0111-0200.pdf - Indian Academy of Sciences

S T U D I E S I N H E L M I N T H O L O G Y *

T r e m a t o d e Paras i t e s os Birds

BY i~AKUND BEHARI LAL, I).Sc.

( Le'ct~trer in Zoology, The University, Luckno'w, India)

l%eceived June 2, 1939

(Communicated by Dr. G. S. Thapar, M.so., Ph.9.)

COi~TI~i~TS

INTRODUCTION AND ACI~.NOWLEDGViENTS . . . . . .

HISTORICAL REVlEW . . . . . . . . . .

COLLEC'I'ION OF IV~ATERIAL . . . . . . . .

TECHNIQIJ~ . . . . . . . . . . . .

CLASSlFICATIOlV . . . . . . . . . . �9 �9

(a) C h a r a c t e r s of S y s t e m a t i c I m p o r t a n c e . . . .

(b) S y s t e m a t i c D e s c r i p t i o n of t h e F a m i l i e s . . . .

~TAMILY i'~OTOCOTu

G e n u s No~ocot.y[us .. G e n u s H i n d i a . .

G e n u s Naviforraia . . Gen•s ]~aramonostomum G e n u s Neo~aramonostomurn

G e n u s Catatropis ..

]F~,MILY CYCLOC(ELID2I~ ..

Genus Hcematotrephus G e n u s Cycloccelum .. G e n u s TyphloŸ G e n u s Contraccelurn G e n u s O~hthalmo2bhagus

I~ 'A~ILu DICROC(ELIID3E

G e n u s ZyŸ243 G e n u s _P latyno s orn um G e n u s Multivitell•

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* Part of the thesis subn¡ and approved for the Degree of Doctor of Science in the University of Lucknow, 1937.

111

B1

112 Makund Behari Lal

I2AMILY OPISTIIORCIIIID,-I~ . .

Genus Opisthorchis . . . .

~A~%s LECITHODENDI<IID2~ . .

Genus Parabascus . . . .

FAMILY ~{ETEROPIIVIDT~ . .

Oenus Ascocotyle . . . .

FAMILY I}IICROPHALLiD.E . .

Genus Levinsenidla . . . .

FAMILY ~EPHALOGONIMIDTE . .

Genus Prosthogonimus . .

FAiM:ILY ]~CHINOSTOi'r ..

Genus Ecl~inostoma . . Genus Echinochasmus . . Genus Stej~hanoŸ . .

Genus Parorchis . . . .

Cenus Paryphos tomum . .

Genus Petasiger . . . .

FA~ILY PSILOSTOMID2E . .

Genus Psilorchis . . . .

FAlVIILY STRIGE];D~ . .

Genus Cyathocotyle . . Genus NeodiŸ Genus Proalaria . . Genus Neoalaria . . Oenus Ph~ryngos tomum

~AMILu S CI:fISTOSOMIDTE . .

Genus Orni~hobilharzia . .

Gen_us Chinhuta . . . .

Oenus Gigantobilharzia . .

HOST-PARASITE I~ELATIONSHIP ..

COPUL&TION IN TREMATODES ..

GEOGI<APIKICAL DISTRIBUTION ..

I~EFEI%ENCES . . . . . .

LlsT oF ABBREVIATIONS . .

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S[udies i~z IIeladTzl/mIo~y 113

INTRODTJCTIOI~ AND ~CI~i~OWL]~DG!%IENTS

BIRDS forro one of the most important economie units of animals, some of them even constituting the delicacies of our table, but very little attention has so faz been paid to the study of their Trematode parasites in India. In view of their economic importante and also in order to remove some existing confusion in the description and classi¡ of these parasites, I detezmined, at the suggestion of Dr. G. S. Tha.par, to work out the morpho- logy and the systematic position of Avian Trematodes from India. In spite of the fact that bi1"ds are migrants, results have not been disappoint- ing and I have been able to collect mate¡ and data that add to our

knowledge of this group.

Part of the work has akeady been published and is referred to at its

propez place in the thesis.

The work was carzied out in the Zoology Depaztment of the Lucknow University during the years 1934-36. I wish to recozd here my deep indebtedness to Dr. G. S. Thapar fox his kindly advice and criticism throughout the progress of my work. He fu~her very kindly allowed me free access to his valuable personal library and plaeed part of his own collection of Avian Treraatodes at my disposal for which I express my sincere thanks.

Several othe1" colleagues rendered valuable help in my work~Mr. J. Dayal in going through the typed manuseript ; Dr. (Miss) ]Ÿ Speer and Dr. A. C. Chatterii in the t~anslation of some German literature ; l%fr. K. S. Iyer in going through some literature in Fzench~and I take this opportunity to put on record my sense of gratitude for their help. I aro also thankful to iVfr. G. l~. i~atu, our artist, for technical assistance in the preparation of illustrations. IVfy thanks are also due to the authorities of the Zoological Survey of India for providŸ facilities to consult their Library. To Pzof. Birbal Sahni, l~.R.s., I aro indebted for the loan of a few iournals from bis p¡ library, and last but not the least I aro thankful to the authorities of the Lucknow University for facflities given to carry out these investiga-

tions. I-IISTORICAL REVlEW

The wolk on AvŸ Trematodes from India is hazdly more than a decade old. The earliest recozd, to whieh I have been able to refer, is that of Bhalerao (1926) where he describes ¡ species of Trernatodes, v4z., Echª corvus, Stephano#rom rey~oldi, Phtzneroibso~us inso~em, LyŸ ka.ke~ and Pl~ttynosomum acuminatum from the Burmese crow, Corvus insolens. Of these, Platynosomum acuminatum has previously been reported from the livel" of a Kestrel from Scotland by ~icoll (191£


Phadke and Gulati (1930) recorded a new genus M~�91 from the gall bladder of the common house crow, Corvus splendens and created a new subfamily Multivitellarin~e for lis reception.

~Ioghe (1932) described two new species Echinostomun~ govindum and Pararnonostomum microstomum from the recial cmca and the small intestine respectively of the Indian Ruff, Philomachus Ÿ

Harshe (1932) described a new species CatatroŸ orientalis from the recial c~eca of the pintail duck, Da fila acuta acuta.

Gogate (1934) recorded four trematodes Echinostoma revolutum, Pary- phostomum testitrifolium, Petasiger minutissimus and Ornithobilharzia sp. from wild ducks in Rangoon.

Srivastava (1935) described a new species, AscocotyIe intermedius, from the intestine of the Indian Fishing Eagle, Halir leucoryphus, and further emended the diagnosis of tire genus Ascocotyle. The same year (1935) he described another species, Catatro2bis indicus from the recial cmea of an Indian fowl, Gallus bankiva murghi.

Bhalerao (1935) descfibed two species, Notocotylus babai from the c~ecum of the common kite, Milvs migrans govinda, and Cycloccelum sharadi from the thoracic cavity of the western yellow-billed magpie, Urocissa favirostris cucullata.

Khan (1935) described eight new species of the genus Cycloco~lum, viz., Cycloccelum lobatum, Cycloccelum nebularium, Cidoccelum straightum Cyclocoelum indicum, Cycloccelum cafiellum, Cycloccelurn mehrii, Cycloccelum allahabadi and Cydoccelum erythropis from three different genera of the Snipes.

Verma (!935) descfibed Echinochasmus bagulai ~rom the small intestine of the Pond heron, Ardeola grayi and the ~ ight heron, Nycticorax nycticorax, and also added Echinochasmus ruficaŸ243 from the intestine of the Indian little grebe of diver, Podiceps ruficollis. In the same paper mention is made of a smM1 vafiety of Prosthogonimus and two other representatŸ of the genus Prosthogonimus, but unfortunately no account of these forros is so lar available. A chambered exeretory bladder was deseribed by Verma in the genus Echinochasmus and the diagnosis of the genus was emended, with a suggestion to split ii up into two sub-genera.

Patwardhan (1935) recorded Ly#erosomum colorosum from the gall bladder of a black-headed mynah, Temenuchus 2bagodarum ; Proalaria alcedensis from the intestJne of the" king-fisher, A lcedo atthis ; and Neodiplostomum tytense from the intestine of a Barn owl, Tyto alba stertens.

Studies i~z 2~ef~ni,~zt~ofogy 115

Thapar and Lal (1935) ,odded a new genus Psilorchis under the family Ps~lostomidŸ from the jntestine of the king-¡ and considered the evolution of several struetures in the family Psilostomidm.

Later, Lal (1935) d› a new species Notocotyh,ts ª from the intestinal c~eca of the wigeon, M~rec~ pen.elope.

Again, Lal (1935) reviewed • genus Notocotylus and discussed the importance of the position of the genital opening. On the basis of this charaeter he created two llew genera, Hi~dia and Navifor~nia.

Lal (1936) described two new species of the genus Pc~ramonostonutm, viz., Paramonr querque&du~~ from the intestinal c~eca of the garganey, Querqued~d~z circia ; and Paramo~~ostom,um cas~rcum from the intestinal c~eca' of the Brahminy duck, Cesarca rut, ila. The value of the position of the genital pore as of generic importanee in the classifieation of the members of the subfamily lXTotocotytin~e was further eonfirmed, a n d a new genus Ne@aramonostomum was created.

Verma (1936) recorded thir ty- three new forros of trematodes from Indian bi§ 5uf gave no diagrams and added very meagre and incomplete deseriptions. Ir is, therefore, not easy to diagnose and inelude these forros with their present confused aceount at this stage, particularly because the author himself, as appears from bis paper, is in doubt regarding the description of many of bis forros. The same year (1936) he also described CyethocoXyle calv,~~si from the intestine of the King Vulture, Torgos c(zlw~s.

Lal (1936) described a new speeies .Parorchis s~,d23is from the cloaca of the common summer snipe, Ÿ162 hypoleucos. The discovery of a speeies of Ÿ from India in the snipe is interesting in so far as this genus has only been recorded from gulls and restricted only to St. Andrews, North- umberland coast, igillport and Ameriea. In the course of a disenssion on the systematic position of the genus a new subfamily Parorchin~e was ereated for its reeeption, a n d a suggestion was put forward regarding the polyphyletic origin of the family Echinostomid~e.

Lal (1936) wkile deseribing a new genus .Typhl@hilus shovd~us from the intestine of a shoveller duek, SŸ clyŸ demonstrated the presence of a wel:-developed muscular ventral sueker in the famity Cycloeadida~ and also discussed the advisabitity of retaining the subfamilies Cyeloccelinm and Typhloecelinm. The signiffcance of a ventral sueker a s a basis for generie

divisions was also pointed out.


Lal (!936) recorded a new species Levinseniella indica and suggested the probable identity of several existing species of the genus with the genus Maritrema owing to the presenee of a cirrus sac.

I r would, thus, appear tha t so far I" only scattered contributions have been made in India on the Avian trematodes and no comprehensive and col- leetive work has been attempted. I have, therefore, endeavoured to present here the results of my investigations, together with summaries of desc¡ tions of other forros given by earlier workers. The latter is indicated by small type to differen•iate it from new and original work.

~OLLECTION OF ~/fATERIAL

A large number of birds were subjeeted to post-mortem examinations in normal salt solution during the years 1934-36, and the Trematodes recovered from them ate enumerated in the following table :

LoeaIity of the host

~[alhaur

Lucknow

J p

Lucknow and ~tapur

Lucknow

Name of host

Casarca rutiIa (Brahminy duck)

To~anus hypoleucos (Summer snipe)

GaIlinago ffallinuIa (Jack snipe)

Spatula clypeata (Shovellor duck)

Mareca peneloTe (The Wigeon)

T otanus f tLscus (Spotted red shank)

~)icrur~s macrzcercu8 (King crow)

Up~pa erops (The Hoopoe)

Ardeola grayf (P~ddy bird)

-=~~~ I I Loc~tion o[

parasi te

Intestinal cmca

Cloaca

Bursa Fabricii

Intestine

Intestinal cseca

Intestine

Name of tlm trematode

Paramanostomum cazar. mtm Lal, 1936

Parorchfs sni2es L~I, 1936

Lev~naeniella indica LaI 1936

TyphIophilus shovdlus Lal, 1936

Notocotylus indicus L~I I935 and Hindia lucio. nowen~is Lal, 1935

SteThanoTrora fusca n. fap.

Intestine

~ 1 7 6

EcMnochazmut megaviteL lus n. sp.

t Since submission of the thesis ]~Iehra (1937) described .5epoderm:z bMd~�91191 LeToderma casarcii, Zepoderma ferm~ginum. Vidyax6hi (1937) described Apharyngostrigea ardeolina, A. indiana, Str~yea orientali and Strigea nephroni~. Ch~tterji (1938) mentions _Prosthogonimus sp. a~ud Catatropfs verrur Pande (1938) describes Crassiphiala stunk~rdii. The characters of these species a te not summarized here.

Sxudies in Helmi~dholoEy 117

Loca]ity of the host

N.~me of hosr �9 {D

0 4 a 5 "~ Location of Name of the treina,tode

Lucknow

Barabanki and Sitapur

Lucknow

Amausi

Luckaow

Luclmow and Sitapur

Ajgain and Chinhut

Kukr~,il, Lucknow

Lucknow

Ardea cinerca rectiros. tris (Common grey heron)

Athcne Brama (Spotted owlet)

Passer domesticus (House sparrow)

K ittacinda ~nelabarica (Sham~)

Milvus migrans (Cerumen kite)

Sarcogyps calcus (King vulture)

Acridotheres tristis (The myn~h)

Querquedula r (The garganey)

Ha[oyen smyrnensis (King-fisher)

Psittacula kran~eri (Green parakr

Ooracias be'nghalens~s (The blue jay)

Corwts ~plendene (Cerumen houEe crow

Cinnyr~ zeylonicus (Sun bird)

Net•ion creces (Cerumen Teal)

Sterba aurantia (Common river tern)

Bulbucus ibis coro-

(Cattle egret)

6 1

11

9 0

7 3

9 4

i

2 1

9 5

6 1

10 0

6 1

10 2

8 0

5 2

8 0

5 1

Intestine

2 Sma.ll intestine

Liver

~ p

Intestine

Bursa fabricii

Intestil]al c~eca

Small intestine

Intr

Intestino

Intestine

Main blood vessels : Kidney, Lung, Liver, etc.

~

Renal vr

Pharyngostomum bag~�91 n, sp.

~Veodiplostomum dilac~- cura n. sp.

Opisthorchis cheelis n. sp.

Ol)~sth~rchis giddhi~ n.sp.

Neoalaria iba, paria n.g., n.sp.

t)rosthogonimua cu~~eatus

Pa• ql�91 quedulum Lal, 1936

Eohinostoma chasma 12 .S ~).

Psilorchis indicus Thal~~r and LM, 1930

NeodiyIostomum sp.

~c]~inor rr n,sp.

Psilvrchis Ajgain{s LaI, 1938

Chinhuta indica Lal, 1937

Giga~zto5ilharzia egret, a Lal, 1937

118 Makund Behari La1

TEci4NIQuE

Most of the Trematode material eollected was ¡ under slight pressure of the coverglass in 90% alcohol. Some forma were fixed in Bouin's Fluid and Chrom-osmic for section-cutting. In certain cases a speeial method of narcotising and killing the worms in 5% alcohol and gradual transfer to 30% alcohol was found very useful. The worms, thus treated, were fixed in a welt-stretched condition without pressure and displacement of organs of injury to parta. This was speeially helpful in examining the ventral glands of the family Notoeotylidae. The material was preserved in all cases in 70% alcohol of sometimes in 70% alcohol-glycerine. The speeimens for whole mounts were stained with Ehrlieh's aeid h~matoxyl in in a dilution of ! : 20 of distilled water and differentiated in tap-water overnight. This proeedure gave a uniformly brilliant stain. Paracarmine stain was also used. Although it stained the structures rather deep, it gave good results in differ- entiating some of the deep-seated struetures. Clearing was done in Clove oil of Xylol. In certain cases sections, both transverse and longitudinal, were eut, and for this double embedding Celloidin-Paraffin method was used. The sections were stained with haematoxylin (Delafield's) and Eosin.

A special glyeerine-aleohol method of clearing was found useful in certain tases, specially ill the Echinostomidae. The unstained material was kept in 70% alcohol-glycerine in proportion of 3 : 1 and left for about hall an hour. Later ir was transferred t o 90~ alcohol-glycerine in proportion of 4=:1 and left overnight. The alcohol gradually evaporates leaving specimens in pure glycerine which also thoroughly penetrates into the specimens. The spines, reproduetive organs, genital pote, etc., were easily seen by this method.

~LASSIFICATION

(oz) Characters of Systematic Import~nce

The author has already published a separate paper on this aspect of the sub~eet and has ffdly discussed the value of different characters in the classification of AvŸ Trematodes (vide these Proceedings, 1937, 5, No. 2,

Sec. ]3, 33-4~).

The systematic aceount of the various families given in the ~ollowing pages is on the lines of the views expressed in the above published paper.

(b) Systen~atic Descr@tion of the Fam~Zies Fam�91 NotocotyZid~e Lfihe, 1909

The family Notocotylidm was formed by Lª (1909) to eontain origin- ally only three genera NotocotyZus, Catatr@is and Paramonostomum.

SLudies .in Helmi~athology 119

Later on several workers added to this list with the result that the family was split up into three subfamilies, Notocotylinm, Nudacotylinae and Ogmo- gasterinm.

The writer in the course of bis investigations on the family observed certain peculiar features, describn later, which necessitates the modifica- tion of the existing diagnosis of the family. The emended diagnosis is us Ÿ : - -

Diagnosis of the family Notocotylidce Lª 1909, emended.--

Trematodes with fiar body, without a head-collar. Skin smooth of thickly covered with fine spines in the anterior part of the body and whole of the ventral surface. Ventral pa~billr very rareIy present. The ventral suriace provided in most cases with regular rows of unŸ glands. Oral sucker simple. Ventral sucker absent. Pharynx not present. Oeso- phagus short. Intestinal c,'~ca simple and end blindly. Excretory pote slightly dorsal. Excretory bladder with a short unpaired median stem and a pair of long limbs which branch and anastomose dorsal to the oesophagus. Testes symmetrical at the posterior end, lateral to the intestinal c~eca. Cir•us sac very ]ong. Vesicu~a serninaIis Zying ex~ernal of j~ar~ly interna~ and part~y extemr Ovary lies between the restes. Shell-gland in front of ovary. Receptaculum seminis absent. Laurer's canal present. Uterus runs in fairly regular transverse loops between the intestinal cmca and in front of the ovary. Vitellaria well-developed and lie in front of the restes, mostly extra-c,'ecal. Genital pote 2bre-equatorial ancl median, behind the intes~ina~ bifurcation or i~, front of it, sometimes lar forwarcl a~ tt, e side of the oral suc,~er of post-equatorial ana lateral. Eggs with long thread-like filaments at both ends. Parasitic in aquatic Birds and ~�91

Of the three subfamilies, Notocotylinm, lq'udacotylinm and Ogmo- gasterin~e, only !qotocotylina~ is recorded from avian hosts in India.

Diago~r of the subfanr Notocotytin~, 14ossack, 1911, emended.--

~otocotylidm, generally with �91 rows of ventral glands on the ventral surface, sometimes absent. Vesicula seminalis lying externa1 or partly interna1 and parfly externa1. Genital pote almost median, pre-equatorial, behind of in front of the intestinal fork. Vitellaria well-developed in the posterior half of the body and lateral to intestinal caca. Testes at the extreme end of the body, symmetrical. Ovary intertesticular, separated from testes by intestinal crura. Uterine coils interca~cal.

Type genus--Notocotylus Sens. Str. B2 F


The author has already made$ elaborate studies on the various genera of the subfamily ~otocotyl inm and as a resnIt of his invest igat ions "che following genera have been reeognized under the subfamily.

Genus Notocoty7us Sens. Str. (Lal, 1935). ,, Hi~zdi~z Lal, 1935.

,, Naviformia Lal, 1935.

,, Paramonos~omum Sens. Str. (Lal, 1936).

,, NeoJ)aramonostomum Lal, !936.

,, Catatrofois Odhner, !905.

Genus Notocotylus Sens. Str. (Lal, 1935)

Monostomes wit.h tu,o to five roze's of ~rotrusible ventral gla~zds ; pha rynx absent ; testes extra-emcal and poster ior ; ovary in between restes ; shell- glands pre-ovarial; uterine loops eonfined to the inter-cmcal region behind the cirrus sac ; reeeptaculum seminis absent ; pa r t of the vesicnla seminalis enclosed withia cirrus sae; cirrus sac #tever exter beyo~~d half of body le•gth ; vagina about �89 t,o as long as cirrus sac ; ger~ita,~ pore situated behind the ir~testinal bifurcation ; vitdline gZands exte~.zd ~ff)to the middZe of the body ; eggs -0!8-.022 mm. long with filaments at either end.

Type species--Notocotylus czttenualus.

Notocotylus indicus Lal, 1935

Monostome measuring 2- 18 mm. x .63 mm. Three rows of protrusible ventral glands, !7, !6, 17. Oral sueker subterminal -12mm. • . ] 3 m m . Oesophagus -075 mm. long. In tes t inal caeea t e rmina te a t a clistance of �9 2 mm. on the right and �9 19 mm. on the left side from the posterior end. Right testis .86 mm. long ; left testis .38 mm. long. Cirrns sae .58 mm. long. Ovary �9 18 mm. X �9 16 mm. Eggs small and numerous with thin shell

and a filament at each end, measuring .018 mm. x .0!I mm. without filaments.

Host.--Mareca pe,~elope (Intest inal cmca).

Locali ty.--Lucknow.

Genus Hindia Lal, 1935

Monostomes with three rows of fbrotrusible veatral glands ; p h a r y n x absent ; restes extra-emeal and poster ior ; ovary in between tes tes ; sheli- glands pre-ovariat ; nterine loops lŸ in the inter-caecal region behind the

$ Vide Proc. Ind. Acad. Sci., 1935, 2, No. 5, ~19-23 ancI ~57-66. Also 1936 , 3, No. 1, 25-34.

Sludies i~z He[mbz1]zo�91 121

cirrus sac; receptaculum seminis absent ; part of the vesicula seminalis eixclosed wi th in the cifras sac ; cirrus sac ezte~di~g about t to �89 body length ; vagina never more than -~ of the cirrus sac ; gota:tal poro at the intestinal fork ; vitelli~,e glands extend -~ to to- of the body length from the posterior end ; eggs �9 014-.0209 mm. long., with ¡ at ei ther end.

Type species--Hindia gibbus.

Hindia l'uck~owensis Lal, 1935

Monostome measuring 2.63 mm. • .74 mm. Ventral glands 16, 15, 16. Oral sucker subterminal .15 mm. in diameter. Oesophagus .13 mm. long. In tes t ina l cmca terminate at a distance of �9 2 mm. on the right and �9 15 mm. on the left sido from the posterior end. Right testis -35 mm. • -175 mm. Left testis -375 mm. • .2 mm. Cirrus sac -825 mm. long. Cirrus about �9 3 mm. long., armed with several rows of spine. Ovary �9 175 mm. x - 15 mm. I2ggs wi th th in shell a n d a filament at each e n d ; measuring .02 mm. x - 01 mm. excluding filaments.

Host.~Mareca pe~zelope (Intest inal crece).

LocaZity.--Lv_eknow.

Hindia babel (Bhalerao, 1935) Lal, 1935

(Syn. Notocol:ylus babel Bhalerao, 1935)

Body smM], elor~g~te, eltipLicle, measur ing 3.83 mm. in l eng th ana .93 in m~ximum breadt, h a~ lit, t le behincl anterior end. Cutiele covered wi th minu to spines. Three rows of un i f o rm ,nmd eversible ventxM glands present, ; lateral ones wit, h 17 glands eactl, and middle one wiŸ 15 gla,nds. Oratl sucker is sub- terminaI and me,~sures -2mm. x �9 16 mm. Oesop]lagus measu~es .12 mm. in lenp;th. .The imtestinM cmea en.d ~~t a dis tante of �9 265 m m . fvom posto•or end.. Testas, elongate, cleeply lobed, symmet r ica l ; extrac0ec&l a~~([ Inea~ure -535--5~t~5 mm. • - 2 d : - .2mm. Cirrus sac, elongated, club-shapecl and measures 1.09 mm. Genital poro media,n, ven t ra l to i~ltestinaI bifurcatioa. Vesicul~ seminMis presmlt inside the cirrus sac. P~~rs prost,atic~~ wide and elongated, measures .435 mm . nar row m~tscul�91 cversible eirrui measures .575 mm. O~zary deeply lobed, median, in between two restes. I t measures .3 x . 3 1 mm. Shell-glaad, pre-ovar iM ana measures .235 mm. x :165 mm. Ovirluct e~larges into uterus wlgch pa,sses aa~erior ly in 18 tr~msverse loops which aro mos t ly inI, ercmcal. Vagina measures ,56 m m . Laure r ' s ca,nal present~. Recep taeu lum semiais absent . Vitel[aria follieul~r, extra-cincel, ~ po r t i on interemeM. They extenr be tween ~ntserior bor&er of testes to an te r io r 3/5 of the body. Yolk reservoir dorsal to shell-glaad. Eggs measure �9 014~--.017 m m. x . 008-- -011 mm. with loag fllaments a t ei ther polo. A short exc re to ry bladder. E x c r e t o r y poro dorsal ~z~d median ne~r the posterior eacl.

Host .~Milvus migrans govinda (C~ecum).

L o caIity.~ Ra~goon.

Remarks.~T~le forro is transferred to the genus Hindia beeause of the

posit ion of its geni tal pote and other charac*ers.

122 Makund Behari La]

Geims Naviformia l.,at, 19~~5

Monostomes with three rows of protrusibIe ve.~ztra! Jauds ; pharynx absent ; restes extra-czecat and poster ior ; ow~ry in between the test es ; shell-glands pre-ovarial ; uterhle coils lie in t he inter-ct~'cal regiou 1)r the cŸ sae ; reeeptaculum seminis absent ; par t of the vesicula scmimllis

is enclosed within the cirrus sac ; cirrus sac exlendi'ng about !~ of lhe hod3, length ; vagina ~ of the cirrus sac ; genital pote in ./)'oŸ191 of the inh:stinal Jbrk ; vitdIine gtands behind mid-body; eggs -6178 mm, • .0208 mm, hmg, with,

filaments at either end.

Type speeies--Navi f or~nia navi formes, No speeies recorded from birds from India .

Genus Paramonostomum Sens. Str. (Lal, 1936)

Monostomes without ventral glands a n d a pha rynx ; in tes t ina l c~cca rtlll

almost upto the posterior end of the an ima l ; restes may of may not be lobed, extra-emeal in the posterior par t of the body ; pa r t of the vesictfla seminalis enclosed within the cirrus sae ; ovary in be tween the testes ; shell- glands pre-ovarial ; uterine loops most ly confined to the inter-c~cal regiml behind the eirrus sac; receptaculum seminis absent ; gemita.l pote always @ens in front of the intestinal fork and beldnd the oral suc#er ; vi[d[i,nc g[a.,nds never end behind the mid-body from the posterior end ; eggs wi th filamellls vt eit]xer end,

Paramonostomum querque&dum Lal, t936.

~onos tome measures 3.398 mm. x !. 11 nxl~l. Oral stmker �9 ] 7 nmI. • �9 12 mm. Oesophagus .!65 mm. long. Right test is -66 mm. • .24 mm. Lei t testis .74 mm. x .235 mm. Cirrus sae .93 mm. long, Ovary .2{~5 mm. x .22 mm. Eggs with th in shell a n d a f i lament at each end ; me'tsuring �9 02 mm. x �9 009 mm. without filaments.

Host,--Querquedula circia (Intesti~ml emea).

Locality.--Amausi.

Paramonostomum casarcum Lal, 1936

Monostome measures 3.8 mm. x .95 mm. Oral sucker .14 mm, x �9 1 mm. Oesophagt~s .14 mm. long. Right testis -8 mm. x �9 15 mm. I,eft

testis . 10mm. x -175mm. Cirrus sae . 9 1 m m . long. Ovary , 3 8 m m , • �9 22 mm. Eggs thin-shelled and provided wi th a f i lament at each p o l e ; measuring . 018 mm. • �9 01 mm. withortt filaments,

Host.~Casarca rutila (Intestinal e~eea).

