STUDIES ON DIGENETIC TREMATODES OF SOME FISHES OF KARACHI COAST

NEELOFER SHAUKAT

Department of Zoology, Jinnah University For Women, Nazimabad, Karachi, Pakistan. 2008 STUDIES ON DIGENETIC TREMATODES OF SOME FISHES OF KARACHI COAST

BY

NEELOFER SHAUKAT M.Sc., M.Phil

THESIS SUBMITED TO JINNAH UNIVERSITY FOR WOMEN FOR FULFILMENT OF THE REQUIRMENT FOR THE DEGREE OF DOCTOR OF PHILOSOPHY (Ph.D.) IN THE SUBJECT OF ZOOLOGY

Department of Zoology, Jinnah University For Women, Nazimabad, Karachi, Pakistan. 2008

TABLE OF CONTENTS

CERTIFICATE………………………………………………..i

DEDICATION………………………………………………...ii

ACKNOWLEDGEMENTS………………………………iii-iv

LIST OF TABLES……………………………………………v

LIST OF FIGURES………………………………………vi-vii

SUMMARY……………………………………………...viii-xii

INTRODUCTION…………………………………………1-18

REVIEW OF LITERATURE………………………...... 19-52

MATERIALS AND METHODS………………………..65-67

- Collection of Specimens………………………………...65-66

- Fixation and Preparation of Permanent slides……….66-67

DESCRIPTIONS OF OF THE GENERA...69-231

1. Pleorchis heterorchis n.sp……………………………...69-76

- Diagnosis………………………………………………...69-71

- Principle Measurements………………………………..71-72

- Etymology…………………………………………………..72

- Remarks…………………………………………………72-76 2. Decemtestis johnii n.sp………………………………...77-82

- Diagnosis………………………………………………...77-78

- Principle Measurements………………………………..78-79

- Etymology…………………………………………………..79

- Remarks…………………………………………………79-82

3. Lecithocladium cybii n.sp……………………………...83-90

- Diagnosis………………………………………………...83-84

- Principle Measurements…………………………………...85

- Etymology…………………………………………………..86

- Remarks…………………………………………………86-90

4. Lecithocladium karachiensis n.sp……………………..91-98

- Diagnosis………………………………………………...91-92

- Principle Measurements…………………………………...93

- Etymology…………………………………………………..93

- Remarks…………………………………………………94-98

5. Lecithocladium magnasoma n.sp…………………….99-106

- Diagnosis……………………………………………….99-100

- Principle Measurements………………………………….101 - Etymology…………………………………………………102

- Remarks………………………………………………102-106

6. Lecithocladium magnavesicula n.sp………………..107-115

- Diagnosis……………………………………………...107-108

- Principle Measurements……………………………..108-109

- Etymology…………………………………………………109

- Remarks………………………………………………109-115

7. Lecithocladium lateropharyngium n.sp…………….116-123

- Diagnosis……………………………………………...116-117

- Principle Measurements……………………………..117-118

- Etymology…………………………………………………118

- Remarks………………………………………………118-123

8. Tubulovesicula olivaceus n.sp………………………124-130

- Diagnosis……………………………………………...124-125

- Principle Measurements……………………………..125-126

- Etymology…………………………………………………126

- Remarks………………………………………………127-130

9. Tubulovesicula microcaudum n.sp………………….131-139

- Diagnosis……………………………………………...131-132

- Principle Measurements……………………………..132-133

- Etymology…………………………………………………133

- Remarks………………………………………………134-139

10. Tubulovesicula magnacirrosa n.sp………………...140-147

- Diagnosis……………………………………………...140-141

- Principle Measurements……………………………..141-142

- Etymology…………………………………………………142

- Remarks………………………………………………142-147

11. Tubulovesicula karachiensis n.sp………………….148-152

- Diagnosis……………………………………………...148-149

- Principle Measurements……………………………..149-150

- Etymology…………………………………………………150

- Remarks………………………………………………150-152

12. Stomachicola muraenesocis Yamaguti, 1934……..153-159

- Diagnosis……………………………………………...153-154

- Principle Measurements………………………………….155 - Etymology…………………………………………………156

- Remarks………………………………………………156-159

13. Prosogonotrema diacanthi Bilqees, 1980………….160-166

- Diagnosis……………………………………………...160-161

- Principle Measurements……………………………..161-162

- Etymology…………………………………………………162

- Remarks………………………………………………162-166

14. Plagioporus karachiensis n.sp……………………..167-177

- Diagnosis……………………………………………...167-168

- Principle Measurements……………………………..168-169

- Etymology…………………………………………………169

- Remarks………………………………………………169-177

15. Stephanostomum gibsoni n.sp……………………..178-196

- Diagnosis……………………………………………...178-179

- Principle Measurements……………………………..180-181

- Etymology…………………………………………………181

- Remarks………………………………………………181-196

16. Bucephalus otolithi n.sp…………………………...197-209

- Diagnosis……………………………………………...197-198

- Principle Measurements……………………………..198-199

- Etymology…………………………………………………199

- Remarks………………………………………………199-209

17. Bucephalus mujibi n.sp……………………………210-217

- Diagnosis……………………………………………...210-211

- Principle Measurements……………………………..211-212

- Etymology…………………………………………………212

- Remarks………………………………………………212-217

18. Prosorhynchus magnacirrus n.sp…………………218-231

- Diagnosis……………………………………………...218-219

- Principle Measurements……………………………..219-220

- Etymology…………………………………………………220

- Remarks………………………………………………220-231

DISCUSSION………………………………………….269-278

REFERENCES………………………………………...279-328

DEDICATION

Dedicated to my adoring and devoted parents to whom I owe everything for what I am today.

ii ACKNOWLEDGEMENTS

Foremost, I gratify almighty Allah who consecrated me with competence and provided the opportunity to consummate the present research work.

It is my privilege to be very thankful to The Vice Chancellor, Dr. Riaz

Ahmed Hashmi, Jinnah University for Women for full cooperation and providing research facilities throughout this research work.

I am really indebted to dear Prof. Dr. Bilqees F. Mujib, Supervisor of my work, Department Of Zoology, Jinnah University for Women, for her diligent supervision, invaluable help, constant encouragement and patience throughout this research.

I feel great honour to express my thanks to my Chairperson, Ms. Bushra

Khalil, Associate Professor, Department of Zoology and fellow faculty members for their continuous cooperation and providing facilities in the department.

iii I would also like to thank Dr. Ali Khan Director CDRI, PARC, Karachi

University Campus and Dr. R.R. Ghazi, Ex-Director, Vertebrate Pest

Control Centre, PARC, Karachi University Campus for reading the manuscript.

I would also like to thank Rehmat Bibi, Research officer, Department of

Zoology, Jinnah University for Women for her help in collection material and typing the manuscript.

Last but not the least I am grateful to my devoted husband, sister-in-laws, my son and my daughter Naureen for helping in various ways and for providing, peaceful and effective environment.

iv LIST OF TABLES

Table-1: List of known trematode species and their hosts in Pakistan

including new species………………………………...... 53-64

Table-2: Number of fishes of various families collected during the

present studies………………………………………………..68

Table-3: Family wise list of fishes showing rate of infection during the

present studies…………………………………………232-233

Table-4: Fish hosts and trematodes identified during the present studies………………………………………………………234

v

LIST OF FIGURES

Fig-1: Pleorchis heterorchis n.sp., holotype, entire………235

Fig-3: Pleorchis heterorchis n.sp., paratype, entire………237

Fig-5: Decemtestis johnii n.sp., holotype, entire…………239

Fig-6: Lecithocladium cybii n.sp., holotype, entire………240

Fig-8: Lecithocladium karachiensis n.sp., holotype,

entire……………………………………………….242

Fig-9: Lecithocladium magnasoma n.sp., holotype,

entire……………………………………………….243

Fig-11: Lecithocladium magnavesicula n.sp., holotype,

entire……………………………………………...245

Fig-13: Lecithocladium lateropharyngium n.sp., holotype,

entire……………………………………………...247

Fig-15: Tubulovesicula olivaceus n.sp., holotype, entire…249

Fig-17: Tubulovesicula microcaudum n.sp., holotype,

entire……………………………………………...251

vi Fig-19: Tubulovesicula magnacirrosa n.sp., holotype,

entire……………………………………………...253

Fig-21: Tubulovesicula karachiensis n.sp., holotype,

entire……………………………………………...255

Fig-22: Stomachicola muraenesocis Yamaguti, 1934…....256

Fig-24: Prosogonotrema diacanthi Bilqees, 1980……...... 258

Fig-26: Plagioporus karachiensis n.sp., holotype, entire...260

Fig-27: Stephanostomum gibsoni n.sp., holotype, entire…261

Fig-28: Stephanostomum gibsoni n.sp……………………262

Fig-31: Bucephalus otolithi n.sp., holotype, entire……….264

Fig-33: Bucephalus mujibi n.sp., holotype, entire………..266

Fig-34: Prosorhynchus magnacirrus n.sp., holotype,

entire……………………………………………...268

vii SUMMARY

Marine fish parasitology is important both from economic and human health point of view. Hardly there is any country where fish is not included in the human diet. Fishes are also zoonotically important, since several diseases are caused in humans by fish parasites including

Anisakiasis, Van Thiel et al., 1960 . Hundreds of fishes suffer due to helminth parasites.

Fish are valuable sources of high grade protein and organic products.

They occupy a significant position in the socio-economic status of the

South-Asian countries by providing the population not only the nutritious food but also income and employment opportunities.

Available literature revealed that the studies so far conducted in Pakistan about the parasites in marine fishes are not adequate, more research is needed since a large number of fishes have not been investigated for trematode parasites. Therefore, this project was undertaken.

During the present investigation (2005-2007) a total of 791 fishes belonging to 14 families, 10 genera and 18 species were collected from

viii different localities of West Wharf, Karachi. The fish species collected include: Pseudosciaena diacanthus Weber and de Beaufort (73),

Muraenesox cinereus Forsskol (50), Stromateus cinereus Bloch (25),

Stromateus sinensis Euph. (20), Chirocentrus dorab Forsskol (16),

Crenidens indicus Forsskol (19), Muraena sp. Forsskol (28),

Platycephalus scabar Laeneans (43), Plectorhynchus cinctus (T.S.)

(50), Cybium gutatum Cuvier (66), Caranx affinis Ruppell (13),

Parastromateus niger Bloch (75), Lates calcarifer Bloch (20),

Pomadasys olivaceum Day (67), Otolithus argenteus Chu, Lo and Wu

(112), Lutjanus johnii Bloch (78), Arius serratus Day (23) and

Psettodes erumei Schneider (13). The gut contents and other organs of these fishes were examined for trematode parasites.

For detail studies, trematodes collected were fixed in AFA solution (a mixture of 70% Ethyl alcohol, Formalin, and Acetic acid in the ratio of

90:7:3) for 24 hours under slight cover glass pressure and then washed several times with 70% alcohol, stained with Mayer’s Carmalum, dehydrated in graded series of alcohols, cleared in clove oil and xylene and mounted permanently in Canada Balsam by usual method.

ix Trematodes were identified with the help of keys and descriptions are compared with the previously described species of various genera.

During present studies 18 trematode species were identified out of which

11 belong to the Family Hemiuridae, including 5 new species of the Lecithocladium Luhe, 1909; 4 new species of the genus

Tubulovesicula, Yamaguti, 1934; 1 species of the genus Stomachicola

Yamaguti, 1934 (Bilqees, 1971) and 1 species of the genus

Prosogonotrema Perez Vigueras, 1940 (Bilqees and Durrani, 1980).

Previously several species of hemiurids were also described from

Pakistan (Bilqees, 1981; Shaukat and Bilqees, 2006).

One new species each of the following genera have also been identified and described: Pleorchis Raillet, 1896; Decemtestis Ozaki, 1925

Stephanostomum Looss, 1899; Plagioporus Stafford, 1904 and

Prosorhynchus Odhner, 1905.

While two new species of the genus Bucephalus Baer, 1827 are also reported.

x As mentioned above during the present studies 18 trematodes species are identified including Pleorchis heterorchis n.sp. from the fish Lutjanus johnii and Otolithus argenteus; Decemtestis johnii n.sp. from the fish

Lutjanus johnii; Lecithocladium cybii n.sp. from the fish Cybium guttatum; Lecithocladium karachiensis n.sp. from the fish

Parastromateus niger; Lecithocladium magnasoma n.sp. from the fish

Stromateus sinensis; Lecithocladium magnavesicula n.sp. from the fish

Pomadasys olivaceum; Lecithocladium lateropharyngium n.sp. from the fish Stromateus sinensis; Tubulovesicula olivaceus n.sp. and

Stephanostomum gibsoni n.sp. from the fish Pomadasys olivaceum;

Tubulovesicula microcaudum n.sp. from the fish Otolithus argenteus;

Tubulovesicula magnacirrosa n.sp., Tubulovesicula karachiensis n.sp. and Prosogonotrema diacanthi Perez Vigueras, 1940 (Bilqees and

Durrani, 1980) from the fish Pseudosciaena diacanthus; Stomachicola muraenesocis Yamaguti, 1934 (Bilqees, 1971) from the fish Muraenesox cinereus; Plagioporus karachiensis n.sp from the fish Caranx affinis;

Bucephalus otolithi n.sp. from the fish Otolithus argenteus; Bucephalus mujibi n.sp from the fish Cybium guttatum and Prosorhynchus magnacirrus n.sp. from the fish Psettodes erumei.

xi The new species of various genera described here are distinguished from other species of the genera on the basis of differences in the diagnostic features. More emphasis is given on morphological variations and position of various organs, genital pore, sucker-width ratio and egg sizes rather than body sizes because it may differ with the age of the specimens as well as during fixing and processing. As observed during the present studies the rate of infection is high in carnivorous fishes probably because these fishes feed on other fishes and invertebrate which act as intermediate host.

This work will be continued as numerous fishes of Karachi coast are still uninvestigated for trematode parasites, the knowledge of all helminthes and protozoan parasites for fishes of Karachi Coast is limited and require more investigation especially the protozoan parasite.

xii INTRODUCTION

Pakistan is located at the gateway of the Persian Gulf. It is close to Iran,

Oman, India and the land-locked Central Asian countries. Pakistan is thus provided with a good opportunity to benefit from its geo-strategic position. However, the increasing pollution levels in the coastal regions and degradation of marine resources of the country are emerging as important environmental threats needing immediate actions.

Karachi is located at North-east of Arabian Sea in the Oriental region. It is surrounded by Palearctic region in the North, Arabian Sea in the South,

India in the East and Arabia in the West.

Karachi is well equipped not only in having a large Fish Harbour but also serves as a regular pathway for migratory fishes between the surrounding

Oceans (Indian Ocean and Pacific Ocean).

About one thousand species of fishes are found in the marine and freshwaters of Pakistan. Mostly edible, not only flesh of fishes but eggs and air bladder are also of good market value (Farooq & Sadiq, 1992).

1 The marine world extends from the ocean to the deepest bed and from the shore to distant shore. Fish is important for human life, hardly there is any country where the fish is not included in the human diet.

Fishes are zoonotically important, since several diseases are caused in human beings by fish parasites. Hundreds of fishes suffer presence of parasites in their bodies resulting as carrier to the human body. The most important human diseases from fish to human is Anisakiasis Van Thei et al., 1960 Several species of fish parasites have been identified as harmful

(Bilqees et al., 2003).

Fishes are the most numerous and diverse of the major vertebrate groups.

They dominate the waters of the world through a marvelous variety of morphological, physiological, behavioural patterns. Their diversity is reflected in the large number of living species. Few years ago compilation of fish species (Eschmeyer, 1998) lists 23,250 species with valid descriptions, the estimated number actually is around 25,000.

Because 200 new fish species are being described each year, the total may reach 30,000 or more (Joseph & Moyle, 2000).

2 Fish are valuable sources of high grade protein and organic products.

They occupy a significant position in the socio-economic status of the

South-Asian countries by providing the population not only the nutritious food but also income and employment opportunities (Joseph & Moyle,

2000).

The fish also as a taxonomic group, generated unlimited curiosity of the naturalists and zoogeographers from the period as early as that of the great Aristotle (Talwar and Jhigran, 2001).

By volume, 97% of all water is in the oceans and 0.0093% is in fresh water lakes and streams (Horn, 1972), 58% of the modern fish species are marine while 41% are freshwater inhabitants and 1% move on a regular basis between the two environments (Cohen, 1970).

More than half of the world’s population has an insufficient and nutritional unbalanced diet. An adequate diet requires an average of about

44g. per day, protein being about 10-12g. of the total caloric intake. As mentioned above, fish is an excellent source of protein, containing all the

10 essential amino acids in desirable concentrations for human beings and available at cheaper rates (Agrawal, 1999).

3

Fish is a renewable source of wealth. Unlike minerals and other fixed resources which do not have self-generating capacity to replenish the loss due to natural causes and human exploitation.

The importance of fishing in the economy of Pakistan, has by no means diminished, despite the rapid pace of industrialization in urban areas.

Fishing is the main source of livelihood for the people living in the coastal areas. It is estimated that over 200,000 people are engaged in fishing, its processing and marketing. This sector earns a foreign exchange worth Rs. 1,485 million ($ 90 million) or 3.4% of total export earnings (Syed, 1985). The territorial sea water of the coast of Sindh and

Baluchistan has the potential to provide sea food to as many as twice the present population of Pakistan.

The total sea coast of Pakistan is 682 miles, of which Baluchistan and

Sindh have 477 and 205 miles, respectively. The Baluchistan coast, which is about 70% of the total coast line, Mekran has 367 miles and

Lasbela district of Kalat division 110 miles. Although Pakistan is quite rich in fish resources, they have remained rather un-exploited (Syed,

1985).

4

At present, exporting fish and shrimps worth $ 90 million annually

(Syed, 1985). However, there is sufficient scope for fish export in future.

In a country like Pakistan where protein availability is scarce and now aquaculture is in progress to fill up this protein gap, it is essential for getting good quality of protein from fisheries resources and these should be free of diseases. Besides other factors optimum concentration of various elements in water for fish life needs to be investigated.

Even modest doses of sea foods, equivalent to one or two fish dishes a week, may be of some value for the prevention of coronary heart diseases and other parasitic infections (Fins fisheries news pak.agr.1985). It is known that it also reduces the tendency of blood to clot and helps lower cholesterol levels in the blood.

As mentioned above fish are the important group of vertebrates and have a great importance and significance in the life of mankind. These are most common and widely distributed almost in all parts of the world.

Fisheries is of great importance to human beings and in addition to providing food, most of the fishing industries yield a number of by-

5 products of commercial importance. The total present world catch of fishes and other aquatic organisms reaches the enormous figure of about

30 million metric tons. In various countries fish forms 17-83% of the protein intake of mankind. Fish play most important role in the economies of many countries. Nearly all freshwater and marine fishes are edible and have been an important source of protein. The white meat of fish, contains 16-29% of protein and has a food value of 300-1600 calories per pound. Important marine food fishes are Salmon, Cod,

Herring, Eels, Tuna, Mackerel and Sardines (Syed, 1985).

Fish in terms of good value rank and mostly superior having less amount of fat and more or equal protein as compared to other fleshes consumed as food by human being. As mentioned above one thousand species of fishes are found in marine and freshwaters of Pakistan. Majority of these are edible. Fishes can be divided into two major groups, cartilaginous and bony fishes. A good number of elasmobranch fishes are edible but only a few are consumed locally. “Mohr Mongra” is the most popular fish in this group, salted fishes of this category are exported to some countries like Sri-Lanka. Among the bony fishes, the flesh of the ‘Clupeid’ is good to eat, but full of fine bones. Imported Clupeids are

6 Shads, Herrings, Sparts, Palla is the most valuable fish in this group. Its flesh is oily and excellent in taste. Another economically valuable fish is the Cat fish.

Most fish species have at least some direct monetary value, are part of food webs that support economically important species, or live in environments whose good health is essential for the support of fisheries.

Furthermore, commercial and subsistence fisheries provide a good share of the protein needed by humans. Despite this high value, fish populations are in decline everywhere. It has been estimated that marine fisheries alone can yield, on a sustained basis, between 90 and 100 million metric tons of fish per year (Rounsefell, 1975). This figure reached for a brief period, but present catches are now around 80 million metric tons per year (including 15 million tons from freshwater) and the amount is declining (O’ Bannon, 1994). Major fisheries around the world, such as the Salmon fisheries of the Pacific North-West and the

Atlantic Cod fisheries have collapsed. Other declining economic value of wild-caught fish include fish collected for the aquarium trade and sport fisheries, both of which depend on large populations of healthy fish in clean water. In most cases the loss of fisheries and fish population is

7 predictable, and often it is blatantly taken in time despite the economic, social and ecological losses that are caused by parasites.

Currently the cultural values of fish include scientific research and educational values. The scientific values of fish are obvious to

Ichthyologists who find the study of fish essentially interesting.

The educational values of fish are also obvious. They are the one group of vertebrates in which it is really possible to maintain large populations in the laboratory for observation and experimentation. Many varieties are readily available for dissection, giving an easy overview of the several types of adaptations possible in vertebrates. Most important is that they can typically be found in large numbers and diversity in waters fairly close to most educational institutions, so that many types of ecological demonstrations can be made. Fish have the right combination of availability, size and ease of handling to make them the most desirable group of vertebrates for comparative classroom studies of ecology, behaviour, evolution, and anatomy, and to be very valuable in other areas such as genetic and physiology as well.

8 In August, 2004 the Marine Fisheries Department at the Wetland Center,

Karachi and the World Wide Fund for Nature, Pakistan, jointly hosted a workshop. The purpose of the event was to identify factors contributing to the country’s failure in achieving the required growth of fisheries, one factor could be parasitic infections. The experts discussed various aspects of the issue at the consultative workshop on 5-year plan for Fisheries

Sector. Stakeholders in fisheries sector, experts from the Federal and

Provincial Departments of Marine Fisheries, National Institute of

Oceanography, Baluchistan Coastal Department Authority, as well as academicians and representatives from various NGOs, and community- based organizations participated and provided their inputs.

Combination of all the factors described above, including mismanagement of marine resources and lack of enforcement of environmental laws, has initiated a process leading to an unsustainable situation. In addition, the communities of migratory fishermen residing on the islands are very poor and their only source of livelihood is subsistence fishing. They heavily depend on the mangroves for their fuel and fodder requirements. Hence the fisheries and the mangroves of the area are heavily over-exploited. Diversification of the livelihood base of

9 the local communities will be required for rehabilitation of the ecosystem.

Like all other animals fishes are also affected by various parasites and diseases. The internal helminth parasites of the fishes include trematodes, cestodes, nematodes and acanthocephala.

Marine parasitology is important in aquatic sciences. Fish parasites play a major role in marine biodiversity, infecting hosts at all different trophic levels. The growth of marine aquaculture, concerns about the effects of pollution on fish health, and the possible use of parasites as biological indicator has led to a steady increase interest in this topic.

More over parasites are an integral part of every ecosystem, representing a major factor in global biodiversity. Host parasite checklists suggest that on average, there are at least 3-4 metazoan parasites per studied marine fish species within a specific environment. This led to a conservative estimate by Klimpel et al. (2001), of 20,250 – 43,200 marine metazoan fish parasites, calculated on the basis of the 13,500 currently known fish species inhabiting brackish or marine waters. Fish parasites clearl

10 constitute a major part of the living species within the world’s oceans (http://www.marine parasitology.com/marine parasitology).

Besides infection from living parasites, pathogens that are already dead or their remains within fish tissues might harm the consumers by causing allergic reactions. Parasites can also significantly contribute to financial losses for fishery industry.

Parasites as disease causing agents can have various effects on the health of fish. They can infest all organs, depending on the site and intensity of infestation, they are more or less harmful. A high number of negative consequences does not only influence the fish itself but is assumed to also have a larger impact on whole ecosystems.

Fish parasites are important biological indicators to describe migration patterns of phylogenetic interactions as well as pollution and eutrophication. Within fisheries biology, fish parasites were successfully used to separate different fish stocks. Long living species such as endoparasitic helminths can give information on the seasonal migrations

11 of their hosts and migration habits of different age groups while short living species, combining a direct life cycle and high reproduction rates

such as protozoan, ectoparasites and monogenean trematodes can give information on environmental conditions. It can be expected that with an increasing knowledge of the species diversity, life cycle biology and ecological needs of various marine fish parasites, further information becomes available for their potential use as biological indicators. This tool will surely provide a better ecological understanding of the whole marine ecosystem (http://www.marineparasitology.com/marine parasitology).

Marine fish parasitology contributes to various scientific disciplines.

Zoological studies, such as those dealing with the , classification, morphology and phylogeny of marine fish parasites are still needed, if our attempts to provide a natural animal system including the fish parasites are to be successful. Besides utilizing classical methodologies such as morphometrical studies or using scanning and transmission electron microscopy, modern techniques in biochemistry and molecular biology offer a wide range of new applications to better

12 understand the phylogenetical characteristic of the different species. In addition, topics concerning the fish hosts with their parasites interactions

(host and site specificity, immunology, pathogenicity) have not yet been fully explored. Even the coelacanth Latimeria chalummnea Smith, 1939 is known to harbour several different marine fish parasite. Therefore, modern marine fish parasitology is firmly anchored within the different biological disciplines, and can offer various topics for integrated research activities (http://www.marine parasitology.com/marine parasitology).

Trematodes are one of the common parasites of fishes of Karachi coast.

Several species have been identified and described so far from 22 species of fishes (Bilqees, 1981; 1985; 2003; 2004 a-d Shaukat & Bilqees 2006).

A large number of fishes have not been investigated for trematode parasites. Therefore, this project is undertaken to collect, identify and report these important fish parasites in addition to those already known.

Thousands of trematode species have been described from various parts of the world. Trematodes belong to the Phylum Platyhelminthes. These flukes are leaf-like unsegmented flat worms. These vary in size from 1 mm to several centimeters in length. The most characteristic external structures are two suckers – one oral through which digestive tract opens

13 and the other ventral (acetabulum) for attachment. Each individual worm is hermaphrodite (monoecious) except the schistosomes which are unisexual. The body is covered with integument, which often bears spines. Body cavity is absent. Alimentary canal is present but incomplete.

It consists of a mouth surrounded by oral sucker, a muscular pharynx and the oesophagus which usually bifurcates in front of the ventral sucker into a pair of blind intestinal caeca. Anus is absent. Reproductive system is highly developed, with the exception of schistosomes, the reproductive organs are hermaphroditic. The genital organs lie between the two branches of the intestine. Trematodes are oviparous and lay eggs which, with the exception of schistosomes, are operculated.

The clinical picture of trematode infections depend upon size and number of worms present in the host and organs or tissues parasitized. The pathogenic lesions produced may be local or systemic, usually both. The former consists of ulceration, sloughing of tissue and abscess formation.

Systemic manifestations are usually due to absorption of antigenic by- products of the worms which frequently provoke a generalized leucocytosis, hyper-eosinophilia and allergic manifestations.

14 The digenetic trematodes, or flukes are among the most common and abundant of parasitic worms, second only to nematodes in their distribution. They are parasites of all classes of vertebrates, specially marine fishes and nearly every organ of the vertebrate body can be parasitized by some trematode as adult or juvenile. The adult parasite produces eggs and the eggs are passed in the definitive host’s faeces.

Several species cause economic losses to society through infections of domestic animals and others are medically important parasites of humans. Because of their importance Digenea have stimulated vast amounts of research and the literature of this group is immense.

It can be easily estimated that trematode species are more in number than all the animal species in whole animal kingdom including species in all the vertebrate and invertebrate phyla. Because each and every animal species, both vertebrates and invertebrate is infected with one or more species of trematodes. These can infect each and every organ of the animals and man. The organs involved are mostly intestine, stomach, liver, heart, lungs, blood, eyes, brain, gonads and kidneys, subcutaneous tissues. In fishes mostly adult trematodes are found in the intestine and stomach. Some are found in liver, bile ducts and other organs. Fish can act as definitive host, intermediate host, transport host, paratenic host or

15 reservoir host for various trematode species. Some zoonotic infections are also known in which animal trematode infect man and cause diseases.

Therefore, significance of fish trematodes can not be ignored specially because fish is a valuable food commodity. Only healthy fish can provide good quality of meat, protein and vitamins. It is equally good for healthy and sick persons. Mortality of fishes due to infections specially in young ones is a great economic loss. Researches on fish parasites specially in country like Pakistan is most essential for better knowledge of parasitic diseases but very little attention is being paid on this subject in our country.

As mentioned above trematodes are one of the important parasites infecting fishes throughout the world and these are also common in fishes of Karachi coast (Bilqees, 1981; Shaukat & Bilqees, 2005). Previously trematodes known from various fish species belonging to different families have been reported (Shaukat and Bilqees, 2005). Revised check- list including present species is also prepared and presented here.

During the present studies eighteen trematode species have been identified belonging to the genera Pleorchis Railliet, 1896; Decemtestis,

Ozaki 1925; Lecithocladium Luhe, 1901; Tubulovesicula Yamaguti,

16 1934; Stomachicola Yamaguti 1934 (Bilqees, 1971); Prosogonotrema

Perez Vigueras 1940 (Bilqees and Durrani, 1980); Plagioporus Stafford

1904; Stephanostomum Looss, 1899; Bucephalus Baer, 1827; and

Prosorhynchus Odhner, 1905. A total of 18 species have been identified and described here including 16 new species and 2 already known species. The species identified are Pleorchis heterorchis n.sp. from the fish Lutjanus johnii and Otolithus argenteus; Decemtestis johnii n.sp. from the fish Lutjanus johnii; Lecithocladium cybii n.sp. from the fish

Cybium guttatum; Lecithocladium karachiensis n.sp. from the fish

Parastromateus niger; Lecithocladium magnasoma n.sp. from the fish

Stromateus sinensis; Lecithocladium magnavesicula n.sp. from the fish

Pomadasys olivaceum; Lecithocladium lateropharyngium n.sp. from the fish Stromateus sinensis; Tubulovesicula olivaceus n.sp. and

Stephanostomum gibsoni n.sp. from the fish Pomadasys olivaceum;

Tubulovesicula microcaudum n.sp. from the fish Otolithus argenteus;

Tubulovesicula magnacirrosa n.sp., Tubulovesicula karachiensis n.sp. and Prosogonotrema diacanthi Perez Vigueras 1940 (Bilqees and

Durrani, 1980) from the fish Pseudosciaena diacanthus; Stomachicola muraenesocis Yamaguti, 1934 from the fish Muraenesox cinereus;

Plagioporus karachiensis n.sp. from the fish Caranx affinis; Bucephalus otolithi n.sp. from the fish Otolithus argenteus; Bucephalus mujibi n.sp

17 from the fish Cybium guttatum and Prosorhynchus magnacirrus n.sp from the fish Psettodes erumei. All these are figured and compared with the previous species of their respective genera.

This project will be continued to identify more trematodes of fishes of

Karachi coast.

18 REVIEW OF LITERATURE

Trematodes are one of the common parasites in fishes of Karachi coast.

Previously species of the genera reported from Pakistan include:

Acanthocolpus Luhe, 1906; Pseudacaenodera Yamaguti, 1970;

Tormopsolus Poche, 1926; Stephanostomum Looss, 1899;

Acanthostomum Looss, 1899; Karachitrema Bilqees, 1973;

Parapolylekithum Bilqees et al., 2004; Alcicornis MacCallum, 1917;

Bucephalus Baer, 1827; Telorhynchus Crowcroft, 1947; Bucephalopsis

(Dies, 1855); Prosorhynchus Odhner, 1905; Callodistomum Odhner,

1902; Mehrailla Srivastava, 1939; Paradiscogaster Yamaguti, 1934;

Pseudohypertrema Bilqees, 1976; Diplobulbus Yamaguti, 1934;

Paradiplobulbus Bilqees, 1972; Tergestia Stossich, 1899; Laruea

Srivastava, 1937; Ectenurus Looss, 1907; Erilepturus Woolcock, 1935;

Lecithocladium Luhe, 1901; Magnacetabulum Yamaguti, 1934;

Mecoderus Manter, 1940; Tubulovesicula Yamaguti, 1934;

Uterovesiculurus Skrjabin et Guschanskaja, 1954; Hysterolecitha

Linton, 1910; Qadriana Bilqees, 1971; Lecithochirium Luhe, 1901;

Prosorchis Yamaguti, 1934; Sterrhurus Looss, 1907; Stromaturus

Bilqees and Khatoon, 2003; Acerointestinecola Jahan, 1970;

Allostomachicola Srivastava, 1939; Allostomachicola Yamaguti, 1958; 19 Cameronia Bilqees, 1971; Cestodera Bilqees, 1971; Segmentatum

Bilqees, 1971; Stomachicola Yamaguti, 1934; Trifoliovarium Yamaguti,

1940; Crassicutis Manter, 1936; Lampritrema Yamaguti, 1971;

Aephnidiogenes Nicoll, 1915; Anterodiscus Bilqees, 1974; Bianium

Stunkard, 1930; Bicaudum Bilqees, 1971; Multiovarium Bilqees, 1974;

Orientodiploproctodaeum Bhutta and Khan, 1970; Lepidapedon

Stafford, 1900; Tritesticulum Bilqees, 1971; Notoporus Yamaguti, 1938;

Pseudolepidapedon Yamaguti, 1938; Complexobursa Oshmarin et

Mamaev, 1963; Buckleytrema Gupta, 1956; Monodhelmis Dollfus,

1937; Proctotrematoides Yamaguti, 1938; Pseudohurleytrema

Yamaguti, 1954; Monorchis (Monticelli, 1893) Looss, 1902; Enenterum

Linton, 1910; Neoenenterum Bilqees and Khatoon, 2004;

Olivacreadium Bilqees, 1976; Pseudocoitocaecum Bilqees, 1972;

Allopodocotyle Pritchard, 1966; Hamacreadium Linton, 1910;

Helicometrina Linton, 1910; Plagioporus Stafford, 1904; Podocotyle

(Dujardin, 1845); Podocotyloides Yamaguti, 1934; Thynotrema Bilqees et al., 2007; Thynstenopera Bilqees and Khatoon, 2004;

Pseudoplagioporous Yamaguti, 1938; Opistholebes Nicoll, 1915;

Pleorchis Railliet, 1896; Prosogonotrema Perez Vigueras, 1940;

Neodiptherostomum Bilqees et al., 2003 and Waretrema Srivastava,

1937 (Table 1&2). These trematodes show great biodiversity (Attached).

20 Here only those genera are reviewed which have been identified during the present studies including Pleorchis Railliet, 1896; Decemtestis

Ozaki, 1925; Lecithocladium Luhe, 1901; Tubulovesicula Yamaguti,

1934; Stomachicola Yamaguti, 1934 (Bilqees, 1971); Stephanostomum

Looss, 1899; Bucephalus Baer, 1827; Prosorhynchus Odhner, 1905;

Plagioporus Stafford, 1904 and Prosogonotrema Perez Vigueras, 1940

(Bilqees and Durrani, 1980).

Family: Pleorchiidae (Poche, 1926) Yamaguti, 1971 GENUS PLEORCHIS Railliet, 1896

The genus Pleorchis syn. Polyorchis (Stossich, 1888) nec. Polyorchis

Agassiz, 1862 was proposed by Railliet, 1896 to accommodate the type- species. Pleorchis polyorchis (Stossich, 1888) Stiles, 1896 from the fish

Sciaena sp. Later on several species have been added to the genus

(Yamaguti, 1971; Bartoli et al., 2004; Bilqees, 1977; Madhavi, 1985;

Shen, 1983; Gupta and Gupta, 1978). Recently the type-species

Pleorchis polyorchis has been redescribed from the fish Sciaena sp. from

Western Mediterranean Sea (Bartoli et al., 2001). Here a new species of this genus P. heterorchis is described from the fish Lutjanus johnii and

Otolithus argenteus. As mentioned above, previously only one species of

21 the genus Pleorchis ghanensis Fischthal & Thomas, 1968 (Bilqees,

1981) has been reported from Pakistan.

Family: Opecoeliidae Ozaki, 1925. GENUS DECEMTESTIS Ozaki, 1925

The genus Decemtestis was established by Yamaguti, 1934 with type- species D. sillagonis and other two species D. callionymi and D. ditrematis which have ten testes and non-filamentous eggs with or without polar prolongation. Yamaguti, 1934 also transferred

Helicometrina azumae Layman, 1930 to Decemtestis on the basis of non-filamented eggs and position of genital pore different from that in

Helicometrina Linton, 1910. Manter, 1933 placed H. azumae under the genus Rhagorchis Manter, 1931 firstly, because of its non-filamented eggs, and secondly, because of its morphological similarity to

Rhagorchis odhneri Manter, 1931 and its occurrence in a related host.

Srivastava, 1936 added three more species to Decemtestis: D. brevicirrus, D. mehrai and D. biacetabulata, all from fishes of the Bay of Bengal. The last named species has two concentric acetabula, one enclosed within the other. Accordingly, Srivastava, 1936 gave an emended diagnosis of Decemtestis. Later more species were added by

Yamaguti (1938, 1951, 1959); Park (1939) and Manter (1954).

22 The genus Decemtestis Yamaguti, 1934 contains relatively few species.

Most of the species are reported from Japan (Yamaguti, 1934, 1938,

1951, 1971) few from India and other regions (Srivastava, 1936; Park,

1939; Manter, 1954). But there is no report of genus from fishes of

Karachi coast, Pakistan. Present is the first report of a new species of the genus Decemtestis johnii from the fish Lutjanus johnii (Lutianiidae) of

Karachi coast. The species name refers to the fish species.

The genus Decemtestis was proposed by Yamaguti, 1934 for D. sillagonis Yamaguti, 1934 from a sillaginid fish in Japan and now contains about 25 nominal species parasitic in marine fishes. It is easily recognized by the combination of 9 or 10 testes and eggs without filaments. It is presumably close to Helicometrina, from which it differs only in the absence of filaments on the eggs (although eggs of D. sillagonis are described as having polar prolongations, most species are described as being without any protuberance) and in having a sinistral rather than median genital pore. Allodecemtestis Hafeezullah, 1970 was proposed by Hafeezullah, 1970a for Decemtestis biacetabulatus

Srivastava, 1936 and D. pseudolabri Manter, 1954; trematodes related to

Decemtestis in which there were ‘two ventral suckers’, ‘one concentrically enclosed within other’. Three further species have been

23 proposed: A. bilqeesae Ahmad, 1990; A. skrjabini Ahmad, 1988 and A. odeningi Ahmad, 1988. The structure of the concentric ventral suckers has never been explained adequately, but the interpretation of the presence of two suckers is not plausible. These species are all considered species of Decemtestis, requiring the new combinations D. bilqeesae

Ahmad, 1990 n. comb.; D. odeningi Ahmad, 1988 n. comb. and D. skrjabini Ahmad, 1988 n. comb.

Family: Hemiuridae Looss, 1899, Luhe, 1901

GENUS LECITHOCLADIUM Luhe, 1901

Lecithocladium Luhe, 1901 is one of the largest genera of digenea belonging to the family Hemiuridae. There are at least 83 nominal species as mentioned by Bray and Cribb, 2004 or may be more (including those in the genera considered synonymous by Gibson, 2002). Most of the species are characterized by an infundibuliform oral sucker, an elongated, cylindrical pharynx and a large fusiform seminal vesicle (Bray and Cribb, 2004). Previously several species of the genus have been described from fishes of Karachi coast (Bilqees, 1981; Bilqees et al.,

2005; Farooq & Khanum, 1980; Zaidi & Khan, 1977).

24 Lecithocladium Luhe, 1901 contains a large number of species described from various parts of the world including Pakistan. (Yamaguti, 1971;

Bilqees et al., 2005; Bray & Cribb, 2004; Gibson, 2002; Gupta & Gupta,

1983; Al-Yamani & Nanhas, 1981; Gupta & Singh, 1983; Srivastava,

1942; Gupta & Puri, 1981). 13 species of the genus are known from fishes of Pakistan (Bilqees, 1981, Farooq & Khanum, 1980). In Pakistan most of the Lecithocladium species described are from fishes of family

Stromateidae (Bilqees et al., 2005, Farooq & Khanum, 1980).

The genus Lecithocladium Luhe, 1901 was proposed by Yamaguti, 1971 with type-species L. excisum (Rudolphi, 1819) Luhe, 1901 from

Scomber scombrus. Other species of the genus Lecithocladium Luhe,

1901 reported from different parts of the world are: L. aegyptense

Fischthal et Kuntz, 1963 from Pomadasys olivaceum and Monodactylus argentatus; L. angustiovum Yamaguti, 1953 from Scomber kanagunta,

Rastrelliger chrysozonus and Caranx affinis; L. annulatum Chauhan,

1945 from Stromateus cinereus; L. apolecti Velasquez, 1962 from

Apolectus niger; L. brevicauda Srivastava, 1942 from Chrysophrys bifasciata; L. bulbolabrum Reid, Coil et Kuntz, 1966 from Rastrelliger kanagunta; L. carultum Chauhan, 1945 from Sciaena carulta and

Harpodon nehereus; L. chingi Manter et Pritchard, 1960 from

25 Acanthurus mata, Naso sp. and Melichthys vidua; L. crenatum (Molin,

1859) Looss, 1907 from Centrolophus pompilius syn. of L. excisum

(Dawes, 1946), Nahhas and Short, 1965. Also from Arnoglossus bosci and Peprilus poronotus; L. cristatum (Rudolphi, 1819) Looss, 1907 from

Dactylopterus volitans; Stromateus fiatola syn. of L. excisum Dawes,

1946; L. excisiforme Cohn, 1902 from Scomber scomber; L. glandulum

Chauhan, 1945 from Lutjanus johnii and Mugil speigleri; L. gulosum

(Linton, 1901) Looss, 1907 from Porober scombrus syn. of L. excisum

Nahhas and Short, 1965; L. harpodontis Srivastava, 1942 from

Chrysophrys datnia and Scomber colias; L. magnacetabulum Yamaguti,

1934 from Psenopsis anomala; L. megalaspis Yamaguti, 1953 from

Megalaspis sp. and Megalaspis cordyla; L. pagrosomi Yamaguti, 1934 from Pagrosomus major; L. pampi Lebedev, 1968 from Pampus argenteus; L. parviovum Yamaguti, 1953 from Scomber kanagunta; L. psenopsis Yamaguti, 1934 from Psenopsis anomala; L. scomberi

Yamaguti, 1953 from Scomber kanagunta, Scomber microlepidotus and

Scomber brachypomus; L. seriolellae Manter, 1954 from Seriolella brama and Cyttus australis; L. gazzai Pandey et al., 2000 from Gazza minuta and L. bengalensis Saxena and Prakash, 2006 reported from

Gazza minuta.

