UNIVERSITI PUTRA MALAYSIA A SANGUINICOLID BLOOD …psasir.upm.edu.my/11777/1/FPSS_1992_2_A.pdf · Morfologi cruoricola lates N. Gen., N. sp. daripada saluran darah ikan siakap ternakan

UNIVERSITI PUTRA MALAYSIA

A SANGUINICOLID BLOOD FLUKE IN SEA BASS (LATES CALCARIFER BLOCH) IN COASTAL PENINSULAR MALAYSIA

BRETT W. HERBERT

FPSS 1992 2

A SANGUINICOLID BLOOD FLUKE IN SEA BASS (LATES CALCAR/FER BLOCH) IN COASTAL PENINSULAR MALAYSIA

By

BRETT W. HERBERT

Thesis submitted in partial fulfilment of the requirements for the Degree of Master of Science in the

Faculty of Fisheries and Marine Science, Universiti Pertanian Malaysia.

June 1992

ACKNOWLEDGEMENTS

I wis h to thank my supervisor , Dr . Faizah S haharom , who

provided guidance , support and as s istance at a l l t imes

during the course of this study , and who , as course

c oordinator , eased the period of ad justment into a new

country .

My s incere thanks to Dr . Has san Daud and Dr . Rohana

Subas inghe on the supervisory committee , who a l so

provided their time , comments and suggestions where

necessary . I also wish to thank the technical staff at

the Facu lty of Fisheries and Marine Sc ience_ , UPM , for

their assistance , particu larly Mrs . Kartini Mohamad who

did the electron microscopy , and Mr . Ros l i As l im , who

taught me the finer points of histological procedures and

photography . Als o , thanks to Mr . Soh Keh Seng , Mr . Z ubir

Baharuddin , Mis s Noraini Abu Has san , and Mis s Jariah

Sulaiman who ass isted in the col lection of sea bas s . I

am most grateful to Mr . Yap Khiam Leong and Mr . Jeremy

Yap , who provided fish and transport to their net c ages

at Pulau Ketam gratis , and who were mos t helpfui in

providing information and were very co-operative at a l l

times .

ii

Thanks also to Prof . J . C . Pearson and Dr . R . M .

Overstreet who gave he lpful comments and s uggest ions on

the description part of the study , and to Dr . Has s an Daud

and Dr . G . Nash , who as s isted in interpretation of some

o f the histological material . I am very grate ful to Dr .

I . G . Anderson who arranged for supply of sea bas s and

working fac i l ities at Oonoonba veterinary Laboratory in

Austra l ia , and who gave me acce s s to his histologic a l

material from both Malays ian and Austral ian sea bas s

infected with sanguinicol ids . Thanks also to Dr . Leong

Tak Seng who furnished fac il ities at univers iti S ains

Mal aysia for examinat ion of sea bas s from Penang , and who

provided acces s to his col lect ion of paras ites o f sea

bas s .

Fina l ly , my sincere thanks to my fami ly , friends and

course mates , whose support and encouragement throughout

the course of this study were inva luable .

III

TABLE OF CONTENTS

Page ACKNOWLEDGEMENTS • • . • . • • . • . • . • • . . . . . . . • . . . . . . . . . . • . . • i i

L IST OF TABLES • • • . • • • • • . • . . . . . . . . . . . . . . . . . . . . . . • . . • . vi

LIST OF FIGURES • • . . . • . . . . . . . . . . . . . . . . . . . . . . • . . . . . . • . vii

L I ST OF PLATES • . . . • • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix

L IST OF ABBREVIATIONS • • . • • • . • . . . . . . • • • • . . . . . . . . . • . . . xi

ABSTRACT • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • . • • • • • • • • xii

ABSTRAK • • • • • • • • • • • • • • • • • • • • • • . • . . . • • • • • • • . . • • • • • • • • • xv