Locality.~Malhaur, Lucknow.

Slz~dz'es i~z ]--f elmint]~o/o~y 123

Genus Neopammo~osLo~rmm I,al, t936

1V~onostomes w,i~ho~~t ventral g~,z~ds and pharynx ; intestinal c~eca run aImost upto the posterior end of the animal ; testes extra-c~cal and posterior and generally lobed; part of the vesicula seminalis enclosed within the cirrus sac; ovary in between the testes; shell glandspre-ovarial; uterine loops mostly confined behind the cirrus sac; Ÿ seminis absent ; receptaculum semi~is ~terinum may be prese~~t ; genital pote always opens behind i~~testinal fork ; vitelli~e gla~~ds e~,d behi~zd mid-body from the posterior e~d; eggs with filaments at either end.

Type species~Neop~ramonostomum io~r

(Syn. _Paramonostomum ionorne Travassos)

Neopammonostomum mierostomu~n (Moghe 1932) Lal, 1935

(Syn. P~ramo~r omum microstomum 2r 19 32) B ody elonga~ed, 1 -348 ruin.--1-62~t- nlm. long. M ~ x i m u m wiclth -375 m m - .409 mm.

an te r io r to restes. Or~l sucker, small, t e rmina l , -063 m m . in d i ame te r . Ventra l suckcr absent. Very short oesophagus; ph~rynx and prepharynx absent. Intestina/ cmc~ extend up to .05 mm. from posterior oncl. Testes, extra-crac,uf at the posterior end of anim~l, elongated, and lobed, -13--.l£ mm. long. Cirrus sac in anterior half of body, between the intestin,~l cmca, -.ll3 mm. long. Ir cont~dns vesicula seminMis straight and n~rrow, ~nd opons at genita/pore, i~umodiŸ below the intestin~l bifur- o~tion. Ov~ry lies in the medi~n line botween the restes, intercmcal, shell-gI~md posterior to ov~ry. • wi th 13-15 loops on e i the r sido. Vagin~ as long Ÿ cirrus sac. Ute rus ex tends Mong more th~n .{ of the body . Vitelh~ri~ e x t e n d f r o m ~nLerior end of res tes to abou t middle of the uterus region. Eggs provi&ed w i th lo~g polaz' filaments Ÿ measure -051 mn:t. long.

Host.~PhŸ pug~~ax (Small intestine).

Locality.--Nagpur.

Remarks.--It is rather peculiar that the she11-gland has been describid by ~oghe posterior to the ovary, a condition which is unique in the family lXlotocotylidm. This apparently appears to be a case of misrepresentation of structures as has been pointed out by some 0ther authors also. But in case the post-ovarial position of she11-gland is confirmed in other specimens, ir would not only affect our concept of the family Notocotylidm but would also place ir sti11 nearer the family Pronocephalidse.

Genus CatatroŸ Od.hner, 1905

~onostomes with three rows of weakly developed ~o~r ventral gMnds ; the middle row of ventral gl~nds m,zy be placed on a median keel of ,z ridg~ ; pharynx absent ; testes extra-ca~cal and posterior ; ovary intertesticular; shell-gland pre-ovarial; vesicula seminalis is divisible into an


external and an internal portion ; receptaculum seminis absent; va gi~m is very much devel@ed, abou~ as long as cirrus sac ; ge~zital pore situated behª the i~testinal fork; eggs with ¡ at either pole.

Type species~Catatropis verrucosa.

Catatr@is orientalis I<arshe, 1932

Remarks.--t-Iarshe while deseribing this new species does not mention any~hing about the nature of the ventral glands present, whether they are protrusible or non-protrusible. The question of protrusibility of non- protrusibility is a vital one, as it is the most important dii~erence 5:�91 which Catatropis is distinguished from Notocotylus, ttindia and Naviformia. This species, therefore, should in the opinion of the w¡ be kept as a species i~r till more light is thrown by a reinvestigation of the ~orm and hence the characters of Catatr@is orientalis, sp. inq. ate not summarised here.

Catatropis i~~dicus S¡ 1935

Worms lighL brown in colour, 4 .6 m m . long and 1.2 mm. broad. Dorsa l surf~ce com, ex, ~'entral surface concave wi th 3 longitudinM rows of non-pro~rusible unicellular g lands- -g lands in median row conf, iguous, those in laterM rows d is t inc t and 10-12 in number in each row. Exc re t roy sy s t em similar to Lhat seen in Catab'opis. Oral sucker �9 14--2 mm . in diameter ; oesophagus .2- .26 mm. long ; phaxynx absent . Testes cleeply lobed, -75- .99 mm. • -2- .3 mm. , extrac~ecM ; vesicula seminMis enormous, outside the cirrus sac. Cirrus sac flask-shaped, -87-1-2 m m . long anct . 17 - . 2 m m . broaB conta in ing cone-shaped pars prosta• - 3 5 m m . • *09 m m . su r round› b y prosta te ceHs. Genital pote close be}¡ oral sucker. Ova ry lobed, -26--35 m m . size, intercmcmt, in leve] wi th restes a m d ' p o s t e r i o r to shetI-gland. L~ure r ' s c~n~l presenfi. RecepSaculum seminis ahsent . Vitella~da, i r regular follicles. I%ecep~~cul~im seminis u t e ¡ present . Llte~als in tr~xxsverse coils, l~[etraLerm m u s c u l a r as long as cirros sac. Eggs thin-shelled, w i th long polgr ¡ T h e y me~sure -017- .02 mm. x -00 �91 mm. wi thout ¡

Host ~ Gallus bankiva murghi (Recta1 cmca).

Remarks.~This forro is remarkable in having the genital pore Ÿ forwards at the posterior margin of the oral sucker. This position of the gential pore appears to be unique for the genus. The vesicula seminalis also is wholly external in Catatropis indicus The importance of this character was emphasizecl by Chatterji (1933) who created a new genus Ganada in the subfamily Lepodermatinm on the presence of an external vesŸ seminalis, besides the usual internal vesicula seminalis. Owing to the unique position of the genital pore, the absence of a median ridge or keel for bearing ventral glands and lastly in the absence of an internal vesicula seminalis the w¡ feels inelined to remove this species from the

Sfudies in ]u191 125

genus Cat~tropis. Probably a re-study of more material of this forro will necessitate the creation of a new genus for this species.

Remarks on the family Notoco~ylidce.--In the preceding pages and also earlier in the pages of these Proceedi~zgs* the writer has given an account of the subfamily Notocotylin~e. The sub-families Nudacotylin~e and Ogmogasterin~e possess such distinctive characters when compared with Notocotylina that there could be no doub* for their retention as distinct subfamilies. Yamaguti (1933) described a new genus Cymbiforma from the intestine of a maroma1, Sika nippon and created the subfamily Cymbiformin~e for its reception. The genus shows af¡ with the members of the subfamily Nudacotylina~ but differs in the relative position of the cirrus sac and uterine coils. Yama£ also mentions, tha t Barker (1915), in describing the subfamily lXTudacotylinm, covered variations in these charaeters as we11. Ir would, thus, appear tha t the difference in the relative position of uterine coils and cirrus sac should not forro a sufficient basis for the erection of a new subfamily Cymbiforminm. The question of a mammalian host lodging this form may rise a little doubt b u t a s already discussed ea~lier in the Host-parasite relationship (vide these Proaeedings, 1937, 5, No. 2, Sec. B, 33-44) this poin* may be negligible. The subfamily Cymbiformime may, therefore, be dropped and be regarded a s a synonym of the subfamily 2qudacotylinm, and the genus 6"ymbiforma~ r comes under the subfamily 1~udacotylinm.

FamiIy Cyciocceli&e Kossack, 1911

Stossich (!902) crezted a subfamily Cycloccelinm for the reception of four genera, CyclocoeTum, H~ma�91237 OŸ and TyŸ Kossack (I911) raisecl ii to the status of a family, Cycloccelid~e and added to ir two genera, HyŸ and ~a~r t-Ie aiso inclu4ed the genus Bothriogaster under this family, t-Iarrah (!922), while revising the monostomes of North America, divided the family into th:ee subfamilies : - -

(1) Cycloc~linse with C2clocaehtm, Ha~matotre2bhus and Hyptiasmus.

(2) Typhloccelinm with Ty13hloccclum and Tracheophilus.

(3) Ophthalmophaginse with 0!bhthalmophagus , BothrŸ an4 Spaniometra.

Witenberg (1926) fused the subfamŸ Ophthalmophagina~ and Cyclo- cceHnm, leaving only two subfamilies, Cyclocoelin~e and Typhloco~linm. The

* V i d e 1935, 2, ~o . 5 ; ~nd 1936, 3, No. 1.


main point on which Witenberg maintained only two absence or presence of ca'.cal diverticula.

Family diagnosis.~Trematodes, large to meditan muscular fiar body; small ventral sucker sometimes terminal of subterminal, oral sucker zudimexttary ;

subfamilies is the

size with strong present. Mouth mostly al~st,nt.

Pharynx non-muscular. Intestinal cmca join each other at the posterior end. They are either simple of provided with Iateral diverticula on their inner margin. Excretory pote dorsal and short, situated at the posterior end. Genital pote median, not very much behind the mouth opening. Copulatory organ present but feebly devel£ Seminal vesicle lies insidc the cirrus sac. Vitellaria, lateral, between body-wall and intestine, somc- times run continuously along theintestine and meet at the posterior end. Gonads interc~ecal. Testes two, simple or pressed, and lie near cach other, Ovary, always entire and spherical. Laurer's canal and receptaculum seminis exceptionally present. Uterus very strongly developed with regular peculiarly pointed loops and extends from the posterior end up to the intestinal bifurcation. Eggs numerous, without polar filaments and prodnc- ing miracidia with two eyespots while still inside the uterus. Generally parasitic in the body cavity, air-sacs and nasal opening of aquatic birds. Exceptionally present in the intestinal canal.

Type genus--Cycloc~lum.

The family is divided into two subfamilies : ~

(1) CycloCcelin~e without lateral intestii~al diverticula.

(2) Typhloc0elinm__ with lateral intestinal diverticula.

The AvŸ Trematodes reported from India under the subfamily Cyclo- ccelinm belong to the following genera :--

7. Hcematotre:ht~us.

2. Cycloco~tum.

Key for the identi¡ of the Indian genera of the subfamily Cyclo- coelinm :

Ovary in front of both the testes which ate not separated by uterine coils-- H cematot r ep h us.

Ovary in between two restes which are always separated by uterine coils~C y clo ccel um.

oCtudies in Hekszinthofo~y 127

Genus H:vrnatotre#hus Stossich, 1902 Only one specles has been reported from avian host in India.

H~em~totre~bhus ~ebulari,~m (Khan, !935)

(Syn. C yc~oc(elum nebularium K h a n , 1935)

Trematode• with medium size, 10-13 mm. long and 2 -3 .5 mm. broad. Ph,%rynx welNdeveloped. Oesophagus S-stmped of str~ight. Intes t inal cmc~~ devoid of lateral diverticula a nd meet postenorl 7 in ah aro -285 mm. in f ront of the posterior end. Excre tory bladder between the intest inal arc and posterior end of the body, with dorsally placed excretory pote. The two round restes ly[ng obliquely behind the ova ry measure 0-7-1 mm. in diameter and ~re not separated 'by u t e ¡ coils. Cirrus sac tubular, dilated posteriorly, extending from the genital pore to the anter ior encl of the intestinM bifnrcation. Ov~ry rounded, smaller than test.~s, and measures .31-- -41 mm. in diaraeter. Laurer's canal absent . Uterus in t ransvere loops with large thiek-shelled eggs. Vitellaria, confinecl to fihe ext reme edges of the body. A sm,%ll yolk reservoir is present. Eggs, without operculum showing miracidia with characterist ic double eyespot, and measuring -12 mm. x .0S7 mm. in size.

Host.--Glottis nebulari~ ( A b d o m i n a l a i r - sacs ) .

Locality.--Allahabad.

Remarks.--Khan w h o i n c l u d e d t h i s s p e c i e s u n d e r t h e g e n u s Cycloccdum has described a small structure lying inside the ovary which he calls a

reeeptaculum seminis. This position of receptaculum seminis is rather

peculiar. Ir is further interesting to note that Khan does not ¡ any

sperms in it and he mentions that Morishita (1924) ¡ termed a similar

structure as 8otype. The present w¡ considers that the structure is possibly not of the nature of receptaculum seminis which evidently is absent

in this form. Owing to the pretesticular position of the ovary, the species

is r e m o v e d t o t h e genus Hr

Oenus Cydocce~um Brandes, 1892

ikTine species of this genus have so lar been recorded from Avian Jaosts

in India. Out of these one Cyc[oc~l~m ~ebul~rium has just been allocated to the genus Hwmato~reph~s. Of the remaining eight, seven have been

described to possess a receptaculum seminis which is absent in the genus

C ycloccdurn.

Cycloccelum ery~hro2bis K h a n , 1935

l_~ng6h 7.5-17 mm. Maximum breaclth 1 .7 -2 .3 mm. OrM sueker feeble -]5 mm. x .1 mm. Pha, rynx . 15 - .25 mm. in diameter. Oesophagus more or less strai~ht. Intestinal aro . 15 mm. in front of the posterior end. Excretory bl~dder �9 !2---15 mm. by -19--34 mm. in size. Excretroy pote median, dorsally situ~ted at hincier end. Gon~ds in posterior fif~h of the body. Testes sphe¡ ; aaterior testis separated by four or ¡ uterk~e coils from posterior tes• Vesicula sen¨ straight


a n d inside the cirrus sac. C~enital pore , ~~entral, j u s t behincl t he p h a r y n x . Cirrus sac c lub-shaped, reactdng rniadle of i n t e s t i n a l b i fu rca t ion . O v a r y v e r y smal l �9 1 9 - - 2 9 m m . in d i ame te r ; shell-gland mass spher ica l ; u t e ru s runs f o r w a r d in c losely s i t u a t e d coils ; m e t r a t e r m short . Vitel lar ia l a t e ra l ly pressed aga ins t bocly-wall . Th in-she l led o r a .

Host.--Tringa ervthro~us.

LocaIity.--Allahabad. Rem arks.--This speeies definitelF confortas to the characters of the

genus Cycloccehtm. Ir does not possess a receptaculum seminis and differs in ttlis important feature from the other species which have been described to possess a receptaculum seminis.

Cycloc~vlum sharadi Bhalerao, 1935

Trematocles wi th body t a p e r i n g t o w a r d s b o t h e n d s ; m e d i u m size, m e a s u r i n g I0-5-II mm. in length ancl 4--1.5 mm. in maxhnum bre~dth. Oral sucker, muscular measuring .44 mm. • .32 mm. Prephar~uxx short. Pharynx well developed, muscular, and measur› -425 mm. x .38 mm. Oesophagus short. ]~xcretory bladder, simple, ¡ sac, excretory pote opens dorsally by a short cluct, l%erve ganglia on either side of the ph~rynx, ~ dorsal nerve-band ~oins them. Genit~~I gl~nds in posterior fourth of the body. Testes separated by ufierine coils. Posterior testis in eontact with intestinal ate and measures 2.15 mm. x 1-32 mm. Anterior testis measures 2.07 mm, x i -7 mm. and is in cont&et with left intestina/c,eeu]n. CilTus sac slnall with cirrus in the forro of horizontal S. Vesicula semina]is lŸ fflls up the cirrus sac. Pars prostatica non- eellular ; prostate glands present. Male opening in the genital atriuln. Genital atrium large and situated posterior to the pharynx, genital pote slightly behind pharynx. Ovary, between two testes, oval, measures .5 mm. • .43 mm. Shell-gland, ]arger than ovary on the leffl of the latter. Receptaculum seminis, oval, on the left of ovary between ir and testis. Ir measures -36 mm. x .275 mm. Vitellaria between cmca and marg i n of body, extending u p t o hinder border of p h a r y n x . L a u r e r ' s cana l a b s e n t . 7Jterine coils in pos ter ior ~rd, intercmcal , m e t r a t e r m s t r o n g l y n m s c u l a r , s u r r o u n d e d w i t h gland-ce]ls. Long and el l ipt ical eggs -123 - . 14 m m . x . 0 6 - . 0 8 1 m m .

Host.--Urocissa f~virostris cucullat~ (Thoracic cavity).

Locality.~Muktesar.

Cyctoccelum allahabadi Khan, 1935

Size 17 m m . x 2 .5 -3 m m . Oral sucker flat. P h a r y n x -28 m m . in d i a m e t e r . Oesophagus S-shaped. Testes sep~ra ted f r o m each o the r b y u t e r ine eoils ; Geni ta l po t e a t pos te r ior end of p h a r y a x . Cirrus sac c lub-shaped, e x t e n d i n g u p t o midd le of in tes t ina l b i furcat ion. Ovary , - 3 - . 3 5 m m . x .2--25 m m . in size. R e c e p t a - cu lum seminis inside the o v a r y ancl well developed, l%eceptaculum seminis u t e r i n u m absen t . Shell-gland poster ior a n d lef t t o ova ry . U te ru s fills up the b o d y b e t w e e n pos te r io r tes t is a n d in tes t ina l b i fu rca t ion . Vitel lar ia , e x t e n d i n g o v e r c e c a aL p laces a n d reaching uter ine coils. O v a th in -she l led ~nd opercu ta te , �9 119 m m . • -08 m m .

Host.--Tring~ erythroJ~us (Thoradc air-sac).

Locality.---~lahabad.

Studies b~ lldmimhobgy 129

Cycgocaegum ca, Ÿ Khan, 1935

I l o d y 17-25 m m . • 2.5-~1.6 m m . Oral sueke r r m l i m e n t a r y . P h a r y n x -275 m m . in d i ame te r . Oesophagus S-shaped of sgraight . E x e r e t o r y pore dorsM a n d t e rmina l . Ÿ sepa ra t e d l:rom each oLher b y u te r ine eoils ; a n t e r i o r tes t is to the r igh t side and ,8 m m . • . 6 - . 8 m m . in size ; poster ior , med ian , much- lobed , 1-1 m m . x . 7 - . 88 m m . in size. Gen i t a l po te a l pos te r io r m a r g i n of p h a r y n x . Girrus sac f lask-shaped and ha rd ly r e a c h i n g int~;stinal b i furca t ion . O v a r y to t h e left s i c I e . - 3 7 - . 5 5 m m . x �9 37- .5 m m . Shel l -gland b:dfind ow~ry. ] g e e e p t a c u l u m s e m i n i s pea r - shaped on the

inner s ide of t h e o v a r y . Igect:l)taeulum seminis uge r inum filled ~~dth sperms. �8 t h r o w n irfi;o doub le loops, never ove r l app ing c:eca. Vite l lar ia , i r regular in extens ion on two sides ; y o l k r e se rvc i r pvesent , . 15 m m . long. O r a wit, h ful ly develoiiecI miracidia , -t~] lxll~t. X .0(IS Inln.

Host.--Capeila gallinago (Cervical air-sacs).

Locali~y. ~Allataabad.

Cydoca~{um straightum Khan, 1935

13ody long, 25 m m . x ~-3 m m . in size ; oral sucker f eeb ly developed, ven t r a l sucker absen t , l ' h a r y n x -344' m m . x .425 m m . Oesophagus S-shaped or s traigh~. Excre - t o r y p o t e dorsa l a n d t e rmina l . Tes tes s e p a r a t e d f r o m each o the r b y u ter ine coils ; an t e r i o r t o t t le r i gh t side amt t . 0 5 m m . x .68 m m . ; pos te r io r mesia l , 1-25 m m . x -99 m m . Cirrus sae ne~~rly flask-shape~t r each ing t o .15 m m . in f ron t of in tes t ina l bifurcN,ion, l~ars pro~~batiea, prostt~te g]~nds a n d eirrus n o t observed. Ovary , median , �9 45 m m . x -41 m m . in size. She[1-glamt to left. of o v a r y ; r ecep tacu lum seminis imme- d ia t e ly behinr o v a r y , Laure r ' s camal absen t . R e c e p t a c u l u m seminis ufierinum large, filled w i t h s pe rm s . Ute rus filling up the space u p t o i n t e s t i n a l b i furca t ion . Vitel laria f r o m midd l e of i n t e s t ina l b i f l l rea t ion to e x c r e t m , y . b l a d d e r . A smal l yo lk reservoir p re sen t . Ow~, th ick-shel lcd, non-opcrcu la te , w i th ful ly deve loped mirac id ia , -136 m m . X .068 : m m .

Host.--Glottis nebularia (Abdominal air-sac).

Locality.~Phulpur, Allahabad.

Cyclocoelum indicum Khan, 1935

L e n g t h 2 0 - 2 7 m m . Brea~lth � 9 1 R u d i m e n t a r y oral sucker . P h a r y ~ x �9 28 m m . in d i a m e t e r . Ocsophagus s t r a igh t . E x c r e t o r y pote dorsal a t hincler end.

Tes t e s s e p a r ~ t e d f r o m each o the r b y u~arine t o t s ; an t e r io r , to the l e f t a n d �9 85 m m . in d i ~ m e t e r ; pos te r ior , in ttte i n t e s t ina l arc . 85 - -93 m m . in d i ame te r . Cirrus sac c tub-shaped . Geui t~ l pore belfind ph~rynx , med ian . O v a r y - 5 - . 6 m m . in d i a m e t e r ; shell-glan(1 n e a r r i g h t wa]l of owt ry ; r e cep t acu lum semin t s e longated inside the o w r y . R e c e p t a c u l u m semŸ u t e r i n u m presen t . U te ru s wi th w ide ly sepa ra t ed loops, e x t e n d i n g o v e r cmca ah& reach ing b o d y walI. ~i telI~l~a i r regu la r in ex~ension on t h e two s ides . O r a , t h in - she l l ed w i t h fuI1y deve loped miracidi~, m e a s u r e 0-12 m m . x .068 m m .

Host.~Glo#is nebularia (Body cavity).



Cydoccelurn rnehrii Khan, 1935

L e n g t h 18-28 mm. , b r e a d t h 3 . �91 m m . O~'~,1 sueke r v e r y ruclimer~~ary. I ' h a r y n x muscub~r 0-27 m m . in d iamete r . Oesoph,~gus S-sh,~ped. E x c r e t o r y p o t e p o s t e r i o r a n d aorso-medi~m. Tes tes s e p a r a t e d f rom eaeh o the r b y u t e r i ne coils ; ar~terior, l~~eral, a n d .85 m m . • 0 . ~ - 1 m m . ; pos te r io r in i n t e s t i na l ~rc, . 9 - 1 . 3 m m . in d i a m e t e r . Cirrus sac re to r t - shaped , exten(liI~g beh ind a n t e r i o r w~ll of i n t e s t i n a l b i fu r ca t i on . Geni ta l pore, v e n t r a l Lo middle of p h ~ r y n x . O r a r F , 4 - . 6 m m . in d i a m e t e r ; she]l. g land elose behŸ ovarF ; r e cep t~cu lum seminis , smal l p e a r - s h a p e d ins ide t he ova ry . ]~ecep tacu lum seminis u t e r i n u m large. Ute rus , w i t h w e l l - s e p a r a t e d loops, ove r l~pp ing ca~2a a t v,~rious places. Vite l lar ia , dense, e x t e n d i n g f r o m midd l e of i n t e s t i n a l b i fu rca - ~ion •o e x c r e t o r v bl~dder. Yo lk re se rvo i r smal l . Ov~, thin-shellect , w i t h fu l ly deve]oped mir~eid ia , �9 12 m m . • .068 m m .

Host.~Capella gallinago gallinago.

Loc~z~ity.--?

Cyc[occslum [obatum I(han, 1935

L e n g t h 1 3 m m . , bre~d[th 2 . 4 m m . Oral sucker n o t v i s ib le ii1 who le m o u n t . P h ~ r y n x muscu la r , .27 m m . in d i ame te r . Oesoph~gus s t r a i g h t . E x c r e t o r y pore mecli~n ~nd dorso - t e rmina l . Tes tes s e p a r a t e d f r o m each o t h e r b y uLerilxe coils ; a n t e r i o r tes t is , l a t e ra l ~Ild equa l in size ~o t h e p o s t e r i o r ; pos t e r io r m e d i a n , -6 .... 75 m m , X -55- .7 m m . Geni ta l pote a t m i dd l e of p h ~ r y n x . Cirrus sac m o r e or less f lask-

shape~ wi~h t u b u l a r an te r io r end and sae- l ike pos te r io r p~ r t r e ach ing j u s t beh ind a n t e r i o r e~xd of in tes t in~] .bifurc~tiom Ov~ry, . 4 - . 5 m m . • -35 - -55 m m . SheLl-gland prese~xt. I~ecep t acu lum seminis inside t he o v a r y . R e c e p b a c u t u m semin i s ube r inum abseI~¨ U t e r u s bebween pos te r io r end of shel l -g land arAd i n t e s t i n a l b i fu rca t ion . Vitell,~ri~ f r o m pos te r io r end of cirrus sac ~o exerebory b ladder . O r a , bhin-shel led � 9 1 non-opercu la te , w i t h ful ly deve loped mi rac id ia , .119 m m . • .068 m m .

Host~Glottis nebul~tria (Thoracic cavity).

L ocal i t y . ~ Allaha bad.

Remarks on the genus Cycloccslum.~The species Cycloccelum sharadi, Cycloccelurn allhabadi, Cycloccdum straightum, Cycloccelum caŸ Cyclo- ccelum indicum, Cycloccelum mehrii ancl Cyclocoelum lobatum have been described ~o 2ossess a receptaculum semirtis. This structure is no~ present in the type species o~ the geuus Cycloccelum. In view of the opinion already expressed ( tal , 1937) ort the systematic value of this charac~er ir may be considered desirable to exclude all these species whŸ possess a recepta- cuhlm seminis from the genus Cycloccelum. I t is suggested that a new genus ReceŸ characterised by the presence of receptaculum seminis, may be created for* their reception.

Subfamily Typhloccelinr Harrah, 1922

Under the subfamily Typhloccelinm, the author discovered a genus TyŸ LaI, 1936, fuli details of which have already been published in a previous number of these .Proceedings (vide 1936, 4, No. 1, Sec. B).

Studies in Hdminzho/or 131

Genus Typhlophilus Lal, 1936

Distome, with a fiar ribbon-like body, and witI1 ah extremely feeble funnel-shaped oral sucker a n d a small muscular ventral sueker. Curved prepharynx, globular pharT~nx and extremely small ~sophagus. Intestinal c~eea, provided with abont 10 diverticula on the inneŸ margin, meet in the middle posteriofly to forro the Intestinal Bow. ~xcreŸ bladder creseent- shaped, excretory pore dorsal and sub-terminal. Testes, two, in grape-like bunehes separated from each other by the first eoil of the uterus. The anterior testis lies on the left and near intestinal c~ecum, the posterior fills the arch near the Intestinal Bow. Ovary oval, on the fight side at the level of the left testis. Ootype, shell-gland a n d a small receptactflum seminis present between the ovary and the posterior testas. Genital pore situated ventral to the intestinal bifureation, immediately behind the pharynx. Eggs small, thin-shelled, withot~t filaments.

Type species--TyŸ237 shovellus.

TyphlopAilus shovellus La1, 1936

Ribbon-like and grey coloured trematode ; 3.6 mm. long and 1-15 mm. broad. Oral sucker extremely feeble, -19 mm. • .195 mm. Ventral sacker circtdar and muscular, abou~ the middle of the body, �9 125 lllm. in diameter. Prepharynx .2 mm. long, Pharynx thiek-walled, .19 mm. • �9 21 mm. Oesolohagus extremely small. In tes t ina l c~eca provided with 10 diverticula on their inner margin and join posteriorly. Excretory bladder .35mm. loug. Anterior testis . !25mm. x -06 mm. Posterior testis -!5 mm. x .07 mm. Cirrus sac .25 mm. long. Ovary at the level of the leŸ testis, .055 mm. x .07 mm. Uterus full of small eggs, eggs measure .02 mm. X .01 mm. with thin shell aud not provided with any

polar filament.

Host.--Spatula clypeata (Small intestine).