26 Thirteen species of the genus Lecithocladium Luhe, 1901 reported so far from fishes of Pakistan include: L. psenopsis Yamaguti, 1934 from

Stromateus sinensis; L. hexavitellarii Bilqees, 1971; L. anteporus

Bilqees, 1971; L. microductus (Bilqees, 1971); L. arabiana, Bilqees,

1971; L. microcaudum Bilqees, 1971; L. octovitellarii Bilqees, 1971 all from Stromateus sp., of Karachi coast; L. karachii Zaidi and Khan, 1977 from Carangoides presustus; L. pakistanensis Zaidi and Khan, 1977 from Carangoides malabaricus; L. stromatei Farooq and Khanum, 1980 from Stromateus niger; L. arabicum Farooq and Khanum, 1980 from

Caranx djedaba; L. thynensis Bilqees and Nighat, 1985 from Thynnus sp. and L. arii Bilqees et al., 2005 from the fish Arius serratus. These species are also from fishes of Karachi coast. All species except L. karachii, L. arabicum and L. thynensis are from fishes of family

Stromateidae. All these species are from fishes of Karachi coast.

Numerous species are also known from other parts of the world

(Yamaguti, 1971; Bray & Cribb, 2004; Gibson, 2002; Gupta and Gupta,

1983; Al-Yamani and Nahhas, 1981; Gupta and Singh, 1983; Srivastava,

1942; Gupta and Puri, 1981). During the present studies five new species of the genus Lecithocladium; L. cybii, L. karachiensis, L. magnasoma,

L. magnavesicula and L. lateropharyngium have been identified.

27 Originally Bilqees, 1972 described the genus Magnapharyngium in family Hemiuridae having large pharynx. Later, she synonymized this genus with Lecithocladium Gibson and Bray, 2004 also considered

Magnapharyngium as synonym of Lecithocladium.

Family: Hemiuridae Looss, 1899

GENUS TUBULOVESICULA Yamaguti, 1934

The genus Tubulovesicula Yamaguti, 1934 was described with type- species T. spari Yamaguti, 1934 from Sparus macrocephalus and

Pagrosomus unicolor syn. of T. lindbergi (Sogandares, 1959) Zhukov,

1960. Other species of the genus Tubulovesicula reported from different parts of the world are: T. lindbergi (Layman, 1930) Yamaguti, 1934;

Siddiqi and Cable, 1960 from Leptocephalus conger, Synodus sp. and unidentified eel; T. madurensis Nigrelli, 1940 syn. of T. lindbergi

Manter, 1954 from Scorpaena madurensi; T. magnacetabulum

Yamaguti, 1939 from Epinephelus akaara; T. marsupialia Oshmarin,

1965 from Saurida tumbil; T. muraenesocis Yamaguti, 1934 from

Muraenesox cinereus, Trachinocephalus myops, Lophius litulon,

Inimicus japonicus, Conger myriaster, Hamazima, Muraenesox coniceps;

T. muraenesocis from Trachinocephalus; T. nanamoensis Mc Farlane,

1936; Manter, 1947 from Scorpaenichthys marmoratus; T. pinguis

28 (Linton, 1940; Manter, 1947 syn Distomum grandiporum Rud. of

Linton, 1898; D. rufoviride Rud. of Linton, 1898; D. tornatum Rud. of

Linton, 1905 from Anguilla rostrata, Cynoscion regalis, Hemitripterus americanus, Merluccinus bilinearis, Menidia notata, Menticirrhus saxatilis, Opsanus tau, Paralichthys dentatus, Merlinus carolinus,

Sphyraena borealis and Synodus foetens; T. californica Park, 1936 from

Enophryns bison; T. diacopae Nagaty et Abdel-Aal, 1962 from Diacope sp.; T. hebrae Nagaty et Abdel-Aal, 1962 from Diacope sp.; T. pseudorhombi Yamaguti, 1938 from Pseudorhombus pentophthalmus and T. serrani Nagaty, 1956 from Serranus sp.

Members of the family Hemiuridae Looss, 1899 are among the most frequently encountered digeneans in teleosts. A variety of genera including Tubulovesicula have been described from various parts of the world including Pakistan (Bilqees and Nighat, 1981; Bhutta and Khan,

1975; Linton, 1898, 1905, 1940; Nagaty, 1956; Nagaty et Abdel-Aal,

1962; Park, 1936; Siddiqi and Cable, 1960; (Nicoll, 1914) Yamaguti,

1934, 1938, 1939; Zaidi and Khan, 1977).

Species of the genus Tubulovesicula Yamaguti, 1934 reported previously from fishes of Karachi coast, Pakistan are T. spari Yamaguti,

29 1934 (Bilqees, 1981; Zaidi and Khan, 1977) from Muraenesox cinereus

(Forsk) (Congridae); T. anguillae Yamaguti, 1934 (Zaidi and Khan,

1977) from Harpodon nehereus (Har) (Synodidae); T. magna Bilqees and Nighat, 1981 from Pomadasys olivaceum (Day) (Pomadasyidae) and

T. anguisticauda Nicoll, 1914; Yamaguti, 1934; (Bhutta and Khan,

1975; Bilqees, 1981) from Muraenesox cinereus (Forsk);

(Muraenesocidae). During the present studies four new species

Tubulovesicula olivaceus, T. microcaudum, T. magnacirrosa and T. karachiensis have been identified.

Faimly: Hemiuridae Looss, 1899

GENUS STOMACHICOLA Yamaguti, 1934

Type-species: S. muraenesocis Yamaguti, 1934 syn. Lecithocladium longicaudum Shen Tseng, 1935 in Muraenesox cinereus. Species reported from different parts of the world are: S. magnus (Manter, 1931)

Manter, 1947 syn. Dinurus magnus Manter, 1947; Pseudostomachicola magnus (Manter, 1947) Skrjabin et Guschanskaja, 1954 in Cynoscion nebulosus; S. rubeus (Linton, 1910) Manter, 1947 syn. Dinurus rubeus

Linton, 1910 in Lycodontis moringa, L. funebris, Gymnothorax vicinus and Synodus foetens and S. pritchardae Pandey et al., 2000 in

Acanthurus triostegus.

30 Jahan, 1970 established genus Acerointestinecola with the type-species

A. karachiensis. Bilqees, 1971 described three genera namely, 1.

Cameronia: type-species C. octovitellarii and C. pakistani; 2.

Segmentatum: type-species S. karachiensis and S. gadrii; S. cimerensis;

S. magnaesophagustum; 3. Cestodera: type-species C. gastrocecus and other species C. unicecus. Sinclair et al., 1972 synonymized S. magnum with S. rubea. Gupta and Sharma, 1973 established the genus

Indostomachicola with species I. kinnei. Gibson and Bray, 1979 synonymized Acerointestinecola and Indostomachicola with

Stomachicola.

Hafeezullah, 1980 suggested species described under the genus

Cameronia, Segmentatum and Cestodera all are confirmed to original description of Stomachicola muraenesocis. Therefore, he presumptly suppressed Cameronia, Segmentatum and Cestodera with

Stomachicola. Gupta and Singh, 1985 transferred S. lepturusi and S. rauschi to Allostomachicola Yamaguti, 1958. Though, Verma, 1973 and

Gupta and Ahmad, 1978 were against the validity of Allostomachicola and considered it congeneric to Stomachicola.

31 Biqees, 1971 reported separate male and female genital openings and 8 vitellaria for Cameronia octovitellarii. The hermaphroditic pouch and duct are very small and are not observed clearly. This information is based on the material studied by Gupta and Gupta, 1991. Bhalerao, 1971 considered the number of vitellaria as a variable character. Bilqees, 1971 has also mentioned that in trematodes from Muraenesox cinereus the number of vitelline tubules vary from 8-10. Hafeezullah and Gupta also did not considered number of vitelline tubules as generic character.

Bilqees, 1971 reported Segmentatum karachiensis with posteriorly fused caeca and S. magnaesophagustum with post-acetabular union of caeca and its condition in the middle of tail in addition to posterior union. The same condition is observed by Gupta and Gupta, 1991 in their specimens.

When examined the section of that region, it was clear that two limbs overlapping each other instead of fused. Sometimes these overlapped in such a fashion that they appeared one. In addition to the united type of caeca S. magnaesophagustum also have ventral cuticular expansion in the pre-acetabular region, oral sucker with cuticular spines and cuticular expansion at the acetabular region. Cestodera unicecus Bilqees, 1971 has a transverse branch between two caeca just above the ovary. In the opinion of Gupta and Gupta, 1991 this result is due to union of deep loop

32 of caeca. This information is true because if the loop of right caeca above the ovary in Cameronia pakistani Bilqees, 1971 moved more forward, the same condition will also appear for this species as in C. unicecus.

Absence of pre-oral lobe and union or non-union of excretory arms in pharyngo-oral region seem to be false microscopical observations due to highly motile anterior region, Gibson and Bray, 1979 suggested that uterine seminal receptacle is the commonest form of seminal receptacle present in Hemiuroidea.

Family: Prosogonotrematidae Perez Vigueras, 1940

GENUS PROSOGONOTREMA Perez Vigueras, 1940

The genus Prosogonotrema Perez Vigueras, 1940 was proposed by

Yamaguti, 1971 with type-species P. bilabiatum Perez Vigueras, 1940 in

Ocyurus chrysurus from Cuba and Jamaica. Species of the genus

Prosogonotrema diacanthi reported from different parts of the world are:

P. abalisti Parukhin, 1964 in Abalistes stellaris from Gulf of Tonkin; P. carangis Velasquez, 1961 in Caranx sp. from Phillipines; P. clupeae

Yamaguti, 1952 in Clupea (Amblygaster) clupeoides syn. of P. labiatum

Manter, 1969 from Celebes; P. subequilatum Pritchard, 1963 in Naso unicornis, N. hexacanthus from Hawaii and P. symmetricum Oshmarin,

1965 in Pristipomoides microlepis, P. sieboldii from Hawaii.

33 Two species reported from Karachi, Pakistan are Prosogonotrema diacanthi Bilqees and Durrani, 1980 from Pseudosciaena diacanthus and

P. karachiense Bilqees and Durrani, 1980 from Lutjanus johnii.

During the present studies P. diacanthi has been identified from the fish

Pseudosciaena diacanthus of Karachi coast. This species was originally described by Bilqees and Durrani, 1980 from the same fish and locality.

Family: Opecoeliidae Ozaki, 1925

GENUS PLAGIOPORUS Stafford, 1904

Species of the genus Plagioporus occurs in both marine and freshwater teleosts throughout the world. Species reported from different parts of the world are: Plagioporus acanthogobii Yamaguti, 1951 in Acanthogobius hasta and Ostracion tuberculatum; P. acerinae (Pigulewskii, 1931)

Price, 1934 syn. Lebouria acerinae Price, 1934 in Acerina cernua; P. alacer (Looss, 1901) Price, 1934 syn. Lebouria acerinae (Looss, 1901) in Labrus maculatus, L. merula, Crenilabrus pavo, C. quinquemaculatus,

C. griseus and Callionymus lyra; P. angusticollis (Hausmann, 1896)

Dobrovolny, 1939 in Cottus gobio and Salmogairdneri; P. apogonichthydis Yamaguti, 1938 in Apogonichthys carinatus; P. beringi

Mamaev, 1965 in Coryphaenoides sp.; P. branchiostegi Yamaguti, 1937

34 in Branchiostegus japonicus; P. calotomi (Yamaguti, 1934) in

Calotomus japonicus; P. choerodontis (Yamaguti, 1934) Yamaguti, 1938 syn. Lebouria choerodontis in Choerodon azurio; P. congeri Yamaguti,

1970 in Conger sp.; P. cooperi (Hunter et Bangham, 1932) Price, 1934, syn. Lebouria cooperi Hunter et Bangham, 1932 in Notropis,

Rhinichthys, Erinemus, Opsopoeodus, Ericymba, Ammocrypta and

Rheocrypta; P. crassigula (Linton, 1910) Price, 1934 syn. Lebouria crassigula Linton, 1910 in Calamus calamus, C. bayonado, Decodon puellaris, Diplodus holbrookii; P. dactylopagri Manter, 1954 in

Dactylopagrus macropterus; P. diacopae (Nagaty et Abdel Aal, 1962) n. comb. syn. Hamacreadium diacopae Nagaty et Abdel Aal, 1962 in

Diacope (=Lutjanus) fulviflamma and Lethrinus sp.; P. gerridis Fischthal et Thomas, 1970 in nigri; P. glomeratus Roitman, 1963 in

Rhodeus sericeus, Acanthorhodeus asmussi, Phoxinus ozekanowskii, P. lagowski, Brachymystax lenok, Thymallus arcticus grubei; P. hypophthalmichthys Akhmerov, 1959 in Hypophthalmichthys molitrix;

P. idoneus (Nicoll, 1909) Price, 1934 syn. Lebouria idoneus Nicoll,

1909 in Anarhichas lupus and Callionymus lyra; P. indicus in Lutjanus fulviflamma; P. interruptus Manter, 1954 in Pseudolabrus coccineus; P. ira Yamaguti, 1940 in Choerodon azurio; P. isaitschikowi (Layman,

1930) Price, 1934 syn. Lebouria isaitschikowi Layman, 1930 in

35 Sebastodes schlegelii, Sebasticus albofasciatus and Lethrinus microdon;

P. japonicus Yamaguti, 1938 in Neopercis sexfasciata, N. multifasciata,

N. aurantiaca, N. muronis, Plotosus anguillaris and Lethrinus microdon;

P. krusadaiensis (Gupta, 1956) n. comb., syn. Hamacreadium krusadaiensis Gupta, 1956 in marine cat fish; P. kyusen Yamaguti, 1959 in Halichoeres poecilopterus; P. lobatus (Yamaguti, 1934) Yamaguti,

1938 in Chelidonichthys kumu; P. longicirratus Manter, 1963 in trigger fish (thum); P. longisacculus Yamaguti, 1970 in Pristipomoides microlepis; P. longisaccus Fischthal et Kuntz, 1964 in Choerodon anchorago; P. longivesicula Yamaguti, 1952 syn. of Hamacreadium interruptum Nagaty, 1941- Fischthal and Kuntz, 1965 in Lethrinus sp.;

P. macassarensis Yamaguti, 1952 in Lethrinus sp.; P. macrouterinus

Haderlie, 1953 in Ptychocheilus grandis; P. maorum Allison, 1966 in

Octopus maorum; P. mormyri (Stossich, 1885) n. comb., syn. Distomum mormyri Stossich, 1885 in Pagellus mormyrus; P. multilobatus

Travassos, Freitas et Buhrnheim, 1966 in Haemulon sp.; P. myoxocephalis Akhmerov, 1960 in Myoxocephalus platycephalus; P. niloticus Vercammen-Grandjean, 1960 in Tilapia nilotica; P. obductus

(Nicoll, 1909) Price, 1934 in Bairdiella chrysura; P. obovatus (Molin,

1859) n. comb. syn. Allocreadium obovatus (Molin, 1859) Stossich,

1901 in Chrysophrys aurata; P. occidentalis Szidat, 1944 in Gobio

36 fluviatilis; P. pacificus Yamaguti, 1938 in Neopercis multi-fasciata, N. aurantiaca and N. muronis; P. parvus Manter, 1963 in Lethrinus variegates; P. pennelli (Leiper et Atkinson, 1914) Byrd, 1963, syn.

Podocotyle pennelli Leiper et Atkinson, 1914 in Trematomus bernachii and Notothenia neglecta; P. polymixiae Yamaguti, 1970 in Polymixia japonica; P. pratti Wu, 1933 in Nocomis micropogon; P. preporatus

Manter, 1954 in Chelidonichthys kumu; P. rooseveltiae Yamaguti, 1970 in Rooseveltia brighani; P. siliculus Sinitzin, 1931 in Salmo clarki,

Goniobasis plicifera silicula and Potamobius sp.; P. sillagonis

Yamaguti, 1938 in Sillago sihama; P. stefanski Slusarski, 1958 in

Salomo trutta m. fario; P. tohei Yamaguti, 1970 in Conger sp.; P. truncatus (Linton, 1940) Yamaguti, 1954 in Cynoscion regalis,

Menticirrhus saxatilis, Morone Americana and Synodus foetens; P. ula- ula Yamaguti, 1970 in Etelis marshi and P. varitus (Nicoll, 1910) Price,

1934 syn. of P. alacris (Looss, 1901) Manter, 1954 in Callionymus lyra.

Plagioporus was proposed by Stafford, 1904 for P. serotinus Stafford,

1904 from a Canadian freshwater fish. At least 100 species have been placed in this genus, many of which have subsequently been moved to other genera. At least nine genera have received species first placed in

Plagioporus. This is because Plagioporus and its near relatives embody the basic characteristics of plagioporines and the differences that can be

37 used to separate them are relatively hard to detect. The modern concept of the genus was refined, in particular by Gibson and Bray, 1982, who restricted it to species from freshwater fishes, in which the excretory vesicle extends only to the posterior testis. Gibson and Bray, 1982 proposed Macricaria for marine species in which the excretory vesicle is more anteriorly extensive. Macricaria has received at least 15 species described originally in Plagioporus. Other significant recipients of species of Plagioporus are Allopodocotyle (two species),

Gaevskajatrema Gibson and Bray, 1982 (two species), Neolebouria

(three species) and Podocotyle (two species). Plagioporus may still have the largest number of species of any opecoelid genus but many other new combinations await proposal, so that the final number of species in this, the type-genus for sub-family, may ultimately be quite small. Its concept is here expanded to include certain species in which the vitelline follicles are restricted to the hindbody; these are presently incorporated in

Allopodocotyle and are discussed under that genus. Baikalotrema

Layman, 1951 was established by Layman, 1951 for Allocreadium polymorpha Layman, 1933. The species and the genus have not been considered in the literature. Yamaguti, 1971 moved the species to

Podocotyle but did not propose that the genus was a synonym of

Podocotyle. The original description shows clearly that the ovary is

38 entire. The morphology is thus consistent with either Plagioporus or

Allopodocotyle, depending on the length of the excretory vesicle, which is not known. Because the species is from a freshwater fish, it is here combined, tentatively, with Plagioporus as P. polymorpha Layman,

1933 n. comb., and Baikalotrema is considered a tentative synonym of

Plagioporus (Bray and Gibson, 2005).

Two species of the genus Plagioporus found in Pakistan in marine fishes are Plagioporus (caudotestis) mujibi Bilqees, 1972 reported from

Crenidens indicus and Plagioporus heterorchis Bilqees, 1977 from the fish Pomadasys olivaceum. One species is also known from freshwater.

During present studies a new species Plagioporus karachiensis from the fish Caranx affinis has been identified.

Family: Acanthocolpidae Luhe, 1906

GENUS STEPHANOSTOMUM Looss, 1899

Species of Stephanostomum Looss, 1899 are parasites of marine fishes and are present throughout the world (Yamaguti, 1970, 1971; Bilqees,

1981; Bartoli and Bray, 2001, 2004; Bray and Cribb, 2003).

Stephanostomum is a genus consisting of species possessing, in

39 particular a ‘crown’ of spines around the oral opening. It is included in the family Acanthocolpidae Luhe, 1906 which was originally characterized by spined female terminal genitalia. This genus is one of the largest in number of described species having more than 112 nominal species (Saoud et al., 2002; Bray and Cribb, 2003).

The genus Stephanostomum Looss, 1899 was proposed by Yamaguti,

1971 with type-species S. cesticillus (Molin, 1858) Looss, 1899 from

Lophius piscatorius. Other species reported from different parts of the world are: S. dentatum Stunkard, 1961; S. anisotremi Manter, 1940 from

Anisotremus scapularis; S. aulostomi Nahhas et Cable, 1964 from

Aulostomus maculatus; S. australe Manter, 1954 from Chelidonichthys kumu; S. baccatum (Nicoll, 1907) Manter, 1934 from Hippoglossus vulgaris and Cottus scorpius; S. californicum Manter et Van Cleave,

1951 from Umbrina roncador; S. carangis (Yamaguti, 1951), Caballero,

1952 from Caranx equula; S. coryphaenae Manter, 1947 from

Coryphaena hippurus; S. edentulum Linton, 1910 from Neomaenis griscus; S. elongatum (Park, 1939) Hanson, 1950 from Areliscus joyneri;

S. hispidum (Yamaguti, 1934) Manter, 1940 from Seriola sp.; S. lineatum Manter, 1934 from Urophycis regis; S. indicum Srivastava,

1937 from Pristis cuspidatus; S. longisomum Manter, 1940 from Caranx

40 hippos; S. megacephalum Manter, 1940 from Caranx hippos; S. megacephalum Perez Vigueras, 1955 from Elagatis bipinnulatus; S. microstephanum Manter, 1934 from Epinephelus niveatus; S. naucrotis

Nagaty, 1957 from Naucratus sp.; S. numi Yamaguti, 1970 from

Aulostomus chinensis; S. pacificum Yamaguti, 1951 from Caranx equula; S. platacis Nagaty, 1957 from Platax sp.; S. pristis

Deslongchamps in Lamoureux, 1824 from Gadus euxinus, G. minutus, G. melangus and G. callarias; S. provitellosum Sogandares-Bernal, 1959 from Balistes naufragium; S. pseudocarangis Sogandares-Bernal, 1959 from Holocentrus ascensionis; S. robustum (Mac Callum, 1917),

Caballero, 1952 from Conger conger; S. sentum (Linton, 1910), Manter,

1940 syn. of S. minutum Caballero, 1952 from Calamus calamus; S. sobrinum (Levinsen, 1881) Looss, 1899 from Cottus scorpius; S. solontschenki Parukhin, 1968 from Merluccius merluccius; S. triglae

(Lebour, 1908) Dawes, 1946 from Trigla gurnadus; S. tristephanum Mc

Farlane, 1936 from Ophiodon elongatum; S. adinterruptum

Hafeezullah, 1971 from Fistularia villosa; S. egypticum Ramadan, 1983 from Lethrinus mahsena; S. fistulariae Yamaguti, 1940 from Fistularia petimba; S. casum Siddiqi and Cable, 1960 from Lutjanus argentimaculatus; S. microsomum Madhavi, 1976 from Rachycentron canadum; S. pseudoditrematis Madhavi, 1976 from Rachycentron

41 canadum; S. sphyraenae Wang, 1982 from Sphyraena pinguis; S. lalandi

(as S. dorsalis Manter, 1940; S. mazatlana Sogandares-Bernal, 1959 and

S. [now Seriolina] nigrofasciata (in Parukhin, 1966, 1976) from Seriola sp.; S. dorsalis Lamothe-Argumedo et al., 1997 from Seriola sp.; S. attenuatum Hafeezullah, 1971 from Scomberoides tala; S. argyrosomi

Shen in Shen and Qiu, 1995; S. ceylonicum Luhe, 1906; S. simhai Gupta and Ahmad, 1979; S. trompeteri Zhukov, 1983; S. lucknowensis Gupta and Jahan, 1977 from Triacanthus strigilifer; S. qatarense Saoud et al.,

2002; S. triacanthi Saoud et al., 2002; S. tenue McGladdery, 1999 and several others.

Species reported so far from fishes of Karachi coast include

Stephanostomum ditrematis (Yamaguti, 1939; Manter, 1947) from

Pseudosciaena diacanthus (Bhutta and Khan, 1975) and S. dicotylus

Bilqees, 1972 from Cybium guttatum. Presently a new species of the genus S. gibsoni is described from the fish Pomadasys olivaceum.

Family: Bucephalidae Poche, 1907 GENUS BUCEPHALUS Baer, 1827

Bucephalus was initially established to represent the earliest known bucephalid B. polymorphous Baer, 1827 on the basis of a cercaria from

42 the European unionid (freshwater) Unio pictorum and Anodonta sp. Von

Siebold, 1848 erected the genus Gasterostomum for all adult trematodes with a ventral mouth, believing that a European adult bucephalid that he named Gasterostomum fimbriatum von Siebold, 1848 and which

Yamaguti, 1958 a transferred to the genus Bucephalus.

Species reported from various parts of the world are: B. allahabadensis

Srivastava, 1963 in Bagarius bagarius; B. aoria Verma, 1936 in Aoria aoria; B. australis (Szidat, 1961) in Urophycis brasiliensis; B. bagarius

Srivastava, 1963 in Bagarius bagarius; B. barina Srivastava, 1938 in

Scatophagus argus; B. blanchardi (Stossich, 1898) in Labrax lupus; B. brevitentaculatus Corkum, 1967 in Trichurus lepturus; B. carangis

Yamaguti, 1970 in Caranx lugubris; B. carangoides Yamaguti, 1970 in

Carangoides sp., Sphyraena barracuda; B. chilensis Szidat, 1963; cercaria in Brachyodontes rodriguezi; B. clupeae Beneden, 1870 in

Clupea sprutus; B. confusus Velasquez, 1959 in Sphyraena borealis; B. crux Levinsen, 1881 in Modiolaria discors; B. cynoscion Hopkins, 1956 in Cynoscion nebulosus; B. elegans Woodhead, 1929 in Ambloplites rupestris; B. fimbriatus Siebold, 1848 in Perca fluviatilis; B. fragilis

Velasquez, 1959 in Megalaspis cordyla; B. gadorum in Gaddus morrhua; B. gangeticus Srivastava, 1938 in Pseudotropius athenoides,

43 Macrones seenghala; B. gorgon Linton, 1905 in Seriola lalandi, S. dumerili, S. zonata; B. heterotentaculatus Bravo-Hollis et Sogandares-

Bernal, 1956 in Scomberomorus sierra; B. indicus Srivastava, 1938 in

Macrones seenghala; B. intermedius in Anodonta cellensis; B. introversus Manter, 1940 in Seriola sp. and Caranx hippos; B. jagannathai Verma, 1936 in Cybium guttatum; B. kaku Yamaguti, 1970 in Sphyraena barracuda; B. kathetostomae (Manter, 1934) Manter, 1940 in Kathetostoma albigutta; B. labracis Paggi et Orecchia, 1965 in

Morone labrax; B. leognathi Velasquez, 1959 in Seriola nigrofasciata;

B. longicornutus (Manter, 1954) Howell, 1966 in Kathetostoma gigarteum; B. margaritae Ozaki et Ishibashi, 1934 in Pictada mertensii;

B. marinus Vlasenko, 1931 in Onos tricirrhatus; B. markewitschi

Koval, 1949 in Lucioperca and Esox; B. mytili in Mytilus edulis; B. octotentacularis Kakaji, 1969 in Wallagonia attu; B. paraheterotentaculatus Velasquez, 1959 in Seriola nigrofasciata; B. polymorphus Baer, 1827 in Unio and Anodonta; B. priacanthi Manter,

1940 in Priacanthus arenatus; B. pseudovaricus Velasquez, 1959 in

Caranx sp.; B. retractilis Yamaguti, 1952 in Caranx sp.; B. scorpaenae

Manter, 1940 in Scorpaena plumieri; B. sebastichthydis Yamaguti, 1959 in Sebastichthys pachycephalus; B. sextentaculatus Yamaguti, 1970 in

Caranx sexfasciatus; B. skrjabini Akhmerov, 1963 in Siniperca chuatsi;

44 B. solitarius Kohn, 1966 in Caranx crysos; B. sphyraenae Yamaguti,

1952 in Sphyraena sp.; B. tridentacularia Verma, 1936 in Aoria aoria,

Macrones seenghala and Bagarius bagarius; B. tritentacularis

Srivastava, 1963 in Bagarius bagarius; B. ulua Yamaguti, 1970 in

Carangoides ferdau; B. uranoscopi Yamaguti, 1934 in Uranoscopus japonicus; B. urophyci Szidat, 1961 in Urophycis brasiliensis; B. viperae

(Beneden, 1870) Yamaguti, 1958 in Trachinus vipera and B. vivae

(Beneden, 1870) Yamaguti, 1958 in Trachinus draco.

Only one species previously reported from Pakistan is B. varicus

(Manter, 1940) Zaidi and Khan, 1977 from the fish Sphyraena obtusata.

The primary generic characters for Bucephalus are pre-testicular ovary and features of the rhynchus. The rhynchus consists of a sucker that is topped by a ring of 6-21 retractable tentacles, depending on the species.

The tentacles may have one or two lobes extending from the posterior surface and retractable papillae may be present on the terminal end of tentacles. Most species have a relatively thin-walled cirrus sac (<5µm thick). The male duct is straight and consists of a pars prostatica and a distal thin-walled ejaculatory region; the pars prostatica may have two or three distinct regions (but no seminal duct), with different types of

45 prostatic cells. The ejaculatory duct opens into the genital atrium in close association with the protrusible genital lobe (Gibson and Bray, 2004).

Here two new species Bucephalus otolithi and Bucephalus mujibi have been identified from the fishes Otolithus argenteus and Cybium guttatum respectively.

Family: Bucephalidae Poche, 1907

GENUS PROSORHYNCHUS Odhner, 1905

There are over 70 named species of Prosorhynchus Odhner, 1905, about

60 of which are marine and 10 are from freshwaters. These are described from China (Pan, 1988), South-Eastern Asia (Moravec and Sey, 1989) and South America (Thatcher, 1999). As pointed out by Cribb et al.

2002, 13 of the marine species are found in members of the fish family

Serranidae, particularly in the sub-family Epinephelinae. Some hosts were not fully identified and Cribb et al., 2002 did not consider the parasites from these. Cribb et al., 2002 detected 16 species reported in what are definitely or probably epinephelines. These are P. atlanticus

Manter, 1940; P. bulbosus Kohn, 1961; P. caudovatus Manter, 1940; P. chorinemi Yamaguti, 1952; P. epinepheli Yamaguti, 1939; P. freitasi

Nagaty, 1937; P. gonoderus Manter, 1940; P. jupe (Kohn, 1967); P. 46 longisaccatus Durio et Manter, 1968; P. macintoshi (Velasquez, 1959);

P. ozakii Manter, 1934; P. pacificus Manter, 1940; P. platycephali

(Yamaguti, 1934); Srivastava, 1938; Bilqees, 1981; P. promicropsi

Manter, 1940; P. serrani Durio et Manter, 1968 and P. thapari Manter,

1953 (Manter, 1934; 1940a, b; 1953; Yamaguti, 1934; 1939; 1952;

Nagaty, 1937; Velasquez, 1959; Kohn, 1961; 1967; Durio et Manter,

1968).

Some other species of the genus Prosorhynchus reported from different parts of the world are: P. aguayoi Perez Vigueras, 1955 in Rypticus saponaceus; P. apertus McFarlane, 1936 in Ophiodon elongatus; P. arabianus Srivastava, 1937 in Synaptura pan; P. atlanticus Manter,

1940 in Mycteroperca bonaci, M. microlepis and M. venenosa; P. berycis

Yamaguti, 1970 in Beryx decadactylus; P. bulbosus Kohn, 1961 in

Garrupa sp.; P. caudovatus Manter, 1940 for P. crucibulum Eckmann,

1932 in Epinephelus sp., Epinephelus goreensis and Lutjanus maltzani;

P. chasmoechinatus Komiya et Tajimi, 1941 in Pseudorasbora parva; P. chorinemi Yamaguti, 1952 in Chorinemus moadetta; P. congeri

Yamaguti, 1970 in Conger sp.; P. costai Travassos, Artigas et Pereira,

1928 syn . of P. crucibulum Nagaty, 1937 in Acestrorhamphus sp.,

Aspranax fasciatus and Salminus hilarii; P. crucibulum (Rud., 1819)

47 Odhner, 1905 syn. of Monostomum crucibulum Rud., Gasterostomum armatum Molin, G. crucibulum Olsson, in Conger sp., Conger myriaster,

Muraenesox cinereus, Serranus sp., Polynemus opercularis, Sparus berda and Eumicrotremus pacificus; P. crucibulum japonicus Yamaguti,

1958 in Conger myriaster; P. elongatus (Pigulewsky, 1931) Kniskern,

1952 syn. Mordriikovia elongatus Pigulewsky, 1931, in Esox lucius; P. epinepheli Yamaguti, 1939 in Epinephelus akaara and Epinephelus sp.;

P. facilis (Ozaki, 1924) Eckmann, 1932 syn., Gotonius facilis Ozaki,

1924 in Intimicus japonicus, Seriola aureovittata and Lethrinus haematopterus; P. freitasi Nagaty, 1937 in Serranus guttatus,

Epinephelus and Plecapomus; P. gonoderus Manter, 1940 in

Epinephelus analogus; P. grandis Lebour, 1908 syn. of P. crucibulum

(Rud.) Nagaty, 1937 in Gadus morrhua and G. merlangus; P. kahala

Yamaguti, 1970 in Seriola dumerili; P. longicollis Yamaguti, 1953 in

Sphyraena sp.; P. longisaccatus Durio et Manter, 1968 in serranid; P. longus Velasquez, 1959 in Psettodes erumei; P. luzonicus Velasquez,

1959 in Lates calcarifer; P. macintoshi (Velasquez, 1959)n. comb., syn.

Neidkartia macintoshi Velasquez, 1959 in Epinephelus bleekeri; P. manteri Srivastava, 1938 in Tetrodon oblongus; P. ovatus (Linton, 1900)

Linton, 1940 syn. Gasterostomum ovatum Linton, 1940 in Lobotes surinamensis; P. ozakii Manter, 1934 in Epinehelus niveatus,

48 Mycteroperca sp., Epinephelus analogus and Garrupa sp.; P. pacificus

Manter, 1940 syn. P. atlanticus Manter, 1940 - Hanson, 1940 in

Mycteroperca sp., Sebastopyr ruberrimus, Trisotropis venenosus apua and Epinephelus analogus; P. paracrucibulus Velasquez, 1959 in

Ambassis buruensis; P. platycephali (Yamaguti, 1934) Srivastava, 1938, syn. Gotonius platycephali Yamaguti, 1934 in Platycephalus indicus; P. polydactyli Yamaguti, 1970 in Polydactylus sexfilis; P. promicropsi

Manter, 1940 in Promicrops itaiara and Mycteroperca bonaci; P. rotundus Manter, 1940 in Rypticus safronaceus bicolor; P. scapellum

McFarlane, 1936, syn. P. crucibulum Nagaty, 1937 in Scorpaenichthys marmoratus; P. serrani Durio et Manter, 1968 syn. P. crucibulum

(Rud.) of Nagaty, 1937 in Serranus louti; P. stunkardi Siddiqi et Cable,

1960 in Scomberomorus sp.; P. thapari Manter, 1953 syn. P. facilis

(Ozaki) of Nagaty, 1937 in Plectropoma maculatum; P. triangularis

Tubangui et Masilungan, 1944 in Glossogobius giurus; P. triglae Nicoll,

1914 syn. P. crucibulum Nagaty, 1937 in Trigla gurnardus; P. truncatus

Verma, 1936 in Arius jatius and P. tsengi Chin, 1933 syn. P. platycephali Yamaguti, 1934 in Platycephalus indicus.

Odhner, 1905 established the genus Prosorhynchus as bucephalids that had a rhynchus lacking a sucker and named P. squamatus Odhner, 1905

49 as the type-species. He considered P. squamatus to be distinct from specimens identified as Gasterostomum armatum by van Beneden, 1870 and used the name P. aculeatus (van Beneden, 1870) for those specimens. Odhner, 1905 also transferred G. crucibulum (Rudolphi,

1819) which Rudolphi, 1819 had described as Monostomum crucibulum into his genus, creating P. crucibulum (Rudolphi, 1819). Nagaty, 1937 and Jones, 1943 believed P. squamatus and P. crucibulum to be synonyms and therefore considered P. crucibulum (Rudolphi, 1819) the type, however, Brinkmann, 1957 showed clear evidence for separating all three of Odhner’s species.

Odhner, 1905 made the combination Prosorhynchus aculeatus Odhner,

1905 for specimens described as Gasterostomum crucibulum Beneden,

1870. He considered these specimens to be conspecific with those reported as G. armatum, but Issaitschikow, 1928 thought that the erection of genus Skrjabiniella was warranted to accommodate P. aculeatus, as S. aculeatus, because the opening of the pharynx was so far posterior and because the testes were oblique rather than tandem.

Jones, 1943 later redescribed S. aculeatus and advocated that elongate species in Prosorhynchus having tandem testes and the vitellarium in two separate lateral groups be placed in Prosorhynchus (sensustricto),

50 while those species having elliptical body with oblique testes and the vitellarium in one anterior arching field be placed in Skrjabiniella.

Yamaguti, 1958a; 1971 relegated Issaitschikow’s, 1928 genus to sub- generic status and included P. magniovatus Yamaguti, 1938 and P. uniporus Ozaki, 1924 (as a synonym of P. aculeatus) in the subgenus

Skrjabiniella. Manter, 1953 believed that the generic level features promoted by Issaitschikow, 1928 were of specific level value only, and considered Skrjabiniella a junior synonym of Prosorhynchus.

Brinkmann, 1957, Kniskern, 1952 and even Skrjabin and Guschanskaja,

1962 also considered Skrjabiniella a junior synonym of Prosorhynchus

(Gibson and Bray, 2004).

Cribb et al., 2002 mentioned that there were 13 species of

Prosorhynchus reported from epinephaline serranids and this figure now stands at 16 since new species have been added. The names are P. maternus Bray and Justine, 2006; P. longisaccatus Durio et Manter,

1968 and P. lamprelli Bott et Cribb, 2005. Prosorhynchus maternus belongs to a group of species which share the character of the uterus being restricted to the post-ovarian region, a relatively unusual feature in this genus. Other members of this group are P. atlanticus Manter, 1940;

P. bulbosus Kohn, 1961; P. gonoderus Manter, 1940; P. longicollis

51 Yamaguti, 1953; P. macintoshi (Velasquez, 1959); P. pacificus Manter,

1940 and P. promicropsi Manter, 1940. Of parasites of non- epinephelines only P. mizellei Kruse, 1977 from the smooth lump sucker, the cyclopterid Aptocyclus ventricosus (Pallas), in the Bering Sea and P. kahala Yamaguti, 1970 from the great amberjack, the carangid Seriola dumerili have this character (Yamaguti, 1970; Kruse, 1977).

The only species so far reported from Epinephelus malabaricus is P. pacificus (or its synonym P. atlanticus). These records are by

Hafeezullah and Siddiqi (1970) from Karwar, India and Leong and Wong

(1988, 1990) from juvenile E. malabaricus imported from the Philippines for culture in Malaysia.

Species of Prosorhynchus reported from Pakistan are: P. platycephali

(Yamaguti, 1934; Srivastava, 1938; Bilqees, 1981) from Platycephalus scabar; P. thapari (Manter, 1953) Bilqees, 1981 from Plectorhynchus cinctus; P. longus (Velasquez, 1959); Bilqees, 1981 from Psettodes erumei and P. erumenis Bilqees, 1976 from Psettodes erumei.

During present studies a new species Prosorhynchus magnacirrus has been identified from the fish Psettodes erumei.

52 MATERIALS AND METHODS

Data presented in this investigation were obtained from February, 2005 to July, 2007.

A total of 791 fishes belonging to 14 fish families, 10 genera and 14 species were collected from different points of West Wharf, Karachi

(Table 2). The fish species collected include Pseudosciaena diacanthus,

Otolithus argenteus, Muraenesox cinereus, Stromateus cinereus,

Stromateus sinensis, Chirocentrus dorab, Crenidens indicus, Lutjanus johnii, Arius serratus, Lates calcarifer, Muaena sp., Psettodes erumei,

Platycephalus scabar, Plectorhynchus cinctus, Cybium guttatum,

Stromateus niger, Caranx affinis and Pomadasys olivaceum.

COLLECTION OF SPECIMENS

The fishes were dissected from the ventral side and the gut was exposed.

It was then placed in the large Petri dish, stretched out and cut into three

regions i.e. the fore-gut (stomach), the mid-gut (small intestine) and the hind-gut (large intestine with the rectum). The liver and gonads were also

65 separated and were found negative. The sections of the gut were opened

with a longitudinal cut, and the whole inner surface was slightly scraped

to remove the parasites with mucous. These were then washed several

times with tap water and counted by sorting from the mucous under a low

powered microscope. The individual specimens were then removed from

the gut wall for further study under the microscope.

FIXATION AND PREPARATION OF PERMANENT SLIDES

The parasites were fixed in AFA solution (a mixture of 70% Ethyl

Alcohol, Formalin, and Acetic acid in the ratio of 90:7:3) for 24 hours

under slight cover glass pressure and then washed several times with

70% alcohol, stained with Mayer’s Carmalum, dehydrated in a graded

series of alcohols, cleared in clove oil and xylene and mounted

permanently in Canada Balsam by usual method.

Measurements are given length by width in millimeters. Diagrams were

made with the help of a Camera lucida. Photographs of some of the

specimens to show peculiar diagnostic features, which were taken with a

Nikon photomicroscope (Optiphat-2) using fuji colour film. Type

66 specimens are in the collection of the department of Zoology, Jinnah

University for Women, Karachi.

67 DESCRIPTIONS

SPECIES OF THE GENUS PLEORCHIS PLEORCHIS HETERORCHIS N.SP. (Figs. 1-4)

Family: Pleorchiidae (Poche, 1926) Yamaguti, 1971

Sub-Family: Pleorchiinae Caballero, 1952

Genus: Pleorchis Railliet, 1896

Host: Lutjanus johnii (Lutianidae) and Otolithus argenteus (Sciaenidae)

Location: Intestine

Locality: West Wharf, Karachi.

No. of specimens: 1 from each host, 41 specimens of first host and 73 of second host were examined.

Holotype: JUW – T1

Paratype: JUW – T2

Diagnosis

Body flat, longitudinally oval, elongate, and medially broad, slightly truncate posteriorly with mid-terminal notch. Entire surface is smooth.

Ventral sucker is round, embedded at anterior 1/3 of body length.

Prepharynx is well developed, the posterior part of the prepharynx is widened in the form of a funnel into which projects the muscular pharynx. Oesophagus is short and broad at the base. Intestinal caeca

69 H-shaped with anterior arms much shorter than the posterior, intestinal bifurcation almost in the middle of forebody, anteriorly caeca wide and short extending as far as anterior limit of pharynx.