CHAPTER

I INTRODUCTION

I I

Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Sanguinicol ids . . . . . . . . . . . . . . . . . . . . . . . . • . . 2 C lassification and Taxonomy . . . . . . . . . . . . . . 9 Pathological E f fects on the Host . . . . . . . . . 1 2 Sea Bas s Cu lture in Ma lays ia . . . . . . . . . . . . . 2 2

DESCRIPT ION OF A NEW BLOOD FLUKE , CRUORICOLA LATES N . G . , N . SP . ( D IGENEA : SANGU INICOL IDAE ) , FROM CULTURED SEA BASS, LATES CALCARIFER BLOCH 1 7 9 0 ( CENTROPOMIDAE ) •

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Materials and Methods . . . . . . . . . . . . . . . . . . .

Genus Cruoricol a gen . nov . . . . . . . . . . . . . . .

C:-uoric,?l a l ates sp . nov . . . . . . . . . . . . . . . .

D1Scus s 10n . . . . • . . . . . . . . . . . . . . . . . . . . • . . . .

S unun.ary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2 4 2 5 2 6 2 7 3 6 4 6

I I I H I STOPATHOLOGY AND HAEMATOLOGY OF LATES CALCARIFER INFECTED WITH SANGU INICOL ID BLOOD FLUKES Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Materials and Methods . . . . . . . . . . . . . . . . . . .

Res ul ts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . �

Discus s ion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Conc lus ion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Summary

iv

4 8 5 1 5 3 7 2 9 1 9 2

CHAPTER Page

IV DISTRIBUTION OF BLOOD FLUKES OF CULTURED SEA BASS IN MALAYS IA Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 4 Materials and Methods . . . . . . . • . . . . • • . . . . • 9 5 Res u Its . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 7 Discu s s ion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 3 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 8

V CONCLUS ION AND FUTURE DIRECT IONS Conc lus ion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 0 9 Future Directions . • . . . . . • • • • • . . . . . • • . • • • 1 1 3

BIBLIOGRA.PHY . . . . . . • • • . . . • . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 1

APPENDIX . . . . . • . . • . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 3 3

BIOGRA.PHICAL SKETCH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 4 3

v

LIST OF TABLES

Table Page

1 Selected Characteristics of Marine Sanguinicolid Trematodes . . . . . . . . . . . . . . . . 4

2 Distribution of Adult and Juvenile Cruoricol a in Lates cal cari fer from Pu lau Ketam as Determined from Histological Studies . . . . . . . . . . . . . . . . . . . . 54

3 Dis tribution of Cruoricol a Eggs in La tes cal cari fer Tissues as Determined f rom Histological Studies . . . . . . . . . . . . . . 55

4 In fection and Preva lence Rates of Cultured Sea Bas s Examined for Cruori cola la tes in various Locat ions in Malaysia . . . . . . . . . . . . . . . . . . . 98

vi

LIST OF FIGURES

Figure Page

1 Cruoricola lates n . g . , n . sp . Holotype . Dorsal View . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

2 Termina l Genitalia of C. lates n . g . , n . sp . Slightly Convent iona lised to s how Relationships of Ducts and Auxiliary Seminal Vesic le . . . . . . . . . . . . . . . . . . . . . 34

