi, 2011 (Fig. 9.33)
Host: Elopidae: Elops saurus (Linnaeus)
Locality: PR, BOB
Reference: Singh & Gupta (1984)
Description: Body elongate, aspinose, 4580–4990 long (including ecsoma),
1250–1470 wide. Ecsoma 1640–2200 long; 1080–1900 wide. Oral sucker 210–240
long, 270–280 wide. Intestinal caeca simple, extend into ecsoma for more than half
of its length. Pit absent. Ventral sucker 450–510 long, 460–520 wide. Sucker ratio:
1.6–1.9. Testes symmetrical, contiguous or apart, lying in anterior half of body, close
to ventral sucker. Seminal vesicle tripartite, thin-walled extends obliquely either
to anterior margin of ventral sucker or just posterior to it. Pars prostatica short.
Hermaphroditic duct tubular, enclosed in a thick-walled sinus-sac. Genital pore
submedian, extracaecal or on dextral caecum. Ovary entire, pre-equatorial. Uterus
extends into ecsoma, descending to just anterior to middle of ecsoma. Vitellarium
compact, consisting of seven digitiform lobes. Eggs 16–17 long, 8–9 wide.
Remarks: This species is characterised by the thin-walled tripartite seminal vesicle,
symmetrical testes, smaller ventral sucker in relation to body length.

pritchardae (after Singh &
Gupta, 1984)

testelobatus (after Surekha &
Vijayalakshmi, 2005b)
Lecithochirium testelobatus Surekha & Vijayalakshmi, 2005 (Fig. 9.34)

Host: Synodontidae: Saurida undosquamis (Richardson)
Locality: VSK, BOB
No. 5
Reference: Surekha & Vijayalakshmi (2005a)
Description: Body triangular, 2550–2710 × 500–740. Ecsoma 600–950 long. Oral
sucker 13–170 × 110–130. Ventral sucker 210–310 × 250–300 in size. Sucker ratio
1:1.5. Caeca not extending into ecsoma. Testes two, large, lobed, each testis with lat-
eral lobate extension. Seminal vesicle tripartite, thin-walled and convoluted present
in between intestinal bifurcation and ventral sucker. Pars prostatica elongate, sur-
rounded by numerous prostatic cells. Hermaphroditic duct straight. Genital atrium
deep, tubular, genital pore median, immediately posterior to oral sucker. Ovary oval,
post-testicular. Vitellarium 6-lobed, lobes are short, post ovarian. Uterine coils not
extending into ecsoma. Eggs 22–25 × 15–17.
Remarks: The important features of the species are: Body triangular, ecsoma broad,
thick-walled, testes lobed located in the equatorial plane of hindbody.
Lecithochirium tumbilense Surekha & Vijayalakshmi, 2005
Locality: VSK, BOB
Reference: Surekha & Vijayalakshmi (2005b)
Description: Body elongate, 4050–4200 long 760–910 wide. Ecsoma half as long
as body, 1300–2100 long. Oral sucker 190–250 × 230. Ventral sucker at midbody,
350–450 × 340–440. Sucker ratio 1:2. Caeca extend into ecsoma, terminate just ante-
rior to terminal region. Testes round to oval. Seminal vesicle triprtite, pars prostatica
long surrounded by prostate gland cells. Ejaculatory duct present in the sinus-sac
and linked anterodorsally with pars prostatica. Ovary rounded, located in ecsoma.
Vitellarium with 4 and 3 deep lobes, overlapped by ovary within the ecsomal region.
Uterine coils distributed both in ecsoma and body proper. Eggs 20–25 × 15–18.
Remarks: The special character of this species is the location of ovary and vitellarium
in the ecsoma instead of the soma. This character needs verification since it is unsual
to find the gonads located in the ecsoma.
Subfamily Plerurinae
Diagnosis: Body small, spindle-shaped, ecsoma reduced or well developed, body

surface smooth. Pharynx well developed, Caeca terminate in ecsoma. Testes preovar-
ian, symmetrical to tandem, usually oblique. Seminal vesicle elongate, constricted
into two or three or four sections, thin-walled. Pars prostatica vesicular or tubu-
lar, may be partly enclosed by muscles of Sinus-sac apparently absent or poorly
developed. Sinus organ present or absent. Hermaphroditic duct commonly vesicular
proximally and tubular distally. Genital atrium usually deep. Genital pore midven-
tral, in forebody. Ovary entire or lobed. Juel’s organ present. Uterus extends into
ecsoma. Vitellarium post ovarian, composed of two masses, lobed. Eggs numerous.
Excretory vesicle Y-shaped, arms united anteriorly. Type genus Plerurus.
Genus Plerurus Looss, 1907

[Syn. Paraplerurus Fischthal & Kuntz, 1963; Voitrema Yamaguti, 1971]
Diagnosis: With characters of the subfamily. Sinus organ absent. Vitelline lobes
tubular or digitiform. Type species: P. digitatus (Looss, 1899).
Plerurus digitatus (Looss, 1899) Looss, 1907 (Fig. 9.35)

[Syn. Plerurus ashiae Gupta & Sayal, 1979; Lecithochirium carangis Yamaguti,
1942; Lecithochirium lobatum Yamaguti, 1952; Lecithochirium longicaudatum
(Yamaguti, 1953) Gibson & Bray, 1979; Plerurus alectis Yamaguti, 1970; Pleru-
rus ashiae Gupta & Sayal, 1979; Plerurus atulis Shen, 1990; Plerurus carangi
Parukhin, 1966; Plerurus carangis (Yamaguti, 1942) Manter & Pritchard, 1960;
Fig. 9.35 Plerurus digitatus

(After Bray, 1990)
Plerurus cynoglossi Wang, 1982 Plerurus hainanensis Gu & Shen, 1981 Pleru-
rus kawakawa Yamaguti, 1970; Plerurus longicaudatus (Yamaguti, 1953); Plerurus
opakapaka Yamaguti, 1970; Plerurus pomadasydis Shen & Tong, 1985; Plerurus
sauridae (Fischthal & Kuntz, 1963); Plerurus scomberomori Shen, 1990; Plerurus
spatulocirrus Gu & Shen, 1981; Plerurus sphyraenae Yamaguti, 1970; Plerurus
trichiuri Gu & Shen, 1981; Sterrhurus longicaudatus Shen, 1987 (Reference: Bray
(1990c)].
Host: Scombridae: Acanthocybium sp.
Reference: Gupta, N.K. & Sayal (1979); Bray (1990c)
Locality: Kavaratti (Laccadive Islands)
Distribution: Mediterranean sea; Atlantic, Indian Ocean: BOB, Western Australia,
Pacfic, South China Sea; Taiwan, China; Pacific, Central Thailand, Philippines,
Pacific Hawaii.
Description: Body elongate. 4190–4680 in total length including ecsoma, the lat-
ter 850–1380 long and constricted from body proper; maximum width of body
1070–1240. Oral sucker 300 × 220–300. Intestinal caeca long, terminate at pos-
terior end of ecsoma. Ventral sucker large, 960–1000 × 960–1040. Sucker ratio
1:3.1–3.0. Testes two, ovoid, diagonal. Seminal vesicle tripartite, thin–walled. Pars
prostatica vesicular, with distinct glandular sheath. Short ejaculatory duct joins with
metraterm to form cylindrical, narrow hermaphroditic duct. Sinus-sac absent. Gen-
ital pore mid–ventral, just post-bifurcal. Ovary reniform. Uterine coils may or may
not extend into ecsoma, main bulk of uterus lies posterior to vitellarium, metraterm
distinct. Vitellarium in two masses, with digitiform lobes, three lobes on left and four
on right. Eggs 18–24 × 6.8.
Remarks: Bray (1990c) discussed in detail the morphology, hosts, geographic dis-
tribution and synonymies of this species based on material collected from different
species of fish and geographic localities, and also the holotype slides of relevant
species and synonymized several species of hemiurids with Plerurus digitatus. The
only Indian species of the genus, P. ashiae Gupta & Sayal, 1970 collected from
Acanthocybium sp. from Laccadive Islands is also considered as a synonym of P.
digitatus. As a result of these synonymies P. digitatus has emerged as the most
cosmopolitan having wide geographic range and euryxenic hemiurid species. The
hosts and geographic localities for the species were given by Bray (1990c) and valid
reasons were given for synonymizing the various species. The Indian reports are:
P. ashiae Gupta & Sayal, 1970 from Acanthocybium sp. from Laccadive Island;
Plerurus longicaudatus from Euthynnus affinis from VSK, BOB by Madhavi & Rao
(1974) and Saurida undosquamis from MS, BOB by Nellaiappan & Ramalingam
(1980) and Satyanarayana (1982).
Subfamily Hemiurinae Looss, 1899

[Syn. Pseudoprosorchiinae Gupta & Jain, 1992]
Diagnosis: Ecsoma present. Body surface plicated. Testes tandem or oblique. Sem-
inal vesicle oval or bipartite, thin-walled, in hindbody. Pars prostatica tubular, pro-
static cells numerous. Hermaphroditic duct enclosed in sinus-sac. Sinus organ absent.
Ovary single. Vitellarium composed of one or two oval masses. Utrerine coils exten-
sive. Excretory arms united in forebody. Type genus Hemiurus Rudolphi, 1809.
Genus Parahemiurus Vaz & Pereira, 1930

[Syn. Anahemiurus Manter, 147; Bapatina Srivastava & Sahai, 1977; Daniella Sri-
vastava & Sahai, 1977; Dentiacetabulum Srivastava & Sahai, 1977; Engraulitrema
Gupta & Jain, 1992, Lecithomonium Shen, 1987; Trilecithotrema Gu & Shen, 1981]
The genus Parahemiurus is a fairly large genus comprising nearly 22 species. It is
characterized by the following features: presence of ecsoma, plicated tegument, oval
seminal vesicle situated in hindbody, long pars prostatica surrounded by numerous
prostatic cells, presence of a hermaphroditic duct but absence of sinus organ, and
vitellarium composed of two oval or lobed masses. The type species Parahemiurus
merus (Linton, 1910) has a wide geographic distribution confined to temperate region
and is highly euryxenic. However, criteria for distinguishing species in this genus
are not clearly specified, as a result confusion prevails in determining the valid-
ity of the species reported under it. Fourteen species of Parahemiurus have been
reported from Indian marine fish, of which only five species are regarded as valid: P.
engraulisi Gupta & Jahan, 1975; P. gastricus (Sahai & Srivastava, 1977) Hafeezullah,
1990; P. madrasensis (Sahai & Srivastava, 1977) Bray, 1990; P. hamiltoni (Gupta
& Jain, 1992) and P. yanamensis Hafeezullah, 1980. The synonymized species are:
P. cameroni Gupta & Ahmad, 1979; P. puriensis Ahmad, 1981; P. simhai Gupta &
Gupta, 1978 and P. triacanthusi Gupta & Puri, 1982 with P. engraulisis Gupta &
Jahan, 1975 and P. dussumieriai Hafeezullah, 1981 and P. indicus Ahmad, 1981
with P. madrasensis (Sahai & Srivastava, 1977) (Reference: Hafeezullah, 1990a;
Bray, 1990a. Bray (1990a) reviewed the genus Parahemiurus and provided a key for
the separation of valid species of the genus. The possibility exists that ultimately all
the species may prove to be synonymous to P. merus, the type species.
Parahemiurus cameroni Gupta & Ahmad, 1979
[Syn of P. engraulisi Gupta & Jahan, 1975] (Reference: Bray, 1990a)
Host: Engraulidae: Thryssa hamiltoni (Gray)
Locality: PR, BOB
Reference: Gupta & Ahmad (1979)
Parahemiurus dussumieriai Hafeezullah, 1981
[Syn. of P. madrasensis (Sahai & Srivastava, 1977)] (Reference: Bray, 1990a)
Host: Clupeidae: Dussumieria acuta (Val.)
Locality: South Indian Ocean
Parahemiurus engraulisi Gupta & Jahan, 1977 (Fig. 9.36)

[Syn. P. cameroni Gupta & Ahmad, 1979; P. simhai Gupta & Gupta, 1978; P. puriensis
Ahmad, 1981; P. triacanthusi Gupta & Puri, 1982] (Reference: Hafeezullah 1990)
Host: Clupeidae: Ilisha filigera (Valenciennes); Englaulidae: Thryssa hamiltoni
(Gray), and Thryssa· mystax (Schneider); T. dussumierii; Carangidae: Carangoides
malabaricus. Triacanthidae: Triacanthus strigilifer
Location: Stomach
Localities: Visakhapatnam (Bay of Bengal), Tuticorin (Gulf of Mannar), and Madras
(Bay of Bengal)
Reference: Gupta, V & Jahan (1977); Hafeezullah (1980b); Gupta & Ahmad (1979);
Ahmad (1981a)
Description: Body cylindrical, ecsoma well developed, 3620–5800 long, 650–1000
wide. Ecsoma fully evaginated, 530–2360 long, 540–740 wide. Oral sucker 280–360
long, 240–350 wide. Intestinal caeca extend into ecsoma to near its posterior end.
Ventral sucker equal or larger than oral sucker, 310–360 by 330–380 wide. Genital
pore median, at posterior end of oral sucker. Testes equatorial or pre-equatorial, diag-
onal or tandem, contiguous. Seminal vesicle subspherical or elongated, undivided
and thick-walled sac, extends up to anterior end of anterior testis. Pars prostatica
long and narrow, surrounded by prostate gland cells. Hermaphroditic duct narrow,
thick-walled extends from genital pore to middle of ventral sucker or just anterior to
it. Ovary equatorial or post equatorial. Uterus extends to anterior two third length of
ecsoma Eggs 21–10 long, 10–20 wide. Vitellarium two masses, deeply divided into
5–7 lobes, immediately post ovarian.
Fig. 9.36 Parahemiurus

engraulisi (= P. puriensis)
(after Ahmad, 1981)
Remarks: This species is distinguished from all the other species of the genus by
the digitiform nature of the vitelline masses.
Parahemiurus gastricus (Sahai & Srivastava, 1977) Hafeezullah, 1990
[Syn. Dentiacetabulum gastricusSahai & Srivastava, 1977]

Host: Clupeidae: Dussumieria acuta (1) Sardinella sirm (Walbaum) (2) Locality:
Madras, BOB; (1) Tuticorin (Gulf of Mannar) (2).
Reference: 1. Sahai & Srivastava (1977); 2. Hafeezullah (1990a)
Remarks: Sahai & Srivastava (1977) described Dentiacetabulum gastricus from

Dussumieria acuta from Madras (Bay of Bengal). Hafeezullah (1990a) collected
the same species from Sardinella sirm from Tuticorin and synonymized Dentiac-
etabulum with Parahemiurus Vaz & Pereira,1930. P. gastricus in having a short
and saccular sinus-sac close to the pharynx, resembles Parahemiurus yanamense
Hafeezullah, 1980, but the latter has a small ecsoma in comparison with the size of
the soma, whereas in the former the ecsoma is only slightly shorter than the soma
proper, secondly, the sides of the body in the latter are almost parallel whereas in the
former this is not the case, being widest at the junction of the body proper and the
ecsoma. Bray (1990a) expressed the view that it may be a synonym of P. madrasensis
(Sahai & Srivastava, 1977).
Parahemiurus hamiltoni (Gupta & Jain, 1992) nec P. hamiltoni Hafeezullah, 1980
Syn: Engraulitrema hamiltoni Gupta & Jain, 1992
Engraulidae: Thryssa hamiltoni (Gray)
Locality: PR, BOB
Reference: Gupta, V & Jain (1992)
Parahemiurus indicus Ahmad, 1981

Synonymized with P. madrasensis (Sahai & Srivastava, 1977) by Bray (1990a)
Host: Clupeidae: Dussumieria acuta (Cuv. & Val.)
Locality: PR, BOB
No. 38
Reference: (Ahmad, 1981)
Parahemiurus madrasensis (Sahai & Srivastava, 1977) Bray, 1990
[Syn: Daniella madrasensis Sahai & Srivastava, 1977; Parahemiurus merus of Gupta
& Sehgal (1971); P. indicus Ahmad, 1981; P. dussumieriai Hafeezullah, 1981. Ref-
erence:Hafeezullah (1981), Bray (1990a)]
Host: Clupeidae: Sardinella albella (Val.) (1); Dussumieria elopsoides (Bleeker) (2);
D. acuta (3,4)
Locality: BOB (2,3,4), Southern Indian Ocean (1)
Reference: 1. Sahai & Srivastava (1977); 2. Gupta N.K. & Sehgal (1970a); 3. Ahmad
(1981a); 4. Hafeezullah (1981)
Remarks: This species is distinguished by its elongated body shape, plications

extending through the entire body, position of the thin-walled seminal vesicle and
the position of ovary and vitelline masses.
Parahemiurus puriensis Ahmad, 1981

[Synonymized with P. engraulisi Gupta & Jahan, 1977 by Bray (1990)]
Host: Engraulidae: Thryssa dussumieri (Val.)
Locality: PR, BOB
Parahemiurus simhai Gupta & Gupta, 1978

[Syn. of P. engraulisi Gupta & Jahan, 1977 Reference: Hafeezullah, 1980]
Host: Carangidae: Caranx malabaricus (Cuv. & Val.)
Locality: Quilon, Kerala, AS
No. 5
Parahemiurus triacanthusi Gupta & Puri, 1982

[Synonymized with P. engraulisi Gupta & Jahan, 1977 by Bray (1990])
Host: Triacanthidae: Pseudotriacanthus strigilifer Cantor
Locality: PR, BOB
Reference: Gupta, V & Puri (1982a)
Parahemiurus yanamensis Hafeezullah, 1980
Host: Engraulidae: Thryssa hamiltoni (Gray)
Locality: Godavari Estuary
No. 4
Description: Body cylindrical, ecsoma partially or fully extended. Body proper

3644–4276 long, 481–522 wide. Ecsoma 412–550 long. Tegumental plications to
level of vitellarium. Oral sucker terminal, 151–175 × 96–165, smaller than ventral
sucker. Ventral sucker 192–283 in diametre, situated near anterior end of body. Sucker
ratio 1:1.4–2.1. Caeca extend into ecsoma or not. Testes tandem, separated by uterine
coils. Seminal vesicle midway between ventral sucker and anterior testis, unipartite,
sac-like, thin-walled. Pars prostatica long and winding, prostatic cells well developed,
surround pars prostatica to dorsal to ventral sucker. Sinus-sac short, wide, dextral to
pharynx, contains hermaphroditic duct. Genital pore at base of oral sucker. Ovary
kidney-shaped. Vitellarium two compact bodies immediately post ovarian, indented.
Uterus extends into ecsoma and to level of pharynx. Eggs 17–19 × 9–10.
Remarks: P. yanamensis is characterized by having a cylindrical pharynx, a sinus-

sac which is quite wide and barrel-shaped and a pars prostatica that is only partially
surrounded by prostatic cells. It closely resembles P. engraulisi Gupta & Jahan, 1977
but differs from it in the cylindrical pharynx’ the barrel-shaped sinus-sac, the indented
vitelline bodies and the pars prostatica partly surrounded by prostatic cells.
Subfamily Opisthadeninae Yamaguti, 1970

[Syn. Intuscirrinae Skrjabin & Guschanskaja, 1959]
Diagnosis: Body elongate, ecsoma absent. Tegument smooth. Ventral sucker in ante-
rior half of body. Oesophagus short. Caeca terminate bindly. Testes tandem to oblique,
situated well posterior to ventral sucker, not separated from ovary by uterine coils.
Seminal vesicle tubular to saccular, in fore or hindbody. Pars prostatica long or short.
Ejaculatory duct present or absent. Sinus-sac present, encloses hermaphroditic duct.
Sinus organ usually absent. Ovary oval, close to testes. Blind seminal receptacle
present. Uterus mainly preovarian or postovarian. Vitellarium forms two entire or
slightly indented masses, poterolateral to ovary. Excretory arms short, united anteri-
orly. Type genus: Opisthadena Linton, 1910.
Genus Neopisthodena Machida, 1980
Neopisthadena habei Machida, 1980 (Fig. 9.37)
Host: Kyphosidae: Kyphosus cinerascens Forsskal, Ashen drummer
Locality: Tuticorin (Gulf of Mannar).
Description: Body 5.0–9.75 mm long, 1.21–0.96 mm wide. Tegument thick. Ventral
sucker 692–1075 long, 708–1108 wide, globular, situated at 966–1608 from anterior
and of body. Oral sucker 250–383 long, 350–550 wide, subterminal, withdrawn
in anterior part of body. Sucker width ratio about 1:1.6. Proximal parts of caeca
form shoulder, then turning posteriad, extending to posterior end of body. Testes
transversely oval, tandem, posterior to ventral sucker. Seminal vesicle long, narrow,
Fig. 9.37 Neopisthadena

habei (after Hafeezullah,
1990)
tubular and coiled anterior to testes. Pars prostatica long, narrow and curved posterior
to ventral sucker, surrounded by well-developed prostatic gland cells. Aglandular
part of pars prostatica straight, entering sinus-sac anterior to ventral sucker. Sinus-
sac oval, muscular, containing muscular and coiled hermaphroditic duct, opening
just posterior to intestinal bifurcation into shallow genital atrium. Genital cone or
sinus organ absent. Ovary kidney-shaped or transversely elongate. Blind seminal
receptacle present. Vitellarium compact, two separate masses, transversely elongate,
parallel to each other, postovarian. Post-vitelline region of body filled with coils of
uterus. Eggs 30–43 × 10–14.
Remarks: Machida (1980) described this species from Kyphosus cinerascens from
Ki Peninsula, Japan, and Bray & Cribb (2002) reported it from K. sydneyanus from
the Indian Ocean off Western Australia. So far this is the only species reported in the
genus Neopisthadina.
Family Accacoeliidae Odhner, 1911
Accacoeliidae comprises a small group of digeneans easily recognized by the fairly

large highly muscular cylindrical body, well-developed suckers with ventral sucker
usually on a peduncle and presence of an anterior extension to the pharynx which
penetrates the base of pharynx.
The family was first recognized by Odhner (1911) as a subfamily and first used
at the family level by Dollfus (1923). Yamaguti (1975) supported the erection of
superfamily Accacoeloidea by Dollfus (1960) for inclusion of this family. However,
commenting on the terminology, systematic and evolution of the Hemiuroidea, Gib-
son and Bray (1979) have supported the inclusion of the family Accacoeliidae under
Hemiuroidea Looss, 1899. Different classifications were proposed for the family but
the one proposed by Gibson and Bray (1979), with two subfamilies, Accacoeliinae
Odhner, 1911 and Paraccacladiinae Bray & Gibson, 1977 is followed here.
Diagnosis: Body large, cylindrical, smooth, without ecsoma. Oral and ventral suck-
ers well developed, latter usually protuberant. Pharynx with an anterior extension pro-
jecting into oral sucker. Caeca H-shaped, usually with uroproct. Testes two, oblique
or in tandem usually at midbody. Seminal vesicle thin-walled may or may not extend
into hindbody. Pars prostatica tubular, prostatic cells few, sinus-sac and sinus organ
present. Uterine coils occupy entire hindbody. Vitellarium comprised of numerous
filamentous tubules. Excretory vesicle Y-shaped with arms united in forebody. Type
genus: Accacoelium Monticelli, 1893.
Subfamily: Accacoeliinae Odhner, 1911
Diagnosis: Body elongate. Oesophagus long and narrow. Caeca H-shaped, uroproct
present. Sinus-sac and sinus organ present or absent. Vitellarium filamentous with
single main collecting duct on right and single system of ramifying branches in fore
or hindbody.
Family Accacoeliidae Odhner, 1911 183
Genus Tetrochetus Looss, 1911

[Syn. Paratetrochetus Hanson, 1955]
Diagnosis: Body cylindrical with smooth surface. Ventral sucker pedunculate.

Caeca H-shaped. Pars prostatica small, sinus-sac and sinus organ absent. Male and
female ducts open together into shallow genital atrium. Uterine coils extensive. Vitel-
larium tubular, in hindbody, branched. Type species: T. raynerii (Nardo, 1833).
Three species of Tetrochetus Looss have been recorded from Indian marine fishes
(Radhakrishnan & Nair, 1981; Hafeezullah, 1988; Madhavi et al. 1993).
Tetrochetus coryphaenae Yamaguti, 1934 (Fig. 9.38 )
Hosts: Diodontidae: Diodon hystrix L (1, 3); Coryphaenidae: Coryphaena hippurus
L. (2, 3)
Locality: Kerala, AS (1); Tuticorin, BOB (2); VSK, BOB (3)
Distribution: Japan, Mediterranean, Spanish Northeast Atlantic coast, Brazil.
Reference: 1. Radhakrishnan & Nair (1981b); 2. Hafeezullah (1988); 3. Madhavi
et al. (1993)
Description: Body elongate, 2624–13040 long, 256–1152 wide. Tegument thick,

muscular, smooth. Oral sucker 176–480 × 176–432 in size. Ventral sucker pedun-
culate, 208–432 × 208–416. Pharynx oval, extending anteriorly into oral sucker.
Oesophagus long, divides into caeca just in front of ventral sucker. Caeca H-shaped,
Fig. 9.38 Tetrochetus

coryphaenae (Original)
anterior caecal limbs reach almost to level of pharynx. Six characteristic finger-like
projections occur at junction of oesophagus with caeca. Caeca long open into excre-
tory bladder. Testes longitudinally oval, obliquely tandem, situated at mid-hindbody.
Seminal vesicle sinuous, extends well posterior to ventral sucker. Pars prostatica sur-
rounded by prostatic cells. Sinus-sac and sinus organ present. Genital pore sinistral
to pharynx. Ovary post-testicular, spherical. Uterine coils occupy entire hindbody.
Vitellarium comprised of numerous filamentous tubules. Excretory vesicle Y-shaped
with arms united in forebody.
Remarks: This species was first recorded by Yamaguti (1934) from C. hippurus
from off Japan and subsequently from Hawaii (Yamaguti (1970). Nahhas and Cable
(1964) added a new host Diodon hystrix from Curacao and Jamaica. Dyer et al. (1997)
reported its occurrence in the dolphin fish C. hippurus in Puerto Rico. There are three
reports of the species from India. Diodon hystrix from Kerala and Visakhapatnam
(Radhakrishnan and Nair, 1981; Madhavi et al., 1993) and Coryphaena hippurus
from Tuticorin, BOB (Hafeezullah, 1988). Radhakrishnan and Nair (1981) studied
the seasonal occurrence and histopathology of the parasite in D. hystrix. Madhavi
(et al., 1993) reported the larval stage of the parasite from Lepas sp. and commented
on its life cycle pattern.
Tetrochetus aluterae (Hanson, 1955) Yamaguti, 1958 (Fig. 9.39)
[Syn. Paratetrochetus aluterae Hanson, 1955]

Host: Balistidae: Balistes capistratus Shaw; Exocoetidae: Exocoetus volitans Lace-
pede,

aluterae (after Hafeezullah,
1988, Records of Zool.
Survey of India, 85, p 352.)
Family Accacoeliidae Odhner, 1911 185
Locality: Gopalpur and Paradip (Bay of Bengal) and Tuticorin (Gulf of Mannar, Bay
of Bengal)
Description: Length 3.64 mm, width 764. Sucker length ratio 1:1.29–1.48. Anterior
testis 227–354 long, 140–291 in depth; posterior testis 224–372 long, 140–300 in
depth; Ovary 105–242 long, 140–301 in width. Eggs 21–35 × 14–16.
Remarks: Hanson (1955) established the genus Paratetrochetus with P. aluterae
as the type species based on a single specimen collected from Aluterus scriptus.
Yamaguti (1958) reduced Paratetrochetus to synonymy with Tetrochetus. Siddiqi &
Cable (1960) reported T. aluterae from C. hippurus and Acanthocybium solandri
(Cuvier) from Puerto Rico and pointed out that T . coryphaenae and T. aluterae may
be identical. The two species are identical except for a small difference in egg size.
Tetrochetus macrorchis Yamaguti, 1970 (Fig. 9.40)
Host: Diodontidae: Diodon hystrix Linn.,
Locality: Trivandrum (Arabian Sea).
Number of specimens; 2
Description: Body 8.167–11.00 mm long, 677–1.980 mm wide. Ventral sucker
borne on short peduncle, 605–688 long, 567–618 wide. Oral sucker 371 long 398–481
wide. Testes two, antero-dorsal and postero-ventral near body wall, diagonally dis-
posed in middle region of hindbody, 825–1196 long, 618–948 wide. Ovary 495–756
long, 536–577 wide. Mehlis’ gland anterior to ovary. Vitelline reservoir near ovary;
vitellarium tubular, branched, between ovary and base of peduncle. Proximal part
of ascending uterus forms uterine seminal receptacle, it then descends to slightly
short of posterior end of body, crosses transversely and then ascends again, distal
part forming metraterm ventral to pars prostatica. Both the ducts open separately into
small genital atrium. Eggs 21–31 × 14–24.

macrorchis (after
Hafeezullah, 1988, Records
of Zool. Survey of India, 85,
p 353)
Remarks: This species is robust, quite large and broadest of all the species described
so far. The body is greatly elongated with almost parallel or subparallel sides. The
testes are very large, sometimes forming slight bulges in the body wall. With these
characters, this species can be distinguised from the remaining species. It was origi-
nally described by Yamaguti (1970) from the Hawaiian fishes Xanthichthys ringens,
Diodon hystrix and Sphaeroides cutaneus.
Key to species of Tetrochetus reported from Indian marine fishes.

1. Body narrow, slender., elongated; sides almost parallel; testes small as com-
pared to body length and width; ventral sucker larger than oral sucker ..……...
T. coryphaenae Yamaguti, 1934
Body broader, not slender and narrow; testes medium-sized or large ventral sucker
tending to be equal to oral sucker …………………………………………...… 2
2. Body usually attenuated from pregonadal level to base of peduncle; testes
medium-sized; ventral sucker almost equal to oral sucker ………. T. aluterae
Hanson, 1955
Body robust, broader and usually large as compared to body length, ventral sucker
definitely larger than oral sucker ……………………. T. macrorchis Yamaguti,
1970
Family Dictysarcidae Skrjabin & Guschanskaja, 1955
[Syn. Cylindrorchiidae Pohe, 1926; Aerobotrematidae Yamaguti, 1958; Peloro-

helminthidae Fischhthal & Kuntz, 1964; Albulatrematidae Yamaguti, 1965;
Tetrasteridae Oshmarin, 1965; Dollfustravassossidae Teixeira de Freitas & Kohn,
1967]
Diagnosis: Body oval, ecsoma absent. Tegument smooth. Oral and ventral suckers
well developed. Pharynx well developed. Oesophagus short. Caeca terminate blindly
near posterior end of body. Testes two large, oval or elongate, preovarian, posterior
to ventral sucker. Seminal vesicle tubular, in forebody, pars prostatica tubular, sinus-
sac well developed or poorly developed, sinus organ absent. Genital atrium small or
absent. Hermaphroditic duct well developed or small. Ovary oval or with four short
or elongate lobes, separated from testes by loops of uterus. Juel’s organ and uter-
ine seminal receptacle present. Uterus with many loops in preovarian area, mostly
confined to hindbody. Eggs small, numerous. Vitellarium with six to eight oval or
digitiform lobes or two compact multilobate masses, postero or anterolateral or pos-
terior to ovary. Excretory arms united in forebody. Type species: Dictysarca Linton,
1910.
Subfamily Albulatrematinae Yamaguti, 1965
[Syn. Pelorohelminthinae Fischthal & Kuntz, 1964; Tetrasterinae Oshmarin, 1965]

Family Dictysarcidae Skrjabin & Guschanskaja, 1955 187
Genus Elongoparorchis Hanumantha Rao, 1961

[Syn. Pelorohelmins Fischthal & Kuntz, 1964; Tetraster Oshmarin, 1965; Dollfus-
travassossius Teixeira de Freitas & Kohn, 1967]
Elongoparorchis pneumatis Hanumantha Rao, 1961 (Fig. 9.41)
[Syn. E. palawanensis Fischthal & Kuntz, 1964; E. siamensis (Oshmarin, 1965);
Pelorohelmins palawanensis Fischthal & Kuntz, 1964; Tetraster siamensis Osh-
marin, 1965]
Host: Ariidae: Arius jella Day.
Locality: VSK, BOB
Reference: Rao (961)
Description: Body flat, 9200 long 3500 wide. Tegument smooth. Oral sucker 600 in
diameter, Ventral sucker in anterior quarter of body, 770 in diameter. Pharynx opens
into thick-walled glandular stomach. Caeca sinuous, reach posterior end of body.
Testes elongated, symmetrical, arcuate with dilatations, situated postero-lateral to
ventral sucker. Genital pore median, immediately behind oral sucker. Seminal vesi-
cle sinuous, may reach dorsally to ventral sucker. Pars prostatica short, sinus-sac
poorly developed. Sinus organ small, papilla-like. Hermaphroditic duct relatively
long, formed as continuation of metraterm, with ejaculatory duct entering laterally.
Ovary in posterior half of hindbody, ovarian lobes digitiform. Uterus mainly preo-
varian. Eggs small, 13.2 × 6.6 in size. Vitellarium arranged in two groups of three
and four lobulated radiating branches, situated behind ovary.
Remarks: Rao (1961) erected the genus Elongoparorchis with E. pneumatis Rao,
1961 from the air bladder of the marine catfish Arius jella collected from Bay of
Bengal as the type species. The genus was included in the family Isoparorchiidae.
Fig. 9.41 Elongoparorchis

pneumatis (after
Hanumantha Rao, 1960
courtesy Cambridge
University Press, J.
Helminth. 35, p. 303)
Hafeezullah & Siddiqi (1973) redescribed the terminal genitalia of E. pneumatis

and suggested inclusion of Elongoparorchis together with the genera Pelorohelmins
and Dollfustravassossius in the family Pelorohelminthidae Fischthal & Thomas,
1968. Ultimately the genus is transferred to the family Dictysarcidae Skrjabin &
Guschanskaja, 1954, subfamily Albulatrematinae Yamaguti, 1965 (Gibson, 2002).
The genera Pelorohelmins Fischthal & Kuntz, 1964; Tetraster Oshmarin, 1965; Doll-
fustravassossius Teixeira de Freitas & Kohn, 1967 were regarded as synonyms of
Elongoparorchis (Jones & Khalil, 1984). Pelorohelmins palawanensis Fischthal &
Kuntz, 1964 and Tetraster siamensis Oshmarin, 1965 were further regarded as syn-
onyms of E. pneumatis. E. pneumatis thus occurs in the air bladder of catfishes of Bay
of Bengal, Palawan Island and Pacific coast off Siam. Rao and Rao (1970) studied
the seasonal occurrence of the parasite in Arius thalassinus from Bay of Bengal.
Family Didymozoidae Monticelli, 1888
[Syn. Koellikeriidae Ishii, 1935; Didymodiclinidae Pozdnyakov, 1960]

Didymozoids are the most complicated group of digeneans possessing several
unique features quite untypical of the group. Adult didymozoids are tissue para-
sites of marine fishes and usually occur encapsulated in pairs. They exhibit varying
degrees of sexual dimorphism either completely gonochoristic or partially or totally
hermaphroditic. The pattern of the life cycle of this group and hence its relationships
with other digeneans remaines obscure. Tentatively it was included by Gibson & Bray
(1979) under the Hemiuroidea and this was supported by molecular evidence (Barker
et al. 1993). Several classifications have been suggested for this family. Pozdnyakov
& Gibson (2008) provided an overview on the morphology of didymozoids.
The Didymozoids are particularly common parasites of scombrids especially the
tunas of Indo-pacific region. The most extensive studies on didymozoids are those
of Yamaguti (1938, 1970) from the coasts of Japan and Hawaii. Few reports exist
from India (Job, 1961–1968; Madhavi, 1982; Madhavi & Rao, 1983; Murugesh &
Madhavi, 1994, 1995; Murugesh et al. 1992). According to Pozdnyakov & Gibson
(2008), the Didymozoidae is one of the most difficult, probably the most complicated
group of digeneans to deal with.
Diagnosis: Hermaphroditic or gonochoristic forms, found free in tissues or encap-

sulated in pairs. Body divided into two regions or not. Oral sucker present, pharynx
and ventral sucker present or not. Digestive system somewhat reduced. Gland cells
present over pharynx, oesophagus and anterior region of caeca. Testes usually two,
rarely one, of variable shape and location. Terminal genitalia reduced. Genital pore
in forebody, near oral sucker or pharynx. Ovary and vitellarium thread-like, branched
or not. Seminal receptacle uterine or blind sac. Juel’s organ present or absent. Uter-
ine coils extensive. Eggs oval, miracidium with anterior crown of spines. Excretory
vesicle Y-shaped.
Family Didymozoidae Monticelli, 1888 189
Type genus: Didymozoon Taschenberg, 1879

Division of the family into subfamilies is complicated because of the extreme
diversity in the organization, lack of information on their biology and the phyloge-
netic relationships. There is no uniformity in the classifications proposed by different
authors. Yamaguti (1971) recognized 23 subfamilies, most of them including a single
genus. Pozdnyakov (1993) recognized two families under order Didymozoidea: the
family Didymozoidae including all hermaphroditic forms comprising five subfami-
lies and the family Didymodiclinidae Podznyakov, 1993 including all gonochoristic
forms and comprising three subfamilies, each subfamily further divided into tribes.
The same classification with slight modification was followed by Pozdnyakov &
Gibson (2008). They merged the two families into only one the Didymozoidae and
recognized five subfamilies under it: Didymozoinae Monticelli, 1887, Didymodi-
clininae Pozdnyakov, 1993, Gonapodasminae Ishii, 1935, Koellikerinae Ishii, 1935
and Nematobothriinae Ishii 1935. The latter classification is followed here and the
didymozoid genera reported from Indian region fall into four subfamilies. No tribes
are recognized in this account. To date 49 species of didymozoids distributed over
27 genera are reported from India. Arranging them according to any of the earlier
classifications proved difficult. These 27 genera are tentatively grouped here into
four subfamilies, the main purpose is to simplify the identification process: Didy-
mozoinae Moticelli, 1888; Colocyntotrematinae Yamaguti, 1958; Didymodiclininae
Podznyakov, 1993 and Nemaobothriinae Ishii, 1935.
Key to Subfamilies:
1. Hermaphroditic forms, single or encysted inpairs.…………………………….. 2
Completely or incompletely gonochoristic forms ………. Didymodiclininae
Podznyakov, 1993
2. Body long, slender, worm-like usually not encysted………..……. Nematoboth-
riinae Ishii, 1935
Body divided into two regions, a narrow forebody and wider hindbody
….…..………………. 3
3. Hindbody of fellow occupants not fused………………………Didymozoinae
Monticelli, 1888
Posterior region of fellow occupants partially or completely fused
......…………………..................................... Colocyntotrematinae Yamaguti,
1958
Subfamily Didymozoinae Monticelli 1888
[Syn. Adenodidymocystinae Yamaguti, 1979; Neodidymozoinae Yamaguti, 1970]
Diagnosis: Encysted in pairs. Body divided into fore and hind bodies. Forebody
elongate, narrow; hindbody cylindrical, spindle-shaped, disc-like, or spherical with
smooth or lobed surface. Pharynx and oesophagus present. Caeca extend into hind-
body. Testes two in hindbody. Ovary and vitellarium filamentous. Uterus long, coiled.
Type genus: Didymozoon Taschenberg, 1879.
Key to genera
1. Hindbody entire …………………………………………………….….......... 2

Hindbody possessing secondary branches with protrusions and indentations
……….12
2. Posterior region of body cylindrical……………………………..………........ 3
Posterior region of body not cylindrical…………….……………………...... 4
3. Forebody attached to anterior tip of hindbody …….……………Didymozoon
Taschenberg, 1879
Forebody attached to about midlevel of hindbody….…….….. Allodidymozoon
Yamaguti, 1959
4. Posterior region of body flattened on one or both sides, vitellarium radially
branched……………………………………………………………….…….....
Platocystoides Madhavi, 1982
Hindbody not flattened, vitellarium not radially branched
………………….………. 5
5. Hindbody with numerous regularly or irregularly alternating lobes
……………….. 6
Hindbody without lobes …….………………………………………….......... 9
6. Hindbody dichotomously branched, testis single ……………Renodidymocystis
Madhavi, 1982
Hindbody not dichotomously lobed, testes double ………..………………… 7
7. Hindbody with numerous irregular lobes or digitiform protrusions, testes dou-
ble, unbranched ……………………………………………… 8
Hindbody triangular, lobed on convex side only, testes and vitellarium branched
….. ……………………………………………………… Lobatocystis Yam-
aguti, 1965
8. The lobes or protrusion are confined to posterior half of body, ovary and vitel-
larium branched …………………………………………………………….
Kamegaia Yamaguit, 1970
The hindbody covered with digitiform protrusions all over, ovary and vitellarium
each with two branches …………………………………………………………
Metadidymozon Yamaguti, 1970
9. Hindbody long, slender, elongate, forebody attached terminally… Indodidymo-
zoon Madhavi, 1982
Hindbody disc-like, hemispherical or bell-shaped……………………….……
10
10. Hindbody large, disc-like, …………………………………Coeliodidymocystis
Yamaguti, 1970
Hindbody smaller, subglobular or hemispherical ...………………………11
11. Hindbody hemispherical, caeca not filled with black pigment
……………………………… 12
Hindbody disc-like, caeca filled with black pigment …………….. Melanocystis
Yamaguti, 1970
12. Hindbody hemispherical, testes two ………………………………….. Didy-

mocystis Ariola, 1902
Hindbody cylindrical, flattened laterally or tapered posteriorly ……………
………………………………………………….…………. Neometadidymo-
zoon Yamaguti, 1971
Genus Allodidymozoon Yamaguti, 1959
The genus Allodidymozoon was erected by Yamaguti (1959) with A. sphyraenae

from the palate of Sphyraena picuda from the Pacific coast of Japan as the type
species. The genus was differentiated from Didymozoon Taschenerg in two ways:
(1) the position of the genital junction which in Didymozoon is situated at one end
of the hindbody with the entire vitelline filaments lying posterior to it, whereas in
Allodidymozoon the genital junction lies at about the middle of the hindbody and
the vitellarium and ovarian filaments are divided into anterior and posterior groups
at this level. (2) In Didymozoon spp. the vitellarium and ovarian filaments do not
extend into the testicular zone whereas in Allodidymzoon, they extend throughout
the length of hindbody. Additionally, in Didymozoon the forebody arises from the
anterior tip of the hindbody or slightly subterminally, whereas in Allodidymozoon it
arises more posteriorly almost from midlevel of hindbody.
Diagnosis: Encysted in pairs, completely hermaphroditic. Forebody narrow,

attaches to middle region of cylindrical hindbody. Ventral sucker absent. Pharynx
present. Caeca narrow in forebody, distended in hindbody. Testes cylindrical, situ-
ated parallel to one another in anterior region of hindbody. Genital pore at base of
oral sucker. Genital junction at about middle of hindbody. Ovary with two winding
tubes one running anteriorly and the other posteriorly. Vitellarium branched, branches
extending anteriorly and posteriorly, all tubes leading to common tube near genital
junction. Juel’s organ present. Seminal receptacle absent, Eggs small, numerous.
Type species: A. sphyraenae Yamaguti, 1959
Allodidymozoon apharyngi (Job, 1961) Madhavi, 1982
[Syn. Didymozoon apharyngi Job, 1961]

Host: Sphyraenidae: Sphyraena barracuda (Edwards)
Site of infection: Gill arch and Buccal epithelium
Distribution: MS, BOB
Reference: Job (1961a); Madhavi (1982)
Description: Body 5–10 mm, hindbody straight and cylindrical. Oral sucker
100–150, very prominent, Forebody one-fifth the length of hindbody and is attached
to hindbody at about one-sixth of length from anterior end. Ovarian complex located
close to anterior end of body. Uterus with many eggs. Eggs 12 × 5.
Remarks: Didymozoon apharyngi Job, 1961 has the characters of Allodidymozoon

as defined above and accordingly it was transferred to that genus as A. apharyngi
(Job, 1961) Madhavi, 1982. The description given for the species lacks many details
especially information about the male reproductive system, the ovarian and vitelline
branches, their distribution and the digestive system. It is here regarded as a species
inquirendum.
Allodidymozoon cylindricum Madhavi, 1982 (Fig. 9.42)

Host: Sphyraenidae: Sphyraena obtusata Cuvier, S. barracuda (Edwards)
Site: Mucosa of the stomach wall, buccal epithelium and muscles of the maxilla.
Distribution: VSK, BOB
Reference: Madhavi (1982)
Cysts elliptical, occur embedded in stomach wall, each cyst encloses two identical
individuals, the ventral surfaces of which are in contact with each other with their
forebodies lying in between.
Description: Hermaphroditic. Forebody slender, filiform, 1280–2200 long and

80–160 wide, attached to hindbody at about one third of body length from ante-
rior end, point of attachment lies midway between median line and left body mar-
gin. Oral sucker 40–64 by 24–28. Ventral sucker absent. Pharynx 20–22 in diameter.
Oesophagus slender. Caeca narrow in forebody but distended in hindbody. Hindbody
5.0–17.0 mm long and 1440–2200 wide, typically cylindrical in young and immature
forms, spirally twisted with undulations here and there in mature individuals. Testes
cylindrical, sinuous, situated parallel to one another in anterior region of hindbody.
Vas deferens sinuous, enters forebody, opens through genital pore situated at base of
oral sucker. Genital junction at about middle of hindbody. Ovary cylindrical consists
of two winding tubes, one running from anterior end, other from posterior end to
genital junction; occasionally ovary consists of three tubes two directed anteriorly
and one posteriorly. Laurer’s canal terminates in Juel’s organ. Mehlis’ gland well
developed with its cells extending over proximal uterine coils. Vitellarium with two
Fig. 9.42 Allodidymozoon

cylindricum (after Madhavi,
1982, Courtesy Springer,
Syst. Parasit. 4, p. 102)
anterior branches and three posterior ones. Uterine coils occupy all available space
in hindbody. Eggs 10 × 6.
Allodidymozoon operculare Madhavi, 1982 (Fig. 9.43)

Host: Sphyraenidae: Sphyraena obtusata Cuvier; S. barracuda (Edwards)
Site: Muscles on inner surface of operculum
Locality: Distribution: Kuwait Bay; VSK, MS, BOB
Reference: Abdul-Salam & Sreelatha (1993); Madhavi (1982)
Description: Forebody filiform, 800–960 long, 80–112 wide; attached to ventral

surface of hindbody subterminally either medially or sub-medially. Hindbody flat
ventrally, convex dorsally, 1520–2160 long 860–1216 wide. Oral sucker 47 by 19
in size. Ventral sucker absent. Pharynx present. Oesophagus long. Caeca narrow
in forebody, inflated in hindbody. Ventral sucker absent. Testes paired, club-shaped,
situated parallel to one another in anterior part of hindbody. Genital complex situated
either at midbody or at slightly more posterior level. Number of ovarian filaments
variable, usually single, unbranched, highly convoluted, commencing at anterior
extremity, winding throughout length of hindbody terminating near genital complex,
rarely ovary with two ovarian filaments, joining near genital complex. Vitellarium
usually with two long and one short branches, one of which winds from the anterior
extremity to the genital complex, the other from the posterior extremity to the genital
complex; short vitelline branch arising either independently or as branch of the
longer vitelline tube. Seminal receptacle oval. Uterine reservoir very conspicuous.
Metraterm well differentiated in forebody appearing as thick-walled, winding tube.
Eggs small (10 × 5).
Remarks: This species differs from the other two species by the characteristic shape
of the body and in the number and distribution of vitelline and ovarian branches.
Fig. 9.43 Allodidymozoon

operculare (after Madhavi,
1982 Courtesy, Syst. Parasit.
4, p. 105)
Key to species of Allodidymozoon:

1. Pharynx absent ……………..……………………………..……A. apharyngi
(Job, 1961)
Pharynx present ………………………………………………………….……2
2. Body bilobed, vitellarium with 2 main branches, one anterior and one poste-
rior and a short branch …………………………….………………. A. opercu-
lare Madhavi, 1982
Body cylindrical, vitellarium with 5 branches, 2 anterior and 3 posterior
………………………….……………...................... A. cylindricum Madhavi,
1982
Genus Coeliodidymocystis Yamaguti, 1970
The genus is charactersied by the habitat, the body shape and the distribution of
male and female reproductive organs. Coeliodidymocystis Yamaguti, 1970 was syn-
onymized with Didymocystis Ariola, 1902 by Pozdnyakov (1990) but here it is
retained as valid. The two genera differ clearly in the body shape and the number
and distribution of ovarian and vitelline branches.
Diagnosis: Complete hermaphrodite, encysted in pairs in thin-walled sac. Body

divided into slender forebody and large oval hindbody, flattened on side on which two
cyst occupants are pressed. Oral sucker globular. Pharynx present or absent. Caeca
narrow and parallel in forebody, divergent and arcuate in hindbody. Testes long, tubu-
lar, irregularly winding, continuous end to end, extend along boundary between flat
and convex sides. Ovary tubular, confined to anterodorsal region of hindbody. Sem-
inal receptacle present. Vitellarium tubular, irregularly branched, extending along
convex side of hindbody, ends in large number of tubules. Uterus occupies all avail-
able space of hindbody. Uterine reservoir long, wide, metraterm strongly developed.
Type species: C. kamegaii Yamaguti, 1970
Coeliodidymocystis kamegaii Yamaguti, 1970 (Fig. 9.44)

Host: Scombridae: Katsuwonus pelamis (L.)
Fig. 9.44 Coeliodidymocystis

kamegaii (after Madhavi,
1982 Courtesy Springer,
Site: Serosa of pyloric caeca

Distribution: Hawaii; VSK, BOB
Reference: Yamaguti (1970); Madhavi (1982)
Description: Cysts subglobular up to about 20 mm in diameter. Forebody flat, cylin-
drical, 5.5–6.0 mm long, attached to flat surface of hindbody opposite genital junc-
tion. Hindbody rounded, 20–21.5 × 15–17.5 mm, flat on ventral side, delimited from
convex dorsal side by circular line. Oral sucker 23–25 in diameter. Pharynx absent.
Caeca narrow in forebody, wide, sinuous in hindbody. Testes two, tubular, irregularly
winding contiguous end to end, Vas deferens runs along median field of forebody.
Ovary tubular, consists of a long trunk dividing dichotomously several times. Seminal
receptacle large, divided into two parts connected by a narrow duct. Uterus winding,
initial part surrounded by gland cells, rest of uterus extends through out convex side
of hindbody. Uterine reservoir large. Metraterm sigmoid, thick-walled. Eggs 14–17
by 10–12. Vitellarium trunk bifurcates several times, giving rise to several branches
Remarks: Coeliodidymocystis kamegaii was first described by Yamaguti (1970)
from Katsuwonus pelamis from Pacific coast off Hawaii. Madhavi (1982) reported
it from the same host from the Bay of Bengal off Visakhapatnam coast. It differs
from the type and only other species C. abdominalis Yamaguti, 1970 in the complete
absence of a pharynx and the shorter oesophagus.
Genus Didymocystis Ariola, 1902
[Syns. Didymocystoides Yamaguti, 1970; Univitellodidymocystis Yamaguti, 1970;
Lagenocystis Yamaguti, 1970; Oesophagocystis Yamaguti, 1970; Didymosulcus
Pozdnyakov, 1990]
Diagnosis: Complete hermaphrodites, enclosed in pairs in round, oval or elliptical
cysts. Forebody scoop-shaped, subcylindrical. Hindbody round, oval, hemispherical
or reniform, with or without ventral furrow. Forebody attached to hindbody close to
its anterior end. Pharynx present. Caeca reaching posterior extremity of hindbody.
Ventral sucker absent. Testes tubular, paired, in hindbody close to base of fore-
body. Genital pore near oral sucker or pharynx. Ovary slender, long, with or without
branches. Vitellarium tubular, may or may not extend throughout hindbody. Uterus
occupies all available space in hindbody. Egg reservoir may or may not be present.
Parasitic on gills, operculum, oral cavity, oesophagus, intestine, body cavity, nasal
cavity, orbit, etc. of marine fishes. Type species: Didymocystis thynni (Taschenberg,
1879) Ariola, 1902.
Several didymozoid genera have been synonymized with Didymocystis.
Podznyakov (1990) in a revision of the genus, synonymized the genera Coelio-
didymocystis Yamaguti, 1970, Didymocystoides Yamaguti, 1970, Univitellodidy-
mocystis Yamaguti, 1970 with Didymocystis. Murugesh & Madhavi (1995) further
synonymized Lagenocystis Yamaguti, 1970, Oesophagocystis Yamaguti, 1970 and
Didymosulcus Podznyakov, 1990 with Didymocystis but retained Coeliodidymocys-
tis Yamaguti, 1970 as a valid genus. Fifteen species of Didymocystis were recorded
from marine fish of Indian region, most of them from scombrid fishes.
Didymocystis alalongae Yamaguti, 1938 (Fig. 9.45)
[Syn. Didymocystoides alalongae (Yamaguti, 1938) Yamaguti, 1970, D. bifasciatus

Yamaguti, 1970; Didymocystis bifasciatus (Yamaguti, 1970) Pozdnyakov, 1990]
Host: Scombridae: Thunnus tonggol (Bleeker)
Site: Operculum
Distribution: Japan, Hawaii, VSK, BOB
Reference: Murugesh & Madhavi (1995)
Distribution: Pacific Ocean (Japan, Hawaii); Bay of Bengal.
Description: Cysts flat, yellow, each cyst encloses two hermaphroditic worms of
equal size and shape. Forebody elongate, hindbody rounded. Oral sucker termi-
nal, 40–52 × 28–34. Pharynx present. Oesophagus narrow. Caeca enter hindbody.
Hindbody 3250–4280 × 2420–2800. Testes two, tubular, situated in anterior part of
hindbody, close to base of forebody. Ovary consists of short common stem which
bifurcates into 2 long winding branches. Ovarian branches restricted to peripheral
region of hindbody. Genital junction pre-equatorial, almost at base of forebody.
Vitelline branches 3–7. Uterus strongly convoluted and occupies most of hindbody.
No uterine or egg reservoir. Eggs boat-shaped 18–20 × 10–12.
Remarks: Didymocystis alalongae Yamaguti, 1938 was reported from the gills of
Thunnus tonggol from the Pacific Ocean (Yamaguti, 1938). Murugesh & Madhavi
(1995) reported the species from the same host from Bay of Bengal and considered
D. bifasciatus (Yamaguti, 1970) from Neothunnus macropterus and Parathunnus sibi
off Hawaii as a synonym of this species. The Bay of Bengal and Pacific forms appear
similar in all respects except for the presence of a uterine reservoir in the latter.
Fig. 9.45 Didymocystis

alalongae (after Murugesh &
Madhavi, 1995 Courtesy
Springer, Syst. Parasit. 31,
p. 15)
Didymocystis bifasciata (Yamaguti, 1970) Pozdnyakov, 1990

[Syn. Didymocystoides bifasciatus Yamaguti, 1970]
Host: Carangidae: Scomberoides tala Cuvier
Site: Opercular muscles
Locality: VSK, BOB
Reference: Hussain et al. (1985)
Remarks: This species was synonymized with D. alalongae by Murugesh & Mad-
havi (1982).
Didymocystis bifurcata Yamaguti, 1970 (Fig. 9.46)

[Syn. Didymosulcus bifurcata (Yamaguti, 1970) Pozdnyakov, 1990]
Site: Mouth cavity and inner gill opening
Locality: VSK, BOB
Distribution: Pacific Ocean (Japan, Hawaii) and Bay of Bengal
Distribution: Japan, Hawaii, Bay of Bengal.
Description: Cyst globular, bright yellow, 2300–3000 in diameter, encloses two

hermaphroditic individuals of equal size. Forebody slender attached to one side of
hindbody. Oral sucker terminal, somewhat rounded, 24–28 × 12–14, followed by
pharynx of similar size. Hindbody ovate, convex on outer margin, flattened on inner
margin, 2104–2280 × 2144–2800. Testes two, elongate, at base of forebody. Ovary
with 4–6 terminal branches. Vitellarium consists of several branches, 2–3 in young
specimens and 4–6 in gravid worms. Uterus occupies all available area of hindbody,
ballooning out into large uterine reservoir in middle of hindbody: metraterm present.
Eggs bean-shaped, 15–16 × 9–10.

bifurcata (after Murugesh &
p. 17)
Remarks: More than half a dozen species of the genus Didymocystis have been
reported from Thunnus sp., mainly from the gills, operculum and skin, from differ-
ent regions. Yamaguti (1970) reported D. bifurcata from the inner gill opening of
Parathunnus sibi from Hawaii. This species is distinguished from other species of
the genus by possessing the combination of characters: hindbody convex dorsally,
flat ventrally, fairly large pharynx, voluminous uterine reservoir.
Didymocystis dissimilis Yamaguti, 1938
Syn. Oesophagocystis dissimilis (Yamaguti, 1938) Yamaguti, 1970
Host: Scombridae: Katsuwonus pelamis (Linn.)
Site: Inner surface of oesophagus.
Locality: Japan, VSK, BOB
No: 18
Distribution: Hawaii, Bay of Bengal.
Reference: Yamaguti (1938); Murugesh & Madhavi (1995)
Description: Flukes encysted in pairs with ventral surfaces pressed together. Fore-
body thick, 460–680 × 72–80, blunt anteriorly, arises from anterior flat surface of
hindbody. Oral sucker16–22 × 12–18, followed by weakly muscular pharynx of sim-
ilar size. Oesophagus bifurcates into 2 slender caeca within forebody. Hindbody
cylindrical, roughly quadrangular, 1648–2080 long. Testes 2, tubular, in anterior part
of hindbody, situated along margins in most specimens. Ovary tubular, 24–32 thick,
consists of short common stem and 2 main branches with 4 terminal branches. Juel’s
organ sac-like. Genital complex situated to one side of hindbody, pre-equatorial.
Vitellarium consists of long common stem that divides into 6–10 terminal branches
of unequal length. Uterus occupies all available space in hindbody. Eggs 12–14 ×
8–10. Egg reservoir absent.
Remarks: Yamaguti (1938) described this species from the oesophagus and stomach
of Euthynnus pelamys from Hawaii and later transferred it to the genus Oesophago-
cystis Yamaguti, 1970. Oesophagocystis is now considered a synonym of Didymo-
cystis. D. dissimilis is thus relegated to its original genus, Didymocystis. D. dissimilis
is characterized by the habitat (inner wall of oesophagus), body shape, tubular ovary
confined to the marginal area and vitelline tubules forming a basket enclosing the
uterus.
Didymocystis exigua (Yamaguti, 1970) Pozdnyakov, 1990 (Fig. 9.47)
[Syn. Didymocystoides exiguus Yamaguti, 1970; D. xishaensis Shen Jiwei, 1990]
Host: Scombridae: Auxis thazard (Lacépède), Euthynnus affinis (Cantor)
Site: Intestinal wall
Distribution: Pacific Ocean, Bay of Bengal, China
Reference: Murugesh & Madhavi (1995); Madhavi & Sairam (2000)
Description: Cysts round, deeply embedded in host tissue making it extremely dif-
ficult to separate individuals or cysts from intestinal tissue. Forebody ovoid, 210–280
long. Hindbody globular, 320–800 × 410–912, with flat ventral surface. Testes tubu-
lar, situated along anterior margin of ovary with short common stem, bifurcates

exigua (after Yamaguti,
1970)
into 2 tubular terminal branches located peripherally. Retort-shaped Juel’s organ

present. Vitellarium with very long, undulating common stem, bifurcates into 2
sinuous branches of same thickness; gravid specimens have 3–4 winding termi-
nal branches directed in different directions. Uterus occupies all available space in
hindbody. Eggs embryonated, bean-shaped, 16–20 × 8–10.
Remarks: This species was first described as Didymocystoides exiguus by Yam-

aguti (1970), from the intestinal wall of Euthynnus yaito from Hawaii. Murugesh &
Madhavi (1995) reported the species from the same host and considered Didymo-
cystoides xishaensis Shen, 1990 from the inner wall of the intestine of Auxis thazard
from Chinese waters as a synonym of the species.
Didymocystis guttata Murugesh & Madhavi, 1995 (Fig. 9.48)
Host: Scombridae: Scombeomorus guttatus (Bl. & Schn.)
Site: Junction of maxilla and mandible
Distribution: VSK, BOB

guttata (after Murugesh &
p. 22)
Description+: Forebody scoop-shaped, attached to hindbody on its fiat ventral

surface, 640–1644 × 96–304. Hindbody hemispherical, flattened on ventral side;
1536–4080 × 640–1552. Oral sucker elongate, muscular. Pharynx small. Oesoph-
agus short, bifurcating into 2 broad caeca. Testes 2, tubular, in anterior part of hind-
body. Ovary consisting of 2 stems, one very long and other short and stumpy. Initial
uterine duct thin-walled, provided with dense coat of gland cells for short distance.
Juel’s organ small, retort-shaped. Vitellarium consists of one short curved stem and
6 branches, distributed in anterior part of hindbody. Uterus coiled regularly occu-
pying most of central region of hindbody. Egg reservoir absent. Eggs bean-shaped,
20–24 × 12–14, numerous.
Remarks: This species differs from other species of Didymocystis in many ways,
e.g. the site, body shape, shape of oral sucker and nature of genital junction and
vitellarium.
Didymocystis oesophagicola (Yamaguti, 1970) Pozdnyakov, l990

Syn. Didymocystoides oesophagicola Yamaguti,1970.
Site: Outer wall of oesophagus.
Locality: VSK, BOB.
Distribution: Hawaii, BOB
Description: Each cyst contains 2 hermaphroditic individuals with flattened, broad

forebody and roughly oval hindbody. Forebody 1680–2980 long by 180–265 wide
and bluntly conical at apex, filled with numerous gland cells, especially anteriorly.
Oral sucker small, triangular, 36–44 × 28–32. Pharynx globular. Oesophagus straight,
surrounded by thick zone of gland cells; initial portion of intestine enlarged to form
typical ‘stomach’. Testes 2, tubular, arranged symmetrically in anterior hindbody.
Ovary tubular, winding, divides into 2 main branches, one of which re-divides to
form total of 3 branches distributed laterally in hindbody. Juel’s organ sac-like, very
close to anterior margin of hindbody. Vitellarium divides anteriorly into 6–10 slim
tubules which trace winding course throughout most of mid-hindbody and extend
to posterior tip. Uterine coils convoluted, occupy most available space in hindbody,
balloon out distally to form large, median, uterine reservoir. Eggs 16–18 × 8–10.
Remarks: Didymocystis oesophagicola was first described as Didymocystoides

oesophagicola by Yamaguti (1970) from cysts in the outer wall of the oesopha-
gus of Neothunnus macropterus from Hawaii. Although this species resembles D.
buccalis Yamaguti, 1970 in the branching of the ovary, it differs from the latter in the
more profuse development of the vitellarium, the longer oesophagus surrounded by
gland cells and the stomach-like swelling of the gut close to the intestinal bifurcation.
Didymocystis pinnicola (Yamaguti, 1970) Pozdnyakov, 1990 (Fig. 9.49)
Syn. Didymocystoides pinnicola Yamaguti, 1970.
Host: Scombridae: Katsuwonus pelamis (Linnaeus)

pinnicola (after Murugesh &
p. 20)
Site: Encysted in pairs on dorsal fin membrane.

No. 27
Description: Each bright yellow cyst contains 2 hermaphroditic individuals. Fore-

body subcylindrical, 560–1408 × 85–128. Hindbody flat anteriorly, almost convex
posteriorly, 1450–2600 × 1200–2512. Oral sucker subterminal, cellular, 21–36 ×
28–44. Oesophagus narrow, winding. Caeca thin in forebody but inflated in hindbody
appearing as 2 club-shaped overlapping masses. Testes tubular, winding, situated in
anterior portion of hindbody, sometimes recurved. Ovary tubular, slender, common
ovarian stem runs anteriorly from genital junction and divides twice to form 4 ter-
minal branches of unequal lengths. Vitelline tubules arise from central mass at level
of genital junction to branch into 8–10 terminal branches of similar thickness to
ovary. Juel’s organ oval. Genital junction close to lateral margin of hindbody, post-
equatorial in most mature specimens. Uterus winding throughout most of hindbody,
finally leading into large cylindrical egg reservoir close to ventral side of hindbody
where it tapers to form muscular metraterm which enters forebody as thick tube,
filled with numerous small, oval eggs measuring 14–16 × 10.
Remarks: This species was described as Didymocystoides pinnicola by Yamaguti

(1970) from the dorsal fin membrane of Katsuwonus pelamis off Hawaii. It is char-
acterized by the location, occurring only in the dorsal fin of the host. In addition, the
genital junction is well apart from the base of the forebody.
Didymocystis pseudobranchialis Job, 1964
Host: Sphyraenidae: Sphyraena barracuda (Edwards)
Site: Pseudo branch on the roof of buccal cavity.
Locality: GOM, MS, BOB
Reference: Job (1964)
Description: Hindbody broad at base, narrow anteriorly, dorsal side convex. Fore-
body arises from, narrow end of hindbody at one third of length of hindbody. Fore-
body is usually one third of length of hindbody. Oral sucker, pharynx present. Genital
pore at base of pharynx. Testes appear as paired tubular structures. Genital complex
situated within hindbody close to point of origin of hindbody. Genital pore at base
of oral sucker. Ovary long, coiled extends freely into testicular region. There is no
uterine reservoir. Vitellarium is long and branched.
Remarks: D. pseudobranchialis differs from other species of the genus in the general
structure of fore and hindbodies, the long and coiled ovary extending into testicular
zone, and site specificity being confined to the pseudobranch on the roof of buccal
cavity.
Didymocystis semiglobularis Ishii, 1935
Syn. Didymocystoides semiglobularis (Ishii, 935) Yamaguti, 1970
Host: Scombridae: Scomberomorus guttatus (Bl. & Schn.)
Site: Gill rakers
Locality: VSP, BOB
Distribution: Japanese Pacific and BOB
Description: Cyst globular, encloses two hermaphroditic individuals. Forebody
elongated, thread-like, arises from flattened end of semi globular hindbody. Oral
sucker elliptical. Pharynx muscular, rounded. Oesophagus long. Caeca narrow in
forebody, terminate near posterior end of hindbody. Testes paired, unbranched,
arranged one on each side of anterior part of hindbody. Genital opening at base of
oral sucker. Ovary tubular, elongated with slender branches, situated close to lateral
margins of hind body. Seminal receptacle oval. Vitelline tubules slender, branched,
situated in the median field. Uterine coils occupy entire hindbody, form voluminous
uterine reservoir. Eggs 15 × 9.
Remarks: Ishii (1935) described this species as Didymocystis semiglobularis Ishii,
1935 characterizing it with a semi-globular hindbody and slender forebody. Yamaguti
(1970) transferred it to the genus Didymocystoides as D. semiglobularis (Ishii, 1935)
Yamaguti, 1970. Murugesh & Madhavi (1995) restored it to the genus Didymocystis
as D. semiglobularis.
Didymocystis singularis Job, 1966
Host: Sphyraenidae: Sphyraena jello Cuvier (1); S. obtusata Cuvier (2)
Site: gills
Locality: Pamban, GOM (1); VSK (2), MS (3), BOB
Reference: 1. Job (1966); 2. Hussain et al. (1985a);
Remarks: This species was described by Job (1966) from Sphyraena jello from the
Sea of Pamban, Gulf of Mannar and later Hussain et al., (1985a) reported it from S.
obtusata off the Visakhapatnam coast, Bay of Bengal. This species is characterized
by the short, elongate, narrow forebody which is swollen near the tip and the presence
of a voluminous uterine reservoir.

sphyraenae (after Hussain
et al. 1985)
Didymocystis sphyraenae Hussain, Rao & Shyamasundari, 1985 (Fig. 9.50)

Site: Stomach wall
Locality: VSK, BOB
Reference: Hussain, Rao & Shyamasundari (1985)
Description: Forebody slender, hindbody plump, forebody arises just anterior to

middle of hindbody, 1172 × 82 in size. Hindbody broader at posterior end, narrow at
opposite side, length 5104 × 1056. Oral sucker, pharynx present. Oesophagus narrow,
caeca reach posterior extremity. Testes two, tubular, spirally twisted, situated in
median field of anterior part of hindbody. Ovary double branched, strongly winding,
in posterior region of hindbody, enters into testicular zone. Vitellarium strongly
winding, in posterior region of hindbody some branches even extend into anterior
end. Most of the space in hindbody filled with uterine coils. Genital junction in
middle of hindbody. Eggs 9 × 6. Egg reservoir absent.
Remarks: This species is characterised by the spirally twisted testes, plump shape
of hindbody, highly coiled ovary and vitellarium and absence of uterine reservoir.
Didymocystis superpalati Yamaguti, 1970 (Fig. 9.51)

Syn. Didymosulcus superpalati (Yamaguti, 1970) Pozdnyakov, 1990
Site: Palatal teeth.
Distribution: Hawaiian Pacific, Indian Ocean, Bay of Bengal.
Description: Each bright yellow, hard cyst is rounded with rough, sclerotised cyst
wall. Two hermaphroditic individuals of equal size occur in each cyst. Body dif-
ferentiated into cylindrical forebody and oval hindbody; former usually contracted
into anterior part of latter. Forebody 320–980 × 150–272, with terminal, cellular.
Oral sucker followed immediately by weakly muscular pharynx of same size. Hind-
body 3200–5100 × 1720–2060 with distinct ventral furrow. Testes sausage-shaped,
arranged symmetrically along anterior margin of hindbody. Ovary branched, with

superpalati (after Yamaguti,
1970)
2 main branches and 3–4 side branches of unequal length. Mehlis’ gland and geni-
tal complex pre-equatorial. Juel’s organ oval. Vitellarium with 2 main trunks which
branch 2–3 times in gravid worms to form numerous terminal branches that reach as
far as posterior extremity of hindbody. Uterus occupies all available space in hind-
body; large, comma-shaped uterine reservoir present, leads to muscular, winding
metraterm full of small, bean-shaped eggs, 14–18 × 8–10.
Remarks: Didymocystis superpalati was first described by Yamaguti (1970) from

cysts in teeth of the palate of Neothunnus macropterus. It was characterized by the
main branch of the ovary which bifurcates three times and the vitelline gland which
bifurcates at least twice. D. palati Yamaguti, 1970, also described from the palate of
N. macropterus, is the closest relative of this species, but has a simple branching of
both the ovary and vitellarium. Nikolaeva & Dubina (1985) collected D. superpalati
from Thunnus sp. from the Indian Ocean.
Didymocystis triangularis Hussain, Rao & Shyamasundari, 1985

Site: Gills
Locality: VSK, BOB
Description: Body divided into wide hindbody and slender forebody arising at the
anterior end of hindbody. Hindbody triangular, 4319 long 2656 wide. Forebody
slender 1923 long 112 wide, pointed at tip. Oral sucker rounded. Pharynx small.
Oesophagus narrow. Caeca narrow in forebody, not differentiated in larger hindbody.
Testes club-shaped, slightly winding, extend from hindbody to a distance of 992
posterior to origin of forebody. Vas deferens enters forebody, passes to genital pore
situated close to oral sucker. Ovary slender, tubular, consists of two long winding
tubes extend to anterior half of hindbody. Vitellarium with several tubular branches
extending to other half of hindbody, sometimes overlapping ovarian tubules. Genital
junction with thick shell gland cells, close to origin of forebody. Rest of hindbody
occupied by uterus extending from end to end, highly coiled and forming median
terminal swelling before entering forebody. Eggs 9 × 6.
Remarks: This species closely resembles D. acanthocybii Yamaguti, 1938 and D.

ovata Ishii, 1935 but differs from them in the triangular shape of body, testes slightly
winding and situated near the margin of hindbody.
Didymocystis visakhapatnamensis Hussain, Rao & Shyamasundari, 1985
Site: Stomach wall
Locality: VSK, BOB
Reference: Hussain et al., (1985)
Description: Body divided into fore and hindbodies. Forebody slender and narrow,
1236 long, 132 wide, arises from middle of hindbody. Hindbody inverted V-shaped
2056 × 1568 in size. Oral sucker small followed small round pharynx. Oesophagus
narrow. Caeca narrow in forebody but inflated in hindbody. Testes 2, elongated, club-
shaped, situated at lateral sides of right arm of hind body. Genital pore close to oral
sucker. Ovary and vitellarium strongly winding tubules, slender, extend throughout
hindbody. Ovarian tubules narrow, vitelline tubules broad and are placed opposite
to one another in each lobe, tubules from right and left side meet at junction of
hindbody and forebody where they form the ootype. Bulk of hindbody occupied by
uterus which is highly coiled and strongly convoluted, forms uterine reservoir before
entering forebody. Eggs 9 × 6.
Remarks: It is distinguished from the known members of the genus by the body
shape (invertible V-shaped with left and right arms and round base), in the lateral
arrangement of testes in the anterior half of body, ovary and vitellaria arrangement
opposite to one another.
Didymocystis wedli Ariola, 1902 (Fig. 9.52)

Syns: Didymozoon sp. of Kobayashi, 1921; Didymocystis kobayashii Dollfus, 1926;
Wedlia katsuwonicola Okada, 1926.
Host: Scombridae: Auxis thazard (Lacepede)
Site: Buccal epithelium
Number: 8
Locality: VSK, BOB
Distribution: Japan, Naples and Genoa (Kobayashi, 1921; Dollfus, 1926; Okada,
1926).
Description: Cyst thin-walled encloses two identical worms applied closely against
each other with opposite extremities. Forebody slender, hindbody curved ventrad

wedli (after Madhavi, 1982
Courtesy Springer, Syst.
Parasit. 4, p. 107)
with distinct longitudinal median groove on ventral and anterodorsal side, tapers
to form a pointed tail. Oral sucker pear-shaped, followed by pharynx and slender
oesophagus. Intestinal bifurcation at middle of forebody. Caeca terminate at poste-
rior end of hindbody. Testes 2, sausage-shaped, lie dorsoventrally along the anterior
edge of hindbody. Ovary consists of two long slender tubes extending antero ven-
trally to terminate posterior to testes. Mehlis’ gland complex is situated dorsally at
junction of bilobed anterior and spiral posterior part. Vitellarium composed of paired
anterolateral and postero-lateral stems which give off branches that extend antero-
laterally and postero-laterally. Uterus forms a large spindle-shaped reservoir before
entering forebody. Eggs 18 × 10 in size.
Remarks: The most distinctive characteristic feature of the species is the peculiar
shape of the hind-body which has a bilobed anterior end and spirally twisted at
posterior end.
The various species of Didymocystis show host and site specificity. Of the 15
species of the genus, 9 spp. have been recorded from scombrid fishes, 5 spp. from
Sphyraena spp. and one species from carangids. D. bifasciata (Yamaguti, 1970)
recorded from a carangid fish was synonymized with D. alalongae Yamaguti (1938).
The remaining 14 species differ in the shape of hindbody, the number of branches in
the ovary and the vitellarium and the presence or absence of a uterine reservoir. The
following key is prepared based on these differences.
Key to Indian species of the genus Didymocystis
Species recorded from scombid fishes …………………………….....………… 1
Species recorded from sphyraenid fishes ………………………….....………… 6
1. Hindbody with a deep furrow……………………............................………...... 2

Hindbody without a furrow …………………..……..........................……........ 3
2. Parasitic on operculum and mandible, hindbody bilobed anteriorly
and spirally twisted posteriorly, ovary with 2 main branches, vitellar-
ium with 7–8 posteriorly directed branches, uterine reservoir present
…………………………………......D. wedli Ariola, 102
Parasitic on palatal teeth, hindbody foliate, ovary with 2 main branches and 3–4
side branches of unequal length, vitellarium with numerous irregular branches,
uterine reservoir comma-shaped……………..……………….............……......
D. superpalati Yamaguti, 1970
Parasitic in mouth cavity and inside of gills, hindbody ovate, ovary with back-
wardly directed main branches, vitellarium with 4–6 marginal branches, uter-
ine reservoir present..……………………………………D. bifurcata Yamaguti,
1970
3. Hindbody quadrangular, roughly semicircular or hemispherical …………….. 4
Hindbody round, oval or globular ….……..............................................…….. 5
4. Parasitic on inner surface of operculum, hindbody quadrangular, ovary with 4
terminal branches, vitellarium with 8–10 marginal branches, uterine reservoir
present …………………………………..……… D. dissimilis (Yamaguti, 1938)
Parasitic in the dorsal fin membrane, hindbody roughly triangular
or quadrangular, ovary with 4 terminal branches, vitellarium with
8–10 terminal branches, large cylindrical uterine reservoir present
…………………….....................................…. D. pinnicola (Yamaguti, 1970)
Parasitic at the junction of maxilla and mandible, hindbody hemispherical,
ovary with two unequal branches, vitellarium with 6 marginal branches, uter-
ine reservoir absent ………………………………..………….……… D. guttata
Murugesh& Madhavi, 1995
Parasitic on gill rakers, hindbody semiglobular, ovarian tubules with slen-
der branches, vitelline tubules branched, uterine reservoir voluminous
…...................... D. semiglobularis Ishii, 1935
5. Parasitic in operculum, hindbody rounded, ovary with short common stem and
2 winding branches, vitellarium with 5–7 terminal branches, uterine reservoir
absent ……………….……………..…D. alalongae Yamaguti, 1938
Parasitic on outer wall of oesophagus, hindbody oval, ovary bifurcate with one
arm profusely branched, vitellarium with 6–8 terminal branches, uterine reservoir
present ……………………………………..……………....…..… D. oesophagi-
cola (Yamaguti, 1970)
Parasitic in the intestinal mucosa, hindbody spherical, ovary bifurcate, vitellarium
with 2–3 terminal branches, uterine reservoir absent.……..……....… D. exigua
(Yamaguti, 1970)
6. Parasitic on pseudobranch, ovary long and coiled extends into testicular zone
…………………………………………………………… D. pseudobranchialis
Job, 1964
Parasitic on gills, testes winding situated near margins of hind-
body…………………….. ………………………..................... D. triangularis
Hussain, Rao & Shyamasundari, 1985
Parasitic on stomach wall, body plump, ovary and vitellaria highly coiled
…………… …………………………………. D. sphyraenae Hussain, Rao &
Shyamasundari, 1985
Table 9.5 Indian Species of Didymocystis, their hosts and geographic distribution
Species of Host Geographic References
Didymocystis distribution
D. alalongae Thunnus tonggol Japan, Hawaii, BOB Murugesh & Madhavi
Yamaguti, 1938 (1995)
D. bifurcata Thunnus tonggol Japan, Hawaii, BOB Murugesh & Madhavi
Yamaguti, 1970 (1995)
D. dissimilis Katsuwonus pelamis Hawaii, BOB Murugesh & Madhavi
Yamaguti, 1938 (1995)
D. exigua (Yamaguti, Auxis thazard, Hawaii, China, BOB Murugesh & Madhavi
1970) Euthynnus affinis (1995)
D. guttata Murugesh Scomberomorus BOB Murugesh & Madhavi
& Madhavi, 1995 guttatus (1995)
D. oesophagicola Thunnus tonggol Hawaii, BOB Murugesh & Madhavi
(Yamaguti, 1970) (1995)
D. pinnicola Katsuwonus pelamis Hawaii, BOB Murugesh & Madhavi
(Yamaguti, 1970) (1995)
D. pseudobranchialis Sphyraena barracuda BOB Job (1964)
Job, 1964
D. semiglobularis Scomberomorus Japan, BOB Murugesh & Madhavi
Ishii, 1935 guttatum (1995)
D. singularis Job, Sphyraena jello, S. AS, BOB Job (1966); Hussain
1966 obtusata et al. (1985)
D. superpalati Thunnus tonggol Hawaii, BOB Murugesh & Madhavi
Yamaguti, 1970 (1995)
D. triangularis Sphyraena obtusata BOB Hussain et al. (1985)
Hussain, Rao,
Shyamasundari, 1985
D. Sphyraena obtusata BOB Hussain et al. (1985)
visakhapatnamensis
Hussain, Rao,
Shyamasundari, 1985
D. wedli Ariola, 1902 Auxis thazard Japan, Naples„ Genoa, Murugesh & Madhavi
BOB. (1995)
Parasitic on stomach wall, body inverted V- shaped

………………………………………… ………………………… D. visakhap-
atnamensis Hussain, Rao & Shyamasundari, 1985
Parasitic on gills, forebody short, narrow, elongate, uterine reservoir volu-
minous.…………………………………………………….. D. singularis Job,
1966 (Table 9.5)
Genus Didymozoon Taschenberg, 1879
Diagnosis: Capsule cylindrical or spherical. Forebody long, attached terminally to

hindbody. Hindbody cylindrical, sometimes spiral or with indentations of secondary
orign. Pharynx present. Testes elongate, parallel or slightly interwoven, in anterior
third of hindbody. Genital pore ventral to oral sucker. Genital junction in first or
second third of posterior region. Seminal receptacle present. Ovary and vitellarium
elongate, usually unbranched, rarely with two branches.
Didymozoon diverticulatum Yamaguti, 1970

Host: Scombridae: Euthynnus affinis (Cantor)
Site: Encysted in pairs in subserosa of pyloric caeca
Locality: VSK, BOB
Description: Cyst membranous, semitransparent, with capillary network from host,

encloses two completely hermaphroditic individuals. Young adults cylindrical, but
as they grow, the hindbody becomes elongated and develops irregular out pocket-
ings. Forebody arises from apical depression of hindbody, filiform, scoop-shaped
in oesophageal region. Oral sucker terminal, directly followed by pyriform muscu-
lar pharynx. Oesophagus thin. Caeca not traceable. Ventral sucker absent. Testes
two, tubular, irregularly twisted, near anterior end of hindbody. Ovary and vitelline
gland tubular, narrow, each divided into anterior and posterior branch, parallel, wind-
ing, reach the posterior extremity. Genital junction very large at midbody. Seminal
receptacle retort-shaped. Uterus fills entire hindbody, forms at least three loops. Eggs
13–16 × 9–12.
Remarks: This species is characterized by the strongly diverticulate hindbody and

extremely long and slender forebody.
Didymozoon lobatum Madhavi, 1982 (Fig. 9.67, P1. 1(3&5))

Host: Scombridae: Euthynnus affinis (Cantor).
Location: Gills
Locality: VSK, BOB
Number: 4
Occur encysted in pairs on the gill lamellae.
Description: Completely hermaphroditic. Forebody slender, filiform, 1920–5000
long and 80–112 wide, slightly enlarged at level of oesophagus, attached to anterior
end of hindbody. Latter cylindrical, 6000–11000 long and 510–960 wide, usually flat
on one side, lobed on other side owing to presence of 13–18 crenulations; cylindrical
and unlobed in young adults. Oral sucker pyriform; 48 by 20 in size. Pharynx present.
Ventral sucker absent. Testes cylindrical, paired, asymmetrical, situated overlapping
each other in anterior region of hindbody immediately behind origin of forebody.
Genital junction situated slightly above midbody. Ovary cylindrical, runs close to
smooth edge of hindbody from anterior almost to posterior extremity. Vitellarium
cylindrical, extends along lobed side of hindbody from anterior to posterior end and
gives off one short lateral branch at level of Mehlis’ gland. Uterus fills all available
space in hindbody including that within the lobes. Egg reservoir not prominent. Eggs
small, 15–16 by 8.
Remarks: This species is characterised by the lobed nature of the outer surface of
the hindbody. In this respect it resembles D. diverticulatum Yamaguti, 1970 but in
the latter species, the lobes are irregularly arranged, and the body structure is very
different.
Didymozoon tetragynae Job, 1961

Host: Sphyraenidae: Sphyraena barracuda (Edwards); S. Jello Cuvier
Site: Pectoral fins
Locality: Rameswaram, Tuticorin, Portonovo, Madras, Andaman
Reference: Job (1961)
Description: Cysts long, cylindrical, 12–70 mm long. Forebody slender, long, half
the size of hindbody, arises at one fourth of total length from anterior end. Hindbody
680 thick. Oral sucker 180 × 200. Pharynx absent. Caeca commence just posterior to
oral sucker. Testes paired, nearly one-fifth of total length, club-shaped. Ovary with
four branches. Ovarian and vitelline branches extend anteriorly and posteriorly from
ootype. Ootype located at middle of hindbody. Eggs 1.5 × 1.0.
Remarks: This species is similar to D. apharyngi Job, 1961 in the absence of the
pharynx but differs from it in the extremely long and filamentous forebody, in the
pattern of branching of the ovary and in most of the body proportions.
Genus Indodidymozoon Madhavi, 1982
Diagnosis: Completely hermaphroditic, occur embedded in muscles. Forebody

long, filamentous, attached to hindbody subterminally. Hindbody very long, slender.
Oral sucker terminal. Ventral sucker absent. Pharynx present. Oesophagus simple.
Caeca long, terminate near posterior end of hindbody. Testes cylindrical, long, paired,
extends whole length of hindbody. Vas deferens runs sinuously in median field of
forebody. Genital pore at base of oral sucker. Genital complex near anterior end of
hindbody. Ovary slender, cylindrical, single and unbranched, extends from genital
complex to near posterior end of body. Seminal receptacle absent. Mehlis’ gland
compact. Vitelline gland slender, cylindrical, unbranched, extending whole length of
hindbody. Uterus first descends, forms 5 loops, 2 anterior and 3 posterior. Metraterm
well developed. Eggs small, bean-shaped. Occur embedded in opercular muscles of
teleost fishes. Type species: Indodidymozoon madhaviae Anderson & Cribb, 1994
This genus is characterized by the filamentous body and the origin of the forebody
distant from anterior end of hindbody.
Indodidymozoon madhaviae Anderson & Cribb, 1994 (Fig. 9.53)

[Syn. I. platycephali Madhavi, 1982 nec platycephali (Ku & Shen, 1965) Anderson
& Cribb, 1994]
Host: Platycephalidae: Grammoplites scaber (Linn.)
Location: Opercular muscles
Number: Numerous
Fig. 9.53 Indodidymozoon

madhaviae (after Madhavi,
Description: Body long, slender, filiform, divisible into filamentous forebody

8–38 mm long and 176–192 wide and slightly wider and much longer hind-body mea-
suring 24–50 mm long and 346–560 wide. Forebody attached to hindbody slightly
away from anterior end, resulting in formation of finger-shaped process at anterior
end of hindbody; this process may be either short and blunt or long and filamentous.
Oral sucker elliptical, 52–80 by 40. Ventral sucker absent. Pharynx 40 in diameter.
Oesophagus 320–360 long. Caeca broad, terminate near posterior end of body. Testes
paired, extend throughout length of hindbody, appear as broad ribbon-shaped struc-
tures in young individuals and reduced to thin strands in older ones. Genital complex
in anterior region of hindbody, 1–3 mm from its anterior extremity. Ovary and vitel-
larium cylindrical each represented by single filament running throughout length
of hindbody. Ovarian filament narrow, 16–32 in diameter, commencing 2.5–4.0 mm
from posterior end and terminate near genital complex. Oviduct short. Laurer’s canal
terminates in pouch. Seminal receptacle absent. Mehlis’ gland compact. Vitelline fil-
ament broader than ovarian, 44–62 in diameter, commences at posterior end of hind-
body, terminates near genital complex and leading into prominent vitelline reservoir
posterior to Mehlis’ gland. Uterus first descends describes 5 uterine loops, 2 anterior
and 3 posterior, loops occupying different positions in hindbody, one of two anterior
loops lies anterior to genital complex and other behind; of three descending loops,
proximal lies slightly posterior to midbody, second at about one third of distance
from posterior end and distal loop close to posterior end. Metraterm narrow. tubular,
with thick coat of circular muscles. Eggs 6 × 8.
Remarks: This species was described by Madhavi (1982) as Indodidymozoon platy-
cephali Madhavi 1982 from Platycephalus scaber of Bay of Bengal. Anderson &
Cribb (1994) modified the specific name as I. madhaviae Anderson & Cribb, 1994,
since the original name was preoccupied following transfer of some of the species
from the genus Didymozoon into Indodidymozoon Members of the genus show speci-
ficity towards platycephalid fishes (Plate 1).
Genus Kamegaia Yamaguti, 1970
Diagnosis: Forebody scoop-shaped. Hindbody crenulated, oval in outline, provided

all over with numerous lobular protuberances. Oral sucker rudimentary. Pharynx
muscular. Oesophagus short. Caeca give off number of diverticulae in hindbody.
Testes double, tubular, situated near anterosinistral corner of hindbody. Genital pore
ventrolateral to oral sucker. Ovary with short stem and five irregularly bifurcating
very narrow mini branches extending in greater part of hindbody. Seminal receptacle
present. Vitelline gland divided into five main branches which bifurcate irregularly
and in terminal tubules, more extensive than ovary. Uterus extends in network giving
off one loop or two for each marginal lobe. No egg reservoir. Metraterm strongly
muscular. Eggs very small. Encysted in pairs in muscle wall of intestine of marine
teleosts. Type species: K. kawakawa Yamaguti, 1970
Kamegaia kawakawa Yamaguti, 1970 ((Fig. 9.67, P1. 1(2))

Site: Muscles of intestine
Locality: VSK, BOB
Reference: Madhavi & Sairam (2000)
Genus Lobatocystis Yamaguti, 1965
Diagnosis: Forebody scoop-shaped. Hindbody triangular, with one margin usually

lobed and other two margins straight. Oral sucker contiguous with pharynx. Oesopha-
gus slender. Intestinal bifurcation at level of widest part of forebody. Caeca terminate
near posterior end of hindbody. Testes divided into long unbranched tubules radiating
from junction of vasa efferentia in anterior part of hindbody. Genital pore ventral to
oral sucker. Ovary divided into slender branches reaches to peripheral areas of hind-
body. Seminal receptacle retort-shaped. Vitellarium tubular, very slender, branched,
intertwined among ovary and uterus. Uterus strongly convoluted occupies all avail-
able space of hindbody, without forming egg reservoir. Eggs small, bean-shaped.
Type species: L. yaito Yamaguti, 1965.
Remarks: This genus is characterized by the peculiar shape of the hindbody, the
multiple testes being radially arranged in the anterior part of hindbody and the ovary,
vitellarium and uterus being intertwined throughout the hindbody.
Lobatocystis bengalensis Hussain, Rao & Shyamasundari, 1985

Site: Gills
Locality: VSK, BOB
Reference: Hussain et al., (1985); Madhavi & Sairam (2000)
Description: Body flat, triangular with the convex side flat in middle and 2–3 lobes
on either side each of which divided into 2–3 hemispherical lobes. Posterior end of
body is wedge shaped. Forebody scoop-shaped, arises from middle of flat surface
of hindbody. Forebody 3024 long, 1264 wide. Oral sucker prominent, highly mus-
cular. Pharynx rudimentary. Oesophagus slender, surrounded by thick zone of gland
cells in middle part. Caeca narrow in forebody, inflated in hindbody, terminate near
posterior end of hindbody. Testes divided into 8 long, slender tubular unbranched
lobes, strongly winding arranged radially at base of forebody. Vas deferens thick, runs
anteriorly with metraterm, opens out at base of oral sucker. Ovary tubular, slender,
irregularly branched, in central region, sends slender branches into each marginal
lobe. Seminal receptacle retort-shaped. Vitellarium divided into numerous branches,
extend into terminal lobes, intertwined among ovarian and vitelline tubes. Uterus
strongly convoluted, occupies all available space of hindbody, without forming uter-
ine reservoir. Metraterm strongly muscular. Eggs 12 × 7.
Remarks: This species was differentiated from the type species Lobtocystis yaito
Yamaguti, 1965 by the shape of hindbody and forebody and the presence of thick
zone of gland cells surrounding middle part of oesophagus.
Lobatocystis yaito Yamaguti, 1965 (Figs. 9.54 and 9.67, P1. 1(1))
Site: Gills
Location: VSK, BOB
Reference: Madhavi (1982); Madhavi & Sairam (2000)
Genus Melanocystis Yamaguti, 1970
Diagnosis: Capsule disc-like. Completely hermaphroditic. Enclosed in pairs in flat-

tend oval cysts with hindbodies pressed against each other at the flat surface. Forebody
elongate, attached to midregion of hindbody at flat side. Hindbody disc-like, wider
than long, flat ventrally, convex dorsally. Oral sucker and pharynx present. Caeca
narrow in forebody, but inflated into wide curved cylinders containing dark haematin-
like ingesta. Testes tubular, long, extend along anterior margin of hindbody. Ovary
divided into several winding branches, directed in opposite directions. Vitellarium
tubular, divided into several terminal branches directed posteriorly and ending along
posterior margin of hindbody. Seminal receptacle oval. Uterus occupies all available
space of hindbody. Egg reservoir absent. Eggs bean-shaped.
Fig. 9.54 Lobatocystis yaito

(after Madhavi, 1982
Parasit. 4, p. 102)
Fig. 9.55 Melanocystis

kawakawa (original)
Melanocystis kawakawa Yamaguti, 1970 (Fig. 9.55)

Site: Inner surface of posterior end of oesophagus.
Locality: VSK, BOB
Description: With characters of the genus.
Genus Metadidymozoon Yamaguti, 1970
Diagnosis: Capsule globular, walls with capillary system. Forebody conical. Hind-
body with wide trunk possessing secondary lobes, which end in several terminal
digitiform protrusions. Oral sucker and pharynx present. Oesophagus short. Caeca
narrow. Ventral sucker absent. Testes two, in trunk of hindbody. Genital pore ventral
to oral sucker. Ovary unbranched. Ovary and vitellarium mainly in secondary lobes
and terminal protrusions. Genital junction at middle of hindbody. Seminal receptacle
absent. Hermaphroditic duct short. Vitellarium branched. In gill arches of istiophorid
fishes. Type species: M. branchiale Yamaguti, 1970
This genus is characterized by the light brown or golden yellow colour of the
body and the bean-shaped body flattened between the gill lamellae of the host.
Metadidymozoon branchiale Yamaguti, 1970
Host: Xiphidae: Xiphias gladius (L.)
Site: Gills
Locality: VSK, BOB
Description: With characters of the genus
Genus Neometadidymzoon Yamaguti, 1971
[Syn. Rhopalotrema Anderson, 1998]
Diagnosis: Capsule cylindrical, globular or spherical. Forebody elongate, thin
attached subterminally to hindbody. Hindbody cylindrical, flattened laterally or
tapered towards posterior extremity. Hindbody may be tubercular due to forma-
tion of secondary protrusions and indentations. Pharynx present. Oesophagus long,
surrounded by gland cells. Caeca reach close to posterior extremity. Ventral sucker
present. Testes in both regions of body. Genital pore ventral to pharynx. Ovary and
vitellarium unbranched. Seminal receptacle absent. Mehlis’ gland large. In tissues
of priacanthid fishes. Type species: N. polymorphis (Oshmarin & Mamaev, 1963)
Yamaguti, 1971.
Neometadidymozoon polymorphis (Oshmarin & Mamaev, 1963) Yamaguti, 1971

(Fig. 9.56)
[Syn. Didymzoon polymorphis Oshmarin & Mamaev, 1963]
Host: Priacanthidae: Priacanthushamrur (Forsskål)
Site: Inner wall of operculum
Locality: VSK, BOB
Distribution: Tonkin Bay, North Vietnam
Description: Body consists of two parts, slender forebody and considerably larger
hindbody. Forebody elongated and thread-like, 380–12050 long. Hindbody cylin-
drical and flattened, 3560–18900 by 730–3200 wide. Oral sucker terminal, round.
Pharynx rudimentary. Oesophagus 340–1296 long. Testes conspicuous in forebody,
not easily visible in hindbody. Metraterm conspicuous. Testes elongated, coiled arise
from posterior end of hindbody, extend into forebody. Ovary and vitellarium single,
both convoluted and tubular, reaching close to posterior end of hindbody. Genital
junction 544 from origin of forebody. Uterus first descends forming three loops, one
anterior and 2 posterior, finally extending into forebody as metraterm in intercaecal
field. Genital opening lies midventral to oral sucker. Eggs 12 × 6.
Remarks: Oshmarin & Mamaev (1963) described this under the genus Didymozoon
as D. polymorphis from Priacanthus triacanthus of North Vietnam (Tonkin Bay).
Yamaguti (1971) erected a new genus Neometadidymzoon Yamaguti, 1971 to include
Didymozoon polymorphis.
Fig. 9.56 Neometadidymozoon

polymorphis (after Madhavi,
Genus Platocystoides Madhavi, 1982
Generic diagnosis: Forebody long, slender attached to centre of hindbody. Hind-

body flattened, disc-like, with distinct furrows on ventral side which divide it into
a number of finger-shaped lobes radiating from centre. Oral sucker and pharynx
present. Ventral sucker absent. Caeca narrow in forebody, not traceable in hindbody.
Testes two, tubular, winding, close to one another on right side of hindbody. Ovary
consists of several branches, radiating from genital junction to periphery. Seminal
receptacle present. Vitellarium branched, branches radiating from genital complex,
arranged in two groups of 3 and 4. Uterine coils filling entire hindbody. Uterine reser-
voir small. Eggs small, numerous. Parasitic beneath scales of marine fishes. Type
and only species: P. polyaster (Job, 1962).
Platocystoides polyaster (Job, 1962) Madhavi, 1982 (Fig. 9.57)

[Syn: Platocystis polyaster Job, 1962]
Hosts: Sphyraenidae: Sphyraena obtusata Cuv., S. picuda Bl. & Schn.,
Location: Scales
Number: Numerous
Reference: Job (1962), Madhavi (1982)
Job (1962) described this species as Platocystis polyaster Job, 1962 from the
scales of Sphyraena obtusata from the Pamban area (Bay of Bengal). Madhavi (1982)
based on study of specimens apparently conspecific with P. polyaster and collected
from the scales of S. obtusata and S. picuda caught off the Visakhapatnam coast,
showed that they cannot be assigned to the genus Platocystis Yamaguti, 1938, or to
any other known didymozoid genus. Platocystoides Madhavi, 1982 was, therefore,
erected to include P. polyaster (Job, 1962) as type species. Yamaguti (1971) also
expressed doubts about the inclusion of P. polyaster in the genus Platocystis.
Description: Encysted in pairs in thin-walled cysts underneath scales of fishes, with

flat ventral surface of one individual pressed against ventral surface of second indi-
vidual and with forebodies tucked in between. Body divisible into slender forebody
and disc-like flattened hindbody. Forebody slender 2240–2880 long and 80–1440
wide, attached to central part of hindbody. Hindbody 2100–3000 by 4000–4240,
greatly flattened and expanded, thick medially and thin peripherally, thus fitting
Fig. 9.57 Platocystoides

polyaster (after Madhavi,
contour of scales, ventral surface with numerous deep furrows which radiate from
central region to periphery and divide central part into finger-shaped lobes. Oral
sucker small, pyriform, terminal, 40–60 by 20 in size. Ventral sucker absent. Phar-
ynx globular. Oesophagus very long. Caeca narrow in forebody, not traceable in
hindbody. Initial portions of caeca surrounded by numerous uninucleate gland cells.
Testes two, elliptical, unequal in size, slightly sinuous, situated closely opposed to
one another on right side of disc close to origin of forebody. Genital complex situated
in hindbody near to its junction with forebody. Both ovary and vitellarium appear
as winding filaments radiating from region of Mehlis’ gland to periphery of disc.
Ovary with six filaments, two directed anteriorly, two posteriorly and two on left, all
joining near genital junction. Seminal receptacle situated anterior to Mehlis’ gland.
Vitellarium consists of 7 winding filaments, arranged in two groups, four on right and
three on left. Mehlis’ gland small, compact and enclosed in thin membrane. Uterine
coils numerous, filling all space in hindbody, narrower coils occupying peripheral
part of disc and wider distal coils occupying central part. Metraterm well developed,
forms small reservoir at base of forebody. Eggs small (10 × 6), numerous.
Remarks: There are two reports of this species: Bay of Bengal (Madhavi, 1982) and
Kuwait Bay (Abdul-Salam & Sreelatha, 1993).
Genus Renodidymocystis Madhavi, 1982
Diagnosis: Encysted in pairs in cysts with thin cyst wall. Forebody slender. Hind-
body spherical in young individuals, develops several lobes in gravid individuals.
Oral sucker small, directly followed by pharynx. Oesophagus short. Ventral sucker
absent. Testis single, twisted, unbranched. Ovary and vitellarium consist of long
filaments extending from genital complex into terminal lobes forming one or more
loops in each lobe. Uterine reservoir absent. Eggs bean-shaped. Parasitic in kidneys
of marine teleosts. Type species: Renodidymocystis yamagutii Madhavi, 1982.
Renodidymocystis yamagutii Madhavi, 1982 (Fig. 9.58)
Cysts occur embedded in the kidney tissue, each cyst encloses two worms with
their flat surfaces in contact and forebodies tucked into space between.
Fig. 9.58 Renodidymocystis

yamagutii (after Madhavi,
Description: Completely hermaphroditic. Forebody short, 320–480 long 48–80

wide, slender, delicate, slightly enlarged at oesophageal level and conical anteri-
orly, attached to anterior end of hindbody. Hindbody 800–1920 long, 780–1600
wide, shape varies with degree of maturity, spherical in young and lobed in mature
individuals; lobes develop gradually. Three main lobes develop first and these later
develop primary, secondary and tertiary lobes. Number of lobes increases gradually,
gravid individuals appear highly lobed. Oral sucker circular, small, poorly muscular,
16–20 in diameter. Ventral sucker absent. Pharynx globular, bigger than oral sucker.
Oesophagus short. Caeca not traceable. Testis single, arcuate, in middle of hind-
body in young non-diverticulate individuals, wide and twisted in anterior region of
hindbody in gravid individuals. Genital pore at base of oral sucker. Genital complex
situated in middle of hindbody, ovarian and vitelline filaments radiating from it to
periphery of hindbody. Ovarian filaments wide, few, (3 or 4), unbranched, increas-
ing in size with growth of individuals and extending into some lobes of hindbody.
Vitelline filaments wide, 5–7 in number, very long, either unbranched or giving off
one or two branches, extend into lobes of hindbody, forming loops in one lobe before
entering the next, usually confined to lobes not occupied by ovarian filaments. Semi-
nal receptacle absent. Mehlis’ gland with compact mass of gland cells which extend
along proximal uterine coils. Uterine coils numerous extending into almost all ter-
minal lobes, where they usually form more than one loop. Uterine reservoir absent.
Metraterm poorly developed. Eggs small (8–10 × 4), bean-shaped.
Remarks: Pozdnyakov and Gibson (2008) synonymized the genus Renodidymo-

cystis with Lobatozoum Ishii, 1935 but the genus Renodidymocystis appears quite
distinct in its lobed nature of the body, the occurrence of a single testis that is spirally
twisted and in the absence of a seminal receptacle. The genus Renodidymiocystis is
revalidated here. R. yamagutii is the only species recorded in the genus so far.
Subfamily Colocyntotrematinae Yamaguti, 1958
[Syn. Annulocystinae Yamaguti, 1970; Didymocodiinae Yamaguti, 1970;

Neodiplotrematinae Yamaguti, 1958; Opepherocystinae Yamguti, 1970;
Opepherotrematinae Yamguti, 1958; Osteodidymocodiinae Yamaguti, 1980;
Phacelotrematinae Yamaguti, 1971; Pseudocolocyntotrematinae Yamguti, 1970;
Reniforminae Yamaguti, 1970]
Diagnosis: Worms in capsules fused, but with forebodies free, degree of fusion of
hindbodies varies from fusion at one end only to full union of two or more individuals.
Ventral sucker absent. Pharynx present. Oesophagus long. Caeca extend into posterior
region. Gland cells around oesophagus present or absent. Testes elongate or compact,
unbranched or rarely branched, one or two in each individual. Ovary and vitellarium
branched or not. Seminal receptacle present or absent. Mehlis’ gland well developed.
Excretory vesicle tubular. Type genus Colocyntotrema Yamaguti, 1958
Key to genera
1. Individuals not entirely fused, forebodies free……………. Opepherocystis Yam-
aguti, 1970
Individuals completely fused, forebodies in spherical cavities ……………2
2. Hindbody branched, testes branched………..……………… Didymocodium
Yamguti, 1970
Hindbody not branched, testes not branched……………………………… 3
3. Fused hindbody spherical……………………..……. Pseudocolocyntotrema
Yamaguti, 1970
Fused hindbody skittle-shaped ………….…………. Allopsuedocolocyntotrema
Yamaguti, 1970
Genus Allopseudocolocyntotrema Yamaguti, 1970
Diagnosis: Cyst claviform, attached to inner surface of intestine, contains two sepa-
rate forebodies anteriorly and two hermaphroditic hindbodies which are completely
fused into one. Cyst membrane thin, with capillary network from host fish. Forebody
slender, enclosed in deep ventral pit of hindbody. Oral sucker weakly muscular. Phar-
ynx strongly muscular. Ventral sucker absent. Caeca extend into hindbody. Testes
compact, in two sets of two each in pre-equatorial zone of fused hindbody. Ovary and
vitellarium tubular, bifurcating several times into numerous winding long or short
branches, occupying peripheral area of hindbody, vitellarium more extensive than
ovary. Uterus strongly convoluted in hindbody. Type species: Allopseudocolocyn-
totrema claviforme Yamaguti, 1970.
The genus differs distinctly from Pseudocolocyntotrema in the body shape and in
the testes being more compact.
Allopseudocolocyntotrema claviforme Yamaguti, 1970 (Fig. 9.59)
Fig. 9.59 Allopseudocolocyntotrema

claviforme (after Yamaguti,
1970)
Site: Encysted on inner surface of intestine.

Locality: VSK, BOB
Description: with characters of the genus
Remarks: A claviforme was first reported by Yamaguti (1970) from Euthynnus yaito
from Hawaii. Madhavi& Sairam (2000) reported its occurrence in E. affinis of Bay
of Bengal.
Genus Indoglomeritrema Madhavi & Rao, 1983
Diagnosis: Encysted in pairs in a thick fusiform capsule, contains a capillary net-

work extending into the space between coiled worms. Two fellow occupants not
fused but entangled as compact mass. Body filiform, very long, fragile nearly uni-
form in width. Oral sucker and pharynx present. Oesophagus long. Caeca narrow,
terminate near anterior margin of testes. Ventral sucker present. Testes paired, long,
tubular, symmetrical, originate near caecal ends, extend to proximal end of ovary.
Ovary single, tubular, narrow, extends from behind testes to near genital junction.
Seminal receptacle absent. Uterus descends to posterior extremity then ascends to
genital pore. Vitellarium single, tubular, extends between genital junction and poste-
rior extremity. Parasitic on gills of marine fishes. Type and only species: I. epinepheli
Madhavi & Rao, 1983
Indoglomeritrema epinepheli Madhavi & Rao, 1983

Host: Serranidae: Epinephelus tauvina (Forskål)
Site of infection: gills
Locality: VSK, BOB
Reference: Madhavi & Rao (1983)
Description: Cysts fusiform, 6.0–9.5 mm long and 1.76–2.0 mm wide and attached
lengthwise along lower surface to gill filament. Cyst wall thick, contains two layers:
outer thin hyaline layer containing numerous vesicles and inner thick layer formed
of several layers of connective tissue. Cyst wall is supplied by host capillaries which
extend all over cyst wall and enter into cyst cavity into space between compact
coils of worms. Each cyst encloses two worms. Worms long, filamentous, fragile,
coiled and entangled forming compact mass. Body length exceeds several metres,
more or less uniformly wide, 80–120 in width. Oral sucker circular, 24 in diameter,
poorly developed. Ventral sucker circular, 28 in diameter, situated at distance of 380
from anterior end. Pharynx rudimentary but present. Oesophagus 200 long. Intestinal
bifurcation midway between suckers. Caeca short take zigzag course to level of testes
and·terminate shortly after entering testicular zone, proximal part narrow, widened
distally, surrounded by gland cells. Testes paired, ribbon-shaped, symmetrical, extend
over considerable area from caecal ends to terminate short of ovary. Seminal vesicle
long takes sinuous course between caeca and joins metraterm before opening into
genital atrium. Genital pore at base of oral sucker. Ovary single, tubular, extends from
below testes to level of Mehlis’ gland. Seminal receptacle and Laurer’s canal absent.
Vitellarium single, tubular, exceeds length of ovary, extends from genital junction to
near posterior end. Mehlis’ gland well developed, its cells distributed over ootype
and proximal uterine coils to considerable length. Uterus first descends to posterior
end, then takes ascending turn finally to open into genital atrium. Eggs small, bean-
shaped, numerous, golden yellow in colour, 18–20 by 8 in size. Metraterm well
differentiated.
Remarks: The genus Indoglomeritrema has characters of the subfamilies Nema-

tobothriinae and Glomeritrematinaeas given by Yamaguti (1971). Except for the
presence of pharynx, the structure of the worm very closely fits into that of Allone-
matobothrium Yamaguti, 1965 of the subfamily Nematobothriinae. However, the
important features which separate the genus from Allonematobothrium and from all
members of the Nematobothriinae are: (1) the presence of a thick capsule formed
of several layers of connective tissue. Some nematobothriines do occur encysted but
in a cyst with very thin and delicate wall; (2) the extension of the capillary network
of host origin into the cyst wall and subsequently into the cyst cavity for supplying
nutritive material to the parasite. Among genera having a capillary network the genus
that bears the closest resemblance to Indoglomeritrema is Glomeritrema Yamaguti,
1942 of the subfamily Glomeritrematinae. However, in this genus the cyst wall is
thin and the cyst cavity is divided by septa, the structure of the worm is also different
in that a ventral sucker is absent and there is a single testis. There are also differences
in the arrangement of gonads.
Genus Didymocodium Yamaguti, 1970
Diagnosis: Cyst wall thin, without any capillary network of host origin. Two fore-
bodies free, enclosed at base in oval slit on midventral surface of hindbody. Two
hindbodies completely fused and divided into several primary lobes which in turn
divide dichotomously into a number of secondary and primary lobules. Oral sucker
and pharynx present. Ventral sucker absent. Caeca terminate blindly in hindbody.
Testes in two sets of several tubular branched lobes, each set situated lateral to base
of forebody. Ovary and vitellarium long, branched, ovary strongly winding around
testicular area and extending into every primary and some tertiary branches of hind-
body. Vitelline branches largely in peripheral area of each lobule, encircling uterine
coils. Genital junction just lateral to outer end of testes. Uterus coiled mostly lon-
gitudinally in each tubule, leaving entire testicular area free. No uterine reservoir.
Eggs numerous, bean-shaped, small.
Type species: D. euthynni Yamguti, 1970

Didymocodium euthynni Yamaguti, 1970
site: VSK, BOB
Genus Opepherocystis Yamaguti, 1970
Diagnosis Capsule globular. Forebodies elongate at some distance from each other.
Posterior region spherical, contains vestibular canal. Oesophagus short. Caeca reach
posterior region where they are greatly enlarged. Testes elongate, one in each fused
individual. Genital pore ventral to oral sucker. Ovary and vitellarium unbranched.
Seminal receptacle present. In wall of pyloric caeca of scombrids. Type species: O.
kawakawa Yamaguti, 1970
Opepherocystis kawakawa Yamaguti, 1970
Scombridae: Euthynnus affinis (Cantor)
VSK, BOB
Genus Opepherotrematoides Yamaguti, 1970
Diagnosis: Cysts rounded contain fused hindbody and two forebodies attached to
hindbody Each individual contains one set of reproductive organs. Forebody enclosed
in cylindrical canal opening midventrally and extending deeply into parenchyma
Oral sucker terminal, rounded, cellular. Pharynx present. Caeca narrow in forebody,
diverticulated in hindbody. Testes two, on each side of hindbody at level of mid-
ventral pore. Common genital pore ventral to oral sucker. Ovary and vitellarium
tubular, long, very narrow, branched dichotomously into several terminal branches
distributed in peripheral area of hindbody. Vitellarium more extensive than ovary,
with more numerous branches. Seminal receptacle present. Uterine coils profuse
in hindbody, metraterm well developed, extend from genital pore to near genital
junction to testicular region of each side, convoluted before entering forebody. Eggs
numerous, bean-shaped.
Type species: O. multitubularis Yamaguti, 1970
Remarks: This genus is characterized by: the presence of a midventral pit in the
hindbody, one pair of testes on each side, extremely narrow and branched ovary and
vitellarium, caeca which terminate in the hindbody in numerous irregular diverticles.
Opepherotrematoides multitubularis Yamaguti, 1970 (Fig. 9.67, P1. 1(4))
Site: Serosa of stomach
Locality: VSK, BOB
Genus Pseudocolocyntotrema Yamaguti, 1970
Diagnosis: Cyst round, contains two individuals equal in size, and fused completely
by hindbody. Cyst membrane thin with blood capillaries from host. Forebody slender,
enclosed in ventral pit of hindbody. Hindbody globular. Oral sucker poorly developed.
Ventral sucker absent. Caeca extend into hindbody. Testes winding, tubular or bent
back on themselves close to ventral surface of hindbody. Ovary and vitellarium
tubular, branched, extend in peripheral area of hindbody, occupying two quadrants
alternately. Uterus coiled intricately in hindbody internal to ovary and vitellarium.
Metraterm well differentiated. Genital pore ventral to oral sucker.
Type species: P. yaito Yamaguti, 1970

Pseudocolocyntotrema yaito Yamaguti, 1970 (Figs. 9.60 and 9.67, P1. 1(6))
Site: Intestinal wall
Distribution: VSK, BOB; Hawaii
Description: Two hermaphroditic individuals fused together in form of pumpkin

and enclosed in thin hyaline capsule of host origin with forebodies bent on them-
selves in ventral pit of hindbody. Forebody 300–700 in length, 90–240 in maximum
width. Hindbody 1400 × 1300 in size, smooth, without meridional septa. Mouth ter-
minal, oral sucker 37–50 × 33–47, contains abundant cellular mass. Pharynx mus-
cular. Oesophagus narrow, bifurcates behind widest part of forebody. Caeca narrow
in forebody, wider in hindbody, terminate blindly near in front of dorsal dome.
Testes winding, situated longitudinally one on each side along ventral margin of
hindbody. Vas deferens running alongside metraterm in forebody, genital opening
ventral to oral sucker. Ovary irregularly dichotomous situated close to left margin
in left ventral quadrant and right dorsal quadrant of hindbody. Vitellarium tubular
and irregularly branched, extend close to body wall in right ventral quadrant and left
dorsal quadrant of hindbody. Uterus largely internal to ovary and vitelline branches,
occupies all hindbody except dome-shaped region. Metraterm well differentiated.
Eggs 12–14 × 7.5–9.3.
Remarks: This species was first reported by Yamaguti (1970) from the tunas, Euthyn-
nus yaito and Auxis thazard of Hawaii. Later Madhavi (1982) and Madhavi & Sairam
(2000) recorded it from Euthynnus affinis off Visakhapatnam coast, Bay of Bengal.
The cysts occur attached to intestinal wall and appear as prominent yellowish masses.
Fig. 9.60 Pseudocolocyntotrema

yaito (after Madhavi, 1982,
Parasit. 4, p. 122)
Subfamily Didymodiclininae Pozdnyakov, 1993
Diagnosis: Completely gonochoristic. Sexual dimorphism weakly expressed. Body

of both sexes filiform. Oral sucker present. Ventral sucker present or absent. Pharynx
present. Oesophagus simple. Caeca simple, blind. Male: Testes two, long, parallel
reach close to posterior extremity. Vasa efferentia short. Ejaculatory duct long. Rudi-
ments of female reproductive system usually absent. Female: Genital junction at
about midbody. Ovary and vitellarium usually unbranched, rarely branched, ovary in
anterior half of body and vitellarium in posterior half of body. Uterus forms numerous
loops. Metraterm well developed. Eggs small. Type genus: Didymodiclinus Pozd-
nyakov, 1993
Key to genera
1. Body filamentous……………………………………………..............……… 2
Female body divisible into fore and hindbodies…………………………………
………….…. Neogonapodasmius Radhakrishnan & Nair, 1981
2. Uterus with a single loop ……………………….…………….. Gonapodasmius
Ishii, 1935
Uterus with numerous loops ………………………….. Didymodiclinus Pozd-
nyakov, 1993
Genus Didymodiclinus Pozdnyakov, 1993
Diagnosis: Encapsulated or not. Body filiform, long. Ventral sucker present.

Oesophagus long or short. Caeca reach close to posterior extremity. Male: Smaller
than female. Testes two, parallel, reach close to posterior extremity. No rudiments
of female reproductive system. Female: Larger than male. Genital junction approx-
imately at midbody. Ovary long, narrow, curved, in anterior half of body. Seminal
receptacle present or absent. Uterus with single loop. Vitellarium similar in form to
ovary, in posterior half of body. Type species: D. pacificus (Yamaguti, 1970) Pozd-
nyakov, 1993.
Didymodiclinus branchialis (Yamaguti, 1970) Pozdnyakov, 1994

[Syn. Gonapodasmius branchialis Yamaguti, 1970]
Host: Serranidae: Epinephelus latifasciatus (Temn. & Schl.)
Site: Gill lamellae
Locality: VSK, BOB
Distribution: Hawaii, South China Sea, BOB
Description: Worms encapsulated along length of gill lamellae. At each location one
long, bright yellow female occurs in association with one small opaque male. Female
extends along several adjacent lamellae tracing deep tunnel-like route along longi-
tudinal axis but never extending to tip of gill lamella. Male within coils of female.
Rarely, two females occur associated with one male. Female digs into gill bone
constructing connection between adjacent lamellae. Male: Body filiform, opaque,
6.40–18.90 mm long. Oral sucker 48–62 × 35–50. Mouth terminal. Pharynx rudi-
mentary. Oesophagus present. Caeca extend along entire length of body, initial part
covered by gland cells. Ventral sucker larger than oral sucker, 105–148 in diame-
ter. Sucker ratio 1:2. Testes long, tubular, paired, originate at different levels from
posterior extremity. Vas deferens narrow, median, muscular, opening ventrally to
oral sucker. Female genitalia represented by empty, rudimentary metraterm. Female:
Body filiform. Oral sucker 62–74 × 48–52. Oesophagus long. Digestive system sim-
ilar to that in male. Ovary tubular, winding in anterior part of body. Juel’s organ
small, bean-shaped. Genital junction divides body in 1:1.8 ratio. Initial descending
uterus contains abundant spermatozoa forming uterine seminal receptacle. Uterus
loops at distance of 72–140 from posterior end from where it forms ascending arm
which transforms into metraterm anterior to ventral sucker and opens through com-
mon genital pore ventral to oral sucker. Vitellarium single, tubular, arising 180–218
from posterior tip. Male genitalia represented by a poorly formed vas deferens. Eggs
small, 16–20 × 8–12, bean-shaped.
Remarks: Yamaguti (1970) described this species as Gonapodsmius branchialis

from the gill lamellae of an unknown sea bass and from the olfactory organ of
Epinephelus quernus. Later it was recorded from Epinephelus latifasciatus in the
Bay of Bengal (Madhavi, 1982) and Cephalopholis platycentron from the China
Sea (Gu & Shen,1983). Podznyakov (1994) included it in the genus Didymodiclinus
Pozdnyakov, 1994 as D. branchialis (Yamaguti, 1970).
Didymodiclinus reticulum (Murugesh & Madhavi, 1994)

[Syn. Gonapodasmius reticulum Murugesh & Madhavi, 1994]
Host: Serranidae: Epinephelus radiatus (Day)
Site: VSK, BOB
Habitat: The bright chrome yellow worms are long and filiform and burrow deep into
the host tissue forming an anastamosing (reticulate) network. The males are much
smaller than the females and lie between the body coils of the elongated females.
Description: Male: Filiform, opaque, grey. Body slender, 49–52 mm long, 320–450
wide, rounded anteriorly enlarged at level of ventral sucker, blunt posteriorly. Oral
sucker terminal, small, ovoid, 60–84 by 60–66. Ventral sucker 130–134 × 90–102.
Pharynx non-muscular. Oesophagus simple, lined initially by gland cells. Intestinal
bifurcation just anterior to ventral sucker. Caeca lined with prominent gland cells
initially, becoming glandular posteriorly, extending to posterior extremity where they
end blindly. Testes paired, tubular, originate at different levels from posterior extrem-
ity, anteriorly running almost parallel to each other. Vas deferens sinuous, in central
field of body, opens out through genital pore ventrolateral to oral sucker. Female
organs absent except for the presence of narrow rudimentary metraterm lying anteri-
orly to ventral sucker, adjacent to vas deferens. Female: Filiform, yellow, thread-like,
67–90 mm long, 440–490 wide; maximum width at level of intestinal bifurcation.
Oral sucker ovoid, 90–104 × 64–66, non-muscular. Pharynx oval. Oesophagus sim-
ple anteriorly, glandular posteriorly. Caeca lined initially by thick layer of gland
cells. Ventral sucker 112–140 × 92–104. Ovary single, tubular, posteriorly directed,
originating in anterior part of body. Juel’s organ sac-like, posterior to Mehlis’ gland.
Vitellarium tubular, single, forwardly directed joining ovary at Mehlis’ gland com-
plex. Genital junction situated in posterior part of anterior third of body. Uterus first
descends, loops posteriorly and ascends forwards forming metraterm. Metraterm
thick, with numerous, bean-shaped eggs 18–20 × 10–12.
Didymodiclinus spilonotopteri (Yamaguti, 1970) Pozdnyakov, 1994

[Syn. Gonapodsmius spilonotopteri Yamaguti, 1970]
Host: Exocoetidae: Cypselurus comatus (Mitchill)
Locality: VSK, BOB
Distrition: Hawaii, BOB
Description: Male: Body very narrow, 6.3–21 mm long 100–350 wide, anterior
extremity bluntly pointed, posterior extremity blunt. Oral sucker 35–107 × 32–88.
Ventral sucker 33–54 in diameter. Pharynx small. Oesophagus long. Caeca termi-
nate at posterior extremity. Testes parallel to each other, originate at different levels
short of posterior extremity. Genital pore postero-lateral to oral sucker. Female:
Body filiform, 56–141 mm long, 280–380 wide. Digestive system as in male. Ovary
tubular, comparatively wide, originates 6.0–10.0 mm posterior to anterior extremity.
Seminal receptable cylindrical, directed forward from genital junction. Genital junc-
tion divides body in approximately ratio of 1:4–8. Uterus descends, then ascends,
Metraterm not very distinct. Eggs bean-shaped 20–23 × 12–16. Vitellarium up to 70
wide, extends between posterior extremity and genital junction.
Remarks: Yamaguti (1970) reported this species from Cypselurus spilonotopteri

and C. spilopterus, Hawaii. It differs from D. branchialis in the poorly developed
ventral sucker, situated far post-bifurcal. Madhavi (1982) reported its occurrence in
Cypselurus comatus off the Visakhapatnam coast, Bay of Bengal.
Genus Neogonapodasmius Radhakrishnan & Nair, 1981
Diagnosis Completely gonochoristic. Female large, with slender filamentous fore-

body and cylindrical hindbody. Male small, cylindrical. Oral sucker and oesopha-
gus present. Pharynx and ventral sucker absent. Caeca reach to posterior extremity.
Genital pore at base of oral sucker. Male: Testes two ribbon-shaped, juxtaposed,
extend close to posterior extremity. Rudiments of female reproductive organs present.
Female: Ovary with two branches, tubular, winding, anterior to genital junction. Vitel-
larium with two branched tubes, running posterior to genital junction. Uterus with
ascending and descending coils. Egg reservoir present. Type species: N. hemiramphi
Radhakrishnan & Nair, 1981.
Neogonapodasmius hemiramphi Radhakrishnan & Nair, 1981 (Fig. 9.61)

Host: Hemiramphidae: Hyporamphus xanthopterus (Val.)
Site: Dorsal fin.
Locality: Trivandrum coast (AS)
Reference: Radhakrishnan & Nair (1981a)
Description: Completely gonochoristic with pronounced sexual dimorphism Oral
sucker terminal, 320–480 × 285–448. Pharynx absent. Oesophagus long, simple.
Caeca narrow, tubular, extend more or less to posterior extremity. Male: Small cylin-
drical transparent, without demarcation into fore and hindbodies, 8.0 mm in length
and 0.58 mm in width. Testes paired, ribbon-shaped, juxtaposed running through
greater part of hindbody, confluent anteriorly. Genital pore at base of pharynx. Rudi-
ments of female reproductive organs present. Female: Female long, yellowish with
body demarcated into fore and hindbodies. Forebody filamentous, 8.5–14.0 mm long,
176–320 wide. Hindbody cylindrical, 14–15 mm long and 1.44–3.2 mm wide. Ovary
consists of two tubular slender, coiled branches confined to anterior part of hindbody,
to area anterior to genital junction. Genital junction situated far anterior to middle of
hindbody. Mehlis’ gland large. Uterus with ascending and descending coils running
in zigzag manner to either side. Egg reservoir large, occupies entire middle part of
hindbody. Metraterm well developed. Vitellarium with two main branches giving
off numerous sub-branches, branches narrow, extend from genital junction to near
posterior end.
Remarks: Radhakrishnan & Nair (1981a) distinguished Neogonapodasmius from
the related genera Gonapodasmius Ishii, 1935 and Paragonapodasmius Yamaguti,
1938, by the division of the female body into distinct fore and hindbodies and the
branched nature of ovary and vitellarium.
Fig. 9.61 Neogonapodasmius

hemiramphi (after
Radhakrishnan & Nair,
1981)
Subfamily Nematobothriinae Ishii, 1935
Diagnosis Hermaphroditic, free in host tissues. Body not divided into regions,
fusiform, tape-like. Oral sucker present. Ventral sucker present or absent. Pharynx
present or absent. Gland cells around oesophagus and anterior part of caeca present or
absent. Oesophagus and caeca simple. Testes elongate, usually two, parallel, oblique
or in tandem. Genital pore ventral to pharynx or oral sucker. Ovary and vitellarium
very elongate, usually unbranched. Ovary in anterior half of body. Mehlis’ gland
well developed. Seminal receptacle and Juel’s organ present. Uterus long with one
to three loops. Metraterm well developed. Hermaphroditic duct present, sometimes
absent. Eggs small, numerous. Type genus: Nematobothrium van Beneden, 1858
Key to Genera
1. Body long, filiform……………………………………………………………2
Body short, fusiform ……………………………… Skrjabinozoum Nikolaeva &
Parukhin, 1969
2. Uterus with three loops………………………………………..………..………3
Uterus with single loop……………………….…………………………...…….6
3. Ventral sucker absent……………………………………..………………………5
Ventral sucker present …………………………………………………………4
4. Ovary and vitellarium unbranched……………………… Nematobothrium van
Beneden, 1858
Ovary and Vitellarium each with 2 branches ……….…… Metanematobothrium
Yamaguti, 1938
5. Testes double, oesophagus with branches and gland cells, eggs oval
………………………………………………………………Neonematobothrium
Yamaguti, 1965
Testis single, oesophagus without branches, eggs unipolar
………………………………………………Oculonematobothrium Murugesh,
Sairam & Madhavi, 1992
6. Testes long, extend to posterior end of body ……………. Metanematobothri-
oides Yamaguti, 1965
Testes confined to anterior third of body ……………….……Allonematobothrium
Yamaguti, 1965
Genus Nematobothrium van Beneden, 1858
Diagnosis Body thin, long. Pharyx present. Oesophagus simple. Caeca terminate
close to posterior extremity. Gland cells around anterior part of caeca absent. Testes
elongate, not long. Genital pore at tip of genital sucker, ventral to oral sucker or
pharynx. Genital junction at about midbody. Ovary long, unbranched, in anterior half
of body. Seminal receptacle present. Uterus with three loops. Vitellarium similar in
form to ovary, in posterior half of body. Type species: N. filarina van Beneden, 1858.
Fig. 9.62 Nematobothrium

megalaspium (after
Murugesh, Sairam &
Madhavi, 1992)
Nematobothrium megalaspium Murugesh, Sairam & Madhavi, 1992 (Fig. 9.62)

Host: Carangidae: Megalaspis cordyla L.
Site: Free in body cavity
Locality: VSK, BOB
Reference: Murugesh et al. (1992)
Description: Body long, slender filiform, 960–1820 mm long 192–640 wide, ante-
rior region acute, posterior end rounded. Oral sucker terminal 128–144 × 60–70.
Pharynx replaced by dense mass of gland cells situated immediately posterior to oral
sucker. Oesophagus simple, narrow. Intestinal bifurcation halfway between suckers.
Caeca long surrounded by gland cells in the anterior part, continuing posteriorly as
convoluted winding arms, ending posteriorly. Ventral sucker larger than oral sucker.
Testes paired, tubular, obliquely tandem originating at different levels, Ovary tubu-
lar, single, originating posterior to anterior testis. Genital junction at middle of body.
Uterus with 3 loops. Vitellarium single, tubular runs posteriorly from genital junction.
Eggs small 16–20 × 12.
Remarks: This species differs from all the other species in the genus in the absence
of a pharynx, the presence of well-developed gland cells in the area immediately
posterior to the oral sucker and in the initial portions of caecal arms being lined by
well-developed nucleated gland cells.
Genus Allonematobothrium Yamaguti, 1965
Diagnosis: Body slender, very long. Oral sucker strongly muscular, pharynx absent.
Oesophagus long surrounded by gland cells, Caeca simple reach posterior extremity.
Ventral sucker present. Testes paired, originate posterior to genital junction, terminate
just anterior to ovary. Ovary single, tubular, unbranched, intertesticular. Uterus first
descending, looping at posterior extremity. Vitellarium tubular, unbranched, extends
between genital junction and posterior extremity. Type species: Allonematobothrium
epinepheli Yamaguti, 1965
Allonematobothrium epinepheli Yamaguti, 1965

Host: Serranidae: Epinephelus tauvina (Forskål)
Site: Opercular muscles
No. 7
Locality: VSK, BOB
Reference: Madhavi (1982); Murugesh et al. (1992)
Description: With characters of the genus Allonematobothrium. Body long, worm-

like, 76–206 mm long, 580–800 wide. Genital junction situated at midbody, dividing
the body in the ratio of 1:1. Testes commenced at different levels behind genital
junction. Eggs 20–24 × 8–10.
Remarks: Yamaguti (1965) reported the species from Epinephelus quernus of

Hawaii. The BOB specimens while agreeing fully with the Hawaiian specimens,
differ slightly in the position of genital junction, position of ovary and testes and the
egg size.
Genus Metanematobothrium Yamaguti, 1938
Diagnosis: Not encapsulated. Body thin, long, pharynx present. Caeca termi-
nate close to posterior extremity. Testes long, parallel, extend almost along entire
body length. Genital junction at about mid-hindbody. Ovary filiform, thin, with
two branches one branch directed anteriorly, other posteriorly. Seminal receptacle
present. Uterus with three loops. Vitellarium fiiform, usually with two branches.
Type species: M. guernei (Moniez, 1891) Yamaguti, 1938.
Metanematobothrium bivitellatum Mamaev, 1968

Locality: VSK, BOB
Ref. Madhavi & Sairam (2000)
Description: Worms elongate, of uniform width, 21–60 mm long, 1800–2690 wide.

