Parasit. Hung. 6. 1973.

Trematodes of Bats in India *

Dr. István M A T S K Á S I

Zoological Department of the Hungarian Natural History Museum, Budapest

"Trematodes of Bats i n I n d i a " - Matskási, I . - P a r a s i t . Hung. 6. 77 -98 . 1973.

I n the course of h e l m i n t h o l o g i c a l i n v e s t i g a t i o n s of 278 bat specimens r e p r e s e n t i n g 36 species c o l l e c t e d i n I n d i a , 22 trematode species were i d e n t i f i e d . Three new species (Neohe- terophyes topali,Prosthodendrium robustum, Parabascus g h a t i ) are described. The paper submits the f a u n i s t i c a l data of the c o l l e c t e d trematodes and discusses some species of u n c e r t a i n taxonomical status or i n t e r e s t i n g from the taxonomic and systematic p o i n t s of view.

I n 1967, d r . Gy. TOPÁL, mammalogist of the Hungarian N a t u r a l H i s t o r y Museum, has been on a z o o l o g i c a l c o l l e c t i n g t r i p t o I n ­d i a . During h i s c o l l e c t i n g s he preserved the d i g e s t i v e systems of the c o l l e c t e d bats i n a l c o h o l , i n order t o f u r t h e r p a r a s i t o -l o g i c a l i n v e s t i g a t i o n s . The f i x e d d i g e s t i v e systems were d i s ­sected i n the P a r a s i t o l o g i c a l Laboratory of the Hungarian Natu­r a l H i s t o r y Museum. I should l i k e t o express my g r a t i t u d e t o Gy. TOPÁL f o r the m a t e r i a l of trematodes a l s o i n t h i s p l a c e .

The present paper submits the r e s u l t of sy s t e m a t i c a l i n v e s t i g a ­t i o n s conducted on trematodes obtained by d i s s e c t i o n of the bat g u t s . The bats i n f e c t e d w i t h trematodes o r i g i n a t e d from the f o l l o w i n g s i x d i s t r i c t s of I n d i a :

1 . Kashmir: Srinagar ( S r . ) - Bumzov Cave (Bu.)

The S c i e n t i f i c Results of Dr. Gy. TOPÁL's C o l l e c t i n g s i n I n d i a , Part 7 .

2. Western I n d i a : Bhaja Caves (Bha.) - Elephant a Caves (E— leph . ) E l l o r a Caves ( E l l . ) - K a r l a Caves (Ka.) - Maila­ble shwar (Ma.)

3. Central I n d i a : R i c c h a i ( R i . ) - Jabalpur (Ja.) 4 . Orissa: D a i t a r i (Da.) - Bhubaneswar (Bhu.) - Konarak(Ko.)

- U d a y g i r i (Ud.) 5. Himalaya: Ghum (Gh.) 6. West-Bengal: C a l c u t t a (Ca.) - B i r s i v p u r ( B i . ) - Nalbani

(Ha.) - Salt Lake (Sa.) - Bar k a l i k a p u r (Ba.) - Cherrapun-jee (Che.)

I use the l o c a l i t y names henc e f o r t h i n an abbreviated form (see i n brackets) .

The c o l l e c t i n g method of the helminths - obtained by d i s s e c t i o n of the guts conserved i n a l c o h o l - caused many subsequent d i f ­f i c u l t i e s . A p a r t of the f l u k e s were f i x e d i n a con t r a c t e d s t a t e , the g r e a t e s t p a r t of them was h a r d l y coloured, and many specimens were impossible t o clean of the gut contents w i t h o u t damage . Therefore the measurements of the body, the apprecia­t i o n of the topography of the organs, and the d e t e r m i n a t i o n i n p a r t of the m a t e r i a l remain u n c e r t a i n . The p o s t - f i x a t i o n was obtained i n Bouin.

The i n v e s t i g a t i o n s of the I n d i a n f l u k e fauna p a r a s i t i z i n g bats was launched by PANDE (1935a, b, c ) and f o l l o w e d by BHALERAO (1936) and GUPTA et BHARDWAJ (1958). I n the neighbouring coun­t r i e s GROSCHAPT et TENORA, 1971 ( A f g h a n i s t a n ) and BHALERAO, 1926a,b (Burma) c a r r i e d out bat h e l m i n t h o l o g i c a l i n v e s t i g a t ­i o n s . These papers submitted a meagre m a t e r i a l of occasional c o l l e c t i n g s , t h e r e f o r e the treatment of Gy. TOPÁL's m a t e r i a l o r i g i n a t i n g from wide-ranging c o l l e c t i n g s was u s e f u l i n every r e s p e c t , despite the bad c o n d i t i o n of the trematodes.