3 Transverse Sect ions of C. lates n . g . , n . sp . 36

4 Sea Bas s Culture Sites in Peninsu lar Malaysia Sampled for C. lates • . . . . . • . . . . . . . . • . 96

5 Numbers of C. lates in Lates calcarifer stocked in May 1991 at Pu lau Ketam . . . . . . . . . . . . 100

6 Regres s ion of Haematoc rit on Weight- Pulau Ketam . . . . . . . . . . . . . . . . . . . . . . . . . . . 135

7 Regres s ion of Serum Prote in on Weight- Pulau Ketam . . . . . . . . . . . . . . . . . . . . . . . . . . . 135

8 Regres s ion of Plasma Prote in on

9

Weight- Pulau Ketam . . . . . . . . . . . . . . . . . . . . . . . . . . . 136

Regre s s ion of Haematocrit on Number of Worms - Pulau Ketam 136

10 Regres s ion of Serum Protein on Number of Worms - Pu lau Ketam . . . . . . . . . . . . . . . . . . 137

11 Regre s s ion of Plasma Protein on Number of Worms - Pulau Ketam . . . . . . . . . . . . . . . . . . 137

12 Regres s ion of Haematocrit on Weight- Pulau Acheh . . . . . . . . . . . . . . . . . . . . . • . . . • • 138

13 Regre s s ion of Serum Protein on Weight- Pulau Acheh . . . . . . . . . . . . . . . . . . . . . . . . . . . 138

14 Regre s s ion of Plasma Protein on We ight- Pulau Acheh . . . . . . . . . . . . • . . . . . . . . . . . . . . 139

VII

Figure Page

15 Regres s ion of Haematoc rit on Number of Worms - Pu lau Acheh . . . . . . . . . . . . . . . . . . . . . . . . 139

16 Regre s s ion of Serum Prote in on Number of Worms - Pu lau Acheh . . . . . . . . . . . . . . . . . . . . . . . . 14 0

17 Regres s ion of Plasma Protein on Number of Worms - Pu lau Acheh . . . . . . . . . . . . . . . . . . . . . . . . 140

18 Regre s s ion of Haematocrit on Weight- Setiu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

19 Regres s ion of Serum Protein on Weight- Setiu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 1

20 Regres s ion of Plasma Protein on Weight- Setiu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142

21 Regres s ion of Haematoc rit on Number of Worms - Setiu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 2

viii

LIST OF PLATES

Plate Page

1 Ventrolateral Submarginal Spine s of Cruoricola lates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 4

2

3

4

5

6

7

Subterminal Mouth of C. lates

Two Adu lt C. lates in Wa l l of Rectum of L. calcarifer . . . . . . . .. . . . . . . . . .. . . .

Dorso-ventral duct-like Structures in the Ovary and vite l laria of C. lates . . . . . . . . . . . . .

Separate Genital Pore s of Cruoricola lates .. .

Four Cruoricola lates in a Hepat ic vein of Lates calcarifer . . . . . . . . . . . . . . . . . . . .

Transverse Section of C. lates in Pericardial Vein and a Degenerated Egg in Fibrocytic Reaction in ventric le ......... .

8 Necrot ic Juveni le Worm Encaps u lated in Fibrocytic React ion in Hepatopancreas

34

37

3 9

4 1

56

56

of Sea Bas s . . • • . . . . • . . . . . . . . . . . . . . . . . . . . . . . • . 57

9 Necrotic Juvenile Pres umed C. lates in Connective Tissue at Base of Gi l l Filament . . . . 57

10 Adu lt C. lates in Mesenteric Venule of Sea Bas s . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

11 Live C. lates Eggs Adhered to Af ferent Filamental Artery Wa ll in Gi l l of Lates calcarifer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60

1 2 Two Miracidia Prior to Escape from primary Lame l lar Epithe l ium . . . . . . . . . . . . . . . . . . . . . . . . . . . 60

1 3 S ingle Miracidium Immediate ly Prior to Es cape from Gil l . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 1

IX

plate

1 4

1 5

1 6

1 7

1 8

1 9

2 0

2 1

2 2

2 3

2 4

2 5

Mirac idium Escaping from Gi l l Epithe l ium o f Sea Bas s . . . . . . . . . . . . . . . . . . . . . .

Les ion Le ft by Escaped Mirac idium in Gill Epithel ium . . . . . . . . . . . . . . . . . . . . . . . . . .

Inflammatory Reaction Induced by Presence of Mu ltiple Mirac idia in One Location . . . . . . •

Haemorrhage and Inflammation around Mu ltiple Miracidia in Gi l l Filament

Eggs of C. lates Ins ide Af ferent Filamental Artery of Sea Bas s . . . . . . . . . . . . . . .