Locality.-- Ltxcknow. Remarks on the family Cycloccelidce.--The retention of the two sub-

families Cycloccelin~e and Typhlocr b~ Witenberg (1926) on the basis oi c~ecal diverticula was questioned by Joyeux and Baer (1927) who regard the c~ecal diverticula of only generic importance in the same way as the relative t~osition of gonads. The writer (Lal, 1936) has previously shown that the various genera of Cycloccelidse can be arranged under two d~stinct grout~s on the basis of cmcal diverticula. Both groups have forros which show variations in the relative position of the gonads, the nature of utefine


coils and the position of genital pore, and thus the presence or absencc of cmcal diverticula seem to forro the natural subdivisions of the family. Cameron (1934) and other earlier workers have considered this character i11 the division of the family Fasciolida~ into two subfanlilies, Fasciolinm alld Fasciolopsinm. It is, therefore, desirable to retain here Witenberg's subdivisions into Cycloccelinae and Typhloccelinae.

The subfamily TyphloccMinae has three distinct genera but, as mell- tioned by ~Vitenberg (19-o6), the subfamily CycloccMinm has got a vast assemblage of forros. Joyeux and Baer (1927) llave been rather conser- vative and do not consider the various geliera of Witenberg as vMid. They suggest the fusion of all the genera of the subfamily Cycloccelina~ into two forros, viz., Cycloccelu~~, and SŸ which they distinguish by the relative position of the ovary ancl restes. The genus Cydoca~lum, according to Joyeux and Baer, contains all those forms which have their ovary either pre-testicnlar of intertesticular. Although Witenberg's classitication of the family into genera is not free from defects the writer does not agree with Joyettx and Baer in their eontention. As already stated the relative position of the gonads has beeli greatly emphasised in several cases and should here be regarded as a character of at least generic importante. Ÿ position of the nterine eMls is another charaeter of sufficient importance and eombined with the relative position of the ovary merits eonsideration in establishing the genera of the subfamily Cyeloccelin~e. Therefore, the members of the subfamily Cycloe0elin~e can be gronped into loar distinct divisions, as follows :--

1. Ovary in front of the two testes, not separated by t~terine coils, e.g., Hr

Other forms ate Wardia~lum, HcenwŸ237 Corfl@yrum alld Uvitellina.

9. Ovary in between the two restes separated by uterine eoils, e.g., C),cloca~[um.

Other forros ate Cycloprimum, Harrahium, ProhypEasmus, Ityptias- mus, All@yge and Transccelum.

3. Ovary behind both the testes, not separated by uterine coils, e.g., Co~~y~�91191

4. Ovary behind both the testes which are separated by uterine eoils, e.g., OŸ The genas Sflaniome~m also comes nnder tMs.

.~tudies i~z f:(efmin[~ofa~y 133

Each division is represented by a well-estabHshed genus and the other previously known genera put under eaeh group may be regarded as mete synonyms. Now, there remaiu only two genera, viz., BothriogcŸ and Morishitium which have not been acconnted for here. Both of these possess ventral sueker and ir would be better to keep them as distinct gellera till more information is available about them.

Diagnosis of the ge~zus Ha~matotr@hus Stossich, 1902, emended.--

(Syn. Wardianum Witenberg, 1926 ; Hwmatop'rimu~n Witenberg, 1926 ; CorŸ Witenberg, 1926 and Uvitdlin~t Witenberg, 1926.)

Cyclocoelin~e with ¡ body ; anterior end narrower than the poste¡ ; oral sucker not well developed, subterminal ; veutral sucker absent, intestinal czeea smooth, without any lateral projections, extend up to the p6sterioŸ end of the animal, meeting in the middle line ; testes two, ahnost spherical, lying in the posterior broad end of the animal and not separated by uterine eoils ; ovary spherical, smaller than testes, always in front of both testes, vitelline glands extend along the intestinal c~eca.

Diagnosis of the genus Cycloccdum Brandes, 1892, emended.--

(Syn. Cydoprimum Witenberg, 1926; Harrahium Witenberg, 1926; _Prohyptiasmus Witenberg, 1926; Hyptiasmus Kossack, ].911 ; AlloŸ

Johnston, 1913 and T,ransccclum Witenberg, 1926.)

Cycloccelinm with flat body ; anterior end narrower or may be as broad as posterior; oral sucker more or less funnel-shaped, weakly developed; ventral sueker absent; intestinal c~eca, smooth without projections, run almost up to the posterior end of animal where they meet toge~her in the middle ; testes two, almost spherical, the posterior one lying in the " In- testinal Bow," the anterior further forwards, separated flora each otller by uterine coils; ovary spherical, smaller than restes, lying always in between the two restes ; vitelline glands extend along the intestinal emca.

Diagnosis of the ge~r Contraccdum Witenberg, 1926, emended.--

Cycloccelin~e with ¡ spindle-shaped body ; anterior and posterior ends bote narrower than the middle ; oral sueker quite terminal ; ventral s•cker absent ; intestinal creen, smooth without aay lateral proj eetion, ruu almost

* The n a m e Bothriogaster is n o t ava i l~ble fo r t he gener ic n ~ m e since ir is preoccupied b y Bothriogaster Ssel ivanow, 1879 ( l~yr i~pod~) . I n v iew of th is , t h e n a m e will have to b e changecl.

134 Makund BeharŸ Lal

up to the posterior end of the animal where they meet together in the middle ; testes two, almost spherical, lying side by side, more or less contiguous and not separated by uterine coiIs, in the middIe of the body , ovary oval, smaller than testes, lying behind both testes in the " Intest inal Bow " vitelline glands extend along the intestinal c~eca.

Diagnosis of the ge~us Ophtha~mophagus Stossich, 190~, emended.--

(Syn. SŸ162 Kossack, 1911.)

Cyclocmlinm with flat body; anterior end narrower o r a s broad as the posterior end ; oral sucker weakly developed ; ventral sucker absent ; intestinal cmca smooth, without lateral projections, run almost up to the posterior end of animal where they meet in the middle ; testes two, almost spherical situated in the interc~ecal region and separated by interyention of uterine coils ; ovary spherical, smaller than the testes, lying always behind both the testes ; vitelline glands extend along the intestinal cmca.

FamiIy Dicroccdii&e Looss, 1907

The family Dicrocmliidm contains numerous genera and has been a subject of investigation by a large number of workers. The Iatest comprehensive work is tha t of Poche (1925) where he has recognised not less than ~0 different genera. The main features on which the generic division is based in this family is the relative position of the restes. The distinction and arrangement of the various genera under the family is ra ther confused and a revision of the family wi11 have to be done. The family Dicroeceli- idm has the following diagnosis :--

Trematodes small to medium in size with a more or less elongated, flattened, translucent and non-muscular body. Pha rynx and r present. The intestinal cmea are simple and do not quite reach the posterior end of the body. The excretory bladder is tubular of sac-like. The genital pore is situated in the middle line between the oral and ventral suekers. The testes ate situated in leve1 with of behind the ventral sucker and in front of the ovary, opposite to each other or one behind the other. The cirrus sac is sma11 and does not extend bekŸ the anterior margin of the ventral sucker. The ovary is sit•ated behind the testes. The vitelline glands ate we11 developed and are lateraI1y situated, par t ly overlapping the intestinal cmca. The uterus occupies most of the space behind the genital glands. Eggs relatively small.

Type genus--Dicroc~lum.

Sfudies in Hdminlhofogy 135

Among birds in India, the family is represented by three genera :--

1. Genus L>,j~erosomum Looss, 1899.

9,. Genus Platynosomum Looss, 1907.

3. Genus Multivitellaria Phadke and Gulati; 1930.

Oenas Lyperosomum Looss, 1899

Looss creafed the genus Lyperosornum for the reception of L. porrectum, L. longicauda, L. stro=gSosum aud L. plesiostomum. Braun (1901) added three species to the genus, viz., L. corrigia, L. salebrasum and L. rudectum. Von Linstow (]906) descfibed L. squa~natum but this was later removed from the genus by Skrjabin (1913). Looss again (1907) ineluded under the genus, L. lobatum, L. a~ssoni and L. clathratum, the last one, however, was assigP, ed later to the genus Pla~2nosomum. Skrjabin (]913) added L. fili- forme. Nicoll (1914) described L. scitulum and L. direptum of which the latter was transferred later to the genus Oswaldoia by Travassos (1919). lohns ton (1917) descfibed three species L. parvium, L. megastomum and L. harrisoni. In the same year, T=avassos (1917) added L. obliqum, L. tram- versum, L. rarum, L. lari and L. sinuosum, tsaitsehicoff (]919) descfibed L. dor~icum and L. attenuatum. Layman (1922) added three species L. ncagnitestiun~, L. vane~�91 aud L. transverso-geniŸ The same author (1923) described L. fringillce and aga in (1926) L. lanicola, L. asowi, L. loossi and L. aI,zudce. Baylis (1927) transferred Distoma vitta as L. vitta. The latest comprehensive work is that of Skr~abin and Udinzew (1930) who also described a new species L. papabejani and besides hkve given a useful key for the identification of the va¡ species. Pfice (1935) deseribed a peculiar species L. ~nonenteron which shows only a single intestinal cmc~~m. He thinks that probably this condition of the intestine may be present in some other species. The wfiter considers thfs feature as very important and thinks that such forros should not be included in the genus. In fact, ir appe~rs necessary to create a separate genus for the forro described by

Price.

Diagnosis of the genus Lyperosomunr Looss, 1899.--

DŸ elongated ribbon-sha2ed; spherical in section. Testes tandem behind the verttral sucker. Vitellaria beh/nd the lestes. Other characters as of the family.

From India two species have so far been described under the genus, one by Bhalerao (1926) and the other by Patwar�91 (!935).

B8


Lyperosomum kakea Bhalerao, 1926

Body elongated, t aper ing a t bo th ends ; euticle w i t h o u t spines. Lengt, h 3 .36 mm. and b read th (max in lum a t the level o[ ventra~ sucker) -33 m m . Oral sucker te rmina l , �9 13 mm. X -15 m m. Ventra l sucker, s i tua ted a t l [ 5 th of the d i s t an te f rom the an te r io r

end, measlu'es .2 mm. in diameter . P r e p h a r y n x abs~nt. P h a r y n x , . globular , measur - ing -06 mm.* in diameter . I n t e s t i n a l cmca up to the pos ter ior end o[ body . Testes , oval, one behind the other some d[stanc~ behind ven t ra l , sucker plaeed c~ntra l ly wi th the i r axes oblique and measure . 1 5 - - 1 8 m m . x , 1 2 - . 1 2 5 m m . Geni ta l pore some dis tance behind p h a r y n x . Cirrus sac pear-shaped, lies cent ra l ly , a b e u t .28 m m . anter ior to vent ra l sucker ancl conta ins a coHed vesicula seminalis . Para pros ta t ica , ductus ejaeulatorius and cirrus small. O v a r y ovoid, b sh ind testes, in cent ra l line and nleasures -165 mm. x 12 m m. Recep t aeu lum seminis i m m e d i a t e l y behind i~. L~urer 's canal present . Shell-gland, post-ovari,~l. U~erine coils fill up poster ior pa r t of b o d y behind the ovary. Vitel laria small, ex t end ing a l i t t le beh ind ova ry . Eggs, oval, operculate measur ing -028- .03 mm. • . 0 t 6 - - 0 1 8 m m .

HosL~Corvus insolens (Liver).

LocaIity.--lZangoon .

Lyperosomum colo?osum Patwardhan, 1935

Body elongatecl, tapering at both ends and measures 1.7-3.0mm. in length. IVlaximum breadth .192-.34 mm. at the level o~ acetabulum. Integument smooth ; parenchyma with scattered pigment particles. Oral sucker, subterminal, elliptical, �9 076--13~i mm. x �9 092-.16 nlm. Pharynx muscular and spherical, �9 063-.076 mm. in d~ameter. Oesophagus .06--Imm. long. Intestinal c~ca extend upto 4[5th of the worm posteriorly. Acetabulum -138-.24 mm. i~ diameter, and is .356--63 mm. behind the anterior encl of the body. Testes, two, post-acetabular, one behind the other. Anterior testis sphezical .127-.22 mm. in cliamet~r. Posterior testis, oval, separa ted f rom anter ior by a ut~rine loop a~d measures . 108- .208 mm. , -123- �9 2 1 7 m m . Genital poro m i d w a y b3tween ac~ tabu lum a n d an te r io r end of body. Cirrus pouch elongated, py r i fo rm -12-.21 m m . • . 04 - -07 m m . I r con ta ins large vesicula seminalis. Ovary spherical, small~r t h a n restes, .0~t m m . behind poster ior testis, in th3 m e t a n line. I t measures .08~- .1~8 m m . in dŸ R e c e p t a c u l u m seminis dorsal ancl la tera l to o v a r y and measu re s - 0 2 3 - - . 0 4 mm. in d iameter . Shell-gland is s imilar in pos i t ion bu t s l ight ly ven t ra l . Vitellaria, post- ovarial , in two la teral rows of 6-7 follicles on each side. U te rus ex t ends backwards as l a r ~~ pos t~ ¡ end of body. ~Eggs large, ell iptical, th ick-she l led and measure �9 0 1 2 5 - - .022 mm. x . 0 2 5 - - .04 mm.

Host.~Temenur p~zgodamm.

Locality.--Nagpur.

Key for the identification of the species of Lyperosomum"

Ovary larger than both testes~L, kakea.

Ovary smaller than both testes--L, colorosum.

* I n the original paper ir is men t ioned aa . 6 mm, wh ich is p r o b a b l y a mispr in t .

Studies in Helmin/]zo/ogy 137

Genus Pl~~ynosomum Looss, 1907

The genus .PIety~osomum was created by Looss (1907) for a trematode, obtained from Circatus gallicus, which he named P. $emifuscum. Later on several authors added to this genus. Kossack (19!0) described 2o. fastesum, lXTicoll (!915) reeorded _P. acuminatum from Kestrel and also transferred two species of Dicroccelium as P. deflectens and P. fletiola~um. Travassos ( !916)added _P. microchis and later again (1918) P. arietis. Tubangui (i 928) reported P. phil~iŸ237 The latest work on the genus i s that of Cameron (1928) where he deseribed P. planic@ites from a tiger cat.

.Diagnosis of [.he ge#us Platy~wsomum Looss, 1907.--

Dicrocceliidr lancet-shaped, greatest width at the level of the testes. Anterior end more pointed than posterior. Suckers almost of equal size. Testes symmetrical, at equal height behind ventral sucker. Cirrus sac plump, sac-shaped. Ovary lies behind the restes. Other characters as of the family.

The only forro recorded from India is P. acuminatum by Bhalerao (1926) which was previonsly obtained by Nicolt (1915) from Kestrel in Scotland.

.Pla.ty~osomum acumina~um Nicoll, 1915

Hos~.--Corvus insolens (Liver).

Locat#y.--Rangoon.

Remarks.--Bhalerao gives no description or figure but mentions that the only variations from lficoll's desc¡ that he finds in his specimens, ate in the climensions of the body and the posterior ex~ension of vitellaria. Since the writer has no specimens, he cannot adcl any observations on the subject. A summary of ~icoll's deseription is, however, given below :--

Lengt,h 6.3 mm., rn~x~mum brea, dth just behind ven~r~l sucker is 1 -O mm. Oral s u c k e r tw i s t ed to righ~, -45 m m . x -40 m m . VentrM sucker is -6 m m . x .75 m m . , si~uated 1.97 mm. from the ~nterior end. Ph~rnyx co~tiguous w•th oral sucker and naeasures -i7mm. x-15mm. Ossophagus short. Intestinal diver~icul~ long and ~xarrow. Geni ta l pore over the p h a r y n x , tw i s t ed to r ight . Clrrus pouch -7 m m . • �9 16 m m . cont~ins vesicul~ seminalis , p~~rs p ros ta t i c~ and duc tus e jaculator ius . TesSes

s y m m e t r i c M , i m m e d i a t e l y behind ven t r a l sucker and measu re - 2 5 m m . x - $ 6 m m . O v a r y beh i nd lef t tes t is , oval. Vitel lar i~ la teral , b~ tweea level of restes , and 2-3 m m . f r o m pos te r io r e n d ; follicles small . U~erus flHs up grea~er por~ioa of pos t -~cetabul~r s p ~ c e ; convo]u t ions en t i re ly confined beh ind ven t ra l sucker . Vagin~ weak ly deve loped . Eggs numerous , measure . 033- .039 m m . • -018- .02 m m .


Genus Multivitellaria Phadke and Gulati, 1930

Body thick and opactue , ventrally convex. Pharynx present. Ovary post-testicnlar and median above the fork of the excretory bladder. Tess lateral and lying under the cmca. Twenty-eight pairs of vitellaiia lying exterior to the cmca. Folticles of the vitel]ine glands tubular. Genital pore sueker-like close to ancl in front of the acetabulum and tying behind the gut-fork. I~xcretory bladder u

Type-species--Muttivitellaria hewletti.

M~dtivitellc~ria hewletti Phadke and Gulati, 1930

]3ody thick and op~que ancl venSrally convex. Length 6 .0 -16 .6 mm., b r ead th 4 - 0 - 8 . 7 m m . Oral sucker smMIer ancl less promin~nt t,h~n ac3tabulum. Pharyn_x present. Oesophagus -5mm. long; intest inal creta, s t ra ight of sIightly curved in a zigzag m~nner. Testns, two, lateral, pre-ovarial, lying uncler the cmca. Longi- tudinM secbions of the fluke showed the presente of smninal vesicle ancl pa,rs prosb~tic~ in cloae p roŸ of the cirrus sac. Genibal pote, suck~r-ltke, close to and in f ront of acetabulum, and lying b~hincl the iut~stimal fork. Ovary, situ~tecl meclially ~djacent to shell-gland, rec~pt~culum semtnis, and L.~urer's c~nal. Vitell~ri~, 28 pMrs, lying exterior to cmca. Excre tory bladder Y-sh,~ped. Ub~rus with loops ; eggs ~s-ss~ by 3~-53t~.

Host.--Corvus sŸ (Gall bladder).

Locaiity.--Bonlbay.

Remarks.--This forro shows the usual eharacters of the family Dicro- cceliid~e but differs from all the genera in possesshlg a s and fleshy body, much greater extension of the vitellaria, and a sucker-Iike genital pote. The writer, therefore, fully agrees with the authors in regarding ttfis forro as a distinct genus. Phadke and Gulati have also suggested a new subfamily Multivitellarime for the reeeptiou of t•is forro.

Key for the identifieation of AvŸ genera of the family DicrocMiid~e reported from India :

I. Testes tandem .. Ly~erosomum.

Testes connubial .. 2

2. Vitellaria extencl throughout the body length--genital sucker

present . . . . M~ltivitdlari~.

Vitelta¡ confined to the middle of the body--geni ta l sucker absemt . . . . . . Platynosomum.

S/udies in f-fefmint/wfoyy 139

Family OŸ Braun, 1901

The falnily Opisthorchiid~e contains about adozen genera and attempts w e r e made to divide the family into subfamilies but i~iorgan (1927) has pointecl out the ulldesirability of this course of action owing to absence of ally constant differences. Further, IViorgan (1927) has suggested the syno- n y n l y of 7Vo~~ulr to Opisthorchis and the writer agrees with him. In fact severa l other genera, viz., Amphimerus and Cyclorchis which ate recognised by 1Viorgan as distinct from Opisthorchis also do not show those distinctive f ea tu re s which shonld be taken as characters of generic importance. 1Viorgan (1927) further suggests the fusion of the genera Clonorchis and Opisthorchis otl t he basis of the branching of the testes but the w¡ thinks that the difference in the excretory bladder in the two cases is rather mnch pro- nounced. Ir would, therefore, be desirable to keep the two genera as d i s t inc t . Recently u (1933) described a genus Oesophagicola f r o m the oesophagus of a marine sllake and created a new snbfamily Oeso- phagicolinm for its reception, under the family Opisthorchiidse. The genus, in qnestion, no dotlbt shows important differences from the members of the family Opisthorchiid~e with which ir resembles in the absence of a cirrus sac and the relative position of the gonads. Ir is, however, not possible, for the writer at present to make any comments on the systematic position

of this genns.

Diagnosis of the .family Opisthorchiidce Braun, 1901; emended 1VLorga~,

1927.--

Body flat and transparent, sometimes much thickened, narrowing aliteriorly. Suckers near each other (except in Microtmm~), often only moderately developed of atrophied. Pharynx present and usually followed by a short oBsophagus. Intestinal cmca long and unbranched, not always reaching the posterior end. Excretory vesicle u with proportion- ately long stem and short branches, opening at the posterior end or on the ventral surface in the region of the testes. In the former case the stem winds S-shaped between the testes or may lie dorsal to the testes. The genital opening is immediately in frollt of the Ventral sucker. A cirrus sac is absent. The testes lie near the hind end of the body, obliquely of directly behind one another ; sometimes they lŸ laterally. They are simple, lobed of dendritic. The ovary lies immediately in front of the testes and may be simple, lobed of multilobed. Laurer's canal present ; receptaculum seminis usually strongly developed. Vitellada moderately well developed, divided q acini of continuous and lŸ between the cseca and the margins of the body. Uterine folds in front of the restes and mostly in front of the ovary,

140 Mal~und Behari La l

extending to or in front of the ventral sucker. Eggs numerous, small and light-brown in colour. In the gallbladder and bile ducts of mamnlals, birds, reptiles and fish.

The family is represented among birds in India by a singl› gentls

OŸ Gentts Opisthorchis Blanchard, 1898

A very large number of speeies have been described under the genus and their diagnostic characters ate based on the nature of the euticle, ratio of oral to ventral sueker, disposition of the gonads, nature of the excretory bladder, the position of the ventral sueker, etc. In fact, there ate sorae species which do not show any clear differentiation and will probably be proved as synonymous. ]3ut in the present state of our knowledge and also in the absenee of data on the life-history, ir is not possible yet to suggest any ehange in their nomenelature, although Pfice (1932), on morphological grounds alone, has emphasised the identity oŸ several species.

Morgan (!927) deseribed O. dendriticus from the liver of a Sarus crane, Antigone imported into the London Zoologieal Gardens from India. The writer has been able to eolleet two species of the genus from the avian hosts in India ; and although he is fully aware of the undesirabili ty of multiplication of species he finds ir neeessary to do so as the speeimens colleeted present eertain important differences from the existing species of the genus. They ate, therefore, being described here as new species. The diagnosis of the genus OŸ given by 1Viorgan (1927)is also slightly emended.

Diagnosis of the genus O~isthorchis emended.--

Opisthorchiid~e; body distinetly, often very greatly elongated, sometimes pear-shaped, anterior end attenuated; posterior end broader. Skin generally smooth. ]~xcretory vesicle usually •-shaped with long sigmoid stem winding between the testes of passing dorsal to them. Copulatory organs absent. Testes in the posterior portion of the body, and placed either obliquely of directly behind one another. Ovary simple of Iobed. Laurer's canal present. Receptaculum seminis promŸ I7te¡ coils extend from the ovary to the ventral sucker, sometimes slightly overlap the intestinal cseca. Vitellaria moderately developed, lateral of the intestinal cmca and not extending anterior of the ventral sucker, ending poste¡ at the level of the ovary. Vitella¡ do not always forro one region.

Type species--O2bisthorchis felineus.

Studies i~~ t-telrzintho[o gy 141

Opisthorchis gliddhis n. sp. (Figs. 1 and 2)

About a dozen specimens of this t r ema to �91 were obtMned from the liver of Sarcogyps caIv,us shot by the wri ter irt Badshahbagh, Lucknow, The parasites appeared pink in fresh eondi t ion and the i r anter ior end was very contraet i le and movable.

The body is more of less pear-shaped, and measures 3,9 mm. • 1.5 mm. The oral sucker is ven t ro- te rmina l and measures �9 15 mm. • -2 mm. The

ventral sucker lies at a distanee of 1.275 mm. f rom the anter ior end and measures . 175 mm. x .2 mm. The mou th leads into a short prepharynx �9 05 mm. in length. The pha rynx Ÿ ell iptical and measures .15 mm. x �9 l l mm. The cesophagus is th ick-wal led and measures -3 mm. in l en~h . The intes t inal c~eca extend almost up to the posterior end of the body.

w'6

I

Fzo. 2

l ~m. 1

Fin. 1,--OpisthorcMs giddhis n. sp. , e n t i r e w o r m - - d o r s a l v i e w ,

Fin, 2,--OpisthorcMs giddhis--eggs.

The excretory bladder is Y-shaped and opens by the excretory pore at the postero-dorsal end of the body. The med ian s tem of t he ' Y ' passes in between the testes and bifurcates into the l~tera! horns between the

anterior testis and the r ecep tacu lum seminis.


The testes ate ovoid bodies with notched margin and lŸ obliquely one

behind the other at the posterior end of th• body. The anterior testis

which lies on the left side measures .4 mm. • �9 15 mm. The posterior testis

measures -475 mm. • �9 16 mm. The vasa deferentia unite to forro a coiled

vesicula seminalis in the region of the metraterm. The vesicula seminalis

runs along the right margin of the ventral sucker and opens immediately

in front of it at the genital pote.

The ovary is ah elongated, flattened, transversely placed body measuring

�9 42 mm. • .I mm. A well-developed receptaculum seminis lies behind

the ovary and measures .2 mm. • -15 mm. The duct from the recepta-

culum seminis forros a loop near the left end of the ovary before entering

the 5otype. A small oviduct leads from the ovary to open into the 8otype

which lies in front of the ovary and is surrounded by sma11 unicellular shell-

glands.

The vitellaria consist of small follicles which s t re tch extra-cmcally in the region of the uterus. The vitel l ine zone on the left side is broken up into groups while on the r ight side i t is more oz less cont inuous except for a slight anterior part. The transverse vitelline ducts run in front of the

ovary and meet to forra a very small yolk reservoir from which a small

duct opens into the 5otype.

The uterus arises from the left of the 5otyloe and runs forward in peculiar

transverse loops. It ends in ah elongated metraterm and opens in front of

the ventral sucker. The eggs are numerous but small and measure �9 0225mm. X -O!mm.

Remarks.--This forro di~ffers from all the known species of the genus

023isthorchis in having a pecul iar ly thick-walled oesophagus. The nearest al ly of this forro is O. o3sequens from which ir differs in possessing dist inct prepharynx, very feebly Iobed restes, nature and extent of vi te l lar ia and much smaller eggs. I t is, therefore, regarded as a new species.

OpistT, orchis cheelis n. sp. (Figs. 3 & 4)

A large number of these trematodes were collected by Dr. G. S. Thapar

from the liver of the common kite, Milvus migrans. The body of the animal

is elongated and leaf-like and measures 6-8 mm. x 1.5 mm. The cuticle

is thin and sparingly spinose, specially in the anterior hall of the body.

The oral sucker is terminal and measures �9 2 mm. in diameter. The

ventral sucker which lies at a distance of !. 65 mm. from the anterior end is �9 2mm. x .I3mm. in size.

S/~tdies in Helmi~~fho�91 143

iLgl.

,0,5 mm.

FIG. 4

FIG.

FIG.

FIG. 3

S.--Op~s~horet�91191 el�91 n. sp., entire worm,~veg t r a l view. no t shown.

4.--OpŸ d~eeb�91

]3ody spines

The mouth leads into an extremely small prepharynx which is followed by a globular pharynx .095 mm. • .085 mm. in size. The oBsophagus is thin-walled and measures �9 3 mm. The two intestinal cmca extend upto t¡ posterior end of the body.

The excretory bladder is u The median stem is not S~shaped but slightly sinuous and runs in between the two restes. I t divides into the two horns just behind the receptaculum seminis. The excretory pote is dorsal and terminal.


The two restes ate obliquely placed behind each other. The anterior

testis which is situated slightly towards the left is fo~r-lobed and measures

�9 48 mm. • �9 42 mm. The posterior testis Ÿ ovoid and measures .4 mm. x

�9 3 mm. The rasa deferentia unite to form the vesŸ seminalis which is

very slightly coiled and runs along the right border of the ventral sucker

to open at the genital pote which lies immediately in front of it.