Posteriorly caeca reach close to posterior end of the body with no lateral out pocketings.

Male reproductive system consists of 44 testes, intercaecal, arranged in two parallel rows, right row having 12 pairs of testes and left 10 pairs of testes (Fig. 1). Testes sub-globular, entire to slightly or distinctly irregular, almost all of them of the same size, extending immediately from posterior of the ovary to anterior of the excretory vesicle. Cirrus pouch overlaps the ventral sucker, extends into hindbody, terminating above the ovary, contains bi-partite seminal vesicle, pars prostatica and ejaculatory duct. Proximal part of seminal vesicle is spherical, dorsal, smaller than the distal part which is antero-ventral. Ejaculatory duct opens into the genital atrium. Genital atrium spherical, wide, thin-walled. Genital pore is very small situated just behind the bifurcation of the intestinal caeca and above the ventral sucker.

Female reproductive system consists of ovary in the form of 16 small, unequal, irregular follicles, scattered mainly in ventral parenchyma,

70 lying above and close to the testes, more nearer to the anterior part of the body. Uterus is pre-ovarian, intercaecal, coils and fills the space between ovary and ventral sucker, ventral to cirrus sac. Eggs are numerous, thin-shelled. Vitellaria lateral, follicular, follicles small, very numerous, extending from post-bifurcal to posterior extremity of body and to lateral body margins. Excretory vesicle is tubular, relatively small, located posteriorly between the intestinal caeca reaching to the posterior level of last pair of testes. Excretory pore is terminal, within terminal notch of the body.

Principle measurements of P. heterorchis n.sp. (In millimeters)

Entire body size: 3.562–3.900 x 1.40–1.50

Forebody: 0.81–0.85 x 0.70–0.85

Hindbody: 3.10–3.25 x 1.39–1.40

Oral sucker: 0.21–0.25

Ventral sucker: 0.20–0.26

Sucker width ratio: 1:1.4

Prepharynx: 0.19–0.22 x 0.10–0.15

Pharynx: 0.15–0.23 x 0.11–0.16

Cirrus sac: 0.71–0.92 x 0.04–0.15

Anterior testis: 0.11–0.12 x 0.10–0.11

71

Posterior testis: 0.12–0.125 x 0.12–0.13

Seminal vesicle: 0.15–0.18 x 0.14–0.15

Ovarian follicles: 0.03–0.07 x 0.05–0.08

Eggs: 0.022–0.04 x 0.024–0.043

Excretory vesicle length: 0.71–0.73

Etymology: The present new species Pleorchis heterorchis is named due to the variation in the size and number of testes.

Remarks

Genus Pleorchis was originally proposed by Railliet, 1896. The type- species described was P. polyorchis from Corvina nigra. Recently it is redescribed from the fish Sciaena from Western Mediterranean Sea

(Bartoli et al., 2004). Several other species were described from various parts of the world, including: P. americanus Luhe, 1906 from

Cynoscion regalis reported from off Woods Hole, U.S.A.; P. californensis Manter and Van Cleave, 1951 in fish Cynoscion nobilis,

California; P. ghanensis Fischthal and Thomas, 1968 from the fish

Cynoscion micrognathus reported from Ghana; P. magnaporus Arai,

1963 from Cynoscion parvipinnis and Urobatis maculatus reported from Baja, California; P. sciaenae Yamaguti, 1938 from Sciaena

72 albiflora reported from East China Sea; P. uku Yamaguti, 1970 from

Apsion virescens reported from Hawaii; P. puriensis Gupta and

Ahmad, 1976 from fish Sciaena vogleri; P. mamaevi Parukhin, 1974 from fish Ichnius sp. reported from Red Sea (Indian Ocean); P. psettodesai Gupta and Gupta, 1978 from Psettodes erumei from India;

P. indicum Gupta and Puri, 1980 from Epinephelus (=Serranus) diacanthus from India; P. arabicus Al-Yamani and Nahhas, 1981 from fish Otolithus rubber from Arabian Gulf off Kuwait; P. nibeae

Shen, 1983 from fish Nibea albiflora off Hebei, China and P. hainanensis Shen, 1983 from fish Pennahia anea from off

Guangdong, China.

Karachi coast has rich fish fauna, about 1000 species of fishes are found but trematodes of only 48 species of fish (Bilqees et al., 2005) have been reported.

Digenetic trematodes are the largest group among monozoic platyhelminths. They require one or two intermediate hosts to complete their life cycles. Almost all of them are endo-parasites.

Several species of trematodes have been reported from fishes of

Karachi coast including one species of genus Pleorchis Railliet, 1896;

P. ghanensis Fischthal & Thomas, 1968 (Bilqees, 1981).

73

P. ghanensis was synonymized with P. sciaenae by Madhavi and

Narasimhulu, 1985, while Bray, 1986 considered that this and most species with about 44 testes, as probable synonyms of the latter species.

The present new species P. heterorchis is described from the fish

Lutjanus johnii (Lutianidae) and Otolithus argenteus (Sciaenidae).

Most of the species of the genus Pleorchis are found in the fish of the family Sciaenidae. Previously only one species of the genus, P. ghanensis Fiscthal & Thomas, 1968 has been reported in the fishes,

Pseudosciaena diacanthus, Otolithus argenteus (Sciaenidae) and

Pomadasys olivaceum (Pomadasyidae) of Karachi coast, Pakistan

(Bilqees, 1977).

The present new species has 44 testes arranged in 4 longitudinal rows,

20 on one side and 24 on the other side. The species which have 44 testes are P. magnaporus, P. psettodesai, P. indicum and P. puriensis. In P. indicum lateral diverticula on the posterior caeca are absent as in the present species. In P. magnaporus the caeca are shorter which terminate well apart of the posterior extremity. In this respect the present species is close to the above mentioned species but the anterior limit of vitelline fields are different. In P. puriensis

74 vitelline fields are confluent in the forebody. It also lacks anterior caeca but posterior caeca are devoid of lateral diverticula as in the present species. The vitellaria in P. puriensis is also interrupted at the ovarian level and reaches as far as posterior margin of the pharynx, while in the present species vitellaria does not reach beyond the anterior level of ventral sucker.

The excretory vesicle in the present species is tubular and shorter reaching only to posterior level of the posterior testes. The post- testicular space is also relatively longer. The testes in the holotype are slightly irregular or rounded in the specimen from Otolithus argenteus. The ovary in the present form consists of 16 irregular follicles of different sizes while ovary in all other species is either irregular or lobed.

The present species is therefore, separated from other species in having a combination of characters such as oral sucker equal or slightly smaller than ventral sucker, anterior caeca are prominent, forebody much shorter than hindbody, vitellaria extending from anterior level of ventral sucker to posterior extremity of the body, confluent in the post-testicular region. Ovary consists of 16 irregular and unequal follicles anterior to testes. Testes are 44 in number,

75 irregular or smooth, cirrus sac is extending much posterior to ventral sucker enclosing bilobed seminal vesicle, posterior lobe being larger.

Pars prostatica is small. Ejaculatory duct is long and genital opening immediately post-bifurcal. On the basis of such distinguishing characters from other known species the present form is considered to be a new species for which the name Pleorchis heterorchis n.sp. is proposed.

76 SPECIES OF THE GENUS DECEMTESTIS DECEMTESTIS JOHNII N.SP. (Fig. 5)

Family: Opecoeliidae Ozaki, 1925.

Sub-family: Plagioporinae (Dujardin, 1845)

Genus: Decemtestis Yamaguti, 1934

Host: Lutjanus johnii (Lutianiidae)

Location: Intestine

Locality: West Wharf, Karachi.

No. of specimens: 4 from 1 fish, 37 fishes were examined.

Holotype: JUW – T3

Paratype: JUW – T4

Diagnosis

Relatively small trematodes, body smooth, flattened with narrow forebody and broader hindbody. Anterior end is bluntly rounded and posterior end is broadly rounded, greatest width is in the middle. Oral sucker is sub-terminal, rounded. Pre-oral lobe is very small.

Prepharynx is not present. Pharynx is pear-shaped. Oesophagus is relatively long. Caeca are simple, long reaching to near posterior extremity of body. Acetabulum is rounded, larger than oral sucker, highly muscular. Genital pore is immediately above the intestinal

77 bifurcation at the base of oesophagus. Cirrus sac is long, extending from the anterior level of acetabulum to the base of oesophagus containing bilobed seminal vesicle at the base, a small pars prostatica and a long ejaculatory duct ending into a small, shallow genital atrium. The male and female ducts open separately in the genital atrium. Testes are 10 in two rows, 5 in each row, spherical in shape, almost smooth. Ovary is in posterior half of body, deeply lobed, slightly post equatorial with 7-8 prominent lobes, 5 lobes are large and

2-3 small projections, close to anterior first pair of testes. Uterus small, coiled anterior to ovary. Eggs are numerous, oval in shape.

Vitellaria consist of numerous rounded follicles extending anteriorly, anterior to acetabulum, terminating posteriorly to intestinal bifurcation, confluent in the posterior region and reaching to posterior extremity of the body. Excretory pore is terminal. Excretory vesicle is tubular, extending anteriorly to the level of anterior pair of testes.

Principle measurements of D. johnii n.sp. (In millimeters)

Body size: 2.65–2.71 x 0.86–0.89

Oral sucker: 0.20–0.21

Oesophagus: 0.31–0.33 x 0.90–0.10

Ventral sucker: 0.41–0.43 x 0.29–0.3

78

Sucker width ratio: 1:1.4

Pharynx: 0.11– 0.12 x 0.130–0.133

Testes: 0.07–0.09 x 0.11–0.12

Ovary: 0.35–0.37 x 0.32–0.33

Cirrus pouch: 0.645–0.67 x 0.05–0.07

Seminal vesicle: 0.40–0.48 x 0.14–0.16

Eggs: 0.04–0.07 x 0.011–0.020

Excretory vesicle: 0.91–0.93

Posterior testes from posterior extremity: 0.49–0.51

Etymology: The present new species Decemtestis johnii refers to the host.

Remarks

The present specimens are included in the genus Decemtestis

Yamaguti, 1934 as these are small trematodes with lanceolate body, unarmed. Hindbody with slightly crenulated lateral margins. Oral sucker is sub-terminal, medium-sized. Oesophagus is bifurcating at about middle of forebody. Caeca terminating near posterior extremity.

Testes are ten in number, in two more or less irregular longitudinal rows in posterior half of the body. Cirrus pouch is elongated,

79 extending over acetabulum slightly overlapping the anterior border, enclosing the winding seminal vesicle. Genital pore is sub-median.

Ovary is immediately pre-testicular. Uterus coiled between testes and acetabulum. Eggs are non-filamented. Vitellaria are extending in lateral fields from posterior extremity to intestinal bifurcation, confluent in post-testicular area. Excretory vesicle is tubular reaching to level of anterior testis.

The genus was erected by Yamaguti (1934) with type-species D. sillagonis in Sillago sihama from Japan. Later on several other species have been described including: D. azumae (Layman, 1930)

Yamaguti, 1934; D. bera Yamaguti, 1938; D. callionymi Yamaguti,

1934; D. ditrematis Yamaguti, 1934; D. goniistii Yamaguti, 1934; D. megacotyla Yamaguti, 1938; D. neopercis Yamaguti, 1938; D. pagrosomi Yamaguti, 1938; D. parapercis Yamaguti, 1938; D. spari

Yamaguti, 1938; and D. takanoha Yamaguti, 1951. The species reported from India are D. biacetabulatus Srivastava, 1936; D. brevicirrus Srivastava, 1936; D. mehrai Srivastava, 1936. Other species of the genus reported are D. kobayashii Park, 1939 and D. pseudolabri Manter, 1954 from Korea and New Zealand.

80 Species of the genus Decemtestis have not been reported previously from fishes of Karachi coast. Present is the first report describing a new species D. johnii. The present new species is different from the

Japanese species in the size of body, sucker-width ratio, anterior extension of vitellaria, shape of ovary, shape of testes, position of genital opening. In D. neopercis, D. goniistii, D. bera, D. spari and D. pagrosomi, the testes are mostly transversely elongated and are relatively large. In D. megacotyla acetabulum is large and transversely elongated, the testes are also distinctly transversely elongated.

In all these species the genital opening is at the base or anterior level of pharynx while it is at the base of intestinal bifurcation in the present new species. The ovary in the above mentioned species is irregular or 4 lobed while it is 7-8 lobed in the present new species.

The cirrus sac in D. pagrosomi and D. spari is also extending to posterior level or posterior to acetabulum while it is terminating at the anterior level or slightly overlapping the acetabulum in the present new species.

The species D. sillagonis and D. callionymi are different from the present species in having the genital opening at the level of pharynx and ovary with 3-4 lobes. In D. callionymi the vitellaria are also

81 interrupted at the acetabular level and the cirrus sac is extending far posterior to acetabulum. In D. ditrematis the arrangement of testes is different, two longitudinal rows of 4 each and one anterior and posterior in the center. D. pseudolabri is much smaller (0.539–0.616 x

0.2–0.246) than the present species. D. biacetabulatus is also smaller with bilobed acetabulum and from a different fish. D. brevicirrus is from a different fish host, has small cirrus and small body. In the two above species vitelline extension is also variable as compared to the present species.

82 SPECIES OF THE GENUS LECITHOCLADIUM LECITHOCLADIUM CYBII N.SP. (Figs. 6-7)

Family: Hemiuridae Looss, 1899, Luhe, 1901

Sub-family: Diurinae Looss, 1907

Genus: Lecithocladium Luhe, 1909

Host: Cybium guttatum (Cybidae)

Common name: Surmai

Location: Stomach

Locality: Karachi coast, Pakistan

No. of specimens: 9 specimens from 2 hosts, 53 fishes were examined.

Holotype: JUW – T5

Paratype: JUW – T6

Diagnosis

Body divided into soma and ecsoma. Soma is large and wide at the level of ventral sucker. Ecsoma is smaller than soma becoming narrower towards the tail ending into a knob. Oral sucker is large, terminal, flattened, oral opening wide, transversely elongated.

Prepharynx is absent. Pharynx is large, elongated, strongly muscular, oesophagus prominent, anterior part of which is sac-like. Intestinal

83 bifurcation is anterior to ventral sucker. Ventral sucker is smaller than oral sucker. Testes are two, diagonal, sub-globular, one behind the other, situated almost in the middle of hindbody. One posterior to seminal vesicle and one lateral to it. Seminal vesicle elongate, pointed at both ends, anterior extremity recurved back over proximal part, leading to pars prostatica. Pars prostatica is long, winding reaches anteriorly to the posterior level of ventral sucker. Cirrus sac is long, narrow, extending from the level of ventral sucker to forebody.

Junction of cirrus sac and genital atrium is at the mid-level of pharynx. Genital atrium is also long, tubular, narrow, genital pore ventral to anterior part of oral sucker. Ovary post-testicular, well separated from posterior testes, in posterior fourth-quarter of soma, transversely flat. Seminal receptacle is immediately posterior to ovary and smaller than it. Vitellaria consist of 7, long tubes, all directed posteriorly. Uterus reaches posteriorly to middle of ecsoma, then passes forward coiling between ovary and testes, dorsal to testes and seminal vesicle, opening into base of cirrus sac. Eggs are numerous, small, oval to elongate. Excretory pore is terminal on ecsoma.

Excretory vesicle is rounded posteriorly, bifurcates at the level of posterior testis, arms unite dorsal to oral sucker.

84 Principle measurements of L. cybii n.sp. (In millimeters)

Body size: 6.9–6.91 x 0.59–0.6

Soma length: 3.70–3.71

Soma width: 0.59–0.6

Ecsoma length: 3.20–3.21

Forebody: 1.31–1.32

Hindbody: 5.7–5.72

Oral sucker: 0.4–0.41 x 0.51–0.52

Pharynx: 0.415–0.42 x 0.31–0.32

Ventral sucker: 0.37–0.38 x 0.39–0.4

Sucker width ratio: 1: 0.6

Seminal vesicle: 0.35–0.36 x 0.61

Ventral sucker to seminal vesicle: 0.6–0.61

Ventral sucker to anterior testis: 0.88–0.89

Anterior testis: 0.12–0.125 x 0.1–0.110

Posterior testis: 0.12–0.175 x 0.141–0.145

Posterior testis to ovary: 0.30–0.31

Ovary: 0.28–0.29 x 0.10–0.11

Eggs: 0.03–0.037 x 0.017–0.025

85 Etymology: The present new species Lecithocladium cybii refers to the host.

Remarks

The species of the genus Lecithocladium Luhe, 1909, described previously from Pakistan are L. psenopsis Yamaguti, 1934 (Bilqees,

1981) from Stromateus sinensis; L. hexavitellarii (Bilqees, 1971); L. anteporus (Bilqees, 1971); L. microductus (Bilqees, 1971); L. arabiana, (Bilqees, 1971); L. microcaudum (Bilqees, 1971); L. octovitellarii (Bilqees, 1971) all from Stromateus sp., of Karachi coast; L. karachii Zaidi and Khan, 1977 from Carangoides presustus;

L. pakistanensis Zaidi and Khan, 1977 from Carangoides malabaricus; L. stromatei Farooq and Khanum, 1980 from

Stromateus niger; L. arabicum Farooq and Khanum, 1980 from

Caranx djedaba and L. thynensis Bilqees and Nighat, 1985 from

Thynnus sp. and L. arii Bilqees et al., 2005 from the fish Arius serratus. These species are also from fishes of Karachi coast. All species except L. karachii, L. arabicum and L. thynensis are from fishes of family Stromateidae.

Present species is different from the previously described species as well as from the species from other parts of the world. The present

86 species is characterized by having infundibuliform oral sucker, large muscular pharynx longer than oral sucker, anterior part of oesophagus sac-like, ventral sucker smaller than oral sucker, testes diagonal, anterior testis overlapping seminal vesicle dorsally at the posterior level. Seminal vesicle elongate, pointed at both ends. Pars prostatica winding reaching to the posterior level of ventral sucker. Sinus sac is long, tubular, extending to the middle of pharynx joining the tubular genital atrium which opens at the anterior level of ventral sucker.

Ovary is in the posterior fourth-quarter of soma, larger than the testes.

Seminal receptacle is immediately behind ovary. Vitellaria are 7, long tubes, 4 of which extend posteriorly into ecsoma. Uterus is reaching to the middle of ecsoma. Excretory vesicle is knob-like posteriorly, anteriorly extending into a tube to the posterior level of testes.

The anterior sac like portion of oesophagus has not been described in any of the previous species. Although ventral sucker is smaller than oral sucker is found in several other species. Species having oral sucker larger than ventral sucker are L. exisum Luhe, 1901; L. brevicaudum Srivastava, 1942; L. annulatum Velasquez, 1962; L. anguistiovum Gibson and Bray, 1986; L. parvioum Yamaguti, 1953;

L. scombri Yamaguti, 1953; L. apolectis Velasquez, 1962; L. aegyptensis Fischthal and Kuntz, 1963; L. bulbolabrum Reid, Coil &

87 Kuntz, 1966; L. stromatei Gupta & Gupta, 1983; L. arabicum Farooq

& Khanum, 1980; L. inglisi Gupta et Ahmed, 1977; L. lutiani Gu et

Shen, 1978; L. keralense Gupta & Gupta, 1983; L. indicum Gupta &

Gupta, 1978, L. elongatus Gupta and Puri, 1981; L. bengalensis

Gupta & Gupta, 1983; L. jagannathi Ahmed, 1981; L. sulphuriusi

Gupta & Gupta, 1983; L. microlepidotus Gupta & Gupta, 1983; L. caranxi Gupta & Gupta, 1983; L. siddiqui Gupta & Gupta, 1978 as in the present species but these are different in body size, relative length of soma and ecsoma and shape, size of seminal vesicle and ovary and presence of sac-like anterior part of oesophagus.

Species having oral sucker smaller or equal to ventral sucker are L. magnacetabulum Yamaguti, 1934; L. carultum Gupta & Gupta,

1983; L. megalaspis Gibson & Bray, 1986; L. chauhani Chauhan,

1945; L. triacanthi Gupta & Gupta, 1983; L. manteri Gupta and Puri,

1981; L. psenopsis Gupta & Gupta, 1983; L. glandulum Chauhan,

1945; L. chingi Manter & Pritchard, 1960; L. falklandicum Gupta &

Gupta, 1983; L. purense Gupta & Gupta, 1978; L. thapari Gupta &

Gupta, 1983; L. puriensis Gupta & Gupta, 1983; L. singhi Gupta et

Singh, 1983; L. guptai Gupta & Gupta, 1983; L. seriolellae Gupta &

88 Gupta, 1983; L. tewarii Gupta & Gupta, 1987 are different from the present species.

L. gazzai Pandey et al., 2000 differs from the present new species L. cybii in having oral sucker larger than the ventral sucker while in L. gazzai the oral sucker is almost equal to the ventral sucker.

L. gazzai is median, ventral to the oral sucker. Seminal vesicle is spindle-shaped, small as compared to the present new species. Ovary is median, tri-lobed, post-testicular lying a little anterior to the posterior end of soma separated from posterior testis by uterine coils.

Vitellaria post-ovarian, consisting of 10 long, slender, winding tubules confined up to a little anterior to posterior end of soma,while all the new species described here consist of 7 vitelline tubules. L. gazzai resembles with L. chaetodipteri in having oral sucker smaller of same size as ventral sucker and the presence of 10 vitelline tubules but differs from it in having pharynx not overlapping anterior margin of ventral sucker, seminal vesicle is not extending to posterior margin of ventral sucker and large size of body.

89 L. gazzai differs from L. cybiumi and L. hanumanthai in having genital pore at anterior part of oral sucker, tri-lobed ovary, in the presence of pre-oral lobe and 10 vitellaria Pandey et al., 2000.

L. bengalensis Saxena and Prakash, 2006 differs from previously described species in having genital pore sub-median, lying in the middle of the oral sucker while resembles with other Lecithocladium species in having same number of vitelline tubules i.e. 7.

90 LECITHOCLADIUM KARACHIENSIS N.SP. (Fig. 8)

Host: Parastromateus niger (Stromateidae)

Location: Stomach

Locality: Karachi coast, Pakistan.

No. of specimens: 17 from 3 fishes, 63 fishes were examined.

Holotype: JUW – T7

Paratype: JUW – T8

Diagnosis

Body is long, soma cylindrical, widest in the region of the ventral sucker. Ecsoma is also long, or slightly shorter and narrower than the soma with bluntly pointed tail. Oral sucker is sub-terminal, wider than long, sub-globular, larger than ventral sucker. Prepharynx is absent.

Pharynx is elongated to oval. Ventral sucker is situated anteriorly, and posterior to intestinal bifurcation. Oesophagus is a saccular chamber terminating into a small tube. Caeca showing uniform thickness and wider at the posterior extremity of ecsoma.

Testes are two, equal in size, globular, smaller than the ovary, situated immediately posterior and close to seminal vesicle. Seminal vesicle thin-walled, oval, pointed anteriorly and rounded posteriorly, far

91 posterior to ventral sucker, anterior extremity of the seminal vesicle joins the pars prostatica. Pars prostatica is anterior to seminal vesicle, uncoiled, not extending into the forebody. Sinus sac is long, tubular, extending into anterior forebody, joins the genital atrium at the posterior level of pharynx. Genital opening is at the base of oral sucker.

Ovary is post-testicular, well separated from testes, in the posterior- third of soma. Seminal receptacle is smaller than ovary and immediately posterior to it. Uterus reaches to ecsoma a shorter distance to mid-level, then passes forward coiling between ovary and testes joining the sinus sac immediately anterior to ventral sucker.

Vitellaria consist of 7, long, delicate, tubules, radiating at the point between ovary and seminal receptacle, all directed posteriorly, reaching at the junction of soma and ecsoma but not extending posteriorly into ecsoma. Eggs are numerous, small. Excretory pore is terminal. Excretory vesicle is tubular.

92 Principle measurements of L. karachiensis n.sp. (In millimeters)

Entire size: 8.8–8.98 x 1.33–1.39

Soma length: 4.7–4.75

Soma width: 0.78–0.80

Ecsoma size: 4.1–4.23 x 0.55–0.59

Forebody: 1.43–1.45 x 0.6–0.62

Hindbody 7.00–7.10 x 0.69–0.70

Oral sucker: 0.49–0.5 x 0.75–0.77

Ventral sucker: 0.48–0.49 x 0.49–0.5

Sucker width ratio: 1: 0.6

Pharynx: 0.49–0.5 x 0.38–0.4

Seminal vesicle: 0.5–0.51 x 0.21–0.22

Sinus sac length: 1.9–2.0

Anterior testis: 0.20–0.21 x 0.18–0.19

Posterior testis: 0.14–0.15 x 0.16–0.163

Posterior testis to ovary: 0.09–0.1

Ovary: 0.21–0.22 x 0.24–0.25

Eggs: 0.028–0.034 x 0.016–0.020

Etymology: The present new species Lecithocladium karachiensis refers to its locality.

93 Remarks

There is a long list of species of the genus Lecithocladium from several localities including India, Pakistan and other parts of the world

(Ahmad, 1981; Gupta & Mehrotra, 1970; Gupta & Singh, 1982;

Gupta & Govind, 1985; Gupta & Gupta, 1978; Gupta & Ahmad,

1977; Gupta & Puri, 1981; Hafeezullah, 1975; Gupta & Gupta, 1983;

Gupta & Jain, 1992; Bilqees, 1981; Zaidi & Khan, 1977; Farooq &

Khanum, 1980; Gaevaskaya & Kovaleva, 1978; Gu & Shen, 1978;

Bashirullah et D’Silva, 1973; Machida & Uchida, 1990; Toman, 1992;

Korotaeva, 1974; Bray & Cribb, 2004; Chambers et al., 2001).

The species of the genus Lecithocladium Luhe, 1909, described previously from Pakistan are L. psenopsis Yamaguti, 1934 from

Stromateus sinensis; L. hexavitellarii (Bilqees, 1971); L. anteporus

(Bilqees, 1971); L. microductus (Bilqees, 1971); L. arabiana

(Bilqees, 1971); L. microcaudum (Bilqees, 1971); L. octovitellarii

(Bilqees, 1971) all from Stromateus sp., of Karachi coast; L. karachii

Zaidi and Khan, 1977 from Carangoides presustus; L. pakistanensis

Zaidi and Khan, 1977 from Carangoides malabaricus; L. stromatei

Farooq and Khanum, 1980 from Stromateus niger; L. arabicum

Farooq and Khanum, 1980 from Caranx djedaba and L. thynensis

94 Bilqees and Nighat, 1985 from Thynnus sp., L. arii Bilqees et al.,

2005 from the fish Arius serratus. These species are also from fishes of Karachi coast. All species except L. karachii, L. arabicum and L. thynensis are from fishes of family Stromateidae.

The present species is regarded new and shows some characters common with several other species described from Pakistan such as ventral sucker smaller than oral sucker. The species having ventral sucker smaller than the oral sucker are L. psenopsis, L. hexavitellarii,

L. anteporus, L. tetravitellarii, L. microductus, L. stromatei, L. arabicum and L. thynensis, but the present species is different in having a saccular anterior portion of oesophagus and straight pars prostatica, genital opening at the base of oral sucker and vitellaria not extending into ecsoma.

In the species described previously from Pakistan, the testes are contiguous, except in L. microductus, L. hexavitellarii, L. microcaudum, L. arabicum and L. thynensis, the ovary is very close or contiguous with the posterior testis. In L. stromatei, L. arabicum,

L. octovitellarii, the uterus reaches to part of ecsoma but vitellaria do not enter into it. In L. hexavitellarii, L. tetravitellarii, L. stromatei, L.

95 arabiana, L. thynensis and L. arii Bilqees et al., 2005 the uterus and vitellaria both extend posteriorly to some distance in ecsoma.

Number of vitelline tubules also vary. These are 8 in L. octovitellarii,

L. karachii and L. pakistanensis; 6 in L. hexavitellarii, L. microcaudum, L. arabicum, L. arabiana and L. anteporus; 4 vitelline tubules are found in L. tetravitellarii and L. microductus and

7 tubules are present in L. psenopsis, L. stromatei, L. thynensis and

L. arii as in the present new species but other characteristics are variable. Other species having ventral sucker smaller than oral sucker are L. exisum Luhe, 1901; L. brevicaudum Srivastava, 1942; L. annulatum Velasquez, 1962; L. anguistiovum Gibson and Bray,

1986; L. parvioum Yamaguti, 1953; L. scombri Yamaguti, 1953; L. apolectis Velasquez, 1962; L. aegyptensis Fischthal and Kuntz, 1963;

L. bulbolabrum Reid, Coil & Kuntz, 1966; L. stromatei Gupta &

Gupta, 1983; L. arabicum Farooq et Khanum, 1980; L. inglisi Gupta et Ahmed, 1977; L. lutiani Gu et Shen, 1978; L. keralense Gupta &

Gupta, 1983; L. indicum Gupta & Gupta, 1978, L. elongatus Gupta and Puri, 1981; L. bengalensis Gupta & Gupta, 1983; L. jagannathi

Ahmed, 1981; L. sulphuriusi Gupta & Gupta, 1983; L. microlepidotus Gupta & Gupta, 1983; L. caranxi Gupta & Gupta,

1983; L. siddiqui Gupta & Gupta, 1987 as in the present species but

96 these are different in body size, relative length of soma and ecsoma and shape, size of seminal vesicle and ovary and presence of sac-like anterior part of oesophagus.

Present species is different from all these species in relative sizes of soma and ecsoma and in other characteristics of diagnostic value. The species from India and other localities including L. glandulum

Chauhan, 1945; L. lutjanii Gu et shen, 1978; L. dawesi Bashirullah et

D’Silva, 1973; L. hanumanthi Gupta & Jain, 1992; L. inglisi Gupta et Ahmad, 1977; L. purense Gupta et Gupta, 1978 and L. singhi

Gupta et Singh, 1983 are also different from the present specimens.

Bray and Cribb (2004) regarded L. dawesi and L. hanumanthi as synonym of L. megalaspis. While L. inglisi, L. purense and L. singhi are also the synonym of L. megalaspis (Gibson & Bray, 1986). Gupta

& Gupta also suggested the following synonym: L. harpodontis

Srivastava, 1942 with L. exisum Luhe, 1901; L. psettodi Srivastava,

1942 with L. manteri Gupta & Puri, 1981; L. sciaeni Gupta & Puri,

1981 with L. elongatus Gupta & Puri, 1981; and L. srivastavai Gupta

& Puri, 1981; L. stromateusi Gupta & Puri, 1981 with L. aegyptensis

Fischthal & Kuntz, 1963 and L. fotedari Gupta & Singh, 1982 with L. jagannathi Ahmad, 1981. All these species are different from the present species in a combination of characters such as sucker-width

97 ratio, egg sizes, soma and ecsoma relative sizes and relative sizes of suckers and gonads.

98 LECITHOCLADIUM MAGNASOMA N.SP. (Figs. 9-10)

Host: Stromateus sinensis (Stromateidae)

Location: Stomach

Locality: Karachi coast, Pakistan

No. of specimens: 10 specimens from 3 hosts, 75 fishes were examined.

Holotype: JUW – T9

Paratype: JUW – T10

Diagnosis

Soma is very long and cylindrical, widest posteriorly at the junction with ecsoma. Ecsoma narrow, short, bluntly pointed posteriorly.

Plications on both sides of ecsoma are clearly visible. Oral sucker terminal, cup-shaped, wide anteriorly, narrow posteriorly, much larger in size than the ventral sucker. Ventral sucker is globular, smaller than the oral sucker, situated anteriorly. Prepharynx is absent. Pharynx is strongly muscular, rectangular, larger than oral sucker. Oesophagus is prominent. Intestinal bifurcation is pre-acetabular.

Testes are two, tandem, close together, rounded, almost equal in size, situated in the posterior half of soma. Seminal vesicle anterior to

99 anterior testes, far behind ventral sucker, thick-walled, elongated, narrow anteriorly, posterior end rounded, anterior extremity leading to pars prostatica. Pars prostatica is long, slightly coiled posteriorly, not reaching in forebody. Sinus sac narrow, extended anteriorly into the forebody, reaching to posterior level of pharynx, joining the long, narrow genital atrium at the base and lateral to pharynx. Junction of sinus sac and genital atrium is at the base of pharynx. Genital pore is ventral, at the posterior border of oral sucker.

Ovary post-testicular, situated almost in the middle of posterior half of soma, transversely oval. Seminal receptacle just posterior to ovary, vitellaria are 7, long tubules, not reaching into ecsoma. Uterus reaches posteriorly, just behind the junction of soma with ecsoma, passes forward coiling between ovary and dorsal to testes and seminal vesicle, joining the sinus sac in the post-acetabular region. Eggs are oval, small and numerous. Excretory pore is terminal, excretory vesicle is tubular.

100 Principle measurements of L. magnasoma n.sp. (In millimeters)

Body size: 7.1–7.13 x 0.69–0.7

Soma length: 5.30–5.33

Soma width: 0.70–0.72

Ecsoma length: 1.8–1.83

Ecsoma width: 0.40–0.41

Forebody length: 3.64–3.66

Hindbody length: 4.8–4.82

Oral sucker: 0.53–0.54 x 0.59–0.61

Pharynx: 0.62–0.65 x 0.30–0.35

Ventral sucker: 0.42–0.45 x 0.48–0.49

Sucker width ratio: 1: 0.803–0.813

Seminal vesicle: 0.15–0.17

Ventral sucker to seminal vesicle: 0.6–0.61

Ventral sucker to anterior testis: 1.02–1.023

Anterior testis: 0.19–0.20 x 0.21–0.22

Posterior testis: 0.20–0.205 x 0.21–0.22

Posterior testis to ovary: 0.28–0.29

Ovary: 0.172–0.180

Eggs: 0.022–0.05 x 0.020–0.025

101 Etymology: The species name Lecithocladium magnasoma refers to large anterior part of the body, the soma.

Remarks:

The species of the genus Lecithocladium Luhe, 1901, described previously from Pakistan are L. psenopsis Yamaguti, 1934 from

Stromateus sinensis; L. hexavitellarii (Bilqees, 1971); L. anteporus

(Bilqees, 1971); L. microductus (Bilqees, 1971); L. arabiana,

(Bilqees, 1971); L. microcaudum (Bilqees, 1971); L. octovitellarii

(Bilqees, 1971) all from Stromateus sp., of Karachi coast; L. karachii

Zaidi and Khan, 1977 from Carangoides presustus; L. pakistanensis

Zaidi and Khan, 1977 from Carangoides malabaricus; L. stromatei

Farooq and Khanum, 1980 from Stromateus niger; L. arabicum

Farooq and Khanum, 1980 from Caranx djedaba; L. thynensis

Bilqees and Nighat, 1985 from Thynnus sp., and L. arii Bilqees et al.,

2005 from the fish Arius serratus. These species are also from fishes of Karachi coast. All species except L. karachii, L. arabicum and L. thynensis are from fishes of the family Stromateidae.

Present species are different from the previously described species from Pakistan as well as species from other parts of the world. The present species is characterized by having cup-shaped,

102 infundibuliform oral sucker, large, rectangular pharynx, longer than oral sucker, ventral sucker smaller than oral sucker. Testes are two, tandem, close to each other, equal in size, seminal vesicle elongated, almost pointed anteriorly and rounded posteriorly, leading to pars prostatica. Pars prostatica is slightly coiled posteriorly, extending anteriorly from the anterior of the seminal vesicle, not reaching in the forebody, sinus sac tubular, joins the genital atrium laterally near the base of pharynx, genital pore at the posterior margin of oral sucker.

The present species is different from species having ventral sucker larger than oral sucker. These include L. karachii Zaidi and Khan,

1977; L. octovitellarii Bilqees, 1971 and L. pakistanensis Zaidi and

Khan, 1977.

The species having ventral sucker smaller than the oral sucker are L. psenopsis Yamaguti, 1934; L. hexavitellarii Bilqees, 1971; L. anteporus Bilqees, 1971; L. tetravitellarii Bilqees, 1971; L. microductus Bilqees, 1971; L. stromatei Farooq and Khanum, 1980;

L. arabicum Farooq and Khanum, 1980 and L. thynensis Bilqees and

Nighat, 1985, in this character the present specimens are close to the above mentioned species but is different in a combination of

103 characters such as sucker-width ratio, relative sizes of soma and ecsoma, position of testes and ovary, extension of vitellaria and uterus in the ecsoma.

The species having oral sucker smaller than ventral sucker are different from present species. These include L. magnacetabulum

Yamaguti, 1934; L. psenopsis Yamaguti, 1934; L. carultum Gupta &

Gupta, 1983; L. glandulum Chauhan, 1945; L. megalaspis Gibson &

Bray, 1986; L. seriolellae Gupta & Gupta, 1983; L. chingi Manter et

Pritchard, 1960; L. chauhani Chauhan, 1945; L. falklandicum Gupta

& Gupta, 1983; L. purense Gupta et Gupta, 1978; L. thapari Gupta &

Gupta, 1983; L. triacanthi Gupta & Gupta, 1983; L. puriensis Gupta

& Gupta, 1983; L. manteri Gupta and Puri, 1981; L. singhi Gupta et

Singh, 1983; L. guptai Gupta & Gupta, 1983; L. tewarii Gupta &

Gupta, 1987 and L. bengalensis Saxena and Prakash, 2006.

The species having oral sucker larger than ventral sucker are L. excisum Luhe, 1901; L. brevicaudum Srivastava, 1942; L. annulatum Velasquez, 1962; L. anguistiovum Gibson and Bray,

1986; L. parvioum Yamaguti, 1953; L. scombri Yamaguti, 1953; L. apolectis Velasquez, 1962; L. aegyptensis Fischthal and Kuntz, 1963;

104 L. stromatei Gupta & Gupta, 1983; L. arabicum Farooq & Khanum,

1980; L. inglisi Gupta et Ahmed, 1977; L. lutiani Gu et Shen, 1978;

L. keralense Gupta & Gupta, 1983; L. indicum Gupta & Gupta, 1978;

L. elongatus Gupta and Puri, 1981; L. bengalensis Gupta & Gupta,

1983; L. jagannathi Ahmed, 1981; L. sulpuriusi Gupta & Gupta,

1983; L. microlepidotus Gupta & Gupta, 1983; L. caranxi Gupta &

Gupta, 1983 and L. siddiqui Gupta & Gupta, 1978, as in the present species but other differences are quite prominent.

The present species is close to L. psenopsis Yamaguti, 1934 from

Stromateus sinensis, in gross morphology, in having oral sucker larger than ventral sucker, elongated seminal vesicle and broad base of ecsoma but differences are found in the posterior extent of uterus which does not enter into the ecsoma in the present species. Ovary in the present species is far posterior to testes. In L. psenopsis the anterior testis overlaps most part of the posterior testis and the seminal vesicle overlaps the anterior testis, while in the present species the anterior testis does not overlaps the posterior testis and the seminal vesicle is also separated. The above mentioned characteristic differences are sufficient to separate the present species from L. psenopsis and all other species of the genus.

105 In L. psenopsis the vitellaria are very short and do not reach the posterior end of the soma, while vitellaria are very long reaching near about posterior end of the soma in the present species.

L. gazzai Pandey et al., 2000 has oral sucker nearly equal to the ventral sucker prepharynx absent. Genital atrium is paramedian, ventral to anterior part of oral sucker. Ovary is tri-lobed, post- testicular lying little anterior to posterior end of soma. Receptaculum seminis indistinct. Vitellaria post-ovarian consisting of 10, long, slender winding tubules.

106 LECITHOCLADIUM MAGNAVESICULA N.SP. (Figs. 11-12)

Host: Pomadasys olivaceum (Pomadasyidae)

Location: Stomach

Locality: Karachi coast, Pakistan

No. of specimens: 10 specimens from 4 hosts, 39 fishes were examined.

Holotype: JUW – T11

Paratype: JUW – T12

Diagnosis

Body is elongated, smooth, widest at the base of soma. Ecsoma is slightly smaller and narrower than the soma. Tail is flattened. Oral sucker is larger than the ventral sucker. Prepharynx is absent. Pharynx is large, longer than the oral sucker. Oesophagus is absent, intestinal bifurcation immediately anterior to ventral sucker.

Testes are two, close, slightly, transversely oval, posterior to seminal vesicle. Seminal vesicle is large, elongated, bluntly spindle-shaped, close to ventral sucker. Pars prostatica is simple, short, lateral to ventral sucker. Sinus sac is a small tube in the forebody, joining the

107 genital atrium at the base of pharynx. Genital pore is at the base of oral sucker.

Ovary situated posteriorly, at the base of soma, rounded anteriorly, flat posteriorly. Vitellaria consist of 7, long tubules extending from the base of ovary to the posterior end of soma. Uterus reaches to about more than two-third of ecsoma then extends forward between ovary and testes and anterior to testes joining the sinus sac at the base of pharynx. Excretory vesicle is tubular, arms joining at the base of oral sucker anterior to genital opening.

Principle measurements of L. magnavesicula n.sp. (In millimeters)

Body size: 6.06–6.12 x 0.75–0.76

Soma length: 3.05–3.10

Soma width: 0.70–0.82

Ecsoma length: 3.01–3.03

Ecsoma width: 0.75–0.76

Forebody length: 1.03–1.036

Hindbody length: 4.04–4.06

Oral sucker: 0.59–0.6 x 0.82–0.85

Pharynx: 0.6–0.9 x 0.4–0.51

108

Ventral sucker: 0.49–0.50 x 0.54–0.55

Sucker width ratio: 1: 0.830–0.831

Seminal vesicle: 0.83–0.85 x 0.30–0.35

Ventral sucker to seminal vesicle: zero

Ventral sucker to anterior testis: 0.69–0.7

Anterior testis: 0.25–0.26 x 0.26–0.27

Posterior testis: 0.2–0.21 x 0.24–0.25

Posterior testis to ovary: 0.29–0.3

Ovary: 0.28–0.29 x 0.34–0.35

Eggs: 0.012–0.02 x 0.023–0.036

Etymology: The present new species Lecithocladium magnavesicula refers to its large-sized seminal vesicle.