A n c h i t r e m a t i d a e Mehra, 1935 Anchitrema sanguineum (Sonsino, 1894) Hosts: (the a b b r e v i a t i o n o f the l o c a l i t i e s i n b r a c k e t s ) : Mega-derma l y r a ( B i . ) , Taphozous longimanus (Ko.), (Ca.), P i p i s t r e l -

F i g s . 1-4 1 = Neoheterophyes brevicorpus Matskási, 1973, 2 = N. t o p a l i sp. n. 3 = Lecithodendrium l i n s t o w i D o l l f u s , 1931, 4 - L.macro-stomum (Ozaki, 1929). ( F i g s . 1-4 o r i g . )

l u s mimus (Ca.,Sa.), Rhinolophus r o u x i (Ud.), Rh.lepidua (Da.), Myotis adversus (Ca.); L o c a l i z a t i o n : rectum.

The f a m i l y Anchitrematidae was es t a b l i s h e d as a d i s t i n c t f a m i ­ly-group by MEHRA i n 1935 ( s u b f a m i l y A n c h i t r e m i n a e ) . CABALLERO ( 1 9 6 1 ) gave i t f a m i l y s t a t u s . On the basis of the I n t e r n a t i o n a l Codex of Z o o l o g i c a l Nomenclature the c o r r e c t author of the taxon i s MEHRA, w i t h the date 1 9 3 5 .

I t i s remarkable t h a t t h i s species very common i n the t r o p i c s , was found i n I n d i a o n l y i n E a s t - I n d i a n bats, and on l y few hosts were i n f e c t e d w i t h i t . I n I n d i a i t has been found by PANDE ( 1 9 3 5 a ) .

P l a g i o r c h i i d a e Lühe, 1 9 0 1 P l a g i o r c h i s v e s p e r t i l i n n i g (Müller, 1784) Hosts: Megaderma l y r a ( B i . ) , Scotophilus temmincki (Ca.),Myotis adversus (Ca.), M. longi p e s (Bumz.); L o c a l i z a t i o n : rectum. The measurements and the r a t i o of suckers of the c o l l e c t e d f i v e specimens correspond w i t h the data of the cosmopolitan P. ves- p e r t i l i o n i s .

M i c r o p h a . l l i d a e Travassos, 1920 Renschetrema malayi Rohde, 1964 Hosts: Myotis adversus (Ca.), M. mur i c o l a (Gh.), M . s i l i g o r e n s i s (Gh.); L o c a l i z a t i o n : i n t e s t i n u m . ROHDE ( 1 9 6 4 ) described t h i s f l u k e species from Malaysian ba t s , l a t e r the author found i t i n North Vietnam (MATSKÁSI, 1 9 7 3 ) . I t i s r a t h e r s t r i k i n g , however, t h a t t h i s species l i v i n g i n Malaysia i n t r o p i c a l c l i m a t e , o c c u r r e d i n I n d i a - as i n North Vietnam - i n the high mountains but never i n the p l a i n s w i t h a t r o p i c a l c l i m a t e . Only one specimen p a r a s i t i z e d the small i n ­t e s t i n e of a Myotis adversus c o l l e c t e d i n C a l c u t t a , but 6 and 12 specimens, r e s p e c t i v e l y , were found i n Myotis m u r i c o l a and M. s i l i g o r e n s i s c o l l e c t e d i n Ghum ( l y i n g between the ranges of the Himalaya). The c o l l e c t e d f l u k e s concur both as t o body s t r u c t u r e and measurements w i t h the Malaysian and Vietnamese exemplars.

H e t e r o p h y i d a e Odhner, 1914 Neoheterophyes brevicorpus Matskási, 1973 - ( P i g . 1 ) Hoata: Myotia m u r i c o l a (Gh.), M. a i l i g o r e n a i a (Gh.), l a i o (Che.); L o c a l i z a t i o n : amall i n t e a t i n e . This apeciea was found f i r a t i n North Vietnam by the author (MATSKÁSI, 1973) .The apecimena c o l l e c t e d i n I n d i a agree i n mor­phology and measurements w i t h those described from Vietnam. The species occurs i n the high mountains (Himalaya) i n I n d i a as i n Vietnam. The bat l a i o c o l l e c t e d i n Cherrapunjee ( i n the f o r e ­ground of the Himalaya)was i n f e c t e d w i t h one trematode specimen o n l y , w h i l e the two Myotis species were i n f e c t e d w i t h 4-6 f l u ­kes .