Oral sucker muscular, small, 190–200 in diameter. Ventral sucker rudimentary. Phar-
ynx small. Oesophagus surrounded by dense zone of gland cells, forms stomach-like
swelling. Caeca unequal, extend up to posterior end of body. Testes two, tubular,
extend to near posterior end of body, winding. Ovary with two branches, anterior
branch longer. Juel’s organ present. Vitellarium with 2 branches, an anterior and a
posterior. Uterus with four arms and three loops. Eggs boat-shaped, 18–20 × 12–16.
Remarks: Metanematobothrium bivitellatum was descried by Mamaev (1968) from

the pericardium, blood vessels of the gills, liver, kidneys and body cavity of E. affinis
and A. thazard, It was differentiated from the only other species under the genus viz.,
M. quernii (Moniez, 1891) Yamaguti, 1938 in having a two branched vitellarium and
a large Laurer’s canal.
Genus Metanematobothrioides Yamaguti, 1965
Diagnosis: Body slender, very long. Oral sucker well developed. Ventral sucker
distinct, some distance from intestinal bifurcation. Pharynx small. Caeca surrounded
by gland cells at beginning, terminate at posterior extremity. Testes paired, tubular,
sinuous, extend almost entire length of body. Genital pore ventral to oral sucker.
Ovary filiform, winding, unbranched, originating some distance behind distal ends
of testes, and descend to genital junction. Uterus first descending, turning anteriorly
at posterior extremity. Eggs elliptical, small. Vitellarium single, filiform, extends
between ovary and posterior extremity. Type species: M. opakapaka Yamaguti, 1965.
Metanematobothrioides branchialis Madhavi, 1982 (Fig. 9.63)

Host: Paridae: Pristipomoides typus Bleeker
Site: Muscles of branchial region
No. 4
Locality: VSK, BOB
Description: Body long, ribbon-like, 285 mm long 960 wide. Oral sucker ellipti-
cal, highly muscular, appearing as plug-like structure with terminal mouth opening
400 × 285 I size. Ventral sucker very small, 58 × 62, situated close behind oral sucker.
Pharynx rudimentary. Oesophagus long, surrounded by thickly arranged gland cells
Intestinal bifurcation around midway between suckers. Caeca narrow, extend to pos-
terior end of body. Testes 2, cylindrical, originate at different levels anterior to genital
junction. Genital pore ventral to oral sucker. Genital junction at midbody dividing
the body in ratio of 1:1.3. Ovary cylindrical, extends anteriorly from genital junction,
partly intrudes into testicular zone. Seminal receptacle absent. Vitellarium single, nar-
row, thread-like extends from genital junction to near posterior end of body. Uterus
with only one loop, descends first to posterior end. Metraterm well developed. Eggs
20 × 8–10.
Fig. 9.63 Metanematobothrioides

branchialis (after Madhavi,
Remarks: Yamaguti (1965) erected the genus Metanematobothrioides for

nematobothrium-like forms with paired testes, a single ovary, a single vitellarium
and a single uterine loop. M. branchialis differs from the type species.
Genus Neonematobothrium Yamaguti, 1965
Diagnosis: Body long, slender. Oral sucker and pharynx rudimentary. Oesophagus
unusually long, with numerous diverticles surrounded by gland cells, bifurcates in
middle region of body into unequal branches each of which passes into a narrow con-
voluting caecum terminating near posterior extremity. Ventral sucker absent. Testes
narrow, twisted, parallel to each other for the most part, originating at different levels.
Genital pore post pharyngeal. Ovary single, tubular, unbranched. Seminal receptacle
present. Uterus first descending to near posterior end of body, then ascending and
turning backward a short distance from anterior extremity forming another U-turn
near posterior extremity before taking its final ascending course. Eggs small. Vitellar-
ium single, narrow, tubular, extending between ovary and posterior extremity. Type
species N. kawakawa Yamaguti, 1965.
Remarks: This genus is characterized as follows: Oesophagus with branches and

surrounded by gland cells, uterus with three U-turns, intestinal caeca very narrow
and strongly convoluted, terminate near posterior extremity, ventral sucker absent.
Neonematobothrium kawakawa Yamaguti, 1965
Site: Free in subcutaneous tissue of opercular region
Locality: VSK, BOB
Reference: Madhavi & Sairam (2000), Yamaguti (1965)
Distribution: Hawaii, VSK, BOB
Description: Body flattened, slender, 32–48 mm long by 370–650 wide. Oral sucker
and pharynx rudimentary. Oesophagus simple, slender anteriorly, but profusely diver-
ticulate on each side and surrounded by numerous gland cells for its greater posterior
portion. Caeca long unequal, strongly twisted, narrow, tubular, terminate near poste-
rior extremity. Ventral sucker absent. Testes two, narrow, strongly twisted, parallel to
each other for most part but originate at different levels. Vas deferens winds forward
in median field close to final ascending limb of uterus. Ovary narrow, twisted, origi-
nates 2.0–3.7 mm posterior to anterior end, terminates at a distance of 20.5 mm from
anterior extremity. Seminal receptacle oval, situated dorsal to genital junction. Gen-
ital junction posterior to midbody. Uterus forms three loops, one at a short distance
from head end and two near posterior extremity. Distal portion of uterus differenti-
ated into metraterm, which runs forward nearly in median field long with vas deferns.
Eggs 14–21 × 7.5–12. Vitellarium narrow, tubular, strongly twisted throughout. Eggs
14–21 by 7.512.
Neonematobothrium dorsale Yamaguti 1970

Site: Free in subcutaneous tissue of dorsal body folds flanking dorsal fin
Distribution: Hawaii, Bay of Bengal

Remarks: This species differs from N. kawakawa in lodging in a special location.
Genus Oculonematobothrium Murugesh, Sairam& Madhavi, 1992
Diagnosis: Hermaphroditic. Body ribbon-shaped. Oral sucker small. Pharynx rudi-

mentary or absent. Ventral sucker absent. Caeca lined with thick zone of gland cells
initially, getting simple later on, extend to posterior extremity. Genital complex just
anterior to midbody. Testis single in anterior third of body, terminates anterior to
genital junction. Genital pore near oral aperture. Ovary single, tubular, anteriorly
directed. Vitellarium single extends between genital junction and posterior extrem-
ity. Uterus with four unequal arms and three loops, first descending and running
backwards. Eggs oval, unipolar. Occur tangled together in orbital cavity of marine
teleosts. Type species: O. orbitum Murugesh, Madhavi & Sairam, 1992.
Oculonematobothrium orbitum Murugesh, Sairam & Madhavi, 1992 (Fig. 9.64)

Host: Stromateidae: Pampus argenteus Euphrasenite: Orbital cavity
Locality: VSK, BOB
Reference: Murugesh et al., (1992)
Description: Body ribbon-like, 6.4–7.8 mm long, 432–688 wide. Oral sucker coni-
cal, 44 × 46 by 35–36 in size. Ventral sucker absent. Pharynx rudimentary. Oesoph-
agus long, leads into pouch. Caeca long, initial portion covered with thick zone of
gland cells, rest of caeca simple, thrown into folds, terminate near posterior end of
Fig. 9.64 Oculonematobothrium

orbitum (after Murugesh,
Sairam & Madhavi, 1992)
body. Testis single occupies anterior third of body, terminates anterior to genital junc-
tion. Ovary single, narrow, tubular winds in anterior third of body. Genital junction
located in posterior part of anterior half of body. Seminal receptacle not clear. Uterus
with two long and two short arms and three loops, descending loop with thick zone
of gland cells, first and third loops lie close to posterior end of body, second loop at
16–20 mm from anterior tip. Metraterm well developed. Eggs filamented, 32–34 by
8–12. Egg filament long, Vitellarium single, tubular, strongly winding originates at
variable distance from posterior end, terminates at genital junction.
Remarks: The distinguishing features of the genus Oculonematobothrium are the

complete absence of a pharynx and ventral sucker, the single testis, the uterus with
three loops and filamented eggs. So far O. orbitum is the only species reported under
the genus.
Genus Skrjabinozoum Nikolaeva & Parukhin, 1969
Diagnosis: Not encapsulated. Body elongate, filiform. Pharynx present. Oesoph-

agus short. Caeca reach close to posterior extremity. Testes elongate, parallel or
interwoven in anterior third of body. Vas deferens short. Ejaculatory duct long. Gen-
ital pore ventrolateral to oral sucker. Genital junction at about middle of body. Ovary
and vitellarium long, unbranched, in anterior and posterior half of body, respec-
tively. Seminal receptacle present or absent. Type species: S. exocoetidi Nikolaeva
& Parukhin, 1969
Skrjabinozoum waltairense Hussain, Rao & Shyamasundari, 1984 (Fig. 9.65)

Host: Ariommatidae: Ariomma indica (Day) (= Psenes indicus (Day)
Site: Cysts in the connective tissue of operculum and orbit.
Locality: VSK, BOB
Number: 8
Description: Body elongate, not differentiated into fore and hindbodies, 2030–9900
long and 550–1370 wide. Testes well developed in younger individuals and female
reproductive organ not differentiated. A minute male individual occurs together with
Fig. 9.65 Skrjabinozoum

waltairense (Hussain, Rao &
Shyamasundari, 1984)
a fully developed hermaphroditic individual inside each cyst. Oral sucker muscular,
round. Pharynx absent. Oesophagus long, narrow, with layer of gland cells. Caeca
simple. Ventral sucker absent. Testes two, juxtaposed, tubular, slightly branched
situated near anterior end of body. Ovary single, tubular, strongly winding originates
260–290 below the testes, extends up to Mehlis’ gland complex. Genital junction
1970–3210 from anterior end. Seminal receptacle small. Uterus runs forward as a
complicated tube close to ovary near posterior end of forebody and turns back on
itself forming a loop anteriorly. It descends to posterior extremity close to vitellarium
to take its final ascending course forming two loops one anterior and one posterior,
terminates below oral sucker as metraterm. Eggs 7 x 2.
Remarks: S. waltairense differs from the type species S. exocoetidi Nikolaeva &
Parukhin, 1974 in the size of oral sucker, oesophagus with gland cells on either side,
branched testes and in the egg size.
Skrjabinozoum yamagutii Gupta & Gupta, 1986 (Fig. 9.66)

Host: Nomeidae: Psenes cyanophrys (Val.)
Locality: PR, BOB
No. 1
Reference: Gupta, P.C. & Gupta, V.C. (1986a)
Remarks: This species differs from S. waltairense in the larger body size 15.83 ×
1.27 mm, position of testes far behind the intestinal bifurcation instead of close to it,
ovary not extending anteriorly and in the shape of body.
Fig. 9.66 Skrjabinozoum

Gupta, 1986)
Fig. 9.67 Plate 1. Didymozoid trematodes from tunas of Bay of Bengal
Family Hirudinellidae Dollfus, 1932
[Syns. Botulidae Guiart, 1938; Lampritrematidae Yamaguti, 1941; Mediolecithidae

Oshmarin, 1968; Profundiellinae Skrjabin, 1958]
Hirudinellidae are very large hemiuroids occurring usually in the stomach of large
marine teleosts. They are unique among hemiuroids in that they lack a hermaphroditic
duct instead possess a cirrus-sac and cirrus which however, are not homologous to
those of other digeneans.
Family Hirudinellidae Dollfus, 1932 237
Diagnosis: Body large, highly muscular, ecsoma absent. Body surface smooth. Oral
and ventral suckers well developed. Gut caeca terminate blindly or form uroproct.
Testes two, oblique or tandem in hindbody. Seminal vesicle tubular, sinuous. Pars
prostatica well developed. Ejaculatory duct long, surrounded by cirrus-sac, opens into
genital atrium through cirrus. Genital pore median in forebody. Ovary post-testicular.
Seminal receptacle uterine. Uterine coils occupy most of hindbody. Vitellarium com-
posed of numerous branched tubules, mainly postovarian. Eggs Y-shaped.
Type genus Hirudinella Blainvile, 1828.

Genus Hirudinella Blainvile, 1828.
Diagnosis: With characters of the family. Type species: H. ventricosa (Pallas, 1774)
Hirudinella ventricosa (Pallas, 1774) Baird, 1859 (Fig. 9.68)

[Syn. Uroproctinella attenuata Hafeezullah, 1971; U. cochinensis Sahai & Srivas-
tava, 1978 (Ref. Gibson & Bray, 1977; Madhavi, 2011)]
Hosts: Scombridae: Scomberomorus commersoni Lacepede (1); Euthynnus affinis
Canttor (2); Thunnus albacores (Bonnaterre) (3). Carangidae: Scomberoides tala
Cuvier (4)
Locality: VSK, BOB (1,2); Veeraval, AS (3); Cochin, AS (4)
Reference: 1. Murugesh & Madhavi (1990); 2. Madhavi & Sairam (2000); 3.
Hafeezuhllah (1971b); 4. Sahai & Srivastava (1978b)
Description: Body very large, 43 mm × 5 mm, attenuated behind ventral sucker.

Tegument very thick, smooth. Oral sucker 1411 × 1176 in size. Ventral sucker 2020
by 2446. Sucker ratio 1:2. Pharynx ear shaped, overlapped by oral sucker. Oesoph-
agus short. Caeca long, sinuous, extend to posterior end of body and open into
excretory vesicle forming uroproct. Testes oval, slightly diagonal, posterior to ven-
tral sucker. Seminal vesicle in forebody. Pars prostatica long, winding, surrounded
Fig. 9.68 Hirudinella

ventricosa (after Gibson &
Bray 1977)
by thin layer of prostatic gland cells. Ejaculatory duct enters genital bulb containing
male and female ducts, both of them opening ventrally into tubular genital atrium.
Genital pore post-bifurcal. Ovary subglobular, well removed posteriorly from testes.
Seminal receptacle absent. Uterus extends posteriorly up to attenuated part of body.
Uterine seminal receptacle prominent. Eggs 15–24 × 12–17. Vitellarium branched,
narrow, very long, highly convoluted tubules extend from anterior level of anterior
testis to attenuated part of body. Excretory vesicle Y-shaped, stem short, arms thin
thrown into complicated coils, meet dorsal to oral sucker. Excretory pore terminal.
Remarks: Gibson (1976) and Gibson and Bray (1979) discussed the history, syn-
onyms, hosts and geographic distribution of H. ventricosa. They considered the
genus Hirudinella as monospecific and synonymized all the species reported under
this and the genus Uroproctinella with H. ventricosa. The Indian reports of the genus
include those of Raju (1962) in several scombrid fishes at Tuticorin and Minicoy;
Hafeezullah (1971b) from Thunnus albacores from Veeraval, AS; Murugesh & Mad-
havi (1990) from Scomberomorus commersoni from VSK, BOB; Madhavi & Sairam
(2000). from Euthynnus affinis from VSK, BOB; Sahai & Srivastava (1978b) from
Scomberoides tala from, Cochin, AS. Murugesh & Madhavi (1990) described the
miracidium of this species.
Family Lecithasteridae Odhner, 1905
Lecithasterids are primarily parasites in the gut of marine fishes. Initially, they were
included in a subfamily under Hemiuridae Looss, 1899 as Lecithasterinae Odhner,
1905. It was given a full family status by Skrjabin & Guschankaja (1957) and this
concept was accepted by Gibson & Bray (1979) in the review on Hemiuriodea.
Members of this family have an oval body, lack an ecsoma and the vitellarium
consists of six to eight oval or digitiform lobes arranged in a rosette. Molecular
results indicate that the Lecithasteridae is not distinct from the Hemiuridae.
Diagnosis: Body oval, ecsoma absent. Tegument smooth. Oral and ventral suck-
ers well developed. Caeca usually terminate blindly, occasionally unite forming
cyclocoel. Testes two, tandem, oblique or symmetrical in hindbody. Seminal vesicle
oval, tubular or constricted into portions, in forebody or hindbody. Pars prostat-
ica usually tubular occasionally vesicular. Ejaculatory duct long or short or absent.
Hermaphroditic duct present. Permanent sinus organ absent but hermaphroditic duct
often protrudes to form sinus organ. Ovary post-testicular, oval or 4-lobed. Blind
seminal receptacle or Juel’s organ present, large, thick-walled, situated anterodorsal
to ovary. Uterus varies from mainly postovarian to entirely preovarian. Eggs numer-
ous, small. Vitellarium commonly seven-lobed, often in rosette arrangement, usually
immediately postovarian, occasionally preovarian or at level of ovary.
Family Lecithasteridae Odhner, 1905 239
Type genus Lecithaster. Luhe, 1901.

Some genera of lecithasterines erected to include species from Indian marine fish
have been invalidated: Bilqeesotrema Gupta & Jain, 1991, based on a single speci-
men was considered a synonym of Aponurus Looss, 1901 and Anadichodena Dutta,
1995 was synonymized with Lecithaster Luhe, 1901 (Gibson, 2005). Another genus
Cryptodiscus Srivastava & Sahai, 1998 was synonymized with Prolecitha Manter,
1961 by Abdul-Salam & Khalil (1987). The family is now represented in Indian
marine fish by 14 species, spread over genera belonging to four subfamilies.
Key to subfamilies
1. Uterine seminal receptacle present; Juel’s organ present …………………….

………………………………………………….. Hysterolecithinae Yamaguti,
1958
Uterine seminal receptacle absent, blind seminal receptacle present ……....2
2. Uterus entirely preovarian, sinus-sac weak, membranous
…………………………..…. Trifoliovariiinae Yamaguti, 1958
Uterus partly postovarian, sinus-sac well developed ……………………. 3
3. Caeca terminate blindly, genital atrium not sucker-like, vitellarium postovarian
…………………………………………………………………. Lecithasterinae
Luhe, 1901
Caeca form cyclocoel, genital atrium sucker-like, vitellarium preovarian
……………………………………..……………………….. Prolecithinae Yam-
aguti, 1971
Subfamily: Lecithasterinae Luhe, 1901
Diagnosis: Body small, oval. Caeca terminate blindly. Testes two. Seminal vesicle
in forebody. Pars prostatica short. Ejaculatory duct short or absent. Sinus-sac well
developed. Sinus organ absent. Ovary entire or four-lobed. Seminal receptacle large.
Uterus reaches postovarian region. Vitellarium seven oval to digitiform lobes in
rosette, immediately postovarian. Excretory arms united in forebody or not.
Type genus: Lecithaster Luhe, 1901

Key to genera
Ovary lobed, vitelline lobes petal-shaped……………………………………..…….

Lecithaster Luhe, 1901
Ovary globular, vitelline lobes globular………………………………………….….
Aponurus Looss, 1907
Genus Aponurus Looss, 1907
[Syn: Brachadena Linton, 1910; Bilqeesotrema Gupta & Jain, 1991]
Diagnosis: Body oval. Testes two, tandem to oblique. Seminal vesicle saccular, in
forebody. Pars prostatica well developed. Hermaphroditic duct tubular. Sinus-sac
oval to elongate. Ovary oval, seminal receptacle small to large. Vitellarium with
seven globular to elongate lobes arranged in two groups of three and four, lying
lateral and posterior to ovary. Excretory arms united anteriorly.
Type species: A. lagunculus Looss, 1907
Seven species belonging to this genus have been recorded from marine fishes of
India: A. breviformis Srivastava 1939, A. bengalensis Srivastava, 1939, A. drepan
Hafeezullah, 1990 A. laguncula Looss, 1907, A. orientalis Ahmad, 1981 and A. wal-
tairensis. Bilqeesotrema Gupta & Jain 1991 was synonymized with Aponurus and
its single species became A. theraponi (Gupta & Jain, 1991) Madhavi, 2011. All
these seven species appear similar in morphology and three of them A. breviformis,
A. theraponi and A. orientalis were recorded from Terapon spp. Bray & Macken-
zie (1990) expressed the view that many of the species reported under the genus
would fall synonyms of A. laguncula the type species. They examined the validity
of the various species reported under the genus and discussed the characters used
for separating them from A. laguncula. The most significant features for separa-
tion of species are considered to be the size of sinus-sac, sucker proportions and
the alignment of gonads. Two of the Indian species namely A. waltairensis and A.
drepanei have already been synonymized with A. laguncula. The remaining species
were not well differentiated from A. laguncula and may eventually fall synonyms of
A. laguncula. The reported occurrence of cryptic species among A. laguncula com-
plex reported from Mediterranean (Carreras-Aubets et al. 2011) indicates the need
to exercise caution for taking any decision regarding the validity of various Indian
species of the genus. In this account the full description is given only for A. laguncula
whose validity is unquestionable, and all the remaining species are tentatively kept
as species inquirendum.
Aponurus bengalensis Srivastava, 1939 (Fig. 9.69)
Host: Ariidae: Plicofollis dussumieri (Val.)
Locality: PR, BOB
Reference: Srivastava (1939)
Status: Species inquirendum
Remarks: According to Bray and Mackenzie (1990b), A. bengalensis is similar to

A. laguncula but has symmetrical testis and a smaller sinus-sac.
Aponurus breviformis Srivastava, 1939 (Fig. 9.70)
Host: Terapontidae: Terapon puta (Cuv.) (1); Pristigasteridae: Ilisha elongata (2),
Ilisha filigera, (Val.) H. ilisha (Hamilton Buchanan) (3); Carangidae: Parastromateus
niger Bloch (3)
Fig. 9.69 Aponurus

bengalensis (after Srivastava,
1969)
Fig. 9.70 Aponurus

breviformis (after Srivastava,
1939)
Locality: PR, BOB (1); Hughli Estuary (2); Digha coast (3)
Reference: 1. Srivastava (1939); Zhukov (1977); Hafeezullah & Dutta (1995)
Description: Body small, elongated 830–1060 long, 160–220 wide, Tegument

smooth. Ventral sucker pre-equatorial. Oral sucker smaller than ventral sucker. Caeca
extend to posterior end of body. Testes two, situated in middle third of body, tan-
dem. Seminal vesicle saccular, in forebody, pars prostatica long, tubular, surrounded
by prostatic cells, hermaphroditic duct, short, enclosed in sinus-sac. Genital pore
immediately posterior to intestinal bifurcation. Ovary globular, post-testicular. Sem-

inal receptacle present. Vitellarium consists of seven lobes posterior to ovary. Uterus
posterior to seminal vesicle. Eggs 20–23 by 12.
Remarks: This species is characterized by the small size of eggs measuring 20–23
by12.
Aponurus drepani Hafeezullah, 1990 (Fig. 9.71)

[Syn. A. laguncula Looss, 1907]
Host: Family Drepanidae: Drepane punctata (L.)
Locality: Tuicorin (GOM)
No. 4
Description: Body 1140–1610 long, 290–420 wide. Tegument moderately thick,

smooth. Ventral sucker 192–225 long, 203–225 wide. Oral sucker 87–118 long,
91–115 wide. Sucker width ratio about 1:2.1. Caeca simple, reaching almost to pos-
terior end of body. Testes 2, globular, oblique with dextral anteriormost, equatorial.
Seminal vesicle pyriform, anterodorsal to ventral sucker, placed obliquely. Pars pro-
statica long, curved, surrounded by well-developed prostate cells. Sinus-sac oval,
muscular, extending posteriorly to slightly beyond intestinal bifurcation, opening at
base of oral sucker, encloses hermaphroditic duct. Ovary globular. Seminal recep-
tacle dorsal to posterior testis. Vitellarium comprises 7 rounded lobes, in stellate
fashion, all lobes separate, posterior to ovary. Uterus voluminous, extends to poste-
rior extremity; metraterm present. Eggs 21–25 × 7–14.
Remarks: According to Hafeezullah (1990b) this species resembles A. breviformis

in sucker ratio and egg size, but differs from it in having a more anterior genital pore,
the pars prostatica curved and longer than the seminal vesicle and in the position
Fig. 9.71 Aponurus drepani

and size of the seminal receptacle. From A. orientalis it differs in the prebifurcal
position of the genital pore and in having a pars prostatica curved and longer than the
seminal vesicle and in the much smaller eggs. Bray & Mackenzie (1990b) regarded
the species as a synonym of A. laguncula.
Aponurus laguncula Looss, 1907 (Fig. 9.72)

[Syn. A. waltairensis Hussain, Rao & Shyamasundari, 1984: A. drepani Hafeezullah,
1990)]
Host: Scombridae: Rastrelliger kanagurta (1); Acanthuridae: Acanthurus mata
(Cuvier); Drepanidae: Drepane punctata (L.) (2)
Locality: VSK, BOB
Reference: 1. Madhavi & Trivenilakshami (2011), Hussain, Rao & Shyamasundari
(1984)
Description: Body 500–705 × 167–242. Tegument unarmed. Oral sucker 68–81 ×

68–85. Ventral sucker immediately anterior to midbody, 123–151 × 126–144. Sucker
ratio 1:1.57–1.85. Caecal bifurcation at midway between suckers. Caeca long, ter-
minate blindly near posterior end of body. Testes ovoid, oblique, in anterior hind-
body, Seminal vesicle subglobular, in posterior forebody. Pars prostatica narrow,
curved, with relatively few gland cells. Sinus-sac thin-walled, contains fibrous con-
nective tissue and scattered gland cell. Genital pore median, in anterior forebody.
Ovary oval to subtriangular, postero-lateral to posterior testis. Seminal receptacle
lies between ovary and posterior testis. Uterus fills most of hindbody. Eggs numer-
ous, 32–42 × 16–21. Vitellarium consists of seven irregular follicles immediately
posterior to ovary.
Remarks: Bray and Mackenzie (1990b) furnished details of the hosts, geographic
distribution and earlier reports of A. laguncula. This species is highly euryxenic and
has a wide geographic distribution. A notable feature of the species is the wide range
Fig. 9.72 Aponurus

laguncula (original)
in egg size, 32–42 × 16–21. The egg shape is also considered distinctive with narrow
more or less pointed anopercular end. Madhavi & Trivenilakshmi (2011) reported
A. laguncula from Rastrelliger kanagurta and stated that infection with the species
has also been noted in other marine fish like carangids, lutjanids, polynemids, Arius
sp. Therapon spp and Nemipterus sp. confirming the euryxenic nature of the species.
However, in view of the morphological variations that occur in different specimens
of A. laguncula especially in the egg size and shape, the occurrence of cryptic species
cannot be ruled out.
Aponurus orientalis Ahmad, 1981 (Fig. 9.73)

Locality: PR, BOB
Description: Body 1000–1235 long by 220–235 wide. Oral sucker 66–74 long
by 88–96 wide. Ventral sucker spherical, pre-equatorial, 163–195 in diameter, at
242–256 from anterior end of body. Sucker ratio 1:2.09–2.12. Intestinal bifurcation
dorsal, nearer to oral sucker than to the ventral sucker. Caeca simple, in lateral fields,
terminating close to posterior extremity. Testes directly tandem, at 438–472 from
anterior end of body. Seminal vesicle retort-shaped, in forebody, Pars prostatica short,
surrounded by numerous prostate gland cells. Sinus-sac enclosing hermaphroditic
duct. Genital pore at intestinal bifurcation. Ovary postequatorial, larger than the
testes. Vitellarium postovarian, consisting of 7 rounded follicles. Uterus running
between ventral sucker and caecal ends. Eggs 30–35 long by 17–19 wide.
Fig. 9.73 Aponurus

orientalis (after Ahmad,
1981)
Remarks: A. orientalis resembles A. laguncula Looss, 1907 but differs from it in

having a retort-shaped seminal vesicle, a very short pars prostatica, a smaller and
subglobular sinus-sac, the testes separated from the ventral sucker, the ovary larger
than the testes and the uterus extending to the caecal ends and from A. breviformis in
having the genital pore at the intestinal bifurcation, the ovary larger than the testes
and the comparatively large ventral sucker.
Aponurus theraponi (Gupta & Jain, 1991) Madhavi, 2011

[Syn. Bilqeesotrema theraponi Gupta & Jain}]
Host: Teraponidae: Terapon puta (Cuv.)
Locality: PR, BOB
Reference: Gupta V & Jain 1991
Status: Species inquirenda
Bilqeesotrema Gupta & Jain, 1991 is similar to Aponurus Looss, 1907 differ-
ing according to the authors only in having symmetrical instead of diagonal testes.
Madhavi (2011) synonymized the genus with Aponurus and B. theraponi Gupta &
Jain, 1991 became A. theraponi. Detailed study of A. theraponi may prove it to be a
synonym of A. orientalis Ahmad, 1981 both sharing the same host genus and locality.
Genus Lecithaster Luhe, 1901

[Syn. Anadichadena Dutta, 1995]
Diagnosis: Body oval or elongate. Testes two, obliquely symmetrical, oval or lobed.
Seminal vesicle saccular to elongate, sinuous, in forebody or dorsal to ventral sucker.
Sinus-sac oval. Genital atrium short. Ovary four-lobed. Seminal receptacle saccular,
large, dorsal to ovary. Vitellarium with seven radiating tear-shaped or globular lobes.
Excretory arms not united anteriorly. Type species: T. confusus Odhner, 1905
Four species of Lecithaster have been recorded from India, two of them L.
extralobus Srivastava, 1935 and L. indicus Srivastava, 1935 are from fresh water
fishes and the remaining two L. bombayensis Ahmad, 1983 and L. chrysophrysi
Gupta & Ahmad, 1976 are from marine fishes. Another species L. binoyi (Dutta,
1995) (Syn. Anadichadena binoyi Dutta, 1995) was regarded as species inquiren-
dum.
Lecithaster bombayensis Ahmad, 1983 (Fig. 9.74)

Host: Paralichthyidae: Pseudorhombus diacanthus (Bloch.)
Locality: BOM, AS
Number: 24 specimens
Description: Body 2650–2735 long. 140–792 wide. Oral sucker 130–117 in diame-
ter. Ventral sucker 261–284 long. 290–334 wide, situated in posterior part of anterior
third of body. Sucker width ratio 1:1.88–2.23. Intestinal bifurcation dorsal, mid-
way between suckers. Caeca simple, long. Testes two, deeply four-lobed diagonally
Fig. 9.74 Lecithaster

bombayensis (after Ahmad,
1983)
tandem, anterior testis immediately posterior to ventral sucker. Seminal vesicle sac-
cular, posterior to ventral sucker. Pars prostatica tubular, slightly sinuous, overlaps
right edge of ventral sucker, surrounded by numerous prostate gland cells. Sinus-
sac saccular, lies anterior to ventral sucker, encloses hermaphroditic duct. Genital
pore ventral, median, immediately post-bifurcal. Ovary deeply four-lobed, lobes with
rounded ends. Seminal receptacle oval, postovarian. Vitellarium with seven claviform
lobes, postovarian. Uterus descends posteriorly as far as posterior edge of vitellar-
ium, then ascends anteriorly to open at base of sinus organ. Eggs 38–42 long, 20–24
wide.
Remarks: This species differs from all the species of Lecithaster, excepting L. testilo-
batus Manter, 1969 in having deeply lobed testes. L. testilobatus differs from this
species in having symmetrical testes lying at the posterior edge of the ventral sucker,
the seminal vesicle lying dorsal to the ventral sucker and the ovary and vitellarium
lying in middle third of the body.
Lecithaster chrysophrysi Gupta & Ahmad, 1976 (Fig. 9.75)
Host: Sparidae: Rhabdosargus sarba (Forsskal)
Locality: PR, BOB
Number: One specimen
Description: Body elongate, 2540 long, 214 wide. Oral sucker subspherical, sub-
terminal, 110 long, 178 wide. Pharynx ovoid. Intestinal caeca simple, long. Ventral
sucker cup-shaped, smaller than oral sucker, pre-equatorial, 163 long, 115 wide.
Genital pore on lateral side near intestinal bifurcation. Testes entire, subspherical,
tandem, contiguous, well separated from ventral sucker. Seminal vesicle saccular,
extends up to middle of anterior testis. Pars prostatica tubular, sinuous, surrounded
by large number of prostate gland cells. Hermaphroditic duct tubular, enclosed in
oval hermaphroditic pouch. Ovary tetra-lobed, post-testicular, postequatorial. Sem-
inal receptacle on right side of ovary. Vitellarium consists of seven rosette-shaped
Fig. 9.75 Lecithaster

chrysophrysi (after Gupta &
Ahmad, 1976)
lobes, postovarian. Uterus extends up to posterior end of body, anteriorly opens at

base of hermaphroditic pouch. Eggs oval, 38–45 long 21–23 wide.
Remarks: This species differs from all the known species of the genus in having
the ventral sucker smaller than the oral sucker, tandem and contiguous testes and
in having the lateral, bifurcal genital pore. This latter character is most unusual for
hemiuroids and is probably an artefact of fixation or flattening. L. bombayensis differs
from L. chrysophrysi in the oval shape of the body, the deeply lobed and obliquely
arranged testes, in the location of the ovary and vitellarium close to the posterior
extremity and in the ventral sucker being much larger than the oral sucker.
Subfamily Hysterolecithinae Yamaguti, 1958
Diagnosis: Body small, elongate. Ventral sucker in middle or anterior half of body.
Caeca end blindly near posterior extremity. Testes two, oval, symmetrical or obliquely
tandem, preovarian. Seminal vesicle tubular or saccular, in forebody. Pars prostat-
ica tubular or vesicular, short. Sinus-sac present. Permanent sinus organ absent.
Hermaphroditic duct present within sinus-sac. Ovary oval, in anterior or posterior
half of hindbody. Juel’s organ and uterine seminal receptacle present. Uterus almost
entirely in hindbody. Vitellarium seven oval to digitiform lobes, postovarian.
Type genus Hysterolecitha Linton, 1910.
The systematic position of the Hysterolecithinae within Lecithasteridae is yet
to be proved. Morphologically they fit well into the family but molecular studies
of Blair et al. (1998) indicated no relationship. Tentatively they are retained under
Lecithasteridae.
Key to genera
Eggs may be filamented, seven digitiform vitelline lobes ……………….. Thulinia

Gibson & Bray, 1979
Eggs never filamented, vitellarium 2–7 entire or irregular masses..…. Hys-
terolecithoides Yamaguti, 1934
Genus Hysterolecithoides Yamaguti, 1934
Diagnosis: Body fusiform to oval. Tegument unarmed. Ventral sucker in about mid-
dle of body. Caeca reach towads posterior extremity, blind. Testes 2, subcylindrial,
symmetrical. Seminal vesicle elongate, tubular, mainly in posterior forebody. Pars
prostatica wih sheath of large gland cells. Sinus-sac oval, genital atrium distinct with
blind sac projecting antero-dorsally. Genital pore median, about midwy between
suckers. Ovary oval, post-testicular or overlapping testes. Juel’s organ present. Uter-
ine seminl receptacle present. Bulk of uterus post-ovarian. Vitellarium 2–7 subglob-
ular masses, entire or irregular, immediatley post-ovaria. Excretory arms end blindly
at level of oral sucker. Type species: H. epinepheli Yamaguti, 1934
Hysterolecithoides frontilatus, (Manter,1969) Yamaguti, 1971 (Fig. 9.76)
[ Syn. Theletrum frontilatum Manter, 1969; Neotheletrum frontilatum (manter, 1969)

Gibson & Bray, 1979; Machidatrema fontilatum (Manter, 1969) Leon-Regagnon,
1998]
Hosts: Siganidae: Siganus canaliculatus (Park); S. javus (L.)
Locality: Andaman Isles
Distribution: Indo-Pacific (Bay of Bengal), Moreton Bay, Queensland, New Caledo-
nia, Lizard island, Great Barrier Reef
Fig. 9.76 Hysterolecithoides

fontilatus (after Hafeezullah
& Dutta, 1980, Rec. Zool.
Surv. India, 77, p. 76)
Reference: Hafeezullah & Dutta (1980); Hafeezullah (1990b)

Description: Body 4211 to 6077 long, 1278 to 1938 wide. Oral sucker 357–440
long, 343–398 wide. ventral sucker 825-1045 long, 728-866 wide. Sucker ratio 1:
1.8 to 2.28. Caeca simple reaching posterior end of body. Testes two, ovoid, sym-
metrical, immediately behind ventral sucker. Seminal vesicle tubular, long, coiled
posteriorly reaching ventral sucker. Pars prostatica long, winding, surrounded by
numerous prostatic cells. Sinus-sac muscular, pear-shaped, encloses straight tube of
sinus duct, and filled with gland cells. Genital pore midway between the two suck-
ers. Genital arium short. Ovary post-testicular. Vitelline lobes three, situated between
poserior end of body and ventral sucker. Uterus coiled, confined to hindbody. Eggs
21-28 by 10-14. Excretory vesicle I-shaped, dividing near anterior border of venral
sucker, branches not uniting dorsal to ventral sucker.
Remarks: This species was originally described as Theletrum frontilatum Man-
ter, 1969. Yamaguti plaed this species in Hysterolecithoides which was accepted
by Hafeezullah & Dutta (1980) and Hafeezullah (1990). Machida (1989) erected
another genus Neotheletrum with N. akeh Machida as the type species. Leon-Regagon
et al (1998) included both the species, N. frontilatus and N. akeh, in a new genus
Machidtrema which is close to Hysterolecithoides but differs in the lack of a blind
sac projecting from genital atrium, the tandem testes the coiling of uterus between
the testes and the ovary. In a later study Bray & Cribb (2000) who discussed in
detail the hysterolecithasterine and bunocotyline genera retained the original status
as Hysterolecithoides frontilatus. H. frontilatus is mainly a parasite of Siganus spp.
Important diagnostic featues of the species are: preovarin uterus, testes overlapping
the venral sucker, blind excretory arms, presence of a Juel’s organ, vitellarium divdied
into three masses, distal part of pars prostatica loops dorsal to the sinus-sac.
Hysterolecithoides yamagutii Gupta & Dwivedi (2006) (Fig. 9.77)

Host: Serranidae: Epinephelus flavocaeruleus (Lacepede)
Number : 2
Reference: Gupta, V & Dwivedi 2006)
Description: Body elongate, large, 3200 to 4360 long, numerous dermal glands
present in the anterior half of body. Oral sucker subterminal, prepharynx absent.
Caeca simple reach to posterior extremity. Ventral sucker large, in middle third of
body, 550–650 by 580–680 in size. Testes symmetrical, far apart from each other.
Seminal vesicle tubular, winding in front of ventral sucker. Pars prostatica long, sur-
rounded by numerous prostatic ells. Hermaphroditic duct small. Genital atrium pear
shaped. Genital pore median, a little posterior to intestinal bifurcation. Ovary oval,
post-testicular. Seminal receptcle large. Uterus highly coiled, largely in hindbody.
Eggs 20–30 by 40–50 in size. Vitellarium with 3 lobes, immediatley postovarian.
Remarks: This species differs from all the other species in the genus in the presence
of dermal gland cells in forebody.
Fig. 9.77 Hysterolecithoides

Dwivedi, 2006)
Genus Thulinia Gibson & Bray, 1977
Diagnosis: Body elongate. Ventral sucker in anterior half of body. Caeca end blindly
near posterior extremity. Seminal vesicle saccular or tubular. Sinus-sac well devel-
oped, sinus organ present. Ovary in posterior half of hindbody, separated from testes
by uterine loops. Vitellarium seven digitiform loops. Excretory arms united in fore-
body. Type species: T. microrchis (Yamaguti, 1934). Only one species of this genus
is reported in Indian waters, under the name Hysterolecitha sigani Mnter, 1969.
Thulinia microrchis (Yamaguti, 1934) Bray, Cribb & Barker, 1993 (Fig. 9.78)
[Syn. Hysterolecitha microrchis Yamguti, 1934; Hystrolecitha sigani Manter, 1969;

Hysterolecitha xesuri yamaguti, 1938; Hysterolecitha tinkeri Manter & Pritchard,
1960; Thulinia tinkeri (Manter & Pritchard, 1960) Gibson &Bray, 1979]
Hosts: Ephippidae: Platax tiera Forsskål; Siganidae: Siganus oramin (Scneider);
Sctophagidae: Scatophagus argus (Linnaeus)
Localities: Tuticorin (Gulf of Mannar, Karwar) (AS)
Number of specimens: 6 + 14 + 20 respectively
Distribution: Japan, Hawaii, New Caledonia, Micronesi, Queensland, Australia,

Fiji, and it appears to have reached Mediterranean sea, presumably by Lessepsian
migration (Fischthal, 1980)
Description: Body 1764 to 4200 long, 420 to 876 wide, elongate. Oral sucker
119–216 long, 137–271 wide, ventral sucker 274–516 long 271–492 wide, spherical.
Sucker ratio 1: 1.83 to 2.28. Drusenmagen present near bifurcaton. Caeca reaching
almost posterior end of body. Testes entire or slightly indented diagonal, in hind-
body, separated by uterine coils. Seminal vesicle saccular or tube-like in forebody.
Fig. 9.78 Thulinia

microrchis (after
Hafeezullah, 1990)
Pars prostatica short, surrounded by well developed prostate gland cells. Sinus-sac
wide globlar, large immediatley postbifurcal; hermaphroditic duct wide, appearing
to be divided into three regions, protrusible, opens into genital atrium. Genital pore
wide, immediately post-bifurca. Ovary subglobular near posterior end of body. True
seminal receptacle absent, but uterine seminal receptacle present. Vitellarium digi-
tate, in two masses, right one with 3 lobes and left with 4 lobes, postovarian. Uterus
voluminous, extends short of posterior end of body. eggs 21–24 x 9–12.
Remarks: Bray et al (1993a) synonymized H. sigani with Thulinia microrchis
although they could not detect filaments on the eggs of the type specimens. Per-
sonal observation (RAB) have shown that it is often not possible to detect filaments
in all the specimens of this species. It is now known from 24 host species from
throughout the Indo-Pacific region (Cribb et al. 2014)
Subfamily ProlecithinaeYamaguti, 1971
[Syn. Follicovitellotrematinae Gupta, A. N. and Sharma, 1972]
Diagnosis: Body spindle-shaped. Ventral sucker large, prominent. Cyclocoel
present. Testes two, symmetrical, at level of ventral sucker. Seminal vesicle saccular,
in forebody. Pars prostatica short, vesicular. Terminal genitalia complex. Sinus-sac
present, muscular, asymmetrical, anterior wall forms muscular sucker or papilla-
like structure. Permanent sinus organ absent. Genital atrium sucker-like. Ovary
four-lobed. Large globular seminal receptacle posterior to ovary, close to poste-
rior extremity of body. Uterus mainly anterior to gonads. Vitellarium seven rounded
lobes, immediately preovarian. Excretory arms not united in forebody. Type genus:
Prolecitha Manter, 1961.
Genus Prolecitha Manter, 1961

[Syn. Lobatovitelliovarium Yamaguti, 1965; Follicovitellosum Gupta, A. N. and
Sharma, 1972; Cryptodiscus Srivastava and Sahai, 1978]
Diagnosis: With characters of the subfamily. Type species: P. obesa Manter, 1961
Prolecitha obesa Manter, 1961 (Fig. 9.79)
[Syn. P. beloni Nagaty and Abdel Aal, 1962; Lobatovitelliovarium fusiforme Yam-
aguti, 1965; Follicovitellosum indicum Gupta & Sharma, 1972; Cryptodiscus indicus
Srivastava and Sahai, 1978]
Host: Belonidae: Belone strongylura, needlefish, B. crocodila, giant needlefish
Localities: Rameswaram (Palk Bay) and Tuticorin (Gulf of Mannar), respectively.
No. of specimens: 8 + 1, respectively
Reference: Hafeezullah (1980, 1990b); Abdul-Salam & Khalil (1987)
Description: Body spindle-shaped, 1400–1500 long, 470–560 wide. Oral sucker
150–170 in diameter. Caeca wide, undulating, united posteriorly. Ventral sucker sit-
uated in middle of body, 280–370 × 390–450, eversible. Sucker ratio 1:2.61–2.67.
Testes symmetrical, near posterior margin of ventral sucker. Seminal vesicle oval,
immediately anterior to ventral sucker. Pars prostatica leads into prostatic vesicle,
surrounded by numerous prostatic cells, joins metraterm forming hermaphroditic
duct surrounded by strong sinus-sac. Ovary divided into four oval lobes, near pos-
terior end of body. Vitellarium divided into 7 compact tubes, preovarian, between
ovary and testes. Uterus forms compact transverse loops, joins hermaphroditic duct.
Eggs 16–18 × 9–11.
Fig. 9.79 Prolecitha obesa

(after Hafeezullah, 1980,
Bulletin Zool. Surv. India, 3,
p. 75)
Remarks: Manter (1961) established the genus Prolecitha for P. obesa from a belonid
fish from Hawaii. Manter (1969) regarded P. beloni Nagaty & Abdel-Aal (1962)
as a synonym of P. obesa and synonymized the genus Prolecitha with Dichadena
Linton, 1910. Yamaguti (1971) regarded Prolecitha as distinct from Dichadena.
Abdul–Salam and Khalil (1987) provided a review on P. obesa and listed the species
that have been synonymized with this: P. beloni Nagaty & Abdel Aal (1962),
Lobatovitelliovarium fusiforme Yamaguti, 1965; Follicovitellosum indicum Gupta
& Sharma, 1972; Cryptodiscus indicus Srivastava & Sahai, 1978. From India, P.
obesa has been reported by Hafeezullah (1977, 1980, 1990b).
Subfamily Trifoliovariinae Yamaguti, 1958
Diagnosis: Body elongate. Suckers well developed. Ventral sucker in anterior third
of body. Caeca long, terminate blindly. Testes two, in anterior hindbody, separated
from ovary by uterus. Seminal vesicle in fore or hindbody, oval tubular, connected to
pars prostatica by short aglandular duct. Ejaculatory duct short, enters hermaphroditic
duct, which is a continuation of uterus. Sinus-sac present. Sinus organ absent. Ovary
and vitellarium close to posterior extremity, ovary four-lobed. Seminal receptacle
large. Uterus preovarian. Vitellarium with seven or eight claviform lobes, immedi-
ately postovarian. Excretory arms united in forebody. Type genus: Trifoliovarium
Yamguti, 1940.
Genus Trifoliovarium Yamaguti, 1940
[Syn. Cladoolecithotrema Ichihara, 1970; Pseudolecithaster Campbell & Munroe,

1977]
With characters of the subfamily. Mainly characterized by the vitellarium consist-
ing of seven claviform lobes arranged in a rosette, lobes may be branched distally.
Type species: T. acanthocepolae Yamaguti, 1940
Trifliovarium bombayensis Ahmad, 1983 (Fig. 9.80)
Terapontidae: Terapon theraps (Cuv.):
Locality: BOM, AS
No. 16
Description: Body 4720–4815 long 400–452 wide. Oral sucker 228–267 in

diameter. Ventral sucker 180–226 long, 287–318 wide, protuberant. Sucker ratio
1:1.18–1.39. Caeca simple, terminate near posterior end of body. Testes two, rounded,
contiguous, directly tandem, equal in size. Seminal vesicle tubular, extends slightly
posterior to ventral sucker. Pars prostatica long, surrounded by numerous prostate
gland cells. Sinus-sac saccular, encloses tubular hermaphroditic duct. Genital pore
ventral, sinistral, just post-bifurcal. Ovary 4–lobed, near posterior end of body. Sem-
inal receptacle retort-shaped, ventrolateral to ovary. Vitellarium consisting of seven
elongate distally swollen lobes, connected centrally. Uterus between ovary and ven-
tral sucker. Eggs 52–58 long. 28–35 wide.
Fig. 9.80 Trifoliovarium

1983)
Remarks: Trifoliovarium bombayensis is distinguished from the closely related

species in having a spherical oral sucker, an oval pharynx, the slightly post-bifurcal
genital pore, the seminal vesicle reaching slightly posterior to the ventral sucker, the
saccular sinus-sac, the contiguous testes distantly separated from the ovary. It differs
from the other Indian species T. triacanthi (Parukhin, 1964) in having a spherical
oral sucker rather than funnel-shaped, an oval pharynx instead of barrel-shaped, a
comparatively longer oesophagus, the slightly post-bifurcal genital pore instead of
at the level of the pharynx or oesophagus, the seminal vesicle extending slightly pos-
terior to the ventral sucker, the saccular sinus-sac instead of tubular, the contiguous
testes instead of separated and eggs of a larger size.
Trifoliovarium triacanthi (Parukhin, 1964) Bray & Cribb, 2000 (Fig. 9.81)
[Syn. Hysterolecitha triacanthi Parukhin, 1964, Trifoliovarium triacanthi Bilqees,
1973, Trifoliovarium triacanthusi Gupta & Ahmad, 1976]
Host: Triacanthidae: Triacanthus strigilifer (Day) (1); T. brevirostris Temn. &
Schlegel (2)
Fig. 9.81 Trifoliovarium

triacanthi (after Parukhin,
1964)
Localities: Puri, Machilipatnam, Kakinada (Bay of Bengal), Turicorin (Gulf of Man-

nar) and Calicut (Arabian Sea).
No. of specimens: 3 + 5 + 10 + 3 + 3, respectively, total 124 on 7 slides
Reference: 1. Gupta & Ahmad (1976b); 2. Hafeezullah (1990b)
Description: Body long, 4240–4930 long, 540–680 wide. Oral sucker terminal,
funnel-shaped, 220–240 long, 240–290 wide. Ventral sucker subspherical, 190–290
long, 280–410 wide, at 960–1290 from anterior extremity. Intestinal caeca simple,
extend up to posterior extremity. Genital pore submedian, just posterior to pharynx.
Testes oval, obliquely tandem, in middle third of body, distant from ventral sucker.
Seminal vesicle elongated, slightly winding extends either to anterior margin of ven-
tral sucker or to middle of it. Pars prostatica narrow surrounded by many prostate
gland cells. Sinus-sac long. Ovary tetralobate, near posterior end of body. Seminal
receptacle retort-shaped. Vitellarium consists of seven elongate lobes, extending ven-
trally and posterior to ovary. Uterus winding, extends posteriorly as far as posterior
part of ovarian complex and anteriorly opens at base of sinus-sac. Eggs 25–39 long,
11–015 wide.
Discussion: Bilqees (1973) described T . triacanthi, from the fish Triacanthus bre-
virostris from off Karachi (Arabian Sea), differentiating it from T. acanthocepolae
Yamaguti, 1940 mainly by the presence of a relatively larger ventral sucker (Sucker
width ratio 1:1.4), a shorter posterior extent of the seminal vesicle with respect to the
ventral sucker and a smaller egg size. Gupta and Ahmad (1976b), obviously unaware
of Bilqees’ work, described T. triacanthusi from the fish Triacanthus strigilifer from
Puri (Bay of Bengal). Hafeezullah (1990b) regarded T . triacanthusi as conspecific
with T triacanthi. This synonymy has also been indicated by Bray & Cribb (2000),
who considered that Hysterolecitha triacanthi Parukhin, 1964 from T. brevirostris
from the Gulf of Tong King was the oldest available name for this species and placed
it in the genus Trifoliovarium.
Family Sclerodistomidae Odhner, 1927
(Syn. Bhaleraoiidae Srivastava, 1948, Mabiaramidae Teixeira de Freitas & Kohn,

1967)
Sclerodistomids are parasitic in marine teleosts. Odhner (1927) included this
group as a subfamily under Hemiuridae. It was elevated to a full family status by
Dollfus (1932). The family Prosogonotrematidae Vigueras, 1940 and the subfamily
Prosorchiinae Yamaguti, 1934 were considered by Gibson & Bray (1977) as subfam-
ilies under Sclerodistomidae. A characteristic feature of the family is the presence
of Manter’s organ, an accessory excretory organ which is attached to the excretory
vesicle into which it opens. The families Bhaleraoiidae Srivastava, 1948 and Mabi-
aramidae Teixeira de Freitas & Kohn, 1967 are regarded as synonyms of this family.
Diagnosis: Body stout, muscular. Ventral sucker at middle of body. Drusenmagen

present. Testes symmetrical to oblique, in forebody or anterior hindbody. Sinus-
sac present. Sinus organ conical, fills entire sinus-sac. Genital pore midventral in
forebody. Ovary post-testicular. Laurer’s canal opens into Juel’s organ. Canalicular
or blind seminal receptacle absent. Uterus pre- or postovarian. Eggs small, numerous.
Vitellarium composed of convoluted filamentous tubules, usually four to seven, either
entirely in forebody or entirely in hindbody. Manter’s organ single or double. Type
genus: Sclerodistomum Looss, 1912.
Four subfamilies are recognized under Sclerodistomidae: Indian species are con-
fined to the three subfamilies: Prosogonotrematinae, Prosorchiinae Yamaguti, 1934,
Sclerodistominae Odhner, 1927.
Key to subfamilies
1. Manter’s organ double, testes symmetrical or oblique in hindbody
………………………………………………………….... Sclerodistominae
Odhner, 1927
Manter’s organ single, testes in forebody or dorsal to ventral suck-
er………………………….…….2
Family Sclerodistomidae Odhner, 1927 257
2. Ovary in forebody, uterus and vitellarium mainly preovarian…. Proso-

gonotrematinae Perez-Vigueras, 1940
Ovary in hindbody, uterus and vitellarium mainly postovarian …. Prosorchi-
inae Yamaguti, 1934
Subfamily Sclerodistominae Odhner, 1927
Only one species, Sclerodistomum chirocentrusi Gupta & Jain, 1991 from Chirocen-
trus dorab of the coast of Puri, Bay of Bengal, has been reported from India.
Diagnosis: Body stout. Testes in anterior hindbody. Pars prostatica long, convoluted.
Distinct sinus-sac present. Sinus organ conical, stout. Ovary post-testicular, about
mid-hindbody. Seminal receptacle rudimentary. Uterus entirely within hindbody,
mostly post-gonadal, reaches posterior extremity. Vitellarium composed of seven
(usually) convoluted, tightly helical, unbranched, filamentous tubules, three on one
side of body, four on other, post-testicular. Manter’s organ double, symmetrical. Type
genus Sclerodistomum Looss, 1912.
Genus Sclerodistomum Looss, 1912
Diagnosis: With characters of subfamily. Type species S. italicum (Stossich, 1893)

Sclerodistomum chirocentrusi Gupta & Jain, 1991) (Fig. 9.82)
Host: Chirocentridae: Chirocentrus dorab (Forsskål).
Locality: BOB
Reference: Gupta & Jain (1991b).
Fig. 9.82 Sclerodistomum

chirocentrusi (after Gupta &
Jain, 1991)
Description: Body elongated, 6100 long, 1750 wide. Ventral sucker subspherical,
larger than oral sucker, pre-equatorial, 1100 × 1050. Oral sucker subterminal, spheri-
cal, 350 × 360. Pharynx sphericai. Oesophagus short. Caeca extend to just anterior to
posterior extremity. Testes in hindbody, oblique, extracaecal, entire. Seminal vesicle
tubular, in hindbody. Pags prostatica long, sinuous. Ejaculatogy duct short, joining
metraterm at base of genital cone to form hermaphroditic duct. Sinus organ 300 ×
100. Genital poge ventral to intestinal bifurcation. Ovary post-testicular, submedian.
Vitellarium tubular, comprised of six tubes, three on each side, radiating posteriorly
from level of ovary. Uterus intercaecal, fills much of hindbody Eggs 9–10 × 6–7.
Excgetogy poge terminal. Excgetogy vesicle V-shaped.
Remarks: This species, based on a single specimen, is apparently distinguished by
its 6-fold (rather than 7-fold) vitellarium and the tiny eggs.
Subfamily Prosogonotrematinae Perez-Vigueras, 1940
Important features of the family are: Body stout. Ventral sucker large, in middle of
body. Drusenmagen absent. Testes symmetrical, in forebody. Sinus-sac well devel-
oped encloses conical sinus organ. Ovary in forebody or at level of ventral sucker.
Juel’s organ well developed. Uterus preovarian, not extending into hindbody. Vitel-
larium with about three convoluted and filamentous tubules on each side of body,
situated in forebody. Manter’s organ single. Type genus Prosogonotrema Vigueras,
1940.
Genus Prosogonotrema Perez-Vigueras, 1940
[Syn. Bhaleraoia Srivastava, 1948]
With characters of the subfamily. Type species: P. bilabiatum Perez-Vigueras, 1940.
Sixteen species have been reported in the genus. Seven of these have been consid-
ered synonyms of P. bilabiatum by Nasir (1973). Additional nine species have been
reported after Nasir’s work. The validity of many of these species remains to be
assessed.
Seven species of Prosogonotrema were reported from India: P. pritchardae
Hafeezullah, 1970 from Gastrophysus spadiceus and Pristipomoides argyrogrm-
micus from Arabian Sea (Hafeezullah, 1970 and from Stromateus niger from Rat-
nagiri coast, AS, (Karyakarte & Yadav, 1973); P. caranxi Velasquez, 1961 from
Monacanthus monoceros (Day) from Visakhapatnam coast, BOB‘ (Hussain & Rao.
1980); P. arabicum Yadav, 1980 from Apolectus niger, Ratnagiri coast, AS; P. pavesi
Lokhande, 1990 from Apolectus niger, Maharashtra, AS; P. posterouterina Yadav,
1980 from Apolectus niger, RT, AS; P. zygaenae Ali & Bhagwan, 1971 from
Zygaena malleus at Bombay and P. piscicola (Srivastava, 1948) Gibson, 2002 (Syn.
Bhaleraoia piscicola Srivastava, 1948) from a marine fish at AS. Of these P. caranxi,
P. pritchardae and P. zygaenae were regarded as synonyms of P. bilabiatum (Ref.
Madhavi, 2011). The validity of the remaining species is to be determined. Consid-
ering the reasons given by Nasir (1973) for synonymizing the various species with
P. bilabiatum, there is every possibility that some of the Indian species erected after
1973, would fall synonyms of P. bilabiatum. In this account the description and other
details are provided only for P. bilabiatum whose validity is well established.
Prosogonotrema bilabiatum Perez-Vigueras, 1940 (Fig. 9.83)
[Syn. Prosogonotrema abalisti Parukhin, 1964; Prosogonotrema carangi Velasquez,

1961; Prosogonotrema clupeae Yamaguti, 1952; Prosogonotrema pritchardae
Hafeezullah, 1971; Prosogonotrema subequilatum Pritchard, 1963; Prosogonotrema
zygaenae Ali & Bagwan, 1971] (Reference: Nasir, 1973; Sey et al., 2003).
Hosts: Monacanthidae: Monacanthus monoceros (Day) (1); Sphyrnidae: Zygaena
malleus (Linn.) (2) Tetraodontidae: Gastrophysus spadiceus (Richardson) (3);
Lutjanidae: Pristipomoides argyrogrammicus (Richardson); (3); Carangidae:
Parastromateus niger (Bloch) (4).
Localities: VSK, BOB (1); BOM (AS) (2); RT (AS) (3, 4)
Reference: 1. Hussain & Rao (1980); 2. Ali & Bhagwan (1971); 3. Hafeezullah
(1971b); 4. Karyakarte & Yadav (1973)
Description: Body elongated, 5632–7152 long, 1488–1568 wide. Tegument thick,

muscular. Preoral lobe present. Suckers prominent. Oral sucker 320–400 × 372–468.
Ventral sucker 1120–1208 × 990–1184. Sucker ratio 1:3 to 1:3.2. Pharynx oval.
Caeca sinuous, long. Testes two, symmetrical, midway between oral and ventral
suckers, separated by seminal vesicle and proximal uterine coils. Seminal vesicle
tubular, coiled, pretesticular followed by tightly coiled pars prostatica surrounded by
dense layer of prostatic cells. Ejaculatory duct convoluted, embedded in muscular
tissue, joins metraterm to form hermaphroditic duct surrounded by cup-shaped geni-
tal cone. Genital pore median, at base of oral sucker. Ovary globular, anterodorsal to
ventral sucker. Uterus preovarian, occupies space in forebody. Eggs 23–31 × 11–19.
Vitellarium in the form of slender tubules, anterior to ovary, five tubules on right
sideand four on left. Excretory vesicle Y-shaped, accessory excretory vesicle single.
Fig. 9.83 Prosogonotrema

bilabiatum (after
Perez-Vigueras, Hussain &
Rao, 1980)
Remarks: P. bilabiatum was first described by Perez-Vigueras (1940) from Ocyurus

chrysurus at Hawana, Cuba. Subsequently it has been recorded from a wide range
of localities including Celebes (as P. clupeae Yamaguti, 1952), Gulf of Tonkin (as
P. abalisti Parukhin, 1964), Philippines (as P. carangi Velasquez, 1961), Hawaii
(as P. subequilatum Pritchard, 1963), Tonkin Bay (as P. symmetricum Oshmarin,
1955), Bay of Bengal (as P. caranxi by Hussain & Rao, 1980) and Arabian Sea
(as P. zygaenae Ali & Bhagwan, 1971, as P. pritchardae by Hafeezullah, 1971 and
Karyakarte & Yadav, 1973), Qatari waters of the Arabian Gulf (Kardousha, 2016),
estuarine waters of Bangladesh (Latifa et al. (2008). It has also been recorded from a
wide range of hosts belonging to the families Carangidae, Haemulidae, Polynemidae,
Stromateidae Tetraodontidae, Lutjanidae and Sphyrnidae.
Subfamily Prosorchiinae Yamaguti, 1934
Diagnosis: Body very long slender. Oral sucker subterminal. Preoral lobe present.
Oesophagus with posterior diverticula. Testes in forebody. Seminal vesicleelongate.
Ovary ventral, a little behind. middle of body. Seminal receptacle small. Laurer’s
canal present. Uterine coils convoluted behind ovary. Uterine seminal receptacle
present. Uterine coils extend up to posterior end of body. Excretory system Y-shaped,
uniting in front. Eggs thick-shelled. Type genus ProsorchisYamaguti, 1934.
Genus Prosorchis Yamaguti, 1934
Diagnosis: Body long, slender, attenuated. Oral sucker subterminal. Preoral lip
present. Oesophagus very short, with posterior diverticula. Intestinal caeca simple,
terminating at posterior extremity of body. Ventral sucker prominent, large, near oral
sucker. Testes dorsal, directly tandem, partly overlapping, just anterior to ventral
sucker. Seminal vesicle elongate. Pars prostatica well differentiated. Hermaphroditic
duct present. Genital atrium opening ventrally near anterior border of oral sucker.
Ovary ventral, just posterior to middle of body. Seminal receptacle small, immedi-
ately posterior to ovary. Uterus extending to near posterior end. Vitellarium tubular,
long, convoluted, extending from ovary to posterior end of body. Eggs numerous,
thick-shelled. Excretory system Y-shaped, uniting anteriorly. Type species: Prosor-
chis psenopsisYamaguti, 1934.
Prosorchis breviformis Srivastava, 1936 (Fig. 9.84)

Host: Carangidae: Elagatis bipinnulatus (Quoy & Gaimard) (= Seriolichthys bipin-
nulatus Quoy & Gaimard)
Locality: PR, BOB
Reference: Srivastava (1936a)
Description: Body subcylindrical, 4500–5400 × 750–960. Preoral lip long, roughly

triangular. Oral sucker ventrally placed, transversely oval. Ventral sucker spherical,
situated at the junction of first and second third of body length, about twice the
size of oral sucker. Pharynx oval, well developed. Oesophagus extremely small with
oesophageal diverticulum. Caeca long, sinuous, extending to posterior end. Testes
small, spherical, obliquely tandem, intercaecal, about halfway between intestinal
Fig. 9.84 Prosorchis

breviformis (after Srivastava,
1936)
bifurcation and ventral sucker. Seminal vesicle lying between testes and intestinal
bifurcation. Pars prostatica short, tubular, surrounded by numerous prostate gland
cells, continuing anteriorly to join ejaculatory duct which joins terminal part of uterus
to form hermaphroditic duct. Genital atrium median, ventral, at the level of posterior
third of oral sucker, enclosing tiny small genital papilla. Ovary transversely ovoid,
in posterior half of body. Seminal receptacle small, spherical. Uterine coils extend
up to posterior end. Eggs numerous, small, 33–38 × 18–25. Vitellarium consists of
two to four longitudinal, highly convoluted, tubes extending from ovary to hinder
end, main tubes giving off secondary branches which anastomose. Excretory bladder
V-shaped, main stem sinuous, bifurcating just posterior to ventral sucker, into two
lateral cornua, which anastomose dorsal to pharynx.
Remarks: The species is recorded to differ from the type-species of the genus P.
psenopsis Yamaguti, 1934, in the shape and much smaller size of body, the compar-
atively posterior position of the ventral sucker; the position of the testes, and the
character and disposition of the vitellarium.
Chapter 10
Superfamily Apocreadioidea Skrjabin,
1942
Molecular studies (Olson et al., 2003; Pulis et al., 2014; Littlewood et al., 2015) have
shown that apocreadiids are not lepocreadioids, and that they are distinct enough to
warrant a separate superfamily.
Family Apocreadiidae Skrjabin, 1942
[Syn: Homalometridae Cable & Hunninen, 1942; Schistorchiidae Yamaguti, 1942;

Sphincterostomatidae Yamaguti, 1958, Trematobrienidae Dollfus, 1950]
The Apocreadiidae was reviewed by Cribb & Bray (1999) and is considered
to include genera recognized previously within the families Apocreadiidae, Homa-
lometridae, Schistorchiidae, Sphincterostomatidae and Trematobrienidae. Key fea-
tures of the family are extensive vitelline follicles, eyespot pigment dispersed in the
forebody, the I-shaped excretory vesicle, the lack of a cirrus-sac and the genital pore
opening immediately anteriorly to the ventral sucker (usually) or immediately poste-
rior to it. Three subfamilies are recognized within the Apocreadiidae. The subfamilies
are Apocreadiinae, Schistorchiinae and Postporinae.
Key to subfamilies of Indian species:
Oral sucker with partial muscular sphincter encircling oral aperture …… Schistorchi-
inae Yamaguti 1942
Oral sucker without partial sphincter at aperture ………… Apocreadiinae Skrjabin,
1942
Subfamily Apocreadiinae Skrjabin, 1942
[Sy. Homalometrinae Cable & Hunninen, 1942; Eocreadiinae Mehra, 1962;

Microcreadiinae Mehra, 1962]
© Crown 2018 263

https://doi.org/10.1007/978-94-024-1535-3_10
264 10 Superfamily Apocreadioidea Skrjabin, 1942
Diagnosis: Tegument spinose or unspined. Oral sucker with or without papillae.

Ventral sucker with or without lamellar lips or papillae. Prepharynx usually very
short. Caeca blind or form cyclocoel. Genital pore median immediately anterior to
ventral sucker. Testes two, tandem, oblique or symmetrical. Ovary entire. Vitelline
follicles extensive. Excretory vesicle I-shaped extends to level of testes or ovary.
Key to genera
Vitelline follicles restricted to hindbody …………………. Homalometron Stafford,
1904
Vitelline follicles extend into forebody …………. Neoapocreadium Siddiqi & Cable,
1964
Genus Homalometron Stafford, 1904
[Syn. Apocreadium Manter, 1937; Austrocreadium Szidat, 1956; Barbulostomum
Ramsey, 1965]
Diagnosis: Tegument spined. Oral sucker without papillae. Ventral sucker unspe-
cialized. Prepharynx longer than pharynx. Oesophagus short. Caeca blind, extend
close to posterior end of body. Testes tandem. Genital pore median, immediately
anterior to ventral sucker. Ovary entire. Vitelline follicles restricted to hindbody.
Excretory vesicle I-shaped. Lymphatic vessel present or absent. Type species: H.
pallidum Stafford, 1904.
Homalometron vinodae (Ahmad, 1985) Cribb & Bray, 1999 (Fig. 10.1)
[Syn. Apocreadium vinodae Ahmad, 1985a]
Host: Balistidae: Sufflamen chrysopterum Bl. & Schn. (= Hemibalistes chrysoptera
(Bl. Schn.))
Locality: BOM, AS
Number: 51
Reference: Ahmad (1985a); Cribb & Bray (1999)
Fig. 10.1 Homalometron

vinodae (after Ahmad 1985,
Pakistan J. Zool. 17, p. 161)
Family Apocreadiidae Skrjabin, 1942 265
Description: Body elongate, 1620–1670 long by 450–470 wide. Entire tegument

spinose. Eyespot pigment present. Oral sucker cup-shaped, terminal, 116−125 long
by 146−155 wide. Ventral sucker nearly spherical, 142−150 in diameter, situated
at junction of anterior and middle third of body. Sucker ratio 1:1.0 − 1.2. Intestinal
bifurcation dorsal, nearer to oral sucker than to ventral sucker. Caeca end blindly
near posterior end of body. Testes two, entire, smooth, tandem, contiguous, sube-
qual in size, in posterior half of body. Cirrus-sac absent. Seminal vesicle tubular,
winding, extends posteriorly to just anterior to anterior testis. Pars prostatica sur-
rounded by few prostate gland cells. Genital atrium tubular. Genital pore median,
ventral, immediately anterior to ventral sucker. Ovary spherical, submedian, to right
of median line, immediately pretesticular. Seminal receptacle saccular, immediately
pretesticular. Uterus confined between anterior testis and ventral sucker. Eggs elon-
gate, operculate, 55–61 long by 25–28 wide. Vitelline follicles extend from mid-level
of ventral sucker to posterior end of body, confluent in post-testicular area. Excre-
tory vesicle tubular extends anteriorly as far as posterior border of posterior testis.
Excretory pore terminal. Lymphatic system with two pairs of narrow longitudinal
vessels extends most of body length; occasionally branching.
Remarks: This species was initially included in the genus Apocreadium as A. vinodae
Ahmad, 1985. Cribb and Bray (1999) in a review of Apocreadiidae transferred the
species into the genus Homalometron as H. vinodae (Ahmad, 1985). Homalometron
is a large genus comprises over 20 species. H. vinodae is the only species of the
genus known from India.
Genus Neoapocreadium Siddiqi and Cable, 1960
This genus very closely resembles Homalometron, but differs only in the vitellarium
extending anterior to the ventral sucker. Lymphatic vessels are also present. Type
species: N. angustum (Sogandares-Bernal, 1959).
Neoapocreadium caballeroi Ahmad, 1987 (Fig. 10.2)
Host: Sillaginidae: Sillago sihama (Forsskal)
Fig. 10.2 Neoapocreadium

caballeroi (after Ahmad
1987, Pakistan J. Zool. 19,
p. 328)

No.27
Description: Body elongate, 1749–2270 long, 650–720 wide. Tegument spined.

Oral sucker 190–240 in diameter, subterminal. Ventral sucker 200–380 in diame-
ter, spherical, situated at junction of middle and anterior third of body. Sucker ratio
1:1.52–1.58. Prepharynx very short, with band of circular muscles at anterior end.
Pharynx oval. Oesophagus long. Intestinal bifurcation between suckers. Caeca sim-
ple, end blindly near posterior end of body. Testes longitudinally sigmoid, contigu-
ous, in posterior half of body, occupy most of hindbody. Cirrus-sac absent. Seminal
vesicle tubular, dorsal to ventral sucker, extends posteriorly as far as mid-level of
ovary, or one-third of it. Pars prostatica short, with few prostatic gland cells. Geni-
tal pore median, immediately anterior to ventral sucker. Ovary entire, pretesticular.
Seminal receptacle postovarian. Vitelline follicles rounded, intrude into intercaecal
space extend from level of intestinal bifurcation, to posterior end of body, confluent
in post-testicular region, Uterus scanty, between anterior testis and ventral sucker,
opens into hermaphroditic duct. Eggs few, 110–120 × 60–68 in size. Excretory vesi-
cle I-shaped, terminates near posterior border of ovary. One pair of lymphatic vessels
extends along inner side of each caecum.
Remarks: This species differs from all the other species of the genus in having
roughly sigmoid testes occupying most of the hindbody, the post-testicular field is
very short and the excretory vesicle extends anteriorly as far as posterior level of
ovary.
Neoapocreadium imtiazi (Ahmad 1984) Cribb and Bray 1999 (Fig. 10.3)
[Syn. Crassicutis imtiazi Ahmad, 1984]
Host: Drepanidae: Drepane punctata (Linn.)

imtiazi (after Ahmad 1984)
No. 17
Description: Body elongate, 2200–3500 long, 715–820 wide, elongate. Oral sucker
255–300 in diameter. Ventral sucker 350–416 in diameter, preequatorial. Sucker
ratio 1:1·37–1·38. Prepharynx absent. Pharynx oval. Oesophagus short. Intestinal
bifurcation between suckers. Caeca simple, end blindly near posterior end of body.
Testes oval, contiguous, in posterior half of body. Cirrus-sac absent. Seminal vesi-
cle saccate, dorsal to ventral sucker, extends posteriorly as far as anterior margin
of ovary, or one-third of it. Pars prostatica short with few prostatic gland cells.
Hermaphroditic duct short. Genital pore median, immediately anterior to ventral
sucker. Ovary tri-lobed, pretesticular. Seminal receptacle postovarian. Vitelline fol-
licles rounded, intrude into intercaecal space, extend from posterior border of oral
sucker, confluent anteriorly and in post-testicular region, interrupted opposite ventral
sucker. Uterus scanty, between anterior testis and ventral sucker. Eggs few, 64–50 by
46–55 in size. Excretory vesicle I-shaped, terminates near posterior border of rear
testis.
Remarks: The most important character of the species is the interruption of the
vitelline fields at the level of the ventral sucker. In this respect, it differs from all the
species of Neoapocreadium.
Neoapocreadium karwarense (Hafeezullah 1970) Cribb and Bray 1999 (Fig. 10.4)
[Syn. Crassicutis karwarensis Hafeezullah, 1970]
Host: Gerreidae: Gerres filamentosus Cuv.
No. 20
Distribution: AS and Australia

karwarense (after
Hafeezullah, 1970)
Description: Body 1290–3000 long, 379–980 wide, elongate. Tegument thick. Eye-
spot pigment present. Oral sucker 143–295 in diameter, spherical, subterminal. Ven-
tral sucker 215–360 in diameter, spherical, preequatorial. Sucker ratio 1:1·22–1·52.
Caeca simple, reaching posterior end of body. Testes entire, tandem, in posterior half
of body. Cirrus-sac absent. Seminal vesicle sac-like, postero-sinistral and dorsal to
ventral sucker. Ejaculatory duct curving along anterior margin of ventral sucker. Gen-
ital pore median, slightly anterior to ventral sucker. Ovary subglobular, submedian
to right, between anterior testis and ventral sucker. Seminal receptacle club-shaped,
dorsal to ovary, occasionally tubular. Vitellarium consists of large follicles, invariably
from level of intestinal bifurcation to posterior end of body, confluent in forebody
and post-testicular region. Uterus not observed in any specimen. Only one collapsed
egg measuring ‘81–57’.
Remarks: Reported by Cribb and Bray (1999) from Gerres subfasciatus from Aus-
tralia.
Subfamily Schistorchiinae Yamaguti 1942
Diagnosis: Tegument spined or not. Oral sucker unspecialized or with reduced mus-
culature and highly glandular, with a U-shaped sphincter half encircling the oral
opening. Ventral sucker in anterior half of body. Prepharynx and pharynx present.
Intestinal bifurcation in mid-forebody. Caeca open to exterior through separate ani
or through uroproct. Testes two, tandem or multiple. Genital pore median, imme-
diately anterior to ventral sucker. Ovary entire. Vitelline follicles may be restricted
to hindbody or enter forebody. Excretory vesicle I-shaped. Type genus: Schistorchis
Lühe, 1906.
Genus Schistorchis Lühe, 1906
Diagnosis: Tegument spined. Oral sucker and ventral sucker unspecialized. Caeca
open independently through ani. Testes multiple, usually 11, in one or two columns.
Vitelline follicles profuse and usually restricted to hindbody. Type species: S. carneus
Lühe, 1906.
Four species of Schistorchis were reported from marine fishes of Indian coastal
waters.
Schistorchis carneus Lühe, 1906 (Fig. 10.5)
[Syn. Pleorchis oligorchis Johnston, 1913; Schistorchis oligorchis (Johnston 1913)]
Host: Tetraodontidae: Arothron hispidus Linnaeus; Gastrophysus lunaris (Bloch)
Locality: Mandapam, GOM; Kalingapatnam coast, BOB
Distribution: Sri Lanka, Queensland
Reference: Hafeezullah (1981b); Madhavi et al. (1986)
Description: Body rectangular, 5596–6000 long, 2076–3052 wide. Oral sucker

1196–1306 in diameter, mouth opening small, surrounded by strong circular muscles,
margin of anterior part of musculature notched or indented, remaining part of oral
Fig. 10.5 Schistorchis

carneus (after Madhavi
et al., 1986)
sucker highly cellular. Ventral sucker 343–522 in diameter, weakly muscular, situ-
ated near anterior end just post-bifurcal. Sucker ratio 1:0.43. Pharynx close behind
oral sucker, large. Oesophagus not present. Caeca arcuate extend to posterior end of
body, ani formed. Testes 11, in two lateral groups of 5 and 6 on either side of mid-line,
intercaecal, unequal, entire. Cirrus-sac absent. Seminal vesicle pear-shaped extends
posterior to ventral sucker, continues as pars prostatica surrounded by prostatic cells.
Ejaculatory duct short. Genital pore immediately anterior to ventral sucker. Ovary
pretesticular, globular. Seminal receptacle adjacent to ovary. Uterus short, preovarian.
Eggs 56–59 × 38–42. Vitelline follicles extend to level of ventral sucker.
Remarks: S carneus has previously been recorded from Sri Lanka and Queensland
(Luhe, 1906; Johnston, 1913). The species shows specificity towards tetraodontidae
fishes. From India, there are two reports: that of Hafeezullah (1980) from Arothron
hispidus of Gulf of Mannar and of Madhavi et al. (1986) from Gastrophysus lunaris of
Bay of Bengal. The Bay of Bengal specimens were much larger (12,100–14,400 ×
4,580–4,800) than those from the Gulf of Mannar (5,596–6,000 × 2,076–3,052).
There is a marked difference in the egg size also (78–80 × 28–32 against 56–59 ×
38–42). In other characters, there is close similarity between the forms from the two
localities.
Schistorchis longivesiculurus Hafeezullah, 1981 (Fig. 10.6)
Host: Siganidae: Siganus oramin Schneider
Locality: Tuticorin (Gulf of Mannar) and Karaikal (Coromandel coast)
Number: Numerous

longivesiculurus (after
Hafeezullah 1981, 1981,
Bull. Zool. Surv. India. 3,
p. 168)
Description: Body 7658–15482 long, 1100–1306 wide, elongate with parallel sides.
Oral sucker 508–868 in diameter, globular. Ventral sucker 453–825 in diameter, close
to anterior end of body. Sucker ratio 1:0.81–0.94. Pharynx squarish. Prepharynx and
oesophagus absent. Intestinal bifurcation anterior to ventral sucker. Caeca long, ani
not clearly seen (obscured by vitelline follicles). Testes follicles 11, in one row along
median line from posterior to ovary to near posterior end of body. Seminal vesicle
long, winding, with thick muscular wall, extends far posterior to ventral sucker.
Pars prostatica surrounded by prostatic cells present towards anterior end of male
duct. Cirrus-sac absent. Genital pore immediately anterior to ventral sucker. Ovary
pretesticular, globular, smaller than testes. Seminal receptacle present. Uterus short,
terminal part joining male duct to form hermaphroditic duct. Eggs 42–77 by 28–51.
Vitelline follicles extend to level of ventral sucker.
Remarks: This species is characterized by the presence of 11 testes follicles arranged

in a single longitudinal row and the long winding tubular seminal vesicle.
Schistorchis manteri Gupta & Tandon 1984 (Fig. 10.7)
Host: Carangidae: Alepes djedaba (Forsskal)
Locality: PR, BOB
No. 2
Reference: Gupta, S.P. and Tandon (1984)
Description: Body elongate, 9050 long, 1750 wide. Oral sucker 510 × 650. Ventral
sucker subspherical, 550 long. 600 wide, almost equal to oral sucker. Prepharynx

manteri (after Gupta &
Tandon, 1984)
small. Pharynx muscular. Oesophagus short. Intestinal bifurcation just anterior to

ventral sucker. Caeca extend to posterior end of body, without opening or ani. Genital
pore on lateral side of ventral sucker. Testes 11 in number, arranged in zigzag manner
in one median row. Seminal vesicle free in parenchyma between ventral sucker and
ovary extends as far as anterior end of ovary. Ovary ovoid, submedian, pretesticular.
Seminal receptacle flask-shaped, lateral to ovary. Uterus coiled, lying between ovary
and genital pore. Eggs numerous, ovoid, 45–65 long, 25-40 wide. Vitellarium follic-
ular, mainly lateral and ventral to caeca, extend from just posterior to ventral sucker
to slightly anterior to posterior end of body, confluent in post-testicular region.
Remarks: The distinguishing features of this species are: The suckers subequal, the
11 testes are arranged in a single zigzag manner, the seminal vesicle is large, and the
ovary is situated far apart from the ventral sucker.
Schistorchis skrjabini Parukhin, 1963 (Fig. 10.8)
Host: Triacanthidae: Triacanthus biaculeatus (Bloch)
Locality: Coromandel coast, BOB
No. 6
Distribution: South China Sea
Description: Body 1925−6682 long, 536−1906 wide. Oral sucker larger than ven-
tral sucker, differentiated into smaller muscular portion and highly cellular portion.
Ventral sucker 137−275 × 110−357. Sucker ratio 1:0.36 –0.46. Caeca long, open
through ani. Testes 11, arranged in 2–3 rows in middle part of body. Cirrus-sac absent.
Seminal vesicle globular extends beyond ventral sucker posteriorly. Pars prostatica
surrounded by prostatic cells. Ejaculatory duct, narrow, tubular. Genital pore imme-
diately anterior to ventral sucker. Ovary globular, smaller than testes, pretesticular.

skrjabini (after Hafeezullah
1981b, Bull. Zool. Surv.
India. 3, p. 172)
Uterus preovarian. Vitellarium follicular extends from posterior margin of ventral

sucker to posterior end of body.
Remarks: S. skrjabini was first described by Parukhin (1963) from Triacanthus bre-
virostris and Abalistes stellaris from the South China Sea. In specimens reported by
Hafeezullah (1980) from Triacanthus biaculeatus from Coromandel coast, the oral
sucker is differentiated into a small muscular part around the short mouth opening
and the rest of it is highly cellular. In some specimens, there is a tendency to form
muscular lips or small lobes around the mouth. This differentiation in the structure of
the oral sucker has not been mentioned in Schistorchis skrjabini by Parukhin (1963).
Chapter 11
Superfamily Transversotrematoidea
Witenberg, 1944
Family Transversotrematidae Witenberg, 1944
[Syns. Transversotrematinae Witenberg, 1948, Circuitocoelidae Wang, 1941; Squa-

macolidae Pan & Wang, 1985]
The Transversotrematidae is the smallest family of digeneans comprising only

four genera and about 23 species. Members of this family are distinctive in that they
occur as ectoparasites residing underneath the scales of freshwater and marine fishes.
Morphologically also the family exhibits many unusual features such as transversely
elongate body, the absence of an oral sucker, the cyclocoel gut, etc. The life cycle
pattern is also distinctive in that the cercariae are almost progenetic and directly
attack the definitive hosts. The life cycle involves only two hosts. The family does
not show affinities with any other digenean family. Hence, the superfamily Transver-
sotrematoidea Witenberg, 1944 was erected to include this family.
Genus Transversotrema Witenberg, 1944
Diagnosis: Body transversely elongated, spined, pair of eyespots present. No oral

sucker. Ventral sucker median. Prepharynx, pharynx and oesophagus present. Gut
cyclocoel. Testes symmetrical, lobed. Seminal vesicle tubular. Cirrus-sac and pars
prostatica absent. Genital pore at anterior margin. Ovary sinistral to left testis, lobed.
Uterus short. Eggs large. Vitellarium with widely distributed follicles. Excretory
vesicle I-shaped. Excretory pore at small notch in middle of posterior margin. Type
species: T. haasi Witenberg, 1944.
Tranversotrema patialense (Soparkar, 1924) Crusz & Sathananthan, 1960 (Fig. 11.1)
[T. koliensis (Olivier, 1947) Yamaguti, 1971; T. chackai (Nadakal, Mohandas & Sun-
deraraman, 1969) Mohandas, 1973; T . laruei Velasquez, 1958; T. soparkari (Pandey,
1971) Pandey & Shukla, 1972] Ref. Cribb et al. 1992.]
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https://doi.org/10.1007/978-94-024-1535-3_11
274 11 Superfamily Transversotrematoidea Witenberg, 1944
Fig. 11.1 Transversotrema

patialense (original)
Host: Mugilidae: Mugil cephalus L

Locality: VSK, BOB
Description: With characters of genus. Body transversely elongated. Tegument

spined, with one pair of prominent eye spots. Ventral sucker median, at midbody.
Testes entire or lobed, symmetrical, intercaecal. Seminal vesicle tubular. Genital pore
median, near anterior body margin. Ovary entire, submedian, anterior to left testis.
Uterus very short, eggs rarely found. Egg oval, 132–136 × 76–81 in size Vitelline
follicles numerous forming thick lateral extracaecal zones, continuous posteriorly.
Remarks: There are several reports of the occurrence of T. patialense underneath the
scales of fishes from India and adjacent regions (Crusz et al. 1964; Rao and Ganap-
ati, 1967; Murty & Rao, 1968; Betterton, 1979; Rekharani & Madhavi, 1985). Apart
from T. patialense, few other species of cercariae and adults of Transversotrema have
been recorded from India, namely T . chackai (Nadakal, Mohandas & Sunderraman,
1973) Mohandas, 1973; T. soparkari (Pandey, 1971) Pandey & Shukla, 1972; and
T. chauhani Agrawal & Singh, 1981. Cribb et al. (1992) in a review of the genus,
Transversotrema, synonymized the former two species with T. patialense. The exis-
tence of cryptic species among Transversotrema spp. has also been indicated (Hunter
& Cribb, 2012; Cribb et al. 2014). In the life cycle of the species, the snail Thiara
tuberculata serves as the intermediate host (Crusz, 1964). The cercaria is furcocer-
cous, with accessory structures at the junction of the body and tail. The reproductive
organs are fully formed. The cercaria attaches to the fish, moves to the space beneath
the scales and develops into an adult within a week. Apart from the life cycle, infor-
mation is also available on the chromosomes (Madhavi & Ramanjaneyulu, 1986)
and the miracidium (Madhavi & Jhansilakshmibai, 1968) of T. patialense. All these
studies and the molecular studies indicated that the family Transversotrematidae
deserves an independent status in the phylogenetic tree of the Digenea.
Chapter 12
Superfamily Haploporoidea Nicoll, 1914
Key to families
Testis single, intestine with one or two caeca ………………………. Haploporidae
Nicoll, 1914
Testes two, intestine with two caeca ………………..……… Atractotrematidae Yam-
aguti, 1939
Family Haploporidae Nicoll, 1914
[Syns. Hyporhamphitrematidae Machida & Kuramochi, 2000; Waretrematidae Sri-

vastava, 1937; Megasolenidae Skrjabin, 1942]
Members of the family Haploporidae Nicoll, 1914 are parasites of marine, estuar-
ine and freshwater teleost fishes. The superfamily Haploporoidea Nicoll, 1914 was
erected by Mehra (1961) for the families Haploporidae, Waretrematidae Srivastava,
1937, Haplosplanchnidae Poche, 1925 and Megaperidae Manter, 1934. Later Over-
street and Curran (2005) discussed the relationships of these various families and
included only two families under this superfamily: the Haploporidae and the Atrac-
totrematidae. The Waretrematidae was reduced to a subfamily level, Waretrematinae
under Haploporidae. The Haplosplanchnidae is included in a separate superfamily
Haplosplanchnoidea Poche, 1925 (see Madhavi, 2005). The Megaperidae has been
transferred to Lepocreadioidea Odhner, 1905 (see Bray, 2005a).
Haploporidae is mainly characterized by the presence of a hermaphroditic sac and
usually a single testis. An external seminal vesicle is also present in many species.
Reviews on the family were provided by Overstreet & Curran (2005) and Blasco-
Costa et al. (2009a, 2009b).
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276 12 Superfamily Haploporoidea Nicoll, 1914
Diagnosis: Tegument spinous. Forebody with dispersed eyespot pigment. Oral and
ventral suckers present. Pharynx well developed. Intestine single or bifurcated into
two caeca, extent variable. Testis single. Hermaphroditic sac presents enclosing
hermaphroditic duct of variable length, pars prostatica, internal seminal vesicle and
terminal part of the uterus. External seminal vesicle present. Genital pore median,
anterior to ventral sucker. Seminal receptacle usually present. Ovary pretesticular.
Extent of uterus varies. Eggs enclose miracidia with or without eyespots. Vitellar-
ium variable in shape and distribution. Excretory vesicle Y or I-shaped. Type genus:
Haploporus Looss, 1902.
Overstreet and Curran (2005) recognized four subfamilies: Haploporinae Nicoll,

1914, Chalcinotrematinae Overstreet & Curran 2005, Megasolininae Manter, 1935
and Waretrematinae Srivastava, 1937. Bray et al. (2014), Andres et al. (2014, 2015,
2016) and Andres-Gomez et al. (2017) based on molecular evidence transferred
Cadenatellinae from Lepocreadiidae to this family. The following key serves to dif-
ferentiate these subfamilies:
1. Hermaphroditic sac well-developed ……………………...2
Hermaphroditic sac atrophied …………………… Cadenatellinae Gibson &
Bray, 1982
2. Vitellarium not extensive composed of one or two condensed masses ………….
Haploporinae Nicoll, 1914
Vitellarium composed of groups of follicles……………………………….…..3
3. Vitellarium follicles elongate, distributed around testis or irregularly dispersed
in hindbody. Uterus occupies entire hindbody. Chalcinotrematinae Overstreet &
Curran, 2005
Vitelline follicles variously shaped, occupy large area of hindbody.
Uterus confined to median region between hermaphroditic sac and ovary
……………………………….. 4
4. External seminal vesicle cylindrical, elongate, testes one or two ….…………
Megasolininae Manter, 1935
External seminal vesicle sac or club-shaped, testis single …………………….
Waretrematinae Srivastava, 1937
Subfamily Haploporinae Nicol, 1914

[Syns. Discogasterinae Yamaguti, 1958, Unisaccinae Martin, 1973]
Diagnosis: Body fusiform or elongate, with delicate spined tegument. Intestine with
a single caecum or two caeca, extend to various levels in body. Testis single, oval.
External seminal vesicle sac or club-shaped. Seminal receptacle canalicular or uter-
ine. Uterus extensive usually confined to hindbody. Eggs containing miracidia with
eyespots. Vitellarium one or two compact masses. Excretory vesicle Y or I-shaped.
Type genus Haploporus Looss, 1902.
Eight species belonging to this subfamily and spread over the genera Haploporus
Looss, 1902 (2 spp.), Lecithobotrys Looss, 1902 (2 spp.), Pseudounicoelium Ahmad,
1987 (2 spp.) Saccocoelium (1 sp.) and Unisaccus Martin, 1973 (1 sp.) have been
Family Haploporidae Nicoll, 1914 277
recorded from marine fish of India. Of the two species reported under the genus
Lecithobotrys, L. mugilis Rekharani and Madhavi, 1985 was transferred to the genus
Unisaccus Martin, 1973 and L. vitellosus Sharma & Gupta, 1970 as well as Sacco-
coelium tripathi Dutta & Manna, 1998 have been regarded as species inquirendae
(see Blasco-Costa et al. 2009). The genus Pseudounicoelium Ahmad, 1987 was syn-
onymized with Unisaccus by Overstreet & Curran (2005).
Key to the three genera under which species were recorded from India
1. Intestine sac-like, pharynx well developed …………………. Unisaccus Martin,
1973
Intestine with two caeca, pharynx not so well developed…………………….2
2. Caeca relatively short ……………………………… Saccocoelium Looss, 1902
Caeca cylindrical, relatively long ………..…………..Haploporus Looss, 1902
Genus Haploporus Looss, 1902
[Syn. Neohaploporus Manter, 1963]
Diagnosis: Body elongate with diffuse eyespot pigment. Ventral sucker smaller
than oral sucker. Caeca two, medium or long. Testis single. Hermaphroditic sac
elongate, shorter or longer than ventral sucker, hermaphroditic duct long occupies
approximately half length of hermaphroditic sac. Internal seminal vesicle smaller
or larger than external seminal vesicle. Ovary pretesticular. Uterus in hindbody.
Eggs relatively small. Vitelline gland in the form of two compact symmetrical lobes,
situated immediately behind ovary. Type species: H. benedeni (Stossich, 1887) Looss,
1902.
Two species belonging to this genus have been recorded from India: H. indicus
Rekharani & Madhavi, 1985 and H. pseudoindicus Rekharani & Madhavi, 1985.
Haploporus indicus Rekharani& Madhavi, 1985 (Fig. 12.1)
Locality: VSK, BOB
No. 14
Description: Body 752–1120 long, 162–208 wide, spindle-shaped. Tegumental

spines large. Oral sucker funnel-shaped, 39–58 × 58–89, ventral sucker slightly
smaller than oral sucker, 39–58 × 39–46. Sucker-ratio 1: 0.5–0.8. Caeca long, slen-
der, terminate at testicular level. Genital pore median, in between intestinal bifur-
cation and ventral sucker. Testis single, elliptical, situated in middle third of body.
Hermaphroditic sac well developed, claviform, curves round ventral sucker. Genital
atrium shallow, unarmed. External seminal vesicle club-shaped, between ovary and
hermaphroditic sac. Ovary small, oval, pretesticular. Uterus occupies most of hind-
body. Eggs numerous, elliptical, 31–35 × 11–15, enclose miracidia with prominent
eyespots. Vitelline gland in form of two small compact symmetrical lobes, situated
immediately posterior to ovary. Seminal receptacle small. Excretory vesicle elongate
extends to mid-post-testicular space.
Fig. 12.1 Haploporus

indicus (after Rekharani &
Madhavi, 1985)
Remarks: According to Rekharani & Madhavi (1985) H. indicus closely resembles

H. benedeni Looss, 1902 from Mugil cephalus and M. chelo from the Gulf of Naples
and H. lateralis Looss, 1902 from M. auratus and M. labrosus from Trieste. It differs
from these species in the position of the testis in the middle of the hindbody rather
than close to the ventral sucker, in the position of the intestinal bifurcation anterior
to the ventral sucker, in the shape of the hermaphroditic sac and its extension to the
level of the ovary and in the egg size.
Haploporus pseudoindicus Rekharani & Madhavi, 2005 (Fig. 12.2)
Host: Mugilidae: Liza macrolepis (Smith)
Locality: VSK, BOB
No. 1
Description: Body elongate, 394–407 long, 135–195 wide. Oral sucker cup-shaped,
80 long by 128 wide. Ventral sucker 564 by 80, situated in anterior third of body.
Sucker-ratio 1: 0.63. Prepharynx absent. Pharynx small, Oesophagus long. Intesti-
nal bifurcation near posterior border of ventral sucker. Caeca elliptical, parallel,
reach to level of testis Genital pore median, near anterior border of ventral sucker.
Testis single, large, oval, in posterior half of body. Hermaphroditic sac cylindrical,
overlaps ventral sucker anterolaterally, External seminal vesicle club-shaped extends
to posterior border of ventral sucker. Ovary small, spherical, pretesticular. Uterine
coils narrow occupy space between ovary and ventral sucker, extend posteriorly to
midlevel of testis. Eggs small, 19 by 15, miracidia non-oculate. Vitellarium two
tandem compact masses, just posterior to ovary.
Fig. 12.2 Haploporus

pseudoindicus (after
Rekharani & Madhavi, 1985)
Remarks: H. pseudoindicus differs from H. benedeni and H. lateralis in the sucker-

ratio, the position of the caecal bifurcation, the position of the testis and the extent of
the uterus. It differs from H. indicus in the size and shape of the body and the much
smaller eggs enclosing miracidia apparently lacking eyespots.
Key to Indian species of Haploporus

1. Eggs large, enclose miracidia with eyespots …………………………………H.
indicus Rekharani & Madhavi, 1985
Eggs smaller, enclose miracidia lacking eyespots ……………………....………
H. pseudoindicus Rekharani & Madhavi, 1985
Genus Lecithobotrys Looss, 1902

Lecithobotrys vitellosus Sharma & Gupta, 1970
Host: Mugilidae: Liza parsia (Ham.)
Locality: MS, BOB
Reference: Sharma & Gupta (1970)
Remarks: Regarded as species inquirendum by Blasco-Costa et al. (2009)
Genus Saccocoelium Looss, 1902
Saccocoelium tripathi Dutta & Manna, 1998
Host: Mugilidae: Mugil sp.
Locality: Chilka lagoon
Reference: Dutta & Manna (1998a)
Remarks: Considered as species inquirendum by Blasco-Costa et al. (2009).
Genus Unisaccus Martin, 1973

[Syn. Pseudounicoelium Ahmad, 1987]
Genus Unisaccus is characterized by having a single undivided caecum, uterus
extending up to posterior end of body and vitellaria confined to one side of body
only.
Diagnosis: Body pyriform to fusiform. Eyespot pigment diffuse. Oral sucker termi-
nal. Pharynx relatively large. Caecum sac-like with or without bifurcation posteriorly.
Testis spherical, in posterior half of body. Hermaphroditic sac twice as long as wide.
Hermaphroditic duct relatively long, armed. Metraterm relatively short. Ovary at
or slightly anterior to level of testis. Uterus in hindbody. Eggs with miracidia hav-
ing single eyespot. Vitellarium in two distinct lateral clusters of follicles, lateral to
or posterior to intestine. Excretory vesicle Y-shaped. Type species: U. brisbanensis
Martin, 1973.
Overstreet and Curran (2005) synonymized Pseudounicoelium Ahmad, 1987 with

Unisaccus. The genus is represented by four species in Indian marine fishes, including
the two species reported under Pseudounicoelium Ahmad, 1987 and one species
transferred from the genus Lecithobotrys.
Unisaccus guptai (Ahmad, 1987) Overstreet & Curran, 2005 (Fig. 12.3)
[Syn: Pseudounicoelium guptai Ahmad, 1987]
Host: Mugilidae: Mugil cephalus Linn.
Locality: PR, BOB
References: Ahmad (1987b); Overstreet and Curran (2005)
Description: Body fusiform, 1020–1360 long, 310–380 wide. Oral sucker 100–125
in diameter. Ventral sucker smaller than oral sucker, 60–75 in diameter, at junction
of anterior and middle third of body. Sucker-ratio 1:0.6. Caecum single, undivided,
bottle-shaped, in middle third of body. Testis single, near posterior end of body.
Hermaphroditic sac saccular, extends to anterior margin of ventral sucker. External
Fig. 12.3 Unisaccus guptai

(after Ahmad, 1987, Pakistan
J. Zool.19, p. 338)
seminal vesicle tubular, extends posteriorly to just posterior to mid-body. Genital

pore median, at level of pharynx, surrounded by muscular genital sucker. Ovary
spherical, distinctly pretesticular. Vitellarium consists of 7–9 follicles on each lateral
side confined to ovarian zone. Uterus runs between ovary and ventral sucker. Eggs
few, elongate, contain developing miracidia with prominent eyespots. 68–75 long
31–36 wide.
Remarks: This species differs from the remaining three species of Unisaccus from
the Indian region, in the location of the testis very close to posterior extremity, the
very short post-testicular space, the ovary well separated from the testis instead of
close to it, the much larger eggs and the position of the genital pore anterior to the
pharynx rather than at the level of the ventral sucker.
Unisaccus martini Ahmad, 1986 (Fig. 12.4)
Host: Mugilidae: Liza vaigiensis (Quoy and Gaimard)
Locality: Panjim coast, Goa.
No. 12
in diameter. Ventral sucker smaller than oral sucker, 68–90 in diameter pre-equatorial
Sucker-ratio 1: 0.60–0.68. Caecum undivided. Testis single, spherical, near poste-
rior end of body. Hermaphroditic pouch saccular, extends slightly posterior to ventral
sucker. Hermaphroditic duct provided with spirally arranged pads bearing spines, and
terminal portion of uterus. External seminal vesicle saccular, extends to caecal ends.
Genital pore ventral, midway between pharynx and ventral sucker. Ovary spherical,
immediately pretesticular, submedian, to left of median line. Uterus descends to pos-
terior end of body. Eggs operculate, containing developing miracidia with prominent
Fig. 12.4 Unisaccus martini

(after Ahmad, 1986)
eyespots, 95–110 long, 55–67 wide. Vitelline follicles few, 6–7 in number, lateral,
on one side of body only, confined to ovarian zone.
Remarks: This species differs from the other species of the genus, in having the oral
sucker much larger than the ventral sucker, in the much larger eggs and in the fairly
large ovary.
Unisaccus mugilis (Rekharani & Madhavi, 1985) Blasco-Costa et al. 2009 (Fig. 12.5)
[Syn. Lecithobotrys mugilis Rekharani & Madhavi, 1985]
Hosts: Mugilidae: Mugil cephalus Linn., Liza macrolepis (Smith); Valamugil cun-
nesius (Val.)
Number: Numerous
Description: Body small, length 394–407, width 135–195. Oral sucker 39–51 ×
41–54. Ventral sucker 54–68 × 39–54. Sucker-ratio 1:1.0–1.2. Caeca two, short,
cylindrical, terminate just posterior to mid-body. Genital pore median, in between
pharynx and ventral sucker. Testis single, elliptical, situated in posterior half of
body. Hermaphroditic sac well developed, curves round ventral sucker separated
from ovary by uterus. External seminal vesicle saccular situated in between ovary
and ventral sucker. Ovary pretesticular, overlapping anterior border of testis. Seminal
receptacle present. Vitelline follicles large, distributed along lateral fields between
ovary and ventral sucker with 6–8 follicles on each side. Uterus confined to hind-
body, transforms anteriorly as metraterm. Eggs large, 78–79 × 31–37, contain oculate
miracidia.
Remarks: Rekharani & Madhavi (1985) included this species in the genus
Lecithobotrys as L. mugilis but Blasco-Costa et al. (2009) transferred it to the genus
Unisaccus Martin, 1973. U. mugilis differs from the other species of the genus in the
Fig. 12.5 Unisaccus mugilis

(after Rekharani & Madhavi,
1985)
smaller size of the body and the organs. In addition, the egg size ranges of the other
species of Unisaccus, do not overlap those of U. mugilis, they are either bigger or
markedly smaller.
Unisaccus overstreeti (Ahmad, 1987) Overstreet & Curran, 1987 (Fig. 12.6)
[Syn: Pseudounicoelium overstreeti Ahmad, 1987]
Host: Mugilidae: Mugil cephalus L.
Locality: Goa, AS
References: Ahmad (1987b); Overstreet and Curran (2005)
Description: Body fusiform, 710–940 long, 250–330 wide. Oral sucker spherical,
75–130 in diameter. Ventral sucker pre-equatorial, 135–208 in diameter. Sucker-ratio
1:1.6–1.8. Prepharynx long. Pharynx spherical. Oesophagus relatively short. Caecum
single, undivided, bottle-shaped extends to mid-hindbody. Testis single, oval, some
distance anterior to posterior end of body, smaller than ovary. Hermaphroditic duct
situated in forebody, extends to midlevel of ventral sucker. Genital pore median in
front of ventral sucker, surrounded by muscular sucker. Ovary spherical, larger than
testis, pretesticular, partly overlaps testis. Vitellarium consists of 10–12 follicles on
each lateral side of body, confined to preovarian zone. Uterus runs between ovary
and ventral sucker. Eggs few, large, 82–100 × 52–60 wide.
Remarks: Ahmad (1987b) included this species in a new genus Pseudounicoelium

as P. overstreeti Ahmad, 1987. Overstreet and Curran (2005) synonymized the genus
with Unisaccus. U. overstreeti (Ahmad, 1987b) is characterized by the ventral sucker
being larger than the oral sucker and the large ovary which overlaps anterior part of
testis.
Fig. 12.6 Unisaccus

overstreeti (after Ahmad,
1987, Pakistan J. Zool.19,
336)
Key to the four Indian species of Unisaccus

1. Testis situated almost at posterior extremity of body, ovary situated far anterior
to testis ……………………………………………….U. guptai (Ahmad, 1987)
Testis situated a little away from posterior end of body, ovary immediately pretes-
ticular …………………………………………………………………………2
2. Ventral sucker larger than oral sucker, ovary larger than testis, overlaps anterior
part of testis ………………………………………U. overstreeti (Ahmad, 1987)
Ventral sucker smaller than or subequal to oral sucker, ovary smaller than testis,
pretesticular ……………………………………………………………………3
3. Ventral sucker smaller than oral sucker, sucker-ratio 1: 0.60–0.68, eggs 95–110 ×
55–67 …………………………………………….… U. martini Ahmad, 1986
Ventral sucker subequal to oral sucker, sucker-ratio 1: 1.0–1.2, eggs 78–79 by
31–37 ………………………………………………….…. U. mugilis Rekharani
& Madhavi, 1985
Subfamily Chalcinotrematinae Overstreet & Curran, 2005
This subfamily was erected by Overstreet and Curran (2005) to accommodate
the genera possessing vitelline follicles forming a patch surrounding the testis rather
than in two lateral groups, the uterus extending throughout the hindbody and into the
forebody and a short prepharynx.
Diagnosis: Body fusiform. Oral sucker simple. Ventral sucker in anterior half of
body. Intestine as a single caecum or two caeca. Testis single. Hermaphroditic sac
oval. Ovary pretesticular. Uterine coils throughout hindbody. Eggs large, enclose
miracidia with or without eyespots. Vitellarium in two fields surrounding testis.
Type genus Chalcinotrema Freitas, 1947.
Genus Paralecithobotrys Freitas, 1947
Diagnosis: Body elongate, eyespots present. Oral sucker terminal. Ventral sucker
in anterior third of body. Caeca cylindrical terminate near posterior end of body.
Hermaphroditic sac elongated. Testis single in middle of hindbody. External seminal
vesicle longer than internal seminal vesicle. Ovary immediately pretesticular. Uterus
occupies most of hindbody. Eggs large, with oculate miracidia. Vitellarium with
subspherical to lobed follicles, which surround testis. Type species: P. brasiliensis
Freitas, 1948.
Paralecithobotrys overstreeti Ahmad, 1985
Host: Mugilidae: Liza vaigiensis (Quoy and Gaimard).
No. 58
Reference: Ahmad (1985d)
Description: Body cylindrical, 1910–2245 long, 415–445 wide. Oral sucker oval,
subterminal, 155–180 long, 185–195 wide. Ventral sucker spherical, prebifurcal,
larger than the oral sucker, 210–250 in diameter, at junction of anterior and middle
third of body. Sucker-ratio 1: 1.12–1.28. Intestinal bifurcation dorsal, between ovary

and ventral sucker. Caeca terminate about one-fourth body length from posterior
end. Testis single, longitudinally elongate, in posterior part of middle third of body.
Hermaphroditic sac saccular, in forebody Genital pore ventral, median, at midlevel of
pharynx, surrounded by genital sucker. Ovary spherical, immediately pretesticular.
Uterine seminal receptacle present. Uterus in long loops from near posterior end of
body to hermaphroditic sac. Eggs thin-shelled, operculate, containing non-oculate
miracidia, 90–125 µm long, 55–65 µm wide. Vitellarium in two lateral clusters of
6–8 follicles each, confined to ovarian zone.
Remarks: The relationship of this species to Paralecithobotrys is doubtful. Over-
street & Curran (2005) mentioned the identifying features of Paralecithobotrys as:
vitellicles follicles distributed in a patchy manner rather than in two distinct groups
and a uterus extending beyond the hermaphroditic sac. Neither of these features is
satisfied by P. overstreeti. Moreover the two other species of the genus have been
recorded from freshwater fishes. Overstreet & Curran (2005) in their analysis of the
genus, did not mention this species. It is here regarded as a taxon inquirendum.
Genus Saccocoelioides Szidat, 1954
[Syn. Lecithobotrioides Thatcher & Dossman, 1974]
Diagnosis: Body elongate. Eyespot pigment in forebody. Oral sucker terminal. Ven-
tral sucker at mid-body, larger or smaller than oral sucker. Prepharynx short. Oesoph-
agus longer than pharynx. Caeca sac-like. Hermaphroditic duct ellipsoidal. External
seminal vesicle longer or smaller than internal seminal vesicle. Ovary close to testis.
Uterus fills most of hindbody. Eggs with miracidia having eyespots. Vitellarium fol-
licles in lateral fields on both sides of gonads, extend posteriorly beyond testis. Type
species: S. nanii Szidat, 1954.
Two species of this genus S. martini Madhavi, 1979 and S. chilkaensis Dutta, 1995
have been recorded from Indian waters. S. martini was transferred to the genus
Pseudohapladena Yamaguti, 1952 as P. martini (Madhavi, 1979a) Overstreet &
Curran, 2005.
Saccocoelioides chilkaensis Dutta, 1995 (Fig. 12.7)
Host: Mugilidae: Mugil sp.
Locality: Chilka lake
No. 100
Description: Body small, 810–1007 long, 320–350 wide. Ventral sucker equatorial,
70–90 × 100–110. Oral sucker 70–90 × 110–120. Sucker-ratio 1:1.0. Prepharynx
long. Oesophagus short. Intestinal caeca short, stumpy, reaching anterior level of
testis. Testis single, entire, situated at posterior end of body. Hermaphroditic sac
formed dorsal to ventral sucker. External seminal vesicle present, extending up to
testis. Genital pore anterior to ventral sucker. Ovary entire, small, pretesticular. Sem-
inal receptacle absent. Uterus between ventral sucker and testis. Eggs 53–69 × 36–49
contain miracidia. Vitellarium follicular, situated posterior to caecal ends.
Fig. 12.7 Saccocoelioides

chilkaensis (after Dutta,
1995)
Remarks: According to Dutta (1995) this species differs from all the known species
of the genus Saccocoelioides in the long prepharynx, the short oesophagus, the
extent of external seminal vesicle to testis and the number of vitelline follicles. In the
illustration the vitellarium appears as a single V-shaped mass overlapping the testis.
Species of Saccocoelioides are usually found in freshwater fishes.
Subfamily Megasoleninae Manter, 1935
[Syn. Scorpidicolinae Yamaguti, 1971]
Members of this subfamily are characterized by vitelline fields extending through-

out the hindbody and an elongate and narrow external seminal vesicle.
Diagnosis: Body elongate, tegument thick with minute spines. Oral sucker terminal
or subterminal. Prepharynx shorter or longer than pharynx. Ventral sucker peduncu-
late or not. Caeca cylindrical, extend into hindbody, usually to posterior end. Testes
single or two in tandem or slightly oblique. External seminal vesicle cylindrical.
Uterus confined to medial region between caeca. Vitelline fields distributed laterally
usually between ovary and hermaphroditic sac. Eggs large. Type genus: Megasolena
Linton, 1910.
Genus Vitellibaculum Montgomery, 1957
[Syn. Allomegasolena Siddiqi & Cable, 1960]
Diagnosis: Body elongate, with spined tegument. Eyespot pigment present or

absent. Pharynx well developed. Ventral sucker pedunculate or not. Caeca cylindrical
extend into hindbody. Testes single or two, in tandem or slightly oblique. External
seminal vesicle cylindrical. Uterus confined to medial region between caeca. Uter-
ine seminal receptacle present. Eggs operculate or not, miracidium lacks eyespots.
Vitelline fields extend throughout hindbody. Type species: V. girella Montgomery,
1957.
Vitellibaculum fischthali Ahmad 1986 (Fig. 12.8)
Host: Lutjanidae: Lutjanus fulviflamma (Forskal)
Fig. 12.8 Vitellibaculum

fischthali (after Ahmad,
1986)

No.: 10
Description: Body elongate, 1480–2615 long, 270–435 wide. Oral sucker spher-
ical, subterminal, 115–160 in diameter. Ventral sucker nearly same size as oral
sucker, prebifurcal, between pharynx and intestinal bifurcation, 110–165 in diame-
ter. Sucker-ratio 1:1. Prepharynx present. Pharynx barrel-shaped. Oesophagus long.
Intestinal bifurcation dorsal, posterior to ventral sucker. Caeca reach posterior end of
body. Testes two, tandem, close together, in posterior third of body, almost equal in
size. Hermaphroditic sac saccular. External seminal vesicle saccular, extends to level
of intestinal bifurcation. Genital pore ventral, at anterior level of pharynx. Ovary
spherical, immediately pretesticular, median. Uterine seminal receptacle present.
Uterus preovarian. Eggs elongate, operculate, 110–130 long, 65–82 wide. Vitellar-
ium follicular, follicles almost rounded, 22–28 follicles on each side, extend from
midway between intestinal bifurcation and ovary to caecal ends. Excretory vesicle
V-shaped, arms extend to midlevel of anterior testis. Three pairs of longitudinal lym-
phatic vessels present on each side extending from oral sucker to posterior end of
body.
Remarks: The presence of two testes situated in tandem near the posterior end of the
body and the fairly well developed and laterally distributed vitelline follicles suggest
the inclusion of this species in the genus Vitellibaculum. V. fischthali differs from
all the known species of the genus, in having a genital pore at the anterior level of
the pharynx instead of between the pharynx and the ventral sucker, in the extension
of the external seminal vesicle to the level of the intestinal bifurcation, instead of
extending well posterior, and in having eggs of double the size than the other species.
Subfamily Waretrematinae Srivastava, 1937

[Syn. Carassotrematinae Skrjabin, 1942; Spiritestinae Yamaguti, 1958.
Diagnosis: Body pyriform to elongate. Tegument delicate. Oral sucker with or with-
out associated lobes. Ventral sucker near mid-body or in anterior half of body. Intes-
tine either sac-like or cylindrical, extends to approximately mid-body. Testis sin-
gle. External seminal vesicle sac or club-shaped. Uterus confined to median region
between caeca and extends between hermaphroditic sac and ovary. Eggs opercu-
late or not. Miracidium lacks eyespots. Vitelline follicles surround testis. Lymphatic
system absent. Excretory vesicle Y or I-shaped. Type genus Waretrema Srivastava,
1937.
Five species distributed over four genera Carassotrema, Pseudohapladena, Skrjbi-

nolecithum and Waretrema have been recorded from Indian marine fish.
Key to genera of Waretrematinae:
1. Oral sucker with muscular lobed projections ……………… Waretrema Srivas-
tava, 1937
Oral sucker without muscular lobed projections ……………………….…….2
2. Caeca two, not sac-like, terminate in post-testicular space ……Carassotrema
Park, 1938
Caeca one or two, short or moderately long, terminate in post-testicular space
…………3
3. Body elongate, caeca terminate at mid-body or in anterior half of body
………………………………………….…. Pseudohapladena Yamaguti, 1954
Body fusiform, caeca terminate near mid-body or in posterior half of body
………………………………………... Skrjabinolecithum Belous, 1954
Genus Carassotrema Park, 1938
Diagnosis: Body oval to pyriform. Oral sucker subterminal. Ventral sucker roughly
of same size as oral sucker. Oesophagus usually as long as pharynx. Caeca cylindri-
cal, terminate in post-testicular space. Testis irregular or lobed, elongate, in posterior
half of hindbody. Hermaphroditic sac elliptical or arcuate. External seminal vesicle
elongate, sac-like. Ovary immediately pretesticular. Seminal receptacle canalicu-
lar. Uterus confined to region between testes and hermaphroditic sac. Eggs non-
operculate. Miracidium lacks eyespots. Vitellarium follicular or tubular in lateral
fields of hindbody. Excretory system Y-shaped. Type species: C. koreanum Park,
1938.
Carassotrema bengalense Rekharani & Madhavi, 1985 (Fig. 12.9)
Host: Mugilidae: Mugil cephalus L.
Locality: VSK, BOB, Chilka lagoon
References: Rekharani & Madhavi (1985); Shameem & Madhavi (1991)
Description: Body bottle-shaped 760–880 long, 288 wide. Oral sucker 56–80 by
80–112. Ventral sucker at one-third of body length from anterior end, 96–128 in
Fig. 12.9 Carassotrema

1979)
diameter. Sucker-ratio 1: 1.0–1.3. Prepharynx present. Pharynx globular. Oesoph-

agus of same length as prepharynx. Intestinal bifurcation near anterior border of
ventral sucker. Caeca long, terminate near posterior end of body. Genital pore imme-
diately posterior to pharynx. Testis single, large, occupies most of intercaecal space
in hindbody. Hermaphroditic sac situated obliquely anterior to ventral sucker. Exter-
nal seminal vesicle situated postero-lateral to ventral sucker, connected to internal
seminal vesicle through long narrow duct overlying ventral sucker. Ovary triangular,
or ovoid, situated anterolateral to testis, Seminal receptacle lateral to ovary. Vitelline
follicles large, form wide zones along lateral fields, reach anterior border of ventral
sucker. Uterus confined to space between ventral sucker and ovary with one or two
eggs. Eggs 80–88 by 58–64.
Remarks: C. bengalense differs from all the species included in the genus Caras-
sotrema in having an ampullaceous body, the intestinal bifurcation anterior to the
ventral sucker, a smaller hermaphroditic sac situated completely in the forebody, a
globular external seminal vesicle connected to the internal one through a narrow duct,
more numerous vitelline follicles and a very short uterus capable of holding one or
few eggs. Shameem and Madhavi (1991) elucidated the life cycle of C. bengalense
and found the snail Stenothyra blancordiana (Neville) of Chilka lagoon serving as
the molluscan host for the species.
Genus Pseudohapladena Yamaguti, 1952
[Syn. Parasaccocoelium Zhukov, 1971]
Diagnosis: Body fusiform to elongate. Eyespot pigment present. Oral sucker sub-
terminal. Ventral sucker at anterior third of body. Oesophagus usually as long as phar-
ynx. Caeca sac-like, terminate at middle of hindbody. Testes sub spherical, near mid-
dle or posterior portion of hindbody. Hermaphroditic sac elongate. Hermaphroditic
duct occupies major part of sac. Ovary small, pretesticular. Seminal receptacle
canalicular. Uterus confined to region between testes and hermaphroditic sac. Eggs
large. Miracidium lacks eyespots. Vitellarium follicular or tubular in lateral fields
of hindbody, extend to posterior end of body in some species. Excretory vesicle
Y-shaped. Type species: P. scatophagi Yamaguti, 1952.
Pseudohapladena martini (Madhavi, 1979) Overstreet & Curran, 2005 (Fig. 12.10)
[Syn. Saccocoelioides martini Madhavi, 1979]
Hosst: Mugilidae: Mugil cephalus (Linn.); Liza macrolepis (Smith); Valamugil cun-
nesius ((Val.) Cichlidae: Etroplus suratensis (Bloch) and Clupeidae: Tenualosa ilisha
(Hamilton).
Locality: Gosthani Estuary (Visakhapatnam), Chilka lake
References: Madhavi (1979a); Rekharani & Madhavi (1985); Shameem & Madhavi
(1991)
Description: Body oval 688–1178 long 320–400 wide. Oral sucker poorly devel-
oped, 58–125 by 78–136. Ventral sucker 86–136 by 88–136. Sucker-ratio 1: 1.0–1.2.
Pharynx with a distinct sphincter. Intestinal bifurcation at or posterior to level of
ventral sucker. Caeca short saccular, terminate at midway between ventral sucker
and ovary. Genital pore in level with anterior margin of pharynx. Testis single, sit-
uated close to posterior end of body. Hermaphroditic sac anterior or anterolateral
to ventral sucker. External seminal vesicle club-shaped lies dorsal to ventral sucker.
Ovary small. Uterus between testis and hermaphroditic duct. eggs large, 74–78 by
43–47. Miracidium lacks eyespots. Vitelline follicles large, distributed in lateral fields
between caecal ends and posterior edge of testis.
Fig. 12.10 Pseudohapladena

martini (original)
Remarks: This species was initially included in the genus Saccocoelioides Szidat,
1954 as S. martini Madhavi, 1979. Overstreet & Curran (2005) transferred it to the
genus Pseudohapladena Yamaguti, 1952 as P. martini (Madhavi, 1979a). The life
cycle of P. martini has been elucidated by Shameem & Madhavi (1991) from Chilka
lake, with the prosobranch gastropod Stenothyra blancordiana (Neville) serving as
the molluscan intermediate host.
Genus Skrjabinolecithum Belous, 1954
[Syn. Malabarotrema Zhukov, 1972]
Skrjabinolecithum bengalense (Madhavi, 1979) Overstreet and Curran, 2005
(Fig. 12.11)
[Syn. Pseudohapladena bengalense Madhavi, 1979]
Locality: VSK, BOB
Description: Body elongate, 3520 in length, 520 in width, Tegument thin, densely
spined in anterior half. Eyespot pigment present. Oral sucker small, poorly muscular,
funnel-shaped, 144 by 200. Ventral sucker 200 in diameter, situated in anterior third
of body. Caeca saccular, lie closely applied to one another, terminate at midway
Fig. 12.11 Skrjabinolecithum

