UNIVERSITI PUTRA MALAYSIA A SANGUINICOLID BLOOD FLUKE...
Transcript of UNIVERSITI PUTRA MALAYSIA A SANGUINICOLID BLOOD FLUKE...
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 .
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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 .
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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
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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
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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