Remarks

Several species of the genus Lecithocladium are known from

Pakistan. As compared to the Pakistani species, the present new species L. magnavesicula has oral sucker larger than ventral sucker as in L. psenopsis Yamaguti, 1934; L. hexavitellarii Bilqees, 1971; L. anteporus Bilqees, 1971; L. tetravitellarii Bilqees, 1971; L. microductus Bilqees, 1971; L. stromatei Farooq and Khanum, 1980;

109 L. arabicum Farooq and Khanum, 1980 and L. thynensis Bilqees and

Nighat, 1985.

The other differential characters are as follows: in L. psenopsis ecsoma is small. Soma is very long. Plications are in ecsoma region only. Seminal vesicle is overlapping the anterior testis, far posterior to ventral sucker. Ovary is post-testicular, contiguous with posterior testis. Uterus does not reach into ecsoma. Vitellaria are 7 tubules, pars prostatica is in hindbody.

In L. hexavitellarii soma is long, ecsoma small, seminal vesicle contiguous with anterior testis, lying far posterior to the ventral sucker. Ovary is contiguous with posterior testis, uterus reaches posteriorly into anterior-third of ecsoma, vitellaria consist of 6 tubules reaching in the ecsoma.

In L. tetravitellarii soma is short and stout, curved ventrally. Ecsoma is long. Plications are in anterior half of ecsoma. Pharynx is elongated, cylindrical and flat reaching dorsal to ventral sucker. Seminal vesicle is much posterior to ventral sucker. Pars prostatica is long, coiled.

Sinus sac is small, swollen. Genital pore is median, ventral to anterior border of oral sucker. Ovary is contiguous with posterior testis, larger than testes. Vitellaria are 4, long, coiled tubules of equal length.

110

In L. microductus soma is stout, curved ventrally. Plications are between hindbody to near middle of ecsoma, absent in posterior region of ecsoma. Ecsoma short with broadly, pointed posterior extremity. Testes flattened, contiguous. Seminal vesicle is thin- walled, bulbous, slightly overlapping anterior border of anterior testis not reaching the ventral sucker and far from it. Pars prostatica is long, coiled, sinuous. Sinus sac is bulb-like. Genital pore is median, ventral to posterior part of oral sucker. Ovary is in the posterior half of soma.

Uterus does not reach into ecsoma. Vitellaria consist of 4 tubules.

L. arabiana has soma ventrally curved. Plications are prominent in the hindbody. Oesophagus is present. Seminal vesicle is not reaching the ventral sucker. Pars prostatica bilobed, overlapping the seminal vesicle ventrally, not reaching into forebody. Uterus not reaching into ecsoma and vitellaria consist of 4-6 long tubules.

In L. anteporus soma is long. Plications are in the region of ecsoma.

Seminal vesicle is far posterior to the ventral sucker. Pars prostatica is saccular, bilobed, composed of masses of cells, male and female ducts run separately for a long distance, anterior to ventral sucker meeting behind oral sucker in a short sinus sac. Genital pore is large, ventral to

111 anterior margin of the oral sucker. Vitellaria consist of 6 convoluted tubules, slightly reaching ecsoma.

In L. stromatei, soma is cylindrical. Ecsoma is smaller than soma.

Oral sucker is larger than the ventral sucker. Oesophagus is short.

Testes two, transversely elongated. Seminal vesicle is elongated, narrow, thin-walled, overlapping anterior testis, not reaching to ventral sucker, anterior extremity recurved back over proximal part.

Sinus sac extends into forebody. Genital atrium is long and narrow.

Genital pore is median, ventral to middle of oral sucker. Vitellaria consist of 7 tubules not extending into ecsoma. Excretory pore is terminal, arms are not prominent.

In L. microcaudum soma is elongated, smooth and delicate. Ecsoma is small, one-fourth of the soma length. Seminal vesicle is large, pre- testicular, and separated, at a distance posterior to ventral sucker, thin- walled, anteriorly leading to pars prostatica. Pars prostatica is elongated, slightly overlapping ventral sucker. Genital atrium is tubular. Genital opening is ventral to oral opening. Ovary is rounded, separated from testes and larger than these. Uterus does not reach ecsoma, passes between the vitelline tubules, then dorsal to ovary and

112 testes, overlapping the seminal vesicle, joining the sinus sac at the anterior level of ventral sucker.

In L. thynensis soma and ecsoma both are elongated, with plications over the complete surface of soma. Ecsoma ventrally curved with prominent dorsal ala. Seminal vesicle is far behind the ventral sucker, smaller than the testes. Sinus sac is tubular. Genital atrium is also long. Junction of sinus sac and genital atrium is at mid-level of pharynx. Vitellaria consist of 7 tubules radiating from the base of ovary.

In L. arii soma and ecsoma are long, almost equal in size. Plications are in the hindbody. Pars prostatica is long, curved into 3-4 loops.

Genital atrium is long, narrow. Genital pore is ventral to anterior part of oral sucker.

The species which have ventral sucker larger than oral sucker are L. karachii Zaidi and Khan, 1977; L. octovitellarii Bilqees, 1971 and L. pakistanensis Zaidi and Khan, 1977 these are different from the present species and also different in the above mentioned diagnostic features. In L. karachii, uterus does not reach into ecsoma and vitellaria consist of 8 tubules. In L. pakistanensis vitellaria also

113 consist of 8 tubules and uterus does not reach into ecsoma. L. octovitellarii also has 8 vitelline tubules but uterus is extending into ecsoma and vitelline tubules terminate anteriorly in the soma. While in the present new species seminal vesicle is very large reaching to the base of ventral sucker, pars prostatica very small at the side of ventral sucker, ovary consist of 16 separate follicles at the base of soma and the vitelline tubules are 7, completely located in the ecsoma, uterus also reaches into three-fourth of ecsoma. These characteristics separate the present species from the species described previously from Pakistan as well as from other parts of the world.

The species from India and other localities which have ventral sucker smaller than oral sucker, including L. glandulum Chauhan, 1945; L. lutjanii Gu et shen, 1978; L. dawesi Bashirullah et D’Silva, 1973; L. hanumanthi Gupta & Jain, 1992; L. inglisi Gupta et Ahmad, 1977; L. purense Gupta et Gupta, 1978 and L. singhi Gupta et Singh, 1983 are also different from the present specimens. Bray and Cribb (2004) regarded L. dawesi and L. hanumanthi as synonym of L. megalaspis.

While L. inglisi; L. purense and L. singhi are also the synonym of L. megalaspis (Gibson & Bray, 1986). Gupta & Gupta also suggested the following synonym: L. harpodontis Srivastava, 1942 with L. exisum

Luhe, 1901; L. psettodi Srivastava, 1942 with L. manteri Gupta &

114 Puri, 1981; L. sciaenid Gupta & Puri, 1981 with L. elongatus Gupta

& Puri, 1981; and L. srivastavai Gupta & Puri, 1981; L. stromateusi

Gupta & Puri, 1981 with L. aegyptensis Fischthal & Kuntz, 1963 and

L. fotedari Gupta & Singh, 1982 with L. jagannathi Ahmad, 1981.

All these species are different from the present species in a combination of characters such as sucker-width ratio, egg sizes, soma and ecsoma relative sizes and relative sizes of suckers and gonad, seminal vesicle large, close to ventral sucker, ovary at the base of soma and vitelline tubules in the ecsoma.

Four species have sub-globular oral sucker and pharynx including L. chingi Manter and Pritchard, 1960; L. ilishae Mamaev, 1970 nec

Bashirullah and D’Silva, 1973, L. ilishae Bashirullah and D’Silva,

1973 nec Mamaev, 1970 and L. invasor Chambers et al., 2001.

Therefore, these are different from the present species and are from different fish hosts and localities. L. aegyptense Fischthal and Kuntz,

1963; L. harpodontis Srivastava, 1942 although have an infundibuliform oral sucker, an elongated pharynx and relatively small ventral sucker, are also different from the present species.

115 LECITHOCLADIUM LATEROPHARYNGIUM N.SP. (Figs. 13-14)

Host: Stromateus sinensis (Stromateidae)

Location: Stomach

Locality: West Wharf, Karachi coast, Pakistan.

No. of specimens: 4 specimens from 2 hosts, 20 fishes were examined.

Holotype: JUW – T13

Paratype: JUW – T14

Diagnosis

Body is elongated. Soma is very long, widest at the region of acetabulum. Ecsoma is very short. Oral sucker is larger than the ventral sucker. Pharynx is elongated, situated on the lateral side of ventral sucker. Ventral sucker is globular.

Testes are two, diagonal, situated in the middle of the body. Seminal vesicle elongated, close to the testes, anteriorly leading to pars prostatica, which runs from the lateral side of the ventral sucker to its anterior margin. Sinus sac is short, opening into the genital atrium in the middle of the oral sucker.

116 Ovary is post-testicular, very close to the testes almost in posterior- third quarter of the soma. Seminal receptacle is immediately post- ovarian. Vitellaria consist of 7, long unequal tubules not extending into ecsoma and not even reaching to the base of soma. Excretory vesicle is tubular, reaching anterior to ovary, arms are uniting at the middle of oral sucker.

Principle measurements of L. lateropharyngium n.sp. (In millimeters)

Body size: 3.58–3.60 x 0.55–0.56

Soma length: 2.77–2.78

Soma width: 0.60–0.61

Ecsoma length: 0.8–0.81

Ecsoma width: 0.21–0.22

Forebody length: 0.79–0.80

Hindbody length: 2.65–2.67

Oral sucker: 0.48–0.6 x 0.82–0.85

Pharynx: 0.6–0.9 x 0.4–0.51

Ventral sucker: 0.49–0.495 x 0.42–0.43

Sucker width ratio: 1: 0.830–0.84

Seminal vesicle: 0.45–0.46 x 0.14–0.15

Ventral sucker to seminal vesicle: 0.20–0.21

117

Ventral sucker to anterior testis: 0.50–0.51

Anterior testis: 0.20–0.21 x 0.21–0.215

Posterior testis: 0.185–0.190 x 0.183–0.185

Posterior testis to ovary: Negligible

Ovary: 0.10–0.11 x 0.20–0.21

Eggs: 0.017–0.023 x 0.01–0.011

Etymology: The present new species Lecithocladium lateropharyngium refers to the presence of pharynx on the lateral side of the body.

Remarks

There are thirteen species of the genus reported from fishes of Karachi coast. The present species can be distinguished from all other species of the genus Lecithocladium Luhe, 1901 in having pharynx lateral to ventral sucker. Ventral sucker very close to oral sucker and is smaller than the oral sucker.

Seminal vesicle and pars prostatica posterior to ventral sucker, genital opening ventral to middle of oral sucker or slightly anterior to it.

Vitellaria and uterus terminate much anterior to base of ecsoma. Tail is short, seminal vesicle, testes and ovaries are close. Vitellaria consist

118 of 7 convoluted tubules, while in all other species pharynx is anterior to ventral sucker. In the present species the oral sucker is pear-shaped and much larger than ventral sucker as in L. psenopsis Yamaguti,

1934 from Stromateus sinensis; L. hexavitellarii (Bilqees, 1971;

Bilqees, 1981); L. anteporus (Bilqees, 1971; Bilqees, 1981); L. microductus (Bilqees, 1971; Bilqees, 1981); L.arabiana (Bilqees,

1971; Bilqees, 1981); L. microcaudum (Bilqees, 1971; Bilqees,

1981); L. octovitellarii (Bilqees, 1971; Bilqees, 1981) all from

Stromateus sp., of Karachi coast; L. karachii Zaidi and Khan, 1977 from Carangoides presustus; L. pakistanensis Zaidi and Khan, 1977 from Carangoides malabaricus; L. stromatei Farooq and Khanum,

1980 from Stromateus niger; L. arabicum Farooq and Khanum, 1980 from Caranx djedaba; L. thynensis Bilqees and Nighat, 1985 from

Thynnus sp. and L.arii Bilqees et al., 2005 from the fish Arius serratus.

Species having oral sucker larger than the ventral sucker are L. exisum Luhe, 1901; L. brevicaudum Srivastava, 1942; L. annulatum

Velasquez, 1962; L. anguistiovum Gibson and Bray, 1986; L. parvioum Yamaguti, 1953; L. scombri Yamaguti, 1953; L. apolectis

Velasquez, 1962; L. aegyptensis Fischthal and Kuntz, 1963; L.

119 bulbolabrum Reid, Coil and Kuntz, 1966; L. stromatei Gupta and

Gupta, 1983; L. arabicum Farooq et Khanum, 1980; L. inglisi Gupta et Ahmed, 1977; L. lutiani Gu et Shen, 1978; L. keralense Gupta and

Gupta, 1983; L. indicum Gupta and Gupta, 1978; L. elongatus Gupta and Puri, 1981; L. bengalensis Gupta and Gupta, 1983; L. jagannathi

Ahmed, 1981; L. sulphuriusi Gupta and Gupta, 1983; L. microlepidotus Gupta and Gupta, 1983; L. caranxi Gupta and Gupta,

1983; L. siddiqui Gupta and Gupta, 1978 as in the present species but these are different in body size, relative length of soma and ecsoma and shape, size of seminal vesicle and ovary and vitellaria and uterus terminating much anterior to the base of soma.

In most species uterus reaches into ecsoma, while it terminates much anterior to base of ecsoma. The present species is different from other species in having ventral sucker larger than oral sucker as in L. karachii Zaidi and Khan, 1977; L. octovitelarii Bilqees, 1971 and L. pakistanensis Zaidi and Khan, 1977.

Species having oral sucker smaller or equal to ventral sucker are L. magnacetabulum Yamaguti, 1934; L. carultum Gupta and Gupta,

1983; L. megalaspis Gibson and Bray, 1986; L. chauhani Chauhan,

120 1945; L. triacanthi Gupta and Gupta, 1983; L. manteri Gupta and

Puri, 1981; L. psenopsis Gupta and Gupta, 1983; L. glandulum

Chauhan, 1945; L. chingi Manter and Pritchard, 1960; L. falklandicum Gupta and Gupta, 1983; L. purense Gupta and Gupta,

1978; L. thapari Gupta and Gupta, 1983; L. puriensis Gupta and

Gupta, 1983; L. singhi Gupta et Singh, 1983; L. guptai Gupta and

Gupta, 1983; L. seriolellae Gupta and Gupta, 1983; L. tewarii Gupta and Gupta, 1978 are different from the present species in the above mentioned diagnostic features.

Previously the species described from Pakistan are mostly from family Stromateidae. The present species is also found in the fish

Parastromateus sinensis of the same family. The present species is separated from the above mentioned species in the main diagnostic feature as mentioned above. Therefore, a new species is proposed and the name Lecithocladium lateropharyngium is given.

The new species of Lecithocladium reported here differ from L. gazzai Pandey et al., 2000 in having oral sucker larger than the ventral sucker while in L. gazzai the oral sucker is almost equal to the ventral sucker.

121

Genital pore in L. gazzai is median, ventral to the oral sucker. Seminal vesicle is spindle-shaped, small as compared to the present new species. Ovary is median, tri-lobed, post-testicular lying a little anterior to the posterior end of soma separated from posterior testis by uterine coils.

Vitellaria post-ovarian, consisting of 10, long, slender, winding tubules confined up to a little anterior to posterior end of soma, while all the new species described here consist of 7 vitelline tubules. L. gazzai resembles with L. chaetodipten in having oral sucker smaller or same size as ventral sucker and the presence of 10 vitelline tubules but differs from it in not having pharynx overlapping anterior margin of ventral sucker, seminal vesicle is not extending to posterior margin of ventral sucker and large size of body.

L. gazzai differs from L. cybiumi and L. hanumanthai in having genital pore at anterior part of oral sucker, tri-lobed ovary, in the presence of pre-oral lobe and 10 vitellaria.

122 L. bengalensis Saxena and Prakash, 2006 differs from previously described species in having genital pore sub-median, lying in the middle of the oral sucker while resembles with other Lecithocladium species in having same number of vitelline tubules i.e. 7.

Thus on account of the above mentioned differences as against all those described earlier, all the present specimens of Lecithocladium deserves the status of new species.

123 SPECIES OF THE GENUS TUBULOVESICULA TUBULOVESICULA OLIVACEUS N.SP. (Figs. 15-16)

Family: Hemiuridae Looss, 1899

Sub-family: Diurinae Looss, 1907 (Syn. Stomachicolinae Yamaguti, 1958)

Genus: Tubulovesicula Yamaguti, 1934 Syn. Lecithurus Pigulewsky, 1938

Host: Pomadasys olivaceum (Pomadasyidae)

Location: Stomach

Locality: Karachi coast, Pakistan

No. of specimens: 4 specimens from 2 hosts, 15 fishes were examined.

Holotype: JUW – T15

Paratype: JUW – T16

Diagnosis

Body long, tail relatively small, rounded posteriorly, greatest width is at the region of ventral sucker. Lateral cuticular thickenings present near posterior end of soma, more prominent at the right side. Oral sucker is sub-terminal, pre-oral lobe present, prepharynx absent, pharynx small, intestinal caeca radiating directly from its base, oesophagus is absent. Intestinal caeca long, reaching near about the

124 posterior end of ecsoma. Ventral sucker is very large as compared to oral sucker, situated anteriorly in anterior-quarter of the body. Testes are unequal, rounded, symmetrical, immediately behind ventral sucker, left testis smaller, close to ventral sucker, seminal vesicle pre- testicular, long, distinctly marked off posteriorly, becoming tubular anteriorly. Pars prostatica small, hermaphroditic duct enclosed in muscular pouch, genital opening immediately behind intestinal bifurcation. Ovary is almost rounded in the middle of soma, vitelline tubules coiled, 7, radiating posteriorly, far anterior to ecsoma then extending forward in the forebody joining the hermaphroditic duct.

Uterus not reaching the tail terminating far anterior to ecsoma. Eggs numerous, small. Excretory pore terminal, excretory vesicle tubular, arms uniting dorsally at the base of pharynx.

Principle measurements of T. olivaceus n.sp. (In millimeters)

Entire body size: 3.8–3.9 x 0.62–0.625

Soma length: 3.0–3.05

Soma width: 0.65–0.66

Ecsoma length: 0.89–0.90

Ecsoma width: 0.36–0.37

Forebody length: 0.49–0.50

125

Hindbody length: 3.09–3.10

Oral sucker: 0.11–0.12 x 0.14–0.145

Pharynx: 0.05–0.055 x 0.043–0.044

Ventral sucker: 0.35–0.355 x 0.38–0.385

Sucker width ratio: 1: 0.03–0.031

Pars prostatica length: 0.19–0.20

Seminal vesicle length: 0.45–0.46

Anterior testis: 0.08–0.09 x 0.078–0.80

Posterior testis: 0.13–0.14 x 0.10–0.11

Posterior testis to ovary: 0.5–0.51

Ovary: 0.25–0.24 x 0.21–0.25

Eggs: 0.022–0.04 x 0.017–0.02

Etymology: The present new species Tubulovesicula olivaceus refers to its host.

Remarks

Species of the genus Tubulovesicula Yamaguti, 1934 reported previously from Karachi coast, Pakistan, are T. spari Yamaguti, 1934;

(Bilqees, 1981); Zaidi and Khan, 1977 from Muraenesox cinereus

(Forsk) (Congridae); T. anguillae Yamaguti, 1934; (Zaidi and Khan,

126 1977) from Harpodon nehereus (Har) (Synodidae); T. magna Bilqees and Nighat, 1981 from Pomadasys olivaceum (Day) (Pomadasyidae) and T. anguisticauda Nicoll, 1914; Yamaguti, 1934; (Bhutta and

Khan, 1975); Bilqees, 1981 from Muraenesox cinereus (Forsk)

(Muraenesocidae).

In the present species T. olivaceus n.sp., pars prostatica is smaller than the seminal vesicle, ecsoma is much smaller than the soma and vitelline tubules are 7 in number and uterus does not extend to ecsoma.

Manter (1954) has shown that number of vitelline tubules in a species of Tubulovesicula may be either 7 or 8 arranged in various positions.

T. serrani is similar to T. anguillae Yamaguti, 1934 which has a very long pars prostatica; in T. anguillae it is much longer than the seminal vesicle; the uterus extends rather far into the ecsoma and eggs are

0.034 x 0.023–0.024. T. serrani differs from T. anguisticauda

(Nicoll, 1914) in having much longer ecsoma and more posterior extent of the pars prostatica.

Position of pars prostatica in T. pseudorhombi and T. anguisticauda is anterior to the ventral sucker, very similar to the present specimen; while in T. spari and T. anguillae the pars prostatica is well

127 developed and extends from the middle of the acetabulum to near the internal bifurcation. In T. serrani the length of pars prostatica appears to be slightly less, than the length of seminal vesicle and hermaphroditic duct.

In T. pseudorhombi seminal vesicle is moderately long in close spirals on dorsal side of posterior half of acetabulum; while in T. spari seminal vesicle is broad at the base, runs sinuously on the postero- dorsal side of the ventral sucker. In T. anguillae the position of seminal vesicle is different. It is S-shaped lying in front of right testis.

In T. magna and T. anguisticauda the seminal vesicle is tubular, more or less winding, pre-testicular in position. In T. serrani the seminal vesicle is sinuous, dorsal to posterior half of acetabulum and is slightly larger in length than the pars prostatica, while in the present specimen T. olivaceus n.sp., the seminal vesicle is long, pre-testicular, distinctly marked off posteriorly.

In T. serrani the ovary is ovoid, almost twice as large as one testis to right mid-line, posterior to right testis and overlapped dorsally by the right caecum. The position of ovary in T. pseudorhombi, T. spari, T. anguillae, T. anguisticauda and T. serrani is compared, the testes

128 and the ovary is close to the ventral sucker. But in the present species

T. olivaceus n. sp. the ovary is far posterior from the ventral sucker.

In T. pseudorhombi, T. magna and T. serrani the uterus does not reach in the ecsoma as in the present species; while in T. spari the position is entirely different. Here the uterus coils down on the tail where it turns to the right and passes forward on the ventral side of the right caecum. In T. anguillae the tail region is a little longer than the body proper so the uterine coils extend into the tail as far as its middle. In T. anguisticauda the uterus is extensive and occupies a major portion of the body space between the ventral sucker to the proximal part of the tail.

T. magnacetabulum Yamaguti, 1939 is extraordinary different from all the other species in having four times large acetabulum than the oral sucker. Pars prostatica is very long as in T. anguillae. Seminal vesicle is posterior to ventral sucker. One of the two testes is contagious with the ovary which is at a negligible distance to the acetabulum. Uterus is extending into the anterior part of the tail region.

129 T. pseudorhombi Yamaguti, 1938 in Pseudorhombus pentopthalmus; regarded by Manter (1954), as a syn., of T. spari despite difference in egg size.

The present new species, therefore, is distinctly different from the previously described species of the genus in having ventral sucker at least three times larger than the oral sucker for which a new species T. olivaceus is proposed.

130 TUBULOVESICULA MICROCAUDUM N.SP. (Figs. 17-18)

Host: Otolithus argenteus (Sciaenidae)

Location: Stomach

Locality: Karachi coast, Pakistan

No. of specimens: 2 specimens from a single host, 19 fishes were examined.

Holotype: JUW – T17

Paratype: JUW – T18

Diagnosis

Body is spindle-shaped, differentiated into soma and ecsoma. Soma is very long covering almost the entire length. Ecsoma is very small, extensible to varying degrees. Oral sucker is sub-terminal, very small as compared to ventral sucker. Pharynx is tubular in size, adjacent to the oral sucker, pre-pharynx absent. Intestinal caeca long reaching the posterior end of ecsoma, ventral sucker about one-fourth of body length from anterior end. Small genital atrium is followed by large- sized genital pore.

Testes are unequal, rounded, symmetrical, away from the ventral sucker. Left testis is little larger than the right lying at its posterior

131 border, right separate. Seminal vesicle pre-testicular, long, extending posterior to ventral sucker and situated dorsal to it, distinctly marked off curved dorsally, becoming tubular anteriorly pars prostatica very long extending from dorso-lateral to ventral sucker to the muscular pouch, hermaphroditic duct enclosed in muscular pouch leading into prominent cirrus sac. Genital pore is immediately below the intestinal bifurcation.

Ovary is oval, kidney-shaped in the middle of soma, vitellaria consist of 7, long, tubular lobes, 3 on one side and 4 on the other, lying immediately behind ovary, mostly directed backward. The uterus passes in close coils, slightly intruding anterior of ecsoma and then passes forward between the testes above or beside the acetabulum and pre-acetabular area. Eggs are thick-shelled and of moderate size.

Excretory pore is terminal.

Principle measurements of T. microcaudum n.sp. (In millimeters)

Body size: 5.4–5.42 x 1.11–1.15

Soma length: 4.4–4.42

Soma width: 1.2–1.21

Ecsoma length: 1.3–1.32

132 Ecsoma width: 0.5–0.51

Forebody length: 0.8–0.83

Hindbody length: 3.1–3.13

Oral sucker: 0.21–0.22 x 0.27–0.28

Pharynx: 0.09–0.10 x 0.10–0.105

Ventral sucker: 0.56–0.57 x 0.56–0.561

Sucker width ratio: 1: 0.035–0.037

Pars prostatica: 1.05–1.10

Seminal vesicle: 0.41–0.42

Ventral sucker to seminal vesicle: 0.14–0.15

Ventral sucker to anterior testis: 0.1

Anterior testis: 0.1–0.16

Posterior testis to ovary: 0.39–0.4

Ovary: 0.15–0.20

Eggs: 0.029–0.05 x 0.03–0.038

Etymology: The present new species Tubulovesicula microcaudum due to its short tail.

133 Remarks

Tubulovesicula is a poorly known genus in Pakistan and only four species have been reported from Marine Fishes. These include

Tubulovesicula spari Yamaguti, 1934; (Bilqees, 1981) from the fish

Muraenesox cinereus; T. magna Bilqees and Nighat, 1981 from

Pomadasys olivaceum; T. anguillae Yamaguti, 1934; (Zaidi and

Khan, 1977) from Harpodon nehereus and T. anguisticauda (Nicoll,

1914); Yamaguti, 1934; Bhutta and Khan, 1975 from Muraenesox cinereus.

During the present studies species of the genus Tubulovesicula has been recovered from the intestine of Otolithus argenteus which appears totally different from the species described from Pakistan and other parts of the world and is regarded a new species and the name T. microcaudum n.sp. is proposed.

The new species is different from the species found in Pakistan in having relatively small-sized body with soma very long and ecsoma very small, curved tail. Seminal vesicle is pre-testicular, long, extending posterior to ventral sucker. Hermaphroditic duct enclosed in muscular pouch. Uterus occupy extensive part of soma, slightly

134 intruding into the ecsoma, while in T. anguisticauda the tail is very long, tubular seminal vesicle is not enclosed within muscular pouch, posteriorly it extends upto the posterior margin of the ventral sucker, uterus is extensive and occupies a major portion of the body space between the ventral sucker to the proximal part of the tail. Excretory vesicle is Y-shaped.

In T. spari body is spindle-shaped, fusiform, smaller than the present species, broadest at the level of vitellaria. Seminal vesicle is long and slender, as in the present specimen T. microcaudum n.sp., broad at the base, runs sinuously on the postero-dorsal side of the acetabulum.

Pars prostatica is surrounded by numerous prostatic cells.

Hermaphroditic pouch is pear-shaped and muscular. Uterus coils down on the left side half way into the tail which shows that the T. spari is different from the present specimen.

The diagnostic characters of T. anguillae are very different from the specimens of T. microcaudum n.sp. The body is equally divided into soma and ecsoma in which uterus reaches the middle of the ecsoma.

Seminal vesicle is S-shaped lying in front of the right testis in T. anguillae.

135 In T. magna the tail is knob-like, more or less longer than the present specimen in which the tail is straight and curved. Seminal vesicle is tubular, more or less winding, pre-testicular in position. Testes are almost symmetrical in position. Uterus is not reaching the ecsoma.

While in the present specimen the position of the testes and ovary are different and uterus does not reach the ecsoma.

Species from other parts of the world include T. muraenesocis

Yamaguti, 1934 which is closely related to T. anguisticauda Nicoll,

1915 but differs markedly in the size of the eggs and in the posterior extent of the vesicula seminalis and pars prostatica. The uterus is confined to body proper and the tail region is very long, which shows that this specimen is fairly different from the present specimen.

T. pseudorhombi Yamaguti, 1938 is similar to the present specimen in having short tail but other diagnostic features are very different. In

T. pseudorhombi, the body is fusiform, pointed at both extremities, very broad in the middle of ecsoma. Tail is short and straight. Seminal vesicle is long, spirally coiled, while in T. microcaudum n.sp., soma is much longer and tubular as compared to T. pseudorhombi. Tail is short and curved. Seminal vesicle is tubular, curved dorsally. In the

136 present specimen, uterus slightly enters the ecsoma, while in T. pseudorhombi the uterus terminates a little above the soma.

T. lindbergi (Layman, 1930) Yamaguti, 1934 (is the synonym of T. spari as reported by McCauley, 1960) is different from the present specimen in having anterior end broadly rounded, having prominent ecsoma. Prepharynx present (absent in the present specimen). Seminal vesicle is tubular, sinuous at about the level of ventral sucker.

Prostatic vesicle tubular, long, with some convolutions, partially or completely surrounded with prostatic gland cells, its posterior end usually, though not always within the zone occupied by ventral sucker. Ovary close to the posterior testis, while in the present species ovary is far away from the ventral sucker and posterior testis. Uterus with descending and ascending arms much coiled. Excretory bladder is Y-shaped.

T. pinguis (Linton, 1940) Manter, 1947 is also different from the present specimen in having cervical glands into its anterior portion and posteriorly it communicates with two lateral pouches, termed

‘stomachs’. The stomachs are slightly larger than the oesophagus. The caeca are lined with tall epithelium and extend the length of the body, terminating blindly near the posterior end. Testes are large, close to

137 ventral sucker, seminal vesicle reaching near about the anterior testis.

Vitellaria are not extending to the ecsoma, uterus slightly protruding in the ecsoma. Ovary and vitellaria are close to the testes. While in the present species there is no stomach. Testes are small, seminal vesicle long, terminating at the anterior level of the left testis. Ovary and vitellaria are not close to the testes.

The other species namely T. madurensis Nigrelli, 1940; T. magnacetabulum Yamaguti, 1939; T. marsupialia Oshmarin, 1965;

T. nanamoensis Mc Farlane, 1936; T. californica Park, 1936; T. diacopae Nagaty et Abdel-Aal, 1962 and T. serrani Nagaty, 1956 are also different from the present species showing differences in the important diagnostic features.

In Tubulovesicula the degree to which prostate gland cells enclose the prostatic vesicle appears to be a function of maturity and not a taxonomic character as suggested by McCauley, 1960.

The new species described above is T. olivaceus which is different from the present species T. microcaudum n.sp., in having long ecsoma and longer tail as compared to T. microcaudum, in which the tail is very short and curved. The ventral sucker in T. olivaceus n.sp., is larger than the ventral sucker of T. microcaudum n.sp. Seminal

138 vesicle in T. olivaceus n.sp., is broader than long while in T. microcaudum n.sp., it is longer than broad. The two testes are far away from the oral sucker in T. microcaudum n.sp., while in T. olivaceus n.sp. they are close to the posterior border of the ventral sucker. In T. microcaudum n.sp. cirrus sac is larger in size as compared to T. olivaceus n.sp. Ovary is kidney-shaped with moderate-sized eggs in T. microcaudum n.sp., while in T. olivaceus n.sp., the ovary is globular and eggs are small-sized. Uterus is long, terminating far anterior to ecsoma in T. olivaceus n.sp., while in T. microcaudum n.sp. it passes in close coils slightly intruding anterior of ecsoma.

139

TUBULOVESICULA MAGNACIRROSA N.SP. (Figs. 19-20)

Host: Pseudosciaena diacanthus (Sciaenidae)

Location: Stomach

Locality: Karachi coast, Pakistan

No. of specimens: 3 specimens from a single host, 17 fishes were examined.

Holotype: JUW – T19

Paratype: JUW – T20

Diagnosis

Body is spindle-shaped, differentiated into soma and ecsoma. Soma is very long, ecsoma small. Oral sucker is terminal, small as compared to ventral sucker. Pharynx is globular, adjacent to the oral sucker.

Prepharynx is absent. Intestinal caeca are long, reaching to the posterior end of ecsoma. Ventral sucker is large situated in the anterior-fourth of the body.

Testes are two, small, almost equal in size, far behind the ventral sucker. Seminal vesicle is tubular, extending posteriorly to the posterior margin of ventral sucker. Pars prostatica is long, with

140 numerous prostatic cells. Sinus sac is pressed out, it is large, globular.

Genital opening is pre-acetabular.

Ovary is at a considerable distance posterior to testes. Vitellaria consist of 7 tubules radiating posteriorly from the ovary, post- equatorial in soma and far anterior to ecsoma. Uterus is occupying most part of the body not reaching into the ecsoma and passes forward beside the acetabulum and pre-acetabular area joining the hermaphroditic duct. Excretory pore is terminal.

Principle measurements of T. magnacirrosa n.sp. (In millimeters)

Body size: 9.3–9.35 x 1.4–1.42

Soma length: 6.3–6.5

Soma width: 1.40–1.42

Ecsoma length: 2.8–3.0

Ecsoma width: 0.41–0.42

Forebody: 1.54–1.55 x 0.87–0.88

Hindbody length: 7.2–7.25

Oral sucker: 0.4–0.41 x 0.38–0.381

Pharynx: 0.10–0.11 x 0.20–0.21

Ventral sucker: 0.78–0.79 x 0.54–0.541

141

Sucker width ratio: 1: 0.051–0.053

Pars prostatica: 1.11–1.30 x 0.37–0.38

Seminal vesicle: 0.20–0.21 x 0.21–0.22

Ventral sucker to seminal vesicle: Negligible

Ventral sucker to anterior testis: 0.55–0.65

Anterior testis: 0.2–0.24 x 0.20–0.22

Posterior testis to ovary: 0.50–0.51

Number of vitelline tubules: 7

Ovary: 0.240–0.241 x 0.22–0.23

Eggs: 0.051–0.061 x 0.051–0.080

Etymology: The name given to the new species T. magnacirrosa is due to the large cirrus sac.

Remarks

The present species is characterized by having a long soma, short and narrow ecsoma and sucker-width ratio. Uterus is not reaching into ecsoma. Ovary situated almost in the middle of the soma from ecsoma. Vitellaria are far anterior to ecsoma. Testes are almost at equal distance between ovary and ventral sucker. Seminal vesicle is long, tubular, reaching posteriorly at the level of base of ventral

142 sucker. Pars prostatica large, containing numerous prostatic cells.

Hermaphroditic duct is small, sinus sac or sinus pouch is large, rounded anteriorly. Genital opening is almost at equal distance of intestinal bifurcation and ventral sucker.

The above mentioned two species namely T. olivaceus n.sp. and T. microcaudum n.sp., are also different from each other in having different position of testes, ovary and morphology of sinus sac.

The position of testes in T. olivaceus n.sp., is close to the ventral sucker. In T. microcaudum n.sp., it is away from the ventral sucker, while in the present specimen T. magnacirrosa n.sp., the position of testes is far behind the ventral sucker.

Ovary is kidney-shaped in the middle of the soma in T. microcaudum n.sp. In T. olivaceus n.sp., it is in the middle of the soma, while in T. magnacirrosa n.sp., the ovary is at a considerable distance, posterior to testes.

In T. microcaudum n.sp., the sinus sac is elongated, tubular, dorso- ventral to the ventral sucker, pyriform with strong muscular sheath, traversed by hermaphroditic duct formed by the junction of prostatic vesicle and metraterm at base of sinus sac. As compared to T.

143 microcaudum n.sp., the cirrus sac in T. olivaceus n.sp. is different showing small, tubular sinus sac, hermaphroditic duct enclosed in muscular pouch, ventrally at the anterior part of the ventral sucker, finally reaching the large genital opening which is immediately behind the intestinal bifurcation. In T. magnacirrosa n.sp., the sinus sac is entirely different, it is large and pressed out of the body, showing globular shape anteriorly.

The Pakistani species and species found in different parts of the world are compared with the present specimen of T. magnacirrosa n.sp. T. spari is different in having small body. In T. anguillae the body is equally divided into soma and ecsoma. In T. pseudorhombi the body is fusiform, pointed at both ends, very broad in the middle of ecsoma.

Tail is short and straight. T. magna has knob-like tail. In T. lindbergi the anterior end is broadly rounded while in the present specimen T. magnacirrosa the body is very long and spindle-shaped. Soma covering almost entire length. In T. serrani Nagaty, 1954 the body is very long, slender, ecsoma is equal in length to that of body proper, about half of ecsoma extended. The above length of the specimen is seen in T. anguillae which also has a very long ecsoma (equal to body length), while all the new species described are different in length from T. serrani. In T. serrani the sinus sac is ovoid, thick-walled

144 containing hermaphroditic vesicle and hermaphroditic duct, genital pore immediately anterior to intestinal bifurcation. Ovary is also ovoid as large as one testis. Vitellaria consist of 8 tubules. Uterus is between ovary and ecsoma.

Manter (1954) has shown that number of vitelline tubules in a species of Tubulovesicula may be either 7 or 8 arranged in various positions.

The main difference is the very long pars prostatica in T. anguillae where it is much longer than the seminal vesicle, also uterus extend far into the ecsoma in T. anguillae, while in all other species the position of uterus is quite different. In T. olivaceus n.sp. and T. microcaudum n.sp., the uterus do not extend into the ecsoma, also vitelline tubules are 7 in number in all new species described above.

If we compare the seminal vesicle of different species, it shows the following differentiations: T. anguisticauda has tubular seminal vesicle and not enclosed in muscular pouch. In T. spari seminal vesicle is long and slender, broad at the base, runs sinuously on the postero-dorsal side of the acetabulum. In T. anguillae seminal vesicle is S-shaped. In T. magna the seminal vesicle is tubular, more or less winding, pre-testicular in position. In T. pseudorhombi seminal vesicle is long, spirally coiled. In T. lindbergi it is tubular, sinuous at

145 about the level of ventral sucker. In T. pinguis the seminal vesicle reaches near about the anterior testis while in the present specimen T. magnacirrosa n.sp. the seminal vesicle is tubular, extending to posterior margin of the ventral sucker.

The position of uterus in T. anguisticauda is different occupying a major portion of the body space between the ventral sucker to the proximal part of the tail. In T. spari uterus coils down on the left side half way into the tail. In T. anguillae the position of uterus is entirely different from other specimen of Tubulovesicula, which reaches the middle of the ecsoma only. In T. muraenesocis the uterus is confined to the body proper only. In T. pseudorhombi uterus slightly enters the ecsoma. In T. lindbergi uterus shows descending and ascending arms, much coiled. In T. pinguis uterus slightly protrudes into the ecsoma.

While in the present specimen T. magnacirrosa n.sp. the uterus occupies most part of the body not extending into the ecsoma and passes forward beside the acetabulum and pre-testicular area.

The third new species T. magnacirrosa n.sp. is comparable with the first T. olivaceus n.sp., and second T. microcaudum n.sp. in having following differential characteristics. Both T. olivaceus n.sp. and T. microcaudum n.sp. are smaller in size as compared to T.

146 magnacirrosa n.sp. In T. magnacirrosa n.sp., not only the entire body size is greater but also the tail is long. The cirrus sac and sinus sac are also large in size as compared to T. olivaceus n.sp. and T. microcaudum n.sp. The testes are relatively far from the ventral sucker in T. magnacirrosa n.sp. than T. olivaceus n.sp. and T. microcaudum n.sp.

147 TUBULOVESICULA KARACHIENSIS N.SP. (Fig. 21)

Host: Pseudosciaena diacanthus (Sciaenidae)

Location: Stomach

Locality: Karachi coast, Pakistan

No. of specimens: 2 specimens from a single host, 17 fishes were examined.

Holotype: JUW – T21

Paratype: JUW – T22

Diagnosis

Body of the worm is medium-sized, more or less spindle-shaped with very short ecsoma as compared to the soma or body proper. Soma is very long. Oral sucker is sub-terminal, smaller than the ventral sucker.

Pharynx is rounded, immediately behind the oral sucker. Caeca do not extend into the tail region. Ventral sucker is in the anterior-third of the soma.

Testes are two, almost equal in size, parallel to each other. The right testis is close to the seminal vesicle. Seminal vesicle is tubular, slightly twisted, posteriorly extending to the base of ventral sucker, anteriorly joining the large pars prostatica. Hermaphroditic duct is

148 small, enclosed in small muscular pouch. Cirrus sac is very prominent. Genital pore is ventro-lateral to pharynx.

Ovary is rounded, almost in the middle of soma. Vitellaria consist of

7, long, tubules directed posteriorly from the base of the ovary far from posterior end of soma. Seminal receptacle is not prominent. The uterus reaches slightly in ecsoma then turns upwards passing between the testes, joining the hermaphroditic duct anteriorly. Eggs are relatively large, thick-shelled. Excretory pore is terminal, excretory tube reaching to posterior part of soma, arms extending to the level of pharynx.

Principle measurements of T. karachiensis n.sp. (In millimeters)

Body size: 5.4–5.5 x 7.4–7.50

Soma length: 4.19–4.20

Soma width: 7.79–7.81

Ecsoma length: 1.13–1.14

Ecsoma width: 0.4–0.41

Forebody: 1.0–1.01 x 0.501–0.511

Hindbody length: 3.88–3.89

Oral sucker: 0.23–0.231 x 0.311–0.312

149 Pharynx: 0.10–0.11 x 0.18–0.181

Ventral sucker: 0.50–0.51 x 0.48–0.49

Sucker width ratio: 1: 0.0460–0.0462

Pars prostatica: 0.611–0.601 x 0.160–0.161

Seminal vesicle: 0.59–0.60 x 0.11–0.12

Ventral sucker to seminal vesicle: 0.0510–0.0511

Ventral sucker to anterior testis: 0.180 – 0.181

Anterior testis: 0.151–0.152 x 0.181–0.179

Posterior testis to ovary: 0.19–0.195

Number of vitelline tubules: 7

Ovary: 0.200–0.201 x 0.20–0.21

Eggs: 0.015–0.025 x 0.016–0.018

Etymology: The present new species Tubulovesicula karachiensis refers to its locality.