1979)
between ventral sucker and ovary. Genital pore median, anterior to ventral sucker.
Testis single, elliptical, situated close to posterior end of body. Hermaphroditic sac
elliptical, curves round right side of ventral sucker, encloses coiled seminal vesicle,
small pars prostatica and long hermaphroditic duct covered with papillae. External
seminal vesicle long, slender, tubular. Ovary small, pretesticular. Uterus pretesticular,
confined to space between testis and hermaphroditic duct. Metraterm short. Eggs
fairly large, 71 by 39. Miracidium lacks eyespots. Vitelline follicles long slender
tubules, filling most of space between testis and hermaphroditic sac.
Remarks: The morphological features of the species fit into those of the genus Pseu-
dohapladena as defined above. Overstreet and Curran (2005) included the species
in the genus Skrjabinolecithum Belous, 1954 as S. bengalense (Madhavi, 1979)
Overstreet and Curran, 2005. The status of various species in Pseudohapladena and
Skrjabinolecithum require further study. Besprozvannyk et al. (2016) retained this
species in the latter genus.
Skrjabinolecithum indicum (Zhukov) Overstreet & Curran, 2005
[Syn. Malabarotrema indicum Zhukov, 1972]
Host: Cichlidae: Etroplus suratensis (Bloch)
Locality: Off Kochi, Arabian Sea
Reference: Zhukov (1972)
Distribution: Vietnam
Remarks: The species was described as Malabarotrema indicum Zhukov 1972 from
Etroplus suraensis from Arabian Sea off Kochi Coast. Overstreet and Curran (2005)
regarded Malabarotrema as a junior synonym of Skrjabinolecithum. Besprozvannykh
et al. (2016) undertook a study of molecular analysis of a number of haploporids
including S. indicum collected from Valamugil seheli from Vietnam.
Genus Waretrema Srivastava, 1937
Diagnosis: Body elongate, forebody with eyespot pigment. Oral sucker terminal,
with anterior retractile muscular lobes. Prepharynx long. Oesophagus longer than
pharynx. Caeca moderately long, terminate just posterior to middle of hindbody.
Testis single, elongate near posterior end of body. Hermaphroditic sac elongated,
arcuate. External seminal vesicle narrow and elongate. Ovary pretesticular. Uterine
seminal receptacle present. Uterus confined between ovary and hermaphroditic sac.
Eggs non-operculate, miracidium lacks eyespots. Vitelline follicles irregular in shape,
fairly large, in lateral fields of ovario-testicular zone. Type species: W. piscicolum
Srivastava, 1939.
Waretrema piscicolum Srivastava, 1939 (Fig. 12.12)
Host: Mugilidae: Liza vaigiensis (Quoy and Gaim).
Locality: Karachi, Arabian Sea.
Reference: Srivastava (1939g)
Fig. 12.12 Waretrema

piscicolum (original)
Description: Body fusiform, 2300–4600 long by 840–1230 wide. Oral sucker

funnel-shaped, 260–390 × 330–440, divided anteriorly into six conical projections,
two larger dorso-lateral, two lateral and two ventrolateral. Prepharynx long. Phar-
ynx oval. Oesophagus long. Intestinal bifurcation posterior to ventral sucker. Caeca
extend to middle of posterior third of hindbody. Ventral sucker weakly developed,
260–470 × 370–520. Sucker-ratio 1: 1.1–1.5. Testis single, longitudinally elongate,
in posterior half of hindbody. External seminal vesicle elongated, terminates pos-
teriorly to near ovary. Hermaphroditic sac club-shaped, extends posterior to ventral
sucker. Genital pore median, in zone of pharynx. Ovary ovoid, situated in contact
with testis. Seminal receptacle absent. Uterus preovarian, Eggs thin-shelled, 64–73 ×
41–49, miracidia without eyespots. Vitelline follicles tubular extend from ovarian
zone to posterior end of body.
Remarks: This species was initially described by Srivastava (1939 g) based on mate-
rial collected from Liza waigiensis of Karachi, and later it was reported from Scatoph-
agus argus of Philippines (Velasquez, 1961a; Bilqees, 1980); Crenimugil crenilabis
from Japan (Machida, 1996); and Scatophagus argus from China (Liu & Yang 2003).
Pulis & Overstreet (2013) considered none of these subsequent reports as valid. The
material from Japan was renamed as Spiritestis machidai Pulis & Overstreet, 2013,
whereas the specimens listed in the other reports were considered to belong to the
genus Capitimitta Pulis & Overstreet, 2013.
Subfamily Cadenatellinae Gibson & Bray, 1982
Diagnosis: Body elongate. Oral sucker with 8–10 lobes on dorsal and dorso-lateral
margins. Prepharynx and oesophagus present. Intestinal bifurcation in posterior
region of forebody. Uroproct present. Testis single, in anterior to mid-hindbody.
Hermaphroditic sac absent. Seminal vesicle long, sinuous, extends well into hind-
body. Pars prostatica short, vesicular, ejaculatory duct short. Genital pore median,
in forebody. Ovary round. Uterine seminal receptacle present. Eggs thin-shelled.
Vitelline fields restricted to hindbody. Excretory vesicle reaches ovary or ventral
sucker. Type genus: Cadenatella Dollfus, 1946.
Genus Cadenatella Dollfus, 1946

[Syn. Jeancadenatia Dollfus, 1946]
Diagnosis: With characters of subfamily. Type species: C. cadenati Dollfus, 1946.
Molecular studies by Bray et al. (2014), Andres et al. (2014, 2015, 2016) and
Andrade-Gomez et al. (2017) have shown that this species is to be placed within
the superfamily Haploporoidea.
Cadenatella dollfusi (Hafeezullah, 1980) Bray & Cribb, 2001 (Fig. 12.13)
[Syn. Jeancadenatia dollfusi Hafeezullah, 1980]
Host: Kyphosidae: Kyphosus cinerascens (Forskal)
Locality: Tuticorin (GOM)
No. 2
Description: Body long, filiform, 7665 long, 343 wide. No ventral accessory suck-
ers. Tegument spinose. Eyespot pigment present. Oral sucker 147 × 123, with ten oral
lobes, two pairs of long pointed anterolateral lobes and three pairs of short lobes,
one pair dorso-median and two pairs dorso-lateral. Ventral sucker 284 in diame-
ter, situated in anterior region of body. Pharynx long. Prepharynx and oesophagus
distinct. Caeca united posteriorly, anus opening near posterior end of body. Testis
single, situated in posterior part of body. External seminal vesicle very long, extends
from dorsal to ventral sucker to middle of space between ventral sucker and testis,
consists of elongated posterior part and very long narrow anterior part, surrounded
by thick muscular wall. Hermaphroditic sac absent. Seminal vesicle tubular, free
in parenchyma. Pars prostatica surrounded by few prostatic cells. Ejaculatory duct
short. Ovary globular, pretesticular, separated from testis. Uterine seminal receptacle
large. Uterus preovarian. Metraterm differentiated behind ventral sucker. Vitellarium
follicular, extensive post-testicular. Eggs 59 to 63 × 35. Excretory vesicle tubular,
extends to ovary.
Remarks: C. dollfusi is characterized by the presence of two pairs of long pointed

horn-like ventrolateral and three pairs of short oral lobes on the oral sucker and the
absence of accessory suckers. This species was initially included in the genus as
Fig. 12.13 Cadenatella

dollfusi (after Hafeezullah,
1980, Bull. Zool. Soc. India.
2, p. 149)
Jeancadenatia Dollfus, 1946, which was synonymized with Cadenatella Dollfus,

1946 by Nahhas and Cable (1964). This synonymy was accepted by Overstreet
(1969), Bray & Cribb (2001) and Bray (2005).
Family Atractotrematidae Yamaguti, 1939
This is a small family with members from the intestine or gall bladder of marine
fishes. The diagnostic features of the family include presence of two symmetrical
or slightly oblique testes, vitelline fields with follicles interconnected as elongate
lobes, a hermaphroditic sac enclosing both male and female terminal genitalia and a
Y-shaped excretory vesicle.
Two genera of this family each including a single species were reported from
marine fish of India.
Key to genera
1. Testes in forebody or on either side of ventral sucker, ovary post-testicular
…..……….Atractotrema Goto & Ozaki, 1929
Testes in hindbody, ovary intertesticular ……..….…… Isorchis Durio & Manter,
1969.
Genus Atractotrema Goto & Ozaki, 1929
Diagnosis: Body fusiform. Ventral sucker anterior to mid-body. Caeca narrow,

extend to posterior third to fifth of body. Testes in forebody or on either side of ven-
tral sucker. Ovary submedian, post-testicular in mid-body. Eggs moderately large.
Vitelline follicles in lateral fields, sometimes confluent. Lymphatic system present.
Excretory vesicle with long arms. Type species: A. fusum Goto & Ozaki, 1929.
Atractotrema kuntzi Ahmad, 1985 (Fig. 12.14)
Host: Siganidae: Siganus argenteus (Quoy & Gaimard) (=Teuthis rostratus Val).
No. 15
Fig. 12.14 Atractotrema

kuntzi (after Ahmad, 1985)
Description: Body ovoid, plump. Tegument spined. Eyespot pigment granules

present. Oral sucker 95–120 in diameter. Ventral sucker 70–90 in diameter, in ante-
rior part of middle third of body. Sucker ratio 1: 0.73–0.75. Intestinal bifurcation
nearer to ventral sucker than to oral sucker. Caeca end blindly about 1/3–1/4 body
length from posterior end. Testes two, rounded, symmetrical, far apart, lateral to
caeca. Hermaphroditic sac globular, in forebody. External seminal vesicle saccular,
overlaps ventral sucker. Genital atrium shallow. Genital pore ventral, bifurcal. Ovary
spherical, postero-lateral to ventral sucker, larger than testes. Vitellarium follicular,
fields extend from posterior level of ovary to near caecal ends. Uterus not extensive,
intercaecal, reaches to ends of caeca or slightly beyond. Eggs operculate, 46–65 long,
25–32 wide. Excretory vesicle V-shaped, with short stem, and diverging arms extend
to level of ventral sucker. Excretory pore terminal.
Remarks: A. kuntzi differs from the other two species of the genus namely A. fusum
Goto and Ozaki, 1929 and A. sigani Durio and Manter, 1969 in the smaller egg size,
in the oral sucker being larger than ventral sucker and other minor differences.
Genus Isorchis Durio & Manter, 1969

[Syn. Krusadaitrema Zhukov, 1972]
Diagnosis: Body fusiform. Oral sucker terminal. Ventral sucker at mid-body. Caeca
terminate at testicular or post-testicular level. Testes symmetrical or nearly sym-
metrical, immediately post-equatorial. Hermaphroditic sac containing conspicuous
hermaphroditic duct. Ovary spherical to triangular, at testicular level. Eggs few, large.
Vitellarium in lateral fields extending from near pharynx to post-testicular region.
Type species: I. parvus Durio & Manter, 1969.
Isorchis skrjabini Ahmad, 1985 (Fig. 12.15)
Host: Chanidae: Chanos chanos (Forskal).
No. 17
Fig. 12.15 Isorchis

skrjabini (after Ahmad,
1985)
Family Atractotrematidae Yamaguti, 1939 297
Description: Body small, plump, 665–1065 long 370–600 wide. Oral sucker cup-
shaped, 90–140 long, 82–120 wide. Ventral sucker 75–108 in diameter, equatorial.
Sucker-ratio 1: 0.90–0.91. Intestinal bifurcation dorsal, nearer to ventral sucker than
to oral sucker. Caeca ending blindly, not reaching to posterior end of body. Testes
two, symmetrical, well separated from one another in hindbody. Hermaphroditic
sac saccular, between intestinal bifurcation and ventral sucker. External seminal
vesicle saccular, extends posteriorly up to anterior margin of right testis. Genital
atrium shallow into which uterus and cirrus open separately. Genital pore ventral, at
intestinal bifurcation. Ovary triangular, intertesticular. Uterus descends posteriorly
to slightly posterior to testes. Eggs 65–77 long, 30–40 vide. Vitellarium follicular,
fields extend from level of intestinal bifurcation to caecal ends, non-confluent in
forebody and post-testicular. Excretory vesicle Y-shaped, arms short. Excretory pore
terminal.
Remarks: Durio and Manter (1969) erected the genus Isorchis with I. parvus as
its type species collected from Chanos chanos from New Caledonia. Martin (1973)
added the second species, I. manteri Martin, 1973 from Mugil cephalus from Queens-
land, Australia. Ahmad (1985d) included this species in a new genus, Pseudisorchis
Ahmad, 1985 and considered the genus Krusadaitrema Zhukov, 1971 as a synonym
of Isorchis. The single species K. chanosi Zhukov, 1971 from Chanos chanos from
Krusadai Island became I. chanosi (Zhukov, 1971). I. skrjabini Ahmad, 1985 differs
from the other species in having cup-shaped oral sucker that is larger than ventral
sucker and in having vitellarium extending to the level of the intestinal bifurcation
rather than to the pharynx.
Isorchis chanosi (Zhukov, 1971) Ahmad, 1985
[Syn. Krusadaitrema chanosi Zhukov, 1971]
Host: Chanidae: Chanos chanos (Forsskal).
Locality: Krusadai Island, S. India (Gulf of Mannar)
Reference: Zhukov (1971)
Remarks: This species is characterized by: the fusiform body and the occurrence
of a pair of muscular diverticula associated with the proximal portion of the
hermaphroditic duct and attached to the wall of hermaphroditic sac. It differs from
I. skrjabini in having the oral sucker smaller than the ventral sucker, in the oval
ovary and the vitellarium extending from the level of intestinal bifurcation up to
the posterior end rather than up to caecal ends.
Chapter 13
Superfamily Lepocreadioidea Odhner,
1905
Members of this superfamily are mainly parasites in the gut of marine teleosts. Mor-
phologically they exhibit similarities to Allocreadioidea Looss, 1902, but an obvious
difference is the spined tegument as against smooth tegument in allocreadiids. Differ-
ences also occur in the life cycle pattern and the cercarial characters between the two
groups. In a recent reorganization of the superfamily utilizing the phylogenetic esti-
mates inferred from molecular sequences, Bray and Cribb (2012) split the old family
Lepocreadiidae into three major families: Lepocreadiidae, Aephnidiogenidae and
Lepidapedidae. They also recognized the families Enenteridae, Gyliauchenidae and
Gorgocephalidae. The status of two families Apocreadiidae and Acanthocolpidae are
not considered lepocreadioid. The former is now considered a distinct superfamily
Apocreadioidea Skrjabin, 1942, (see Olson et al. 2003), and the latter is considered a
member of the superfamily Brachycladioidea Odhner, 1905, (see Curran et al. 2006).
The classification proposed by Bray & Cribb (2012) is adopted in the present account.
Key to Families Found in Indian Waters
1. Body usually amphistomatous, pars prostatica external to cirrus-sac

……………………………………… Gyliauchenidae Ozaki, 1933
Ventral sucker equatorial or pre-equatorial, pars prostatica enclosed in cirrus-sac
….….….…. 2
2. External seminal vesicle present ………………..………….…..………….… 3
External seminal vesicle absent ………………………… Enenteridae Yamaguti,
1958
3. Ovary close to testes ……………………………… Lepocreadiidae Odhner,
1905
Ovary separated from testes ………………………… Aephnidiogenidae Yam-
aguti, 1934
© Crown 2018 299

https://doi.org/10.1007/978-94-024-1535-3_13
300 13 Superfamily Lepocreadioidea Odhner, 1905
Family Lepocreadiidae Nicoll, 1935
Lepocreadiidae is one of the major families found in the intestine of marine fishes
in all oceans. The important features of the family are: the spined tegument, well-
developed vitelline follicles, the presence of an external seminal vesicle, a cirrus-sac
enclosing an internal seminal vesicle and the I-shaped excretory vesicle.
Diagnosis: Body elongate or transversely elongate. Tegument spinous. Eyespot pig-

ment often present. Oral and ventral suckers well developed. Prepharynx, pharynx,
oespophagus usually present. Caeca usually terminate blindly but may form cyclo-
coel or uroproct or separate ani. Testes usually two, lobed or smooth. External seminal
vesicle present. Cirrus-sac present, encloses internal seminal vesicle, pars prostat-
ica, prostatic cells and cirrus. Genital pore in posterior part of forebody. Vitelline
follicles extensive, in lateral fields of body. Excretory vesicle I-shaped. Type genus:
Lepocreadium Stossich, 1903.
Several systems have been proposed for division of the family Lepocreadi-
idae into subfamilies (Yamaguti 1958, 1971; Mehra 1960; Bray 2005a). Yamaguti
(1971) recognised as many as 24 subfamilies but Bray (2005) discussed the valid-
ity of various subfamilies and restricted their number to only three: Lepocreadiinae
Odhner, 1905; Aephnidiogeninae Yamaguti, 1934 and Lepidapediinae Yamaguti,
1958. Recently, Bray and Cribb (2012) utilizing phylogenetic estimates inferred from
molecular sequences, reorganized the superfamily Lepocreadioidea and elevated the
subfamilies Aephnidiogeninae and Lepidapedinae to family level as Lepocreadiidae,
Aephnidiogenidae Yamaguti, 1934, and Lepidapedidae Yamaguti, 1958. As a result,
the family Lepocreadiidae retains only one subfamily, the Lepocreadiinae.
The constituent genera in the family Lepocreadiidae have been listed by Bray
& Cribb (2012). The various lepocreadiid species reported from India fall into 19
genera (Madhavi, 2011).
Key to Genera of Lepocreadiidae
The key proposed by Bray (2005a) has been modified to include only the genera
reported from Indian marine fishes.
1. Genital pore dorsal …………………….…………Neohypocreadium Machida
& Uchida, 1987
Genital pore ventral ………………………………………………………… 2
2. Testes numerous ………………..……………………………………3
Testes two ……………………………………………………………4
3. Body wider than long or as long as wide, caeca open as ani, testes numerous
……………………………… Neomultitestis Machida, 1982
Body transversely oval, caeca blind, testes 6–24… Transversocreadium
Hafeezullah, 1970
4. Genital pore marginal………………………………………………………5
Genital pore submedian or nearly so ……………..………………………10
Family Lepocreadiidae Nicoll, 1935 301
5. Genital pore postero-lateral to ventral sucker ……..…...…. Opisthogonoporus

Yamaguti, 1937
Genital pore in forebody……………………………….………………… 6
6. Body squat or subtriangular, genital pore marginal, beside oral sucker
………………………………………………… Lepocreadioides Yamaguti, 1936
Body oval, genital pore submarginal, in mid-forebody ……………………… 7
7. Body rhomboidal, no anterior scoop, caeca blind ………………………….
Pseudolepocreadioides Hafeezullah, 1970
Lateral edges of body folded ventrally to form a scoop, caeca apparently open
out through anal pores............. 8
8. Scoop incomplete …………………………..…… Bianium Stunkard, 1930
Scoop complete, joined posteriorly ……………………………………… 9
9. Anterior part of body forms scoop, testes tandem ………….… Diploprocto-
daeum La Rue, 1926
Body cup-shaped, testes symmetrical ……………..….………. Cotylocreadium
Madhavi, 1972
10. Genital pore postero-lateral to ventral sucker ……………. Opisthogono-
poroides Madhavi, 1972
Genital pore antero-lateral to ventral sucker …………………………………11
11. Body oval ………….………………Lobatocreadium Madhavi, 1972
Body elongate, cylindrical or disc-like…………………….………12
12. Oral sucker surrounded by ring of lobes, cyclocoel present …………
…………………………………. Karyakartia Hafeezullah, 1980
Oral sucker lacking lobes, caeca terminate blindly or form uroproct or ani
……..……… 13
13. Ovary multilobed ………………………..……Echeneidocoelium Simha &
Pershad, 1964
Ovary oval or with few lobes ……………………………………… 14
14. Excretory pore dorsal, body a flattened disc ……………………..……..…. 15
Excretory pore terminal, body oval to elongate……………….……………..16
15. Oral sucker aperture ventral …………………………. Hypocreadium Ozaki,
1936
Oral sucker aperture dorsal ………………….. Lutianotrema Singh, Singh &
Singh, 2004
16. Uroproct present ………………………….……………………. Opechona
Looss, 1907
Caeca end blindly…………………………………………………………… 17
17. Pseudoesophagus present ………………….………………… Prodistomum
Linton, 1910
Pseudoesophagus absent …………………………………………… 18
18. Ovary 3–4 lobed, uterus usually sinistral ………….…………. Preptetos
Pritchard, 1960
Ovary oval, uterus mainly median ………………………… Lepocreadium
Odhner, 1905
Genus Bianium Stunkard, 1930

[Syn. Diplocreadium Park, 1942; Diploporetta Strand, 1942; Amarocotyle Travas-
sos, Freitas & Buhrnheim, 1965; Anterovitellosum Gupta, 1967]
The genus Bianium erected by Stunkard (1939) is characterized by the presence
of two anal pores and a scoop-like feature in the anterior region which is incomplete
in the sense that the two edges of the scoop do not fuse posteriorly. Several genera
like Diploporus Ozaki, 1928; Diplocreadium Park, 1942; Diploporetta Strand, 1942,
Anterovitellosum Gupta, 1967, have been regarded as synonyms of Bianium. Genus
Diploproctodaeum La Rue, 1926, was also regarded as a synonym of this genus, but
Bray et al. (1996) differentiated the two genera by the structure of the scoop which
is incomplete in Bianium and complete in Diploproctodaeum.
Diagnosis: Body oval, scoop incomplete; lateral flaps may spread laterally or be
folded over ventrally. Ventral sucker pre-equatorial. Caeca long, abut body wall and
may form ani. Testes oblique to tandem, in mid-hindbody. Cirrus-sac claviform,
mainly in hindbody. Genital pore sinistral to ventral sucker. Ovary multilobed to
trilobed. Uterus pretesticular, metraterm distinct. Vitellarium extends from posterior
end to level of ventral sucker or intestinal bifurcation or restricted to small patches
lateral to ventral sucker.
Six species of Bianium have been reported from marine fishes of India. Two
of these species Bianium bombayensis Gupta, 1968, and B. rameswarensis Gupta,
Sharma and Jain, 1970, have been transferred to the genus Diploproctodaeum as
D. bombayense (Gupta, 1968) Sey 1995 and D. rameswarense (Gupta, Sharma and
Jain, 1970) Bray, Cribb and Barker, 1996. Hafeezullah (1970b) regarded B. madrasi
Gupta, 1968, as a synonym of B. plicitum (Linton, 1928). Most species of Bianium
and its close relative Diploproctodaeum have been described with ani, but Shimazu
(1994) and Bray et al. (1996) reported that the caeca abut the body wall, but do not
necessarily perforate the wall to form ani. Shimazu summarized the situation thus
‘(1) the posterior ends of the caeca were in contact with the tegument in the posterior
part of the body; (2) there was a body pit lined with tegument at each point of contact;
(3) the ceca usually end blindly, but rarely opened into the body pits, thus forming
the so-called ani; and (4) neither sphincter nor gland cells were seen around the body
pits’.
Bianium indicum (Gupta, 1967) Bray, Cribb & Barker, 1996 (Fig. 13.1)
[Syn: Anterovitellosum indicum Gupta, 1967]
Host: Tetraodontidae: Tetraodon viridipunctatus (Gunther),
Locality: PR, BOB
References: Gupta, (1967); Bray, Cribb and Barker (1996)
Description: Body 990 × 570, spined, anterolateral margins of body raised in form
of collar. Anterior region of body with numerous gland cells. Oral sucker subterminal,
75 × 105 in size. Ventral sucker pre-equatorial, 225 in diameter, sucker ratio 1: 1.4.
Pharynx and oesophagus present, latter very short. Caeca broad, abut body wall,
may form ani on either side of excretory pore. Testes in posterior half of body,
Fig. 13.1 Bianium indicum

(after Gupta, 1967);
tandem, sub-rectangular. Cirrus-sac well developed, lies obliquely in front of ventral

sucker. Seminal vesicle saccular, pars prostatica well developed, ejaculatory duct
cylindrical. Genital pore submedian close to anterior border of vental sucker. Ovary
pre-equatorial, follicular, between testis and posterior border of cirrus-sac. Uterus
with few eggs. Eggs 44 × 30. Vitellarium restricted to small lateral patches at level
of cirrus-sac.
Remarks: Gupta (1967) included this species in a new genus and new species as
Anterovitellosum indicum Gupta, 1967. However, the dendrogram of Bray et al.
(1988) indicated the close relationship of this species with B. plicitum, and based on
this, Bray et al. included the species in the genus Bianium. The restricted vitelline
distribution is characteristic of the species.
Bianium plicitum (Linton, 1928) Stunkard, 1931 (Fig. 13.2)
Syn: Psilostomum plicitum Linton 1928; Bianium madrasi Gupta, 1968
Host: Tetraodontidae: Gastrophysus lunaris (Bloch); Lagocephalus spadiceus
(Richardson); 3. Spheroides sp. 4. Takifugu oblongus (Bloch); 5. Tetraodon
viridipunctatus (Gunther)
Site: Intestine
Locality: Madras (1, 5), VSK (2), Digha Coast, BOB (3, 4)
References: 1. Hafeezullah (1970b); 2. Madhavi (1972); 3. Kumari and Srivastava
(1976); 4. Hafeezullah and Dutta (1998); 5. Gupta, A. N. (1968f)
Distribution: Widely distributed in Atlantic, Pacific and Indian Ocean, tropical and
subtropical waters.
Fig. 13.2 Bianium plicitum

(original)
Description: Body elongate or foliate, 1680–1830 long 540–640 wide. Lateral mar-
gins expanded, expansions not meeting ventrally. Oral sucker larger than ventral
sucker. Ventral sucker at about posterior end of anterior third of body. Prephar-
ynx indistinct. Pharynx well developed, anterior margin lobed. Oesophagus short.
Intestinal bifurcation anterior to ventral sucker. Caeca abut body wall, may form ani
on either side of excretory pore. Testes tandem or slightly oblique, post-equatorial.
External seminal vesicle saccular. Cirrus-sac clavifom, extends posterior to ventral
sucker, encloses internal seminal vesicle, pars prostatica surrounded by prostatic
cells and cirrus. Genital pore close to anterior margin of ventral sucker, submedian,
sinistral. Ovary multilobed, pretesticular, median, almost equatorial. Seminal recep-
tacle present. Uterus between ovary and cirrus-sac. Metraterm well developed. Eggs
50–66 × 29–42. Vitelline follicles extend from level of ventral sucker to posterior
end of body. Excretory vesicle tubular, excretory pore terminal.
Remarks: This species is widespread, and many species of Bianium were syn-
onymized with it. Kumari and Srivastava (1976) and Hafeezullah and Dutta (1998)
recorded B. plicitum from Spheroides spp. from Digha coast and Zhukov (1977)
reported it from Torquigener oblongus from Hooghly Estuary. Manter (1947)
observed extensive intraspecific variations in various structures of this species in
the light of which Hafeezullah (1970b) considered B. madrasi Gupta, 1968, as a
synonym of B. plicitum. Bilqees (1974) also observed variations in the species and
concurred with the proposed synonymy.
Bianium purii Gupta, 1968 (Fig. 13.3)

Host: Ariidae: Sciades sona (Hamilton) (= Arius sona Cuv. and Val.)
Locality: PR, BOB
Number: 36
Reference: Gupta, A.N. (1968f)
Description: Body 900–930 by 360–420, elongate, anterior edge of body produced

into thin, narrow fold continuous on either side with lateral folds whose edges turned
over ventrally but do not form the scoop. Anterior part of body filled with gland cells.
Oral sucker subterminal 60–75 × 90–95. Ventral sucker rounded, equal to oral sucker,
65 in diameter. Prepharynx small. Pharynx globular, its anterior region wavy, and wall
divided longitudinally into 9–10 long flaps. Oesophagus short. Caeca abut body wall,
may form ani. Genital pore to left of median line, opposite oesophagus. Testes post-
equatorial, tandem, overlapping. Cirrus-sac, club-shaped, extends obliquely from
genital pore to just anterior to ovary. Seminal vesicle small. Pars prostatica and ejac-
ulatory duct long. External seminal vesicle extends to halfway between cirrus-sac and
anterior testis. Ovary lobed, 10–15 lobes, pretesticular. Seminal receptacle present.
Uterus occupies space between ovary and cirrus-sac. Metraterm well developed.
Eggs 30–35 by 45–48. Vitellarium profuse, follicular, distributed laterally on either
side, confluent behind testes.
Fig. 13.3 Bianium purii

after Gupta, 1968
Remarks: This species was differentiated from B. plicitum in the following features:
the winding disposition of the ejaculatory duct, the meeting of the two sides of the
body fold in the ventral sucker region, the presence of lobes at the anterior region of
pharynx, the slightly smaller size of the ventral sucker compared to the oral sucker
and in the smaller eggs.
Key to Indian species of Bianium

1. Vitelline follicles restricted to small patches at level of cirrus-sac…. B. indicum
(Gupta, 1967)
Vitelline follicles reach to posterior extremity ………………………..……2
2. Eggs smaller than 50 long …..…………………………… B. purii Gupta, 1968
Eggs longer than 50 long …………….…………B. plicitum (Linton, 1928)
Genus Cotylocreadium Madhavi, 1972
The genus Cotylocreadium, erected by Madhavi (1972) to include Diplocreadium
triacanthi Hafeezullah, 1970, from Triacanthus bilineatus, is characterized by the
cup-shaped body and the peculiar winding of the caeca.
Diagnosis: Body cup-shaped. Oral sucker rounded. Ventral sucker at about mid-
body. Intestinal bifurcation in mid-forebody. Caeca diverge to body margins then pass
medially, then posteriorly, terminate blindly at level of testes. Testes two, symmet-
rical, near posterior extremity. External seminal vesicle saccular, extends into hind-
body. Cirrus-sac claviform, reaches ventral sucker, encloses seminal vesicle, pars
prostatica and cirrus. Genital pore sinistrally submedian, bifurcal. Ovary between
anterior margins of two testes. Vitellarium extensive. Type species: C. triacanthi
(Hafeezullah, 1970).
Cotylocreadium triacanthi (Hafeezullah, 1970) Madhavi, 1972 (Fig. 13.4)
[Syn: Diplocreadium triacanthi Hafeezullah, 1970; Diploproctodaeum triacan-
thi (Hafeezullah, 1970) Sey, 1995]
Fig. 13.4 Cotylocreadium

triacanthi (after Madhavi,
1972, courtesy Allen Press,
J. Parasitol, 58, p. 222);
Hosts: Triacanthidae: Triacanthus biaculeatus(Bloch) (1), Pseudotriacanthus strig-

ilifer (Cantor) (2)
Locality: Calicut (1), VSK (2)
Reference: 1. Hafeezullah (1970b), 2. Madhavi (1972)
Description: Body 1190–1360 long, 960–1290 wide, vase-shaped, formed by ven-

tromedial fusion of lateral expansions, with expanded anterior region and rounded
posterior part. Anterior region with numerous gland cells. Tegument thin. Eye-
spot pigment present. Oral sucker 81–110 by 116–155, subterminal. Ventral sucker
122–155 in diameter, feeble, almost equatorial. Sucker ratio 1: 1·07–1·23. Prepharynx
short. Pharynx spherical. Oesophagus longer than pharynx. Caeca diverge to body
margins then pass medially, then posteriorly, terminate blindly near lateral margins
at or near junction of middle and posterior thirds of body. Testes entire, close to
posterior end of body, slightly diagonal or symmetrical. External seminal vesicle
saccular. Cirrus-sac club-shaped, disposed obliquely between intestinal bifurcation
and ventral sucker; encloses internal seminal vesicle, prostatic complex and protrusi-
ble cirrus. Genital pore sinistral, bifurcal. Ovary median, entire, occasionally slightly
irregular in outline, postequatorial, pretesticular. Seminal receptacle saccular, large,
lateral to ventral sucker and ovary. Uterus scanty, between testes and ventral sucker.
Metraterm highly muscular, thick-walled, stout, long, anterolateral to ventral sucker.
Eggs 63 by 48–54. Vitellarium follicular, follicles extend from level of intestinal
bifurcation to posterior end of body, confluent posterior to testes and penetrate into
folding lateral expansions of body. Excretory vesicle tubular, extends to ovary.
Remarks: Hafeezullah (1970b) included this species in the genus Diplocreadium

Park, 1939 as D. triacanthi Hafeezullah, 1970. However, Madhavi, (1972) erected
a new genus Cotylocreadium Madhavi, 1972, for inclusion of this species, charac-
terizing it by the cup-shaped body, the peculiar course of the caeca, the position of
cirrus-sac anterior to ventral sucker and the entire ovary. Sey (1995) included it in
the genus Diploproctodaeum as D. triacanthi (Hafeezullah, 1970b) Sey 1995.
Genus Diploproctodaeum La Rue, 1926

This genus differs from Bianium in that the scoop is complete being formed by
the fusion of median edges. This genus is represented by two species in Indian
marine fish; both were initially included under Bianium but transferred to the genus
Diploproctodaeum by Bray, Cribb and Barker (1996) based on the above character.
Most species of Diploproctodaeum have been described with ani, but Shimazu (1994)
and Bray et al. (1996) reported that the caeca abut the body wall, but do not necessarily
perforate the wall to form ani (see also discussion of Bianium).
Diploproctodaeum bombayense (Gupta, 1968) Bray, Cribb & Barker, 1996
(Fig. 13.5)
[Syn: Bianium bombayensis Gupta, 1968]
Host: Sciaenidae: Dendrophysa russelli (Cuvier) (1), Tetraodontidae: Takifugu
oblongus (Bl.) (2)
Locality: BOM (AS) (1); MS, BOB (2)
References: 1. Gupta, A.N. (1968f); 2. Hafeezullah (1970b)
Fig. 13.5 Diploproctodaeum

bombayense (after
Hafeezullah, 1970)
Description: Body elongate, 1425–1905 × 720–945 . Lateral folds of forebody

turned ventrally with its anterior edges projected beyond oral sucker. Oral sucker
240–255 in diameter. Prepharynx small. Pharynx cup-shaped with wavy anterior
margin. Oesophagus very small. Caeca abut body wall, may form ani. Genital pore
sinistral to posterior margin of ventral sucker. Testes ovoid, subequal, overlapping,
posterior testis slightly bigger than anterior. Cirrus-sac hockey stick-shaped, over-
lapping ventral sucker. External seminal vesicle between ventral sucker and ovary.
Internal seminal vesicle long, narrow, continues as narrow ejaculatory duct in dis-
tal portion of cirrus-sac. Prostatic cells present. Ovary lobed. Seminal receptacle
present. Uterus coils between ventral sucker and ovary. Metraterm runs parallel to
cirrus-sac. Eggs 45–60 × 65–76 Vitellarium follicular, present more or less through-
out postpharyngeal region.
Remarks: This species is characterized by the profusely developed vitelline follicles.

Diploproctodaeum rameshwarensis (Gupta, Sharma & Jain, 1970) Bray, Cribb &
Barker, 1996 (Fig. 13.6)
[Syn: Bianium rameshwarensis Gupta, Sharma & Jain, 1970]
Locality: Rameswaram, BOB
No. 10
References: Gupta, Sharma and Jain (1970); Bray, Cribb and Barker (1996)
Fig. 13.6 Diploprctodaeum

rameshwarensis (after
Gupta, Sharma and Jain,
1970)
Description: Body oblong, 1680–1750 long 980–1150 wide. Lateral body margins
in anterior part of body form longitudinal ridge. Parenchymatous gland cells pro-
fusely developed. Oral sucker 280–300 × 266–270. Prepharynx absent. Pharynx with
8–10 thick knob-like protuberances on its anterior margin. Oesophagus short. Caeca
well separated from lateral margins, abut body wall, may form ani. Ventral sucker
equatorial, smaller than oral sucker, 170–190 × 210–220. Genital pore submedian,
sinistral, bifurcal. Testes oblique, close to each other, in posterior part of body. Cirrus-
sac well developed, dorsal to ventral sucker. Ovary lobed. Seminal receptacle absent.
Uterus preovarian. Eggs 32 × 48. Vitelline follicles small, extend on each side from
intestinal bifurcation to posterior extremity, confluent in post-testicular space.
Remarks: The position of the ventral sucker well posterior to the intestinal bifurca-
tion, the presence of 8–10 lobes on the anterior margin of the pharynx and the bifurcal
anterior extent of vitellarium are the diagnostic features of D. rameshwarensis.
Genus Echeneidocoelium Simha & Pershad, 1964
Diagnosis: Body elongate. Tegument thick. Oral sucker subterminal, retractable.

Forebody short. Prepharynx, pharynx and oesophagus present. Caeca terminate near
posterior end of body. Genital pore sinistral to ventral sucker. Testes sigmoid, tandem,
intercaecal, in posterior region of hindbody. Cirrus-sac containing internal seminal
vesicle, prostatic vesicle, and cirrus. External seminal vesicle long. Prostate cells
external and internal to cirrus-sac. Ovary follicular, pretesticular. Seminal receptacle
present. Metraterm short. Eggs large. Vitellarium extensive, occupying lateral fields
of most of hindbody. Excretory vesicle I-shaped. Type and only species: Echeneido-
coelium indicum Simha and Pershad, 1964.
Echeneidocoelium indicum Simha & Pershad, 1964 (Fig. 13.7)

Host: Echeneidae: Echeneis remora Linn. (1), E. naucratesLinn. (2)
Locality: VSK, BOB
Distribution: BOB, Israel coast, Mediterranean (Fischthal, 1982), Heron Island,
Queensland (Bray and Cribb, 1998); New Caledonia (Bray and Justine, 2012)
References: Simha and Pershad, 1964 (1), Madhavi, 1970 (2), Lakshmi and Rao
(1978),
Description: Body elongate, 3000–9000 long, 720–1200 wide. Tegument spined.

Oral sucker subterminal 97–160 long, 97–200 wide. Ventral sucker 210–500 in diam-
eter. Sucker ratio 1:2.1–2.5 Intestinal bifurcation slightly anterior to ventral sucker.
Intestinal caeca wide with short anterior dilatations near bifurcation, extend almost to
posterior end of body, ends of caeca impinging on excretory vesicle but not opening
into it. Genital pore sinistral, at midlevel of ventral sucker. Testes large, sigmoid,
with three loops, tandem, almost contiguous, in intercecal region of posterior half of
hindbody. Posterior testis longer, attenuated posteriorly. Cirrus-sac curved, extends
posterior to ventral sucker, encloses long seminal vesicle, oval prostatic vesicle,
short cirrus and prostatic cells. External seminal vesicle long, slender, sinuous, ter-
Fig. 13.7 Echeneidocoelium

indicum (after Madhavi,
1970 courtesy Allen Press, J.
minates slightly anterior to ovary, connected to internal seminal vesicle by narrow

duct surrounded by external prostatic cells. Ovary follicular, pretesticular, separated
from testis by seminal receptacle. Seminal receptacle saccate. Uterus preovarian.
Metraterm short. Eggs large, oval, thin-shelled, yellowish, 53–66 long by 35–42
wide. Vitelline follicles extensive, circumcaecal, in lateral fields from level of cirrus-
sac to posterior limit of posterior testis. Excretory pore dorsal, subterminal with a
sphincter. Excretory vesicle I- shaped, extends to level of ventral sucker.
Remarks: This species was first described by Simha and Pershad (1964) from Ech-
eneis remora from the Bay of Bengal. Simha and Pershad (1964) placed the genus in
the family Acanthostomidae. Madhavi (1970) redescribed and redefined the species,
showing that it belongs to the family Lepocreadiidae. It is characterized by the long
slender body with a short forebody and long hindbody, the multilobed ovary, long
elliptical testes arranged in tandem in the posterior half of the body and extensive
vitelline follicles.
Genus Hypocreadium Ozaki, 1936

In their review of the genus Hypocreadium Ozaki, 1936, Bray and Cribb (1996)
summarized the characters of the genus as being ‘broadly oval to rounded body, with
a weakly lobed or entire ovary and with the dorsal excretory pore at the level of
caecal ends.
Diagnosis: Body flat, rounded or wider than long, appears as narrow disc. Tegument
spines sparse or absent. Oral sucker subglobular. Ventral sucker rounded, in middle
of body. Prepharynx present or absent. Pharynx oval. Oesophagus distinct. Caeca
undulating in arc around gonads, terminate blindly posterior to ovary. Testes two,
oval, symmetrical, in anterior part of hindbody. External seminal vesicle saccular or
elongate. Cirrus-sac claviform encloses globular internal seminal vesicle, elongate or
bipartite pars prostatica and long ejaculatory duct. Genital pore sinistral, in forebody.
Ovary intertesticular, entire or weakly lobed. Uterus usually preovarian. Vitelline
follicles surround gonads with lateral borders lacking follicles. Excretory vesicle
I-shaped. Type species: H. symmetrorchis Ozaki, 1936.
Hypocreadium cavum Bray & Cribb, 1996 (Fig. 13.8)
[Syn: Hypocreadium indicum of Hussain et al. (1986)]
Fig. 13.8 Hypocreadium

cavum (after Hussain, Rao
and Shymasundari, 1986)
Host: Balistidae: Abalistes stellatus (Anonymous)1

Locality: VSK, BOB
Distribution: Great Barrier Reef (Bray & Cribb, 1996, Bray et al., 2009); Red Sea
(Al-Barakati, 2011)
References: Hussain et al. (1986b)
Description: Body transversely oval to round, flattened dorsoventrally with median

notch at anterior end. Body with more or less frilled edges turned over ventrally,
broader than long, 1408–2336 × 1680–2624. Oral sucker subterminal, rounded,
144–192 × 160–224. Ventral sucker 128–240 × 128–240 in size, situated at mid-
body. Sucker ratio 1:1.2–1.4. Prepharynx not distinct. Pharynx oval. Oesophagus
long. Caeca arcuate, slightly winding, terminate close to each other near poste-
rior extremity. Testes symmetrical, intercaecal at midbody with smooth or irregular
margins. Genital pore immediately to left of pharynx and posterior margin of oral
sucker. Cirrus-sac claviform, disposed obliquely along median line of right caecum
to level of middle of ventral sucker, not touching ventral sucker. Internal seminal
vesicle small, prostatic vesicle sinuous, cirrus long and sinuous. External seminal
vesicle dextral to ventral sucker. Ovary slightly lobed, intertesticular, immediately
behind ventral sucker. Seminal receptacle long. Uterus preovarian forming thick-
walled metraterm anterior to ventral sucker, opening near pharynx. Eggs oval, 64 ×
36. Vitellarium follicular, profusely developed, extend from level of pharynx, leav-
ing peripheral area free, confluent between caecal ends, both lateral and median to
caeca.
Remarks: H. indicum (Madhavi, 1972) of Hussain et al. (1986b) was regarded as

identical to H. cavum by Bray and Cribb (1996). The latter was differentiated from
H. indicum by the presence of a notch at the anterior median region of the body just
anterior to the oral sucker and the location of the excretory pore on the dorsal surface
between the caecal ends. This species shows some specificity towards Abalistes
stellatus and has been recorded from this host from the Great Barrier Reef and the
Red Sea apart from BOB.
Hypocreadium indicum (Madhavi, 1972) Hussain, Rao & Shyamasundari, 1986
(Fig. 13.9)
[Syn: Pseudocreadium indicum Madhavi, 1972]
Hosts: Monacanthidae: Paramonacanthus choirocephalus (Bleeker) (1); Balistidae:
Abalistes stellaris (Bloch) (2)
Number: 33 mature and 6 immature from 3 hosts.
Locality: VSK, BOB
References: 1. Madhavi (1972); 2. Hussain et al. (1986b)
1 Following Bray et al. (2009), we are using the specific name Abalistes stellatus (Anonymous) here
for the starry triggerfish, rather than A.stellaris (Bloch and Schneider), a commonly used name for
the same fish, in recognition of the clarification of the nomenclatural confusion by Matsuura and
Yoshino (2004).
Fig. 13.9 Hypocreadium

J. Parasitol, 58, p. 222);
Description: Body greatly flattened, broader than long, appearing as disc sometimes
with folded lateral edges; length 992–1504; width 1200–1600, spines probably lost.
Oral sucker 128–156 in diameter. Ventral sucker 140–163 in diameter, in anterior
third of body. Sucker ratio 1:1.1–1.2. Forebody 320–484 with scattered granules of
eyespot pigment. Prepharynx absent. Pharynx oval. Oesophagus as long as pharynx.
Caeca arcuate with inwardly turned posterior ends, terminate about halfway between
ovary and posterior end of body. Testes elongate, symmetrical, intercaecal, at mid-
body, with smooth or irregular margins. Genital pore immediately to left of posterior
end of oesophagus. External seminal vesicle dextral to ventral sucker. Cirrus-sac lies
obliquely along median side of right caecum, never touches ventral sucker. Inter-
nal seminal vesicle small, prostatic vesicle sinuous, cirrus long and sinuous. Ovary
compact or slightly lobed, intertesticular, separated from ventral sucker by uterus.
Seminal receptacle long, extends to left of ventral sucker. Vitellarium extends from
level of pharynx along each caecum, confluent in post-testicular space, situated both
lateral and median to caeca but absent along edges of body. Uterus preovarian. Forms
thick-walled metraterm about half as long as cirrus-sac, anterior to ventral sucker.
Eggs large, oval, 62–66 long and 35–39 wide.
Remarks: Madhavi (1972) collected these worms from Monacanthus choiro-

cephalus from BOB, and included them in the genus Pseudocreadium as P.indicum
Madhavi, 1972. Hussain et al. (1986) included the species in the genus Hypocrea-
dium as H. indicum (Madhavi, 1972) and redescribed the species based on material
collected from ‘Abalistes stellaris’ from BOB. However, Bray and Cribb (1996) con-
sidered the worms reported by Hussain et al. as H. indicus, to be similar to H. cavum
Bray and Cribb, 1966.
Hypocreadium patellare Yamaguti, 1938 (Fig. 13.10)
Host: Acanthuridae: Sufflamen fraenatum (Latrielle)
Locality: Tuticorin, AS
Distribution: Japan (Yamaguti, 1938); Great Barrier Reef (Bray and Cribb, 1996;
Bray et al. 2009); New Caledonia (Bray and Justine, 2012)
References: Hafeezullah (1970b)
Fig. 13.10 Hypocreadium patel-

lare (after Yamaguti,
1938)
Description: Body circular with a distinct anterior notch. Oral sucker oval, subter-
minal. Ventral sucker bigger than oral sucker, just preequatorial. Intestinal bifurca-
tion in mid-forebody. Caeca narrow, arcuate around, converge posteriorly, terminate
blindly near midline. Testes 2, irregularly oval, symmetrical, in anterior half of hind-
body. External seminal vesicle saccular. at level of ventral sucker. Cirrus-sac large,
situated obliquely across posterior forebody and mid-ventral sucker. Internal seminal
vesicle oval, pars prostatica bipartite, ejaculatory duct long, folded. Genital atrium
large. Genital pore sinistral at level of bifurcation. Ovary oval, intertesticular. Sem-
inal receptacle large, saccular. Uterus passes into post-ovarian region. Eggs 66 ×
34. Vitellarium follicular, follicles numerous, surround gonads, well separated from
body margins.
Remarks: Hafeezullah (1970b) identified the forms collected from Sufflamen frae-
natum as H. patellare. According to him, except for certain body measurements,
shape of cirrus-sac and slight difference in the position of genital opening, the forms
agreed fairly well with Yamaguti’s description, based on material collected from
Japan. Bray et al. (2009) found morphological variations, designated as Atypical
A and Atypical B, in the forms collected from Great Barrier Reef and called for
molecular studies to determine whether these variations are host induced or genetic.
H. patellare is characterised by the presence of a distinct anterior notch, the uterus
reaches into post-ovarian region, and the ovary is intertesticular (Bray et al. 2009).
Key to Indian species of Hypocreadium:
1. Anterior region of body with a distinct anterior notch ……………………….. 2
Anterior region of body without any notch …………………….....……….. H.
indicum (Madhavi, 1972)
2. Uterus passes into post-ovarian region ………………………H. patellare Yam-
aguti, 1938
Uterus preovarian ……………………………….. H. cavum Bray and Cribb,
1996
Genus Karyakartia Hafeezullah, 1980
Generic diagnosis: Body elongate. Tegument spinose. Oral sucker terminal, fun-
nel or bell-shaped, surrounded by ring of tentacles or lobes around mouth. Ventral
sucker pre-equatorial. Prepharynx and pharynx present. Oesophagus short. Caeca

simple, forming cyclocoel near posterior end; anus or cloaca not formed. Testes two,
tandem, post-equatorial. External seminal vesicle present. Cirrus-sac antero-dorsal
to ventral sucker, encloses internal seminal vesicle, pars prostatica, surrounded by
well-developed prostate cells and cirrus. Genital pore near intestinal bifurcation.
Ovary pretesticular. Seminal receptacle present. Vitellarium follicular, extensive.
Uterus preovarian. Metraterm present. Eggs few. Excretory vesicle tubular; pore ter-
minal. Intestinal parasites of marine fishes. Type species: Karyakartia pambanense
(Karyakarte, 1968).
Karyakartia pambanense (Karyakarte, 1968) Hafeezullah, 1980 (Fig. 13.11 a, b)
Syn: Acanthostomum pambanense Karyakarte, 1968
Host: Terapontidae: Terapon theraps (Cuvier); T. jarbua (Forskal), T. puta Cuv.
Locality: Pamban, GOM; Machilipatnam, BOB
References: Karyakarte(1968); Hafeezullah and Chakrabarti (1994)
Description: Body 2200–3000 long, 240–520 wide, cylindrical. Tegument spinose.

Oral sucker 185–210 long, 161–168 wide at anterior end of body, funnel or bell-
shaped, terminal. Oral lobes or tentacles 19–21, arise from rim of mouth, anteriorly
directed. Ventral sucker 147–161 in diameter, spherical, situated at one-fourth of
body from anterior end. Intestinal bifurcation between pharynx and ventral sucker.
Caeca simple, reach posterior end of body, uniting to form cyclocoel. Anus or cloaca
not formed. Testes two, entire, tandem, separated from each other by vitelline folli-
cles in some specimens, post-equatorial. External seminal vesicle tubular, winding,
extends from posterior part of ventral sucker to well posterior to it. Cirrus-sac bul-
bous, anterodorsal to ventral sucker, encloses internal seminal vesicle, pars prostatica
surrounded by well-developed prostate cells and muscular cirrus. Genital pore near
intestinal bifurcation. Ovary globular, pretesticular. Seminal receptacle immediately
Fig. 13.11 a Karykartia (a)

pambanense (after
Hafeezullah & Dutta, 1998)
b Karyakartia pambanense (b)
Oral sucker
posterior to ovary. Vitellarium follicular, extensive, extend from midway between

ventral sucker and ovary to posterior end of body, may intrude between two testes,
and anterior testis and ovary. Uterus scanty, preovarian, winding. Metraterm not so
well differentiated. Eggs few, 70–73 × 45–52, yellowish, thick-walled.
Remarks: This form originally described by Karyakarte (1968) from Terapon puta
from Pamban, BOB as Acanthostomum pambense Karykarte, 1968, was redescribed
and placed in the new genus Karyakartia by Hafeezullah (1980a) who collected the
forms from Terapon jarbua (Forskal) from Machilipatnam coast, BOB. The form
shows characters intermediate between the families Enenteridae and Lepocreadi-
idae. Hafeezullah (1980a) preferred inclusion under Enenteridae while Bray (2005)
and Abdou et al. (2006) placed it under Lepocreadiidae. Gibson and Bray (1982)
expressed the view that Godavaritrema marina Karyakarte and Yadav, 1976 recorded
from Terapon theraps at Karwar (AS) as a possible synonym of K. pambanense but
Bray (1986) stated that it ‘bears little resemblance to Karyakartia’.
Genus Lepocreadioides Yamaguti, 1936
[Syn. Bicaudum Bilqees, 1971]
Diagnosis: Body broadly oval, may be notched posteriorly. Ventral sucker in anterior
half of body. Prepharynx short. Pharynx large. Intestinal bifurcation in mid-forebody.
Caeca long. Testes two, oval, oblique to symmetrical, in mid-hindbody. External
seminal vesicle tubular or oval in forebody or overlaps ventral sucker. Cirrus-sac
claviform, in forebody or overlaps ventral sucker, encloses oval seminal vesicle, pars
prostatica and long ejaculatory duct. Genital pore sinistrolateral to oral sucker. Ovary
trilobed, anterior or anterosinistral to anterior testis. Uterus pretesticular, Vitelline
fields extend from oesophageal level to posterior extremity of body. Excretory vesi-
cle saccular, passes between testes, reaches bifurcal level. Type species: L. zebrini
Yamaguti, 1936.
Four species belonging to this genus have been recorded from Indian marine
fishes: L. indicus Srivastava, 1941; L. orientalis Park, 1939; L. srivastavai Gupta and
Mehrotra, 1970 and L. thapari Gupta and Govind, 1983. Hafeezullah and Chakrabarti
(1994) based on extensive intraspecific variations found in individuals of L. orientalis,
especially between young and old individuals, synonymized the remaining three
Indian species of the genus with L. orientalis.
Lepocreadioides orientalis Park, 1939 (Fig. 13.12)
[Syn: L. indicus Srivastava, 1941, L. srivastavai Gupta and Mehrotra, 1970, L. thapari
Gupta and Govind, 1984; Bicaudum otolithi Bilqees, 1970]
Hosts: Platycephalidae: Platycephalus indicus (1); Cynoglossidae: Cynoglossus lida
(Bleeker) (2–6, 8); C. bilineatus (Bloch)(4,8); C. dubius (Day) (4,6); C. lingua (Ham-
Busch.) (4); C. macrolepidotus (Bleeker) (4); C. puncticeps (Richardson) (4); C.
sindensis (Day)(4); C. arel (6); C. cynoglossus (7); C. macrostomus Norman(6);
C. puncticeps (Richardson) (6); Soleidae: Aesopia cornuta Kaup; Soleidae: Zebrias
altipinnis (Alcock); Z. synapturoides (Jenkins) (6)
Fig. 13.12 Lepocreadioides

orientalis (after Hafeezullah
and Chakrabarti, 1994, Bull.
Zool. Surv. India, 94, 5)
Locality: East and West coasts of India

References: 1. Srivastava (1941d); 2. Madhavi (1972); 3. Madhavi et al. (1986), 4.
Hafeezullah (1970b); 5. Hafeezullah and Chakrabarti (1994); 6. Bijukumar (1997);
7. Gupta, N.K. and Mehrotra (1970c); 8. Gupta, P.C. and Govind (1983)
Distribution: Indo-Pacific
Description: Body foliate, 1220–2940 long, 1580–1440 wide, bluntly pointed ante-
riorly, broadly rounded posteriorly, lateral margins crenulated. Tegument with minute
spines in anterior region. Oral sucker 80–130 long 90–190 wide. Ventral sucker usu-
ally smaller than or equivalent to oral sucker. Oesophagus short, narrow. Caeca
terminate blindly anterior to posterior end of body. Genital pore marginal, near oral
sucker on left side. Genital atrium shallow to slightly tubular. Testes ovate or ellip-
tical, postequatorial, asymmetrical, intercaecal. Cirrus-sac club-shaped with elon-
gate neck, extends obliquely from genital atrium to about level of anterior margin
of ventral sucker, encloses internal seminal vesicle, pars prostatica surrounded by
prostate gland cells, ejaculatory duct and cirrus. External seminal vesicle varies in
shape, saccular to tubular, between posterior part of cirrus-sac and ventral sucker.
Ovary 3-lobed, sinistral, anterior to posterior testis. Uterine coils few, between ven-
tral sucker and testes. Metraterm well developed. Seminal receptacle dorsal to ovary.
Vitelline follicles lateral, along caeca, anterior limit varying between ventral sucker
and intestinal bifurcation, surrounding caeca posteriorly in post-testicular region.
Remarks: Yamaguti (1936) erected the genus Lepocreadioides with L. zebrini as

the type species described from a soleid fish Zebrias zebrinus (Temm. et Schleg.)
from the East China Sea and the Sea of Japan. The genus is mainly characterized by
the marginal position of the genital pore on left side and near the oral sucker. Other
species reported in the genus are: Lepocreadioides indicum Srivastava, 1941, in Platy-
cephalus insidiator from the coasts of Puri and Karachi; subsequently reported by
Hafeezullah (1970b) and Madhavi (1972) in Cynoglossus spp. from Arabian Sea and
Bay of Bengal; Lepocreadioides srivastavai Gupta and Mehrotra, 1970, in Cynoglos-
sus cynoglossus and Cynoglossus lingua from Ernakulum coast, Arabian Sea; Lep-
ocreadioides sp. of Karyakarte and Yadav (1976d), in Cynoglossus oligolepis from
Ratnagiri coast, Arabian Sea. Gupta and Govind (1983) described Lepocreadioides
thapari from the fish Cynoglossus lida from the Puri coast. Hafeezullah from a study
of five populations of Lepocreadioides specimens, from Cynoglossus sp. from the
Arabian Sea and the Bay of Bengal found morphological variations in respect of
many characters and concluded that there are no pronounced morphological dif-
ferences between these species and considered them as possible synonyms of L.
orientalis. Apart from Indian ocean, L. orientalis has been recorded from Iranian
marine waters (Bannai and Muhammad, 2015), China and Australia (see Bray and
Cribb, 1998).
Genus Lepocreadium Stossich, 1903 Three species of Lepocreadium have been

reported from Indian marine fish, including L. guptai Gupta & Gupta, 1987 from
Rastrelliger kanagurta (Cuvier) from Puri Coast; L. panajiense Gupta and Sayal,
1979 from Gerres abbreviata Bleeker from Goa, AS and L. puriensis Gupta &
Gupta, 1987 from Rastrelliger brachysoma (Bleeker). Bray & Gibson (1990) while
reviewing the genera Opechona and Prodistomum, considered L. guptai as a synonym
of Opechona bacillaris (Molin), L. puriense as a synonym of Prodistomum orientalis.
L. panajiense was regarded as a species inquirendum, but later thought it possibly
to be an apocreadiid of the genus Neoapocreadium Siddiqi & Cable, 1960 by Bray
(2005a).
Genus Lobatocreadium Madhavi, 1972

The genus includes Lepocreadium-like species with a squat oval body, tandem to
slightly oblique overlapping testes situated close to the posterior extremity of the
body, the uterus mainly between the ovary and the testes and a follicular ovary
consisting of two small lateral bunches.
Diagnosis: Body squat, oval. Oral sucker subterminal. Ventral sucker in about mid-
body. Pharynx with lobed anterior margin. Caeca long. Genital pore sinistral, bifur-
cal. Testes tandem, overlapping dorsoventrally, close to posterior extremity. External
seminal vesicle saccular. Cirrus-sac claviform, encloses seminal vesicle, pars prostat-
ica and cirrus. Ovary follicular, in two lateral groups joined by isthmus, pretesticular.
Uterus pretesticular. Vitellarium follicular, extensive. Eggs oval, operculate. Excre-
tory bladder reaches to testes. Type species: L. manteri Madhavi, 1972.
Four species of this genus have been reported from India: L. manteri Madhavi,
1972; L. gupti Ahmad, 1981; L. indicum Ahmad, 1983; L. balistes Hussain, Rao and
Shyamasundari, 1986.
Lobatocreadium balistes Hussain, Rao & Shyamasundari, 1986 (Fig. 13.13)
Host: Balistidae: Abalistes stellatus (Anonymous)
Locality: VSK, BOB
Fig. 13.13 Lobatocredium

balistes (after Hussain, Rao
& Shyamasundari, 1986)
Description: Body elongated, plump, 2384–2488 in length and 1168–1600 in width.

Oral sucker 368–400 × 416–512. Ventral sucker large, 400–592 × 432–608, anterior
to the midlevel of body. Sucker ratio 1:1.1–1.9. Prepharynx short. Pharynx large with
eight lobe-like structures projecting from anterior margin into lumen. Oesophagus
short and wide. Intestinal bifurcation between two suckers. Caeca broad extend to
posterior extremity. Genital pore median at level of intestinal bifurcation, surrounded
by circular muscles. Testes two, ovoid, tandem, slightly overlapping in posterior half
of body. Cirrus-sac elongated, clavate, straight, between intestinal bifurcation and
ventral sucker, encloses small saccular internal seminal vesicle, elongated prostatic
vesicle and eversible cirrus. External seminal vesicle elongate, saccular, narrow ante-
riorly, situated dorsal to ventral sucker, extends along its length. Ovary near posterior
margin of ventral sucker, contains oval follicles connected by narrow stalks arranged
in two groups, 5–7 follicles grouped on left side, 3–5 follicles on right side. Seminal
receptacle elongate, post-ovarian. Vitelline follicles, small, extensive, from level pos-
terior to pharynx to posterior extremity, more confluent posteriorly. Uterus pretestic-
ular between ovary and testis, overlapping ventral sucker dorsally. Metraterm narrow
becoming bulbous before opening into genital atrium. Eggs 64–80 × 48–64.
Remarks: L. balistes resembles the type species L. manteri Madhavi, 1972, in many
respects of its organization but differs from it in the sucker ratio, the ventral sucker
being larger than the oral sucker, in having a median genital pore situated closely
posterior to the intestinal bifurcation, a straight cirrus-sac between the pharynx and
the ventral sucker and tandem testes not close to the posterior extremity.
Lobatocreadium gupti Ahmad, 1981(Fig. 13.14)

Host: Pomacanthidae: Pomacanthus annularis (Bloch.)
Locality: BOM, AS
No. 9
References: Ahmad (1981c)
Description: Body flattened, broadly rounded at each end, 1920–2075 long and
1200–1245 wide at level of ovary. Entire body spinose. Eyespot pigments present.
Oral sucker spherical, subterminal l260–285 in diameter. Ventral sucker median, near
midbody, same size as oral sucker. Pharynx with eight lobe-like structures projecting
from anterior margin into lumen. Oesophagus absent. Intestinal bifurcation dorsal,
nearer to oral sucker than to ventral sucker. Caeca simple, bow outward, then back-
ward, end blindly at midlevel of anterior testis or anterior margin of posterior testis.
Testes two, smooth, tandem, contiguous, near hind end of body. Post-testicular space
absent. Cirrus-sac claviform, disposed obliquely between genital pore and ventral
sucker. External seminal vesicle saccular, extends posteriorly as far as anterior mar-
gin of ventral sucker. Genital pore sinistral, just post-bifurcal. Ovary follicular, oval
follicles connected by narrow stalk, arranged in two groups, 4–5 follicles grouped
on right side and 5–6 follicles on left side. Seminal receptacle saccular, preovarian.
Uterus runs between anterior border of posterior testis and cirrus-sac. Metraterm
tubular, thick–walled. Eggs oval, operculate, 45–56 long and 23–34 wide. Vitelline
follicles small, rounded, in two broad lateral zones, extend from intestinal bifurcation
to just anterior to caecal ends. Excretory vesicle short, reaches to testis; excretory
pore terminal.
Fig. 13.14 Lobatocreadium

gupti (after Ahmad, 1981)
Remarks: This species was differentiated from the type species L. manteri in having
suckers of equal size, the vitellarium extending from the intestinal bifurcation to just
anterior to the caecal ends and the smaller eggs.
Lobatocreadium indicum Ahmad, 1981 (Fig. 13.15)
Host: Triacanthidae. Triacanthus biacetabulatus(Bloch)
Locality: PR, BOB
References: Ahmad (1981c)
Description: Body flattened, 1972–2270 long by 1180–1256 wide at level of ovary.

Eyespot pigment present. Oral sucker oval, 274–320 long by 330–387 wide. Ventral
sucker near midbody, 200–239 in diameter. Sucker ratio 1: 0.67–0.70. Prepharynx
absent. Pharynx oval, muscular, with 8 lobe-like structures projecting from anterior
margin into lumen. Oesophagus very short. Intestinal bifurcation dorsal, nearer to
oral sucker than to ventral sucker. Caeca bowing outward, then backward, ending
blindly at posterior edge of anterior testis. Testes two, smooth, tandem, intercaecal,
near hind end of body, posterior testis overlapping posterior border of anterior testis.
Post-testicular space absent. Cirrus-sac clavifom, disposed obliquely between genital
pore and ventral sucker, extends posteriorly to just anterior to ventral sucker. Genital
pore extracaecal, lateral to left caecum, at bifurcal level. Ovary follicular, follicles
90–120 long by 40–55 wide, oval follicles connected with narrow stalks, arranged in
two groups, 4–6 follicles grouped on left side and 6–9 follicles on right side. Seminal
receptacle saccular, postovarian. Vitellarium small, in two broad lateral zones, extend
from well posterior to intestinal bifurcation to posterior level of ovary. Uterus runs
between cirrus-sac and anterior testis. Metraterm undifferentiated. Eggs thin-shelled,
39–44 long by 22–25 wide. Excretory vesicle wide and short, reaches to testes; pore
subterminal.

indicum (after Ahmad, 1981)
Remarks This species differs from L. manteri in the smaller sucker ratio, the tandem
testes, the longer external seminal vesicle, the postovarian seminal receptacle and
the extracaecal genital pore.
Lobatocreadium manteri Madhavi, 1972 (Fig. 13.16)

Host: Balistidae: Sufflamen fraenatum (Latrielle); S. chrysopterum(Bl. and Schn.)
Locality: VSK, BOB
No. 20
References: Madhavi (1972)
Description: Body plump, broadly rounded at both ends, 720–1280 long and
510–976 wide; Tegument spined. Eyespot pigment not evident. Oral sucker 117–284
long by 156–296 wide, subterminal. Ventral sucker near midbody, 144–230 in diam-
eter; sucker ratio 1:0.76–0.87. Prepharynx short. Pharynx large, highly muscular,
spherical or broader than long, with 8 lobe-like structures projecting from ante-
rior margin into lumen. Oesophagus short and wide. Intestinal bifurcation midway
between suckers or slightly more anterior. Caeca broad, bowing outward, then inward,
ending blindly near testes. Genital pore ventral to left caecum, bifurcal; surrounded
by circular muscles. Testes ovoid, broader than long, overlapping, near posterior
end, touching posterior edge of body. Cirrus-sac clavate, situated obliquely or hor-
izontally in space between ventral sucker and intestinal bifurcation, encloses oval
seminal vesicle, elongated prostatic vesicle, and eversible cirrus; external seminal
vesicle ovoid, situated dorsal to ventral sucker. Ovary near posterior margin of ventral
sucker, oval follicles connected by narrow stalks, arranged in 2 groups, 5–7 follicles
grouped on left side and 3–4 follicles on right side. Seminal receptacle oval, pre-
ovarian, partly overlapping ventral sucker. Uterus pretesticular. Metraterm narrow,
becoming bulbous before opening into genital atrium. Eggs large, 58–62 long by 27
wide. Vitelline follicles small, in 2 broad lateral zones from level of oral sucker to
level of testes; prominent vitelline duct lies between 2 ovarian groups.

manteri (after Madhavi,
J. Parasitol, 58, p. 218)
Key to the species of Lobatocreadium

1. Vitelline fields extending from anterior region to posterio extremity
…………………. 2
Vitelline fields not extending beyond testes ………………......................……. 3
2. Genital pore median, at base of pharynx ……………L. balistes Hussain, Shya-
maundari and Rao,1986
Genital pore bifurcal, ventral to left caecum …………………....… L. manteri
Madhavi, 1972
3. Sucker ratio 1:1, eggs 45–56 × 23–34. …………………………… L. gupti
Ahmad, 1981
Sucker ratio 1: 0.67–0.70, eggs 39–44 × 22–25 …………………….....….........
L. indicum Ahmad, 1983
Genus Lutianotrema Singh, Singh & Singh, 2004
Two species are described in the genus Lutianotrema erected by Singh, Singh
& Singh (2004), including the type species L. quinqueliai Singh, Singh & Singh,
2004 and L. kanyakumariae Chaubey, Singh & Jaiswal, 2006, both from the snap-
per, Lutjanus quinquelineatus (Bloch), from the Bay of Bengal (Singh et al. 2004;
Chaubey et al. 2006). The characters of Lutianotrema fit into the genus Hypocre-
adium in almost all details except for the dorsal location of oral sucker which is
an unusual feature. Bray et al. (2009) pointed out the similarity of Lutianotrema to
Hypocreadium but needed confirmation of the dorsal position of oral sucker before
proposing the synonymy. In view of this controversy, details of the two species of
Lutianotrema are not included in this account.
Genus Neohypocreadium Machida & Uchida, 1987
The genus Neohypocreadium erected by Machida & Uchida (1987) is mainly
characterized by the dorsal location of the genital pore. Bray et al. (1994) examined
the voucher specimens of Preptetos chaetodoni Madhavi, 1972 and found the gen-
ital pore to be dorsal in location and accordingly transferred P. chaetodoni to the
genus Neohypocreadium as N. chaetodoni (Madhavi, 1972). They further regarded
P. overstreeti Ahmad, 1984 as a synonym of this species.
Diagnosis: Body round to oval. Tegument spined. Oral sucker subterminal. Ventral
sucker pre-equatorial. Pharynx moderately developed. Caeca arcuate, terminate near
posterior extremity of body. Testes round to oval, symmetrical, in anterior region
of hindbody. External seminal vesicle tubular, mainly in forebody. Cirrus-sac elon-
gate oval, may reach into hindbody. Internal seminal vesicle oval. Pars prostatica
oval, ejaculatory duct long. Genital pore dorsal, sinistral to pharynx. Ovary trilobed,
intertesticular. Uterus short, pretesticular. Vitelline follicles between level of oesoph-
agus or intestinal bifurcation and posterior extremity. Excretory vesicle inverted L-
shaped. Type species: N. longisaccatum Machida & Uchida, 1987.
Neohypocreadium chaetodoni (Madhavi, 1972) Bray, Cribb & Barker, 1994

(Fig. 13.17)
[Syn: Preptetos chaetodoni Madhavi, 1972; Preptetos overstreeti Ahmad, 1984]
Fig. 13.17 Neohypocreadium

chaetodoni (after Madhavi
Host: Chaetodontidae: Chaetodon pictus Forsskal

Number: 94 from 6 of 8 hosts
Locality: VSK BOB (1); Panjim, Goa, AS (2)
References: 1. Madhavi (1972); 2. Ahmad (1984b); Bray et al. (1994).
Description: Body fusiform, 1488–2640 long and 832–1552 in maximum width.

Tegument spined to level of ovary, spines often lost. Oral sucker 78–117 long by
86–137 wide. Ventral sucker in anterior third of body, 94–203 by 98–168. Sucker
ratio 1:1.2–2.0. Prepharynx absent. Pharynx muscular. Oesophagus long. Intestinal
bifurcation about midway between suckers. Caeca narrow anteriorly, broader pos-
teriorly terminate blindly at posterior end. Genital pore to left of caecum at level of
intestinal bifurcation, dorsal. Testes large, smooth or crenulated, longer than wide,
equal or subequal, separated by excretory vesicle, situated in middle of hindbody.
External seminal vesicle large, sinuous, tubular, extends posterior to ventral sucker to
about midway between ventral sucker and ovary. Cirrus-sac stout, extends obliquely
in forebody, contains small seminal vesicle, oval prostatic vesicle, and muscular pro-
trusible cirrus. Genital atrium large. Ovary median, deeply lobed, to left of anterior
testis or slightly more anterior. Seminal receptacle large, between ovary and pos-
terior testis. Uterus preovarian, confined to left side, not extending beyond caeca,
modified terminally into short, thick metraterm opening into genital atrium. Eggs
large, oval, 59–66 long by 35–43 wide. Vitelline follicles moderately large, extend
from level of intestinal bifurcation to posterior end of body, circumcaecal, fill most of
post-testicular space except region occupied by excretory bladder. Excretory bladder
inverted L-shaped, long, extends between testes to anterior margin of ventral sucker
and then curves round anterior margin of ventral sucker.
Remarks: N. chaetodoni (Madhavi, 1972) is the only species of the genus reported
from India. Preptetos overstreeti Ahmad, 1984, also recorded from Chaetodon pictus
but from Goa coast, was regarded as a synonym of this species by Bray et al. (1994).
Genus Neomultitestis Machida, 1982
Diagnosis: Body transversely oval. Eyespot pigment present. Ventral sucker trans-
versely oval, in midbody. Prepharynx, pharynx and oesophagus present. Intestinal
bifurcation in posterior part of forebody. Caeca arcuate, open as ani near posterior
extremity. Testes numerous, 13–80, oval to subglobular, arranged in single group
or two lateral groups at level of ventral sucker. External seminal vesicle saccular,
overlaps ventral sucker. Cirrus-sac claviform, lies transversely in posterior part of
forebody. Internal seminal vesicle oval, ejaculatory duct long. Genital pore sinis-
tral at level of pharynx or intestinal bifurcation. Ovary follicular or multilobate,
mainly post-testicuar. Uterus forms narrow coils, may pass into postovarian region.
Metraterm tubular or infundibuliform, with gland cells. Eggs thin-shelled. Vitelline
fields reach from pharyngeal level to posterior extremity. Excretory vesicle Y-shaped
or saccular. Type species: N. palauensis Machida, 1982.
Neomultitestis bengalensis (Madhavi, 1972) Machida, 1982(Fig. 13.18)
[Syn: Multitestis bengalensis Madhavi, 1972]
Host: Ephippidae: Platax teira (Forskal)
No: 13
Locality: VSK, BOB
References: Madhavi (1972); Machida (1982)
Distribution: Bay of Bengal, Hainan Island (China)
Description: Body transversely ovate, 800–1010 long and 910–990 wide. Oral
sucker 93–117 long by 136–167 wide. Ventral sucker larger than oral sucker, wider
Fig. 13.18 Neomultitestis

bengalensis (after Madhavi,
than long, discoidal, 192–288 long by 480–576 wide. Sucker ratio 1:2.7–3.5. Eye-
spot pigment present on either side of pharynx. Prepharynx short. Pharynx ovoid.
Oesophagus as long as pharynx. Intestinal bifurcation immediately anterior to ventral
sucker. Caeca arch around ventral sucker and open through anal pores just anterior
to posterior extremity. Genital pore median, close to left body margin, at level of
posterior end of pharynx. Testes more than 80 follicles, occupy middle third of body
dorsal and lateral to ventral sucker, extend slightly lateral to caeca. External seminal
vesicle long, tubular. Cirrus-sac long, slender, lying almost transversely and touching
anterior margin of ventral sucker. Ovary follicular, dorsal to posterior third of ventral
sucker, slightly intrudes into testicular field. Seminal receptacle large, preovarian
or postovarian. Uterus extends slightly posterior to ovary, mostly dorsal to ventral
sucker. Metraterm long, thick-walled, anterolateral to ventral sucker. Eggs oval, 58
long and 28 wide. Vitelline follicles small, numerous, fill entire body except central
area dorsal to ventral sucker and lateral to oral sucker.
Remarks: The genus Neomultitestis Machida, 1982, is characterized by the follow-

ing combination of characters: the caeca open dorsally through ani; the numerous
testes form a single group or two lateral groups, the discoidal ventral sucker and
the transversely oval body. Three species have been reported under the genus: N.
palaunensis Machida, 1982, N. bengalensis (Madhavi, 1972) and N. aspidogastri-
formis Bray & Cribb, 2003. Species of the genus show specificity towards Platax
teira (Ephippidae). N. bengalensis has also been recorded from Hainan Island, China
(Shen, 1990).
Genus Opechona Looss, 1907

The definition of the genus Opechona Looss, 1907 as given by Bray & Gibson
(1990) is adopted here. According to these authors, the genera Opechona Looss,
1907 and Prodistomum Linton, 1910 differ only in one character, the caeca form a
uroproct in Opechona while they terminate blindly in Prodistomum. Following this
definition, all the species of Opechona, described as having blind caeca were either
transferred to the genus Prodistomum or considered as species inquirendae pending
further study of the nature of the caecal termination. From marine fish of India, nine
species of Opechona have been reported. Of these O. buckleyi Gupta & Ahmad,
1977 was regarded as a synonym of Lepidapedon nelsoni Gupta & Mehrotra, 1969.
Opechona waltairensis Madhavi, 1972, was included in the genus Prodistomum as
P. waltairense (Madhavi, 1972) Bray & Gibson, 1990. O. bacillaris (Molin, 1859)
was reported by Madhavi (1972) from Rastrelliger kanagurta. O. ahmadi where
caecal termination is not clear is tentatively retained in the genus Opechona. We are
transferring the remaining six species of the genus, where the caeca are reported as
terminating blindly, into the genus Prodistomum.
Diagnosis: Body oval to elongate. Oral sucker infundibuliform or rounded. Prephar-

ynx distinct. Pharynx oval. Oesophagus short to long. Pseudoesophagus short to long.
Intestinal bifurcation in forebody. Caeca reach posterior extremity. Uroproct present.
Testes two, oval, smooth or lobed, tandem, in mid or posterior hindbody. External
seminal vesicle saccular. Cirrus-sac claviform, thick-walled. Internal seminal vesicle
saccular, pars prostatica oval, ejaculatory duct long, sinuous. Genital atrium distinct,
genital pore close to ventral sucker, sinistral. Ovary lobed or entire, pretesticular.
Uterus preovarian. Metraterm distinct. Vitelline fields from posterior extremity to
anterior region of hindbody. Excretory vesicle I-shaped, extends into forebody. Type
species: O. bacillaris (Molin, 1859) Looss, 1007.
Opechona ahmadi Gaevskaya, 1990 (Fig. 13.19)
[Syn.O. siddiqii Ahmad 1986 nec O. siddiqi Ahmad, 1984; O. pritchardae Ahmad,
1991]
Host: Haemulidae: Pomadasys maculatus (Bloch)
Locality: Goa, AS.
Number: 7
References: Ahmad (1986)
Description: Body elongate, 790–940 long, 135–180 wide; widest at level of testes.
Tegument spinose to ovary. Oral sucker 50–70 long, 58–80 wide. Ventral sucker
smaller than oral sucker, pre-equatorial, 40–60 in diameter. Sucker ratio 1: 0.80–0.85.
Oesophagus short. Pseudoesophagus of similar length to oesophagus. Intestinal
bifurcation in mid-forebody. Caeca simple, long, termination not known. Testes two,
lobed, contiguous, in posterior half of body. External seminal vesicle saccular. Cirrus-
sac club-shaped, extends from genital pore posteriorly usually halfway between
ventral sucker and ovary. Genital pore submedian, anterior to anterior margin of ven-
tral sucker. Ovary globular, submedian, to right of median line, pretesticular. Seminal
Fig. 13.19 Opechona

ahmadi (after Ahmad, 1986)
receptacle saccular. Uterus preovarian, with few eggs. Metraterm thick–walled.

Eggs 80–90 × 45–55. Vitellarium follicular, confined to hindbody. Excretory vesicle
tubular, extends anteriorly as far as intestinal bifurcation. Excretory pore terminal.
Remarks: The specific name of this species O. siddiqii Ahmad, 1986, is a junior
homonym of O. siddiqii Ahmad, 1984. Gaevskaya (1990) named it as O. ahmadi,
while Ahmad (1991) renamed it as O. pritchardae. This species differs from most
other speces of Opechona, in the lobed gonads, the very short post-testicular space
and fewer eggs in the uterus.
Opechon bacillaris (Molin, 1859) Dollfus, 1927(Fig. 13.20)
Syn: Lepocreadium guptai Gupta & Gupta, 1987 (Vide Bray & Gibson,1990)
Locality: VSK(1, 2) and Puri (3), BOB
Distribution: Widely distributed in tropical and subtropica seas
References:1. Madhavi (1972); 2. Madhavi et al. (1986); 3. Gupta, P.C. & Gupta,
V.C. (1987b)
Fig. 13.20 Opechona

bacillaris (after Madhavi &
TriveniLakshmi, 2011)
Description: Body elongate, 1600 × 4,30 long 280–540 wide. Oral sucker large,
infundibuliform, 159–379 × 178–284. Ventral sucker smaller than oral sucker,
90–221 × 130 × 216. Sucker ratio 1: 0.74–0.76. Prepharynx short. Pharynx large doli-
iform. Oesophagus short. Pseudoesophagus long. Intestinal bifurcation in posterior
forebody. Caeca narrow, reach to posterior end of body, form uroproct. Testes oval,
tandem, in posterior half of hind body. External seminal vesicle saccular, recurved.
Cirrus-sac claviform, reaches well into hindbody. Genital pore close to or anterior
to ventral sucker. Ovary pretesticular, trilobed. Seminal receptacle large, between
ovary and testis. Uterus between ovary and cirrus-sac. Metraterm as long as cirrus-
sac. Eggs 79 × 29. Vitelline follicles extend anteriorly as far as posterior border of
ventral sucker, confluent ventrally between testes and in post-testicular space. Excre-
tory vesicle I-shaped reaches level of bifurcation.
Remarks: O. bacillaris is cosmopolitan in distribution. Bray & Gibson (1990) fur-
nished details of hosts, geographic distribution and synonyms of O. bacillaris and
provided a key for separation of valid species of the genus. They regarded Lepocre-
adium guptai Gupta & Gupta, 1987 as a synonym of O. bacillaris. Køie (1975)
described the life cycle of O. bacillaris and reviewed the available literature on the
species. O. bacillaris has been reported from R. kanagurta of Visakhapatnam and
Kaligapatnam coasts (Bay of Bengal) by Madhavi (1972) and Madhavi et al. (1986).
Almost all the reports of O. bacillaris are from scombrid fishes and thus the species
is stenoxenic to scombrids (Bray & Gibson. 1990).
Opechona buckleyi Gupta & Ahmad, 1977
[Synonymized with Lepidapedon nelsoni Gupta & Mehrotra, 1969 by Bray & Gibson
(1990)]
Host: Stromateidae: Pampus argenteus (Euphrasen) (1); Scombridae: Thun-
nus albacares (Bonnaterre) (2)
Locality: Puri, BOB (1); Bombay, AS (2)
References: 1. Gupta, V & Ahmad, (1977), 2. Ahmad (1985a)
Genus Opisthogonoporoides Madhavi, 1972
Diagnosis: Body ovoid. Tegument spined. Ventral sucker large. Caeca extend to
level of testes. Genital pore submedian, posterior to ventral sucker. Testes two sym-
metrical, on opposite sides of body. External seminal vesicle present. Cirrus-sac situ-
ated obliquely posterior to ventral sucker, encloses seminal vesicle, prostatic vesicle,
cirrus and numerous prostatic cells. Ovary lobed, pretesticular. Seminal receptacle
large, intertesticular. Vitellarium extends from oral sucker to ends of caeca, confluent
anteriorly. Uterus pretesticular. Excretory vesicle I-shaped.
Opisthogonoporoides was erected by Madhavi (1972) with O. hanumanthai Mad-
havi, 1972, as the type species. The genus is characterized by the genital pore lying
postero-lateral to ventral sucker, the symmetrical testes situated close to the pos-
terior end of the body and the distribution of vitelline follicles in the lateral fields
between oral sucker and testes. Apart from the type species, two more species of
the genus have been reported from India. These are: O. acanthuri Hussain, Rao &
Shyamasundari, 1986 and O. thapari Ahmad, 1981.
Opisthogonoporoides acanthuri Hussain, Rao & Shyamasundari, 1986 (Fig. 13.21)

Host: Acanthuridae: Acanthurus mata Bennet
Locality: VSK, BOB
No. 2
Description: Body elongate oval, 1008–1120 in length and 576–704 in maximum

width. Oral sucker circular in outline 140–160 × 176–192. Ventral sucker very large
compared to oral sucker, slightly broader than long, equatorial, 336–352 × 320–480.
Sucker ratio 1:3. Pharynx small. Intestinal bifurcation near to ventral sucker. Caeca
long narrow, terminate just pretesticular. Genital pore in hindbody slightly posterior
to ventral sucker, midway between left edge of ventral sucker and left body wall.
Testes elongated, symmetrical but divergent with longitudinal axis lying parallel
to corresponding edges of body. External seminal vesicle saccular, anterior to left
testis, between ventral sucker and left testis. Cirrus-sac elongate, oval to round,
slightly narrow posteriorly and broader anteriorly lying oblique at postero-sinistral
margin of ventral sucker. Internal seminal vesicle small, ovoid. Pars prostatica long,
tubular, extends along anterior edge to almost anterior end of cirrus-sac. Ejaculatory
duct directed posteriorly, forms cirrus. Numerous prostatic cells fill space inside
cirrus-sac. Ovary trilobed, anterior to right testis. Seminal receptacle large, oval to
round intertesticular. Uterus pretesticular, extends along right side of ventral sucker to
intercaecal space of forebody, then turning backwards occupying space anterior to left
testis and opening into genital atrium through short metraterm. Eggs, yellowish, oval
64–80 in length and 80–112 in maximum width. Vitellarium follicular, follicles large,
extend from oral sucker to ends of caeca, mostly extracaecal, sometimes forming
confluent bands one to right anterior to intestinal bifurcation and other posterior to
left caecal end. Excretory vesicle ‘I’ shaped extends to level of ovary.
Fig. 13.21 Opisthogonoporoides

acanthuri (after Hussain,
Rao & Shyamasundari,
1986)
Remarks: O. acanthuri differs from the type species O. hanumanthai Madhavi, 1972,
in the much larger size of the ventral sucker (sucker ratio 1: 3), the shorter intestinal
caeca and in the distribution of vitelline follicles.
Opisthogonoporoides hanumanthai Madhavi, 1972 (Fig. 13.22)
Host: Siganidae: Siganus canaliculatus (Park); Acanthuridae: Acanthurus mata Gun-
ther, Ctenochaetus strigosus (Bennett)
No. Numerous
References: Madhavi (1972)
Description: Body ovate, 1040–1960 long and 640–800 wide. Tegument armed with
scale-like spines. Oral sucker subterminal, 117–176 in diameter. Ventral sucker large,
highly muscular, 195–246 long by 253–273 wide, with muscular plug-like structure
filling lumen. Sucker ratio l: 1.5–1.9. Pharynx and oesophagus present. Intestinal
bifurcation nearer to oral sucker than to ventral sucker. Caeca long, narrow, terminate
in level with anterior margin of testes. Genital pore in hindbody, slightly posterior to
ventral sucker, midway between left edge of ventral sucker and left body wall. Testes
2, elliptical, smooth or irregular, situated in middle of hindbody, symmetrical but
divergent with longitudinal axes lying parallel to corresponding body edges. External
seminal vesicle oval. Cirrus-sac narrow posteriorly and broader anteriorly, lying
obliquely at postero-sinistral margin of ventral sucker. Internal seminal vesicle ovoid,
pars prostatica long, tubular, extends along anterior edge and almost to anterior end
of cirrus-sac; ejaculatory duct directed posteriorly and opening into cirrus; numerous
prostatic cells fill space inside cirrus-sac. Ovary 3- or 4-lobed, anterior to right testis.
Seminal receptacle large, intertesticular. Uterus pretesticular, extends along right side

hanumanthai (after Madhavi
of ventral sucker to intercaecal space of forebody, then turns backwards occupying

space anterior to left testis and opening into genital atrium through short metraterm.
Eggs large, yellowish, oval, 58 long by 25–27 wide. Vitelline follicles large, extend
from oral sucker to ends of caeca, mostly extracaecal, confluent at level of intestinal
bifurcation, forming band anterior to ventral sucker. Excretory bladder I-shaped,
extends to level of ovary, passes between testes, containing hard spherules.
Remarks: This species is characterized by the presence of a plug-like structure filling

the lumen of the ventral sucker and the fairly large ventral sucker,
Opisthogonoporoides thapari Ahmad, 1981(Fig. 13.23)
Host: Scaridae: Scarus ghobban Forsskal
Locality: BOM, AS
References: Ahmad (1981b)
Description: Body 1760–2250 long. 1030–1085 wide at level of ovary, oval, taper-
ing posteriorly and rounded anteriorly. Oral sucker 180–230 in diameter. Ventral
sucker 260–296 in diameter, pre-equatorial, without muscular plug-like structure fit-
ting lumen; aperture oval. Sucker ratio 1:1.28–1.44. Pharynx oval. Oesophagus long.
Intrestinal bifurcation dorsal, usually closer to pharynx than to ventral sucker. Caeca
long, narrow, terminate at anterior margin of testes. Testes two, smooth in outline,
symmetrical, post-equatorial, in middle of posterior third of body. External seminal
vesicle saccular, extends posteriorly to caecal ends or just anterior. Cirrus-sac broad

posteriorly, narrow anteriorly, lying obliquely at postero-sinistral margin of ventral

sucker. Genital pore ventral. sub-median, usually midway between posterior edge
of ventral sucker and left body wall. Ovary multilobed, submedian, immediately
anterior to right testis. Seminal receptacle oval, post-ovarian. Uterus pretesticular,
opens into genital atrium through short, thick-walled metraterm. Eggs large oval, yel-
lowish, 44–50 long, 26.29 wide. Vitelline follicles large, lateral, extracaecal, extend
from posterior level of pharynx to caecal ends, not forming band anterior to ventral
sucker. Excretory bladder tubular, extends to level of ovary, without conspicuous
concretions; pore terminal.
Remarks: This species differs from O. hanumanthai Madhavi, 1972 in having a

smaller sucker ratio, a ventral sucker without a plug-like structure, a multilobed
ovary, the vitellarium extending from the posterior level of the pharynx and not
forming band anterior to the ventral sucker.
Key to species:
1. Plug-like structure fills the lumen of ventral sucker …O. hanumanthai Madhavi,
1972
Plug-like structure absent ………………………………………………….. 2
2. Ventral sucker very large, ovary 3-lobed…..…… O. acanthuri Hussain, Rao &
Shyamasundari, 1986
Ventral sucker of moderate size, ovary multilobed ……….……. O. thapari
Ahmad, 1981
Genus Opisthogonoporus Yamaguti, 1937
Diagnosis: Body elongate. Eye spot pigment present. Ventral sucker in anterior third
of body. Prepharynx short. Pharynx large. Oesophagus distinct. Intestinal bifurca-
tion dorsal to ventral sucker. Caeca reach close to posterior extremity, blind. Testes
two, oval, tandem, contiguous, in posterior half of hindbody. External seminal vesi-
cle narrow, tubular, almost reaches ovary. Cirrus-sac claviform or saccular. Internal
seminal vesicle saccular. Pars prostatica oval, ejaculatory duct long. Genital pore
sinistolateral, distinctly posterior to ventral sucker. Ovary lobed, entire or slightly
indented, pretesticular. Uterus preovarian. Vitellarium extends slightly anterior to
ventral sucker. Type species: O. amadai Yamaguti, 1937.
Opisthogonoporus indicus Mulay, 1976
Host: Carangidae: Caranax carangus (Bloch)
Locality: Maharashtra, AS
References: Mulay (1976)
Remarks: O. indicus differs from O. amadai Yamaguti, 1937, the only other species
in the genus, in that the vitellarium extends into the forebody and the ovary is unlobed.
Bray (2005) retained the species within the genus pro tem.
Genus Preptetos Pritchard, 1960
Diagnosis: Body elongate. Eyespot pigment present. Oral sucker subglobular. Ven-
tral sucker in middle third of body. Prepharynx short to long. Pharynx long. Oesoph-
agus long. Caeca long. Testes two, oval, symmetrical or oblique, inmiddle third of
body. External seminal vesicle without gland cells. Cirrus-sac small, confined to fore-
body, or long extending into hindbody. Internal seminal vesicle oval to subglobular,
pars prostatica oval, ejaculatory duct long or short. Ovary pretesticular, lobed. Uterus
anterior to ovary. Vitelline follicles reach ventral sucker or extend into forebody. Type
species: P. caballeroi Pritchard, 1960.
Four species belonging to this genus have been recorded from India from marine
fishes: P. chaetodoni Madhavi, 1972; P. madrasensis Ahmad, 1981; P. overstreeti
Ahmad, 1984; P. pritchardae Ahmad, 1984. Bray et al. (1994) included P. chaetodoni
in the genus Neohypocreadium Machida & Uchida, 1987 as N. chaetodoni (Madhavi,
1972) and considered P. overstreeti Ahmad, 1984 as a synonym of this species.
Preptetos madrasensis Ahmad, 1981 (Fig. 13.24)
Host: Fistulariidae: Fistularia petimba (Lacepede)
Fig. 13.24 Preptetos

madrasensis (Ahmad, 1981)
Description: Body 2750–3460 long, 1200–1257 wide at level of genital pore,

fusiform, blunt at each end. Tegument spinose throughout. Oral sucker 330–339 in
diameter. Ventral sucker 200–236 in diameter, smaller than oral sucker, intercaecal,
pre-equatorial. Sucker ratio 1: 0.65–0.69. Pharynx oval. Oesophagus long. Intestinal
bifurcation dorsal. Usually closer to pharynx than to ventral sucker. Caeca termi-
nate blindly at posterior end of body. Testes two, smooth, intercaecal, longer than
wide, diagonal, separated by excretory bladder, situated in anterior part of posterior
third of body. External seminal vesicle saccular extends posteriorly up to anterior
border of ventral sucker. Cirrus-sac club-shaped, well anterior to ventral sucker. Gen-
ital pore ventral, immediately post-bifurcal. Ovary 3–lobed, pretesticular, to left of
anterior testis, separated by excretory bladder, post-equatorial, Seminal receptacle
spherical, postovarian. Uterus preovarian, confined to left side, terminally modified
into short thick metraterm, opening into genital atrium. Eggs large, oval 49–55 long,
30–33 wide. Vitellarium extends from some distance post-bifurcal to caecal ends,
circumcaecal, fill post-testicular space except region occupied by excretory bladder.
Excretory bladder tubular, extending to anterior margin of ventral sucker and then
curving round anterior margin of ventral sucker; pore terminal.
Remarks: The following combination of characters serve to identify the species:
the external seminal vesicle extending anterior to the ventral sucker, the sucker ratio
1:0.65–0.69 and the vitellarium extends from the intestinal bifurcation to the posterior
end.
Preptetos pritchardae Ahmad, 1984 (Fig. 13.25)
Locality: AS
Description: Body fusiform, 3625–3715 long 1410–1480 wide. Tegument spinose
throughout. Eyespot pigment granules present on each side of pharynx. Oral sucker
340–375 in diameter. Ventral sucker smaller than oral sucker, 240–265 in diameter,
situated at junction of anterior and middle thirds of body. Sucker ratio 1: 0.7–0.71.
Intestinal bifurcation nearer to oral sucker than to ventral sucker. Caeca long, termi-
nate blindly near posterior end. Testes smooth, oval, diagonal, in posterior half of
body, posterior testis larger than anterior testis. Cirrus-sac club-shaped in forebody,
disposed obliquely between intestinal bifurcation and ventral sucker. External sem-
inal vesicle saccular, extends posteriorly as far as middle of ventral sucker. Genital
pore submedian, bifurcal. Ovary four-lobed, immediately anterior to posterior testis.
Uterus preovarian. Metraterm as long as cirrus-sac. Eggs 50–72 × 44–54. Vitelline
follicles large, mainly extracaecal, extend from posterior border of oral sucker to
posterior end of body.
Remarks: This species differs from all the other species of Preptetos in the follow-
ing respects: the extension of the mainly extracaecal vitellarium from the posterior
border of the oral sucker to the posterior end of the body, the position of the ovary
immediately anterior to the posterior testis, the preovarian seminal receptacle, the
oral sucker being larger than the ventral sucker and the metraterm being as long as
the cirrus-sac.
Fig. 13.25 Preptetos

pritchardae (after Ahmad,
62)
Genus Prodistomum Linton, 1910

[Syn. Acanthocolpoides Travassos, Freitas & Buhrnheim, 1965
Diagnosis: Body elongate. Oral sucker small. Ventral sucker rounded in anterior
third of body. Prepharynx long. Pharynx oval. Oesophagus distinct with tegumental
lining. Pseudoesophagus short to long. Intestinal bifurcation in mid-forebody. Caeca
extend to posterior end of body and terminate blindly. Testes two, oval, entire or lobed,
tandem, in mid to posterior part of hindbody. External seminal vesicle saccular or
coiled. Cirrus-sac claviform. Internal seminal vesicle subglobular. Pars prostatica
oval. Ejaculatory duct usually long, wide. Genital atrium distinct. Genital pore close
to ventral sucker, anterosinitral. Ovary entire to lobed, pretesticular or anterolateral
to anterior testis. Uterus preovarian. Metraterm distinct. Vitelline fields confined to
hindbody or extend into forebody. Excretory vesicle tubular, passes dextral to gonads,
terminates near intestinal bifurcation. Type species: P. gracile Linton, 1910.
The genus is represented by 7 species in marine fish of India, all transferred from
the genus Opechona.
Prodistomum gaevskayae (Ahmad, 1991) n.comb (Fig. 13.26)

[Syn. Opechona gaevskayae Ahmad, 1991]
Host: Fistulariidae: Fistularia petimba Lacepede
Locality: Goa, AS
No: 14
Description: Body elongate, 3730–4550 long by 550–640 wide. Tegument spined.
Eyespot pigment granules present at level of pharynx. Oral sucker 170–200 in diam-
eter. Ventral sucker spherical, same size as oral sucker, pre-equatorial, Sucker ratio
1:1. Pharynx barrel-shaped, provided with three anteriorly directed finger-like almost
transparent thin-walled processes. Oesophagus smaller than pseudo-oesophagus.
Intestinal bifurcation usually midway between suckers. Caeca terminate at poste-
rior end of body. Testes two, tandem, separated from one another, situated in anterior
part of posterior third of body. Cirrus-sac club-shaped, extends well beyond ventral
sucker. External seminal vesicle saccular. Genital atrium oval. Genital pore ven-
tral, submedian, towards right of anterior margin of ventral sucker. Ovary spherical,
pretesticular, separated from anterior testis by seminal receptacle. Seminal recep-
tacle saccular. Uterus preovarian. Eggs 95–115 × 45–55. Vitelline follicles extend
from base of external seminal vesicle up to posterior end of body, confluent in
post-testicular space. Excretory vesicle tubular, extends anteriorly up to the level
of intestinal bifurcation; pore terminal.
Fig. 13.26 Prodistomum

gaevskayae (after Ahmad,
1991)
Remarks: The most distinguishing feature of the species is the presence of thin
finger-like pharyngeal processes. In this respect, it resembles Opechona pharyn-
godactylus Manter, 1940, but differs from it in the larger body size, the extent of
vitellarium, the larger eggs, a larger post-testicular space and a smaller sucker ratio.
Prodistomum mohsini (Ahmad 1984) n. comb (Fig. 13.27)
[Syn. Opechona mohsini Ahmad 1984]
Host: Carangidae: Scomberoides tala (Cuvier).
Locality: BOM, AS
Number: 19
Description: Body elongate, 5180–7205 long, 745–930 wide. Body spinose. Eye-
spot pigment granules present on each side dorsally in pharynx–oesophagus region.
Oral sucker infundibuliform, 500–640 long, 320–370 wide. Ventral sucker pre-
equatorial, smaller than oral sucker, 245–300 in diameter. Sucker ratio 1: 0.76–0.81.
Pseudoesophagus as long as oesophagus. Intestinal bifurcation dorsal, nearer to ven-
tral sucker than to oral sucker. Caeca simple, long, terminate blindly near hind end of

mohsini (after Ahmad 1984,
Pakistan J. Zool, 16, p. 56)
body. Testes two, tandem, separated from each other, equal in size, located in mid-
dle part of hindbody. Cirrus-sac club-shaped, extends beyond ventral sucker nearly
the diameter of it. External seminal vesicle saccular. Genital pore ventral, subme-
dian, to left of median line, anterior to ventral sucker. Ovary spherical, pretesticular,
separated from anterior testis by seminal receptacle. Seminal receptacle saccular,
postovarian. Uterus preovarian. Eggs 100–130x, 58–75. Vitellarium follicular, in
lateral fields, extend from posterior margin of ventral sucker to hind end of body,
confluent in post-testicular area. Excretory vesicle tubular, reaching to posterior level
of pseudo-oesophagus. Excretory pore terminal.
Remarks: Distinguishing features of this species are the large body size, large eggs
and globular ovary.
Prodistomum orientalis (Layman, 1930) Bray and Gibson, 1990 (Fig. 13.28)
[Syn. Phryngora orientalis Layman, 1930; Opechona orientalis (Layman, 1930); O.
scombri Yamaguti, 1938; Lepocreadium puriensis Gupta & Gupta, 1987 (see Bray
and Gibson 1990)]
Host: Scombridae: Rastrelliger brachysoma (Bleeker); R. kanagurta (Cuvier)
Locality: PR, BOB
References: Gupta, P.C. & Gupta, V.C. (1987); Madhavi & Triveni Lakshmi (2011)

orientalis (after Madhavi &
Triveni Lakshmi, 2011)
Description: Body elongate, somewhat broad, 1138–1350 long 224–336 wide Tegu-
ment spined. Eyespot pigment granules present at level of pharynx. Oral sucker
60–68 × 52–64 in size. Ventral sucker spherical, larger than oral sucker, 80–104 in
diameter, pre-equatorial. Pharynx oval, Oesophagus and pseudoesophagus small.
Intestinal bifurcation usually midway between suckers. Caeca terminate at posterior
end of body. Testes two, broader than long, tandem, situated in posterior third of
body. Cirrus-sac club-shaped, extends well beyond ventral sucker. External seminal
vesicle saccular. Genital atrium oval. Genital pore ventral, submedian, towards right
of anterior margin of ventral sucker. Ovary spherical, pretesticular, separated from
anterior testis by seminal receptacle. Seminal receptacle saccular. Uterus preovarian.
Eggs 64–68 × 40–44. Vitelline follicles extend from base of ventral sucker up to
posterior end of body, confluent in post-testicular space. Excretory vesicle tubular,
extends anteriorly up to the level of intestinal bifurcation, pore terminal.
Remarks: This species originally included under the genus Opechona has been trans-
ferred to Prodistomum as P. orientalis by Bray & Gibson (1990) based on the absence
of an uroproct. Bray & Gibson (1990) furnished details of the synonyms, the hosts
and geographic distribution as well as the characteristic features of the species.
P. orientalis has been recorded from R. kanagurta from Aden (Indian Ocean) by
Parukhin (1976). Records from the related hosts include those from Sulawesi (N-W
Pacific ocean) by Yamaguti (1953) and North-west Australia by Korotaeva (1974) and
Bay of Bengal by Zhukov (1960) and Madhavi & Trivenilakshmi (2011). Taking into
consideration the preference to scombrid hosts and also the morphological features,
Bray and Gibson (1990) considered Lepocreadium puriensis Gupta & Gupta, 1987
from R. brachysoma from Puri coast, Bay of Bengal as a synonym of P. orientalis.
Prodistomum siddiqi (Ahmad 1984) n. comb. (Fig. 13.29)
[Syn. Opechona siddiqi Ahmad, 1984]
Host: Scombridae: Rastrelliger kanagurta (Cuvier).
Locality: BOM, AS
Number: 7
Ref. Ahmad (1984a)
Description Body elongate, 2955–3530 long, 410–610 wide; widest at level of

testes. Body spinose up to ovary. Oral sucker 250–270 long, 187–225 wide. Ven-
tral sucker smaller than oral sucker, pre-equatorial, 128–195 in diameter located at
junction of anterior and middle-third of body. Sucker ratio 1: 0.68 Pseudoesoph-
agus long. Caecal bifurcation dorsal, nearer to ventral sucker than to oral sucker.
Caeca simple, long, end blindly near hind end of body. Testes two, slightly lobed,
longer than wide, separated from each other by vitelline follicles, located in posterior
half of body. Cirrus sac club-shaped, extends from genital pore posteriorly usually
halfway between acetabulum and ovary; internal seminal vesicle saccular. Prostatic
vesicle surrounded by prostate gland cells. Cirrus thick–walled, External seminal
vesicle saccular, Genital pore ventral, submedian, to the left of median line, oppo-
site anterior margin of ventral sucker. Ovary distinctly 3–10 lobed, pretesticular,

siddiqi (after Ahmad, 1984,
separated from anterior testis by a distinct gap. Seminal receptacle saccular, pos-
tovarian. Vitellarium follicular, overlap caeca dorsally, intrude between the testes,
extend from level of pseudoesophagus to hind end of body, confluent in forebody and
in post-testicular space. Uterus preovarian, with few eggs. Metraterm thick walled.
Eggs elongated, operculated, 100–120 long, 60–70 wide. Excretory vesicle tubular,
extending anteriorly up to posterior level of pharynx. Excretory pore terminal.
Remarks: This species is characterized by possessing the following combination

of characters: Oral sucker bigger than the ventral sucker, the lobed testes and the
vitellarium extending well anterior to ventral sucker.
Prodistomum travassossi (Ahmad, 1985) n. comb. (Figure 13.30)

[Syn. Opechona travassossi Ahmad, 1985]
Host: Cynoglossidae: Cynoglossus lida (Bleeker)
Locality: BOM, AS
No. 14

travassossi after Ahmad,
1985, Pakistan J. Zool, 17,
p. 148)
Description: Body elongate, 2500–4380 long by 530–580 wide. Tegument spined

to posterior end of body. Oral sucker spherical, subterminal, 160–205 in diameter.
Ventral sucker spherical, nearly same size as oral sucker, 158–210 in diameter, pre-
equatorial. Sucker ratio 1: 1. Pseudoesophagus long. Intestinal bifurcation dorsal,
midway between suckers. Caeca end blindly near posterior end of body. Testes two,
deeply lobed, tandem, intercaecal, occupying almost all available intercaecal space,
situated in posterior third of body, close together but separated by band of vitelline fol-
licles, equal in size. External seminal vesicle saccular. Cirrus-sac cup-shaped, curved,
extends posteriorly into hindbody. Genital pore median, immediately anterior to ven-
tral sucker. Ovary spherical, median, intercaecal, immediately pretesticular. Seminal
receptacle saccular. Vitellarium consists of numerous follicles, extend from anterior
level of pharynx to posterior end of body, confluent between ovary and anterior testis,
between testes and in post-testicular space. Uterus preovarian, between ovary and
ventral sucker. Metraterm thick-walled. Eggs elongate, operculate, 100–120 long by
50–60 wide. Excretory vesicle tubular, extends anteriorly up to level of intestinal
bifurcation.
Remarks: Important features of the species are the lobed testes, the spherical ovary,
the extent of the vitelline follicles from the anterior margin of the pharynx to the
posterior end and the large eggs.
Prodistomum vinodae (Ahmad, 1984) n. comb. (Figure 13.31)
[Syn. Opechona vinodae Ahmad, 1984]
Host: Stromateidae: Pampus argenteus (Euphrasen)
Locality: Goa, AS
Status: Species inquirendae
Description: Body elongate 810–1140 long 165–240 wide, Entire body spinose.
Eyespot pigment granules present. Oral sucker-cup-shaped, 52–75 in diameter. Ven-
tral sucker of same size as oral sucker, in middle third of body. Sucker ratio 1:1.
Pseudo-esophagus as long as oesophagus. Caeca end blindly near posterior end of
body. Testes deeply multilobed, equal in size, in posterior third of body. Cirrus-sac
extends to halfway between ventral sucker and ovary. Genital pore median, immedi-
ately anterior to ventral sucker. Ovary trilobed. Vitelline follicles from anterior end
of pharynx to hind end of body, confluent in forebody. Uterus with 1–5 eggs. Eggs
110–118 by 62–68 in size. Excretory vesicle extends to intestinal bifurcation.

vinodae (after Ahmad, 1984
Remarks: This species is identified by the combination of the following characters:

the vitellarium extending to the level of the pharynx, the deeply lobed testes, the
trilobed ovary and the larger eggs.
Prodistomum waltairense (Madhavi, 1972) Bray and Gibson, 1990 (Fig. 13.32)
Syn: Opechona waltairensis Madhavi, 1972
No: 4
Locality: VSK, BOB
Description: Body elongated, slender with parallel sides and rounded extremities,
2030–3120 long and 448–640 wide. Tegument spined to level of ventral sucker.
Oral sucker terminal, 78–100 long by 78–124 wide. Ventral sucker just anterior to
midbody, subequal to or slightly larger than oral sucker, 97–148 in diameter. Sucker
ratio 1: 1.0–1.3. Oesophagus long; pseudoesophagus usually longer than oesophagus,
bifurcation about midway between pharynx and ventral sucker. Caeca long, termi-
nate blindly near posterior end. Genital pore slightly to left of median line, opposite

waltairense (after Madhavi
anterior margin of ventral sucker. Testes 2, lobed, tandem, close together, situated in
posterior third of body. Cirrus-sac extends posterior to ventral sucker. External sem-
inal vesicle long, sinuous, extends almost to ovary. Ovary indistinctly three-lobed,
immediately pretesticular, submedian; seminal receptacle globular, postero-lateral to
ovary; uterus pretesticular; metraterm well developed, to left of cirrus-sac, about half
as long as cirrus-sac. Eggs large, yellowish, 78 long by 31–39 wide. Vitelline follicles
in lateral fields, anterior limit varying from level of intestinal bifurcation to middle of
pseudoesophagus, filling post-testicular space except region occupied by excretory
bladder. Excretory bladder long, I-shaped; extending to intestinal bifurcation.
Remarks: Madhavi (1972) included the species in the genus Opechona as

O.waltairensis but based on the absence of a uroproct, it was transferred to the
genus Prodistomum by Bray & Gibson (1990). P. waltairense is characterized by
the long slender body, lobed gonads arranged close together near posterior region of
body.
Key to Indian species of Prodistomum
1. Vitellarium restricted to hindbody…………………………………………...... 2
Vitellarium extends into forebody ………………………..……………….….. 4
2. Pharynx provided with three finger-like process ……………….. P. gaevskayae
Ahmad, 1991
Pharynx without any finger-like processes………………………………......... 3
3. Oral sucker normal,……………………………..………..……P. orientalis
Gaevskaya, 1990
Oral sucker infundibuliform……………………… ……………….. P. mohsini
Ahmad, 1984
4. Oral sucker larger than ventral sucker……..….. P. siddiqii Ahmad, 1984
Oral sucker of same size as ventral sucker ………………………………….... 5
5. Ovary lobed …………………………………………………….……….......... 6
Ovary spherical ……………………………..…………….. P. travassossi
Ahmad, 1985
6. Vitellarium extends to level of pharynx, eggs 110–118 by 62–68 ………..... P.
vinodae Ahmad, 1984
Vitellarium extends to intestinal bifurcation, eggs 78 × 31–39 …...…. P. wal-
tairense (Madhavi, 1972)
Genus Pseudolepocreadioides, Hafeezullah 1970.

The genus Pseudolepocreadiodes was erected by Hafeezullah (1970) with P. sym-
metrorchis Hafeezullah, 1970 as the type species. The genus is closely related to
Lepocreadioides Yamaguti, 1936 but differs in having a post-testicular ovary and
symmetrical testes in the ventral sucker zone. Four species including the type species
are known in the genus, all recorded from Indian waters.
Diagnosis: Body rhomboid. Tegument probably spined anteriorly. Eyespot pigment

present. Oral sucker terminal or subterminal. Ventral sucker median, equatorial.
Prepharynx, pharynx and oesophagus present. Caeca simple, extend to posterior
end of body. Testes symmetrical, on either side of ventral sucker. External seminal
vesicle saccular. Cirrus-sac club-shaped, contains internal seminal vesicle, prostatic
complex and cirrus. Genital pore marginal, near oral sucker. Ovary lobed, median,
post-testicular. Seminal receptacle present. Uterus scanty, metraterm distinct. Vitel-
larium reaches forebody, fills lateral wings of body. Excretory vesicle I-shaped. Type
species: P. symmetrorchis Hafeezullah, 1970
Pseudolepocreadioides bombayensis Ahmad, 1981 (Fig. 13.33)
Host: Carangidae: Scomberoides tol (Cuv. & Val.)
Locality: BOM, AS
No. 42
Reference: Ahmad (1981c)
Description: Body 1540–1908 long, 1650–1719 wide, rhomboid. Oral sucker
167–190 in diameter. Ventral sucker 200–235 in diameter, larger than oral sucker,
pre-equatorial. Sucker ratio 1:1.18–1.23. Intestinal bifurcation dorsal; usually closer
to pharynx than to ventral sucker level. Caeca simple, end blindly at posterior end
of body. Testes two. spherical, extracaecal, symmetrical, on either side of ventral
sucker. External seminal vesicle saccular, lies between ventral sucker and seminal
receptacle. Cirrus-sac club-shaped, lies obliquely between genital pore and posterior
border of ventral sucker, overlaps left or right margin of ventral sucker. Genital pore
ventral, marginal, sinistral to oral sucker. Ovary highly lobed, median, intercaecal,
post-testicular. Seminal receptacle immediately preovarian, transversely elongate.
Uterus scanty, coils between ovary and ventral sucker. Metraterm indistinct. Eggs
34–38 long, 24–27 wide. Vitellarium consists of tubular follicles. extending from
posterior border of oral sucker to posterior end of body, filling wing-like sides of
body, confluent in post-ovarian region. Excretory vesicle I-shaped, reaches to ovary:
pore terminal.
Remarks: This species differs from P. symmetrorchis Hafeezullah, 1970 and P.
secundus Ahmad, 1979 in the extension of the vitellarium from the posterior bor-
der of the oral sucker to the posterior end of the body, in the cirrus-sac extending
posteriorly as far as the posterior border of the ventral sucker and the smaller eggs.
Fig. 13.33 Pseudolepocreadioides

1981)

secundus (after Ahmad,
1979)
Pseudolepocreadioides secundus Ahmad, 1979 (Fig. 13.34)

Host: Pomacanthidae: Pomacanthus annularis (Bloch)
Locality: PR, BOB
Description: Body 1230–1780 long, 1320–1850 wide, rhomboidal. Oral sucker

spherical. 150–193 in diameter. Ventral sucker equal to oral sucker, 150–193 in diam-
eter, pre-equatorial. Caeca simple, straight, reaching posterior end of body. Testes
oval, symmetrical, on either side of ventral sucker. External seminal vesicle saccular,
extends to middle of ventral sucker. Cirrus-sac club-shaped, lying obliquely between
ventral sucker and genital pore. Genital pore marginal, sinistral to oral sucker. Ovary
trilobed, median, post-testicular. Seminal receptacle oval, between ovary and ventral
sucker. Uterus coiled between ovary and cirrus-sac; metraterm distinct. Eggs 84–91
by 39–50. Vitellarium consisting of large follicles, reaching from level of testes
to posterior end of body, filling wing-like sides of body.
Remarks: P secundus differs from the type species P. symmetrorchis Hafeezullah,

1970 in having suckers of equal size, a trilobed rather than multilobed ovary, the
testes are well separated from the ventral sucker instead of close to it, the vitellarium
extends between the anterior border of the testes instead of from the level of the
intestinal bifurcation and the eggs are larger.
Pseudolepocreadioides symmetrorchis Hafeezullah, 1970 (Fig. 13.35)
Host: Drepaneidae: Drepane punctata L
Locality: Veeraval, AS (1); Kalingapatnam, BOB (2)
Reference:1. Hafeezullah (1970b); 2. Madhavi et al. (1986)
Description: Body 1150–1740 long, 1370–1570 wide. Tegument thick; spines prob-
ably lost in processing. Oral sucker 137–200 by 152–233, subspherical, subterminal.
Ventral sucker 165–277 in diameter, spherical slightly postequatorial. Sucker ratio
1: 1·15–1·25. Caeca simple, straight, reaching posterior end of body. Testes oval,
symmetrical, on either side of ventral sucker. External seminal vesicle saccular,
dorso-dextral to ventral sucker, extends beyond its posterior margin. Cirrus-sac club-
shaped, lying obliquely between ventral sucker and genital pore; encloses internal

symmetrorchis (after
Hafeezullah, 1970, courtesy
Cambridge University Press.
Parasitology 61, p. 346)
seminal vesicle, well developed pars prostatica, and cirrus; cirrus protrusible. Genital
pore marginal, sinistral to oral sucker. Ovary highly lobed, median, post-testicular.
Seminal receptacle. saccular, between ovary and ventral sucker. Uterus scanty, coils
between ovary and cirrus-sac. Metraterm distinct. Eggs 51–60 by 33–48. Vitellar-
ium consists of small follicles occasionally becoming tubular, from level of intestinal
bifurcation to slightly short of posterior end of body, filling wing-like sides of body.
Uterus scanty, coils between ovary and cirrus-sac; metraterm distinct. Eggs 51–60
by 33–48
Remarks: This species is characterized by the following combination of characters:
the oral sucker is smaller than ventral sucker, ovary is deeply lobed, cirrus-sac lies
obliquelsy between genital pore and ventral sucker,
Pseudolepocreadioides thapari Ahmad, 1981 (Fig. 13.36)
Host: Gerreidae: Gerres filamentosus (Cuv.)
Locality: BOM, AS
No. 14
Description: Body rhomboidal, 1560–1675 long and 1610–1720 wide. Oral sucker
spherical, subterminal, 200–230 in diameter. Ventral sucker smaller than oral sucker,

pre-equatorial, 140–165 in diameter. Sucker ratio 1:0.7–0.71. Intestinal bifurcation

dorsal, usually midway between suckers. Caeca simple, straight, terminate near pos-
terior end of body. Testes two, multilobed, equal in size, symmetrical, extracaecal,
well separated from ventral sucker. External seminal vesicle saccular, extends pos-
teriorly either up to anterior margin of ovary or half-way between ovary and ventral
sucker. Cirrus-sac club-shaped, straight, lies obliquely between ventral sucker and
genital pore. Genital pore marginal, sinistral to oral sucker. Ovary entire, wider than
long, post-equatorial, intercaecal. Seminal receptacle oval, postovarian. Vitelline fol-
licles large, tubular, extend from posterior border of pharynx to posterior end of body,
filling wing-like sides of body, confluent in forebody and non–confluent in postovar-
ian zone. Uterus runs between ovary and intestinal bifurcation. Metraterm indistinct.
Eggs elongate, operculate, 42–51 long and 26–32 wide.
Remarks: P. thapari Ahmad, 1981, was differentiated from P. symmetrorchis
Hafeezullah, 1970, and P. secundus Ahmad, 1979, in having the oral sucker larger
than the ventral sucker, a multilobed testes, an unlobed ovary and the vitellarium
extending from the posterior border of the pharynx and in the posterior extent of
external seminal vesicle.
Key to Indian species of Pseudolepocreadioides Hafeezullah, 1970
1. Oral sucker larger than ventral sucker, testes lobed………………….….…P. tha-
pari Ahmad, 1979
Oral sucker equal to or smaller than ventral sucker, testes spheri-
cal………………….…………… 2
2. Cirrus-sac large, reaches posterior border of ventral sucker…..……… P. bom-
bayensis Ahmad, 1981
Cirrus-sac in forebody ………………………….3
3. Suckers subequal, ovary trilobed…………………….…… P. thapari Ahmad,
1981
Oral sucker smaller than ventral sucker, ovary deeply lobed…. P. symmetrorchis
Hafeezullah, 1970
Genus Transversocreadium Hafeezullah, 1970
Diagnosis: Body much wider than long. Oral sucker subterminal. Ventral sucker
spherical. Prepharynx indistinct. Pharynx and oesophagus present. Caeca arcuate.
Testes 11, in two groups on either side of ventral sucker. External seminal vesi-
cle present. Cirrus-sac with internal seminal vesicle, prostatic complex and cirrus.
Genital pore marginal, near oral sucker. Ovary lobed. Seminal receptacle present.
Uterus scanty; metraterm present. Excretory vesicle tubular. Type and only species:
T. cablei.
Transversocreadium is similar to Multitestis Manter, 1931, particularly M. rotun-
dus Sparks, 1954, and to Rhagorchis Manter, 1931, in the number of testes but differs
from these genera in having a much wider than long body, the horizontally arranged
testes, the marginal genital pore near the oral sucker and a distinctly sinistral ovary.
Five species of the genus have been recorded so far, all from Indian waters.
Fig. 13.37 Transversocreadium

cablei (after Hafeezullah,
1970, courtesy Cambridge
University
Press. Parasitology, 61,
p. 349)
Transversocreadium cablei Hafeezullah, 1970 (Fig. 13.37)

Locality: Calicut (AS) (1); VSK (BOB) (2)
No. 5
References: 1. Hafeezullah (1970b); 2. Madhavi (1972)
Description: Body 470–650 long, 1120–1580 wide, transversely elongated. Oral

sucker, 51–68 in diameter, subterminal, feeble. Ventral sucker 59–68 in diameter,
spherical, feeble, in centre of body. Sucker ratio 1: 1·13–1·24. Prepharynx indistinct.
Pharynx 44–53 in diameter. Oesophagus short. Caeca arcuate, terminate blindly
near posterior end of body. Testes arranged horizontally, five on right and six on
left side of ventral sucker. External seminal vesicle present. Cirrus-sac club-shaped,
disposed obliquely from right to left, extends posteriorly to about midlevel of ven-
tral sucker, encloses internal seminal vesicle in swollen base, prostatic complex in
tubular distal part. Genital pore marginal, in notch left of oral sucker. Ovary deeply
multilobed, antero sinistral, sinistral or postero-sinistral to ventral sucker. Seminal
receptacle large, saccular, near ovary. Uterus scanty, from level of intestinal bifurca-
tion to slightly posterior to ventral sucker; metraterm present. Vitellarium follicular,
disposed along course of caeca. Eggs collapsed, 41–62 by 27–41. Excretory vesicle
tubular; excretory pore terminal.
Remarks: This species was described by Hafeezullah (1970b) from Triacanthus

biaculeatus from Calicut. Madhavi (1972) later recorded the species from the same
host from Visakhapatnam coast, Bay of Bengal. The species is characterized by the
presence of five testes on the right side and six on the left side and the multilobed
ovary.
Transversocreadium fotedari Ahmad & Dhar, 1987 (Fig. 13.38)
Locality: Goa, AS
References: Ahmad & Dhar (1987a)
Description: Body transversely elongate, 1120–1240 long 2800–2856 wide. Oral

sucker 140–165 in diameter. Ventral sucker 170–200 in diameter, sucker ratio 1: 1.2.
Intestinal bifurcation midway between suckers. Caeca arcuate, approaching each
other near posterior end of body. Testes 24 in number, in two groups with 12 on each

fotedari (after Ahmad and
Dhar, 1987)
side. External seminal vesicle saccular, extends to posterior margin of ventral sucker.
Cirrus-sac club-shaped, disposed obliquely from left to right, extends posteriorly up
to anterior margin of ventral sucker, internal seminal vesicle oval, prostatic vesicle
surrounded by prostate gland cells, cirrus thick-walled. Genital pore marginal, in
notch on left side of body wall near oral sucker. Ovary multilobed, lateral to ventral
sucker. Seminal receptacle postovarian. Uterus well developed occupies all available
space between intestinal bifurcation and posterior end of body. Metraterm thick-
walled. Eggs 70–85 X43–50. Vitelline follicles disposed along the course of caeca.
Excretory vesicle I-shaped extends anteriorly up to anterior margin of ventral sucker.
Remarks: This species differs from the remaining four species of the genus in the
following respects: The greater number (24) of testes, larger eggs and the uterus
descends posteriorly as far as the posterior end of body.
Transversocreadium manteri Ahmad, 1981 (Fig. 13.39)
Host: Sparidae: Argyrops spinifer. (Forskal)
Locality: BOM, AS
No. 26
Description: Body transversely elongate, 1130–1245 long and 2730–2870 wide.