The ovary Ÿ somewhat bilobed and median in position. It measures

.4mm. • .2?5mm. The recepiaculum seminis is ah elongated body,

slightly consiricted in the middle and lies immediately behind the ovary.

It measures .3 mm. • �9 1 mm. A small duct leads from the ovary into the

8otype which is situated a little to the left of the ovary and is surrounded

by unicellular shell-glands.

The vitellaria are more of less continuous follicles, extra-c~ecal, and lie in the middle region of the body. The vitelline zone on the right side is slightly smaller t han the left. The vitelline ducts from the two sides run backwards in the forro of a ' V ' and meet to forro a very small yolk reservoir from which a narrow duct leads into the 6otype.

The uterus arises from the right side of the 5otype and runs forward in inter-emcal transverse toops and opens at the genital pote immediately in front of the ventral sucker. I t contains a large number of eggs which measure .025 mm. • .01 mm.

Re~narhs.--This speeies differs from all the existing species of the genus except O. viverrini, O. piscicola, O. ca~~inus, O. noverce and O. peclicellata in having its cuticle armed with minute and sparingly arranged spines.

From O. viverrini ir differs, however, in having the oral sucker slightly larger than the ventral, much longer cesophagus, slightly lobed ovary, disproportionately developed vitellaria and smaller eggs.

From O. Piscicola it differs in having the ventral sucker smaller than the oral, oblique position of testes, disproportionate vitel la¡ confined to almost middle th i rd of the body, and in having larger eggs.

From O. caninus it ditiers in having a non-pedieled ventral sucker si tuated at double the greater distance from anterior end, a very small prepharynx, mueh longer cesophagus, extension of intestinal cmca up to the posterior end, more or less straight (not coiled) vesŸ seminalis, very slight lobation of ovary, elongated and post-ova¡ reeeptaculum seminis and the non-pedicled genitat pote.

From O. noverce it ditters in having a very long oesophagus, extension of intestinal cmca to the posterior end, anterior testis lobed and posterior

gtudies iJz t-lell~int~olog), 145

with a]most entire margin, ovary ahead of the horns of the excretory bladder, much smaller range of vitellaria and much smaller eggs.

From O. fSedicellata ir differs in having smaller ventral sucker, non- pedicled gelHtal pote, absence of looped terminal ends of the male and female ducts, extra-cacal vitellaria, elongated receptaculum seminis and smaller eggs.

I4iey for the identi¡ of the avŸ species of the genus O~islhorchis reported from Indian hosts :

i. Vitellaria in middle of body, restes slightly lobed .. 2

Vitellaria confined to posterior hall of the body, testes dendritic O. dendriticus.

2. Oesophagus thickwalled, euticle non~spiny . . O. giddhis.

Oesophagus thinwalled, cuticle spiny . . O. cheelis.

F a ~ i l y LeciZhodendriid~. Odhner, 1910

Owing to the close af¡ between Lecithodendriina and Pleuro- genetin~e, Odhner (1910) created the family Lecithodendriida for their recepiion. This has not been questioned, but some confusion has been created abo•t the grouping of genera into subfamilies. It may be pointed out here that this is not based on any de¡ grounds and apparently dif~erent characters have been considered in this connection ; e.g., the length of the intestinal cmca, the extent of the vitellaria and the position of the testes as well as the position of the genital pore ancl the host-relationship.

Mehra (1935) attached undue inlportance to trivial characters and thus added confusion by ft~rther dividing the family Lecithodendriida into six snbfamilies, viz., 7knchitremina N. Stlbfam. Lecithodendriina, P1enro- genetin~e, Phaneropsolina N. Stlbfam. ; ]~xotidendriina i~. Snbfam., and ~Eumegacetina N. Snbfam. On a review of bis work ir appears that too much stress has been laid on the posterior extension of intestinal caca and position of cirrtts sac or on the median of lateral position of the genital pote.

The present writer considers that all these features show variations in the family and shonld not forro the basis of subfamily divisions. The only character which appears to be constant is the cirrns sac. It is either present of absent. The presence or absence of r sac should, as has already been discussed (Lal, 1937), forro a character of snbfamily importance.


I t is, therefore, eonsidered desirable to keep only two subfamilies, vi::., Lecithodendriinae with forros in which the cirrus sae is absent and I~ genetinee eontaining forma which possess a cirrus sac. The various subfamilies of 5iehra can easily be accommodated under these two heads. 2r emphasis has been laid on keeping the genus Ganeo in the sttbfamiIy Pleurogenetinae although ir laeks a cirrus sae. The chief a ,gument advanced is the position of the genital pote which is sinistral in Ganeo as in other Plenrogenetinm while it is median in Lecithodendriinae. The writer considers that the shifting of the genital pore from median to margilml posit ion in about the same Ievel is not a charaeter of stlbfamily importanee, a t any late, eertainly not of a greater signifieance than the presente of absenee of a cirrus sae. Ir would, therefore, be desirable to transfer the gemls Ganeo to the sttbfamily Lecithodendriinm as suggested earlier by Travassos (1930).

Diagnosis of ~t~e family Le~ithode,~~drii&~ Odhner, 1910.--

Digenea, with variable shape of body. Body spines either present or absent. Ventral sucker at about the middle of the body. Pharynx , oeso- phagtls and illtestinal cacen of variable length. Exeretory bladder V - o f Y-shaped. Testes generally symmetrical, sometimes, elose together, at variable leve1 in the body. Cirrus sae present of absent. Ovary, dorsal, mostly dextral, at varying levels of the body. Reeeptaeulum seminis smatl. Laurer's canal present. Vitellaria sitttated on either side, branehed follieles, dendritic mostly in front of the middle body. Uterus well-developed, with regular ioops, mostly pressed to the hinder end of the body. Geni tal pote in the anterior part of the body, median of left-sided and sometim› dorsal. Eggs nnmerot~s, sma[1. Parasitic in intestine of ~amlnals , 13irds, ReIotiles and Amphibia.

The family is divided into two svtbfamilies : - -

1. Leeithode~ldriinae--without a cirrus sae.

9,. Plem'ogenetinae--with a cirrus sac.

The onty avian species tmder this family reported so lar f rom Ind ia comes under the st, bfamily Plettrogenetina~ as it possesses a cirrus sae.

Genus Parabascus Looss, 1907

(Syn. Pleur@solus Mehra, 1935)

DŸ small, abottt 1 mm. long with blunt ends. Skin strongly and thiekly spinose, Oral sucker sttbterminal, globular. Aeeta- bulum almost pre-equatorial, about twice as large as oral sucker. ~Pharynx small, oesophagns long. Cacen extend post-testicttlar. Excre tory pote

Smdies in Arebnint~o�91 147

ca~tdo-• Coputatory organ muscular. Cirrus sac targe, thick, ciub- shaped, extends to left around acetabulum. Ductus ejaculatofius, pars prostatica and prostatic cells present. Genital pore at the side of acetabulum. Testes two, 1urge, oval, post-acetabular. Ovary elongate pre-testieular, ¡ of median line. Shellgland post-acetabular. Vitellaria chiefly pre-acetabular. Uterine coils broad, ahnost entirely post-acetabular. I~ggs numerous, light-brown and operculate.

Parabascus insolens (Bhalerao, 1926)

(Syn. Phaner@solus insolens Bhalerao, 1926 ;

P[eur@solus i~solens Mehra, 1935) B o d y pear - s l l aped , cut ic le cove red wi th spines . L e n g t h . a 8 5 - . 7 3 m m . B r e a d t h

- 2 9 - -365 m m . , m a x i m u m be ing a l t h e level of t:mbes. Oral sucker .1 t5 m m . • -1 m m . V e n t r a l sucker smMler t h a n oral, ,1 m m . ilx d [amete r . P h a r y n x , g lobular , m u s c u l a r -(}3- .035 m m . in dk~meter . I n t e s t i n a l c~cca sho r t and s o m e w h a t a n t e r i o r to the

t,~stes. Tes tes , ovM w i t h ent i re marg ins , lŸ s y m m e t r i c a l l y , and measu ra 0 . 0 9 7 - 0 - 1 2 m m . * • -0~- .093 m m . Cirrus sac, z igzag of orden horse-shoe-sh,~ped. I t is do r sa l ~o v e n t r a l sucker , p a r t l y an tm ' i e r to ir, l y ing b e t w a e n ir and the in t e s t ina l fork. On i t s pos t e r io r side i t con ta ins a la rge vesŸ semin~lis ancl p~rs p r o s t a t i c a . Duc tus e] acu la to r i us sma l l followe(t by f~irly leng m u s c u l a r p~nis c~p~bl~ of p r o t r u s i o a t h r o u g h gelxital pote , s i tuatecl immec t i~ te ly ~~nterior to t h e ven t r a l sucker , on t h e lef t side of m i d d l e line. O v a r y oval , m e a s u r e s . 6 - 7 m m . x . 0 t - . 0 5 m m . , r i gh t of the v e n t r a l s u c k e r and is o f t en ove r l~pped b y t h e r ight t e s t i s . Shell-gh~nd beh ind v e n t r a l sucker in t h e cen t ra l l ine. l l e c e p t a c u l u m seminis va rŸ in size. I r lies on the inner sicle of t h e r i gh t t e s t i s . I , a u r e r ' s camal p re sen t . U t e r u s long, coil~ m o s t l y pos t - t e s t i cu l~r . E g g s , brown, ope rcu la t ed , . 0 1 8 - . 0 2 2 m m . x .097- .0105 m m . The v i t e l l a r i a are s i tua tec l ,~t the sides of of o v e r l ~ p p i n g i n t e s t i n a l cmca; follicles, Small, 8-12 on eaeh side. ]~]xcretory pore a l t he h ind enr of b o d y leads in to V-shaped b l adde r whose ~rms d i v c r g e and e n d s l igh t ly beh ind t e s t e s .

Host.~Corvus insolens (Intestine).

LocaIity.--Rangoon. Remarks.~Bhalerao (1926) descfibed this species under the genus

_Phaneropss Mehra (1935), on small size of cirrus sac and acetabular position of genital pote, suggested the creation of a new genus Pleuropso~us for it. On looking through the figure given by Bhalerao ir appears that the size of the cirrus sac is not small but owing to its bent character, it does not extend beyond the intestinal bifurcation. The position of the genital pore in the present form is pŸ and this is certainly different from the position of genital pote in Phaner@solus where it is lar forwards just behind the ph~ryllx. The present forro also resemb!es the genus Pambascus in the shape of cirrus sac, the pre-acetabular position of genital pore, in the

* I n the o r ig ina l a ccoun t ir is " O . 97 -0 .12 m m . " which ev iden t ly is ~ m i s p r i n t .


extent of vitellaria and in the position of the excretory pore. It differs, however, from Parabascus in having shorter intestinal c~ea, smaller ventral sucker and ah avŸ host-features which are not of very great importanee.

I t is t rae that the speeies shonld be removed from the genus Phaner@- sobas, bttt it shonld not be taken asa basis f£ the creation of a new genus Pleur@so~u.s as suggested by Mehra (1935). It is suggeted to place ir under the existing genns _Pambascus with which ir resembles in many important poŸ discussed before.

Family Heter@hyidce Odhner, 1911

Odhner (1914) suggested the name I-leterophyid~e to replace the older names Cotylogonimidm and Coenogonimid~e. Ciurea divided the family into five snbfamilies, w;z., I-teterophyinm, 1Vfetagoniminm, Centrocestinm, Apophallina ~ , and Cryptocotylin~e. This division has subsequently been modified by ~icoll (1923), Faust and Nishigori (!924), Poche (1925) and Witenberg (1926). The family I-leterophyidse as ir stands to-day has the following diagnosis :--

5iinnte forros, t~snally not exceeding 2 mm. in length. Anterior portion of body thinner and more movable than the posterior portion. Skin covered with small scale-like spines that are rednced posteriorly and may even disappear towards the posterior end of the body. Intestinal caca simple, usnally extending to the posterior end of the body. Genital pore in the immediate neighbonrhood of the acetabulmn, genital ducts nsttally open into a genital sinus, which may be varionsly modified and contains a cirrns-like body of gonotyl (genital sttcker). Acetabulum usnally median bnt may be displacedto the right of the median line; sometimes it is partially of completely atrophied and enclosed in the genital sinus. Cirrus sac absent ; seminal vesicle well developed, I7- of S-shaped ; the vas deferens snrrounded proximally by • mass of prostatie cells. Testes two of one, oval, globular, or slightly lobed, near the posterior end of the body, side by side, of obliqnely one in front of the other. Ovary oval, globular of slightly lobed, generally pre-testicnlar sometimes post-testicular nsnally to the right of the median line. Seminal receptacle and Laurer's canal present near the ovary, usnally near its poste¡ border. Vitellaria, mainly in the lateral fields, may extend anteriorly to of beyond the genital apertttre. Uterus usnally restricted to the intercmcal field between the ovary and genital pote, bttt may extend to posterior end of body. Adults parasitic in the intestine of birds and mammals.

Type genus--Heter@hyes Cobbold, 1866.

S~u�91 i~z tteh.i~~~/zoZo gy 149

Only one genus Ascoco~y~e of the family tIeterophyid~e has been reported from birds iil India.

Ascocotyle Looss, 1899, eme~zded Srivastava, 1935

Diag~~osis.~Siinute distomes, body thickly spinose ; oral sueker armed with a single of double crown of straŸ cylindrieal spines. Ora1 sucker continued posteriorty into a distinct appendage ; prepharynx long, pharynx well developed and muscular, msophagus present or absent, intestinal cmca long or short. Acetabulam median, situated in assoeiation with the genital sinus in a depression of the ventral body surfaee. Testes situated one on each side at the hinder end of body ; vesieula seminalis and ejaculatory duet well developed. Cilrus sac is absent. Ovary median of slightly to one side, pre-testicular ; receptaeulum seminis large, situated in leve1 with ovary of behind it. Vitellaria lateral, usually post-aeetabular sometimes extending as lar forward as the pharynx and meeting mesially near the intestinal bifurcation. Uterus usualIy post-acetabMar, rarely extending as lar forward as the pharynx ; eggs large, opereulate, measuring 0. 015-0. 035 mm. x 0-008-0.017 mm. in size. Parasitic in birds and mammals.

The only aviail speeies recorded from India is Ascoco~yle #zter~Jzedius SrŸ 1935.

~Body py r i fo rm, . 6 - - . 9 m m . • . 2 - - . 3 8 m m . in size wit, h b a c k w a r d ] y directed sp ines . Oral sucker t m ' m i n d , .04. - ,05 mm. in di~unet, er, surrountled b y double erown of cy l ina r i ca l spines ,~bou~ 28-30 in numb~r . Ven t ra l sucker feeble, - 0 6 6 - - . 0 7 7 m m . in d i a m e t e r , in midd le of b o a y , ly lng togegher w i t h the geni ta l pore in a shMlow depres - s i o n - - t h e ver~~rogen[tM sinus. Geni ta l ope~ing gual:�91 b y t, he gonotyls . E x e r e t o r y p o r e t e rminM. Oral sucker has a,n orM cmeum, which l[es dorsal to p r eph~rynx . Pre - p h ~ r y n x long. I~harynx .0:3-- .0~ m m . • -02-- -03 m m . Oesophagua shorb. I n t e s t i n a l cm, ca, termin,$te in lev› w i t h t he an te r io r ma rg in of ovary . Testes , poster ior , s y m m e t r i e a l , m eas u re . 12 - - .17 m m . • . 0 7 - - - 1 2 m m . VesŸ seminMJs, enor- m o u s , re to i~-shaped. Cirru:~ sac absent,. E j a c u l a t o r y duc t .ioins wi th u te rus fo rming a gen i t a l sinus. O v a r y be t ween righg ~estis ancl v e n t r N sucker i r regular and measures �9 0 8 - - . 1 m m . X 1 1 - - .1.I m m . l / e c e p t a c u l u m semtnis, yo lk reservoi r aud Lau re r ' s c~nM present . Vi te l lar ia ex t e~d up to p h a r y n x . U~erus ne ver ex tend ing be?-ond gen i tM sinus. Eggs, operc ª . 03 - - .035 m m . x - 0 1 5 - .017 m m .

Host.--Hgl�91 Zeucory~hus (Intestine).

LocalŸ y . ~ A liaha ba d.

Remarhs.--The genus AscocotyZe was createdby Looss in 1899. Faust (1920) described the genus BhagicoZa but subsequently (1926) considered it as a species of Ascocotyle. Meanwhile Stunkard and Haviland (192t)spli t up the genus AscocotyZe into two sub-genera AscocotyZe and P,zrascoco~yle.

150 Makund BeharŸ Lal

Witenberg (1929)recognized Pargscocogy;e of Stunkard and I-laviland and suggested that a redeseription of new material of PhagicoIa pi~heco- fihagicole could settle the question whether Pe;'a.scocotyle is synonymous with Phagicola or they both ate valid genera. Price (193£ proved the identity of Phagicola and ParascocotyZe and eonsidered tha t P]~.agicola and AscocotSe should be regarded as distinct genera, differing from each other in the arrangement of spines on the collar in one of two rows, presence of absence of oesophagus, iength of intestinal c~eca, position of vitelIaria and the extent of the uterus. Srivastava (1935) in deseribing a species Asco- co~2~e (Phagicola) intermedius has pointed out tha t this speeies connects the two genera of Price as regards the extent of vitellaria. I-Ie has, therefore, regarded Phag~coZa and Ascoco~yZe as sub-genera of Ascocotyle and based the separation of the species Ascocotyle intermedius under the sub-genus Phagi- cola on the grounds that it possesses a long oesophagus, long in�91 c~eca, and has its uterus con¡ behind genital sinus. This does not appear to be sound. In fact in his description Srivastava says (page 271), " The wide intestinal c~eca ate moderately long." In his diagram of the species a few eggs ate shown a littIe ahead of the region of the genital sinus. There appears, therefore, to be some confusion in the matter . Owing to the aŸ of oral spines the extent of vitellaria and spination of the body, Price (1986) still holds that the sub-genera Ascocotyle and Phegicola should be given a generic statns. There is, in raer, a great deal of variability in the extent of vitellaria and in the size and extent of oesophagus and intestinal c~eea throughout the genns, and in the absence of constant characters, the writer considers tha t this differentiation into genera and sub-genera be abandoned.

Family Micr@helZidce Travassos, 1920

Ward (1901) created a subfamily Mierophallinm for the reception of the two genera M` and Levinseniella and kept it close to the subfamilies BrachycaeHn~e and Pleurogenetina~. A few other isolated genera, viz., Mo~socaecum Stafford, SŸ Jagerskiold, Sfldof)halIus Jagerskiold and Maritrerna Nicoll were added to this subfamily. Odhner (19!1) transferred the sabfamily ~icrophallinm to the family I-Ieterophyidm and was supported by lgicoll (1924), Poche (!926), Fuhrmann (1928), Stunkard (1929), Faust (!929) and Sprehn (1932). Other workers, wiz., Ransom (!920), Ciurea (!924), VŸ (1924), Wigenberg (1929) and 1Viiielter and Van Cleave (1952) emphasized its exclusion from I-Ieterophyidm. Travassos (1920) while agreeing with these latter workers raised ir to the status oŸ a family Nicrophallidm, to which view the present writer fully subsc~ibes.

Studies ha Hdming;olo gy 151

The fami ly Microphallidm as it stands to-day has the following diagnosis : - -

Trematodes, wi th small oval or ell iptical of pear-shaped or conical body with b lun t apex. Oral sucker, te rminal or subterminal. Ventral sucker s i tuated in the mid-body or hind-body. Pharynx present, prepharynx present or absent. Oesophagus long; intes t inal troca, with one caecum sometimes longer t h a n the other, very short a n d never reaching behind ven t ra l sucker. Excre to ry bladder V-shaped. Genital opening either ad- }aeent of away Irom the ventra l sucker, oftell provided with a genital atr ium w h i c h may have sometimes thimble-shaped pockets on its lateral walls. Cirrus sac present or absent. Pars prostat iea lying free of covered when cirrus sac present. Vitell ine glands, in separate follieles either in groups of irregular of in the forro of a horse-shoe-shaped band. Testes behind ovary ; the two restes ate almost on the same level. Ovary near the acetabulum. Receptaculum seminis absent. Laurer 's canal present. Uterus fills up the posterior broad por t ion of the body. Eggs, small and numer-

OUS.

Type genus--MicrophaIZus.

Only one genus of the family, viz., Levinseniella has been reported from

India . Genus Levir~seniella Stiles and Hassa11, 1901

Diagnosis.--Microphallidw.; excretory bladder small and not reaching up to restes. Vitel larŸ in groups of 7-8 follicles on each side. Genital pore close to the ven t ra l sucker. Genital a t r ium shows poeket-like thickenings. There are always four th imble-shaped papillm associated with the male apparatus. The " Female papillae " are thin-walled, between the

ven t ra l sucker and the male a tdum.

Type species-- Levinseniel[a brachysoma.

Levinse~r indica Lal, 1936.*

Body roughly t r iangular , covered over with minute spines upto the level of the vent ra l sucker ; measures .98 mm. • mm. Oral sucker �9 l mm. x -12 mm. Ventral sucker -08 mm. x .078 mm. Prepharynx �9 02 mm. long. Pha r ynx -056 mm. x .055 mm. Oesophagus .22 mm. iong. Right testis �9 105 mm. • - 058 mm. Left testis �9 12 mm. X - 06 mm. Vesicula seminalis .1 mm. • .065 mm. Pars prostat ica .1~5 mm.

* u Proc. Ind. Acad. Sci., 1936, 4, No. 2, Sec. ]3, 92-98. ]B4


long. Nale genitaI opening lies at about �9 09 mm. from the ve:ltral sueker. Ovary -10,5 mm. • -06 mm. Eggs numerous with thick shell and without filaments, measure .017 mm. • .006 mm. Vitelline glands in groups of

of 6 just behind testes.

Host--Gallinago gallinula (Bursa Ÿ

Locality--Lueknow.

Remarks on the family Microphallidce.--The subfamily Microphallin~e Ward (1901) was raised to the status of a family ~Iicrophallid~e by Travassos (1920). This family shows distinctive differences from the family Hetero- phyid~e in the copulatory apparatus, the absence of a reeeptaculum seminis and the nature of the genital pote. I t has, therefore, been rightly excluded from the laŸ Iamily and kept as a separate family.

The type genus of the subfamily ~icrophallin~e is MicroŸ which is chraeterised by the absence of a cirrus sae. The genus Maritrerna has also been included under the same subfamily al though it shows a well- developed cirrus sae. The presenee of a cirrus sae alone, as indicated earlier (Lat, 1937), is suffic{ent for the exclusion of the genus Maritremc~ from this subfamily. The w¡ therefore, considers it desirable to retain the subfamily IV[ierophallinm for only those forms in which a eirrus sac is absent, and to ereate a new subfamily lVIaritremin~e for the reception of other forros which possess a cirrus sae.

The family ~ierophallidm is divided into two sub-families thus : - -

1. Subfamily ~icrophallin~e.

2. Subfamily ~ari t ieminm. N. Subfam.

Diagnosis of the subfamily Micropha~~ina~.--

Mierophallidm without a cirrus sac. VesŸ seminalis and pars prostatica lying free in the parenchyma. Vitellaria in groups of 5-7 follicles on each side behincl the restes.

Type genus--Micr@ha[~us.

Diagnosis of the subfami[y Maritremina~. N. Subfam.

lVficrophallid~e with a cirrus sac. Vesicula seminalis and pars prostatica inside the cirrus sae. Vitellaria in the forro of a band.

Type genus--Maritrema.

Shtdies bz Z-felmi~~thoZog'y 153

Family Cephalogonimidce Nico11, 1914

NicoIt (19!4) erected the family Cephalogonimid~e for the reception of the subfamily Cephalogonimin~e Looss, 1899. Pande (1932) and Bhalerao (1936) think tha t the family should be brought back to the subfamily status and should be included in Plagiorchidm. In support of this view, Bhalerao (1936) has pointed out t5a t Nicoll (1935) has himself included it in Plagi- orchidm. He has also tried to show that the position of the genital pote in some genera of Plagiorchidm, viz., Renifer, Ochetosoma and Lechriorchis is lateral to pharynx and forros a near approach to the condition met with in Cephalogonimid~e and further tha t theŸ is a similarity in the nature of the excretory bladder in the two tases. The writer, however, does not agree with Bhalerao in bis contention. The mete faet, that the original author does not mention CephMogonimidm as a distinct family a t a Iater stage, should not be enough to prove the non-eMstence of the family. The condition of the geni talpore and cirrus sae as also some other characters ate very distinctive in Cephalogonimidm and ii would be more appropriate to regard ir as a distinct family. Some of the genera of Plagi- orchidm mentioned by Bhalerao should be simply regarded as connecting links showing a possible transition from one family to the other. This does not, however, invalidate the existence of two distinct families, Cephalogoni- midm and Plagiorchid~e.

Diagnosis of the family Cephalogonimidce Nicoll, 1914.--

Skin spiny. Prepharynx and pharynx present; oesophagus variable, crura �89 to more than ~th length of the body. ]Excretory vesicle u sometimes with a caudal vesicle. The median stem of the ' Y' is shorter than the branehes. Genital pote marginM, dorsal or anterior to the oral sucker. The cirrus sac long and sinuous of saccular, extending to or beyond the intestinalbifuŸ Testes behind acetabulum and ovary. Ovary close to the acetabulmn and on the right side of the median line of the body. Receptaculura semin.is and Laurer's canal present. Uterine coils in ascending and descending rami, chiefly in posterior region behind the testes. Eggs thick-walled and numerous.

Type genus--Cephalogonimus.

The family is represented by the genus P'rosthogonimus among Mrds i11 India.

Genus Prosthogonimus Lª 1899

Diagnosis.--Body small or medium in size, flattened, with the maxi- muro breadth behind the middle. Skin spiny. Prepharynx short. Oesophagus


of medium size. Excretory bladder u Testes symmetrŸ bet~ind ovary and acetabttlmn, Ovary close to acetabulam. Cirrus sac Iong, slender ah4 sin~ous with convoluted vesicula seminatis. Genital pore marginal at the leve1 of the oral sucker. Receptaculum seminis and Laurer's canal present. Vitellaria extra-csecal. Uterus in coils in posterior part of the body, chiefly behind the restes. Eggs numerous.

Rema~ks or the ge~r162 Prosthogo~r genus Prosthogonimus was transfe~red to the family Plagiorchida by Poche (1925). Fuhrinann (!928) an4 Spreim (1932) also keep ir under Plagiorchidm. Sinha (1932) has discussed at length the clifferences between the genus Prosthogonimus and the family Plagiorchid~e and removed the geiius to the family Cephalo- gonimid~e to which view the w~iter fully subseribes.

Prosthogo~r cuneatus (Rudoll0hi, 1809) Braun, 1901

A single specimen of tiiis parasite was obtained by Dr. G. S. Thal0ar from the 5ursa fab¡ of tiie common mynah, Acridot/beres tristis.

The specimen measures 4.95 mm. • 2.375 mm. The oral sucker is te~xuinal and measures -2 mm. • -25 mm, The phaŸ measures -175 mm. X -I mm. This is followed by a lona oesophagus -3 mm. in

size which bifurcates at its posterior end into the two intestinal c~eca. The

ventral sucker is well develope4 and lies at a distance of l : 8 mm. from the

~nterior end. Ir has a circular shape and measures .6 mm. in diameter.

The restes lŸ at the same level behin4 the ovary and the ventral sucker.

They ate �9 6 mm. apart from each other and oval in outline. They nleasure

�9 75 mm. • -4 mm. The cirrus sac is a slender an4 sinuous structure,

containing coiled vesicula seminalis. It opens at the lateral margin of the

body by the side of the oral s~cker.

The ovary is greatly lobecl ancl lies in beiween the right testis and the

ventral sucker. Ir measures -6 mm. • .4 mm. The receptaculum seminis

is a more or less sloherical body lying j~tst behincl the ovary on the inner

borcler of the restes. The 5otype oceupies a small atea just behind the

ventral sucker.

The vŸ are con¡ almost to the middle third of the body.

They are extra-cmcal, ancl consist of a large number of distinct small ancl

zounded follicles.