Remarks

The present new species of T. karachiensis is differentiated from previously described species from Pakistan and also from other parts of the world. The four previously described species are T. spari, T. magna, T. anguillae and T. anguisticauda. The Pakistani species are different from the present new species T. karachiensis n.sp., in having

150 tubular seminal vesicle not enclosed in muscular pouch in T. anguisticauda. In T. spari seminal vesicle is long and slender, broad at the base, runs sinuously on the postero-dorsal side of the acetabulum. In T. anguillae the seminal vesicle is S-shaped, lying in front of the right testis. In T. magna the tubular seminal vesicle is more or less winding, pre-testicular in position. In T. pseudorhombi the seminal vesicle is spirally coiled. In T. lindbergi seminal vesicle is sinuous, at about the level of ventral sucker. In T. pinguis the seminal vesicle is antero-dorsal to the testes.

In the present species pars prostatica is curved at the side of the ventral sucker. In T. anguisticauda, T. magna, T. muraenesocis, T. lindbergi and T. pinguis all have tubular pars prostatica with small hermaphroditic duct. Sinus sac is pyriform while in T. karachiensis n.sp. it is tubular. The position of the uterus in the present specimen is that it reaches slightly into the ecsoma similar to T. pseudorhombi, while in T. anguisticauda the uterus is extensive and occupies a major portion of body space reaching the proximal part of the tail. In T. spari it reaches half way into the tail. T. anguillae has uterus reaching the middle of the ecsoma. In T. magna the uterus does not reach the ecsoma. In T. lindbergi the uterus has descending and ascending arms, much coiled. In T. pinguis the uterus is in the form of close

151 coils often reaching the ecsoma. Vitellaria in almost all the species of

Tubulovesicula are 6-7 in number.

In T. olivaceus n.sp., tubular sinus sac is present, hermaphroditic duct is enclosed in muscular pouch ventrally at the anterior part of ventral sucker, finally reaching the large genital opening which is immediately behind the intestinal bifurcation. In T. magnacirrosa n.sp. the sinus sac is entirely different, it is large and is pressed out of the body, showing globular shape anteriorly. T. karachiensis n.sp. is also different from the above mentioned new species in having slightly twisted seminal vesicle reaching posterior to the ventral sucker close to testes. Pars prostatica is curved at the side of ventral sucker. Sinus sac is tubular. Genital opening is at the level of pharynx.

T. karachiensis n.sp. is the fourth species of the genus

Tubulovesicula recovered during the present studies from the marine fish of Karachi coast. It differs from other 3 described species in having different diagnostic features including the size of ventral sucker, position of seminal vesicle, size of testes and ovaries and position of uterus. A new species T. karachiensis is therefore proposed.

152 SPECIES OF THE GENUS STOMACHICOLA STOMACHICOLA MURAENESOCIS YAMAGUTI, 1934 (Figs. 22-23)

Family: Hemiuridae Looss, 1899

Sub-family: Stomachicolinae Yamaguti, 1958

Genus: Stomachicola Yamaguti, 1934

Host: Muraenesox cinereus (Muraenesocidae)

Location: Stomach

Locality: Karachi coast, Pakistan.

No. of specimens: 13 specimens from 2 hosts, 50 fishes were examined.

Holotype: JUW – T22

Paratype: JUW – T23

Diagnosis

Body is long, broadest at the level of ventral sucker, almost uniformly broad from below the ventral sucker, with body constriction in the post-ovarian region. Tail is very long, about three times longer than soma. Forebody is short, slightly curved ventrally. Oral sucker is small, sub-terminal, separated above from the pre-oral lobe.

Oesophagus is very short. Caeca with dark, enormous contents, extend through the uterine coils to the posterior extremity of the body.

153 Ventral sucker much larger than the oral sucker, rounded, strongly muscular.

Testes are two, anterior testis is large, oval, ventrally located just behind the acetabulum. Posterior testis is oval, smaller than the anterior testis, on the dorsal side of the body. Seminal vesicle is saccular, transversely elongated and post-acetabular in between and above the two testes. Pars prostatica is winding dorsal to acetabulum.

Hermaphroditic duct enclosed in muscular pouch, opening into wide genital atrium with its aperture just behind the oral sucker.

Ovary is ventral, triangular, convex anteriorly and concave posteriorly, post-testicular in position. Uterine seminal receptacle is much larger than the ovary situated near the ecsoma. Vitellaria consist of 7 tubules, partly running through uterine coils, reaching a considerable distance posterior to ovary, leaving posterior two-third of entire body length or more free. Uterine coils are not only occupying inter-caecal field but also turning round, caeca extending into the tail as far as posterior end of vitellaria. Eggs are small. Excretory vesicle is tubular, excretory arms united anteriorly.

154 Principle measurements of S. muraenesocis (In millimeters)

Entire body size: 21.0 x 21.51

Soma: 4.3 x 4.5

Ecsoma: 16.7 x 16.9

Oral sucker: 0.24–0.25 x 0.28–0.29

Pharynx: 0.29–0.30 x 0.30–0.31

Ventral sucker: 0.84–0.85 x 0.850–0.851

Sucker width ratio: 1: 0.028

Pars prostatica: 2.4–2.41 x 0.011–0.012

Seminal vesicle: 0.74–0.75 x 0.35

Anterior testis: 0.530–0.531 x 0.30–0.31

Posterior testis: 0.5–0.35 x 0.331–0.332

Oral sucker from anterior extremity: 0.29–0.03

Ventral sucker from anterior end of the body: 1.25–1.27

Posterior testis to posterior extremity: 16.09–16.10

Ovary: 0.25–0.24 x 0.21–0.25

Number of vitelline tubules: 7

Eggs: 0.05–0.08 x 0.051–0.07

155 Etymology: The present specimen is identified as Stomachicola muraenesocis Yamaguti, 1934.

Remarks

Type-species: of the genus is S. muraenesocis Yamaguti, 1934 syn.

Lecithocladium longicaudum Shen Tseng, 1935 in Muraenesox cinereus. Other species reported from different parts of the world are

S. magnus (Manter, 1931) Manter, 1947 syn. Dinurus magnus

Manter, 1947, Pseudostomachicola magnus (Manter, 1947). Skrjabin et Guschanskaja, 1954 in Cynoscion nebulosus; S. rubeus (Linton,

1910) Manter, 1947 syn. Dinurus rubeus Linton, 1910 in Lycodontis moringa, L. funebris, Gymnothorax vicinus and Synodus foetens and

S. pritchardae Pandey et al., 2000 in Acanthurus triostegus.

Stomachicola muraenesocis Yamaguti, 1934 is previously known from the fish Muraenesox cinereus of Karachi coast, Pakistan (Zaidi and Khan, 1977). From the same fish and locality, specimens of this species were also recovered during the present studies. These specimens exhibit some morphological variations. Present specimen of Stomachicola muraenesocis recovered is not as long as that of the

156 previously described species by Yamaguti, 1934 (Zaidi and Khan,

1977).

Seminal vesicle in the present specimen is larger in size than in the previously described specimens. Both the testes are far from the ventral sucker, while in the previously described specimens the testes are very close to the ventral sucker. Pars prostatica is long, tubular, sinuous, surrounded by prostrate gland cells winding dorsal to the acetabulum. Ductus hermaphroditicus is short and tubular, enclosed within the hermaphroditic pouch. Vitelline follicles in the present specimens are 7 in number similar to species described previously by

Yamaguti, 1934.

Ovary is triangular, almost equal in size than the previously described species in which it is kidney-shaped. The seminal receptacle in both the previous and present specimens is large in size.

During the survey of digenetic trematode parasites of marine fishes from Puri coast, Balasore coast (Orissa) and Digha coast (West

Bengal), India. A total 48 fishes of the families Symbranchidae and

Muraenidae were examined, out of which only 10 were found infected with 35 mature and 16 immature digenetic trematode parasites. These parasites referred to the genus Stomachicola Yamaguti, 1934 are

157 placed by Gibson and Bray, 1979 in the sub-family Stomachicolinae of the family Hemiuridae. Yamaguti, 1971 listed only three species under this genus reported from Japan, Florida and Carolina respectively and after that Indian workers made the contribution of eight new species upto date. In between this period, five new genera, one from India and four from Pakistan have been described and placed in the sub-family Stomachicolinae.

Yamaguti, 1934 established the genus Stomachicola of which he listed following species viz., S. muraenesocis Yamaguti, 1934; S. rubea (Linton, 1910) Manter, 1947 and S. magnum (Manter, 1931)

Manter, 1947. After that following species reported from India have been added namely, S. mastacembeli Verma, 1973; S. polynemi

Gupta and Gupta, 1974; S. lepturusi Gupta and Gupta, 1974; S. bayagbonai Siddiqi and Hafeezullah, 1975; S. singhi Gupta and

Ahmad, 1979; S. rauschi Gupta and Ahmad, 1979; S. chauhani

Pandey and Tewari, 1984 and S. chauhani Gupta and Singh, 1985. A new name S. guptai (in honour of its senior author) is being proposed for S. chauhani Gupta and Singh, 1985 as its synonym of S. chauhani Pandey and Tewari, 1984.

158 Due to great diversity in the internal organs of this parasite, Gupta and

Gupta, 1991 considered S. rubea (Linton, 1910) Manter, 1947; S. mastacembeli Verma, 1973; S. polynemi Gupta and Gupta, 1974; S. chauhani Pandey and Tewari, 1984 and S. guptai (in honour of its senior author) is being proposed for S. chauhani Gupta and Singh,

1985 nom. nov., conspecific to S. muraenesocis Yamaguti, 1934.

Another species S. pritchardae Pandey et al., 2000 has been described from marine fishes of Puri coast, Orissa from the fish Acanthurus triostegus which is characterized by having sub-cylindrical, elongated body. Oral sucker is very small as compared to ventral sucker. Genital pore is at the base of oral sucker.Testes are two, posterior testis is smaller than the anterior testis. Seminal vesicle is elongated, saccular, lying free in parenchyma. Ovary is tetra-lobed, a little posterior to the testes. Vitellaria are 7, long tubules. Excretory pore is terminal.

Present specimens are regarded S. muraenesocis with slight morphological variations. Previously it has been reported (Bilqees,

1981) from the same fish host and locality of Pakistan.

159 SPECIES OF THE GENUS PROSOGONOTREMA PROSOGONOTREMA DIACANTHI BILQEES, 1980 (Figs. 24-25)

Family: Prosogonotrematidae Perez Vigueras, 1940

Sub-family: Prosogonotrematinae Perez Vigueras, 1940

Genus: Prosogonotrema Perez Vigueras, 1940

Host: Pseudosciaena diacanthus

Location: Stomach

Locality: Karachi coast

No. of specimens: 3 specimens from a single host, 39 fishes were examined.

Holotype: JUW – T24

Paratype: JUW – T25

Diagnosis

Body is robust, elongated, rounded anterior end and posterior end is bluntly pointed, greatest width at acetabular region. Body covered with thick cuticle. Forebody very long, comprising all the organs of the body. Hindbody very short, containing the last part of the intestinal caeca. Oral sucker is very small. Prepharynx is absent.

Pharynx is present. Acetabulum is much larger than oral sucker, situated close to posterior extremity than to anterior extremity.

160

Testes are pre-ovarian, very far above the acetabulum, almost in between the oral sucker and ventral sucker, rounded in shape. Seminal vesicle is tubular, lying close to the anterior testis. Pars prostatica is a winding tube, ending in a genital pore at the level of pharynx.

Ovary is small and spherical, situated a little above the anterior level of the acetabulum. A small elongated, pointed seminal receptacle is present. Vitellaria are forming slender, convoluted tubules 4 on one side and 3 on the other side, arising from the base of ovary. Uterus is coiled transversely between the testes and the ovary. Excretory pore is dorsal at the posterior extremity.

Principle measurements of P. diacanthi (In milimeters)

Body Size 11.90–12.00 x 3.49–3.50

Forebody 8.47–8.48 x 3.40–3.50

Hindbody 3.40–3.41 x 2.40–2.41

Oral sucker 0.80–0.81 x 0.81–0.82

Ventral sucker 2.09–2.10 x 1.90–1.92

Sucker width ratio 1:0.0038 x 0.0039

Pars prostatica 1.52–1.60 x 0.32–0.38

161 Pharynx 0.20–0.21 x 0.29–0.30

Anterior Testis 0.50–0.51 x 0.51–0.51

Posterior testis 0.54–0.55 x 0.60–0.61

Ovary 0.39–0.40 x 0.48–0.49

Genital cone 0.82–0.83 x 0.30–0.31

Seminal receptacle 0.70–0.71 x 0.38–0.39

Number of vitelline tubules 7

Eggs 0.021–0.027 x 0.013–0.016

Etymology: The present specimen is identified as Prosogonotrema diacanthi Bilqees, 1980.

Remarks

Two species of the genus Prosogonotrema reported from Karachi,

Pakisan are Prosogonotrema diacanthi Bilqees and Durrani, 1980 from Pseudosciaena diacanthus and P. karachiense Bilqees and

Durrani, 1980 from Lutjanus johnii. Both the Pakistani species differ from the previously described species in body size, acetabulum and sucker-width ratio.

162 P. diacanthi Bilqees, 1980 is characterized by having a more posterior position of the acetabulum, a greater sucker-width ratio, well developed receptacular seminis uterinum, relatively small forebody and larger operculate eggs.

P. karachiense Bilqees, 1980 is characterized by smooth pre-oral lobe. Ovary is median, large antero-dorsal to acetabulum. Eggs are relatively smaller than P. diacanthi. Pars prostatica is enormously developed and a prominent excretory vesicle.

Present species is different from other species and is compared below:

P. bilabiatum Perez Vigueras, 1940 is characterized by having two pre-oral lobes, or simple pre-oral lobe. Oesophagus is short.

Acetabulum is near the posterior extremity. Seminal vesicle is tubular, winding. Genital pore is ventral to pharynx. Ovary is antero-lateral to acetabulum. Receptacular seminis is absent. Eggs are small.

P. clupeae Yamaguti, 1952 is characterized by very prominent pre- oral lobe. Ovary is relatively large, antero-dextral to acetabulum, just in front of the equator, smaller than P. diacanthi and P. karachiense.

Acetabulum is prominent, occupying most of the middle-third of the body with its center a little behind equator. P. clupeae differs from P.

163 bilabiatum in body size, in the absence of bi-lobate pre-oral lip in the length of eggs.

The body length of P. diacanthi is close to P. clupeae and P. symmetricum Oshmarin, 1965. In P. diacanthi the eggs are slightly larger, the sucker width ratio is different and the forebody is much larger than in P. clupeae and P. symmetricum. In P. diacanthi the position of acetabulum is similar to P. clupeae and P. subequilatum but P. diacanthi differs from P. clupeae and P. subequilatum in the position of testes, in sucker-width ratio and in the presence of receptacular seminis uterinum.

P. symmetricum has oral sucker sub-terminal, smaller in size than the present specimen, oesophagus is present. Mostly other diagnostic features show little variations with P. diacanthi. In P. symmetricum the genital cone is entirely enclosed in genital atrium situated ventral to pharynx and oesophagus which is also the characteristic feature of

P. diacanthi. The vitelline tubules (some anteriorly directed) may even reach the level of genital cone. Excretory vesicle is Y-shaped.

The above species differs from the most closely related P. bilabiatum in the sucker-width ratio 1: 2.6 – 3.0 instead of 1: 3.3 and from P.

164 subequilatum in the acetabulum being little more posterior and in smaller egg size. In P. subequilatum the sucker-width ratio is 1: 1.6 –

1.9 and the uterine coils extend into the extra caecal fields.

The species from Pakistan P. karachiense is larger in size as compared to P. diacanthi. P. karachiense is peculiar among the species of the genus in having a relatively large ovary immediately anterior to and overlapping the anterior periphery of acetabulum and in the enormous development of pars prostatica. P. karachiense is larger in size than P. bilabiatum, P. clupeae, P. subequilatum, P. abalisti and P. symmetricum. As far as body size is concerned P. karachiense is close to P. carangis Velasquez, 1961, but differs in other morphological characteristics. As far as egg size is concerned P. karachiense has larger eggs than all other species except P. subequilatum which has egg sizes close to P. karachiense. The sucker width ratio in P. karachiense is greater than all other species of the genus. The ovary in P. subequilatum is dorsal to acetabulum.

The present species P. diacanthi is different with some variations as compared to the original description in having tubular, winding seminal vesicle present behind the ovary. Pars prostatica is tubular, winding. Ejaculatory duct convoluted in muscular tissue at the base of

165 genital cone. Genital atrium is enclosing bi-partite genital cone opening at the level of pharynx. Ovary is antero-dorsal to acetabulum.

Excretory vesicle is Y-shaped reaching the level of oral sucker.

166 SPECIES OF THE GENUS PLAGIOPORUS PLAGIOPORUS KARACHIENSIS N.SP. (Fig. 26)

Family: Opecoeliidae Ozaki, 1925

Sub-family: Plagioporinae Dujardin, 1845

Genus: Plagioporus Stafford, 1904

Host: Caranx affinis (Carangidae)

Location: Intestine

Locality: West Wharf, Karachi.

No. of specimens: 4 specimens from a single host, 13 fishes were examined.

Holotype: JUW – T26

Paratype: JUW – T27

Diagnosis

Body is medium in size, oral region narrower, posterior body region flat. Oral sucker is terminal, fairly small as compared to ventral sucker which is large, slightly broader than long with strongly muscular slit- like opening followed by a bulbous pharynx tapering at both ends.

Oesophagus is distinct and clearly leads into intestinal bifurcation.

Intestinal caeca are reaching to posterior end of the body. Cirrus pouch is large, saccular, more or less pear-shaped, slender near the

167 pore and thick posteriorly, containing elongated seminal vesicle, poorly developed pars prostatica, opening at the intestinal bifurcation situated antero-dorsal to ventral sucker. Cirrus sac internally contains a large, broad seminal vesicle and few prostatic cells. The cirrus pouch extends dorsal to the acetabulum occupying the space between the intestinal bifurcation and acetabulum.

Testes are two, separate, irregular, almost tandem in the posterior part of the body. Ovary is rounded to oval, pre-testicular, a little behind the ventral sucker. Vitellaria consist of numerous, prominent follicles extending in lateral fields from the posterior level of pharynx to near the posterior end, confluent in the pre-acetabular and post-testicular region totally covering the caeca and larger follicles intruding posterior to ovary. The uterus is pre-testicular, reaching anteriorly to genital opening filled with large eggs. Excretory vesicle is lateral on the left side of body behind testes.

Principle measurments of P. karachiensis n.sp. (In milimeters)

Body Size 2.30–2.31 x 8.20–8.22

Maximum width of body 0.85–0.86

Oral sucker 0.10–0.11 x 0.131–0.132

168 Acetabulum 0.40–0.41 x 0.50–0.51

Sucker width ratio 1:0.025 x 0.026

Pharynx 0.125–0.130 x 0.135–0.136

Anterior Testis 0.19–0.20 x 0.26–0.27

Posterior testis 0.20–0.21 x 0.26–0.27

Cirrus pouch 0.19–0.20 x 0.24 – 0.25

Ovary 0.20–0.21 x 0.21–0.22

Genital pore 0.01–0.02

Seminal vesicle 0.21–0.22 x 0.075–0.080

Eggs 0.04–0.05 x 0.01–0.02

Etymology: The present new species Plagioporus karachiensis refers to the locality of host.

Remarks

Two species found in Pakistan in marine fishes are Plagioporus

(caudotestis) mujibi Bilqees, 1972 reported from Crenidens indicus and Plagioporus heterorchis Bilqees, 1977 from the fish Pomadasys olivaceum. Here a third new species P. karachiensis is identified and described from the marine fish Caranx affinis. Only one species P.

169 gonii Bilqees and Khan, 1988 has been reported from the freshwater fish Labeo gonius.

P. (caudotestis) mujibi has body large, posterior, wide and rounded.

Oral sucker is sub-terminal. Both oral and ventral sucker show little difference in size. Pharynx is large. Oesophagus is not distinct. Cirrus pouch weakly developed containing internal seminal vesicle. Testes are two, transversely elongated, lying in the posterior part of the body.

Ovary is pre-testicular, between the anterior testis and posterior part of the acetabulum. Vitelline follicles lateral. Uterus is with few coils, eggs are few. Excretory vesicle is Y-shaped. Excretory pore is terminal.

P. heterorchis has elongated, spindle-shaped body. Oral sucker is three times smaller than the ventral sucker. Cirrus sac is slightly curved and slightly extending to the posterior part of the acetabulum.

Seminal vesicle is large with scattered gland cells. Vitelline follicles are arranged laterally extending from the base of the pharynx to the posterior end of the body. Uterus is pre-ovarian. Excretory vesicle is tubular, extending to the posterior testis. Excretory pore is terminal.

170 Species found in different parts of the world are different from

Pakistani species described above are P. macassarensis Yamaguti,

1952 from the small intestine of Lethrinus species having distinct oesophagus, is not prominent in P. heterorchis. Seminal vesicle in P. heterorchis extends posteriorly to the acetabulum but in P. macassarensis it is above the acetabulum. Uterus in P. macassarensis is descending from right to left as far as anterior testis and then winding forward along cirrus pouch. Vitelline glands divided on each side into about 10 grape like bunches, extending along the intestine on the left from genital pore to posterior extremity. Excretory vesicle is terminal, tubular, with swollen anterior end.

P. longivesicula Yamaguti, 1952 from the fish Lethrinus sp. is blunt anteriorly, rounded posteriorly. Oral sucker is sub-terminal, leading into a short prepharynx and pharynx with long oesophagus. Cirrus pouch is claviform. Seminal vesicle is tubular, sigmoid in P. serotinus

Stafford, 1904 but constructed into two portions as in P. macassarensis. Genital pore is ventral to the left caecum. Ovary consisting of a central part and three rounded lobes of nearly the same size, situated in front of posterior testis by right side of anterior testis.

Uterus winding from side to side between anterior testis and acetabulum, finally running along left side of cirrus pouch. Vitellaria

171 on each side 8 grape like bunches of small follicular acini, extending all round caeca from level of genital pore to posterior extremity.

Excretory vesicle is long and tubular.

P. (caudotestis) synagris Yamaguti, 1952 reported from synagris sp. has fusiform body with blunt ends. Testes are globular, contiguous, oblique. Cirrus pouch is thin-walled, extending along left side of acetabulum with its posterior end between acetabulum and anterior end of receptaculum seminis. Seminal vesicle is filling entire posterior portion of cirrus pouch, with its tapered anterior part turned back on itself two times at level of middle of acetabulum. Genital pore is in left median line from anterior extremity. Ovary is ovoid, entire, situated obliquely on the right median line just inside caecum at posterior part of middle-third of body. Uterus forms a loop in front of the anterior testis and then proceeds forward along the side of the cirrus pouch. There are only two collapsed eggs beside the few abortive ones. Vitelline follicles are extending from level of genital pore to posterior extremity, surrounding caeca on all sides. Excretory vesicle is tubular, with terminal pore.

In P. macassarensis the excretory vesicle is tubular, forming bulbous dilation wide just in front of its terminal opening, much more widely

172 swollen at its anterior end reaching to acetabulum, giving off a pain of lateral collecting vessels between ovary and testes, each collecting vessel divided in neck region into an anterior and posterior tubule.

P. longivesicula may turn out to be identical with P. pallensicus

(Shipley et Hornell, 1905) may be confused with P. macassarensis but they are quite different in the anterior extent of the excretory vesicle, in the origin of the Laurer’s canal and position of its dorsal opening in the lobed ovary, etc.

P. synagris is characterized by long cirrus pouch reaching beyond the acetabulum and the enormous receptaculum seminis being much larger than the ovary. The size of oral sucker and acetabulum is very similar to the present specimen. In the present specimen P. karachiensis n.sp. the cirrus pouch is heavy, pear-shaped containing a large seminal vesicle internally. While in P. synagris the shape of the seminal vesicle is quite different from the present specimen P. karachiensis n.sp. as described above.

P. congeri Yamaguti, 1970 reported from the fish Conger sp. has elongated body, tapered anteriorly in forebody but nearly cylindrical

173 in the hindbody. Oesophagus is very long. Cirrus pouch is claviform.

Seminal vesicle is elliptical. Cirrus sac is smooth, conical, projecting out of genital pore. Genital pore is near left margin of body at level of intestinal bifurcation. Ovary is heart-shaped. Uterine coils confined to inter-caecal field between ovary and acetabulum. Vitellaria are forming H-shaped pattern. Excretory vesicle is tubular, reaching to dorsal side of ovary. Genital pore is terminal. This species resembles

P. isaitschkowi (Layman, 1930) most closely in shape and size of body and internal anatomy, but differs from it in the posterior position of the genital pore and posterior extent of the intestinal limbs. While the present specimen P. karachiensis n.sp. is also different from P. congeri and other species in having different shape and size of the cirrus pouch and other diagnostic features are also different.

P. longisacculus Yamaguti, 1970 from the fish Pristipomoides microlepis has ventro-terminal oral sucker. Cirrus pouch is very long, reaching to the level of posterior end of acetabulum containing a tubular seminal vesicle winding muscular ejaculatory duct everted out of the genital pore as a smooth stout cirrus sac. Ovary is 3 or 4 lobed, post-equatorial, separated dorsally from anterior testis by anterior end of excretory vesicle. Uterine coils occupying entire caecal field between ovary and acetabulum. Vitelline follicles are circum-caecal.

174 Excretory vesicle tubular, sigmoid anteriorly and reaching to ovary, opening at mid-terminal notch. P. longisacculus differs from the known members of the genus Plagioporus as well as the present species P. karachiensis n.sp. in the posterior extent of the cirrus pouch P. longisacculus bears resemblance to P. longivesicula

Yamaguti, 1952 from Lethrinus sp. of Macassar in the length of excretory vesicle.

P. polymixiae Yamaguti, 1970 reported from the fish Polymixia japonica having lanceolate body, little difference in the size of oral sucker and ventral sucker. The species differs from the known members of the genus Plagioporus as well as the present species P. karachiensis n.sp. in the seminal receptacle being so much reduced that it can hardly be detected. It is also worth noted that the ovary and testes are very irregularly lobed.

P. ula-ula Yamaguti, 1970 reported from the fish Etelis marshi. This species differs from the present specimen P. karachiensis n.sp., P. apogonichthydis Yamaguti, 1938 and P. sillagonis Yamaguti, 1938 or any other Plagioporus species with a similar extent of the cirrus pouch in the ovary being 4 or 5 lobed. The specific name refers to the

175 local name of the host that the Laurer’s canal opens dorsal to the posterior end of the acetabulum is worth noting.

P. rooseveltiae Yamaguti, 1970 reported from the fish Rooseveltia brighani. This species closely resembles P. ula-ula from Etelis marshi, but differs from it and from the present new species described above in the anterior extent of the vitellaria and in the length of the

Laurer’s canal. In P. ula-ula this canal opens dorsal to the posterior end of the acetabulum, but in P. rooseveltiae it opens some distance posterior to the acetabulum.

P. tohei Yamaguti, 1970 reported from the fish Conger sp. (Local name “Tohei”) having plump body. This species differs from the most closely related P. congeri from the fish Conger sp. in the anterior extent of the vitellaria and in the length and course of the Laurer’s canal. It is also noted that the measurements of both species do not coincide. The present species P. karachiensis n.sp. also differs from

P. tohei in having medium-sized body with tapering ends, while P. tohei is plump having sub-terminal oral sucker. Genital pore is sub- lateral in P. tohei, while in P. karachiensis n.sp. the genital pore is terminal. Ovary is irregular in shape with 3-4 lobes in P. tohei, while

176 the present specimen P. karachiensis n.sp. has rounded, oval, pre- testicular ovary. Excretory vesicle lateral on the left side of the ovary behind the testes in the present specimen P. karachiensis n.sp., while in P. tohei the excretory vesicle is tubular, reaching to dorsal side of ovary with dorsal terminal pore.

P. (caudotestis) thalassomatis Yamaguti, 1942; Yamaguti, 1970 reported from the fish Thalassoma ballieni has cuticle with extremely fine, transverse serrations on the surface. Caeca are terminating blindly at level of ovary in front of left testis. Seminal vesicle is N- shaped. Genital pore to left of oesophagus at varying levels, may be near pharynx or intestinal bifurcation. Ovary is sub-globular. Uterus is winding in post-acetabular, inter-caecal field, intruding into space between ovary and anterior testis.

Present new species P. karachiensis is separated from the previously described species in the main diagnostic features described above.

177 SPECIES OF THE GENUS STEPHANOSTOMUM STEPHANOSTOMUM GIBSONI N.SP. (Figs. 27-29)

Family: Acanthocolpidae Luhe, 1906

Sub-family: Acanthocolpinae; Stephanostominae

Genus: Stephanostomum Looss, 1899

Synonyms: Stephanochasmus Looss, 1900; Lechradena Linton, 1910;

Echinostephanus Yamaguti, 1934; Monorchistephanostomum Vigueras, 1942

Host: Pomadasys olivaceum (Pomadasyidae)

Location: Intestine

Locality: Karachi coast, Pakistan

No. of specimens: 2 specimens from 2 hosts, 13 fishes were examined.

Holotype: JUW – T28

Paratype: JUW – T29

Diagnosis

The body of the worm is long with tapering anterior and posterior ends. Narrow at the region of the cirrus sac and wider at the level of the region of the gonads. Tegument is spinous, densely spined in the forebody. Circum oral spines are about 38-41 in two alternate circular

178 rows. Oral sucker terminal, muscular, longer than wide. Prepharynx is long. Pharynx is pyriform. Oesophagus is distinct. Intestinal bifurcation is in the middle of the body.

Gonads are in posterior-third of the body. The testes are situated in the middle of posterior-quarter of the body, oval to globular. Posterior testis is bilobed, larger than the anterior testis. Cirrus sac is very long, reaching in the posterior half of the body, close to seminal receptacle, containing bi-partite seminal vesicle, posterior lobe rounded, anteriorly elongated and sigmoid. Pars prostatica well developed.

Ejaculatory duct is long. Cirrus sac is leading to genital atrium lying at the anterior level of the dorsal side of the ventral sucker. Genital opening is dorsal, at the anterior level of ventral sucker.

Ovary is rounded, smaller than the testes. Seminal receptacle is anterior, close to the ovary and contagious with the uterus. Uterus is anterior to ovary. The follicular vitellaria in the posterior half of the body, extend from posterior part of cirrus sac to posterior extremity, confluent in the post-testicular region.

179 Principle measurements of S. gibsoni n.sp. (In millimeters)

Body size: 6.8–6.9 x 0.8–0.81

Forebody length: 0.5–0.59 x 0.4–0.41

Hindbody length: 5.88–5.90 x 0.8–0.79

Circum oral spine number: 38–41

Oral sucker: 0.2–0.22 x 0.16–0.17

Pharynx: 0.24–0.25 x 0.10–0.11

Prepharynx: 0.28–0.29 x 0.08–0.09

Ventral sucker: 0.4–0.45 x 0.39–0.40

Sucker width ratio: 1: 0.004–0.0041

Ventral sucker to cirrus sac: 1.5–1.49

Ventral sucker to anterior testis: 4.2–4.25

Anterior testis: 0.5–0.55 x 0.34–0.35

Posterior testis (Bi-lobed)

Lobe 1: 0.31–0.315

Lobe 2: 0.29–0.30

Posterior testis to posterior extremity: 0.68–0.7

Cirrus sac: 2.49–2.5 x 0.09–0.10

Seminal vesicle (Anterior part): 1.0–1.1 x 0.15–0.16

Seminal vesicle (Posterior part): 0.20–0.21 x 0.19–0.195

180 Ovary: 0.24–0.25 x 0.23–0.24

Seminal receptacle: 0.3–0.29 x 0.21–0.22

Excretory vesicle: Not traced

Tail: 0.41–0.42 x 0.4–0.41

Eggs: 0.06–0.12 x 0.05–0.07

Etymology: The present new species Stephanostomum gibsoni is named in the honour of Dr. D.I. GIBSON a well known parasitologist who has been associated with Zoology Department, Natural History

Museum, London.

Remarks

Luhe, 1906 erected the sub-family Acanthocolpinae Luhe, 1906 to include the genus Stephanochasmus Looss, 1900 (now considered the junior synonym of Stephanostomum Looss, 1899).

Cable and Hunninen, 1942 redefined the family, removed the

Deropristidae Cable and Hunninen, 1942 as a separate family and included Stephanostomum, Dihemistephanus Looss, 1901 (now considered a Lepocreadiid) Echinostephanus Yamaguti, 1934 (now considered a synonym of Stephanostomum). The distinguishing

181 feature is the presence or absence of circum oral spines (Yamaguti,

1958).

Stephanostomum Looss, 1899 is the best known genus in the family with over 112 nominal species. Until recently, the type-species, S. cesticillum Molin, 1858 has not been redescribed to modern standards, but unlike many species, it lacks a uroproct (Bartoli &

Bray, 2001). There has been some discussion as to whether the presence or absence of a uroproct should be used to split the genus

(Bray, 1985), but Bartoli & Bray, 2001 concluded that the difficulty in detecting the uroproct in some forms and its apparent occurrence at random among similar forms (see in Bray & Cribb, 2003b) argue against this.

Names usually considered synonyms of Stephanostomum include:

Stephanochasmus Looss, 1901; Lephradena Linton, 1910;

Echinostephanus Yamaguti, 1934 and Monorchistephanostomum

Vigueras, 1942. Stephanochasmus was erected by Looss, 1900 as a replacement name for Stephanostomum. The type-species of

Lechradena is now considered a synonym of Stephanostomum casum Linton, 1910 is known to have a uroproct (Manter, 1947) and would, therefore, probably be the valid name for the genus with the

182 uroproct if it was ever considered a good idea to split

Stephanostomum. Echinostephanus was erected by Yamaguti, 1934 and was said to differ from Stephanochasmus in the possession of a uroproct.

Monorchistephanostomum Vigueras, 1942 was synonymized with

Stephanostomum by Yamaguti, 1954. Vigueras, 1955 divided

Stephanostomum into four sub-genera; he did not include the type- species, S. cesticillus in Stephanostomum, but placed it in

Schistostephanostomum.

Originally genus Stephanostomum was proposed by Looss, 1899. As mentioned above this is a large genus containing numerous (more than

112) species, but only two species are known from fishes of Karachi coast. Present is the third species from the fish of same locality.

The numbers of circum oral spines are one of the diagnostic features in the genus. The circum oral spines in the present new species are in two alternate circular rows each with 38-41 circum oral spines, while there are 30-34 circum oral spines in S. madhaviae Madhavi, 1976 from Caranx ignobilis; 31 in S. bicoronatum (Stossich, 1883)

Fuhrmann, 1928 from Sciaena umbra; 33-38 in S. bicoronatum

183 Stossich, 1883 from Argyrosomus hololepidotus; 33-38 in S. votonimoli Bray and Cribb, 2003 from Scomberoides lysan; 33-38 in

S. nyoomwa Bray and Cribb, 2003 from Caranx sexfasciatus; 35 in S. cesticillum (Molin, 1858) Looss, 1899 from Lophius piscatorius; 35-

36 in S. pagrosomi Yamaguti, 1939 from Lethrinus nebulosus; 36 in

S. cobia Bray and Cribb, 2003 from Rachycentron canadum; 36 in S. pristis (Deslongchamps, 1824) Looss, 1899 from Phycis phycis; 36 in

S. minutum (Looss, 1901) Manter, 1940 from Uranoscopus scaber;

36 in S. gaidropsari Bartoli and Bray, 2001 from Gaidropsarus mediterraneus; 36 in S. ditrematis Yamaguti, 1939 from Seriola dumerili; 36-39 in S. pacificum Yamaguti, 1951 from Pseudocaranx wrighti; 36-39 in S. aaravi Bray and Cribb, 2003 from Lethrinus miniatus; 37-42 in S. seriolae Yamaguti, 1970 from Seriola dumerili;

42 in S. petimba Yamaguti, 1970 from Fistularia petimba; 40-46 in S. petimba (Bray and Cribb, 2003) Yamaguti, 1970 from Seriola hippos;

43-46 in S. filiforme Linton, 1940 from Seriola lalandi; 48 in S. caducum (Looss, 1901) Manter, 1934 from Gadus minutus; 48-50 in

S. caducum Koie, 1984 and Karlsbakk, 1993; 44-56 in S. baccatum

Nicoll, 1907 from Hipoglossus vulgaris and 49-51 in S. euzeti Bartoli and Bray, 2004 from Seriola dumerili.

184

Six species of Stephanostomum reported from same host

Rachycentron canadum have 34 circum oral spines in S. cloacum

Srivastava, 1938; 30-34 in S. rachycentronis Shen, 1990; 33-34 in S. imparispine Linton, 1905; 36 circum oral spines in S. microsomum

Madhavi, 1976 and 54-58 circum oral spines in S. dentatum Linton,

1900.

24 circum oral spines in two alternating rows are found in S. cesticillum (Molin, 1858) Looss, 1899 from Lophius piscatorius; 24 in S. lebourae Caballero, 1952 from Gaddus sp.; 28 in S. bulbposum

Ramadan, 1983 from Lethrinus mahsena; 32 in S. keilbachi Reimer,

1983 from Uranoscopus archionema; 30-34 in S. rachycentronis

Shen, 1990 from Rachycentron canadum; 32-42 in S. gaabooli

Nagaty et Abdel-Aal, 1962 from Therapon gerbua; 33 in S. bicoronatum (Stossich, 1883) Fuhrmann, 1928 from Umbrina cirrhosa; 34 in S. jesseni Reimer, 1983 from Seriola nigrofasciata; 34 in S. imparispine (Linton, 1905) Manter, 1940 from Rachycentron canadum; 35 in S. hawaiiense Yamaguti, 1970 from Caranx sexfasciatus; 35 in S. uku Yamaguti, 1970 from Aprion virescens; 36 in S. ditrematis (Yamaguti, 1939) Manter, 1947 from Ditrema temmincki; 36 in S. minutum (Looss, 1901) Manter, 1940 from

185 Uranoscopus scaber; 36-40 in S. casum (Linton, 1910) McFarlane,

1934 from Lutjanus sp; 37-42 in S. seriolae Yamaguti, 1970 from

Seriola dumerili; 40-42 in S. tenue Overstreet, 1969 (Linton, 1898)

Linton, 1934 syn. Distomum tenue tenuissime Linton, 1898 from

Roccus lineatus; 40-44 in S. kawalea Yamaguti, 1970 from Sphyaena helleri; 42 are in S. petimba Yamaguti, 1970 from Fistularia petimba;

46 in S. japanocasum (Yamaguti, 1934) Manter et Van Cleave, 1951 from Sascottus setiger; 44-48 in S. filiforme Linton, 1940 from

Seriola lalandi; 50 are in S. promicropsi Manter, 1947 from

Promicrops itaira; 52-56 are in S. yagara Yamaguti, 1970 from

Fistularia petimba; 54-58 in two rows are in S. dentatum (Linton,

1900) from Paralicthys dentatus; Circum oral spines in double rows are in S. hystrix (Dujardin, 1845) Looss, 1899 from Pleuronectes maximus; 56 circum oral spines in two rows are found in S. pagrosomi (Yamaguti, 1939) Manter, 1947 from Pagrus auratus; 42 are in S. polymixiae Yamaguti, 1970 from Polymixia japonica; 80 in

S. multispinosum Manter, 1940 from Mycteroperca olfax and 80-84 circum oral spines in S. trachinoti Fischthal et Thomas, 1968 from

Trachinotus glaucus.

Few species have only three rows of circum oral spines, double row dorsally and a single row on the ventral side such as in S. interruptum

186 Spark and Thatcher, 1958; 34 circum oral spines in two alternating rows dorsally and a single row ventrally in S. cloacum (Srivastava,

1938) Manter et Van Cleave, 1951 from Lates calcarifer; S. lineatum

Manter, 1934 from Fistularia petimba is also different from present species in having 50-52 circum oral spines in three rows and 70-80 circum oral spines are also in three rows in S. admicrostephanum

Perez Vigueras, 1955 from Epinephelus mystacinus and is also different.

The present specimens are also different from the species having interrupted rows of circum oral spines such as 23-24 circum oral spines in S. interruptum Spark and Thatcher, 1958 from Bairdiella chrysura, Micropogon undulates, Cynoscion nebulosus and Ocyurus chrysurus; 38 in two alternating rows interrupted ventrally in S. manteri Perez Vigueras, 1955 from Elaphotoxon sp. and 50-52 circum oral spines in three rows in S. lineatum Manter, 1934 from

Fistularia petimba.

The present species have two uninterrupted rows of circum oral spines. Several other species having similar arrangement of circum oral spines but are different in other diagnostic features such as number of circum oral spines and position of tegumentary spines. Of

187 more than 112 nominal species of Stephanostomum, 26 are quoted with a spine number ranging 35 or 36. Of these, 22 have uninterrupted rings of circum oral spines: S. adinterruptum Hafeezullah, 1971; S. argyrosomi Shen in Shen and Qiu, 1995; S. attenuatum Hafeezullah,

1971; S. aulostomi Nahhas and Cable, 1964; S. carangi Liu, 1998; S. carangis Yamaguti, 1951; S. casum Linton, 1910; S. ceylonicum

Luhe, 1906; S. coryphaenae Manter, 1947; S. ghanensis Fischthal and Thomas, 1968; S. lucknowensis Gupta and Jahan, 1977; S. microsomum Madhavi, 1976; S. minutum Looss, 1901; S. pacificum

Yamaguti, 1951; S. pristis Deslongchamps, 1824; S. pseudocarangis

Sogandares-Bernal, 1959; S. pseudoditrematis Madhavi, 1976; S. rhombispinosum Lebour, 1908; S. sentum Linton, 1910; S. simhai

Gupta and Ahmad, 1979; S. trompeteri Zhukov, 1983 and S. uku

Yamaguti, 1970. Out of these, two S. pristis and S. minutum have a double uninterrupted crown of 18 spines.

The present specimens are different from the species having uninterrupted rows of circum oral spines these include: S. cabanum having 32 circum oral spines in two uninterrupted alternating rows

Perez Vigueras, 1955 from Elaphotoxon rubber; 23-26 in S. lopezneyrai Perez Vigueras, 1955 from Anisotremus sp.; 30-34 in S. ghanensis Fischthal et Thomas, 1968 from Trachinotus goreensis; 34

188 in S. mediovitellarium Perez Vigueras, 1955 from Calamus bajonudo;

36-38 in S. rhombispinosum (Lebour, 1908) syn. Stephanochasmus from Gadus merlangus (36 spines are in 2 uninterrupted alternating rows); 48-50 in S. gracile Perez Vigueras, 1942 from Sphyraena barracuda and 46 in S. nipponicum Yamaguti, 1957 from

Pagrosomus auratus.