Oral sucker spherical, subterminal, 130 160 in diameter. Ventral sucker 190–225 in
diameter, median, larger than the oral’ sucker,’ equatorial. Sucker ratio 1: 1.4–I.46.
Intestinal bifurcation dorsal, usually between suckers. Caeca simple, arcuate, ending
blindly near hind end of body. Testes 14 in number, arranged horizontally in two rows;

manteri (after Ahmad, 1981)
six on left side and eight on right side. External seminal vesicle saccular, extends
to posterior border of ventral sucker. Cirrus-sac club-shaped, disposed obliquely
from left to right, extending from genital pore to anterior border of ventral sucker
encloses ovoid internal seminal vesicle in swollen base, prostate complex in tubular
distal part, thin-walled cirrus. Genital pore marginal, in notch, left to oral sucker.
Ovary deeply multilobed, sinistral to ventral sucker. Seminal receptacle saccular,
postovarian. Vitelline follicles small, rounded, disposed along course of caeca. Uterus
descends to level of caecal ends. Metraterm tubular, thick-walled, Eggs elongate,
operculate, 35–45 by 26–30. Excretory vesicle tubular, reaches to ventral sucker.
Remarks: This species was differentiated from T. cablei by the following features:
Fourteen testes arranged in two rows instead of 11 testes arranged in a single row, the
cirrus-sac extends to the anterior border of the ventral sucker rather than the middle
of it and the larger sucker ratio.
Transversocraedium secundus Ahmad, 1983 (Fig. 13.40)
Host: Gerreidae: Gerres filamentosus (Cuvier)
Locality: PR, BOB
No. 30
Reference: [Ahmad, 1983a]
Description: Body transversely elongate, 1165–1380 long by 2360–2612 wide. Oral
sucker spherical, subterminal, 170–226 in diameter. Ventral sucker smaller than oral
sucker, postequatorial, 140–158 in diameter. Sucker ratio 1: 0.7–0.82. Intestinal bifur-
cation dorsal, usually halfway between the suckers. Caeca arcuate, end blindly near
hind end of body. Testes 6 in number, arranged horizontally, three on each side of
ventral sucker. External seminal vesicle saccular, extends to middle ofventral sucker.
Cirrus-sac club-shaped, disposed obliquely from left to right, extends posteriorly to
just anterior to anterior border of ventral sucker. Genital pore marginal, in notch,
left of oral sucker. Ovary deeply multilobed, sinistral or postero-sinistral to ventral
sucker. Seminal receptacle preovarian. Uterus descends posteriorly to level of caecal
ends. Metraterm thick-walled, tubular. Eggs elongated, operculated, 38–55 by 29–35.
Vitellarium follicular, disposed along course of caeca, confluent near posterior end
of body. Excretory vesicle tubular, reaches to ventral sucker.
Remarks: This species was differentiated from T. cablei Hafeezullah, 1970 by
the following features: the oral sucker is larger than the ventral sucker; three testes
on each side rather than five on the right and six on the left and vitelline fields are
confluent near the posterior end of the body.

1983)

yamagutii (after Ahmad,
1983)
Transversocreadium yamagutii Ahmad, 1983 (Fig. 13.41)

Host: Balistidae: Sufflamen fraenatum (Latriella)
Locality: BOM, AS
No. 25
Description: Body transversely elongate, 1326–1630 long by 2640–2780 wide. Oral

sucker 165–211 in diameter. Ventral sucker spherical, 402–465 in diameter, median,
larger than oral sucker, equatorial in position. Sucker ratio 1: 2.12–2.43. Intestinal
bifurcation dorsal, usually halfway between suckers. Caeca arcuate, ending blindly
near hind end of body. Testes 10, elongate, arranged horizontally, five on each side
of ventral sucker. External seminal vesicle saccular, extends posteriorly as far as pos-
terior border of ventral sucker. Cirrus-sac club-shaped, elongate disposed obliquely
from right to left, extends posteriorly as far as middle of ventral sucker. Genital pore
marginal, in notch, right to oral sucker. Ovary spherical, sinistral to ventral sucker.
Seminal receptacle saccular, postovarian. Metraterm indistinct. Uterus descends pos-
teriorly halfway between ventral sucker and caecal ends. Eggs elongated, opercu-
lated, 32–39 long by 23–25 wide. Vitellarium follicular, disposed along course of
caeca, confluent in forebody and in posterior end of body, extend from level of
oesophagus. Excretory vesicle tubular, reaches to ventral sucker.
Remarks: T yamagutii differs from T. cablei and T. secundus in having a very

large ventral sucker, a spherical ovary, vitelline fields confluent in the forebody,
smaller eggs and 10 testes.
Species differentiation The various species of Transversocreadium can be differ-

entiated mainly by the number of testes:11 in T. cablei; 14 in T. manteri; 24 in T.
fotedari, 10 in T. yamagutii and 6 in T. secundus. The ovary is multilobed in all the
species except in T. yamagutii where it is spherical. Further only in T. secundus the
oral sucker is larger than the ventral sucker.
Family Aephnidiogenidae Yamaguti, 1934
Diagnosis: Body elongate. Tegument spinous. Eyespot pigment present or absent.

Oral sucker subglobular, subterminal. Ventral sucker rounded, of similar size to or
smaller than oral sucker; in anterior third of body. Prepharynx short. Pharynx oval.
Oesophagus distinct. Intestinal bifurcation in forebody. Caeca reach close to posterior

extremity, blind. Testes two, entire or lobed, tandem or oblique; in mid-hindbody.
External seminal vesicle long, coiled or absent. Cirrus-sac claviform, extends into
hindbody may be absent. Pars prostatica vesicular. Ejaculatory duct short to long-,
thin- or thick-walled. Genital pore sinistrally submedian, post-bifurcal. Ovary oval,
pretesticular, usually separated from anterior testis, maybe close to ventral sucker.
Uterus intercaecal, often mainly between anterior testis and ovary. Metraterm distinct,
muscular. Vitellarium follicular; fields extend from posterior extremity to variable
levels in anterior hindbody and forebody. Excretory vesicle I-shaped, reaches ovarian
or bifurcal level; pore terminal. In marine and freshwater teleosts.
Type genus: Aephnidiogenes Nicoll, 1915.

Four species belonging to this family have been reported from marine fishes of India.
Genus Aephnidiogenes Nicoll, 1915
Diagnosis: Body elongate, narrow. Tegument spinose. Oral sucker subglobular. Ven-
tral sucker of similar size to or smaller than oral sucker. Prepharynx short. Pharynx
oval. Oesophagus distinct. Intestinal bifurcation in forebody. Caeca reach close to
posterior extremity, open out through ani. Testes two, entire, tandem, separated, situ-
ated in mid-hindbody. Cirrus-sac absent. Seminal vesicle long, coiled. Male duct wall
close to genital atrium thick, muscular, perforated by ducts from prostate gland cells.
Genital pore immediately anterior to ventral sucker. Ovary oval, pretesticular, sepa-
rated from anterior testis, close to ventral sucker. Uterus intercaecal mainly between
anterior testis and ovary. Meraterm distinct. Vitelline fields extend from posterior
extremity to level of ovary. Excretory vesicle I-shaped, reaches ovarian level. Type
species: A. barbarous Nicoll, 1915.
Three species of this genus were reported from India.
Aephnidiogenes indica Ahmad, 1978
Host: Sparidae: Rhabdosargus sarba (Forskal)
Locality, BOB
Remarks: Bray & Cribb (1977) regarded the species as of uncertain status.
Aephnidiogenes senegalensis Dollfus & Capron, 1958 (Fig. 13.42)
Locality: Veraval, AS
Description: With characters of the genus.
Remarks: A. senegalensis shows specificity to haemulid fishes and has been recorded
from these hosts from a wide range of localities.
Aephnidiogenes vinodae Gupta & Singh, 1985
Host: Sparidae: Rhabdosargus sarba (Forskal)
Locality, PR, BOB,
Reference: Gupta, PC. & Singh (1985)
Family Aephnidiogenidae Yamaguti, 1934 355
Fig. 13.42 Aephnidiogenes

senegalensis (after
Hafeezullah, 1970, courtesy
Genus Neolepocreadium Thomas, 1960
Diagnosis: Body elongate. Ventral sucker in anterior quarter of body. Pharynx oval.
Oesophagus long. Caeca long, blind. Testes two, entire, tandem, in mid-hindbody.
External seminal vesicle long, coiled. Cirrus-sac elongate, oval. Internal seminal
vesicle large, saccular. Pars prostatica vestigial. Ejaculatory duct long or short. Gen-
ital pore submedian, immediately anterior to ventral sucker. Ovary oval or lobed,
pretesticular, separated from ventral sucker. Uterus intercaecal, preovarian. Vitel-
larium reaches from posterior extremity to about level of ovary. Excretory vesicle
reaches ovarian level. Type species: N. caballeroi Thomas, 1960.
Neolepocreadium trachinoti Madhavi, Narasimhulu & Shameem, 1986 (Fig. 13.43)
Host: Carangidae: Trachinotus blochi (Lacepede)
Locality: Kalingapatnam coast, BOB
Reference: Madhavi et al. (1986)
Description: Body long, cylindrical, 4800–8000 long, 576–900 wide. Tegument

spined. Eyespot pigment present in forebody. Suckers poorly developed. Oral sucker
76–101 by 78–117. Ventral sucker 260–320 in diameter, in anterior fifth of body,
Fig. 13.43 Neolepocreadium

trachinoti (after Madhavi,
Narasimhulu, Shameem,
1986)
Prepharynx absent. Pharynx small. Oesophagus long. Caeca long terminate blindly
near posterior extremity of body. Genital pore median, situated immediately ante-
rior to ventral sucker. Testes large, elliptical, tandem, separated by vitelline follicles,
in posterior half of body. External seminal vesicle long, sinuous, extends posterior
to ventral sucker, surrounded by prostatic cells. Cirrus-sac well developed, situated
transversely in front of ventral sucker, encloses elongated seminal vesicle, short
prostatic duct, few prostatic cells and short cirrus. Genital atrium shallow, highly
muscular. Ovary spherical, pretesticular. Seminal receptacle large, flask-shaped, sit-
uated postero-lateral to ovary. Uterus occupies space between ovary ventral sucker,
describing numerous compact transverse coils. Eggs thin-shelled, 50–58 by 127–35.
Excretory vesicle I-shaped, extends to level of ovary. Vitelline follicles large, irreg-
ularly shaped, extend from level of intestinal bifurcation to posterior end of body.
Remarks: Thomas (1960) erected the genus Neolepocreadium with N. caballeroi

from Trachinotus goreensis and T. glaucus as the type species. Yamaguti (1971)
synonymised the genus with Aephnidiogenes Nicoll, 1915, but later workers (Kruse,
1978, Machida, 1982, Bray, 2005b) regarded the genus as valid. Molecular studies by
Bray et al. (2009) led Bray & Cribb (2012) to include Neolepocreadium in the family
Family Aephnidiogenidae Yamaguti, 1934 357
Aephnidiogenidae, based on sequences obtained from specimens identified as N.

caballeroi from Trachinotus blochii from the Great Barrier Reef. N. trachinoti is said
to differ from N. caballeroi in the body shape, in having a longer cirrus-sac enclosing
an elongated rather than an oval seminal vesicle, and in the well developed prostatic
complex and in the egg size.
Family Enenteridae Yamaguti, 1958
Members of this family are parasitic mainly in herbivorous fishes. The family Enen-
teridae was reviewed by Bray & Cribb (2001) with keys to the genera and species
and diagnoses of the family and genera, based on a cladistic analysis utilizing 44
characters. Subfamilies are not recognized.
Diagnosis: Body oval to elongate. Tegument spinose. Oral sucker often with lobes.
Caeca form cyclocoel with or without anus. Testes two, tandem or oblique in hind-
body. Cirrus-sac present. Seminal vesicle internal only. Genital pore in posterior
forebody. Ovary pretesticular. Seminal receptacle canalicular. Vitellarium follicular,
mainly in hindbody. Excretory vesicle I-shaped. Type genus Enenterum Yamaguti,
1958.
Key to Indian genera of Enenteridae

1. Cyclocoel formed but no anus ………………………………. Proenenterum Manter,
1954
Cyclocoel and anus present ………………………………………2
2. Oral sucker infundibuliform, with up to ten lobes…………….… Enenterum
Linton, 1910
Oral sucker subglobular, lacking lobes ……………………………. Koseira
Nagaty, 1942
Genus Enenterum Linton, 1910
Diagnosis: Body oval. Eyespot pigment present. Oral sucker infundibuliform with
up to ten lobes. Ventral sucker rounded, pre-equatorial. Prepharynx distinct. Phar-
ynx oval or pyriform. Oesophagus short. Intestinal bifurcation in forebody. Caeca
form cyclocoel, open through single anus. Testes two, tandem in hindbody. External
seminal vesicle absent. Cirrus-sac oval. Genital pore in posterior forebody, may be
associated with accessory sucker or genital papilla. Ovary oval, rounded or lobed,
pretesticular. Seminal receptacle canalicular. Uterus preovarian. Eggs operculate.
Vitellarium follicular, mainly in hindbody. Excretory vesicle I-shaped. Type species:
E. aureum Linton, 1910.
Two species belonging to this genus E. minutum Yadav, 1977, and E. mannarense
Hafeezullah, 1980, have been reported from Indian marine fish. E. minutum is
regarded as ‘of doubtful status’ by Bray (1986a).
Enenterum mannarense Hafeezullah, 1980 (Fig. 13.44)

Host: Kyphosidae: Kyphosus cinerascens (Forsskal)
Locality: Gulf of Mannar
No. 1
Distribution: South Africa (Bray, 1986a), Western Australia (Bray & Cribb, 2001)
Description: Body 9439 long, 1238 wide, elongate, Tegument spinose in anterior
region. Oral sucker funnel-shaped, 701 long 481 wide, produced into five basic
muscular conical broad-based lobes, one ventral, two lateral and two dorsal, ventral
lob notched into two to give total of six lobes, Ventral sucker 413 long 468 wide.
Prepharynx long. Pharynx pear-shaped. Oesophagus absent. Intestinal bifurcation
abuts pharynx. Caeca wide, extend laterally united near posterior end of body, form
single anus. Testes two, entire, tandem, separated, situated in posterior part of body.
No external seminal vesicle. Cirrus-sac thin-walled, ovate, anterodorsal to ventral
sucker contains saccular seminal vesicle becoming narrow and coiled anteriorly,
pars prostatica surrounded by well-developed prostate gland cells and protrusible
cirrus. Genital pore posterior to intestinal bifurcation, submedian, to left. Accessory
sucker present behind genital pore. Ovary globular, entire, almost in middle of body,
Fig. 13.44 Enenterum

mannarense (after
Hafeezullah, 1980, Bull.
Zool. Soc. India. 2, p. 146)
Family EnenteridaeYamaguti, 1958 359
separated from anterior testis. Seminal receptacle large, postero-dorsal to ovary.

Uterus preovarian, coiled behind ventral sucker. Metraterm muscular. Vitellarium
follicular, extensive from posterior end of body to posterior margin of ventral sucker,
follicles intruding between gonads. Eggs 56–63 × 42–45.
Remarks: A single specimen of this species was collected by the author, and there
are no further reports of this species from India. Thus, the infection is extremely rare.
However, this species has been recorded from Kyphosus sp. from other countries
such as South Africa (Bray, 1986a) and Ningaloo coral reef (Western Australia)
(Bray & Cribb, 2001). E. mannarense is characterized by the presence of five basic
anteriorly directed conical oral lobes with broad bases and only the ventral lobe is
subdivided into two by a deep notch making a total of six lobes. In this respect it
differs from all the other species of the genus.
Enenterum minutum Yadav, 1977
Host: Sciaenidae: Otolithes ruber (Bl. & Schn.)
Locality: Ratnagiri, AS
Reference: Yadav (1977c)
Status: Species inquirendum.
Genus Koseiria Nagaty, 1942
Diagnosis: Body elongate. Eyespot pigment present. Oral sucker subglobular or

infundibuliform, lacking lobes. Pharynx and oesophagus present. Caeca form cyclo-
coel posteriorly, giving rise to single terminal anus. Testes two, tandem, in mid-
hindbody. Cirrus-sac oval. Internal seminal vesicle tubular, coiled. Pars prostatica
distinct, Ejaculatory duct short. Genital pore median, in posterior region of forebody,
may be associated with accessory sucker. Ovary oval or lobed, Seminal receptacle
canalicular. Uterus preovarian. Vitellarium follicular, mainly in hindbody. Excretory
vesicle I-shaped. Type species E. tahmeli Nagaty, 1942.
Nagaty (1942) established the genus with K. tahmeli as the type species. Two
species of this genus have been recorded from India: K. nagatyi Ahmad, 1984 and
K. manteri Ahmad, 1987.
Koseiria manteri Ahmad, 1987 (Fig. 13.45)
Host: Hemiramphidae: Rhynchorhamphus georgii (Val.)(= Hemiramphus leu-
copterus (Val.))
No. 14
Description: Body elongate, 3120–4470 long, 560–690 wide. Eyespot pigment

granules scattered in pharynx region. Oral sucker spherical, 180–250 in diameter.
Ventral sucker smaller than oral sucker, 130–145 in diameter, situated at junction of
anterior and middle-third of body. Sucker ratio 1: 0.58–0.72. Prepharynx present.
Pharynx barrel-shaped. Oesophagus long. Intestinal bifurcation midway between
suckers. Caeca unite posteriorly, open ventro–subterminally by anus. Testes two,
spherical, tandem, contiguous, intercaecal, in anterior part of posterior third of body.
Fig. 13.45 Koseira manteri

J. Zool, 19, p. 333)
Cirrus-sac ovoid, in forebody, lies horizontally. Seminal vesicle tubular, coiled. Pars
prostatica short, tubular, surrounded by prostate gland cells. Ejaculatory duct short.
Genital pore ventral, median or submedian, in forebody. Ovary oval, pretesticular.
Seminal receptacle saccular, preovarian. Vitellarium with numerous follicles, over-
laps caeca ventrally, few intrude into intercaecal space, extends from posterior border
of ventral sucker to posterior end of body, confluent in post-testicular field. Uterus
runs between ovary and ventral sucker. Eggs elongate, operculate, 84–97 long, 47–60
wide. Excretory vesicle tubular, reaches to posterior margin of ovary.
Remarks: This species differs from K. nagatyi Ahmad, 1984, in the shape of the oral
sucker, the barrel-shaped pharynx, the intestinal bifurcation being midway between
the suckers, the postequatorial position of the ovary and the larger eggs.
Koseiria nagatyi Ahmad, 1984 (Fig. 13.46)
Host: Kyphosidae: Kyphosus cinerascens (Forskal).
Locality: BOM, AS.
Number: 9 specimens from 2 hosts out of 21 examined.
Fig. 13.46 Koseira nagatyi

J. Zool,16, p. 51)
Description: Body elongate, 2970–4360 long, 560–690 wide. Tegument unspined.

Eyespot pigment granules scattered in pharynx region. Oral sucker inverted bell-
shaped, 305–382 long, 205–290 wide. Ventral sucker spherical, smaller than oral
sucker, median, 170–245 in diameter, situated at junction of anterior and middle-
third of body. Sucker ratio 1: 0.82–0.84. Prepharynx present. Pharynx flask-shaped.
Oesophagus long. Intestinal bifurcation dorsal, nearer to ventral sucker than to oral
sucker. Caeca unite posteriorly, open ventro–subterminally by anus. Testes two, ellip-
tical, tandem, contiguous, intercaecal, in posterior half of body. Cirrus-sac ovoid, in
forebody, occupies entire space between intestinal bifurcation and ventral sucker.
Seminal vesicle tubular, coiled. Pars prostatica short, tubular, surrounded by prostate
gland cells. Ejaculatory duct short, joining metraterm at base of genital pore. Genital
pore ventral, median or submedian, just right of median line, immediately post-
bifurcal. Ovary elliptical, immediately pretesticular, contiguous with anterior testis.
Seminal receptacle saccular, preovarian. Vitellarium follicular, overlaps caeca dor-
sally, intrudes into intercaecal space, commence from posterior to ventral sucker,
extend to posterior end of body, confluent in post-testicular field. Metraterm short.
Eggs oval, operculate, 55–70 long, 38–45 wide. Excretory vesicle tubular, reaches
to anterior margin of ovary. Excretory pore terminal.
Remarks: K. nagatyi differs from other species in having an inverted bell shaped
oral sucker, a pyriform pharynx, elliptical and contiguous gonads and in egg size.
Genus Proenenterum Manter, 1954
The genus Proenenterum was established by Manter (1954)to include P. iso-
cotylum Manter, 1954 (type species) and P. ericotylum Manter, 1954. The genus is
characterized by a cyclocoel without an anus and the unlobed oral sucker.
Diagnosis: Body elongate oval. Eyespot pigment absent. Oral sucker subglobular.
Prepharynx and pharynx present. Oesophagus short or absent. Caeca form cyclocoel
posteriorly but no anus. Testes two, tandem, in mid-hindbody, entire or lobed. Exter-
nal seminal vesicle absent. Cirrus-sac oval. Internal seminal vesicle tubular, coiled.
Pars prostatica distinct, Ejaculatory duct short. Genital pore submedian, in posterior
region of forebody. Ovary oval or lobed. Seminal receptacle canalicular. Uterus pre-
ovarian. Vitellarium follicular, mainly in hindbody. Excretory vesicle I-shaped. Type
species: P. isocotyleum Manter, 1954.
Proenenterum manteri Ahmad 1985 (Fig. 13.47)
Host: Carangidae: Tachysurus platystomus (Richardson).
Fig. 13.47 Proenenterum

manteri (after Ahmad, 1985)
No. 21
Description: Body elongate, 780–1222 long, 225–370 wide. Entire body spinose.
Oral sucker 72–130 in diameter. Ventral sucker spherical, larger than the oral sucker,
155–205 in diameter. Sucker ratio 1: 2.15.2.34. Prepharynx short, pharynx oval.
Oesophagus present. Intestinal bifurcation dorsal midway between suckers. Caeca
unite near posterior end of body, forming cyclocoel; anus lacking. Testes two, tan-
dem, deeply lobed, close together, intercaecal, in posterior third of body. Cirrus-sac
saccular, filling space between ventral sucker and intestinal bifurcation, may or may
not overlap anterior margin of ventral sucker; encloses tubular, much coiled seminal
vesicle, a short pars prostatica with a few prostate gland cells and a short cirrus.
Genital pore ventral, slightly to right of median line, just posterior to intestinal bifur-
cation. Ovary triangular, immediately pretesticular, median. Seminal receptacle sac-
cular, postovarian. Vitellarium follicular, lateral, dorsal and ventral to caeca, filling
post-testicular space, extending from some distance posterior to intestinal bifurca-
tion to posterior end of body. Uterus preovarian, running between ovary and ventral
sucker. Metraterm lacking. Eggs thin-shelled, operculate, 70–86 long, 43–50 wide.
Excretory vesicle I-shaped.
Remarks: P. manteri is the only species of the genus reported from India. It is
characterized by possessing the following combination of characters: Fairly large
ventral sucker, vitelaria terminating a little behind ventral sucker and larger eggs.
Family Gyliauchenidae Fukui, 1929
The Gyliauchenidae comprises approximately 30 species, which are almost entirely

restricted to herbivorous fish and to the Indo-West Pacific region. Gyliauchenids are
essentially characterized by specializations of the digestive system: the oral sucker
has been lost and functionally replaced by the pharynx. The oesophagus is long and
convoluted and often provided with a bulb. The ventral sucker is usually placed at
posterior end. The family was regarded by some to be close to the Lepocreadiidae and
by others to the Paramphistomidae. Phylogenetic analysis using 18SrRNA sequences
by Blair and Barker (1993) gave strong support for placing Gyliauchenidae close to
the Lepocreadiidae and no support to the alternative hypothesis. Bray et al. (2009)
studied the phylogenetic relationships with the superfamily Lepocreadioidea and
produced more evidence that gyliauchenids are lepocreadioids. Bray & Cribb (2012)
considered the Gyliauchenidae one of the six families in the superfamily. Al-Jahdali
& Hassanine (2012) based on the evidence obtained from a study of the life cycle of
Gyliauchen volubilis Nagaty, 1956 strongly supported the phylogenetic relationship
of Gyliauchenidae and Lepocreadiidae inferred from molecular phylogenetic studies.
Extensive reviews on the family were given by Nahhas & Wetzel (1995) and Hall &
Cribb (2005b).
Diagnosis: Body elongate or pyriform, smooth. Oral sucker absent, ventral sucker
usually at posterior extremity. Pharynx surrounds mouth. Oesophagus well devel-
oped, often highly coiled, surrounded by well developed gland cells, oesophageal
bulb usually present. Caeca long. Testes two, tandem, oblique or symmetrical. Cirrus-
sac well developed. Seminal vesicle tubular, partly external to cirrus-sac. Pars prostat-
ica prominent. Ejaculatory duct well developed. Prostate gland cells profuse, external
to cirrus sac. Genital pore usually median. Genital atrium prominent. Ovary entire
or lobed, inter-, pre- or post-testicular. Uterus passes anteriorly from ovary usually
in coils. Vitelline follicles profuse, may fill entire posterior part of body. Excretory
vesicle I-shaped. Lymphatic ducts present. Type genus Gyliauchen Nicoll, 1915.
Ten genera are included in the family (Hall & Cribb, 2005b). From India, only
two species of the genus Gyliauchen Nicoll, 1915 have been reported: G. ozakii,
Srivastava, 1938 and G. indicum Gupta & Tandon, 1983
Genus Gyliauchen Nicoll, 1915
Diagnosis: Body elongate to pyriform. Ventral sucker close to posterior extrem-

ity. Oesophagus convoluted. Caeca long, broad. Testes rounded, tandem or oblique
or symmetrical. Cirrus-sac well developed. Ovary entire, between or anterior to
testis. Vitelline follicles profuse along lateral fields of body. Uterus preovarian. Type
species: G. tarachodes Nicoll, 1915
Gyliauchen ozakii Srivastava, 1938 (Fig. 13.48)
Host: Synodontidae: Harpodon neherius (Hamilton) (1); Siganidae: Siganus canalic-
ulatus ( Park)(2)
Locality: AS (1), Mandapam, BOB (2)
Ref. 1. Srivastava (1938b); Gupta N.K. & Sehgal (1971)
Fig. 13.48 Gyliauchen

ozakii (original)
Family Gyliauchenidae Fukui, 1929 365
Description: Body plump, 5015 long, 1309 in breadth across the middle of the body.
Tegument smooth. Pharynx oval. Oesophagus tubular, coiled, surrounded by promi-
nent gland cells all along its course. Oesophageal bulb well developed. Caeca small,
flattened. Posterior sucker at posterior end of body, 510–520, in diameter. Testes
two in posterior part of body anterior to posterior sucker, sometimes overlapping
each other. Seminal vesicle highly developed, divided into two parts by constriction.
Pars prostatica small, swollen and surrounded by numerous prominent and closely
aggregated prostate gland cells. Ejaculatory duct broad, tubular, lined internally by
irregular plaques. Genital sinus peculiar, spherical, reversible, opening guarded by
several fairly broad triangular chitinous processes. Genital pore close to left cae-
cum just anterior to its blind end. Ovary small, spheroidal and pretesticular. Seminal
receptacle between ovary and anterior testis. Vitellarium follicular, restricted mostly
to intestinal zone but some of them reaching anteriorly, as far as level of oesophageal
bulb. Uterus small. Eggs 68–85 x 34–51.
Remarks: G ozakii is characterized by the presence of a chitinous process in the

genital atrium. Such a structure has not been reported for any other species.
Gyliauchen indicum Gupta S.P. & Tandon, 1983 (Fig. 13.49)
Host: Engraulidae: Coilia ramkarati (Hamilton)
Locality: PR, BOB
No. 5
Ref: Gupta S.P. and Tandon (1983)
Description: Body crescent-shaped, convex dorsally and concave ventrally,

2110−2400 long, 720−880 wide, prolonged into conical process postero-dorsal to
ventral sucker. Tegument aspinose. Pharynx pyriform, terminal, with diverticula,
Fig. 13.49 Gyliauchen

indicum (after Gupta S.P. &
Tandon, 1983);
150−230 long, 170−195 wide. Oesophagus long, coiled. Oesophageal bulb muscu-
lar. Caeca muscular, extend to gonads. Posterior sucker spherical, ventro-terminal or
subterminal, 330−400long, 290−390 wide. Genital pore ventrolateral, at midlevel
or posterior end of intestinal caeca. Testes obliquely tandem or symmetrical, sepa-
rate or close to each other in posterior part of body. Genital atrium large, extends to
posterior end of caeca or just posterior to it, encloses internal seminal vesicle, pars
prostatica and ejaculatory duct. Space around. seminal vesicle and pars prostatica
surrounded by prostate gland cells. External seminal vesicle not reaching to gonads.
Ovary pretesticular, spherical, 110−220. Seminal receptacle postovarian, larger than
ovary, between ovary and testes. Vitellarium follicular, extends from just anterior to
oesophageal bulb to just posterior to intestinal bifurcation, confluent in region of
oesophagus. Uterus short with few coils between testes and genital pore. Eggs large,
40−80 long, 30−46 wide. Excretory vesicle small pyriform, extending anteriorly as
far as posterior end of seminal receptacle. Pore at tip of conical process.
Remarks: This species is characterized by the complete absence of oesophageal

gland cells. The only other species of Gyliauchen lacking oesophageal gland cells is
G. zancli Nahhas & Wetzel, 1995. G. zancli differs from G. indicum in the smaller
body size, relatively larger pharynx and smaller testes. G. indicum differs from the
other Indian species G. ozakii Srivastava, 1938, in the extent of the external seminal
vesicle not reaching the gonads, in having testes close to the posterior sucker instead
of well separated, and in the absence of a chitinous process in the genital atrium.
Chapter 14
Superfamily Brachycladioidea Odhner,
1905
Family Acanthocolpidae Lühe, 1906
[Syn. Pleorchiidae Poche, 1926; Pseudolepidapedinae Yamaguti, 1938, Stephanos-

tominae Yamaguti, 1958].
The Acanthocolpidae Lühe, 1906, is a fairly large family of digenetic trematodes
that infect the gastrointestinal tract of marine teleost fishes. They are cosmopolitan in
distribution with representatives occurring in all the oceans. Members of the family
are characterised by a spinous tegument, the lack of an external seminal vesicle and
the presence of a uterine seminal receptacle (Bray, 2005). Lühe (1906) erected the
subfamily Acanthocolpinae Lühe, 1906, to include the genera Acanthocolpus Lühe,
1906, Stephanochasmus Looss, 1900 (=Stephanostomum Looss, 1899), and Dero-
pristis Odhner, 1902. The subfamily was elevated to family rank by Lühe (1909).
The genus Deropristis was removed and included in the separate family Deropris-
tidae Cable & Hunninen (1942). Many genera have since been added to the family
Acanthocolpidae. According to the keys to acanthocolpid genera presented by Bray
(2005), there are at present 17 genera in the family. The well-known genera in the
family are Acanthocolpus Lühe, 1906, and Stephanostomum Looss, 1899. Bray et al.
(2005) assessed the relationship within Acanthocolpidae using LSU SSUW rDNA
sequences.
Diagnosis: Body oval to elongate. Tegument spined. Oral sucker may be provided
with one, or two or three rows of enlarged circum-oral spines. Eyespot pigment may
be present. Oral sucker subglobular or infundibuliform. Ventral sucker pre-equatorial.
Intestinal bifurcation in forebody. Caeca occasionally H-shaped, terminate blindly at
posterior extremity of body or open into excretory bladder forming a uroproct. Testes
usually two, tandem or oblique, occasionally numerous, arranged in 2–4 longitudinal
rows. Cirrus-sac present. Internal seminal vesicle oval, bipartite, tripartite or tubular.
Pars prostatica present. Ejaculatory duct may be armed with spines. Genital atrium
tubular, short or long, may be armed with spines. Genital pore median or submedian,
immediately anterior to ventral sucker. Ovary pretesticular, entire. Uterine seminal
© Crown 2018 367
https://doi.org/10.1007/978-94-024-1535-3_14
368 14 Superfamily Brachycladioidea Odhner, 1905
receptacle present. Uterus preovarian or pretesticular. Metraterm distinct or indis-

tinct, may be armed with spines. Eggs large, operculate. Vitellarium follicular, in
lateral fields extend from posterior extremity to level of uterine field or occasionally
to pharynx. Excretory vesicle I-shaped. Type genus: Acanthocolpus Lühe, 1906.
Although the family contains 17 genera, the Indian species fit into 7 genera that
can be separated by the following key:
1. Intestine H-shaped, testes numerous …………………Pleorchis Railliet, 1896
Intestine lacking anterior caeca, testes two ………………………………2
2. With rows of enlarged oral spines ………………………………..…… … 3
Lacking oral rows of spines …………………………………………………5
3. Single row of enlarged oral spines…………………Monostephanostomum
Kruse, 1979
Two or more rows of oral spines ………………..…………………………….4
4. Two rows of enlarged oral spines…………….,… Stephanostomum Looss, 1899
Three to five rows of enlarged oral spines ….Stephanostomoides Oshmarin &
Mamaev, 1966.
5. Caeca terminate blindly……………………………………………6
Uroproct present ………………………………………Tormopsolus Poche,
1926
6. Genital atrium long, narrow, metraterm and ejaculatory duct may
be lined with spines, ventral sucker protuberant or pedunculate.
…………………………Acanthocolpus Lühe, 1906
Genital atrium short, genital atrium, ejaculatory duct and metraterm lacking
spines.………………………………… Pseudolepidapedon Yamaguti, 1938
Genus Acanthocolpus Lühe, 1906

[Syn. Podocotyle (Indopodocotyloides) Gupta & Sayal, 1970; Pseudacanthocolpus
Rabie & Ahmed, 2000]
Diagnosis: Body elongate. Tegument spined. No oral spines. Eyespot pigment

present. Oral sucker terminal. Ventral sucker in anterior third of body, protuberant or
pedunculate. Prepharynx long. Pharynx oval. Oesophagus short. Intestinal bifurca-
tion in forebody. Caeca long, blind. Testes two, oval, tandem or oblique, in posterior
half of hindbody. Cirrus-sac claviform usually reaches into hindbody. Seminal vesi-
cle internal, saccular, bipartite, or tripartite. Pars prostatica narrow. Ejaculatory duct
long, spined or not. Genital atrium bears spines or unarmed. Ovary subglobular,
close to anterior testis. Uterus preovarian, intercaecal. Metraterm long, spined or
not. Vitelline fields reach to cirrus-sac or beyond. Type species: A. liodorus Lühe,
1906.
Twenty species belonging to this genus have been reported from India. For as
many as eight species, the host is Chirocentrus dorab (Forsskal) (Madhavi, 2011).
However, in course of time many of these species have been invalidated and some
species have been transferred to other genera. For instance, A. guptai Gupta & Puri,
1980, and A. inglisi Gupta & Gupta, 1982, were regarded as synonyms of A. liodorus
Family Acanthocolpidae Lühe, 1906 369
Lühe, 1909; A. lucknowensis Gupta & Puri, 1980, and A. puriensis Gupta & Puri,
1980, were considered as synonyms of A. caballeroi Gupta & Sharma, 1972, by
Hafeezullah (1991). Four species A. dorabi Gupta & Ahmad, 1981, A. microtesticu-
latus Lokhande, 1990, A. srivastavai Varma & Singh, 1987, and A. thapari Gupta and
Ahmad, 1981, were put under the category of taxon inquirendum by Bray & Justine
(2012). A. orientalis Srivastava, 1939, was transferred to the genus Stephanostomum
as S.orientalis (Srivastava, 1939) (Madhavi, 1976a). Similarly, A. tenuis Manter,
1963, is included under the genus Stephanostomoides as S. tenuis (Manter, 1963).
Two recently erected species A. amravatai Mishra, Chandra & Saxena, 2013, and A.
durghai Mishra, Chandra & Saxena, 2013, lack the body spines and the genital pore
is located posterior to the ventral sucker and the presence of spines in the metraterm
and the cirrus-sac was not mentioned. From the general characters, it appears that
they may be opecoelids. The two species should be considered as taxa inquirenda.
There has been much discussion about the validity of A. luehei and A. caballeroi
as distinct from the type species A. liodorus. Caballero (1952), Yamaguti (1958) and
Manter (1963) considered A. luehei as synonymous with A. liodorus. Mamaev &
Oshmarin (1966) and Hafeezullah (1971b, 78) have disagreed with this synonymy.
Hafeezullah (1991) revalidated A. luehei and A. caballeroi differentiating them from
A. liodorus by differences in the extent of the vitellarium and presence or absence
of a pedunculate ventral sucker as follows:
A. luehei: Ventral sucker non-pedunculate. Vitellarium overlaps the cirrus-sac.
Cirrus-sac extends from base of pharynx to midway between ventral sucker and
ovary.
A. liodorus: Ventral sucker pedunculate or situated on a lateral protrusion. Cirrus-
sac extends posteriorly to anterior level of vitellarium. Vitellarium not overlapping
cirrus-sac.
A. caballeroi: Ventral sucker pedunculate, Cirrus-sac extends posteriorly beyond
anterior level of vitellarium. Vitellarium overlaps the cirrus-sac.
For the purpose of present account, all the three species are tentatively considered
as distinct. In view of the confusion prevailing about the status of different species
in the genus, only three species of Acanthocolpus whose validity is probable are
described in detail and the remaining species are simply listed. Future investigations
should concentrate on the study of validity of these various species employing modern
tools in taxonomy. The genus thus requires a focused study.
Acanthocolpus caballeroi Gupta & Sharma, 1972 (Fig. 14.1)
[Syn: A. puriensis Gupta & Puri, 1980; A. lucknowensis Gupta & Puri, 1980]
Locality: RT, AS (1); PR, BOB (2,3) Digha coast, BOB (4)
Reference: 1. Gupta, A.N. & Sharma (1972a); 2. Gupta, V& Puri (1980); 4. Hafeezul-
lah (1991)
Fig. 14.1 Acanthocolpus

caballeroi (after Gupta &
Sharma, 1972)
Description: Body cylindrical, 2805–11175 long 285–615 wide, unspined. Oral

sucker lacks circumoral spines, smaller than ventral sucker. Ventral sucker at tip of
peduncle, near anterior end of body. Caeca end blindly near posterior end of body.
Testes two ovate, directly tandem, post-equatorial. Cirrus-sac very long, extends dis-
tinctly posterior to anterior level of vitellarium, encloses bipartite seminal vesicle,
pars prostatica and very long spined ejaculatory duct. Genital pore at base of pedun-
cle. Ovary globular, small, immediately pretesticular. Uterus with few eggs, preo-
varian. Metraterm long, spined. Eggs 60 × 30. Vitelline follicles extend anteriorly
almost to midbody covering an appreciable posterior part of cirrus-sac. Excretory
bladder Y-shaped.
Remarks: The species was originally described by Gupta, A. N & Sharma (1972a)
from Chirocentrus dorab from Ratnagiri coast (AS). Hafeezullah considered it a
synonym of A. luehei Srivastava, 1939. Later, Hafeezullah (1991) revalidated it. It
was differentiated from A. luehei by the pedunculate nature of ventral sucker and
the very long cirrus-sac and the vitellarium overlapping a larger part of cirrus-sac.
Hafeezullah (1991) regarded A. puriensis Gupta & Puri, 1980, and A. lucknowensis
Gupta & Puri, 1980, as synonyms of A. caballeroi.
Acanthocolpus liodorus Lühe, 1906 (Fig. 14.2)
[Syn: A. guptai Gupta & Puri, 1981; A. inglisi Gupta & Gupta, 1980]
Locality: East & West coasts of India
References: Hafeezullah (1971b, 78); Madhavi (1976a); Zhukov (1977); Gupta, V
& Puri (1980), Gopal et al. (1985); Lakshmi & Rao (1978)
Description: Body cylindrical, with a peduncle near anterior end. 2240–2700 long
410–450 wide, unspined. Ventral sucker borne on peduncle. Oral sucker subtermi-
nal, without circumoral spines. Prepharynx long, pharynx pear-shaped, oesophagus
shorter than prepharynx. Caeca terminate blindly near posterior end of body. Testes
two, elongate, directly tandem, postequatorial. Cirrus-sac extends posterior to pedun-
cle of ventral sucker up to anterior level of vitellarium, encloses seminal vesicle, pars

liodorus (after Luhe, 1906)
prostatica and spined ejaculatory duct. Genital pore near intestinal bifurcation. Ovary
spherical, immediately in front of anterior testis. Uterus between ovary and peduncle,
metraterm spined. Eggs 62 by 33. Vitelline follicles from posterior level of cirrus
pouch to posterior end of body.
Remarks: A. liodorus has been recorded from C. dorab from many regions along
east and west coasts of India (Madhavi, 1976a; Zhukov, 1977; Kumari & Srivastava,
1976; Gopal et al. 1985; Hafeezullah, 1991). Acanthocolpus inglisi Gupta & Gupta,
1980, and A. guptai Gupta & Puri, 1980, both reported from C. dorab from Puri
coast were considered as synonyms of A. liodorus by Hafeezullah (1991)
Acanthocolpus luehei Srivastava, 1939 (Fig. 14.3)
Host: Chirocentridae: Chirocentrus dorab Forskal (1, 4); Gerreidae: Gerres punctatus
Cuvier (2); Carangidae: Carangoides armatus (Bl. & Schn.) (3)
Locality: East and West coasts of India
References: 1. Srivastava (1939i); 2. Hafeezullah (1971b, 1978); 3. Gupta, V& Jahan
(1975); 4. Gupta, A.N. & Sharma (1972a). A.N. & Sharma (1972a).
Description: Body elongate, 3200–3900 by 500–680 in size. Entire tegument

spined, thickly in forebody. Oral sucker very small, 60–80 by 90–100. Ventral
sucker well developed 240–280 in diameter, located at 1/8th of body length. Prephar-
ynx long. Pharynx oval. Oesophagus small. Caeca extend to posterior end of body.
Testes oval, intercaecal, behind ovary. Cirrus-sac extends to midway between ventral
sucker and ovary, encloses bipartite seminal vesicle, small, narrow pars prostatica,
long spined ejaculatory duct. Genital atrium small. Genital pore anterior to ven-
tral sucker at base of intestinal bifurcation. Ovary 120–160 in diameter, situated just

luehei (original)
post-equatorial. Uterus preovarian. Eggs 70–80 × 30–40. Metraterm well developed,

tubular, sinuous armed with minute spines. Vitellarium composed of a large number
of small follicles extending from posterior to anterior quarter of body to posterior
end, confluent in post-testicular space.
Remarks: A luehei differs from A. liodorus in the non-pedunculate nature of the ven-
tral sucker and the cirrus-sac extending beyond the anterior limit of the vitellarium.
Other species reported from India
Acanthocolpus amravatai Mishra, Chandra & Saxena, 2013
Host: Leiognathidae: Leiognathus daura (Cuvier)
Locality: PR, BOB
Reference: Mishra et al. (2013)
Remarks: This form should be considered a taxon inquirendum. Characters such as

the smooth tegument, the lack of eyespot pigment, the seminal vesicle lying free in
the parenchyma and the extensive vitellarium indicate that it may be an opecoelid.
Acanthocolpus apoorvae Saxena & Prakash, 2003
Information on this species is not available.
Acanthocolpus chorinemusi Gupta & Srivastava, 1989
Host: Carangidae: Scomberoides lysan (Forsskal)
Locality: Rameswaram, GOM
Reference: Gupta, S.P. & Srivastava (1989)
Acanthocolpus dorabi Gupta & Ahmad, 1981

Host: Chirocentridae: Chirocentrus dorab (Forsskal).
Locality: PR, BOB
Reference: Bray & Justine (2012); Gupta, V & Ahmad (1981)
Remarks: This form is retained as taxon inquirendum and probably belongs to the
genus Stephanostomoides.
Acanthocolpus durghai Mishra, Chandra & Saxena, 2013
Locality: PR, BOB
Reference: Mishra et al. (2013)
Remarks: This species should be considered as a taxon inquirendum. Characters such

as the smooth tegument, the lack of eyespot pigment, the seminal vesicle lying free
in parenchyma, the extensive vitelline follicles indicate that it may be an opecoelid.
Acanthocolpus equulai Gupta & Srivastava, 1989
Host: Leiognathidae: Leiognathus daura Cuvier
Locality: PR, BOB
Acanthocolpus guptai Gupta & Puri, 1980
Host: Chirocentridae: Chirocentrus dorab (Ruppell)
Locality: PR, BOB
Remarks: Synonymized with A. liodorus by Hafeezullah (1991)
Reference: Gupta, V & Puri (1980)
Acanthocolpus inglisi Gupta& Gupta, 1980
Host: Carangidae: Carangoides malabaricus (Bl. & Schn.)
Locality: PR, BOB
Reference: Gupta, S.P. & Gupta, R.C. (1980a)
Remarks: Synonymized with A. liodorus by Hafeezullah (1991)
Acanthocolpus lucknowensis Gupta & Puri, 1981
Host: Gerridae: Gerres punctatus (Cuv. & Va1.).
Locality: PR, BOB.
Remarks: Synonymized with A. caballeroi Gupta & Sharma, 1972 by Hafeezullah
(1991)
Acanthocolpus lutjanusi Gupta & Srivastava, 1989
Host: Lutjanidae: Lutjanus fulvus (Bl. & Schn.))
Locality: PR, BOB
Acanthocolpus manteri Pandey & Tewari, 1984
Host: Scombridae: Euthynnus sp.
Locality: BOM, AS
Reference: Pandey & Tewari (1984)
Remarks: Hafeezullah (1991) considered this species as a synonym of A. luehei.
Acanthocolpus microtesticulatus Lokhande, 1990

Host: Chircentridae: Chircentrus dorab Forskal
Locality: PR, BOB
Reference: Lokhande (1990a)
Remarks: Bray & Justine (2012) regarded the species as taxon inquirendum
Acanthcolpus orientalis Srivastava, 1939
Host: Carangidae: Alepes djedaba (Forsskal) (1), Leiognathidae: Leiognathus daura
Cuvier (2)
Locality: PR, BOB
Remarks: Transferred to the genus Stephanostomum as S. orientalis (Sivastava,

1939) Madhavi, 1976
Reference: 1. Srivastava (1939i); 2. Gupta, V & Ahmad (1981); Madhavi (1976a)
Acanthocolpus puriensis Gupta & Puri, 1980
Host: Carangidae: Alepes djedaba (Forsskal)
Location: Intestine.
Locality: PR, BOB
Remarks: Synonymized with A. caballeroi Gupta & Sharma, 1972 by Hafeezullah
(1991)
Acanthocolpus srivastavai Varma & Singh, 1983
Host: Chirocentridae: Chirocentrus dorab (Forskal)
Locality: Trivandrum, Kerala, AS.
Reference: Varma & Singh (1983)
Remarks: Regarded as species inquirendum (Bray & Justine, 2012)
Acanthocolpus thapari Gupta & Ahmad, 1981
Host: Chirocentridae: Chirocentrus dorab (Forskal)
Locality: PR, BOB
Reference: Gupta, V & Ahmad (1981)
Remarks: This form is regarded as a species inquirendum. It probably belongs in the
genus Stephanostomoides according to Bray & Justine (2012)
Acanthocolpus valiyathurai Varma & Singh, 1983
Host: Carangidae: Alepes djedaba (Forskål)
Locality: Trivandrum, Kerala, AS
Reference: Varma & Singh (1983)
Genus Monostephanostomum Kruse, 1979
Monostephanostomum is characterized by the presence of a single row of circum-

oral spines, a condition which contrasts with the two rows of Stephanostomum Looss,
1899, and three to five rows of Stephanostomoides Mamaev et Oshmarin, 1966 (Bray,
2005). Reimer (1983) also suggested that the relatively small number of oral spines,
a distinct narrow neck devoid of spines just posterior to the oral sucker, the presence
of uroproct and the position of the ventral sucker at about the middle of the body
constitute characters of the genus. All but the last character remained valid.
Diagnosis: Body claviform to elongate. Oral sucker with single row of few large
circum-oral spines. Ventral sucker in middle or anterior third of body. Caeca open
via uroproct. Testes oval, tandem, in posterior part of hindbody. Cirrus-sac elongate.
Seminal vesicle oval. Pars prostatica weekly developed. Ejaculatory duct long, lined
with knob-like spines or tubercles. Genital atrium small. Genital pore median. Ovary
oval, pretesticular. Uterus pretesticular. Metraterm long, with or without spines, Vitel-
larium reaches into forebody or not. Excretory vesicle I-shaped, reaches ovary. Type
species: M.manteri Kruse, 1979.
The genus is represented by two species in marine fishes of India, M.

mesospinosum (Madhavi, 1976) and M. loossi (Pandey & Tewari, 1984) which were
initially included under the genus Stephanostomum Looss, 1899. Bray et al. (2007)
found molecular evidence that at least one Monostephanostomum species (M. nolani
Bray & Cribb, 2007) is embedded within the genus Stephanostomum. As this is not
the type species of the genus, Monostephanostomum, it is retained here as distinct.
Monostephanostomum loossi (Pandey &Tewari, 1984) Saoud, Nahhas, Al Kuwari
& Ramadan, 2002
Syn. Stephanostomum loossi Pandey & Tewari, 1984
Host: Scombridae: Euthynnus sp.
Locality: BOM, AS
Description: This species is characterised by claviform body, 32 circumoral spines;

contiguous gonads, vitellarium restricted to hindbody.
Reference: Pandey & Tewari (1984).
Monostephanostomum mesospinosum (Madhavi, 1976) Bray & Cribb, 2002
(Figs. 14.4 and 14.5)
[Syn. Stephanostomum mesospinosum Madhavi, 1976]
Host: Carangidae: Carangoides malabaricus (Bloch)
Number: 12
Locality: VSK, BOB
Reference: Madhavi (1976a); Bray & Cribb (2002)
Description: Body filiform, slender, 6640–8320 long, 224–256 wide. Tegument

spined. Eyespot pigment present. Oral sucker small, terminal, 78–85 by 98–117,
circumoral spines 18, rhomboidal, arranged in single uninterrupted circle, ventral
spines smaller, 31–39 long, gradually increase in size, mid-dorsal spines 54–62 long.
Ventral sucker 124–156 × 136–175. Sucker ratio 1:1.2–1.4. Forebody small, about
7% of total body length. Prepharynx long. Pharynx pyriform. Oesophagus short.
Intestinal bifurcation just anterior to ventral sucker. Caeca broad, long, open into
excretory bladder. Testes 2, elliptical, tandem, in posterior half of body, separated
by vitelline follicles. Cirrus-sac long, narrow, extends to slightly more than halfway
between ovary and ventral sucker, joins metraterm slightly posterior to ventral sucker.
Fig. 14.4
Monostephanostomum
mesospinosum (after
Madhavi, 1976)
Fig. 14.5
Monostephanostmum
mesospinosum, oral spines
Ovary spherical, pretesticular. Uterus preovarian, intercaecal. Metraterm long, thick-

walled, spined, surrounded by gland cells. Eggs 54–56 by 23–27. Vitellarium extends
from base of cirrus-sac to posterior end of body, interrupted at level of ovary and
testes.
Remarks: This species is characterized by the long slender body, the oral sucker
with 18 oral spines, the small forebody, the vitellarium interrupted at the level of the
gonads and the testes and ovary separated by vitelline follicles.
Genus Stephanostomum Looss, 1899
The genus Stephanostomum Looss, 1899, has a wide distribution. Its members infect
marine teleosts particularly those of warm waters. According to Cribb et al. (2002),
Stephanostomum is second only to Phyllodistomum Braun, 1889, in the number of

described species among digeneans of fishes. There are over 107 nominal species
of Stephanostomum of which 19 have been reported from Indian region. The large
size of the genus complicates the identification of species. Specific determination
is based on the number, shape and arrangement of circum-oral spines, the relative
sizes of the suckers, the extent and arrangement of vitellarium follicles, the length of
cirrus-sac and genital atrium, the distance between gonads, the presence or absence
of a uroproct and the egg size (Stunkard, 1961).
Diagnosis: Body elongate. Tegument spined. Eyespot pigment present. Oral sucker
with two rows of enlarged oral spines. Ventral sucker in anterior third of body.
Prepharynx long. Pharynx oval. Oesophagus short. Intestinal bifurcation in fore-
body. Caeca long, open via uroproct or blind. Testes two, oval, tandem or oblique,
may overlap, in posterior half of hindbody. Cirrus-sac claviform. Seminal vesicle
internal, saccular. Pars prostatica narrow. Ejaculatory duct long, usually reaches into
hindbody, bears spines or unarmed. Genital atrium short to long, may be spined.
Ovary subglobular, close to anterior testis. Uterus preovarian, intercaecal. Metraterm
long, spined or not. Vitelline fields reach to cirrus-sac. Excretory vesicle I-shaped.
Type species: S. cesticillus (Molin, 1858) Looss, 1899.
The nineteen species of Stephanostomum reported from India, their hosts and
localities are listed by Madhavi (2011). S. lucknowense Gupta & Jahan, 1975,
was synonymized with S. triacanthi Madhavi, 1976. S. chauhani Gupta & Ahmad,
1979, was considered a synonym of S.orientalis by Hafeezullah & Dutta (1998). S.
mesospinosum Madhavi, 1976, and S. loossi Pandey & Tewari, 1984, were moved
to the genus Mesostephanostomum Kruse, 1979, by Bray & Cribb (2002) and Saoud
et al. (2002). The Indian species of the genus have not so far been subjected to any
review. Bray & Cribb (2002) listed the known species of Stephanostomum. Informa-
tion available on the various Indian species is provided below:
Stephanostomum adinterruptum Hafeezullah, 1971 (Fig. 14.6)
Host: Fistulariidae: Fistularia petimba Lacépède
Locality: VSK, BOB
Reference: Hafeezullah (1971a); Madhavi (1976a)
Description: Body elongate, 2540–2570 long 320–380 wide. Oral sucker

140–143 by 173–185, oral spines 36, in two complete circles, dorsal spines large
and strong, lancet like, ventral spines small and feeble. Ventral sucker 145–168 in
diameter, 612–708 from anterior end. Sucker ratio 1:0.83–0.93. Testes elongate,
oval, contiguous, near posterior end of body, post-testicular space 125–128. Cirrus-
sac long, reaches more than halfway between ventral sucker and ovary. Seminal
vesicle club-shaped. Cirrus unarmed. Genital atrium short. Ovary contiguous with
testis. Vitelline follicles extend from posterior edge of ventral sucker to posterior end
of body, not contiguous in post-testicular space. Uterus preovarian. Metraterm long,
unarmed. Eggs 63–72 × 39–45.
Fig. 14.6 Stephanostomum

adinterruptum (after
Hafeezullah, 1971 Courtesy
J. Helminth, 45, p. 85)
Remarks: This species closely resembles S. casum (Linton, 1910) but differs from
it in the number of spines which are ventrally interrupted, the smaller sucker ratio
and in the long unarmed cirrus.
Stephanostomum anterospinosum Khan & Karyakarte, 1987
Host: Chirocentridae: Chirocentrus dorab (Forsskål)
Locality: RT, AS
Reference: Khan & Karykarte (1987b).
Stephanostomum attenuatum Hafeezullah, 1971 (Fig. 14.7)
Host: Carangidae: Scomberoides tala (Cuv.)
Locality: MS, BOB
Description: Body elongate, slender, attenuated. Oral sucker 90–101 × 125–155,

cup-like or funnel-shaped with 36 peribuccal spines in two complete rings. Ventral
sucker 209–283, rounded, 924–1,068 from anterior end. Sucker ratio 1:1.67–1.84.
Prepharynx and pharynx present. Caeca long. Testes oval, contiguous in poste-
rior fourth of body, post-testicular space short. Cirrus-sac long and slender, not
reaching midbody, encloses long seminal vesicle, pars prostatica and short cirrus
armed with spines. Genital atrium short. Genital pore immediately anterior to ven-
tral sucker. Ovary contiguous with anterior testis. Uterus scanty. Metraterm spined.
Eggs 57–72 × 39–42. Vitelline follicles from slightly post-equatorial level to poste-
rior end of body, sparse and contiguous anterior to ovary

attenuatum (after
Hafeezullah, 1971, Courtesy
J. Helminth, 45, p. 85)
Remarks: Important features of the species include the attenuated body, the oral
sucker being smaller than the ventral sucker, the vitellarium not reaching the cirrus-
sac, the contiguous anterior testis and ovary and the scanty uterus.
Stephanostomum bhuneshwari Gupta & Puri, 1986 (Fig. 14.8)
Host: Mullidae: Parupeneus macronemus (Lacepede)
Locality: PR, BOB
Description: Body 5750–7280 long by 530–740 wide. Oral spines 40, in two unin-
terrupted rows. Sucker ratio 1:2.50–4.19. Testes unequal, separated by vitelline folli-
cles, in posterior third of body. Cirrus-sac long. Cirrus spined. Ovary separated from
testis by uterus. Vitellarium extends from midlevel of cirrus-sac to posterior end of
body. Metraterm long. Eggs 50–70 by 25–40.
Remarks: This species resembles S. triacanthi Madhavi, 1976, but differs from it in
the smaller pharynx and the absence of a uroproct.

bhuneshwari (after Gupta &
Puri, 1986)
Stephanostomum casum (Linton, 1910) McFarlane, 1934 (Fig. 14.9)

[Syn. Lechradena edentula Linton, 1910; L. edentulum Linton, 1910; Stephanos-
tomum casus Linton, 1910; Stephanostomum edentulum (Linton, 1910) Yamaguti,
1953. (Reference: Caballero, 1952)
Lutjanidae: Lutjanus malabaricus (Schn.)
Locality: VSK, BOB
Number: 9
Distribution: Tortugas, Mexican Pacific, Galapagos, Bermuda, Bahama Islands,
Puerto Rico, Curacao, Red Sea, Palawan Island, New Caledonia, Ghana
Description: The specimens from Bay of Bengal measured 5,760–6,400 long
672–720 wide. Sucker ratio is 1.6–2.0. Egg size 62 by 39–41.
Remarks: Characterised by somewhat elongate body form, presence of 36 oral

spines, cirrus-sac extending to halfway between ventral sucker and ovary, contiguous
gonads situated in posterior half of hindbody and vitellarium extending to the level of
posterior margin of ventral sucker. This species has a wide geographic distribution,
and in most regions, the hosts are Lutjanus spp.

casum (after Linton,1910)
Stephanostomum chauhani Gupta & Ahmad, 1979

Host: Lutjanidae: Lutjanus rivulatus (Russell).
Locality: PR, BOB
Number: 4
Reference: Gupta, V & Ahmad (1979a)
Remarks: Regarded as a synonym of S. orientalis (Srivastava, 1939) Madhavi, 1976

by Hafeezullah & Dutta (1998).
Stephanostomum cloacum (Srivastava, 1938) Manter & Van Cleave, 1951
(Figs. 14.10)
[Syn. Stephanochasmus cloacum Srivastava, 1938]
Host: Latidae: Lates calcalifer Bloch (1); Rachycentridae: Rachycentron canadum
L. (2), Triacanthidae: Triacanthus biaculeatus (Bloch)
Locality: Karachi, AS (1); Kakinada, PR, BOB (2); Chilka Lagoon (3)
References: 1. Srivastava (1938c); 2. Hafeezullah (1978); 3. Madhavi & Shameem
(1993)
Description: Body elongate, 2640–2720 by 416–480, spined. Oral sucker 112–128

in diameter. Circumoral spines 34, arranged in two alternate rows without any gap.
Oral spines 32 × 6, aboral spines 20 × 4, lateral spines 28 × 6. Ventral sucker
192–208 × 208–224. Forebody narrow with scattered eyespot pigment. Intestinal
bifurcation slightly anterior to ventral sucker. Caeca long open into excretory bladder.
Genital pore median, anterior to ventral sucker. Testes two, tandem, in posterior
Fig. 14.10 (adult)

Stephanostomum cloacum
(after Madhavi & Shameem,
1993)
third of body. Cirrus-sac long, extends posterior to ventral sucker, nearly halfway
between ovary and ventral sucker, encloses seminal vesicle, pars prostatica, unspined
ejaculatory duct. Ovary spherical, pretesticular. Vitellarium follicular, distributed in
lateral fields from posterior margin of ventral sucker to posterior end of body filling
entire post-testicular space. Uterus preovarian. Metraterm short, unspined, lateral to
ventral sucker. Eggs 60–70 × 36–40.
Remarks: Srivastava (1938c) first reported the species from Rachycentron canadus
L. from Karachi coast, Arabian Sea. Later, Hafeezullah (1978) reported S. cloacum
in Rachycentron canadus from Kakinada coast, Bay of Bengal, and its metacercaria
from the gobiid fish Minous monodctylus. Madhavi and Shameem(1993) reported the
species from Triacanthus strigilifer from Chilka lagoon and provided details of the
life cycle of the species with Nassarius orissaensis serving as first intermediate host
and the bivalve Modiolus undulatus as the second intermediate host. The species has
34 circum-oral spines, the vitellarium overlaps the cirrus-sac, the forebody is short,
and ejaculatory duct is long.
Stephanostomum ditrematis (Yamaguti, 1939) Manter, 1947 (Fig. 14.11)

[Syn. Echinostephanus ditrematis Yamaguti, 1939; S. longisomum Manter, 1940; S.
filiforme Linton, 1940; S. manteri Perez-Vigueras, 1955; S. cubanum Perez-Vigueras,
1955; S. seriolae Yamaguti, 1970
Host: Carangidae: Megalaspis cordyla L
Locality: VSK, BOB
Distribution: Inland Sea of Japan, Florida, Curacao & Jamaica, Cuba, Woods Hole;
Massachusettes, Bimini; British West Indies, Tropical American Pacific, Bay of
Bengal.
Description: Body slender, 3297–8540 long 202–539 wide. Tegument region imme-
diately posterior to oral sucker unspined, rest of body heavily spined, Eyespot pig-
ment present. Oral sucker bowl-shaped, surrounded by double ring of 36 alternating
circum-oral spines, not interrupted ventrally, ventral spines shorter than dorsal spines,
spines stout, not sharp. Ventral sucker prominent, circular, at end of first sixth of body
length. Sucker ratio 1:1.9–2.7. Prepharynx very long. Pharynx elongate. Oesophagus
sinuous. Intestinal bifurcation just anterior to ventral sucker. Caeca long, uroproct
present. Testes 2, oval, contiguous or well separated from each other, situated very
near posterior end of body. Cirrus-sac long, slender. Seminal vesicle tubular. Pars
prostatica sinuous, joined to ejaculatory duct by short muscular duct. Ejaculatory
duct long, coiled, tubular. Genital atrium very long, lined with villous-like struc-
tures. Genital pore median, post-bifurcal at anterior margin of ventral sucker. Ovary

ditrematis (after Bartoli &
Bray. Courtesy Springer, 58.
p. 46)
spherical, slightly posterior to middle of hindbody. Uterine seminal receptacle large.

Uterus preovarian, between ovary and proximal end of cirrus-sac. Eggs large, thin-
shelled, collapsed, 48–62 × 21–33. Vitelline follicles filling post-testicular space,
confluent in dorsal preovarian field, extend to near level of middle of cirrus-sac.
Excretory vesicle I-shaped.
Remarks: S ditrematis has been reported from various hosts from a wide range of
geographic regions showing some preference for carangid hosts. Characterised by
the presence of 36 oral spines arranged in two rows, with ventral oral spines shorter
than dorsal spines, the presence of a uroproct and the vitellarium confluent between
the testes and between the testis and the ovary and extending to the level of the middle
of the cirrus-sac.
Stephanotomum equula Gupta & Puri, 1985 (Fig. 14.12)
Locality: PR, BOB
Ref: Gupta, V& Puri (1985b)
Description: Body 5200–6800 long 470–710 wide. Oral spines 40, in two unin-
terrupted alternating rows. Sucker ratio 1:3.07–3.80. Testes widely separated by
vitelline follicles, situated in posterior third of body. Cirrus-sac extends slightly pos-
terior to ventral sucker. Cirrus armed with few spines. Ovary close to testis. Uterus
with few eggs. Eggs 45–75 × 20–40. Vitellarium extends from posterior end of
cirrus-sac to posterior end of body.

equula (after Gupta & Puri,
1985)

madhaviae (after Madhavi
1976)
Remarks: This species resembles S. triacanthi Madhavi, 1976, very closely and may
be synonymous to that.
Stephanotomum gazzai Gupta & Puri, 1985
Host: Leiognathidae: Gazza minuta (Bloch)
Locality: PR, BOB
Reference: Gupta, V& Puri (1985b)
Remarks: The description of the species based on a single specimen lacks many
details. It is here considered as species inquirendum
Stephanostomum lucknowense Gupta & Jahan, 1975
Host: Triacanthidae: Triacanthus strigilifer (Day)
Locality: PR, BOB
Number: 4
Remarks: Synonymized with S. triacanthi Madhavi, 1976 by Saoud et al. (2002)
Stephanostomum madhaviae Bray & Cribb, 2013 (Figs. 14.13 and 14.14)
[Syn. S. orientalis (Srivastava, 1939) of Madhavi, 1976]

madhaviae oral spines
Hosts: Carangidae: Caranx sexfasciatus Quoy & Gaimard; Carangoides malabaricus

(Bloch & Schneider), Carangoides chrysophrys Cuvier
Location: VSK, BOB
Distribution: Queensland, off Hope Island.
Ref: Madhavi (1976a); Bray & Cribb (2003)
Description: Body elongate, 2606–3938 by 220–353. Tegument spined, unarmed
patch immediately posterior to oral sucker. Oral sucker terminal, circumoral spines
30–34, in double ring with distinct ventral hiatus, spines decrease in size towards
hiatus. Ventral sucker in anterior third of body. Caecal wall contiguous with excre-
tory bladder wall, uroproct formation not clear. Testes 2, oval, tandem, contiguous,
post-testicular space short. Cirrus-sac reaches well into hindbody. Seminal vesicle
saccular. Pars prostatica elongate, lined with anucleate cell-like bodies. Ejaculatory
duct short, lined with closely packed rows of cupolas (spines). Genital atrium short
reaching anterior to mid-level of ventral sucker. Genital pore median, slit-like, imme-
diately anterior to ventral sucker. Ovary oval, contiguous with anterior testis. Uterus
narrow, intercaecal, preovarian. Eggs 67–73 × 34–43. Vitellarium follicular, reaches
almost to posterior end of cirrus-sac, lateral, dorsal and ventral to caeca.
Remarks: Bray & Cribb (2013) described this species from the carangid Caranx
ignobilis from Queensland and considered the species identified by Madhavi (1976a)
as S. orientalis (Srivastava, 1939) Madhavi, 1976, to be identical to this. S. madhaviae
is characterized by: an oral sucker with 30–34 circum-oral spines arranged in a
double ring, with a distinct ventral hiatus and spines decreasing in size gradually
from dorsal surface towards the hiatus. Other diagnostic features of S. madhaviae
are the contiguous gonads, the vitellarium which overlaps the cirrus-sac and the very
short post-testicular space.
Stephanostomum microsomum Madhavi, 1976 (Figs. 14.15 and 14.16)
Host: Rachycentridae: Rachycentron canadum (Linnaeus)
Locality: VSK, BOB
Number: Numerous
Description: Body small, spindle shaped, 1280–1760 long; 256–304 wide at level of
testes. Eyespot pigment on either side of pharynx. Oral sucker 85–117 in diameter,
armed with 36 spines arranged in two uninterrupted rows, spines slender, needle-
like, oral spines 39 in size, aboral spines smaller 23 long. Ventral sucker large,
145–195 in diameter, sucker ratio 1:1.4–1.6. Forebody one third of body length.
Intestinal bifurcation immediately anterior to ventral sucker. Caeca long extend to
near posterior end of body. Genital pore median, immediately anterior to ventral
sucker. Testes 2, oval, tandem, contiguous, situated close to posterior extremity.
Cirrus-sac slender, sinuous, extends well posterior to ventral sucker, almost reaching
ovary, contains coiled seminal vesicle, small pars prostatica and long ejaculatory
duct. Ovary globular, close in front of anterior testis. Uterus short, occupies space
between ovary and ventral sucker. Vitelline follicles small, extend in lateral fields
between posterior margin of ventral sucker and posterior end of body. Eggs 68–78
by 39–49.
Remarks: This species is characterized by its very small body size and 36 oral spines
arranged in two rows. In these respects, it has been considered as closely related to
S. minutum (Looss, 1901) Manter, 194, but is differentiated from S. minutum by the
extent of the vitellarium, the longer forebody, the longer and sinuous cirrus-sac and
the larger eggs.

microsomum (after Madhavi
1976) entire

microsomum oral spines
Stephanostomum orientalis (Srivastava, 1939) Madhavi, 1976 (Fig. 14.17)

[Syn. Acanthocolpus orientalis Srivastava, 1939 S. chauhani Gupta & Ahmad, 1979]
Hosts: Carangidae: Alepes djedaba (Forsskal) (1), Megalaspis cordyla (L) (3), Leiog-
nathidae: Leiognathus daura (Cuvier) (2); Lutjanidae: Lutjanus johni (Bloch) (3)
Locality: PR, Digha Coast, BOB
Reference: 1. Srivastava, (1939i); 2. Gupta, V& Ahmad(1979a); 3. Hafeezullah &
Dutta (1998)
Description: Body elongate, slender, 4350–6090 long, 390–740 wide, spined. Oral
sucker funnel-shaped, circum oral spines 32, arranged in two alternate rows with a gap
on ventral side. Ventral sucker situated just behind intestinal bifurcation. Prepharynx
very long. Pharynx pyriform. Oesophagus short. Intestinal bifurcation anterior to

orientalis (after Srivastava,
1936)
ventral sucker. Caeca extend to posterior end of body. Testes two, ovate, tandem, may
or may not be separated by vitelline follicles. Cirrus-sac very long, tubular, sinuous
extends far posterior to ventral sucker, encloses bipartite seminal vesicle, tubular pars
prostatica surrounded by prostatic gland cells and long and spined cirrus. Genital
atrium unspined, between ventral sucker and intestinal bifurcation. Ovary globular,
pretesticular. Seminal receptacle absent. Uterus preovarian. Metraterm differentiated.
Eggs 57–82 by 47–55. Vitelline follicles circumcaecal extend from posterior level
of cirrus-sac to posterior end of body. Excretory vesicle Y-shaped.
Remarks: Srivastava (1939i) originally described this species as Acanthocolpus ori-

entalis Srivastava, 1939, from the carangid fish Alepes djedaba from Puri Coast.
Madhavi (1976a) collected specimens from adjacent Visakhapatnam coast that are
identical in all respects to Srivastava’s specimens except for the presence of oral
spines. On the basis of this close similarity, Madhavi presumed that circum-oral
spines were lost in Srivastava’s specimens due to maceration and transferred Acan-
thocolpus orientalis to the genus Stephanostomum. Hafeezullah & Dutta (1998)
accepted this change in status and further synonymised S. chauhani Gupta & Ahmad,
1979, from the fish Lutjanus rivulatus with this species.
Stephanostomum polynemi Madhavi, 1976 (Fig. 14.18)
Host: Polynemidae: Leptomelanosoma indicus Shaw
Locality: VSK, BOB
Description: Body elongate, 6880–8320 long, 512–704 wide. Eye spot pigment
present on either side of pharynx. Oral sucker small, oral spines 32 needle-like,
arranged in two uninterrupted rows, ventral aboral spines smallest, 39 long, ventral
oral spines 60 long, lateral and dorsal spines 70 long. Ventral sucker 290–320 in diam-
eter. Sucker ratio 1: 2.0–1:2.3. Forebody small 1/9th to 1/10th of total body length.
Prepharynx long. Pharynx spherical. Oesophagus short. Caeca long. Testes large,
elliptical, smooth, tandem, close together but not contiguous, situated in posterior
third of body. Cirrus-sac small, extends only slightly posterior to ventral sucker. Semi-
nal vesicle long, occupies most of space inside cirrus-sac. Pars prostatica small. Cirrus
armed. Genital atrium short. Genital pore anterior to ventral sucker. Ovary spheri-
cal, pretesticular, separated from anterior testis by short distance. Uterus preovarian.
Metraterm tubular thick-walled, unarmed, nearly as long as cirrus-sac. Vitelline fol-
licles numerous, extend from posterior end of cirrus-sac to posterior end of body,
confined to lateral fields, fill all post-testicular space. Eggs 58–62 by 31–39.
Remarks: This species closely resembles S. cloacum Srivastava, 1939, but differs
from it in the smaller cirrus-sac and vitellarium extending more anteriorly.

polynemi (after Madhavi
1976)
Stephanostomum pseudoditrematis Madhavi, 1976 (Fig. 14.19)

Host: Rachycentridae: Rachycentron canadum (Linn.)
Locality: VSK, BOB
Number: Numerous
Description: Body long, slender, 5440–10080 long, 256–480 wide. Oral sucker
claviform, oral spines 36, in two uninterrupted rows. Oral spines slightly bigger
than aboral spines. Sucker ratio 1: 1.0–1.4. Forebody one sixth of body length.
Caeca form uroproct. Testes large, elliptical, separated by few vitelline follicles, sit-
uated near posterior region. Cirrus-sac long, slender, extends to one third of distance
between ovary and ventral sucker, contains elongated seminal vesicle, small pars
prostatica and unarmed cirrus. Genital atrium long, unarmed. Ovary separated from
testis by vitellarium. Metraterm long, narrow, sinuous. Vitellarium extends from base
of cirrus-sac to posterior end of body, fills post-testicular area and space between
gonads.

pseudoditrematis (after
Madhavi 1976)
Remarks: This species differs from S. ditrematis (Yamaguti, 1939) in the mid-ventral
oral spines not being smaller than the remaining spines, and the unspined nature of
the cirrus and metraterm.
Stephanostomum simhai Gupta & Ahmad, 1979
Host: Sparidae: Rhabdosargus sarba (Forsskal)).
Locality: PR, BOB
No: 8
Description: Body elongated, spinose, a disc-like anterior and blunt posterior end,
6170–7400, 520–750 wide. Oral sucker funnel-shaped, 110–140 long, 200–260 wide.
Circum-oral spines 36, arranged in two alternate rows of 18 spines each, first row
spines smaller, 37–48 long. Ventral sucker pre-equatorial, 270–350 long, 260–390
wide. Genital pore anterior to ventral sucker. Testes oval, tandem unequal, separated
from one another by vitelline follicles, in anterior part of posterior third of body.
Cirrus-sac elongated, extends up to half way between ventral sucker and ovary.
Genital atrium unspined, short. Ovary oval, pretesticular, separated from anterior
testis. Seminal receptacle just posterior to ventral sucker up to hind end of body.
Uterus preovarian, extending between ovary and ventral sucker. Metraterm muscular,
unspined. Genital atrium unspined, short. Eggs oval, operculated, 52–80 long, 43–51
wide.
Remarks: Presence of ani and the long seminal vesicle reported in this species is
unusual for members of the genus Stephanostomum and the family Acanthocolpidae.
The description lacks details of vitelline distribution, and the figures are of poor
quality. Additional specimens of this species have to be examined to determine its
taxonomic status.
Stephanostomum srivastavai Gupta & Gupta, 1980 (Fig. 14.20)
Locality: PR < BOB
No. 13
Reference: Gupta S.P. & Gupta, R.C (1980a)
Description: Body elongate, attenuated 2800–5600 long, 310–540 wide. Tegument
spinose. Oral sucker cup-like, terminal, 100–150 long, 170–210 wide, circumoral
spines 32, spindle shaped, uninterrupted, arranged in two alternate rows of 16 spines
each, ventral spines 50–75 long, dorsal spines 50–80 long. Ventral sucker subspheri-
cal, larger than oral sucker, 150–210 long, 140–200 wide. Prepharynx long. Pharynx

srivastavai (after Gupta &
Gupta, 1980)
elongate. Caeca extend to posterior end of body. Genital pore immediately anterior
to ventral sucker. Testes oval, tandem, median, subequal, separated from each other
by vitelline follicles, in anterior part of posterior third of body. Cirrus-sac elongate,
reaches less than half way between ventral sucker and ovary. Cirrus spiny. Genital
atrium unspined, short. Ovary separated from anterior testis by vitelline follicles,
90–210 long, 80–130 wide. Uterus preovarian; winding between ovary and ventral
sucker. Metraterm unarmed. Eggs 72–90 long, 28–42 wide. Vitellarium follicular,
confluent between testes and between testis and ovary extend from near base of
cirrus-sac up to hind end of body.
Remarks: Differentiated from S. attenuatum Hafeezullah, 1971, in having 32 circu-

moral spines instead of 36 and in having gonads separated instead of contiguous.
Stephanostomum theraponi Gupta & Ahmad, 1979 (Figs. 14.21 and 14.22)
Host: Terapontidae: Terapon theraps (Cuv. Et Val.)
Locality: PR, BOB
No. 2
Reference: Gupta, V& Ahmad (1979a)

theraponi (after Gupta &
Ahmad, 1979)

theraponi oral spines
Description: Body elongated, slender, with an attenuated neck, 4940–5580 long,

370–480 wide. Tegument spinose. Oral sucker flat funnel-shaped, terminal, 140–165
long. 220–260 wide. Circumoral spines 28, arranged in two uninterrupted alternate
rows of 14 each, first row spines longer, 59–77 long, second row spines smaller. 44–55
long. Ventral sucker spherical, larger than oral sucker, 185–200 in diameter. Genital
pore immediately anterior to ventral sucker. Testes oval, tandem, separated from
one another by vitelline follicles, near posterior end of body. Cirrus-sac elongated,
extends well into hindbody. Cirrus spined, eversible, Genital atrium short. Ovary
oval, pretesticular, separated from anterior testis by vitelline follicles. Uterus extends
between ovary and ventral sucker. Metraterm thin-walled. Eggs 59–88 by 50–58.
Vitellarium follicular, confluent between testes, testis and ovary, anterior to ovary
and posterior to posterior testis, extend from just posterior to base of cirrus-sac to
hind end of body.
Remarks: According to the authors, this species closely resembles S. attenuatum

Hafeezullah, 1971, in having ventrally uninterrupted oral spines, the vitellarium
reaching to the base of the cirrus-sac but differs from the latter in having 28 rather,
than 36 oral spines and gonads separated from one another instead of contiguous.
Stephanostomum triacanthi Madhavi, 1976 (Fig. 14.23)
[Syn. S. lucknowense Gupta & Jahan, 1977]
Localities: VSK, PR, Digha coast (BOB)
Reference: Madhavi (1976a); Gupta & Jahan, (1977); Hafeezullah & Dutta (1998);
Saoud et al. (2002)
Description: Body elongate, 5690 long, 576 wide. Tegument spinose. Oral sucker
terminal, surrounded by two alternating complete rings of circumoral spines, 40 in
number. Oral spines 34 long, aboral spines 54 long. Ventral sucker globular, situated
at about one fourth of body length from anterior end. Testes two, post-equatorial,
elliptical, tandem, separated by vitelline follicles. Cirrus-sac claviform, short, just
reaching into hindbody, encloses undivided seminal vesicle, pars prostatica and a long

triacanthi (after Madhavi
1976)
spined cirrus, joins genital atrium near posterior margin of ventral sucker. Genital
atrium long opening near anterior margin of ventral sucker. Ovary globular, pretes-
ticular, separated from anterior testis. Uterus preovarian, metraterm long, spined.
Eggs 62 × 31–35. Vitelline follicles circumcaecal, extend from posterior level of
cirrus-sac to posterior end of body. Excretory vesicle Y-shaped, pore terminal.
Remarks: This species is distinguished from other species of the genus by the fol-
lowing combination of characters. Oral spines 40, in 2 rows, sucker ratio 1:2.0; testes
wide apart separated by vitelline follicles, located in posterior third of body, small
cirrus-sac, vitelline follicles extending to level of cirrus-sac.
Bray & Cribb (2003) prepared a digest for differentiating various species of
Stephanostomum using characters such as extent of vitellarium, number of oral spines
providing information on their hosts and geographic occurrence. The same digest is
simplified here to represent only the Indian species of the genus
Digest of Stephanostomum spp. from marine fishes of India (COS - circum-

oral spines; H - type-host)
1. Vitellarium anterior extent less than 10% (i.e. % of hindbody devoid of
follicles).
32COS; H-Euthynnus sp. Bombay coast; S. loossi Pandey & Tewari, 1984
32 COS; H-Leptomelanosoma indicus, VSK Coast, BOB. S. polynemi Madhavi,
1976
36 COS; 2 ani; H-Rhabdosargus sarba, PR, BOB. S. simhai Gupta & Ahmad,
1979
40 COS; H-Parupeneus macronemus, PR, BOB. S. bhuneshwari Gupta & Puri,
1986
40 COS; H-Triacanthusbiaculeatus VSK, BOB. S. triacanthi Madhavi 1976
2. Vitellarium anterior extent greater than 10% (i.e. % of hindbody devoid of
follicles).
24 COS; H-Chirocentrus dorab, Ratnagiri coast, AS, S. anterospinosum Khan
& Karyakarte, 1989
30-34 COS; H-Caranx sexfasciatus, Carangoides malabaricus, C. chrysophrys,
VSK, BOB.S. madhaviae Bray & Cribb 2003. Syn. S. orientalis (Srivastava,
1939) of Madhavi, 1976
32 COS; H-Lutjanus rivulatus, PR, BOB, S. chauhani Gupta & Ahmad, 1979.
32 COS:H-Pampus argenteus, PR, BOB.S. srivastavai Gupta & Gupta, 1980
34 COS; H.-Lates calcalifer, Rachycentron canadum, AS, BOB, S. cloacum
(Srivastrava, 1939)
34 COS; H-Gazza minuta, PR, BOB. S. gazzai Gupta & Puri, 1986
36 COS; H-Scomberoides tala, MS, BOB. S. attenuatum Hafeezullah, 1971
36 COS; H- Rachycentron canadum. VSK, BOB. S. pseudoditrematis Madhavi,
1976
36-40 OS; H-Megalaspis cordyla.VSK, BOB. S. ditrematis (Yamaguti, 1939)
3. Vitellarium anterior extent at level of ventral sucker
36 COS. H-Fistularia serrata.VSK, BOB. S. adinterruptum Hafeezullah, 1971
36 COS. H-Rachycentron canadum.VSK, BOB. S. microsomum Madhavi, 1976
Genus Stephanostomoides Oshmarin & Mamaev, 1966
Diagnosis: Body elongate. Oral sucker infundibuliform, with 3–5 rows of enlarged
oral spines. Ventral sucker close to oral sucker. Hindbody long. Caeca form uroproct.
Testes two, oval, in posterior half of body. Cirrus-sac long, reaches to about mid-
hindbody. Internal seminal vesicle, oval, bipartite or tripartite. Pars prostatica elon-
gate. Ejaculatory duct long, spined. Genital atrium deep armed with scattered spines.
Ovary subglobular, pretesticular. Uterus pretesticular. Metraterm long, spined. Vitel-
larium restricted to posterior two thirds of hindbody. Type species: S. tenuis (Manter,
1963) Oshmarin & Mamaev, 1966
The genus Stephanostomoides was erected by Mamaev & Oshmarin (1966) with
S. dorabi Mamaev & Oshmarin, 1966, from Chirocentrus dorab in North Vietnam
Bay as the type species. Hafeezullah (1978) considered this species a synonym of S.
tenuis (Manter, 1963) Hafeezullah, 1978, which hence became the type species of
the genus. Three species belonging to this genus have been recorded from India but
only one species S. tenuis is retained as valid.
Stephanostomoides tenuis (Manter, 1963) Hafeezullah, 1978 (Figs. 14.24 and 14.25)
[Syn. Acanthocolpus tenuis Manter, 1963; S. dorabi Mamaev & Oshmarin, 1966; S.
indicus Karyakarte & Yadav, 1976; S. sharmai Gupta & Gupta, 1980]
Host: Chirocentridae: Chirocentrus dorab (Forsskal))
Fig. 14.24
Stephanostomoides tenuis
entire (after Madhavi, 1976)
Fig. 14.25
Stephanostomoides tenuis
oral sucker
Localities: Mandapam (Gulf of Mannar), Gopalpur, Puri, Pondicherry, Visakhapat-

nam (Bay of Bengal) Kozhikode, Mangalore, Karwar (Arabia sea)
Distribution: Fiji, North Vietnam, New Caledonia, India
References: Manter (1963); Hafeezullah (1971a, 1978);
Description: Body elongate, slender, spinose, 8460–11420 long, 690–900 wide.
Oral sucker funnel–shaped, terminal, 320–440 long, 190–300 wide, with circumoral
spines in 5 complete alternating rows of 19 each. Prepharynx 180–280 long. Pharynx
elongate. Oesophagus short. Intestinal caeca extend to posterior extremity, open into
excretory bladder forming uroproct. Ventral sucker protuberant, subspherical, near
anterior extremity. 250–390 long, 320–480 wide. Genital pore immediately anterior
to ventral sucker. Testes elongate, tandem, subequal, contiguous, in posterior third
of body. Cirrus-sac elongate, tubular extends slightly anterior to ovary. Seminal
vesicle entire. Pars prostatica surrounded by a large number of prostate gland cells.
Cirrus spiny. Ovary spherical, pretesticular, contiguous with anterior testis. Uterus
preovarian. Metraterm tubular, spiny. Eggs oval, operculated. 68–83 long. 37–45
wide. Vitellarium follicular, extend from level of metraterm to posterior end of body,
confluent in post-testicular space.
Remarks: S. tenuis is reported as a common species of C. dorab from many localities:
North Vietnam Bay (Mamaev, 1970); Visakhapatnam coast, Bay of Bengal (Madhavi,
1976; Satya Gopal et al., 1985); Sofala Bank (Parukhin, 1976); Orissa coast, Bay of
Bengal (Gupta & Ahmad, 1977); Krusadai Island, Gulf of Mannar (Zhukov, 1977),
South-west coast of India (Radhakrishnan & Nair, 1979); New Caledonia (Bray &
Justine, 2012). Hafeezullah (1991) reported this species from six localities in the
Bay of Bengal, the Arabian Sea and the Gulf of Mannar and synonymized S. indicus
Karyakarte & Yadav, 1976 from C. dorab off Karwar, Arabian Sea and S. sharmai
Gupta & Gupta (1980) from off Puri, Bay of Bengal with S. tenuis.
Genus Tormopsolus Poche, 1926
Tormopsolus is very similar to Stephanostomum being distinguished from the latter
by the lack of enlarged circum-oral spines.
Diagnosis: Body elongated to attenuate. Eyespot pigment present. Oral sucker sub-
terminal, enlarged oral spines absent. Ventral sucker rounded in anterior third of
body. Prepharynx and pharynx present. Oesophagus short or absent. Caeca extend
to posterior end, open via uroproct. Testes two, oval, entire, tandem, in posterior
half of hindbody. Cirrus-sac elongate, claviform. Seminal vesicle bipartite or saccu-
lar. Pars prostatica long. Ejaculatory duct long, spined. Genital atrium deep. Genital
pore median. Ovary rounded, pretesticular. Uterus preovarian, intercaecal. Metraterm
long, spined. Vitellarium restricted to hindbody. Excretory vesicle I-shaped. Type
species T. osculatus (Looss, 1901) Poche 1926.
Fourteen species have been mentioned in the genus of which only eleven species
are regarded as valid. Most species have been recorded from carangid fishes. Two
species of Tormopsolus, T . spatulatum Bilqees, 1972, and T. filiformis Sogandares-
Bernal & Hutton, 1959, were reported from India, but T. spatulatum has been syn-
onymized with T. filiformis (see Madhavi, 1976a; Hafeezullah, 1991).
Tormopsolus filiformis Sogandares-Bernal & Hutton, 1959 (Fig. 14.26)

[Syn. T. rhachicentri Parukhin, 1965; T. spatulatum Bilqees, 1972]
Host: Rachycentridae: Rachycentron canadum (L.)
Locality: Kakinada Bay (1); VSK, BOB (2); Vaco-da-Gama, AS (3)
Distribution: Madeira, Florida, Gulf of Mexico, South China sea, Karachi Coast,
BOB, AS
Ref. 1. Hafeezullah (1978, 1991); 2. Madhavi (1976a); 3. Hafeezullah (1991),
Description: Body filiform, 4800–5650 long, 200–360 wide. Tegument spined.

Eyespot pigment present. Oral sucker 40 × 50–60. Ventral sucker 90 × 90–100
in size, in anterior quarter of body. Forebody 900–1300 long. Prepharynx long. Phar-
ynx well developed. Oesophagus absent. Caeca form uroproct. Genital pore median,
anterior to ventral sucker. Genital atrium extends from genital pore to mid-level of
ventral sucker. Testes 2, tandem, in posterior fourth of body. Cirrus-sac extends well
into hindbody, encloses bipartite seminal vesicle, heavily spined cirrus. Ovary inter-
caecal, pretesticular. Uterus preovarian. Metraterm heavily spined. Eggs 74 by 31.
Vitellarium from posterior tip of cirrus-sac to posterior end of body.
Fig. 14.26 Tormopsolus

filiformis (after
Sogandares-Bernal &
Hutton, 1959)
Remarks: Madhavi (1976b) reported T. filiformis Sogandares-Bernal & Hutton,

1959, from Rachycentron canadum from VSK, BOB and synonymised T. spatulatum
Bilqees, 1972, reported from the same host from Karachi coast, with this species.
Hafeezullah (1991) further synonymized T. rhachycentri Parukhin, 1965, from R.
canadum from South China Sea with this species. The species shows specificity
towards Rachycentron sp. Hafeezullah (1991) cautioned that the spatulate condition
of the prepharyngeal part of the body and the size of the oesophagus depend on the
degree of relaxation of the specimens at the time of fixation. These features cannot
be used as valid criteria for the separation of species.
Genus Pleorchis Railliet, 1896
[Syn. Parapleorchis Al-Yamini & Nahhas (1981)
The distinguishing features of the genus Pleorchis are its multiple testes, H-shaped
caeca and acinous ovary. Poche (1926) erected the family Pleorchiidae to include the
genera Pleorchis and Schistorchis. However, Caballero & Caballero (1952) noted
that the genus Pleorchis has many features in common with the acanthocolpids such
as the spined tegument, the follicular vitellarium, the preovarian uterus, the lack of
an external seminal vesicle and the presence of a distinct cirrus-sac. Based on these
similarities, the genus was placed under Acanthocolpidae (Caballero & Caballero,
1952; Bray, 2005). Six species belonging to this genus have been recorded from
marine fish in India:
P. sciaenae Yamaguti 1938 by Madhavi & Narasimhulu 1986
P. puriense Gupta & Ahmad, 1976
P. indicum Gupta & Puri, 1979
P. psettodesai Gupta & Gupta, 1988
P. srivastavai Saxena, Johri & Gupta, 2010
P. santoshai Gupta, Johri, Chandra & Saxena, 2011
A new genus Parapleorchis was erected by Al-Yamini & Nahhas (1981) to include
Pleorchis puriensis Gupta & Ahmad, 1976, characterizing the genus by the complete
absence of anterior caeca. A second species P. keshavai Gupta, Johri, Chandra &
Saxena, 2011, was added to the genus subsequently (Gupta et al., 2011). However,
Madhavi & Narasimhulu (1986) expressed the view that P. puriensis was based
on a single specimen and suggested that examination of more specimens of this is
needed to confirm the absence of anterior caeca. They synonymized Parapleorchis
with Pleorchis and considered P. puriensis as a synonym of P. sciaenae Yamaguti,
1938. This view was endorsed by Bray (1985). Bray (1985) regarded P. psettodesai
Gupta & Gupta, 1988, also as a synonym of P. sciaenae. Among the other species, P.
indicus Gupta & Puri, 1979, based on a single specimen collected from Epinephelus
diacanthus resembles P. sciaenae in most details; more specimens are to be studied
to determine its status as distinct from P. sciaenae. In the world register of marine
biota, P. keshavai Gupta, Johri, Chandra & Saxena, 2011, is also listed as a synonym
of P. sciaenae. The validity of two recently described species P. srivastavai Saxena,
Johri & Gupta, 2010, from Parascorpaena picta from Deegha coast, BOB and P.
santoshai Gupta, Johri, Chandra & Saxena, 2011, from Parascorpaena picta (Cuvier)
(Scorpaenidae) and Trachinotus botla (Shaw) (Carangidae) from Digha coast remains
to be judged. It is to be noted that descriptions for most of the species were based
on just one or two specimens. According to Madhavi & Narasimhulu (1986) and
Bartoli et al. (2004), the key characters for species differentiation in this genus are
the shape of the caeca, the wide variations in the number of testes and the extent of
the vitellarium. The genus evidently requires a thorough revision. Only one species
of Pleorchis namely P. sciaenae Yamaguti, 1934, whose validity is unquestionable
is included in this account.
Pleorchis sciaenae Yamaguti, 1934 (Fig. 14.27)
[Syn. P. puriense Gupta & Ahmad, 1976; P. psettodesai Gupta & Gupta, 1981; P.
keshavai Gupta, Johri, Chandra & Saxena, 2011; Parapleorchis puriensis (Gupta &
Ahmad, 1976) Al-Yamini & Nahhas, 1981; P. ghanensis Fischthal & Thomas, 1968]
(Ref. Bray (1985)
Hosts: Paralichthyidae: Pseudorhombus diacanthus (Bloch) (1); Sciaenidae: Sciaena
spp (2)
Psettodididae: Psettodes erumei (Bl. & Schn.) (3,4)
Localities: PR, VSK, Digha coast (BOB); Quilon, AS
Fig. 14.27 Pleorchis

sciaenae
Distribution: Japan, BOB, AS, Tonkin Gulf, Vietnam; Kuwait

References: 1. Hafeezullah & Siddiqi (1970); 2. Madhavi & Narasimhulu (1984); 3.
Gupta & Ahmad (1976b); 4. Gupta & Gupta (1981); Al-Yamini & Nahhas (1981),
Nahhas et al. (1998).
Description: Body flat, leaf like, 2530–2540 long, 720–770 wide. Oral sucker spher-
ical, 170–220 in diameter. Ventral sucker spherical, pre-equatorial smaller than oral
sucker, 140–160 in diameter. Prepharynx long. Pharynx ovoid with conspicuous
anterior circular muscle ring. Oesophagus short. Intestinal bifurcation some dis-
tance anterior to ventral sucker. Caeca terminating at posterior end of body; a pair
of caecal diverticula extend anterolaterally just anterior to pharynx. Genital pore
immediately anterior to ventral sucker. Testes 40–44, in intercaecal space on each
side of excretory bladder, in two ventral and two dorsal longitudinal rows. Cirrus-sac
claviform, extends posteriorly well into hindbody. Seminal vesicle bipartite. Pars
prostatica surrounded by many prostate gland cells. Cirrus muscular. Ovary multi-
lobed, median, pretesticular, wider than long. Uterus runs between ovary and genital
pore. Metraterm long. Eggs ovoid, operculated, 68–72 long, 22–30 wide. Vitellarium
follicular, extends from anterior margin of ventral sucker to posterior end of body.
Remarks: This species occurs in a wide range of hosts and has been reported from
several localities in Indo-Pacific.
Other acanthocolpid species
Genus Pseudolepidapedon Yamaguti, 1938
Diagnosis: Body elongate. Eyespot pigment present. Oral sucker subterminal.
Ventral sucker in anterior half of body. Prepharynx long. Pharynx large. Oesophagus
short. Intestinal bifurcation in posterior part of forebody. Caeca long, blind. Testes
two oval, tandem. Cirrus-sac claviform encloses large, convoluted internal seminal
vesicle, poorly developed pars prostatica and long unspined ejaculatory duct. Exter-
nal seminal vesicle absent. Genital atrium small, unarmed. Ovary oval, pretesticular.
Uterus preovarian, intercaecal. Metraterm unarmed. Eggs oval. Vitelline follicles
confined to hindbody or may reach into forebody. Excretory vesicle I-shaped. Type
species: P. paralichthydis Yamaguti, 1938.
Pseudolepidapedon indica Gupta & Ahmad, 1976
Host: Mullidae: Upeneus macronemus (Playfair)
Locality: Puri, Orissa
No. 3
Description: Body plum-shaped, spinose, 2000–3295 long 550–920 wide. Oral
sucker 230–300 in diameter. Ventral sucker larger than oral sucker, in anterior third
of body, 290–450 in diameter. Prepharynx absent. Pharynx ovoid. Oesophagus
short. Caeca extend to posterior extremity. Genital pore median, at level of posterior
third of ventral sucker. Testes oval, contiguous, tandem, subequal, in posterior half
of body. Cirrus-sac saccular, extends either to just anterior to ovary or to its middle.
Seminal vesicle bi-lobed. Pars prostatica surrounded by many prostate gland cells.
Ejaculatory duct short. Ovary spherical or subspherical, pretesticular. Seminal
receptacle absent. Vitellarium follicular, confluent anterior to ventral sucker, extend
from middle of pharynx to posterior extremity. Uterus running between testis and
genital pore. Eggs ovoid, operculated, 94–105 long, 75–86 wide.
Pseudolepidapedon, which has ten described species, was considered an acan-
thocolpid genus by Skrjabin (1954). In the overall morphology, in particular,
in having an alimentary system with long prepharynx, large pharynx and short
oesophagus, they resemble acanthocolpids. Indeed, eight of the described species
(P. paralichthydis Yamaguti, 1938, P. balistis Manter, 1940, P. kobayashii Yamaguti,
1938, P. lethrini Yamaguti, 1952, P. mordvinovae Parukhin, 1978, P. pudens (Linton,
1900), P. sinaloense Bravo-Hollis, 1956 and P. laticaecale Reimer, 1981) have an
overall morphology which is markedly similar to that of a typical acanthocolpid. But
the Indian species, P. indica Gupta & Ahmad, 1976, has its genital pore posterior to
the aperture of the ventral sucker and lacks a prepharynx (Gupta & Ahmad, 1976)
and clearly belongs to another genus. According to Bray et al. (1996), knowledge
of life cycle or larval information, or molecular phylogenetic studies, are needed
to shed more light on the status of this form. The species is temporarily retained in
the genus Pseudolepidpedon. The subfamily Pseudolepidapedinae is subsumed into
the family Acanthocolpidae. Curran & Pulis (2014) found that molecular evidence
placed P. balistis within the genus Stephanostomum.
Lepidauchen nicolli Yadav, 1977
Host: Haemulidae: Pristipoma furcatus (Bl. & Schn.)
Locality: RT, AS
Remarks: Regarded as species inquirendum.
Chapter 15
Superfamily Opecoeloidea Ozaki, 1925
Family Opecoelidae Ozaki, 1925
The Opecoelidae is a large family of digeneans with many species recorded from
marine fishes. There are several reports dealing with descriptions of opecoelids from
marine fishes of India. So far 98 species belonging to 20 genera have been recorded
(Madhavi, 2011). However, considering the richness of the fish fauna of the region and
the common occurrence of these parasites, it is evident that may more species remain
to be detected. The life cycle of only one species Helicometra gibsoni Murugesh,
Madhavi, Swarnakumari, 1993 has so far been elucidated from India (Murugesh et al.
1993).
Diagnosis: Body oval or elongate. Tegument smooth. Oral and ventral suckers
present, the latter may be pedunculate. Digestive system simple. Caeca long, blind, or
unite to form a cyclocoel or open via anus or form uroproct. Genital pore in forebody,
median or submedian. Testes two or numerous, situated in tandem or symmetrically
in hindbody. Cirrus-sac present or absent. Genital pore in forebody. Ovary between
testes and ventral sucker. Vitellarium extensive, occupy wide zones on lateral sides
of body. Excretory bladder I-shaped.
In the recent review of the Opecoelidae (Cribb, 2005a), four subfamilies were
recognized in the family. Recent molecular studies (Shedko et al., 2015; Bray et al.,
2016; Martin et al. 2017) have indicated that the current subfamily arrangement is not
reflected in the postulated phylogeny, in that the Plagioporinae is paraphyletic. Bray
et al. (2016) initiated a re-organization of the subfamilies in erecting a new subfam-
ily, the Helicometrinae, so there are now five nominal subfamilies. The opecoelids
recorded from Indian marine fish, however, fall into four of these five subfamilies.
The Opecoelinae Ozaki, 1925, Plagioporinae Manter, 1947, Stenakrinae Yamaguti,
1970 and Helicometrinae Bray, Cribb, Littlewood & Waeschenbach, 2016 that can be
differentiated by the presence or absence of a well-developed cirrus-sac, a canalicular
seminal receptacle and filamented eggs.
© Crown 2018 405

https://doi.org/10.1007/978-94-024-1535-3_15
406 15 Superfamily Opecoeloidea Ozaki, 1925
Key to subfamilies
1. Eggs filamented ……. Helicometrinae Bray, Cribb, Littlewood & Waeschenbach,

2016
Eggs without filaments …………………………………............……….......... 2
2. Cirrus-sac membranous, poorly developed or absent, seminal vesicle external or
internal ………………..…….………………………….….Opecoelinae Ozaki,
1925
Cirrus-sac well developed, encloses seminal vesicle, pars prostatica and ejacu-
latory duct …………….. ………………………….......................................... 3
3. Canalicular seminal receptacle absent…………………..Stenakrinae Yamaguti,
1970
Canalicular seminal receptacle present ………….…………Plagioporinae Man-
ter, 1947
Subfamily Opecoelinae Ozaki 1925
[Syn: Horatrematinae Srivastava, 1942, Notoporinae Srivastava, 1942, Genitocotyli-

nae Skrjabin & Koval, 1958]
Diagnosis: Body oval to elongate. Ventral sucker often pedunculate, unspecialized

or bears papillae or tentacular processes. Accessory sucker occasionally present.
Caeca blind or unite and form cyclocoel or unite and open through anus or form
uroproct or open through separate ani. Testes two tandem or oblique. Cirrus-sac com-
pletely absent or reduced and enclosing only terminal part of male terminal genitalia.
Seminal vesicle normally external. Ovary pretesticular, entire or lobed. Canalicular
seminal receptacle absent. Uterine seminal receptacle present. Uterus preovarian.
Eggs large, numerous, without filaments. Vitelline fields usually restricted to hind-
body, occasionally extend into forebody, normally confluent in post-testicular region.
Eight genera belonging to this subfamily have been recorded from marine fish in
India. They can be separated by the key provided below;
Key to genera of Opecoelinae.
1. Caeca blind or form uroproct ………………………………..............................2

Caeca united posteriorly ………………………….….….….….….….….….….6
2. Ventral sucker pedunculate, bears papillae………………….…Paropecoelus
Pritchard, 1966
Ventral sucker not pedunculate…………………………………………………3
3. Body elongate, caeca form uroproct……………….……Pseudopecoeloides
Yamaguti, 1940
Body fusiform, caeca blind……………………………………………………...4
Subfamily Opecoelinae Ozaki 1925 407
4. Testes symmetrical, pars prostatica strongly developed……………………….5

Testes tandem, pars prostatica not strongly developed….……………….
…………Pseudopecoelus Von Wicklen, 1946
5. Genital pore opposite pharynx …………………………….…….. Manteriella
Yamaguti, 1955
Genital pore submarginal at level of posterior margin of ventral suck-
er……………………………Notoporus Yamaguti, 1938
6. Accessory sucker present in forebody ………………………….…Anisoporus
Ozaki, 1928
No accessory sucker……………………………7
7. Caeca united and open through anus………………………..…Opegaster Ozaki,
1928
Caeca cyclocoel, no anus…………………...........………….Coitocaecum
Nicoll, 1915
Genus Anisoporus Ozaki, 1928
Generic diagnosis: Body elongate. Ventral sucker pedunculate, larger than oral
sucker, with complex papillae at aperture. Accessory sucker present in forebody.
Caeca unite and open via anus situated close to posterior end of body. Testes tandem,
entire. Cirrus-sac totally absent. Genital pore sinistral, in forebody. Ovary entire.
Uterus restricted to area between ovary and ventral sucker. Vitelline follicles do not
extend into forebody. Excretory vesicle extends to level of ovary. Type species: A.
cobraeformis Ozaki, 1928
Anisoporus orientalis Madhavi, 1975 (Fig. 15.1)
Host: Dactylopteridae: Dactyloptena orientalis (Cuvier)
No: 16
Locality: VSK, BOB
Description: Body elongate, 4640–6480 long, 480–720 wide. Forebody with an

accessory sucker situated on right side slightly below level of bifurcation. Oral sucker
136–175 in diameter. Ventral sucker pedunculate, 216–264 in diameter, possesses
both apertural and peripheral papillae. Sucker ratio 1:1.45–1.6. Prepharynx present.
Pharynx oval. Oesophagus long. Intestinal bifurcation immediately anterior to pedun-
cle of ventral sucker. Caeca join at posterior end of body, open out through anus. Anal
pore ventral and terminal. Genital pore sinistral, opposite posterior border of phar-
ynx. Testes oval, intercaecal, tandem, situated in middle third of hindbody. Seminal
vesicle long, tubular, sinuous, extends to midway between ovary and ventral sucker.
Ovary ovoid, pretesticular, separated from anterior testis. Uterine coils between ovary
and ventral sucker continue anteriorly as narrow tube. Vitelline follicles circumcae-
cal, commence at level of ovary intrude into intergonadal space, filling post-testicular
space. Eggs 39–43 by 23–31. Excretory vesicle I-shaped extends to level of ovary.
Fig. 15.1 Anisoporus

orientalis (after Madhavi
1975)
Remarks: This species differs from the type and only other species of the genus, A.
cobraeformis Ozaki, 1928 from Daicoccus peterseni from Japan in the absence of a
genital sucker, more anterior position of genital pore, longer hindbody and smaller
eggs.
Genus Coitocaecum Nicoll, 1915
[Syn. Ozakia Wisniewski, 1934; Paradactylostomum Zhukov, 1972, Pseudocoito-
caecum Bilqees, 1972]
Coitocaecum was erected by Nicoll (1915) for C. gymnophallum Nicoll, 1915 and
is characterized by the combination of a cyclocoel gut, absence of papillae on the
ventral sucker and a greatly reduced cirrus-sac. The genera Ozakia Wisniewski, 1934;
Paradactylostomum Zhukov, 1972 and Pseudocoitocaecum Bilqees, 1972 have been
regarded as synonyms of this genus (Bray, 1987a, Cribb, 2005a).
Generic diagnosis: Body elongate-oval. Ventral sucker sessile, without papillae,

larger than oral sucker. Caeca united posteriorly. Testes tandem to oblique, entire
or lobed, situated close to posterior end of body. Cirrus-sac small, seminal vesicle
naked, restricted to forebody or enters hindbody. Genital pore sinistral, usually in
forebody. Ovary entire or lobed. Uterine seminal receptacle present. Uterus restricted
to area between ovary and genital pore. Vitelline follicles restricted to hindbody or
may extend well into hindbody. Excretory vesicle I-shaped.
Type species: G. gymnophallum Nicoll, 1915
The genus at present consists of over 40 species represented in freshwater as

well as marine fishes. Geographically, it is cosmopolitan in distribution but prin-
cipally in the Indian and Pacific Oceans. From the Indian waters, six species have
been described under the genus Coitocaecum, two species under Pseudocoitocae-
cum and one species under Paradactylostomum. Merging of these genera caused
many homonyms and synonymies of the species. For instance, C. indicum has
three homonyms: C. indicum Ahmad, 1980 from Engraulis mystax; C. indicum
Ahmad, 1983 from Sufflamen fraenatus (=C. capistratus) and C. indicum (Gupta
and Ahmad, 1978) syn Pseudocoitocaecum indicum Gupta & Ahmad, 1978 from
Upeneus macronemus.
Bray (1987) regarded C. indicum Ahmad, 1980 and Paradactylostomum indicum
Zhukov, 1972 as conspecific to C. thrissocles (Bilqees, 1972) Bray, 1987. Of the
remaining two homonyms, C. indicum (Gupta & Ahmad, 1978) is here renamed as
C. longivesiculum and C. indicum Ahmad, 1983 is retained as such. Similarly, C.
manteri Ahmad, 1980 is a junior homonym of C. manteri Parukhin 1971, necessitat-
ing renaming of the species. The replacement name suggested here is C. purius. Fol-
lowing these alterations, the genus Coitocaecum is now represented by nine species
in Indian marine waters.
Coitocaecum bengalense (Madhavi, Narasimhulu & Shameem, 1986) (Fig. 15.2)
[Syn. Paradactylostomum bengalense Madhavi, Narasimhulu & Shameem, 1986
Host: Engraulidae: Thryssa mystax (Bl. & Schn.)
Locality: Kalingapatnam coast, BOB.
Reference: Madhavi et al. (1986)
Description: Body spatulate, broad posteriorly, narrow anteriorly, 1376–1920 long

and 256–400 wide. Oral sucker 46–76 by 78–104. Ventral sucker 117–136 in diam-
eter situated slightly anterior to midbody. Sucker ratio 1:1.2–1.5. Prepharynx small.
Pharynx oval. Oesophagus fairly long. Caeca long, form cyclocoel. Anus absent.
Genital pore median in between caecal bifurcation and ventral sucker. Testes deeply
Fig. 15.2 Coitocaecum

Narasimhulu & Shameem,
1986)
lobed, tandem in posterior third of body. Cirrus-sac very small, saccular, thin-walled,
in forebody, encloses ejaculatory duct. Seminal vesicle long, winding, extends well
posterior to ventral sucker. Prostatic cells absent. Ovary lobed, postequatorial, pretes-
ticular, submedian. Uterus preovarian. Uterine seminal receptacle in proximal coils
of uterus. Eggs 55–58 by 27–30. Vitelline follicles large, irregularly shaped extend
from intestinal bifurcation to posterior end of body, confluent posteriorly, most of
follicles extracaecal. Excretory vesicle I-shaped extends to level of ovary.
Remarks: This species was initially included in the genus Paradactylostomum

Zhukov, 1972, which was synonymized with Coitocaecum by Bray (1987). P. ben-
galense Madhavi, Narasimhulu & Shameem, 1986, became C. bengalense (Madhavi,
Narasimhulu & Shameem, 1986). C. bengalense is characterized by the spatulate
body, the lobed gonads situated in the posterior third of the body, the long winding
seminal vesicle extending well posterior to the ventral sucker and the extension of
the vitellarium from the intestinal bifurcation to the posterior end of the body.
Coitocaecum bombayense (Ahmad, 1983) Bray, 1987 (Fig. 15.3)
[Syn. Pseudocoitoaeum bombayensis Ahmad, 1983]
Host: Muraenidae: Gymnothorax undulatus (Lacépède)
Locality: BOM, AS
Description: Body elongate, 1320–1690 long, 320–430 wide at level of testes.

Oral sucker oval, 77–93 long, 106–119 wide; ventral sucker larger than oral sucker,
135–145 in diameter, 70–110 post-bifurcal. Prepharynx short. Pharynx oval. Oesoph-
agus tubular. Intestinal bifurcation dorsal, nearer to ventral sucker, in posterior part

bombayense (after Ahmad,
1983)
of anterior third of body. Caeca forming cyclocoel near posterior end of body. Testes
two, transversely elongate, directly tandem, contiguous, intercaecal, smooth in out-
line, in posterior third of body. Seminal vesicle shaped like a slender club, extends
posteriorly well beyond posterior border of ventral sucker. Cirrus-sac fusiform, thin-
walled encloses a small pars prostatica with few prostate gland cells and a short cir-
rus. Genital pore ventral, median, immediately post-bifurcal. Ovary entire, smooth in
outline, submedian, contiguous with anterior testis. Vitellarium follicular, follicles
rounded, circumcaecal, confluent in forebody and posterior to testis, extend from
middle of oesophagus to posterior end of body. Excretory vesicle tubular, reaches to
ovary; pore terminal.
Remarks: This species was initially included in the genus Pseudocoitocaecum
Bilqees, 1972, which was synonymized with Coitocaecum by Bray (1987). Important
features of C. bombayense are: the unlobed gonads, the seminal vesicle extending
beyond the ventral sucker, the immediately post-bifurcal genital pore and the vitel-
larium extending anteriorly as far as the mid-level of the oesophagus.
Coitocaecum chaetodoni Ahmad, 1983
Host: Chaetodontidae: Chaetodon pictus (Forskål)
Locality: BOM, AS
No. 46
Description: Body elongate, flattened, forebody more gradually narrowing, poste-
rior end rounded, 1150–1360 long, 382–431 wide. Oral sucker oval, 110–122 long,
130–135 wide. Ventral sucker 190–236 long, 165–188 wide, 90–110 post-bifurcal,
situated at junction of anterior and middle third of body, aperture transversely elon-
gate. Sucker ratio 1:1.2–1.4. Prepharynx absent. Pharynx ovoid. Oesophagus 48–56
long. Intestinal bifurcation dorsal, usually midway between suckers. Caeca form
cyclocoel, near posterior end of body. Testes two, transversely elongate, tandem,
close together, in posterior third of body; anterior testis kidney-shaped, posterior
testis distinctly bi-lobed. Seminal vesicle slender club-shaped, straight, overlaps
right margin of ventral sucker extends up to mid-level of ventral sucker. Cirrus-sac
fusiform, thin-walled, encloses small pars prostatica with few prostate gland cells and
small cirrus. Genital pore ventral, sinistral, opposite posterior border of oral sucker.
Ovary kidney-shaped, contiguous with anterior testis. Uterus preovarian, with few
large eggs, runs between ovary and ventral sucker. Metraterm undifferentiated. Eggs
few, large, oval, operculate, 76–91 long, 47–69 wide. Vitellarium follicular, follicles
large, circumcaecal, extend from posterior level of pharynx to posterior end of body,
confluent in post-testicular area. Excretory vesicle tubular, reaches to ovary.
Remarks: This species was differentiated from all the known species of the genus
Coitocaecum in having the posterior testis bi-lobed and the anterior testis and ovary
kidney-shaped. Other important features of the species are: the location of the genital
pore at the posterior border of the oral sucker, the vitellarium extending from the
posterior level of the pharynx to the posterior end of the body and the seminal vesicle
being a slender tube extending posteriorly to the mid-level of the ventral sucker.
Coitocaecum indicum Ahmad, 1983

Host: Balistidae: Sufflamen fraenatus (Bl. & Schn.)
Locality: BOM, AS
Description: Body elongate, flattened, 1280–1636 long, 395–425 wide at level of

ventral sucker. Oral sucker 113–129 long, 127–143 wide. Ventral sucker larger than
oral sucker, immediately post-bifurcal, 227–243 long, 210–225 wide, at 410–480
from anterior end of body; aperture oval. Sucker ratio 1:1.57–1.65. Prepharynx
absent. Pharynx ovoid. Oesophagus long, Intestinal bifurcation dorsal, immediately
anterior to ventral sucker. Caeca form cyclocoel, near posterior end of body. Testes
two, tandem, transversely elongate, contiguous, equal in size, in posterior half of
body. Seminal vesicle saccular, narrowing anteriorly, extends posteriorly as far as
posterior margin of ventral sucker, overlaps right margin of it. Cirrus-sac fusiform,
thin-walled, encloses small pars prostatica with few prostate gland cells and short
cirrus. Genital pore ventral, sinistral, at level of mid-oesophagus. Ovary transversely
elongate, immediately pretesticular, slightly submedian. Vitellarium follicular, folli-
cles large, extend from mid-level of oesophagus to posterior end of body, confluent in
post-testicular area. Uterus preovarian, with 3–5 eggs. Metraterm undifferentiated.
Eggs oval, operculate, 75–86 long, 44–52 wide. Excretory vesicle tubular, reaching
to ovary.
Remarks: This species differs from other Indian species of the genus in the club-
shaped seminal vesicle extending to the posterior margin of the ventral sucker, the
testes and ovary being transversely elongate and contiguous, the vitellarium com-
mencing at the level of the intestinal bifurcation and the small eggs.
Coitocaecum longivesiculum nom. nov.pro Pseudocoitocaecum indicum Gupta &
Ahmad, 1978 (Fig. 15.4)
Locality: PR, BOB
No: 15
Reference: Gupta & Ahmad (1978c)
long, 90–115 wide. Ventral sucker larger than oral sucker, 130–170 in diameter.
Prepharynx short. Pharynx ovoid. Oesophagus short. Caeca form cyclocoel. Genital
pore in forebody, at 390–520 from anterior extremity. Testes entire, oval contiguous,
subequal, in posterior third of body. Seminal vesicle elongate, tubular, extends to
anterior margin of ovary. Cirrus-sac thin-walled, in forebody, encloses indistinct
pars prostatica and cirrus. Ovary multi-lobed, submedian, pretesticular. Uterus runs
between ovary and genital pore. Eggs oval, operculated, 50–64 long, 30–40 wide.
Vitellarium follicular, circumcaecal, confluent anterior to ventral sucker, extend from
some distance posterior to intestinal bifurcation to posterior end of body.
Remarks: This species was initially included in the genus Pseudocoitocaecum

Bilqees, 1972 as P. indicum Gupta & Ahmad, 1978 Bray synonymized the genus

longivesiculum (Gupta &
Ahmad, 1978, Acta. Parasit,
26, p. 230)
Pseudocoitocaecum with Coitocaecum. P. indicum became C. indicum (Gupta &

Ahmad, 1978) which is a homonym of C. indicum Ahmad, 1980. It is here renamed
as C. longivesiculum nom. nov. This species is characterized by the long seminal
vesicle extending past the ventral sucker to the level of the ovary, the multi-lobed
ovary, the genital pore anterior to the ventral sucker and distinctly post-bifurcal, the
very short oesophagus and the vitelline follicles being confluent in the forebody.
Coitocaecum purius nom. nov. pro C. manteri Ahmad, 1980 nec C. manteri
Parukhin, 1971 (Fig. 15.5)

purius (after Ahmad, 1980)
Host: Pomacentridae: Glyphidodon bengalensis (Cuv.& Val.).

Locality: PR, BOB
No: 14
Description: Body short, with rounded extremities, 1022–1300 long, 431–460 wide.
Oral sucker subspherical, subterminal, 100–131 long, 130–180 wide. Ventral sucker,
at junction of anterior and middle third of body, larger than oral sucker, 165–195
long, 200–220 wide. Prepharynx short. Pharynx ovoid. Oesophagus long. Caeca form
cyclocoel near posterior end of body. Testes entire, tandem, transversely elongated,
in posterior third of body. Seminal vesicle saccular, extends to posterior margin of
ventral sucker. Cirrus-sac thin-walled, encloses indistinct pars prostatica and cirrus.
Genital pore at level of intestinal bifurcation. Ovary transversely elongated, pretes-
ticular. Uterus extends between ovary and ventral sucker. Eggs 87–96 long 50–73
wide. Vitellarium follicular, extend from middle of oesophagus to posterior end of
body.
Remarks: This species is characterized by a combination of characters such as the

genital pore at the level of the intestinal bifurcation, the extension of the seminal
vesicle to the posterior margin of the ventral sucker, the vitellarium up to middle of
the oesophagus and considerably large eggs.
Coitocaecum scombri Ahmad, 1980 (Fig. 15.6)
Locality: PR, BOB
No: 8

scombri (after Ahmad, 1980,
Acta. Parasit, 27, p. 426)
Description: Body elongate, with rounded extremities. Oral sucker subterminal.