Miracidium in Ventric le of Heart Surrounded by Macrophages . . . . . . . . . . . . . . . . . . .

Dead Mirac idium in ventric le of Heart

Nec rotic Eggs and Dead Juvenile Worm in Pancreatic Tis sue of Sea Bas s . . . . . • . . . • . .

Pigmented Macrophage Aggregate Surrounded on Two Sides by Necrotic Eggs of C. lates ........................... .

Replacement of Pancreatic Tis s ue in Me sentery by C. lates Eggs . . . . . . . . . . . . . . . . . .

MMC s , Me lanomacrophage s and Necrot ic C. lates Eggs in Head Kidney of Sea Bas s

Formation of Apparent MMCs in Cauda l Kidney of L. calcarifer

x

Page

6 1

6 2

6 2

6 3

6 4

6 5

6 5

6 7

6 7

6 8

7 0

7 0

LIST OF ABBREVIATIONS

ant . - anterior

c - caeca

c i - c irrus

co - anterior commissure

cp - c irrus pouch

EGC - eos inophilic granu lar cell

F . - female genital pore

FeR - food convers ion ratio

fp - female pore

g - gland cells

M . - ma le genital pore

m - Mehlis' gland

MMC - melanomacrophage centre

n - nerve c anal

o - ovary

oe - oes ophagus

ov - oviduct

00 - ootype

PER - prote in eff ic iency ratio

post - posterior

s - sperm

sd - sperm duct

sp - spine

sv - s eminal ves ic le

t - testis

u - uterus

ue - uterine egg

�l - micrometre

v - vite l laria

vd - vite l l ine duct

vr - vite l l ine reservoir

xi

Abstract of thesis submitted to the Senate of Universiti Pertanian Malaysia in partial fu l fi lment of

the requirements for the degree of Mas ter of Science .

A SANGUINICOLID BLOOD FLUKE IN SEA BASS (LATES CALCARIFER BLOCH) IN COASTAL PENINSULAR MALAYSIA

By

BRETT W. HERBERT

June 1 9 92

Chairman : Dr . Faizah Shaharom

Faculty : Fisheries and Marine Sc ience

Cruoricola lates N . Gen . , N . Sp . from the blood ves s e l s

of cu ltured s e a bas s (Lates calcarifer Bloch 1 7 9 0 ) i s

described . It 1S a lanceolate s anguinicolid with a

s ingle column of submarginal , ventral s pines; extens ive

vite l larium ; and X-s haped intest ine. The s ingle , lobed

testis extends laterally to the caeca. The cirrus l ies

dorsal to the spherical seminal ve s icle . Auxi l iary

seminal ves icle present. The uterus is post-ovarian ,

part ly s ituated between the lobes o f the ovary ; mid-

portion is thick . Cruoricola lates N . Gen . , N. Sp . is

found in all sea bas s over lSg we ight in the type

local ity , Pulau Ketam .

XII

The adults of C. lates primarily inhabit the venous

circulat ion of Lates calcarifer. Eggs were found in the

kidney , l iver , ventric le of the heart and g i l l s o f a l l

f i s h examined three months after stocking .

Cruoricola lates eggs in tis sues evoke a cel lular immune

response cons isting of encapsulation by either act ivated

macrophages and /or endothe l ial cells. In the heart this

is accompanied by macrophage infiltration. In the

kidneys , encapsulation of eggs is fol lowed by p igment

depos ition in and around the capsu le . The main foci of

pathological e f fect are the panc reat ic ac inar t is sue ,

head kidney , and intertubu lar caudal kidney tis sue .

Cruoricola lates egg deposition in these tissues may have

a negative e f fect on growth through reduct ion in food

convers ion ratio and depres s ion of immunological

capabi l ity .