The uterus ¡ up the entire sl0ace behind the restes. Ir has both

ascending and descending convolutions which pass in between the fwo

Sl2,ldies in He{minthology 155

restes. I n f ront of the ventra l sucker the u•erus runs more Gr less s t ra ight to fog~ the met ra te rm whieh runs along the cirrus sac and opens a t the genital pote. The eggs ate numerous but small and measure .02 mm. •

�9 O1 mm. Fa~nily Echilr Looss, 1902

The family Echinostomida was erected by Looss (!902) for the reception of bis subfamily EchinostominŸ and some other genera. Dietz (!909) described a large number of fornls from avian hosts. Odhner (1910) attempted to classify the family into three subfamilies, ]~chinostomin~e, Himasthlin~ and Echinochasmina. A large number of genera have been describ~d under the family which are not included in any of the existing subfamilies. Recently the writer (Lal, !936) erected a new subfamily Parorchin~ foŸ the reception of the genus Pazorch{s. The family Echino- stomida has the following diagnosis :--

Body more Gr less elongated, small Gr very large, usually much flattened anteriorly, less so, Gr even cylindrical posteriorly. Oral sueker small and weak, surrounded dorsally and lateral ly, bn t not ventral ly, by a collar-like fold, bearing one Gr two rows of spines, which are cont inued la tera l ly to ventral corners, the corner spines sometimes large Gr speeialized ; acetabu- tum large and powerful, usual ly pre-equatorial and sometimes near oral sucker. Cuticle usual ly spinose, specially anteriorly. ]Exeretory vesicle u wi th la teral twig-like branches. Pharynx and oesophagus present; intestinal caca extend to posterior end of body. Genital ape~ture pre- acetabular ; genital sinus present Of absent. Cirrus pouch nsually present. Testes post-equatorial, "Ÿ Gr connubial in ppsition. Ovary pre-testi- ctdar, usnally to right of median line; Laurer's canal present. Recepta- culum seminis present Gr absent. Vitellaria lateral, rarely extending anterior to acetabulum ; sometimes confined to middle and consist of distinct and big follicles. Uterus in Ÿ coils, rarely extending beyond intercmcal ¡ Parasites of intestines Gr bile ducts of vertebrates.

The family is divided into four subfamilies :--

I. Echinostominse Looss, !899--Type subfamily.

2. IIimasthlina Odhner, !910.

3. Echinochasmina Odhner, 1910.

~. Parorchina Lal, 19�91

Subfamily Echi~ostomirt~ Looss, 1899

Diag~osis.--]F, chinostomida, eirrus sac generally extends up to the centre of the ventral sucker aild is never pushed behind. Cirrus long,


generally not covered with spines and when contracted ir appears to be cofled. SemŸ vesicle cofled and undivided. Collar spines hi one o1" two rows, confinuous on the dorsal side.

Type genus--EchiHostonz~.

Echinostoma revolu~um Looss, 1899

H ost--A Has Ÿ (Intestine).

Localitv.~Rangoon. Re~narks.--Gogate (1934) obtained this spedes from the duck, A~,as

pcecilorhyncha but gives no description or diagram. He mentions tha t the only feature in which hŸ forro differs from the descriptioll of Lª (1909) is the size. The biggest of them is only a little longer than the smallest mentioned hi the desc¡ The writer has no specimens of this species in his possession and cannot add to the description. A summary of Lfihe's des.cription is, however, given below : - -

" Bocly 1 0 - 2 1 m m . long. M o u t h sucke r -25- - .50 m m . P h a r y n x .21 - - . 3 5 m m . Ven t r a l sucker .67-1 .69 mm. Cut ic le sp iny . Col lar sp ines "~7. T h e r e are 27 ' R andsfiacheln ' and 5 ' E c k s t a c h e l n ' on e i t he r side. S o m e t i m e s sp incs 30 i n n u m b e r . Testes of ve ry var iab le shape, in p o s t - e q u a t o r i a l body . V i t e ] l a r i a c x t e n d u p to t h e level of t h e ace t abu lum f rom the p o s t e r i o r end . They do n o t m e e t in t h e m i d d ] e i n t h e region of u te rus or tes tes . Eggs . 0 9 7 - - .126 m m . • - 0 5 9 - - . 0 7 1 m m . "

Echinostomum govindum Moghe, !932

B o d y 4 . 6 - ~ . 9 4 m m . • .92 m m . Cut ic le w i t h sp ines n u m e r o u s ,~n te r io r ly b u t absen t in the pos t - t e s t i cu la r region. Ora l sucke r -1~9 m m . x -133 m m . , s u r r o u n d e d b y ren i fo rm collar of 32 spines. T h e r e a t e 4 sp ines on e~ch siete p l aced c]ose t o g e t h e r . Shor t prephar~nax. Muscular p h a r y n x -22 m m . • -36 m m . fo l lowed b y l ong oeso- p h a ~ . Intesti~xat cmca run u p t o "14 m m . f rom pos t e r i o r end . Ventral1 s u c k e r s u b - globul~r l ies a b o u t .96 mm. f rom an te I ' i o r e l ld a n d m e a s u r e s -6 m m . in d i ,~meter . Testes , ovoid , pos t -ovar ia l , .4 mm. • -24 m m . in size, lŸ one b e h i n d t h e o the r . C i r rus sac ovoid , l ies be tween i n t e s t i n a l b i f u r c a t i o n a n d a n t e r i o r b o r d e r of v e n t r a l s u c k e r , i5 con ta ins t r an sve r se ly e longa ted vesŸ semina l i s s u r r o u n d e d b y p r o s t a t i c g h m d s . Geni ta l pote bek ind the i n t e s t i n a l b i fu rc~ t ion . O v a r y pre- tes t icuL~r , . 12 m m . in di~- meter . Shel l -glaud lies be tween o v a r y ancl a n t e r i o r t e s t i s . U t e r u s runs up 6o v e n t r a l sucker . Vag ina runs dorsa l to v e a t r a l sucke r a~d c i r rus sac. Vitel larh~, r o u a d e d follicles, e x t e n d f rom pos te r ior end of i n t e s t i n a l cmca to a n t e r i o r l eve l of v e u t r a l s u c k c r . Eggs -047 m m . • -023 mm.

Host.~PhiIomachus 23ugn~x (Rectal c~eca).

Locality.--Nagpur.

Echi~r chasma n. sp.

Two immature individuals of this species were obtained from the smal 1 intesti=e of the garganey, Querquedul~ circia at Amausi. The most remarkable feature of this form is the excretory bladder which is great ly

Sfztd/es in HeAni~2t/m�91 157

sinuous and showed peculiar periodical expansions and contractions of its

stem.

The body which has spiny cuticle is elongated with a broad posterior end and measures 2.58 mm. • �9 64 mm. The head with collar of spines is marked off clearly from the rest of the body. The collar spines ate 34 in number and ate arranged in one regular row without break on the dorsal

side. The dorsal spines measure .075 mm. and the lateral and marginals �9 05 m m . in l eng th .

a i

�9 2 m m .

laxo. 6

~I~. 5. P ie . 6.

lVlG. 5 Echinostoma chasm(, n.sp., en[~re worm~vent r~ l view. Bo&y spines no~ shown. Eehinostoma chasrna, head collar showlzLg spin~s. (Sketched from glycerine

mount).

T h e ora l sucker is .15 m m . in d i ame te r . T h e r e is a smal l p r e p h a r y n x ,

a n d a m u s c u l a r p h a r y n x m e a s u r i n g .175 m m . • 125 m m . T h e oesophagus

is .25 m m . long a n d b i fu rca te s a t i ts pos t e r io r e n d i n to t w o i n t e s t i n a l c~eca

158 Makund Behar i L a l

which are wide and llave great ly crenated margins and run close b y the side of the acetabuhlm up to the posterior end of the animal.

The ventral sucker is .45 mm. • .41 mm. and lies at a dis tance of �9 65 mm. from the anterior end.

The restes ate ovoid bodies, the anterior measures �9 13 mm. • . 15 mm. and the posterior .11 mm. • .15 mm. The cirrus sac, which par t i a l ly overlaps the ventral sucker on the left side, eontains an incompletely bilobed vesŸ seminalis. The cirrus is fa i r ly long and opens at the geni ta l pore just in front of the acetabulum and behind the in tes t inal bifurcation.

The ovary is oval and pretesticular and measures .85 mm. • �9 55 mm. The shell-gland, 6otype, etc., could not be observed. The uterus runs in more or less sinuous but straight course dorsal to ven t ra l sucker in the median line and opens at the genital pore. No eggs are present in the uterus.

The vitellaria ate not ful ly developed and consist of small follicles of diffuse nature, extending from the posterior end up to the level of the middle of the ventral sucker.

Remarks.--The forro described above comes under the genus Echino- storna owing to its having a collar of spines tmin te r rup ted on the dorsal side, cirrus sac anterior to the ventral sucker a n d a non-spiny cirrus. But the forro is peculiar in possessing an incomplete ly bilobed seminal vesicle, a feature in which ir differs from all the exis t ing species of the genus and resembles the subfamily I~chinochasmin~e. Although greater details of anatomy are not known owing to immature state of the specimens, the writer feels inclined to regard ir a s a new species of the genus Echinostoma showing a transit ional stage in the lobation of the vesŸ seminalis.

SubfamiIy Himasthtin~e Odhner, 1910

No representative of this subfamily has been reported so avŸ hosts in India.

far from

Subfamily Echinochasminr Odhner, 1910

Echinostomi&e.--Cirrus sae is pear-shaped and does not reach even up to the centre of the ventral sucker. Seminal vesicle is not coiled and is sharply divided into two parts. Pars pros ta t ica present. Ductus ejaculatorius is extremely short. Collar spines genera l ly in one row but some show arrangement in a double row. They are discont inuous on the dorsal side and 20-26 in number.

Type genus--Echinochasmus.

Studies in He/mi~'~tho�91 159

Echinochasmus megavite~~us n. sp.

A large number of these parasites were obtained from the intesthie of the paddy bird, Ardeola grayi. The worms exhibited slow movements of their body when liberated ili llormal salt golution.

t.gL

Fin. 7

~IG. 7. EcMnochasmus megavi~elIu,~ n. sp., enttire worm--ven~r~t view. :Bo~ly spines not show~.

The body is elongated oval and measures 1.05 mm. • .45 mm. The euticle bears spŸ The anterior end has a prominent head which bears a collar of spines. The collar spines nurabering 24, ate arranged in one single row, Ieaving a l i t t le space on the dorsal side. They a te arranged in two groups of 12 spines each and ate of nn i form size measur ing .025 mm. long.

160 M a k u n d Behar i La l

|

c ' i m m . "

Fin. 8

Fin. 8. Echinochasmus megav~tell~s n. sp.--Head collar showing spines. (Sketehe4 ~rom glycerine moun~.)

The oral sucker is sub-ventra l and measures .06 m m • .05 mm. The mouth leads into a thin, prepharynx .075 mm. in size. This is fol lowed by a muscular pharynx measuring -09 mm. • -1 mm. The oesophagus is shor t and bifurcates at its posterior end into the two in tes t inal c8eca which run up

to the posterior end of the restes. Their t e rmina l ends ate curved inwards

and come near each other but do not meet.

The ventral sucker is strong and powerful and measures �9 18 mm, •

�9 17 mm. I r lies a t a d is tan te of -38 mm. from the anter ior end and

�9 09 mm. behind the intest inal bifurcation.

The restes lŸ very elose to eaeh other in the posterior th i rd of the body and are interc~~caI in position. The anter ior testis measures .2 mm. •

�9 ]1 mm. and the posterior .17 mm. • -112 mm. The vesŸ seminal is lies in front of the ventral sucker and is a bilobed body of . 1 mm. length.

I r ends in a small duetus ejaeulatorius which is sur rounded by pros ta te gland cells. The vesŸ seminalis, ductus ejaculator ius , a short cirrus and prostate gland cells ate all enclosed in a th in -wal led cirrus sae whieh 1les in a transverse position just in front of the vent ra l sucker.

The ovary which lies on the righe side of the med ian line in f ront of the two restes is an ovoid s t ructure and measures .08 mm. • .07 mm.

Stz~dies in Hel,mi~zt/zoloKy 161

The 5otype lies in the median planea little to the left of the ovary in

front of the anterior testis and is surrounded by small unicellular shell-

glands. There is no receptaculum seminis in these forros. The uterus

arises from the left side of the 6otype. Ir is a short tube, thick in the middle, and eontains very few eggs. It is con¡ between the 6otype and the ventral sucker and ends in a thick-walled metraterm which

runs along the left border of the ventral sucker and oloens at the genital

pote which lies at a distance of .02 mm. in front of the ventral sucker.

The eggs ate fairly large and thiek-shelled and measure �9 07 mm. x - 05 mln.

The vitellaria consist of very large and comloact follicles lying in the

posterior hall of the body behind the ventral sucker. They ate extra-

c~eeal and meet together behind the testes filling up all the post-testicular space. The two transverse vitelline ducts run just in front of the anterior

testis and open together into the 8otype.

Remarhs.--The present forro differs from all the desc¡ species of the genus Echinoch~smus except E. mordax, besides severa1 other fea tures , in the position of its eirrus sac whieh is transversely placed and is slightly

ahead of the ventral sucker. From E. mordc, x it differs in possessing t w o extra collar spines, ratio of oral to ventral sueker, in having bigger t e s t e s and smaller ovary and in having big massive extra-cmcal vitellaria. T h e present forro, therefore, appears to be new to science and because of i t s large vitelIaria Ÿ named Echiqr ~~egavite~~us, n. sp.

Ec]Knoch~sm#s ~'e#~ov~~~s 11. sp. (I~igs. 9 & t0)

Several specimens of this Trematode were recovered from the intestine

of the common house crow, Corvos S~~e~de~s. The worms ate more of less

oval in shape and measm'e 1-17mm. x -425 mm. The cuticle bears

minute spines which extend up to the anterior testis. The head is very

prominent ancl possesses a well-developed collar beset with spines. The

collar spines ate 2~ in number and ate arranged in two groups of 12 spines

each leaving a short space on the dorsal side in the region of the oral sueker,

They show a slight variation in size increasing in leng~h towards the dorsal

surfaee and forma single row except for ~ pairs of spines towards the

pharynx which show ah alternating arrangement.

The oral sncker is .l mm. x -095 mm. The prepharynx is extremely

small. The pharynx measures .09 mm. x -06 mm. and leads into ah

oesophagus of moderate size which bifurcates into the two intestinal c~ca

at its posterior end.


t

~ m r n .

Fiel. I0 .

Fle. 9

FIG. 9. Eehinochasmus ~'en'�91 n. sp., entire worm--ventral view. (Bocly spines not shown.)

FIe. 10. Ec~inochc,~~mus reniovarus n. sp. He~d collar showing spines. (Sketched from gl7cerine mount.)

The ventral sucker, larger than the oral, is circular in outline and

measures -16 mm. in diameter. It lies at a distance of .46 mm. from the

anterior end and -13 mm. behind the intestinal bifurcation.

The excretory bladder is peculiar inasmuch as it shows five tubular

chambers opening one after the other into the bladder. These are just the

lateral branches of the stem of the bladder which have become dilated at

their proximal ends. This gives a chambered appearance to the excretory

b ladder . T h e e x c r e t o r y p o t e is dorsa l a n d s u b - t e r m i n a t .

s225 in ZJefmim/zolozy 163

The testes, two in number, much broader than long, ate eontiguous and occupy the posterior third of the body. The anterior testis nleasures �9 25 mm. X -!4 mm. and the posterior -26 mm. X -145 111111.

The vesicula seminalis is d i s t inc t ly bi lobed and lies on the right side of the median line immedia te ly in f ront of the ven t ra l sucker. The cirrus is short and Ÿ enelosed in a eirrus sac which also encloses the vesŸ seminalis and prostate gland cells. The geni ta l pote lies at a distance of �9 016 mm. in front of the ace tabu lum in the micldle line.

The ovary is pre-testieular, k idney-shaped and dextral in position. I r measnres .1! mm. • .05 mm. The 5otype eomplex lies on the left of the ovary just in front of the anter ior testis . The uterus is short and contains a few eggs whieh ate thiek-shel led, large and measure .075 mm. •

�9 045 mm.

The vi tel lar ia which eonsist of smalI 1ounded foIlicIes extend from the posterior end of the body r ight up to the level of the pharynx. They meet bo th anteriorly and poster ior ly and forro a wrea th of foIlieles aeross the body.

Remarks.--The present forro differs from all the species of the genus except E. bagulai in possessing a chambered arrangement of the excretory bladder�9 But it differs from E. bagulai in the greater anterior extension of the vitellaria and shape of the ovary and restes. In its general external appearance i t stands nearest to E. corvus f~onl which i t differs in the position of the cirrus sac and possessing a d i s t inc t ly ehambered excretory bladder.

I t is, therefore, regarded a s a new speeies which is designated E. renio-

varus beeause of its possessing a k idney-shaped ovary.

Echi~~och~smus corvus Bhalerao, 1926

Bocly 1 ,06 -1 .08mm. • � 9 Collar k idney-shaped with 2,i spiues, disposed in one row, broken dorsally by a shor~ sp~ce. 3 i- 3 'P]okstache]n ' and 9 + 9 ' l~andstt~chehx '. Oral suoker -076-.086 mm. in diameI~el'. Ventr;~l sucker measures �9 13-- .14= mm. in dio%meter. Preph&rynx present, s]{ort �9 mm. in lengf, h. Pharynx -058 mm. Intestinal caca simple, extend to f, he poM;e¡ end of the body. Testos, one behind the other, separated by ~~ broad ba,ud o[ vitellaria from the posterior end. Anterior testis, transversely ova], entire nt~rgin and nmasures .~(i mm. x .]Smm. Posterior testis somewha/ round but … anteriorly, and measures -23 mm. x �9 175 mm. Cirrus sac lies dorsal to ventral sucker, in the median line, and extends posf, eriorly almos% to the hind rnargin of ventral sucker. (~enital pote, pre-acetabuL%r and behind intestinal bifurcation. Vesicula seminalis well developed. Pars prostatica and cluctus ejaculatorius very small. Ovary immediate]y anterior to testas, dextral, oval, me&sures .1-.13 mm. x .08-.i05 mm. l%ecepIsa~ulum seminis round, present on its innerside. Laurer's cŸ present, shell-gland in front of anterior testis. IJterus srnall, col~tailaing very few eggs, between anterior testis a~d vent~~l sucker.


Eggs ovat, opercul~ted, with brown shdl , oa~d measure ,0"/-.085 Imn. • ,OIO-.06 mm. Vitel lada consist of large ronnded follieles, si tntt ted dorsal ly Mong t he side of body , Vitellaria meeg cen~rally in f ron t of ventra l sucker and pos te r io r ly beh ind the tesLis. The?, do not extend up t,o pharynx.

Hod.--Corvus i~zsolens (Intestine).

Loca[[email protected].

Echi~,ocka~smus ruficGbensis Verma, 1935

]3ocly Iength 2.5 - - 3 -5 mm . ; lnaxi lnum bveadth, in f r o n t of ~~eeLM)utum, 0 4 - 0-6 mm. Ot'al suckee spherical 0-16 mm. of oval 0 -15 mm. • 0 -1 m m . Collar la, rge, t ransversely elong~ted, reni form of tria,ngular, 0 , 3 - 0 - 4 6 m m . • 0 . 5 - 0 . 6 5 m m . ; collar spines 24, dorsal ly i n t e r rup ted in one regular row on eaeh side, ventr .M-most of each side sm~ller, o thers near ly equal. P r e p h a r y n x 0 . 1 - 0 - 2 m m . ; p h a r y n x ov~~l, 0.15 mm. • 0 - I 7 mm. ; oesophagus long, 0 .2S-O.£ m m . Ventral sucker sub- globular.. 0 -36 mm . in diameter , a t one-third the body leng th f rom an te r io r end. O v a r y small, oval, a long mid-tramsverse line of sligl:itty ahead of iL ; t e s t e s me~i~n, close behincl one anogheI', in poster ior hMf of body, lnaegin smoo th ; an te r io r s u b q u a d r a t e of ovo- sphericaJ, broader t h a n long 0 -18-0 .22 mm. x 0 . t 0 - 0 . 3 5 m m . ; poster ior ovo- sphericM of sub- t . ¡ 0 - 1 8 - 0 . 2 5 m m . x 0 . l S - 0 . 2 m m . O r a 0 , 0 q - 0 . 0 8 m m . x 0.089-0.05'2. mm.

Host.--Podiceps ruficollis.

LocaliZy.--Atlahabad.

Echiuoch~smus bagul~i Verma, !935.

Length 1.0 mm. t,o 1 .76 m m. ; m a x i m u m width , abou t middle oE body , 0 ,25 mm. t~ 0.5 mm. OrM sueker 0 .07 mm. in diameter . Oolla.r r en i fo rm wi th 24 spines, in terrupted clorsMly, in two rows, a l ternat ing wi th one a n o t h e r ; spines of outer row 0.025-0.035zrma. long, of inner row 0 . 0 g I - 0-027 mm. long. P e e p h a r y n x ~~boufi 0.05 mm. long ; p h a r y n x 0 . 0 4 2 - 0 . 5 9 mm. in d iamete r ; oesophagus 2~ to 3 t imes ~~s lo~g as phar~qax. VengraI sueker 0 .2 mm. ilx cliamefler. Oval, y ~~d test, es in post;et'ic.,r l ia[f of body ; ov~ry roun&e&, 0 -05 mm . in cliamete~', to r igh t of med i~n line ; l;este.s m e d ª close behind one ano the r ; t r~nsversely elongated, variable in forro ~~n(t outline, ; ,~nterior ~est.is usuMly somewha t smMIer in dimensions t h a n the poster ior . Eggs lm'ge, q to 15 or more and measure 0 .067 -0 .0766 mm. x 0 .050 -0 .0588 m m .

Host<--Ardeola gmyi & Nycticorax nycticorax (Small intestine).

Locality.--Allahabad & Nagpur.

SteŸ n. sp,

Two spedmens of this trematode weze obtained froni the small intestine of the spotted red-shank, Tota~zusfuscus. The worms are elongated and cylindrical in shape and measure 4. 775 mm. • �9 65 mm., the greatest width being a li t t le behind the posterior testis. The head is marked off froin the rest of the body and bears a collar of 22 spines. The spines ate arranged in two groups of !I each, leaving a short space on the dorsal side in the region

S/ud;es i~~ HeImi~~Ÿ 165

of the oral sucker. The spines ate of uniform slze, measuring �9 05 mm. •

�9 02 mm. and ate arranged in one row except for four pairi of spines towards

the pharynx which show ah arrangement in double row.

.0

~.,o.

les.

v;e.9l.

/! FzG. 12

E

.~ mm.,

FzQ. II.

Fz~. 12.

FI(~. 11

Stephanoprora fusca r~. sp., enbire worm--vent0ral vfew.

Stephanoprora fusca--hea4 collar showir~g spines. (Sketched from glycerine motmt.)


The oral sucker is sub-ventral and measures -1 mm. in diameter. There is a small prepharynx measuring .15 mm. and this teads into a muscular pharyllx which measures -125mm. • .1 mm. This is followed by un oesophagus .275 mm. long which bifureates at its posterior end into the two intestinal caeca which run up to the posterior end of the body.

The ventral sueker Ÿ strong and muscular and measures .35 mm. in diameter. I t 1tes a t a d is tante of �9 9 mm. from the anterior end and �9 3 mm. behin& the intest inal bifurcation.

The testes are oval with ent i ie margin and lŸ at about the middle of the body. The anterior testis measures .375 mm. • mm. a n d the posterior .4 mm. • .22 mm. The vesicula seminalis is bilobed and lies on the right side of the median line. I t is enelosed in a thin-wal led cirrus sac which also eontains a small cirrus surrounded by prosta te gland cells. The eirrus sae measures �9 • mm. in length and bears the geni ta l aperture at its anterior tip.

The ovary is si tuated at a distanee of .75 mm. behind the ventral sucker in front of the two testes in the median line. I t is oval in shape and measures �9 15 mm. • �9 11 mm. The 8otype complex lies be tween the ovary and the anterior testis. There ate a large number of small unieeltular shell-glands present aro'~ncl the 8otype. The nterus arises from the left side of the 8otype, curves behind and after forming a loop runs dorsal to ovary towards the anterior end. Tkere ate a large number of eggs present in the uterus which measure .05 mm. • �9 025 mm.

The vitellaria eonsist of elliptical follieles and ex tend from in between the testes to the posterior end. The fotlicles of t he two sides approach each other but do not meet in the middle. The two vitel l ine ducts run parallel to the body in a longi tudinal clireetion to open into a very small yolk reservoir. A sma11 and narrow dnct from the yo lk reservoir leads into the 8otype.

Remar/~s.--The forro deseribed a~ove differs from al1 the existing species of the genus Stephano~rora. I t differs from S. fbolycestus and S. reyno~di in the absence of a reeeptaculum seminis. Je'roto S. pseudo- ecl~f~~atus, S. ornatab ancl S. spfnosa it differs in the posi t ion of the eirrus sae. I t stands nearest, however, to S. meru~r S. denticulatus and S. conciliatus. But i t differs from S. r~eru[r in ¡ smaller eggs, well convoluted uterus and in the absenee of a reeeptaculum seminis u ter inum. F rom S.. daenti- cuIat~~s it differs in possessing smaller eggs and in no t having its eirrus sac contiguous with the ventral sucker. I t cliffers f rom S. co~,ci~ia~us in having

S~udies be t-Id~ningho/o~y 167

a dextral position of the cirrus sac, smaller eggs, longer prepharynx and non- contiguous restes. I r is therefore, a new speeies for which the name Steph~noprora fusca n. sp. is proposed.

Stephan@rora reynoldi Bhalerao, 1926

B o d y elongated, cuticle covered w i t h spines up t,o the poster ior m a r g i n of anterior testis. Collar kidney-sha,ped wi th 22 spines in one row and b roken dorsal ly. 2 ' Ecks ta - c h e l n ' and 9 ' 12~anclstacheln' on each side. Oral sucker .21 n~m. • .68 mm. Pre- p h a r y n x short . Testes w i th ent i re m~~rgin o~e beh ind the other, in the midcUe of body . Anter ior tes t is .39 mm. • .33 m m . , pos te r io r .50 m m . x .33 mm. Cirrus sac s i tu~ted sinistral to ven t r a l sucker a~ld ex t ends ~rom the in tes t ina l fork to the poster ior end of ventra l sucker, pm~r-sh,~ped and contMns two- lobed vesŸ seminMis. Pa r s prost~t ica an d ductus ej,~cula.torius small. Ovary , au te r io r t o restes, oval, s l igh t ly dextrM and measures .15 mm . • . l l S m m . R e c e p t a c u l u m seminis post-ovar ia l , and shell-gl~nd behind ir. Laure r ' s ca,nal present . Uterus , w i th few ova, be tween an te r ior test is a nd vent ra l sucker. Eggs measure .056- .0895 m m . • .031- -049 m m . Vitellaria consist of large folHeles. Yolk reservoir present . Vitellm:i~ ex tend f rom the centre oZ the anter ior test is t o the poster ior end and run 1,~terM to the body, ~pproach ing e~eh o ther closely a t the poster ior end. E x c r e t o r y pore ternSnal .

Host.--Corvus insolens (Intestine).

Locality.~Rangoon.

Subfamily -Parorc]~in~ La1, 1936"

Genus Parorchis Nicoll, 1907

This genus {s of rare occurrence and is recorded for the first time in India and the species Parorchis snipis Lal, 1936, forros the tkird valid species of the genus.