In the present specimens the spines are not present on the entire body, large spines are densely packed in the forebody. In S. madhaviae tegument is spinous, unarmed patch immediately posterior to oral sucker, large spines densely packed in forebody, anterior spines in mid-ventral region long, longest spines; spines smaller and sparser in hindbody, none seen posteriorly to ovary (Bray and Cribb, 2003).

In S. bicoronatum the position of spines is similar to S. madhaviae except that spines in the forebody are not densely packed but they are in regular array in the forebody, anterior spines in the mid-ventral region 19-41 (30) long, longest spines 34-49 (39) long; spines smaller, more irregularly arrayed and sparser in hindbody, none seen posteriorly to middle of posterior testis.

189

S. votonimoli also has position of spines similar to S. madhaviae except that spines in the hindbody are also densely packed as compared to the spines which are sparse in hindbody in case of S. madhaviae. Anterior spines in S. votonimoli in mid-ventral region are

9-14 (12) long, longest spines 14-19 (17) long; spines smaller and sparser in mid-hindbody, none seen posteriorly to ovary.

In S. nyoomwa the position of spines is same as described in S. madhaviae, except that anterior spines in mid-ventral region are 14-20

(18) long, longest spines 22-28 (25) long; spines smaller and sparser in hindbody; none seen posteriorly to ovary.

In S. cobia again the same position of spines is present as in S. madhaviae, except that anterior spines in mid-ventral region 14-20

(17) long, longest spines 20-31 (25) long. Here the spines reach the posterior extremity. In none of the above described species the spines reach the posterior extremity.

In S. petimba position of spines is same as described in S. madhaviae, except that anterior spines in mid-ventral region 5-18 (13) long,

190 longest spines 18-34 (25) long. Like S. cobia here the spines reach the posterior extremity.

In S. pacificum spines in mid-ventral region 11-17 (15) long, longest spines 23-32 (27) long; spines smaller in hindbody; reach to posterior extremity.

In S. aaravi the position of spines is again the same as in S. madhaviae, except that anterior spines in mid-ventral region are 21-34

(28) long, longest spines 42 long; spines smaller in hindbody; reach to posterior extremity.

S. pagrosomi again has the same position, anterior spines in mid- ventral region are 4-14 (8) long, longest spines 9-18 (13) long; reach to posterior extremity.

The position of spines described by Bartoli and Bray (2001) for S. cesticillum is entirely different from those specimens described so far.

Here, the area immediately posterior to oral sucker is unspined dorsally and ventrally. Posterior to this area tegument is heavily spined. Spines are strong, with enlarged base, long acuminate, not scale-like, slightly recurved in exposed distal region, posterior to level

191 of ventral sucker, size of spines decrease slowly. Tegument is unspined from level of anterior testis to posterior extremity.

S. pristis and S. minutum have entire surface spined, except margin of ventral sucker aperture (Bartoli and Bray, 2001).

S. ditrematis and S. filiforme have region immediately posterior to oral sucker unspined, while the rest is heavily spined. Spines heavily developed along neck, long acuminate, slightly recurved in exposed distal region, with enlarged and rounded base. Size of spines decreases slowly along hindbody, but still found at posterior extremity

(Bartoli and Bray, 2004).

S. euzeti and S. petimba have unspined regions immediately posterior to oral sucker, around margin of aperture of ventral sucker and join anterior to genital aperture, rest of the position and shape of spines is the same as in S. ditrematis (Bartoli and Bray, 2004).

In the present species the seminal vesicle is bi-partite, posterior part is saccular, anterior long and sigmoid. Cirrus sac is very long reaching in posterior-third of body close to seminal receptacle. While in S. madhaviae, S. votonimoli, S. cobia, S. pacificum, S. aaravi and S.

192 pagrosomi the seminal vesicle is saccular proximally, narrow distally, with thicker wall distally and with distinct sphincter.

S. cesticillum has seminal vesicle undivided and coiled. In S. bicoronatum, S. pristis and S. minutum it is undivided and rectilinear, while S. gaidropsari has proximally coiled seminal vesicle

(Bartoli and Bray, 2001). S. ditrematis and S. filiforme have internal seminal vesicle tubular, undivided, its distal part sometimes sinuous, also sphincter is seen between seminal vesicle and pars prostatica

(Bartoli and Bray, 2004).

In the present species the genital atrium is near the dorsal anterior level of the ventral sucker, while Bray and Cribb, 2003 have described that in S. madhaviae the genital atrium is relatively short, reaching anterior to mid-level of ventral sucker. In S. bicoronatum genital atrium is of moderate depth, reaches to level of posterior part of ventral sucker; S. votonimoli has genital atrium of moderate length, reaching to level of posterior region of ventral sucker or extreme anterior hindbody. In S. nyoomwa genital atrium is short, anterior to ventral sucker. In S. cobia genital atrium is of moderate length, reaches to posterior part of ventral sucker or just into hindbody. In S. petimba and S. pacificum the genital atrium is long, reaches well into

193 hindbody. S. aaravi and S. pagrosomi has short genital atrium which reaches dorsal to anterior part of ventral sucker.

Bartoli and Bray (2001) have described that genital atrium in S. cesticillum is short, thin-walled, devoid of sclerified structures; while in S. bicoronatum, genital atrium is long, thin-walled, apparently devoid of sclerified structures; S. pristis has long genital atrium, walled with villi and scarce, scattered, sclerified structures. In S. minutum genital atrium is short, thin-walled, reaches to about middle of ventral sucker; S. gaidropsari has wide genital atrium, long, thick- walled, apparently devoid of sclerified structures. S. minutum has very long genital atrium, its wall bearing numerous, tiny, unsclerified, villous like structures, devoid of cupolas; S. filiforme also has very long genital atrium in all specimens upto 3.5 times ventral sucker length in some specimens.

Previously, Sogandares-Bernal and Hutton (1959) reported immature

Stephanostomum sp. from Acanthostracion tricornis. This is the first record of a Stephanostomum immature from a pericardial membrane of a fish in New Caledonia. S. casum, S. cesticillum, S. ceylonicum,

S. coryphaenae, S. pristis, S. sentum, S. nipponicum and S. minutum are the only species of the genus known from this continent.

194

S. promicropsi Manter (1947) reported from Promicrops itaira (Jew fish) from Tampa, Bay Florida. Also known from Tortugas, Florida.

An outstanding feature of S. promicropsi is that the vitellaria is ventral to the caeca and are interrupted at the level of testes.

In S. japanocasum reported from Epinephelus sp., oral sucker is smaller than the ventral sucker, 40-44 oral spines are in two rows, largest spines 45-53 microns long; genital pore is median, immediately pre-acetabular cirrus sac extending 1/2 - 2/3rd distance between acetabulum and ovary containing sac-like seminal vesicle, pars prostatica is short, spines with spherical base, cirrus sac joining metraterm dorsal to acetabulum, genital atrium unarmed.

This species is unusual having 46 or more spines, vitellaria extending anterior to the acetabulum. S. casum is similar to S. japanocasum in many respects, although its vitellaria are less extensive. The number of oral spines are fewer and the metraterm longer.

S. provitellosum has interrupted vitellaria, larger oral sucker and long unspined metraterm. S. pseudocarangis has 36 oral spines, oesophagus longer than pharynx, rounded testes, eggs 50-51 microns long; S. microstephanostomum has 150 oral spines in three rows.

195

The present specimens have very long cirrus sac reaching posteriorly close to seminal receptacle in posterior half of the body, bi-partite seminal vesicle, well developed pars prostatica, long ejaculatory duct and genital atrium is also moderately long, genital opening is dorsal, near the anterior level of ventral sucker and can be separated from all the previous species of the genus.

196 SPECIES OF THE GENUS BUCEPHALUS BUCEPHALUS OTOLITHI N.SP. (Figs. 30-31)

Family: Bucephalidae Poche, 1907

Sub-family: Bucephalinae Nicoll, 1914

Genus: Bucephalus Baer, 1827 (Syn. Gasterostomum Siebold, 1848) (Bucephalus Diesing, 1855)

Host: Otolithus argenteus (Sciaenidae)

Location: Intestine

Locality: West Wharf, Karachi.

No. of specimens: 2 specimens from a single host, 20 fishes were examined.

Holotype: JUW – T30

Paratype: JUW – T31

Diagnosis

Body is slender, rounded off behind, covered all over with very fine spines. Rhynchus weakly muscular with 6-7 tentacular appendages, each of which tapers to a rounded point and provided with two (large proximal and small distal) forwardly directed pointed processes.

Mouth opening is centrally located almost in the middle of the body.

Intestine is saccular, relatively large antero-ventrally.

197

Testes are two, close to intestine, sub-globular to elliptical, obliquely tandem, largely in the post-equatorial region. Cirrus pouch is long, cylindrical, extending anteriorly, reaching mid level of ovary, seminal vesicle oval. Pars prostatica is weakly developed and small. Genital lobe is weakly muscular. Genital atrium is prominent. Genital pore is sub-terminal in the posterior region.

Ovary is triangular, dorsally situated on the right side of the second half of the body anterior to the intestine. Uterus coiled mainly in vitellarian zone and on the right side of the cirrus pouch. Eggs are small, numerous, rounded. Vitelline follicles 6-7 on either side arranged in two lateral groups i.e. 12-14 in total number. Pre- equatorial and pre-ovarian in position. Excretory vesicle is large, opening terminally.

Principle measurements of B. otolithi n.sp. (In millimeters)

Entire length: 4.0–4.03

Breadth: 0.49–0.50

Rhynchus: 0.2–0.21 x 0.16–0.165

Stomach: 0.19–0.20 x 0.22–0.23

198 Anterior testis: 0.20–0.21 x 0.13–0.135

Posterior testis: 0.13–0.14 x 0.12–0.125

Cirrus pouch: 0.55–0.56 x 0.08–0.09

Seminal vesicle: 0.11–0.115 x 0.09–0.10

Ovary: 0.12–0.13 x 0.12–0.13

Excretory vesicle: 0.09–0.10

Eggs: 0.011–0.005 x 0.010–0.004

Etymology: The present new species Bucephalus otolithi refers to its host.

Remarks

Species of Bucephalus are known from various parts of the world but only two species B. varicus Manter, 1940 (Zaidi and Khan, 1977) in

Trachynotus bailloni, Caranx sp., Platax sp., Serranus sp.,

Chloroscombrus and Seriola and B. hexalobatus Bilqees et al., 2006 are known from fishes of Karachi coast.

The present species of the genus Bucephalus otolithi can be separated from the previously described species in having 7 tentacles. In B. tritentacularis Srivastava, 1963 tentacles are 3 in number; in B. bhartica tentacles are also 3. In B. gangeticus Srivastava, 1938; B.

199 elacatus and B. keni Agrawal, 1988 tentacles are 4. In B. allahabadensis Srivastava, 1963; B. barina Srivastava, 1938 and B. purshottami there are 5 tentacles. In B. indicus Srivastava, 1938 and

B. jagannathai Verma, 1936 6 tentacles are present. B. octotentacularis Kakaji, 1969 has 8 tentacles and B. aoria Verma,

1936 has 14-22 tentacles. B. tridentacularia Verma, 1936 Kakaji,

1969 has tentacles with lateral processes. B. bagarius Srivastava,

1963 has 7 tentacles as in the present species but this is different in number of vitelline follicles, anterior extent of cirrus sac and position of pharynx and caecum in relation to ovary and testes.

The genus Bucephalus was proposed by Baer, 1827 with Bucephalus polymorphus as type-species. The present new species Bucephalus otolithi reported from Pakistan is different from other species in many ways. B. varicus Zaidi and Khan, 1977 is different in having rhynchus well developed, sucker-like, wedge-shaped anteriorly. Mouth is opening in the middle-third of the body on the left side of the ovary.

Cirrus sac tubular, curved, elongated reaching the middle portion of the posterior testes. Vitelline follicles are rounded, 20 in number, arranged laterally in the first half to the middle of the body. The uterine coils extend to the anterior first part of the body. While in the present specimen the rhynchus is weakly developed, cirrus sac is

200 curved, stomach large post-ovarian and vitelline follicles are large 6-7 arranged pre-equatorially.

B. hexalobatus reported from Pomadasys olivaceum off Karachi coast also differs from the present species in having 6 lobed rhynchus, weakly developed. Vitelline follicles are relatively larger, centrally located anteriorly, nearer to rhynchus 26-28 in number in the anterior- third of the body.

B. barina Srivastava (1938) is the synonym of B. polymorphus described by Gupta and Mehrotra, 1970 from Therapon jarbua and

Pomadasys maculatus from Bay of Bengal should be regarded as identical to B. varicus Mahavi, 1974. The species of B. barina occurred in all the sciaenids recorded by Srivastava, 1938 in

Scatophagus argus.

B. carangis Yamaguti (1970) reported from Caranx lugubris is different from the present specimen in having slender body. Mouth opening is in the posterior part of middle-third of the body. Both the oesophagus and pharynx are short. Seminal vesicle is far away from the posterior testis. Cirrus pouch is confined to posterior half of caudal-third of the body. Pars prostatica is well developed. Ovary is ovoid between pharynx and anterior testis. Vitelline follicles are 15 on

201 each side. B. carangis differs from the present specimen mainly because of the characteristic feature having oesophagus joining the intestine about its middle. In this respect it resembles B. sextentaculatus but differs from it in the tentacles of the rhynchus being 7 instead of 6.

B. carangoides Yamaguti (1970) reported from Sphyraena barracuda is plum, spinose, tapering towards bluntly pointed posterior extremity.

Rhynchus is globular with 7, long tentacles, two nodular protuberances near its base; while in the present specimen the tentacles on the rhynchus are not very long with protuberances. In B. carangoides the vitellaria form 10-16 follicles i.e. 24-30 in total number extending from behind the rhynchus to ovarian zone; while in the present specimen the vitelline follicles are 6-7 in number.

B. kaku Yamaguti (1970) reported from Sphyraena barracuda as compared to the present specimen shows fan-shaped, strong rhynchus with 11 horn-like tentacular appendages along its anterior margin.

Testes are oval, separated by uterus in the anterior part of posterior- third of the body. Cirrus pouch is sub-cylindrical. Seminal vesicle is elliptical. Ovary is post-equatorial. Vitelline follicles are 24-32 in total number on each side. B. kaku closely resembles B. sphyraenae but

202 differs distinctly in tentacles of the rhynchus being more numerous and better developed in the oesophagus, opening into the intestine near its anterior end and in the intestine extending further backwards from the pharynx then forward. The above characteristic feature of B. kaku separates it from the new species of B. otolithi described.

B. sphyraenae Yamaguti (1952) has conical, muscular appendages of different shapes, they are pointed, conical, stout and slender. Cirrus pouch is sub-cylindrical, provided with well developed oblique muscle fibres reaching to posterior testis. Genital atrium opens ventrally by a funnel-shaped passage. Ovary is ovoid, post-equatorial on the right side of the intestine. Uterus reaching as far as anterior- quarter of the body. Vitelline follicles are on either side of the body in middle-third, 13 on the right and 19 on the left. Excretory vesicle tubular, reaching a little further forward than anterior extent of uterus.

B. sextentaculatus Yamaguti (1970) reported from Caranx sexfasciatus is different from the present specimen in having massive rhynchus flattened at the apex but rounded behind with 6 pointed tentacles each with two small processes near its base. Uterus is filling all available space of body, posterior to vitellaria, forming a long, longitudinal loop reaching nearly to rhynchus. Vitellaria are divided

203 into asymmetrical groups of 11-17 each totaling 24-31, extending on each side anterior to pharyngo-intestinal region. In the anatomy of digestive organs it resembles B. kaku but the rhynchus always bears 7 tentacles instead of 11. The above features differentiate it from the present new species.

B. gorgon (Linton, 1905) Eckmann, 1932 reported from Chile:

Seriola lalandi (=Seriola mazatlana). Body is comma-shaped, strongly muscular, spines prominent, rhynchus occupying anterior- third part of the body, broad anteriorly and narrow posteriorly, mouth located ventrally in the middle of the body leading into a pouch-like intestine through a short oesophagus. Anterior testis is larger than the posterior testis lying close to the cirrus sac. Cirrus sac elongated, large, occupying more than posterior half of the body, reaching the level of the ovary, enclosing small seminal vesicle. Uterus coils, occupying large part of the body, almost reaching the posterior part of the anterior rhynchus. Genital pore is terminal. 12 vitelline follicles in a single group situated in the middle of the body. B. gorgon is referred to as B. introversus Manter, 1940. B. introversus was considered synonym of B. gorgon by Bartoli et al., 2005.

204 B. ulua Yamaguti (1970) is reported from Hawaii from the fish

Carangoides ferdau. Body small, plump, bluntly pointed posteriorly where a conspicuous notch is often seen at the excretory pore.

Rhynchus is very strongly muscular with semi-circular row of 7 tentacles. Mouth is in the equatorial region. Cirrus pouch sub- cylindrical reaching to level of intestine. Ovary is ovoid, lateral to the intestine. Uterus is coiled mainly in vitellarian zone and on the right of cirrus pouch. Vitelline follicles are 24-29 in number, i.e. 11-16 in two lateral groups. Excretory vesicle inflated anteriorly, reaching to the level of anterior end of vitellaria or posterior end of the rhynchus.

Genital pore is terminal. This species is very similar to B. carangoides in all respect except the rhynchus in B. ulua which is very strongly muscular. While in the present specimen B. otolithi n.sp. the rhynchus is weakly muscular.

B. margaritae Ozaki et Ishibashi (1934) reported from Brazil in the fish Caranx crysos, Caranx hippos, Caranx latus, Chloroscombrus chrysurus, Menticirrhus americanus, Menticirrhus littoralis,

Micropogonias furnieri, Oligoplites palometa, Oligoplites saliens,

Oligoplites saurus and Urophycis brasiliensis. B. margaritae has spinous body, triangular rhynchus with 2 tentacles on both sides.

205 Small mouth in the centre of the body opens into the small, thin, tubular oesophagus leading into a pouch-like bulbous intestine. The intestine overlaps the posterior, large testis. The two testes are contagious, situated far from the cirrus sac above the mid-dorsal region of the body. Cirrus sac elongated. Uterus and genital pore sub- terminal. Vitelline follicles are 20 in number, arranged laterally in the first half of the middle of the body.

Bucephalus varicus was considered synonym of Bucephalus margaritae Bray, 1984.

B. solitarius reported from Venezuela from fish Caranx crysos has elongated, tubular body with small rhynchus having 5 tentacles.

Mouth is present. Pharynx is small, rounded leading into an elongated intestine through a very small oesophagus. Both the testes are contiguous, equal in size, at the level of the pharynx. Cirrus sac is in posterior part of body, elongated, tubular enclosing an oval vesicula seminalis. Ovary is larger in size, below the mouth and close to it.

Uterus is occupying larger space reaching the anterior second part of the body. Vitelline follicles are 18 in number in groups on either side of the body. Genital pore is sub-terminal.

206

B. urophyci Szidat (1961, 1963) is reported from Argentina from the fish Urophycis brasiliensis Szidat, 1961. Body with prominent, muscular rhynchus having 7 tentacles out of which 3 are pointed backwards. Mouth and pharynx are in the posterior middle part of the body reaching the small intestine. Testes are large, contiguous.

Anterior testis is close to the pharynx. Cirrus sac is elongated, enclosing small seminal vesicle. Ovary is above the pharynx in the middle part of the body. Uterus is with eggs seen in the anterior second part till the posterior part of the body. Vitelline follicles are numerous, not countable. Excretory pore is sub-terminal. While in the present species B. otolithi n.sp. the rhynchus is weakly muscular and vitelline follicles are 12-14 in number.

B. australis (Szidat, 1961) Yamaguti, 1971 reported from Conger orbigniamus (Etchegoin et al., 2005) and Urophycis brasiliensis

(Szidat, 1961). Etchegoin et al., 2005 transferred B. australis to the genus Prosorhynchus based on specimens collected from the type- host from Argentina, considering morphological features not included in the original description.

207

Srivastava (1938) suggested B. varicus Manter, 1940 and B. barina

Srivastava, 1938 as synonym of Bucephalus polymorphus Baer, 1827

Gupta and Mehrotra, 1970.

Species reported from marine fishes of India by Srivastava and

Chauhan, 1973 are B. aoria, B. tridentacularia, B. jagannathai, B. indicus, B. gangeticus, B. barina and B. indicus is the synonym of B. tritentacularis Srivastava, 1963.

Srivastava (1963) described three species under the genus Bucephalus viz. B. bagarius, B. tritentacularis and B. allahabadensis.

Kakaji (1969) described B. octotentacularis under the genus

Bucephalus. The authors have observed a wide range of variations in the individuals of the species described above.

According to Poche (1907) synonyms from other parts of the world are Rhipidocotyle papillosum Wood head, 1929 reported from

Micropterus salmoides is the synonym of B. papillosus.

Gasterostomum is a synonym of Bucephalus.

208

The present new species is distinctly different from other species of the genus in having different morphology of rhynchus, number of tentacles or projections, position of stomach, testes, morphology of cirrus sac, shape and size of seminal vesicle and pars prostatica, position of ovary, number of vitellaria and saccular excretory vesicle.

209 BUCEPHALUS MUJIBI N.SP. (Figs. 32-33)

Host: Cybium guttatum (Cybidae)

Location: Intestine

Locality: West Wharf, Karachi.

No. of specimens: 3 specimens from a single host, 13 fishes were examined.

Holotype: JUW – T32

Paratype: JUW – T33

Diagnosis

Body is elongated, spined anteriorly, rhynchus with small, 7 unequal projections, narrow anteriorly at the pre-vitellarian region ending into a muscular tentacular rhynchus, bluntly pointed posteriorly, widest at the middle of the body. Mouth is almost in the middle of the body.

Pharynx is spherical, close to anterior level of anterior testis. Caecum is tubular, extending anteriorly to post-vitelline region.

Testes are two, tandem, large, spherical, lying close to each other, reaching in the pre-equatorial body region, anterior testis slightly overlapping the posterior testis. Cirrus sac is little far from posterior end, elongated, containing rounded seminal vesicle. Long pars

210 prostatica and slightly large genital atrium, extending anteriorly posterior to posterior testis.

Ovary is rounded, situated laterally, in the middle of the body, close to the anterior testis and the pharynx. Uterus fills most of the posterior three-fourth of the body region reaching anteriorly at the level of vitellaria and posteriorly near about the posterior body region. Eggs occupy large space within the body. Vitellaria consist of 2 lateral fields of large follicles on each side, situated far from the rhynchus.

Excretory opening is terminal.

Principle measurements of B. mujibi n.sp. (In millimeters)

Entire length: 2.85–2.90

Breadth: 0.75–0.80

Rhynchus: 0.20–0.21

Rhynchus to caecum: 1.15–1.16

Rhynchus to vitellaria: 0.77–0.78

Number of follicles in each set: 13

Caecum: 0.27–0.28

Mouth: 0.13–0.14

Anterior testis: 0.25–0.255

211 Posterior testis: 0.22–0.23

Cirrus sac: 0.69–0.695

Seminal vesicle: 0.15–0.155

Ovary: 0.21–0.22

Excretory vesicle: 0.30–0.31

Genital atrium: 0.025–0.051

Eggs: 0.02–0.021 x 0.029–0.030

Etymology: Species name is in the honour of late Dr. K.A. Mujib, of

Department of Zoology, University of Karachi, who was expert in fish craniology.

Remarks

Species of Bucephalus are known from various parts of the world but only two species B. varicus Manter, 1940 (Zaidi and Khan, 1977) in

Trachynotus bailloni, Caranx sp., Platax sp., Serranus sp.,

Chloroscombrus and Seriola and B. hexalobatus Bilqees et al., 2006 are known from fishes of Karachi coast.

The two new species of Bucephalus described here are B. otolithi n.sp. and B. mujibi n.sp., resembles in having rhynchus with 7 tentacular appendages. The other marine and freshwater species found

212 in different parts of the world having 6-7 appendages on the rhynchus are B. sphyraenae Yamaguti, 1952; B. retractilis Yamaguti, 1952; B. varicus Manter, 1940; B. polymorphus Baer, 1827; B. carangoides

Yamaguti, 1970 and B. sextentaculatus Yamaguti, 1970 but other characters which separates the present specimen from other Pakistani species are the presence of mouth which is located centrally in the middle of the body.

In B. mujibi n.sp. cirrus sac is not very long nor reaching the level posterior to ovary, while in B. otolithi n.sp. the cirrus sac is long reaches the level posterior to ovary. Seminal vesicle is round in B. mujibi n.sp. while oval in B. otolithi n.sp. Genital pore is sub-terminal in B. otolithi n.sp. while in B. mujibi n.sp. the genital pore is terminal.

Vitelline follicles are 13 in number in B. mujibi n.sp. while in B. otolithi n.sp. they are 6-7.

B. retractilis also has 7 tentacles at the rhynchus; B. tridentacularia

Srivastava, 1963 has rhynchus with 3 tentacles, smooth, cuticular spines in anterior half of body; B. bhartica has rhynchus with minute spines at the base, cuticular spines extend beyond equatorial region of body; B. gangeticus Srivastava, 1938 has rhynchus with 4 tentacles, 2 dorsal; B. elacatus has rhynchus with 4 tentacles, 2 ventral, short

213 tentacles without spines; B. keni has rhynchus with 4 tentacles, each tentacle tridient provided with two, pointed arrow-headed lateral processes; B. allahabadensis Srivastava, 1963 has rhynchus with 5 tentacles, ovary anterior to intestinal sac; B. barina Srivastava, 1938 has rhynchus with tentacles having rose thorn-like spines arranged radially; B. purshottami has rhynchus with tentacles having simple spines at the base and tip; B. indicus Srivastava, 1938 has rhynchus with 6 tentacles, cirrus sac extends upto anterior limit of posterior testis; B. jagannathai Verma, 1936 has rhynchus with 6 tentacles, cirrus sac extends upto anterior margin of anterior testis; B. bagarius

Srivastava, 1963 has 7 tentacles at the rhynchus; B. octotentacularis

Kakaji, 1969 has 8 tentacles at the rhynchus without lateral processes, cirrus sac extend upto posterior end of ovary; B. tridentacularia

Verma, 1936 has tentacles with lateral processes, cirrus sac extends upto middle of anterior testis and B. aoria Verma, 1936 has 14-22 tentacles at the rhynchus.

In the species B. varicus (Zaidi and Khan, 1977) found in Pakistan the rhynchus is well developed. In B. hexalobatus Bilqees et al., 2006 the rhynchus has 6 lobes, weakly developed tentacles. While the present specimen B. mujibi n.sp. consist of small, 7 unequal projections with muscular rhynchus.

214

In B. varicus mouth is in the middle-third of the body, while in B. hexalobatus it is centrally located anteriorly. In B. retractilis the mouth is situated in the mid-body at level of anterior testis or ovary.

In B. carangis mouth opening is in the posterior part of middle-third of body. In B. gorgon and B. introversus mouth is located ventrally in the middle of the body leading into a pouch-like intestine through a short oesophagus. In B. margaritae small mouth is in the center of the body. In B. solitarius mouth is again in the middle of the body. In B. sphyraenae the mouth is post-equatorial. In B. urophyci mouth and pharynx are in the posterior middle part of the body. While in the present specimen B. mujibi n.sp., the mouth is almost in the middle of the body.

The position of cirrus sac in B. varicus and B. barina is tubular, curved and elongated reaching the middle portion of the posterior testis. In B. hexalobatus the cirrus sac is prominent, elongated, slightly curved, relatively large. In B. sphyraenae and B. kaku the cirrus pouch is sub-cylindrical. In B. retractilis it is cylindrical. In B. carangis cirrus pouch is confined to posterior half of caudal-third of the body. In B. gorgon and B. introversus it is elongated, large, occupying more than posterior half of the body reaching the level of

215 the ovary. In B. margaritae, B. urophyci and B. solitarius cirrus pouch is elongated. While in the present specimen B. mujibi n.sp., the cirrus sac is elongated and little far from posterior end of the body.

The position of vitelline follicles show quite difference in the location as in B. varicus and B. barina they are large 6-7 arranged pre- equatorially. In B. hexalobatus the number of vitelline follicles exceeds very much which reaches up to 26-28 centrally located in anterior-third part of the body. In B. carangis the vitelline follicles are

15 on each side. In B. carangoides and B. kaku the follicles are 10-16 i.e., total number is about 24-30. In B. sphyraenae the vitelline follicles are 13 on the right and 19 on the left. In B. sextentaculatus the vitellaria is divided into asymmetrical groups of 11-17 each totaling 24-31, extending on each side anterior to pharyngo-intestinal region. In B. gorgon and B. introversus 12 vitelline follicles in a single group situated in the middle of the body. In B. margaritae the number of follicles are same as in B. varicus i.e. 20 in number arranged laterally in the first half of the middle of the body. In B. solitarius vitelline follicles are 18 in number in groups on either side of the body. In B. urophyci they are uncountable. While in the present specimen B. mujibi n.sp., the vitelline follicles are in groups of 13 on each side of the body.

216

B. keni Agrawal, 1988 reported from the freshwater fish Mystus aor

(Ham.) fairly differs from all the known species of Bucephalus in the absence of cuticular spines all over the body. It resembles B. gangeticus and B. elacatus in having 4 tentacles but differs in shape of tentacles which are arrow-headed and provided with two lateral processes. Mouth is small in the posterior second part of the body.

Cirrus sac elongated, one-fourth of the body length reaching anteriorly up to middle of anterior testis. Vitelline follicles are 13-15 in number on each side anterior to intestine. Excretory pore is sub-terminal. All the above diagnostic features separate them from the present specimen

B. mujibi n.sp.

The present species shows great variations as compared to previously described species in the morphology of rhynchus and associated structures, position and number of vitelline follicles, anterior extension of cirrus sac.

217 SPECIES OF THE GENUS PROSORHYNCHUS PROSORHYNCHUS MAGNACIRRUS N.SP. (Fig. 34)

Family: Bucephalidae Poche, 1907

Sub-family: Prosorhynchinae Nicoll, 1914

Genus: Prosorhynchus Odhner, 1905

Host: Psettodes erumei (Psettotidae)

Location: Intestine

Locality: West Wharf, Karachi.

No. of specimens: 2 specimens from a single host, 13 fishes were examined.

Holotype: JUW – T34

Paratype: JUW – T35

Diagnosis

Body is oval, plum, pointed at both ends, widest at the middle, anteriorly showing small rhynchus. Rhynchus tegument spinosed, mouth in the anterior one-third of the body. Pharynx is spherical, caecum is oval, reaching anteriorly to the level of the vitellaria.

Testes are two, oval to rounded, clearly separated from each other, situated antero-lateral to the ovary. Cirrus sac is very large, reaching in the pre-equatorial body region, proximal end at the level of

218 posterior vitelline follicles. Seminal vesicle is also large, rounded.

Pars prostatica is straight with gland cells on either side of elongated tube. Ejaculatory duct is short, opening inside the genital atrium.

Genital atrium is very large, muscular, enclosing genital lobe. Genital pore is sub-terminal. Excretory pore is terminal.

Ovary is rounded, larger than the testes, mainly antero-lateral to cirrus sac and close to the anterior testis. Uterus fills most part of the body with large, operculate, tanned eggs, almost occupying the whole body except the rhynchus and genital atrium. Vitellaria consist of 2 lateral fields of large follicles, usually symmetrical, having 7 follicles on each side, slightly irregularly arranged, extending anteriorly far behind the rhynchus to the level of the anterior part of the cirrus sac.

Excretory vesicle is terminal, anterior extent of vesicle obscured by eggs.

Principle measurements of P. magnacirrus n.sp. (In millimeters)

Entire length: 1.18–1.19

Breadth: 0.635–0.64

Rhynchus: 0.08–0.081 x 0.05–0.051

Rhynchus to caecum: 0.013–0.0131

219 Rhynchus to vitellaria: 0.014–0.017

Number of vitelline follicles: 7 follicles on each side

Caecum: 0.09–0.10 x 0.077–0.08

Pharynx: 0.04–0.05

Mouth: 0.009–0.010

Ovary: 0.22–0.225 x 0.21–0.22

Anterior testis: 0.10–0.11 x 0.091–0.10

Posterior testis: 0.115–0.12 x 0.10

Cirrus sac: 0.43–0.45 x 0.11–0.12

Seminal vesicle: 0.14–0.139 x 0.08–0.09

Genital atrium: 0.149–0.150 x 0.150–0.154

Eggs: 0.050–0.052 x 0.030–0.031

Etymology: The present new species Prosorhynchus magnacirrus refers to its large-sized cirrus sac.

Remarks

Species of Prosorhynchus reported from Pakistan are P. platycephali

(Yamaguti, 1934; Srivastava, 1938; Bilqees, 1981) from

Platycephalus scabar; P. thapari Manter (1953) Bilqees, 1981 from

Plectorhynchus cinctus; P. longus Velasquez (1959) Bilqees, 1981

220 from Psettodes erumei and P. erumenis Bilqees, 1976 from Psettodes erumei.

The present specimen P. magnacirrus n.sp., is different from P. platycephali in having oval body, pointed at both ends, while P. platycephali is long and slender. In P. platycephali mouth is in the middle, cirrus sac is one-fifth of the whole body, containing pear- shaped seminal vesicle at the base. Vitelline follicles are 13-17, while in the present specimen mouth is more anterior, cirrus sac is very large, vitelline follicles are 7 in number on both lateral sides. Eggs are numerous, scattered through out the body except at the rhynchus and genital atrium.

The other species reported from Pakistan is P. thapari, here the mouth is in middle of body. Cirrus sac is one-fourth of the total body length containing elongated seminal vesicle. Vitelline follicles are 12-14 arranged laterally at the side of the intestine and is also distinct from the present new species.

The third species reported from Pakistan is P. longus which is very long, equally broad through out its length. Mouth is in the middle, cirrus sac is sigmoid, seminal vesicle oval and genital atrium is large.

221 The number of vitelline follicles are 14-20 and are situated posterior to pharynx or intestine in the posterior half of the body and is also different from the present species.

The fourth species reported from Pakistan is P. erumenis which is also separated from present species in having sub-cylindrical body, mouth situated in the second part of the anterior region. Ovary is large, inter-testicular, cirrus sac is cylindrical, muscular, large containing large seminal vesicle. Vitelline follicles are large, arranged in two lateral rows, one of 17 and other of 22 follicles.

Differences in other species of Prosorhynchus are also noted. P. australis originally described by Szidat, 1961 from Urophycis brasiliensis Kaup, 1858 redescribed by Etchegoin et al., 2005 from the fish Conger orbignianus. P. australis is different from the present specimen in having elongate body, conical rhynchus with two lateral and two central bands of dorso-ventral muscles surrounding the central funnel. Mouth is in the middle of the body surrounded by uterus. Cirrus sac is thick-walled, sinistral, containing tubular, looped seminal vesicle, long pars prostatica and numerous prostatic cells. A small pre-testicular ovary is located to the right of slightly diagonal

222 testes. Vitelline follicles are 24-32, large in size, arranged in the form of an arch in the anterior part of the body.

P. australis most closely resembles P. longisaccatus Durio et Manter,

1968 from which it can be readily differentiated by the latter having the ovary to the right or partly posterior to the anterior testis, instead of pre-testicular, the testes are clearly diagonal and the cirrus sac larger relative to body length. A small pre-testicular ovary is located to the right of slightly diagonal testes, which together with the shape of the rhynchus precludes its identification as either P. australis or P. longisaccatus.

Yamaguti, 1971 transferred P. australis to genus Bucephalus but

Etchegoin et al., 2005 did not agree with the transfer of P. australis to genus Bucephalus Baer, 1827 because P. australis lack a ring of retractile tentacles in the rhynchus which is a diagnostic feature of

Bucephalus (Overstreet and Curran, 2002).

In P. chorinemi (Yamaguti, 1952) from Scomberoides tala, rhynchus shows two distinct regions, anterior disc-like part and a posterior conical part, which is different from the type material. In P. indicus

(Madhavi, 1974) rhynchus is described as a pad-like structure. P. indicus has a slender, elongate body with long neck. Rhynchus is

223 small, triangular in shape. Anterior part is disc-like, produced laterally into small papillae, posterior part conical. Mouth opening is median, slightly behind mid-body, cirrus sac reaching level of posterior testis containing ovoid seminal vesicle. Straight pars prostatica and numerous prostatic cells. Vitelline follicles 12-13 on the right side and

16-19 on the left, confined to posterior half of body, commencing at level of Mehlis’ gland, extending anteriorly beyond level of gut.

The species of Prosorhynchus having elongate body and tandem distribution of gonads are P. facilis (Ozaki, 1924); P. tsengi Chin,

1933 and P. platycephali (Yamaguti, 1934) these are very similar to

P. indicus as described above.

P. platycephali is synonymized with P. tsengi. P. indicus differs from

P. tsengi and P. facilis in having long neck not occupied by uterine coils, the gut extending posteriorly from the oesophagus, the more posteriorly situated gonads and the much smaller eggs, all the diagnostic characters separate the present new species P. magnacirrus n.sp., from the above described species.

Hanson (1950) considered P. atlanticus a synonym of P. pacificus on the basis of variations in the size of eggs in specimens from Bermuda.

Winter, 1960 and Overstreet, 1969 accepted this synonymy although

224 Nahhas and Cable, 1964 were of the opinion that the two species are distinct. The specimens had eggs measuring 27-33 by 16-20 in size and support the synonymy. Egg sizes reported for P. pacificus after accepting the synonymy, range 24-36 by 12-24. P. luzonicus

Velasquez, 1959 can be differentiated from P. pacificus on the basis of slightly larger eggs (30-39 x 17-24). In view of the variation noted in the egg size of P. pacificus, P. luzonicus is considered a synonym of P. pacificus. Hafeezullah and Siddiqi, 1970 reported this species as

P. atlanticus from Epinephelus tauvina from Karwar (Arabian Sea).

P. crucibulum and P. aculeatus are common parasites of the Conger eel. Of the various morphological features, the scaly tegument is particularly important, as it is typical of bucephalids such as in

Prosorhynchus sp., (Mathews, 1973). Several previous authors have referred to the presence of spines on the tegument of these two species of Prosorhynchus (Nicoll, 1910; Jones, 1943; Bray, 1973; Mathews,

1973) but detailed study is scarce. The spines on the anterior body and rhynchus were the largest in P. aculeatus (Jone, 1943) while in P. crucibulum the spines are apparently shorter and exhibit a different shape, being wider than long. Another interesting feature of these two species of Prosorhynchus is that their spines are of a similar size. P.

225 crucibulum has no spine in the middle of the ventral indentations of the rhynchus.

As compared with the present specimen P. magnacirrus n.sp., P. crucibulum is different in having flattened, elongate body, rhynchus with five indentations, body armed with spines, having a mid-ventral mouth (typical of bucephalid), muscular pharynx and a sac-like intestine. Prepharynx and pre-pharyngeal pouch are present associated with pre-pharyngeal glands. Most obvious structure is a papilla-like structure located on the anterior pharyngeal wall. The same structure was not seen in P. aculeatus for which Mathews, 1973 reported that the pre-pharyngeal glands are weakly developed.

P. manteri reported from Trichiurus naumela, is also different from the present new species in having the excretory vesicle extending to the level of the ovary.

In P. maternus reported from Epinephelus malabaricus rhynchus is elongated with 5-6 muscular lobes. Mouth is at the level of ovary or anterior testis, well inside anterior half of body. Cirrus sac is elongated, parallel-sided, never reaching posterior testis. Pars prostatica long in two distinct parts, proximal part coiled at level of seminal vesicle, with few gland cells, distal part wider surrounded by

226 dense layer of gland cells and ovary is oval, mainly antero-lateral to anterior testis.

P. maternus when compared with the present species is different in many features. In P. magnacirrus n.sp. the rhynchus is simple without lobes. Mouth is far from the ovary and the seminal vesicle. Cirrus sac is very large, proximal end reaching the level of ovary and posterior testis. Vitelline follicles are 7 in number on each side as compared to the 13 follicles in P. maternus. Uterus covers the whole body with large eggs as seen in the present specimen but in P. maternus it is restricted only to the post-ovarian region.

P. pacificus has more posterior mouth, larger cirrus sac and larger eggs. Vitellaria reach the rhynchus and the cirrus sac reaching to or almost to the posterior testis (Madhavi, 1974).

Hanson (1950) considered P. atlanticus a synonym of P. pacificus and Winter, 1960; Overstreet, 1969 and Madhavi, 1974 agreed P. atlanticus differs from P. maternus. In P. atlanticus the vitellarium reaches the rhynchus, the cirrus sac reaches to or almost to the posterior testis, the mouth is more posterior, the rhynchus is relatively longer, the cirrus sac is relatively larger and several other ratios do not

227 overlap (Bray and Justin, 2006). The above diagnostic feature also separates it from the new specimen P. magnacirrus n.sp.

In other species namely, P. luzonicus Velasquez, 1959 the vitellaria reach the rhynchus and the cirrus sac reaches almost to the posterior testis; while in P. gonoderus the position of mouth is more anterior, vitellaria and the gonads are also more anterior than the present species (Manter, 1940a).

P. longicollis has tiny button-like rhynchus, the mouth is situated well in the posterior-third of the body, the testes are relatively close to the posterior extremity and the cirrus sac overlaps the gonads as far as the ovary (Yamaguti, 1953).

Other species differ from P. maternus in the following characters: In

P. mcintoshi the cirrus sac overlaps the posterior testis and the mouth is equatorial (Velasquez, 1959). P. bulbosus differs in being narrower, with a more posterior vitellarium, an equatorial mouth and testes in the posterior half of the body (Kohn, 1961; Kohn, 1967). P. promicropsi differs in much more posteriorly situated vitellarium, with the caecum reaching distinctly anterior to it and more posteriorly

228 situated mouth and gonads (Manter, 1940b). P. mizellei from a cyclopterid, is a squat worm, with the relatively very large cirrus sac reaching to the ovary, a small rhynchus and eggs 32-40 µm long

(Kruse, 1977). P. kahala, from a carangid, is an elongate worm, with all the gonads in the posterior half of the body, the mouth is in the posterior-quarter of the body and the cirrus sac overlapping the anterior testis (Yamaguti, 1970).