Neoheterophyes t o p a l i sp. n. - ( P i g . 2 ) Host: Myotis s i l i g o r e n s i s (Gh.) L o c a l i z a t i o n : small i n t e s t i n e ; I n t e n s i t y : 1 exemplar Measurements i n yu: Body l e n g t h : 1920; maximal w i d t h : 512; o r a l sucker: 192x176; v e n t r a l sucker: 57x83; v e n t r a l sucker + acces­sory sucker l e n g t h : I 6 6 ; t e s t e s : 243x147, and 236x132; ovary: 192x217; c i r r u s - s a c : 608x96; eggs: 19x10.

Diagnosis: A medium sized trematode w i t h elongated body. Tegu­ment covered w i t h v e r y f i n e s p ines. V e n t r a l sucker s u b t e r m i n a l . Pharynx f o l l o w e d by short oesophagus. The two caeca approaching o r a l surface of t e s t e s . V e n t r a l sucker connected w i t h accessory appendages c h a r a c t e r i s t i c of the genus. The two s p h a e r i c a l t e s ­tes s i t u a t e d l a t e r a l l y i n the second p a r t o f the body.Ovary im-m e d i a t e l l y behind accessory sucker. V i t e l l a r i a s i t u a t e d on both sides of body. F o l l i c l e s commencing a n t e r i o r l y of ovary and t e r m i n a t i n g at t e s t e s . Shape of c i r r u s - s a c a s p e c i f i c f e a t u r e : the elongated, curved, s a c c u l i f o r m organ begins close by the ovary, arching beside v e n t r a l sucker, and t e r m i n a t i n g at pre-acetabular g e n i t a l pore; i t s d i s t a l p a r t t h i n n e r than proximal one. Uterus w i t h r e l a t i v e l y few whorls, f i l l i n g space between ovary and end of body.

Pour p a r a s i t i c trematodes of the f a m i l y Heterophyidae were h i t ­h e r t o known from b a t s , namely Heterophyes heterophyes ( S i e b o l d ,

1853), Neoheterophyes bychowskyi Khotenovsky, 1970, H. b r e v i ­ corpus Matskási, 1973, and N. huynhi Matskási, 1973. This spe­c i e s , discovered i n I n d i a , belongs t o the genus Neoheterophyes, supported by the g l a b r o u s , c h a r a c t e r i s t i c a l l y shaped accessory sucker, the l e n g t h of the caeca and the p o s i t i o n of the v i t e l l -ary glands (KHOTENOVSKY, 1970; MATSKÁSI, 1973).

D i f f e r e n t i a l d i a g n o s i s : N. t o p a l i sp. n. d i f f e r s from N. by- chowskyi by the s i g n i f i c a n t l y d i f f e r e n t measurements of the body, the d i f f e r e n t r a t i o of the suckers (the two organs are equal i n N. bychowakyi, while the o r a l sucker of N. t o p a l i i s l a r g e r than i t s acetabulum), the p o s i t i o n of the ovary (the ovary i n N. t o p a l i i s s i t u a t e d behind the acetabulum and not p a r a a c e t a b u l a r l y ) as w e l l as by the p o s i t i o n and the shape o f the c i r r u s - s a c . The new species d i f f e r s from N.bre­ vicorpus by the d i f f e r e n t measurements and shape of the bo­dy, the p o s i t i o n o f the ovary and v i t e l l a r i e s , as w e l l as by the shape and p o s i t i o n of the c i r r u s - s a c .

Holotype: s l i d e 1/29 ( L o c a l i t y : Ghum - North I n d i a , c o l l : Gy. TOPÁL; date of c o l l e c t i n g : 21 October, 1967), deposited i n the P a r a s i t o l o g i c a l C o l l e c t i o n of the Hungarian N a t u r a l H i s t o r y Mu­seum, Budapest. D e r i v a t i o nominis: I dedicate the new species to György TOPÁL, mammalogist, the c o l l e c t o r of the m a t e r i a l .