Ventral sucker subspherical, larger than oral sucker, at junction of anterior and the
middle third of body, 262–276 long, 245–251 wide. Prepharynx absent. Pharynx
ovoid. Oesophagus 100–120 long. Caeca form cyclocoel near posterior end of body.
Testes entire, transversely elongated, separated from each other, postequatorial. Sem-
inal vesicle saccular extends to anterior margin of ventral sucker. Cirrus-sac thin-
walled, 100–120 long, encloses in distinct pars prostatica and cirrus. Genital pore
at level of intestinal bifurcation. Ovary bi-lobed, immediately pretesticular. Uterus
extends between ovary and ventral sucker. Eggs ovoid, operculated, 60–80 long
30–46 wide. Vitellarium follicles circumcaecal, extend from middle of oesophagus
to posterior end of body.
Remarks: The important characters of this species include the location of the gen-
ital pore at bifurcal level, the separated testes, the bi-lobed ovary, the vitellarium
extending to the bifurcal level and the extension of the seminal vesicle to the anterior
margin of the ventral sucker.
Coitocaecum thapari Ahmad, 1980 (Fig. 15.7)
Host: Synodontidae: Trachinocephalus myops (Forster)
Locality: PR, BOB
Number: 9
Description: Body elongate, 3360–3471 long, 865–877 wide. Oral sucker spherical,
subterminal, 210–223 in diameter. Ventral sucker larger than oral sucker, close behind
pharynx, 270–293 long, 320–342 wide. Prepharynx absent. Pharynx ovoid. Caeca
form cyclocoel near posterior end of body. Testes entire, tandem, subequal, trans-
versely elongate, separated from each other, in middle third of body. Seminal vesicle

thapari (after Ahmad, 1980,
Acta. Parasit, 27, p. 427)
short, winding, extends to one-third length of ventral sucker. Cirrus-sac thin-walled

encloses indistinct pars prostatica and cirrus. Genital pore post-pharyngeal. Ovary
triangular, immediately pretesticular, pre-equatorial. Uterus voluminous extends
between ovary and ventral sucker. Eggs oval, operculated, 62–67 long, 38–50 wide.
Vitellarium follicular, circumcaecal, extend from level of posterior margin of ventral
sucker to posterior extremity. Excretory bladder tubular; excretory pore terminal
Remarks: The most notable distinguishing features of C. thapari are: the separated,
oval testes, the vitellarium restricted to the hindbody, the genital pore at the level
of the pharynx, the short, winding seminal vesicle extending to the middle of the
ventral sucker.
Coitocaecum thrissoclesis (Bilqees, 1972) Bray, 1987
[Syn. Pseudocoitocaecum thrissoclesis Bilqees, 1972; Paradactylostomum indicum
Zhukov, 1972; C. indicum Ahmad, 1980]
Host: Engraulidae: Thryssa mystax. (Bloch & Schneider)
Locality: PR, BOB
No. 12
Reference: Bilqees (1972); Ahmad (1980b)
Description: Body short, 1960–2130 long, 441–478 wide. Oral sucker subspheri-
cal, 182–196 long, 128–132 wide. Ventral sucker at junction of anterior and mid-
dle third of body, 240–290 long, 228–241 wide. Prepharynx short. Pharynx ovoid.
Oesophagus long. Caeca form cyclocoel near posterior end of body. Testes bi-lobed,
tandem, subequal, close together, in posterior third of body. Seminal vesicle saccular,
extends slightly beyond posterior margin of ventral sucker. Cirrus-sac thin-walled,
90–110 long, enclosing indistinct pars prostatica and cirrus. Genital pore median,
post-bifurcal. Ovary tri-lobed, contiguous with anterior testis, post equatorial. Uterus
confined to region between ovary and ventral sucker. Eggs few, oval, operculated,
60–82 long, 40–51 wide. Vitellarium follicular, circumcaecal, confluent anterior to
ventral sucker, extend from region posterior to pharynx to posterior extremity. Excre-
tory bladder tubular; excretory pore terminal.
Remarks: According to Bray (1987a), Pseudocoitocaecum thrissoclesis Bilqees,
1972, from Thrissocles purava from the Karachi coast of Pakistan, Paradactylosto-
mum indicum Zhukov, 1972, from Setipinna phasa from the Bay of Bengal and
Coitocaecum indicum Ahmad, 1980, from Thryssa mystax of the Puri Coast, Bay of
Bengal, are conspecific and to be referred as Coitocaecum thrissoclesis (Bilqees,
1972) Bray, 1987. This species shows specificity towards engraulid fishes. It is char-
acterized by the somewhat lobed gonads, post-bifurcal genital pore, the saccular
seminal vesicle extending slightly posterior to the ventral sucker and the vitellarium
extending anteriorly to the ventral sucker to the intestinal bifurcation.
Bray (1987a) arranged the various species of Coitocaecum into five groups. The
Indian species from marine hosts are distributed in the first three groups as follows:
Group A: species with uninterrupted vitelline follicles extending into forebody, and
the seminal vesicle not extending posteriorly to the ventral sucker: C. chaetodoni,
C. purius, C. scombri
Group B: species with vitelline follicles extending into the forebody and the seminal
vesicle extending posteriorly to the ventral sucker into the hindbody: C. bengalense,
C. bombayense, C. indicum, C. thrissoclesis, C. longivesiculum
Group C: species with vitelline follicles restricted to hindbody and the seminal vesicle
not extending posteriorly to ventral sucker: C. thapari
Key to Indian species of the genus Coitocaecum
1. Genital pore at level of pharynx ………………………........................................2

Genital pore near intestinal bifurcation ……………………….….……..............4
2. Testes separated, vitellarium extends up to posterior margin of ventral sucker…...
C. thapari Ahmad, 1980
Testes contiguous, vitellarium extends anterior to ventral sucker………………3
3. Seminal vesicle short extends to one-third length of ventral sucker, sucker ratio
1:1.2–1.4………………………C. chaetodoni Ahmad, 1983
Seminal vesicle extends to posterior margin of ventral sucker, sucker ratio
1:1.57–1.65…………………………. C. indicum Ahmad 1983
4. Seminal vesicle extends to near posterior margin of ventral sucker or just beyond
………………………………………………………………………………….5
Seminal vesicle not extending posterior to ventral sucker …………………….6
5. Ovary deeply lobed ………………………………………………………….....7
Ovary entire ……………………….C. bombayense (Ahmad, 1983)
6. Testes and ovary lobed……………………..…… C. thrissoclesis (Bilqees, 1972)
Testes entire, transversely oval………………………………..………………..8
7. Testes lobed …………………….. C. bengalense (Madhavi, Narasimhulu &
Shameem 1986)
Testes entire, not lobed …………………………………… C. longivesiculum
nom. nov.
8. Testes contiguous, seminal vesicle extends to near anterior margin of ventral
sucker ……………………………C. scombri Ahmad, 1980
Testes separated, seminal vesicle extends up to posterior margin of ventral sucker
or beyond……………………………………………………………………….9
9. Testes deeply lobed ………………. P. bengalense (Madhavi, Narasimhulu &
Shameem, 1986)
Testes oval ……………………………………… C. purius nom. nov.
Genus Manteriella Yamaguti, 1958

Manteriella was proposed by Yamaguti (1958) for M. crassa (Manter, 1947). It is
recognized by the combination of a ventral sucker without papillae, blind caeca, the
complete absence of a cirrus-sac, the genital pore close to the pharynx, the testes
strongly oblique or symmetrical, the large and prominent pars prostatica and the
vitelline follicles extending well into the forebody.
Diagnosis: Body oval or elongate. Ventral sucker simple, larger than oral sucker.
Caeca terminate blindly close to posterior end of body. Testes diagonal or tandem,
entire or lobed, well separated from posterior end of body. Cirrus-sac absent. Naked
pars prostatica unusually large, prominent. Genital pore sinistrally submarginal at

level of anterior margin of ventral sucker. Ovary entire or lobed. Uterus restricted
to area between ovary and genital pore. Vitelline follicles enter forebody to level
of pharynx. Excretory vesicle extends to level of ovary. Type species: M. leiognathi
Yamaguti, 1938.
A single species M. yamagutii Ahmad, 1990 has been reported from marine fish from
India.
Manteriella yamagutii Ahmad, 1990 (Fig. 15.8)
Host: Siganidae: Siganus javus (Linn.)
Locality: Panjim, Goa, AS
Description: Body plump, 1230–1480 long by 660–730 wide. Oral sucker spherical,
subterminal, 110–160 in diameter. Ventral sucker 310–360 long by 390–480 wide,
pre-equatorial, situated at junction of anterior and middle third of body. Sucker ratio
l:3.0–3.54. Prepharynx absent. Pharynx oval. Intestinal bifurcation closer to ventral
sucker than to oral sucker. Caeca terminate at posterior end of body. Testes two, multi-
lobed, diagonally tandem, confined to posterior half of body, subequal in size. Post-
testicular space 140–220 long. Cirrus-sac absent. Seminal vesicle tubular, extends
just posterior to posterior margin of ventral sucker. Pars prostatica thick-walled, bul-
bous, surrounded by prostate gland cells. Cirrus short. Genital pore ventral, opposite
anterior margin of pharynx. Ovary spherical, submedian, dextral, immediately ante-
rior to posterior testis. Uterus confined between anterior margin of anterior testis and
ventral sucker. Eggs elongate, operculate, 52–62 long by 30–40 wide. Vitellarium
consist of numerous follicles, lateral and ventral to caeca, few invading intercaecal
space, extend from level of intestinal bifurcation to posterior end of body. Excre-
Fig. 15.8 Manteriella

yamagutii (after Ahmad,
1990)
tory vesicle tubular, sigmoid, reaches anteriorly up to anterior testis. Excretory pore
terminal.
Remarks: This species was differentiated from the type species M. crassa (Manter,
1947) Yamaguti, 1958, in having multi-lobed testes, the seminal vesicle extending
posterior to the ventral sucker, the vitellarium non-confluent anteriorly and other
minor differences.
Genus Notoporus Yamaguti, 1938
[Syn: Horatrema Srivastava, 1942]
The genus Notoporus was proposed by Yamaguti (1938) with N. leiognathi Yamaguti,
1938 as the type species. It is characterized by the sinistral genital pore, a prominent
pars prostatica, the absence of a cirrus-sac and accessory sucker, blind caeca and
the ventral sucker without papillae. The most significant feature of the genus is the
unusually large and prominent pars prostatica. Horatrema Srivastava, 1942 with the
type species H. pristipomatis Srivastava, 1942 was regarded as a synonym of this
genus (Ahmad, 1985a).
Diagnosis: Body oval or elongate. Ventral sucker sessile, without papillae. Caeca
terminate blindly close to posterior extremity. Cirrus-sac absent. Pars prostatica
unusually large and prominent. Genital pore sinistrally submarginal at level of ante-
rior margin of ventral sucker. Ovary entire to tri-lobed. Uterus restricted to area
between ovary and genital pore. Vitelline follicles enter forebody to level of pharynx.
Excretory vesicle tubular, extends to ovary. Type species: N. leiognathi Yamaguti,
1938.
Apart from N. pristipomatis (Srivastava, 1942), three species belonging to this
genus have been recorded from marine fish in India. N. stunkardi Ahmad, 1985; N.
gibsoni Ahmad, 1987 and N. fotedari Ahmad & Dhar, 1988. According to Cribb
(2005a), the various species reported under the genus show significant differences
from the type species and may eventually be moved to other genera.
Notoporus fotedari (Ahmad & Dhar, 1988) Madhavi, 2011 (Fig. 15.9)
[Syn. Neonotoporus fotedari Ahmad & Dhar, 1988]
Host: Chaetodontidae: Chaetodon vagabundus (L.)
Locality: PR, BOB
No. 14
Reference: Ahmad & Dhar (1988)
Description: Body elongate, 1305–1470 long, 235–320 wide at level of genital

pore. Oral sucker inverted bell-shaped, terminal, 160–190 long and 175–205 wide.
Ventral sucker pre-equatorial, 175–205 long and 150–170 wide, situated in anterior
part of middle third of body. Sucker width ratio 1:0.82–0.85. Prepharynx, pharynx,
oesophagus present. Intestinal bifurcation dorsal, roughly midway between suckers.
Caeca simple, terminate at posterior end of body. Testes two, entire, intercaecal,
diagonally tandem separated from one another or contiguous, situated in posterior
third of body, equal in size. Cirrus-sac club-shaped, curved, dorsal to ventral sucker,
extends posteriorly as far as midway between ovary and anterior testis. Seminal
Fig. 15.9 Notoporus

fotedari (after Ahmad &
Dhar, 1988)
vesicle tubular, winding. Prostatic vesicle bulbous, surrounded by prostate gland

cells. Cirrus short. Genital pore marginal, towards left margin of body, at anterior
level of ventral sucker. Ovary three-lobed, pretesticular, submedian, dextral. Uterus
confined to intercaecal field between antenor testis and ventral sucker. Metraterm
thin-walled opening into genital pore. Eggs elongate, operculate 90–100 by 50–60
wide. Vitellarium follicular, fields interrupted at ventral sucker level on left side
only, extend from posterior margin of pharynx to posterior end of body, confluent
in forebody. Excretory vesicle tubular, sigmoid reaches anteriorly as far as level of
anterior testis. Excretory pore terminal.
Remarks: This species is characterized by the inverted bell-shaped oral sucker, the
entire testes, the three-lobed ovary, the long cirrus-sac extending to midway between
the ovary and testes and the vitellarium interrupted at ventral sucker level on one
side. The long cirrus-sac indicates that this species is in the wrong subfamily.
Notoporus gibsoni Ahmad, 1987 (Fig. 15.10)
Host: Drepanidae: Drepane punctata (L.)
Locality: Goa, AS
No. 29
Description: Body elongate, 1370–l825 long and 260–305 wide at level of testes.
Oral sucker saucer-shaped terminal, 125–148 long and 185–218 wide. Ventral sucker
150–165 long and 110–135 wide, situated in anterior part of middle third of body.
Sucker ratio 1:0.59–0.61. Prepharynx absent. Pharynx oval. Intestinal bifurcation
midway between suckers. Caeca terminate near posterior end of body. Testes two,
entire, longitudinally elongate, oblique, close together, subequal in size, situated in
posterior third of body. Cirrus-sac absent. Seminal vesicle tubular, slightly winding,
Fig. 15.10 Notoporus

gibsoni (after Ahmad, 1987,
Pakistan J. Zool. 19, 322)
curved anteriorly, extends posteriorly up to about midway between ovary and ven-
tral sucker. Pars prostatica thick-walled, bulbous, lined internally with tall epithelial
cells, surrounded by prostate gland cells. Cirrus short. Genital pore towards left body
margin, midway between ventral sucker and intestinal bifurcation. Ovary pyriform,
submedian, to right of median line, post–equatorial, intercaecal. Uterus confined
between anterior margin of posterior testis and ventral sucker. Eggs elongate, oper-
culate, 110–115 long and 55–63 wide. Vitellarium follicular, fields interrupted at
ventral sucker level on both sides, extend from bifurcal level to posterior end of
body, confluent in forebody. Excretory vesicle tubular, curved, reaches to anterior
level of anterior testis.
Remarks: This species most closely resembles N. stunkardi Ahmad, 1985 in the
elongate body, the vitelline fields interrupted at the level of the ventral sucker on
both sides, the entire gonads, the seminal vesicle extending posteriorly to midway
between the ovary and the ventral sucker. It differs from N. stunkardi as follows: the
saucer-shaped oral sucker which is larger than ventral sucker, rather than spherical
and smaller; the pharynx and the eggs larger.
Notoporus pristipomatis (Srivastava, 1942) Ahmad, 1985 (Fig. 15.11)
[Syn.Horatrema pristipomatis Srivastava, 1942]
Host: Haemulidae: Pomadasys commersoni (Lacepede); Leiognathidae: Leiognathus
daura (Cuvier); L. bindus (Val.); Secutor insidiator (Bloch); Gazza minuta (Bloch)
Locality: AS, BOB
Reference: Srivastava (1942b); Madhavi (1975a); Ahmad (1985a)
Description: Body pear-shaped with a narrow anterior region and broad leaf-like
posterior region, both demarcated by a constriction. Length 1250–1290 by 300–330.
Tegument smooth. Oral sucker 130–150 by 140–190 in diameter. Ventral sucker

pristipomatis (after Ahmad,
1985)
larger, 160–180 in diameter. Sucker ratio 1:1.20–1.25. Intestinal bifurcation almost

at midway between oral and ventral suckers. Caeca long. Testes spherical, diagonal,
in posterior third of body. Seminal vesicle long, sinuous tube extends well posterior
to ventral sucker. Pars prostatica voluminous, cirrus small. Genital pore post-bifurcal,
sinistral, situated ventral to left caecum. Ovary transversely oval, in front of right
testis. Vitellarium follicles asymmetrical in distribution extend from posterior end to
level of intestinal bifurcation on right side and to ventral sucker on left side. Uterus
with few eggs in between ventral sucker and ovary. Eggs large, 88 × 42–46.
Remarks: This species was first described by Srivastava (1942) as Horatrema pris-
tipomatis from Pomadasys commersoni. Madhavi (1975) reported it from three
species of leiognathid fishes of Visakhapatnam coast. Ahmad (1985a) after examin-
ing the specimens collected from the type host from Bombay Coast synonymized the
genus Horatrema with Notoporus. N. pristipomatis is characterized by the leaf-like
body with a constriction, the asymmetrical distribution of vitelline follicles and the
large eggs.
Notoporus stunkardi Ahmad, 1985 (Fig. 15.12)
Host: Tetraodontidae: Gastrophysus lunaris (Bl. Schn.)
Locality: BOM, AS
No: 17
Description: Body elongate, 1170–1430 long by 240–300 wide. Oral sucker 32–40
in diameter. Ventral sucker larger than oral sucker, 88–100 in diameter, situated in
anterior part of middle third of body; aperture transversely elongate, provided with
anterior and posterior muscular lips. Sucker ratio 1:2.5–2.67. Prepharynx absent.

stunkardi (after Ahmad,
157)
Pharynx barrel-shaped. Oesophagus long. Intestinal bifurcation dorsal, nearer to

ventral sucker than to oral sucker. Caeca terminate blindly near posterior end of
body. Testes two, entire, diagonal, close together, postequatorial, situated in posterior
third of body. Cirrus-sac absent. Seminal vesicle tubular, curved anteriorly, extends
posteriorly to about midway between ventral sucker and ovary. Pars prostatica very
wide, surrounded by prostate gland cells. Cirrus long. Genital pore marginal, dextral,
midway between ventral sucker and intestinal bifurcation. Ovary entire, submedian,
to left of median line, oval, immediately pretesticular. Seminal receptacle absent.
Uterus confined to intercaecal field between anterior testis and ventral sucker. Eggs
elongate, operculate, 75–79 long by 32–42 wide. Vitellarium extends from posterior
third of oesophagus to posterior end of body, interrupted at ventral sucker level on
both sides, confluent in forebody and post-testicular zone. Excretory vesicle tubular,
curved, reaches to level of anterior testis.
Remarks: The distinguishing characters of the species are: the very long oesophagus,
presence of muscular lips on the ventral sucker and the interruption of vitellaria
opposite ventral sucker on both the sides.
Key to Indian species of the genus Notoporus
1. Vitellarium interrupted at ventral sucker level on both sides…………………..2

Vitellarium interrupted at ventral sucker level on one side
only……………………………………………………………………………..3
2. Ventral sucker much larger than oral sucker, sucker ratio 1:2.5–2.67…………
…………………………………………N. stunkardi Ahmad, 1985
Ventral sucker smaller than oral sucker ratio 1:0.59–0.61…………….….………

……… N. gibsoni Ahmad, 1987
3. Body flat, leaf-like with a constriction at level of ventral sucker, vitellarium
extend to level of bifurcation …………………………..…… N. pristipomatis
(Srivastava, 1942)
4. Body elongate, with out any constriction, vitellarium extends to level of pharynx
…………………………………N. fotedari (Ahmad & Dhar, 1988)
Genus Opegaster Ozaki, 1928
The genus is characterized by the combination of caeca uniting to open through a
common anus, simple papillae on the aperture of the ventral sucker, highly reduced
cirrus-sac and vitelline follicles extending well into the forebody. The genus appears
very similar to Opecoelus Ozaki, 1925, the anterior extent of vitellarium being the
only character separating the two genera. Aken’Ova (2008) synonymized the two
genera. However, this is not adopted here, following Cribb (2005).
Generic diagnosis: Body elliptical. Ventral sucker sessile or protuberant, bears 3–5
pairs of small papillae at aperture. Caeca unite, open out through common anus near
posterior end of body. Testes tandem to oblique entire or deeply lobed. Cirrus-sac
small. Seminal vesicle naked, usually restricted to forebody, may enter hindbody.
Genital pore sinistral, in forebody. Ovary entire or lobed. Uterus restricted to area
between ovary and genital pore. Vitelline follicles extend into forebody. Excretory
vesicle extends up to ovary or testes. Type species: O. ovatus Ozaki, 1928.
Two species belonging to this genus have been reported from India: O. ditrema-
tis Yamaguti, 1942 and O. paramacrorchis Hafeezullah, 1971, the latter was syn-
onymized with O. ditrematis by Madhavi (1975a).
Opegaster ditrematis Yamaguti, 1942 (Fig. 15.13)
[Syn: O. paramacrorchis Hafeezullah, 1971.]
Hosts: Psettodidae: Psettodes erumei (Bl. & Schn.); Paralichthyidae: Pseudorhom-
bus triocellatus (Bl. & Schn.); P. micrognathus Norman; Synodontidae: Saurida
tumbil Bloch; S. undosquamis Richardson; Trachinocephalus myops (Bl. & Schn.);
Diodontidae: Diodon hystrix L.; Sciaenidae: Johnius aneus Bloch; Otolithes ruber
(Bl. & Schn), Protonibea diacanthus Lacépède; Priacanthidae: Priacanthus tyenus
Richardson; Terapontidae: Terapon jarbua Forskål (1) Apogonidae: Apogon ellioti
(Day) (2) Location: Intestine Locality: VSK, BOB. Reference: Madhavi (1975) (1);
Hafeezullah (1971) (2) Distribution: N.W. Pacific (Yamaguti, 1942) Hawaii (Manter
& Pritchard, 1960, Pritchard, 1966); Great Barrier Reef (Bray & Cribb, 1989); Indian
Ocean (Hafeezullah, 1971; Madhavi, 1975a).
Description: Body elongate oval. Oral sucker rounded. Ventral sucker protuberant,
larger than oral sucker, anterior and posterior lips bear 5 short dome-shaped papillae.
Prepharynx distinct. Oesophagus long. Intestinal bifurcation in posterior forebody.
Caeca narrow, unite posteriorly, open out through a single anal pore situated ventro-
terminally. Testes tandem, lobed. Seminal vesicle saccular, reaches into hindbody
Fig. 15.13 Opegaster

ditrematis (after
Hafeezullah, 1971 Courtesy
Cambridge Univ. Press,
or to about posterior edge of ventral sucker. Cirrus-sac membranous, contains few

prostatic cells and narrow cirrus. Genital pore sinistral, at level of posterior or mid-
oesophagus. Ovary oval or indented anteriorly. Uterine seminal receptacle present.
Uterus preovarian, intercaecal. Metraterm thin-walled, narrow. Eggs oval, 56–64 ×
26–32. Vitelline follicles lie dorsal, lateral and ventral to caeca, extend anterior to
ventral sucker, up to bifurcation.
Remarks: The species is a generalist occurring in a wide range of taxonomically

unrelated marine fish and exhibits morphological variations (Manter & Pritchard,
1960; Madhavi, 1975a). Variations were noted in the shape of testes, position of
genital pore, extent of seminal vesicle and vitellarium, sucker ratio and egg size.
Madhavi (1975a) considered O. paramacrorchis Hafeezullah, 1971, as a synonym
of O. ditrematis, considering that the only feature differentiating the two species
namely the extent of excretory vesicle may not be an important taxonomic character
for differentiation of species in the genus.
Genus Paropecoelus Pritchard, 1966
The genus was proposed by Pritchard (1966) for P. sogandaresi Pritchard, 1966.
It is characterized by combination of the following characteristics: caeca united
posteriorly to open via an anus, a cirrus-sac enclosing only the cirrus and an indistinct
pars prostatica and a ventral sucker that bears complex branching papillae at margin
and aperture (Cribb, 2005a).
Diagnosis: Body elongate. Ventral sucker with eight or 16 papillae along margin and
aperture. Caeca unite and open through anus close to posterior end of body. Testes
tandem, entire or lobed, in posterior half of hindbody. Cirrus-sac small. Genital pore
sinistral, in mid-forebody. Ovary entire or lobed, pretesticular, well separated from
testis. Uterus restricted to area between ovary and ventral sucker. Vitelline follicles
restricted to hindbody. Excretory vesicle extends to level of ovary
Type species: P. sogandaresi Pritchard, 1966.

Seventeen species belonging to this genus have been recorded from marine fishes.
According to Rohner and Cribb (2013), the various species are readily distinguished
by the number of marginal papillae on the ventral sucker, the proportions of the body,
the extent of vitelline follicles and the position of genital pore. They included the
17 species of the genus in 3 groups chiefly based on the number of papillae on the
ventral sucker. Nine species of Paropecoelus have been recorded from Indian marine
fish, of which three species P. bhabai Gupta and Puri, 1985, P. filiformis Ahmad,
1978 and P. manteri require reconsideration according to Rohner and Cribb (2013).
Following their view, the three species are considered here as species inquirendae. P.
theraponi Gupta and Ahmad, 1976 also falls into this category since the description
of the species based on a single specimen lacks many details including the number
of papillae on the ventral sucker. The remaining five species are distributed among
the groups of Rohner & Cribb (2013) as follows:
1. Species with eight marginal papillae. Eg. P. dollfusi Ahmad, 1983; P. indicus
Madhavi, 1973; P. overstreeti Ahmad, 1983; P. sciaeni Ahmad, 1983.
2. Species with 16 marginal papillae. Eg. P. pritchardae Ahmad, 1983
Of the four species included in the group 1, in P. dollfusi the vitellarium extends only
to the level of ovary, where as in the remaining three species they extend anteriorly
to midway between the ovary and the ventral sucker. P. indicus is distinct in having
lobed gonads. In P. sciaeni, the vitellarium is interrupted opposite the gonads as is
the case with P. pritchardae.
Brief descriptions for the five Indian species are given below:
Paropecoelus dollfusi Ahmad, 1983 (Fig. 15.14)

Host: Mullidae: Upeneus sulphureus (Cuvier)
Locality: PR, BOB
No. 32
Description: Body elongated, nearly parallel sides with rounded extremities,

1915–2322 long by 287–322 wide. Oral sucker terminal, 68–92 in diameter; oral
opening triangular, anteroventral. Ventral sucker pedunculate, larger than oral sucker,
112–140 long by 135–168 wide, with four uniramous peripheral papillae one on each
anterolateral and each postero-lateral margin; apertural papillae absent Prepharynx
present. Pharynx oval. Oesophagus long. Intestinal bifurcation dorsal, immediately
anterior to ventral sucker. Caeca united posteriorly opening subterminally via anus.
Testes two, deeply bi-lobed, tandem, median, intercaecal, in middle of posterior half
Fig. 15.14 Paropecoelus

dollfusi (after Ahmad, 1983)
of body. Seminal vesicle tubular, sinuous, extends slightly posterior to ventral sucker
peduncle. Cirrus-sac small, encloses indistinct pars prostatica and cirrus. Genital pore
sinistral, at anterior level of oesophagus. Ovary four-lobed, post equatorial, imme-
diately pretesticular, median, intercaecal, in posterior part of middle third of body.
Uterine seminal receptacle present. Vitellarium follicular, overlap caeca dorsally, in
lateral fields, extend from anterior border of ovary up to posterior extremity, filling
most of post-testicular space. Uterus fairly long, coiled, running between ovary and
genital pore. Eggs oval, yellowish, 40–50 long by 28–35 wide. Excretory vesicle
I-shaped.
Remarks: The important features of P. dollfusi are considered to be: the lobed gonads,
the vitellarium extending anteriorly as far as the ovary level, the ventral sucker with
eight marginal papillae and sinuous seminal vesicle which extends only slightly
posterior to ventral sucker.
Paropecoelus indicus Madhavi, 1975 (Fig. 15.15)
Host: Mullidae: Upeneus sulphureus Cuvier
Locality: VSK, BOB
Description: Body elongate, rounded posteriorly, tapering anteriorly, 2320–2880

long, 352–448 wide at level of ovary. Forebody 12–14% of total length. Oral sucker
66–78 in diameter. Ventral sucker on short pend uncle, 117–156 in diameter, with four
biramous papillae one on each anterolateral and each postero-lateral margin and four
smaller apertural papillae two on anterior lip of aperture interlocking with two similar

indicus (after Madhavi 1975)
papillae on posterior lip of aperture. Prepharynx short. Pharynx oval. Oesophagus

98 long. Intestinal bifurcation anterior to peduncle. Caeca narrow, united posteri-
orly and opening through common ventro terminal anus. Genital pore sinistral at
mid-oesophageal level. Testes deeply indented on anterior and posterior margins,
tandem, close together, in posterior half of hindbody, post-testicular space 426–624.
Cirrus-sac small, in conspicuous. Seminal vesicle long, tubular, sinuous, extends
slightly posterior to ventral sucker. Ovary four-lobed, pretesticular. Seminal recep-
tacle absent. Uterus fairly long, with numerous coils in preovarian area. Eggs oval,
yellowish, collapsed, 54–58 by 27–31. Vitelline follicles irregular, in lateral fields
commencing midway between peduncle and ovary, slightly intrude into intergonadal
space, filling most of post-testicular area. Excretory vesicle I-shaped extending to
ovary.
Remarks: The important features of the species are considered to be the deeply lobed
testes, the seminal vesicle which extends only slightly posterior to the ventral sucker
and in the extension of the vitellarium to midway between the ventral sucker and the
ovary and the presence of eight papillae on the ventral sucker.
Paropecoelus overstreeti Ahmad, 1983 (Fig. 15.16)
Host: Terapontidae: Terapon theraps (Cuv. & Val)
Locality: BOM, AS

1983)
Description: Body elongate, 2200–2300 long by 279–312 wide at level of posterior

testis. Oral sucker spherical, terminal, 105–128 in diameter. Ventral sucker peduncu-
late, oval, 127–150 in diameter, larger than oral sucker, with four biramous peripheral
papillae one on each anterolateral and each postero-lateral margin; apertural papil-
lae absent. Prepharynx, pharynx and oesophagus present. Intestinal bifurcation dor-
sal, midway between pharynx and ventral sucker. Caeca united posteriorly opening
ventro-subterminally via anus. Testes two, elliptical, intercaecal, contiguous, median,
tandem, postequatorial. Post-testicular space 13–15% of total body length. Seminal
vesicle thin-walled, tubular, winding, extends slightly posterior to ventral sucker.
Cirrus-sac small, thin-walled, encloses indistinct pars prostatica and cirrus. Genital
pore sinistral, at level of mid–pharynx. Ovary spherical, postequatorial, in posterior
part of middle third of body, contiguous with anterior testis. Uterine seminal recep-
tacle present. Uterus intercaecal, running between ovary and genital pore. Eggs oval,
yellowish, 49–58 long by 34–38 wide. Vitellarium follicular, lateral, overlapping
caeca, extending from some distance posterior to ventral sucker to posterior end of
body, filling most of post-testicular space. Excretory vesicle I-shaped, reaching to
ovary.
Remarks: The distinguishing features of the species are the oval and contiguous
gonads, the vitellarium which extends to midway between the ovary and ventral
sucker, the ventral sucker larger than the oral sucker, the eight marginal papillae
on the ventral sucker and the seminal vesicle extending just posterior to the ventral
sucker.
Paropecoelus pritchardae Ahmad, 1983 (Fig. 15.17)
Host: Lutjanidae: Lujanus fulviflamma (Forskål)

pritchardae (after Ahmad,
1983)
Locality: PR, BOB

No. 68
Description: Body elongated, 2280–2300 long by 278–310 wide. Oral sucker

110–130 long by 100–115 wide. Ventral sucker pedunculate, 130–150 long by
140–160 wide, larger than the oral sucker, with pair of four-pronged lateral tentacles
one on each anterolateral and each postero-lateral margin and two apertural papillae
on anterior margin of aperture may interlock with two similar papillae on posterior
margin making a total of 16 peripheral papillae. Intestinal bifurcation dorsal to ventral
sucker. Caeca united posteriorly opening ventro-subterminally via anus. Testes two,
elongate, median, tandem, equal in size, intercaecal, separated by vitelline follicles, in
posterior half of body. Seminal vesicle tubular, winding, extending slightly posterior
to ventral sucker stalk. Cirrus-sac thin-walled, encloses indistinct pars prostatica and
cirrus. Genital pore at level of mid-pharynx. Ovary tri-lobed, lobes spherical, inter-
caecal, equatorial. Uterine seminal receptacle present. Uterus runs between ovary
and genital pore. Eggs oval, 40–49 long by 22–29 wide. Vitellarium follicular, in lat-
eral fields, extends from slightly posterior to ventral sucker to posterior end of body,
interrupted at level of gonads, fill most of post-testicular space. Excretory vesicle
I-shaped, reaching to ovary; pore terminal.
Remarks: This species differs from all the other species of the genus in having 16
papillae on the ventral sucker, smooth testes, a lobed ovary and vitelline fields that
extend to the posterior level of the peduncle.
Paropecoelus sciaeni Ahmad, 1983 (Fig. 15.18)

Host: Sciaenidae: Johnius dussumieri (Cuvier)
Locality: PR, BOB
No. 15
Description: Body elongate, 2240–3115 long by 312–338 wide. Oral sucker 88–110
in diameter. Ventral sucker pedunculate, retracted in most of the specimens, l70–192
long by 220–247 wide, with four biramous peripheral papillae one on each antero-
lateral and each postero-lateral margins and four apertural papillae two on anterior
lip of aperture of ventral sucker interlocking with two similar papillae on posterior
lip of aperture. Total number of papillae 16. Intestinal bifurcation dorsal, between
pharynx and ventral sucker. Caeca united posteriorly opening ventro-subterminally
via anus. Testes two, spherical, equal in size, median, tandem, intercaecal, postequa-
torial, separated from one another, 143–173 in diameter. Anterior testis lying at the
junction of middle and posterior third of body. Posterior testis lying in anterior part of
posterior third of body. Seminal vesicle an ovoid sac, intercaecal, dextral, mostly in
hindbody, leading to sinuous duct extending forward and left. Cirrus-sac thin-walled,
small enclosing indistinct pars prostatica and cirrus. Genital pore sinistral, at ante-
rior level of pharynx. Ovary four-lobed, pretesticular, median, tandem, intercaecal,
postequatorial, separated from anterior testis, lies in posterior part of middle third
of body. Vitellarium follicular, lateral, interrupted at level of gonads, extends from
anterior l/3rd distance between ventral sucker and ovary to posterior end of body,
filling most of post-testicular space. Uterus runs between ovary and genital pore.
Eggs oval, 52–56 by 29–42. Excretory vesicle I-shaped, reaches to ovary.

sciaeni (after Ahmad, 1983)
Remarks: This species closely resembles P. indicus Madhavi, 1975, but differs from
it in having a genital pore at the anterior level of the pharynx, a saccular seminal
vesicle, entire and spherical testes, vitelline fields interrupted at the level of the
gonads and extending from the anterior 1/3rd of the distance between the ventral
sucker and the ovary and a smaller post-testicular space. Unlike other Indian species
of Paropecoelus, this species possesses 16 marginal papillae on the ventral sucker.
Key to Indian species of Paropecoelus Pritchard, 1966
1. Ventral sucker with 16 marginal papillae …………………….… P. pritchardae

Ahmad, 1963
Ventral sucker with eight marginal papillae …………………………...............2
2. Testes lobed ………………………………………………………….……..….3
Testes oval …………………………………………….………………………..4
3. Vitellarium extends to level of ovary ………………………..……. P. dollfusi
Ahmad, 1983
Vitellarium extends to midway between ovary and ventral sucker …….. P. indicus
Madhavi, 1975
4. Ovary four-lobed, vitellarium interrupted at level of gonads ………… P. sciaeni
Ahmad, 1983
Ovary spherical, vitellarium not interrupted opposite gonads ….... P. overstreeti
Ahmad, 1983
Genus Pseudopecoeloides Yamaguti, 1940

The genus was erected by Yamaguti (1940) for P. tenuis Yamaguti, 1940, from the
priacanthid Pseudopriacanthus nipponicus (Cuv. & Val.). Important characters of
the genus include elongate body, with a pedunculate non-papillate ventral sucker
and a uroproct.
Generic diagnosis: Body elongate. Ventral sucker pedunculate without papillae.

Caeca open into excretory bladder forming uroproct. Testes tandem, contiguous or
separate, entire or lobed, well separated from posterior end of body. Cirrus-sac absent
or small. Seminal vesicle naked, extends well into hindbody. Genital pore sinistral,
close to pharynx. Ovary entire or irregularly lobed. Uterus restricted to area between
ovary and genital pore. Vitelline follicles restricted to hindbody. Excretory bladder
extends to ovary. Type species: P. tenuis Yamaguti, 1940.
The genus is fairly large comprising over 25 species. Bartoli et al. (2003) devised
a key to the then known 20 species. Aken’Ova et al., (2009) added seven species
from fishes of Australia and reviewed the genus. The genus is closely related to
Pseudopecoelus Von Wicklen, 1946 and the only character separating the two is the
lack of a uroproct in Pseudopecoelus. Four species belonging to this genus have
recorded from India.
Pseudopecoeloides orientalis Gupta & Ahmad, 1978

Locality: PR, BOB
No. 1
Reference: Gupta, V. & Ahmad (1978c)
Description: Body elongate, slender, 3328 long, 320 wide. Oral sucker 220 long,
240 wide. Ventral sucker pedunculate, smaller than oral sucker. Pharynx ovoid.
Oesophagus long. Caeca extend to posterior extremity, form uroproct. Genital pore
median, immediately posterior to pharynx. Testes spherical, tandem, in middle third
of body, separated from each other by vitelline follicles. Seminal vesicle tubular,
winding, extends to just posterior to base of ventral sucker peduncle. Cirrus-sac
short, thin-walled, encloses indistinct pars prostatica and cirrus. Ovary spherical,
pretesticular, pre-equatorial, separated from anterior testis by vitelline follicles. Uter-
ine coils mainly between ovary and seminal vesicle. Eggs oval, operculated, 32–46
long, 15–27 wide. Vitellarium follicular, interrupted at level of gonads, confluent
anterior to ovary, between ovary and testes, extending from some distance posterior
to seminal vesicle to posterior end of body
Remarks: The validity of the species needs to be confirmed by examining more

specimens collected from the type host and type locality
Pseudopecoeloides puriensis Ahmad, 1978
Locality: PR, BOB
Reference: (Ahmad, 1978b)
Remarks: This species was differentiated from the closely related species of the
genus by the position of the genital pore, the shape of the testes, the nature and
extent of the vitelline follicles and the relative sizes of the suckers
Pseudopecoeloides scomberi Hafeezullah, 1971 (Fig. 15.19)

Syn: Pseudopecoelus scomberi (Hafeezullah, 1971) Madhavi, 1975
Host: Carangidae: Scomberoides tol (1, 2); Bothidae: Engyprosopon grandisquamis
(Temn. & Schl.) (3)
Locality: VSK, BOB (1,2); Kerala coast, AS (3)
Distribution: BOB, Queensland, Malaysia
Reference: 1. Hafeezullah (1971c); 2. Madhavi (1975a); 3. Bijukumar (1997)
Description: Body 6828–7056 long, 504–583 wide, elongate, forebody short, hind-
body long. Oral sucker 212–221 long 206–221 wide. Ventral sucker 218–236 long, on
short peduncle, without papillae. Sucker length ratio 1:1. Prepharynx short. Pharynx
120–135 × 123–143, globular. Oesophagus indistinct. Intestinal bifurcation slightly
anterior to peduncle of ventral sucker. Caeca form uroproct near posterior end of
body. Testes oval, entire, tandem, separated from each other, in middle third of
hindbody. Cirrus-sac inconspicuous. Seminal vesicle tubular, long, winding. Pars
prostatica indistinct. Cirrus long protrusible. Genital pore sinistral to median line, at
Fig. 15.19 Pseudopecoeloides

scomberi (after Hafeezullah,
1971 Courtesy Cambridge
Univ. Press, Parasitology, 62,
p. 325)
posterior level of pharynx. Ovary entire, median, anterior to anterior testis. Seminal
receptacle absent. Vitellarium follicular, continuous, reaching slightly posterior to
ventral sucker peduncle, confluent posterior to testis. Uterine coils preovarian. Eggs
54–72 × 30–42. Excretory vesicle tubular, extends to ovary.
Remarks: P scomberi was first described by Hafeezullah (1971c) and was included
in the genus Pseudopecoeloides based on the presence of a uroproct, the only
feature that separates this genus from Pseudopecoelus. Madhavi (1975a) examined
apparently individuals of the same species collected from the same host and locality
but could not find evidence for a uroproct either in the whole mounts or in the
sections, hence transferred the species into the genus Pseudopecoelus as P. scomberi
(Hafeezullah, 1971c). Bray (1987), who examined specimens collected from
Malaysia, could not find a uroproct, and referred to the species as Pseudopecoelus
scomberi. Bijukumar (1997) also included the species in the genus Pseudopecoelus.
Later Aken’Ova et al. (2009) based on observations made on specimens collected
from S. lysan from Queensland, ‘concluded that the present species has a uroproct,
based on observations on the single laterally mounted specimen’.
Pseudopecoeloides tenuoides Martin, 1960 (Fig. 15.20)
Hosts: Scombridae: Rastrelliger kanagurta (Cuvier)
Locality: Digha coast, BOB
Reference: Kumari and Srivastava (1976)
Genus Pseudopecoelus Von Wicklen, 1946
[Syn: Neopecoelus Manter, 1947]
Genus Pseudopecoelus, erected by von Wicklen (1986) for Cymbephallus vulgaris
Manter, 1934, contains several species, possessing typical opecoelid features such
as an elongate body, a pedunculate ventral sucker and blind caeca. The only feature
Fig. 15.20 Pseudopecoeloides

tenuoides (after Martin,
1960)
differentiating this genus from Pseudopecoeloides is the blind termination of

the caeca instead of opening into excretory bladder forming the uroproct. Other
important features of the genus include the ventral sucker lacking papillae, the small
or absent cirrus-sac, the sinuous seminal vesicle and the sinistral genital pore.
Generic diagnosis: Body elongate-oval. Oral sucker simple. Ventral sucker sessile
or protuberant or pedunculate, without any papillae or lobes. Caeca terminate blindly
close to posterior end of body. Testes tandem, entire or deeply lobed, well separated
from posterior end of body. Cirrus-sac small or absent. Seminal vesicle winding usu-
ally extends into hindbody. Genital pore sinistral, usually close to pharynx. Ovary
entire or lobed. Uterus occupies area between ovary and genital pore. Vitelline fol-
licles confined to hindbody. Excretory vesicle extends to ovary.
Type species: P. vulgaris Von Wicklen, 1946.
Five species belonging to this genus have been reported from India.
Pseudopecoelus bilqeesae Ahmad & Dhar 1987
Host: Carangidae: Carangoides malabaricus (Bloch.)
Locality: PR, BOB
Number: 16
Reference: Ahmad & Dhar (1987b)
Description: Body elongate, 2160–3200 long and 240–370 wide. Oral sucker
120–150 in diameter. Ventral sucker pedunculate, larger than oral sucker, near ante-
rior extremity, 140–180 long and 180–230 wide. Sucker ratio 1:1.5–1.53. Phar-
ynx oval. Oesophagus very short. Intestinal bifurcation dorsal, immediately ante-
rior to ventral sucker. Caeca blind, terminate at posterior end of body. Testes two,
transversely elongate, slightly indented, separated from one another by distance of
200–240, directly tandem, subequal in size, equatorial. Cirrus-sac absent. Semi-
nal vesicle tubular, winding, extends posteriorly to midway between ventral sucker
and ovary. Ovary triangular, median, intercaecal. Uterine seminal receptacle present.
Uterus between ovary and ventral sucker. Eggs 44–53 long and 23–29 wide. Vitelline
follicles extend from posterior margin of pharynx to posterior end of body, interrupted
at level of ventral sucker on both sides, confluent in post-testicular area.
Remarks: P bilqeesae differs from other species of the genus by possessing the
following combination of characters. The vitellarium is interrupted at the level of the
gonads, the widely separated gonads, the transversely oval testes, indented laterally,
the triangular ovary and the vitellarium extending anteriorly as far as the level of the
pharynx.
Pseudopecoelus brayi Madhavi & Trivenilakshmi, 2010 (Fig. 15.21)

Host: Holocentridae: Myripristis kuntee Valenciennes
Locality: VSK, BOB
Reference: Madhavi & Trivenilakshmi (2010)
Description:
Description: Body elongate with parallel sides, rounded posteriorly, 2816–4576

long, 848–1136 wide. Oral sucker rounded subterminal, 152–232 × 160–240 in size.
Oesophagus long, distinct, twice the length of pharynx. Caeca long, narrow, termi-
nate blindly at posterior end of body. Intestinal bifurcation midway between phar-
ynx and ventral sucker. Ventral sucker larger than oral sucker, 232–304 × 232–296,
sucker width ratio 1:1.2–1.5. Testes two, deeply lobed, each with 4–5 lobes, located
Fig. 15.21 Pseudopecoelus

brayi (after Madhavi &
Trivenilakshmi, 2010)
in tandem in posterior half of body; separated by vitelline follicles. Genital pore

submedian, to left of oesophagus. Genital atrium surrounded by concentric ring of
circular muscles creating a sucker-like appearance. Seminal vesicle tubular, naked,
long, sinuous to convoluted at anterior end, elongate to saccular at posterior end,
Cirrus-sac short, cylindrical, 320–352 × 16, with thick fibrous wall, surrounded by
dense zone of prostatic cells. Cirrus long, muscular. Ovary lobed, usually with four
lobes, pretesticular. Uterine seminal receptacle present. Uterine coils preovarian, in
intercaecal space between ovary and ventral sucker. Eggs 52–56 × 28–30. Vitellar-
ium follicular, follicles extend from intestinal bifurcation to posterior end, overlap
caeca dorsally, lateral fields mainly lateral to caeca, confluent posterior to testis and
between gonads.
Remarks: This species most closely resembles P. sewelli Bray, 1987 reported from
Hoplostethus atlanticus off southeastern Australian waters, in the general morpho-
logical features especially in having elongate body and deeply lobed gonads but
differs from this species in the following characters: (i) in the non-pedunculated ven-
tral sucker, (ii) in the seminal vesicle not extending posterior to the ventral sucker,
(iii) in the presence of a prominent club-shaped, thick-walled cirrus-sac enclosing
the cirrus and surrounded by thick zone of prostatic gland cells, (iv) in the densely
distributed vitelline follicles and (v) much smaller eggs (52–56 × 28–30 vs. 66–93 ×
29–60).
Pseudopecoelus dollfusi Ahmad & Dhar, 1987 (Fig. 15.22)
Host: Terapontidae: Terapon theraps (Cuvier)
Locality: PR, BOB
No. 17
Reference: Ahmad & Dhar (1987b)

dollfusi (after Ahmad &
Dhar, 1987)
Description: Body elongate, narrow, 1760–2360 long and 260–360 wide. Tegu-
ment thin, unspined. Oral sucker 120–160 long and 110–140 wide. Ventral sucker
pedunculate, of similar size as oral sucker, situated near anterior extremity, 115–130
long and 120–140 wide. Sucker ratio 1:1.0–1.09. Forebody 290–350 long. Hindbody
1380–1930 long. Intestinal bifurcation dorsal in posterior forebody. Caeca blind, ter-
minate at posterior end of the body. Testes two, transversely elongate, constricted
medially, indented laterally, directly tandem, separated from one another by a dis-
tance of 200–270, situated in the posterior half of body, equal in size.. Seminal
vesicle saccular, extends only to posterior margin of ventral sucker. Prostatic vesi-
cle surrounded by prostate gland cells. Ejaculatory duct long. Genital pore ventral,
submedian at mid-level of pharynx. Ovary oval, pretesticular, situated in posterior
part of middle third of body. Uterine seminal receptacle present. Uterus between
ovary and ventral sucker. Eggs 46–50 long and 23–25 wide. Vitellarium follicular,
extending from posterior level of pharynx to posterior end of body, interrupted at
level of gonads, confluent in post-testicular space. Excretory vesicle tubular, reaches
posterior margin of ovary.
Remarks: P. dollfusi differs from other species of the genus by possessing the fol-
lowing combination of characters. The vitellarium is interrupted at the level of the
gonads, the transversely oval testes, constricted medianly, indented laterally, the
vitellarium extending anteriorly as far as the level of the pharynx and the seminal
vesicle extends posteriorly to midway between the ventral sucker and the ovary.
Pseudopecoelus odeningi Ahmad, 1987 (Fig. 15.23)
Host: Sciaenidae: Johnius belengeri (Cuvier)
Locality: PR, BOB
No. 17

odeningi (after Ahmad,
1987)
Description Body elongate, narrow, 1730–1945 × 24–255 in size. Oral sucker

120–128 × 110–112 in size. Ventral sucker pedunculate, larger than oral sucker,
140–152 × 190–208 in size. Sucker ratio 1:1.72–1.80. Pharynx oval. Oesophagus
very short. Intestinal bifurcation dorsal, immediately anterior to ventral sucker. Caeca
terminate blindly at posterior end of body. Testes two, entire, longitudinally elon-
gate, tandem, close together, equal in size, in middle third of body. Cirrus-sac absent.
Seminal vesicle saccular at base, then narrows anteriorly towards right side, extends
posteriorly only to posterior margin of ventral sucker. Pars prostatica surrounded by
prostate gland cells. Cirrus short, muscular. Genital pore submedian, ventral, to left
of median line, at mid-level of pharynx. Ovary four-lobed, immediately pretesticu-
lar, median, intercaecal, pre-equatorial. Uterine seminal receptacle present. Uterus
preovarian, restricted to area between ventral sucker and ovary. Vitelline follicles
extend from midway between ventral sucker and ovary to posterior end of body,
confluent in post-testicular field. Eggs 38–50 × 22–25. Excretory vesicle tubular,
reaches anteriorly as far as to posterior margin of ovary.
Remarks: This species differs from other Indian species of the genus by possessing
the following combination of characters: Vitellarium not extending anterior to ven-
tral sucker, seminal vesicle club-shaped extends to near posterior margin of ventral
sucker.
Pseudopecoelus pritchardae Gupta & Sayal, 1979 (Fig. 15.24)
Host: Percidae: Holocentrus sp
Locality: Kavaratti (Laccadive Islands), India.
No. 3
Reference: Gupta, N.K. & Sayal (1979)

pritchardae (after Gupta &
Sayal, 1979)
Description: Body elongate. 3080–3000 × 720–943, uniformly broad except the

anterior end which is narrower. Oral sucker terminal, 171 × 189–222. Ventral sucker
rounded, 231–217 240–217, situated at posterior part of anterior third of body. Sucker
ratio 1:1.35–1.5. Caeca terminate blindly at posterior end of body. Testes large, tan-
dem, unequal in size, irregularly lobed, post-equatorial. Seminal vesicle long, narrow,
extends posteriorly beyond ventral sucker. Cirrus-sac fusiform, contains pars prostat-
ica and ejaculator v duct. Genital pore submedian, at level of intestinal bifurcation.
Ovary four-lobed, immediately pretesticular, slightly dextral. Uterus winding in inter-
caecal field between ovary and ventral sucker. Metraterm runs along male terminal
duct. Vitellarium follicular, commence at level of intestinal bifurcation, extend all
along caecal length, confluent in post-testicular region. Eggs 34–60 × 25–40.
Remarks: This species is characterized by possessing the following characters: the

vitellarium extends to the level of the intestinal bifurcation, the seminal vesicle
extends posterior to the ventral sucker and the genital pore is bifurcal.
Pseudopecoelus stunkardi Ahmad, 1990 (Fig. 15.25)
Host: Lactaridae: Lactarius lactarius (Schnelder)
Description: Body elongate, with nearly parallel sides, extremities rounded,

1930–2470 long by 250–315 wide. Oral sucker subterminal, 120–135 long by
110–120 wide. Ventral sucker pedunculate, nearly same size as oral sucker, close
to anterior extremity, 110–130 in diameter, peduncle 180–365 long. Sucker ratio

1990)
1:1.0–1.08. Pharynx massive, nearly same size as oral sucker. Intestinal bifurcation
dorsal, at level of ventral sucker. Caeca terminate at posterior end of body. Testes two,
bi-lobed, directly tandem, separated from one another by distance of 30–50, in poste-
rior half of body, equal in size. Seminal vesicle tubular, winding, extends posteriorly
to 1/3rd length between peduncle and ovary. Pars prostatica thick-walled, bulbous,
surrounded by prostate gland cells. Cirrus short. Genital pore ventral, submedian,
immediately posterior to oral sucker. Ovary four-lobed, median, intercaecal, pretes-
ticular, situated in posterior part of middle third of body. Uterine seminal receptacle
present. Uterus preovarian, running between ovary and ventral sucker. Eggs 45–50
long by 32–28 wide. Vitellarium consisting of numerous follicles, extend from base
of ventral sucker peduncle to posterior end of body, confluent in post-testicular space.
Excretory vesicle tubular, reaches posterior margin of ovary.
Key to species of Pseudopecoelus from Indian marine fishes
1. Vitellarium extend anterior to ventral sucker…………………………………...2
Vitellarium do not extend anterior to ventral sucker…………………………...5
2. Seminal vesicle club-shaped, extends to posterior margin of ventral sucker
………………………. P. dollfusi Ahmad & Dhar, 1987
Seminal vesicle sinuous, tubular, extends posterior to ventral sucker …………3
3. Vitellarium interrupted at level of ventral sucker, …………..P. bilqeesae Ahmad
& Dhar, 1987
Vitellarium not interrupted, …………………….......................…………..…. 4
4. Genital pore at level of oesophagus ………………… P. brayi Madhavi & Trive-
nilakshmi, 2010
Genital pore at level of bifurcation…………………….…… P. pritchardae
Gupta & Sayal, 1979
5. Seminal vesicle club-shaped, extends to near posterior margin of ventral
sucker..……………………………… P. odeningi Ahmad, 1987
Seminal vesicle sinuous, tubular, extends posterior to ventral sucker ……….....
P. stunkardi Ahmad, 1990
Subfamily Plagioporinae Manter, 1947
Diagnosis: Body oval to elongate. Oral sucker rounded or funnel-shaped. Ventral

sucker occasionally pedunculate, usually without lobes or projections. Caeca blind
or unite and form cyclocoel or unite and open through anus or form uroproct or open
through separate ani. Testes usually two, tandem or oblique or symmetrical, rarely
up to ten. Cirrus-sac well developed contains seminal vesicle and pars prostatica.
Genital pore median or submedian near level of bifurcation. Ovary pretesticular,
entire or lobed. Canalicular seminal receptacle present. Uterus usually restricted to
area between ovary and genital pore, rarely extends posterior to testis. Eggs oval.
Vitelline follicles extensive, may or may not extend anterior to ventral sucker. Excre-
tory vesicle tubular, anterior extent variable.
Type genus: Plagioporus Stafford, 1904

Key to genera
The species of plagioporines recorded from Indian marine fishes are distributed over
nine genera that can be separated by the following key.
1. Testes ten………………………………. 2
Testes two…………..........……………..3
2. Ventral sucker simple………………………Decemtestis Yamaguti, 1934
Ventral sucker biacetabulate, arranged concentrically one inside the other
……………………….………Allodecemtestis Hafeezullah, 1970
3. Caeca unite posteriorly and open via common anus………….Vesicocoelium
Tang, Hsu, Huang, & Lu, 1975
Caeca terminate blindly…………........................…………4
4. Ventral sucker pedunculate………………………….Podocotyloides Yamaguti,
1934
Ventral sucker not pedunculate…………………………….5
5. Vitelline follicles extend into forebody……………………… 6
Vitelline follicles restricted to hindbody……………………………7
6. Ovary deeply lobed…………………………..……..Podocotyle Dujardin, 1845
Ovary entire…………………………….…Allopodocotyle Pritchard, 1966
7. Ovary deeply lobed………………………….Hamacreadium Linton, 1910
Ovary entire…………………………….8
8. Testes symmetrical, uterus extends posterior to testis…….……Phyllotrema Yam-
aguti, 1934
Testes tandem, uterus preovarian …………………………Macvicaria Gibson &
Bray, 1982
Genus Allodecemtestis Hafeezullah, 1970

The genus Allodecemtestis was proposed by Hafeezullah (1970a) to include species
of Decemtestis Yamaguti, 1934 provided with two concentric ventral suckers, one
lying inside another. Allodecemtestis biacetabulatus (Srivastava, 1936) from Sillago
sihama was designated as the type species. Except for the concentric nature of ventral
sucker, Allodecemtestis and Decemtestis appear similar. Cribb (2005a) regarded the
two genera as congeneric. However, following the criteria suggested by Hafeezullah
(1970a), the two genera are here treated as distinct.
Generic diagnosis: Body elongate. Ventral suckers two, one concentrically within
another. Caeca simple, long. Testes ten, in two longitudinal rows in posterior half
of body. Cirrus-sac claviform, overlaps ventral sucker, encloses coiled seminal vesi-
cle, tubular pars prostatica, prostatic cells, ejaculatory duct and cirrus. Genital pore
situated halfway between pharynx and oesophagus. Ovary lobed, pretesticular. Sem-
inal receptacle large. Vitelline follicles extensive. Uterus between ovary and ventral
sucker. Excretory vesicle I-shaped.
Type species: Allodecemtestis biacetabulatus (Srivastava, 1936)

Subfamily Plagioporinae Manter, 1947 443
Apart from the type species, five species from India have been assigned to this genus
namely A. carangai (Gupta & Puri, 1982) Ahmad, 1988 (syn. D. carangai Gupta
& Puri, 1982); A. drepanei (Gupta & Puri, 1982) Ahmad, 1988 (Syn. D. drepanei
Gupta & Puri, 1982); A. odeningi Ahmad, 1988; A. skrjabini Ahmad, 1988 and A.
bilqeesae Ahmad, 1990. Ahmad (1988) considered A. carangai as a synonym of A.
drepanei.
Allodecemtestis biacetabulatus (Srivastava, 1936) Hafeezullah, 1970 (Fig. 15.26)
[Syn Decemtestis biacetabulatus Srivastava, 1936]
Host: Scombridae: Rastrelliger kanagurta (Cuvier) (1); Sillaginidae: Sillago sihama
(Forsskal) (2); Sillago lutea (McKay); S. chondropus (Bleeker) (3)
Locality: VSK, PR, BOB
Reference: 1. Srivastava (1936c); 2. Hafeezullah (1970a); Sujatha & Madhavi (1997)
Description: Body elongate, subcylindrical, extremities broadly rounded. Oral

sucker subterminal, smaller than ventral sucker. Ventral suckers two, one concen-
trically enclosed with in other, almost equatorial. Prepharynx and pharynx present.
Caeca simple, wide, almost reaching posterior end. Testes ten, spherical or trans-
versely elongated, usually in two longitudinal rows in posterior half of body or
amassed together irregularly. Cirrus-sac elongated, claviform, somewhat overlapping
ventral sucker, encloses an elongated, coiled seminal vesicle, tubular pars prostatica
surrounded by prostatic gland cells, ejaculatory duct and cirrus. Genital pore situ-
ated halfway between pharynx and oesophagus. Ovary lobed, pretesticular, median
or slightly submedian. Seminal receptacle large, dorsal to ovary. Vitellarium follic-
ular, usually reaching from intestinal bifurcation to posterior end of body, confluent
or not in acetabular zone. Excretory vesicle I-shaped.
Remarks: Sujatha & Madhavi (1997) in a comparative study of the digenean fauna
of sillaginid fishes from inshore and offshore waters off Visakhapatnam (Bay of
Bengal) found the infection to be confined to inshore sillaginids.
Fig. 15.26 Allodecemtestis

biacetabulatus (after
Srivastava, 1937)
Allodecemtestis bilqeesae Ahmad, 1990 (Fig. 15.27)

Host: Cynoglossidae: Cynoglossus bilineatus (Bloch)
Description: Body elongate, 1235–1685 long by 475–557 wide; widest at level

of testicular zone. Oral sucker spherical, 95–110 in diameter. Ventral suckers two,
median, one concentrically enclosed within other. Outer sucker oval, wider than long,
immediately post-bifurcal, 215–240 long by 270–290 wide, pre-equatorial. Inner
sucker oval, 100–115 long by 130–155 wide. Sucker ratio 1:2.63–2.84. Prepharynx
absent. Pharynx oval. Oesophagus long. Intestinal bifurcation dorsal, nearer to ventral
sucker than to oral sucker. Caeca terminate blindly at posterior end of body. Testes
ten in number, oval, arranged more or less in two longitudinal rows, confined to
posterior half of body. Cirrus-sac club-shaped, extends posteriorly as far as anterior
margin of ventral sucker. Seminal vesicle coiled with one backward coil. Genital
pore submedian, to right of median line, at anterior level of pharynx. Ovary three-
lobed. Seminal receptacle saccular, lateral to ovary. Uterus extends posteriorly and
intrudes into intertesticular space as far as penultimate testis. Eggs. 70–80 long by
41–42 wide. Vitelline follicles numerous, extend from level of intestinal bifurcation
to posterior end of body, non-confluent in forebody, confluent in post-testicular zone.
Excretory vesicle tubular, reaches anteriorly to ovary.
Remarks: This species was differentiated from all the known species of the genus
Allodecemtestis in that the uterus extends posteriorly intruding into the intertesticular
space as far as the penultimate testis rather than being confined to the space between
the anterior testis and the ventral sucker.

bilqeesae (Ahmad, 1990)
Allodecemtestis carangai Gupta & Puri, 1982

Location: PR, BOB
Number: One
Syn.: (1988)
Allodecemtestis drepanei Gupta & Puri, 1982
[Syn: A. carangi Gupta & Puri 1982]
Host: Drepanidae: Drepane punctata (Cuv. &Val.)
Locality: PR, BOB
No. One
Remark: The description is based on a single specimen and is inadequate, lacking in
many details.
Allodecemtestis odeningi Ahmad, 1988 (Fig. 15.28)
Locality: PR, BOB
Description Body elongate, 1845–2535 long by 392–455 wide. Oral sucker oval,
110–130 long 125–145 wide. Ventral sucker double, one concentrically enclosed
within other. Outer sucker subspherical, wider than long, situated in posterior part of
anterior third of body, 225–275 long by 305–335 wide. Inner sucker oval, 105–135

odeningi (after Ahmad,
1988)
long by 135-165 wide. Sucker length ratio 1:2.31–2.44. Prepharynx very short. Phar-
ynx oval, muscular. Oesophagus long. Intestinal bifurcation dorsal, nearer to ventral
sucker than to oral sucker. Caeca terminate blindly at posterior end of body. Testes
10 in number, oval, arranged more or less in two longitudinal rows, confined to pos-
terior half of body. Cirrus-sac club-shaped, entirely in forebody, extends posteriorly
as far as anterior margin of ventral sucker. Seminal vesicle coiled with one back-
wardly directed loop. Pars prostatica surrounded by prostate gland cells. Genital pore
marginal, to left of median line, at mid-level of pharynx. Ovary spherical, subme-
dian, postequatorial. Seminal receptacle saccular, lateral to ovary. Uterus between
ovary and ventral sucker. Eggs 61–68 × 31–38. Vitelline follicles numerous, extend
from posterior level of pharynx to posterior end of body, non-confluent in forebody
confluent in post-testicular zone, interrupted at level of ventral sucker on both sides.
Excretory vesicle tubular, reaches anteriorly up to ovary.
Remarks: This species differs from all the other species of Allodecemtestis, in that
its ovary is unlobed rather than lobed and its vitellarium is interrupted at the level
of the ventral sucker, rather than continuous. It differs from D. biacetabulatus in the
smaller sucker ratio, from D. drepanei in the larger sucker ratio, the more anterior
location of the ventral sucker, in the extent of vitellarium to the level of the pharynx
and position of the genital pore at the mid-level of the pharynx.
Allodecemtestis skrjabini Ahmad, 1988 (Fig. 15.29)

Host: Cynoglossidae: Cynoglossus bilineatus (Bloch)
Locality: PR, BOB
Number of specimen: 19

skrjabini (after Ahmad,
1988)
Description: Body elongate, 1672–1922 long, by 775–892 wide oral sucker

151–185 in diameter. Ventral sucker two, one concentrically enclosed within other.
Outer sucker 272–305 in diameter, situated in anterior part of middle third of body,
pre-equatorial. Inner sucker spherical, 110–135 in diameter. Sucker ratio 1:1.64–1.80.
Prepharynx absent. Pharynx oval. Oesophagus long. Intestinal bifurcation dorsal,
about midway between suckers. Caeca terminate at posterior end of body. Testes
ten in number, oval, confined to posterior half of body, arranged in two more or
less longitudinal rows. Cirrus-sac club-shaped, extends posteriorly as far as anterior
margin of ventral sucker. Seminal vesicle coiled with one backwardly directed loop.
Pars prostatica surrounded by prostate gland cells. Genital pore submedian, to right
of median line, at mid-level of oesophagus. Ovary three-lobed, submedian, to left
of median line. Seminal receptacle saccular. Vitelline follicles extend from anterior
level of ventral sucker to posterior end of body, confluent in post-testicular region.
Uterus between anterior testis and ventral sucker. Eggs 71–82 long by 35–41 wide.
Excretory vesicle tubular, reaches anteriorly as far as ovary. Excretory pore terminal.
Remarks: This species differs from all the other species of Allodecemtestis, in that the
vitellarium extends only to the anterior margin of the ventral sucker. It further differs
from A. biacetabulatus in the smaller sucker ratio, in the more posterior location of
the genital pore and in the elongated eggs; from A. drepanei in the pre-equatorial
location of ventral sucker, the larger sucker ratio, the three-lobed ovary and the larger
eggs.
Genus Allopodocotyle Pritchard, 1966
The genus was proposed by Pritchard (1966) for A. plectopomi (Manter, 1963)
Pritchard, 1966. It is characterized by a combination of characters such as testes
tandem or oblique and well separated from the posterior end of the body; the genital
pore is sinistral in the mid-forebody, the ovary is entire, and the vitelline follicles are
restricted to the hindbody.
Diagnosis: Body elongate. Oral sucker unspecialized. Ventral sucker larger than oral
sucker. Caeca long terminate blindly at posterior end of body. Testes two, tandem or
oblique, far away from posterior end of body. Genital pore sinistral in mid-forebody.
Cirrus-sac well developed. Ovary entire. Uterus between ovary and ventral sucker.
Vitelline follicles usually confined to hindbody. Excretory vesicle extends to level of
ovary. Type species: A. plectopomi (Manter, 1963) Pritchard, 1966.
Six species of Allopodocotyle have been reported from marine fishes of Indian
waters. However, Bray & Justine (2009) have transferred A. yamagutii Gupta &
Ahmad (1977) to the genus Macvicaria as M. yamagutii (Gupta & Ahmad, 1977)
Bray & Justine, 2009. Further, Bray (1987) indicated that this species may have
affinities with Podocotyloides. It is probably a synonym of Podocotyloides paru-
penei according to Bray & Cribb (1989). Also, A. conius was included as taxon
inquirendum (World Register of Marine Species, Editor, Gibson, D.I). As a result of
these adjustments, only three Indian species remained in the genus: A. pritchardae
Madhavi, 1975; A. argyropsi Madhavi, 1975 and A. lutianusi Gupta & Ahmad, 1976
Allopodocotyle argyropsi Madhavi, 1975 (Fig. 15.30)

Syn: A. lutjanusi Gupta and Ahmad, 1976
Host: Sparidae: Argyrops spinifer (Forsskal)
Locality: VSK, BOB
Description: Body elongate, 1296 to 1840 long and 416 to 480 wide at level of
ventral sucker. Oral sucker 94–117 in diameter. Ventral sucker 176–179 by 195–214.
Forebody 20–25% of body length. Sucker ratio 1:1.8–2.0. Prepharynx very short.
Pharynx 66–78 long by 78 wide. Oesophagus short. Intestinal bifurcation immedi-
ately anterior to ventral sucker. Caeca broad, terminate near posterior end of body.
Genital pore sinistral, at level of oesophagus. Testes rounded, 156–187 by 144–175,
tandem, contiguous, in middle of hindbody. Cirrus-sac slender, arcuate, extends well
posterior to ventral sucker. Seminal vesicle long, tubular, straight, occupies most of
space inside cirrus-sac. Prostatic vesicle prominent. Cirrus short and wide. Ovary
spherical, to pyriform, median, pretesticular, separated from anterior testis. Seminal
receptacle small, between ovary and anterior testis. Vitelline follicles extend from
posterior margin of ventral sucker, fill post-testicular space. Uterus preovarian, short.
Eggs large, yellow, 66–78 by 33–39. Excretory vesicle extends to level of ovary.
Remarks: A argyropsi resembles A. mecoperae (Manter, 1940) and A. pedicellata
(Stossich, 1887) in having tandem testes and a cirrus-sac extending posterior to the
ventral sucker, but differs from both the species in the body shape, in the longer and
more slender cirrus-sac, the more anterior genital pore and vitellarium extending to
the level of the ventral sucker.
Fig. 15.30 Allopodocotyle

argyropsi (after Madhavi
1975)
Allopodocotyle coniusi Singh, 2009

Host: Plotosidae: Plotosus canius (Hamilton)
Locality: Kanyakumari (BOB)
Remarks: Regarded as taxon inquirendum
Reference: Singh (2009)
Allopodocotyle lutianusi Gupta & Ahmad, 1976
Host: Lutjanidae: Lutjanus rivulatus (Russell)
Location: Intestine
Locality: PR, BOB
No. one
Reference: Gupta, V & Ahmad (1976c)
Remarks: This species appears very similar to A. argyropsi Madhavi, 1976, with
which it was not compared. The description based was on a single specimen and is
inadequate. The species is here considered a synonym of A. argyropsi.
Allopodocotyle pritchardae Madhavi, 1975 (Fig. 15.31)
Host: Lutjanidae: Lutjanus lunulatus (Park, 1797)
Locality: VSK, BOB
No. 25
Description: Body fusiform with indentation at level of ventral sucker, 2640–3280
long and 1152–1264 wide. Oral sucker subterminal, 214–242 by 234–242. Ven-
tral sucker large, broader than long, 400–480 by 480–576, with transverse aperture,
779–910 from anterior end. Sucker ratio 1:2.0–2.12. Oesophagus longer than phar-
ynx. Intestinal bifurcation midway between suckers. Caeca terminate blindly near
posterior end. Genital pore to left of median line, at level of posterior end of pharynx.
Testes rounded, close together in third quarter of body. Cirrus-sac clavate, with base
Fig. 15.31 Allopodocotyle

pritchardae (after Madhavi
1975)
overlapping ventral sucker, contains oval seminal vesicle and cirrus. Ovary spheri-
cal, to right of anterior test. Seminal receptacle oval, situated in space in between
gonads. Uterus intercaecal, between anterior testis, ovary and ventral sucker. Eggs
74–78 by 27–31. Vitelline follicles in two broad lateral zones from ventral sucker to
posterior end, confluent in post-testicular space. Excretory bladder I-shaped, extends
to ventral sucker.
Remarks: Allopodocotyle pritchardae resembles A. lethrini (Yamaguti, 1942) and

A. tamame (Yamaguti, 1942) but differs from the min the body shape, in the oval
seminal vesicle and in the position of the seminal receptacle posterior to the ovary.
Allopodocotyle upeneusi Gupta & Ahmad, 1976
Host: Mullidae: Upeneus macronemus (Cuv. & Val)
Location: Intestine
Locality: PR, BOB
Reference: Gupta, V & Ahmad (1976c)
Remarks: It is considered as species inquirendum but regarded as a probable syn-

onym of Podocotyloides parupenei (Manter, 1963) Pritchard, 1966 (Reference: Bray,
1987; Bray & Cribb, 1989)
Genus Decemtestis Yamaguti, 1934
This genus was proposed by Yamaguti (1934) with D. sillagonis Yamaguti, 1934
as the type species. It is characterized by the presence, usually, of ten testes arranged
either in two rows or clumped together in the hindbody, a submedian genital pore,
usually anterior to the intestinal bifurcation and extensive vitelline fields.
Generic diagnosis: Body elliptical. Oral and ventral suckers simple, latter larger
than oral sucker. Caeca extend to posterior end of body, terminate blindly. Testes
nine or ten, in two irregular columns in hindbody, well separated from posterior
end of body. Genital pore sinistral or dextral. Ovary entire to deeply lobed. Uterus
between testes and ventral sucker. Eggs without filaments. Vitelline follicles enter
forebody, extend to posterior end of body. Excretory vesicle extends to ovary.
Type species: D. sillagonis Yamaguti, 1934.
Many species belonging to the genus have been reported from Indian marine
fishes that are taxonomically quite unrelated. Some of the species of the genus were
relegated to the genus Allodecemtestis Hafeezullah, 1970. The first species to be
reported from India were D. mehrai Srivastava, 1936 and D. brevicirrus Srivastava,
1936. D. mehrai has also been reported from several fish hosts from different localities
(Hafeezullah, 1970a, Gupta & Mehrotra, 1970; Gupta & Jahan, 1975; Sujatha &
Madhavi, 1997; Bijukumar, 1997). Hafeezullah (1970a) also reported D. brevicirrus
and D. kobayashi Park, 1939. Karyakarte & Yadav (1977) reported D. kobayashi and
added two new species D. marginoacetabulatus Karyakarte & Yadav, 1977 and D.
cynoglossi Karyakarte & Yadav, 1977. They did not accept the genus Allodecemtestis
and transferred its two species back into the genus Decemtestis. Ahmad (1983c)
added two species D. dollfusi Ahmad, 1983 and D. singhi Ahmad, 1983. Other
species reported include D. srivastavai Gupta & Jahan, 1975; D. varmai Gupta &
Gupta, 1988 and D. mystusi Dhanumkumari, 1999. Thus, ten species are known in
the genus. So far, no attempt has been made to review the genus and examine the
validity of various species reported under the genus.
Decemtestis brevicirrus Srivastava, 1936 (Fig. 15.32)
Host; 1. Sillaginidae: Sillago sihama(Forsskal); 2. Therapontidae: Terapon jarbua
(Forsskal) Locality: PR, BOB (1); MS (BOB) (2)
Reference: 1. Srivastava (1936c); 2. Hafeezullah (1970a)
Description: Body elongated, oval, long, 520–680 wide. Oral sucker 150–180 in
diameter. Ventral sucker 200–230 by 280, situated in anterior half of body. Sucker
ratio 3:4. Prepharynx, pharynx short. Caeca broad, extend to level of posterior testis.
Testes ten in number, oval, arranged in two irregular rows on either side of median
line in posterior half of body. Cirrus-sac club-shaped, extends obliquely from the
genital atrium to slightly less than half distance between intestinal bifurcation and
ventral sucker, encloses slightly coiled, elongated, bulb-shaped seminal vesicle, small
tubular pars prostatica surrounded by sparsely developed prostate gland cells, short
ejaculatory duct and knob-shaped cirrus. Ovary four-lobed, pretesticular. Seminal
receptacle oval, adjacent to ovary. Vitelline follicles extend from pharynx to posterior
end of body, interrupted at level of ventral sucker. Uterine coils distributed between
ventral sucker and ovary. Eggs 50 × 37.
Remarks: This species is characterized by the short cirrus-sac restricted to the fore-
body, and the vitellarium is interrupted opposite the ventral sucker.
Fig. 15.32 Decemtestis

brevicirrus (after
Srivastava,1936)
Decemtestes cynoglossi Karyakarte & Yadav, 1977

Host: Cynoglossidae: Cynoglossus oligolepis (Gunther)
Locality: RT, AS
No: 4
Reference: Karyakarte & Yadav (1977a)
Description: Body elongate, 1620–1820 in length, 520–640 in width. Oral sucker

terminal, 120–160 by 180–200. Ventral sucker larger than oral sucker, 240–260 by
210–230, situated on left side in anterior half of body. Sucker ratio 1:1.14–1.50.
Prepharynx absent. Pharynx globular. Oesophagus short. Caeca long extend to pos-
terior end of body. Testes ten, arranged in two longitudinal rows, six on right, four
on left, in last third of body. Cirrus-sac elongated extends to anterior border of ven-
tral sucker, encloses seminal vesicle, pars prostatica surrounded by prostate gland
cells and short cirrus. Genital pore submedian and located posterior to intestinal
bifurcation. Ovary club-shaped, pretesticular on left side of body just anterior to left
longitudinal row of testes. Seminal receptacle on left side of ovary. Uterus preo-
varian, contains only few eggs. Eggs 72–74 by 26–28. Vitelline follicles extend in
lateral fields from level of intestinal bifurcation to posterior end of body, confluent
in forebody, not in post-testicular region. Excretory bladder Y-shaped.
Remarks: Closely resembles D. brevicirrus Srivastava, 1936 but differs from it in

the posterior extent of the cirrus-sac to anterior border of ventral sucker, in the
club-shaped rather than lobed ovary, in the extent of the vitellarium to the intestinal
bifurcation and in the post-bifurcal position of the genital pore.
Decemtestis dollfusi Ahmad, 1983 (Fig. 15.33)
Host: Scombridae: Thunnus albacares (Bonnaterre)
Locality: PR, BOB

dollfusi (after Ahmad, 1983)
Description: Body elongated, 1012–1126 long by 325–370 wide at level of ovary.

Oral sucker spherical, terminal, 110–127 in diameter; Ventral sucker spherical,
median, 130–150 in diameter, 110–135 post-bifurcal, immediately pre-equatorial,
385–426 from anterior end of body, larger than oral sucker. Sucker ratio 1:1.18.
Forebody 37–38% of body length. Prepharynx very short. Intestinal bifurcation dor-
sal, halfway between suckers. Caeca terminate blindly at posterior end of body. Testes
ten in number, spherical, irregularly arranged near hind end of body. Post-testicular
space 40 to 60 long. Cirrus-sac club-shaped, overlaps right margin of ventral sucker,
extends just posterior to ventral sucker containing inverted ‘N’-shaped seminal vesi-
cle occupying 51–52% of cirrus-sac length, prostate complex and ejaculatory duct.
Genital pore median, post-bifurcal. Ovary oval, submedian, immediately posterior to
ventral sucker. Seminal receptacle retort–shaped, postovarian. Uterus between testes
and ventral sucker. Metraterm thick-walled, opening into genital atrium. Eggs ellip-
tical, yellow, 54–60 long by 34–37 wide. Vitelline follicles in eight pairs of separate
lateral clusters lying dorsal to caeca, extending from just post-bifurcal to just anterior
to caecal ends. Excretory vesicle tubular, reaching to testes; pore terminal.
Remarks: This species differs from all the known species of the genus Decemtestis
in having the vitellarium divided into eight pairs of separate lateral clusters and
extending from a just posterior to the intestinal bifurcation to just anterior to the
caecal ends, the post-bifurcal genital pore and the ovary close to the ventral sucker
and well separated from the testes.
Decemtestis kobayashii Park, 1939 (Fig. 15.34)
Hosts: Cynoglossidae: Cynoglossus bilineatus (Bloch); C. oligolepis Lutjanidae:
Lutjanus johni Bloch; Tetraodontidae: Gastrophysus lunaris (Bloch)
Locality: Karwar, RT, AS
Reference: Hafeezullah (1970a); Karyakarte & Yadav (1977a)

kobayashii (after
Hafeezullah 1970)
Description: Body spindle-shaped, 1460–1540 long 480–540 wide. Oral sucker

terminal or subterminal, 100–120 by 140–180. Ventral sucker in anterior half of body,
230–250 by 200–220. Sucker ratio 1:1.40–1.56. Prepharynx absent. Oesophagus
short. Caeca reach to posterior end of body. Testes ten, situated in posterior half
of body, arranged in two rows, four on right and six on left. Cirrus-sac elongated,
reaches anterior border of ventral sucker, encloses seminal vesicle, well-developed
pars prostatica, surrounded by prostate gland cells and short cirrus. Genital pore
sinistral, bifurcal. Ovary weakly lobed, pretesticular, anterior to right row of testes.
Seminal receptacle anterior to ovary. Vitelline follicles extend from level of intestinal
bifurcation to posterior end of body, interrupted at level of ventral sucker, confluent
in post-testicular region. Uterus preovarian contains limited number of eggs. Eggs
54–60 by 34–38. Excretory bladder Y-shaped.
Remarks: D. kobayashii was described by Park (1939) from Areliscus joyneri and A.
purpurmaculatus, Korea. Hafeezullah (1970a) recorded the species for the first time
in India from Cynoglossus bilineatus, Lutjanus johnii and Gastrophysus lunarisat
Karwar, Madras and Bay of Bengal. Later Karyakarte & Yadav (1977a) collected the
species from Cynoglossus oligolepis at Ratnagiri, West Coast of India. This species
is characterized by the interruption of vitellarium at the level of the ventral sucker.
Decemtestis mehrai Srivastava, 1936 (Fig. 15.35)
Hosts: Sillaginidae: Sillago sihama (Forskål) (1, 3); S. lutea (McKay); S. indica
McKay, Dutt & Sujatha;S. chondropus (Bleeker) (4); Triacanthidae: Triacanthus
biaculeatus (Bloch) (2); Ariidae: Plicofollis dussumieri (Val.) (2); Cynoglossidae:
Cynoglossus lida (Bleeker) (5)
Location: Gut
Localities: Ernakulam, Kerala, AS, BOB
References: 1. Srivastava (1936c); 2. Gupta, N.K. & Mehrotra (1970); 3. Gupta, V.
& Jahan (1975); 4. Sujatha & Madhavi (1997); 5. Bijukumar (1997)
Distribution: Bay of Bengal and Arabian Sea

mehrai (after Srivastava
1936)
Description: Body elongated, 1590–2130 long, 620–670 wide. Oral sucker ter-
minal, 120–190 in diameter. Pharynx muscular. Oesophagus long. Intestinal caeca
simple, extend to posterior end of body. Ventral sucker larger than oral sucker, spher-
ical or oval, pre-equatorial, 220–250 long, 260–290 wide. Genital pore submedian,
at mid-level of oesophagus. Testes 10 in number, postequatorial, spherical or oval,
in two or three longitudinal rows. Cirrus-sac elongate, extends to anterior margin or
up to middle of ventral sucker. Seminal vesicle tubular, inverted ‘N’-shaped. Pars
prostatica tubular. Ejaculatory duct 30–40 long. Space around pars prostatica and
distal portion of seminal vesicle surrounded by large number of prostate gland cells.
Ovary entire, bi-lobed or tri-lobed, median or submedian, postequatorial, immedi-
ately pretesticular. Seminal receptacle elongated. Uterine coils in region between
testes and ventral sucker. Eggs oval, 42–49 × 42–56. Vitellarium follicular, extends
from intestinal bifurcation to posterior end of body.
Remarks: This species has been recorded from a wide range of hosts and localities.
Hafeezullah (1970a) studied individual variations in D. mehrai and concluded that
such characters as the extent of the vitellarium and the size and arrangement of testes
should not be relied upon to distinguish the species in this genus.
Decemtestis marginoacetabulatus Karyakarte & Yadav, 1977
Host: Carangidae: Trachinotus ovatus (Linn.)
No: 24
Reference: Karyakarte & Yadav (1977a)
Description: Body triangular, 1660–1780 long, 640–800 wide. Oral sucker 240–260
by 180–200 in size. Ventral sucker larger than oral sucker, 270–300 in diameter.
Prepharynx short. Pharynx short. Oesophagus short. Caeca long, terminate close
to posterior end of body. Testes ten, located in last third of body, arranged in two
longitudinal rows, located on right side of body. Cirrus-sac tubular. Genital pore
marginal on left side of body, situated at level of anterior part of pharynx. Ovary
rounded, pretesticular, situated just anterior to left longitudinal row of testes. Uterus
preovarian. Eggs 46–48 by 36–42. Vitelline follicles extend from pharyngeal level
and reach to posterior end of body.
Remarks: In possessing a cirrus-sac not extending posteriorly to the ventral sucker,

D. marginoacetabulatus comes close to D. brevicirrus. However, it differs from the
other species of the genus in several respects, such as the triangular body, the extent of
the cirrus-sac to the anterior margin of the ventral sucker, the extent of the vitellarium
and in the marginal position of the genital pore.
Decemtesis singhi Ahmad, 1983 (Fig. 15.36)
Host: Sciaenidae: Otolithus ruber (Bl. & Schn.)
Locality: PR, BOB
Ref: Ahmad (1983c)

singhi (after Ahmad, 1983)
Description: Body elongated with rounded extremities, 986–1125 long by 320–347

wide at level of ventral sucker. Oral sucker 143–167 in diameter. Ventral sucker spher-
ical, 110–127 in diameter, median, smaller than oral sucker, 135–167 post-bifurcal,
at 463–489 from anterior end of body, equatorial. Sucker ratio 1:0.761. Intestinal
bifurcation in mid-forebody. Caeca terminate at posterior end of body. Testes ten,
spherical, near posterior end of body. Post-testicular space very short. Cirrus-sac
club-shaped extends posteriorly to just post-bifurcal level, containing inverted N-
shaped seminal vesicle prostate complex and ejaculatory duct. Genital pore sinistral,
at posterior border of oral sucker. Ovary tri-lobed, pretesticular, submedian. Seminal
receptacle ventrolateral to ovary. Uterus lies between testes and intestinal bifurca-
tion. Eggs elliptical, 50–54 long by 30–37 wide. Vitellarium follicular, extends from
anterior border of ventral sucker to posterior end of body. Excretory vesicle tubular,
reaches to testes, pore terminal.
Remarks: This form differs from all the known species of the genus Decemtestis in
having the oral sucker larger than the ventral sucker, the vitellarium extends from
the anterior border of the ventral sucker to the posterior end of the body, the uterine
coils pass between the ventral sucker and the intestinal bifurcation and the equato-
rial ventral sucker. In the posterior extent of the cirrus-sac, this species resembles
D. brevicirrus, but the latter species differs from this species in having its genital
pore at mid-pharynx level, a four-lobed and immediately pretesticular ovary, testes
occupying a large space and the caeca terminating at posterior end of body.
Decemtestis srivastavai Gupta & Jahan, 1975 (Fig. 15.37)
Host: Sisonidae: Mystus gulio Hamilton
Number: 6

srivastavai (after Gupta &
Jahan, 1975)
Locality; PR, BOB

Reference: Gupta & Jahan (1975)
Description: Body elongated 1660–1870 long, 440–550 wide. Oral sucker 110–120
long, 120–160 wide. Ventral sucker larger than oral sucker, pre-equatorial, 180–200
long, 200–210 wide. Pharynx muscular, subglobular, 60–70 × 90–110. Oesophagus
70–80 long. Caeca extend to posterior end of body. Genital pore median or sub-
median, at mid-level of oesophagus. Testes ten, postequatorial, spherical or oval, in
two longitudinal rows. Cirrus-sac elongate, extends from mid-level of oesophagus to
anterior margin or middle of ventral sucker. Seminal vesicle inverted N-shaped. Pars
prostatica tubular, followed by ejaculatory duct. Space around pars prostatica and
distal portion of vesicula seminal is surrounded by large number of prostate gland
cells. Ovary entire or tri-lobed, median, postequatorial, immediately pretesticular.
Seminal receptacle elongate. Uterus runs between testes and ventral sucker. Eggs
oval, 70–90 × 30–40. Vitellarium follicular, extends either from pharynx region or
from intestinal bifurcation to posterior end of body, interrupted lateral to ventral
sucker.
Remarks: This species differs from all the known Indian species of Decemtestis
excepting D. kobayashii in the interruption of vitellarium at the level of the ventral
sucker.
Decentestis varmai Gupta & Gupta, 1988
Host: Haemulidae: Pomadasys commersonni (Lacepede) (=Pristipoma operculare
(Playfair))
Locality: Bay of Bengal and Puri Coast, Orissa, India.

Prevalence: Single specimen from one host out of three examined.
Remarks: The description based on a single specimen agrees in all details with D.
mehrai. The differences mentioned have no specific significance.
Differentiation of the species of Decemtestis: of the various species, D. kobayashii
and D. srivastavai stand distinct in having vitelline follicles interrupted opposite the
ventral sucker. D. brevicirrus and D. mehrai can be differentiated by the criteria
reported by Hafeezullah (1970a). D. dollfusi is distinct in having its testes clumped
together close to the posterior end of the body, the ovary well separated from testis
and lying close to the ventral sucker, the median genital pore lying between the
ventral sucker and the intestinal bifurcation, the cirrus-sac extending posterior to the
ventral sucker and the host, a tuna. All these features in fact warrant the erection of
a new genus.
D. singhi Ahmad, 1983 differs in that the ventral sucker is smaller than the oral
sucker, the cirrus-sac restricted to far anterior to the ventral sucker and the genital
pore is sinistral to the posterior border of the oral sucker. D. marginoacetabulatus
Karyakarte & Yadav 1977 is distinctive in having a triangular body. D. kobayashi has
its vitellarium interrupted at ventral sucker level. D. cynoglossi Karyakarte & Yadav,
1977 is very similar to D. mehrai and is here synonymized with it. D. varmai Gupta
& Gupta, 1988, based on a single specimen, agrees in all details with D. mehrai. The
differences specified have no specific significance.
D. cynoglossi, D. srivastavai and D. varmai are very similar to D. mehrai and are
probably synonymous to that species. Hafeezullah found that some of the species,
such as D. mehrai and D. kobayashii, show extensive intraspecific variations in
respect of many characters, especially the anterior extent of the vitellarium, the
interruption of the vitellarium at the level of the ventral sucker, the size of testes
and their arrangement in two regular or irregular rows and amassed together near
posterior end of body. These characters alone cannot therefore be used for erection of
new species in the genus. Hence, care should be exercised not to erect new species on
the basis of one or two specimens. Keeping the above facts in view, it is necessary to
subject the genus to a thorough revision. Hence, no attempt is made here to develop
a key for separation of species in the genus.
Genus Hamacreadium Linton 1910
[Syn: Olivacreadium Bilqees, 1976]
The genus was proposed by Linton (1910) with H. mutabile Linton, 1910 as the type
species. Many species have been added to the genus from a wide range of hosts and
geographical locations. However, confusion prevails in the relationships of the genus
with other genera and the validity of various species reported in the genus. A review
of the genus was provided by Martin et al. (2017) wherein an analysis was made on
the validity of various species. The genus is considered to differ from Podocotyle
and Plagioporus in having oblique testes.
Generic diagnosis: Body elongate. Ventral sucker usually larger than oral sucker.
Caeca narrow, blind, end close to posterior end of body. Testes two usually oblique,
in posterior third of body. Genital pore submedian, in mid-forebody. Ovary deeply
lobed, pretesticular. Uterus confined to space between testis and ventral sucker. Eggs
without filaments. Vitellarium enter forebody and extend beyond testes to posterior
end of body. Excretory vesicle extends well into forebody. Type species: H. mutabile
Linton, 1910.
Six species of Hamacreadium have been recorded from marine fishes of India
of which two species H. leiperi Gupta, 1956 and H. indicum Gupta & Tewari,
1984 from Chirocentrus dorab of MS, BOB are considered incertae sedis (Mar-
tin et al. 2017). Hamacreadium krusadaiensis Gupta, 1956 was transferred to the
genus Neolebouria by Gibson (1976) and H. leiperi Gupta, 1956 to Neolebouria by
Martin et al. (2017). H. leiognathi is included in the genus Neonotoporus as N. leiog-
nathi (Hafeezullah 1971) Ahmad, 1985. H.chilkai (Chatterji, 1958) is considered
synonym of Allocreadium fasciatusi Kakaji, 1969 by Madhavi (1978) and Martin
et al. (2017). Maculifer spiralis Soota, Srivastava & Ghosh, 1970 was considered a
synonym of Hamacreadium hainanense Shen, 1990 by Martin et al. (2017).
Hamacreadium mutabile Linton, 1910 (Fig. 15.38)
Host: Lethrinidae: Lethrinus nebulosus (Forsskål) (1); Lutjanidae: Lutjanus fulvi-
flamma (Forsskål) (1, 2); L. rivulatus (Cuvier) (1, 2), L. quinquelinearis (Bl.) (1)
Locality: VSK, BOB
Distribution: NW Atlantic; E. Pacific Ocean; W. Pacific Ocean; Indian Ocean
References: Hafeezullah (1971c); 2. Madhavi (1975); Hafeezullah & Dutta (1980);
Bray & Cribb (1989)
Description: Body elliptical, 1760–2512 long and 720–832 wide. Oral sucker
96–144 by 136–175 in size. Ventral sucker situated in anterior third of body, larger
Fig. 15.38 Hamacreadium

mutabile (after Hafeezullah
p. 326)
than oral sucker, 136–195 by 156–210 in size. Sucker ratio 1:1.53. Intestinal bifur-
cation just anterior to ventral sucker. Caeca long, narrow. Genital pore submedian,
sinistral, opposite caecal bifurcation. Testes rounded, oblique, contiguous in poste-
rior half of hindbody. Cirrus-sac small, narrow, running obliquely between anterior
margin of ventral sucker and genital pore, encloses seminal vesicle, pars prostatica
and cirrus. Ovary three-lobed, to right of anterior testis. Seminal receptacle saccu-
lar. Uterus occupies space between ovary and ventral sucker. Eggs 60–62 by 28–34.
Vitelline follicles numerous, extend from level of intestinal bifurcation to posterior
end of body, confluent posterior to testes.
Remarks: H mutabile has a wide geographic distribution having been recorded from
many species of marine fish from different geographic locations. It shows specificity
towards lutjanid fishes (Martin et al. 2017). From India, it was first reported by
Hafeezullah (1971a) from Lethrinus nebulosus and subsequently by Madhavi (1975)
from Lutjanus fulviflamma and L. rivulatus. Hafeezullah & Dutta (1980) considered
Maculifer spiralis Soota, Srivastava &Ghosh, 1970 as a synonym of H. mutabile
and reported H. interruptum from an unidentified marine fish from Andaman sea.
Bray & Cribb (1989) agreed with the synonymy proposed by Hafeezullah and further
considered H. interruptum Nagaty, 1941 and H. indicum Gupta and Tewari, 1985
from Chirocentrus dorab as synonyms of H. mutabile. Maculifer spiralis Soota,
Srivastava & Ghosh, 1970 was, however, considered a synonym of Hamacreadium
hainanense Shen, 1990 by Martin et al. (2017). Priority suggests that the name
should be Hamacreadium spiralis (Soota, Srivastava & Ghosh, 1970) n. comb. It
is distinguished from H. mutabile by the structure of the excretory vesicle and the
anterior vitelline extent (Martin et al. 2017). Martin et al. (2017) recognized H.
interruptum ‘as a distinct species, distinguishable from H. mutabile morphologically,
biogeographically, and by infection in lethrinids rather than lutjanids.’
Genus Macvicaria Gibson & Bray, 1982

Macvicaria Gibson & Bray, 1982 was erected by Gibson & Bray (1982) to include
species of Plagioporus reported from marine fish and possessing typical plagioporine
characters such as blind caeca, an excretory vesicle extending anterior as far as the
testes, the genital pore ventro-sinistral in the mid-forebody, an entire ovary and the
vitellarium extending into the forebody. The genus has been thoroughly reviewed by
Bray & Justine (2009).
Diagnosis: Body elongate oval, smooth tegument, oral and ventral suckers simple,
ventral sucker usually larger than the oral. Caeca blind, extend posteriorly past testes.
Testes two, tandem to oblique, separated from posterior end. Genital pore sinistral,
in mid-forebody. Ovary entire. Uterus restricted to area between ovary or testes and
ventral sucker. Eggs without filaments. Vitelline follicles enter forebody and extend
posteriorly beyond testes. Excretory vesicle extends well anterior to anterior testis.
Type species: M. alacris (Looss, 1901) Gibson & Bray, 1982.
The genus now contains over 40 species. Bray & Justine (2009) provided a very
useful key for the separation of various species. Ten species of Macvicaria have
been recorded from marine fish of India. But some of these species have been syn-
onymized: M. deeghaensis (Gupta& Gupta, 1988) with M. jagannathi (Reference:
Bray, 1990b; Aken’ Ova, Cribb and Bray, 2008), M. filamentusi (Gupta & Govind,
1984) with M. yamagutii (Gupta & Ahmad, 1977) (syn: Allopodocotyle yamagutii
(Gupta & Ahmad, 1977). M. yamagutii (Gupta & Ahmad, 1977) is considered as a
possible synonym of M. jagannathi by Bray & Justine 2009). Further M. puriensis
Gupta & Govind, 1984 based on a single specimen appears to be another synonym
of M. jagannathi. Details of the remaining species from Indian marine fish are given
below:
Molecular studies have shown that the genus as at present constituted is not mono-
phyletic (Antar et al., 2015; Bray et al. 2016; Faltýnková et al., 2017; Rima et al.
2017).
Macvicaria chilkai (Gupta & Govind, 1984) Bray & Justine, 2009 (Fig. 15.39)
Syn: Plagioporus chilkai Gupta & Govind, 1984
Host: Trichiuridae: Trichiuris salva (Bleeker.) (sic. Trichiurus savala Cuvier)
Locality: Chilka lake, Orissa
Reference: Gupta, P.C. & Govind (1984b)
Description: Body elongate, 1890–2100 long, 430–540 wide. Oral sucker subter-
minal, 100–120 long, 140–190 wide. Ventral sucker larger than oral sucker, 240–250
long, 250–260 wide. Sucker ratio 1:1.4–1.8. Prepharynx absent. Pharynx subglob-
ular. Oesophagus, nearly equal to pharynx. Intestinal bifurcation nearer to ventral
sucker than to oral sucker. Caeca end blindly near posterior end of body. Genital
pore submedian, sinistral to mid-oesophagus. Testes entire, subequal, tandem, sep-
arated by vitelline follicles, in posterior half of body. Cirrus-sac elongate, curved,
extends beyond ventral sucker, either just posterior to ventral sucker or just posterior
Fig. 15.39 Macvicaria

chilkai (after Gupta &
Govind, 1984)
to middle of ovary and ventral sucker; encloses long and slender seminal vesicle,
long, tubular pars prostatica, surrounded by a large number of prostate gland cells
and tubular, thick-walled ejaculatory duct. Ovary entire, subspherical, median, equa-
torial, pretesticular. Seminal receptacle ovoid, smaller than ovary, lies postero-lateral
to ovary, overlapping its margin dorsally. Uterus coiled between ovary and ventral
sucker, opens into genital atrium through thick-walled metraterm. Eggs oval, few,
operculate, 70–80, long, 32–40 wide. Vitellarium follicular, rounded, interrupted
along testes, extend from level of intestinal bifurcation to hind end of body, conflu-
ent in post-testicular area.
Remarks: This species is characterized by the cirrus-sac extending just into the
hindbody, the vitellarium extending to the level of the bifurcation, the long slender
seminal vesicle and the submedian genital pore lying sinistrally at mid-oesophagus
level.
Macvicaria cynoglossi (Madhavi, 1975) Bray, 1990 (Fig. 15.40)
Syn. Plagioporus cynoglossi Madhavi, 1975
Host: Cynoglosidae: Cynoglossus lida (Bleeker) (1); C. arel (Bl. & Schn.); Soleidae:
Aesopia cornuta Kaup; Zebrias altipinnis (Alcock); Z. synapturoides (Jenkins)(2)
Locality: VSK, BOB (1); Trivandrum, AS (2)
Reference: 1. Madhavi (1975a); 2. Bijukumar (1997)
Description: Body fusiform, with slight indentation at level of ventral sucker,
3440–4000 long and 750–960 wide. Oral sucker subterminal, 200–215 in diame-

cynoglossi (after Madhavi,
1975)
ter. Ventral sucker 360–410 in diameter, ratio of sucker widths 1:1.6–2.0. Prephar-
ynx absent. Pharynx globular. Oesophagus one and half to twice length of pharynx.
Intestinal bifurcation nearer oral sucker than ventral sucker. Caeca narrow, terminate
near posterior extremity. Genital pore sinistral, at level of posterior end of oesoph-
agus. Testes slightly lobed, obliquely tandem to tandem, intercaecal, in posterior
half of hindbody. Cirrus-sac slender, somewhat dilated at distal end, extends to ante-
rior margin of ventral sucker or slightly overlapping ventral sucker, encloses long
seminal vesicle, small prostatic vesicle and short cirrus. Ovary spherical, pretesticu-
lar, approximately at middle of hindbody. Seminal receptacle small, inconspicuous,
dorsal to ovary. Mehlis’ gland large. Uterus intercaecal, between ovary and ventral
sucker. Vitelline follicles irregular in shape, fields from level of pharynx to posterior
end, mostly extracaecal, confluent in post-testicular area and in forebody. Numerous
deeply staining gland cells smaller than vitelline follicles occur along edges of body.
Eggs 66–71 by 47–50. Excretory vesicle I-shaped, extends to ovary.
Remarks: The body shape, the extent of cirrus-sac only to the anterior margin of the
ventral sucker, the position of the genital pore sinistral to the oesophagus and the
lobed nature of the testes are the important features of M. cynoglossi.
Macvicaria deeghaensis (Gupta & Gupta, 1988) Aken-Ova, Cribb & Bray, 2008
Syn. of M. jagannathi (Gupta and Govind, 1984) Bijukumar, 1997
Host: Sparidae: Pagrus spinifer (Cuv. & Val.); Argyrops spinifer Forskal
Locality: Bay of Bengal, Digha coast, West Bengal, India.
No. 5
Remarks: Synonymized with M. jagannathi by Bray & Justine (2009)

Reference: Gupta, P.C. & Gupta, V.C (1988)
Macvicaria filamentusi (Gupta & Govind, 1984) Bray & Justine, 2009
[Syn. of M. yamagutii (Gupta &Ahmad, 1977) Bray & Justine, 2009:]
Host: Gerreidae: Gerres filamentosus (Cuv. & Val.)
Locality: PR, BOB
Reference: Gupta, P.C.& Govind(1984b)
Macvicaria indica (Gupta & Sehgal, 1970)
Host: Lutjanidae: Lutianus fulviflamma (Forskal)
Locality: Mumbai, AS
Reference: Gupta, N. K.& Sehgal (1970)
Remarks: The description of the species based on a single specimen is inadequate.

For instance, details of the cirrus-sac were not provided. The genital pore is mentioned
as dextral. The presence of a seminal receptacle was not mentioned. The body shape
is very different from that of other species.
This species may belong to some other genus

Macvicaria jagannathi (Gupta & Singh, 1985) Bijukumar, 1997 (Fig. 15.41)
Syn: Plagioporus jagannathi Gupta & Singh, 1985; P. deeghaensis Gupta & Govind,
1984; M. deeghaensis (Gupta & Govind, 1984)
Host: Labridae: Pseudodax moluccanus (Bleeker); Sparidae: Argyrops spinifer
(Forsskål), Nemipteridae: Nemipterus furcosus (Valenciennes)
Locality: Puri coast, Bay of Bengal; New Caledonia
Reference: Gupta & Singh (1985a); Gupta, P.C. & Gupta, V.C. (1988); Bijukumar
(1997); Bray & Justine (2009)
Description: Body elongate, 2510–2730 × 530–580. Oral sucker subterminal,

100−130 × 140−170. Ventral sucker larger than oral sucker, 290−300 × 280–300.
Sucker ratio 1:1.6–2.3. Prepharynx absent. Pharynx subglobular. Oesophagus short.
Caeca simple, extend to just anterior to posterior end of body. Testes entire, sub-
spherical, tandem, subequal, separate, in posterior half of body. Cirrus-sac elongate,
slender, extends beyond ventral sucker. Seminal vesicle elongate. Pars prostatica
tubular, surrounded by many prostate gland cells. Ejaculatory duct tubular, thick-
walled. Genital pore submedian, sinistral to pharynx. Ovary entire, subspherical,
submedian, pretesticular. Seminal receptacle ovoid, longer than ovary, lies postero-
lateral to ovary. Uterus between ovary and ventral sucker. Eggs ovoid, operculated,
63–71 × 43–53. Vitellarium follicular, extends either from bifurcal level or just pos-
terior to posterior end of body, non-confluent inforebody.

jagannathi (after Gupta &
Singh, 1985, Acta Parasitol.
42, p. 1)
Remarks: This species was originally included in the genus Plagioporus as P. jagan-
nathi Gupta & Singh, 1985. Bijukumar (1997) transferred it to the genus Macvicaria.
The shape of the body, the long post-testicular space, the cirrus-sac extending poste-
riorly to the ventral sucker, the position of the genital pore sinistral to the pharynx and
the extent and distribution of vitelline follicles distinguishes M. jagannathai from
other Indian species of Macvicaria.
Macvicaria longicaudus (Hafeezullah, 1971) Bijukumar, 1997 (Fig. 15.42)
Syn. Plagioporus longicaudus Hafeesullah, 1971
Hosts: Cynoglossidae: Cynoglossus lida (Bleeker) (1); C. dubius Day; C. macrosto-
mus Norman; Soleidae: Synaptura commersoniana (Lacépède) (2)
Localities: Tuticorin, GOM (1); Kerala, AS (2)
Reference: 1. Hafeezullah (1971c); 2. Bijukumar (1997)
Description: Body 2611–3552 long, 612–800 wide, elongate, subcylindrical. Oral
sucker 173–237 in diameter, subterminal, smaller than ventral sucker. Ventral sucker
266–330 in diameter, spherical, at 753–1059 from anterior end of body. Sucker
width ratio 1:1.3–1.5. Prepharynx indistinct. Pharynx globular. Oesophagus long.
Intestinal bifurcation nearer to oral sucker than to ventral sucker. Caeca not quite
reaching posterior end of body. Testes oval, entire or slightly irregular, tandem,
immediately postequatorial. Cirrus-sac claviform, not reaching ventral sucker pos-
teriorly, encloses coiled seminal vesicle, pars prostatica with gland cells and cirrus.
Genital pore sinistral at about mid-oesophageal level. Ovary globular, immediately
pretesticular, slightly right of median line. Seminal receptacle dorsal to and much
larger than ovary, to right of median line. Uterus between ovary and genital pore;
metraterm differentiated. Eggs 74–82 × 38–47. Vitelline follicles extend from intesti-
nal bifurcation to posterior end, may be continuous or not at level of ventral sucker,
confluent in post-testicular space.
longicaudus (after
Hafeezullah 1971 Courtesy
Cambridge Univ. Press,
Parasitology, 62, p. 322)
Remarks: The shape of the body, the appreciably longer post-testicular space, the
cirrus-sac extending only to the anterior margin of the ventral sucker and the extent
and distribution of vitelline follicles distinguishes this species from all the other
Indian species of the genus Macvicaria.
Macvicaria puriensis (Gupta & Govind, 1984) Bray & Justine, 2009
Syn, Plagioporus puriensis Gupta & Govind, 1984
Host: Lutjanidae: Lutjanus waigiensis (Quoy & Gaimard)
Locality: PR, BOB
Reference: Gupta & Govind (1984b)
Remarks: The description based on a single specimen lacks details and the extent of
intraspecific variations exhibited by the species is not known. It is here regarded as
a species inquirenda.
Macvicaria yamagutii (Gupta & Ahmad, 1977) Bray & Justine, 2009 (Fig. 15.43)
Syn: Allopodocotyle yamagutii (Gupta & Ahmad, 1977)
Location: Intestine
Locality: Puri, Orissa
Reference: Gupta, V& Ahmad (1977a)
in diameter. Ventral sucker larger than oral sucker, 255–320 long, 330–390 wide.
Pharynx well developed. Oesophagus long. Caeca terminate near posterior extremity.
Testes oval, tandem, subequal in middle third of body, separated from each other.
Cirrus-sac elongate, extends from genital pore to beyond posterior margin of ventral
Ahmad, 1977)
sucker. Seminal vesicle long, tubular. Pars prostatica surrounded by many prostate
gland cells. Ejaculatory duct 50–80 long. Genital pore submedian just posterior to
pharynx. Ovary oval, median, pretesticular, separated from anterior testis. Seminal
receptacle globular, overlapping right side of ovary. Uterus winding between ovary
and genital pore. Eggs elongate, operculated, 61–94 long, 30–51 wide. Vitelline
follicles extend from level of intestinal bifurcation or just posterior to it to hind end
of body.
Remarks: This species was first described as Allopodocotyle yamagutii by Gupta

and Ahmad (1977) and was transferred to the genus Macvicaria as M. yamagutii by
Bray and Justine (2009). Bray & Justine (2009) expressed the view that it might fall
a synonym of M. jagannathi.
The five valid Indian species of Macvicaria can be separated by the following
key:
1. Testes lobed……………………………………..M. cynoglossi(Madhavi, 1975)

Testes entire……………………………………….…….2
2. Cirrus-sac extending to ventral sucker…………..….M. longicaudus (Hafeezul-
lah, 1971)
Cirrus-sac extending beyond ventral sucker…………………………3
3. Cirrus-sac reaching into posterior part of ventral sucker to ovary dis-
tance…….………………………..M. yamagutii (Gupta & Ahmad, 1977)
Cirrus-sac reaching into anterior part of ventral sucker to ovary dis-
tance………….……….4
4. Genital pore sinistral to mid-oesophagus………………….….M. chilkai (Gupta
& Govind, 1984)
Genital pore sinistral to pharynx……………M. jagannathi (Gupta & Singh,
1985)
Genus Phyllotrema Yamaguti, 1934
Generic Diagnosis Body oval, posterior extremity rounded or with 4–5 distinct
pointed projections. Oral sucker simple. Ventral sucker larger than oral sucker. Caeca
long. Testes two, symmetrical, well separated from ventral sucker. Genital pore sinis-
tral or dextral, in mid-forebody. Ovary lobed, pretesticular. Uterus passes between
testes to posterior half of hindbody. Vitelline follicles enter forebody and extend pos-
teriorly beyond testes to posterior end of body. Excretory vesicle extends to ovary.
Type species: T. bicaudatum Yamaguti, 1934.
Phyllotrema tetracaudatum Hussain, Rao & Shymsundari, 1986 (Fig. 15.44)
Host: Congridae: Uroconger lepturus (Richardson)
Locality: VSK, BOB
Description: Body elongate, foliate, 4000–4768 long, 1648–2176 broad, with

rounded anterior end and tetrafid posterior extremity. Oral sucker 400–416 ×
Fig. 15.44 Phyllotrema

tetracaudum (after Hussain,
Rao & Shymsundari, 1986)
400–432 in size. Ventral sucker large, 624–704 × 624–720 in size, situated at begin-
ning of middle third of body. Caeca simple, long. Testes globular, unequal, symmet-
rical, just post-equatorial. Cirrus-sac largely in forebody. Seminal vesicle strongly
recurved, occupies greater part of cirrus-sac. Genital pore at level of oesophagus.
Ovary smaller than testes, between ventral sucker and testes. Seminal vesicle elon-
gate, just anterior to ovary. Uterus pretesticular. Eggs 80–96 × 32–48. Vitelline fol-
licles extend from level of intestinal bifurcation to posterior end of body, forming
bunches of follicles. Excretory vesicle simple, bifurcating into lateral horns imme-
diately posterior to ovary.
Remarks: Yamaguti (1934) erected the genus Phyllotrema, included it in the fam-
ily Allocreadiidae and indicated its affinity with the family Fellodistomidae in the
disposition of testes, ovary and uterus. Yamaguti (1971) erected a new subfamily
Phyllotrematinae in the family Lepocreadiidae to include this genus. P. tetracau-
datum differs from the type species P. bicaudatum Yamaguti, 1934, in having four
lobes at the posterior end, in the cirrus-sac situated in the forebody, and the uterus
not extending posterior to the testes.
Genus Podocotyle Dujardin, 1845
[Syn. Sinistroporus Stafford, 1904; Psilolintonum Oshmarin, 1964; Neopodocoty-
loides Pritchard, 1966]
This is one of the oldest and largest opecoelid genera. Over one hundred nominal
species have been described in the genus, but Bray & Campbell (1982) in a detailed
analysis of the genus transferred many of the species into other genera and retained
only 20 valid species. The important features diagnostic of the genus are: genital
pore sinistral at the level of the oesophagus, ovary three- to four-lobed, vitelline
follicles usually not entering forebody, but extending posteriorly beyond the testes to
the posterior end of the body, and uterus restricted to the area between ovary, ventral
sucker and genital pore.
Remarks: Three species of Podocotyle have been described from marine fish of
India: P. dorabi Gupta & Puri, 1979 from Chirocentrus dorab, P. indica Pandey &
Tewari, 1989, from a marine fish and P. simhai Gupta & Sayal. 1979 from C. dorab.
Of these, P. indica was regarded as species inquirendum, and the remaining two
species are considered as probable synonyms of Acanthocolpus liodorus Luhe, 1906
(Reference: World Register of Marine species by Gibson, 2014)
Genus Podocotyloides Yamaguti, 1934
Syn: Pedunculacetabulim Yamaguti, 1934; Pedunculotrema Fischthal & Thomas,
1970.
This genus includes Podocotyle-like forms with a round ovary and a pedunculate
ventral sucker. Aken’Ova (2003) provided a review on the genus.
Generic diagnosis: Body elongate. Oral sucker simple. Ventral sucker pedunculate,
usually larger than oral sucker. Caeca blind, extend close to posterior end of body.
Testes two, tandem, separated from posterior end of body. Genital pore sinistral
usually in mid-forebody. Uterus restricted to area between ovary and ventral sucker.
Eggs without filaments. Vitelline follicles may or may not extend into forebody,
extend posteriorly beyond posterior end of testes to posterior end of body. Type
species: P. petalophallus Yamguti, 1934
Details of the five species of the genus reported from marine fish of India are given
below:
Podocotyloides dorabus Lokhande, 1990
Host: Cynoglossidae: Cynoglossus oligolepis (Bleeker)
Locality: RT, AS
Reference: Lokhande(1990d)
Podocotyloides parupenei (Manter, 1963) Pritchard, 1966 (Fig. 15.45)
[Syn. Podocotyle parupenei Manter, 1963]
Host: Terapontidae: Terapon puta (Cuv. & Val.) (1); T. jarbua Forskål (2); T. theraps
(Cuvier) (1); Nemipteridae: Nemipterus japonicus (Bloch) (1)
Locality: Tuticorin GOM (1); VSK, BOB (2)
Reference: 1. Hafeezullah (1971c); 2. Madhavi (1975a)
Distribution: Fiji, Northern Great Barrier Reef.
Description: Body slender, elongate, 2185–4168 long 368–402 wide. Oral sucker
longer than wide, 107–140 in diameter. Ventral sucker protuberant, 214–368 in diam-
eter, sucker ratio 1:1.9–2.0. Forebody 10–14% of body length. Oesophagus short.
Caeca narrow not quite reaching posterior end of body. Genital pore sinistral, at level
of pharynx. Testes elongate, smooth, tandem, separated by vitelline follicles, anterior
testis near midbody, post-testicular space usually 25–33% of body length. Cirrus-sac
sinuous, extends well posterior to ventral sucker to midway between ventral sucker
and ovary, contains straight seminal vesicle, with single constriction near base, distal
Fig. 15.45 Podocotyloides

parupenei (after Hafeezullah
p. 322)
portion of cirrus-sac protrusible. Ovary smooth, spherical, separated from anterior

testis by vitelline follicles. Seminal receptacle flask-shaped. Uterus preovarian with
few eggs. Eggs large, 72–88 by 38–57. Vitelline follicles large, extend from slightly
posterior to base of ventral sucker, interrupted at level of gonads, contiguous dor-
sal to cirrus-sac and between gonads, largely filling post-testicular space. Excretory
vesicle extends to level of ovary, pore terminal.
Remarks: This species was first described as Podocotyle parupenei from Parupenus
indicus (Shaw) by Manter (1963) from Fiji. Pritchard (1966) transferred it to the
genus Podocotyloides on the basis of its unlobed ovary and pedunculate ventral
sucker. Hafezullah (1971c) recorded it in Terapon puta Cuv. & Val., T. theraps
Cuvier and Nemipterus japonicus (Bloch) from the Indian region. Later, Madhavi
(1975a) recorded it from Terapon jarbua from Bay of Bengal. According to Madhavi
(1975a), the eggs lacked the knob-like protrusion at one end noted by Manter (1963)
in the specimens from Fiji.
Podocotyloides pedicellatus (Srivastava, 1938) Pritchard, 1966
[Syn. Peduculacetabulum pedicellatum Srivastava, 1937]
Host: Hemiscylliidae: Chiloscyllium indicum (Gmelin)
Reference: Srivastava (1938d)
Podocotyloides pentavitellosus Yadav, 1981
Host: Chirocentridae: Chirocentrus dorab Forsskål

Podocotyloides yamagutii Khan & Karyakarte, 1987

Host: Cynoglossidae: Cynoglossus oligolepis (Gunther)
Locality: RT, AS
Reference: Khan & Karyakarte (1987a)
Genus Vesicocoelium Tang, Hsu, Huang & Lu, 1975
[Syn. Godavaritrema Karyakarte & Yadav, 1976]
Diagnosis: Body oval. Oral sucker simple. Ventral sucker unspecialized, larger than
oral sucker. Caeca unite to open via common anus. Testes two, tandem, in posterior
half of body. Genital pore sinistral at level of oesophagus. Ovary entire. Uterus
preovarian. Eggs oval, without filaments. Vitelline follicles extend well into forebody
and posteriorly beyond testes. Excretory vesicle extends to level of ovary. Type
species: V. solenophagum Tang, Hsu, Huang& Lu, 1975.
Vesicocoelium marinum (Karyakarte & Yadav, 1976) Cribb, 2005 (Fig. 15.46)
[Syn. Godavaritrema marinum Karyakarte & Yadav, 1976]
No. 4
Reference: Karyakarte & Yadav (1976).
Description: Body elongated, 2100–3140 long, 780–1200 wide. Oral sucker

160–200 in length and 120–160 in width. Ventral sucker larger than oral sucker,
measuring 200–220 by 240–260, at junction of anterior and middle third of body.
Sucker ratio 1:1.50–1.70. Prepharynx short. Oesophagus fairly long. Caeca unite
and open out through common anal pore close to posterior end of body. Testes trans-
versely elongated, tandem, in posterior third of body. Cirrus-sac long extends to
Fig. 15.46 Vesicocoelium

marinums (After Karykarte
& Yadav 1976)
near posterior margin of ventral sucker, encloses oval seminal vesicle, few prostatic
cells and short cirrus. Ovary elongated, pretesticular, submedian. Seminal receptacle
uterine. Vitelline follicles extend from level of oesophagus to posterior end of body.
Uterus preovarian. Eggs 64–68 × 34–38.
Remarks: Karyakarte & Yadav (1976) erected the genus Godavaritrema to include
G. indica (type species) from the freshwater fish Macrones seenghala and G. marina
from the marine fish Terapon theraps. Cribb (2005) synonymized the genus Godavar-
itrema with Vesicocoelium Tang, Hsu, Huang& Lu, 1975. The genus Vesicocoelium
closely resembles the opecoeline genus Opegaster Ozaki, 1928 but differs from it
mainly in lacking papillae on the ventral sucker and in the presence of a cirrus-sac.
Subfamily Stenakrinae Yamaguti, 1970
This subfamily shows characters intermediate between subfamilies Opecoelinae

and Plagioporinae. The Stenakrinae possess a well-developed cirrus-sac but lack
a canalicular seminal vesicle. Species recorded under this subfamily from Indian
marine fish belong to two genera: Neonotoporus Srivastava 1942 and Pseu-
dopecoelina.
Neonotoporus: genital pore is nearly marginal, lateral to ventral sucker, pars prostat-
ica strongly developed, Caeca blind.
Pseudopecoelina: genital pore sinistral, almost marginal, pars prostatica not so well
developed. Caeca open into excretory bladder to form uroproct.
Neonotoporus Srivatava, 1942
The genus was proposed by Srivastava (1942b) for Notoporus trachuri Yamaguti,
1938 and now contains about 12 species of which 5 species have been recorded from
marine fish of India.: N. gibsoni Ahmad 1990; N. overstreeti Ahmad & Dhar, 1987
and N. srivastavai Ahmad & Dhar, 1987; N. skrjabini Ahmad, 1987; N. leiognathi
(Hafeezullah, 1971) Ahmad, 1985.
Generic diagnosis: Body elongate or oval. Oral sucker oval or funnel-shaped. Ven-
tral sucker unspecialized. Caeca blind, terminate close to posterior end of body. Testes
oval, oblique or almost symmetrical. Pars prostatica strongly developed. Genital pore
usually sinistral, anywhere from posterior to anterior margin of ventral sucker. Ovary
entire or distinctly lobed. Uterus extends posteriorly as far as to posterior margin of
testis. Vitelline follicles extend well into forebody.
Neonotoporus gibsoni Ahmad, 1990 (Fig. 15.47)
Locality: Goa, AS
No. 7
Subfamily Stenakrinae Yamaguti, 1970 473
Fig. 15.47 Neonotoporus

gibsoni (Ahmad, 1990)
Description: Body elongated, 1440–1642 long by 260–295 wide. Oral sucker fin-
ger bowl-shaped, terminal, 132–140 long by 180–220 wide. Ventral sucker larger
than the oral sucker, occupies almost entire body width, 192–220 long by 240–270
wide, pre-equatorial. Sucker ratio 1:1.22–1 33. Intestinal bifurcation dorsal, midway
between suckers. Caeca terminate at posterior end of body, ending blindly. Testes
two, deeply multi-lobed, oblique, close together, situated in posterior third of body,
subequal in size. Cirrus-sac club–shaped, dorsal to ventral sucker, curved anteriorly,
extends posteriorly just anterior to midway between ventral sucker and ovary. Sem-
inal vesicle tubular. Prostatic vesicle bulbous, surrounded by prostate gland cells.
Cirrus short, muscular. Genital pore marginal, towards right margin, at anterior level
of ventral sucker. Ovary three-lobed, submedian to left of median line, pretesticular,
partly overlaps left caecum. Uterus confined to intercaecal field between anterior bor-
der of posterior testis and ventral sucker. Eggs elongate, operculate, 70–80 by 40–48.
Vitellarium consists of numerous follicles, interrupted at level of ventral sucker on
both sides, extend from posterior margin of pharynx to posterior end of body, conflu-
ent in forebody and post-testicular zone. Excretory vesicle tubular, sigmoid, reaching
anteriorly as far as anterior level of anterior testis. Excretory pore terminal.
Remarks: The diagnostic features of this species are: oral sucker is finger bowl-
shaped, all the gonads are lobed, and the vitellarium is interrupted on both sides of
ventral sucker.

leiognathi (after Hafeezullah
p. 328)
Neonotoporus leiognathi (Hafeezullah, 1971) Ahmad, 1985 (Fig. 15.48)

[Syn: Hamacreadium leiognathi Hafeezullah, 1971]
Reference: Hafeezullah (1971c)
Description: Body 1506–1741 long, 353–435 wide at testicular level, elongate. Oral
sucker 87–111 disc-like, deep, 161–190 wide, terminal. Ventral sucker 152–205 ×
161–208, spherical, pre-equatorial. Sucker width ratio 1:1–1·2. Intestinal bifurcation
almost halfway between pharynx and ventral sucker. Caeca ending just short of pos-
terior end of body. Testes slightly lobed, diagonal with left more anterior, in posterior
third of body. Cirrus-sac very long, extends well posterior to ventral sucker, contains
coiled, tubular seminal vesicle; pars prostatica with prostatic gland cells, ejaculatory
duct and cirrus. Genital pore post-bifurcal, sinistral, lies ventral to left caecum or
lateral to it. Ovary 2–4-lobed, right of median line, postequatorial, pretesticular. Sem-
inal receptacle antero–dorsal to ovary. Vitellarium follicular, from level of intestinal
bifurcation or slightly anterior to it to posterior end of body, discontinuous on left
side in ventral sucker zone. Uterus scanty, proximal coils filled with sperm, becoming
postovarian in posterior extent; metraterm indistinct. Eggs 75–90 × 56–66.
Remarks: This species was initially included in the genus Hamacreadium as

H. leiognathi Hafeezullah, 1971 and was transferred to the genus Neonotoporus
by Ahmad (1985). The important features of the species are: the pear-shaped body,
disc-like oral sucker and the vitellarium interrupted opposite the ventral sucker on
one side.
Neonotoporus overstreeti Ahmad & Dhar 1987 (Fig. 15.49)
Locality: PR, BOB

overstreeti (after Ahmad &
Dhar 1987)
No. 10
Reference: Ahmad & Dhar (1987c)
Description: Body elongate, with nearly parallel sides, expanded at ventral sucker
level, extremities rounded, 1980–2450 long and 485–602 wide at level of the ven-
tral sucker. Oral sucker 170–200 in diameter. Ventral sucker larger than the oral
sucker, 240–270 long and 220–250 wide, 262–305 post-bifurcal, situated in anterior
part of middle third of body. Sucker ratio 1:1.25–1.30. Prepharynx absent. Intestinal
bifurcation dorsal, nearer to the oral sucker than to ventral sucker. Caeca extend to
posterior end of body. Testes two, deeply multi-lobed, diagonally tandem, postequa-
torial, close together, equal in size. Cirrus-sac club-shaped, curved anteriorly, extends
posteriorly as far as anterior margin of ovary. Seminal vesicle tubular. Prostatic vesi-
cle bulbous, lined internally with tall epithelia, surrounded by prostate gland cells.
Genital pore marginal, towards left body margin, at anterior level of ventral sucker.
Ovary entire, transversely elongate. Seminal receptacle absent. Uterus confined to
intercaecal field between anterior border of posterior testis and ventral sucker. Eggs
elongate, operculate, 70–82 long and 35–40 wide. Vitellarium consists of numerous
follicles, interrupted at level of ventral sucker on one side of body only, extend from
anterior level of intestinal bifurcation to posterior end of body, confluent in forebody
and in post-testicular zone. Excretory vesicle tubular sigmoid, reaches anteriorly up
to the anterior margin of the anterior testis
Remarks: The distinguishing features of the species are: oral sucker funnel-shaped,
testes lobed, ovary entire, vitellarium interrupted on one side of ventral sucker, cirrus-
sac extends to near posterior margin of ventral sucker.
Neonotoporus skrjabini Ahmad, 1987

Locality: PR, BOB
No. 24
Description: Body elongate with nearly parallel sides, 2020–2070 × 260–310 in

size. Oral sucker finger bowl-shaped, 125–175 × 100–230. Ventral sucker oval,
median, larger than oral sucker 215–240 × 230–270, pre-equatorial. Sucker ratio
1: 1.14–1.21. Prepharynx absent. Pharynx oval. Oesophagus long. Intestinal bifurca-
tion midway between suckers. Caeca terminate at posterior end of body. Testes two,
oval, oblique, close together, subequal in size, postequatorial. Cirrus-sac club-shaped,
dorsal to ventral sucker, curved anteriorly, extends posteriorly to anterior margin of
ovary. Seminal vesicle tubular; prostatic vesicle bulbous, lined internally with tall
epithelia, cirrus short. Genital pore marginal towards left margin, midway between
ventral sucker and intestinal bifurcation. Ovary spherical, pretesticular, submedian.
Seminal receptacle absent. Vitelline follicles numerous, extend from bifurcal level to
posterior end of body, interrupted at level of ventral sucker on both sides, confluent
anterior to ventral sucker. Uterus descends to anterior margin of posterior testis. Eggs
88–92 × 42–50.
Remarks: The distinguishing features of this species are: vitelline follicles are inter-
rupted on both sides of ventral sucker, gonads are spherical, cirrus-sac very long,
extends to level of ovary.
Neonotoporus srivastavai Ahmad & Dhar, 1987 (Fig. 15.50)
Host: Trichiuridae: Eupleurogrammus muticus (Gray)
Locality: PR, BOB
No. 4
Reference: Ahmad & Dhar (1987c)
Description: Body elongate, 2080–2440 long and 290–350 wide. Oral sucker
funnel-shaped, terminal, massive, 260–300 long and 175–192 wide; anterior border
of oral opening provided with two labia. Ventral sucker larger than the oral sucker,
250–270 long and 228–240 wide, equatorial in position. Sucker ratio 1:1.25–1.30.
Intestinal bifurcation in posterior forebody. Caeca terminate at posterior end of the
body. Testes two, deeply multi-lobed, oblique, close together, postequatorial, situated
in middle of posterior third of body, longer than wide, subequal in size. Cirrus-sac
roughly C-shaped, dorsal to ventral sucker, extends posteriorly as far as posterior
margin of ventral sucker. Seminal vesicle saccular, straight. Prostatic vesicle bul-
bous, lined internally with tall epithelia, surrounded by prostate gland cells. Genital
pore marginal, towards left body margin, at anterior level of ventral sucker. Ovary
spherical, pretesticular, to right of median line, partly overlapping right caecum.
Seminal receptacle absent. Uterus conformed to intercaecal field between anterior
margin of posterior testis and ventral sucker. Eggs 80–95 long and 40–45 wide.
Vitelline follicles, interrupted at level of ventral sucker on both sides, extend from

srivastavai (after Ahmad &
Dhar, 1987)
anterior bifurcal level to posterior end of body, confluent in forebody and in post-
testicular zones. Excretory vesicle tubular, sigmoid, reaches anteriorly up to level of
the anterior testis.
Remarks: The distinguishing features of this species are: the oral sucker is funnel-
shaped, vitellarium is interrupted at level of ventral sucker on both sides, testes
multi-lobed, ovary spherical, cirrus-sac sac extends to near posterior margin of ventral
sucker.
Key to Indian species of the genus Neonotoporus:
1. Vitellarium interrupted at level of ventral sucker on both sides…………….…2

Ovary spherical, vitellarium interrupted at level of ventral sucker on one side
only ……..3
2. Ovary spherical ……………………………..... N. skrjabini Ahmad, 1957
Ovary lobed……………………...... N. gibsoni Ahmad, 1990
3. Cirrus-sac extends posteriorly to level of ovary…………………………….. 4
Cirrus-sac extends to posterior border of ventral sucker……………………….
N. srivastavai, Ahmad & Dhar, 1987
4. Testes deeply lobed, ovary entire ………………….……. N. overstreeti Ahmad&
Dhar, 1987
Testes slightly lobed, Ovary 2–4-lobed ……………………… N. leiognathi
Hafeezullah, 1971
Genus Pseudopecoelina Yamaguti, 1942

This genus was erected by Yamaguti (1942) for P. dampieriae Yamaguti, 1942. It
is characterized by the absence of any papillae on the ventral sucker, in having the
ventral sucker larger than the oral sucker and the cirrus-sac large and extending well
into the hindbody. Three species belonging to this genus have been recorded from
Indian marine fish: P. elongata Hafeezullah, 1971; P. puriensis Ahmad, 1978 and
P. stunkardi Ahmad, 1978. Information is available to the author for only the first
species.
Diagnosis: Body elongate, with almost parallel sides. Oral and ventral suckers sim-
ple, latter larger or smaller than the former, may be protuberant. Caeca open into
excretory vesicle to form uroproct. Testes tandem, well separated from posterior end
of body. Genital pore sinistral, ovary entire or slightly lobed, uterus restricted to
area between ovary and anterior testis and genital pore. Vitelline follicles may enter
forebody. Excretory vesicle extends to ovary. Type species: S. dampieriae Yamaguti,
1942
Pseudopecoelina elongata Hafeezullah, 1971 (Fig. 15.51)

Host: Mullidae: Upeneus bensasi (Schlegel); U. taeniopterus Cuv
Locality: Mandapam, GOM, MS, BOB
No. 5
Reference: Hafeezullah (1971c)
Description: Body 2531–3396 long, 204–240 wide, elongate narrow, with short
peduncle or protuberance bearing ventral sucker near anterior extremity. No acces-
Fig. 15.51 Pseudopecoelina

elongata (after Hafeezullah,
1971)
sory sucker in forebody. Ventral sucker 152–179 × 164–178, spherical, without papil-
lae. Oral sucker 110–120 long, 83–113 deep, subglobular, ventro terminal. Length
ratio of suckers 1:1.38–1.62. Caeca simple becoming obscured posteriorly by vitel-
larium, probably joining excretory vesicle near posterior end to form cloaca. Testes
oval or spherical, tandem, separated by vitellarium, in middle of posterior half of
body. Cirrus-sac long, tubular, extends far back of ventral sucker contains long, tubu-
lar seminal vesicle, short pars prostatica and ejaculatory duct. Genital pore median,
ventral, post-pharyngeal. Ovary spherical, median, pretesticular, separated from ante-
rior testis by vitellarium. Seminal receptacle absent. Shell gland preovarian. Vitelline
follicles from halfway between ventral sucker and ovary to short of posterior end of
body, discontinuous in testicular and sparse in ovarian zones. Uterus scanty, preo-
varian. Eggs collapsed, 69–76 × 30–39. Excretory vesicle I-shaped.
Remarks: It differs from the type species in the shape and size of the body, the longer
post-testicular space and the larger eggs.
Pseudopecoelina puriensis Ahmad, 1978
Host: Terapontidae: Terapon theraps (Cuv.)
Locality: PR, BOB
Remarks: This species was not properly described, but was said to differ from other
Pseudopecoelina species by ‘having gonads contiguous, posterior testis much larger,
shorter post-testicular space, different sucker’s ratio and vitellarium [sic] extending
from the level of pharynx up to posterior extremity’ (Ahmad, 1978).
Pseudopecoelina stunkardi Ahmad, 1978
Host: Sciaenidae: Pseudosciaena sina (Cuvier)
Locality: PR, BOB
Remarks: This species was not properly described but was said to differ from other
Pseudopecoelina species ‘in the extension of vitellarium from the level of intestinal
bifurcation up to posterior extremity. It further differs from P. dampieriae in having
ovary pre-equatorial, different shape and size of body, a long post-testicular space
and circus [sic] sac reaching up to a little anterior to ovary; from P. elongata in
not having vitellarium discontinuous in testicular and sparse in ovarian zone, ovary
close to anterior testis, different sucker’s ratio and from P. puriensis in having gonads
separated, different sucker’s ratio and in not having posterior testis much larger’
(Ahmad, 1978).
Subfamily Helicometrinae Bray, Cribb, Littlewood &

Waeschenbach, 2016
Diagnosis: Body oval to elongate. Oral sucker unspecialized or distinctly funnel-

shaped. Ventral sucker larger or smaller than oral sucker. Caeca extend close to
posterior end of body, blind or form ani. Testes two or several (3–9), usually deeply
lobed, occasionally entire, tandem to oblique; well separated from posterior end
of body. Genital pore median or slightly submedian, bifurcal or prebifurcal. Ovary
usually distinctly lobed, occasionally almost entire. Seminal receptacle canalicular.
Uterus distinctly helical, between ovary and anterior testis and genital pore. Eggs with
long unipolar filaments. Vitellarium reaches into forebody or restricted to hindbody,
post-testicular fields reach to posterior extremity. Excretory vesicle I-shaped, extends
as far as ovary. In marine fishes, cosmopolitan.
Bray et al. (2016) found molecular and morphological evidence for the recognition
of a new subfamily. The subfamily was accepted by Rima et al. (2017) and Martin
et al. (2017).
Type genus: Helicometra Odhner, 1902
1. Testes two…………..………………………. Helicometra Odhner, 1902
Testes 3 to 9…………………………. Helicometrina Linton, 1910
Genus Helicometra Odhner, 1902
[Syn.Stenopera Manter, 1933]
Diagnosis: Body oval to elongate, oral sucker circular or funnel-shaped. Ventral

sucker larger or smaller than oral sucker. Caeca blind, extend close to posterior
end of body. Testes two, tandem to oblique, well separated from posterior end of
body. Genital pore median. Ovary lobed. Uterus helical, between ovary and anterior
testis and genital pore. Eggs with polar filament. Vitelline follicles extensive, may
extend into forebody. Excretory vesicle extends to ovary. Type species: H. pulchella
(Rudolphi, 1819,) Odhner, 1902.
The genus Helicometra is large comprising about 40 species. The recent review
by Blend & Dronen (2015) should be consulted for keys and illustrations to most
species.
Helicometra filamentosa Madhavi, 1975 (Fig. 15.52)

Host: Lutjanidae: Lutjanus sp.
Locality: VSK, BOB
No. 1
Description: Body elongate, 1840 long and 640 wide, rounded posteriorly, anterior
end pointed. Oral sucker 168–203. Ventral sucker one-third body length from anterior
end, 280 in diameter. Sucker ratio 1:1.4. Prepharynx short. Pharynx 101 long by 117
wide. Oesophagus 125 long. Intestinal bifurcation midway between pharynx and
ventral sucker. Caeca terminate lateral to posterior testis. Genital pore slightly to
left of midline at oesophagus level. Testes tandem, contiguous, rounded, close to
posterior end. Cirrus-sac thin-walled, straight, overlapping anterior third of ventral
sucker, containing sinuous seminal vesicle, few prostatic cells and short cirrus. Ovary
immediately pretesticular, submedian, four-lobed, 203 long by 203 wide. Seminal
receptacle small, dorsal to ovary. Uterus coils between ovary and ventral sucker,
then extends forward to genital pore. Vitelline follicles reach from level of intestinal
Subfamily Helicometrinae Bray, Cribb, Littlewood & Waeschenbach, 2016 481
Fig. 15.52 Helicometra

filamentosa (after Madhavi,
1975)
bifurcation to posterior end of body, confluent in forebody and posterior to testes.