Haemato logical parameters (haematocrit , serum protein ,

plasma protein) were so variable that no re lationship

between them and infect ion with blood f lukes cou ld be

de scribed .

XIII

Cruori col a lates was present in sea bas s cu l ture s ites

s ampled in Penang , Johore , Pahang and Terengganu .

Ke lantan s ites appeared not to have high incidence of

infection , probably due to the fres hwater influence .

Intens ity and prevalence of infect ion appear to increase

with intens ity of culture .

Histological and dissect ion techniques are complementary

in giving a comprehens ive picture of the locat ion of

worms and eggs in the host . As Cruori cola lates is

readily avai lable , and as sea bass are eas i ly maintained

under laboratory conditions , there is wide scope for

further studies on this worm and its relations hip with

the host .

XIV

Abstrak tes is yang dikemukakan kepada Senat univers iti Pertanian Malaysia , sebagai memenuhi sebahagian daripada

keperluan untuk mendapat I j azah Master Sains .

FLUK DARAH SANGUIHICOLID DALAM IKAN S IAKAP (LATES CALCARIFER) DI PANTAI SEMEHANJUHG MALAYS IA

oleh

BRETT W. HERBERT

Jun 1 9 9 2

Pengerus i : Dr . Faizah Shaharom

Fakulti : Perikanan dan Sains Samudera

Morfologi cruoricol a l a t es N . Gen . , N . sp . daripada

saluran darah ikan s iakap ternakan ( La t es calcari fer

B loch 1 7 90 ) dihuraikan . Ia adalah f luk darah

sanguinicol id yang berbentuk daun dengan s atu deretan

spina bawah pinggiran di bahagian ventral , vi telarium

yang luas dan usus berbentuk silang . Testis tunggal

berlobus meman j ang kebahagian s i s i sekum . S irus

meman j ang di bahagian dorsal ves ike l sperma yang

berbentuk s fera . Terdapat ves ike l sperma auks i l iari .

Uterus terdapat di bahagian be lakang ovari dan

sebahagiannya terletak antara lobus ovari ; bahagian

tengah uterus adalah tebal . Cruori col a l ates d i j umpai

dalam semua ikan s iakap yang beratnya me lebihi 1 5 gram di

lokas i tertentu iaitu Pulau Ketam , Selangor . Cruoricol a

xv

l ates yang matang tinggal di dalam peredaran vena Lates

cal cari fer . Telur di j umpai di dalam ginj a l , hati ,

ventrike l j antung dan insang pada semua ikan yang

diperiksa t iga bulan se lepas pe lepas an . Telur C. l a t es

dalam tisu merangsangkan tindakan imun selular iaitu

termasuk pengkapsu lan sarna ada oleh makrofaj terakt i f

dan/ atau sel endote l ium . Di dalam j antung tindakan ini

disertai dengan penyusupan makrofaj . Di dalam ginjal

pengkapsu lan telur diikuti oleh endapan pigmen di dalam

dan di seke l i l ing kapsul . Kesan patologi terutamanya

terdapat di tisu as inar pankreas , kepala antara tubu l

ginjal dan tisu ginja l kauda . Ada kemungkinan telur C .

l a tes d i dalam tisu ini mempunyai ke san negat if k e atas

pertumbuhan L . calcari fer me lalui pengurangan dalam

nisbah pertukaran makanan dan penurunan keupayaan

keimunan . Parameter hematologi ( hematokrit , protein

serum , protein plasma ) sangat berbe za hingga perhubungan

antara parameter ini dan jangkitan f luk darah t idak dapat

dihuraikan .

Cruori col a l a tes terdapat di dalam ternakan ikan s iakap

yang disampe l dari Pu lau Pinang , Johor , Pahang dan

Terengganu . Kawasan di Ke lantan tidak mempunyai ins iden

j angkitan yang tinggi mungkin disebabkan o leh pengaruh

XVI

air tawar . Keamatan j angkitan meningkat dengan

peningkatan keamatan ku ltur ikan. Teknik histologi dan

pembedahan adalah sal ing membantu dalam membuat gambaran

menyeluruh tentang lokasi cacing dan telur dalam perumah .