L e c i t h o d e n d r i i d a e Odhner, 1910 Lecithodendrium l i n s t o w i D o l l f u s , 1931 - ( P i g . 3 ) Hosts: Megaderma l y r a ( E H . ) , Scotophilus teramincki (Ca.), Sec— tozous dormeri ( J a . ) , Taphozous melanopogon (Ud.), T. longima-nus (Ca.), Miniopterus sehreibersi(Ma . ) , Myotis l o n g i p e s (Bu.), P i p i s t r e l l u s ceylonicus (Ka.), P. k u h l i ( S r . ) , P. mimus (Na., Ca.,Sa.,Ba.), Hipposideros a t e r ( K o . ) , Rhinolophus r o u x i (Ud.), Rh. l e p i d u s (Da.); L o c a l i z a t i o n : small i n t e s t i n e . L. l i n s t o w i occurred i n every one of the c o l l e c t i n g l o c a l i t i e s (except the Himalaya). The e x t e n s i t y and i n t e n s i t y of the i n ­f e c t i o n were very h i g h . PANDE (1935c) published t h i s species f i r s t from I n d i a under the name Mesodendrium mödlingeri, l a t e r

P i g s . 5-8 5 = Prosthodendrium mehrai (Pande, 1935), 6 and 8 = P. a s c i d i a (Van Beneden, 1873), 7 = P. minor Groschaft et Tenora, 1971. ( P i g s . 5-8 o r i g . )

GUPTA et BHARDWAJ (1958) described t h i s species as Lecithoden-drium minutum.

Lecithodendrium macrostomum (Ozaki, 1929) - ( P i g . 4) Hosts: Miniopterus s c h r e i b e r s i (Ma.), P i p i s t r e l l u s mimus (Ha., Ba., Ca.), P. ceylonicus (Ka., Bha.), Scotophilus temmincki (Ca., Ko.), Rhinolophus r o u x i (Ud., Ma.), Rh. le p i d u s (Da.); L o c a l i z a t i o n : small i n t e s t i n e .

L. macrostomum was found more r a r e l y than L. l i n s t o w i . PAHDE (1935c) described t h i s f l u k e species f i r s t from I n d i a under the name L. elongatum.

Prosthodendrium chilostomum ( M e h l i s , 1831) Hosts: Scotophilus temmincki (Ca.), P i p i s t r e l l u s mimus (B a . ) , P . k u h l i ( S r . ) , P. ceylonicus ( K a . ) , I a i o (Che.); L o c a l i z a t i o n : i n t e s t i n e .

Prosthodendrium longiforme (Bhalerao, 1926) Hosts: Megaderma l y r a ( B i . ) , Taphdzous melanopogon (Bhu.,Ud.), T. longimanus (Ko., Ca.), Scotophilus temmincki (Ca.), Sc. hea-t h i (Na.), Myotis l a n i g e r (Che.), Miniopterus f u l i g i n o s u s (Ma.), Rhinolophus r o u x i (Ud.), Hipposideros f u l v u s (Ud. ) . L o c a l i z a t i o n : i n t e s t i n e . PAHDE (1935b) found t h i s species f i r s t i n I n d i a , and pu b l i s h e d i t under the name Lecithodendrium longiforme v a r , a l l a h a b a d i .

Prosthodendrium mehrai (Pande, 1935) - ( P i g . 5) Host: P i p i s t r e l l u s ceylonicus (Bha., Ka.), L o c a l i z a t i o n : i n t e s ­t i n e . Pande found t h i s f l u k e f i r s t i n the small i n t e s t i n e of the bat Scotozous dormeri c o l l e c t e d near Allahabad. Since i t s f i r s t de­s c r i p t i o n the species was not found again.There are 3 specimens i n our m a t e r i a l from two l o c a l i t i e s i n Western I n d i a .

Prosthodendrium minor Groschaft et Tenora, 1971 - ( P i g . 7) Hosts: Myotis adversus (Ca.), Scotophilus h e a t h i (Ca.), P i p i ­s t r e l l u s k u h l i ( S r . ) ; L o c a l i z a t i o n : i n t e s t i n e . 84

P i g s . 9-12 9 - Prosthodendrium brachyurna Groschaft et Tenora, 1971, 10, 11 and 12 = P. l o o s s i (Pande, 1935). ( P i g s . 9-1 2 o r i g . J

Measurements i n yu: Body l e n g t h : 256 -320; g r e a t e s t w i d t h : 153-176; o r a l sucker: 32-44 x 44; v e n t r a l sucker: 19-28 x 28 -32 ; t e s t e s : 51-57 x 25-39 and 38-57 x 39-44; ovary: 38-57 x 32-38; p r o s t a t i c mass: 32-51 x 48 - 5 1 ; eggs: 19-25 x 9 -12 .