Egg 51 by 27, filament about 15 times as long as egg.
Remarks: H. filamentosa is most similar to H. aposinuata Pritchard, 1966, from
Holocentrus scythrops and Thalassoma duperry from Hawaii. It differs, however, in
that the caeca do not extend posterior to the testes, the testes are smaller and more
posterior in position, the cirrus-sac is straight, and the filament of the egg is very
long.
Heicometra gibsoni Murugesh, Madhavi & Swarnakumari, 1993 (Figs. 15.53
and 15.54)
[Syn.H. fasciata of Madhavi, 1975]
Host: Scorpaenidae: Scorpaenopsis cirrhosus (Thunberg); Terapontidae: Terapon
jarbua (Forsskal)
Locality: VSK, BOB
Reference: Murugesh et al. (1993)
Description: Body elliptical, 1440–3232 long, 480–928 wide. Oral sucker
140–260 × 140–252. Ventral sucker larger than ventral sucker, 192–376 × 192–372.
Sucker ratio 1:1.4. Prepharynx short. Pharynx cylindrical. Oesophagus fairly long.
Caeca long, narrow, terminate blindly close to posterior extremity. Testes post-
equatorial, lobed, closely tandem. Cirrus-sac elongate, broad posteriorly, narrow
anteriorly, extends posteriorly to mid-level of ventral sucker, encloses coiled sem-
inal vesicle, pars prostatica surrounded by numerous prostatic cells and eversible
cirrus. Genital pore median, at mid-oesophagus level. Ovary contiguous with ante-
rior testis, with 4–6 unequal lobes. Seminal receptacle retort-shaped, lateral to ovary.
Uterus coiled spirally between ovary and ventral sucker. Metraterm one-third length
of cirrus-sac. Eggs oval, large, 58–64 × 28–30, with unipolar filament. Vitelline fields
reach from mid-oesophageal level to posterior extremity of body. Excretory vesicle
extends to level of ovary.

gibsoni (after Murugesh
et al. 1993) Courtesy
Springer, Syst. Parasit, 25,
67)

gibsoni coloured
Remarks: Murugesh et al. (1993) elucidated the life cycle of H. gibsoni, with the
snail Anachis terpsichore serving as the first intermediate host, the shrimp Alpheus
malabaricus as the second intermediate host and Terapon jarbua as the experimental
definitive host. The cercariae possessed all typical opecoelid characters apart from its
long extensile tail. Natural infections with adult flukes were reported in the scorpaenid
fish Scorpaenopsis cirrhosus. Morphologically, the adult of H. gibsoni is similar to
H. fasciata (Rudophi, 1819) but the cercarial stages of the two species showed
many differences especially in the tail being long and extensile in H. gibsoni and
microcercous in H. fasciata (Reference: Reversat, 1990). Murugesh et al. (1993) also
considered H. fasciata of Madhavi (1975) from Scorpaenopsis cirrhosus a synonym
of H, gibsoni.
Genus Helicometrina Linton 1910
Diagnosis: Body elongate. Ventral sucker usually larger than oral sucker. Caeca
narrow, blind, end close to posterior end of body. Testes 3–9, in two longitudinal
parallel rows in hindbody. Genital pore submedian, in mid-forebody. Ovary deeply
lobed, pretesticular. Uterus forms concentric coils, confined to space between testis
and ventral sucker. Eggs filamented. Vitellarium enters forebody and extends to
posterior end of body. Excretory vesicle extends to level of ovary. Type species: H.
nimia Linton, 1910.
Helicometrina nimia Linton, 1910 (Fig. 15.55)
[Syn. H. orientalis Srivastava, 1936; H. septorchis Srivastava, 1936; H. elongata
Noble & Park, 1937; H. hexorchis Gupta & Sehgal, 1970; H. otolithi Bilqees, 1972;
H. karachiensis Bilqees, 1972; H. delicatulus Bilqees, 1972]
Hosts: Sciaenidae: 1, 2. Otolithus ruber (Bl. & Schn.), Pennahia anea (Bloch);
Kathala axillaris (Cuv.), J. glaucus (Dy); J. sina (Cuv.), 2, Sillaginidae: Sillago
sihama (Forskål):2. Serranidae: Epinephlus maculatus (Bloch); E. undulosus (Quoy
Fig. 15.55 Helicometrina

nimia (original)
et Gaim.); Lutianidae: Lutianus johnii (Bloch); L. quinquelinearis (Bleeker); L. rivu-

latus (Cuv); Platycephalidae: Grammoplites indicus; Platycephalus scaber; Haemul-
idae: Pomadasys furcatus (Bl. & Schn.); Psettodidae: Psettodes erumei (Shneider);
Terapontidae: Terapon jarbua (Forskal), T. puta (Cuv. et Val.); Polynemidae: Poly-
dactylus sextarius (Bl. & Schn.) (3)
Localities: AS, BOB
Distribution: Widely distributed, refer the map no. adapted from Roumbedakis et al.
(2014)
References: 1. Srivastava (1936b), 2 Hafeezullah (1971a), 3. Mani & Anuprasanna
(2010)
Description: Body elongate, with rounded extremities, 1724–4160 long, 496–1576

wide. Oral sucker terminal, subspherical, 112–273 by 110–276. Ventral sucker larger
than oral sucker, spherical, pre-equatorial 192–331 by 234–70. Prepharynx small.
Pharynx oval. Oesophagus fairly long. Caeca simple, extend to posterior end of body.
Genital pore submedian, bifurcal. Testes nine in number, postequatorial, spherical or
subspherical, in two longitudinal rows of 5 and 4, respectively. Cirrus-sac elongated,
extends from level of intestinal bifurcation to left of ventral sucker to its posterior
end. Seminal vesicle tubular. Pars prostatica narrow. Ejaculatory duct long. Space
around anterior portion of vesicular seminalis and pars prostatica surrounded by
many prostate gland cells. Ovary lobed, submedian, pretesticular. Seminal recepta-
cle pear-shaped, anterior to ovary, eggs oval, operculated with long unipolar fila-
ments, 54–58 × 27–31. Vitellarium follicular, fields reach from well into forebody
to posterior extremity.
Remarks: Helicometrina nimia has been recorded from a wide range of hosts in
Atlantic, Pacific and Indian oceans. Over 50 species of fishes belonging to diverse
families have been recorded as hosts for this species. From India, Srivastava (1936b)
described two species of Helicometrina, H. orientalis Srivastava, 1936, and H.
septorchis Srivastava, 1936. Subsequently, H. hexorchis Gupta & Sehgal, 1970;
H. scomberi Gupta & Jahan, 1975, H. unica Gupta & Puri, 1985 and H. indica
Dhanumkumari, 1999, were recorded from various species of marine fishes from
India. Deelman (1960) found numerous intraspecific variations in specimens of H.
nimia and synonymized H. orientalis with H. nimia. Later, Hafeezullah (1971c) also
found variations in the number of testes, position of genital pore and posterior extent
of cirrus-sac in the specimens of H. nimia collected from marine fish of Bay of Ben-
gal and Arabian Sea and synonymized H. septorchis with H. nimia. However, several
authors have expressed the view that the differences in the number of testes may be
of taxonomic significance (Mittal & Pande, 2007; Roumbedakis et al. 2014). In view
of this controversy which needs to be resolved, all the Indian species of the genus
are listed below together with a brief account of their diagnostic features as given in
the original publication.
Using molecular and morphological criteria, Oliva et al. (2015) demonstrated that
in the coastal waters of Chile, Helicometrina is represented by two species, for one
of which they retained the name H. nimia and for the other they coined the name
H. labrisomi Oliva, Valdivia, Chavez, Molina & Cárdenas, 2015. It is likely that the
Indian forms of this genus are not conspecific with H. nimia from its original locality
(the Gulf of Mexico) and that there are several cryptic or unrecognized species in
Indian waters.
Helicometrina chauhani Mittal & Pande, 2007
Host: Sparidae: Chrysophrys sarba (Cuv, & Val.)
Locality: PR, BOB
Reference: Mittal & Pande (2007)
Diagnostic features: Eight testes, a cirrus-sac which extends to the mid-level of the
ventral sucker and a vitellarium which extends from bifurcal level to the posterior
end of the body.
Helicometrina hexorchis Gupta & Sehgal, 1970
[Synonym of H. nimia Linton, 1910]
Host: Sciaenidae: Otolithus ruber (Bl. & Schn.), Sillago sihama (Forskål):
Locality: Calicut, AS
No: 4
Reference: Gupta, N.K. & Sehgal (1970b)
Diagnostic features: Six testes, in two rows, a cirrus-sac which extends to the mid-
level of the ventral sucker and a vitellarium which extend from bifurcal level to the
posterior end of the body.
Helicometrina indica Dhanumkumari, 1999
Hosts: Bagridae: Mystus cavasius (Hamilton); Terapontidae: Terapon jarbua
(Forsskal)
Locality: Bheemunipatnam estuary
Reference: Dhanumlumari (1999)
Helicometrina orientalis Srivastava, 1936
Host: Scombridae: Rastrelliger kanagurta (Cuvier) (1); Sciaenidae: Sciaena miles
(Cuvier) (2)
References: 1. Srivastava (1936c); 2. Gupta, N.K. & Sayal (1979)
Diagnostic features: Nine testes, a vitellarium which extends from the bifurcal level
to the posterior end of the body and a cirrus-sac which extends posteriorly to mid-
dorsal region of ventral sucker.
Helicometrina quadrorchis Manter & Pritchard, 1960
Host: Sciaenidae: Otolithus ruber (Bl. & Schn.)
Locality: Calicut, AS
Number: 1
Reference: Gupta, N.K. & Sehgal (1970b)
Diagnostic features: Four testes, a clavate cirrus-sac, a submedian genital pore at the
level of the posterior end of the pharynx, tri-lobed ovary and a vitellarium which
reach from the anterior end of pharynx to the posterior end of the body.
Helicometrina scomberi Gupta & Jahan, 1975.

Scombridae: Rastrelliger kanagurta (Cuvier)
Locality: PR, BOB
Number: 1
Reference: Gupta, V & Jahan (1975)
Diagnostic features: Nine testes, in two rows and a vitellarium which extends from
the anterior margin of the ventral sucker to the posterior end of the body.
Helicometrina septorchis Srivastava, 1936
Sciaenidae: Otoliothus ruber (Bl. & Schn.); Sillaginidae: Sillago sihama (Forskål):
Locality: 1. PR, BOB; 2. Calicut, AS
Reference: 1. Srivastava (1936b); 2. Gupta, N.K. & Sehgal (1970b)
Number: 1
Remarks: Synonymized with H. nimia by Deelmann (1960)
Diagnostic features: Seven testes in two longitudinal rows, a cirrus-sac which extends
to the anterior margin of the ventral sucker, a tri-lobed ovary and a vitellarium which
extends from the level of the pharynx to the posterior end of the body.
Helicometrina unica Gupta & Puri 1985

Host: Sparidae: Rhabdosargus sarba (Forskål)\
Locality: PR, BOB
Reference: Gupta, V. & Puri (1985)
Diagnostic features: Seven testes, four in left row and three in right row, a just
post-bifurcal submedian or median genital pore, a cirrus-sac which extends to the
mid-region of the ventral sucker, a bipartite seminal vesicle and a vitellarium which
extends from the bifurcal region to the posterior end of the body.
Family Opistholebetidae Nicoll, 1915
Opistholebetidae comprises a small group of digeneans closely related to opecoelids.

They show some degree of specificity towards tetraodontid fishes. A combination
of four features distinguishes opistholebetids from opecoelids and from other dige-
neans: the usual position of the ventral sucker in the posterior half of the body often
at the posterior end, the presence of a post-oral ring, the presence of pigment granules
and the occurrence in tetraodontid and diodontid fishes. Molecular studies indicate
that this taxon is embedded in the Opecoelidae and is considered a subfamily (pro
tem) by some workers (e.g. Bray et al. 2016; Martin et al. 2017).
Diagnosis: Body elongate to pyriform. Tegument smooth. Parenchyma usually with

scattered pigment granules. Oral sucker subglobular. Ventral sucker usually in pos-
terior half of body sometimes at posterior end of body. Post-oral ring muscular,
prominent. Prepharynx short. Pharynx well developed. Caeca blind extend close
to posterior end of body. Testes two, globular, symmetrical or tandem or oblique.
Cirrus-sac encloses seminal vesicle, pars prostatica and ejaculatory duct. Genital
Family Opistholebetidae Nicoll, 1915 487
pore median, in forebody just near intestinal bifurcation. Ovary spherical, pretes-
ticular. Canalicular seminal receptacle present. Uterus intercaecal. Eggs yellowish,
operculate. Vitelline follicles profuse, fill body from pharynx to posterior end of
body. Excretory vesicle I-shaped.
Type genus Opistholebes Nicoll, 1915
The family comprises only three genera: Opistholebes Nicoll, 1915, Maculifer
Nicoll, 1915 and Heterolebes Ozaki, 1935. Genus Alloheterolebes Hafeezullah, 1971
proposed as a replacement name for Pseudoheterolebes Gupta, 1968 which was
preoccupied by Pseudoheterolebes Yamaguti, 1959 was regarded as a synonym of
Maculifer Nicoll, 1915 by Cribb (2005).
Key to genera:
1. Ventral sucker at posterior end of body …………………..…Opistholebes Nicoll,
1915
Ventral sucker at some distance from posterior end of body ………………… 2
2. Body elongate, testes tandem ………………….…. Maculifer Nicoll, 1915
Body rounded, testes oblique or symmetrical…………………..Heterolebes
Ozaki, 1935
Genus Heterolebes Ozaki, 1935
Diagnosis: Body rounded. Ventral sucker distinctly separated from posterior end of
body, entirely restricted to posterior half of body. Gonads posterior to ventral sucker.
Heterolebes buckleyi (Gupta, 1968) Cribb, 2005 (Fig. 15.56)

[Syn. Opistholebes buckleyi Gupta, 1968]
Host: Tetraodontidae: Tetraodon viridipunctatus(Gunther)
Locality: BM, AS
Number: 4
Reference: Gupta, A.N. (1968b)
Description: Body pear-shaped, broadly rounded at anterior end, flat at posterior

edge, 735–975 long, 555–875 broad. Oral sucker 116–120 long, 125–140 wide; post-
oral ring 75 thick, 185–190 wide. Ventral sucker subterminal, larger than oral sucker,
Fig. 15.56 Heterolebes

buckleyi (after Gupta, 1968)
180–185 × 187–190 in size. Sucker ratio 1: 1.5. Disc, muscular ridge or any special
musculature around ventral sucker absent. Prepharynx absent. Pharynx muscular.
Oesophagus absent. Caeca simple, margins roughly indented, extend to posterior
end. Testes rounded, symmetrical, close to each other, dorsal to ventral sucker, pos-
terior margin may coincide with that of ventral sucker. Cirrus-sac well developed,
claviform, extends from bifurcal point to ventral sucker, encloses small internal sem-
inal vesicle pars prostatica and ejaculatory duct. Genital pore ventral, bifurcal. Ovary
ovoid, submedian, overlaps ventral sucker. Seminal receptacle present. Uterus inter-
caecal. Metraterm absent. Eggs ovoid, 30 × 60 in size. Vitellarium with large follicles
extending on each side from level of oral sucker to posterior end, and meeting only in
bifurcal region. Pigment spots interspersed with vitelline follicles, mostly in lateral
regions. Excretory vesicle transversely elongated.
Remarks: This species was initially included in the genus Opistholebes as O. buckleyi
Gupta, 1968. Cribb (2005) included it in the genus Heterolebes as H. buckleyi (Gupta,
1968) because of the non-terminal position of the ventral sucker and the position of
the testis and ovary postero-dorsal to the ventral sucker.
Heterolebes spindalis Gupta, 1968 (Fig. 15.57)
Host: Tetraodontidae, Tetraodon immaculatus (Cantor)
Locality: PR, BOB
Number: 4
Ref: Gupta, A.N. (1968c)
Description: Body 1448–1701 in length, 726–882 in maximum breadth. Oral sucker
175–231 in diameter, immediately followed by a post-oral muscular ring, oral sucker
is surrounded by a few cephalic glands laterally. Ventral sucker transversely elon-
gated, in posterior half of body, 278– 325 × 375–419. Sucker ratio 1: 3. Prephar-
Fig. 15.57 Heterolebes

spindalis (after Gupta, 1968,
Acta Parasitol. 15, p. 362)
ynx short, thin-walled. Pharynx muscular, elongated. Oesophagus short. Caeca arch
around ventral sucker, converge towards posterior end. Testes two, transversely ovoid,
symmetrical, close to posterior margin of ventral sucker. Cirrus-sac club-shaped
extends obliquely from bifurcal point almost to ventral sucker, encloses saccular
seminal vesicle, pars prostatica and ejaculatory duct. Genital pore submedian, post-
bifurcal. Ovary between testis and ventral sucker, overlaps posterior part of ventral
sucker. Seminal receptacle small, oval. Uterus short, runs anteriorly dextrally to
cirrus-sac. Eggs few, yellowish, non-operculate, 28–31 × 49–61 in size. Vitellar-
ium with numerous follicles, extend from level of pharynx to posterior end of body;
confluent in forebody and in posterior region of the post-testicular zone.
Remarks: H spindalis differs from H. buckleyi in the oval body shape, the much
larger ventral sucker and in the extent of the vitellarium into the post-testicular zone.
Genus Maculifer Nicoll, 1915
[Syn: Pseudoheterolebes Gupta, 1968 nec Yamaguti, 1959; Alloheterolebes
Hafeezullah, 1971]
Generic diagnosis: Body elongate. Ventral sucker separated from posterior end of
body, at middle or anterior half of body. Gonads posterior to ventral sucker.
Type species: M. subequiporus Nicoll, 1915.
Maculifer indicus (Gupta, 1968) Cribb, 2005 (Fig. 15.58)
[Syn: Pseudoheterolebes indicus Gupta, 1968; Alloheterolebes indicus (Gupta,
1968) Hafezullah, 1971; Diplobulbus vitellosus Bilqees, 1972; Maculifer vitellosus
(Bilqees, 1972) Cribb, 2005]
Fig. 15.58 Maculifer

indicus (after Gupta, 1968)
Host: Tetraodontidae: Tetraodon viridipunctatus (Guinther).

Number of specimens: four mature, three immature
Ref. Gupta, A, N. (1968b)
Description: Body spindle-shaped, 3900–4000 long and 1065–1710 in maximum
breadth. Tegument thick, smooth. Oral sucker terminal, 450–480 in diameter. Ventral
sucker lies ventrally at junction of anterior two-third and posterior one-third of body,
660 by 405 in size. Prepharynx, pharynx present. Caeca fairly wide, extend posteri-
orly as far as level of ventral sucker. Testes rounded or ovoid, symmetrically, closely
posterior to ventral sucker or slightly overlapping it. Cirrus-sac club-shaped, lies ante-
rior to ventral sucker, major part occupied by bipartite seminal vesicle. Ejaculatory
duct short. Genital pore median, post-bifurcal. Ovary oval, dorsal to ventral sucker,
at level of testes. Seminal receptacle saccular, closely preovarian. Uterus ascends
anteriorly to open through genital pore. Metraterm not clearly differentiated. Eggs
60 × 50 in size. Vitellarium consist of small follicles, extend from bifurcation up to
posterior end.
Remarks: Maculifer indicus is the only species of the genus reported from India.
Maculifer spiralis Soota, Srivastava & Ghosh, 1970
Host: Marine fish
Locality: Nicobar island
Remarks: This species was considered a synonym of the opecoelid Hamacreadium
mutabile Linton, 1910, by Hafeezullah & Dutta (1980), Bray & Cribb (1989), Cribb
(2005) and a synonym of H. hainanense Shen, 1990, by Martin et al. (2017).
Genus Opistholebes Nicoll, 1915
Diagnosis: Ventral sucker at posterior end of body. Gonads anterior to ventral sucker.
Type species: O. amplicoelus Nicoll, 1915
Opistholebes indicus Gupta, 1968 (Fig. 15.59)
Host: Tetraodontidae: Tetraodon immaculatus (Cantor)
Locality: MS, BOB
No: 30
Ref: Gupta, A.N. (1968b)
Description: Body pear-shaped, anterior end broadly pointed, posterior end with
dome-like bulge or truncated; 1210–1950 long, 630–1270 wide at ovarian level.
Oral sucker 75–215 × 150–315 in diameter, with a distinct post-oral ring. Ventral
sucker subterminal, extends into dome-shaped bulge, 135–285 × 180–375, posterior
margin separated from margin of body by muscular ridge. Pharynx well developed,
with wavy anterior margin. Caeca broad, arcuate, reach anterior margin or equator of
ventral sucker. Testes two, inner margin flattened, globular or subglobular, symmet-
rical, intercaecal, postovarian. Cirrus-sac claviform. Internal seminal vesicle large,
coiled. Pars prostatica and prostate gland cells present. Ejaculatory duct small. Geni-
tal pore median, distinct, just post-bifurcal. Ovary oval, slightly submedian, anterior
Fig. 15.59 Opistholebes

indicus (after Gupta, 1968,
Acta Parasitol. 15, p. 356)
to right testis. Seminal receptacle larger than ovary. Uterus short, sinistral to ovary
and cirrus-sac, intercaecal. Eggs 60 × 45. Vitelline follicles rounded to oval, mostly
extracaecal, extend from pharynx to ventral sucker, some follicles overlap caeca
while others enter into intercaecal field.
Remarks: Differs from the type species in having a small oesophagus, in not having
caeca diverging horizontally after emergence from the intestinal bifurcation; also in
its shape and size of testes and ovary.
Opistholebes tetradontis Gupta, 1968 (Fig. 15.60)

Host: Tetradontidae, Tetraodon immaculatus (Cantor)
Locality: MS, BOB
No: 30
Ref: Gupta, A.N. (1968b)
Description: Body more or less oval, anterior extremity rounded, posterior extrem-
ity bulged to accommodate ventral sucker, 890–937 long, 574–688 wide in middle
region. Oral sucker subterminal, 118–130 long, 135–156 wide; post-oral ring present.
Bunch of unicellular glands on each side of oral sucker. Ventral sucker subtermi-
nal, mostly occupies posterior bulge, larger than oral sucker, 153–162 × 197–220
in size. Sucker ratio 1:1.7. Prepharynx small, feebly muscular. Pharynx well devel-
oped, broader than long, anterior margin wavy. Oesophagus very short. Caeca sim-
ple, proximal portion widely separated from body margin, distal portion close to it;
extend more or less to equatorial level of testes. Testes symmetrical, anterior to or
may overlap ventral sucker. Cirrus-sac claviform, extends from intestinal bifurcation
to anterior to ovary. Internal seminal vesicle long, coiled, tubular. Prostate glands
present. Ejaculatory duct short. Ovary oval to elliptical, submedian, anterior to left
Fig. 15.60 Opistholebes

tetradontis (after Gupta,
1968)
testis. Seminal receptacle spherical, immediately preovarian, close to its left margin.
Vitelline follicles in lateral fields, mostly extracaecal, extend from pharyngeal level to
posterior margin of testes. Genital pore median, immediately post-bifurcal. Uterus
intercaecal, filling entire intercaecal space on right side of ovary and cirrus-sac.
Metraterm absent. Eggs non-operculate, 21–33 × 49–61 in size. Excretory vesicle
tubular.
Remarks: This species differs from O. indicus in having a smaller prepharynx, in

the more anterior location of the genital pore and in the presence of unicellular gland
cells on either side of the oral sucker.
Chapter 16
Superfamily Monorchioidea Odhner,
1911
Family Monorchiidae Odhner 1911
Members of the Monorchiidae are parasites in the digestive tract of marine and
freshwater fishes. They are characterized by the possession of a spiny tegument,
complex terminal genitalia armed with spines, restricted vitelline follicles and well-
developed uterine coils. According to the recent review provided by Madhavi (2008),
the family contains 40 nominal genera distributed over 6 subfamilies.
Molecular results indicate that the Monorchiidae, along with the freshwater family
Lissorchiidae, are best placed in a superfamily, the Monorchioidea Odhner 1911 and
a suborder Monorchiata, as a sister taxon of the Xiphidiata (Olson et al., 2003;
Littlewood et al. 2015). The relationships between the various genera and the supra-
generic subfamilies have also not been well established, although molecular results
are showing an emerging pattern (Atopkin et al. 2017). Manter and Pritchard (1961)
and Mamaev (1968) recognized six subfamilies under the family. However, Yamaguti
(1971) split the family into 12 subfamilies. The one proposed by Madhavi (2008)
with six subfamilies is adopted in the present account.
The most important feature of the family is the presence of a terminal organ, a
sac-like structure, that is actually a continuation of the uterus. The terminal organ
may be saccular or bipartite, spined wholly or partially or only in the anterior part.
Details of the structure of the terminal genitalia, number of testes, the distribution of
the vitelline follicles and the position of the genital pore are useful for differentiation
of genera in this family.
Diagnosis: Body small or medium sized, oval to elongate or long and slender. Tegu-
ment spined. Oral sucker rounded or funnel-shaped, rarely with ring of spines. Ventral
sucker usually pre-equatorial. Prepharynx present or absent. Pharynx well developed.
Oesophagus of variable length. Caeca long. Testis single or double, symmetrical or
tandem, usually in hindbody. Cirrus-sac well developed, encloses simple or bipartite
or sinuous seminal vesicle, pars prostatica, numerous prostatic cells and spined cirrus.
Genital atrium well developed, may be spined. Genital pore median or submedian in
© Crown 2018 493
https://doi.org/10.1007/978-94-024-1535-3_16
494 16 Superfamily Monorchioidea Odhner, 1911
fore or hindbody. Ovary spherical, prestesticular. Mehlis’gland and Laurer’s canal

present. Seminal receptacle uterine. Uterus usually occupies entire hindbody opens
through tubular metraterm into distal or middle part of terminal organ. Terminal
organ simple or bipartite entirely spined or only in anterior part. Eggs oval, opercu-
late, may have polar filaments. Vitellarium usually forms symmetrical bunches or
follicular or tubular acini in fore or hindbody. Excretory vesicle saccular or V- or
Y-shaped. In gastrointestinal tract or fishes. Type genus: Monorchis Looss, 1902.
About seventy species of monorchiids belonging to 42 genera have been reported
from Indian marine fishes (Madhavi, 2011). The descriptions given for some of the
species are incomplete lacking in details of terminal genitalia. Many of them were
based on a single specimen or poorly prepared forms. It was a difficult task trying to
arrange them in an order and differentiate the various species especially those under
the genera Lasiotocus Odhner, 1911, and Opithomonorcheides Parukhin, 1966. A
few species whose descriptions were inadequate or for which the literature was not
available were not included in this account.
Key to subfamilies
1. Eggs without a polar filament……………………………………………2
Eggs with a long polar filament………………….……………..Hurleytrematinae
Yamaguti, 1958
2. Body small, oval, vitelline follicles limited…………………………………3
Body long, slender, vitelline follicles extensive……..….Opisthomonorcheidinae
Yamaguti, 1971
3. Genital pore in forebody………………………………………Monorchiinae
Odhner, 1911
Genital pore in hindbody………………………………….Opistomonorchiinae
Yamaguti, 1970
Subfamily Monorchiinae Odhner, 1911
[Syn: Ancylocoelinae Skrjabin & Koval, 1957, Lasiotocinae Yamaguti, 1958;
Postmonorcheidinae Yamaguti, 1958; Pseudoproctotrematinae Yamaguti, 1958;
Telolecithinae Yamaguti, 1958; Neopisthomonorchiinae Varma, 1982]
Diagnosis: Body small, oval or elongate. Tegument spined. Oral sucker rounded
or funnel-shaped, rarely with ring of spines. Caeca long or short. Testis single or
double, symmetrical or oblique or tandem when double, usually in hindbody. Cirrus-
sac extends into hindbody or not, encloses oval or sinuous seminal vesicle, prostatic
complex and spined cirrus. Genital atrium spined or not. Genital pore median, post-
bifurcal. Ovary submedian, smooth or lobed. Terminal organ oval or bipartite, usually
spined. Uterine coils extensive, may or may not extend posteriorly beyond testis.
Eggs oval. Vitelline follicles in lateral fields of forebody or hindbody. Type genus
Monorchis Looss, 1902.
Key to genera
1. Testis single…………………………………………………………2
Testes double…………………………………………………………… 10
Family Monorchiidae Odhner 1911 495
2. Oral sucker with a ring of spines……….……………..………… 3

Oral sucker without a ring of spines ………………… 5
3. Cirrus-sac long, extends to level of testis, vitelline follicles at level of ventral
sucker…………………………Huridostomum Mamaev, 1970
Cirrus-sac claviform, vitelline follicles at level of ovario-testicular
zone…………………. 4
4. Terminal organ unipartite……………Ametrodaptes Bravo-Hollis, 1956
Terminal organ bipartite………………Pseudametrodaptes Trivenilakshmi &
Madhavi, 2008
5. Testis near posterior extremity of body, uterus not extending posterior to
testis……………………………….Leiomonorchis Mamaev, 1970
Testis in middle third of body, uterus filling post-testicular
space …..………………………6
6. Genital atrium spined …………………………Genolopa Linton 1910
Genital atrium not spined………………………………………7
7. Terminal organ unipartite ……………………Proctotrema Odhner, 1911
Terminal organ bipartite …………………..……………………8
8. Genital atrium with spiny diverticulum …………………… Proctotrematoides
Yamaguti, 1938
Genital atrium without spiny diverticulum ……..……………….………9
9. Vitellarium follicles in ovario-testicular zone or at level of ventral sucker
…………………………Lasiotocus Looss in Odhner 1911
Vitellarium follicles distributed in lateral fields of forebody…………….
Monorchis Looss, 1902
10. Caeca short, vitellarium follicles on either side of pharynx
……………………………Allobacciger Hafeezullah & Siddiqi, 1970
Caeca long, vitellarium not reaching pharynx ………………… 11
11. Testes obliquely tandem …………..………….....…………… 12
Testes symmetrical ………………………………….……… 13
12. Vitellarium forming two compact masses at level of ovary……...…Timonia
Bartoli & Prevot, 1966
Vitellarium follicular, at level of ventral sucker….………….………Neotimonia
Madhavi, 2008
13. Uterus not extending posteriorly to testes…..……..……….….Monorcheides
Odhner, 1905
Uterus extending well posteriorly to testes………………… Paramonorcheides
Yamaguti, 1938
Genus Allobacciger Hafeezullah & Siddiqi, 1970

[Syn: Pseudomonorcheides Wang, 1982]
Diagnosis: Body small, plump. Oral sucker subterminal, large. Ventral sucker small,
in mid-region of body. Pharynx small. Oesophagus short. Intestinal bifurcation in
mid-forebody. Caeca short, inflated, end in anterior half of body. Testes two, symmet-
rical, immediately posterior to caecal ends. Cirrus-sac claviform contains saccular
seminal vesicle, prostatic complex and armed cirrus. Genital atrium small. Genital
pore just post-bifurcal. Ovary deeply lobed, dextral to ventral sucker. Uterus fills
most of post-testicular space and region to left of cirrus-sac. Terminal organ saccu-
lar. Vitelline follicles form symmetrical bunches on either side of pharynx. Excretory
vesicle V-shaped; pore terminal. In intestine of marine teleost fishes; off China, India,
Japan. Type species: A. macrorchis Hafeezullah & Siddiqi, 1970.
Allobacciger macrorchis Hafeezullah & Siddiqi, 1970 (Fig. 16.1)
[Syn: Monorcheides macrorchis Machida, 2005]
Host: Nemipteridae: Scolopsis vosmeri (Bloch)
Locality: Calicut, VSK (AS, BOB)
References: Hafeezullah & Siddiqi (1970b); Machida (2005)
Description: Body 750–870 long 440–590 wide, ovate. Tegument thin. Oral sucker
96–125 in diameter. Ventral sucker 68–75 in diameter roughly at midbody. Sucker
ratio 1: 0.58–0.73. Prepharynx present. Pharynx oval. Oesophagus short. Caeca short
reach testicular region. Testes oval, lateral, symmetrical, equatorial. Cirrus-sac pyri-
form, dorsal to ventral sucker encloses unipartite seminal vesicle, short pars prostat-
ica, diffused prostatic cells and long muscular protrusible cirrus. Ovary three-lobed,
lobes distinct, large, spherical, anterodorsal to right testis. Seminal receptacle present.
Uterus voluminous, fills most of hindbody, reaching anteriorly to level of oesophagus.
Metraterm short, muscular. Terminal organ saccular filled with sperm. Eggs 15–22
by 12. Vitellarium in two lateral clusters of large follicles in forebody, extracaecal,
reach level of posterior border of oral sucker.
Remarks: The genus Allobacciger erected by Hafeezullah and Siddiqi (1970) with
A. macrorchis Hafeezullah and Siddiqi, 1970 as the type species, was originally
placed in the family Fellodistomidae. Machida and Uchida (2001) described the
Fig. 16.1 Allobacciger

macrorchis (after
Hafeezullah & Siddiqi, 1970)
second species of Allobacciger, A. centropygis, from Centropyge heraldi from Japan

and transferred Allobacciger from the family Fellodistomidae to Monorchiidae.
Madhavi (2008) included the genus in this family and considered Pseudomonorchei-
des Wang, 1982 as a synonym of this genus. However, Machida (2005) considered
the genus Allobacciger as a synonym of Monorcheides Odhner, 1905 and trans-
ferred A. macrorchis to this genus as M. macrorchis (Hafeezullah and Siddiqi,
1970b). Madhavi (2008) retained Allobacciger as a valid genus separating it from
Monorcheides by the position of testes closely posterior to the ventral sucker, the
uterine coils extending posterior to the testis, and the I-shaped excretory vesicle.
McNamara et al. (2014) used A. centropygis sequences in their study of cryptic
speciation in monorchiids.
Genus Ametrodaptes Bravo-Hollis, 1956

[Syn. Neopisthomonorchis Varma, 1982]
Diagnosis: Body elongate. Oral sucker large, armed with alternating rows of spines
on dorsal side. Prepharynx as long as oesophagus. Pharynx well developed. Intestinal
bifurcation in posterior forebody. Caeca long. Testis single, elliptical in posterior third
of body. Cirrus-sac claviform, contains saccular seminal vesicle, prostate complex
and spined cirrus. Genital atrium with diverticulum, bears numerous spines. Genital
pore immediately anterior to ventral sucker. Ovary pretesticular. Uterus occupies
greater part of hindbody. Metraterm with distinct sphincter, opens into proximal
portion of terminal organ. Terminal organ bipartite, proximal part spinous. Vitelline
follicles in lateral fields in gonadal region. Excretory bladder tubular, reaches ovary.
In marine fishes. Type species: A. mexicanus Bravo-Hollis, 1956.
Genus Ametrodaptes is characterized by the presence of a ring of spines surrounding

the oral aperture and a bipartite terminal organ. Trivenilakshmi and Madhavi (2008)
regarded the genus Neopisthomonorchis Varma, 1982 as a synonym of Ametrodaptes
and the single species N. pristopomatis Varma, 1982 became A. pristopomais (Varma,
1982). Four more species of Ametrodaptes have been reported from marine fish from
India, but a separate genus Pseudametrodaptes Triveni Lakshmi and Madhavi, 2008,
was erected to include these four species. The latter genus was differentiated from
Ametrodaptes in the absence of spines in the genital atrium and the unipartite nature
of the terminal organ with the metraterm joining its distal end (Trivenilakshmi &
Madhavi, 2008). As a result of these adjustments, the genus Ametrodaptes is left
with only one Indian species A. pristipomatis (Varma, 1982).
Ametrodaptes pristipomatis (Varma, 1982) Trivenilakshmi & Madhavi, 2008
[Syn: Neopisthomonorchis pristipomatis Varma, 1982]
Host: Haemulidae: Pomadasys furcatus (Bl. & Schn.)
Locality: Kerala coast, AS
No. 6
References: Varma (1982), Trivenilakshmi & Madhavi (2008)
Description: Body elongated 1200–3106 long, 370–475 wide, heavily spined. Oral
sucker funnel-shaped, 192–195 long, 210–235 wide. Circumoral spines short, stout,
almond-shaped, arranged in three uninterrupted alternating rows dorsally and two

medially interrupted alternating rows ventrally, with 17–19 spines in each dorsal row
and 11–13 spines in each ventral row. Ventral sucker spherical, pre-equatorial, smaller
than oral sucker, 173–195 long, 180–191 wide. Sucker ratio 1: 0.84. Prepharynx
tubular. Pharynx oval. Oesophagus short. Caeca extend to posterior end of body.
Genital pore submedian, posterior to ventral sucker. Testis postequatorial, ovoid,
in posterior half of hindbody. Cirrus-sac large, extends obliquely from genital pore
to anterior edge of ovary. Seminal vesicle oblong. Pars prostatica surrounded by
numerous prostatic cells. Cirrus large, heavily spined, spines of two types, short and
thorn-like on entire surface, long and needle-like on lateral margin. Genital atrium
short, spinous, atrial spines appear as continuation of spines of cirrus and terminal
organ. Ovary pretesticular, ovoid. Uterus occupies all available space between cirrus-
sac and hind end of body. Terminal organ well developed, consists of thin-walled
unspined basal chamber and muscular heavily spined distal chamber. Eggs 16–19 ×
8–11. Vitellarium follicular in two lateral groups of 6–8 widely separated follicles
on each side, extend from just anterior to ovary to just poserior to testis.
Remarks: This monorchiid was included by Varma (1982) in a new genus
Neopisthomonorchis Varma, 1982, and a new subfamily Neopisthomonorchiinae
Varma, 1982. Trivenilakshmi & Madhavi (2008) noted the close similarity of the
genus to Ametrodaptes Bravo-Hollis, 1956, the only difference being the loca-
tion of the genital pore posterior to the ventral sucker in Neopisthomomonorchis.
Pending verification of the position of the genital pore, Trivenilakshmi & Madhavi
(2008) synonymized Neopisthomonorchis with Ametrodaptes and the subfamily Neo-
pithomonorchiinae with Monorchiinae. A. pristipomatis is the only Indian species
under the genus Ametrodaptes.
Genus Genolopa Linton 1910
This genus closely resembles the genera Proctotrema and Lasiotocus but differs from
both of these genera mainly in the presence of spines in genital atrium.
Diagnosis: Body elongate, spined. Oral sucker subterminal, ventral sucker pre-
equatorial. Prepharynx short. Oesophagus long. Intestinal bifurcation in forebody.
Caeca long. Testis submedian, pre-equatorial. Cirrus-sac large, extends well into
hindbody, encloses oval seminal vesicle, numerous prostatic cells and eversible
spined cirrus. Genital atrium spined. Genital pore median, anterior to ventral sucker.
Ovary compact or lobed, pretesticular. Uterus occupies entire post-testicular region.
Metraterm opens into proximal end of terminal organ. Terminal organ club-shaped,
armed with prominent spines. Vitelline follicles form bunches in ovarian zone. Type
species: G. ampullacea Linton, 1910.
Genolopa ampullacea Linton, 1910 (Fig. 16.2)
Host: Marine fish
Locality: DC, BOB
Reference: Dutta & Manna (1998)
Distribution: Florida, Puerto Rico, Venezuela, Bahamas, Curaçao, Jamaica, Gulf of
Mexico, Rio de Janeiro.
Fig. 16.2 Genolopa

ampullacea (after Linton,
1910)
Remarks: This is the type species of the genus originally described from Haemulon
sciurus off the USA and reported in other species of fish from India, Puerto Rico,
Venezuela, Bahamas, Curaçao, Jamaica and Brazil. From India, it was reported by
Dutta & Manna (1999) from a marine fish at DC, BOB. This species is characterized
by triple spination in the terminal genitalia. The genital atrium is armed with a circle of
conspicuous very long pointed blade-like spines. The cirrus spines are thorn-shaped;
the metraterm spines are very fine and needle-like.
Genolopa bychowskii Zhukov, 1977
Host: Carangidae: Apolectus niger (Bloch)
Locality: DC, BOB
Reference: Hafeezullah & Dutta (1998)
Description: Body 940–1220 by 160–230, fusiform. Oral sucker smaller than ven-
tral sucker. Ventral sucker pre-equatorial. Prepharynx present. Pharynx well devel-
oped. Oesophagus very long. Caeca extend to posterior end of body. Testis single,
ovate, in posterior part of body. Cirrus-sac very long, extends well posterior to ventral
sucker. Genital pore immediately anterior to ventral sucker. Genital atrium spined.
Ovary immediately anterior to testis, tri-lobed. Uterus coiled, largely between ovary
and ventral sucker. Eggs 14 × 9.
Remarks: This species is characterized by the position of gonads in the posterior

part of body, the very long oesophagus and the very long cirrus-sac extending well
posterior to the ventral sucker.
Genolopa trifolifer Nicoll, 1915

Host: Haemulidae: Pomadasys hasta (Bloch)
Locality: BOM, AS
Distribution: Queensland, Australia
Reference: Hafeezullah (1971b, 1984).
Genus Huridostomum Mamaev, 1970
Diagnosis: Body elongate. Oral sucker cup-shaped with ring of enlarged spines,
Ventral sucker poorly developed. Pharynx small. Oesophagus long. Caeca long. Testis
single, in posterior quarter of body. Cirrus-sac long, sinuous, extends well posterior to
ventral sucker almost to level of testis, encloses a long seminal vesicle, few prostatic
cells and long, spined eversible cirrus. Genital pore median, in forebody. Ovary
elliptical, indented, pretesticular. Uterus confined to pretesticular region. Terminal
organ simple, sac-like, armed with spines. Vitelline follicles small, form a single
band in front of ventral sucker. Type species: H. formionis Mamaev, 1970.
Huridostomum formionis Mamaev, 1970 (Figs. 16.3 and 16.4)
Host: Carangidae: Parastromateus niger (Bloch)
Locality: VSK, BOB
Reference: Trivenilakshmi & Madhavi (2008)
Description: Body elliptical, broad at midbody, anterior part extensile, appears
as narrow neck, terminates in expanded funnel-like oral sucker region, poste-
rior region cylindrical, terminal part muscular appears as finger-like knob, length
1248–2320, maximum width 336–480. Entire tegument spinose. Oral sucker cup-
shaped, expanded anteriorly, 128–224 by 192–256 with vertical slit-like opening,
Fig. 16.3 Huridostomum

formionis Mamaev, 1970
(after Trivenilakshmi &
Madhavi 2008)
Fig. 16.4 Huridostomum

formionis Mamaev, 1970
Madhavi 2008)
anterior dorsal margin with group of 36–40 enlarged spines arranged in two rows.
Ventral sucker in second quarter of body, much smaller than oral sucker, 66–72
in diameter. Sucker ratio 1:0.35–0.38. Prepharynx small. Pharynx oval. Intestinal
bifurcation well anterior to ventral sucker. Caeca long, narrow, terminate just short
of posterior extremity. Genital pore median, anterior to ventral sucker. Testis single,
elliptical with irregular outline, situated in posterior half of hindbody. Cirrus-sac
long, narrow sinuous, extends well posterior to ventral sucker to almost mid-level
of testis. Seminal vesicle elliptical, prostate cells few, cirrus very long, eversible
armed with numerous triangular spines. Ovary entire or lobed, anterolateral to testis.
Seminal receptacle uterine. Uterus extensive occupies entire hindbody except for
knob-like portion at posterior region. Distal part of uterus modified into muscular
metraterm. Terminal organ lies dorsal to anterior part of cirrus-sac, small, oval to
elliptical, thick-walled, spined along its entire length. Metraterm joins distal part of
terminal organ. Vitelline follicles in two lateral groups in front of ventral sucker,
each group with numerous small follicles, commence from bifurcation point extend
slightly posterior to ventral sucker, confluent at level of bifurcation.
Remarks: Mamaev (1970) erected the genus Huridostomum with H. formionis
Mamaev, 1970 from Parastromateus niger from the Gulf of Tonkin as the type
species. Trivenilakshmi & Madhavi (2008) reported the species from the same host
from Bay of Bengal. H. formionis is characterized by the following combination of
characters: peculiar body shape, presence of enlarged spines along the anterodorsal
margin of oral sucker, very long cirrus-sac extending almost to the level of testis,
unipartite terminal organ with metraterm joining it distally and extensive uterine
coils. The species shows extensive intraspecific variations with regard to body shape
and extent of cirrus-sac.
Genus Lasiotocus Looss, 1907
[Syns. Ancylocoelium Nicoll, 1923; Chrisomon Manter & Pritchard, 1961]
The three genera of monorchiids Proctotrema Odhner, 1911, Lasiotocus Looss,
1907, and Chrisomon Manter and Pritchard, 1961, reflect many similarities in their
organization, and validity of these genera as distinct has been much discussed. How-
ever, the problem of the differentiation of Proctotrema and Lasiotocus was resolved
by Bartoli & Prevot (1966) who after examination of type material of Proctotrema
showed that the two genera can be differentiated by the structure of terminal geni-
talia, Proctotrema possessing a simple oval terminal organ with metraterm joining
it proximally while in Lasiotocus the terminal organ is bipartite and the metraterm
joins its middle part. Dove and Cribb (1998) suggested the separation of the gen-
era Lasiotocus and Chrisomon by the nature of vitellarium, the latter possessing a
vitellarium composed of clusters of tubular acini as against follicular vitellarium in
Lasiotocus. However, Bartoli and Bray (2004) showed that in the type species of the
genus Chrisomon, namely C. tropicus Manter & Pritchard, 1961, the vitellarium is
follicular and accordingly Madhavi (2008) erected a new genus Parachrisomon Mad-
havi, 2008, to include the remaining species of Chrisomon possessing a vitellarium
as clusters of tubular acini. The three genera can be differentiated as follows:
Proctotrema Odhner, 1911: Terminal organ unipartite, uterus joining its distal end.
Vitellarium follicular.
Lasiotocus Looss, 1907: Terminal organ bipartite, uterus joining its middle region.
Vitellarium follicular.
Parachrisomon Madhavi, 2008: Terminal organ as in Lasiotocus. Vitellarium in the
form of tubular acini.
Dove and Cribb (1998) defined the genus Lasiotocus as follows:
Diagnosis: Body plump to elongate, tegument spinose. Oral sucker subterminal.
Ventral sucker in middle third of body, small. Prepharynx present. Pharynx small.
Testis single in posterior half of body. Cirrus-sac encloses saccular seminal vesicle,
pars prostatica surrounded by prostatic cells and spiny eversible ejaculatory duct.
Genital atrium unspined. Genital pore anterior to ventral sucker. Ovary anterior to
testis. Uterine seminal receptacle present. Laurer’s canal present. Vitellarium com-
posed of two lateral bunches of follicles in region between ventral sucker and testis.
Terminal organ bipartite, proximal portion with few or no spines, distal portion spined
and receiving uterus partway along its length. Excretory vesicle saccular to tubular.
Type species: Lasiotocus mulli (Stossich, 1883) Looss, 1907
Twelve species of Lasiotocus have been recorded from marine fishes of Indian
waters, including two spp. transferred from the genus Chrisomon. Not all these
12 species appear to be valid. The status of these various species of Laiotocus is
discussed below:
Lasiotocus bengalensis Ahmad & Gupta, 1985, from Cephalopholis sonnerti
(Val.) from PR, BOB. This species resembles very closely L. hastai Madhavi, 1974.
The characters mentioned by Ahmad and Gupta (1985) for the differentiation of the
two species such as slight differences in the shape of the oral sucker, the extent of
the caeca, the extent of the cirrus-sac and the shape and location of the testis are all
variable features. L. bengalensis is considered here as a synonym of L. hastai.
Lasiotocus engraulisi Gupta & Gupta, 1976, from Stolephorus indicus (van Has-
selt) from PR, BOB: This species reflects many similarities to L. maculatus Madhavi,
1994, differing only in the morphometrics, L. engraulisi being slightly bigger. The
ovary is triangular in L. engraulisi rather than three-lobed. In the absence of any
major differences, it is regarded here as a synonym of L. maculatus. Lasiotocus
jagannathi Ahmad & Gupta, 1985, from Cybium guttatum (Bl. & Schn.), from PR,
BOB: The description based on a single specimen is inadequate. Some of the features
like extensive vitelline follicles, tubular genital atrium preclude its inclusion in the
genus Lasiotocus. More specimens are required to determine its exact taxonomic
position. Presently, it is considered as taxon inquirendum.
Lasiotocus rainai Gupta & Jain, 1985, from Serranus indicus (Cuv. et Val.) from
PR, BOB: The description of the species is based on only two specimens and lacks
many details needed for the identification of the species. From the figure, it appears
that the specimens were in a contracted state. The species is considered here as
species inquirenda.
Lasiotocus sunderbanensis (Dutta, Hafeezullah & Manna, 1994) Dove & Cribb,
1998, from Polynemus paradiseus and L. polynemi (Dutta, Hafeezullah & Manna,
1994) Dove & Cribb, 1998, from Eleutheronema tetradactylum were originally
included by Dutta, Hafeezullah & Manna (1994) under the genus Chrisomon as
C. sunderbanensis and C. polynemi. Dove & Cribb (1998) while separating the two
genera Chrisomon and Lasiotocus by the nature of vitelline follicles, the former
having a vitellarium in the form of follicles in contrast to tubular nature of the vitel-
larium in Chrisomon, included C. sunderbanensis as well as C. polynemi in the genus
Lasiotocus. But unlike all other species of Lasiotocus the uterus in these two species
does not extend posteriorly to the testis. The taxonomic status of these two species
remains to be determined.
After this analysis the genus Lasiotocusis left with six species recorded from
Indian marine fish.
Lasiotocus hastai Madhavi, 1974 (Fig. 16.5)

[Syn. Lasiotocus bengalensis Ahmad & Gupta, 1985 (new syn.)]
Hosts: Haemulidae: Pomadasys hasta (1); Serranidae: Cephalopholis sonnerati
(Val.) (2)
References: 1. Madhavi (1974a); 2. Ahmad & Gupta (1985); 3. Gupta & Jain (1992a).
Description: Body elongate, 2240–2990 long and 455–585 wide at level of ventral
sucker. Tegument spined. Oral sucker subterminal, hexagonal, 132–172 by 195–224.
Ventral sucker poorly developed 112–122 in diameter, at a distance of 820– 962 from
anterior end. Sucker ratio 1:0.52–0.60. Intestinal bifurcation nearer to oral sucker than
to ventral sucker. Caeca long terminate at about middle of post-testicular area. Testis
single, elliptical, in middle third of body. Genital pore median, just anterior to ventral
sucker. Cirrus-sac long, straight or arcuate, reaches level of ovary, encloses oval
seminal vesicle, long pars prostatica and cirrus armed with bristle-like spines. Genital
atrium large, unarmed, supported by muscle fibres. Ovary three-lobed, dextral, close
to testis. Uterus extends to near posterior extremity. Terminal organ well developed
about half as long as cirrus-sac, bipartite, posterior part unarmed, filled with granular
Fig. 16.5 Lasiotocus hastai

material, anterior part with thick muscular wall and much reduced lumen filled with
closely set bristle-like spines, its basal part modified into sphincter. Uterus joins
middle of anterior part of terminal organ. Eggs 20–23 × 12. Genital atrium spacious.
Gland cells occur over genital atrium and anterior part of terminal organ. Vitelline
follicles arranged in two groups of 5–7 follicles each, situated on either side of ovary.
Remarks: This species closely resembles L. maculatus but differs from it in the much
larger body size, more elongated body, different shape of oral sucker and the muscular
terminal organ provided with a sphincter. Gupta and Jain (1992a) considered this
species as a synonym of P. maculatus, but the two species are retained as valid based
on the differences mentioned above. Hafeezullah (1984) suggested that this species
might be a synonym of Genolopa trifolifer Nicoll, 1915, but examination of large
numbers of specimens available to the author did not reveal the presence of any spines
in the genital atrium, as such it is retained in the genus Lasiotocus as L. hastai.
Lasiotocus maculatus Madhavi, 1974 (Fig. 16.6)
[Syn. L. engraulisi Gupta & Gupta, 1976 (new syn.)]

Hosts: Haemulidae: Pomadasys maculatus (Bloch), P. argyreus (Val.), Rhoncis-
cus furcatus (Bl. & Schn.) (1); Engraulidae: Stolephorus indicus (van Hasselt) (2);
Cynoglossus dubius Day (3)
Fig. 16.6 Lasiotocus

maculatus (after Madhavi,
1974)
References: 1. Madhavi (1974a); 2. Gupta, S.P & Gupta, R.C (1976); 3. Gupta, V &
Jain (1992a);
Description: Body elongate, anterior end truncate, posterior end rounded, 922–1360
long and 272–384 wide. Tegument spined. Oral sucker funnel-shaped, 98–180 by
148–273. Ventral sucker small, 47–90 by 58–70 in size, situated in middle third of
body. Sucker ratio 1: 0.28–0.38. Intestinal bifurcation nearer to oral sucker than to
ventral sucker. Caeca long terminate at about middle of post-testicular area. Testis
single, elliptical, in middle third of body. Genital pore median, just anterior to ventral
sucker. Cirrus-sac long, arcuate, reaches level of testis encloses oval seminal vesicle,
long pars prostatica surrounded by numerous prostatic cells and cirrus armed with
bristle-like spines. Genital atrium spacious, unarmed. Ovary three-lobed, dextral,
close to testis. Seminal receptacle uterine. Uterus fills most of post-testicular space.
Terminal organ bipartite, muscular, surrounded by gland cells, situated to left of
ventral sucker, posterior vesicle unarmed, anterior part tubular, thick-walled armed
with bristle-like spines similar to those of cirrus. Uterus joins middle of anterior
tubular part of terminal organ. Eggs small, numerous, 20–23 by 12. Vitelline follicles
large, arranged in two groups of 5–7 follicles each, situated on either side of ovary.
Remarks: L. maculatus most closely resembles L. longitestis Durio and Manter,

1968, reported from Plectorhynchus sp. from New Caledonia but differs in its less
elongate oral sucker, the smaller testis, the three-lobed rather than four-lobed ovary
and the smaller eggs (24–28 by 14–15 in L. longitestis). Gupta and Jain (1992a)
reported L. maculatus from Cynoglossus dubius (Day) from Puri Coast, BOB.

odhneri (after Srivastava,
1939)
Lasiotocus odhneri (Srivastava, 1939) Thomas, 1959 (Fig. 16.7)

[Syn: Genolopa odhneri Srivastava, 1939J]
Host: Leiognathidae: Leiognathus daura Cuvier
Locality: Arabian Sea
Reference: Srivastava (1939j)
Remarks: Some of the diagnostic features of the species are: body size 2,000–2,400
by 580. Sucker ratio 1: 0.7. Caeca extend up to posterior end of body. Cirrus-sac
extends to anterior margin of ovary. Genital pore submedian. Terminal organ-shaped
like an inverted U, anterior to cirrus-sac.
Lasiotocus overstreeti Gupta & Gupta, 1990 (Fig. 16.8)

Host: Lutjanidae: Lutjanus monostigma (Cuvier)
Locality: DC, BOB
No. of specimens: 2
References: Gupta, P. C. and Gupta, V. C. (1990)
Description: Body elongate, 1170–1230 by 360–420. Oral sucker funnel-shaped,

110–120 × 130–140. Caeca end blindly close to posterior extremity. Ventral sucker
poorly developed 20–30 in diameter. Testis oval, pre-equatorial, 150–170 × 120–130.
Cirrus-sac elongate, extends to posterior margin of testis, encloses elongated seminal
vesicle, tubular pars prostatica, numerous prostatic cells. Cirrus long armed with
spines. Genital atrium unarmed. Ovary tri-lobed, overlaps testis. Terminal organ
tubular, bipartite, posterior part unarmed 240–340 long, anterior part 40–60, armed
with spines. Uterus fills post-testicular space. Eggs 20–22 by 8–9. Vitelline follicles
in two lateral groups, 4–9 on left, 4–7 on right side.

overstreeti (after Gupta &
Gupta, 1990)
Remarks: This species closely resembles L. hastai but differs from it in having a
funnel-shaped rather than oval oral sucker, the absence of gland cells over the anterior
part of terminal organ, the absence of a sphincter in the terminal organ.
Lasiotocus polynemi (Dutta, Hafeezullah and Manna, 1994) Dove and Cribb,
1998
Syn: Chrisomon polynemi Dutta, Hafeezullah and Manna, 1994
Host: Polynemidae: Eleutheronema tetradactylum (Shaw)
Locality: DC, BOB
No. of specimens: 3
Reference: Dutta, et al. (1994)
Lasiotocus puriensis Ahmad and Gupta, 1985 (Fig. 16.9)
Host: Serranidae: Cephalopholis sonnerati (Val.)
Locality: PR, BOB
No. 3
Reference: Ahmad and Gupta, V (1985)
Description: Body elongate, 1870–3180 long, 280-432 wide at level of testis. Entire
tegument spinose. Eyespot pigment absent. Oral sucker funnel-shaped, terminal,
110–220 long, 90–110 wide. Ventral sucker in anterior part of middle third of body,
150–180 in diameter. Sucker width ratio 1:0.63–1.66. Intestinal bifurcation dorsal,
usually midway between pharynx and ventral sucker. Caeca extend to posterior end
of body. Testis oval, in anterior part of posterior third of body. Cirrus-sac club-shaped,
extends posteriorly to just anterior to ovary. Seminal vesicle saccular. Cirrus thick-
walled, with bristle-like spines, opening into genital atrium. Genital atrium unspined,
overlaps posterior margin of ventral sucker. Ovary spherical, immediately pretestic-
ular. Uterus occupies all available space posterior to testis, then runs anteriorly and
opens medially into terminal organ. Terminal organ bipartite, 1/3rd to 1/2 of cirrus-
sac length; basal part unspined, filled with viscous substance; anterior part tubular,
thick-walled, armed with bristle-like spines; opening into genital atrium. Eggs ovoid,

puriensis (after Ahmad and
Gupta, 1985)
operculate, 14–16 long, 9–12 wide. Vitelline follicles lateral, 8–10 on each side, in
ovario-testicular zone. Excretory vesicle tubular, extends to testis; pore terminal.
Remarks: L. puriensis closely resembles Lasiotocus hastai from Pomadasys hasta

(Bloch.) in having the funnel-shaped oral sucker and unlobed ovary but differs from
it in having the genital atrium triangular and spacious, a more elongate oral sucker,
the prepharynx and seminal vesicle being comparatively longer and the cirrus-sac
extending posteriorly to just anterior to the ovary.
Lasiotocus srivastavai Mittal & Pande, 2007

Host: Serranidae: Serranus indicus (Cuv. & Val.)
Locality: PR, BOB
No. 15
Reference: Mittal & Pande (2007)
Description: Body small, elongate, 1425–1760 long by 425–445 wide. Tegument

spinose. Oral sucker 175–240 by 21–24 in size. Ventral sucker smaller than oral
sucker, 100–125 wide. Prepharynx absent. Pharynx globular. Oesophagus absent.
Caeca extend up to mid-length of post-testicular space. Genital pore submedian, in
forebody, surrounded by spacious genital atrium. Testis single, 145–200 long by
100–130 wide. Cirrus-sac large, club-shaped, extends obliquely from genital pore
up to just posterior to ovary or posterior end of testis. Seminal vesicle saccular. Pars
prostatica tubular surrounded large number of prostatic cells. Cirrus spiny. Ovary
three-lobed, submedian, overlaps testis or separated. Vitelline follicles in two lateral
groups of four follicles each, extend from middle of cirrus-sac to just posterior to
ovary. Uterus fills all available space in hindbody, opens into terminal organ.
Remarks: This species differs from all the Indian species of the genus in the extension
of the cirrus-sac as far as posterior end of the testis or just posterior to ovary.
Genus Leiomonorchis Mamaev, 1970

[Syn: Caballerotrematoides Madhavi, 1977]
Diagnosis: Body elongate. Tegument spinose. Suckers well developed. Forebody

short. Prepharynx ands pharynx present. Intestinal bifurcation in posterior part of
forebody. Caeca long. Testis single, near posterior end of body. Cirrus-sac long,
claviform encloses saccular seminal vesicle, pars prostatica surrounded by prostatic
cells and armed cirrus. Genital atrium unarmed. Genital pore anterior to ventral
sucker. Ovary lobed, pretesticular, separated from testis by uterine coils. Uterine coils
fill space between testis and ventral sucker, not extending posteriorly beyond testis.
Terminal organ bipartite, distal portion spined and receives uterus partway along
its length. Vitellarium composed of acinous tubules forming star-shaped clusters
in lateral fields of middle third of body. Genital atrium unspined. Type species: L.
leiognathi Mamaev, 1970.
Leiomonorchis mamaevi (Madhavi, 1977) Madhavi, 2008 (Fig. 16.10)

[Syn: Caballerotrematoides leiognathi Madhavi, 1977]
Fig. 16.10 Leiomonorchis

mamaevi (after Madhavi,
1977)
Locality: VSK, BOB

Reference: Madhavi (1977; 2008)
Number: 9
Description: Body elongate, 2240–3200 long 384–576 wide. Tegument spined in

anterior region. Oral sucker small, 58–85 by 66–117. Prepharynx small or absent.
Pharynx 43–77 by 58–78. Oesophagus nearly twice the length of pharynx, Intestinal
bifurcation immediately anterior to ventral sucker. Caeca long terminate near pos-
terior end of body. Ventral sucker 110–144 in diameter. Genital pore median, near
anterior border of ventral sucker. Testis single, large, elliptical, situated very close
to posterior extremity. Cirrus-sac small, clavate, extends slightly posterior to ventral
sucker, encloses oval seminal vesicle, small pars prostatica surrounded by numerous
prostatic cells and cirrus armed with needle-like spines. Ovary four-lobed, pretes-
ticular, separated from testis by uterine coils. Uterine coils extensive, occupy the
space between testis and ventral sucker, not extending posterior to testis. Terminal
organ small, half as long as cirrus-sac, bipartite, posterior part ovoid, anterior part
thick-walled, spined. Genital atrium tubular unarmed, fairly long but contracted in
some specimens. Eggs small, numerous, non-filamented, 20 by 12 in size. Vitelline
follicles large, acinous, situated in lateral fields of middle third of body from anterior
limit of testis to just anterior to ovary.
Remarks: Madhavi (1977) erected the genus Caballerotrematoides with C. leiog-

nathi Madhavi, 1977 as the type species. One more species C. secundus Ahmad,
1981 was added to the genus. The genus was characterised by the vitellarium
composed of acinous tubules forming star shaped clusters in the middle third of
body and the uterus not extending posterior to the testis. Madhavi (2008) syn-
onymized the genus Caballerotrematoides Madhavi, 1977, with Leiomonorchis
Mamaev, 1970, erected by Mamaev (1970) with L. leiognathi Mamaev, 1970, as
the type species. Madhavi (2008) redesignated C. leiognathi as L. mamaevi (Mad-
havi, 1977) since L. leiognathi was preoccupied. Caballerotrematoides secundus
Ahmad, 1981 becameLeiomonorchis secundus (Ahmad, 1981).
Leiomonorchis secundus (Ahmad, 1981) Madhavi, 2011 (Fig. 16.11)

[Syn: Caballerotrematoides secundus Ahmad, 1981]
Host: Carangidae: Scomberoides tala (Cuv.): deep queenfish.
Locality: PR, BOB
Description: Body elongate, 1760–1950 long. 270–330 wide. Oral sucker 120–165
in diameter. Ventral sucker 87–110 in diameter, pre-equatorial. Sucker ratio
1:0.66–0.725. Caeca simple, extend posteriorly up to 1/3rd of testis. Testis sin-
gle, elongated, near hind end of body. Cirrus-sac extends beyond ventral sucker to
halfway between ventral sucker and ovary, contains tubular seminal vesicle, prostate
complex and long cirrus armed with thorn-shaped spines, opening into a shallow
Fig. 16.11 Leiomonorchis

1981)
genital atrium. Genital pore immediately anterior to ventral sucker. Ovary three-
lobed, median, intercaecal, pretesticular. Uterus descends posteriorly as far as mid-
dle of testis, enters medial side of terminal organ. Terminal organ bipartite, posterior
part oval, unspined; anterior part tubular, spined, spines thorn-shaped. Eggs non-
filamented, 33–39 long, 19–23 wide. Vitelline follicles large, rod-shaped, lateral,
extend from base of cirrus-sac to anterior border of testis. Excretory vesicle tubular,
reaches to ovary.
Remarks: This species differs from L. mamaevi (Madhavi, 1977) in having its oral
sucker larger than the ventral sucker, a longer prepharynx and oesophagus, the intesti-
nal bifurcation more posteriorly placed, the caeca extending up to 1/3 of the testis
and the ventral sucker located more posteriorly.
Genus Monorcheides Odhner 1905

This genus is characterized by the presence of two testes situated symmetrically
close to posterior end of body, the uterus not extending posterior to the testes, the
vitellarium consisting of two symmetrical bunches situated in the lateral fields near
the ventral sucker.
Diagnosis: Body oval. Oral sucker terminal. Ventral sucker small, pre-equatorial.
Intestinal bifurcation in forebody. Caeca long. Testes two, symmetrical, one on each
side close to posterior end of body. Cirrus-sac small or large, may or may not extend
posterior to ventral sucker, encloses simple seminal vesicle, prostatic complex and
spined cirrus. Genital pore median, post-bifurcal. Ovary deeply lobed, pretesticular.
Uterus fills hindbody, does not extend posterior to testes, opens into terminal organ at
its base. Terminal organ simple, spined. Vitellarium follicles in two lateral bunches
at level of ventral sucker or in forebody. Excretory vesicle Y-shaped. Type species:
M. diplorchis Odhner, 1905.
Monorcheides hafeezullahi Bijukumar, 1997 (Fig. 16.12)

Host: Paralichthyidae: Pseudorhombus triocellatus (Bl. & Schn.)
Reference: Bijukumr (1997)
Fig. 16.12 Monorcheides

hafeezullahi (after
Bijukumar, 1997, Acta
Parasit. 42, p. 154)
Description: Body suboval, widest in hindbody, slightly narrowing anteriorly,

875–1750 long by 625–800 wide, somewhat reddish in colour. Tegument spined.
Suckers poorly developed. Oral sucker rounded 120–133 in diameter. Ventral sucker
133–147, slightly postequatorial, sucker ratio 1:1.1–1.8. Intestinal bifurcation nearer
to oral sucker than to ventral sucker. Caeca narrow, long. Testes two, elliptical, poste-
quatorial, symmetrical. Genital pore median, midway between suckers. Cirrus-sac
large, extends slightly posterior to ventral sucker, encloses oval seminal vesicle, short
pars prostatica, surrounded by numerous prostatic cells and long protrusible cirrus
armed with numerous long needle-like spines. Genital atrium unarmed. Ovary 2–4-
lobed, situated between testes, slightly submedian. Uterus voluminous, occupies all
available space between testes and ventral sucker, terminal organ bipartite, distal part
spined. Vitellarium in form of lateral bunches of irregularly lobed follicles, situated
in forebody at level of genital pore. Vitelline ducts prominent, converge towards
vitelline reservoir, situated near ovary. Eggs oval, 8–12 by 7–8 in size. Excretory
vesicle Y-shaped.
Remarks: M. hafeezullahi Bijukumar, 1997, closely resembles M. decapteri Manter
and Pritchard, 1961, in general appearance but differs in the robust body, in having
a four-lobed rather than three-lobed ovary, in the character and distribution of the
vitelline follicles and in the nature of the armature of the terminal genitalia.
Genus Monorchis Looss, 1902
This genus possesses typical monorchiid features such as small oval body, spined
tegument, two suckers, short caeca, a single testis, a long cirrus-sac enclosing a
seminal vesicle, a prostatic complex and a spined cirrus, a median genital pore anterior
to the ventral sucker, a spherical ovary, well-developed uterine coils and a bipartite
terminal organ, with the anterior part spined.
Two species of this genus have been recorded from marine fish of India.
Monorchis bengalensis Dutta & Manna, 1997
Host: Haemulidae: Pomadasys hasta (Bloch)
Locality: Port Canning, West Bengal

No. 9
Reference: Dutta & Manna (1997)
Description: Body small, 810–1000 long, 510–680 wide. Oral sucker subterminal,
120–140 × 140–150 in size, larger than ventral sucker. Ventral sucker 90–110 ×
80–110. Sucker ratio 1: 0.68. Caeca extend up to posterior end of body. Testis more
or less at middle of hindbody. Cirrus-sac very large, pretesticular. Seminal vesicle
oval, pars prostatica long surrounded by numerous prostatic cells, cirrus spined.
Genital pore near pharynx. Ovary pretesticular, bean-shaped. Vitellarium follicles
oval, extracaecal, form bunches on each side of body, each bunch consisting of
12–14 follicles. Uterus fills most of hindbody. Terminal organ bipartite, anterior part
thick-walled, spined, eggs 12–14 × 8–12.
Remarks: This species differs from M. minutus Madhavi, 1977, the only other
species of the genus known from India, in having oral sucker larger than ventral
sucker, the number of vitelline follicles in each bunch and the shape and position of
ovary.
Monorchis minutus Madhavi, 1977 (Fig. 16.13)

Location: VSK, BOB
No. Numerous
Description: Body small, oval, 387–560 long, 234–320 wide. Tegument covered
with minute spines. Oral sucker subterminal, 50–86 by 56–192. Ventral sucker 59–92
in diameter. Sucker ratio 1: 0.85–1.0. Intestinal bifurcation at midway between suck-
ers. Caeca long narrow, terminate close to posterior extremity. Genital pore median,
midway between intestinal bifurcation and anterior margin of ventral sucker. Testis
single, spherical, located slightly below midbody. Cirrus-sac long, extends well pos-
terior to ventral sucker to reach mid-level of testis or slightly beyond, encloses oval
Fig. 16.13 Monorchis

minutus (after Madhavi,
1977)
seminal vesicle, long pars prostatica, prostatic cells and cirrus armed with needle-
like spines. Genital atrium small, unarmed. Ovary small, broadly ovate, pretesticular,
contiguous with anterior margin of testis, dextral to ventral sucker. Vitelline follicles
small, oval, forming two lateral bunches in between bifurcation point and anterior
margin of ventral sucker, each bunch consists of 8–11 follicles. Uterine coils exten-
sive, occupy entire post-testicular space. Terminal organ half as long as cirrus-sac,
bipartite, anterior part thick-walled, armed with needle-like spines, metraterm joins
it laterally. Eggs 18–23 by 8–10. Excretory vesicle I-shaped, reaches level of ventral
sucker.
Remarks: This species is characterized by the small body size and the subequal suck-
ers. It is distinguished from the type species with which it shows close resemblance,
apart from the above features, in the smaller and more anteriorly located gonads, the
more posteriorly distributed vitelline follicles and the more extensive uterine coils.
Genus Neotimonia Madhavi, 2008
Diagnosis: Body elliptical. Suckers small subequal. Ventral sucker pre-equatorial.

Caeca long, narrow, terminate at level of testes. Testes two elliptical, obliquely tan-
dem, in middle third of hindbody. Cirrus-sac arcuate, extends well into hindbody,
encloses oval seminal vesicle, well-developed prostatic complex and spined cirrus.
Genital atrium short unarmed. Genital pore median, just anterior to ventral sucker.
Ovary lobed, to right of anterior testis. Uterine coils extensive, fill entire hindbody.
Metraterm opens into proximal region of terminal organ. Terminal organ bipartite,
anterior part with triangular spines. Eggs not filamented. Vitellarium consists of
irregularly shaped follicles arranged in two groups at level of ventral sucker. Type
species: N. caballeroi (Madhavi, 1977).
Neotimonia caballeroi (Madhavi, 1977) Madhavi, 2008 (Fig. 16.14)

[Syn: Timonia caballeroi Madhavi 1977]
Host: Polynemidae: Polydactylus sextarius (Bl. & Schn.)
Locality: VSK
Reference: Madhavi (1977, 2008)
Description: Body oval, 1080–1920 wide. Tegument spined. Suckers poorly devel-
oped. Oral sucker small 58–66 by 58–78. Ventral sucker 63–97 in diameter, at a dis-
tance of 320–560 from anterior end. Prepharynx small. Pharynx spherical. Oesoph-
agus fairly long. Intestinal bifurcation nearer to oral sucker than to ventral sucker.
Caeca narrow with thick walls, terminate near posterior extremity of body. Geni-
tal pore median, immediately anterior to ventral sucker. Testes two, oval, situated
obliquely in midbody. Cirrus-sac large, clavate, curves around right margin of ventral
sucker to reach level of ovary, encloses oval seminal vesicle, small pars prostatica,
numerous prostatic cells and highly muscular eversible cirrus lined with closely set
needle-like spines. Ovary three-lobed, pretesticular or to right of anterior testis more
Fig. 16.14 Neotimonia

caballeroi (after Madhavi,
1977)
or less at midbody. Uterine coils extensive, fill entire hindbody. Terminal organ bipar-
tite, posterior part unarmed, anterior part with triangular spines differing in size and
shape from those of cirrus. Uterus opens into anterior region of terminal organ. Eggs
small, 16 by 12. Vitellarium consists of many irregularly shaped follicles arranged in
two groups at level of ventral sucker, the two groups tend to meet medianly at level
of ventral sucker.
Remarks: The genus Neotimonia Madhavi, 2008, was proposed by Madhavi (2008)
for Timonia caballeroi Madhavi, 1977, which differs from other species of Timonia
Bartoli & Prevot, 1966, by the bipartite terminal organ, the vitellarium in the forebody
and the uterine coils descending into the post-testicular region. N. caballeroi is the
type and only species of the genus.
Genus Paramonorcheides Yamaguti, 1938
Diagnosis: Body elliptical, Suckers weakly muscular. Caeca long, terminate at some
distance anterior to posterior extremity. Testes two, longitudinally elongate, sym-
metrical or not. Cirrus-sac extends well posterior to ventral sucker, contains oval
seminal vesicle, well enveloped prostatic cells and armed eversible cirrus. Genital
atrium small. Genital pore median anterior to ventral sucker. Ovary lobed, pre- or
intertesticular. Uterus occupies all available space in hindbody. Metraterm opens into
terminal organ at junction of two parts. Terminal organ bipartite, proximal part with
spines. Vitellarium in two lateral groups of follicles in region between ventral sucker
and testis. Excretory vesicle saccular. Type species: P. awatati Yamaguti, 1938.
Paramonorcheides selaris Trivenilakshmi & Madhavi, 2009 (Figs. 16.15 and

16.16)
Host: Carangidae: Selar crumenophthalmus (Bloch)
Locality: VSK, BOB
Reference: Trivenilakshmi & Madhavi (2009)
Description: Body elliptical, 1408–1760 long, 384–640 wide. Forebody flat, exten-
sile, hindbody cylindrical. Tegument spined. Suckers weekly muscular. Oral sucker
42–60 by 60–68. Ventral sucker rounded pre-equatorial, 80–88 in diameter, sucker
ration 1: 1.2–1.25. Intestinal bifurcation at midway between suckers. Caeca narrow
terminate in posterior quarter of body. Testes two, sausage-shaped, situated symmet-
rically in posterior region of body, with posterior ends in contact. Cirrus-sac large,
curved, extends well posterior to ventral sucker, reaching ovarian zone, encloses
oval seminal vesicle, short pars prostatica, numerous prostatic cells. Cirrus long,
armed with thorn-like spines, tip of cirrus additionally provided with 6–8 very long
spike-like spines. Genital atrium unarmed, genital pore median, just anterior to ven-
tral sucker. Ovary spherical or oval anterolateral to right testis. Uterine coils occupy
most of hindbody, extend posterior to testis. Metraterm narrow, muscular. Terminal
organ bipartite, with oval unarmed basal part and narrow proximal portion armed
with densely arranged acicular spines. Eggs oval, 20 × 12 in size. Vitelline follicles
irregular, arranged in two lateral groups between genital pore and ventral sucker.
Excretory bladder saccular.
Remarks: This species is closest to the Australian species, P. pseudocaranxi Dove

& Cribb, 1988, but differs in its shorter cirrus-sac extending only to the level of the
ovary rather than to the level of the testes, in lacking eyespot pigment and in details
of the armature of terminal genitalia.
Fig. 16.15 Paramonorcheides

selaris (after
Triveni Lakshmi & Madhavi,
2009)
Fig. 16.16 Paramonorcheides

selaris, terminal genitalia
Madhavi, 2009)
Genus Proctotrema Odhner, 1911
Diagnosis: Body elongate. Tegument spinose. Oral sucker funnel-shaped. Ventral

sucker small, pre-equatorial. Caeca not reaching posterior extremity. Testis single
in middle or posterior third of body. Cirrus-sac encloses saccular seminal vesicle,
pars prostatica surrounded by prostatic cells and spiny eversible ejaculatory duct.
Genital atrium unarmed. Genital pore anterior to ventral sucker. Ovary lobed or
not, anterior to testis. Uterine seminal receptacle present. Uterus occupies most of
hindbody. Terminal organ oval, undivided, spined. Vitellarium composed of two
groups of follicles situated on either side of ovary. Type species: P. bacillovatum
Odhner, 1911.
Proctotrema guptai Ahmad & Dhar, 1987 (Fig. 16.17)

[Syn: Lasiotocus guptai (Ahmad & Dhar, 1987) Dove and Cribb, 1998)]
Host: Haemulidae: Pomodasys maculatus (Bloch)
No. 7
Reference: Ahmad & Dhar (1987a)
Description: Body elongate, with rounded extremities, 2150–2670 long, 470–560

wide at level of ovary. Tegument thin, spined. Oral sucker funnel-shaped, 280–330
long 200–230 wide. Ventral sucker smaller than oral sucker, 130–150 in diame-
ter; situated at junction of anterior and middle third of the body. Sucker ratio 1:0.65.
Prepharynx 55–80 long. Pharynx 60–70 long, 70–80 wide. Oesophagus long. Intesti-
nal bifurcation dorsal, nearer to ventral sucker than to oral sucker. Caeca terminate just
posterior to testis. Testis single, elliptical, post-equatorial. Cirrus-sac club-shaped,
straight, extends posteriorly to midway between ventral sucker and ovary. Seminal
vesicle saccular. Pars prostatica surrounded by prostate gland cells. Cirrus thick-
walled, armed with thorn-shaped spines. Genital atrium oval, unspined. Genital pore
median, immediately anterior to ventral sucker. Ovary three-lobed, pretesticular to
left of median line. Uterus descends to posterior end of body, then ascends anteriorly
Fig. 16.17 Proctotrema

guptai (after Ahmad & Dhar,
1987)
and opens into terminal organ at its base. Terminal organ simple, saccular, unipartite.
Vitellarium 6–8 follicles on each side, confined to ovarian zone. Excretory vesi-
cle tubular, reaching anteriorly as far as posterior margin of testis. Excretory pore
terminal.
Remarks: Dove and Cribb (1998) transferred this species into the genus Lasiotocus
as L. guptai (Ahmad & Dhar, 1987). Much confusion prevails in the differentiation
of the genera Lasiotocus and Proctotrema. Bartoli and Prevot (1966) considered the
presence of a coiled seminal vesicle, the small size and the structure of terminal
organ and the opening of the uterus at the proximal end of terminal organ are the
characters separating the genus Proctotrema from Lasiotocus. Nahhas & Powell
(1965) consider the type of terminal organ and the point of uterine junction with it
as of generic value. As per these suggestions, the morphology of the form described
by Ahmad and Dhar (1987a) fits into the genus Proctotrema. Hence, it is retained
here as P. guptai.
Genus Proctotrematoides Yamaguti, 1938
The four monorchiid genera including Genolopa Linton, 1910, Lasiotocus Looss,
Proctotrema Odhner, 1911, and Proctotrematoides Yamaguti, 1938, appear very
similar. Various views were expressed regarding the validity of these genera. Finally,
all the four genera were retained as valid differing mainly in the structure of the
terminal genitalia and the nature of their spination. The genus Proctotrematoides
is considered as distinct from the other three genera in possessing the following
combination of characters: Presence of spines in the cirrus, terminal organ and genital
atrium, presence of a muscular spiny atrial pouch, spherical ovary, extent of vitelline
follicles from anterior level of ventral sucker to the level of posterior end of testis,
bipartite terminal organ with uterus entering its spiny anterior part, excretory vesicle
extending to near intestinal bifurcation point.
Four species belonging to this genus have been reported from Indian marine fish.
For two of these species, namely P. thapari Ahmad, 1980, from Stromateus cinereus
from PR, BOB and P. stromateusi Gupta & Ahmad, 1976, from the same host and
locality, only abstracts were published. Full details of the morphology of the species
are yet to be published. The other two species are P. indicus Ahmad & Gupta, 1985,
and P. yamaguti Dutta & Manna, 1998.
Proctotrematoides indicum Ahmad & Gupta, 1985 (Fig. 16.18)

Host: Stromateidae: Pampus argenteus (Euphrasen) (=Stromateus cinereus Bloch
Locality: PR, BOB
Number: 9
Reference: Ahmad & Gupta (1985)
Fig. 16.18 Proctotrematoides

indicum (after Ahmad &
Gupta, 1985)
Description: Body short, oculate, forebody attenuated, posterior end rounded,

1440–1870 long, 272–280 wide. Entire tegument spinose. Oral sucker 61–130 in
diameter. Ventral sucker spherical, equatorial, larger than oral sucker, 81–140 in
diameter. Sucker ratio 1: 1.07–1.30. Intestinal bifurcation dorsal, nearer to ventral
sucker than to oral sucker. Caeca simple, terminate at posterior end of body. Testis
single, ovoid, in posterior third of body. Cirrus-sac elongate, club-shaped, straight or
curved, extends posteriorly as far as ovary or just anterior. Seminal vesicle saccular,
pars prostatica surrounded by numerous prostate gland cells. Cirrus thick-walled,
armed with bristle-like spines, opens into genital atrium. Atrial diverticulum short,
armed with bristle-like spines. Genital pore ventral, median, between intestinal bifur-
cation and ventral sucker. Ovary three-lobed, immediately pretesticular. Seminal
receptacle absent. Uterus descends posteriorly to just posterior to testis, then coiling
anteriorly and joins anterior end of terminal organ medially. Terminal organ bipar-
tite, basal part unspined, filled with viscous substance; anterior part tubular, armed
with bristle-like spines, opens into genital atrium. Eggs broadly oval, operculate,
13–15 long, 8–12 wide. Vitelline follicles small, rounded, intrude into intercaecal
space dorsally and ventrally, extend laterally from level of genital pore to hind end
of cirrus-sac. Excretory vesicle tubular, reaching to testis.
Proctotrematoides yamagutii Dutta & Manna, 1998 (emended) (Fig. 16.19)

Host: Ophichthidae: Pisodontophis boro (Hamilton)
Locality: Port Canning, West Bengal, BOB
No. 9
Reference: Dutta & Manna (1998b)
Description: Body 1550–2380 long, 460–600 wide, elongated and pointed posteri-
orly. Tegument spinose. Suckers subequal. Oral sucker terminal 90–140 in diameter.
Ventral sucker postequatorial, 90–140 in diameter; sucker ratio 1: 0.96. Prepharynx
small. Pharynx oval. Oesophagus long. Caeca extend just posterior to testis. Testis
single, oval. Cirrus-sac intercaecal, tubular, curved. Seminal vesicle unipartite. Pars
Fig. 16.19 Proctotrematoides

yamagutii (after Dutta &
Manna, 1998)
prostatica small surrounded by prostate gland cells. Genital opening ventral to ventral
sucker. Ovary oval, pretesticular, dextral. Vitellarium follicular, situated laterally in
two groups along ovario-testicular zone. Uterus occupies post-testicular space. Ter-
minal organ unipartite and armed with triangular spines. Eggs with unipolar spines,
16–20 by 12–16. Excretory vesicle I-shaped.
Remarks: This species differs from all the other species of the genus in that the eggs
are provided with a polar spine. Otherwise, it is very similar to P. pisodontophis
Yamaguti, 1938, reported from Pisodonophis cancrivora from Inland sea of Japan.
Genus Pseudametrodaptes Trivenilakshmi & Madhavi, 2008

The genus Pseudametrodaptes was erected by Triveni Lakshmi & Madhavi (2008)
to include the four Indian species of the genus Ametrodaptes Bravo-Hollis, 1966,
characterized by the absence of spines in the genital atrium and presence of a uni-
partite terminal organ with the metraterm joining its distal region.
Diagnosis: Body elongate without eyespot pigment. Tegument spined. Oral sucker
funnel-shaped with ring of enlarged spines. Ventral sucker in anterior third of body.
Intestinal bifurcation anterior to ventral sucker. Caeca long, end near posterior
extremity of body. Testis single, elliptical, equatorial or postequatorial. Cirrus-sac
slender, extends well into hindbody, encloses bipartite or saccular seminal vesicle,
well-developed prostatic complex and spined cirrus. Genital atrium tubular, thin-
walled, unarmed. Genital pore median in forebody. Ovary spherical or indented,
pretesticular. Uterus fills most of hindbody. Eggs small, oval. Terminal organ unipar-
tite, armed with triangular spines, metraterm joins its distal end. Vitelline follicles
on lateral fields of ovario-testicular zone or extend little anteriorly. Type species: P.
secundus (Madhavi, 1977) Triveni Lakshmi & Madhavi, 2007 (Syn. Ametrodaptes
secundus Madhavi, 1977).
Pseudametrodaptes bravoae (Ahmad, 1985) Trivenilakshmi & Madhavi, 2008

(Fig. 16.20)
[Syn: Ametrodaptes bravoae Ahmad, 1985]
Host: Terapontidae: Terapon puta (Cuv. and Val.)
Locality: BOM, AS
No. 9
References: Ahmad (1985); Triveni Lakshmi & Madhavi (2008)
Description: Body elongate, with nearly parallel sides, 2056–2510 long by 300–350
wide. Tegument spined. Oral sucker funnel-shaped, terminal, 105–136 long by 87–98
wide. Circumoral spines 54–56 in number, arranged in two alternating rows dorsally
and in a single row ventrally, spines of first dorsal row 9–12 long; spines of second
dorsal row 6–8 long and spines of ventral row 12–16 long. Ventral sucker spherical,
larger than oral sucker, 130–150 in diameter, pre-equatorial, situated in posterior
part of middle third of body. Sucker ratio 1: 1.4–1.5. Intestinal bifurcation dorsal,
midway between suckers. Caeca simple, lateral, end blindly near posterior margin of
testis. Testis single, elliptical, very large, situated in middle third of body. Cirrus-sac
Fig. 16.20 Pseudametrodaptes

bravoae (after Ahmad, 1985)
club-shaped, extends posteriorly as far as anterior margin of testis. Seminal vesi-

cle bipartite. Pars prostatica long, surrounded by prostate gland cells. Cirrus thick-
walled, armed with thorn-shaped spines. Genital atrium tubular. unspined. Genital
pore median, ventral. Ovary four-lobed, overlaps testis anteriorly. Vitellarium con-
sisting of 18–23 follicles on each lateral side, confined between ventral sucker and
ovary. Uterus descends posteriorly to halfway of post-testicular space, then ascends
anteriorly and opens into terminal organ at its base. Terminal organ simple, unipar-
tite, spined, spines thorn-shaped. Eggs operculate, 15–20 × 10–13 in size. Excretory
vesicle saccular, short.
Remarks: This species differs from P. secundus (Madhavi, 1977) in having a bipartite
seminal vesicle and in the much larger oesophagus.
Pseudametrodaptes caballeroi (Ahmad, 1985) Triveni Lakshmi & Madhavi, 2008
(Figs. 16.21 and 16.22)
[Syn: Ametrodaptes caballeroi Ahmad, 1985]
Host: Lactariidae: Lactarius lactarius (Bl. & Schn.)
Locality: BOM, AS
References: Ahmad (1985b); Triveni Lakshmi & Madhavi (2008)
Description: Body elongate, 2200–2850 long by 405–475 wide. Tegument spined.
Oral sucker funnel-shaped, 120–130 long by 80–95 wide. Circumoral spines 52–54 in

caballeroi (after Ahmad,
1985)

caballeroi (after Ahmad,
1985)
number, arranged in two alternating rows dorsally and in a single row ventrally, spines
on ventral row larger than those on dorsal side Ventral sucker spherical, 135–180 in
diameter, situated in anterior part of middle third of body. Sucker ratio 1: 1.53–1.59.
Intestinal bifurcation nearer to ventral sucker than to oral sucker. Caeca simple,
terminate blindly near posterior edge of testis. Testis single, elliptical, very large,
situated in anterior part of middle third of body. Cirrus-sac club-shaped, extends
posteriorly up to midway between ovary and ventral sucker. Seminal vesicle saccular.
Pars prostatica surrounded by prostate gland cells. Cirrus thick-walled, armed with
thorn-shaped spines. Genital atrium tubular, unspined. Genital pore median, ventral,
situated just anterior to ventral sucker. Ovary three-lobed, immediately pretesticular.
Seminal receptacle uterine. Uterus descends posteriorly to halfway of post-testicular

space, then ascends anteriorly and opens into terminal organ at its base. Terminal
organ simple, unipartite, spined, spines thorn-shaped, 16–18, long. Eggs 15–20 long
by 10–13 wide. Vitellarium consists of 6–7 follicles on each lateral side, confined to
ovarian zone. Excretory vesicle saccular, reaches anteriorly to posterior margin of
testis. Excretory pore terminal.
Remarks: This species differs from P. secundus (Madhavi, 1977) as follows: ovary
is tri-lobed rather than unlobed, and eggs are larger. The ventral sucker is located far
posterior to the intestinal bifurcation.
Pseudametrodaptes fischthali (Ahmad & Dhar, 1987) Trivenilakshmi & Mad-

havi, 2008
[Syn: Ametrodaptes fischthali Ahmad & Dhar, 1987]
Host: Haemulidae: Pomadasys maculatus (Bloch.)
Locality: PR, BOB
No. 8
References: Ahmad & Dhar (1987b); Triveni Lakshmi & Madhavi (2008)
Description: Body elongate, slender, 2380–3310 long, 270–410 wide at level of

ovary. Tegument spined. Eyespot pigment absent. Oral sucker funnel-shaped, termi-
nal, 130–190 long and 124–160 wide. Circumoral spines 52–54 in number; arranged
in two alternating rows dorsally and ventrally; spines unequal in size; anterior row
larger, 14–17 long; posterior row smaller, 10–13 long. Ventral sucker spherical, larger
than oral sucker, 160–220 in diameter, median, situated in anterior part of middle
third of body. Sucker ratio 1: 1.29–1.37. Prepharynx long. Oesophagus short. Intesti-
nal bifurcation immediately anterior to ventral sucker. Caeca terminate at anterior
level of testis. Testis single, elliptical, median, situated in middle part of posterior
third of body. Cirrus-sac club-shaped, curved anteriorly, extends posteriorly as far
as anterior margin of ovary. Seminal vesicle bipartite, pars prostatica surrounded
by prostate gland cells. Cirrus thick-walled, armed with thorn-shaped spines. Gen-
ital atrium tubular, unspined. Genital pore median, immediately anterior to ventral
sucker. Ovary three-lobed, median, immediately pretesticular. Uterine seminal recep-
tacle present. Uterus fills entire hindbody, opens into terminal organ at its base. Ter-
minal organ simple, saccular, unipartite, armed with thorn-shaped spines. Eggs oval,
9–14 long and 8–11 wide. Vitellarium consists of 7–10 follicles on each lateral side,
confined to ovario-testicular zone. Excretory vesicle tubular, extends anteriorly as
far as anterior margin of ovary. Excretory pore terminal.
Remarks: This species differs from the type species P. secundus in the three-lobed
ovary, bipartite seminal vesicle and the larger cirrus-sac extending posteriorly up to
anterior margin of ovary.
Pseudametrodaptes secundus (Madhavi, 1977) Trivenilakshmi &Madhavi, 2008

(Fig. 16.23)
[Syn: Ametrodaptes secundus Madhavi, 1974]
Locality: VSK, BOB
Number: Numerous
References: Madhavi (1977), Triveni Lakshmi & Madhavi (2008)
Description: Body elongate, 1920–4640 long 256–438 wide. Tegument spined, in

regular rows in forebody. Oral sucker poorly developed, funnel-shaped, 130–150
by 136–195 in size. Circumoral spines 48–52, arranged in two rows dorsally and
single row ventrally, spines small, 10–16 in size. Ventral sucker 136–164 in diameter
situated in anterior third of body. Intestinal bifurcation immediately anterior to ventral
sucker. Caeca long terminate near posterior end. Testis single, elliptical situated in
posterior quarter of body, post-testicular space 192–800 long. Cirrus-sac slender,
clavate, extends to midway between ovary and ventral sucker, contains saccular
seminal vesicle, prostatic complex and cirrus armed with triangular spine. Genital
pore median, immediately anterior to ventral sucker. Ovary spherical or elliptical,
or indented, pretesticular, contiguous with testis. Uterine coils fill all space between
cirrus-sac and ventral sucker, except that to right of testis. Terminal organ simple,
undivided, elliptical, half as long as cirrus-sac armed with triangular spines. Uterus
joins it near distal end. Eggs 16–20 by 9–12. Vitelline follicles large, spherical,
confined to lateral fields in testiculo-ovarian zone.

secundus (after Madhavi,
1977)
Remarks: This species is characterized by the spherical ovary, saccular seminal

vesicle and cirrus-sac extending to midway between ventral sucker and ovary.
Key to Indian species of Pseudametrodaptes:

1. Seminal vesicle saccular, ovary spherical ……………P. secundus (Madhavi,
1977)
Seminal vesicle bipartite, ovary 3- or 4-lobed………………………… 2
2. Cirrus-sac extends to midway between ventral sucker and ovary… P. caballeroi
(Ahmad, 1985)
Cirrus-sac extends to level of ovary …………..……........…………… 3
3. Prepharynx very long …………………………… P. fischthali (Ahmad & Dhar,
1987)
Prepharynx absent ………………...........................………… P. bravoae Ahmad
(1985)
Genus Timonia Bartoli & Prevot, 1966
Diagnosis: Body elongate. Suckers small. Intestinal bifurcation midway between

suckers. Caeca long. Testes two, oblique, in posterior half of body. Cirrus-sac of
variable size may or may not extend posterior to ventral sucker. Seminal vesicle
oval. Cirrus spined. Genital pore median, anterior to ventral sucker. Ovary lobed
or not, pretesticular. Uterus not extending posterior to testis. Terminal organ oval,
spined, receives uterus laterally. Vitelline follicles in lateral fields between cirrus-sac
and postovarian region. Type species: T. mediterranea Bartoli & Prevot, 1966.
Five species of Timonia have been described from Indian marine fish. However,
Madhavi (2008) erected a new genus Neotimonia Madhavi, 2008, for inclusion of
Timonia caballeroi Madhavi, 1977.
Timonia indica Madhavi, 1977 (Fig. 16.24)
Host: Polynemidae: Leptomelanosoma indicum (Shaw)
Locality: VSK, BOB
Fig. 16.24 Timonia indica

Number: 15
Description: Body small, oval, 848–1280 long, 256–320 wide. Tegument spined.
Oral sucker, 50–66 by 74. Ventral sucker in anterior region of middle third of body,
small, circular, 58–78 in diameter. Prepharynx small. Pharynx oval. Oesophagus
long. Intestinal bifurcation midway between suckers. Caeca broad, terminate near
posterior end of body. Testes two, spherical, small, situated obliquely one behind
other near posterior end of body. Genital pore median, slightly in front of ventral
sucker. Cirrus-sac long, slender, C-shaped, extends from mid-level of ventral sucker
to slightly anterior to genital pore, encloses long seminal vesicle, small pars prostat-
ica, numerous prostate cells and slender cirrus armed with triangular spines. Ovary
three-lobed, situated at about middle of hindbody. Vitellarium formed of large folli-
cles distributed in lateral fields between ventral sucker and testis with 8–10 follicles
on each side. Uterine coils occupy space between testes and ventral sucker, not
extending posterior to testis. Terminal organ small, simple, lies anterior to ventral
sucker, armed with triangular spines, uterus joins it laterally. Eggs small, oval, 16 by
8 in size.
Remarks: This species differs from the type species in the smaller testes, the smaller
cirrus-sac not extending past the ventral sucker, the three-lobed ovary, more extensive
vitelline follicles and the smaller eggs.
Timonia nahhasi Ahmad, 1985 (Fig. 16.25)

Locality: BOM, AS
No.: 16
Description: Body elongate, with blunt anterior and rounded posterior extremity,
1230–1880 long by 435–490 wide. Eyespot pigment absent. Oral sucker funnel-
shaped, terminal, 160–220 long by 160–220 wide. Ventral sucker spherical, smaller
Fig. 16.25 Timonia nahhasi

J. Zool. 17, p. 152)
than oral sucker, 70–75 in diameter, median, pre-equatorial, 92-140 post-bifurcal.

Sucker ratio 1: 0.34–0.43. Prepharynx short. Pharynx oval. Oesophagus surrounded
by numerous gland cells. Intestinal bifurcation dorsal, midway between suckers.
Caeca simple, lateral, terminate blindly near posterior end of body. Testes two, entire,
longitudinally elongate, diagonal, separated by uterine coils, subequal in size, in pos-
terior third of body. Post-testicular space very short. Cirrus-sac club-shaped, curved
anteriorly, extends from genital pore to well posterior to ventral sucker, roughly
midway between ventral sucker and ovary or more posterior. Seminal vesicle sac-
cular. Pars prostatica surrounded by prostate gland cells. Cirrus thick-walled, armed
with thorn-shaped spines; spines 21–24 long. Genital pore median, ventral, midway
between intestinal bifurcation and ventral sucker. Ovary entire, pyriform, immedi-
ately pretesticular, submedian, to right of median line. Uterus descends posteriorly up
to just anterior to posterior extremity, then ascends anteriorly, opening into terminal
organ medially. Terminal organ bipartite, spined entirely; spines thorn-shaped. Eggs
oval, operculate, 17–22 × 10–13 wide. Vitellarium consists of 8–9 follicles on each
side between posterior end of cirrus-sac and anterior end of anterior testis. Excretory
vesicle tubular, extends anteriorly up to anterior level of anterior testis. Excretory
pore terminal.
Remarks: T. nahhasi was differentiated from the other two species of the genus,
namely T. mediterranea and T. indica, in having a funnel-shaped oral sucker, in the
presence of a prepharynx, in the presence of gland cells around the oesophagus and
in having a tubular excretory bladder.
Timonia stunkardi Ahmad, 1985 (Fig. 16.26)
Host: Mullidae: Upeneus vittatus (Forsskal)
Locality: BOM, AS
No. 51
Description: Body slender, elongate, 2200–2850 long, 405–475 wide. Oral sucker
funnel-shaped, 90–120 by 105–130 in size. Ventral sucker longitudinally elongate,
Fig. 16.26 Timonia

1985)
smaller than oral sucker, 44–56 long 38–40 wide. Sucker ratio 1: 0.30–0.32. Prephar-
ynx long. Pharynx oval. Oesophagus short. Intestinal bifurcation nearer to ventral
sucker than to oral sucker. Caeca terminate blindly near posterior edge of poste-
rior testis. Testes two, oval, diagonal, well separated from one another by uterine
coils. Cirrus-sac club-shaped, curved, extends from genital pore posteriorly either
up to anterior margin of ovary or almost to it. Seminal vesicle oval. Pars prostatica
long, Cirrus thick-walled armed with thorn-like spines. Genital pore median, post-
bifurcal. Ovary spherical, pretesticular. Vitellarium consists of 30–35 follicles con-
fined to pretesticular zone and confluent medially anterior to ventral sucker. Uterus
fills post-testicular space. Terminal organ bipartite, posterior part saccular, unspined,
anterior part tubular, spined. Eggs 28–35 by 9–14. Excretory bladder tubular, reaches
anteriorly up to anterior margin of anterior testis.
Remarks: T. stunkardi closely resembles T. caballeroi but differs from it in the fol-
lowing characters: oral sucker is funnel-shaped instead of subspherical, prepharynx is
longer, oesophagus is very short, intestinal bifurcation is nearer to ventral sucker than
to oral sucker, intestinal caeca terminate at posterior edge of posterior testis rather
than near the posterior end of body, ovary is spherical, postequatorial, submedian
anterior to the anterior testis rather than tri-lobed, equatorial, close to the anterior
testis, the testes are well separated rather than close together, eggs are larger, cirrus
spines are triangular rather than needle-like, terminal organ spines are needle-like
rather than triangular, and oral sucker is much larger than the ventral sucker instead
of being the same size or smaller.
Timonia vinodae Ahmad, 1987 (Fig. 16.27)
Host: Engraulidae: Thryssa mystax (Bloch & Schn.)
Locality: Goa, AS
Description: Body elongate, fusiform, 840–1125 long, 285–355 wide. Tegument

spined. Oral sucker funnel-shaped, 70–82 long, 75–90 wide. Ventral sucker slightly
larger than oral sucker, 85–98 in diameter, pre-equatorial. Openings of mouth and
Fig. 16.27 Timonia vinodae

J. Zool. 19, p. 325)
ventral sucker surrounded by well-developed sphincters. Sucker ratio 1: 1.08–1.12.

Intestinal bifurcation nearer to oral sucker than to ventral sucker. Caeca terminate just
posterior to testis. Testes two, diagonal, situated in posterior half of body. Cirrus-sac
C-shaped, extends posteriorly as far as anterior margin of anterior testis, encloses sac-
cular seminal vesicle, pars prostatica, numerous prostatic cells, cirrus thick-walled,
spined, spines needle-like. Genital pore median, anterior to ventral sucker. Ovary tri-
angular, close to anterior testis. Uterine coils numerous, fill post-testicular space. Ter-
minal organ bipartite, posterior part saccular unspined, anterior part tubular spined,
spines thorn-shaped. Eggs 14–18 by 11–13 in size. Vitellarium consists of numer-
ous follicles confined between posterior margin of pharynx and base of cirrus-sac.
Excretory vesicle saccular.
Remarks: This species is characterized by the following features: the oral and ventral
openings are surrounded by well-developed sphincter muscle, oral sucker smaller
than ventral sucker, vitellarium confined between pharynx and base of cirrus-sac.
These characters distinguish it from T. stunkardi Ahmad, 1985, with which it shows
close similarity.
Subfamily Hurleytrematinae Yamaguti, 1958
Diagnosis: Body oval, elongate or long and slender. Tegument spined. Oral sucker
small, Ventral sucker pre-equatorial. Prepharynx present or absent, pharynx globular,
oesophagus long, caeca long. Testes single or double, oval, symmetrical, oblique or
tandem when double, in middle or posterior third of hindbody. Cirrus-sac contains
saccular or bipartite seminal vesicle, numerous prostate cells, surrounding narrow
pars prostatica and spined cirrus. Genital pore anterior to ventral sucker, median
or submedian. Ovary pretesticular, entire or lobed. Uterus fills most of hindbody.
Terminal organ present or absent, unipartite or bipartite, armed with spines. Eggs
filamented. Vitellarium forms symmetrical bunches of follicles or tubular acini of
variable extent. Excretory vesicle saccular or tubular. Type genus: Hurleytrema Sri-
vastava, 1939.
Key to genera of Hurleytrematinae

1. Testis single ………………………………………………………………2
Testes two ………………………………………………........................…3
2. Seminal vesicle undivided ……...………………………...…… Hurleytrema
Srivastava, 1939
Seminal vesicle bipartite………………………………………Hurleytrematoides
Yamaguti, 1954
3. Testes close to posterior extremity, uterus not extending posterior to testes
…………………………………………Helicometroides Yamaguti, 1954
Testes at middle of hindbody, uterus extending posterior to testes
…………………………. Diplohurleytrema Nahhas & Cable, 1964
Genus Diplohurleytrema Nahhas & Cable, 1964
Diagnosis: Body fusiform. Suckers well developed. Pharynx oval. Oesophagus

short. Intestinal bifurcation in forebody. Caeca terminate just posterior to ventral
sucker. Testes diagonal in middle of hindbody. Cirrus-sac long, claviform, encloses
bipartite seminal vesicle, well-developed prostatic complex and spined cirrus. Gen-
ital pore median, anterior to ventral sucker. Ovary entire, dextral, postero-dorsal
to ventral sucker. Uterus in testicular and post-testicular region, fills post-testicular
space. No terminal organ. Vitelline follicles extend laterally from level of pharynx
to ovary. Excretory vesicle tubular. Type species D. brevicaecum Nahhas & Cable,
1964.
Diplohurleytrema nahhasi Ahmad, 1981 (Fig. 16.28)
Type host: Lutjanidae Pristipomoidesargyrogrammicus (Val.), snapper;
Locality: BOM, AS
Description Body elongated, 1710–1842 long, 350–420 wide at level of ventral

sucker. Oral sucker 87–115 in diameter, spherical terminal; ventral sucker 170–200
long, 202–231 wide, larger than oral sucker, median, at junction of anterior and
middle third of body. Sucker ratio 1:2.0–2.3. Caeca simple, terminate blindly near
anterior margin of posterior testis. Testes two, equal in size, elongated, directly tan-
dem, contiguous, in posterior half of body. Cirrus-sac club-shaped, extends beyond
ventral sucker to level of ovary, contains bipartite seminal vesicle, prostate complex
and cirrus armed with thorn-like spines, opening into genital atrium. Genital pore
immediately anterior to ventral sucker. Ovary four-lobed, submedian, pretesticular.
Seminal receptacle postovarian. Uterus descends to posterior end of body, opens into
posterior part of terminal organ. Terminal organ undivided, armed with bristle-like
spines, opens into genital atrium. Eggs 25–30 × 12–14, with unipolar filaments 2–4
times length of eggs. Vitellarium follicular, follicles large, lateral, overlap caeca,
Fig. 16.28 Diplohurleytrema

nahhasi (Ahmad, 1981)
extend between ventral sucker and anterior testis. Excretory bladder tubular, reaches
to anterior testis; pore terminal.
Remarks: Ahmad (1981) differentiated Diplohurleytrema nahhasi from D. brevi-

caecum Nahhas and Cable, 1964, the type and only other species of the genus,
through differences in sucker proportions, extent of caeca, extent of cirrus-sac and
distribution of vitelline follicles apart from other minor differences.
Genus Hurleytrema Srivastava, 1939
Diagnosis: Body medium sized, narrow, spinose. Oral sucker larger than ventral
sucker situated at junction of first and middle third of body length. Prepharynx,
pharynx and oesophagus present. Caeca narrow, reaching posterior end. Testis single,
compact, just posterior to middle third of body length. Cirrus-sac well-developed,
elongated, extending from genital atrium to last third of middle third of body, encloses
oval seminal vesicle, pars prostatica with prostate glands and armed cirrus. Genital
atrium close to ventral sucker. Ovary tri-lobed, pretesticular. Seminal receptacle
absent. Uterus occupies most of hindbody. Eggs triangular with long polar filament.
Vitellarium follicular, follicles extend from genital atrium almost to posterior end of
body. Type species: H. ovocaudatum Srivastava, 1939.
Remarks This genus differs from all the genera of the subfamily in the extent of the
vitellarium, which extends from the level of the ventral sucker to almost the posterior
end.
Hurleytrema ovocaudatum Srivastava, 1939 (Fig. 16.29)
Host: Siganidae: Siganus fuscescens (Bl. & Schn.)
Locality: AS
Reference: Srivastava (1939j)
Description: Body elongate, 3000 long, 420 wide. Tegument spinose. Oral sucker
subterminal 110 × 130 in size. Prepharynx long. Pharynx oval. Oesophagus fairly
long, narrow. Caeca slender, ending blindly at posterior end. Ventral sucker 76–80
in diameter, situated at the end of first third of body. Sucker ratio 3:2. Testis single,
pear-shaped, situated just posterior to middle third of body. Cirrus-sac club-shaped,
extends from genital pore to last third of middle third of body, encloses elongated
seminal vesicle, tubular pars prostatica, armed cirrus. Genital atrium situated close
to left margin of ventral sucker. Ovary three-lobed, pretesticular. Seminal recep-
tacle absent. Uterus well developed occupies whole of intercaecal space between
metraterm and blind end of caeca. Metraterm well developed, armed with minute
spines. Vitellarium with elongated pear-shaped small follicles extend laterally from
level of ventral sucker to last fifteenth part of body length. Eggs triangular with an
opercular polar filament, 19–30 × 11–15.
Fig. 16.29 Hurleytrema

ovocaudatum (after
Srivastava, 1939)
Genus Helicometroides Yamaguti, 1934

[Syn. Hysterorchis Durio & Manter, 1968]
Genus Helicometroides Yamaguti, 1934, was initially included under the family
Opecoelidae. It was shifted from Opecoelidae to Monorchiidae based on the pres-
ence of spined tegument, nature of the terminal genitalia and well-developed uterine
coils. The genus Hysterorchis, Durio and Manter, 1968, was synonymized with Heli-
cometroides (Ref. Madhavi, 2008). Molecular results reported by Searle et al. (2014)
confirm the position of this genus in the Monorchiidae.
Diagnosis: Body elongate. Tegument spined. Eyespot pigment present. Oral sucker
terminal. Ventral sucker pre-equatorial. Oesophagus long. Intestinal bifurcation
immediately anterior to ventral sucker. Caeca long. Testes two, elongate, symmetri-
cal, close to posterior extremity. Cirrus-sac claviform, curved around ventral sucker,
contains bipartite seminal vesicle, few prostatic cells and spined cirrus. Genital pore
submedian, anterosinistral to ventral sucker. Uterus occupies space between testes
and ventral sucker. Terminal organ tubular, unspined. Vitelline follicles fairly large,
extend from mid-level of oesophagus to anterior margin of testes, confluent anteri-
orly. Type species: H. longicollis Yamaguti, 1934.
Two species belonging to this genus have been recorded from India.
Fig. 16.30 Helicometroides

leiperi (after Ahmad, 1981)
Helicometroides leiperi (Ahmad, 1981) Madhavi, 2008 (Fig. 16.30)

[Syn. Hysterorchis leiperi Ahmad, 1981]
Host: Sciaenidae: Otolithus ruber (Bloch &Schneider)
Locality: PR, BOB
References: Ahmad (1981b); Madhavi (2008)
Description: Body 1468–1542 long, 580–633 wide, pyriform. Entire tegument

spinose. Eyespot pigments present on either side behind pharynx. Oral sucker 75–90
in diameter. Ventral sucker 130–188 in diameter spherical larger than oral sucker.
Sucker ratio 1: 1.733–2.088. Intestinal bifurcation dorsal at junction of anterior and
middle third of body. Caeca simple lateral, ending blindly anterior to testes. Testes
two, oval, symmetrical, slightly overlapping each other, near posterior end of body.
Cirrus-sac arcuate, on left side of ventral sucker, extends slightly beyond posterior
margin of ventral sucker, contains bipartite seminal vesicle, prostate complex and
cirrus armed with short thorn-shaped spines. Genital pore immediately post-bifurcal.
Ovary three-lobed, median, intercaecal, halfway between ventral sucker and testes at
junction of middle and posterior third of body. Vitelline follicles small, lateral, over-
lap caeca dorsally and ventrally, invade intercaecal space, extend from just posterior
to intestinal bifurcation to anterior edge of ovary. Uterus with single vertical coil
descends to anterior border of testes, joins posterior end of terminal organ. Terminal
organ simple, unspined, thick-walled. Eggs yellow, 55–61 long, 26–29 wide with
long, filament at an opercular end. Excretory bladder Y-shaped, reaches to testes,
pore terminal.
Remarks: This species differs from H. pseudovitellosus in having the vitelline fields
non-confluent anterior to the ventral sucker and in the larger eggs.
Helicometroides pseudovitellosus (Madhavi, 1974) Madhavi, 2008 (Fig. 16.31)
[Syn: Hysterorchis pseudovitellosus Madhavi, 1974]
Fig. 16.31 Helicometroides

pseudovitellosus (after
Madhavi, 1974)
Host: Lutjanidae: Lutjanus sp.

Locality: VSK, BOB
No. 11
References: Madhavi (1974a); Madhavi (2008)
Description: Body pyriform, 960–1040 long 384–560 wide. Tegument spined. Eye-
spot pigment present. Oral sucker 78–89 by 86–117. Ventral sucker circular, 78–88 in
diameter somewhat pre-equatorial. Sucker ratio 1:0.83– 0.88. Testes two, large, sym-
metrical, ovoid, situated in posterior third of body close to posterior end. Cirrus-sac
long, arcuate, extends to near posterior margin of ventral sucker, encloses bipar-
tite seminal vesicle, short tubular pars prostatica and eversible cirrus armed with
triangular spines. Genital pore median just anterior to ventral sucker. Ovary three-
lobed, anterior to right testis. Seminal receptacle absent. Vitelline follicles numerous,
arranged in two wide lateral zones, extend from intestinal bifurcation to mid-level
of testes. Uterus with single vertical coil, runs posteriorly into intertesticular area.
Metraterm small, tubular, not spined. Eggs 31–39 by 19, with long filament at one
end.
Genus Hurleytrematoides Yamaguti, 1954

This genus is characterized by the following features: Elongate body with spined
tegument and eyespot pigment in forebody. Caeca long. Testis single. Cirrus-sac
small, encloses bipartite seminal vesicle, weakly developed prostatic complex and
spined cirrus. Genital pore median, anterior to ventral sucker. Ovary globular or
lobed, pretesticular. Uterus fills entire hindbody. Terminal organ unipartite, spined.
Vitellarium in lateral fields between ovary and cirrus-sac. Type species:H. chaetodoni
(Manter, 1942) Yamguti, 1954.
Hurleytrematoides filiformis Madhavi, 1974 (Fig. 16.32)
Host: Chaetodontidae: Chaetodon pictus (Forsskal)
No. 18
Locality: VSK, BOB
Reference: Madhavi, (1974a)
Fig. 16.32 Hurleytrematoides

filiformis (after Madhavi,
1974)
Description: Body slender, filiform, of uniform width throughout, 2000–3040 long

and 160–240 wide. Suckers poorly developed. Oral sucker 85–117 by 105–117 in
size. Ventral sucker 88–94 in diameter. Sucker ratio 1: 0.80–0.86. Forebody one-fifth
to one-sixth of total length. Eyespot pigment present on either side of oesophagus.
Caeca long terminate some distance from posterior extremity. Genital pore median,
just anterior to ventral sucker. Testes at about middle of hindbody, elliptical, con-
stricted in middle in some. Cirrus-sac claviform, thin-walled extends well posterior to
ventral sucker, encloses bipartite seminal vesicle, prostatic vesicle and cirrus armed
with bristle-like spines. Ovary ovoid, slightly anterior to testis, separated from testis
by uterine coils. Seminal receptacle absent. Uterine coils vertical, sigmoid, fill all
postovarian space, modified terminally into muscular thick-walled metraterm free
of spines. Terminal organ absent. Eggs filamented, 31 by19. Vitelline follicles large,
rounded, in two lateral groups separated by uterine coils, extend from level of pos-
terior end of cirrus-sac to slightly in front of ovary.
Remarks: Nahhas and Powell (1965) grouped Hurleytrema-like forms into four gen-
era based on simple or bipartite seminal vesicle and simple or divided terminal organ
and restricted the genus Hurleytrematroides to the species possessing bipartite sem-
inal vesicle and a simple terminal organ. H. filiformis is characterized by the long
slender body, fairly large testes, anteriorly placed vitellarium and separated gonads.
McNamara & Cribb (2011) considered this species as one of the two most elon-
gate members of the genus, the other being H. loi McNamara & Cribb, 2011, from
Chaetodon spp. on the Great Barrier Reef.
Subfamily: Opisthomonorcheidinae Yamaguti, 1971
Diagnosis: Body long, slender, delicate. Tegument spined. Forebody short, hind-
body long, Suckers poorly developed. Ventral sucker in anterior quarter of body.
Intestinal bifurcation in forebody. Caeca long, terminate near posterior extremity of
body or some way short. Testis single or double close to posterior extremity. Cirrus-
sac small extends posterior to ventral sucker, encloses oval seminal vesicle, prostatic
complex and spined cirrus. Genital pore median in posterior forebody. Ovary pretes-
ticular, separated from testis by uterine coils. Uterus fills intercaecal space between
ovary and ventral sucker, may extend into post-testicular space, joins terminal organ
at proximal or distal end. Terminal organ oval or bipartite. Eggs oval, numerous, not
filamented. Vitelline follicles small, in space between testis and cirrus-sac. Excretory
vesicle I-shaped. Type genus: Opisthomonorcheides Parukhin, 1966.
Key to genera:
1. Testis single ……………………… Opisthomonorcheides Parukhin, 1966
Testes two ………………………………….................................…….…… 2
2. Terminal organ simple, uterus pretesticular ………….. Opisthodiplomonorchis
Madhavi, 1974
Terminal organ bipartite, uterus extends into post-testicular space
………………………… Neolasiotocus Ahmad, 1991
Genus Neolasiotocus Ahmad, 1991
Diagnosis: Body elongate, without eyespot pigment. Tegument spined. Prephar-

ynx, pharynx, oesophagus present. Caeca not reaching posterior extremity. Testes
two, directly tandem, in hindbody. Cirrus-sac club-shaped, encloses seminal vesicle,
pars prostatica and spined cirrus. Genital pore median, in forebody. Ovary lobed,
pretesticular. Vitellarium follicular, confined between base of cirrus-sac and anterior
testis. Uterus filling hindbody. Terminal organ bipartite, with spined anterior part.
Eggs non-filamented. Excretory vesicle saccular. Type species: N. brayi Ahmad,
1991.
Neolasiotocus brayi Ahmad, 1991 (Figs. 16.33 and 16.34)
Host: Terapontidae: Terapon theraps (Cuv.)
No. 11
Description: Body elongate, 3260–5125 long by 490–610 wide at the level of testes.
Eyespot pigment absent. Tegument spined. Oral sucker spherical, 180–245 in diam-
eter. Ventral sucker larger than oral sucker, 245–280 in diameter, situated at junction
of anterior and middle third of body. Sucker ratio 1:1.14–1.36. Caeca terminate just
posterior to posterior testis. Testes two, elongate, directly tandem, close together,
nearly equal in size situated in middle of posterior half of body. Cirrus-sac elongate,
club-shaped, curved, dorsal to ventral sucker, extends posteriorly well beyond ven-
tral sucker. Seminal vesicle saccular. Pars prostatica tubular, surrounded by prostate
gland cells. Cirrus thick-walled, muscular, armed with thorn-shaped spines. Genital
atrium cup-shaped, unspined. Genital pore median, immediately anterior to ven-
tral sucker. Ovary three-lobed, post equatorial. Uterine seminal receptacle present.
Uterus fills hindbody, opens into terminal organ medially at junction of unspined
and spined portions. Terminal organ bipartite, thick-walled, posterior vesicle sac-
cular, unspined; anterior part tubular, armed with thorn-shaped spines. Vitellarium
consists of 28–35 follicles on each lateral side, confined between the base of the
Fig. 16.33 Neolasiotocus

Fig. 16.34 Neolasiotocus

cirrus-sac and anterior margin of anterior testis. Eggs oval, operculate, 17–22 long
by 11–13 wide. Excretory vesicle saccular, short. Excretory pore terminal.
Remarks: Neolasiotocus most closely resembles the genera Timonia Prevot, 1966
and Paratimonia Prevot & Bartoli 1967, in possessing two testes, bipartite terminal
organ, median genital pore situated in forebody and non-filamented eggs but differs
from these genera in the testes being tandem and in the uterus descending to posterior
end of body. So far N. brayi Ahmad, 1991 is the only species in the genus.
Genus Opisthodiplomonorchis Madhavi, 1974
This genus synonymized with the deropristid genus Skrjabinopsolus Ivanov in Ivanov
& Murygin, 1937, by Hafeezullah (1984) was resurrected as valid by Choudhury &
Dick (1998).
Diagnosis: Body elongate. Ventral sucker in anterior third of body. Caeca extending
to posterior end of body. Testes oval, tandem, near posterior extremity of body.
Cirrus-sac small. Genital atrium tubular thin-walled. Genital pore median, anterior
to ventral sucker. Ovary multi-lobed, pretesticular, separated from testis by uterus.
Uterus occupies space between testis and ventral sucker. Terminal organ simple,
sac-like, lined by triangular spines. Metraterm joins its distal end. Eggs small, oval.
Vitelline follicles small, distributed between ovary and posterior end of cirrus-sac,
arranged in two lateral groups of pre and postovarian follicles. Type species: O.
elongatus Madhavi, 1974.
Opisthodiplomonorchis elongatus Madhavi, 1974 (Fig. 16.35)
[Syns. Skrjabinopsolus elongatus (Madhavi, 1974) Hafeezullah, 1984; S. indicus

Gupta & Ahmad, 1976; S. kurotchkini Parukhin, 1976 (Reference: Hafeezullah,
1984); O. paurosoma (Wang, 1987) (Reference: Choudhury & Dick (1998)
Fig. 16.35 Opisthodiplomonorchis

elongatus Madhavi, 1974
Hosts: Psettodidae: Psettodes erumei (Bloch& Schneider); Polynemidae: Polydacty-

lus sextarius Bloch& Schneider (1); Pomacanthidae: Abudefduf bengalensis (Bloch)
(2) PortoNovo
Locality: VSK, BOB, (1) Chilka lagoon (2) PortoNovo, DC (BOB), Veraval (AS)
(3)
Distribution: Kuwait coast of Arabian Gulf (Sey & Nahhas, 1997); South China
(Wang, 1987)
References: 1. Madhavi (1974a); 2. Gupta, V & Ahmad (1974b); 3. Hafeezullah
(1984)
Description: Body elongate 2885–3750 long, 420–570 wide. Tegument spinose.