Parorchis sn{Ÿ I, al, 1936"

Bocly boset with spŸ for the anterior 2/3rci l)art ; 2- 58 mm. x 1�9 11 mm. Head ineonspieuously marked, rŸ collar with single row of miz~ute spines. Four pairs of spines at the laterM margin of collar la]~ger, �9 mm. lo~g. Oral sucker �9 21 mm. • .27 mm. Ventral sucker �9 mm. x .55 mm. ]?repharynx .07 mm. Pharynx �9 12 mm. x �9 18 mm�9 O~sophagus �9 19 mm�9 long. Excretory bladder .26 mm. x �9 3mm. l~ight testis .35mm. x .27mm. Left testis .36mm. • .225mm. VesŸ semi~is bahincl the ventral sucker .2 mm. x . 13 mm. Cirrus sac .09 mm. long. Ovary .185 mm . • .14 m m . R, e c e p t a c u l u m seminis behinc1 o v a r y .035 mm. • �9 032 mm. Shel l -gland post -ovar iM. ~Iale an(t femMe ope~ings lie tog£ in a genitM a t r i u m behind in tes t inM b[furc~t ion. Eggs in a s[ngle row, show segmenta t ion , and measure . 0 5 - . 0 6 mm. • .025 m m . Vitellir, e glands, abou t a dozen follicles on e~ch side, pa r t l y extra-cmeM a n d p a r t l y i n t e rc~ca l .

Host.--Totanus 7~yŸ (Cloaca).

Locality.~Lucknow. B5


The author also discussed the systematic position of the genus Parorchis and carne to the conclusion tha t this genus should be placed in a separate subfamily Parorehinm LaI, 1936, in the family Echinostomidee. In the same communication* evolution of Echinostomes was discussed and a polyphyletic origin of Echinostomid~e was suggested.

IsoZated Genera of the Family Echiqwstomidr

'1'he genera Paryphostomum and Petasiger ate not included in any of the existing subfamilies of Echinostomidm. The writer is not in possession of material of these forma and cannot definitely express his opinion. But on looking up the available literature, he rinda tha t both these genera should be ineluded in the subfamily Eehinostorninee, because of the arrangement of collar spines in ah uninterrupted row and non-extension of its cirrus sac behind the ventral sucker.

Paryphostomum testitrifolium Gogate, 1934

Bocly e longa ted 3 - 5 - 5 m m . , a r m e d w i t h spines in f r o n t of v e n t r a l suck~r. Y[axi- muro w id th in the reg ion of t e s t e s . 5 5 5 - . 9 5 2 m m . H e a d col lar s t r o n g l y deve loped single row, 27 spines, u n b r o k e n dorsa] ly . V e n t r a l l y col lar sp ines g~therect i n to t w o ' end ~ffroups' of ~ spines each. Ora l s u c k e r s u b - t e r m i n M , m e a s u r e s -115 m m . x �9 128 m m . P r e p h a r y n x smalI, .027 m m . P h a r y n x g lobu la r .135 m m . • .093 m m . 0 e s o p h a g u s -464 m m . long. I n t e s t i n a l cmca r u n to p o s t e r i o r end of b o d y . V e n t r a l sucker .517 m m . x .539 m m . enclos ing a spac ious cup-sh~pecl c a v i t y . Gen i t a l po t e in b e t w e e n ven t r a l sucker and i n t e s t i n a l b i fu rea t ion , m e d i a n . Tes t e s , b r a n e h e 4 , t r i - foliate, t w o an te ro - l a t e ra l ~nd one p o s t e r o - m e d i a ~ b r a n c h e s ly ing one b e h i n d t h e o the r ; p o s t e ¡ ]arger t h a n an ter ior . Cirrus pouch �9 197 m m . • .0945 m m . , dorsal and oblique to v e n t r a l sucker. O v a r y globular , -17 m m . x .165 m m . , p r e - t e s t i c u l a r . Shell-gland in be tween o v a r y a n d a n t e r i o r t e s t i s , l ~ e c e p t a c u I u m seminis , pos te r io r to ova ry . Vi te l lar ia e x t e n d b e t w e e n v e n t r a l sueke r a n d p o s t e ¡ end of bccly, filling the pos t - t e s t i cu l a r p o r t i o n w i t h loosely p a c k e d follicles. U te rus , s h o r t n o t m u c h coiled, pre- tes t icul~r , intercme~l. V a g i n a is do r sa l to v e n t r a l sucker . O r a , n u m e r o u s , smaU .0771 m m . x .0409 m m .

Host.--Dendrocygna javanic~z (Intestiae).

Zocality.--l?.angoon.

Petasiger minutissimus Cogote, 1934

B o d y small , e longa ted .948-1-3S8 m m . ~Vell-developed heact col lar w i t h 23 spines, of which 17 a te arrangecl in a s ingle dorsa l ly u n i n t e i r u p t e d row, m e a s u r i n i .0368 - . 0 6 1 3 m m , x . 0 1 - . 0 1 7 i m m . R e m a i ~ i n g 6 spines, in t w o v e n t r a l ' e n d g roups ' eaeh w i th 3 spines, m e a s u r i n g . 0 5 5 2 - . 0 Ÿ m m . • - 015 - .0225 m m . 5 I a x i m u m w i d t h

* Vide Proc. Ind. Acad. Sc~., 1936, 4, No. 1, Sec. B, 27-35 .

Sfudies in Ztelmin, t/mfogy 169

behind vent ra l sucker -29t- .349 mm. Body anterior to~ventrat sucker covered with spines. Oral sueker .045--053mm. Preph-~rynx -05-.06 mm. P h a r y n x globular �9 0495-.05 mm. Oesophagus thin .3--3B mm. Intestinal c~ecal ends hidden in the

vitellaria. Ventral sueker .177- 195 mm. x .198-.226 mm. GenitM pore, ventral to inLestinal bifureation. 'restes oval, overlapping, with loI~g axes a t right angles. Cirrus pouch �9 113 mm. x -121 mm. dorsal to ventrM sucker and h~lf por t ion extencling bebween ir and the intestinal bifureation. Ovary ovoid, -06~4--064 mm. x -066 mm. elose to ventral sucker, laterM. Recaptaculum seminis and shell-gland obscured b y vitelline follicles. Vitelline fo[ticles behind posterior margin of vertgral sueker to posterior end of body and filting loosely the po~t-testieular regiolx. Uterus, pre- tes~icular, short. Ova, [ew large .0592-.093 mm. x -0368-.0-18 mm.

Host.--Dendrocygna javanica ( In t e s t ine ) .

Locality.--Rangoon.

K e y fo r l h e i d e n t i f i e a t i o n of t h e A v Ÿ G e n e r a a n d Spee i e s of t h e f a m i l y

E c h i n o s t o m i d e e r e p o r t e d f r o m I n d i a :

! . T e s t e s c o m m b i a l . . . . . . Parorchis sn@is. ' r e s t e s t a n d e m . . . . . . 2

2. S p i n e s o n co l la r b r o k e n on t h e do r sa l

s i d e . . . . . . . . 3

S p i n e s o n co l la r u n i n t e r r u p t e d on

d o r s a l s ide . . . . . . 4

3. Co l la r s p Ÿ 24 . . . . . .

Col la r s p i n e s 22 . . . . . .

4. S e m i n a l ves ic le u n d i v i d e d . .

S e m i n a l ves ic le i n c o m p l e t e l y b i l obed .

5. Co l l a r s p i n e s 27 . . . . . .

Col la r s p i n e s 23 . . . . . .

Collar s p i n e s 32 . . . . . .

Co l l a r s p i n e s 37 . . . . . .

6. I ~ x c r e t o r y b l a d d e r w i t h c h a m b e r e d

d i v e r t i c u l a . . . . . .

E x c r e t o r y b l a d d e r w i t h o u t c h a m b e r e d

d i v e r t i c u l a . . . . . .

7. V i t e l l a r Ÿ e x t e n d p r e a c e t a b u l a r . .

V i t e l l a r i a b e h i n d a c e t a b u l u m . .

8. V i t e l l a r i a m e e t in t h e p r e - a c e t a b u l a r

r e g i o n . . . . . .

V i t e l l a r i a b e h i n d a c e t a b u l u m . .

Echinochasmus (see 6).

SteŸ (see 10).

5 Echinostoma chasma.

Paryphostomum Ÿ Petasiger minutissimus. Echinostomum govindum. Echinsotoma revoIutum.

7

8

Echinochasmus reniovarus. Echinochasmus bagulai.

Echinochasmus corvus. 9


9. Cifras sac, transverse, ahead of ventral sucker . . . . . .

Cifras sac, vertical, touching ventral sucker . . . . . .

10. Cifras sac dextral . . . . Cifras sac sinistral . . . .

Echinochasmus megavitdlus.

Echinochasmus rr

Stephanoprora fusta. Stephanoprora reynoldi.

Family Psilostomidce Odhner, 1913

Odhner (19!3) raised the subfamily Psilostominm Lª (1909) to the status of the family Psilostornid~e and described several new genera under it. Travassos (192!) added the genus LyperorcMs and Bhalerao (!931) deseribed another genns Testifrondosa~ from the intestine of pig. Thapar and Lal (!935) deseribed frorn King-fisher the genus Psilorchis, the only Avian genus of T~ematode recorded from India under the family. Lal (1938) descfibed the second species, Psilorchis ajgainis, of the genus from ,~~ettion crece�91191

Family Diagnosis.--Worms wŸ flattened leaf-like body of va¡ length. I-l:ead indistinct without any collar. The internal structure is comparable to tha t of the family Echinostomidae. Skin smooth of sometimes covered with scale-like of simple spines. Pharynx strongly developed. Intestinal bifurcation in front of the ventral sucker. Intest inal c~eca reach almost up to the ]~osterior end of the animal. ~Excretory bladder Y-shaped. The two limbs of the bladder ioin each other in the region of the ventral sucker to forro a big unpaired sinus, Ovary is anterior to both the testes. The disposition of the genital apparatus is on the sume plan as tha t in :Echinostomidm. Vitellaria consist of distinct follicles and present on the lateral sides of the body ventral to intestinal caca. They do not always meet together in the middle line posteriorly. :Eggs similar to those of the Echinostor~es, about ,088-.125 mm. long. Parasitic in the intestine of birds and mammals.

Genus Psilorchis Thapar and Lal, 1935

Psilostomidce with leaf-like body. Ventral sucker much larger than the oral. Short Y-shaped excretory bladder. Genital pote, dextral, in front of the ventral sucker, Testes more oz less bean-shaped and tandem ; each testis provided with a well-developed funiculus which leads forward

t Chatterji (1938) mentions th~t Tes~~frondosa cristata ]3haler~o (1931) is synonymous with Paryphostomum su#artyfe,v L~ne (1916) which comes tmdcr Echinostomidm,

S• ilz I-�91191 g y 171

i n t o a v a s d e f e r e n s . R e c e p t a c u l u m s e m i n i s a b s e n t . A y o l k r e s e r v o i r

p r e s e n t . U t e r i n e c o i l s l i e i n f r o n t o f t h e t e s t e s . V e s i c u l a s e m i n a l i s r e t o r t -

s h a p e d , s i t u a t e d i n f r o n t o f t h e v e n t r a l s u c k e r ; c i r r u s s h o r t . V i t e t l a r i a

b e h i n d t h e v e n t r a l s u c k e r a n d d o n o t m e e t t h o s e o f t h e o t h e r s i d e

p o s t e r i o r l y .

T y p e species--PsiZorckis indic~~s.

K e y f o r t h e i d e n t i f i c a t i o n o f t h e s p e c i e s o f t h e g e r m s Psilorchis r e p o r t e d f r o m I n d i a :

C i r r u s s a c a d h e r i n g t o t h e v e n t r a l

s u c k e r ; o v a r y s p h e r i c a l . . . . P. ajgainis.

C i r r u s s a c s e p a r a t e f r o m t h e v e n t r a l

s u e k e r ; o v a r y o v a l . . . . P. indicus.

Psilorchis indicus T h a p a r a n d L a l , 1935

B o d y 8 . 5 7 m m . • 1 .17 m m . ; s m o o t h cugicle w i t h o u t spines . Oral sucker �9 I7 m m . x �9 1 m m . V e n t r a l s u c k e r s i t a a t e d in a n t e r i o r 2/9th p a r t of body , measures �9 75 m m . X -65 m m . P r e p h a r y n x - 18 m m . loag ; P h a r y n x �9 I8 m m . x �9 16 m m . Oeso- phagns �9 09 m m . long. 2~nterior t es t i s �9 75 m m . x . 1.2 m m . Pos te r io r t e s t i s �9 75 m m . X �9 S6 m m . Ov,~ry �9 ~1 m m . X - 25 m m . E g g s Iarge, oval , �9 125-130 m m . X �9 08 - . 1 m m .

Host.--King-fisher ( I n t e s t i n e ) .

Locality.--Lucknow.

_psi~orchis ajgai~is L a l , 1 9 3 8

B o d y 6 . 6 5 m m . • .75 m m . ; cut ic le w i t h o u t sp[nes. Oral sucke r .2 m m . in d iamete r . Ven t r a l sueker -65 m m . x -55 m m . P h ~ r y n x �9 1 m m . i n d i ame te r . Oeso- ph~gus -125 m m . long. 2 in te r ior t e s t i s .65 m m . • .375 m m . Pos t e r io r tes t i s �9 7 m m . x .425 m m . O v a r y .25 m m . in d iameber . E g g s lavge, oval , �9 1 - . 125 m m . x �9 0 4 - . 0 5 m m .

Host.---Ne~~io~ czecca ( I n t e s t i n e ) .

LocaZi~y.--A~gain.

Family Strigeida~ R a i l l i e * , 1 9 ! 9

T h e f a m i l y S t r i g e i d a w a s e r e c t e d b y R a i l l i e t ( 1 9 / 9 ) f o r t h e f o r r o s

previously included under I-Iolostomidm Bianchard, 1847 and I-lemisto- midm Brzndes, 1888. I-Ie further'created a new superfarnily Strigeoidea for the whole group having only one family, Strigeid~e. iVfore reeently a comprehensive study of/che group in the form of a working classi¡


has been given by LaRue (1926). Under the family Strigeiclm, he has listed ¡ subfamilies, Strigein~e, Brauninin~e, Cyathocotylinm, Polycotylinm aad Alariin~e.

Poehe (1925) exeluded the subfamily Cyathocotylin~e from the family Strigeid~e on the basis of presence of a cirrus sac and absence of the division of the body into two regions and erected the family Cyathocotylid~e. Szidat (!936) has further elaborated this view and has divided the family into several subfamilies. On the other hand, Bosma (!931) reeognized the Alari- idm of Tubangui (192'2) a s a distinct family and Dubois (1933) divided ir into three subfamilies.

But in spite of these modifications, the writer is still inclined to main- tain the classification of LaRue (!926) as he considers that the differences between the various subfamilies of StrigeŸ ate not of such a great importance as to warrant their separation into different families.

Diagnosis of the family Strigeidce Railliet, 1919.--

The body is us'ually divided into two regions by a constriction, ah anterior part bearing chiefly the special organs of attachment anda posterior part containing the major portion of the genitalia. The forebody is flattened of incurved ventrally of cup-shaped; the hind body is more or less cylindrical, ovoid of conical. The oral sucker is terminal of sub-terminal ; the acetabulum is generally 10resent but ir is weak. The holdŸ organ is situatecl posterior to the ventral sucker and is sometimes covering the latter. The intestinal cseca are without divertieula, usually extending to the poste¡ end of the body. Cirrus and cirrus sac present of absent. The vitelline glands ate strongly developed. The uterus consists of few coils containing but few eggs.

The writer is giving below a key to the snbfamilies of Strigeid~e Railliet,

1919 :

I. Cirrus sac present . . . . 2

Cirrus sac absent .. 3

2. Body not divided into anterior and posterior part . . . . . . Cyathocotylinm.

Body more of less divided into anterior and posterior loart . . . . Braunininm.

3. Anterior part of the body cup-shaped Strigeinm.

Anterior part of the body ¡

Studies in t]dmint~o/o[y 173

4. Uterus confined to hindbody and does not enter the holdfast organ .. Polycotylin~e.

Uterus usually enters the holdfast organ . . . . . . . . Alar¡

Subfamily CyathocotyIinr Muehling, 1898

Diagnosfs.--The body is short and broad, without division into disdnet anterior and posterior regions. Cirrus and eirrus sae ate present, The genital pote is situated at the posterior end of the body.

Type genus--Cyathocotyle.

Only one species of the genus Cyathocotyle has been reported from India.

Cyathocotvle calvusi Verma, 1936

B o d y 2-06 mal . x 1.02 mm. OrM sucker 0 .15 mm. x 0 .19 mm. , p h a r y a x 0-09 mm. • 0 .097 mm. , oesophagals alXCl p r e p h a r y n x absent . ]E[oldfast o rgan 0 . 3 2 - �9 4 mm. in d iameter , wall covered w i t h v e r y pecu l ia r character is t ic , hair-l ike, ¡ outgrowths. Testes ve ry conspicuous, rnuch e longated ; an ter ior 0 .56 mm. • 0-18 m m . , Mongside holdfas t org~n ; poster ior 0 -65 mm. x 0 .18 mm. in na r rower par t of body , separated f rom the h ind end by abou t hall i ts length . Ova ry roundish, 0-28 mm. in diameter, rnaskecl by vitell[ne follicles, an tero la tera l to h ind testis. Vitel laria grouped in large foUicles of i r regular shape a long intest inM c~ba mee t ing in f ron t . Cirrus sac 0-00 mm. • 0 .14 mm. , con ta ins coi•ecl seminal vesicle, prost~t ic cells and long tubular , eversible cirrus. Genital a t r i u m short , wi th m~le a n d fem~le duc t s opening side by side in to i t ; genital pote postero-berminM. Eggs abou t twice as long as th ick, no t m a n y , 0 . 081 -0 .092 mm . • 0 . 0 4 1 - 0 . 0 4 7 mm.

Host.~Torgos c~Ivus (Intestine).


Subfamily Polycotylince Montieelli, 1892

Diagnosis.~Body divided into two regions, forebody flattened, hindbody cylind¡ Cirrus sae and cirrus absent. I-Zoldfast organ round, elliptieal Gr bulbous. Lateral wing-like suckers present Gr absent. Genital pore posterior. Uterine coils con¡ to hind body and never enter the holdfast organ.

Type genus--Polyco~Se.

�9 Neodi~iostomum dilaccecum n. sp.

A large number of these parasites were obtained from the small intestine of the spotted owlet, Athe~e Bmmg. The body is flat and broad

* A number of specimens of this forro were also obta~ned from Dr. Thapar's co~lec~ion.


and shows a clear division into a long anterior a n d a short posterior part . The posterior margin of the anterior portion of the body is produced into a thin flap which is placed transversely across the body and divides it into two parts.

1L

I~IG. 13 Neodi~lostomum dilacczcum n. sp., enitre worm--ventral view.

The length of the specŸ is 2.35 mm., and its breadth is �9 9 mm.

The anterior po~tion of the body measures I-35 mm. and the !ooste¡ por-

tion 1.0 mm. The antero-ventral oral sucker measures -I mm. • .05 mm.

The mouth leads into a narrow prepharynx which passes into a thick-walled

Stua(ies in Helmi~zt,~olo~y 175

and muscu la r p h a r y n x �9 075 mm. • 05 mm. in size. The oesophagus is short a n d b i fnrca tes into the two in tes t ina l cmca whieh run almost up to the pos te r ior end of the animal t e rmina t ing in grea t ly dilated, club-shaped ends.

T h e ven t r a l sueker is muscular and measures .1 mm. in diameter. The ho ld fa s t organ is a hear t - shaped s t ructure wi th i ts apex point ing an te r io r ly . I t encloses a wide eavi ty in i ts posterior part . There is ah inconspicuous adhesive-gland mass at the posterior end of the hold2ast organ.

The gonads oceupy the posterior hal l of the body. The ovary is elliptical a n d measures -25 mm. • .15 mm. The 5otype with she11-gland lies in be tween the testes. The uterus, s tar t ing from the 5otype, takes an ascending course up to the leve1 of the ovary. I t then turns baek in a descending loop and runs dorsal to the testes to the posterior end where ir opens in to the geni ta l a t r ium. The eggs measure �9 1 mm. • �9 05 mm.

t 2 m ~ .

FIG: 14

Neodiplos~omum dilaco~cum, Reproductive organs enl~rged--dorsM view.


The restes, two in number, are broad ancl transversely elongated boclies. The anterior testis measures .525 mm. x .275 mm. and the posterior mea- sares .5 mm. • .3 mm. The vesicKla seminalis is a thick convolutecl structure lying behind the posterior testis and opening into the genital atr ium along with the female dtlet.

The viteltaria ate d is t r ibuted th roughont the anter ior and the posterior part. Anteriorly they extend a l i t t le beyond the ven t ra l sucker and ¡ up the entire spaee of the holdfast orgam Poster ior ly they s t re teh up to the di lated czecal ends.

Remarks.--The forra described above differs from al1 the species of the genus except N. @athul~for~ne, N. gavi~lis, N. orchilongum, N. cochlear G and N. Thomasi in having a larger broad and fiat anterior part of the body. But it.differs from all these species in several points. From 37. sj~athulce- forme ancl N. gavfalis ir differs in the absence of papill~e around the holdfast orgam Ir differs from N. tyter~se, in greater posterior extension of the vitellaria and the shape of the anterior testis. From 37. orchilo~gum ir differs in the position of the 8otype whic• in N. orchflongum is rather peculŸ situsted in front of both the restes, iVrom 37. cochle~re it is distinguished in having a broad and fiar posterior part of the body and in the absence of a cup-shaped union of the margin of the anterior part of the body. From N. T]r ir differs in the shape of the posterior part of the body, in the shape of the restes, and irt the absence of receptaculum seminis.

Ir is, thus, evident that this forro stands apart from all the known species of the genus. Ir is further distinguishecl by the possession of a heart-shapecl holclfast organ and greatly dilated and club-shapecl caecal ends. Ir is, therefore, designated as a new species.

Neod@lostomum sp.

A single immature speeimen of th is • was obta ined from the intest ine of the blue jay, Coracias benghalensis. The an imal is 1-675 mm. Iong and has the max imum bread th of - 575 mm. in the level of the hotdfast organ. The body is distinctly divided into • parts. An extension of the posterior margin of the anterior portion overllangs the anterior end of the posterior portion of the body. The anterior portion of the bocly is - 905 mm. long including the marginal expansion of �9 05 mm. The posterior parŸ measures �9 82 mm. long.

The oral sucker is oval in outline and measnres .05 mm. • .07 mm. The mouth leacls into a very short prepharynx which passes into a globu/ar

Studies in IIeImbzilwlo~y 177

muscular pharynx �9 031 mm. in diameter. The Oesophagus is rather sma11 and immediately divides into the two intestinal c~eca.

F I o . 15 Neod,�91 sp.--ventrM view.

The ventral sucker, which lies a t a distance of .425 mm. from the anterior end, measures -07 mm. x - 0 7 1 mm. The holdfast organ consists of two hemispherical pieces enclosing a cavity between them and measures .217 mm. • .185 mm.

The ovary lies a t a distance of .08 mm. from the anterior margin of the posterior port ion of the body. I t is an ovoid body, measuring �9 145 mm. X .!1 mm.

The anterior testis which lies close to the posterior is .085 mm. froln

the ovary and measures �9 17 mm. x �9 085 mm. The poste¡ testis measures

�9 185 mm. x .II mm.

The vitellaria extend almost throughout the body of the animal and consist of small follicles. No other details could be made out in the specimen.

Remarks.--=From the description given above, ir is clear tha t the trematode belongs to the genus Neod@lostomum because of the division of the


body into two distinct parts, overhanging margin of the anterior part and absence of any lateral suckers at the anterior end. But in the absence of greater details of structure, ir is not possible to assign any definite specific position to this trematode.

Neod@lostomum tytense Patwardhan, 1935

]3ocly 2-82 m m . long, dividecl into two unequM regions. Foreboc ly f l a t t ened ancl measures 1.62 m m. • 1 -23 mm. I t s lateral lnargins unida pos te r io r ly to forro a spoon- shaped depression, t t i n d bocly cylindrical a ad measures 1-2 m m . • .63 m m . Outicle smooth . Oral sucker smal], and mea sures -056 m m . in 4 i ame te r . Pharyr~x, sma l l globular .062 mm. in d iameter . Oesophagus short . A c a t a b u l u m .087 m m . in d i a m e t e r . ITcldfast organ, sub-circular , sm~ll, .294 mm. in d i a m e t e r and lies .22 m m . b e h i n d vent ra l sucker. O v a r y in f ront of tes tes aL the j u n c t i o n of fore and h i n d body , o r a t e ancl measures .25 m m . • -19 mm. Uterus and eggs n o t 4eveloped . Vi te l la r ia cons i t s of a large n u m b e r of small follicles, scatterecl a r o u a d the ho ld [a s t o rgan , a n d e x t e n d in to the anter ior th i rd of h ind body. Shell-gland s i t u a t e d near t he left side a t a level between the two restes. Testes, one behind the o ther , in the m i d d l e - t h i r d of h i n d body . Anter ior tes t i s pear-shaped, slighbly to leŸ a n d measures -375 m m . • .225 m m . Pos ter ior test is deeplv bilobed, and measures .45 m m . • .225 m m . VesŸ seminMis larga sac behincl pos te r io r t~stis. Genital pore a t pos te r io r end of bo4y .

Host.~Tyto alb~t sterte~~s (Intestino).

Locality.--~agpur.

Proadaria alcedensis Patwardhan, 1935

B o d y 2 .24 m m . long, dividecl in to two par t s . F o r e b o d y f lat tenecl m e a s u r o s �9 72 mm. • "66 m m . H i n d body, cylindrical , and measures 1 .52 m m . • .5 m m . Oral sucker, t e rmina l a n d measures .035 mm. in di~mater . On i t s e i t h e r sida ~re s i tu › red sucker-l ike prominences . No p r e p h a r y a x . P h a r y n x s p h e r i c a l , mtu~cular and .037 mm. in d iameter . Oesophagas short . A c e t a b u l u m , t r a n s v e r s e l y o r a t e , �9 056 mm. • -048 m m . in size. Hold[as t o rgan s l [ght ly ova l and i n v a g i n a t e d pos te r ior -

ly. I t measures -175 m m . • .15 mm. A pair of ~dhas ive glancls a te s i t u a t e d beh ind the hol4fas t organ. O v a r y t ransverse ly oval s i t ua t ed cons iderab ly betf ind the j u n c t i o n of fore and h ind bo4y . I r measures .125 mm. x .1 m m . Vite l lar ia , of c losaly p a e k e d follicles, confmad to an te r ior hall of the hind body. Ute rus extencls a shorb d i s t a n c e in f ron t of the o v a r y and conta ins a few bu t largo eggs me~sur ing .075 m m . • .092 m m . Geni ta l poro a t pos te r ior end of body. Testas lŸ in middle-thircl of h i n d bod• ~n(l occupy ent i re wid th . Anter ior test is measures .5 m m . • -325 m m . and p o s t e r i o r tes t is -4 m m . • -41 m m . VesŸ semi~alis is vo luminous ~nd sac- l ike a n d lies behincl the poster ior test is . Det~ils of cirrus, vesicula seminalis , pars p r o s t ~ t i c a n o t gival'l.

Host.--Alcedo atthis (Intestino).

Locality.--~agpur.

Smdies i~z t-Ielmiu#zolo2"y 179

Neoalaria thaŸ n, g., n. sp.

About hall a dozen specimens of this trematode were obtained from the intestine of the king vulture, SarcogyŸ calvas s t Lucknow. The animals showed active movements of the body whieh is fiar and marked

FIG. 17

]PIs, 16

FIG. 16.---~'eoaZar{a thapar�91 n. g., n. sp . , - -ven t rM view.

Fin. 17.--NeoMaria tkal)aria~rePrOdUCtiVe organs enlal'ged~ve~~6rM view.

of~ into two regions by a very inconspicuous constriction. The posterior end of the animal has a bursa which is shaped like an inverted cup. The writer has not been able to observe any sucker of papill~e inside the bursa.

The length of the specilnen is 2.45 mm. I ts maximum width in the region of acetabulum is �9 9 mm. The anterior par t of the body la roughly square in outline and is much smaller than the barrel-shaped posterior part.

The oral sacker which projects a little beyond the anterior margin of the body measures -15 mm. x -!25 mm. The month leads into an extremely small prepharynx which passes into the muscular pharynx


�9 !25 mm. • .1 mm. The oesophagus is short and thin and bifnrcates into the intestinal c~eca at its posterior end.