Our specimens from I n d i a do not agree i n every respect w i t h the type-specimens described by GROSCHAFT et TEHORA from Afghanis­t a n . While i n specimens discovered i n Afghanistan the o r a l and v e n t r a l suckers are equal, i n the I n d i a n exemplars the v e n t r a l sucker i s smaller than the o r a l one . There i s an other d i f f e ­rence between the i n d i v i d u a l s from Afghanistan and I n d i a , na­mely t h a t the integument of the exemplars from I n d i a i s f i n e l y spinose. Despite the above mentioned d i f f e r e n c e s I l i s t the I n ­dian specimens as Prosthodendrium minor.

Prosthodendrium ovimagnosum (Bhalerao, 1926) Hosts: Megaderma l y r a ( B i . , Ba.), Taphozous longimanus (Ko., Ca.), T. melanopogon (Bhu., R i . , Ud.), Scotophilus" temmincki (Ca., Ko.), Sc .heathi (Na.,Ma.), Miniopterus s c h r e i b e r s i (Ma.), Rhinopoma ha r d w i c k e i (Ud ., Eleph.), Hipposideros f u i v u s (Ud.) . L o c a l i z a t i o n : i n t e s t i n u m . BHALERAO ( 1936 ) , and l a t e r GUPTA et BHARDWAJ ( 1 9 5 8 ) , published the occurrence of t h i s species i n I n d i a . I n our m a t e r i a l , most of the specimens represent t h i s species. The i n t e n s i t y of the i n f e c t i o n extends to 150 specimens. Prosthodendrium ovimagnosum occurred i n every one of the c o l l e c t i n g l o c a l i t i e s , having a wide c i r c l e of hos t s . On the basis of Vietnamese m a t e r i a l , Prosthodendrium hepaticum Chen, 1954 was regarded as a j u n i o r synonym of P. ovimagnosum (MATSKÁSI, 1973) . The study of the In d i a n specimens corroborated t h i s statement, i n so f a r as the m a t e r i a l from I n d i a comprised specimens w i t h an "ovimagnosum" type and a "hepaticum" type of ovary, r e s p e c t i v e l y , and also several t r a n s i t i o n a l forms. The integument of the I n d i a n exem­p l a r s was covered w i t h spines.

Prosthodendrium parvouterus (Bhalerao, 1926) Hosts: Megaderma l y r a ( B i . ) , Taphozous t h e o b a l d i ( R i . ) , T. me-

lanopogon ( E H . ) , Scotophilus temmincki (Ca.); L o c a l i z a t i o n : i n t e s t i n e .

Prosthodendrium a s c i d i a (van Beneden, 1873) - ( P i g s . 6 and 8 ) Hosts: Myotis l o n g i p e s (Bu.), Miniopterus s c h r e i b e r s i (Ma.), Scotozous dormeri ( J a . ) , Rhinopoma hard w i c k e i (Ud.); L o c a l i z a ­t i o n : i n t e s t i n e . Measurements i n yu: Body l e n g t h : 510-610; g r e a t e s t w i d t h : 367-560; o r a l sucker: 96-115 x 89-102; t e s t e s : 96-183 x 108-144; and 108-153 x 115-128; ovary: 89-121 x 76-115; p r o s t a t i c com­ple x : 89-115 x 89-153; eggs: 19-24 x 12.

BHALERAO (1926b) described t h i s species under the name L e c i t h o ­ dendrium dinanatum from Burma. I n DUBOIS's o p i n i o n ( i 9 6 0 ) , Le­cithodendrium dinanatum Bhalerao, 1926,is the j u n i o r synonym of Prosthodendrium a s c i d i a (van Beneden, 1873). Our P. a s c i d i a specimens agree w i t h those described by several European authors (ODEHIHG, 1964, HURK0VÁ, 1963, DUBOIS, I960) and by BHALERAO. There i s a remarkable d i f f e r e n c e between the European and the A s i a t i c specimens i n the p o s i t i o n of the ovary.Whereas i n Euro­pean i n d i v i d u a l s the ovary i s praeacetabular, i t i s para or postacetabular i n Asian m a t e r i a l s .