Oral sucker 100–110 long, 100–130 wide. Ventral sucker pre-equatorial, larger than
oral sucker, 170–180 long, 170–182 wide. Prepharynx long. Pharynx ovoid. Oesoph-
agus tubular. Caeca simple extend to posterior extremity. Genital pore in forebody.
Testes oval, subequal, tandem, contiguous, near posterior end of body. Cirrus-sac
claviform, extends some distance beyond ventral sucker. Seminal vesicle bipartite.
Pars prostatica surrounded by large number of prostate gland cells. Cirrus spined.
Ovary multi-lobed, postequatorial. Seminal receptacle retort-shaped, posterior to
ovary. Uterus coiled, runs posteriorly as far as middle of anterior testis and then runs
anteriorly to open into base of terminal organ. Terminal organ unipartite, spined. Eggs
ovoid, operculated, 37–60 × 14–21. Vitelline follicles in two lateral groups anterior
and posterior to ovary.
Remarks: The genus Opisthodiplomonorchis erected by Madhavi (1974) with O.

elongatus as the type species is characterized by the following combination of char-
acters: Elongate slender body, tandem testes at posterior extremity, unipartite spiny
terminal organ vitelline follicles in two lateral groups anterior and posterior to multi-
lobed ovary. Hafeezullah (1984) synonymized the genus with Skrjabinopsolus and
suggested the new combination as S. elongatus (Madhavi, 1974). He regarded S.
indicus Gupta & Ahmad, 1976, and S. kurotchkini Parukhin, 1976 as synonyms of
S. elongatus. However, Choudhury & Dick (1998) did not accept this synonymy and
restored the original status of the species as O. elongatus. They further considered S.
paurosoma Wang, 1987, as synonymous to O. elongatus. Mani et al. (2014) reported
its occurrence in Polydactylus sextarius from VSK, BOB. The species shows some
specificity towards polynemid fishes.
Genus Opisthomonorcheides Parukhin, 1966
[Syn. Retractomonorchis Madhavi, 1977; Longimonorchis Oshmarin, Mamaev &
Lebedev, 1990; Tritesticularis Bilqees, 1971]
Genus Opisthomonorcheides was erected by Parukhin (1966) with O. decapteri

Parukhin, 1966, from the carangid fishes Decapterus sp. and Selar mate from the
Gulf of Tonkin as the type species. Madhavi (1977) erected another genus Retrac-
tomonorchis Madhavi, 1977, with R. delicatus Madhavi, 1977, from the stromatoid
fishes Pampus chinensis and P. argenteus from VSK, BOB as the type species.
Retractomonorchis was differentiated from Opisthomonorchis by the presence of a
long tubular genital atrium and the position of genital pore anterior to ventral sucker
in contrast to a very short genital atrium and genital pore posterior to ventral sucker
in Opisthomonorcheides. Madhavi (1977) further observed that the genital atrium in
Retractomonorchis is thin and not easily visible and in many specimens the genital
pore appears as if it is located posterior to ventral sucker. She expressed the view that
Retractomonorchis would fall a synonym of Opisthomonorcheides if genital pore in
the latter is proved to be posterior to ventral sucker. Hafeezullah (1984) after exam-
ining large collection of the specimens apparently identical to Opisthomonorcheides
collected from Apolectus niger (Bloch) considered Retractomonorchis as a syn-
onym of Opithomonorcheides. Gupta & Singh (1989) accepted this synonymy while
Ahmad (1984) retained them as distinct genera. Madhavi (2011), Machida (2011)
and Bray et al. (2017) also treated them as congeneric.
Diagnosis: Body elongate, delicate. Forebody short, retractile. Eyespot pigment
present. Oral sucker poorly developed. Ventral sucker well developed. Prepharynx,
pharynx, oesophagus present. Caeca long. Testis single, close to posterior end of
body. Cirrus-sac extends well posterior to ventral sucker, encloses elongate seminal
vesicle, globular pars prostatica, numerous prostatic cells and spined cirrus. Genital
atrium long, tubular, thin-walled, smooth. Genital pore median, anterior to ventral
sucker. Ovary three-lobed, separated from testis by uterus. Uterine coils fill all space
posterior to cirrus-sac, extend well into post-testicular space. Terminal organ small,
spined. Eggs oval, numerous. Vitelline follicles extend from behind cirrus-sac to
level of testis. Type species: O. decapteri Parukhin, 1966.
It is to be noted that merging of the of the two genera Opisthomonorcheides and
Retractomonorchis not only increased considerably the number of species in the
genus Opisthomonorcheides to about 24, but also resulted in several homonyms and
created confusion in the separation of species. For instance, there are three homonyms
for O. indicus and two for O. overstreeti. Hafeezullah (1984) after a careful analysis
of the various species in the two genera, synonymized O. delicatus (Madhavi, 1977),
and all the three homonymous species named as O. indicus with O. karwarensis
Karyakarte& Yadav, 1976, the first described species in the genus from the Indian
marine fishes. Subsequent to Hafeezullah’s publication in 1984, many more species
have been reported and the genus requires a thorough revision. Bray et al. (2017)
provided a review on the genus and considered 27 species of the genus as valid
and 11 easily distinguished and divided them into four groups. They presented a
host–parasite check-list and discussed the host specificity of members of the genus.
In the collection of the senior author, large numbers of flukes identified as O.
karwarensis collected from pomfrets and carangids of VSK, BOB were available.
The specimens exhibited intraspecific variations in several features attributed to the
highly extensile nature of the body, especially the forebody and the slender delicate
body filled with numerous eggs. Evidence indicated that certain features like length
of body, forebody and hindbody ratio, minor variations in sucker proportions, length
of prepharynx, oesophagus, length of genital atrium, position of genital pore, size of
cirrus-sac, metraterm and the shape of ovary (lobed or entire) slight variation in the
extent of the vitellarium cannot be used for differentiation of species in the genus.
In the light of these observations, many species of Opisthomonorcheides would fall

synonyms of O. karwarensis.
In this account, full description is provided only for O. karwarensis and the remain-
ing species are listed providing details of their hosts and localities.
Opisthomonorcheides decapteri Parukhin, 1966
Host: Carangidae: Carangoides armatus (Rüppell); Stromateidae: Parastromateus
niger (Bloch).
Localities: Gulf of Mannar, BOB.
References: Parukhin (1976), Dutta & Manna (1998).
Opisthomonorcheides aspinosus Khan & Karyakarte, 1985
Host: Stromateidae: Pampus argenteus Euphrasen;
Locality: RT, AS
Reference: Khan & Karyakarte (1985).
Opisthomonorcheides chaetodoni Ahmad, 1984
Host: Chaetodontidae: Chaetodon vagabundus Linnaeus (=Chaetodon pictus)
Locality: BOM, AS
Opisthomonorcheides delicatus (Madhavi, 1977) Hafeezullah, 1984
Syn: Retractomonorchis delicatusMadhavi, 1977
Host: Stromateidae: Pampus chinensis (Euphrasen), P. argenteus (Euphrasen)
Locality: VSK, BOB
References: Madhavi (1977); Hafeezullah (1984)
Remarks: Hafeezullah (1984) and Bray et al. (2017) considered this a synonym of
O. karwarensis.
Opisthomonorcheides engraulisi Gupta & Singh, 1989
Host: Engraulidae: Thryssa mystax (Bl./& Schn.)
Locality: PR, BOB
Reference: Gupta, & Singh (1989)
Remark: Bray et al. (2017) considered this a synonym of O. simhai.
Opisthomonorcheides indicus Ahmad, 1977
Host: Apogonidae: Apogon ellioti (Day),
Locality: PR, BOB
Reference: Ahmad (1977).
Remark: This species is a junior homonym of O. indicus Karyyakarte & Yadav, 1976,
and has been renamed four times, O. jamilahmadi Pozdnyakov, 1981, O. puriensis
Ahmad, 1985, O. puriensis Ahmad & Gupta, 1985, and O. apogonichthysi Gupta &
Puri, 1985.
Opisthomonorcheides gibsoni (Ahmad, 1991) Madhavi, 2011
Syn: Retractomonorchis gibsoni Ahmad, 1991
Host: Serranidae: Cephalopholis sonnerati (L.)
Opisthomonorcheides guptai (Ahmad, 1985) Madhavi, 1911

[Syn: Retractomonorchis guptai Ahmad, 1985]
Host: Drepanidae: Drepane punctata (L.)
Locality: BOM, AS
Opisthomonorcheides manteri (Ahmad, 1984) Madhavi, 2011
[Syn. Retractomonorchis manteri Ahmad, 1984]
Host: Carangidae: Caranx sexfasciatus (Quoy & Gaimard)
Locality: BOM, AS
Reference: Ahmad (1984c), Madhavi (2011).
Opisthomonorcheides nahhasi (Ahmad, 1985) Madhavi, 2011
[Syn: Retractomonorchis nahhasi Ahmad, 1984]
Host: Serranidae: Epinephelus undulosus (Quy & Gaimard)
Locality: BOM, AS
Reference: Ahmad (1985), Madhavi (2011).
Opisthomonorcheides karwarensis Karyakarte & Yadav, 1976 (Fig. 16.36)
Syn: O. indicus Karyakarte & Yadav, 1976
O. indicus Ahmad, 1977
O. indicus Gupta & Gupta, 1978
O. delicatus (Madhavi, 1977) Madhavi, 1977
O. malabaricusi Gupta & Puri, 1985
O. apogonichthysi Gupta & Puri, 1984
O. puriensis Ahmad, 1984
Host: Carangidae: Parastromateus niger (Bloch)
Locality: RT, AS; Gopalpur, BOB
Reference: Karyakarte &Yadav (1976b).
Fig. 16.36 Opisthomonorcheides

karwarensis (after Madhavi,
1977)
Opisthomonorcheides madhavae (Ahmad, 1984) Madhavi, 2011

[Syn: Retractomonorchis madhavae Ahmad, 1984]
Host: Serranidae: Cephalopholis sonnerati (Val).
Locality: BOM, AS
Number: 21 specimens from 7 hosts out of 13 examined.
Reference: Ahmad (1984c).
Opisthomonorcheides nigeri Gupta & Puri, 1984
Host: Carangidae: Apolectus niger (Bloch)
Reference: Gupta, V & Puri (1984d).
Remarks: Bray et al. (2017) considered this a synonym of O. karwarensis.
Opisthomonorcheides overstreeti Ahmad, 1985
Host: Stromateidae: Pampus chinensis (Euphrasen)
Locality: PR, BOB
Opisthomonorcheides overstreeti (Ahmad, 1985) Madhavi, 2011
(Syn. Retractomonorchis overtreeti Ahmad, 1985)
Host: Nemipteridae: Scolopsis vosmeri (Bloch).
Locality: Arabian Sea off Mumbai.
Opisthomonorcheides pellonai Gupta & Puri, 1985
Host: Pristigasteridae: Ilisha elongata (Bennett)
Locality: PR, BOB
Reference: Gupta, V& Puri (1985).
Opisthomonorcheides shindei Gupta & Puri, 1985
Host: Apogonidae: Apogon ellioti (Day)
Locality: PR, BOB
Reference: Gupta & Puri (1985).
Opisthomonorcheides simhai Gupta & Puri, 1985
Host: Engraulidae: Thryssa mystax (Bloch & Schn.)
Locality: PR, BOB
Opisthomonorcheides stromatensis Ahmad & Gupta, 1985
Host: Carangidae: Parastromateus niger (Bloch.); Stromateidae: Pampus argenteus
(Euphr.),
Locality: PR, BOB
Ahmad & Gupta (1985b).
Opisthomonorcheides upeneusi Ahmad 1985
Host: Mullidae: Upeneus sulphureus (Cuv.)
Locality: BOM, AS
Opisthomonorcheides yamagutii Gupta & Singh, 1990
Reference: Gupta, P.C.& Singh (1990).
Opisthomonorcheides yamagutinsis Lokhande, 1993

Locality: Maharashtra, AS
Reference: Lokhande (1993).
Subfamily Opisthomonorchiinae Yamaguti, 1952
Diagnosis: Body elongate. Tegument spined. Oral sucker subterminal. Ventral

sucker in middle third of body. Prepharynx, oesophagus present. Caeca long. Testis
single, in posterior third of body. Cirrus-sac long, extends from posterior border of
ventral sucker to level of ovary, encloses oval or sinuous seminal vesicle, numerous
prostatic cells and short unarmed or armed cirrus. Genital atrium spined. Genital
pore median, in hindbody, near posterior margin of ventral sucker.
Ovary submedian, pretesticular. Uterus occupies most of post-testicular area. Ter-

minal organ absent. Eggs with filament at an opercular end. Vitelline follicles few,
form two lateral groups between ovary and ventral sucker or in forebody. Type genus:
Opisthomonorchis Yamaguti, 1952.
Genus Pseudopisthomonorchis Madhavi, 1974
This genus is characterized by the absence of spines on cirrus and metraterm thus
differing from any of the known members of the family Monorchiidae. However, in
other characters such as spined genital atrium, single testis and filamented eggs, the
genus fits into the family Monorchiidae. One other genus of Monorchiidae, namely
Opisthomonorchis Yamaguti, 1951, possesses a genital pore posterior to the ventral
sucker. Hafeezullah (1984) regarded the genus Pseudopisthomonorchis as a synonym
of Opisthomonorchis Yamaguti, 1952, but according to Madhavi (2008) the genus
can be differentiated from Opisthomonorchis by the globular cirrus-sac, absence of
spines in the ejaculatory duct and by the formation of a band of vitelline follicles
across the posterior forebody.
Diagnosis: Body elongated, spined. Eyespot pigment present. Oral sucker terminal.
Ventral sucker pre-equatorial. Prepharynx, oesophagus present. Caeca long. Testis
single. Cirrus-sac posterior to ventral sucker, encloses seminal vesicle, pars prostatica
and unarmed cirrus. Genital pore median, posterior to ventral sucker. Genital atrium
armed with spines. Ovary pretesticular. Eggs filamented. Metraterm small unarmed.
Terminal organ not present. Vitellarium in forebody forming two lateral groups in
front of ventral sucker, overlying caeca. Type species: P. carangi Madhavi, 1974.
Pseudopisthomonorchis carangi Madhavi, 1974 (Fig. 16.37)
Syn: Opisthomonorchis carangi (Madhavi, 1974) Hafeezullah, 1984
Host: Carangidae: Carangoides malabaricus (Bloch)
Locality: VSK, BOB
References: Madhavi (1974a); Hafeezulla & Dutta (1998)
Fig. 16.37 Pseudopisthomonorchis

carangi (after Madhavi,
1974)
Description Body elongate, 1220–2100 long, 256–384 wide, lanceolate, spined.

Eyespot pigment present in oesophageal region. Oral sucker circular. Ventral sucker
larger than oral sucker, pre- or postequatorial. Prepharynx present. Pharynx 25–50
in diameter. Oesophagus longer than prepharynx. Intestinal bifurcation well anterior
to ventral sucker. Caeca extend to posterior end of body. Testes single, subspherical,
postequatorial, situated in anterior part of posterior third of body. Cirrus-sac globular
or elliptical, thin-walled, partially overlapping ventral sucker, separated from testis
by uterine coils, encloses seminal vesicle, pars prostatica and spined cirrus. Genital
atrium spined. Genital pore median, posterior to ventral sucker, some distance from
ventral sucker. Ovary submedian, entire, at level of junction of cirrus-sac and testis.
Seminal receptacle absent. Uterus filling most of postovarian space partially overlaps
testis. Metraterm unarmed, opening into genital atrium directly. Eggs 15–27 by 9–12,
filamented. Vitelline follicular, follicles large and irregular in shape, circumcaecal,
extend from posterior level of ventral sucker to some distance anterior to it, confluent
anteriorly.
Remarks: Madhavi (1974) erected the genus Pseudopisthomonorchis with P. caranxi

Madhavi, 1974, as the type species. The genus is characterized by the position of
genital pore near posterior margin of ventral sucker, absence of spines on the cirrus
and metraterm, the presence of spines in the genital atrium and eggs with filaments.
Pseudopisthomonorchis hanumanthai Gupta & Singh, 1981 (Fig. 16.38)
Host: Carangidae: Carangoides malabaricus (Bl. & Schn.)
Locality: PR, BOB
References: Gupta, P.C. & Singh, RB (1981)
Description Body elongate, 940–120 [sic] long, 170–230 wide with eyespot pig-
ments on either side of oesophagus. Tegument spinose, Oral sucker 40 long 30

hanumanthai (after Gupta &
Singh, 1981)
wide. Ventral sucker subspherical, postequatorial, larger than oral sucker, 50–60
long, 40–50 wide. Sucker ratio 1: 1.3–1.6. Prepharynx 80 long. Pharynx subglobu-
lar. Oesophagus much longer than prepharynx. Caeca simple, extend to posterior end
of testis. Genital pore submedian, posterior to ventral sucker. Testis single, entire,
ovoid, postequatorial. Cirrus-sac saccular. Seminal vesicle occupies proximal part of
cirrus-sac. Pars prostatica tubular, surrounded by prostate gland cells. Cirrus thick-
walled, opens into spinose genital atrium. Ovary entire, just pretesticular, median
or submedian. Uterus coiled, descends to some distance anterior to posterior end of
body, ascends to open at genital atrium. Vitellarium follicular, extend either from
intestinal bifurcation or just anterior to it to either anterior third of ventral sucker or
posterior margin of it. Eggs elongate, 24–28 × 9–11 with long unipolar filament.
Remarks: This species is distinct in the long prepharynx, the very long oesophagus
and in the postequatorial position of the ventral sucker.
Pseudopisthomonorchis secundus Ahmad, 1981
Host: Ephippidae: Platax teira (Forsskal); angel fish;
Locality: BOM, AS
Description: Body 2320–2456 long, 470–535 wide, elongated. Entire tegument

spinose. Eyespot pigments on either sides of oesophagus. Oral sucker 130–180 in
diameter. Ventral sucker 87–120 in diameter. spherical, pre-equatorial, smaller than
oral sucker. Sucker ratio 1:0.66. Intestinal bifurcation midway between pharynx and
ventral sucker. Caeca terminate blindly at middle of testis. Testis single, elongated, in
posterior third of body. Cirrus-sac saccular, curved, well posterior to ventral sucker,
much anterior to ovary, contains saccular seminal vesicle, prostate complex and
muscular cirrus armed with bristle-like spines opening into genital atrium. Genital
atrium spacious, spinose, spines triradiate. Genital pore median, closely posterior to
ventral sucker. Ovary three-lobed, pretesticular. Uterus descends to posterior end of
body, opens into posterior end of terminal organ. Terminal organ undivided, thick-
walled. Eggs oval, 36–42 long, 18–23 wide with long unipolar filament, 3–4 times
of egg. Vitellarium follicular, follicles large, symmetrical, 6–10 on each lateral side
in intestinal bifurcation level not forming transverse band anterior to ventral sucker.
Excretory vesicle tubular, reaches to ovary; pore terminal.
Remarks: Pseudopisthomonorchis secundus differs from P. carangi in several

respects: the ventral sucker smaller than the oral sucker, the longer oesopha-
gus, the caeca ending blindly at mid-testis level, an elongate testis, a shorter post-
testicular space, a three-lobed ovary partly overlapping the right caecum, the larger
eggs and in the absence of a prepharynx.
Pseudopisthomonorchis thapari (Varma & Singh, 1979) Bray & Justine, 2013
(Fig. 16.39)
[Syn: Opithomonorchis thapari Varma & Singh, 1979]
Host: Carangidae: Carangoides armatus (Ruppell) (1), C. chrysophrys (2)
Distribution: 1. Arabian Sea off Quilon; 2. New Caledonia
References: 1. Varma & Singh (1979); 2. Bray & Justine (2013)
Description: Body small, elongate, 1318–1438 by 149–183. Tegument spined. Eye-

spot pigment present. Oral sucker 38–40 × 37–44. Ventral sucker 32–39 × 48–59, at
about midbody. Sucker ratio 1: 1.31–1.33. Prepharynx long, narrow. Pharynx oval.
Oesophagus of similar length to prepharynx. Intestinal bifurcation in anterior fore-
body. Caeca narrow, terminal parts obscured by eggs. Testis single, oval, in anterior
half of hindbody. Cirrus-sac oval, overlaps testis posteriorly. Seminal vesicle large,

thapari (after Varma &
Singh, 1979)
oval, pars prostatica narrow, surrounded by large prostatic cells. Ejaculatory duct
short. Genital atrium narrow, genital pore submedian, closely posterior to ventral
sucker. Ovary rounded, pretesticular. Uterus mostly posterior to ovary. Eggs numer-
ous, small, 21–24 by 10–12, with indistinct aporal filament. Metraterm narrow dis-
tally. Vitellarium follicular, forming two closely adjacent fields in posterior forebody,
follicles few in number, highly irregular in shape.
Remarks: Hafeezullah (1984) synonymized the species with P. carangi Madhavi,

1974. However, Bray and Justine (2013) after examination of specimens of P. carangi
collected from Caranx chrysophrys expressed the view that P. carangi and P. thapari
may be distinguished by differences in body length, sucker ratio, previtelline distance
and postvitelline distance. Varma and Singh (1979) described the cirrus and genital
atrium as spined but Bray and Justine could not find them in their specimens.
Chapter 17
Superfamily Gorgoderoidea Looss, 1899
Family Gorgoderidae Looss, 1901
Members of the family Gorgoderidae Looss, 1901, are well known as endoparasites
of fishes, amphibians and reptiles. As endoparasites, they usually occur in the urinary
bladder of the host, more rarely in other locations such as swim bladder, coelom, gall
bladder, intestine. The diagnostic features of the family are:
Diagnosis: Body elliptical or divisible into narrow forebody and broad disc-like
hindbody. Body size small or large. Tegument smooth. Suckers well developed, oral
sucker smaller than ventral sucker. Pharynx present or absent. Caeca simple or form
cyclocoel. Genital pore in forebody, close to intestinal bifurcation. Gonads in hind-
body. Testes two or multiple. Male and female ducts lead into genital atrium. Male
terminal genitalia with external seminal vesicle, prostatic complex and ejaculatory
duct. Cirrus-sac present or absent. Ovary pretesticular. Seminal receptacle present
or absent. Vitellarium in two compact or dendritic masses. Uterus extends into hind-
body. Eggs small, operculate.
Looss (1901) recognized two subfamilies: the Gorgoderinae Looss, 1899 and
Anaporrhutinae Looss, 1901. The cladistic analysis including various gorgoderid
genera also supported the recognition of two subfamilies (Campbell, 2008). The
subfamilies are distinguished by the presence or absence of a pharynx, the position
of testes and the presence or absence of a cirrus-sac, Laurer’s canal and seminal
receptacle. Recent phylogenetic analysis by Cutmore et al. (2013) based on ITS2
and partial 28SrDNA data also revealed the existence of these two subfamilies and
another subfamily Degenerinae Cutmore, Miller, Curran, Bennett & Cribb, 2013. The
gorgoderids recorded from various Indian marine fish fall under the two subfamilies
Gorgoderinae and Anaporrhutinae
© Crown 2018 551

https://doi.org/10.1007/978-94-024-1535-3_17
552 17 Superfamily Gorgoderoidea Looss, 1899
Key to subfamilies:
1. Testes intercaecal, pharynx absent, seminal receptacle and cirrus-sac absent
.……………………………….…………… Gorgoderinae Looss, 1899
2. Testes overlap and extend laterally to caeca, pharynx present, seminal receptacle
and cirrus-sac present or absent ……………..................…… Anaporrhutinae
Looss, 1901
Subfamily Anaporrhutinae Looss, 1901
Diagnosis: Body usually rounded, flattened, may be divided into narrow forebody
and foliate hindbody. Oral sucker subterminal. Ventral sucker larger than oral sucker.
Pharynx present. Oesophagus short. Caeca long, sinuous, enlarged or diverticulate.
Gonads not restricted to intercaecal space. Testes two, variously lobed, or divided into
numerous follicles, overlapping caeca ventrally. Cirrus-sac absent. Ovary pretestic-
ular, between testes or post-testicular. Seminal receptacle present or absent. Uterus
usually confined to intercaecal space. Vitellarium two, multi-lobed or digitiform
masses, intercaecal or completely lateral to caeca. In urinary bladder or body cavity
of marine fishes. Type genus: Anaporrhutum Ofenheim, 1900.
Campbell (2008) provided a careful analysis of the various genera and species of
gorgoderids, suggested many synonymies and presented useful keys for separation
of valid genera. Later Cutmore et al. (2010) made a major contribution to the group
and subjected the various anaporrhutine genera into detailed analysis. From India,
nine species of anaporrhutine gorgoderids were reported from different species of
elasmobranch fishes occurring in the coastal regions of Bay of Bengal and Arabian
Sea. As a result of the revisions of the subfamily proposed by Campbell (2008) and
Cutmore et al. (2010), some of the Indian species have fallen synonyms of others
and there was also lot of reshuffling of the species from one genus to another. For
example, Staphylorchis gigas Samuel, 1951 and S. scoliodoni Mehra (1960) were
transferred to the genus Anaporrhutum by Campbell (2008), but Cutmore et al. (2010)
considered both these species as synonyms of Staphylorchis cymatodes. The genus
Neoanaporrhutum Gupta &Tewari, 1983 was synonymized with Petalodistomum
and the two species N. hanumanthai Gupta & Tewari, 1983 and N. singhi Gupta
&Tewari, 1983 were moved to this genus. Nagmia yamagutia Tandon, 1969 was
transferred to the genus Petalodistomum (Gupta &Tewari, 1983). Ultimately, the
number of valid Indian species has come down to seven, and they belonged to three
genera Anaporrhutum, Staphylorchis and Petalodistomum.
Key to Genera of Anaporrhutinae
1. Caeca without diverticula …………………… Petalodistomum Johnston, 1913
Caeca with numerous short diverticula …………………………………………2
2. Caeca slightly sinuous, testes follicles distributed in inter and extra caecal
space ………………………………………..Anaporrhutum Brandes in Ofen-
heim, 1900
Caeca strongly winding, testes follicles distributed in extracaecal space
…………………………………………………Staphylorchis Travassos, 1922
Family Gorgoderidae Looss, 1901 553
Genus Anaporrhutum Brandes in Ofenheim, 1900

[Syn. Staphylorchis Travassos, 1922; Winteria Lamothe-Argumedo, 1969 nec
Brauer, 1901]
Diagnosis: Body large, hindbody rounded, broad. Oral sucker large. Ventral sucker
comparatively larger. Prepharynx short. Pharynx muscular. Oesophagus short. Caeca
sinuous, without diverticula. Testes in two groups of spherical follicles distributed
partly intercaecal and partly extracaecal in hindbody. Cirrus-sac absent. Semi-
nal vesicle in forebody, conspicuous. Genital pore median, near intestinal bifur-
cation/pharynx. Ovary pretesticular. Seminal receptacle large. Uterus intercaecal.
Vitelline gland follicular or dendritic. Excretory vesicle I-shaped with long lateral
branches. Type species: A. albidum Brandes in Ofenheim, 1900.
Anaporrhutum albidum Brandes in Ofenheim, 1900 (Fig. 17.1)
Host: Torpedinidae: Narcine timlei (Bl. & Schn.)
Site: Body cavity
Location: MS, BOB
No. 5.
Distribution: Ceylon, Madagascar, Panama, Peru
Remarks: This species is characterized by the following features; caeca strongly

sinuous, testes in the form of numerous small follicles distributed in extracaecal
space.
Anaporrhutum gigas (Samuel, 1952) Campbell, 2008
[Syn. Staphylorchis gigas Samuel, 1952]
Host: Carcharhinidae: Galeocerdocuvier (Peron & Lesueur)
Reference: Samuel (1952)
Remarks: Synonymized with Staphylorchis cymatodes.
Fig. 17.1 Anaporrhutum

albidum (after Hafeezullah,
1971, courtesy Cambridge
univ. Press, J. Helm, 45, p. 81
Anaporrrhutum largum Luhe, 1906

Host: Stegosotmidae: Stegostoma faciatum (Hermann)
Locality: Orissa, BOB
Reference: Lühe (1906)
Anaporrhutum narayani Simha, Rao & Rao, 1971
Host: Myliobatidae: Mobula mobular (Bonnaterre)
Site: Bucco-pharyngeal region
Locality: North Kerala, Arabian Sea.
Reference: Simha et al. (1971)
Remarks: A. narayani differs from A. albidum by being larger, in the more anterior
distribution of testes and the form of the vitelline follicles.
Anaporrhutum scoliodoni (Mehra, 1960) Campbell, 2008
[Syn. Staphylorchis scoliodoni Mehra, 1960]
Host: Carcharhinidae: Scoliodon sorrakowah (Ruppell)
Reference: Mehra (1960)
Remarks: Synonymized with Staphylorchis cymatodes.
Anaporrhutum stunkardi Tandon, 1969 (Fig. 17.2)
Host: Carcharhinidae: Scoliodon dumerilii (Bleeker)
Locality: Bharat, India
Reference: Tandon (1969)
Genus Petalodistomum Johnston, 1913
[Syn. Nagmia Nagaty, 1930; Pernagmia Nagaty & Abdel_Aal, 1961; Nagmioides
Dollfus, 1971, Neoanaporrhutum Gupta &Tewari, 1983 (Reference: Campbell,
2008)]
Fig. 17.2 Anaporrhutum

stunkardi (after Tandon,
1969)
Diagnosis: Body broad, flat. Oral sucker smaller than ventral sucker. Ventral sucker
pre-equatorial. Prepharynx lacking. Pharynx present. Oesophagus short. Caeca with
numerous irregular branches.
Petalodistomum hanumanthai (Gupta & Tewari, 1983) Campbell, 2008 (Fig. 17.3)
[Syn. Neoanaporrhutum hanumanthai Gupta &Tewari, 1983]
Host: Trygonidae: Trygon imbricatus (Day)
No. 2
Reference: Gupta &Tewari (1983)
Description: Body large, lanceolate, flat, 12.97–13.75 mm long, 7.72–9.34
mm wide. Forebody distinctly separated from hindbody. Oral sucker 730–810 long
1000–1230 wide. Ventral sucker pre-equatorial, 1610–1650 in diameter. Pharynx
muscular. Oesophagus short. Caeca wide, undulating, terminate at posterior extrem-
ity. Genital pore immediately post-bifurcal. Testes posterior to ventral sucker, partly
extracaecal, consist of numerous small follicles 11–16 on left and 19–22 on right
side. Seminal vesicle saccular, tripartite, just post-bifurcal. Cirrus-sac absent. Ovary
entire, spherical, median, close to or overlapping ventral sucker. Seminal receptacle
absent. Vitellarium consists of tubular acini in two groups having three acini in each
group, symmetrical, intercaecal and postovarian. Uterus intercaecal, extends up to
posterior end of body. Eggs 32–73 long 32–41 wide. Excretory vesicle Y-shaped
with side branches.
Remarks: This species is characterized by: the symmetrical testes, the grape-like
bunches of numerous small testicular follicles overlapping the caeca in the equatorial
or post-equatorial zone, the absence of a seminal receptacle and the dendritic vitelline
acini situated posterior to the ovary. Based on these characters Gupta & Tewari (1981)
erected a new genus Neoanaporrhutum Gupta & Tewari, 1983 with N. hanumanthai
Gupta & Tewari, 1981 as the type species. However, the genus Neoanaporrhutum
was synonymized with Petalodistomum Johnston, 1913 by Campbell (2008).
Fig. 17.3 Petalodistomum

hanumanthai (after Gupta &
Tewari, 1983)
Fig. 17.4 Petalodistomum

singhi (after Gupta &
Tewari, 1983)
Petalodistomum singhi (Gupta & Tewari, 1983) Campbell, 2008 (Fig. 17.4)
[Syn. Neoanaporrhutum singhi Gupta &Tewari, 1983]
Host: Trygonidae: Trygon imbricatus (Day)
No. 2
Reference: Gupta &Tewari (1983)
Description: Body large, lanceolate, 5.14–5.42 mm long 2.96–3.64 mm wide. Oral

sucker 290–450 long 440–660 wide. Ventral sucker large, pre-equatorial, 630–850 in
diameter. Pharynx muscular. Oesophagus short. Caeca wide, undulating, terminate
just anterior to posterior end. Genital pore immediately post-bifurcal. Testes entirely
extracaecal, consist of symmetrical grape-like bunches of numerous small follicles
17–20 follicles on left and 13–19 on right side. Cirrus-sac absent. Seminal vesicle
tripartite, post-bifurcal. Ovary entire, oval, median, just posterior to ventral sucker.
Seminal receptacle absent. Vitellarium consists of tubular acini in two groups, hav-
ing 2 acini in each group, symmetrical, intercaecal, postovarian. Uterus intercaecal,
extends to posterior end of body. Eggs 18–28 by 18–23.
Remarks: This species closely resembles P. hanumanthai Gupta & Tewari, 1981, but
differs from it in having an oval ovary separated from the ventral sucker, in having
two acini in each vitelline group instead of three and in having the testes entirely
extracaecal.
Petalodistomum yamagutia (Tandon, 1969) Campbell, 2008
[Syn. Nagmia yamagutia Tandon, 1969]
Host: Himantura uarnak (Gmelin)
Locality: Bharat
No. 3
Ref. Tandon (1969a)
Genus Staphylorchis Travassos, 1922

This genus was reduced to subgeneric level under Anaporrhutum by Campbell
(2008), but Cutmore et al. (2010) restored it to its original generic status. Two species
of Anaporrhutum namely A. gigas Samuel, 1952, and A. scoliodoni Mehra, 1960,
were regarded as synonyms of S. cymatodes (Johnston, 1913) Travassos, 1922 the
type species of the genus Staphylorchis.
Diagnosis: Body broadly oval. Caeca strongly sinuous. Testes distributed extra-
caecally throughout entire length of hindbody. Cirrus-sac absent. Type species: S.
cymatodes (Johnston, 1913)
Staphylorchis cymatodes (Johnston, 1913) Travassos, 1922 (Fig. 17.5)
[Syn. S. gigas Samuel, 1952; S. scoliodoni Mehra, 1960; Anaporrutum gigas
(Samuel, 1952) Campbell, 2008; A. scoliodoni (Mehra, 1960) Campbell, 2008
Hosts: Carcharhinidae: Galeocerdo tigrinus Mueller & Henle (1); Scoliodon lati-
caudus Mueller & Henle (2)
Locality: Madras & Nagapattinam, BOB (1); Indian Ocean (2)
References: 1. Samuel (1952), 2. Mehra (1960)
Distribution: Indo-west Pacific, from off India in the Bay of Bengal to Moreton Bay,
in the Coral Sea.
Description: Body circular or elongate, size ranges from 12.8 to 47.4 mm. Suck-
ers more or less subequal, sucker ratio 1: 0.97–1.16. Caeca strongly sinuous, lack
diverticula, terminate close to posterior margin of body. Testes follicular, extracaecal,
95–119 in number, follicles round to oval, extend from level of vitellarium to almost
level with posterior margin of caeca. Seminal vesicle sinuous, situated anterior to
or dorsally to ventral sucker. Ejculatory duct short. Cirrus-sac absent. Ovary lobed,
median, Seminal receptacle variable in size. Vitellarium tubular, in two intercaecal
fields at level of ovary. Uterus extensive, extending posteriorly from ovary to level
with termination of caeca. Metraterm short, opens into genital atrium. Eggs 51–83 ×
34–51.
Fig. 17.5 Staphylorchis

cymatodes (after Samuel,
1952)
Remarks: Cutmore et al. (2010) based on a study of large numbers of specimens

collected from different species of elasmobranchs from Australian waters, found that
S. cymatodes has a high level of morphological variability even between specimens
from a single host and concluded that morphological differentiation based on few
specimens is not reliable. They supported the synonymy of S. scoliodoni and S. gigas
with S. cymatodes.
Subfamily Gorgoderinae Looss, 1899

[Syn. Xystretinae Yamaguti, 1958]
Diagnosis: Body pyriform or divided into narrow forebody and foliate hindbody.
Suckers in anterior part of body. Oesophagus short. Pharynx absent. Caeca terminate
blindly or form cylocoel. Gonads intercaecal. Cirrus-sac absent. Pseudosinus sac may
be present. Ovary pretesticular or paratesticular. Seminal receptacle absent. Uterine
coils confined to hindbody. Vitelline masses intercaecal, preovarian. Type genus:
Gorgodera Looss, 1899.
Key to Genera
1a. Caeca form cyclocoel, pseudosinus sac present, hindbody corrugated and
demarcated by folds ……………………………… Xystretrum Linton, 1910
1b. Caeca not united posteriorly, pseudosinus sac absent, hindbody broad, foliate
………………………………………………………Phyllodistomum Braun, 1899
Genus Xystretrum Linton, 1910
Diagnosis: Forebody tapered anteriorly. Hindbody oval in outline, folded or corru-

gated. Oral sucker terminal. Ventral sucker pre-equatorial. Pharynx absent. Oesoph-
agus short. Caeca forming cyclocoel. Testes symmetrical, in broadest part of hind-
body. Seminal vesicle tubular, Pseudosinus sac present. Cirrus-sac absent. Genital
pore post-bifurcal. Ovary immediately anterior to right testis. Seminal receptacle
absent. Uterus confined to intercaecal area of hindbody. Vitelline masses divided
into digitiform lobes, paired between ventral sucker and testes. Excretory vesicle I-
shaped. In urinary bladder of marine teleost fishes of tropical and subtropical regions.
Type species: X. solidum Linton, 1910
Six species of this genus have been recorded from marine fish of India, of which
two have been recorded from the Bay of Bengal and the rest from the Arabian Sea.
Xystretrum chauhani Ahmad, 1982 (Fig. 17.6)
Host: Balistidae: Sufflanmen capistratus (Shaw)
Locality: BOM, AS
No. 21
Description: Body flask-shaped with narrow forebody and wide hindbody,

1490–1555 long, forebody–hindbody ratio 1: 1.02–1.04. Sides of body with series of
two shallow lateral cup-like indentations beginning just posterior to ventral sucker,
two elevations of body occur on each side between indentations. Oral sucker 160–205
Fig. 17.6 Xystretum

chauhani (after Ahmad,
1982), a Xystretum manteri
(after Ahmad, 1982)
in diameter. Ventral sucker larger than oral sucker, 260–295 in diameter. Sucker ratio
1:1.22–1.44. Oesophagus thick-walled. Intestinal bifurcation nearer to oral sucker
than to ventral sucker. Caeca with prominent shoulders, form cyclocoel. Testes two,
smooth, diagonal, close together in mid-hindbody. Posterior testis larger than ante-
rior testis. Seminal vesicle saccular, in forebody. Pars prostatica surrounded by gland
cells, unites with metraterm to form hermaphroditic duct which penetrates into small
genital papilla that opens into saccular genital atrium via papilla. Genital pore ventro-
median, post-bifurcal. Ovary elongate, entire, anterior to posterior testis. Vitelline
masses compact, diagonally tandem, anterior vitelline mass submedian at mid-level
of ventral sucker, posterior vitelline mass immediately posterior to ventral sucker
and preovarian. Uterus extends posteriorly as far as cyclocoel.
Remarks: In the structure of terminal genitalia, this species resembles X. thapari but
differs from it in having the sides of the hindbody with a series of two lateral indenta-
tions and vitelline glands diagonally placed. Other important features of the species
are the smooth testes, the diagonally placed testes and the compact vitellarium.
Xystretrum manteri Ahmad, 1982 (Fig. 17.6a)
Host: Balistidae: Suffamen capistratus (Shaw)
Locality: Quilon, AS
No. 12
Description: Body flask-shaped, forebody 1490–1600 long, forebody-hindbody
ratio 1: 0.9–0.91. sides of hindbody with series of three lateral indentations, begin-
ning just posterior to ventral sucker, caused by groups of depressions at regular inter-
vals. Three slight elevations occur on each side of body between depressions. Oral
sucker spherical, 170–235 in diameter. Ventral sucker oval, larger than oral sucker, at
junction of fore and hindbody. Sucker ratio 1: 1.31–1.43. Oesophagus thick-walled.
Intestinal bifurcation nearer to oral sucker than to ventral sucker. Caeca from cyclo-
coel. Testes two, symmetrical, transversely elongate, smooth to indented, intercaecal.
Seminal vesicle bipartite, posterior chamber tubular, anterior chamber saccular. Pars
prostatica narrow tube surrounded by gland cells. Cirrus short opens into genital
atrium. Genital pore between ventral sucker and intestinal bifurcation. Ovary trans-
versely elongate, overlaps posterior border of ventral sucker, in between vitelline
glands, pretesticular. Vitellarium symmetrical, posterior to ventral sucker, deeply
lobed. Uterus in narrow coils, extends posterior to testes. Eggs oval, thin-shelled,
55–76 by 40–48.
Remarks: This species differs from X. overstreeti in having a smaller oesophagus,

a smaller sucker ratio, a transversely elongate ovary, a hindbody with three series
of lateral indentations, a bipartite seminal vesicle and the genital pore between the
ventral sucker and the intestinal bifurcation.
Xystretrum overstreeti Ahmad, 1982 (Fig. 17.7)
Host: Triacanthidae: Triacanthus brevirostris (Schlegel)
No. 9
Description: Body foliate, with narrow forebody and wide hindbody, 3804–4121
long. Forebody–hindbody ratio 1:1.08–1.15. Lateral sides of body with series of
four lateral indentations beginning just posterior to forebody caused by groups of
muscles on body wall, resulting in shallow cup-like depression at regular intervals,
fourth depression near posterior end of body, four slight elevations occur on each side
between depressions, transverse striae of body inconspicuous. Oral sucker 256–289
in diameter. Ventral sucker at junction of anterior and middle thirds of body, 600–687
Fig. 17.7 Xystretum

1982)
in diameter. Sucker ratio 1:2.35–2.37. Oesophagus thick-walled. Intestinal bifurca-

tion nearer to ventral sucker than to oral sucker. Caeca form cyclocoel. Testes two,
smooth, diagonal, separated by uterine coils, intercaecal. Seminal vesicle tubular,
anterior to ventral sucker, winding. Pars prostatica narrow, tubular, ventral to pro-
static vesicle. Cirrus short opens into genital atrium. Genital pore at intestinal bifur-
cation. Ovary spherical, immediately posterior to ventral sucker. Vitellarium deeply
lobed, symmetrical, immediately posterior to ventral sucker. Uterus in narrow coils,
extends posteriorly, then anteriorly, overlapping caeca. Eggs 30−33 × 20−23.
Remarks: This species differs from all the other species of the genus in having
the sides of the hindbody with a series of lateral indentations caused by groups of
radiating muscles. Other important features of the species are the very large and
thick-walled oesophagus, the diagonal testes and the bifurcal genital pore.
Xystretrum srivastavai Ahmad, 1982 (Fig. 17.8)
Host: Triacanthidae: Triacanthus brevirostris (Schlegel)
Locality: Goa, AS.
Number: 21
Fig. 17.8 Xystretrum

srivastavai (after Ahmad,
1982)
Description: Body lanceolate, undivided, 1124–1236 long, 365–489 wide, sides

of body with series of 3 cup-like indentations, beginning at level of anterior testis.
Oral sucker 138–182 in diameter, ventral sucker 180 to 230 in diameter, sucker ratio
1: 1.1.26–1.30. Oesophagus thick-walled, caeca form cyclocoel. Testes two, deeply
lobed, diagonal, in middle of hindbody. Seminal vesicle bipartite, posterior chamber
larger, anterior chamber entering prostatic vesicle. Cirrus thick-walled, opens into
genital atrium. Genital pore ventral, median. Ovary spherical, intertesticular. Vitelline
glands two, symmetrical, compact, lateral to ventral sucker. Uterus descending up to
cyclocoel, then ascends anteriorly. Eggs 50–60 by 36–38.
Remarks: This species is closest to X. overstreeti and X. manteri but differs from
both of them in the lanceolate body, testes deeply lobed, vitelline glands compact
and situated lateral to ventral sucker.
Xystretrum thapari Ahmad, 1982 (Fig. 17.9)
Host: Gerreidae: Gerres filamentosus (Cuvier).
Locality: Goa, AS
No. 7
Description: Body lanceolate, 1210–1389 long, 400–490 wide, sides of body with-
out lateral indentations. Oral sucker 160−245 in diameter. Ventral sucker smaller
than oral sucker, immediately pre-equatorial, 135–165 in diameter. Sucker ratio 1:
0.67–0.75. Oesophagus 90–140 long. Intestinal bifurcation nearer to oral sucker
than to ventral sucker. Testes smooth, diagonal, unequal in size, anterior testis sinis-
tral, posterior testis dextral, smaller than posterior testis, near posterior end of body.
Seminal vesicle in forebody. Pars prostatica surrounded by prostatic cells, unites with
Fig. 17.9 Xystretum thapari

(after Ahmad, 1982)
metraterm at base of hermaphroditic duct. Genital papilla small. Genital pore 30 to

35 post-bifurcal. Ovary tetra-lobed, intertesticular. Vitelline glands two symmetrical,
compact, in hindbody. Uterus fills hindbody. Eggs 45−55 × 26−32.
Remarks: This species is characterized by possessing a lanceolate body, having the

oral sucker larger than the ventral sucker, the smooth, diagonal testes and the tetra-
lobed intertesticular ovary.
Xystretrum triacanthi Ahmad & Gupta, 1985 (Fig. 17.10)
Host: Triacanthidae: Pseudotriacanthus strigilifer (Day)
Locality: PR, BOB
No. 2
Reference: Ahmad & Gupta, V (1985)
Description: Body flask-shaped, with narrow forebody, 1320–1760 long, 800–820

wide and rounded hindbody, 1970–2150 long, 1870–2170 wide. Total body
3290–3910 long. Forebody covered with scales. Oral sucker oval, subterminal,
470–550 long, 490–510 wide ventral sucker spherical, pre-equatorial, at junction
of fore and hindbody, larger than the oral sucker, 470–590 in diameter; aperture
eye-drop-shaped. Sucker ratio 1:0.96–1.15. Oesophagus 110–260 long. Intestinal
bifurcation dorsal, nearer to oral sucker than to ventral sucker. Caeca forming cyclo-
coel, near posterior end of body. Testes two, ovoid, symmetrical, well separated but
intercaecal, in middle of hindbody, Seminal vesicle bipartite, in forebody; posterior
part saccular, anterior part tubular, enters side of prostate vesicle. Pars prostatica nar-
row, tubular, surrounded by numerous prostate gland cells. Cirrus thick-walled, short,
opens into genital atrium. Genital papilla absent. Genital pore ventral, post-bifurcal.
Ovary entire, ovoid, submedian, to right of median line, overlapping anterior margin
of right testis. Vitelline glands two, ovoid, compact, symmetrical, median to ovary.
Fig. 17.10 Xystretum

triacanthi Ahmad & Gupta,
1985
Uterus extends posteriorly and fills post-vitelline space within cyclocoel. Metraterm
thick-walled, very long, coiled, opens into genital atrium. Eggs ovoid, 23–27 long,
17–19 wide. Excretory vesicle tubular; pore dorsal, subterminal.
Key to Indian species of the genus Xystretrum

1. Body surface smooth, without lateral indentations .............................……….. 2
Body with cup-shaped lateral indentations ……………............................…… 3
2. Body elliptical in shape, ovary lobed intertesticular ........... X. thapari Ahmad,
1982
Body divisible into fore and hindbodies, ovary pretesticular…… X. triacanthi
Ahmad& Gupta, 1985
3. Body lanceolate, testes deeply lobed ……....................…… X. srivastavai
Ahmad, 1982
Body with fore and hindbodies, Testes smooth
………........................……………… 4
4. Lateral indentations two in number ………..................…… X. chauhani Ahmad,
1982
Lateral indentations more than two ………….......................................………….
5
5. Body with series of lateral indentations, testes symmetrical, seminal vesicle bipar-
tite ………………. X. manteri Ahmad, 1982
Body with four indentations on each lateral side, testes oblique, seminal vesicle
tubular ……………………………… X. overstreeti Ahmad, 1982
Genus Phyllodistomum Braun, 1899
Diagnosis: Forebody narrow, hindbody foliate. Oral sucker terminal. Ventral sucker
pre-equatorial. Pharynx simple, caeca simple, terminate near posterior end of body.
Testes two, symmetrical or diagonal, in wider part of hindbody. Seminal vesicle
saccate, genital pore median, between ventral sucker and intestinal bifurcation. Ovary
pretesticular, Uterus fills most of space of hindbody. Vitellarium masses two, lobed.
Excretory vesicle I-shaped. Type species: P. folium (Olfers, 1816) Braun, 1899.
Species of Phyllodistomum occur in a wide range of marine and freshwater fishes.

It is one of the largest genera of digeneans infecting fishes, comprises over 110 species
including about 28 species recorded from marine fishes. Species of the genus are
characterized by having a more or less foliate hindbody, blind caeca, two testes and
the uterus strongly developed in hindbody (Campbell, 2008). Two species of this
genus were recorded from marine fishes of Indian waters.
Phyllodistomum lancea Mamaev, 1971 (Fig. 17.11)
Locality: VSK, BOB
Fig. 17.11 Phyllodistomum

lancea (Original)
Remarks: P. lancea was first described by Mamaev (1971) from the tunas Euthynnus
affinis and Auxis thazard from the South China Sea. It is distinguished from P. lewisi
Srivastava, 1938 by the large body size and the rounded testes. Madhavi & Sairam
(2000) reported it from the tuna Euthynnus affinis from Bay of Bengal.
Phyllodistomum lewisi Srivastava, 1938
Host: Liza macrolepis (Smith)
Locality: VSK, BOB
Remarks: This species has been reported from freshwater as well as marine fishes in
India, and the hosts belong to Belonidae, Channidae, Mastacembelidae and Mugili-
dae (Srivastava, 1938b; Rekhrani & Madhavi, 1985; Fotedar, 1969).
Chapter 18
Superfamily Microphalloidea Ward, 1910
Family Zoogonidae Odhner, 1902
Zoogonids are gastro-intestinal parasites of marine fishes. They are characterized

by a spinous tegument, a lateral genital pore, a restricted field of vitelline follicles
and the presence of a canalicular seminal receptacle. The family contains about 158
species in 33 genera (Bray 2008). Molecular analysis of a few species showed that
the family is a member of the superfamily Microphalloidea (Ward, 1901) (Cribb et al.
2001; Olson et al. 2003). Two subfamilies are recognized in the family: Zoogoninae
Odhner, 1902, with vitellarium reduced to compact masses and untanned eggs and
Lecithophyllinae Stossich, 1903 with vitelline follicles in lateral fields and tanned
eggs. In the Keys to the Trematoda (Volume 3) the account on Zoogonidae was
dealt by Bray (2008) where keys to subfamilies were provided and the relationships
between various genera were discussed. Earlier Bray (1987a, b) provided reviews on
the subfamilies Zoogoninae Odhner, 1902, and Lecithophyllinae. Molecular results
reported by Olson et al. (2003), Cutmore et al. (2014) and Sokolov et al. (2016)
indicate that the Zoogoninae is sister to the family Faustulidae and that the Lepido-
phyllinae is paraphyletic.
Seven species of zoogonids belonging to six genera have been recorded from the
marine fishes of Indian region, six of them from Visakhapatnam coast of Bay of
Bengal (Madhavi, 1979b).
Diagnosis: Body usually small, elongate or spatulate. Tegument spinose. Oral
sucker globular to infundibuliform. Ventral sucker small to large, simple or may
bear muscular lips, usually situated at midbody. Prepharynx short or long or
practically absent. Pharynx globular. Oesophagus long. Intestinal bifurcation in
forebody or anterior hindbody. Caeca two, rarely one, long and tubular or short and
saccular, terminate in forebody or reach close to posterior extremity. Testes two,
oval or lobed, symmetrical, oblique or tandem, in hindbody or posterior forebody.
Cirrus-sac well developed, saccular or elongate, straight or recurved. Seminal vesicle
internal, short, saccular or tubular, sinuous or bipartite. Pars prostatica prominent.
© Crown 2018 567
https://doi.org/10.1007/978-94-024-1535-3_18
568 18 Superfamily Microphalloidea Ward, 1910
Ejaculatory duct muscular, may form cirrus bearing spines. Genital atrium small
or large, surrounded by gland cells. Genital pore submarginal or marginal, usually
in forebody, occasionally in hindbody. Ovary entire or multi-lobed, in hindbody
or posterior forebody, usually pretesticular. Canalicular seminal receptacle present.
Uterus fills entire hindbody. Eggs may lack tanned capsules, i.e. membranous, or
have thin to thick capsules. Vitellarium small, single or two masses, or divided into
follicles in one median or in two lateral fields, in fore or hindbody. Excretory vesicle
saccular. Type genus Zoogonus Looss, 1901.
Key to subfamilies
1. Vitellarium in the form of one or two compact masses, egg capsule membranous
………………………………………………..……Zoogoninae Odhner, 1902
2. Vitellarium follicles in lateral fields, egg capsule thick, tanned ……………
……………………………..………Lecithophyllinae Stossich, 1903.
Subfamily Zoogoninae Odhner, 1902
Diagnosis: Body small, oval, spined. Ventral sucker size variable, simple or bearing
muscular lips. Testes entire, symmetrical to oblique, in hindbody or lateral to ventral
sucker. Seminal vesicle bipartite, occasionally tubular. Genital atrium small. Genital
pore marginal, in forebody or beside ventral sucker or in anterior hindbody, usually
sinistral. Ovary entire, in hindbody or dorsal to ventral sucker, pre or post-testicular.
Mehlis’ gland weakly developed. Uterus mainly post-gonadal. Egg capsules mem-
branous or weakly tanned. Vitellarium small, single mass or two small masses or
small follicles, in median region of hindbody. Excretory vesicle saccular. Type genus
Zoogonus Looss, 1901.
Key to genera:
1. Vitellarium masses two………………………Diphterostomum Stossich, 1903
Vitellarium mass single …………………………………………………2
2. Caeca short, saccular, divergent ……………………… Zoogonus Looss, 1901
Caeca long, slender, terminate in hindbody Zoogonoides Odhner, 1902.
Genus Zoogonus Looss, 1901
Diagnosis: Body small, pyriform. Tegument spined. Oral sucker infundibuliform,

larger than or similar in size to ventral sucker. Prepharynx distinct. Pharynx oval.
Oesophagus long, reaches posterior margin of ventral sucker. Intestinal bifurcation in
anterior hindbody. Caeca short, saccular, divergent reaching to mid-hindbody. Testes
large, symmetrical in anterior hindbody. Cirrus-sac short, elongate, reaches into hind-
body. Seminal vesicle bipartite. Pars prostatica vesicular. Genital pore marginal,
sinistral or dextral, at about level of ventral sucker. Ovary globular, post-testicular.
Uterus fills hindbody posterior to gonads. Eggs embryonated, egg capsule membra-
nous. Vitellarium forms single compact mass. Type species: Z. rubellus (Olsson,
1868).
Family Zoogonidae Odhner, 1902 569
Zoogonus argentopsi Madhavi, 1979 (Fig. 18.1)

Host: Sparidae: Argyrops spinifer (Forskal)
Locality: VSK, BOB
Description: Body fusiform, deep yellowish, 880–1040 long, 192–272 wide. Tegu-
ment of forebody spined, hindbody smooth. Oral sucker 66–96 by 78–98 in size. Ven-
tral sucker pre-equatorial, 117–136 in diameter. Sucker ratio 1: 1.3–1.7. Prepharynx
long. Pharynx globular. Oesophagus longer than prepharynx. Intestinal bifurcation
just posterior to ventral sucker. Caeca short, saccular, thick-walled, terminate behind
testes. Testes oval, symmetrical, situated near posterior border of ventral sucker. Gen-
ital pore sinistral, opposite anterior margin of ventral sucker. Cirrus-sac elongated,
strongly developed, curves round ventral sucker, from behind ventral sucker to gen-
ital pore on left, encloses bipartite seminal vesicle, long pars prostatica surrounded
by numerous prostatic cells and a long cirrus. Ovary smooth, situated close behind
intestinal bifurcation. Seminal receptacle small, lies posterior to ovary. Uterus con-
fined to post ovarian space. Eggs numerous, oval, eggshell membranous. Metraterm
long with thick muscular wall, opens into genital atrium in front of cirrus-sac. Vitel-
larium small compact mass situated just posterior to ovary.
Remarks: This species is characterized by the small fusiform body, the ventral sucker
much larger than oral sucker, the short caeca and the cirrus-sac extending beyond
posterior margin of ventral sucker.
Fig. 18.1 Zoogonus

argentopsi (after Madhavi,
1979)
Genus Zoogonoides Odhner, 1911
Diagnosis: Body small, pyriform. Tegument spined. Oral sucker globular, smaller or
equal in size to oral sucker. Ventral sucker at middle of body. Prepharynx short. Phar-
ynx globular. Oesophagus long. Intestinal bifurcation in posterior forebody. Caeca
reach to about level of testes or a little beyond. Testes oval, symmetrical, just pos-
terior to or at level of ventral sucker. Cirrus-sac claviform, short to long, straight
or curved. Seminal vesicle bipartite. Pars prostatica oval. Ejaculatory duct short to
long. Genital atrium distinct. Genital pore sinistral, lateral in mid-forebody. Ovary
oval, between testes or slightly posterior. Seminal receptacle present. Vitellarium a
single mass, between ovary and seminal receptacle. Uterus fills much of hindbody.
Metraterm muscular. Miracidia in membranous capsules. Type species: Z. viviparus
(Olsson, 1868).
Zoogonoides pyriformis Pritchard, 1963 (Fig. 18.2)
[Syn: Zoogonoides synodi Yamaguti, 1970]
Host: Labridae: Oxycheilinus cheilinus diagramma (Lacepede) (1),
Locality: VSK, BOB
Geographic distribution: Pacific (Hawaii); Indian ocean (BOB)
References: Madhavi (1979b); 2, Pritchard (1963); 3. Yamaguti (1970)
Description: Body small, pyriform, 489–567 long, by 221–302 wide. Tegument

with sparse spines. Oral sucker rounded, subterminal, 80–112 × 88–112. Ventral
sucker at midbody, 147–208 by 141–188. Prepharynx short, pharynx ovoid, oesoph-
agus long, bifurcation anterior to ventral sucker. Caeca extend to posterior edge of
ventral sucker. Testes subsymmetrical, ovoid, dorsal or dorsolateral to ventral sucker.
Cirrus-sac long, slender, curved, extends obliquely from anterior margin of ventral
Fig. 18.2 Zoogonoides

pyriformis (after Pritchard,
1963)
sucker to genital pore, contains seminal vesicle in two spherical or ovoid parts, pro-
static vesicle, prostatic cells and cirrus. Genital pore sinistral, near level of intestinal
bifurcation. Ovary median, ovoid, near posterior edge of ventral sucker. Seminal
receptacle spherical, immediately postovarian. Uterus fills hindbody, extends for-
ward between testes and ventral sucker, expands into short metraterm. Eggs 37–48 ×
19–26 contain oculate miracidia. Compact vitelline body ovoid, lies ventral to ovary.
Excretory vesicle spherical.
Remarks: This species was originally described by Pritchard (1963) from Malacan-
thus hoedtii from Hawaii. Bray (1987a) considered Z. synodi Yamaguti, 1970, from
Synodus dermatogenys of Hawaii as a synonym of this species. The major distin-
guishing features of the species are the small pyriform body, the large ventral sucker,
the caeca terminating at level of the testes and the smaller eggs.
Bray (1987a) provided a key and constructed a cladogram for species of

Zoogonoides, which indicated a close relationship between Z. viviparus (Olsson,
1868) and Z. pyriformis.
Genus Diphterostomum Stossich 1903
Diagnosis: Body small, pyriform. Tegument spined. Oral sucker globular. Ventral
sucker in middle or posterior half of body, large, muscular or small, weakly devel-
oped. Prepharynx short. pharynx oval. Oesophagus distinct. Intestinal bifurcation in
forebody. Caeca very short, saccular, divergent terminate in posterior forebody or
just overlap ventral sucker. Testes oblique or symmetrical, posterior, lateral or antero-
lateral to ventral sucker. Cirrus-sac elongate, reflexed or not, reaches past anterior
margin of ventral sucker. Seminal vesicle bipartite. Pars prostatica vesicular. Ejac-
ulatory duct long. Genital atrium small, genital pore marginal or submarginal, in
forebody, usually sinistral, occasionally dextral. Ovary globular, to oval, submedian,
just anterior to or between testes. Uterus fills much of hindbody. Miracidium sur-
rounded by sac or very thin shell. Vitellarium forms two subequal masses. Excretory
vesicle small, saccular. Type species: D. brusinae (Stossich, 1888).
Diphterostomum indicum Madhavi, 1979 (Fig. 18.3)
Hosts: Haemulidae: Pomadasys maculatus Bloch, P. argenteus (Forskal), P. furcatus
(Bl. & Schn.); Sciaenidae: Johnius aneus (Bloch)
Locality: VSK, (BOB)
Description: Body small, oval, 768–832 long, 124–176 wide. Tegument spined,
spines thorn-like, prominent, on anterior part of body. Oral sucker funnel-shaped
58–78 by 71–96. Ventral sucker large, highly muscular, 129 to 175 in diameter, lat-
eral sides thickened like lips, protrusible. Forebody long, more than twice length
of hindbody. Pharynx spherical. Oesophagus very long, divides anterior to ventral
sucker into two short saccular caeca terminating in front of ventral sucker. Testes
oval, symmetrical or oblique, immediately posterior to ventral sucker. Genital pore
Fig. 18.3 Diphterostomum

1979)
lateral, usually sinistral occasionally dextral, situated opposite posterior part of ven-
tral sucker. Cirrus-sac large, strongly curved, extends from genital pore to level of
ventral sucker, slightly overlapping anterior part of ventral sucker, encloses bipar-
tite seminal vesicle, vesicular pars prostatica, numerous prostatic cells and long
eversible cirrus. Ovary small, situated dorsal to ventral sucker. Seminal receptacle
small, postovarian. Vitellarium in form of small oval masses, postovarian. Uterus
occupies entire hindbody. Metraterm muscular narrow. Eggs large, 54 by 23, with
membranous capsule. Excretory bladder spherical.
Remarks: The species is amphitypic in that the genital pore is either dextral or
sinistral. It closely resembles the type species D. brusinae (Stossich, 1888) Stossich,
1899, which has been recorded from a wide range of hosts and geographic localities
(see Bray, 1987), but differs from it in the much longer forebody, funnel-shaped oral
sucker and amphitypic nature of genital pore.
Subfamily Lepidophyllinae Stossich, 1903
[Synonyms: Lecithostaphylinae Odhner, 1911, Steganodermatinae Yamaguti, 1934;

Hudsoninae Campbell, 1975]
Diagnosis: Body small to large, oval or spatulate, spined. Intestinal bifurcation in

forebody. Caeca two. Genital pore in forebody. Eggs with tanned operculate shells.
Vitellarium forms two symmetrical fields of globular to subglobular follicles. Excre-
tory system usually I-shaped, occasionally Y-shaped. Type genus: Lepidophyllum
Odhner, 1902.
Key to genera of Lecithostaphylinae

1. Caeca short, saccular, divergent ……………………………. Brachyenteron
Manter, 1934
Caeca long, narrow, extend well into hindbody ……………………………..2
2. Vitellarium extend into forebody ………….………….. Deretrema Linton, 1910

Vitellarium confined to hindbody ………Lecithostaphylus Odhner, 1911.
Genus Deretrema Linton, 1910
Diagnosis: Body oval to pyriform. Tegument spined. Oral sucker smaller than ven-
tral sucker. Pharynx oval. Oesophagus long. Caeca reach to testes or beyond. Testes
oval symmetrical, in anterior hindbody. Cirrus-sac small, claviform. Seminal vesi-
cle tubular, coiled, saccular, bipartite. Pars prostatica weakly developed. Ejacula-
tory duct may form cirrus. Genital pore marginal, in mid-forebody. Ovary entire,
oval to globular, pretesticular. Seminal receptacle and Laurer’s canal present. Uterus
mostly post-gonadal. Metraterm weakly developed. Eggs numerous, tanned, opercu-
late. Vitellarium in posterior half of forebody, with relatively few follicles. Excretory
vesicle saccular. Type species: D. fusillus Linton, 1910.
Deretrema plotosi Yamaguti, 1940 (Fig. 18.4)
[Syns. Deretrema hoplognathus Yamaguti, 1940; D. carangis Yamaguti, 1970; D.
dermacetabulum Madhavi, 1979]
Host: Echeneidae: Echeneis naucrates (Linneaeus)
Locality: VSK, BOB
Distribution: NW Pacific (Japan), Mid-Pacific (Hawaii), Indian Ocean (Bay of Ben-
gal)
Fig. 18.4 Deretrema plotosi

Description: Body elliptical, 3680 by 960. Entire tegument spinose, spines dense
anteriorly. Oral sucker 215 in diameter. Ventral sucker large, 596 × 608, at junction
of middle and anterior third of body, aperture large, circular, tegumental covering
extends over ventral sucker in form of flap so as to form another aperture overlying
aperture of ventral sucker. Sucker ratio 1:2.8. Prepharynx short. Pharynx 117 in
diameter, anterior part covered with minute triangular spines. Oesophagus long.
Intestinal bifurcation midway between suckers. Caeca wide, terminate at middle of
hindbody. Testes spherical, symmetrical, extracaecal, just posterior to ventral sucker.
Cirrus-sac small, thin-walled, extends obliquely from midway between ventral sucker
and bifurcation, to left margin at bifurcal level. Seminal vesicle occupies 2/3rd of
cirrus-sac. Pars prostatica small, prostatic cells few. Ejaculatory duct short narrow.
Genital pore at left body margin at bifurcal level. Ovary subglobular, pretesticular.
Seminal receptacle small. Vitelline follicles large, ovoid, 9 on left and 12 on right,
extend from just post-bifurcal to middle of ventral sucker. Eggs 36–39 by 20–22.
Remarks: This species was originally described as D. dermacetabulum by Madhavi

(1979) characterizing it by the presence of a tegumental flap covering ventral sucker.
Bray (1987b) regarded this species as well as D. haplognathus Yamaguti, 1940,
and D. caranxi Yamaguti, 1970, as synonyms of D. plotosi stating that the size of
the ventral sucker which was used for differentiation of these species is a variable
character for the species.
Genus Lecithostaphylus Odhner, 1911
[Syn. Cypseluritrema Yamaguti, 1970, Manteroderma Skrjabin, 1957; Steganoderma
(Opisthoarchiotrema) Gupta & Sharma, 1974].
Diagnosis: Body oval to elongate oval. Tegument spinous. Oral sucker small, glob-
ular. Ventral sucker rounded, about equal to oral sucker or larger, pedunculate, but
may be withdrawn.
Prepharynx short, distinct. Pharynx small to large. Oesophagus absent to long.

Intestinal bifurcation in mid- to posterior forebody. Caeca pass testes or reach close to
posterior extremity. Testes oval, symmetrical or oblique, in anterior or mid-hindbody.
Cirrus-sac claviform, straight or curved. Seminal vesicle saccular to elongate, straight
or coiled. Pars prostatica short, vesicular. Ejaculatory duct short. Genital atrium
small. Genital pore sinistral, in mid- to anterior forebody, lateral to sublateral. Ovary
globular to irregularly oval, pretesticular, median or submedian, in anterior hindbody
or overlaps ventral sucker. Uterus in hindbody, mainly post-testicular, but may have
significant pretesticular portion. Metraterm muscular. Eggs numerous, operculate,
tanned. Vitelline fields symmetrical, about eight to 16 regular follicles, mainly in
lateral regions of anterior hindbody. Excretory vesicle saccular, short; pore terminal.
Type species L. retroflexus (Molin, 1859).
This genus has been considered a subgenus of the genus Steganododerma but
Bray (2008) in his revised key to the family recognized this genus a separate taxon
characterized by its pedunculate ventral sucker and the caeca reaching past the testes.
Lecithostaphylus nitens (Linton, 1898) Manter, 1947 (Fig. 18.5)

[Syn. Steganoderma (Opisthoarchiotrema) indicus Gupta & Sharma, 1972]
Host: Belonidae: Xenentodon cancila (Hamilton)
Distribution: NW Atlantic (Curacao, Puerto Rico, Florida, Massachussetts), Pacific
(Hawaii, Xisha Island [China]), Indian Ocean
References: Linton (1940), Manter (1947); Yamaguti (1970); Gupta, A.N. & Sharma
(1972b); Gu & Shen (1983)
Description: Body elongate, 4275–4770 long, 900–1050 wide. Tegument smooth.

Oral sucker spherical, 225–300 × 300–375. Ventral sucker bigger than oral sucker,
surrounded by puckered body wall, 425–475 × 595–625 pre-equatorial. Sucker ratio
1:1.6. Prepharynx small. Pharynx globular. Oesophagus short distinct. Caeca wide,
long terminate at posterior end of body. Testes post-equatorial, ventral to caeca,
symmetrical. Cirrus-sac tubular, between intestinal bifurcation and anterior margin
of ventral sucker, encloses elongated seminal vesicle, prostatic vesicle and cirrus.
Ovary pretesticular, median. Uterus fills most of posterior part of body. Vitellarium
follicular, divided into several large follicles arranged laterally mostly in linear fash-
ion, extend from posterior to ventral sucker to post-testicular region, 9–11 follicles
on each side. Eggs 30 × 15.
Remarks: Gupta and Sharma (1972b) described this as a new species in the genus
Steganoderma and erected a new subgenus Opisthoarchiotrema Gupta & Sharma,
1972 as Steganoderma (Opisthoarchiotrema) indicum. In a revision of the Lecitho-
phylinae, Bray (1987) regarded the subgenus Opisthoarchiotrema as a synonym of
Lecithostphylus and synonymized. Steganoderma (Opishorchiotrema) indicus with
Steganoderma (Lecithostaphylus) nitens.
Fig. 18.5 Lecithostaphylus

nitens (=Steganoderma
indicus) (after Gupta &
Sharma, 1982)
Lecithostaphylus parexocoeti (Manter, 1947) Yamaguti, 1971(Fig. 18.6)

[Syn: Steganoderma parexocoeti Manter, 1947, Manteroderma parexocoeti (Manter,
1947) Skrjabin, 1957; Steganoderma (Opisthoarchitrema) parexocoeti Manter, 1947;
Manteroderma skrjabini Parukhin, 1971, Deretrema fusillus of Linton, 1940 in part;
Cypseluritrema robustum Madhavi, 1979 Cypseluritrema prognichthydis Shen &
Li, 2000 (synonym according to Liu et al., 2010) Cypseluritrema spilonotopteri
Yamaguti, 1970 (synonym according to Bray (1986)
Host: Exocoetidae: Cypselurus comatus Mitchill

Locality: VSK, BOB
Distribution: NW Atlantic (Florida, Massachusetts) Pacific (Hawaii, Japan); Indian
ocean (Red Sea, Gulf of Aden; Bay of Bengal)
References: Manter (1947); Bray (2008); Yamaguti (1970); Parukhin (1971); Mad-
havi (1979b)
Description: Body small, plump, triangular, 1520–2000 long, 800–976 wide. Tegu-
ment smooth. Oral sucker 175–234 in diameter. Ventral sucker 218–245 in diameter,
strongly muscular, protrusible, on short peduncle, close to anterior end. Sucker ratio
1: 1.08–1.3. Prepharynx absent. Pharynx globular. Intestinal bifurcation immediately
anterior to ventral sucker. Caeca broad, simple, run parallel to one another, terminate
short of posterior extremity. Genital pore to left of midline, at level of oesophagus.
Testes two, spherical, diagonal, in anterior part of hindbody. Cirrus-sac well devel-
oped, stout, elliptical, runs obliquely from genital pore to just posterior to ventral
sucker, reaches to level of ovary, encloses elongate seminal vesicle, long pars prostat-
ica surrounded by numerous prostatic cells and short cirrus. Ovary spherical, slightly
pretesticular. Seminal receptacle postovarian. Vitellarium with 14–20 large follicles,
arranged in two groups, on either side of ovario-testicular zone. Uterus occupies
entire post-testicular space. Eggs oval, thick-shelled, embryonated, 32–35 × 20–23.
Fig. 18.6 Lecithostaphylus

parexocoeti (after Madhavi,
1979)
Remarks: Madhavi (1979b) identified the forms collected from Cypselurus comatus
from BOB as belonging to a new species under the genus Cypseluritrema Yamaguti,
1970 as C. robustum Madhavi, 1979. Bray (2008) regarded the genus Cypseluritrema
a synonym of Lecithostaphylus Odhner, 1911.
Genus Brachyenteron Manter, 1934
Syn. Cypseluritrematoides Yamaguti, 1970; Prolateroporus Yamaguti, 1971
Diagnosis: Body fusiform to pyriform. Tegument spined or smooth. Oral sucker

infundibuliform. Prepharynx distinct. Pharynx oval. Oesophagus long. Caeca short,
saccular, terminating in forebody or at level of ventral sucker. Ventral sucker in about
mid-region of body, larger or subequal to oral sucker. Testes subglobular, smooth,
symmetrical, close to or overlapping ventral sucker. Cirrus-sac claviform, elongate.
Seminal vesicle tubular, bipartite or straight or coiled, pars prostatica vesicular, oval to
elongate, cirrus long. Genital pore sinistral, in mid-forebody. Ovary globular, pretes-
ticular, overlapping ventral sucker, seminal receptacle and Laurer’s canal present.
Uterus fills most of hindbody. Metraterm muscular. Eggs numerous, tanned, oper-
culate. Vitellarium in two lateral fields of follicles, mainly in forebody or al level of
ventral sucker. Excretory vesicle saccular. Type species: B. peristedioni Manter, 1934.
Brachyenteron parexocoeti Manter, 1947 (Fig. 18.7)
[Syns: Cypseluritrematoides triangularis Yamaguti, 1970; Cypseluritrematoides

longicirratus Madhavi, 1979; C. minor Gu & Shen, 1970]
Host: Exocoetidae: Cypselurus comatus (Mitchill)
Number: one
Locality: VSK coast, BOB
Distribution: NW Atlantic, Mediterranean Sea, Red Sea, Aden Bay, Bay of Bengal,
Arabian Sea, Hawaii, Japan, China.
Description: Body small, triangular, 1200 long, 729 wide. Tegument smooth. Oral
sucker bowl-shaped, 101 × 132 in size. Ventral sucker large, with wide opening, 304
Fig. 18.7 Brachyenteron

parexocoeti (after Madhavi,
1979)
by 292 in size, slightly anterior to equator situated on short peduncle giving body
triangular appearance. Prepharynx absent. Pharynx large, prominent, with indented
anterior border. Caeca short, saccular, terminating at mid-level of ventral sucker.
Genital pore at level of base of pharynx. Testes two, elliptical, anterior testis dor-
sal to ventral sucker, posterior testis slightly below and posterior to ventral sucker.
Cirrus-sac long, slender, extends from genital pore to posterior testis, major part of
it occupied by elongate seminal vesicle, small pars prostatica, few prostatic cells,
long eversible cirrus. Ovary spherical, situated at level of posterior testis near caecal
ends. Vitellarium forming two symmetrical groups of nine follicles, situated sym-
metrically in front of ovary and on either side of ventral sucker. Eggs oval, 31 by 19,
with a thick shell.
Remarks: This species is characterized by a relatively large, protrusible ventral

sucker. Has a wide geographic distribution, showing specificity to exocoetid fishes
as hosts.
Family Faustulidae Poche 1926
(Syn. Baccigeridae Yamaguti, 1958)

Faustulids are parasites of marine, fresh and brackish water fish and are char-
acterized by a spiny tegument, a canalicular seminal receptacle and more or less
extensive uterine coils. The family was for a long time included as a subfamily in the
Fellodistomidae. The molecular study of the family Fellodistomidae Nicoll, 1909,
by Hall et al. (1999) indicated that the family was polyphyletic and recognized one
of its subfamilies at the family level, for which the name Faustulidae Poche, 1926,
was already available.
Diagnosis: Body small, oval or fusiform. Tegument spined or smooth. Oral sucker
smaller or larger than ventral sucker, Prepharynx short. Pharynx globular. Oesopha-
gus distinct. Caeca usually short, reach testes region or just beyond. Testes two, sym-
metrical usually in anterior hindbody. Cirrus-sac thin-walled, occasionally absent.
Seminal vesicle bipartite or saccular. Pars prostatica long surrounded by numerous
prostatic cells. Ejaculatory duct short, narrow. Genital pore median or submedian,
in forebody. Ovary oval or lobed, pre- inter- or post-testicular. Canalicular seminal
receptacle present. Uterus extensive, mostly post-testicular. Eggs numerous. Vitellar-
ium in two clusters, each with one or several follicles. Type genus: Faustula Poche,
1926.
Sixteen species belonging to eight genera were recorded from India.
Key to Genera
1. Genital pore marginal ……………………………. Yamagutia Srivastava, 1937

Genital pore median or submedian…………………………………………… 2
Family Faustulidae Poche 1926 579
2. Testes and ovary in forebody ………………...………... Antorchis Linton, 1911

Testes and ovary in hindbody ………………………………………………….3
3. Ovary multi-lobed ……………………………………….. Faustula Poche, 1926
Ovary entire or with few lobes ……………………………….………………. 4
4. Ventral sucker larger than oral sucker, with lobes or muscular
ridges…………………………………… Paradiscogaster Yamaguti, 1934
Ventral sucker not greatly larger than oral sucker, of normal structure ….…… 5
5. Vitellarium in two groups of indistinct follicles, ovary pre or intertesticular.…. 6
Vitelline follicles condensed into two compact masses, ovary post-
testicular………………………Pronoprymna Poche, 1926
6. Seminal vesicle bipartite, vitelline follicles not closely massed together………7
Seminal vesicle unipartite, vitelline follicles in compact bunches
……Baccigeroides Dutta, 1995
7. Cirrus-sac present …….………………….………….… Bacciger Nicoll, 1914
Cirrus-sac absent ……………………Pseudobacciger Nahhas & Cable, 1964.
Genus Antorchis Linton 1910

[Syn. Parantorchis Yamaguti, 1934; Jonesiella Hafeezullah & Siddiqi, 1970; Neop-
arantorchis Hafeezullah & Siddiqi, 1970]
Antorchis Linton, 1910, was proposed as a replacement name for Mesorchis Linton,
1910, which was preoccupied. Three genera have subsequently been synonymized
with this genus: Parantorchis Yamaguti, 1934, was synonymized by Machida (1975).
Jonesiella proposed by Hafeezullah & Siddiqi (1970b) was found to be preoccupied
and replaced with Neoparantorchis by Hafeezullah & Siddiqi (1971) and then made
a synonym of Antorchis by Machida (1975).
Generic diagnosis: Body fusiform. Tegument armed with small spines. Oral sucker
terminal or subterminal. Ventral sucker in mid- or posterior to midbody. Intestinal
bifurcation in mid-forebody. Caeca short, saccular, reach posterior forebody. Testes
symmetrical usually in forebody. Cirrus-sac oval. Internal seminal vesicle oval, bipar-
tite or coiled. Pars prostatica distinct, prostatic cells numerous. Ejaculatory duct short.
Genital pore median, in forebody. Ovary oval, in posterior forebody. Canalicular sem-
inal receptacle present. Uterus fills most of hindbody. Vitellarium with few follicles
in lateral fields of forebody. Excretory vesicle V- or Y-shaped. Type species: A. urna
(Linton, 1910) Linton, 1911.
Two species of this genus are known from the Bay of Bengal: Antorchis inter-
medius (Madhavi, 1975) and Antorchis pomacanthi (Hafeezullah & Siddiqi, 1970).
Antorchis intermedius (Madhavi, 1975) Cribb, Anderson & Bray, 1999 (Fig. 18.8)
[Syn. Parantorchis intermedius Madhavi, 1975]
Host: Chaetodontidae: Chaetodon pictus Forskål
Locality: VSK, BOB
Number: 2
Fig. 18.8 Antorchis

intermedius (after Madhavi,
1975)
Description: Body oval, 912–1088 long, 442–576 wide at midbody. Tegument

spined. Forebody much longer than hindbody, 448–592 long. Suckers subequal.
Oral sucker 98–117 by 117–128. Ventral sucker slightly pre-equatorial, 137 in diam-
eter. Sucker ratio 1: 1.01–1.17. Prepharynx small. Pharynx and oesophagus present.
Intestinal bifurcation closer to oral sucker than to ventral sucker. Caeca slender, half
long, terminate posterior to testes. Genital pore median, near intestinal bifurcation.
Testes two, spherical, symmetrical, in forebody, almost at midbody. Cirrus-sac large,
thick-walled, placed obliquely from left of ventral sucker to intestinal bifurcation,
encloses small bipartite seminal vesicle, long pars prostatica, numerous prostatic
cells and small cirrus. Ovary ovoid, in forebody, intertesticular, close to right testis.
Seminal receptacle oval, small, lies dorsal to ventral sucker. Uterus occupies entire
post-testicular space forming V-shaped coils. Eggs 35–39 by 16. Vitellarium follicles
large, grouped together into two symmetrical bunches between intestinal bifurcation
and testes.
Remarks: This species is characterized by the combination of following characters:

almost subequal suckers, anteriorly placed gonads, large cirrus-sac, bipartite seminal
vesicle and somewhat large eggs.
Antorchis pomacanthi (Hafeezullah & Siddiqi, 1970) Machida, 1975 (Fig. 18.9)
[Syns. Jonesiella pomacanthi Hafeezullah & Siddiqi, 1970; Neoparantorchis
pomacanthi (Hafeezullah & Siddiqi, 1970b) Hafeezullah & Siddiqi, 1971; Paran-
torchis pomacanthi (Hafeezullah & Siddiqi, 1970b) Madhavi, 1975]
Host: Pomacanthidae: Pomacanthus annularis (Bloch)
Location: VSK, BOB
Distribution: Great Barrier Reef (Cribb et al. 1999);
References: Hafeezullah & Siddiqi (1970b, 1971); Madhavi (1975b); Cribb, Ander-
son & Bray (1999)
in diameter. Ventral sucker 143–218 in diameter, in posterior third of body. Sucker
Fig. 18.9 Antorchis

pomcanthi (after Hafeezullah
& Siddiqi, 1970, Courtesy
Allen Press, J. Parasitol, 56,
p. 934)
ratio 1: 0.70–0.95. Prepharynx indistinct. Pharynx oval. Oesophagus long. Caeca

club-shaped, short, not extending beyond middle of body. Testes spherical symmet-
rical, posterior to caecal ends, in forebody. Cirrus-sac anterior to ventral sucker,
straight or curved at distal end, encloses bipartite seminal vesicle, long pars prostat-
ica, numerous prostatic cells and cirrus. Genital pore post-bifurcal, median. Ovary
in testicular zone, between cirrus-sac and left testis. Seminal receptacle large, dorsal
or dorsolateral to ventral sucker. Uterus fills whole of post-testicular space. Eggs
29–45 × 15–19. Vitellarium follicular, in two lateral clusters from mid-oesophagus
level to caeca, extracaecal, Excretory bladder Y-shaped, arms reaching caecal ends.
Remarks: This species was first described by Hafeezullah & Siddiqi (1970b) from
Pomacanthus annularis from the Bay of Bengal. Later Cribb et al. (1999) reported
it from Pomacanthus semicirculatus and P. sextriatus from Heron Island, Great Bar-
rier Reef and provided a review on the genus Antorchis Linton, 1910, recognizing
seven species in the genus. Cribb et al. discussed the characters that differentiate A.
pomacanthi from the remaining species of the genus.
Genus Bacciger Nicoll, 1924
Bacciger Nicoll, 1924, was reviewed by Bray & Gibson (1980) and Dimitrov &
Bray (1994). Members of this genus show some specificity towards clupeoid fishes.
Two species of this genus have been recorded from India, but one of these has been
transferred to Baccigeroides as B. cochinensis (Hafeezullah & Siddiqi, 1970) leaving
only one Indian species in the genus, B. sardinellae Hafeezullah & Siddiqi, 1970.
Generic diagnosis: Body small. Tegument spinose. Suckers well developed. Phar-
ynx and oesophagus long. Caeca short, divergent, reach ventral sucker or testes.
Genital pore sinistral to intestinal bifurcation. Cirrus-sac present. Seminal vesicle
unipartite. Prostate gland cells present. Testes two, symmetrical, in anterior hindbody.
Ovary entire, pretesticular. Seminal receptacle present. Uterine coils fill two-third
of posterior part of body. Vitellarium follicular, symmetrical in anterior one-third of
body. Type species: Bacciger bacciger (Rudolphi, 1819) Nicoll, 1914.
Fig. 18.10 Bacciger

sardinellae (after
Hafeezullah & Siddiqi, 1970,
Bacciger sardinellae Hafeezullah & Siddiqi, 1970 (Fig. 18.10)

Host: Clupeidae: Sardinella fimbriata (Val.)
Locality: BOM, AS
No. 7
Description: Body small, 680–920 long, 250–400 wide, oval. Oral sucker 91–116
by 111–142 in size, larger than ventral sucker. Ventral sucker 84–99 long, 87–108.
Sucker ratio 1: 0.71–0.93. Caeca simple, reaching middle of body. Testes oval or
subspherical, symmetrical just behind ventral sucker. Cirrus-sac oval, dorsal or
anterodorsal to ventral sucker encloses bipartite seminal vesicle, indistinct pars pro-
statica and ejaculatory duct, mostly filled with prostate gland cells. Genital pore ante-
rior to ventral sucker. Ovary median, intertesticular. Vitellarium two lateral clusters
of small follicles, pretesticular, never reaching level of bifurcation. Uterus volumi-
nous occupying almost whole of hindbody. Eggs 23–30 by 15–18. Excretory vesicle
Y-shaped with short stem and long arms reaching mid-oesophageal level.
Remarks: This species differs from the known valid species of the genus Bacciger by
possessing the following combination of characters: Oral sucker bigger than ventral
sucker, longer caeca, testes and vitelline clusters never reaching ventral sucker.
Genus Baccigeroides Dutta, 1995
[Baccigeroides Manna & Dutta, 1996]
This genus was erected by Dutta (1995) with B. hafeezullai Dutta, 1995, as the type
species. The genus very closely resembles Bacciger Nicoll, 1944, but differs from
it in having a unipartite seminal vesicle situated extracaecally on right side of the
body, the genital pore near the pharynx and the ventral sucker equal in size to the
oral sucker.
Generic diagnosis: Body small. Tegument aspinose. Suckers well developed. Phar-
ynx and oesophagus long. Caeca short. Genital pore sinistral to pharynx. Cirrus-sac
present. Seminal vesicle unipartite. Prostate gland cells present. Testes two sym-
metrical. Ovary entire, pretesticular. Seminal receptacle present. Uterine coils fill
two-third of posterior part of body. Vitellarium follicular, symmetrical, in anterior
one-third of body. Type species B. hafeezullai Dutta, 1995.
Baccigeroides cochinensis (Hafeezullah & Siddiqi, 1970) Dutta, 1995 (Fig. 18.11)
[Syn: Bacciger cochinensis Hafeezullah & Siddiqi, 1970]
Host: Engraulidae: Thryssa mystax (Bl. & Schn.)
No. 2
Description: Body 1120–1190 long, 610–690 long, 610–690 wide. Oral sucker
78–116 in diameter. Ventral sucker 78–87 in diameter, situated 376–388 from ante-
rior end. Sucker ratio 1: 0.67–0.70. Caeca reach to middle of body. Testes entire,
symmetrical, just posterior to ventral sucker. Cirrus-sac ovoid, between pharynx and
ventral sucker, encloses saccular unipartite seminal vesicle, tubular pars prostatica,
indistinct prostatic cells and ejaculatory duct. Genital pore anterior to middle of
oesophagus, submedian. Ovary subglobular, almost equatorial, pretesticular. Semi-
nal receptacle dorsal to ovary and right testis. Uterus voluminous, filling whole of
posterior part of body. Eggs 22–29 by 15–22. Vitellarium in two lateral clusters,
each composed of six large follicles, in caecal zone. Excretory vesicle Y-shaped,
arms wide, reaching mid-level of oesophagus.
Remarks: According to Manna & Datta (1996), Bacciger cochinensis fits into the
genus Baccigeroides by the presence of a prebifurcal genital pore, a prebifurcal
cirrus-sac and the testes well posterior to the ventral sucker. Bray (2008a) expressed
the view that this species is extremely similar to the type species B. hafeezullai Dutta,
1995, and may be a senior synonym.
Fig. 18.11 Baccigeroides

cochinensis (after
Hafeezullah & Siddiqi, 1970,
Fig. 18.12 Baccigeroides

hafeezullai(after Manna &
Dutta, 1996)
Baccigeroides hafeezullai Dutta, 1995 (Fig. 18.12)

[Syn. Baccigeroides hafeezullai Manna & Dutta, 1996]
Host: Engraulidae: Setipinna phasa (Hamilton)
Locality: Chilka Lagoon
Number: 15
Description: Body small, oval, 760–810 long, 440–480 wide. Tegument aspinose.
Oral sucker terminal, 50–60 long, 60–90 wide, almost equal to ventral sucker. Ventral
sucker 60–80 in diameter. Sucker ratio 1:1. Caeca short, reach to level of testes. Testes
two, entire, symmetrical, oval, posterior to ventral sucker. Cirrus-sac ovoid, near
pharynx, and anterodorsal to intestinal bifurcation. Internal seminal vesicle saccular,
unipartite. Pars prostatica surrounded by prostatic gland cells. Ejaculatory duct short.
Genital pore submedian, dextral and pharyngeal in position. Ovary oval, anterior to
right ovary. Vitellarium follicular, forming bunches, eight follicles in each bunch,
symmetrical, anterodorsal to intestinal caeca. Uterus filling most of hindbody. Eggs
20–24 by 6–20.
Remarks: This species differs from B. cochinensis in the absence of a long oesoph-
agus and in the sucker proportions (1: 1.0 versus 1: 0.67–0.70).
Genus Faustula Poche, 1926
[Syn. Eurema MacCallum; Orientophorus Srivastava, 1935]
Diagnosis: Body oval. Tegument armed. Oral sucker subglobular to cup-shaped.
Ventral sucker rounded, in midbody. Prepharynx and pharynx present. Oesophagus
long. Intestinal bifurcation in posterior forebody. Caeca reach into anterior hindbody.
Testes two, symmetrical at ventral sucker level or in anterior hindbody. Cirrus-sac
claviform, in forebody or overlapping ventral sucker. Internal seminal vesicle tubular,
undulating. Pars prostatica not differentiated. Ejaculatory duct short. Genital pore
median, bifurcal or post-bifurcal. Ovary post-testicular, acinous multilobate. Seminal
receptacle canalicular. Uterus fills most of hindbody. Vitellarium follicular, in lateral
fields in region between mid-oesophagus and anterior hindbody. Excretory vesicle
V-shaped, arms extend anteriorly beyond bifurcation. Type species: F. keksooni,

(MacCallum, 1918) Poche, 1926.
Orientophorus Srivastava, 1935, was considered a synonym of Faustula by Yam-
aguti (1958) and the three species included by Srivastava (1935) under the genus
were transferred to the genus Faustula as F. brevichrus (Srivastava, 1935); F. ganget-
ica (Srivastava, 1935) and F. clupeae (Srivastava, 1935). Additional species reported
from marine fish from Indian waters are: F. chauhani Gupta & Srivastava, 1960, F.
basiri Hafeezullah & Siddiqi, 1970, and F. mandapamensis Simha, 1974. Among
these species F. clupeae (Srivastava, 1935) and F. chauhani were regarded as syn-
onyms of F. brevichrus (Reference: Simha, 1964). F. mandapamensis was syn-
onymized with F. gangetica (Reference: Hafeezullah & Dutta, 1998). Only three
Indian species remained valid, namely: F. basiri Hafeezullah & Siddiqi, 1970; F.
brevichrus (Srivastava, 1935) and F. gangetica (Srivastava, 1935). In addition, there
are many species of Faustula reported from freshwater fishes in India.
Faustula basiri Hafeezullah & Siddiqi, 1970 (Fig. 18.13)
Host: Clupeidae: Tenualosa ilisha(Hamilton)
No. 15
Reference: Hafeezullah & Siddiqi (1970)
Description: Body oval, 2140–2580 long, 1030–1100 wide. Tegument spined. Oral
sucker 143–161 long, 200–243 wide. Ventral sucker 203–280 by 195–245 in diame-
ter, almost equatorial. Sucker ratio 1: 0.87–0.9. Caeca simple, arcuate, extend slightly
posterior to midbody. Testes two symmetrical at level of ventral sucker, intercaecal.
Cirrus-sac flask-shaped, between caecal arch and ventral sucker, contains S-shaped
seminal vesicle, pars prostatica, ejaculatory duct and cirrus. Prostate gland cells well
Fig. 18.13 Faustula basiri
(after Hafeezullah & Siddiqi,
1970, Courtesy Allen Press,
developed. Genital pore immediately behind intestinal bifurcation. Ovary deeply

multi-lobed, post-equatorial. Seminal receptacle absent. Uterus occupies whole of
hindbody. Eggs 12–18 by 9–12. Vitellarium follicular, lateral, commencing from
about mid-oesophageal level extend to level of ovary. Excretory vesicle Y-shaped,
arms reaching oesophagus.
Remarks: In having the cirrus-sac anterior to the ventral sucker this species is close
to F. brevichrus but differs from it in its longer oesophagus, post-bifurcal genital pore
and more anterior and posterior extent of the vitellarium.
Faustula brevichrus (Srivastava, 1935) Yamaguti, 1958 (Fig. 18.14)
[Syn. Orientophorus brevichrus Srivastava, 1935; Orientophorus clupeae Srivas-
tava, 1935; F. clupeae (Srivastava, 1935) Yamaguti, 1958;F. chauhani Gupta P.D. &
Srivastava, C.B. 1960]
Host: Clupeidae: Tenualosa ilisha (L)
Reference: Srivastava (1935)
sucker 80–112 long 64–144 wide. Ventral sucker 128–160 in diameter, almost in
midbody. Intestinal caeca extend beyond testes posteriorly. Testes two, symmetrical,
on either side of ventral sucker. Cirrus-sac flask-shaped, overlaps ventral sucker partly
or wholly, encloses seminal vesicle, pars prostatica, ejaculatory duct and cirrus.
Genital pore at about level of intestinal bifurcation, sinistral. Ovary lobed, post-
testicular. Seminal receptacle absent. Uterus fills whole of hindbody. Eggs 15–16 ×
9–10. Vitellarium follicular, lateral, on either side of ventral sucker. Excretory vesicle
Y-shaped, arms reach oesophagus level.
Fig. 18.14 Faustula

brevichrus (after Srivastava,
1935)
Fig. 18.15 Faustula

gangetica (after Hafeezullah
& Siddiqi, 1970, Courtesy
p. 933)
Faustula gangetica (Srivastava, 1935) Yamaguti, 1958 (Fig. 18.15)

[Syn. Orientophorus gangeticus Srivastava, 1935; Faustula mandapamensis Simha,
1974]
Reference: Hafeezullah & Dutta (1998)
Host: Clupeidae: Tenualosa ilisha (Hamilton) 1,2,3; T. sinensis (4,5); Stromateidae:
Stromateus cinereus (6)
Locality: Allahabad (1), BOM; Veraval (2), AS; PR (3), DC (4, 5) BOB; Mandapam
(6)
References: 1. Srivastava (1935); 2. Hafeezullah & Siddiqi (1970b); 3. Gupta &
Ahmad (1977); 4. Kumari & Srivastava (1976); 5. Hafeezullah & Dutta (1998); 6.
Simha (1974).
sucker 124–190 long, 220–320 wide. Ventral sucker 200–350 in diameter, pre-
equatorial. Intestinal caeca extend almost to equatorial level. Testes two, symmetri-
cal at level of or slightly posterior to ventral sucker. Cirrus-sac flask-shaped, extends
beyond ventral sucker posteriorly. Genital pore immediately post-bifurcal. Ovary
lobed, post-equatorial. Seminal receptacle absent. Vitellarium follicular, lateral, com-
mencing from about mid-level of ventral sucker or slightly posterior to it to level
of posterior margin of ovary. Uterus occupies whole of post-testicular area. Eggs
15–17 × 7–11. Excretory vesicle Y-shaped, arms reaching oesophagus.
Differentiation of species of Faustula from Indian marine fish.

1. Genital pore beyond prebifurcal, cirrus-sac extends posteriorly as far as middle or
posterior margin of ventral sucker, vitelline follicles at level of ventral sucker…
…………………………………………..…...... F. brevichrus (Srivastava, 1935)
2. Genital pore immediately post-bifurcal, cirrus-sac extends distinctly posterior
to ventral sucker, vitelline follicles extend from about middle of oesophagus to
posterior margin of ovary ……………………… F. gangetica (Srivastava, 1935)
3. Genital pore immediately post-bifurcal, cirrus-sac extends posteriorly as far as

anterior margin of ventral sucker, vitelline follicles extend from about intestinal
bifurcation to level of ovary ………………..........................................................
F. basiri Hafeezullah & Siddiqi, 1970.
Genus Paradiscogaster Yamaguti, 1934

[Syns. Discogaster Yamaguti, 1934 nec Burmeister, 1835; Discogastroides Strand,
1934; Discogasteroides Srivastava, 1939; lapsus; Odontotrema Hafeezullah & Sid-
diqi, 1970 nec Lindholm, 1927; Odontocotyle Hafeezullah & Siddiqi, 1970; Pseu-
dodiscogasteroides Gupta, 1955] (Reference: Cribb et al. 1999).
Diagnosis: Body fusiform to pyriform. Tegument spined. Oral sucker simple. Ven-
tral sucker specialized as pad-like structure. Prepharynx short. Oesophagus variable
in length. Caeca short, divergent, saccular. Testes two, entire, in hindbody. Cirrus-sac
elongate oval to nearly spherical, internal seminal vesicle bipartite, pars prostatica
long, straight, prominent. Ejaculatory duct short, eversible. Genital pore between ven-
tral sucker and pharynx. Ovary entire, at level of testes. Vitelline follicles in region
of caeca, sometimes anterior to them. Uterus fills posterior half of body. Excretory
vesicle V-shaped, occasionally subglobular or inflated Y-shaped. Type species: P.
pyriformis Yamaguti, 1934.
Cribb et al. (1999) provided a review on the genus Paradiscogaster Yamaguti,

1934, discussed its various synonymies and gave a revised diagnosis of the genus
based on the proposed synonymies. Finally, they included 16 species under the genus
of which four species are known from India: P. arabi (Hafeezullah & Siddiqi, 1970), P.
farooqi Hafeezullah & Siddiqi, 1970, P. indicus Srivastava, 1939, and P. pritchardae
(Gupta & Ahmad, 1978).
Paradiscogaster arabi (Hafeezullah & Siddiqi, 1970) Cribb, Anderson & Bray, 1999
(Fig. 18.16)
[Syns. Odontotrema arabi Hafeezullah & Siddiqi, 1970; Odontocotyle arabi
(Hafeezullah & Siddiqi, 1970) Hafeezullah & Siddiqi, 1971]
Host: Drepanidae: Drepane punctata L.
Locality: Veraval, Calicut (AS) (1); VSK, BOB (3)
Reference: Hafeezullah & Siddiqi, (1970b)
Description: Body 760–2230 long 310–800 wide, elongate, pear-shaped. Tegument

with prominent scale-like spines. Oral sucker 119–262 by 89–221, sunken in anterior
part of body, with pair of tusk-like structures projecting into cavity from anterior
border. Ventral sucker 263–480 × 427–636, discoid, post-equatorial. Caeca short,
club-shaped. Testes oval or spherical, symmetrical, posterior to or postero-dorsal to
ventral sucker. Cirrus-sac elongate or fusiform, lies obliquely between ventral disc
and caecal arch, anterodorsal to ventral sucker, contains bipartite seminal vesicle,
indistinct pars prostatica surrounded by numerous prostatic cells and large muscular
eversible cirrus beset with spines. Genital pore post-bifurcal, submedian, to right.
Ovary spherical, median, pretesticular, postero-dorsal or dorsal to ventral sucker.
Fig. 18.16 Paradiscogaster

arabi (after Hafeezullah &
Siddiqi, 1970, Courtesy
p. 939)
Seminal receptacle oval. Uterus fills most of hindbody. Eggs 24–33 by 15, thick-
shelled. Vitellarium follicular, in two lateral clusters, anterior or anterodorsal to
ventral sucker.
Remarks: This species was originally included in the genus Odontocotyle Hafeezul-
lah & Siddiqi, 1971, which was proposed as a replacement name for the preoccupied
Odontotrema Hafeezullah & Siddiqi, 1970, that had been proposed for O. arabi
Hafeezullah & Siddiqi, 1970. The genus was distinguished from Paradiscogaster
Yamaguti, 1934, by the presence of a pair of tusk-shaped structures projecting into
the oral cavity of the oral sucker. Cribb et al. (1999) noted overall similarity of
this genus with Paradiscogaster and synonymized them. P. arabi has been recorded
from Drepane punctata from Veraval, Calicut (AS) and VSK, BOB (Hafeezullah &
Siddiqi, 1970b; Madhavi, 1975a).
Paradiscogaster farooqii Hafeezullah & Siddiqi, 1970 (Fig. 18.17)
Host: Scatophagidae: Scatophagusargus (L.)
Locality: Tuticorin, GOM
Distribution: India (1), South Africa (2), Kuwait Coast, Arabian Gulf (3), South-east
Queensland, Australia (4)
References: 1. Hafeezullah & Siddiqi (1970b); 2. Bray (1984); 3. Nahhas et al.
(1998); 4. Cribb et al. (1999)
Description: Body 1690–1810 long 460–540 wide, fusiform. Oral sucker 113–128
by 123–141, spherical. Ventral sucker 253–372 by 239–324, discoid, in middle third
of body. Caeca short, saccular, club-shaped. Testes oval or spherical, symmetrical,
or subsymmetrical, postero-lateral to ventral sucker. Cirrus-sac elongate, distal end
slightly curved to right, between intestinal bifurcation and ventral sucker, contains
bipartite seminal vesicle, long pars prostatica and protrusible cirrus, mostly filled

farooqii (after Hafeezullah &
Siddiqi, 1970, Courtesy
p. 933)
with prostate gland cells. Genital pore post-bifurcal, median or submedian. Ovary
spherical, postero-sinistral to ventral sucker, pretesticular. Seminal receptacle large,
spherical. Uterus voluminous, fills most of posterior part of body, metraterm undif-
ferentiated. Eggs 27–31 by 15–18. Vitellarium follicular, lateral, sparsely distributed
from level of intestinal bifurcation to ovary.
Remarks: This species has been recorded from many geographic areas including,
Natal in South Africa, Bay of Bengal in India, the Arabian Gulf and Moreton Bay in
Australia (Bray, 1984). It differs from P. arabi in the nature and sparse distribution
of vitelline follicles.
Paradiscogaster indicus (Srivastava, 1939) Cribb, Anderson & Bray, 1999
(Fig. 18.18)
[Syns Discogasteroides indicus Srivastava, 1939; D. caranxi Srivastava, 1939; Pseu-
dodiscogasteroides indicus (Srivastava, 1939) Gupta, 1955; P. caranxi (Srivastava,
1939) Gupta, 1955]
Host: Carangidae: Alepes djedaba (Hamilton); Triacanthidae: Triacanthus bilineatus
(Bloch); Pseudotriacanthus strigilifer (Cantor)
Locality: VSK, BOB
References: Srivastava (1939e); Gupta, S.P. (1955); Madhavi (1975b); Cribb et al.
(1999)
Description: Body plump, with its lateral margins prominently incised near anterior
margin of ventral sucker, yellowish brown in living condition, 860–980 long 540–680

indicus (after Srivastava,
1939)
wide. Tegument spined. Oral sucker 20–25 × 18, with slit-like opening. Ventral
sucker cup-like 20–28 × 32–40, just pre-equatorial. Caeca small, club-shaped, extend
in straight horizontal line anterior to ventral sucker. Testes two, spherical, 100–140
in diameter, symmetrical on either side of median line, close behind ventral sucker.
Cirrus-sac elongate, median, from just posterior to pharynx to posterior margin of
ventral sucker. Seminal vesicle bipartite, pars prostatica small, ejaculatory duct tubu-
lar, cirrus small. Ovary spherical, in contact with inner margin of right testis. Seminal
receptacle small, retort-shaped. Uterus occupies most of space in hindbody. Eggs
29–30 by 15. Vitellarium confined to space between caeca and middle of oral sucker.
Remarks: Srivastava (1939e) described two species under the genus Discogas-
teroides as D. indicus Srivastava, 1939, and D. caranxi Srivastava, 1939. Yamaguti
(1954, 1958) remarked that D. indicus cannot be assigned to the genus Discogas-
teroides. Gupta (1955) included both these species in a newly erected genus Pseu-
dodiscogasteroides Gupta, 1955, as P. indicus (Srivastava, 1939) Gupta, 1955, and
P. caranxi (Srivastava, 1939) Gupta, 1955, without mentioning the diagnostic fea-
tures of the genus. Later, Skrjabin & Koval (1957) gave diagnostic features of the
genus. Madhavi (1975a) accepted the proposal of Gupta (1955) and distinguished
the genus Pseudodiscogasteroides from Discogasteroides as follows: short plump
body, with a constriction in the middle, oral sucker with a slit-like opening, short
divergent caeca and long cirrus-sac overlying ventral sucker. Madhavi (1975a) syn-
onymized P. caranxi with P. indicus. However, Cribb et al. (1999) synonymized the
genus Pseudodiscogasteroides with Paradiscogaster expressing the view that none
of the characters used for the separation of these genera justifies the erection of a
new genus. P. indicus has so far been recorded from carangids and triacanthids from
Bay of Bengal.
Paradiscogaster pritchardae (Gupta & Ahmad, 1978) Cribb, Anderson & Bray, 1999
(Fig. 18.19)
[Syn. Odontocotyle pritchardae Gupta & Ahmad, 1978]
Host: Carangidae: Alepes djedaba (Hamilton)
Locality: PR, BOB
References: Gupta, V. & Ahmad (1978a); Cribb, Anderson & Bray (1999)

pritchardae (after Gupta &
Ahmad, 1978)
Description: Body elongate, with tubular anterior and broad posterior extremity,
1310–2345 long, 480–690 wide. Oral sucker 115–190 by 85–150 with a pair of tusk-
shaped structures associated with it. Ventral sucker discoid, 230–468 by 340–500.
Sucker ratio 1: 2.0–2.46. Caeca short, club-shaped. Testes two, symmetrical, smooth,
subequal, lateral, posterior to ventral sucker. Cirrus-sac thick-walled, saccular, lies
between ventral sucker and caecal arch. Seminal vesicle bipartite, pars prostatica
indistinct, surrounded by numerous prostatic cell. Cirrus muscular, wide, protrusible.
Genital pore median immediately post-bifurcal or just posterior to it. Ovary spherical,
postero-dorsal to ventral sucker, pretesticular, seminal receptacle postovarian. Uterus
fills most of hindbody. Eggs 37–43 by 13–18. Vitellarium in two lateral groups,
extend from level of intestinal bifurcation to just posterior to anterior margin of
ventral sucker.
Remarks: Cribb et al. (1999) suggested that P. pritchardae may be a synonym of P.

arabi (Hafeezullah & Siddiqi, 1970b).
Genus Pronoprymna Poche, 1926

[Pentagramma Chulkova, 1939; Pseudopentagramma Yamaguti, 1971]
Diagnosis: Body ovoid to elongate. Tegument smooth. Oral sucker terminal or sub-
terminal. Ventral sucker at junction of anterior and middle thirds of body. Prepharynx
absent. Pharynx small. Oesophagus distinct. Intestinal bifurcation in posterior fore-
body. Caeca extend to level of ovary. Genital pore median, post-bifurcal. Testes sym-
metrical, just posterior to ventral sucker. Cirrus-sac thin-walled, dorsal or postero-
dorsal to ventral sucker. Seminal vesicle bipartite. Prostatic cells few. Ovary tri-lobed.
Seminal receptacle and Laurer’s canal present. Vitellarium two compact masses, lat-
eral or just posterior to ventral sucker. Uterus fills most of hindbody. Eggs small.
Excretory vesicle V-shaped, arms reaching to pharynx. Type species: P. ventricosa
(Rudolphi, 1819) Poche, 1926.
This genus was first named as Pentagramma by Chulkova (1939) and included in
the family Steringophoridae. The name Pentagramma Chulkova, has been replaced
by Pseudopentagramma Yamaguti, 1971, but both names have been synonymized
with Pronoprymna Poche, 1926 (Bray & Gibson, 1980).
Pronoprymna petrowi (Layman, 1937) Bray & Gibson, 1980 (Fig. 18.20)
[Syn. Bacciger melanostomum Naidenova, 1970; Orientophorus petrowi (Layman,
1937) Margolis & Ching, 1965; Pseudopentagramma petrowi (Layman, 1937) Yam-
aguti, 1971]
Host: Clupeidae: Sardinella fimbriata (Val.); S. gibbosa (Bleeker)
Locality: VSK, BOB
Distribution: North Pacific rim from 3°N latitude on the coast of Japan to about 66°N
on the Bering Sea coast of the USSR and southward to about 49°N latitude on the
coast of British Columba (Margolis & Ching, 1965)
Description: With characters of the genus Pronoprymna. Body size 630–1459 ×

190–540. Sucker ratio 1: 0.71–1:1.08. Eggs 26–29 × 14–18.
Remarks: P. petrowi has a wide geographic distribution and has been recorded from
clupeoid and salmonoid fishes (Margolis & Ching, 1985). Madhavi (1975a) reported
the species from sardines of Visakhapatnam coast, Bay of Bengal. This constitutes
the only report from Indian Ocean.
Genus Pseudobacciger Nahhas & Cable, 1964
Nahhas & Cable (1964) separated species of Bacciger lacking a cirrus-sac and pro-
posed for them a new genus Pseudobacciger with P. harengulae (Yamaguti, 1938)
(Syn. Bacciger harengulae Yamaguti, 1938) as the type species.
Fig. 18.20 Pronoprymna

petrowi (after Madhavi
1975a)
Diagnosis: Body small. Tegument spinose. Ventral sucker in anterior half of body.
Pharynx and oesophagus present. Caeca short. Testes two, symmetrical, in hind-
body. Ovary intertesticular. Seminal receptacle present. Uterus mainly post-testicular.
Vitellarium in compact lateral masses at level of ventral sucker. Eggs small. Excretory
vesicle V-shaped. Type species: Pseudobacciger harengulae (Yamaguti, 1938).
This genus was differentiated from Bacciger mainly by the absence of a cirrus-sac.
Two species of this genus have been recorded from India: P. harengulae Yamaguti,
1938, and P. cablei Madhavi, 1975, both from the sardines Sardinella fimbriata
and S. gibbosa from Visakhapatnam coast, Bay of Bengal (Madhavi, 1975a). Sun
et al. (2014) sequenced a form they described as Pseudobacciger cheneyae Sun, Bray,
Yong, Cutmore & Cribb, 2014, and found that it appeared to belong on the superfam-
ily Gymnophalloidea and was, therefore, not in the Faustulidae (Microphalloidea).
We are retaining this genus in the Faustulidae until the type species of the genus is
convincingly demonstrated to belong elsewhere.
Pseudobacciger cablei Madhavi, 1975 (Fig. 18.21)

Locality: VSK, BOB
Description: Body ovoid, 528–704 long, 368–512 wide. Tegument smooth. Oral
sucker 39–62 by 58–74 in size. Oral sucker terminal, weakly muscular, 39–62 by
58–74 in size. Ventral sucker pre-equatorial, 215–273 from anterior end, 59–90 in
diameter. Sucker ratio 1: 1.0–1.2. Prepharynx absent. Pharynx 35–59 in diameter.
Oesophagus 73–97 long. Intestinal bifurcation midway between suckers. Caeca short,
wide, terminate in front of ventral sucker. Genital pore median, near intestinal bifur-
cation. Testes smooth, oval, symmetrical, equatorial on either side of ventral sucker.
Cirrus-sac absent. Seminal vesicle bipartite, in forebody, slightly overlapping ventral
sucker. Ovary three-lobed, median, intertesticular, overlaps posterior margin of ven-
tral sucker. Vitelline follicles large, in two extracaecal groups of 5–7 follicles each, at
about level of intestinal bifurcation. Vitelline ducts prominent, converging towards
ovary. Uterus occupies all post-testicular area, overlaps ovary and testes. Eggs small,
23–25 by 12–16. Excretory vesicle V-shaped with arms extending to level of testes.
Fig. 18.21 Pseudobacciger

cablei (after Madhavi 1975a)
Remarks: This species differs from P. harengulae in the larger body size, the more
posteriorly located ventral sucker, the seminal vesicle in the forebody, the larger
gonads, a three-lobed ovary, larger and more anteriorly located vitelline follicles and
the absence of gland cells in the forebody.
Pseudobacciger harengulae (Yamaguti, 1938) Nahhas & Cable, 1964 (Fig. 18.22)
Locality: VSK, BOB
Distribution: Japan, Korea, Florida, USA and Bimini, India, Namibian coast of East-
ern South Atlantic, Black Sea, Sweden.
Description: Body small, oval, 370–1130 long, 280–650 wide. Numerous gland
cells present in parenchyma of forebody. Oral sucker 45–70 in diameter. Ventral
sucker 40–55 in diameter, within anterior half of body. Intestinal bifurcation at level of
anterior margin of ventral sucker. Caeca terminate in pre, mid- or post-testicular level.
Testes ovoid, symmetrical, near caecal ends, slightly below ventral sucker. Seminal
vesicle bi-lobed, at level of mid-ventral sucker, anterior lobe larger than posterior
lobe. Ovary spherical, intertesticular. Vitellarium paired, lobed, pre-equatorial, at
level of ventral sucker. Seminal receptacle spherical, posterior to ovary. Uterus post-
testicular, overlaps testes. Eggs 21–25 by 18–23.
P. harengulae was originally described by Yamaguti (1938) and has been
redescribed by Dimitrov et al. (1999) and Rahimian & Thulin (2003). It has a cos-
mopolitan distribution in clupeoids especially in tropical and subtropical regions of
the Pacific and Indian oceans. Rahimian & Thulin (2003) furnished detailed infor-
mation on this species.
Genus Yamagutia Srivastava, 1939
Diagnosis: Body fusiform, unarmed. Oral sucker subglobular, ventral sucker trans-
versely oval. Intestinal bifurcation in mid-forebody. Caeca short, saccular, divergent,
terminate in forebody. Testes oval, oblique, near posterior extremity of body. Cirrus-
sac oval, transverse, in forebody. Seminal vesicle bipartite. Genital pore submarginal,
dextral, in forebody. Ovary pretesticular. Seminal receptacle present. Uterus occu-
pies much of hindbody. Vitellarium follicular, in lateral fields between bifurcation
and ventral sucker. Type species: Y. lateroporus Srivastava, 1937.
Fig. 18.22 Pseudobacciger

harengulae (after Madhavi
1975)
Fig. 18.23 Yamagutia
lateroporus (after Srivastava,
1939)
Yamagutia lateroporus Srivastava, 1939 (Fig. 18.23)

Host: Clupeidae: Nematolosa nasus (Bloch)
Locality: BOB
Reference: Srivastava (1939c)
Description: Body plump with broadly rounded anterior and posterior ends, 1600
long, 700 wide. Tegument aspinose. Oral sucker 220 × 160. Ventral sucker 450
n diameter, immediately post-equatorial. Prepharynx inconspicuous. Pharynx oval.
Oesophagus narrow, with small oesophageal bulb. Caeca short, saccular. Testes oval,
asymmetrical near lateral sides in hindbody. Cirrus-sac horizontal, club-shaped,
between caeca and ventral sucker. Seminal vesicle bipartite, pars prostatica elon-
gated oval. Ejaculatory duct small, cirrus fairly long. Genital pore just posterior to
anterior third of body length. Ovary spherical, slightly sinistral, partly overlapping
ventral sucker. Seminal receptacle present. Uterus occupies whole space between
genital pore and posterior end. Eggs 34–38 × 15–19. Vitellarium extends laterally
from level of oesophageal bulb to middle of ventral sucker.
Remarks: This is the only species under the genus Yamagutia Srivastava, 1939.
Chapter 19
Superfamily Opisthorchioidea
LaRue, 1957
Family Cryptogonimidae Ward, 1917
Members of the family Cryptogonimidae are usually small and occur in the intestine
and pyloric caeca of marine and freshwater fishes. Cryptogonimids do not possess
any characters that are specific to them. They are recognized by the combination of
characters: The body is small, with a spined tegument and some of them have spines
around the oral sucker. The suckers are well developed. The cirrus-sac is absent, and
a gonotyle may be present. The ventral sucker may be embedded in the genital pit.
The excretory vesicle is Y-shaped. Miller & Cribb (2008) provided a review of the
family.
Diagnosis: Body small to oval. Tegument spined. Eyespot pigment in forebody.
Oral sucker subterminal. Ventral sucker usually embedded in genital atrium. Phar-
ynx present. Oesophagus short. Caeca usually long, sometimes short. Testes usually
two, sometimes single or numerous, intercaecal or extracaecal. Seminal vesicle well
developed. Cirrus-sac absent. Gonotyle sometimes present. Genital pore in forebody.
Ovary pretesticular. Seminal receptacle present. Vitellarium follicular, grouped in
bunches mostly lateral, in middle third of body occasionally in forebody. Uterus
fills most of hindbody. Eggs with or without polar filament. Excretory vesicle V-
or Y-shaped arms reaching pharynx. Parasitic in intestine of marine and freshwater
fishes.
Type genus: Cryptogonimus Osborn, 1903
Eleven species of crytogonimids belonging to eight genera were reported from Indian
marine fish.
Key to genera
1. Circumoral spines absent ……………..…….……………….…………………2
Circumoral spines present………………………………………………………5
2. Testes numerous …………………………………… Polyorchitrema Srivastava,
1939
© Crown 2018 597
https://doi.org/10.1007/978-94-024-1535-3_19
598 19 Superfamily Opisthorchioidea LaRue, 1957
Testes 2 ………………………………………………………3
3. Caeca open out through separate ani …. Orientodiploproctodaeum Bhutta &
Khan, 1970
Caeca terminate blindly ……………………………………………………… 4
4. Body round, squat, uterus extensive in forebody, ovary post-testicular
…Allometadena Madhavi, 1976
Body oval or elongate, uterus occupies most of hindbody, ovary pretesticular
…………..……………………………………… Mehrailla Srivastava, 1939
5. Testes numerous…………………………… Acanthosiphodera Madhavi, 1976
Testes two …………………………………………………………………… 6
6. Ventral sucker similar in size or slightly larger than oral sucker, vitelline follicles
extend well posteriorly beyond testes …………… Neometadena Hafeezullah &
Siddiqi, 1970
Ventral sucker smaller than oral sucker, vitelline follicles do not extend beyond
testes ………………7
7. Gonotyle may be present ………………………… Siphoderina Manter, 1934
Gonotyle absent ……… Euryakaina Miller, Adlard, Bray, Justine& Cribb, 2010
Genus Acanthosiphodera Madhavi, 1976
Diagnosis: Body elongate. Tegument spined. Oral sucker surrounded by enlarged
spines. Ventral sucker embedded in genital atrium. Prepharynx, pharynx, oesoph-
agus present. Intestinal bifurcation in mid-forebody. Caeca long, blind. Testes 9,
distributed in posterior half of body. Gonotyle absent. Seminal vesicle sinuous.
Ejaculatory duct joins uterus before opening into genital atrium. Ovary follicular,
pretesticular. Seminal receptacle present. Uterus extensive. Vitelline follicles in two
lateral groups, posterior to ventral sucker and anterior to testis. Type species: A.
bengalense Madhavi, 1976
Acanthosiphodera bengalense Madhavi, 1976 (Fig. 19.1)
Host: Lutjanus malabaricus (Bloch & Schneider)
Locality: VSK, BOB
No. 22
Description: Body elongate, 1760−2640 × 640−896. Oral sucker 111−136 by
136−195, surrounded by single row of 46−48 spines, spines small, triangular, 10−12
long. Ventral sucker small, embedded in genito-atrial pit, 448−720 in diameter. Fore-
body with numerous deeply staining gland cells. Caeca long, terminate close to pos-
terior end. Testes 9, grouped together irregularly in posterior half of body. Gonotyle
absent. Seminal vesicle narrow, sinuous, commences just posterior to ventral sucker,
extends along left side of ventral sucker just anterior to it, opens into prostatic vesicle.
Ejaculatory duct short unites with metraterm to form hermaphroditic duct opening
into ventrogenital pit. Genital pore in midst of ventrogenital pit. Ovary follicular,
overlaps anterior testis. Seminal receptacle large, preovarian. Vitelline follicles in
two lateral fields, from midlevel of ventral sucker to midovarian level. Uterine coils
narrow. Eggs 16−22 × 8−9.
Family Cryptogonimidae Ward, 1917 599
Fig. 19.1 Acanthosiphodera

1976)
Remarks: The genus Acanthosiphodera Madhavi, 1976 differs from Siphodera (Lin-
ton, 1901) in having enlarged spines around the oral sucker. The testes are grouped
irregularly in Acanthosiphodera and occur in two parallel rows in Siphodera. In all
other respects, the two genera agree. A bengalenese is so far the only species in the
genus.
Genus Allometadena Madhavi, 1976
Diagnosis: Body small, rounded. Forebody with eyespot pigment and numerous
gland cells. Oral sucker large, without oral enlarged spines. Ventral sucker embedded
in parenchyma. Caeca long. Testes symmetrical, equatorial, in anterior hindbody.
Seminal vesicle long, sinuous. Genital pore median, in forebody. Ovary follicular,
follicles forming transverse bands posterior to testes. Seminal receptacle present.
Vitelline follicles dorsal, situated in hindbody. Uterine coils extensive filling forebody
also. Eggs oval. Type species: Allometadena rotundum Madhavi, 1976.
Allometadena rotundum Madhavi, 1976 (Fig. 19. 2)
Host: Lutjanidae: Lutjanus malabaricus (Bl. & Schn.)
Locality: VSK, BOB
No. 3
Description: Body small, round, 608−704 × 624−720. Tegument spined. Fore-

body with gland cells and eyespot pigment. Oral sucker large, 101−124 by 156−163.
Fig. 19.2 Allometadena

rotundum (after Madhavi,
1976)
Ventral sucker 58 in diameter, slightly pre-equatorial, embedded in parenchyma.