There are two wing-like lateral suckers situated by the side of the oral

sucker. They measure .21 mm. in length. The ventral sucker, ahnost circular in shape, is well developed and muscular and measures �9 25 111111. in diameter. The holdfast organ is trough-shaped with thick wall and lies at the level of the demarcation of the body. There is no ad_hesive gland in this trematode.

The gonads ate eonfined to the barrel-shaped posterior part of the body and lie in transverse plane parallel to each other.

The testes ate peculiarly dumb-be11 shaped and lie at a distance of �9 3 mm. apart from each other. The anterior testis measures .5 mm. • �9 25 mm. and the posterior .5 mm. • .275 mm. The vesicula seminalis is thick and convoluted behind the posterior testis and opens into the bursa.

The ovary is elongated and is placed mesially, -15 mm. ahead of the anterior testis. I r measures .3 mm. • .125 mm. The 8otype complex lies in between the two testes. The uterus rnns in ah ascending a n d a descending loop, a l i t t le ahead of the ovary bnt is confined to the posterior body and does not enter holdfast organ. I r opens into the bursa along with the male duct. The eggs ate thick-shelled and measure .07- �9 075mm. • .045--05mm.

The vitellaria also show peculiar distribution. They extend up to the level of oesophagus anteriorly a n d a lit t le behind the testes posteriorly. They ate elosely arranged arotmd the ventral sucker and the holdfast organ but towards the periphery they extend in ehains, cutting clear parenchymatous spaees in between the chains. In the hinder par t they ate eontined to the margins and do not extend into the middie of the body.

To sum up, the genns Neoalaria is characterised t hus :

1. Trematodes with body, not distinetly divided into two regions. There is no tmion of the posterior lateral margins of the forebody. The anterior part of the body is roughly square ; posterior barrel-shaped.

2. Ventral sncker larger than oral; lateral wing-like suckers present.

3. I-Ioldfast organ trough-shaped; adhesive glands absent.

4. Gonads Iie parallel to each other in transverse plane.

5. Testes peeuliarIy dnmb-belI shaped.

6. Bursa withoat any sucker or papillm.

Sludies b~ 7:tdminlhofog'y 181

7. Uterus, with ascending and descending loops, not entering the holdfast organ.

8. Ootype complex in between the testas.

9. Vitellaria marginal in posterior body; in anterior body, dense around ventral sucker and holdfast organ and extending iR transverse chains towards the periphery and cutting clear parenchymatous tissue in between the chains.

I0. ]Eggs, thick-shelled.

larom the above, itis evident that the present forro stands apart from all the genera of St¡ except Alaria and _Proa�91 in possessing lateral wing-like processes at the anterior end. Ir dif~ers from A�91 in having its uterus behind the holdfast organ and in possessing a small trough-shaped holdfast organ and the absence of any union of the posterior lateral margins of the forebody. Ir differs from Proa�91 in the absence of a clear division of the body into two parts and in the absence of adhesive gland. From both these genera ir differs in having peculiar disposition of the vitellaria, dumb-bell shaped restes, btlrsa without sxcker of papilla and peculiar shape of the body.

Ir is, therefore, regarded asa new genus for which the name Neoalaria6 is proposed with Neo~zltzria. ~ALz~~zr~;a es the type species.

As the new genus resembles Proalaria in the disposition of the uterus, it is being included under the subfamily Polycotylinae.

Sub/amily Alariince I-tall and Wigdor, 1918

Body indistinctly or distinctly divided into two portions, anterior part flattened, posterior part broad or cylind¡ Cirrns and cirrus sac absent. IKoldfast organ corclate of elongated oval. Lateral suckers, tentacles of ilaps usually present near the oral sucker. Uterus usnally enters the holdfast organ.

Type genus--Alaria.

P¡ bagu~um i'ff. Sp.

Two specimens of this Trematode were obtained from the intestine of the eastern grey heron, A~'dea ci~r rectirostris. The body shows division into two parts although the constriction is only super¡ The anterior part of the body is fiar and longer than the posterior part which is short and more or less ovoid in shape. The anterior par t measures �9 63 mm. and the posterior �9 35 mm. in length. The lateral margins of the anterior part of the bo6y do not show any union.


FIa. 18

_Pharyngostomum bag~dum n. sp.--ventrM view.

The mouth is surrounded by an oral sucker measuring .05 mm. by �9 03 mm. and leads into ah extremely small prepharynx which is followed by a globular pharynx .05 mm. x .04 mm. in size. The oesophagus is th in and divides at its posterior end into the two intestinal c2eca which run up

to the holdfast organ and ate not visible behind ir.

The holdfast organ is a peculiar thick-walled C-shaped structure with the mouth of the ' C ' directed forwards. The writer has been unable to locate any adhesive gland which is apparently absent.

The ovary is median in position and lies at the constrŸ region of

the body. It measures .085 mm. • mm. Close behind it ate the

two testes, the anterior is elongated and bil0bed and measures .26 mm. •

�9 15 mm., and the posterior is smaller and measures �9 175 mm. • �9 12 mm.

Studies in HelmŸ 183

The vi tel la¡ extend anteriorly up to the level of oesophagus ; posteriorly they are scattered and less numerous behind the restes.

The uterus and vesŸ seminalis open together into a genital atrium which lies a li t t le distance in front of the poste¡ end. A mass of indistinct tissue behind the ovary is probably of the nature of 6otype, shell-gland, etc., but ir has not been properly identi¡ There ate no eggs in the anima1 which appears to be ah immature individual.

Remarks.--The forro described above comes under the family Strigeidm but does not possess either a cirrus sac or cup-shaped anterior body and hence cannot be included under the subfamilies Cyathocotylinse, Brauninise and Strigeinm. The absence of both, clear demarcation into anterior and posterior part of the body and the union of the lateral margin of the forebody as also the presence of a th ick and short posterior body, exclude it from the subfamily Polycotylinae.

The forro, however, resembles the genus Pharyngostomum Ciurea, 1922, of the subfamily Alariinse in having a short and broad posterior body, the absence of united margin of the forebody, in the extension of the vitella¡ up to oesophagus, in the nature of the intestinal cseca, position of the ovary, and a reduced aeetabulum. Although the course of uterus could not be traced in the specimen, it seems probable that the uterus may enter the holdfast organ as it generally extends beyond the ovary which in the present case touches the holdfast organ.

But the present forro differs from the described species of Pharyngo- stomum in having a posterior extension of the vitellaria, in the shape of the holdfast organ and in the almost entire margin of the testes. Ir is, therefore, being described a s a new species.

Key for the identi¡ of the Strigeidse included here :

1. Cirrus sac absent . .

Ckrus sac present . .

2. Uterus does not enter the holdfast organ . . . . . . . .

Uterus enters the holdfast organ . .

3. Lateral suckers present . . . .

Lateral suckers absent . . . .

4. Body not clearly divided into two parts ; adhesive gland absent ; testes dumb-bell shaped . . . .

B5a

. . 2

. . Cyathocotyle catvusi.

3

Pharyngostomum bagulum.

4

Neodiplostomum (see ~o. 5).

Neoalaria dumbdlar


Body elearly divided into two parts ; adhesive gland present . . . .

5. Vitella¡ extend up to the posterior end of the hind body ; anterior testes broad and elongated ..

Vitella¡ extend only up to the anterior third of the hind body; anterior testis pear-shaped ..

Proalaria alcedemis.

Neodiplostomum dilac~vcum.

Neodiploslomum tytense.

Family Schistosomidr Looss, 1899

(Syn. Schistosomatidm Poche, 1907 ; Bilharz¡ Odhner, 1912)

The author has already elaborated on this family in a reccnt paper in the pages of these Proceedings* and has given a detailed history of the family.

Diagnosis of the family Schistosomidce.--

Trematoda.--Sexes separate, pharynx absent ; oesophagus short, terminating posteriorly in a bifurcation to forro intest inal branches or cmca which join caudally at the cmcal union to forro a single, slender intestinal cmcum terminating near the posterior end of the body. Suckers present or absent ; acetabulum, when preseng, cephalad of the genital pore. Body of male may be widened caudad of the acetabulum and have the sides incurved ventrally, forming a gynaecophoric canal in which the female lies. Tcstes consist of four or more follicles. Cirrus pouch present of absent. I:emale more slender than male. Ovary elongate, sometimes spirally curved, and lying cephalad of the caecal union. Laurer's canal present oz absent. Vitellaria extensive, extending from the distal pole of the ovary to the posterior end of the body. Parasitic in the blood-vessels of birds and mammals.

The family Schistosomidae is divided into two subfamilies : - -

!. Schistosominm Stiles and Hassall, 1898.

2. Bilharziellinae Price, 1929.

Key to the subfamilies of the family Schistosomidm :

Cmcal union eaudad of equator of the body . . . . . . . . Schistosomince.

Caecal union cephalad of equator of the body . . . . . . Bgharziellince.

* Vide .Proc. Ind. Acad. Sci., 1937, 6, No. 5, 274-83.

s i~~ Helmb~#~olo2y 18,5

Subfa.mil.y Schist~~somin(e Stiles and ~assall, 1898

Dia.gn,~sis ..... Schistosomid,~'.~Males flattened and with the tateral edges of the body infl~lded ventrally to forro a gyn~cophoric canal. Suckers l~rt~sent. Intest inal cmca lo~~g, usually unifing eaudad of the equator oi the body ; common cmcmu relatively short. '/'estes situated in the anterior of posterior hatf of the body, always cephalad of the cmeal uniml, t~emales slender, thrcad-like, either longer of shorter thau the males. Uterus ustmlly cm~tains nmny eggs.

Type genus-~Schis~osoma Weinland, 1858.

Genus Ornithobilharzia Odhner, 1912

Diagnosis ...... SchŸ162162 shorter tllan male. Male with well- devclol,ed gymccoplmric camd, formed by ah in%lcling of the lateral edges of the body. Stlckers present. Cuticte eovered with spines. Digestive tŸ similar to that of Schistosoma ; intestinM cmca long, showing a tel~deney to forro several anastomoses before finally uniting to forro the common cmctml. Testes numerous, 6(1 of more, commeneing a short distanee c'mdad of aeetabulum, and extending into posterior hall of t11e body. Cirrus pouch rudimentary of absent. Seminal vesicle free in the parenchyma; prostate absent. Genital pote sit~mted immediately betfind the aeetabulum. Female elongate, slender and flatteued. Ovary elongated, loosely of t ightly coiled, alld situated in anterior ttfird of the body. Vitellaria extensive, occupyillg about two-thirds of body length. Laurer's ca~mI present (at least in some species). �8 slmrt, contain~ng only one egg at a time.

Type spccies~OrnithobiIharzia inter~nedia.

Ornithobilharzia si). Gogate, 1931.

Lellgt, h 9 ,5 m m . , brettdt,h .488 m m . ~p[rms a, ntl tubc~reh~s pres~;nt on tht~ bocly. ()r~~l suck~~r ,196 m m , irt di turmter , Vcmtral sueker peduncuh~ted, .472 mm. • ,417 m m . Intx~stin~~l cmc~~ lm~g, joŸ to f om~ a o o m m o n cmcum .,~2 mm,. f~'om t h e posterir end . ~l'estcs a b o u t ~/0, oit, c u b m i 'eebly d eveloped. Cirrtm pouoh r u d i m e n t a r y , t)ontcriol: t:,o vcn t r a [ ~uckcr. (t.ym~~cophoric c~ma[ wc[1 devoloped.

ttost.--Dendrocyg~a javanica (Clotted blood).

Locality.~Rangoon.

RemarŸ237 autiaor of this species found only two immature ~orms and gives no diagram of the~ specimen~, In the absence of an adequate account, the w¡ is unable to comment on this forro.

I76

186 M a k u n d Behar i L a l

Subfamily Bilharzielli~zce Piice, 1929 eme~Med

Schistosomidce.--Suckers present or absent. Gyn~ecopiioric canal absent of imperfectly formed or sometimes well developed. Pa i red in tes t inal cmea short uniting cephalad of the middle of the b o d y , common cmcum iong, with of without lateral denddt ic branches. Testes numerous and si tuated bdi ind the ctecai union along the course of tt le conlmon cmeum. Uterus short, containing a single egg.

Type genus--Bilharzidla Looss, 1899.

Key for the identification of the AvŸ Bilharziellina~ reported from India :

Gynmcophoric canal well developed ; ventral sucker present . . . . Chi~dz,utc~ i~zdica.

Gynmcophorie canal absent ; ven t ra l sueker absent . . . . Gigantobilharzia egreta.

Genus Chinhuta Lal, 1937

3/fale with a well-developed gyn~ecophoric canal, ex tend ing from a little behind the oral sucker up to the posterior end ; female with flattened body and smaller than the male. Suckers present. Oesohagus provided with unicellular oesophageal glands. The two intestinal cmca unite a little in front of the middle of body; common cmcum long, without lateral branches but provided with angular thickenings of its wall. Testes vary between ?0-80 in number and extend from the cmcal union to tlie posterior end of the animal. Cirrus pouch well developed, enclosing a part of vesŸ cula seminalis, prostate gland-cells, pars prostatica and the cirrus. The terminal end of the cirrus sac is slightly bent to the left side. The nlale genital pore lies slightly to the left of the median line near the middle of the body. The ovary is elongated and sinuous, situated at the levcl of the cmcal union. LTterus short and straight. The female genital openŸ is a little behind the aeetabulum. Vitellaria situated on the sides of the common cmcum in distinct follicles. A vitelline reservoir is present.

Type species--Chi~huta i~zdic~.

Chinhuta indica Lal, 1937

Male.--2.95 mm. long, .375 mm. broad ; l a te ra l edges of body rolled inwards to form a deep groove. Oral sucker weak "i mm. ii/ diameteŸ Ventral sucker muscular .15 Inm~ in diameter. Oesophagus .375 mm. long, with unicellular oesophageal glands. Intestinal cmca j oin together

gfudies i~~ Z-f eZ~~2i~~tAofo g'j, 187

in the pre-eqtiatorial region at a d is tante of .625 mm. behind the ventral sucker and 1-275 mm. from the anterior end. Testes 70-80, oval bodies of variable size; largest .12,5 mm. • -0~ mm.; smallest .05 mm. • �9 025 mm. VesŸ seminalis divided into two lobes ; larger �9 117 mm. x. �9 0875 mm. and smaller .087 mm. • .07 mm.

Female.--! .8 mm. long, .19 mm. broad. Oral sucker .0~ mm. in dia- nleter. Ventral sueker .075 mm. in diameter. Oesophagus .225 mm. long with unicellular oesophageal glands. Intes t inal c~eca ioin together at a dis~cance of .675 mml. from the anterior end. Ovary 1tes -27 mm. behind ventral sucker and meas~lres -155 mm. • -05 mm. [No eggs were found in the uterus. Vitellaria dist inet and large follicles posterior to ovary.

Host.--Nettio~ crecca (Main blood-vesels and interna1 organs).

Locality.--Chinhut, Luckllow.

Genes Giga~ztobilharzia Odhner, 1910

Diagrwsis of the gem~s Gigantobilharzia Od]zr162 1 9 i 0 . ~

BiZharzieZZi.m~.--Fenlale cylindrical and shorter than the somewhat flattened male. Posterior ex t remi ty of both sexes provided with lateral lobe-like projeetions, Cutiele wi thont spines of tubercles. Oral sucker present of absent. Ventral sucker always absellt. Gynmcophoric canal absent of redueed to a short groove, s i tuated in anterior par t of the body. Alimentary canal similar to t ha t of BiZharziella. "restes or iginate behind intestinal union and extend almost up to the posterior end of the body. Cirrus pouch present. Genital pote s i tuated between the intest inal bifurca- t ion ana rennion. Ovary moderate ly long and spiral. Vitell ine follicles occupy about nine-tenths of the body Iength. Uterus short, containing a

few eggs.

Type species--Gigantobilharzia aco�91 Odhner, 1910.

Giga~r egre~a Lal, 1987

Male.--38.85 mm. long, .275 mm. broad. No oral oŸ ventral sucker. Oesophagus 1.2 mm. long. Csecal unioll takes place .4 mm. behind c~eeal bifurcation. Testes more than 600, ¡ 35-6 mm. of body ; size varia- able, largest -15 mm. x .1 mm. ancl smallest -075 mm. • .05 mm. VesŸ seminaIis .2 mm. in length.

FemMe.--Not obtained.

Hos~.--Bu[bulcus ibis coroma~dus (Renal vein) .

Locality.--Lucknow.


Remarks.--Only two more spedes of this genus ate recorded G. acotylea by Odhner (1910) from Sweden and G. monacotylea by Szidat (193(/) from East Prussia.

I-rOST- PARASITE RELATIONSHIP

From a review of the habitat of trematodos in hosts, it is clear that the same species of trematode may sometimes infeet more than one host. I t is now a considered fact tha t the host-specificity theory of older hehnintho- logists is no more ah axiom. Cases have been reported of non-specificity of hosts in the trematodes affecting vafious groups of vertebrates. The writer (Lal, 1937) has already emphasised the non-specificity of hosts in trematodes and also referred to a work of similar nature on cestodes by Meggitt (1934). Even in the case of Avian Trematodes there appears to be no specificity of hosts in several cases. During bis own investigations on the subject, the writer has come across examples where one species of trematode recorded from India infects more than one host. Thus, Platynosomum acuminatum has been described from Kestrel by :Nicoll (1915) and from Crow by Bhalerao (1926). Echinochasmus bagulai has been obtained by Verma (1935) Ÿ two different genera of the I-Ierons. Prosthogonimus cuneatus which is now reported from Acridotheres tristis has already been described from a Iarge number of birds from Asia, Africa and Europe. Echinostoma revdutum has also been reported from an exeeedingly large number of different hosts. I r is, thus, evident that the host-speci¡ hypothesis does not strictly hold good for at least the AvŸ Trematodes.

COPULATION IN TREI~TODES

The writer had no opportunity of making observations on the de'velop- ment of the Avian Trematodes deseribed in the present communication, but he has been able to record a case of copulation in the genus Levinseniella.

The process of copulation has been observed in a lazge number of trenla- todes, both ecto-parasitic and endo-parasitic. PrevŸ records reveal that the time of copulation varŸ with the individuals, and climatic conditions do not affect it speeially in those cases where the host happens to be a warm-blooded anima1. Severa1 different methods of copulation have been observed from time to time in the case of Digenetic Trematodes. The fertilization is affected either through cross-copulation of self-copulation. The cross-copulation may be mutual of reciprocal, e.g., in Prosotocus (Looss, !885), Dicroccelium (Linstow, 1890) and Nematostrigea (Nitzsch, 1819), where the terminal portion of the uterus of the met ra te rm serves as the vagina; or it may be one-sided copulation, e.g., in -Paragonimus (Kerbert,

Studies iu Hefmz'uiholos 189

!881), Liolo;he (Cohn, 1902) and Collyridum (Jegan, 1916) in whieh the I, aurer's canal is said to function as vagina. In the case of self-copulation which also has been observed in sorne trematodes, e.g., in AŸ237 (Odhner, 191I) there is the introduction of the cirrus of the individual into its own metraterm.

o s ~

~5 mm.

I~z~. 19

Fro. 19.

Frc~. 20.

~'.$.

,. c~r,

m . 9 . a .

o.$.

gr c~r.

,2 mm~

Fio , 20 L e v i n s e n i e l l a ing@a--eopal ,~ t ing worms .

Zevi, ns~r~iella i n d i c a ~ s h o w i n g r e t r a c t e d cirri and grooves .

In the case of Levinseniella Ÿ (Fig. 19) in which mutual copulation takes place, a pair of individuals were obtained in sexual congress when the bursa fabricii of the Jack snipe, Gallinago galli~ula, was opened in normal salt solution. These trematodes being of small size, the fact of .their recovery in a copulating state could not be definitely ascertained till the peeuliarly shaped speck of tissue was sub]ected to examination under a low power of the mieroseope. The copulating individuals lŸ side by side, with their anterior ends pointing in opposite directions. One individual 1les on its dorsal while the other on its ventral surfaee. The union takes place by the sides of the individuals which also part ial ly ove~lap each other. A complete contact between the couple in the affected region is aeeom- plished through the help of deep undulations in the neighbourhood of the cirrus. These undulations of the skin which take the forro of grooves and papillm ate developed on the affected sides of the individuals in'the region of contact of their genital organs; and they appear to be temporary structures formed only during eopulation as they are entirely absent in other speeimens, t~aeh papilla has one of two farrows on its side and the papilla


of one individual fits into the groove of the other and thus affords a firm hold for the trematodes. The cirrus of these individuals bears a large number of minute spines and those also help in mainta ining the contact during copulation. In the present case the trematodes were so strongly attached to each other that the pressure of the cover-glass did not seem to have any effect.

The cirrus of eaeh individual which is a thick cylindrical body apparently lengthens out very much and is thrust into the metraterm of the other. The male genital atrium lies at the base of the cirrus and during eopulation its cavity, which normally contains the cirrus, is very much reduced by the contraction of its muscular thickenings. This exerts a pressure on the atrium and the eirrus is thereby pushed out.

An attempt was made to fix the specimens in the copulat ing state but the couple separated out and onty the~r half retracted cirri (t:ig. 20) could be observed in the fixed specimens. These observations on the copulation were made for about 20 minutes durŸ wMch a camera lucida diagram was made. Sinee the parasites were recovered already in a copulating state, the author is unable to say anything definite regarding the duration of

eopulation. ~EOGRApHICAL DISTRIBUTION

With regard to the distribution of the various genera of AvaŸ Trema- todes of India in relation to those found in other parts of the world, ir may be mentioned that the distribution seems to be governed by the migration factor. The birds, most of which are migrants, foreign as well as local, can carry with them parasites which are peculiar to certain countries. These cases of hosts as carriers are very common. Certain cases of peculiar distribution are worth mentioning. The genus _Parorchis was reported from Northumberland coast, St. Andrews Millport, and Ameriea. I r has now been obtained in India. Similarly the genus GiganŸ whieh was reported only from Sweden and East Prussia is now being reported from India. The occurrence of Perorchis, Gig~~r and eertain other trematodes in India throws important light on the effect of the migratory hosts on the spread of helminthie infections. Some of these trematodes whieh were more or less endemie in particular eountries have uow been discovered in I n d i a , andas ah explanation of this peculiar distribution ir may be sffid that the parasites were brought to this eountry by the migratory birds. Some of these parasites would in course of t ime produce the eggs whieh may hatch out into miracidia and may infect the snails of a similar or closely similar variety as the original intermediate host. I r is not

S~udies i~a Hdminffm/ogy 191

s u r p r i s i n g t h a t s o m e of t h e s e m i r a c i d i a m a y b e c o m e a d a p t e d t o t h e i r n e w

e n v i ~ o n m e n t a n d w o u l d t h u s p r o d u c e c e r e a r i a w h i e h m a y i n f e c t n e w h o s t s .

T h e u s u a l b a r r i e r s i n c a s e of l a n d a n d s e a a n i m a l s do n o t c o n t r o l t h e i n Ÿ

in c a s e o f b i r d s w h i c h c a n f iy a c r o s s c o n ~ i n e n t s a n d o c e a n s a n d w a n d e r t o

d i s t a n t p l a c e s .

] ~ E F E R E N C E S

B~er, ,1. G.

Barker, F. D.

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13haler~o, G. D.

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Braun, ~r

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Cameron, T. W. 1~.

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�9 . " A new m o n o s t o m e t r e m a t o d e paras i t i c in the musk- r~t w i th ~ key to the paras i tes of the American m u s k - r a t , " Trans. Amer. Micros. Soc., 191($, 35.

�9 . " Nobes on thrc.e l[t t le k n o w n T t~matodes , " Ann. Mag. Nat. H'ist., Londm�91 1927, Ser. IX , 13.

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�9 " On the t r e m a t o d e s of the cligestive t r ac t of a common Ind i~n ~rog , Rana t$grina wi'th a descr ipt ion of Centrovittts adelphi, n.g., n .sp . ," Paras~t., 1926, 18.

. . " On two new monos tomes (Trematoda) f rom avŸ hosts in /3rit.ish Ind ia , " I',~d. Jour. ver. Sci. and Animal Husb., 1935, 5, P a r t 1.

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. . " On Crepidoston�91 far~oni, s, a d i s tomate p~ras[~c o[ t he t r o u t ~nd grayl ing. 1 - - T h e l i fe-his tory," 2aras~& 1927, 29.

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�9 " The in%ernal paras i tes o~ domest ic animalsr London, 19~4.


Chatterji, R, C.

Chen, I I . T.

Ciurea , I.

Cohn, L.

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D�91 ]~.

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�9 " O n Ÿ t r e m a t o d e pg ras i t e s of ,% l%angoon s i luroid fish, Clarias b~�91 B~�91 Acad. Sci., U.P., fndi~~, 1933.

�9 " A n n o t a t e d l i s t o f t h e t l e l m i n t h s r e c o r d e d f r o m d o m e s t i - c ~ t e d a n i m a l s of t l u r m , % " _Proc. N(~t. Acad. Sci., Incli0~, 193S, 8, Part 4.

�9 " A s t u d y of I t a p l o r c h i n m , " Pa,rasit., 193(4 28.

. . " H e t e r o p h y i d e s de la f a u n e p a r a s i t M r e de R o u m a n i e , "

IMd., 192.~, 16.

. . " C o n ~ r i b u t i o n s s l ' › m o r p h o l o g i q a e e t b [ o l o g i q u e de q u e l q u e s s t r i g e i d e s des o i s e a u x I c h t y o p h a g e s de la f a u n e de R o m n ~ n i e , " Arch. I'ath. Exper. Micro., Roumn., 1930, T. I I I .

. . " I t e l m i n t h o l o g i s c h e m i t t e i l u n g e n - - I [ , " Arch. Naturg., 1904, :t.

. . " On a t r e m a t , ode , Ityogonimus Iorum, w i t h n o t e s on t h e o c c u r r e n c e of o t h e r t r e m a t o d e p a r a s i t e s of Talpa e uro2zJaea in t h e A b e r y s t w y t h a t e a , " Par~c~it., 1932, 24.

. . " On ~ c o l l e c t i o n of P a r a m p h i s t o m i d m f r o m M a l a y a wibh r e v i s i o n of t h e g e n e r a Paramphistomum a n d Gastrothy~ax," Ibid., 1936, 28.

�9 . " I ) i e E c h i n o s t o m i d e n d e r V 6 g e ] , " Inaug. 1)iss., 1 9 0 9 ; also Zool. Jahrb., S u2pI., 1910, 12.

. . " R e v i s i o n des ' H e r n i s t o m e s ' " , BMl. Soc. Ne~zch. Sc. Natur., 1931, T. 56.

. . " R e v i s i o n des ' H e m i s t o m e s ' , " Rey, Sui. de Zool., 1933, T. 40.

�9 " On a n e w spec i e s of t h e T r e m a t o d e g e n u s NotocotyIus," Ann. Ma g. Nat. Hi, st., 1931, 7, Ser . lO.

. . " T h e e x o r e t o r y s y s t e m in l ) i g e n e a . I I ~ O b s e r v a t i o n s o n t h e e x c r e t o r y s y s t e m in d i s t o m e c e r c a r i ra ,"

BioI. Bull., 1919 a.

. . " T h e e x c r e t o r y s y s t e m in Digene '~ . I I I - - N o t e s on t h e e x c r e t o r y s y s t e m i a a m o n o s t o m e l a r v a , " lbid., 1919,

36.

. . " C r i t e r i a fin' t h e d i f f e r e n t i a ~ i o n of S c h i s b o s 0 m e l a r v m , "

Jm~r. Paras~,t., 1920, 6.

�9 " T h e e x c r e t o r y s y s t e m in ] ) i g e n e a . I V , " Parasit., 1921,

18.

. . " L a r v a l f lukes f r o m @ e o r g i a , " Trans. Amer. Micros. Soc., 1921, 40.

�9 " N o t e s o n t h e e x c r e b o r y s y s t e m i n AsŸ con- cMola," tM~l., 1922 a, 4,i .

�9 " N o t e s on l a r v a l f lukes f r o m C h i n a , " Pctrasit., 1922 b, 14.

. . " H u m & n H e l m i n t h o l o g y , " L o n d o n , 1930.

Faus t , E. C., a n d 2~ishigori, M.