Oleh kerana c . l a tes mudah diperolehi dan ikan s i akap

mudah dis impan di dalam makmal maka terdapat s kop yang

besar bagi ka j ian lanjut tentang cacing ini dan

perhubungannya dengan perumah .

xvii

CHAPTER I

INTRODUCTION

Foreword

D igenetic trematodes of the fami ly Sanguinicolidae are

parasites which inhabit the circu latory system of marine

and freshwater f ishes . They have been found in the blood

ves se l s , lymphatic system and cardiac tis sues of a

variety o f e lasmobranchs and teleosts . Very l itt l e i s

known about their presence i n f i s h from the South-East

As ian region .

Sanguinicol ids are the only digeneans in which adu lts

are o f economic importance in fish cu lture ( Smith , 1 9 7 2 ;

Bauer et a l . , 1 9 7 3 ) . Most adu lt digeneans l ive in organs

which have direct connection to the outs ide , al lowing

eggs to pas s out with minimal or no e f fect on the host .

As blood f lukes l ive and reproduce within the c ircu latory

system , their presence may cause pathological c hanges in

hos t t i s sues in which the paras ites , or their eggs , have

become lodged . Reports of blood flukes in cu ltured

fis hes are few , probably becau se they are difficult to

f ind and are rarely looked for .

( =Plehniel l a ) has only been

1

The genus San guin i co l a

recorded in f reshwater

2

f i s hes , while a l l other genera of the fami ly have been

reported in marine fishes ( Short , 1 954 ) .

Sanguinicolids

Introduct ion

Fish blood f lukes were f irst described f rom a marine

f i s h in 1 9 0 0 ( Odhner , 1 9 0 0 ) and from fre s hwater fish in

1 9 0 5 ( Plehn , 1 9 0 5 ) . San guinicola spp . were only

recognised as digeneans later ( Odhner , 1 9 1 1 ) , probably

due to their unusual characteristic of having no suckers ,

which are a prominent morphological trait of most other

d igeneans . To date , the fami ly Sanguinicol idae cons ists

o f e ighteen genera ( Overstreet and Koie , 1 9 8 9 ) . For the

purposes o f this discuss ion , " sanguinicol ids" re fers to

members of the families Sanguinicol idae and

Aporocoti l idae of yamaguti ( 1 9 58 , 1 9 7 1 ) , unless otherwise

stated .

Character i s t i c s of Sanguinicolids

Few s anguinicol id life cyc les have been e luc idated and

all , with the exception of Aporocotyl e simpl ex ( a marine

blood fluke ) , are members of the genus San gu i n i col a P lehn

1 9 0 5. Sanguinicola s pp . are exc l u s ively freshwater

( Short , 1 954 ; Yamaguti , 1 958 ) . Characters used in

taxonomy of d igeneans often inc lude the characteristics

o f larval forms , particularly cil iated plate patterns of

miracidia and excretory sys tems in cercariae , as these

3

may indicate phylogenetic relationships ( Chandler and

Read , 1 961 ) . Due to the lack of known life cyc les , most

descriptions of marine sanguinicol ids are only from adu lt

forms .

The habitat within the host is one o f the diagnostic

features o f the sanguinicol ids . As adults , they l ive

e ither in the lymphatic or blood circu latory system or in

the coelom of fish . Most species are found in the blood .

Two species ( Deon tacyl ix ovalis and Pl ehni el l a

( =San guinicol a ) coelomi col a inhabit the coelom ( Manter ,

1 9 4 7 ; S z idat , 1 9 5 1 ) . Koie ( 1 9 8 2 ) found Aporocotyl e

simpl ex in the lymphatic and blood c irculatory sys tems .