In P. freitasi the vitellarium reaches only to the equatorial level and most of the organs are crowded into the posterior half of the body, the rhynchus is relatively tiny and the cirrus sac reaches to the posterior testis (Nagaty, 1937). P. longisaccatus has a relatively large rhynchus, the vitellarium overlaps the rhynchus and the long cirrus sac reaches almost to the equatorial level overlapping the gonads. In

P. serrani the uterus reaches the previtelline region and the cirrus sac overlaps the posterior testis (Durio and Manter, 1968) and it is different from the present new species.

P. bengalensis has elongated rhynchus. Mouth is sub-median, post- equatorial. Pharynx is sub-globular. Cirrus sac is elongated, tubular and extending up to mid-level of anterior testis. Pars prostatica is

229 long, genital atrium pear-shaped, enclosing genital lobes. Vitelline follicles are 10 on the right and 13 on the left side. Uterus is extending far anterior to vitellaria.

The present specimen P. magnacirrus n.sp., is different from P. bengalensis in which the body is oval, mouth is anterior one-third of the body. Testes are clearly separated from each other while in P. bengalensis they are contiguous. In P. magnacirrus n.sp., the vitelline follicles are 7 on each side, while in P. bengalensis they are 10 on right side, and 13 on left side.

The species reported from different parts of the world which have elongated body and tandem position of gonads, are P. crucibulum

(Rud., 1819) Odhner, 1905; P. facilis Ozaki, 1924; P. tsengi Tsin,

1933; P. platycephali Yamaguti, 1934; P. truncatus Verma, 1936; P. arabianum Srivastava, 1933; P. manteri Srivastava, 1938; P. longus

Velasquez, 1959; P. indicus Madhavi, 1974; P. orientalis Gupta and

Ahmad, 1976; P. caballeroi Gupta and Ahmad, 1976; P. erumenis

Bilqees, 1976; P. clarias Wang, 1980 and P. chorinemi Gupta and

Tewari, 1985. While in the present specimen the body is oval and other diagnostic features are entirely different.

230 In P. vinodae again the body is elongated, cylindrical, aspinose. The rhynchus is elongated while in P. magnacirrus n.sp. the rhynchus is small and oval. In P. vinodae the mouth is median or sub-median, pre- equatorial. Cirrus sac is extending up to anterior margin of posterior testis. Vitelline follicles are 14 on right and 16 on left side extending up to anterior margin of ovary. The above diagnostic characters separate it from the new species P. magnacirrus described here.

231

DISCUSSION

The digenetic trematodes or flukes are among the most common and abundant of parasitic worms, second only to nematodes in their distribution.

They are parasites of all classes of vertebrates, specially marine fishes and nearly every organ of the vertebrate body can be parasitized by some kind of trematodes as adult or juvenile. Digeneans development occurs in two hosts.

The first is a mollusk, or very rarely, an annelid. Many species include a second and even a third intermediate host in their life cycles. The adult parasite produces eggs and the eggs are passed in the definitive host’s faeces. Several species cause economic losses to society through infections of domestic animals and others are medically important parasites of human.

This indicates the need to investigate the trematode fauna of this region and a rich trematode fauna of fishes of Karachi Coast awaits serious study.

Because of their importance digenea have stimulated vast amounts of research and the literature of this group is immense.

Trematodes are one of the important parasites infecting fishes throughout the world and these are also common in fishes of Karachi coast. Trematodes show a great diversity in fishes of Karachi coast (Table 1).

269

Thousands of species of trematodes of marine fishes are known and these are world wide in distribution. 75 species of trematodes of marine fishes are already known from Pakistan. These belong to 10 genera and 6 families. In addition to this 18 new species including 2 already known are also identified during the present studies. This is an important addition to scientific literature.

Hemiurids are the most common trematodes revealed during the present studies, as out of 18 new species, 11 are hemiurid trematodes; 5 species are genus Lecithocladium, 4 of Tubulovesicula and 1 species Stomachicola

Yamaguti, 1934 and 1 species of Prosogonotrema Bilqees, 1980 were also recovered during the present studies. Stomachicola muraenesocis reported here appears to be a common species in marine fishes of Pakistan (Bilqees,

1981; Bhutta and Khan, 1975). Previously several species of hemiurids have been described from Pakistan (Bilqees, 1981; Shaukat and Bilqees, 2006).

The genus Decemtestis Ozaki, 1925 was not previously known from

Pakistani marine fishes and is reported here for the first time.

270

5 species of genus Lecithocladium are described here. While previously 13 species (Bilqees et al, 2005) are known from Pakistan. Here it appears to be a more common hemiurid genus.

4 species of the genus Tubulovesicula have been previously reported from

Pakistan (Zaidi and Khan, 1977; Bilqees, 1981). While during the present studies 4 new species have been recovered and described from the Karachi coast.

2 species Prosogonotrema previously reported from fishes of Karachi coast are P. diacanthi Bilqees and Durrani, 1980 and P. karachiense Bilqees and

Durrani, 1980 while another species of Prosogonotrema has also been recovered during the present studies.

Two species found in Pakistan in marine fishes are Plagioporus

(caudotestis) mujibi Bilqees, 1972 and Plagioporus heterorchis Bilqees,

1977; while one species is also known from fresh water. A new species of the genus Plagioporus is reported and described here in detail.

271

Although genus Stephanostomum is one of the largest genus having more than 112 species all over the world (Saoud et al., 2002; Bray and Cribb,

2003) only 2 species are known from Pakistan (Bhutta and Khan, 1977;

Bilqees, 1972) and here a third new species is described.

Species of Bucephalus are known from different parts of the world but only

2 species are known from fishes of Karachi coast (Zaidi and Khan, 1977;

Bilqees et al., 2006). Where as in the present study two new species are being reported.

4 species of Prosorhynchus are known from marine fishes of Pakistan

(Bilqees, 1976; 1981); while during the present studies a new species is reported and described from Karachi coast.

The new species of various genera described here are separated from other species of the genera on the basis of differences in diagnostic features. As mentioned previously more emphasis is given on morphological variations and position of various organs, genital pore, sucker-width ratio and egg sizes rather than body sizes because it may differ with the age of the specimens as

272

well as during fixing and processing. As observed during the present studies the rate of infection in carnivorous fishes was high (Table 3).

20th century has witnessed tremendous advances in basic knowledge of infectious diseases but still many problems remain. Some old plagues have vanished, while others have been routed by rising standards of living and the great advances in preventative medicine. Yet there is no final victory over infection, for elimination of one problem highlights another and the delicate balance between man and micro-organisms remains. More over the spread of air travel is such that infections can readily invade from distant lands.

(Crewe and Haddock, 1985).

Parasitic diseases have always been a major cause of human suffering and their effects have sometimes markedly influenced historical events. Even today parasitic diseases are among the most important causes of human morbidity (Crewe and Haddock, 1985).

There are many indigenous parasites in countries such as Great Britain and many more tropical diseases are imported in people coming from endemic areas. It must be remembered that one can travel half way round the world in a period well within the incubation period of most parasitic diseases, so

273

visitors can easily import parasitic infections into new countries. In addition many thousands of new residents, who may be infected with parasites, currently live in non-tropical countries.

A relatively small number of these parasitic infections can be passed on in the new countries and so could become public health problems rather than just a matter of concern for the infected individual who brought in the disease. Most of them are transmitted by contamination of food and water and in general their spread is restricted by the relatively high standard of personal hygiene in developed countries.

337 generic infectious diseases afflict human beings world wide. 93 of these infections are due to parasites that can be passed on to other humans

(Berger, 2006).

Parasitic infections continue to cause human disease, disfigurement and death on a marine scale in many parts of the world. Parasitic diseases are endemic mostly in the under-privileged countries of the tropics, afflicted with poverty, ignorance, over crowding and poor hygiene, all of which favour survival and spread of parasites and their vectors (Jaypee, 2002).

274

Parasitic infections may remain in-apparent or give rise to clinical disease.

Clinical disease may sometimes be due to trauma inflicted by parasites.

Migration of helminth larvae through the lungs may rupture many pulmonary capillaries and cause considerable extravagation of blood.

Schistosome eggs with their hooks tear vesicle blood vessels and produce haematuria.

A few parasitic infections have been shown to lead malignancy. The liver fluke Clonorchis and Opisthorchis may induce bile duct carcinoma and schistosoma haematolium may pave the way for bladder cancer.

Paragonimus westermani is another trematode causing pulmonary paragonimiasis or chronic cough leading to pulmonary tuberculosis

(Chatterjee, 1981).

Fish born trematode (FBT) infections are a serious public health problem in several parts of the world. It is estimated that about 40 million people are affected world wide (WHO, 1995) endemic in both Europe and Asia.

Situation is of great concern particularly in Thailand, Taiwan, Western

Siberia, Kazakhistan, Ukraine, Vietnam, China, Korea where the first born

275

parasitic diseases are highly prevalent and an estimate of up to 9 million people are infected in the North-eastern region of the country is the major focus of infection (WHO, 1995).

The aquaculture and fishery industry is seriously threatened by the public health concerns due to the trematode infections. The burden of producing fish which is safe for human consumption is on the fish producers and technologists due to the preventable nature of the problem. The eggs of these trematodes enter aquatic system through contamination by excreta of man and animals. If this can be prevented, the fish would be free from the trematodes. Nevertheless, the absence will have to be demonstrated by fish inspecting agencies.

Determining the safety of fish with respect to trematode infections currently involved detailed physical examination of fish for metacercaria of trematodes. This is a rather tedious procedure and requires considerable taxonomic expertise.

Fish is one of the major source of protein for local people in several countries in Asia in particular Thailand, China, Laos and Vietnam. Among

276

the naturally occurring cultured fish, the dominating species successfully breed and reproduce in natural water reservoirs, streams, rivers and are being caught daily. It is estimated that massive amount of fish is consumed not only by villages in communities in some areas but it is being distributed throughout the region. This is one of the essential factor influencing the rate of re-infection in several communities (Sornmavi et al., 1984; Uptham et al.,

1988). It is generally believed that the source of FBT infection are wild. The evidence leading to that believe were, records of the infection and abundance of metacercariae of various trematodes in wild fish species have been reported in Thailand, China and limited information is also available in

Laos (Gibido et al., 1991; WHO, 1995).

The fact that FBT is endemic in under developed and developing countries and the infection creates significant public health problems in several parts of the world is well documented. At the same time, these countries also serve as a major food producers and many of them export aquaculture product particularly fresh water and brackish water fish to feed the rest of the world. Thus there is a high possibility that culture fish reported from

Asia may be contaminated by FBT and transmitting the FBT to consumer both within the country and overseas. With the initiatives of FAO,

277 preliminary studies in Chiangmai, Thailand and in Vietnam demonstrated quite clearly that cultured fish could be a potential source of FBT infection

Khamboonruang et al., 1995; Son et al., 1997. Therefore to tackle this tissue from the root of the problem, it is important to ensure that the endemic and fish producing country be aware of the FBT. To increase the awareness in each endemic country of FBT, it is essential to conduct message under which the effective control strategy can be designed and implemented.

278

REFERENCES

Agrawal V.P, 1999. Recent trends in Aquaculture, Meerut, India. Soc. Bio-sci., pp. 1-3.

Ahmad, J. 1981. Studies on digenetic trematodes of marine fishes from the Bay of Bengal. Part-XVI, Rev. Parasit., 42: 403-413.

Ahmad, J. 1988. Two new species of Allodecemtestis (Digenea: ) from marine fishes of Bay of Bengal. Ang. Parasit., 29: 235-239.

Ahmad, J. 1990. Four new Opecoelid from marine fishes of Arabian Sea off the Panjim coast Goa, India. Proc. Parasit., 9: 49-63.

Akhmerov, A.K. 1959. A new genus of trematode fish, Amurotrema. Trudy Gel’mint. Lab. Akad. Nauk. SSSR., 9: 45-48.

Akhmerov, A.K. 1960. New trematodes from fish in the Amur river. Helminth., 2(3-4): 286-294.

Allison, F.R. 1966. A new species of adult Allocreadidae from Octopus maorum Hutton. Rec. Canterburg Mus., 8(2): 81-85.

Al-Yamani, F.Y. and Nahhas, F.M. 1981. Digenetic trematodes of marine fishes from the Kuwaiti coast of the Arabian Gulf. Kuwait Bull. Mar. Sci., 3: 1-22.

279

Amato, J.F.R. 1983. Digenetic trematodes of percoid fishes of Florianopolis Southern Brasil- Pleorchiidae, Didymozoidae and Hemiuridae, with the description of three new species. Rev. Brasileira Biol., 43: 99-124.

Arora, D.R. and Arora, B. 2005. Med. Parasit. CBS Publishers and distributors. New Delhi. Bangalore (India), pp. vii-viii.

Baer, K.E. VoN 1827. Beitrage zur kenntinis dez neideren. Their Nova Acla Leop-Carol., 13: 523-762.

Bartoli, P. and Bray, R.A. 2001. Contribution to the knowledge of species of the genus Stephanostomum Looss, 1899 (Digenea: Acanthocolpidae) from teleosts of the Western Mediterranean, with the description of S. gaidropsari n.sp. Sys. Parasit., 49: 159-188.

Bartoli, P. and Bray, R.A. 2004. Four species of Stephanostomum Looss, 1899 (Digenea: Acanthocolpidae) from Seriola dumerili (Risso) (Teleostei: Carangidae) in the Western Mediterranean, including S. euzeti n.sp. Sys. Parasit., 58: 41-62.

Bartoli, P, Gibson, D.I. & Bray, R.A. 2004. Redescription of Pleorchis polyorchis (Stossich, 1889) Digenea: Acanthocolpidae, a rare and poorly known parasite of the intestine of Sciaena umbra L. (: Sciaenidae) from the Western Mediterranean Sea. Syst. Parasit., 58: 81- 90.

280

Bashirullah, A.K.M. and D’Silva, J. 1973. Two new parasites of the genus Lecithocladium Luhe, 1901 (Hemiuridae). Jap. J. Parasit., 22: 108-110.

Beneden, van. 1870. Les poisons des cotes de Belgique, leurs parasites et leurs commensaux. Mem. Acad. Roy. Belg., 38: 100 pp.

Berger and Mars, 2006. Human Parasit. Dis. Jones and Bartlett Publ., U.S.A., pp. 11-12.

Bhattia, R. Med. Parasit., Jaypee Publ. Fourth Ed., U.P. India, pp. 11.

Bhutta, M.S. and Khan, D. 1975. Digenetic trematodes of vertebrates from Pakistan. Bull. Dept. Zool. Univ. Punjab, 8: 1-175.

Bilqees, F.M. 1970. Monodhelmis elongatus a trematode from Erethistes elongata of Karachi coast. Pak. J. Zool., 2: 43-45.

Bilqees, F.M. 1971a. Marine fish trematodes of West Pakistan IV. Description of three new genera and species. Pak. J. Sci. Ind. Res., 14: 254-257.

Bilqees, F.M. 1971b. Marine fish trematodes of Pakistan VI. Two new species of the genus Prosorchis Yamaguti, 1934. Pak. J. Sci. Ind. Res., 14: 258-260.

281

Bilqees, F.M. 1971c. Marine fish trematodes of Pakistan VII. Ectenurus crenidensis, a new species from Crenidens indicus (Day), of Karachi coast. Pak. J. Zool., 14: 512-513.

Bilqees, F.M. 1971d. Marine fish trematodes of Pakistan 1. A new species of the genus Podocotyloides Yamaguti, 1934 sensu Pritchard, 1966. Agric. Pak., 22: 253-257.

Bilqees, F.M. 1971e. Marine fish trematodes of West Pakistan. II & III. Description of four new genera and sixteen species (Hemiuridae) with notes on the histology, segmentation, asexual multiplication and regeneration of some of them. Karachi: Agric. Res. Counc. Govt. Pak., pp. 1-55.

Bilqees, F.M. 1972. Marine fish trematodes of West Pakistan VIII. Description of thirteen new species including a new genus Pseudocoitocaecum from fishes of Karachi coast. In: Bilqees, F.M. et al. (Eds) Helminth parasites of some vertebrates chiefly from fishes of West Pakistan. Karachi: Agric. Res. Counc. Govt. Pak., pp. 1-40.

Bilqees, F.M. 1972a. Marine fish trematodes of W. Pakistan X. Tormopsolus spatulatum sp. n. (Acanthocolpidae: Acanthocolpinae) from a fish of Karachi coast. Proc. Helm. Soc. Wash., 39: 138-140.

Bilqees, F.M. 1972b. Marine fish trematodes of W. Pakistan IX. Two new species Plagioporus (Caudotestis) mujibi and Notoporus hystrix (Digenea: Allocreadiidae). Bull. Agri. Res. Counc. Pak., pp. 105-110.

282

Bilqees, F.M. 1972c. Marine fish trematodes of W. Pakistan XI. Helicometrina otolithi sp. n. (Allocreadiidae) and Sterrhurus stromatei sp. n. (Hemiuridae) from Otolithus argenteus (C.V.) and Stromateus sinensis (Euph.). Acta Parasit. Pol., 20: 337-341.

Bilqees, F.M. 1974a. Redescription of the sub-family Oriento- diploproctodaeinae Bhutta and Khan, 1970. Based on four new species including two new genera. Sindh Univ. Res. J., 8: 31-42.

Bilqees, F.M. 1974b. Morphological variations in Bianium plicitum (Linton, 1928) Stunkard, 1930 (Trematodes) from a fish off Karachi coast. Acta Parasit. Pol., 26: 305-310.

Bilqees, F.M. 1976a. Olivacreadium n. gen. (: Opecoelidae) based on two new species from the fishes of Karachi coast. Norw. J. Zool., 24: 33-36.

Bilqees, F.M. 1976b. A new trematode of the genus Helicometrina Linton, 1910 (Opecoelidae) in the porcupine fish Chilomycterus hysterix (L.) of the Karachi coast. Norw. J. Zool., 24: 37-40.

Bilqees, F.M. 1976c. Two trematodes of the genus Prosorhynchus Odhner, 1905 (Bucephalidae) including a new species P. erumenis from the fish Psettodes erumei (Bl. Schn.) of the Karachi coast. Norw. J. Zool., 24: 345-348.

283

Bilqees, F.M. 1976d. A comment on the relationship of Prosorhynchus thapari [Manter, 1953 (Trematoda)] from the fish Plectorhynchus cinctus (T.S.) of the Karachi coast, with a note on its surface ultra-structure. Proc. Pak. Acad. Sci., 13: 29-33.

Bilqees, F.M. 1977a. Pleorchis ghanensis Fischthal et Thomas, 1968 (Trematoda: Pleorchidae, Poche, 1926) from a new fish host Pseudosciaena diacanthus (Lac.) from Karachi coast. Pak. J. Zool., 9: 107.

Bilqees, F.M. 1977b. The synonymy and the surface ultra-structure of the trematode Prosorhynchus platycephali (Yamaguti, 1934) Srivastava, 1938 from Platycephalus scabar off the Karachi coast. Proc. Pak. Acad. Sci., 14: 81-87.

Bilqees, F.M. 1980. Three trematodes including two new species from the fishes of Karachi coast. Zool. Sci., 9: 89-91.

Bilqees, F.M. 1981. Digenetic trematodes of fishes of Karachi coast. Kifayat Academy, Karachi, pp. 207.

Bilqees, F.M. and Durrani, F. 1980. Two species of the genus Prosogonotrema Perez Vigueras, 1940. Pak. J. Zool., 12(2): 205-209.

Bilqees, F.M. and Nighat, Y. 1981. Two hemiurid trematodes from the fishes of Karachi coast. Kifayat Academy, Karachi, pp. 83.

284

Bilqees, F.M. and Nighat, Y. 1985. A new trematode Lecithocladium thynensis, new species (Trematoda: Hemiuridae) from the fish Thynnus sp., off Karachi coast. Pak. J. Zool., 17: 211-214.

Bilqees, F.M. and Khan, A. 1988. Plagioporus gonii nov. sp. (Trematoda: Opecoelidae) from the fish Labeo gonius off Kalri, Lake, Sindh, Pakistan. Angew. Parasit., 29: 37-41.

Bilqees, F.M. and Khatoon, N. 2003. Stromaturus karachiensis n.gen. n.sp. (Digenea: Hemiuridae: Sterrhurinae) from the fish Stromateus niger of Karachi coast. Pak. J. Zool., 35(1): 25-28.

Bilqees, F.M., Khatoon, N. and Hadi, R. 2003. A new species of the genus Stromaturus Bilqees and Khatoon, 2003 from a fish, Otolithus argenteus of Karachi coast. Ind. J. Exp. Zool., 8(2): 435-439.

Bilqees, F.M. and Khatoon, N. 2004. A new trematode species Thynstenopera lobata n.gen. n.sp. (Trematoda: Opecoelidae: Plagioporinae) from the marine fish Thynnus thunnia of Karachi coast, Pakistan. Pak. J. Biol. Sci., 7(8): 1343-1345.

Bilqees, F.M. and Khatoon, S. 2004. Podocotyle (Pedunculotrema) pakistanicus n.sp. (Trematoda: Opecoelidae: Plagioporinae) from the fish Trichiurus savala of Karachi coast. Proc. Parasit., 38: 89-94.

285

Bilqees, F.M., Ghazi, R.R., Khan, A., and Khatoon, N. 2004. Parapolylekithum karachiensis n.gen. n.sp. (Digenea: Allocreadiidae) from the fish Cybium guttatum of Karachi coast. Acta Parasit. Turc., 28(3): 164-166.

Bilqees, F.M., Shaukat, N. and Feroze, R. 2004. Hysterolecitha flaticaudata n.sp. (Digenea: Hemiuridae: Hysterolecithinae) from the fish Engraulis purava of Karachi coast. Proc. Parasit., 38: 95-101.

Bilqees, F.M., Khatoon, N., Shabir, I., Shaukat, N., Muti-ur-Rehman and Khan, A. 2005a. Species of the genus Lecithocladium Luhe, 1909 (Digenea: Hemiuridae Looss, 1899: Elytrophallinae Skrjabin et Guschanskaja, 1954) from fishes of Karachi coast, with description of a new species. Proc. Parasit., 39: 117-148.

Bilqees, F.M., Khatoon, N. and Hadi, R. 2005b. A new species of the genus Stromaturus Bilqees and Khatoon, 2003 from a fish, Otolithus argenteus of Karachi coast. J. Exp. Zool. India, 8(2): 435-439.

Bilqees, F.M., Khatoon, N. and Haseeb, M.F. 2006. Bucephalus hexalobatus n.sp. (Gasterostomata Odhner, 1905: Bucephalidae Poche, 1907: Bucephalinae Nicoll, 1914) from the fish Pomadasys olivaceum of Karachi coast. Int. J. Biol. Biotech., 3(4): 665-667.

Bott, N.J. and Cribb, T.H. 2005. Prosorhynchus lamprelli n. sp. (Digenea: Bucephalidae) from the brassy trevally. Caranx papuensis (Teleostei: Carangidae), from off Lizard Island on the Great Barrier Reef, Australia Zootaxa, 1059: 33-38.

286

Bray, R.A. 1973. Some digenetic trematodes in fishes from the Bay of Biscay and nearby waters. Bull. Brit. Mus. Nat. Hist. Zool., 26: 151-183.

Bray, R.A. 1985. Some helminth parasites of marine fishes of South Africa: Families Gorgoderidae, Zoogonidae, Cephaloporidae, Acanthocolpidae and Lepocreadiidae (Digenea). J. Nat. Hist., 19: 377- 405.

Bray, R.A. 1986. Some helminth parasites of marine fishes of South Africa: Families Enenteridae, Opistholebetidae and Pleorchiidae (Digenea). J. Nat. Hist., 20: 471-488.

Bray, R.A. 2005. Family Acanthocolpidae Luhe, 1906. In: Keys to the Trematoda. Vol. 2. (Eds. A. Jones, R.A. Bray and D.I. Gibson). CABI Publishing and Nat. Hist. Mus., London.

Bray, R.A. and Cribb, T.H. 2002. Monostephanostomum georgianum n.sp. (Digenea:Acanthocolpidae) from Arripis georgianus (Valenciennes) (Perciformes: Arripidae) off Kangroo Island, South Australia, with comments on Monostephanostomum Kruse, 1979 and Stephanostomum Looss, 1899. Sys. Parasit., 53: 61-68.

Bray, R.A. and Cribb, T.H. 2003. Species of Stephanostomum Looss, 1899 (Digenea: Acanthocolpidae) from fishes of Australian and South Pacific waters, including five new species. Sys. Parasit., 55: 159-197.

287

Bray, R.A. and Cribb, T.H. 2004. Species of Lecithocladium Luhe, 1901 (Digenea: Hemiuridae) from Australian marine fishes, with a description of a new species from various hosts off Eastern Australia. Acta Parasit., 49(1): 3-11.

Bray, R.A. and Justine, J.L. 2006. Prosorhynchus maternus n. sp. (Digenea: Bucephalidae) from the Malabar grouper Epinephelus malabaricus (Perciformes: Serranidae) off New Caledonia. Brit. Mus. Nat. Hist. Zool. Folia Parasit., 53: 181-188.

Bravo-Hollis, M. and Sogandares-Bernal, F. 1956. Trematodes of marine fishes of Mexican waters. IX. Four gasterostomes from the Pacific coast. J. Parasit., 42(5): 536-539.

Byrd, M.A. 1963. Helminth parasites of Atlantic vertebrates. Part I. Digenetic trematodes of marine fishes. Proc. Helm. Soc. Wash., 30(1): 129-148.

Caballero, C. E. 1952. Revision de los genero y especies que integran la familia Acanthocolpidae Luhe, 1909 (Trematoda: Digenea). Rev. Med. Vet. Parasit., 11: 1-231.

Cable, R.M. and Hunninen, A.V. 1942. Studies on Deropristis inflata (Molin), its life history and affinities to trematodes of the family Acanthocolpidae. Biol. Bull. Mar. Lab. Woods Hole, 82: 292-312.

288

Chambers, C.B., Carlisle, M.S., Dove, A.D.M. and Cribb, T.H. 2001. A description of Lecithocladium invasor n.sp., (Digenea: Hemiuridae) and the pathology associated with two species of Hemiuridae in acanthurid fish. Parasit. Res., 87: 666-673.

Chatterjee, K.D. 1981. Parasit., 12th edition. Chatterjee Med. Publ. Calcutta, India.

Chauhan, B.S. 1945. Trematodes from Indian marine fishes. Part-IV, on some trematodes of the family Hemiuridae Luhe, 1901, with description of six new species. Proc. Ind. Acad. Sci., 21: 160-173.

Cohen, D. M. 1970. How many recent fishes are there? Proc. Calif. Acad. Sci., 37(17): 341-346.

Cohn, L. 1902. Mitteilungen uber Trematoden. Zool. Anz., 25: 712-718.

Constanza, R. 1991. Ecological economics: The science and management of sustainability. NewYork: Columbia University Press.

Corkum, K.C. 1967. Bucephalidae in fishes of the Northern Gulf of Mexico: Bucephalus Baer, 1827. Tr. Am. Micr. Soc., 86(1): 44-49.

Crewe, W. and Haddock, D.R.W. 1985. Parasites and human diseases. Edward Arnold Pty Ltd. Singapore, pp. (iii-iv) preface.

289

Cribb, T.H. 2005. Family Opecoelidae Ozaki, 1925. In: Keys to the Trematoda. Vol. 2. (Eds. A. Jones, R.A. Bray and D.I. Gibson). CABI Publishing and Nat. Hist. Mus., London, U.K., pp. 443-531.

Cribb, T.H., Chisholm, L.A. and Bray, R.A. 2002. Diversity in the Monogenea and Digenea: does lifestyle matter? Internat. J. Parasit., 32: 321-328.

Cribb, T.H., Bray, R.A., Wright, T. and Pichelin, S. 2002. The trematodes of groupers (Serranidae: Epinephelinae): knowledge, nature and evolution. Parasit., 124: S23-S42.

Crowcroft, P.W. 1947. Some digenetic trematodes from fishes of shallow Transmanian water. Pap. Proc. Roy. Soc. Tasm., pp. 5-25.

Daily, G.C. 1997. Nature’s services. Societal dependence of natural ecosystems. Washington, D.C.: Island Press, pp. 195-214.

Dawes, B. 1946. The trematoda with special reference to British and other European forms. Cambridge: Camb. Univ. Press, pp. 644.

Deslongchamps, E. 1824. Distome scie. In: Lamouroux, M.M., de Saint- Vincent, B. & Deslongchamps, E. (Eds) Encyclopedie methodique. Histoire naturelle des zoophytes, ouanimaux rayonnes, faisant suite a l’histoire naturelle des vers, de Bruguiere. Paris: Mme Veuve Agasse, pp. 281.

290

Dobrovolny, G.C. 1939. A life history of Plagioporus lepomis, a new trematode from fishes. J. Parasit., 25: 461-470.

Dollfus, R.P. 1937. Les Trematodes digenea des selaciens (Plagiostomes). Catalogue par hotes. Distribution geo-graphique. Ann. Par., 15(1-3): 57-73; 164-176; 259-281.

Dollfus, R.P. 1973. Parasitologia mauritanica. Digenetiques de teleosteens. 1 note. Du genre Stephanochasmus A. Looss, 1900. Bull. Institut. Fond. Afriq. Noire, 34: 809-827.

Dujardin, F. 1845. Histoire naturelle des helminthes ou vers intestinaux. Paris: Lib. Encyclop. Roret, XVI, pp. 654.

Durio, W.O. and Manter, H.W. 1968. Some digenetic trematodes of marine fishes of New Caledonia. Part 1. Bucephalidae, Monorchiidae and some smaller families. Proc. Helm. Soc. Wash., 35: 143-153.

Dyer, W.G., Williams, E.H. and Bunkley-Williams, L. 1986. Some trematodes of marine fishes of South-western and North-Western Puerto Rico. Trans. Illinois Acad. Sci., 79: 141-143.

Eckmann, F. 1932. Beitrage zur Kenntnis der Trematoden familie Bucephalidae. Z. Par., 5(1): 94-111.

Emond, R.T.D. 1974. A coloured Atlas of Infectious diseases. Inter. Ed. Toppan Comp., Ltd. Singapore. (Preface).

291

Eschmeyer, W.N., ed. 1998. Catalog of fishes 3 Vols. Sanfrancisco: Cal. Acad. Sci. Sanfrancisco, 2(1-3): 905.

Etchegoin, J.A., Timi, J.T., Cremonte, F. and Lanfranchi, A.L., 2005. Redescription of Prosorhynchus australis Szidat, 1961 (Digenea: Bucephalidae) parasitizing Conger orbignianus Valenciennes, 1842 (Pisces: Congridae) from Argentina. Acta Parasit., 50: 102-104.

Farooq, M. and Khanum, Z. 1980. Two new species of the genus Lecithocladium Luhe, 1901 (Digenea: Hemiuridae) from marine fishes of Karachi coast, Pakistan. Pak. J. Zool., 12: 221-224.

Farooq, M and Sadiq, 1992. FAO, 1992: FAO year book 1991 fishery statistic catches and landing. FAO fisheries series 39 (1993, series 44) series 38: 1-41.

Fischthal, J.H. 1980. Some digenetic trematodes of marine fishes from Israel’s Mediterranean coast and their zoogeography, especially those from Red Sea immigrant fishes. Zool. Script, 9: 11-23.

Fischthal, J.H. and Kuntz, R.E. 1963. Trematode parasites of fishes from Egypt part III, six new Hemiuridae. Proc. Helm. Soc. Wash., 30: 78-91.

Fischthal, J.H. and Kuntz, R.E. 1964. Digenetic trematodes of marine fishes from Palawan Island, Philippines. II. Five Opecoelidae including new species. Proc. Helm. Soc. Wash., 31(1): 40-46.

292

Fischthal, J.H. and Kuntz, R.E. 1965. Digenetic trematodes of fishes from North Borneo (Malaysia). Proc. Helm. Soc. Wash., 32(1): 63-71.

Fischthal, J.H. and Thomas, J.D. 1968. Digenetic trematodes of marine fishes from Ghana: Families Acanthocolpidae, Bucephalidae, Didymozoidae. Proc. Helm. Soc. Wash., 35: 237-247.

Fischthal, J.H. and Thomas, J.D. 1970. Digenetic trematodes of marine fishes from Ghana: Family Opecoeliidae. Proc. Helm. Soc. Wash., 37(2): 129-141.

Froese, R. and Pauly, D. 2002. Fish Base: World Wide Web electronic publication. www.fishbase.org.

Fuhrmann, O. 1928. Trematoda. Kukenthal’s Handb. Zool., pp. 171.

Gaevaskaya, A.V.I. and Kovaleva, A.A. 1978. Data on the trematodes of fish from the South Western Atlantic. Vestnik Zool., 3: 60-66.

Gerald, D. Schmidt and Larry, S. Robert’s, 2000. Foundat. Parasit., 6th ed. Mc. Graw – Hill Book- Co. Singapore, pp. 207.

Giboda, M., Ditrich, O., Scholz, T., Veingsay, T. and Bouaphanh, S. 1991. Human infections in Laos. Transact. Roy. Soc. Trop. Med. Hyg., 85: 534-540.

Gibson, D.I. and Bray, R.A. 1979. The Hemiuroidae: Terminology systematics and evolution. Bull. Brit. Mus. Nat. Hist. Zool., 36: 35-146.

293

Gibson, D.I. and Bray, R.A. 1982. A study and reorganization of Plagioporus Stafford, 1904 (Digenea: Opecoelidae) and related genera, with special reference to forms from European Atlantic waters. Bull. Brit. Mus. Nat. Hist. Zool., 16: 529-559.

Gibson, D.I. and Bray, R.A. 1986. The Hemiuridae (Digenea) of fishes from the North-east Atlantic. Bull. Brit. Mus. (N.H.) Zool., 51: 1-125.

Gibson, D.I. 2002. Family Hemiuridae Looss, 1899. In: Keys to the trematodes Vol. 1 (Eds. D.I. Gibson, A. Jones & R.A. Bray). CAB International Wallingford, pp. 307-340.

Global Biodiversity Assessment. UNEP, 1995, Annex 6, Glossary. ISBN 0-521564816, used as source by “Biodiversity”. Glossary of terms related to the CBD, Belgian Clearing-House Mechanism, retrieved April 26, 2006.

Government of Baluchistan and IUCN- The world Conservation Strategy, 2000 - Baluchistan Conservation Strategy: An Overview.

Government of Pakistan, Finance Division, Economic Advisor’s Wing, 2004 – Pakistan Economic Survey, 2003 – 2004, pp. 102-110.

Government of Pakistan, Ministry of Environment, Local Government and Rural Development, 2002 - Pakistan Country Assessment Report, pp. 102-110.

294

Government of Sindh, Environmental Protection Agency, 1999- Studies on the status of Water Bodies in Rural Sindh and their Effect on Human health.Government of Sindh, Environment Protection Agency and UNDP, 2004 – Status of Water Quality Supplied to Hyderabad City from River Indus, pp. 102-110.

Gu, C. and Shen, J. 1978. Some diurid trematodes (Sub-family: Diurinae Looss, 1907) from marine fishes of economic importance of China. Acta Zool. Sinica., 24: 373-387.

Gupta, N.K. 1956. Studies on the digenetic trematodes of marine fishes from the Gulf of Manaar, India. Res. Bull. Punjab Univ. Zool. Hoshiarpur, (89): 61-83.

Gupta, A.N. and Sharma, P.N. 1973. Studies on the family Hemiuridae (Trematoda: Digenea), with a description of a new genus from marine fish of Indian waters. Mar. Biol., 19: 89-91.

Gupta, N.K. and Mehrotra, V. 1970. A new hemiurid parasite of the genus Lecithocladium Luhe, 1901, with revised key to Indian species of the genus. Res. Bull. Punjab Univ., 21: 73-76.

Gupta, N.K. and Sehgal, S.K. 1967. Two new parasites of digenetic trematodes from some Indian marine food fishes. Ind. J. Helm., 18: Appendix p. 3.

295

Gupta, P.C. and Govind, H. 1985. Studies on trematodes of fishes, on four new species of the genus Lecithocladium Luhe, 1901 from marine fishes of Puri coast. Kanpur Univ. Res. J. (Sci.), 2: (1981): 31-39.

Gupta, P.C. and Gupta, V.C. 1983. Three species of the genus Lecithocladium Luhe, 1901, (Digenea: Hemiuridae) from Bay of Bengal, off Puri coast, Orissa. Kanpur Univ. Res. J. (Sci.), 4: 1-7.

Gupta, P.C. and Singh, R.B. 1982. Two new trematodes of the genus Lecithocladium Luhe, 1901 from a marine fish of Puri coast, Bay of Bengal. Ind. J. Parasit., 6: 325-329.

Gupta, P.C. and Singh, R.G. 1985. Studies on digenetic trematodes of the families Monorchiidae and Hemiuridae from marine fishes of Puri coast, Bay of Bengal. Kanpur Univ. Res. J. (Sci.), 2: (1981), 11-21.

Gupta, R.C. & Gupta, S.P. 1974. On two new species of the genus Stomachicola Yamaguti, 1934 from the stomach of marine fishes from Quilon, Kerala. Ind. J. Zool., 15: 7-10.

Gupta, R.C. & Gupta, S.P. 1978. On four new trematodes from marine fishes. Ind. J. Helm., 28: 114-126.

Gupta, R.C. and Gupta, S.P. 1978. Five new trematodes of the genus Lecithocladium Luhe, 1901 (Hemiuridae) from marine fishes. Folia Parasit., 25: 223-229.

296

Gupta, R.C. and Gupta, S.P. 1980. Three new trematodes of the family Acanthocolpidae Luhe, 1909 from marine fishes of Puri, Orissa. Ind. J. Helm., 32: 61-71.

Gupta, V. & Ahmad, J. 1976. Digenetic trematodes of marine fishes. On four new species of trematodes of marine fishes of Puri, Orissa. Riv. Parasit., 37: 133-142.

Gupta, V. and Ahmad, J. 1977. Digenetic trematodes of marine fishes. On some new and known digenetic trematodes from marine fishes of Bay of Bengal, Puri, Orissa. Riv. Parasit., 38: 181-191.

Gupta, V. & Ahmad, J. 1978. Digenetic trematodes of marine fishes. IX. On two new species of the genus Stomachicola Yamaguti, 1934 (Trematoda: Hemiuridae) from marine fishes of Puri, Orissa. Acta Parasit. Pol., 25: 211-215.

Gupta, V. and Ahmad, J. 1981. Digenetic trematodes of marine fishes. On three trematodes of the genus Acanthocolpus Luhe, 1906 from marine fishes of Bay of Bengal, Puri coast, Orissa. Riv. Parasit., 42: 61-66.

Gupta, V. and Jahan, A. 1977. On some trematodes of marine fishes of Puri. Ind. J. Helm., 27: 46-61.

Gupta, V. & Puri, M. 1980. Four new species of digenetic trematodes of fishes. Ind. J. Helm., 31: 54-64.

297

Gupta, V. and Puri, M. 1981. Six new species of the genus Lecithocladium Luhe, 1901, (Hemiuridae Luhe, 1901) from marine fishes of Puri, Orissa. Ind. J. Helm., 33: 58-78.

Gupta, V. and Jain, N. 1992. On three species of the genus Lecithocladium Luhe, 1901, (Hemiuridae Luhe, 1901) from the stomach of marine fishes off coast Puri, Orissa. Riv. Parasit., 9: 161-172.

Hafeezullah, M. 1970. Digenetic trematodes of marine fishes of India. Part I. Bucephalidae and Cryptogonimidae. Ind. J. Parasit., 22: 1-22.

Hafeezullah, M. 1971. On some new and known digenetic trematodes from marine fishes of India. J. Helm., 45: 73-88.

Hafeezullah, M. 1975. Two new trematodes (Digenea: Hemiuridae) from marine fishes of East India. In: Dr. B.S. Chauhan Commemoration. Vol. Publ. by Zool. Soc. India, 203-210.

Hafeezullah, M. 1978. Acanthocolpid trematodes of marine fishes of India, with considerations on synonymies in the group. Bull. Zool. Surv. India, 1: 29-36.

Hafeezullah, M. 1980. On synonymy of some genera of the sub-families Stomachicolinae Yamaguti, 1958, Diurinae Looss, 1907 and Prolecithinae Yamaguti, 1971 (Digenea: Hemiuridae). Bull. Zool. Surv. India, 3: 69-76.

298

Hafeezullah, M. 1991. Remarks on some known species of digenetic trematodes (Digenea: Acanthocolpidae Luhe, 1909) from fishes of India. Rec. Zool. Surv. India, 88: 105-114.

Hafeezullah, M. and Siddiqi, A.H. 1970. Digenetic trematodes of marine fishes of India. Part 1. Bucephalidae and Cryptogonimidae. Ind. J. Parasit., 22: 1-22.

Hafeezullah, M. and Siddiqi, A.H. 1975. Some digenetic trematodes of marine fishes of Nigeria. Family Hemiuridae Luhe, 1907. In Dr. B.S. Chauhan Commemoration Volume, 1975. Orissa, Publ. Zool. Soc. India, pp. 215-223.

Hanson, M. L. 1950. Some digenetic trematodes of marine fishes of Bermuda. Proc. Helm. Soc. Wash., 17(2): 74-88.

Harderlie, E.C. 1953. Parasites of fresh water fishes of North California. Univ. Calf. Publ. Zool., 57(5): 303-440.

Hausmann, L. 1896. The Life History and Biology of Plagioporus (Trematode: Opecoelidae). J. Parasit., 1978-JSTOR.

Holmes, J. & Bartoli, P. 1993. Spatio-temporal structure of the communities of helminthes in the digestive tract of Sciaena umbra L., 1758 (Teleostei). Parasit., 106: 519-525.

Hopkins, S.H. 1956. Two new trematodes from Louisiana and the excretory system of Bucephalidae. Tr. Am. Micr. Soc., 75(1): 129-135.

299

Horn, M. H. 1972. The amount of space available for marine and fresh water fishes. NOAA Fish. Bull., 70(4): 1295-1297.