Sucker ratio 1: 0.35–0.37. Prepharynx absent. Pharynx spherical. Caeca narrow,
long. Testes 2, ovoid, symmetrical, equatorial, extracaecal, lie close to lateral edges
of body. Seminal vesicle large, sinuous, antero-dextral to ventral sucker. Genital pore
near anterior margin of ventral sucker. Ovary follicular, follicles numerous, of differ-
ent sizes and shape, scattered below testes. Seminal receptacle large. Uterine coils
extensive. Eggs 20 × 10. Vitellarium in two groups, separated by seminal receptacle,
extend from level of ventral sucker to slightly above posterior end.
Remarks: Genus Allometadena Madhavi, 1976 differs from Metadena Linton, 1910,
in the extracaecal testes and the vitellarium being confined to the hindbody. The genus
Exorchis Kobayashi, 1921, also has extracaecal testes, but the vitellarium is confined
to the forebody and the ovary is compact..
Genus Euryakaina Miller, Adlard, Bray, Justine& Cribb, 2010
Diagnosis: Body fusiform, oral sucker with few widely spaced spines. Ventral
sucker smaller than oral sucker. Caeca long. Testes oval, symmetrical or diagonal.
Seminal vesicle bipartite, extends postero-dorsal to ventral sucker. Ovary multilobed.
Uterus voluminous. Vitelline follicles small, extend from intestinal bifurcation to
anterior margin of testes. Type species: E. manilensis (Velasquez, 1961)
Euryakaina marina (Hafeezullah & Siddiqi, 1970) Miller, Adlard, Bray, Justine &
Cribb, 2010 (Fig. 19.3)
[Syn. Centrovarium marinum Hafeezullah & Siddiqi, 1970; Metadena marinum
(Hafeezullah & Siddiqi, 1970) Miller & Cribb, 2008]
Host: Lutjanide: Lutjanus fulviflamma Forsskal
Locality: Tuticorin, AS
No. 4
Reference: Hafeezullah & Siddiqi, (1970a); Miller & Cribb (2008); Miller et al.
(2010)
Description: Body 1364−1811 long, 447–706 wide, spindle-shaped. Tegument

aspinose. Eyespot pigment present. Oral sucker 122 −149 × 128−187. Ventral
Fig. 19.3 Euryakaina

marina (after Hafeezullah &
Siddiqi, 1970)
sucker 68−87 in diameter. Sucker ratio 1:0.5. Prepharyx long. Pharynx oval. Oesoph-
agus short. Caeca pass between testes, reach close to posterior extremity. Testes oval,
symmetrical or diagonal, lateral, postequatorial. Seminal vesicle saccular, bipartite,
extends postero-dorsal to ventral sucker. Prostate complex and cirrus absent. Genital
pore anterior to ventral sucker. Ovary divided into many follicles, median, equato-
rial. Seminal receptacle antero-dorsal to ovary. Uterus voluminous. Eggs13–19 by
9. Vitelline follicles small, between intestinal bifurcation and testes.
Remarks: Hafeezullah and Siddiqi (1970a) included this species in the genus Cen-
trovarium Stafford, 1904 as C. marinum Hafeezullah and Siddiqi, 1970. Miller and
Cribb (2008) based on similarity in morphology and host affinity included it in the
genus Metadena as M. marinum (Hafeezullah & Siddiqi, 1970). Recently, Miller
et al. (2010) based on morphology and ribosomal DNA analysis of the cryptogo-
nimids, transferred the species together with S. manilensis (Velasquez, 1961) to a
new genus Euryakaina as E. marina and E. manilensis. Euryakaina is distinguished
from all other cryptogonimid genera by the combination of a fusiform body, the few
relatively small widely spaced oral spines, a highly lobed ovary anterior to slightly
oblique testes, vitelline follicles that extend from the anterior margin of the testes to
slightly posterior to the intestinal bifurcation and an excretory vesicle that bifurcates
dorsal to the ovary and reunites slightly posterior to the intestinal bifurcation. Fur-
ther, the DNA sequences and ecological niche partitioning provided evidence that
the two are cryptic species.
Genus Mehrailla Srivastava, 1939
Generic diagnosis: Body elongate. Tegument spined. Oral sucker larger than ven-
tral sucker. Ventral sucker in ventrogenital depression, at about junction of anterior
and middle third of body. Prepharynx and pharynx well developed. Oesophagus
short. Caeca simple, reach posterior extremity. Testes 2, lobed, symmetrical, overlap
caeca. Seminal vesicle long and twisted. Pars prostatica ill-defined. Ejaculatory duct
well developed. Genital pore anterior to ventral sucker. Ovary pretesticular, multi-
lobed. Seminal receptacle present. Vitellarium lateral from just post-bifurcal to level
of testes. Uterus extensive, fills most of intertesticular and post-testicular space. Eggs
small, with a polar filament. Excretory vesicle Y-shaped.
Mehrailla ovocaudatum Srivatava, 1939 (Fig. 19.4)
Host: Stromateidae: Pampus argenteus (Euphrasen) (=Stromateus cinereus Bloch),
Lutjanidae: Lutanus johni (Bloch)
Locality: Karwar, Karachi, AS.
Reference: Srivastava (1939 h); Hafeezullah & Siddiqi (1970)
Fig. 19.4 Mehrailla

ovocaudatum (after
Srivastava, 1939 h)
Description: Body 4000–9400 long, 1360 wide. Oral sucker 300 in diameter. Sucker
ratio 3:2. Prepharynx and pharynx well developed. Oesophagus short. Caeca long.
Testes reniform with trilobed outer and entire inner margin, symmetrical, anterior
to posterior third of body, overlap caeca. Seminal vesicle long with three or four
constrictions, to right of ventral sucker, extends from just behind ventral sucker to
one third of distance between ventral sucker and intestinal bifurcation. Ejaculatory
duct wide. Genital pore median, anterior to ventral sucker, lies in small depression of
ventral body wall along with ventral sucker. Ovary multilobed, median, just anterior
to testes. Seminal receptacle spherical. Uterus from ventral sucker to just anterior to
caecal ends. Eggs triangular, 16−23 by 7−10, with long polar filament. Vitellarium
of irregular follicles from bifurcation to level of testes. Excretory vesicle Y-shaped.
Remarks: The generic name Mehracola Srivastava, 1937, was proposed by Srivas-
tava (1937) in an abstract with M. ovocaudatum from Pampus cinereus (=Stromaterus
cinereus) as the type species. Srivastava (1939 h) while giving the full description of
the species changed the generic name as Mehrailla Srivastava, 1838. Mehracola is,
therefore, a nomen nudum. M. ovocaudatum has also been recorded by Hafeezullah
and Siddiqi (1970a) and Hafeezullah & Dutta (1980). The species is characterized
by the lobed testes and the filamented eggs.
Genus Neometadena Hafeezullah & Siddiqi, 1970
[Syn. Neoparacryptogonimus Hafeezullah, 1975]
Generic diagnosis: Body oval. Tegument thick in anterior half of body. Oral sucker
with enlarged oral spines. Ventral sucker unspecialized. Prepharynx and oesophagus
very short. Intestinal bifurcation immediately anterior to ventral sucker. Caeca long,
blind. Testes two, symmetrical, in mid hindbody. Seminal vesicle tubular-saccular.
Gonotyle absent. Ovary with 3–4 large lobes, immediately anterior to testis. Uterus
in hindbody. Vitelline fields in two predominantly extracaecal lateral groups, from
posterior extremity to ventral sucker. Arms of excretory vesicle reach ventral sucker.
Type and only species N. ovata (Yamaguti, 1952).
Neometadena ovata (Yamaguti, 1952) Miller & Cribb, 2008 (Fig. 19.5)
[Syn. Paracryptogonimus ovata Yamaguti, 1952; Neometadena lutiani Hafeezullah
& Siddiqi, 1970; N. ovatus (Yamaguti, 1952) Hafeezullah, 1975]
Host: Lutjanidae: Lutjanus fulviflamma (Forsskal); Haemulidae: Pomadasys argen-
teus (Forsskal)
Locality: Karwar, Tuticotin, AS; VSK, BOB.
Reference: Hafeezullah & Siddiqi (1970a); Madhavi (1976b)
Distribution: Japan, India, Philippines, China, Red sea
Description: Body 3360–3840 long, 1680–2080 wide. Cervical gland cells and
eyespot pigment present. Tegument spined. Oral sucker 167−272 × 225−342. Ven-
tral sucker 190–230 by 202–296 in size, sunk in ventrogenital pit, near anterior
end of body. Sucker ratio 1:0.81−1.0. Prepharynx indistinct. Pharynx subspherical.
Oesophagus short. Intestional bifurcation anterior to ventral sucker. Caeca wide,
reach posterior end of body. Testes symmetrical, postequatorial. Seminal vesicle
Fig. 19.5 Neometadena

ovata (after Yamaguti, 1952)
long, convoluted, extends from posterior to ventral sucker to level of pharynx. Pars
prostatica absent. Distal part of seminal vesicle narrow, opens into genital atrium
situated immediately anterior to ventral sucker in ventrogenital pit. Ovary 3-lobed,
equatorial, pretesticular. Seminal receptacle anterior to ovary. Uterus voluminous.
Eggs 16−21 by 9−12. Vitellarium in lateral fields from posterior level of ventral
sucker to middle of post-testicular space.
Remarks: The genus Neometadena was proposed by Hafeezullah & Siddiqi (1970a)
for N. lutiani Hafeezullah & Siddiqi, 1970. According to Miller & Cribb (2008)
this species is identical to Neoparacryptogonimus ovatus (Yamaguti 1952) (syn.
Paracryptogonimus ovatus, Yamaguti, 1952) except for the absence of oral spines
in N. lutiani which Miller and Cribb felt were lost due to maceration. Miller &
Cribb (2008) regarded Neoparacryptogonimus Hafeezullah, 1975 as a synonym of
Neometadena and N. lutiani as junior synonym of N. ovatus (Yamaguti, 1952). Addi-
tional Paracrytogonimus ovatus Yamaguti, 1952 reports from haemulid, lutjanid and
chirocentrid fishes (Madhavi, 1976b, Leong & Wong, 1989, Shen, 1990, Nahhas
et al 2003) were considered identical to N. ovata. As a result of these synonymies, N.
ovatus has become a widely distributed species whose range includes Japan, India,
Philippines, China and Red Sea.
Genus Orientodiploproctodaeum Bhutta & Khan, 1970

[Syn, Anterodiscus Bilqees, 1994, Cryptocollaritrema Madhavi, 1976, Folliculovar-
ium Singh & Sinha, 1981; Harutrema Mehra & Kharoo, 1975; Multiovarium Bilqees,
1974)
Diagnosis: Body elongate oval, oral sucker surrounded by long collar without
spines. Ventral sucker embedded in ventrogenital sac. Prepharynx, Oesophagus
very short. Intestinal bifurcation immediately posterior to pharynx. Caeca open out
through ani at posterior extremity. Testes 2, oblique in mid hindbody. Seminal vesi-
cle tubular. Gonotyle absent or present as muscular lobe-like structure immediately
posterior to ventral sucker. Ovary deeply lobed between ventral sucker and testes.
Uterus fills entire hindbody. Vitellarium in two lateral groups, extracaecal from close
to posterior extremity to slightly anterior to ventral sucker. Type species: O. diacanthi
Bhutta & Khan, 1970.
Orientodiploproctodaem diacanthi Bhutta & Khan, 1970 (Fig. 19.6)
[Syn. Multiovarium heteroforme Bilqees, 1974; M. interruptum Bilqees, 1974;
Anterodiscus biseminalis Bilqees, 1994; A. triuteri Bilqees, 1974; Cryptocollar-
itrema provesiculatum Madhavi, 1976; Harutrema marinum Mehra & Kharoo, 1975;
Folliculovarium indicum Singh & Sinha, 1981]
Hosts: Lutjanidae: Lutjanus sp. (1); Sciaenidae: Protonibea diacanthus Lacepede(2)
Locality: 1. VSK, BOB; 2. Pondicherry, BOB, Trivandrum and Margao (Arabian
Sea)
Fig. 19.6 Orientodiploproctodaeum

diacanthi (after Madhavi,
1976)

Description: Body cylindrical, 3870−5920 long, 1680−2081 wide, anterior part
thickened as cushion-like structure filled with gland cells. Oral sucker 336−528 ×
608−720. Ventral sucker 240−256 in diameter, at junction of anterior and middle
third of body, embedded in ventrogenital pit. Prepharynx and oesophagus short.
Pharynx oval. Caeca open out through separate anal pores. Testes ovoid, at middle
third of body. Seminal vesicle coiled, largely anterior to ventral sucker, commences
slightly posterior to ventral sucker, runs anteriorly as far as level of pharynx, tightly
coiled, opens into pars prostatica which leads into genital atrium. Gonotyle appears
as muscular pad-like structure attached to posterior margin of genital atrium. Ovary
follicular, pretesticular. Seminal receptacle large, between ovary and seminal vesicle.
Uterine coils extensive. Vitelline follicles small, in two wide lateral zones, from level
of ventral sucker to posterior end. Eggs 14 by 10.
Remarks: The genus Orientodiploproctodaeum is characterized by the presence of
muscular cushion-like structure on the anterior part of body and also the anal pores.
Genus Polyorchitrema Srivastava, 1939
Diagnosis: Body oval. Tegument lacks spines. Oral sucker rounded, without spines.
Ventral sucker larger than oral sucker, embedded in ventrogenital sac. Prepharynx
very short. Oesophagus very short. Intestinal bifurcation in posterior forebody. Caeca
blind, terminate close to posterior end. Testes 8–50 follicles in posterior hindbody.
Seminal vesicle tubulo-saccular. Gonotyle may be present immediately anterior to
ventral sucker. Ovary entire or lobed, pretesticular. Uterus between testes and ventral
sucker. Vitelline follicles in two lateral groups in hindbody from posterior extremity
to slightly posterior to ventral sucker. Type species: P. piscicola Srivastava, 1939.
Polyorchitrema orissai Gupta & Puri, 1985 (Fig. 19.7)
Host: Sparidae: Chrysophrys sarba (Forsskal)
Locality: PR, BOB
No. one
Description: Body 2140 long 755 wide. Oral sucker 270 × 295. Prepharynx short.
Pharynx ovoid. Oesophagus short. Caeca simple, extend to posterior end of body.
Ventral sucker larger than oral sucker, 390 long, 420 wide. Ventrogenital sinus absent.
Genital pore submedian near anterior margin of ventral sucker. Testes 18 in num-
ber, compactly arranged in posterior part of body. Seminal vesicle sinuous, folded.
Pars prostatica long. Ejaculatory duct short. Ovary submedian, pretesticular. Semi-
nal receptacle short, posterior to ovary. Uterus pretesticular. Vitellarium follicular,
lateral, overlapping margins of caeca, extend from posterior end of seminal vesicle
to posterior end of body. Eggs 20–30 by 8–12
Remarks: Species of the genus Polyorchitrema are common in freshwater fishes.
Over half a dozen species have been recorded from these fishes from different fresh-
water bodies in India. The single specimen of P. orissai obtained from a marine fish
may represent an accidental infection.
Fig. 19.7 Polyorchitrema

orissai (after Gupta & Puri
1985)
Genus Pseudoanisocoelium Gupta & Tandon, 1984

This genus was erected by Gupta & Tandon (1984) with P. dorabi from the intestine
of Chirocentrus dorab from Puri, Bay of Bengal as the type species. Miller & Cribb
(2008) kept the genus under the category of genus inquirendum since the description
is inadequate.
Genus Siphoderina Manter, 1934
[ParacryptogonimusYamaguti, 1934; Lappogonimus Oshmarin, Mamaev &
Parukhin, 1961; Pseudallacanthochasnus Velasquez, 1961] Reference: Miller &
Cribb (2008)
Siphoderina Manter, 1934, is the biggest genus in the family Cryptogonimidae
including over 35 species. Miller & Cribb (2008) provided a review on this genus. Two
genera ParacryptogonimusYamaguti, 1934, and Lappogonimus Oshmarin, Mamaev
& Parukhin, 1961, have been synonymized with this genus and all the species in
these two genera were transferred to Siphoderina (Miller & Cribb, 2008). This genus
closely resembles Metadena but differs from it in having oral spines and in the pres-
ence of a gonotyle. Five species belonging to this genus have been reported from
marine fish from Indian waters: Siphoderina americanus (Manter, 1940) Miller &
Cribb, 2008; S. grandispinus (Velasquez, 1961) Miller & Cribb, 2008; Siphoderina
hardayali Pandey & Tiwari, 1983; Siphoderina hirastricta (Manter, 1963) Miller
& Cribb, 2008; Siphoderina sootai (Hafeezullah, 1975) Miller & Cribb, 2008. S.
hardyali reported from an unidentified marine fish from a fish market was inade-
quately described (Pandey & Tewari, 1983) and was regarded as incertae sedis by
Miller & Cribb (2008).
Generic diagnosis: Body oval to elongate oval. Oral sucker with enlarged oral
spines. Ventral sucker embedded in ventrogenital sac. Prepharynx, oesophagus short.
Intestinal bifurcation immediately anterior to ventral sucker or in mid-forebody.
Caeca blind, end close to posterior extremity. Testes two, symmetrical or oblique,
entire or deeply lobed, in mid-hindbody. Seminal vesicle tubulo-saccular. Gonotyle
absent or present. Ovary deeply lobed, overlaps or slightly separated from testes.
Uterus in hindbody, extends close to posterior extremity. Vitelline follicles in two
lateral groups, follicles not extending posteriorly beyond posterior testis. Arms of
excretory vesicle reach pharynx. Type species: S. brotulae Manter, 1934
Siphoderina americanus (Manter, 1940) Miller & Cribb, 2008 (Fig. 19.8)
[Paracryptogonimus americanus Manter, 1940]
Host: Lutjanus johni (Bloch)
Locality: Karwar, AS; VSK, BOB,
Reference: Hafeezullah & Siddiqi (1970a), Madhavi (1976b)
Distribution: Mexico, Panama, N. Carolina, Florida, BOB
Fig. 19.8 Siphoderina

americanus (after Hafezullah
& Siddiqi 1970)
Description: Body thick, plump, 2308–2376 long, 1147–1555 wide. Tegument

spined, Oral sucker 300–307 in diameter with single row of 52–57 spines. Ventral
sucker 187–225 in diameter, deeply embedded in body. Pharynx oval. Oesophagus
lacking. Caeca long. Testes 2, ovoid, symmetrical, far apart, near lateral edges of
body, separated by uterus. Seminal vesicle elongated, curves around left side of ven-
tral sucker. Ovary multilobed, pretesticular. Seminal receptacle large. Uterus extends
from posterior edge of ventral sucker to caecal ends. Eggs 20–25 by 9–10. Vitelline
follicles in two groups.
Remarks: S. americanus has been reported from 11 different host species including
lutjanids (Manter, 1940, Sogandares Bernal, 1969, Overstreet, 1969, Hafeezullah
& Siddiai, 1970, Fischthal, 1977), batrachoidiids (Yamaguti, 1971), centropomids
(Lamothe-Argumedo, 1969) and serranids (Fischthal, 1977) from Eastern Pacific,
Western Atlantic and Indo-Pacific Ocean).
Siphoderina grandispinus (Velasquez, 1961) Miller & Cribb, 2008 (Fig. 19.9)
[Peudallocanthochasmus grandispinus Velasquez, 1961
Host: Haemulidae: Pomadasys argenteus (Forsskal) (=P. hasta (Bloch))

grandispinus
Reference: Velasquez (1961), Hafezullah & Siddiqi (1970a)
Remarks: This species was described as Pseuallacanthochasmus grandispinus

from a snapper from Luzon Island, Philippines and Hafeezullah & Siddiqi (1970a)
from Pomadasys argenteus from Cochin coast. The Indian forms are smaller and
have 28 oral spines instead of 23–25. Miller & Cribb (2008) regarded Pseudalla-
canthochasmus as a synonym of Siphoderina.
Siphoderina hardayali Pandey & Tiwari, 1983
Host: Marine fish
Locality: Fish Market, Lucknow.
Reference: Pandey & Tewari (1983)
Status: Miller & Cribb (2008). considered this incertae sedis and we consider it a
species inquirendum.
Siphoderina hirastricta (Manter, 1963) Miller & Cribb, 2008
[Syn. Paracryptogonimus hitrastrictus Manter, 1963]
Host: Lutjanidae: Lutjanus lunulatus (Park)
Locality: VSK, BOB
Reference: Madhavi 1976b)
Distribution: Fiji, BOB
Siphoderina sootai (Hafeezullah, 1975) Miller & Cribb, 2008 (Fig. 19.10)
Syn: Paracryptogonimus sootai Hafezullah, 1975
Host: Lutjanidae, Lutjanus sp.
Locality: Gopalpur, BOB
No. 12

sootai (after Hafeezullah,
1975)
Description: Body 1114 −1348 long, 688−961 wide, ovate. Tegument thin, spines
probably lost. Oral sucker 77−120 × 196−235, with circlet of preoral spines, easily
lost. Ventral sucker 63−87 in diameter, spherical, at 210–385 from anterior end,
sunken in parenchyma. Sucker ratio 1:0.257 −1: 0.381. Prepharynx and oesopha-
gus absent. Pharynx oval. Intestinal bifurcation abuts pharynx. Caeca not visible.
Testes two, symmetrical, spherical, large, almost equatorial, touching edges of body,
narrowly separated by uterine coils. Seminal vesicle saccular, constricted into two
portions, curves around ventral sucker extends to ovary level. Genital pore in ventro-
genital depression at anterior border of ventral sucker. Prostatic cells and pars prostat-
ica lacking. Ovary multilobed, median, between testes and seminal vesicle. Seminal
receptacle dorsal to ovary. Uterus extends from level of ventral sucker almost to pos-
terior end of body. Eggs 11−18 by 7−8. Vitelline follicles in two lateral bunches,
on either side of ovary.
Remarks: S. sootai resembles S. americanus but differs from it in the very large
testes, the distribution of uterine coils and the sucker proportions.
Chapter 20
Miscellaneous Families
A few digenean species recorded from marine fishes of Indian waters belonged
to families that contain mainly representatives from freshwater fishes. Most of the
reports of these digeneans were based on single specimens and hence may represent
accidental infections. Altogether nine species of digeneans in four families belong
to this category: Cladorchiidae (3 spp), Microscaphiididae (1 sp) Cephalogonimidae
(4 spp), Opisthorchiidae (1 sp)
Superfamily Paramphistomoidea Fischoeder, 1901
Family Cladorchiidae Fischoeder, 1901
Representatives of this family occur in fishes, amphibians, reptiles, and mammals.

For fish parasites, the preferred hosts are freshwater fishes. There are occasional
reports from marine fishes. This family is distinguished from other families of the
superfamily Paramphistomoidea by the presence of paired pharyngeal sacs, two testes
instead of the usual one testis, and the post-testicular ovary. Jones (2005) provided a
review on this family.
Diagnosis: Body small or large. Ventral pouch absent. Ventral sucker close to poste-
rior extremity. Pharynx with paired intra- or extramural pharyngeal sacs. Pharyngeal
bulb absent. Testes two, lobed or not, tandem or symmetrical. External seminal vesi-
cle usually present. Cirrus-sac usually present. Genital sucker present or absent.
Genital pore bifurcal or post or prebifurcal. Ovary usually post-testicular. Uterus
mostly intercaecal. Eggs numerous. Distribution of vitelline follicles varies from
extensive in profuse lateral fields to relatively restricted vitelline fields.
Subfamily Helostomatinae Skrjabin, 1949
Diagnosis: Body small. Ventral sucker simple in structure. Pharynx with extramu-
ral sacs, much longer than pharynx. Oesophageal bulb present. Caeca reach ven-
tral sucker level or not. Genital sucker absent. Genital pore at level of pharyngeal
© Crown 2018 613
https://doi.org/10.1007/978-94-024-1535-3_20
614 20 Miscellaneous Families
sacs. Testes entire, slightly symmetrical. External seminal vesicle present. Cirrus-sac
present. Ovary post-testicular. Uterus mainly intercaecal. Vitelline follicles in lateral
fields along caeca, extent variable. Excretory vesicle saccate. Type and only genus
Helostomatis Fukui, 1929.
Genus Helostomatis Fukui, 1929
Diagnosis: With characters of the subfamily.
Three species of Helostomatis were reported in marine fish from India:
Helostomatis fotedari Gupta & Tandon, 1983 (Fig. 20.1)
Host: Stromateidae: Stromateus cinereus (Bleeker)
No. 1
Description: Body elliptica1, aspinose 3270 long, 1380 wide. Oral sucker terminal,
350 long, 400 wide. Oral diverticula long in form of claviform appendages. Posterior
sucker large 700 in diameter. Oesophagus with oesophageal bulb. Caeca terminate
short of posterior end. Genital pore median just posterior to bifurcation of oral diver-
ticulum, surrounded by genital sucker. Testes spherical, symmetrical, unequal, near
Fig. 20.1 Helostomatis

fotedari (after Gupta &
Tandon, 1983)
Family Cladorchiidae Fischoeder, 1901 615
intestinal bifurcation. Cirrus-sac pear-shaped. Seminal vesicle coiled, surrounded by

many prostate gland cells. Ejaculatory duct short. Ovary submedian, post-equatorial.
Vitellarium follicular, extending in lateral fields from just posterior to oral diverticula
to just posterior to caeca. Uterus coiled, intercaecal, passing between testes to open
at genital pore. Eggs large 90–108 × 60–80.
Remarks: Six species of Helostomatis have been reported from India of which three
were recorded from marine fishes, and the remaining three species were from fresh-
water fishes. H. fotedari is characterized by unlobed ovary and testes, the vitellarium
extending from oral diverticulum to just posterior to caeca.
Helostomatis muelleri Gupta & Puri, 1981
Host: Otolithus ruber (Bl. & Schn.)
Locality: PR, BOB
No. 2
Reference: Gupta & Puri (1981)
Description: Body elliptical, 2480–3220 long, 1140–1410 wide. Oral sucker ter-
minal, 270–370 long, 270–390 wide. Oral diverticles long, in form of claviform
appendages. Posterior sucker large, 590–790 long, 630–820 wide. Oesophageal
bulb 0.18–0.29 mm long, 0.12–0.13 mm wide. Caeca terminating short of posterior
extremity. Genital pore median, slightly posterior to bifurcation of oral diverticu-
lum, surrounded by genital sucker. Testes oval, near intestinal bifurcation. Cirrus-sac
pear-shaped. Ovary median or submedian. Vitellarium follicular, extending in lat-
eral fields from anterior border of oesophageal bulb to just posterior to caecal ends.
Uterus coiled, intercaecal, and extracaecal, passing between testes to open at genital
pore. Eggs large, 117–160 long, 67–80 wide.
Remarks: This species is characterized by its unlobed testes and ovary, the vitellar-
ium extends from the anterior end of the oesophageal bulb and the genital pore is
just posterior to the oral diverticula.
Helostomatis simhai Gupta & Puri, 1981(Fig. 20.2)
Host: Acanthuridae: Acanthurus sandwicensis Streets
Locality: PR, BOB
No. 1
Description: Body elliptical, 2370 long, 1010 wide. Oral sucker 270 × 250. Oral
diverticles long, in form of claviform appendages. Oesophagus long, oesophageal
bulb prominent. Caeca extend to level of posterior sucker. Genital pore slightly
anterior to middle of oesophagus. Testes oval, entire, symmetrical, just posterior to
intestinal bifurcation. Cirrus-sac pear-shaped, overlaps anterior part of oesophagus.
Ovary post-equatorial, near anterior margin of posterior sucker. Uterine coils occupy
space between posterior sucker and genital pore. Eggs 100–140 × 50–75. Vitellarium
follicular extends from middle of cirrus-sac to anterior end of posterior sucker.
Fig. 20.2 Helostomatis

simhai (after Gupta & Puri,
1981)
Remarks: The distinguishing features of the species are: the position of the genital
pore slightly anterior to the middle of the oesophagus and the extent of the vitellarium
between the cirrus-sac and the anterior end of the posterior sucker.
Family Microscaphidiidae Looss, 1900
[Syn. Angiodictyidae Looss, 1902]

Hexangium sigani Goto & Ozaki, 1929 (Fig. 20.3)
[Syn. Aruthur loossia Nagaty, 1954; Hexangium affinum Tubangui & Masilungan,
1944; Hexangium loossi (Nagaty, 1954) Yamaguti, 1958; Hexangium secundum
Annereaux, 1947]
Host: Sciaenidae: Sciaena volgeri (Bleeker)
Locality: PR, BOB
No. 6
Reference: Gupta, S.P. & Tandon (1983)
Distribution: Japan, Kanyan coast, Madagascar, Philippines
Family Microscaphidiidae Looss, 1900 617
Fig. 20.3 Hexangium sigani

(after Goto and Ozaki, 1929)
Description: Body elongate, 4350–6540 long, 750–1600 wide. Oral sucker termi-
nal, without oral diverticula, 200–280 long, 150–300 wide. Ventral sucker absent.
Oesophagus with posterior bulb. Pharynx lacking. Caeca thick, muscular, termi-
nating anterior to testes. Genital pore half way between oral sucker and intestinal
bifurcation. Testes subequal, ovoid, symmetrical, tandem or obliquely tandem, con-
tiguous, pre-ovarian in posterior part of body. Cirrus-sac absent. Seminal vesicle long,
sinuous, tubular. Ovary median or submedian, post-testicular. Vitellarium follicular,
follicles irregular in shape and size, confluent medially, anterior extent well posterior
to intestinal bifurcation extend to just anterior to posterior end of intestinal caeca.
Uterus with many transverse coils between intestinal caeca, becoming rectilinear
near intestinal bifurcation and extending beyond ovary. Eggs numerous, with thick
yellow shell, 80–90 long, 50–60 wide. Excretory pore subterminal, dorsal. Excretory
vesicle heart-shaped, thick-walled, with anterior lateral cornua proceeding into three
long excretory vessels; excretory vessels with numerous bulges or short branches.
Remarks: Gupta & Tandon (1983) reported this species from a sciaenid fish from the
Puri coast, Bay of Bengal, but most records are from species of the genus Siganus.
For example, H. sigani is recorded from Siganus spp. from off Japan (Goto et Ozaki
1929) and the Red Sea (Mohamadain et al. 2017; El-Labadi et al. 2006). It has also
been recorded from this genus of host from the Kenyan coast (Martens & Moens
(1995); Kuwait coast (Al-Yamani & Nahhas, 1981; Sey et al. 2003) and from Holo-
centrus sammara from Madagascar (Razarehelizoa, 1959). There are numerous other
references to this species. Besides H. affinum Tubangui & Masilungan, 1954 from
Siganus (= Amphacanthus) javus from Philippines, H. secundum Anneraux, 1947
from Siganus guttatus (= Teuthis concaterata) from Philippines and H. loossi Nagaty,
1954 from Pseudoscarus harid and Siganus oramin from the Red Sea were regarded
as synonyms of H. sigani by Velasquez (1961). The life cycle of H. sigani unraveled
by Hassanine et al. (2016) revealed the snail Nassarius pullus as the intermediate host.
The cercariae which are monostomate, encyst on vegetation. They also found a close
phylogenetic relationship between the Microscaphidiidae and the Mesometridae.
Superfamily Plagiorchioidea Luhe, 1901
Family Cephalogonimidae Looss, 1899
Genus Masenia Chatterji, 1933

[Syn. Eumasenia Srivastava, 1951]
A characteristic feature of this genus is the presence of enlarged circum-oral

spines: Oral sucker infundibuliform. Genital pore close to anterior extremity. Testes
2. Ovary post-testicular. Cirrus-sac long, extends from genital pore to ventral sucker.
Vitellarium follicular, distributed in two symmetrical fields lateral to ventral sucker
and gonads. Uterine coils extensive fill most of hindbody. Eggs numerous. Occur in
intestine of freshwater fishes; India and Africa. Type species: M. collate Chatterji,
1933.
Four species belonging to this genus have been recorded from marine fishes of
India, three of them from the Puri coast and one from Kerala coast. Details are:
Masenia carangai Gupta & Tandon, 1984 (Fig. 20.4)
Locality: PR, BOB
No. 1
Reference: Gupta & Tandon (1984)
Description: Body small, elongate, 925 long 240 wide. Oral sucker funnel-shaped
140 by 130, surrounded by large number of spines arranged in double alternate rows.
Ventral sucker smaller than oral sucker, 100 in diameter. Prepharynx short. Oesoph-
agus short. Pharynx oval. Caeca long extend to just past anterior testes. Genital pore
submedian, dorsolateral to oral sucker. Testes ovoid, tandem, in hindbody. Cirrus-sac
elongated, extends to anterior end of ovary. Seminal vesicle bipartite. Ovary pretes-
ticular. Uterus fills hindbody. Eggs 18–22 long. Vitelline follicles extend from middle
region of ventral sucker to posterior end of anterior testis.
Family Cephalogonimidae Looss, 1899 619
Fig. 20.4 Masenia carangai

(after Gupta & Tandon,
1984)
Remarks: The species is characterized by the ventral sucker being smaller than the
oral sucker, the lobed ovary, the cirrus-sac extending to the level of the ovary and the
vitellarium extending from the middle of the ventral sucker.
Masenia orissai Gupta & Tandon, 1984 (Fig. 20.5)
Host: Sciaenidae: Protonibea diacanthus (Lacepede)
Locality: PR, BOB
No. 1
Description: Body small, elongated, spinose, 815 long, 280 wide. Oral sucker ter-
minal, funnel-shaped, 130 long, 110 wide, surrounded by many spines arranged in
double alternating rows. Ventral sucker smaller than oral sucker, 900 long, 105 wide.
Prepharynx short, pharynx muscular. Oesophagus indistinct. Caeca simple, extend-
ing to posterior end of anterior testis. Genital pore median, postero-dorsal to oral
sucker. Testes ovoid, tandem. Cirrus-sac elongated, claviform, extending to mid-
dle of ventral sucker. Seminal vesicle bipartite. Ovary spherical, slightly overlapping
posterior end of ventral sucker. Uterus with ascending and descending limbs, extend-
ing up to just anterior to posterior end of body. Eggs ovoid, non-operculated 20–30 ×
Fig. 20.5 Masenia orissai

1984)
10–20. Vitellarium follicular, extending from just anterior to ventral sucker to hind
end of anterior testis.
Remarks: This species differs from M. carangai in having the cirrus-sac extending
only as far as the middle of the ventral sucker
Masenia quiloni (Gupta & Tandon, 1984) Madhavi, 2011 (Fig. 20.6)
[Syn. Eumasenia Gupta& Tandon, 1984]
Host: Engraulidae: Thryssa mystax
Locality: Quilon, Kerala (AS)
No. 1
Description: Body small, 885 long, 231 wide covered with small backwardly
directed spines. Oral sucker funnel-shaped 150 long, 130 wide, surrounded by 29
spines, interrupted dorsally, arranged in double alternative rows. Ventral sucker spher-
ical, pre-equatorial, smaller than oral sucker, 130 in diameter. Caeca simple, extend-
ing laterally as far as middle of anterior testis. Genital pore dorsomedian to oral
sucker. Testes ovoid, equal, obliquely tandem, post-equatorial, just posterior to ven-
tral sucker. Cirrus-sac elongated claviform extends from genital pore to just anterior
Family Cephalogonimidae Looss, 1899 621
Fig. 20.6 Masenia quiloni

1984)
to posterior end of ventral sucker. Ovary ovoid, slightly separated from ventral sucker.
Vitellarium follicular extending from middle of ventral sucker to anterior end of pos-
terior testis. Uterus with ascending and descending limbs, filling posterior part of
body opening at genital pore. Eggs ovoid, non-operculated, 18–30 long, 15–20 wide.
Remarks: This species was initially included in the genus Eumasenia as E. quiloni,
and was transferred to the genus Masenia as M. quiloni (Gupta & Tandon, 1984)
Madhavi, 2011. It is characterized by the funnel-shaped oral sucker surrounded by
29 spines interrupted dorsally, the cirrus-sac extending a little posterior to the ventral
sucker and the vitelline follicles being confined to lateral fields of the middle third
of the body.
Masenia upeneusi Gupta & Puri, 1984
Host: Mullidae: Upeneus macronemus (Lacepede)
Locality: PR, BOB
Superfamily Opisthorchioidea La Rue, 1957
Family Opisthorchiidae Looss, 1899
Subfamily Opisthorchiinae Looss, 1899

Genus Gomtia Thapar, 1930
Gomtia caranxi (Gupta & Jain, 1991) Scholz, 2008
[Syn. Pseudogomtiotrema Gupta & Jain, 1991]
Host: Carangidae: Carangoides malabaricus Bloch
Locality: PR, BOB
Reference: Gupta & Jain (1991b), Scholz (2008)
Remarks: Gupta & Jain (1991b) erected the genus Pseudogomtiotrema Gupta &
Jain 1991 with G. caranxi from Carangoides malabaricus from Puri coast, BOB as
the type species. Scholz (2008) synonymized the genus with Gomtia Thapar, 1930.
According to Scholz (2008), the description of G. caranxi was based on deformed
worms as is obvious from the unnatural shape of the body especially the almost
filiform anterior part, the deformed pre-ovarian region, the anteriorly directed tegu-
mental spines, the artificially everted ventral sucker and other features. The illustra-
tion is of a poor quality. The morphological description provided is inadequate and
erroneous. Moreover, species of the genus Gomtia infect freshwater fishes and it is
unusual to find an opisthorchioid in a marine host. Based on these observations we
are here including G. caranxi under the category of species inquirendum.
Chapter 21
Conclusions
The Bay of Bengal and the Arabian Sea parts of the Indian Ocean are fairly well
represented in the global literature on digenetic trematodes of marine fishes. Several
scientists have contributed to this knowledge on the digenean fauna of marine fishes of
India. An overview of the fauna is presented in the Checklist published on digenetic
trematodes of Indian marine fishes (Madhavi, 2011). The knowledge, however, is
incomplete and several species of marine fish and vast areas along the coasts remain
to be explored for their digenean fauna. The present analysis on the fauna has been
undertaken with a view to gain information on the present status of the knowledge,
identify the areas for future investigations and stimulate research in this subject
employing traditional and modern technologies. The ultimate aim is to convert the
area into a full-fledged and well-explored area for digenean parasites.
All the species of digeneans so far recorded from Indian parts of the Bay of
Bengal and the Arabian Sea and the adjacent islands are covered in this monograph.
Each species of digenean is briefly described together with information on the hosts,
localities, geographic distribution and the relevant literature. The valid status of each
species reported is assessed. This detailed and more or less comprehensive study
permitted an analysis of the aspects such as host specificity, the role of each species
of fish as host for digeneans, a preliminary comparison of the digenean fauna of fish
from the east and west coast of India, the geographic distribution of the parasites and
the extent of endemicity in the fauna. So far there are no reviews on digeneans of
marine fishes of India, except for reviews on Hemiuridae and Bucephalidae published
by Chauhan (1954). Many more species have been added since 1954.
The present analysis revealed that the digenean fauna of marine fishes of Indian
region is rich and comprises many species belonging to diverse families, thus reflect-
ing the richness and diversity of fishes and marine invertebrates of the coastal waters.
From the data presented in this account, it is evident that as many as 648 species of
digeneans belonging to 190 genera and 30 families have been recorded from marine
fishes of India. Over the years some of these species have been invalidated. The num-
© Crown 2018 623

https://doi.org/10.1007/978-94-024-1535-3_21
624 21 Conclusions
ber of species recorded under each family of digeneans and the number of species
invalidated, and the actual number of valid species are presented in Table 21.1. After
this revision, the total number of species has been reduced from the recorded 648 to
479 valid species. As many as 169 species of digeneans have been invalidated so far.
This indicates the lack of a proper scientific approach in dealing with the digeneans of
marine fishes of this region and the need to identify strategies to clear this prevailing
confusion. With the existing knowledge, the actual richness of the system cannot be
predicted. Undoubtedly, there remain many more species to be reported. However,
in spite of the paucity of the information some conclusions have been drawn from
the present analysis.
The number of species recorded under each digenean family is summarized in
Table 21.1. The maximum number of 121 species was recorded in the family Hemi-
uridae, followed by Opecoelidae (84 spp.), Monorchiidae (85 spp.) and Bucephali-
dae (56 spp.). Other digenean families with moderately high numbers of species are
Lepocreadiidae, Acanthocolpidae, Didymozoidae. Surprisingly only two species of
blood flukes (Aporocotylidae) have been recorded: Orchispirium heterovitellosum
from a ray and Paradeontacylix megalaspium from a carangid. Either the infections
with these blood flukes are rare, or they have been overlooked during host exam-
ination. It is of interest to note that didymozoids with as many as 46 species are
well represented, most of them being recorded from tunas. The abundance of these
didymozoids seems to be linked to the richness and diversity of tunas in the Indian
ocean.
The most speciose genera are found to be Prosorhynchus (Bucephalidae), Lecitho-
cladium, Lecithochirium, Erilepturus (Hemiuridae), Acanthocolpus, Stephanos-
tomum (Acanthocolpidae), Opisthmonorcheides (Monorchiidae). However, many
species in these genera have been invalidated by various workers either by syn-
onymizing them with other species or by regarding them as species inquirendae. For
example, in the genus Acanthocolpus only three out of twenty species recorded are
regarded as valid. The situation is similar in many other genera.
With the scanty information available, it would be a difficult task to compare the
digenean fauna of marine fishes from East and West coasts of India, very few studies
being undertaken along the West coast compared to East coast. Many host species
occur in common between the East and West coasts but the prevailing environmental
conditions in the two regions are very different. Geomorphologically, the two coasts
are significantly different. This will be reflected in the digenean fauna of fish in the
two regions. Very interesting results are expected to come out from a comparative
study of the digeneans of all fishes or selected species of fish from the two regions.
The available information on the digenean fauna of fish from the Bay of Bengal and
the Arabian Sea is summarized in Table 21.2. A total of 381 species were recorded
from BOB, 139 species from AS and a few species occur in common in the two
areas. Among the nine coastal states, including four on the east coast and five on
the west coast, most of the reports relate to studies undertaken in the vicinity of
Visakhapatnam and Puri on the east coast. Some data are available from Chennai,
Digha coast, Mumbai, Goa and Trivandrum (Kerala). The data presented in Fig. 21.1
indicates large numbers of digeneans were collected from Visakhapatnam coast (178
21 Conclusions 625
Table 21.1 Number of genera, valid and invalid species in the different families of digenetic trema-
todes recorded from Indian marine fishes
Digenean family Number of
Genera Valid species Invalid Total recorded
Aporocotylidae 2 2 2
Bivesiculidae 3 3 3
Bucephalidae 8 47 9 56
Fellodistomidae 6 13 7 20
Tandanicolidae 3 5 6 11
Haplosplanchnidae 3 6 5 11
Hemiuridae 15 46 75 121
Accacoelidae 3 3 3
Dictysarcidae 1 1 1
Didymozoidae 27 46 46
Hirudinellidae 1 1 1
Lecithasteridae 6 9 6 15
Sclerodistomidae 3 2 6 9
Transversotrematidae 1 1 1
Atractotrematidae 2 2 1 3
Haploporidae 9 14 2 16
Apocreadiidae 3 7 7
Lepocreadiidae 19 42 9 51
Aephnidiogenidae 2 2 1 3
Enenteridae 3 5 1 6
Gyliauchenidae 2 2 2
Acanthocolpidae 6 23 24 47
Opecoelidae 22 76 8 84
Opistholebetidae 3 5 1 6
Monorchiidae 14 58 7 65
Gorgoderidae 6 15 15
Zoogonidae 6 6 6
Faustulidae 8 16 16
Cryptogonimidae 8 12 1 13
Cephalogonimidae 1 4 4
Cladorchiidae 1 3 3
Microscaphidiidae 1 1 1
Opisthorchiidae 1 1 1
626 21 Conclusions
Table 21.2 Geographic distribution of digenean species of Indian marine fishes

Digenean family Endemic Indo-Pacific Cosmopolitan Total
species
Aporocotylidae 2 2
Bivesiculidae 3 3
Bucephalidae 34 9 4 47
Fellodistomidae 9 2 2 13
Tandanicolidae 4 2 6
Haplosplanchnidae 2 3 1 6
Hemiuridae 31 10 5 46
Accacoelidae 2 1 3
Dictysarcidae 1 1
Didymozoidae 31 15 46
Hirudinellidae 1 1
Lecithasteridae 8 1 9
Sclerodistomidae 3 3
Transversotrematidae 1 1
Atractotrematidae 2 2
Haploporidae 13 1 14
Apocreadiidae 7 7
Lepocreadiidae 34 7 1 42
Aephnidiogenidae 1 1 2
Enenteridae 4 1 5
Gyliauchenidae 2 2
Acanthocolpidae 18 1 4 23
Opecoelidae 72 2 2 76
Opistholebetidae 4 4
Monorchiidae 54 3 1 58
Gorgoderidae 12 3 15
Zoogonidae 2 1 3 6
Faustulidae 14 1 1 16
Cryptogonimidae 10 1 1 12
Cephalogonimidae 1 1
Cladorchiidae 3 3
Microscaphidiidae 1 1
Opisthorchiidae 1 1
spp.) followed by Puri coast (155 sp.), Mumbai coast (75 spp.), Gulf of Mannar (45
spp.), Kerala (60 spp.), Ratnagiri (26 spp.)
The host–parasite list comprising 320 species of marine fishes belonging to 88
families, of the Indian region examined so far and the digeneans recorded from them
21 Conclusions 627
Fig. 21.1 Map showing numbers of digenean species recorded from marine fishes of various coastal
areas. Abbreviations DC Digha Coast; PR Puri; VSK Visakhapatnam; MS Madras; BOM Bombay;
RT Ratnagiri
is presented in Table 21.2, updating the host scientific names following FishBase.
As per the survey undertaken by the Zoological Survey of India, it is estimated that
about 2,546 species of marine fish occur in the coastal waters of India. Excluding
the elasmobranchs and the small herbivorous fish, which are not suitable as hosts
for digeneans, there will be more than 1,500 species of marine fish suitable to serve
as hosts for digeneans. The present survey reveals that only one-fifth of available
marine fish have been examined for digeneans so far. Thus, the host–parasite list
gives only a partial picture. However, some conclusions can be drawn from the data
presented.
628 21 Conclusions
The tunas of the coast serving as hosts for 46 species of digeneans and the majority
belonging to Didymozoidae harboured the maximum number of digenean species.
The tuna fishery is quite predominant in Indian waters, and as many as 15 species
of tunas occur in the Indian waters. Only one species of tuna Euthynnus affinis has
been examined in depth for parasites, and an analysis was made of metazoan parasite
communities of this fish (Madhavi & Sairam, 2000). The large body size of tuna, high
vagility, varied diet, long life span and the homeothermic nature seem to promote
digenean infections in these fishes. Among other species of marine fish, Terapon spp.
also served as hosts for many species of digeneans dominated by opecoelids. Terapon
spp. occur in coastal waters in the intertidal zone, and the diet is mostly invertebrates.
The available information on life cycles of opecoelids indicates that intertidal snails
serve as intermediate hosts with crustaceans serving as intermediate hosts. Thus,
most probably the co-occurrence of intermediate and definitive hosts and the diet
and behaviour of the host and the suitable environmental conditions promoted the
infections with opecoelids. This aspect is to be studied in greater detail.
At the host family level, the maximum number of species of digeneans has been
recorded from carangids. Other groups of marine fishes serving as suitable hosts for
digeneans are the mullets and the scombrid fishes. Several species of lutjanid fishes
also occur in the coastal water, forming dense populations during certain seasons but
our list indicates that they are poor hosts for digeneans
Another notable feature of the digeneans recorded from Indian waters is that
majority of them (420 spp.) are endemic, confined to Indian waters and not crossing
the geographic limits. Only 12% of digenean species have a broader geographic
distribution. Even among this latter category, most are restricted to the Indo-Pacific
region sharing their digenean species with fishes of Japan and Hawaii. Few species
have a wider geographic distribution, occurring in tropical and subtropical waters.
Table 21.2 gives details of the geographic distribution of digeneans belonging to
different families. It is evident that in the families Monorchiidae, Haploporidae and
Opecoelidae, most of the species are endemic and very few are non-endemic. In the
family Didymozoidae, there are many species (15 out of 46) that are common to
Indian and Pacific waters. With the limited information available on the life cycles
of these parasites and the host–parasite relationships, it is difficult at this stage to
identify the factors limiting their geographic distribution.
The occurrence of cryptic species at least in some species of digeneans of the
coast exhibiting wide host specificity and geographic distribution cannot be ruled
out. For instance, Aponurus laguncula occurs in a wide range of hosts belonging to
different taxa and shows extensive intraspecific variations in the body organization
(Madhavi & Trivenilakshmi, 2011). The reported occurrence of cryptic species in
Aponurus from the Western Mediterranean (Carreras-Aubets et al. 2011) suggests
that there may be a similar situation prevailing here.
Another example of cryptic species is seen in Helicometra sp. where a form
morphologically similar to H. fasciata has distinctive features of the life cycle, in
particular the cercarial characters, based on which a new name, H. gibsoni, Murugesh,
Madhavi & Swrnakumari, 1993, was given to the Indian species. There is every
21 Conclusions 629
possibility of the occurrence of many cryptic species among species showing wide
geographic range and host specificity.
An aspect that deserves special attention is analysis of host–parasite phylogenetic
relationships employing molecular methods. A close phylogenetic relationship is
known to exist between the host and the parasite, and analysis of this relationship at
molecular level is expected to yield valuable information. So far this type of analysis
has not been attempted in India. Recent methodological advances permit a rigorous
comparison of phylogenetic trees to determine the extent to which these groups have
co-speciated. Linking morphological studies with molecular phylogeny is important
for understanding host–parasite co-evolution. In this context, the tunas and their
didymozoid parasites constitute an ideal model for such studies. The didymozoid
trematodes exhibit a high degree of host specificity and occur embedded in tissues
establishing intimate contact with the host tissue. Future workers should concentrate
on molecular aspects of phylogeny between the host and the parasite.
The main factors determining the fish parasite fauna in the marine environment
as well as their intensity and prevalence of infection in marine fishes can be sum-
marized as being the diet of the host fish, life span of the host, the mobility of the
host throughout its life including the variety of habitats it encounters, its population
density and the size attained, with large hosts providing more habitats suitable for
parasites than small ones (Polyanski, 1966).
Taking into consideration the rich and diverse fish fauna occurring in the coastal
waters of India, and the number of species of digeneans estimated to occur in these
fishes, the number recorded is far from complete. There is vast scope for future
studies which may unravel many interesting and new species of digeneans and
also permit proper understanding of the host specificity, geographic distribution
and host selection. Also, there is an urgent need to undertake molecular charac-
terization/morphometric analysis for the species which would be helpful in clearing
the existing confusion. Information available on the life cycles of digeneans is also
scanty.
Appendix
Host–Parasite list: Indian Marine fish hosts and their digenean parasites in alpha-
betical order
Host taxon Digenean
Phylum: Chordata
(Craniata)
Class Chondrichthyes
Family Dasyatidae
Brevitrygon imbricatus Orchispirium heterovitellatum
Himantura uarnak Petalodistomum yamagutia
Family Carcharhinidae
Galeocerdo cuvier Anaporrhutum gigas, Staphylorchis cymatodes
Galeocerdo tigrinus
Scoliodon dumerilii Anaporrhutum stunkardi
Scoliodon laticaudus Staphylorchis cymatodes
Scoliodon sorrakowah Anaporrhutum scoliodoni
Family Myliobatidae
Mobula mobular Anaporrhutum narayani
Sphyrnidae
Sphyrna zygaenae
Family Stegostomidae Prosogonotrema zygaenae
Stegostoma faciatum Anaporrhutum largum
(Hermann)
Family Torpedinidae Anaporrhutum albidum
Narcine timlei
Family Trigonidae Petalodistomum hanumanthai, Petalodistomum singhi
Trigon imbricatus Lecithocladium excisiforme
Trigon sp.
Class Actinopterygii
Family Acanthuridae
(continued)
© Crown 2018 631

https://doi.org/10.1007/978-94-024-1535-3
632 Appendix
(continued)
Host taxon Digenean
Aanthurus berda Erilepturus berda (=E. hamati), E. orientalis (=E. hamati)
Acanthurus bleekeri Aponurus theraponi
Acanthurus mata Aponurus laguncula, Opisthogonoporoides acanthuri,
Opisthogonoporoides hanumnthai, Pseudocreadium
indicium
Acanthurus sandvicensis Haplosplanchnus stunkardi (=H. caudatus);
Helostomatis simhai
Acanthurus triostegus Haplosplanchnus bengalensis, Haplosplanchnus
caudatus, Haplosplanchnus stunkardi, Helostomatis
simhai, Stomachicola mastacembeli
(=S. muraenesocis); Stomachicola singhi
(= S. muraenesocis)
Acanthurus sp. Hexangium sigani, Opisthogonoporoides hanumanthai
Ctenochaetus strigosus Haplosplanchnus bengalensis
Family Apogonidae
Apogon ellioti Opegaster paramacrorchis (=O. ditrematis),
Opisthomonorcheides indicus,
Opisthomonorcheides shindei
Family Ariidae
Arius arius Rhipidocotyle ligulum
Arius falcarius Lecithocladium acutum
Arius jella Elongoparorchis pneumatis, Monascus filiformis,
(=o Monodhelmis torpedinis)
Cephalocassis jatius Prosorhynchus truncata, Prosorhynchus tsengi,
Prosorhynchus filiformis
Sciades sona Bianium purii
Nemipterus nenga Buckleytrema indica
Ostiogoniosus militaris Monodhelmis militaris
Plicofollis platystomus Monodhelmis thapari
Plicofollis dussumierii Aponurus bengalensis
Family Ariommatidae
Ariomma indica Skrjabinozoum waltairense
Family Bagridae Decemtestes srivastavai, Helicometrina indica
Mystus gulio
Plicofollis dussumieri Aponurus bengalensis, Decemtestis mehrai
Plicofollis platystomus Proenenterum manteri, Monodhelmis thapari,
Monodhelmis torpedinis, Decemtestis mehrai
Family Balistidae
Abalistes stellaris Hypocreadium cavum, Hypocreadium indicum,
Lobatocreadium balistes
Sufflamen capistratus Xystretum chauhani, Xystretum manteri
Sufflamen chrysopterum Homalometron vinodae, Lobatocreadium manteri,
Sufflaman frenatus Coitocaecum indicum, Hypocreadium patellare,
Lobatocreadium manteri, Tetrochetus aluterae
Transversocreadium yamagutii
(continued)
Appendix 633
(continued)
Host taxon Digenean
Family Belonidae
Belone strongylura Prolecitha obesa
Belone crocodila Prolecitha obesa
Belone sp. Neidhartia neidharti
Strongylura strogylura Prosorhynchoides belonea
Tylosurus crocodylus Schikhobalotrema acutum
Xenentodon cancila Prosorhynchoides karvei Lecithostaphylus nitens
(=, Steganoderma nitens)
Family Bothidae
Bothus myriaster Macvicaria jagannathi
Engyprosopon Pseudopecoeloides scomberi
grandisquamis
Family Caesionidae
Caesio coerulaureus Alcicornis multidactylus
Family Carangidae
Alepes djedaba Alcicornis inglisi (sp. inq) Alcicornis thapari (sp. inq.),
Acanthocolpus orientalis (=Stephanostomum
orienalis), Acanthocolpus puriensis (=A. caballeroi),
Acanthocolpus valiyathurai, Bucephalus arabiansis,
Lecithocladium elongatus (=L. parviovum),
Lecithocladium. hanumnthai (=L. megalaspis),
Lecithocladium inglisi (=L. megalaspis),
Paradiscogaster indicus, P. pritchardae,
Prosorhynchus caballeroi, Schistorchis manteri
Alepes melanoptera Bucephalus arabiana
Carangoides armatus Allodecemtestis carangi, Bucephalus arabiana,
Masenia carangi, Opisthomonorcheides decapteri,
Opisthomonorchis armatus, Pseudopisthomonorchis
thapari
Carangoides chrysophrys Alcicornis carangis, Bucephalus margaritae,
Stephanostomum madhaviae
Carangoides malabaricus Acanthocolpus inglisi (=A. liodorus), Alcicornis
carangis, Bucephalus margaritae, Bucephalus
yamagutii, Gomtia caranxi, Monostephanostomum
mesospinosum, Opisthomonorcheides indicus,
Parahemiurus engraulisi Parahemiurus simhai,
Pseudopecoelus bilqeesae, Pseudopisthomonorchis
carangi, Pseudopisthomonorchis hanumanthai,
Stephanostomum madhaviae
Carangoides praestustus Lecithocladium caranxi, Lecithocladium fotedari
(=L. parviovum), Lecithocladium singhi
Caranx carangus Lecithocladium glandulum, M. caranxi,
Opisthogonoporus indicus, Tergestia bengalensis
Caranx gallus Erilepturus caranxi (=E. hamati)
Caranx hippos Tergestia bengalensis
(continued)
634 Appendix
(continued)
Host taxon Digenean
Caranx ire Lecithocladium singhi (=L. megalaspis)
Caranx malabaricus Parahemiurus simhai, Stephanostomum madhaviae
Caranx nigripinnis Bucephalus arabiansis
Caranx sansum Lecithocladium glandulum
Caranx sexfasciatus Alcicornis rauschi (sp. inq), Alcicornis thapari,
Bucephalus margaritae, Lecithocladium sexfasciatum,
Opisthomonorcheides manteri, Stephanostomum
madhaviae
Decapterus russelli Bucephalus margaritae, Lecithocladium bengalense,
Monascus filiformis
Elagatus bipunctata Prosorchis breviformis
Elagatus nigra Bucephalus elacatus
Equula daura Acanthocolpus orientalis (=Stephanostomum
orientalis)
Acanthocolpus chorinemi, Lecithocladium megalaspis,
Nematobothrium megalaspium, Paradeontacylix
megalaspium, Stephanostomum ditrematis,
Stephanostomum orientalis
Genolopa bychowski, Huridostomum formionis,
Megalaspis cordyla Lecithochirium pampi, Lecithocladium annulatum,
Parastromateus niger Lecithocladium excisiforme, Lepidapedon nelsoni,
Monascus filiformis, Opisthomonorcheides decapteri,
Opisthomonorcheides indicus, Opisthomonorcheides
karwarensis, Opisthomonorcheides nigeri,
Opisthomonorcheides stromatense, Prosogonotrema
arabicum, Prosogonotrema bilbiatum,
Prosogonotrema pavasi, Prosogonotrema pritchardae,
Prosogonotrema posterouterinum, Prosorhynchus
madhaviae
Scomberoides lysan Aanthocolpus chorinemusi, Prosorhynchoides
chorinemi, Prosorhynchoides pritchardae,
Prosorhynchoides thapari, Prosorhynhus tewariae
Scomberoides tala Didymocystis bifasciata, Hirudinella ventricosa,
Lecithocladium magnum, Leiomonorchis secundus,
Prodistomum mohsini (=Opechona mosini),
Prosorhynchus chorinemi, Prosorhynchus indicus,
Pseudolepocreadioides bombayensis,
Pseudopecoeloides scomberi, Stephanostomum
attenuatum
Selar crumenophthalmus Paramonorcheides selaris
Trachinotus blochii Neolepocreadium trachinoti
Trachinotus ovatus Decemtestes marginoacetabulatus
Family Chaetodontidae
Chaetodon pictus Antorchis intermedius, Coitocaecum chaetodoni,
Hurleytrematoides filiformis, Neohypocreadium
chaetodoni, Opisthomonorcheides chaetodoni
Chaetodon vagabundus Notoporus fotedari, Opisthomonorcheides chaetodoni
(continued)
Appendix 635
(continued)
Host taxon Digenean
Family Chanidae
Chanos chanos Isorchis skrjabini, Isorchis chanosi
Family Cichlidae
Etroplus suratensis Skrjabinolecithum indicum, Erilepturus chilkai
(=E. hamati)
Family Clupeidae
Clupea fimbriata Aphanurus stossichi
Clupea sp. Rhipidoctyle apapillosa
Dussumieria acuta Aphanurus dussumieri, Elopsium indicum,
Parahemiurus dussumierai, Parahemiurus
gastricus, Parahemiurus indicus, Elopsium indicum
Dussumieria elopsoides Elopsium stunkardi, Haplosplanchnus bengalensis,
Parahemiurus gastricus, Parahemiurus madrasensis
Dussumieria hasseltii Aphanurus dussumierii
Hilsa toli Aphanurus stossichi
Ilisha elongata Aphanurus stossichi
Ilisha filigera Parahemiurus engraulisi
Nematolosa nasus Yamagutia lateroporus
Sardinella albella Parahemiurus madrasensis
Sardinella brachysoma Lecithochirium indicum
Sardinella fimbriata Aphanurus tuberculatus, Bacciger sardinellae,
Pronoprymma petrowi, Pseudobacciger cablei,
Pseudobacciger harengulae
Sardinella gibbosa Pronoprymma petrowi, Pseudobacciger cablei,
Pseudobacciger harengulae
Sardinella longiceps Ahemiurus karachii
Tenualosa ilisha Aphanurus stossichii, Aponurus breviformis, Faustula
basiri, Faustula brevichrus, Faustula gangetica,
Lecithocladium chauhani (=L. harpodontis),
Lecithocladium piscicola, Lecithocladium striatum
Tenualosa sinensis Aphanurus dussumierii, Faustula gangetica
Family: Chirocentridae
Chirocentrus dorab Acanthocolpus caballeroi, Acanthocolpus dorabi
(taxon inq.), Acanthocolpus guptai (=A. liodorus),
A. inglisi (=A. liodorus) Acanthocolpus liodorus,
Acanthocolpus luhei, Acanthocolpus
microtesticulatus (taxon inq), Acanthocolpus
srivastavai (sp. inq), Acanthocolpus
thapari (sp. inq), Allostomachicola rauschi,
Allostomachicola secundus, Anisocoelium dorabi,
Hamacreadium indicum, Hymenocotta mulli,
Neohysterolecitha indica, Paropecoelus filiformis,
Podocotyle dorabi, Podocotyle pentavitellosus,
Podocotyle simhai, Schikhobalotrema mulli,
Sclerodistomum chirocentrusi, Stephanostomoides
dorabi (=S. tenuis), Stephanostomoides indicus
(continued)
636 Appendix
(continued)
Host taxon Digenean
(=S. tenuis), Stephanostomoides sharmai (=S. tenuis),
Stephanostomoides tenuis, Stephanostomum
anterospinosum
Family Congridae
Uroconger lepturus Allostomachicola lepturusi, Phyllotrema
tetracaudatum, Stomachicola chauhni
Family Coryphaenidae
Ariosoma mauritianum Stomachicola muraenesocis
Coryphaena hippurus Dinurus barbatus, Dinurus breviductus, Dinurus
hippuri, Dinurus longisinus, Dinurus tornatus,
Tetrochetus coryphaenae
Family Cynoglossidae
Aesopia cornula Macvicaria cynoglossi, Zebrias altipinnis
Cynoglossus arel Lepocreadioides orientalis, Macvicaria cynoglossi
Cynoglossus bilineatus Allodecemtestis bilqeesae, Allodecemtestis skrjabini,
Decemtestis kobayashii, Lepocreadioides orientalis
Cynoglossus cynoglossus Lepocreadioides srivastavai
Cynoglossus dubius Lasiotocus hastai, Lasiotocus maculatus,
Lepocreadioides indicus, L. orientalis, Macvicaria
longicaudatus
Cynoglossus lida Decemtestis mehrai, Lepocreadioides indicus
(=L. orientalis), Lepocreadioides orientalis,
Lepocreadioides thapari, Macvicaria cynoglossi,
Macvicaria longicaudus, Prodistomum travassossi,
Rhipidocotyle indicus
Cynoglossus lingua Lepocreadioides indicus (=L. orientalis),
Lepocreadioides orientalis
Cynoglossus Lepocreadioides orientalis, Macvicaria longicaudatus
macrostomus
Cynoglossus microlepis Lepocreadioides orientalis, Macvicaria microlepis
Cynoglossus oligolepis Decemtestes cynoglossi, D. kobayashii,
Podocotyloides dorabus, P. yamagutii
Cynoglossus puncticeps Lepocreadioides orientalis
Cynoglossus sindensis Lepocreadioides orientalis
Cynoglossus sp. Lecithochirium cynoglossi
Zebrias synapturoides Mcvicaria cynoglossi
Family Dactylopteridae
Dactyloptena orientalis Anisoporus orientalis
Family Diodontidae
Diodon hystrix Tetrochetus coryphaenae, Tetrochetus
macrorchis, Opegaster ditrematis
(continued)
Appendix 637
(continued)
Host taxon Digenean
Family Drepanidae
Drepane punctata Allodecemtestis drepanei, Aponurus drepani
(=A. laguncula), Apocreadium drepanei, Notoporus
gibsoni, Neoapocreadium imtiazi,
Opisthomonorcheides guptai, Paradiscogaster arabi,
Pseudolepocreadioides symmetrorchis
Family Echeneidae
Echeneis naucrates Diphterostomum plotosi, Echeneidocoelium indicum
Echeneis remora Echeneidocoelium indicum
Family Elopidae
Elops inermis Elopsium puriense
Elops saurus Elopsium secundum, Lecithochirium pritchardae
Family Engraulidae
Setipinna phasa Baccigeroides hafeezullai
Stolephorus indicus Lasiotocus engraulisi, Lasiotocus maculatus
Thryssa dussumieri Parahemiurus puriensis
Thryssa hamiltoni Gyliauchen indicus, Parahemiurus cameronui,
Parahemiurus engraulisi, Parahemiurus hamiltoni,
Parahemiurus yanamensis
Thryssa malabarica Dinurus thapari
Thryssa mystax Baccigeroides cochinensis, Coitocaecum bengalense,
Coitocaecum thrissocles, Erilepturus thrissocli,
Masenia quiloni, Opisthomonorcheides engraulisi,
Opisthomonorcheides simhai. Parahemiurus
engraulisi, Parahemiurus hamiltoni, Timonia vinodae
Thryssa setirostris Thryssatrema hanumanthai
Family Ephippidae
Platax teira Neomultitestis benglensis, Pseudopisthomonorchis
secundus, Thulinia microrchis
Exocoetidae
Cypselurus comatus Brachyenteron parexocoetis, Didymodiclinus
spilonotopteri,
Exocoetus volitans Lecithostaphylus parexocoeti (= Steganoderma
parexocoeti), Tetrochetus aluterae
Family Fistularidae
Fistularia petimba Prodistomum gaevskayae, Preptetos madrasensis,
Stephanostomum adinterruptum, Transversocreadium
secundus
Family Gerreidae
Gerres filamentosus Acanthocolpus lucknowensis (=A. caballeroi),
Macvicaria filamentusi (=M. yamagutii) Macvicaria
yamagutii, Neoapocreadium karwarense,
Pseudolepocreadioides thapari, Transversocreadium
secundus, Xystretum thapari
(continued)
638 Appendix
(continued)
Host taxon Digenean
Family Haemulidae
Pomadasys argenteus Diphterostomum indicum, Neometadena ovata,
Siphoderina grandispinus
Pomadasys argyreus Lasiotocus hastai, Lasiotocus maculatus
Pomadasys furcatus Ametrodaptes pristipomatis, Diphterostomum indicum,
Helicometrina nimia, Lepidauchen nicolli
Pomadasys hasta Qadriana zakiae
Pomadasys maculatus Aephnidiogenes senegalensis Alcicornis indicus,
Bucephalus margaritae, Diphterostomum indicum,
Genolopa trifolifer, Lasiotocus hastai, Lasiotocus
maculatus, Monorchis bengalensis, Monorchis
minutus, Proctotrema guptai, Pseudametrodaptes
fischthali, Pseudametrodaptes secundus
Pomadasys commersoni Notoporus pristipomatis, Decemtestis varmai
Pristipoma furcatum Erilepturus bengalensis
Pristipoma sp. Lecithochirium pristipomi
Family Harpodontidae
Harpodon neherius Lecithocladium carultum, Lecithocladium harpodontis
Family Hemiramphidae
Hemiramphus Alcicornis hemiramphi
brachynopterus
Hemiramphus limbatus Allostomachicola secundus
Hemiramphus marginatus Schikhobalotrema acutum
Hyporamphus Neogonapodasmius hemiramphi
xanthopterus
Rhynchoramphus georgii Schikhoblotrema acutum, Koseira manteri
Rhynchoramphus Koseiria caballeroi
leucopterus
Trachyramphus far Tergestia clonacantha
Family Holocentridae
Holocentrus sp. Pseudopecoelus pritchardae
Myripristis kuntee Pseudopecoelus brayi
Family Kyphosidae
Kyphosus cinerascens Cadenatella dollfusi, Enenterum mannarense, Koseira
nagatyi, Neopisthodena habei
Family Labridae
Halichoerus scapularis Monodhelmis chilkai
Oxycheilinus diagramma Zoogonoides pyriformes
Pseudodax moluccanus Macvicaria jagannathi
Family Lactariidae
Lactarius lactarius Pseudametrodaptes caballeroi, Pseudopecoelus
stunkardi
(continued)
Appendix 639
(continued)
Host taxon Digenean
Family Latidae
Lates calcalifer Hamacreadium chilkai, Stephanostomum cloacum
Family Leiognathidae
Gazza minuta Erilepturus gazzi (=E. hamati), Lecithocladium
bengalense, Lecithocadium gazzi, Lecithocladium
srivastavai (=L. parviovum) Notoporus pristipomatis,
Lecithoclasium purenae (=L. megalaspis),
Stephanostomum gazzai (sp. inq)
Leiognathus bindus Notoporus pristipomatis
Leiognathus daura Acanthocolpus amaravatai (taxon. inq),
Acanthocolpus durghai (taxon. inq) Acanthocolpus
equulai, Lasiotocus odhneri, Leiomonorchis mamaevi,
Neonotoporus leiognathi, Notoporus pristipomatis,
Stephanostomum equulai (=S. triacanthi), S. orientalis
Secutor insidiator Notoporus pristipomatis
Family Lethrinidae
Lethrinus nebulosus Hamacreadium mutabile
Family Lutjanidae
Lutjanus fulviflamma Euryakina marina, Hamacreadium mutabile,
Helicometrina nimia, Macvicaria indica,
Neometadena ovata, Paropecoelus pritchardae,
Vitellibaculum fischthali
Lutjanus fulvus Acanthocolpus lutjanusi
Lutjanus johni Lecithocladium glandulum, Mehrailla ovocaudatum,
Neometadena ovatus, Siphoderina americanus,
Siphoderina elongates, Decemtestis kobayashii,
Stephanostomum orientalis,
Lutjanus leioglossus Elyptophalloides parukhini
Lutjanus lunulatus Siphodera hirastricta, Allopodocotyle pritchardae
Lutjanus malabaricus Sephanostomum casum, Acanthosiphodera
bengalense, Allometadena rotundum, Elytrophalloides
parukhini
Lutjanus marginatus Acanthocolpus lutjanusi
Lutjanus monostigma Lasiotocus overstreeti, Elyptophalloides parukhini
Lutjanus quinquilineatus Hamacreadium mutabile, Lutianotrema
kanyakumariae, Lutianotrema quinqueliai
Lutjanus rivulatus Allopodocotyle argyropsi, Allopodocotyle lutianusi,
Allopodocotyle pritchardae, Hamacreadium mutabile,
Helicometrina nimia, Stephanostomum chauhani,
Lutjanus waigiensis Macvicaria puriensis (sp. inq)
Lutjanus sp. Helicometra filamentosa, Helicometroides
pseudovitellosus, Orientodiploproctodaeum diacanthi,
Paracryptogonimus sootai, Siphoderina sootai
(continued)
640 Appendix
(continued)
Host taxon Digenean
Pristiopomoides Diplohureytrema nahhasi, Lepidapedon manteri,
argyrogrammicus Prosogonotrema pritchardae
Pristipomoides typus Metanematobothrioides branchialis
Family Mastacembelidae
Mastacembalus armatus Stomachicola mastacembali (=S. muraenesocis)
Family Megalopidae
Megalops cyprinoides Elopsium manteri
Family Monacanthidae
Paramonacanthus Hypocreadium indicum, Lintonium pseudovibex,
choirocephalus Prosogonotrema bilabiatum
Family Mugilidae
Liza macrolepis Haploporus pseudoindicus, Haplosplanchnus
caudatus, H. purii, Unisaccus mugilis, Phyllodistomum
lewisi, Pseudohapladena martini, Transversotrema
patialense
Liza parsia Aphanurus microrchis, Lecithobotrys
vitellosus (sp. inq)
Liza vaigiensis Haplosplanchnus caudatus, H. indicus, H. purii,
Paralecithobotrys overstreeti, Uniscus martini,
Waretrema piscicolum
Mugil cephalus Aphanurus harengulae, Carassotrema bengalense,
Haplosplanchnus caudatus, Haplosplanchnus guptai,
Haplosplanchnus vinodae, Pseudohapladena martini
(=Saccocoelioides martini), Unisaccus guptai,
Unisaccus mugilis (=Lecithobotrys mugilis),
Paucivitellosus hanumanthai, Saturnius segmentatus,
Transversotrema patialense, Unisaccus
overstreeti (=Pseudounicoelium overstreeti)
Mugil sp. Lecithochirium mugili, Saccocoelioides chilkaensis,
Saccocoelium tripathi
Valamugil cunnesius Aphanurus harengulae, Haploporus indicus,
Haplosplanchnus caudatus, Lecithocladium
glandulum, Unisaccus mugilis, Pseudohapladena
bengalensis, Pseudohapladena
martini (=Sacocoelioides martini), Skrjabinolecithum
bengalense, Saturnius valamugilis, Transversotrema
patialense
Family Mullidae
Upeneus bensai Pseudopecoelina elongate
Parupeneus macronemus Allopodocotyle upeneusi (sp. inq.), Coitocaecum
thrissocles, Coitocaecum longivesiculum, Masenia
upeneusi, Opisthomonorcheides upeneusi,
Pseudolepidapedon indica, Pseudopecoeloides
elongatus, Pseudopecoeloides
orientalis, Pseudopecoeloides puriensis,
Stephanostomum bhuneswari
(continued)
Appendix 641
(continued)
Host taxon Digenean
Upeneus sulphureus Lecithocladium suphurusi, Paropecoelus dollfusi,
Paropecoelus indicus
Upeneus taeniopterus Pseudopecoelina elongate
Upeneus vittatus Timonia stunkardi
Family Muraenesocidae
Congresox talabanoides Monodhelmis torpedinis, Stomachicola muraenesocis,
Stomachicola serpentina (=S. muraenesocis)
Gymnothorax undulatus Dollfustrema bengalense, Coitocaecum bombayense
Family Nemipteridae
Nemipterus furcosus Macvicaria jagannathi
Nemipterus japonicus Podocotyloides parupenei
Scolopsis vosmeri Allobacciger macrorchis, Opisthomonorcheides
overstreeti
Family Nomeidae
Psenes cyanophrys Skrjabinozoum yamagutii
Family Ophichthyidae
Leptocephalus conger Stomachicola kinnei (=S. muraenesocis)
Pisodontophis boro Lecithochirium fotedari, Proctotrematoides yamagutii
Family Paralichthyidae
Pseudorhombus Lecithaster bombayensis, Pleorchis sciaenae
diacanthus
Pseudorhombus elevatus Macvicaria jagannathi
Pseudorhombus javanicus Macvicaria jagannathi
Pseudorhombus Opegaster ditrematis
micrognathus
Pseudorhombus Monorcheides hafeezullahi, Opegaster ditrematis
triocellatus
Family Platycephalidae
Gramnoplites scaber Helicometrina nimia, Indodidymozoon madhaviae,
Prosorhynchus platycephali, Tubulovesicula
angusticauda
Platycephlus indicus Lepocreadioides orietalis
Family Plotosidae
Plotosus conius Allopodocotyle coniusi
Plotosus lineatus Prosogonarium plotosi
Family Polynemidae
Eleutheronema Lasiotocus polynemi, Erilepturus paralichthydis
tetradactylum (E=hamati)
Leptomelanosoma indicus Lecithochirium polynemi, Monodhelmis torpedinis,
Stephanostomum polynemi, Timonia indica
Polydactylus sextarius Neotimonia caballeroi, Opisthodiplomonorchis
elongatus
(continued)
642 Appendix
(continued)
Host taxon Digenean
Polynemus paradiseus Lecithocladium glandulum
Polynemus plebeius Bucephalus margaritae
Family Pomacentridae
Abudefduf bengalensis Coitocaecum purius, Erilepturus neoorientalis,
Opisthodiplomonorchis elongatus
Family Pomacanthidae
Glyphidodon bengalensis Coitocaecum purius
Pomacanthus annularis Antorchis pomacanthi, Lobatocreadium gupti,
Pseudolepocreadium secundus
Family Priacanthidae
Priacanthus blochi Lecithochirium ernakulamense
Priacanthus tyenus Opegaster ditrematis
Priacanthus hamrur Neometadidymozoon polymorphis
Family Pristigasteridae
Ilisha elongata Aphanurus stossichi, Haplosplanchnus caudatus,
Opisthomonorcheides pellonai, Aponurus breviformes
Ilisha filigera Aponurus breviformes, Parahemiurus engraulisis
Family Psettodidae
Psettodes erumei Helicometrina nimia, Lecithocladium keralense
(=L. parviovum), Lecithocladium psettodi
(=L. glandulum), Neidhartia microrhyncha,
Neonotoporus overstreeti, Opegaster ditrematis,
Opisthodiplomonorchis elongatus,
Opisthomonorcheides yamagutii, Pleorchis sciaenae,
Rhipidocotyle ghanensis
Family Rachycentridae
Rachycentron canadus Bucephalus elacatus, Diphtherostomum indicum,
Lecithocladium jagannathi, Stephanostomum cloacum,
Stephanostomum microsomum, Stephanostomum
pseudoditrematis, Tormopsolus filiformis,
T. spatulum (=T. filiformis)
Family Scaridae
Scarus globban Bivesiculoides scari, Opisthogonoporoides thapari
(sp. inq.)
Family Scatophagidae
Scatophagus argus Bucephalus barina, Monodhelmis orissai,
M. torpedinis, Paradiscogaster farooqii, Thulinia
microrchis
Family Sciaenidae
Dendrophysa russelli Diploproctodaeum bombayense
Johnius aneus Bucephalus barina, Opegaster ditrematis,
Johnius belengeri Bucephalus barina, Prosorhynchoides microcirrus,
Pseudopecoelus odeningi
(continued)
Appendix 643
(continued)
Host taxon Digenean
Johnius carutta Bucephalus barina, Lecithocladium carultum,
Qadriana zakiae
Johnius dussumieri Bucephalus barina, Paropecoelus sciani
(=P. parviovum)
Johnius glaucus Helicometrina nimia
Johnius sina Helicometrina nimia
Kathaila axillaris Helicometrina nimia, Lecithocladium glandulum
Nibea soldado Bucephalus barina, Paropecoelus sciaenae
Otolithus ruber Decemtestis singhi, Enenterum
minutum, Haplosplanchnus otolithi, Helostomatis
mulleri, Helicometrina hexorchis (Syn. of H. nimia),
Helicometrina quadrorchis, Helicometrina septorchis,
Helicometroides leiperi, Opegaster ditrematis
Pennahia anea Helicometrina nimia
Protonibea diacanthus Lecithocladium muthiahi, Masenia
orissai, Orientodiploproctodaeum diacanthi
Pseudosciaena sina Pseudopecoelina stunkardi
Sciaena volgeri Hexangium sigani
Sciaena sp. Monodhelmis torpedinis, Pleorchis sciaenae
Protonibea diacanthus Masenia orissai, Opegaster ditrematis,
Paracryptogonimus sootai, Orientodiploproctodaem
diacanthi
Family Scombridae
Acanthocybium solandri Lecithocladium acutum
Acanthocybium sp. Lecithochirium polynemi, Plerurus digitarus
Auxis thazard Didymocystis exigua, Didymocystis wedli,
Rhipidocotyle pentagonum
Euthynnus affinis Allopseudocolocyntotrema claviforme, Didymocodium
euthynni, Hirudinella ventricosa, Didymocystis
exigua, Didymozoon lobatum, Didymozoon
diverticulatum, Kamegaia kawakawa, Lobatocystis
bengalensis, Lobatocystis yaito, Melanocystis
kawakawa, Metanematobothrium bivitellatum,
Neonematobothrium dorsale, Neonematobothium
kawakawa, Opepherocystis kawakawa,
Opepherotrematoides multitubularis, Phyllodistomum
lancea, Pseudocolocyntotrema yaito, Rhipidocotyle
pentagonum
Euthynnus sp. Acanthocolpus manteri (=A. luhei),
Monostephanostomum loossi
Katsuwonus pelamis Coeliodidymocystis kamegaii, Didymocystis dissimilis,
Didymocystis pinnicola
Rastrelliger brachysoma Prodistomum waltairense, Prosorhynchoides brayi,
Prosorhynchoides orientalis (sp. inq), Renodidymocystis
yamagutii, Pseudopecoeloides tenuides, Hirudinella
ventricosa, Prodistomum orientalis
(continued)
644 Appendix
(continued)
Host taxon Digenean
Rastrelliger kanagurta Alcicornis scomberi, Allodecemtestis biacetabulatus,
Aponurus lagunculuas, Bucephalus kanagurta,
Coitocaecum scombri, Erilepturus hamati,
Helicometrina scomberi, Lecithocladium angustiovum,
Lecithocladium indicum (=L. angustiovum),
Lecithocladium manteri (=L. angustiovum),
Lecithocladium microlepidotus (=L. angustiovum),
Lecithocladium seriolellae (=L. angustiovum),
Lecithocladium siddiquii (=L. angustiovum),
Lecithocladium tewarii L. (=L. angustiovum),
Monascus filiformis, Opechona ahmadi, O. bacillaris,
Prodistomum orierntalis, Renodidymocystis yamagutii
Sarda chiliensis Stomachicola pelamisi (=S. muraenesocis)
Scomberomorus guttatus Bucephalus jagannathai, Didymocystis guttatus,
Didymocystis semiglobularis, Lecithocladium cybiumi
(=L. angustiovum), Prosorhynchoides microcirrus
Thunnus albacares Decemtestis dollfusi, Didymocystis bifurcata,
Didymocystis guttata, Didymocystis oesophagicola,
Didymocystis semiglobularis, Didymocystis
superpalati
Thunnus tonggol Didymocystis alolongae, Didymocystis bifurcata,
Didymocystis oesophagicola, Didymocystis
superpalati
Family Scorpaenidae
Scorpaenopsis cirrhosus Helicometra gibsoni
Family Serranidae
Cephalopholis sonnerati Lasiotocus hastai, Lasiotocus puriensis,
Opisthomonorcheides gibsoni, Opisthomonorcheides
madhavae
Epinephelus bleekeri Prosorhynchus epinepheli
(=Serranus wandersis)
Epinephelus chlorostigma Prosorhynchus epinepheli
Epinephelus diacanthus Prosorhynchus epinepheli
Epinephelus Hysterolecithoides yamagutii
flavocaeruleus
Epinephelus lanceolatus Neoprosorhynchus purius
Epinephelus latifasciatus Didymodiclinus branchialis
Epinephelus maculatus Helicometrina nimia
Epinephelus malabaricus Prosorhynchus atlanticus
Epinephelus radiatus Didymodiclinus reticulum
Epinephelus tauvina Allonematobothrium epinepheli, Indoglomeritrema
epinepheli, Prosorhynchus pacificus
Epinephelus undulosus Helicometrina nimia, Opisthomonorcheides nahhasi,
Prosorhynchus epinepheli
Serranus indicus Lasiotocus srivastavai
(continued)
Appendix 645
(continued)
Host taxon Digenean
Family Siganidae
Siganus argenteus Atractotrema kuntzi
Siganus canaliculatus Gyliauchen ozakii, Hysterolecithoides frontilatus,
Siganus flavoceruleus Hysterolecithoides yamagutii
Siganus javus Hysterolecithoides frontilatus, Manteriella yamagutii
Siganus oramin Hysterolecithoides frontilatus, Opisthogonoporoides
hanumanthai, Schistorchis longivesiculurus, Thulinia
microrchis
Siganus fuscescens Hurleytrema ovocaudatum
Siganus vermiculatus Hexangium loossi
Family Sillaginidae
Sillaginopodys Allodecemtestis biacetabulatus, Decemtestis mehrai,
chondropus Erilepturus hamati, Erilepturus thrissocli (=E. hamati)
Sillago lutea Allodecemtestis biacetabulatus, Decemtestis mehrai,
Erilepturus hamate
Sillago sihama Allodecemtestis biacetabulatus, Decemtestis
brevicirrus, Decemtestis mehrai, Helicometrina
septorchis, Helicomerina hexorchis (syn. of H. nimia)
Neoapocreadium caballeroi, Quadriana zakiae
Sillago soringa Erilepturus hamata (=Erilepturus thrissocli)
Family Sisonidae
Mystus gulio Decemtestis srivastavai
Family Soleidae
Aesopia cornuta Macvicaria cynoglossi, Lepocreadioides orientlalis
Brachiurus orientalis Monascus filiformis, Prosorhynchus arabianum
Synaptura commersoni Macvicaria longicaudus
Zebrias altipinnis Lepocreadioides orientalis
Zebrias synapturoides Lepocreadioides orientalis
Family Sparidae
Acanthopagrus berda Erilepturus berdae, Erilepturus orentalis,
Rhipidocotyle septapapillata
Argyrops spinifer Allopodocotyle argyropsi, Macvicaria jagannathi,
M. deeghaensis (=M. jagannathi),
Transversocreadium manteri, Zoogonus argyropsi
Chrysophrys bifasciata Lecithocladium bicaudum
Rhabdosargus sarba Erilepturus sarbi, Helicometrina chauhani,
Helicometrina unica, Lecithaster chrysophrysi,
Stephanostomum simhai, Polyorchitrema oissai
Family Sphyraenidae
Sphyraena barracuda Allodidymozoon apharyngi, Allodidymozoon
cylindricum, Allodidymozoon operculare,
Didymocystis pseudobranchialis, Didymozoon
tetragynae
(continued)
646 Appendix
(continued)
Host taxon Digenean
Sphyraena obtusata Prosorhynchus bengalensis, Rhipidocotyle khalili,
Rhipidocotye sphyraenae, Allodidymozoon
cylindricum, Allodidymozoon operculare,
Didymocystis pseudobranhialis, Didymocystis
singularis, Didymocystis sphyraenae, Didymocystis
triangularis, Didymocystis visakhapatnamensis,
Didymozoon tetragynae, Erilepturus puriensis,
Lecithocladium thapari (=L. glandulum),
Platocystoides polyaster
Sphyraena picuda Didymocystis pseudobranchialis, Platocystoides
polyester
Sphyraena jello Didymocystis singularis, Didymozoon tetragynae,
Lepidapedon nelsoni
Family Stromateidae
Pampus argenteus Helostomatis fotedari, Lecithocladium annulatum,
Lecithocladium excisum, Lecithocladium glandulum,
Lecithocladium harpodontis, Lecithocladium
stromateusi (=L. parviovum), Lepidapedon nelson,
Mehraiella ovocaudatum, Monascus filiformis,
Oculonematobothrium orbitum, Opechona buckleyi,
Opechona vinodae, Opisthomonorcheides delicatus,
Opisthomonorcheides yamagutinsis,
Proctotrematoides indicum, Prodistomum vinodae
(=Opechona vinodae) Stephanostomum srivastavai,
Transversocreadium fotedari
Pampus chinensis Monascus filiformis, Lecithocladium puriense,
Opisthomonorcheides delicatus
Pampus sp. Monascus filiformis
Stromateus cinereus Faustula gangetica, Helostomatis fotedari,
Lepidapedon nelsoni
Family Synbranchiidae
Minous monodactylus Lecithocladium glandulum
Family Syngnathidae
Monopterus cuchei Stomachicola muraenesocis
Family Syngnathidae
Trachyrhamphus serratus Theledera syngagnathus
Family Synodontidae
Harpodon nehereus Gyliauchen ozakii, Lecithocladium harpodontis,
L. simhai
Saurida undosquamis Opegaster ditrematis, Plerururs digitatus
Saurida tumbil Lecithochirium anisotestes, Lecithochirium jairajpuri
(Sterrhurus jairajpuri), Lecithochirium testelobatum,
Lecithochirium tumbulense Lecithocladium
guptai, Lecithocladium saurida, Lecithochirium
simhai, Opegaster ditrematis, Tubulovesicula
angusticauda, Tubulovesicula lindbergi
(continued)
Appendix 647
(continued)
Host taxon Digenean
Trachinocephalus myops Coitocaecum thapari, Erilepturus yamagutii
(=E. hamati) Opegaster ditrematis
Family Tetraodontidae
Arothron hispidus Schistorchis carneus
Gastrophysus lunaris Bianium plicitum, Decemtestis
brevicirrus, Decemtestis kobayashii, Lintonium
pulchrum, Lintonium puriense, Notoporus stunkardi,
Schistorchis carneus
Logocephalus spadiceus Prosogonotrema bilabiatum
Takifugus oblongus Bianium plicitum, Prosorhynchoides manteri,
Diploproctodaeum bombayense
Tetraodon viridipunctatus Bianium indicum, Bianium plicitum,
Diploproctodaeum rameswarensis, Lintonium
indicum, L. tetradontis
Family Terapontidae
Terapon jarbua Bucephalus margaritae, Helicometrina gibsoni,
Karyakartia pambanensis, Opegaster ditrematis,
Qadriana zakiae, Podocotyloides parupenei
Terapon puta Aponurus theraponi, Aponurus beviformis
Helicometrina nimia, Karyakartia pambanense,
Monodhelmis torpedinis, Podocotyloides parupenei,
Pseudametrodaptes bravoae
Terapon theraps Allodecemtestis odeningi, Aponurus orientalis,
Bucephalus fischthali, Karyakartia pambanensis,
Macvicaria yamagutii, Neolasiotocus brayi,
Neonotoporus skrjabini, Opegaster theraponi,
Paropecoelus overstreeti, Podocotyloides parupenei,
Preptetos pritchardae, Pseudopecoelina purius,
Pseudopecoelus dollfusi, Pseudopecoeloides
parupenei, Rhipidocotyle theraponi, Stephanostomum
theraponi, Timonia nahhasi, Trifoliovarium
bombayense, Vesicocoelium marinum
Family Triacanthidae
Triacanthus biaculeata Cotylocreadium triacanthi, Decemtestis mehrai,
Lobatocreadium indicum, Neonotoporus gibsoni
Schistorchis skrjabini, Stephanostomum cloacum,
Stephanostomum triacanthi, Transversocreadium
cablei, Xystretum triacanthi
Triacanthus bilineatus Paradiscogaster indicus
Triacanthus brevirostris Trifoliovarium triacanthi, Xystretum overstreeti,
Xystretum srivastavai
Pseudotriacanthus Cotyloceadium triacanthi, Lecithocladium triacanthi,
strigilifer Paradiscogaser indicus, Parahemiurus engraulisi,
Parahemiurus triacanthusi, Stephanostomum
lucknowensis (=S. triacanthi), Trifoliovarium
triacanthi
(continued)
648 Appendix
(continued)
Host taxon Digenean
Family Trichiuridae
Eupleurogrammus Alcicornis jairajpuri (taaxon inq,) Bucephalus thapari,
muticus Erilepturus guptajaini, Erilepturus trichiurusi
Lecithochirium agarwali Neoprosorhynchus dayali,
Neonotoporus srivastavai, Prosorhynhus
manteri, Prosorhynchus vinodae (=P. manteri)
Trichiurus lepturus Lecithochirium polynemi, Prosorhynchus manteri
Trichiurus savala Macvicaria chilkai
Trichiurus sp. Lecithochirium acutum
Family Uranoscopidae
Uranoscopus guttatus Bucephalus uranoscopi
Family Xiphidae
Xiphias gladius Metadidymozoon branchiale
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Index
A Alcicornis jairajpuri, 40
Accacoeliidae, 183 Alcicornis multidactylus, 40
Accacoeliinae, 183 Alcicornis rauschi, 41
Acanthocolpidae, 367 Alcicornis scomberi, 42
Acanthcolpus, 368 Alcicornis thapari, 43
Acanthocolpus amravati, 372 Allobacciger, 495
Acanthocolpus caballeroi, 369 Allobacciger macrorchis, 496
Acanthocolpus chorinemusi, 372 Allodecemtestis, 442
Acanthocolpus dorabi, 373 Allodecemtestis bilqeesae, 444
Acanthocolpus durghae, 373 Allodecemtestis carangi, 445
Acanthocolpus equulai, 373 Allodecemtestis drepani, 445
Acanthocolpus guptai, 373 Allodecemtestis odeningi, 445
Acanthocolpus inglisi, 373 Allodecemtestis skrjabini, 446
Acanthocolpus liodorus, 370 Allodidymozoon, 191
Acanthocolpus lucknowensis, 373 Allodidymozoon apharyngi, 191
Acanthocolpus luhei, 371 Allodidymozoon cylindricum, 192
Acanthocolpus lutjanusi, 373 Allodidymozoon operculare, 193
Acanthocolpus manteri, 373 Allometadena, 599
Acanthocolpus microtesticulatus, 374 Allometadena rotundum, 599
Acanthocolpus orientalis, 374 Allonematobothrium, 229
Acanthocolpus puriensis, 374 Allonematobothrium epinepheli, 230
Acanthocolpus srivastavai, 374 Allopodocotyle, 447
Acanthocolpus thapari, 374 Allopodocotyle argyropsi, 448
Acanthocolpus valyithurai, 374 Allopseudocolocyntotrema, 219
Acanthosiphodera, 598 Allopseudocolocyntotrema claviforme, 219
Acanthosiphodera bengalense, 598 Allostomachicola, 129
Aephnidiogenidae, 353 Allostomachicola lepturusi, 130
Aephnidiogenes indica, 354 Allostomachicola rauschi, 131
Aephnidiogenes vinodae, 354 Allostomachicola secundus, 13
Albulatremainae, 186 Ametrodaptes, 497
Alcicornis, 36 Ametrodaptes pristipomatis, 497
Alcicornis carangis, 37 Anaporrhutinae, 552
Alcicornis hemiramphi, 38 Anaporrhutum, 553
Alcicornis indicus, 39 Anaporrhutum albidum, 553
Alcicornis inglisi, 40 Anaporrhutum gigas, 553
© Crown 2018 685

https://doi.org/10.1007/978-94-024-1535-3
686 Index
Anaporrhutum largum, 554 Bucephalus barina, 45

Anaporrhutum narayani, 554 Bucephalus elacatus, 46
Anaporrhutum scoliodoni, 554 Bucephalus jagannathai, 48
Anaporrhutum stunkardi, 554 Bucephalus kanagurtai, 49
Anisoporus, 407 Bucephalus margaritae, 49
Anisoporus orientalis, 407 Bucephalus thapari, 51
Antorchis, 579 Bucephalus uranoscopi, 51
Antorchis intermedius, 580 Buckleytrema, 102
Antorchis pomacanthi, 580 Buckleytrema indica, 102
Aphanurinae, 121 Bucephalus yamagutii, 52
Ahemiurus, 121 Bunocotylinae, 127
Ahemiurus karachii, 121
Aphanurus, 122 C
Aphanurus dussumierii, 122 Cadenatellinae, 293
Aphanurus harengulae, 123 Cadenatella, 294
Aphanurus microrchis, 124 Cadenatella dollfusi, 294
Aphanurus stossichii, 124 Carassotrema, 288
Aphanurus tuberculatus, 126 Carassotrema bengalense, 288
Apocreadiidae, 263 Cephalogonimidae, 618
Apocreadiinae, 263 Chalcinotrematinae, 284
Aponurus, 240 Cladorchiidae, 613
Aponurus bengalensis, 240 Coeliodidymocystis, 194
Aponurus breviformis, 240 Coeliodidymocystis kamegai, 194
Aponurus drepani, 242 Colocyntotrematinae, 218
Aponurus laguncula, 243 Coitocaecum, 408
Aponurus orientalis, 244 Coitocaecum bengalense, 409
Aponurus theraponi, 245 Coitocaecum bombayense, 410
Aporocotylidae, 21 Coitocaecum chaetodoni, 411
Atractotrema, 295 Coitocaecum longivesiculum, 412
Atractotrema kuntzi, 295 Coitocaecum purius, 413
Coitocaecum scombri, 414
B Coitocaecum thapari, 415
Bacciger, 581 Coitocaecum thrissoclesis, 416
Bacciger sardinellae, 582 Cotylocreadium, 306
Baccigeroides, 582 Cotylocreadium triacanthi, 306
Baccigeroides cochinensis, 583 Cryptogonimidae, 597
Baccigeroides hafeezullai, 584
Bianium, 302 D
Bianium indicum, 302 Decemtestis, 450
Bianium plicitum, 303 Decemtestis brevicirrus, 451
Bianium purii, 305 Decemtestis cynoglossi, 452
Bivesiculoidea, 29 Decemtestis dollfusi, 452
Bivesiculidae, 29 Decemtestis kobayashii, 453
Bivesicula, 30 Decemtestis marginoacetabultus, 455
Bivesicula australis, 30 Decemtestis mehrai, 453
Bivesiculoides, 31 Decemtestis singhi, 455
Bivesiculoides scari, 31 Decemtestis srivastavai, 456
Brachyenteron, 579 Decemtestis varmai, 457
Brachyenteron parexocoeti, 577 Deretrema, 573
Bucephalidae, 35 Deretrema plotosi, 573
Bucephalinae, 36 Dictysarcidae, 186
Bucephalus, 44 Didymocodium, 221
Bucephalus arabiana, 44 Didymocodium euthyni, 221
Bucephalus arabiansis, 45 Didymocystis, 195
Index 687
Didymocystis alalongae, 196 Elopsium secundum, 88

Didymocystis bifasciata, 197 Elopsium stunkardi, 89
Didymocystis bifurcata, 197 Elyptophallinae, 149
Didymocystis dissimilis, 198 Elyptophalloides, 149
Didymocystis exigua, 198 Enenteridae, 357
Didymocystis guttata, 199 Enenterum, 357
Didymocystis oesophagicola, 200 Enenterum mannarense, 358
Didymocystis pinnicola, 200 Enenterum minutum, 359
Didymocystis pseudobranchialis, 201 Erilepturus, 139
Didymocystis semiglobularis, 202 Erilepturus hamati, 139
Didymocystis singularis, 202 Euryakina, 600
Didymocystis sphyraenae, 202 Euryakina marina, 600
Didymocystis superpalati, 203
Didymocystis triangularis, 204 F
Didymocystis visakhapatnamensis, 205 Faustulidae, 578
Didymocystis wedli, 205 Faustula, 584
Didymozoidae, 188 Faustula basiri, 585
Didimozoinae, 189 Faustula brevichrus, 586
Didymozoon, 208 Faustula gangetica, 586
Didymozoon diverticulatum, 209 Fellodistomidae, 83
Didymozoon lobatum, 209 Fellodistominae, 84
Didymodiclininae, 224
Didymodiclinus, 224 G
Didymodiclinus branchialis, 224 Genolopa, 498
Didymodiclinus reticulum, 225 Genolopa ampullacea, 498
Didymodiclinus spilonotopteri, 226 Genolopa bychowskii, 499
Dinurinae, 129 Genolopa trifolifer, 500
Dinurus, 133 Gorgoderidae, 558
Dinurus barbatus, 133 Gomtia, 622
Dinurusbreviductus, 134 Gomtia caranxi, 622
Dinurus hippuri, 135 Gorgoderidae, 551
Dinurus ivanosi, 135 Gyliuchenidae, 363
Dinurus longisinus, 136 Gyliauchen, 364
Dinurus thapari, 137 Gyliauchen indicum, 365
Dinurus tornatus, 137 Gyliauchen ozakii, 366
Diphtherostomum, 571
Diphtherostomum indicum, 571 H
Diplohurleytrema, 531 Haploporus pseudoindicus, 278
Diplohurleytrema nahhasi, 531 Haplosplanchnidae, 109
Diploproctodaeum, 307 Haplosplanchninae, 110
Diploproctodaeum bombayense, 307 Haplosplanchnus, 110
Diploproctodaeum rameswarensis, 308 Haplosplnchnus benglensis, 115
Dollfustrema, 68 Haplosplanchnus caudatus, 111
Dollfustrema bengalense, 68 Haplosplanchnus guptai, 112
Haplosplanchnus indica, 115
E Haplosplanchnus orientalis, 115
Echeneidocoelium, 309 Haplosplanchnus otolithi, 115
Echeneidocoelium indicum, 310 Haplosplanchnus purii, 113
Elongoparorchis, 187 Haplosplanchnus stunkardi, 115
Elongoparorchis pneumatis, 187 Haplosplanchnus vinodae, 114
Elopsium, 84 helicometra, 480
Elopsium indicum, 85 Helicometra filamentosa, 480
Elopsium manteri, 86 Helicometra gibsoni, 481
Elopsium puriense, 87 Helicometrina, 483
688 Index
Helicometrina chauhani, 485 K

Helicometrina hexorchis, 485 Kamegaia, 211
Helicometrina indica, 485 Kamegaia kawakawa, 212
Helicometrina nimia, 483 Karyakartia, 314, 315
Helicometrina orientalis, 486 Koseira, 359
Helicometrina quadrorchis, 485 Koseira manteri, 360
Helicometrina scomberi, 486 Koseira nagatyi, 359
Helicometrina septorchis, 486
Helicometrina unica, 486 L
Helicometroides, 533 Lasiotocus, 501
Helicometroides leiperi, 534 Lasiotocus hastai, 503
Helicometroides pseudovitellosus, 534 Lasiotocus maculatus, 504
Hirudinellidae, 236 Lasiotocus odhneri, 506
Hirudinella, 236 Lasiotocus overstreeti, 506
Helostomatinae, 613 Lasiotocus polynemi, 507
Helostomatis, 613 Lasiotocus puriensis, 507
Helostomatis fotedari, 613 Lasiotocus srivastavai, 508
Helostomatis muelleri, 613 Lecithasteridae, 238
Helostomatis simhai, 613 Lecithasterinae, 239
Hemiurinae, 120, 177 Lecithaster, 245
Heterolebes, 487 Lecithaster bombayense, 245
Heterolebes buckleyi, 487 Lecithaster chrysophrysi, 246
Heterolebes spindalis, 488 Lecithobotrys, 279
Hexangium, 616 Lecithobotrys vitellosus, 279
Hexangium sigani, 616 Lepocreadiidae, 300
Hirudinella ventricos, 236 Lecithochiriinae, 165
Homalometron, 264 Lecithochirium, 165
Homalometronvinodae, 264 Lecithochirium acutum, 167
Huridostomum, 500 Lecithochirium durdanae, 168
Huridostomum formionis, 500 Lecithochirium fotedari, 169
Hurleytrematinae, 530 Lecithochirium indicum, 169
Hurleytrema, 532 Lecithochirium jairajpuri, 170
Hurleytrema ovocaudatum, 532 Lecithochirium leiperi, 171
Hurleytrematoides, 535 Lecithochirium polynemi, 172
Hurleytrematoides filiformis, 535 Lecithochirium pristipomatis, 173
Hymenocottinae, 115 Lecithochirium pritchardae, 173
Hymenocotta, 116 Lecithochirium testelobatus, 174
Hymenocotta mulli, 116 Lecithochirium tumbilense, 174
Hypocreadium cavum, 311 Lecithocladium, 150
Hypocreadium indicum, 312 Lecithocladium annulatum, 150
Hypocreadium patellare, 313 Lecithocladium angustiovum, 156
Hysterolecithinae, 247 Lecithocladium apolecti, 157
Hysterolecithoides, 248 Lecithocladium bengalense, 158
Hysterolecithoides frontilatus, 248 Lecithocladium glandulum, 156
Hysterolecithoides yamagutii, 249 Lecithocladium guptai, 159
Lecithocladium harpodontis, 160
I Lecithocladium jagannathi, 161
Indodidymozoon, 210 Lecithocladium megalaspis, 162
Indodidymozoon madhaviae, 210 Lecithocladium parviovum, 163
Indoglomeritrema, 220 Lecithocladium puriense, 164
Indoglomeritrema epinepheli, 220 Lecithostaphylus, 574
Isorchis, 296 Lecithostaphylus nitens, 574
Isorchis skrjabini, 296 Lecithostaphylus parexocoeti, 576
Isorchis chanosi, 297 Leiomonorchis, 509
Index 689
Leiomonorchis mamaevi, 509 Metanematobothrium bivitellatum, 230

Leiomonorchis secundus, 510 Microscaphidiidae, 618
Lepidauchen nicolli, 403 Masenia, 618
Lepidophyllinae, 572 Masenia carangi, 618
Lepocreadioides, 316 Masenia orissai, 619
Lepocreadioides orientalis, 316 Masenia quiloni, 620
Lepocreadium, 318 Masenia upeneusi, 621
Lintonium, 90 Monascus, 93
Lintonium indicum, 91 monascus filiformis, 94
Lintonium pseudovibex, 91 monodhelmis, 103
Lintonium pulchrum, 92 monodhelmis chilkai, 104
Lintonium puriense, 92 monodhelmis militaris, 104
Lintonium tetradontis, 93 monodhelmis orissai, 104
Lobatocreadium, 318 Monodhelmis thapari, 105
Lobatocreadium balistes, 318 Monodhelmistorpedinis, 106
Lobatocreadium gupti, 320 Monorchiidae, 493
Lobatocreadium indicum, 321 Monorchiinae, 494
Lobatocreadium manteri, 322 Monorcheides, 511
Lobatocystis, 212 Monorcheides hafeezullahi, 511
Lobatocystis bengalensis, 212 Monorchis, 512
Lobatocystisyaito, 213 Monorchis bengalensis, 512
Lutianotrema, 223 Monorchis minutus, 513
Monostephanostomum, 374
M Monostephanostomum loossi, 375
Maculifer, 489 Monostephanostomum mesospinosum, 375
Maculiferindicus, 489
Maculifer spiralis, 490 N
masenia, 618 Neidhartia, 79
Masenia carngi, 618 Neidhartia microrhyncha, 79
Masenia orissai, 619 Neidhartia neidharti, 80
Masenia quiloni, 620 Neoapocreadium, 265
Masenia upeneusi, 621 Nematobothrium, 228
Macvicaria, 460 Neoapocreadium caballeroi, 266
Macvicaria chilkai, 461 Neoapocreadium imtiazi, 266
Macvicaria cynoglossi, 462 Neoapocreadium karwrense, 267
Macvicaria deeghaensis, 463 Neogonapodasmius, 226
Macvicaria filamentus, 463 Neogonapodasmius hemiramphi, 227
Macvicaria indica, 463 Neohypocreadium, 323
Macvicaria jagannathi, 464 Neohypocreadium chaetodoni, 323
Macvicaria longicaudus, 465 Neolasiotocus, 537
Macvicaria puriensis, 466 Neolasiotocus brayi, 537
Macvicaria yamagutii, 466 Neolepocreadium, 355
Manteriella, 417 Neolepocreadium caballeroi, 356
Manteriella yamagutii, 418 Neolepocreadium trachinoti, 355
Megasoleninae, 286 Neometadena, 603
Mehrialla, 602 Neometadena ovale, 603
Mehrialla ovocaudatum, 602 Neometadidymozoon, 214
Melanocystis, 213 Neometadidymozoon polymorphis, 214
Melanocystis kawakawa, 213 Notoporus stunkardi, 422
Metadidymozoon, 214 Neomultitestis bengalensis, 325
Metadidymozoon branchiale, 214 Neonematobothrium, 232
Metanematobothrioides, 231 Neonematobothrium dorsale, 232
Metanematobothrioides branchialis, 231 Neonematobothrium kawakawa, 232
Metanematobothrium, 230 Neonotoporus, 472
690 Index
Neonotoporus fotedari, 472 Opisthomonorcheides nigeri, 544

Neonotoporus gibsoni, 473 Opisthomonorcheides overstreeti, 544
Neonotoporus leiognathi, 474 Opisthomonorcheides pelloni, 544
Neonotoporus overstreeti, 474 opisthomonorcheides shindei, 544
Neonotoporus skrjabini, 476 Opisthomonorcheides simhai, 544
Neonotoporus srivastavai, 476 Opisthomonorcheides stromatensis, 544
Neopisthadena, 181 Opisthomonorcheidesupeneusi, 544
Neopisthadena habei, 181 Opisthomonorcheides yamagutii, 544
Neoprosorhynchus, 81 Opisthomonorcheides yamagutinsis, 545
Neoprosorhynchus purius, 81 Opisthorchiidae, 622
Neotimonia, 514 Orchispirium, 22
Neotimonia caballeroi, 514 Orchispirium heterovitellatum, 22
Notoporus, 419 Orientodiploproctodaeum, 604
Notoporus fotedari, 419 Orientodiploproctodaeum diacanthi, 605
Notoporus gibsoni, 420
Notoporus pristipomatis, 421 P
Paradeontacylix, 23
O Paradeontacylix megalspium, 23
Oculonematobothrium, 233 Paradiscogaster, 588
Oculonematobothrium orbitum, 233 Paradiscogaster arabi, 588
Opechona, 326 Paradiscogaster farooqi, 589
Opechona ahmadi, 327 Paradiscogaster indicus, 590
Opechona bacillaris, 328 Paradiscogaster pritchardae, 591
Opechona buckleyi, 329 Pseudopecoelus, 434
Opegaster, 424 Parahemiurus, 177
Opegaster ditrematis, 424 Parahemiurus cameroni, 177
Opepherocystis, 221 Parahemiurus engraulisi, 178
Opepherocystis kawakawa, 222 Parahemiurus gastricus, 177
Opepherotrematoides, 222 Parahemiurus hamilton, 179
Opepherotrematoides multitubularis, 222 Parahemiurus indicus, 179
Opecoelidae, 405 Parahemiurusmadrasensis, 179
Opecoelinae, 406 Parahemiurus puriensis, 180
Opisthadeninae, 181 Parahemiurus simhai, 180
Opisthodiplomonorchis, 539 Parahemiurus triacanthusi, 180
Opisthodiplomonorchis elongatus, 539 Parahemiurus yanamensis, 180
Opisthogonoporoides, 329 Paramonorcheides, 515
Opisthogonoporoides acanthuri, 330 Paramonorcheides selaris, 516
Opisthogonoporoides hanumanthai, 331 Paramphistomoidea, 613
Opisthogonoporoides thapari, 332 Paralecithobotrys, 284
Opisthogonoporus, 333 Paralecithobotrys overstreeti, 284
Opisthogonoporus indicus, 333 Paropecoelus, 425
Opistholebetidae, 486 Paropecoelus dollfusi, 426
Opistholebes, 490 Paropecoelus indicus, 427
Opistholebes tetradontis, 491 Paropecoelus overstreeti, 428
Opisthomonorcheidinae, 536 Paropecoelus pritchardae, 429
Opisthomonorcheides, 540 Paropecoelus sciaenid, 431
Opisthomonorcheides aspinosus, 542 Paucivitellosus, 32
Opisthomonorcheides chaetodoni, 542 Paucivitellosus hanumanthai, 32
Opisthomonorcheides decapteri, 542 Petalodistomum, 554
Opisthomonorcheides gibsoni, 543 Petalodistomum hanumanthai, 555
Opisthomonorcheides karwarensis, 543 Petalodistomum singhi, 556
Opisthomonorcheides madhaviae, 544 Petalodistomum yamagutii, 556
Opisthomonorcheides manteri, 544 Phyllodistomum, 564
Opisthomonorcheides nahhasi, 543 Phyllodistomum lancea, 564
Index 691
Phyllodistomum lewisi, 565 Prosorhynchoides pritchardae, 58

Phyllotrema, 467 Prosorhynchinae, 67
Phyllotrema tetracaudum, 468 Prosorhynchus, 69
Plagioporinae, 441 Prosorhynchus atlanticus, 70
Platocystoides, 216 Prosorhynchus bengalensis, 71
Platocystoides polyester, 216 Prosorhynchus caballeroi, 71
Pleorchis, 401 Prosorhynchus chorinemi, 72
Pleorchis sciaenae, 402 Prosorhynchus epinepheli, 73
Plerurinae, 175 Prosorhynchus indicus, 73
Plerurus, 175 Prosorhynchus madhaviae, 74
Plerurus digitatus, 175 Prosorhynchus manteri, 75
Podocotyloides, 469 Prosorhynchus orientalis, 76
Podocotyloides dorabus, 469 Prosorhynchus pacificus, 76
Podocotyloides parupenei, 469 Prosorhynchus tewariae, 78
Podocotyloides pedicellatus, 470 Prosorhynchus truncatus, 78
Podocotyloides pentavitellosus, 470 Prosorhynchus platycephali, 77
Podocotyloides yamagutii, 471 Prosorhynchus vinodae, 79
Polyorchitrema, 606 Pseudametodaptes, 521
Polyorchitrema orissai, 606 Pseudametrodaptes bravoae, 521
Preptetos, 334 Pseudametrodaptes caballeroi, 522
Preptetos madrasensis, 334 Pseudametrodaptes fischthali, 524
Preptetos pritchardae, 335 Pseudametrodaptes secundus, 525
Proctotrema, 517 Pseudobacciger, 593
Proctotrema guptai, 517 Pseudobacciger cablei, 594
Proctotrematoides, 519 Pseudobacciger harengulae, 595
Proctotrematoides indicum, 519 Pseudocolocyntotrema, 222
Prodistomum, 336 Pseudocolocyntotrema yaito, 223
Prodistomum gaevskayae, 337 Pseudohapladena, 289
Prodistomum mohsini, 338 Pseudohapladena martini, 290
Prodistomum orientalis, 339 Pseudolepidapedon, 403
Prosorchis, 260 Pseudolepidapedon indica, 403
Prosorchis breviformis, 260 Pseudolepocreadioides, 345
Prosorhynchoides, 54 Pseudolepocreadioides bombayensis, 346
Prodistomum travassossi, 341 Pseudolepocreadioides secundus, 347
Prodistomum vinodae, 343 Pseudolepocreadioides symmetrorchis, 347
Prodistomum waltairense, 344 Pseudolepocreadioides thapari, 349
Proeneterum, 362 Pseudopecoelina, 478
Proenenterum manteri, 362 Pseudopecoelina elongata, 478
Prolecithinae, 251 Pseudopecoelina stunkardi, 479
Prolecitha, 252 Pseudopecoeloides, 432
Prolecitha obesa, 252 Pseudopecoeloides orientalis, 433
Pronopyrna, 592 Pseudopecoeloides puriensis, 640
Pronopyrna petrowi, 593 Pseudopecoelus odeningi, 438
Prosogonarium, 107 Pseudopecoelus pritchardae, 439
Prosogonarium plotosi, 107 Pseudopecoelus stunkardi, 440
Prosogonotrematinae, 258 Pseudopisthomonorchis, 545
Prosogonotrema, 258 Pseudopisthomonorchis carangi, 545
Prosogonotrema bilabiatum, 259 Pseudopisthomonorchis hanumanthai, 546
Prosorchiinae, 260 Pseudopisthomonorchis thapari, 548
Prosorhynchoides belonea, 54 Pseudopisthomonorchis secundus, 547
Prosorhynchoides brayi, 55
Prosorhynchoides chorinemi, 56 Q
Prosorhynchoides karvei, 56 Quadriana, 141
Prosorhynchoides microcirrus, 57 Quadriana zakiae, 141
692 Index
R Stephanostomum ditrematis, 383

Renodidymocystis yamagutii, 217 Stephanostomum equula, 384
Rhipidocotyle, 59 Stephanostomum gazzai, 385
Rhipidocotyle apapillosa, 60 Stephanostomum lucknowense, 385
Rhipidocotyle, 60 Stephanostomum madhaviae, 385, 386
Rhipidocotyle indicus, 61 Stephanostomum microsomum, 386–388
Rhipidocotyle khalili, 62 Stephanostomum orientalis, 388
Rhipidocotyle ligulum, 63 Stephanostomum polynemi, 389
Rhipidocotyle pentagonum, 64 Stephanostomum pseudoditrematis, 390, 391
Rhipidocotyle septapapillata, 65 Stephanostomum simhai, 391
Rhipidocotyle sphyraenae, 65 Stephanostomum srivastavai, 392
Stephanostomum theraponi, 393, 394
S Stephanostomum triacanthi, 394, 395
Saccocoelium, 276 Stomachicola, 143
Saccocoelium tripathi, 277 Stomachicola muraenesocis, 144
Saccocoelioides, 285 Skrjabinozoum, 234
Saccocoelioides chilkaensis, 285 Skrjabinozoum waltairense, 234
Saturnius, 127 Skrjabinozoum yamagutii, 235
Saturnius segmentatus, 127
Saturnius valamugilis, 128 T
Schikhobalotrematinae, 117 Tandanicolidae, 101
Schikhobalotrema, 117 Tetrochetus, 183, 186
Schikhobalotrema acutum, 117 Tetrochetus aluterae, 184
Schistorchiinae, 268 Tetrochetus coryphaenae, 183
Schistorchis, 268 Tetrochetus macrorchis, 185
Schistorchis carneus, 268 Thryssatrema, 95
Schistorchis longivesiculurus, 269 Thryssatrema hanumantharaoi, 95
Schistorchis manteri, 270 Thulinia, 250
Schistorchis skrjabini, 271 Thulinia microrchis, 250, 251
Sclerodistomidae, 256 Timonia, 526
Sclerodistominae, 257 Timonia indica, 526
sclerodistomum, 257 Timonia nahhasi, 527
Sclerodistomum chirocentrusi, 257 Timonia stunkardi, 528
siphoderina, 607 Timonia vinodae, 529
Siphoderina americanus, 608 Tormopsolus, 398
Siphoderina grandispinus, 609 Tormopsolus filiformis, 399
Siphoderina hirastricta, 610 Transversocreadium, 349
Siphoderina sootai, 610 Transversocreadium cablei, 350
Skrjabinolecithum, 291 Transversocreadium fotedari, 350
Skrjabinolecithum bengalense, 291 Transversocreadium manteri, 351
Skrjabinolecithum indicum, 292 Transversocreadium secundus, 352
Staphylorchis, 557 Transversocreadium yamagutii, 353
Staphylorchis cymatodes, 557 Transversotrematidae, 273
Stenakrinae, 472 Transversotrema, 273, 274
Stephanostomoides, 396 Transversotrema patialense, 274
Stephanostomoides tenuis, 397 Trifoliovariinae, 253
Stephanostomum adinterruptum, 377 Trifoliovarium, 253
Stephanostomum anterospinosum, 378 Trifoliovarium bombayense, 253
Stephanostomum attenuatum, 378 Trifoliovarium triacanthi, 255
Stephanostomum bhuneswari, 379 Tubulovesicula, 147
Stephanostomum casum, 380 Tubulovesicula angusticauda, 147
Stephanostomum chauhani, 381 Tubulovesicula lindbergi, 148
Stephanostomum cloacum, 382
Index 693
U Xystretum chauhani, 558

Unisaccus guptai, 280 Xystretum manteri, 559
Unisaccus manteri, 281 Xystretum overstreeti, 560
Unisaccus mugilis, 282 Xystretum srivastavai, 561
Unisaccus overstreeti, 283 Xystretum thapari, 562
Xystretum triacanthi, 563
V
Vesicocoelium, 471 Y
Vesicocoelium marinum, 471 Yamagutia, 595
Vitellibaculum, 286 Yamagutia lateroporus, 596
Vitellibaculum fischthali, 286
Z
W Zoogonidae, 567
Waretrema, 292 Zoogoninae, 568
Waretrema piscicolum, 292 Zoogonus, 569
Zoogonus argentopsi, 569
X Zoogonoides, 570
Xystretum, 558 Zoogonoides pyriformis, 570

Digenetic Trematodes of Indian Marine Fishes

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Digenetic Trematodes of Indian Marine Fishes

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Rokkam

Madhavi · Rodney A. Bray

ISBN 978-94-024-1533-9 ISBN 978-94-024-1535-3 (eBook)

Printed on acid-free paper

Visakhapatnam, India Rokkam Madhavi

Part I Systematic Account—Order Diplostomida Olson,

Part II Systematic Account—Order Plagiorchiida La Rue, 1957

Family Didymozoidae Monticelli, 1888 . . . . . . . . . . . . . . . . . . . . . . . 188

Fig. 1.1 Map of India showing the various coastal areas

institutions and universities situated in coastal districts are undertaking investiga-

Fixation is followed by staining in carmalum or haematoxylin, dehydrating in

General Organization and Life Cycles (Fig. 3.1a–e)

atrium is provided with an accessory copulatory organ consisting of a solid anteriorly

Life Cycles of Digenetic Trematodes

Fig. 3.1 (continued)

Family Aporocotylidae Odhner, 1912

[Syn. Sanguinicolidae von Graff, 1907]

Fig. 4.1 Orchispirium

Genus Orchispirium Madhavi & Rao, 1970

Diagnosis: Body lanceolate. Oesophagus long, surrounded by gland cells, enlarged

Description: Body spatulate, 3230–4480 long 990–1280 wide, tapering gradually

Remarks: The important characteristics of O. heterovitellatum are considered to be:

Host: Carangidae: Megalaspis cordyla (Linnaeus)

Fig. 4.2 Paradeontacylix

Fig. 4.3 P. megalaspium

Family Bivesiculidae Yamaguti, 1939

[Syn. Paucivitellosidae Yamaguti, 1971, Treptodemidae Yamaguti, 1971]

Vitellarium follicular, follicles distributed widely, eggshell thick …................. 2

Bivesicula australis Crowcroft, 1947 (Fig. 5.1)

Fig. 5.1 Bivesicula australis

Description: Body 990–1072 long, 385–453 wide, uniform. Eyespot pigment

Fig. 5.2 Bivesiculoides

Genus Paucivitellosus Coil, Reid & Kuntz, 1965.

Description: Body pear shaped 480–592×20–384. Tegument with minute spines.

Fig. 5.3 Paucivitellosus

Remarks: Gnanamani (1989) differentiated P. hanumanthai from P. fragilis Coil,

Gnanamani (1989) showed that the snail Clypeomorus clypeomorus Jusseaume

Family Bucephalidae Poche, 1907

Bucephalidae is a large family of digenean parasites occurring in the gut of marine,

Key to subfamilies of Indian genera of bucephalidae

Subfamily Bucephalinae Poche, 1907

Diagnosis: Body cylindrical, spined. Rhynchus consists of simple sucker with or

Key to Indian genera of Bucephalinae

Diagnosis: Body elongate. Rhynchus either funnel-shaped or short U-shaped struc-

Description: Body elongate 1580–2100 long, 300–400 wide. Rhynchus wedge-

Fig. 6.1 Alcicornis carangis

Alcicornis hemiramphi Gupta & Tandon, 1983 (Fig. 6.2)

Fig. 6.2 Alcicornis

Description: Body elongate, narrow, 3150–5020 long, 330–370 wide at testicular

Fig. 6.3 Alcicornis indicus

Alicornis inglisi Gupta & Jain, 1993

Remarks: This species is here considered as taxon inquirendum.

Fig. 6.4 Alcicornis

Fig. 6.5 Alcicornis

Description: Body long, slender, 2240–2640 long by 192–288 wide. Tegument

Remarks: This species is characterized by the presence of 15 tentacles on the

Alcicornis rauschi Gupta & Jain, 1993

Remarks: The description of the species is based on five macerated specimens.

Alcicornis scomberi Gupta & Ahmad, 1976 (Fig. 6.6)

Description: Body elongate, slender, slightly tapering anteriorly, rounded poste-

Fig. 6.6 Alcicornis

Alcicornis thapari Hafeezullah & Siddiqi, 1970

Description: Body elongate, slightly tapering anteriorly, rounded posteriorly,

Key to Indian species of Alcicornis