Sanguin i cola i nermi s may deve lop in the s kin o f their

carp hosts ( Sommervi l le and Iqbal , 1 9 9 1 ) . . The only

record of a s anguinicolid ( Plehniel l a (=San gu i n i col a )

den tata ) within the lumen o f the intestine o f a fish

( Paperna , 1 9 6 4 a , b ) is open to question , as upon opening

the intestine blood ves sels wi l l be broken , a l lowing

s anguinicol ids in them to escape .

The disti nguis hing morphological features of

s anguinicol ids , and the principal feature that gives them

fami l ial status , is the absence of a pharynx and l ac k o f

wel l developed suckers . Al l other features of the fami ly

as proposed by von Graf f ( 1 9 0 7 ) are absent in one or more

members ( Table 1 ) . Of these , important features pre sent

Table 1. Selected Characteristics of Marine Sanguinicolid Trematodes.

Abbrevia tions are as follows: ant.- anterior; post.- posterior; F- female genital porej M- male genital pore; CP- cirrus pouch; y- cirrus/cirrus pouch presentj N- cirrus/cirrus pouch absent; V-L - ventrolateral.

Genus and Source Uterus Genital Position of Testes Testes Ovary position pores genital pore nwnber position

Aporocotyle preovarian common median, many intercaecal intercaecal (Yamaguti,1958: genital sinistral preuterine

Thulin,1980) atrium

Cardicola postovarian separate M. post.&lat. intercaecal mid-postcaecal (Short,1953; to F.; area: level with

Yamaguti,197l) sinistral; or preuterine postovarian

Chir.;aerohemecus postovarian separate M.dorsal, intercaecal intercaecal (van der sinistral. pre-uterine

Land. 1957) F. ant. to M.

De::g?:acylix extends separace M. post. to F. between postcaecal; (!·lan,er. 1947; preovarian Both right of nerve trunks sinistral

Yamagut i. 1971) medi"ln

f-iype::ar.drot rema postovarian separate M. post. to F. intercaecal most (Naillat"d and Both on left intercaecal.

Ktari,1978) side pre-uter ine. post-testic1l1rir

Merapleh.'1iella postovarian separate M. post. to F. mostly postcaecal;media (Lebedev & F.median, M. postcaecal n

Parukhin.1972) on left pre-uterine

Inte stine Armature CP Cir rus

H-shaped minute, Y Y clumped spines

H-shaped transverse N' y rows of or small V-L N spines

bifurcated ventro- .,. Y lateral margins

H-shaped dorsal and N with lateral diverticules

bifurcated lateral N y

H-shaped fine; N marginal; paired

�

Genus and Source

Neopara cardi cola (Yamaguti,1970;

1971 )

Orchispiriurn ( Madhavi and

Rao,1970)

Paracardicola (Martin,1960)

Paracardicoloides (Mactin, 1974)

paradeontacylix (McIntosh, 1934;

Oqawa & Egusa, 1986)

Pearsonell urn (Overstreet &

Koie,1989)

Plechorchis (Ma rtin , 1975)

Psetcarium (Goto & ozaki,

1929; 1930)

Oterus

e xtends preova rian

extends preovarian

postovarian

preovarian

post ova ria n

extends preovarian

preovarian

postovarian

Table 1 (Continued)

Genital Position of pores genital po re

separate M. post. to Lat marginal notch

common submedian

separate but clos e

common

separate

separate

common

sep arate

postova rian

near mid seminal vesicle level

preovarian, on right side

F. median; H. sinistral and post. to F.

M. dorsa l sinistral; post. to F.; F. a nteromedial .

near posterior on left

F.median; M. post .to and sinistral to F.