Hunter et Bangham, 1932. The digenetic trematodes of marine fishes of Tortugas Florida H.W. Hunter-Amer. Midl. Nat., 38(2): 257-416.

IUCN- The World Conservation Union, 2003 – Environmental Degradation and Impacts on Livelihoods: Sea intrusion A Case Study. www.smarter.com/www.sustrust.org./recentandnew/activities/2003- 2004.html.

IUCN- The World Conservation Union, 2004 – Sindh State of Environment and Development. www.smarter.com/www.sustrust.org./ recentandnew/activities/2003-2004.html.

Issaitschikow, I.M. 1928. Zur Kenntnis der parasitischen Wurmer einiger Gruppen von Wirbeltieren der russischen Arktis. Trudy Morsk. Nauch. Instit. Moskva, 3(2): 5-79.

Jahan, I. 1970. A new trematode parasite Acerointesticola karachiensis n. gen., n. sp. (Sub-family Stomachicolinae Yamaguti, 1934) from the intestine of a fish. Rec. Zool. Surv. Pak., 1: 39-41.

Jaypee, C.K., 2002. CK Jayaram Paniker, Jaypee Publishers Ltd. Med. Parasit., 2002, 5th ed. (Preface).

300

Jones, D.O. 1943. The anatomy of three digenetic trematodes, Skrjabiniella aculeatus (Odhner), Lecithochirium rufoviride (Rud.) and Sterrhurus fusiformis (Luhe) from Conger conger (Linn.). Parasit., 35: 40-57.

Kakaji, V.L. 1969. Studies on helminth parasites of Indian fishes. Part II. Some trematode parasites of fresh water fishes of Uttar Pradesh. Ind. J. Helm., 21(1): 49-80.

Karlsbakk, E. 1993. On the morphology and ecology of some digenean parasites of the four-bearded rockling [Enchelyopus cimbrius (L.)] (Gadidae) in Raunefjorden, Western Norway. Ph.D Thesis. Bergen: Univ. Berg., pp. 92.

Kaup, 1858. Redescription of Prosorhynchus was originally described by Szidat, 1961. Parasitizing Urophysis brasiliensis. Acta Parasit., pp. 102- 104.

Kelvin J. Gaston & John I. Spicer. 2004. “Biodiversity: an introduction”. Blackwell Publishing. 2nd Ed., ISBN 1-4051-1857-1 (pbk.)

Khamboonruang, C., Keawivichit, R., Wongworapat, K.,Suwanranguri, S., Hongpromyart, M., Sukhawat, K., Tonguthai, K. and Lima dos Santos CA. 1995. Application of hazard analysis and critical control point (HACCP) as a possible control measure against Opisthorchis viverrini infection in cultured carp (Puntius gonotius goniotus). Southeast Asian J. Trop. Med. Publ. Health, 28(1): 65-72.

301

Kniskern, V.B. 1952. Studies on the trematode family Bucephalidae. Tr. Am. Micr. Soc., 71(3): 253-266.

Kohn, A. 1961. Um novo Prosorhynchus parasito de Garrupa sp. (Trematoda: Bucephaliformes). Atas Soc. Biol. Rio J., 5: 46-49.

Koie, M. 1984. Digenetic trematodes from Gadus morhua L. (Osteichthyes, Gadidae) from Danish and adjacent waters, with special reference to their life histories. Ophelia, 23: 195-222.

Komiya, Y. and Tajimi, T. 1941. Progenesis in digenea trematodes: Ataxonimic and Synth.J. Camb. Org. Prod. Act. cjo Get Fulltex? Fulltextid=305634.

Korotaeva, V.D. 1974. Helminths of some commercial marine fishes of sub-order Scomberoidei from the Australian region. Izv. Tikh. Nauch. Issl. Skogo Instit. Rylen. Khozy. Okean. (TINRO), 88: 61-66 (In Russian).

Koval, V.P. 1949. A new species of Bucephalus I Dnieper fishes. Dokl. Akad. Nauk. SSSR., 68(1): 205-208.

Kruse, G.O.W. 1977. Some digenetic trematodes from fishes of the Bering Sea with the descriptions of Prosorhynchus mizellei sp. n. (Bucephalidae) and Pseudopecoelus nossamani sp. n. (Opecoelidae). Proc. Helm. Soc. Wash., 44: 73-76.

302

Kruse, G.O.W. 1979. Trematodes of marine fishes from South Australia. 6. Monostephanostomum manteri gen. et sp. n. (Acanthocolpidae). J. Parasit., 65: 921-923.

Lamouthe-Argumedo, R., Garcia-Prieto, L., Osorio-Sarabia, D. and Perez-Ponce de Leon, G. 1997. Catalogo de la coleccion nacional de helmintos. Mexico: Univ. Nac. Auto. Mex., pp. 211.

Layman, E.M. 1930. Parasitic worms from the fishes of Peter the Great Bay. Bull. Pacif. Scient. Fish. Res., 3(6): 1-120.

Layman, E.M. 1933. Einige neue tatsachen Uber die Okologie der Froschtrematoden. Zool. Anz., 101(7-8): 199-201.

Layman, E.M. 1937. Uber die parasitischen Wurmer der Fische des Baikalsees. Trudy Baikal Limn. St., 4: 5-98.

Lebedev, B.I. 1968. New trematodes from pelagic fishes of order Perciformes of Pacific basin. Helminths of animals of Pacific Ocean. Moskva. Nauka, pp. 56-64.

Lebour, M. 1908. Trematodes of the Northumberland coast, No. II. Trans. Nat. Hist. Soc. Northumb., Durh. and Newcastle-upon-Tyne, new series, 2: 1-20.

Leiper, R.T. and Atkinson, E.L. 1914. Helminthes of the British Arctic Expedition, 1910-1913. Proc. Zool. Soc. London, (1): 222-226.

303

Leong, T.S. and Wong, S.Y. 1988. A comparative study of parasite fauna of wild and cultured grouper (Epinephelus malabaricus Bloch et Schneider) in Malaysia. Aquaculture, 68: 203-207.

Leong, T.S. and Wong, S.Y. 1990. Parasites of healthy and diseased juvenile grouper (Epinephelus malabaricus Bloch et Schneider) and seabass (Lates calcarifer Bloch) in floating cages in Penang, Malaysia. Asian Fish. Sci., 3: 319-327.

Levinsen, G.M.R. 1881. Bidrag til Kundskab om Gronlands Trematodfauna. Overs. K. Danske Vidensk. Selsk. Forh., 23(1): 52-84.

Linton, E. 1898. Notes on trematode parasites of fishes. Proc. U.S. Nat. Mus. Wash., 20: 507-548.

Linton, E. 1900. Fish parasites collected at Woods Hole in 1898. U.S. Fish Comm. Bull. 1899, 19: 267-304.

Linton, E. 1901. Parasites of fishes of Woods Hole region, U.S. Fish Comm. Bull. 1899, pp. 460-461.

Linton, E. 1905. Parasites of fishes of Beaufort, North Carolina. Bull. U.S. Bur. Fish. (1904), 24: 321-428.

Linton, E. 1910. Helminth fauna of the Dry Tortugas. II. Trematodes. Pap. Tort. Lab. Carneg. Instit. Wash., 4: 11-98.

304

Linton, E. 1934. A new genus of trematodes belonging to the Sub-family Allocreadiinae. J. Wash. Acad. Sci., 241(2): 81-83.

Linton, E. 1940. Trematodes from fishes mainly from the Woods Hole region, Massachusetts. Proc. U.S. Nat. Mus., 88: 1-172, 26pl.

Linton, S. 1910. Helminth fauna of the Dry Tortugas, II. Trematodes. Carneg. Inst. Wash. Publ. No. 133, 98 pp.

Liu S.-f. 1998. The trematodes of marine fishes from Fujian, China (Acanthocolpidae [sic]). Zool. Res., 19: 468-472 (In Chinese).

Looss, A. 1899. Weitere Beitrage zur Kenntnisder Trematoden fauna Aegypteus, Zugleich versuch einer naturlichen Gliederung des Genus Distomum Retzius. Zool. Jahrb. Syst., 12: 521-784.

Looss, A. 1900. Nachtragliche Bemerkungen Zu den Namen der won mir vorgeschagenen Distomengaltungen Zool. Anz., 23: 601-608.

Looss, A. 1901. Uber die Fasciolidengenera Stephanochasmus, Acanthochasmus und einige andere. Centralblatt fur Bakteriologie, Parasitenkunde und Infektionskrankheiten, 29(14): 595-606, 628-634, 654-661.

Looss, A. 1901. Notizen zur Helminthologie Aegyptens. IV. Uber Trematoden aus Schildkroten der aegyptischen Kusten. Ctbl. Bakt. I., 30(15-16): 555-569, 618-625.

305

Looss, A. 1902. Die Distomen- Unterfamilie der Haploporinae. Arch. Parasit., 6: 9-143.

Looss, A. 1907. Zur kenntnis der Distomenfamilie. Zool. Anz., 31(19- 20): 585-620.

Ludwig, D., Hillborn, R. and Walters, C. 1993. Uncertainity, resources, exploitation and conservation: Lessons from Hist. Sci., 260: 17-36.

Luhe, M. 1901. Uber Hemiuriden. Zool. Anz., 24: 394-403, 473-488.

Luhe, M. 1906. Report on the trematode parasites from the marine fishes of Ceylon. Roy. Soc. Rep. on Pearl Oyster, 5: 97-108.

Luhe, M. 1909. Parasitische Plattwurner .I. Trematoden. Susswasser fauna Deutschl. Heft, 17: 215pp.

MacCallum, G.A. 1917. Some new forms of parasitic worms. Zoopathol., 1: 43-75.

Machida, M. 1984. Trematodes of marine fishes from depth of 200-400m off Yamagata, the Japan Sea. Mem. Nat. Sci. Mus., Tokyo, 17: 101-110.

Machida, M. and Araki, J. 1994. Some trematodes and cestodes in fishes from off Eastern Hokkaido, Northern Japan. Mem. Nat. Sci. Mus., Tokyo, 27: 87-92.

306

Machida, M. and Uchida, A. 1990. Trematodes from unicorn fishes of Japanese and adjacent waters. Mem. Nat. Sci. Mus., 23: 69-81.

Madhavi, R. 1974. Digenetic trematodes from marine fishes of Waltair Coast, Bay of Bengal. Family Bucephalidae. Riv. Parasit., 35: 189-199.

Madhavi, R. 1976. Digenetic trematodes from marine fishes of Waltair coast, Bay of Bengal. Family Acanthocolpidae. Riv. Parasit., 37: 115- 128.

Madhavi, R and Narasimhulu, S.V. 1985. Studies on Pleorchis sciaenae Yamaguti, 1938 (Trematoda: Pleorchiidae) from marine fishes of Bay of Bengal. Riv. Parasit., 46: 185-190.

Mamaev, I.L. 1965. Helminths of fish from Bering Sea. Parasitic worms of domestic and wild animals. Akad. Nauk. SSSR. Vladivostok, pp. 168- 188.

Mamaev, I.L. 1970. Helminths of some commercial fishes in the Gulf of Tong King. In Helminths of animals of South-East Asia (Eds. P.G. Oshmarin & I.L. Mamaev and B.I. Lebedev). Izd. Nauk. Mascow, pp. 127-190 (In Russian).

Mamaev, I.L. and Oshmarin, P.G. 1966. Trematodes of the family Acanthocolpidae Luhe, 1901, in herrings of the North Vietnam Bay. Helm., 7: 155-164.

307

Manter, H.W. 1931. Some digenetic trematodes of marine fishes of Beaufort, North Carolina. Parasit., 23(3): 396-411.

Manter, H.W. 1934. Some digenetic trematodes from deep water fish of Tortugas, Florida. Papers from Tortugas Lab. Carneg. Instit. Wash., 28: 257-345, 15pl.

Manter, H.W. 1936. Some trematodes of Crenote fish from Yucatan. Carneg. Instit. Wash. Publ., 457: 33-28.

Manter, H.W. 1940a. Digenetic trematodes of fishes from the Galapagos Islands and the neighbouring Pacific. Allan Hancock Pacific Expeditions, 2: 325-497.

Manter, H.W. 1940b. Digenetic trematodes of fishes from the Galapagos Islands and the neighbouring Pacific. Allan Hancock Pacific Expeditions, 2: 531-547.

Manter, H.W. 1940. Gasterostomes (Trematoda) of Tortugas, Florida. Papers from Tortugas Lab., 33(1): 1-19.

Manter, H.W. 1947. The digenetic trematodes of marine fishes of Tortugas, Florida. Amer. Midl. Nat., 3: 257-416.

Manter, H.W. 1953. Two new species of Prosorhynchinae (Trematoda: Gasterostomata) from the Fiji Islands. In: J. Dayal and K. Singh (Eds.), Thapar, G.S., Commem. Vol. Univ. Luck., Lucknow, pp. 193-200.

308

Manter, H.W. 1954. Some digenetic trematodes from fishes of New Zealand. Trans. Royal Soc. New Zealand, 82: 475-568.

Manter, H.W. 1963a. Studies on digenetic trematodes of fishes of Fiji, II. Families Lepocreadiidae, Opistholebetidae and Opecoelidae. J. Parasit., 49(1): 99-113.

Manter, H.W. 1963b. Studies on digenetic trematodes of fishes of Fiji, III. Families Acanthocolpidae, Fellodistomatidae and Cryptogonimidae. J. Parasit., 49(3): 443-450.

Manter, H.W. 1969. The occurrence of an accessory excretory vesicle in Prosogonotrema Perez Vigueras, 1940. Parasit. Coll., 24: 173-177.

Manter, H.W. and Van Cleave, H.J. 1951. Some digenetic trematodes, including eight new species, from marine fishes of La Jolla, Calif. Proc. U.S.A. Nat. Hist. Mus., 101: 315-340.

Manter, H.W. and Pritchard, M.H. 1960. Additional Hemiurid trematodes from Hawaiian fishes. Proc. Helm. Soc. Wash., 27: 165-180.

Mathews, R.A. 1973. The life cycle of Prosorhynchus crucibulum (Rudolphi, 1819) Odhner, 1905, and a comparison of its cercaria with that of Prosorhynchus squamatus Odhner, 1905. Parasit., 66: 133-164.

McCauley, James. E. 1960. Some Hemiurid Trematodes of Oregon Marine Fishes. J. Parasit., 46(1): 84-89.

309

McFarlane, S.H. 1936. A study on the endoparasitic trematodes from marine fishes of Departure Bay, B.C. J. Biol. Board Can., 2: 335-347.

McGladdery, S.E. 1999. Stephanostomum tenue in marine aquaculture of rainbow trout (Onchorhynchus mykiss). ICES Identification Leaflets for Diseases and Parasites of Fish and Shellfish, 51: 4pp.

Molin, R. 1858. Prospectus helminthum, quae in prodromo faunae helminthologicae venetae continentur. Sitz.- ber K. Akad. Wiss. Wien, Math.-Naturw. Cl., 19: 189-338.

Molin, R. 1859. Prospectus helminthum, quae in parte secunda prodromi faunae helminthologicae venetae continentur. Sitz.- ber K. Akad. Wiss. Wien, Math.-Naturw. Cl., 33(26): 287-302.

Moravec, F. and Sey, O. 1989. Some trematodes of fresh water fishes from North Vietnam with a list of recorded endohelminthes by fish hosts. Folia Parasit., 36: 243-262.

Moyle, P.B. and Moyle, P.R. 1995. Endangered fishes and economics: Intergenerational obligations. Env. Biol. Fish., pp. 42.

Moyle, P.B., Joseph, J. and Cech, Jr. 2000. Fishes An Introduction To Ichthyology. 4th Ed. Pretice Hall, Inc. U.S.A., pp. 1-5.

Nagaty, H.F. 1937. Trematodes of fishes from the Red Sea. Part 1. Studies on the family Bucephalidae Poche, 1907. Egyptian University, Cairo, 172pp.

310

Nagaty, H.F. 1941. Trematodes of fishes from the Red Sea. II. The genus Hamacreadium Linton, 1910 (Allocreadiidae) with a description of two new species. J. Egypt. Med. Ass., 24(2): 300-310.

Nagaty, H.F. 1956. Trematodes of fishes from the Red Sea. Part 6. On Five distomes including one new genus and four new species. J. Parasit. Cairo, Egypt, 42(2): 151-155.

Nagaty, H.F. 1957. Trematodes of fishes from the Red Sea. Part 8. Five species in the families Schistorchidae, Acanthocolpidae and Heterophyidae. J. Parasit., 43: 217-220.

Nagaty, H.F. And Abdel Aal, T.M. 1962. Trematodes of fishes from the Red Sea. Part XII. On four acanthocolpids including a new species. Parasit., 52: 186-191.

Nagaty, H.F. And Abdel Aal, T.M. 1962. Trematodes of fishes from the Red Sea. Part XVI. On three new species of Hamacreadium (Allocreadiidae). J. Arab Veter. Med. Ass., 22(4): 301-305.

Nahhas, F.M. and Cable, R.M. 1964. Digenetic and aspidogastrid trematodes from marine fishes of Curacao and Jamaica. Tul. Stud. Zool., 11: 169-228.

Nahhas, F.M. and Short, R.B. 1965. Digenetic trematodes of marine fishes from Apalachee Bay, Gulf of Mexico. Tul. Stud. Zool., 12(2): 39- 50.

311

Nicoll, W. 1907. A contribution towards a knowledge of the Entozoa of British marine fishes. Part I. Ann. Mag. Nat. Hist., 19(7): 66-94.

Nicoll, W. 1909. Studies on structure and classification of the digenetic trematodes. Quart. J. Micr. Sc., 53(3): 391-487.

Nicoll, W. 1910. On the Entozoa of fishes from the Firth of Clyde. Parasit., 3: 322-359.

Nicoll, W. 1914. Trematode of fishes from the English Channel. J. Mar. Biol. Ass., 10: 466-505.

Nicoll, W. 1915. The trematode parasites of North Queensland: Parasite of fishes. Parasit., 8(1): 22-41.

Nigrelli, R.F. 1940. Two new species of trematode from the deep sea Scorpion fish, Scorpaena madurensis Cuv. And Val. Zoologica: Sci. Contr. N.Y. Zool. Soc., 25: 264-268.

O’Bannon, B.K. 1994. Fisheries of the United States, 1993. Current Fishery Statistics No. 9300. Silver Spring Md: NOAA/NMFS., 121 pp.

Odhner, T. 1902. Mitteilung Zur Kenntnis der Distomen. Centralbl. Bakt. 1., 31: 58-68.

Olsson, P. 1876. Bidrag till skandinaviens helmith fauna. T. Kongl. Sv. Veter. Akad. Handl., 14(1): 35; 25: 41pp.

312

Olsson, P.D., Cribb, T.H., Tkach, V.V., Bray, R.A. and Little wood, D.T.J. 2003. Phylogeny ad Classification of the Digenea (Platyhelminths: Trematoda). Internat. J. Parasit., 33: 733-755.

Odhner, T. 1905. Die Trematoden des arktischen Gebietes fauna Arctica, 4(2): 291-372.

Oshmarin, P.G. 1965. Two new sub-families of trematodes from fishes in the South China Sea. Helm., 6(2): 99-107. Oshmarin, P.G. and Mamaev, I.L. 1963. New trematodes from fishes of North Vietnam (Tokin Bay). Helm., 4(1-4): 357-365.

Overstreet, R.M. 1969. Digenetic trematodes of marine teleost fishes from Biscayne Bay, Florida. Tul. Stud. Zool. Bot., 15: 119-176.

Overstreet, R.M. and Curran, S.S. 2002. Super-family Bucephaloidea Poche, 1907. In: Keys to the Trematoda. Vol. I (Eds. D.J. Gibson, A. Jones and R.A. Bray). CABI Publishing London, pp. 67-110.

Ozaki, Y. 1924. Gasterostomatous trematodes and three new genera. Zool. Mag., 36(426): 173-201.

Ozaki, Y. 1925. Trematodes from Red Sea fishes Plagioporus Stafford, 1904. Acta Parasit., 50: 276-280.

Ozaki, Y. and Ishibashi, 1934. Petalocotyle nipponica, a new type of trematode family Allocreadiidae. Proc. Imp. Acad., 10(2): 111-114.

313

Paggi, L. and Orecchia, P. 1965. Suunnovo trematode parasita de’ll intestine di Morone Labra: Bucephalus labracis n.sp. Parasit., 7(2/3): 69- 74.

Pan, H. 1988. Studies on digenetic trematodes of fam. Bucephalidae from fresh water fishes of Guang dong Province, with descriptions of two new species. Acta Hydro boil. Sin., 12: 65-69 (In Chinese, English summary).

Pandey, K.C. and Tewari, S.K. 1984. On three new digenetic trematodes from marine fishes of Bombay coast, India. J. Zool. Soc. Ind., 35: 60-65.

Pandey, P.N., Mittal, N. and Jain, N. 2000. Two digenetic trematodes from marine fishes of Puri Coast, Orissa. Flora and Fauna J., 6(2): 107- 109.

Papoutsoglou, S.E. 1976. Metazoan parasites of fishes from Saronicos Gulf Athens-Greece. Thalassographica, 1: 69-102.

Park, J.T. 1936. Two new trematodes, Sterrhurus magnatestis and Tubulovesicula californica (Hemiuridae) from littoral fishes of Pillon’s beach, Holconotus rhodoterus Tr. Am. Micr. Soc., 55(4): 477-482.

Park, J.T. 1939. Fish trematodes from Tyosen.11. Some new digenetic trematode parasites from marine fishes. Keizyo J. Med., 10: 7-18.

Parukhin, A.M. 1964. New species of trematode of the families Prosogonotrematidae and Cephaloporidae from fish from North Vietnam. Uchen. Gork. Gos. Univ., 62: 22-26.

314

Parukhin, A.M. 1966. [On the species composition of the helminth fauna of fishes in the South Atlantic.] Mater. Nauch. Konfer. Vsesoy. Obshch. Gelm., pp. 219-222 (In Russian).

Parukhin, A.M. 1966. [Helminth fauna of carangid fish from the South China Sea]. Biologiya Morya, Kiev [Helminth fauna of animals in southern seas.], pp. 80-96 (In Russian).

Parukhin, A.M. 1968. Helminth fauna of fishes of South Atlantic. Biologiya Morya, Kiev, 14: 96-113 (In Russian).

Parukhin, A.M. 1974. Some new trematode species of the family Acanthocolpidae Luhe, 1909 from fishes of the Red Sea and Indian Ocean. Izvest. Tikho. Nauch. Issledov. Instit. Ryb. Khozy. Okean. (TINRO), 88: 110-116 (In Russian).

Parukhin, A.M. 1976. [Parasitic worms of bottom fishes of the southern seas]. Kiev: Naukova Dumka, pp. 155 (In Russian).

Pigulewsky, S.W. 1931. Neue Arten von trematoden aus Fischen des Dnjeprbassins. Zool. Anz., 96: 9-18.

Pigulewsky, S.W. 1938. Zur Revision der Parasiten – Gattung Lecithaster Luhe, 1901. Livr. Jubil. Prof. Travassos, pp. 191-197.

Poche, F. 1907. Einige Bemerkungen zur Nomenclatur der Trematoden. Zool. Anz., 31: 124-126.

315

Poche, F. 1926. Das System der Platodaria Archiv fur Naturgeschichte, Abt. A, Zool., 91(1925): 1-459.

Price, E.W. 1934. New digenetic trematodes from marine fishes. Smiths. Misc. Coll., 91(7): 8 pp.

Pritchard, M.H. 1963. Studies on digenetic trematodes of Hawaiian fishes, primarily families Lepodermatidae and Zoogonidae. J. Parasit., 49(4): 578-587.

Pritchard, M.H. 1966. A revision of the genus Podocotyle (Opecoelidae) Zool. Jahrb. Syst., 93: 158-172.

Railliet, 1896. Quelques rectifications a La nomenclature des parasites. Rec. Med. Veter., 3: 157-161.

Ramadan, M.M. 1983. A review of the genus Stephanostomum Looss, 1899 (Trematoda: Acanthocolpidae) with description of two species from Red Sea fishes. J. Educ., Ain Shams Univ., 6: 385-397.

Reid, W.A., Coil, W.H. and Kuntz, R.E. 1966. Hemiurid trematodes of Formosan marine fishes. I. Sub-family Diurinae and Stomachicolinae. J. Parasit., 52: 39-45.

Reimer, L.W. 1983. Acanthocolpidae (Digenea) aus Fischen der Kuste von Mocambique. Ang. Parasit., 24: 134-146.

316

Reza, H. Syed. 1985. Prospects of deep sea fishing in Pakistan. Pak. Agric. Fish., pp. 7-8; 58.

Roitman, V.A. 1963. Morphological characteristics of trematode families Allocreadidae Stossich, 1903 and Opecoeliidae Ozaki, 1925. Trudy Gelm. Lab., 13: 301-302.

Rounsefell, G.A. 1975. Ecology, Utilization and Management of Marine fisheries. St. Louis: C.V. Mosby., pp. 516.

Rudolphi, C.A. 1819. Entozoorum synopsis cui accedunt mantissa duplex et indices locupletissimi. Berol., pp. 811.

Sahai, D. and Srivastava, D.D. 1977. Trematodes of Indian fishes, Part 1. Two new genera of hemiurids (Sub-family: Hemiurinae Looss, 1899). Proc. Nat. Acad. Sci. India, 47: 7-12.

Saoud, M.F.A., Nahhas, F.M., Al Kuwairi, K.S.R. and Ramadan, M.M. 2002. Helminth parasites of fishes from the Arabian Gulf: 10. Trematodes of the genus Stephanostomum Looss, 1899(Digenea: Acanthocolpidae Luhe, 1901), with description of Stephanostomum qatarense n.sp. and redescription of Stephanostomum triacanthi Madhavi, 1976. Riv. Parasit., 29(63): 87-103.

Satya, Gopal, K., Vijayalakshmi, C. and Roa, K.H. 1985. Male reproductive system of some digenetic trematodes. Proc. Ind. Acad. Sci. (Animal Science), 94: 145-151.

317

Saxena, A.M. and Prakash, S. 2006. Study of a new species of the genus Lecithocladium from marine fish, Gazza minuta (BL). Flora and Fauna J., 12(1): 21-23.

Shaikh, G.S., Shaikh, R., Shaikh, A.H., Abbasi, A.R. 2002. Sterrhurus tetradontus n.sp. (Trematoda: Hemiuridae: Sterrhurinae) from the fish Tetradon lunaris (BL. SCHIN) of Karachi coast. Proc. Parasit., 35: 81- 86.

Shen, J. 1983. Two new species to the family Pleorchiidae Poche,1926 (Trematoda) from some marine fishes in China. Ocean. Limn. Sinica., 14: 396-401 (In Chinese, English summary).

Shen, J.W. 1990. Description of four new species (Lepocreadiidae and Hemiuridae) and a list of digenetic trematodes of fishes from Yellow Sea. Mar. Sci. Bull., 9: 54-63 (In Chinese).

Shen, J.W. 1990. Digenetic trematodes of marine fishes from Hainan Island. Beijing: Sci. Publ., pp. 228 (In Chinese, English summary).

Shen, J.W. & Qiu Z.Z. 1995. Studies on the trematodes of fishes from the Yellow Sea and the Bo Hai Sea. Beijing: Sci. Press, pp. 207 (In Chinese).

Siddiqi, A.H. and Cable, R.M. 1960. Digenetic trematodes of marine fishes of Puerto Rico. Scientific Survey of Porto Rico and the Virgin Islands, 17: 257-369.

318

Siebold, C.T.E. von 1848. Lehrbucher der vergleichende Anatomie der wirbellosen Thiere Berlin, pp. 679.

Sinclair, N.R., Smith, F.G. and Sullivan, J.J. 1972. The Stomachicola rubea: Tubulovesicula pinguis enigma. Proc. Helm. Soc. Wash., 39: 253- 258.

Sinitzin, D.F. 1931. Uber einige neue und wenig bekannte Organe der digenetischen Trematoden. Zool. Anz., (25): 767-770.

Skrjabin, K.I. and Guschanskaja, L.H. 1954. [Sub-order Hemiurata (Markevitsch, 1951)] in Skrjabin, K.I. [Parasites of Animals and Man], Moskova, 9: 227-650.

Slusarski, W. 1958. The adult Digenea from Salmonidae of the basin of the Vistula and the South Baltic. Acta Parasit. Pol., 6(22): 249-528.

Smith, C.F. 1939. Gene Mitogenomic analysis for coelancanths. Latineria chalumnae- A living fish of Mesozoic type. Linkinghub. elsevier.com/retrive/pii/50378111906006251.

Sogandares-Bernal, F. 1959. Digenetic trematodes of marine fishes from the Gulf of Panama and Bimini, British West Indies. Tul. Stud. Zool., 7: 69-117.

Sogandares-Bernal, F. and Hutton, R.F. 1959. Studies on helminth parasites from the coast of Florida III. Digenetic trematodes of marine fishes of Tampa and Boca Ciega Bays. J. Parasit., 45(3): 337-346.

319

Sogandares-Bernal, F. and Hutton, R.F. 1959. Studies on helminth parasites from the coast of Florida III. Digenetic trematodes of marine fishes of Tampa and Boca Ciega Bays with descriptions of 2 new sp. Bull. Mar. Sc. Gulf and Caribbean. J. Parasit., 9(1): 53-68.

Son, TQ., Hoi, VS., Dan, TD., Nga, C., Toan, TQ., Chau, LV., DE, NV., Son, DT., Luong, PD., Ngoc, TQ. and Lima dos Santos 1997. Application of hazard analysis and critical control point (HACCP) as a possible control measure against Clonorchis sinensis in silver carp, Hypophthalmicthys molitrix. Paper presented as the FAO/NACA/WHO study group on Food Safety Aspects of Products of Aquaculture, Bangkok, Thailand.

Sornmani, S., Vivatanasesth, P., Impand, P., Phatihatakorn, W., Sitabutra, P. and Schelp, FP. 1984. Infection and re-infection rates of opisthorchiasis in the water resource development area of Nam Pong project Khon Kaen Province, Northeast Thailand. Annals Trop. Med. Parasit., 78: 649-656.

Sparks, A.K. 1957. Some digenetic trematodes of fishes of Grand Isle, Louisiana. La. Acad. Sci., 20: 71-82.

Sparks, A.K. and Thatcher, V.E. 1958. A new species of Stephanostomum (Trematoda: Acanthocolpidae) from marine fishes of the Northern Gulf of Mexico. Trans. Amer. Micro. Soc., 77: 287-290.

320

Srivastava, H.D. 1936. New hemiurids from Indian marine fishes.I. A new parasite of the sub-family Prosorchiinae Yamaguti, 1934. Proc. Nat. Acad. Sci. India, 6(2): 175-178.

Srivastava, H.D. 1937. Parasites of the family Acanthocolpidae Luhe, 1909, from Indian marine food fishes. Proc. 24. Ind. Sci. Cong., pp. 297- 298.

Srivastava, H.D. 1938. A new parasite of the family Acanthocolpidae Luhe, 1909, from an Indian host. Ind. J. Veter. Sci. Anim. Husb., 8: 247- 248.

Srivastava, H.D. 1938. Studies on the gasterostomatum parasite of Indian food fishes III. Ind. J. Veter. Sci. & Anim. Husb., 8(4): 317-340.

Srivastava, H.D. 1939. Two new genera of digenetic trematodes from Indian hosts. Proc. 25. Ind. Sci. Cong., pp. 262.

Srivastava, H.D. 1942. New hemiurid trematodes from Indian marine food fishes. Part-III. Two new parasites of the genus Lecithocladium Luhe, 1901 (Sub-family: Diurinae Looss, 1907). J. Parasit., 34: 124-127.

Srivastava, C.B. 1963. On three new species of the genus Bucephalus Baer, 1827 (Bucephalidae) with remarks on the systematic position of the B. indicus Srivastava 1938. Ind. J. Helm., 15(1): 36-44.

321

Srivastava, L.P. 1966. A re-description of Stephanostomum caducum Looss, 1901 (Digenea: Acanthocolpidae) from the intestine of Onos mustelus (L.). Ann. Mag. Nat. Hist., 9: 399-403.

Stafford, J. 1900. Some undescribed trematodes. Zool. Jahrb. Syst., 13: 399-414.

Stafford, J. 1904. Trematodes from Canadian fishes. Zool. Anz., 27(16- 17): 481-495.

Stephen, A. and John, S. Marr, 2006. Human Paras. Dis. Source bk.

Stossich, M. 1883. Brani di elmintologia tergestina. Serie prima. Bol. Soc. Adr. Sci. Nat. Trieste, 8: 111-121.

Stossich, M. 1885. I distomi dei pesci marini e d’aqua dolce. Lavoro Monografico Trieste, pp. 66.

Stossich, M. 1888. Appendice al mio Lavoro distomi dei pescimarini ed’ aqua dolce. Progr. Ginn. Comun. Sup. Trieste V., 25: 14pp.

Stossich, M. 1898. Description of Bucephalus n.sp. (Trematoda: Bucephalidae). A parasite of Eel Anguilla J. Parasit. - bioone.org.

Stossich, M. 1901. The digenetic trematodes of marine fishes of Tortugas, Florida H.W. Hunter-Amer. Midl. Nat., 1947- JSTOR.

322

Stunkard, H.W. 1930. Another trematode with two anal openings. Anat. Rec., 47(3): 363pp.

Stunkard, H.W. 1961. Cercaria dipterocerca Miller and Northup, 1926 and Stephanostomum dentatum (Linton, 1900) Manter, 1931. Biol. Bull. Mar. Biol. Lab. Woods Hole, 120: 221-237.

Szidat, L. 1944. Weitere Unterschungen Uber die Trematoden Fauna einheimischer Susswasser fische. II. Mitteilung Dic Gattung Sphaerostomum (Stiles et hass; 1898) Looss, 1899 und Verwandte. Z. Parasit., 13: 183-214.

Szidat, L. 1961. Versuch einer Zoogeographie des Sud-Atlantik mit hilfe von LeitparaSiten der Meeresfische. Parasit. Schrift., 13: 1-98.

Szidat, L. 1963. Los parasitos de los mitilidos y los danos por ellos causados. Neotropica, 9: 30-85.

Thatcher, V.E. 1999. Two new Bucephalidae (Trematoda) from fishes of the Brazilian Amazon. Acta Amazon., 29: 331-335.

Toman, G. 1992. Digenetic trematodes of marine teleosts fishes from the Seychelles, Indian Ocean. IV. Acta Parasit. Pol., 37: 127-130.

Travassos, L. Artigas, P. and Pereira, C. 1928. Fauna helmintologica dos peixes de agua doce do Brasil. Arch. Inst. Biol. Defesa Agric. Anim., 1: 5- 68.

323

Travassos, L., Freitas, J.F.T. and Buhrnheim, P.F. 1966. Trematodeos de peixes do litoral Capixaba. Plagioporus multilobatus n.sp., parasito de Palmituma. Atas Soc. Biol. Rio Janeiro, 10(2): 35-38.

Tseng, S. 1935. Anatomy of a new appendiculate trematode from the sea eel. Peking Nat. Hist. Bull., 9(3): 171-180.

Tubangui, M.A. and Masilungan, V.A. 1944. Some trematode parasites of fishes in the collection of the University of Philippines. Philipp. J. Sc., 76(3): 57-65.

Upatham, E.S., Viyanant, V., Brockelman, W.Y., Kurathong, S., Lee, P. and Kraengraaeng, R. 1988. Rate of re-infection by Opisthorchis viverrini in an endemic North-east Thai community after chemotherapy. Int. J. Parasit., 18: 643-649.

Velasquez, C. C. 1959. Studies on the family Bucephalidae Poche, 1907 from Philippine fishes. J. Parasit., 44: 449-451.

Velasquez, C. C. 1959. Studies on the family Bucephalidae Poche, 1907 (Trematoda) from Philippine fishes. J. Parasit., 45: 135-147.

Velasquez, C. C. 1961. Some digenetic trematodes of Philippine food fishes. J. Parasit., 47(3): 521-525.

Velasquez, C. C. 1962. Some hemiurid trematodes of Philippine fishes, J. Parasit., 34: 124-127.

324

Vercammen-Grandjean, P.H. 1960. Les trematodes du Lac Kivu Sud (Vermes). Ann. K. Mus. Midden-Africa, Tervuren n. s., (5): 171 pp.

Verma, S.C. 1936. Studies on the family Bucephalidae (Gasterostomata). Part 1. Descriptions of new form of fresh water fishes. Proc. Nat. Acad. Sci. India, 6: 66-89.

Verma, S.L. 1973. Helminth parasites of fresh water fishes. Part XI. On a new trematode Stomachicola mastacembeli n. sp. from the intestine of a fresh water fish Mastacembelus armatus. Zool. Anz., 190: 167-170.

Vigueras, I.P. 1940. Prosogonotrematidae n. fam. Prosogonotrema bilabiatum n.gen. n.sp. Parasito de Ocyurus chrysurus Block. Mem. Soc. Cubana Hist. Nat., 14(3): 249-252.

Vigueras, I.P. 1942. Notas helminthogicas. Monorchistephanostomum gracile n.g. n.sp. (Trematoda: Acanthocolpidae) parasito de Sphyraena barracuda (Pisces). Rev. Univ. Hab., 7(40-42): 193-223 [n.v.].

Vigueras, I.P. 1955. Description de seis especies nuevas de trematodes de la familia Acanthocolpidae division del genero Stephanostomum en sub- generos. Rev. Iber. Parasit. T. Extra., Libro-homenaje al Prof. Lopez- Neyra, pp. 421-441.

Vigueras, I.P. 1955. Description de seis especies nuevas de trematodes de la familie Acanthocolpidae division del genero Stephanostomum en sub- generos. Rev. Iber. Parasit. T. Extra., 23(1): 1-36; 24(1): 17-38.

325

Vigueras, I.P. 1955. Contribution al conocimiento de la fauna Helmintologica Cubana. Mem. Soc. Cubana Hist. Nat., 22: 21-71; 22: 195-233.

Vlasenko, P.V. 1931. Zur Helminthofauna der Schwarzmeerische. Trav. Stat. Biol. Karadagh, 4(4): 88-136.

Wang, P.Q. 1982. Some digenetic trematodes of marine fishes from Fujian Province, China. Ocean. Limn. Sinica, 13: 179-194 (In Chinese).

WHO, 1995. Control of foodborne Trematode infections, WHO technical Report Series 849, Geneva, World Health Organization, pp. 1-5.

Winter, H.A. 1960. Algunos trematodos digeneos de peces marinos de agues del Oceano Pacifico del sur de California. U.S.A. del litoral Mexicano. An. Inst. Biol. Univ. Nac. Auton. Mex., 30: 183-208.

Woodhead, A.E. 1929. Life history studies on the trematode family Bucephalidae I. Trans. Amer. Micr. Soc., 48(3): 256-275.

Wool cock, V., 1935. Digenetic trematodes from some Australian fishes. Parasit., 27(3): 309-331.

Wu, K. 1933. A survey of parasites of fishes in Michigan. Ann. Rep. Michigan Acad. Sci. (1931-1933), pp. 52.

Yamaguti, S. 1934. Studies on the helminth fauna of Japan. Part-II. Trematodes of fishes, I. Japan. J. Zool., 5(3): 249-541.

326

Yamaguti, S. 1937. Studies on the helminth fauna of Japan. Part-I7. Trematodes from a marine fish Branchiostegus japonicus (Houttuyn). Published by author, pp. 15.

Yamaguti, S. 1938. Studies on the helminth fauna of Japan. Part 21. Trematodes of fishes, IV. Published by author, pp. 239.

Yamaguti, S. 1938. Studies on the helminth fauna of Japan. Part 24. Trematodes of fishes, V. Japan. J. Zool., 8(1): 15-74.

Yamaguti, S. 1939. Studies on the helminth fauna of Japan. Part 26. Trematodes of fishes, VI. Japan. J. Zool., 8: 211-230.

Yamaguti, S. 1940. Studies on the helminth fauna of Japan. Part 31. Trematodes of fishes, VII. Japan. J. Zool., 9(1): 35-108.

Yamaguti, S. 1951. Studies on the helminth fauna of Japan. Part 44. Trematodes of fishes, IX. Arb. Med. Fak. Okayama, 7(4): 247-282.

Yamaguti, S. 1951. Studies on the helminth fauna of Japan. Part 44. Trematodes of fishes, X. Japan. J. Zool., 7(4): 315-334.

Yamaguti, S. 1952. Parasitic worms mainly from Celebes. Part I. New digenetic trematodes of fishes. Acta Med. Okayama, 8(2): 146-198.

Yamaguti, S. 1953. Parasitic worms mainly from Celebes. Part III. Digenetic trematodes of fishes. Acta Med. Okayama, 8: 257-295.

327

Yamaguti, S. 1954. Systema helminthes. Part I. Digenetic trematodes of fishes, Satyu Yamaguti, 1953, Tokyo, pp. 405.

Yamaguti, S. 1958. Studies on the helminth fauna of Japan. Part 52. Trematodes of fishes, XI. Public. Seto Mar. Biol. Lab., 6(3): 369-384.

Yamaguti, S. 1958. Systema Helminthum Vol. 1. The Digenetic Trematodes of Vertebrates. Intersci. N.Y., pp. 575.

Yamaguti, S. 1959. Studies on the helminth fauna of Japan. Part 54. Trematodes of fishes, XIII. Public. Seto Mar. Biol. Lab., 7(2): 241-262.

Yamaguti, S. 1970. Digenetic trematodes of Hawaiian fishes. Tokyo: Keigaku Publishing Co., pp. 436.

Yamaguti, S. 1971. Synopsis of Digenetic trematodes of Vertebrates Vol.1 Keigaku Publishing Co. Tokyo, Japan, pp. 1074.

Zaidi, D.A. and Khan, D. 1977. Digenetic trematodes of fishes from Pakitan. Bull. Dept. Zool. Univ. Punjab (New series), 9: 1-56.

Zhukov, E.V. 1960. Endo-parasitic worms of fishes from Sea of Japan and South Kurile shoal. Trudy Zool. Inst. Akad. Nauka SSSR., 28: 1-146.

Zhukov, E.V. 1983. New representatives of the fauna of trematodes from the fishes of the Gulf of Mexico. Parasit., 17: 112-117 (In Russian).

328