Prosthodendrium brachyurna Groschaft et Tenora, 1971 - (Pig.9 ) Host: Myotis s i l i g o r e n s i s (Gh.). L o c a l i z a t i o n : i n t e s t i n e . Measurement i n yu: Body l e n g t h : 558; maximal w i d t h : 372; o r a l sucker: 78 x 78; v e n t r a l sucker: 81 x 75; t e s t e s : 94 x 69 and 90 x 63} ovary: 95 x 70; eggs: 20-21 x 9-10. Measurements of the second damaged exemplar: o r a l sucker: 90 x 55; v e n t r a l su­cker : 75 x 81.

GROSCHAFT and TENORA (1971) discovered t h i s species i n Myotis longipes c o l l e c t e d i n A f g h a n i s t a n . Our specimens d i f f e r from the type specimens i n having a spinose integument.

Prosthodendrium l o o s s i (Pande, 1935) - ( P i g s . 10-12 and 15) Hosts: Megaderma l y r a (Ba.), Scotophilus temmincki (Ca.), Ta­phozous melanopogon (Ud.), Myotis l a n i g e r (Che.), P i p i s t r e l l u s

mimus (Ca., Ba.), P. ceylonicus ( K a . ) , M i n i o p t e r u s s c h r e i b e r s i (Ma.), Rhinolophus r o u x i (Ud.), Rh. l e p i d u s (Logohad P o r t ) ; Lo­c a l i z a t i o n : small i n t e s t i n e .

PANDE (1935b) described t h i s f l u k e from I n d i a . GROSCHAPT and TENORA (1971) recorded P. l o o s s i (Pande, 1935) from bats c o l l ­ected i n Afghanistan, w i t h the remark t h a t i n t h e i r o p i n i o n the f e a t u r e s of the v a l i d species P. l o o s s i c h a r a c t e r i z e a l s o the species c o l l e c t e d and published by ODENING (1968a) as P. urna from Vietnam. They s u b s t a n t i a t e the d i s t i n c t n e s s of P. l o o s s i , drawn i n by DUBOIS ( i 9 6 0 ) as a synonym (a v a r i e t y ) of P. urna (Looss, 1907), by the p o s i t i o n and shape of the p r o s t a t i c com­ple x (pseudocirrus-sac) as w e l l as by the l o c a t i o n of the ovary of the species o r i g i n a l l y described from bats i n I n d i a d i f f e r -r l n g from P• urna (Looss, 1907), as given by PANDE also i n o r i ­g i n a l d e s c r i p t i o n . The p r o s t a t i c complex of P. urna i s preace-t a b u l a r , of P . l o o s s i p a raacetabular. The p o s i t i o n of the ovary, however, i s not a r e l i a b l e f e a t u r e t o d i s t i n g u i s h species since i t i s very v a r i a b l e and i t s form depends on the s t a t e of con­t r a c t i o n of the f l u k e . The measurements of the suckers or other organs do not d i f f e r i n the o r i g i n a l d e s c r i p t i o n s , a l t o u g h the o r a l and v e n t r a l suckers are somewhat smaller i n P. l o o s s i than i n P. urna .

The specimens of our I n d i a n m a t e r i a l are v e r y heterogeneous concerning the p o s i t i o n of the p r o s t a t i c complex and the mea­surements of the organs. The v i t e l l a r i ' a are a l s o very h e t e r o ­geneous. On the basis of the above-mentioned three f e a t u r e s , the m a t e r i a l i s d i v i d e d i n t o two p a r t s . I n the f i r s t group the p r o s t a t i c complex i s paraacetabular, the v i t e l l a r i a are compos­ed of few f o l l i c l e s and d i v i d e d sharply i n t o two groups. The o r a l sucker (4-0-51 x 40-54) and the v e n t r a l sucker (32-40 x 32-41/u) are smaller than those o f the second group ( P i g s . 10, 1 5 ) . I n the second group, the p r o s t a t i c complex i s l y i n g dor-s a l l y of the acetabulum, the v i t e l l a r i a are composed of minute f o l l i c l e s , j o i n i n g i n f r o n t of the acetabulum. The measurements of the o r a l sucker are 49-83 x 55-70 ja, of the v e n t r a l sucker 44-60 x 45-57 ja ( P i g s . 1 1 , 1 2 ) . The integument of both groups

15 16

P i g s . 13-16 13 and 14 = Prosthodendrium robustum sp . n., 15 = P« l o o s s i (Pande, 1935), 16 = Prosthodendrium sp. ( P i g s . 13-16 o r i g . )

i s spinose. The f l u k e s of the f i r s t group are r e l e g a b l e morpho­l o g i c a l l y t o PANDE's type, but on the basis of the measure­ments the second group stay nearer t o the type described by Pande. The exemplars assignable t o the f i r s t group were dissec­ted from several bat species, whereas the specimens of the se­cond group p a r a s i t i z e d two Rhinolophid species o n l y .