F u h r m a n n , O,

Gcgafie, ]3. S.

H a r r a h , E. C.

Har she , K . 1~.

Holl , ]~~. J .

J agersk io ld , L. A.

Jegen , G.

J o h n s t o n , S. ~'.

J o y e u x , Ch., ar~d Baer , J . G.

Kerbe rG C. . .

Khal i l , M. . .

K h a n , M. ~I.

K o b a y a s h i , t i .

Kossack , W.

Studies i'~ f]ekninthology 193

" A P re l im[na ry R e p o r t o n the Life-Cycies of two new H e t e r o p h y i d Flukes , e t c . , " China .~Ied. Jo~r., 192£ 39.

" l~eue T r e m a t o d e n , " CentraI&I. Bakt. Par., 1904, Abt . 1, Orig. 37.

" Notes he lmin thologiques s u i s e s - - I I , " Rey. ,gu~sse de Zool., 1919, 27, No. 2.

" T r e m a t o d a " in K ª Handbuzh der Zoologie, 1928.

" On Tremaflodes f rom wi td ducks in R a n g o o n , " Rec. Ind. Mus., 1934, 36.

" Iqorth Amer ican Monos tomes , " Ill. BioI. Monogr., 1922, l , No. 3.

" On CatatroIgiS oriental@," AlId. Univ. Stud., 1932, 8.

" New t r e m a t o d e s f rom t h e newt , Tritur~ts viridescens," Jour. Helminth., 1928, 4, No. 3.

" Kleine ]3eitrage zur I ( e n n t n i s der Voge l t r ema toden , " CentrL f. Bakt., 1909, 1, Abt . Orig., 48, H e f t 3.

" Collyriclum faba. E i n P a r a s i t der Singvogel, sein B~u und seine Lebensgesch ich te , " Ze~tsch.f~�91 ZooI., 1916, I17.

" On some Queensland trematodes, with anatomicM observ~t ions ancl clescription of new species and gener~," Quart. Jour. "Micros. Sci., 1913, 59.

" Zoogeography o~ Aus~ral ian Trema todes and Cestodes," ~Ied. Jo~zr. Austr., Sydney, 1914, 12.

" T r e m a t o d e s of A u s t r a l i a n :Birds," Jour. Proc. Roy. Soc., Iqew South WMes, 1916, 50.

" Notes sur les 0yclocoel idm (Tr› Bull. de la Soc. Zool. de France, 1927, 52.

" Be i t r ag zur Kenn tn i s s der T r e m a t o d e n , " Arch. f ª 2dikrosk. Anat., 1881, 19.

" On ~ t r e m a t o d e f r o m t h e g~llbladder of Naja bungarus wi th ah emendmer~t of the genus Xenopharynx. Nicoll ," Jour. Helmi~zth., 1923, 1.

" On eight new species of genus Cyclocoelum Brandes f rom l~orth I n d i a n sn ipes , " Proc. U.P. Acad. Sci., 1935.

" ~ rev iew of Yapanese Cerc~ria," Mitt. Med. Acad., Keijo, 1922.

" N e u e D i s t o m e n , " Centralb. f~r Bakt. Para8~t. und Infelr191 1910, 56.

" lYeber Mo~cs~omiden, " Zool. Jahrb. Syst., 1911,

3i .

194 Makund Behari LaI

Lal, 5L B.

Lar~ue, G. R.

LaF~nan, E.

L e b o u r , ~,~. V. , ~ncl E I m h i r s t , 1%.

L e i p e r , 1%. T .

L i n t o n , E .

L i n ~ t o w , v o n . O.

. . " On t h e n l o r p h o l o g y of ~ n e w s p e c i e s o f m o n o s t ; o m e of t h e g e n u s Notocoty[us," Proc. Ind. Aead. SoL, 19:=15, 2, No . 5.

. . " A r e v i e w of t h e g e n u s Notocotylus, w i t h d e s c r i p t i o n of n e w t r e m ~ t o d e p s of Marecapenelope f r o m L u c k n o w , " �91 1935, 2, ~~o. 5.

. . " A r e v i e w o f t h e g e n u s .Paramonostomum w i t h d e s c r i p - t i o n of ~wo n e w s p e c i e s a n d r a m ~ r k s o n t h e g e n e r ~ of t h e s u b f a m i l y N o t o c o t y l i n m , " lbid., 1936, 3, N o . 1.

�9 " O n a n e w s p e c i e s of t h e g e n u s Parorchis f r o m Totanus h!]polei~cos w i t h c e r t a i n r e m a r k s o n t h e f a m i l y , E c h i n o s t o m i d a e , " Ibid., 1936, 4, N o . 1.

. . " O n a n e w g e n u s of t r e m a t o ( l e s of t h e s u b f a m i l y T y p h l o c o e l i n m f r o m t h e s h o v e l l e r d u c k , Spat~tla clypeata," Ib~d., 1936, 4, N o . 1.

�9 " :4 n e w s p e c i e s of t h e g e n u s ZevŸ191 f r o m t, h e j ~ c k s~~ii)e, Galli~~ago gedlinula," Ib~d., 1936, 4, :No. 2.

. . " S t ,ud ies o n t h e T r e m a ~ o d e p a r a s i L e s o f B Ÿ I . V a l u e of different , C h a r a c ~ e r s i n t h e C l a s s i f i c a t i o n of A v i a n T remaLo , � 91 Ibid., 1937, 5.

�9 " SLud ie s o n t h e T r e m a S o d e P a r a s i t e s of ] 3 i r c l s - - P a r t I I . iV[orphology ancl s y s t e m a ~ i c p o s i t i o n o f s o m e n e w b looc l - f l uke s of t h e f a m i l y S c h i s t o s c m i d m , Ibid., 1937, 6 No . 5.

. . " O n a n e w s p e c i e s of PsiIorchis." L i v r o ,Tub i l a r P r o f . T r a v a s s o s R i o de J a n e i r o , B r a z i l , 1938.

. . " S t u d i e s o n ~he ( h ' e m a t o d e F a m i l y S t r i g e i 4 m - - N o . I . _Pharyngostornum cordatttm. N o . I I . Ta~sonomy". TraThs. Amer. Micros. Soc., 1926, 45, 1 - 1 9 .

�9 " s fringiHce n . s p . " Archiv. wiss . pratct. Vet. 1923 (in R u s s i a n ) .

" A c o n t r i b u t i o n t o w a r d s t h e l i f e - h i s t o r y of Parorehi~ acanthus Nico l l , a T r e m a t o d e i n t h e ] K e r r i n g G u l l , " Jour. Mar. BioL Assn., 1 9 1 9 - 2 1 , 1 2 .

. . " l%epor t o n t h e r e s u l t s o f t h e B i l h a r z i a M i s s i o u i n E g y p t " , Jour. Soy. Army. Med. Corps, 1 9 1 5 - 1 8 .

. . " M e d i c a l H e l m i n t h o l o g y : A r e v i e w . " Trop. Diseases BulI., 1923, 20.

�9 " iYotes o n a v i v i p a r o u s d i s t o m e , " Proc. U.S. Nat. Mus. , 1914, 4B.

. . " l ~ o t e s o n T r e m a t o ~ e pa r ,~ s i t e s of ] 3 i r d s , " Ibtd., 1928, 73.

. . " lYeber d e n B a u u n d d i e E n t w i c k l u n g d e s D4stomum cylindraceum Z e d . , " Arch. f . Mi~r. Anat. , 1890, 36.

Sludies in Hdmin/~�91 195

L i n s t o w , r o n . O. . . " H e h n i n t h s f r o m t h e c o l l e c t i o n of C o l o m b o ~ f u s e u m , " Spolia Zeylan., 1906, 3.

L o o s s , A . . . . " l~e[ t r~ge zu r K e n n t n i s d e r T r e m a t o d e n , " Zeitsch. fur ZooI., 1885, 41.

. . " \ V e i t e r e B e i t r g g e z u r K e n n t n i s d e r T r e m a t o d e n - f a u n a A e g $ p L e n s , " Zoo�91 Jahrb., 1899.

. . " On some p a r a s i t e s in t h e M u s e u m of t h e Schoo l of T r o p i c a l M e d i c i n e , L i v e r p o o l , " Ann. Trop. Med. t~arasit., 1907, i .

. . " N o t i z e n z u r H e l m i n t h o l o g i e A e g y p t e n s , " ZbI. Bakt., ] 907, 43.

L ª M. . . " P a r ~ s i t i s c h e P l a t t w ª T r e m a t o d e s , " in D~e Sª 1,'auna De~~tschlands, 1919, H e f t i 7 .

~ [ a n t e r , l i . ~V., a n d " S o m c M o n o s t o m e s f r o m ~qorth A m e r i c a n B i r d s , " W i ] l i a m s , O . L . Eran& Amer. Mic. Soc., 1928, 47.

~r F.J. . . " T h e o r y of h o s t - s p e c i f i c i t y a s a p p t i e d t o c e s t o d e s , " Ann. Trop. Med. and Para~.it:, 1934, 28.

M e h r a , H . ]] . . . . " A n e w g e n u s of t h e f a m i l y L e p o d e r m a t i d e f r o m a •ortoise, w i t h a s y s t c m a t i c d i s c u s s i o n a n d c l a s s i ¡ t i o n of t h e f a m i l y , " 2arasit., 1931, 23.

. . " On •wo n e w spec i e s of t h e g e n u s Astiotrema Looss , belonging to the f~mily Lepode~mat~dm, Ibid., 1931, 23 :

. . " l~ew b l o o d ¡ of t h e f a m i l y Sp i ro rch idse , e tc . ; P ~ r t l i , " BuIl. U.P. Acad. Sci., 1934, 3.

. . " N c w t r e m ~ t o d e s of t h e f a m i l y L e c i t h o d e n d r f i d ~ e w i t h a d i s c u s s i o n o n t h e c l ~ s s i f i c a t i o n of t h e f a m i l y , " Proc. U.P. Acad. Sc i . , ' 1935 , 5.

_ _ . . " C e r t a i n n e w a n d a l r e a A y k n o w n d i s t o m e s of t h e f ~ m i ] y L e p o d e r m a t i d ~ e , " Zeitsch. f ª Parasit., 1937, 9.

1M[ehra, I I . R . , a n d I~egi , P . S. " On a n e w t r e m a t o d e , Eremiorct~is ranarum f rom t h e c o m m o n Ind~~n f rog Rana tigrina," Parasit., 1926,

Mi l l e r , H . 1~.

M o g h e , M. A .

l~ [o r i sh i t a , K.

Montgomery, II. E.

~$.

. . " C o m p a r ~ t i v e stud[es on F u r c o c e r c o u s Ce rca r im , " Itl. Biol. Monogr., 1920, 10.

�9 " T w o n e w spe c i e s of T r e m a t o d e s f r o m ~n I n d i a n Ruf f (Philomachus ~ugnax, G r a y ) , " Parasit., 1932, 24.

. . " N o t e s on t w o n e w M o n o s t o m e s w i t h r u 4 i m e n t a r y v e n t r a l s u c k e r s , " Jouv. Paras~t., 1924, 10.

. . " S o m e A v Ÿ T r e m a t o 4 e s f r o m J a p a n , s p e c i a l l y f r o m F o r m o s ~ , w i t h r e f e r e n c e l i s t of a l l k n o w n J a p a n e s e s p e c i e s , " Annot. ZooI. Japan, 1929, 12.

. . " O b s e r v ~ t i o n s o n b i l h a r z i o s i s ~ m o n g a n i m a l s in I n d i a - - I , " Jouvn. Trop. Ver. Sc., C a l c u t t a , 1906, 1.


.Morgan, D . O . . .

3[Lmller, 5. F. , ancl Cleave, H. J. Van

Nicoll, W. . .

N i t z sch , Ch. L.

Noble, A. :E.

Odtmer, T.

Oiso, T.

Pande, ]9. P.

" Stud[es on t h e f~rmily OpisbhorchiŸ e t c . , " Jmtr. Hebninth., 1927, 5.

" Parasi~es of t h e One ida lake fi~he~, pbs. l - t . " h~,z~~. IVild Life Amz., 1932-31, 3.

" S~u(ties on t h e sf ruc~ure tm~l ch:,,ssi[ica.f&m o~ the d igenet ic $remaf, odes , " Quart. Jota'. illicros. Nci., 1909, 53.

" Some new a n d l i t t le k n o w n T r e m a ~ o d e s , " .Ann,..~[a,{I. Nat. Hist., ] 906, 17.

" Parorehis aea~~thus, t h a f.ype of ,% new genus of t,roma,- todes , " Quart. Jota'. Micro. Sci., 1907, 51.

" On the E n t o z o a of fishes f r o m tiro F i r t h cal C lyde , " Parasit., 1910, 3.

" On two new t rem~~ode p~wa~ites f r o m the I m t b m Cobr~," Proc. Zool. Soc., London , 1912.

" TremŸ p~r~si~es of Nor~h Q u e e n s h m d ~ P a , rt, I , " Parasit., 1914, 6.

" A new Liver - f luke (Platy~�91 acT,minatam) f rom the K e s t r e l , " Proc. Zool. Soc., London , 1915.

" The Bionomics of H e l m Ÿ T rans. So(;. Trop. Med. t tyg..London, 1910, 3.

" .4. referenca list of the t r e m u t o d e p~~r~si~es of Bl'it;ish b i rds , " BarasiL, 1923, 15.

" Ar t ike l A m p h i s t o m ~ , " Allg. Encyclop. d. IViss. u. Kª 1819.

" Two new T r e m a t o d e s f r o m the A m e r i c a n Coo t , " Trans. Amer. Micros. Soc,., 1933, 59,.

" New A v i a n T r e m a t o d e s of the Genus Neodiplostamttm," Jo~�91 Parasit., 1936, 22.

" Z u m Nat, ª S y s t e m der Digenen Trema~od(m, I V , " ZooI. .4~z. , 1911, 38.

Results of the Swedish Zoological Expedition to E~ypt, and White Nile, 1911, 4.

" Z u m N a t ª S y s t e m der Digenen T r e m t r t o d o n V I , " Zool. Anz. , 1913, 42.

" On a new species of A v i t m Schistosom.~ ,~nd i a v e s t i g a - t ions of i t s l i f e -h i s to ry , " Taiwan Igakkzoat Zasskt, 1927.

" Oa two new spacies of the Genus CephalojonŸ .BulL Acad. ScL, Allah~.bad, I932, 2.

" Con t r i ba t i ons t o t he dig~n~Sic t rem~toc les of t h e Microchi rop te r~ of N o r t h e r a Inclh~," Proc. U.P. Acad. Sci., 1935, 5.

P a n d e , B. P .

P a t w a r d h a n , S. S .

P e r M n s , M.

P h a d k e , V. 1~., a n d G u ] a t i , A. N.

P o c h e , F .

P r a t t , H . S .

Pr ' ice , E . W .

B, ansom. B. I I .

S a n 4 g r o u n c t , 3". l i .

Sewel l , R . / 3 . S .

S i n h a , B . ]3.

S k r ] a b i n , K . I .

- - n n d U 4 i n z e w , A . N .

S o p a r k a r , i~ . B . . .

S p r e t m , (3. E . W . . .

Studies in/-felmh~tho/ogy 197

�9 . " A n e w s t r i g e i d T r e m a t o d e of t h e g e n u s CrassipMata f r o m a h I n d i a n K i n g f i s h e r , " P'roc. Nat. Acad. Sc i , I n d i a , 1938, 8, Parfl 4.

. . " T h r e e N e w Spec ies of T r e m a t o d e s f r o m B i r d s , " Proe. Ind. Acad. Sci., 1935, 2.

�9 . " A r e v [ e w of Te lo rch i ina3 , " Paras~t., 1928, 20.

" The ( l e s c r i p t i o n of ~ n~w F l u k e f o u n d in t h e I n d i a n H o u s e - C r o w , " BulI. Imp. Inst. Ayric. Res., P u s a , 1930, 203.

�9 " D a s S y s t e m de r Plat.cdaria,"Arch. Naturgesch., 1925, 2.

�9 . " S y n o p s i s o[ t h e N o r b h A m e r i c a n i n v e r t e b r a t e s , " Amer. Nat., 1902.

�9 . " A s y n o p s i s o[ t h e t r e m a t o d e f a m i l y Sch i s tosonUdm w i t h 4 e s c r i p t i o n of n e w g e n e r a a n d spec i e s , " Proc. U.S. Nat. Mus., 1929, 75.

�9 " A n e w spec ies of t r e m ~ t o d e of t h e f a m i l y H e t e r o p h y i d m , w i t h a n o t e on t h e g e n u s ApophaIl~s a n d r e l a t e d g e n e r a , " Ibfd., 1921, 79.

. . " The T r e m a t o d e Pa ra sŸ es of M ~ ¡ M ~ m m a l s , " Ib�91 1932, 81.

�9 " A n e w h e t e r o p h y i d t r e m ~ t o d e o f t h e g e n ª Ascocoty~e," P,roc. HeImint.h. Soc. Washington, 1936, 3.

. . " S y n o p s i s of t h e t r e m ~ t o d e f ~ m i l y I t e t e r o p h y d i s B w [ t h d e s c r i p t i o n s of a n e w g e n u s a n d t i r e n e w s p e e [ e s " Proc. U.S. Nat. Mus., 1920, 57.

. . " Nema~os~rigea serpe~.s, a, r ~ - d e s c r i p t i o n of HoIostomum serper~s N i t z s c h , " Paras�91 1934, 26.

. . " Cerca r im I n � 9 1 Ind. Jour. Med. Res., 1922, 10.

. . " T h e evohl t ior~ of t h e e x c r e t o r y s y s t e m in c e r r a j a g r o u p s of t h e F u r e o c e r c o u s c e r c a r i m , " l~ec. Ind. M u s . , 1930, 32.

. . " On t h e m o r p h o l o g y ,~n4 s y s t e m a t i c p o s i t i o n of Cephalogonimus m~�91 _4nn. Mc~g. Nat. Hist., 1932, 10.

. . " Trachophih�91 sisov$ n . g . n . s p . , E i n B e i t r a g zu r Sys t e - m ~ t i c d e r G ~ t t u n g Typhlocoe~um S t e s s i c h . , " Centrbl. f. I~akt. u~d. _Parasit., 1913, 69.

. . " V o g e l t r e m a t o d e n a u s l ~ u s s i c h T u r k e s t a n , " Zool. Jahrb. Syts., 1913, 35.

" Two n e w t r e m ~ t o ~ e s f r o m t h e b i H a r y d u c t s of B i r d s f r o m A l ~ n e n i a , " Jour. _Parasit., 1930.

" T h e eerc~ri~, of Schistosomum spindalis," Ind. Jour. Med. t~es., 1921, 9.

" L e h r b u c h d e r t t e l m i n t h o l o g i e , " B e r l i n , 1932.


Srivastava, I-I, D.

St i les , C. W. ,

Stoss ich , M.

S t u n k a r d , t L W.

�9 ~

aad. HassaH, A.

S t u n k a r d , I I . W. , a n d t I a v i l a n d , C. B.

S t u n k a r d , Id. ~V., a n d Cable, 1~,. 3I.

S t u n k a r d , H. W. , a n d D u n i h u e , F . W.

Sz ida t , L.

S z i d a t , L. , a, nd U r s u l a

T a n a b e , B.

T h a p a r , G. S., a n d L~], M. B.

Travassos, L.

�9 " Sgud ies o n ghe f a m i l y H e t e r o p h y i d m ~ P a . r f i I , " Proc. U.I ' . Acad. Sci,., 19a5, 4.

" O n a n e w speeies of Ca&drop~s f r o m a h I n d i a n fowl, Gallus ban,]civa mur(lhi," Ih[.g., I935 , 4.

" I n d e x - c a s of m e d i e a l a n d v e t e r i n a r y Z o o l o g y : " Hyf/. Lab. BuZl., 1908, 37.

. . " I1 Monostmmo?z .mutabLle ele su r f o r m e off iae ," BoII. Soc. Adr'iat. Sci. N(~t. Triest., 1908, 37.

�9 " T h e p a r ~ s i t i c w o r m s collecte& b y ghe .&mer ican M u s e u m of N a t u r a l t ] [ is tory E x p e d [ t [ o n to t h e B e l g t a n C o n g o � 9 1 BulI. Amor. Mus . Nat . Hi,st., 1929, 58.

. . " Furt~her obse rv~ t . ions o n ghe o c e u r r e n e e of ~na l o p e n - i n g s i n d [ g e n e t i c f, r e m a t o d e s , " Zei~sch. ff�91 Para- site,~d~., 1931.

. . " T h e l i f e - h i s t o r y of Cr!IpgoooiyZc Iinf]~a," Jmer. 2~forph. c~)td Ph~jsioI., 1930, 50.

" T r e m a t o d e s f r o m t h e r ag , " Amer. d~It~s. Novga~s., 1924, 1 2 6 .

" T h e l i f e -h i s~ory of l~arorchi.s a~,it~s," BioL Bul l . , 1932, 62.

" N o t e s oIx tremt~Lodes f r o m a L o n g [ s l a n d d u e k w [ t h d z s e r i p t i o n of a n e w speo ies , " Ib~cl., 1931, 60.

. . '~ G.igatdobilha.rzi~~ monacotylea, n . sp . , e i n n ~ u e r BIu5- pat 'as ig aus O s t - P r e u s s i s e h ah W a s s e r v S g e l n , " Zeitsch..f{�91 ParoziteMc., 1930, 2.

�9 " P a r a s i t e n a us S e e s c h w a l b e n . 1. U e b e r n e n e C y a t h o - e o t y l i d e n aus de r D a r m r o n Sternc~ hirundo u n d Sterna para&isea," Ibgd., 1936, 8.

. . " ]3e i t rage z u r K e n n t n i s s de r T r e m ~ t o d e n de r 2r s t o m i d e n G a t t u n g No~ocotyhts," Ze~grabl. Bakt. , 1933, 129.

. . " T h e l i f e - h i s t o r y of a n e w s c h i s t o s o m e , Schistosoma- t�91 path~ocopticum T a n a b e , found in expe r i - m e n t ~ ] l y i n f e c t e d m i c e , " Jo~~rn,. l='arasit., 1923, 9.

" On t h e lVforphology o[' a n e w g e n u s of Trcm~~ocl~~ p a r a s i t e f r o m the K i n g f i s h e r f r o m L u c k n o w , " .Proc. Ind . Acad. ScL, 1935, 2.

�9 " C o n t r Ÿ P a r a C o n h e c i m e n t o :Da l ~ a u n a H e l m i n t o - log ica Bras i l e i r~ , I X , " Arch. de Agric. e Med. Ver., N i c h t h e r o y , 1920, 4.

�9 " t l e p r i n g : I n f o r m a e o e s sobre a f a u n a h e l m i n t o l o g i c a de 3iatto Grosso , " FoU~a -&1ed. Ÿ de Janeiro, 1923, 3.

. . " Cyclocoel idm Br a z i l e i r o s , " Reimpresso do Braz i l Medico, 1925, t A n n . 25, 1.

. . " T r e m a t o d o e s N o v o s , " fbid. , 1935, -4reno 25, 1.

T u b a n g u i , M. A .

Vern l&, S. C.

V e v e r s , G. M.

V i r 1~. D .

W,~rd , H . 13.

W~~rct, t:I. B. , a n d I{ i r sch

W i t e r t b e r g , G.

~ V r i g h t , S .

Y�91 S.

S&dies iu Hehzi~~~holo gy 199

�9 �9 " Tremabocls P~r~%sites of P h i h p p i n e u Philipp. J. Sc L, 1928, 36.

�9 . " T r c m a t o d e P a r a s i t e s of P h i l i p p i n e V e r t e b r a t e s - - - V . F l u k e s f r o m b i rd s , " Ibig., 1932, 47.

�9 . " T r e m a t o d e P a r a s i t e s of P h i l i p p i n e V e r t e b r a t e s - - V I . D e s c r i p t i o n s of n e w species a n d C las s i f i ca t ion , " [bid., 1933, 52.

�9 . " S t u d i e s on t h e I n d i a n Spec ies of t h e Genus Echino- chasmus, P a r t I . a n d on ~n :~ll ied N e w @enus E29istho- chasmus," Z~roc. Ind. Acad. Sci., 1935, 1.

�9 �9 " N o t e s on t r e m ~ t o d e p a r a s t i e s of I n d i a n b i r d s - - P a r t I , " AIld. Uni. Studies, 1936, 12.

. . " A new s$r ige id p a r a s i t e of ~he rare genus Cyatho- cotyle," N~,tttre, Lonclon, 1936, 138, 3~f-35.

. . " 0 b s e r w ~ t i o n s in t h e genus Paragonimus B r a u n w i t h

re-r of P. compa.ch~s," Jour. Helmfnth., 1923, 1.

. . " N e w S t r i g e i d s f r o m I n d i a n B i r d s , " Proc. Nat. Acad. Sci., India, 1937, 7, No . 3.

� 9 " ~qote on t h e P ~ r a s i t e s of t h e L ~ k e F i s h - - 3 , " Trans. Aro. Micro. Soc., 1901, 31.

. . " On t h e s t r u c t u r e ~nd cl~ss i f ic~t ion of ~he p~,rasi~ic w o r m s , " Jour. ParasiL, I917, 4.

. . " Tremat , o d e , " in W a r d an~l W h i p p l e s ' s ]Fresh-water Biologff, N e w Y o r k , 1918.

. . " The spec ies of Paragoni~nus ancl t h e i r d i f fe ren t i a -

t i o n , " Ann. Trop. Med. and ParasŸ 1915, 9.

. . " Die T r e m a t o d e n der F ~ m i h e C y c l o c o e h d m , " ZooI. Jahrb. Abt. Syst., 1926, 52.

. . " N o t e s on Cye locoe l idm," Ann. Mag. Nat. Hist., 1928, 10.

. . " Stuct ies on Trem�91 f a m i l y t I e t e r o p h y i d m , " Ann. Trop. Med. Parasit., 1929, 23.

. . " C o r r e c t i o n s t o m y p~pe r on H e t e r o p h y i d m (1929),"

Ann. Mag. No�91 Hist., 1930, ~[0.

. . " N o t e s on t h e a a a t o m y of MicrophaIlus opac~.s," Trans. Amer. Micr. Soc., 1912, 31.

. . " S~udies on H e ] m i n t h s of : [ a p ~ a - - P a r ~ I , " Jap. Jour. Zool., 1933, 5.

. . " Stucl ies o n t I e l m i n ~ h F a u n a of ~ [ a p a n - - P a r t 3 , "

A v i £ T r e m a t o d e s , " Ibid., 193"~, 5.

" S t u d i e s on I t e l m i n t h F a u n a o2 5 ~ p a n ~ P a r ~ 5, " Ibid., 1935, 6.

. . " S t u d i e s o n ~ e l m i n S h F ~ u n ~ of Y a p a n - - P a r t 16.

T r e m a t o & e s of f i shes ," .Kyoto, 1937, 3.


~)~r.

c~r.

3.8.

COl.

e.bl.

e.eh.

C.2).

f .g.a. g.p.

gr.

h.o.

.~ .c.

1.8.

met.

L I S T OF A B B I ~ E V I A T I O N S U S E D I N TI~IE F I G U R E S

Bursa. m.9.a.

Cirrus. oes.

Cirrus sac. ~ot.

Collar. o.s.

E x c r e t o r y bladrler, ov.

E x c r e t o r y eh:~mbm', ph .

E x c r e t o r y pore. r.s.

Fem~r gen i t a l a t r i u m , tes.

Geni ta l t?ore, uf.

Groove . v.s.

I t o l d f a s t o rgan , ves. sera.

I n t e s t i n a l emca. vit . d.

L a t e r a l sucker , vit. gl.

M e t r a t e r m . vit. ves.

Male g e n i t a l a t r i u m .

O e s o p h a g u s .

( ) o t y p e .

Ora l sucker .

O v a r y .

P h a r y n x .

R e c e p t a c u l u m s~minis.

T e s t e s .

U te ru s .

V e n t r a l sucker .

Ves icul~ semina l i s .

Vi te l l i ae 4uc~.

Vi te l l ine g lands .

Vi te l l ine reservo i r .

0111-0200.pdf - Indian Academy of Sciences

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