Testes number

2

1

2

2

many

1

>100

1 (?) diffuse

Testes position

ant. inter­caecal post.post­caecal

intercaecal transversely coiled tube

postcaecal; preovarian and postovarian

postcaecal; in hindbody

2 media n intercaecal rows

mostly intercaeca1

postcaecal

not clear, extends postovarian

Ovary po sition

postcaecal; between ant. margin of post. testis and right side of body

postcaecal; post-testicular; right of median; mostly pre­uterine

postcaecal preuterine

post uterine; postcaecal, between testes

inter- or postcaecal; pr euteri ne

postcaecal; post-testicular; median

post uterine postcaeca1

postcaecal, 1a tera lly bound by testis; dextral

Intestine

X-shaped with diverticules

bifu rca ted

H-shaped

bifurca t ed

H-shaped

H-shaped

bifurcated

H- sh ape d

Armature

t .... o V-L ro .... s

? possibly lost

la tera lly spined

V-L ban d sheathed in tegument

V-L rows :>f spines; rose thorn hooks posterior

V-L transverse rows

lateral clusters of 3-4

ventro· lateral transverse rows

CP Cir�s

Y Y

y y

Y

y'

y

y

y

N

y

y or N

y

Y

y

VI

Genus and Source Uterus Genital Position of Testes Testes Ovary position Intestine Armature CP Cirrus pores genital pore number position

Pserraroides postovarian separate M.lateral on 1 preovarian postcaecal; H-shaped V-L ? Y (Lebedev & right;F. posttesticular; with transverse

Parukhin, 1972) median & ant. median diverticules rows

Psuedocardlcola Extends separate F. sub-median 5 intercaecal postcaecal, X-shaped ? Y Y (Parukhin, 1985) preovarian & ant. to preovarian sinistral

marginal M.

Selachohemecus postovarian common median 1 preovarian; pre-uterine; four very V-L margin N (Short, 1954) postcaecal posttesticular; short caeca single

median (=X) spines

Cruorlcola postovarian separate sinistral, M. 1 preovarian postcaecal H-shaped V-L single Y Y sinistral & postcaecal posttesticular row lateral. to F. width of body

Table 1 (Con t inued )

1. Except C. congruenta Lebedev and Marnaev, 1 9 6 8 a ppears to have a ci rrus pouch . 2. If cirr a t e .

m

7

in most genera are the X- or H-s haped intestine with s hort

anterior limbs , med ian ovary or ovaries , lac k of Laure r's cana l ,

and Y-shaped exc retory vesic le . Hooks and/or spines , a lthough

not mentioned by von Graf f , have been found in mos t

sanguinicolids desc ribed to date .

Life Cycle

Al l b lood f lukes have a s imp le , two host life cyc le ( Koie ,

1 982 ) . The l ife cyc le of aquatic sangu inicol ids usua l ly involves

a mol lu sc , the intermediate host where cercariae deve lop, and a

fish , which is the definitive host . Most of the sangu inicolid

l i fe cyc les e luc idated to date belong to the genus Sangui n i col a .

The l i fe cyc le of only one marine blood f luke , Aporocotyl e

simpl ex Odhner 1 9 0 0 , has been determined ( Koie , 1 9 8 2 ) .

A l l sanguinicol ids produce eggs , which most often hatch

\orithin the gi l l f i laments . The eggs may or may not be

operculate . In Sanguin i cola spp . the miracidia break through the

g i l l epithelium to the outside ( Bauer et al . , 1 9 7 3 ; Evans 1 9 7 4 a ;

Anderson and Shaharom-Harrison , 1 9 8 6 ) . Presumably , most

sanguinicol id eggs develop in the host and the miracidia active ly

escape , although in some cases the eggs escape from damaged gi l l

fi laments be fore hatching ( Grabda , 1 9 9 1 ) . One sanguinico l id ,

Chimaerohemecus trondheimensi s , is unique in that 9 - 2 0 miracidia

develop inside one egg ( Thulin , 1 9 8 2 ) .

I f the miracidium locates a suitable host it bores in , and

after two generations of sporocysts , cercariae are produced . No