MACY ( 1 9 6 4 ) discovered and described a new trematode species under the name Pycnoporus ramsesi sp. n., from P i p i s t r e l l u s r u p p e l i i n Egypt. On the basis of the d e s c r i p t i o n , the measure­ments and the f i g u r e s , t h i s species i s t o be regarded as a j u ­n i o r synonym of Prosthodendrium l o o s s i (Pande, 1 9 3 5 ) .

Prosthodendrium robustum sp . n. - ( P i g s . 5 , 13, 1 4 ) Host: Myotis m u r i c o l a (Gh.) - L o c a l i z a t i o n : i n t e s t i n u m , I n t e n ­s i t y : 2 exemplars. Measurements ( i n ^ u ) : Body l e n g t h : 800 - 1 0 0 0 ; g r e a t e s t w i d t h : 5 9 2 - 7 3 0 ; o r a l sucker: 128 x 89 and 132 x 1 1 5 ; v e n t r a l sucker: 96 x 128 and 208 x 1 9 2 ; l e f t t e s t i s : I 6O x 102 and 192 x 176; r i g h t t e s t i s : 1 6 0 x 1 1 2 and 256 x 176; ovary: 138 x 99 and I 6 0 x 1 6 0 ; p r o s t a t i c complex: 224 x 179 and 198 x I 4 O ; eggs: 1 9 - 2 2 x 9-12 .

Diagnosis: A medium-sized trematode w i t h p y r i f o r m body. I n t e g u ­ment spinose. Oral sucker s u b t e r m i n a l . Oesophagus absent. Caeca extending t o t e s t e s . Pharynx w e l l developed. Sphaerical t e s t e s l y i n g behind acetabulum. ( P i g . 14 shows a c o n t r a c t e d specimen, t h e r e f o r e the p o s i t i o n of the organs i s not n a t u r a l . ) L a r g e pro­s t a t i c complex s i t u a t e d d o r s a l from acetabulum, i t s o r a l p a r t extending i n t o i n t e r c a e c a l area. G e n i t a l pore p r e a c e t a b u l a r . Ovary postaçetabular. Ovary rounded, smaller than t e s t e s , l o ­cated p o s t a c e t a b u l a r l y , l a t e r a l . V i t e l l a r i a composed of numerous f o l l i c l e s , s i t u a t e d between acetabulum and p r o s t a t i c complex. Uterus very w e l l developed, of numerous whorls, occupying cau­dal p o r t i o n of body.

D i f f e r e n t i a l d i a g n o s i s : Prosibodenarium robustum d i f f e r s from a l l other congeners p r i n c i p i a l l y b y / i t s l a r g e v i t e l l a r y

area. The f o l l i c l e s f i l l out the area between the caeca and the v e n t r a l sucker. The extensive measurements of the o r a l and v e n t r a l suckers are also s p e c i f i c f e a t u r e s .

Holotype: s l i d e 1/240 ( L o c a l i t y : Ghum; date of c o l l e c t i n g : 20 A p r i l , 1 9 6 7 ; c o l l . : Gy. TOPÁL), deposited i n the P a r a s i t o l o g i c a l C o l l e c t i o n of the Hungarian N a t u r a l H i s t o r y Museum, Budapest. Paratype: s l i d e I / 2 4 0 b , deposited as above.

Prosthodendrium sp. - ( P i g . 16) Host: Taphozous melanopogon (Bhu.); L o c a l i z a t i o n : i n t e s t i n u m ; I n t e n s i t y : 1 specimen. Measurements ( i n yu): Body l e n g t h : 1210; g r e a t e s t w i d t h : 800; o r a l sucker: 153x153; v e n t r a l sucker: 96x102; t e s t e s : 249-l60x 208-160; ovary: 153x83; p r o s t a t i c complex: 124x187; eggs: 19x9.

Diagnosis: A medium-sized f l u k e w i t h p y r i f o r m body. Integument covered w i t h spines. Oral sucker s u b t e r m i n a l . Pharynx f o l l o w e d by a long oesophagus. Caeca t e r m i n a t i n g at t e s t e s . Rounded t e s ­tes s i t u a t e d i n f r o n t of acetabulum. Large p r o s t a t i c complex l y i n g between t e s t e s . Sphaerical ovary l o c a t e d p r e a c e t a b u l a r l y , l a t e r a l l y from the acetabulum. V i t e l l a r y glands between o r a l sucker and t e s t e s . U t e r a l whorls occupying p o s t a c e t a b u l a r por­t i o n of body.

This species stands near the bat trematode Prosthodendrium l e p - tocoelium (Ogata, 1942), by the p r e a c e t a b u l a r l y s i t u a t e d ovary, the p o s i t i o n of the v i t e l l a r y glands, the measurements of the suckers, and the g e n i t a l organs. However, the ovary of P. l e p - tocoelium i s i n a median p o s i t i o n , but l a t e r a l l y i n the I n d i a n exemplar. The p r o s t a t i c complex of P. l e p t o c o e l i u m i s f a r from the acetabulum (the ovary i s between the p r o s t a t i c complex and the acetabulum), i n our I n d i a n specimen the v e s i c u l a seminalis i s s i t u a t e d d o r s a l l y of the v e n t r a l sucker, and touches i t . The measurements of eggs d i f f e r a l s o , because the eggs of P. l e p t o ­ coelium are 26-27x 15 , the eggs of the I n d i a n exemplar 23x12 u.

Hosts: Scotophilus temmincki (Ca.), Taphozous longimanus (Ca.), P i p i s t r e l l u s mimus (Ca.), P. k u h l i ( S r . ) , P. ceylonicus (Ka.); L o c a l i z a t i o n : small i n t e s t i n e .

PAHDE (1935b) described t h i s species from I n d i a under the name Pycnoporus l o o s s i . I n DUBOIS's o p i n i o n ( i 9 6 0 ) i t i s synonymous w i t h P. acetabulatus Looss, 1899-

Pycnoporus macrolaimus (Linstow, 1894) Host: Hipposideros a t e r (Ko.); L o c a l i z a t i o n : i n t e s t i n e . Only one damaged specimen found.

P l e u r o g e n i d a e Looss, 1899 Maxbraunium b a e r i Roh.de, 1964 - ( P i g . 17) Host: Myotis s i l i g o r e n s i s (Gh.); L o c a l i z a t i o n : i n t e s t i n u m . Rohde (1963) described t h i s species from Malaysia. I n d i a i s the second l o c a l i t y of t h i s f l u k e .

Parabascus g h a t i sp. n. - (P i g s . 18-20) Host: Megaderma l y r a ( E l l o r a Caves); L o c a l i z a t i o n : small i n ­t e s t i n e ; I n t e n s i t y : 14 specimens.

Comparative measurements i n u of Parabascus g h a t i sp.n. Body l e n g t h 64O 608 540 704 768 64O Body w i d t h 540 64O 480 576 672 528 Oral sucker 80x112 80x96 57x99 96x105 86x118 64x96 V e n t r a l sucker 121x121 105x112 112x112 128x140 132x160 121x121 L e f t t e s t i s 128x112 128x224 96x160 144x160 169x166 128x144 Right t e s t i s - 144x192 - 128x160 - 128x166 Ovary 128x76 172x105 112x89 140x160 179x70 144x144 Cirrus-sac 128x112 - 108x108 176x176 179x144 140x140 Egg 19x19 20x9 19x10 22x9 19x11 19x11

Diagnosis: A s m a l l - s i z e d trematode, g e n e r a l l y t a p e r i n g toward o r a l sucker. Integument spinose. Oral sucker t e r m i n a l or sub-te r m i n a l . Muscular pharynx f o l l o w e d by short oesophagus. Two caeca, extending along two sides of body, reaching caudal end of t e s t e s , a r c h i n g t o middle of body. V e n t r a l sucker l a r g e r than o r a l one. Two l a r g e t e s t e s s i t u a t e d l a t e r a l l y , s ymmetrically at

l e v e l of acetabulum, or p o s t a c e t a b u l a r l y . Shape of ovary i r r e ­g u l a r , r a r e l y l o b e d . Ovary l o c a t e d between r i g h t t e s t i s and acetabulum, i n area between caecal branches. Uterus occupying postacetabular p a r t of body. Cirrus-sac p a r a a c e t a b u l a r . G e n i t a l pore adjacent t o acetabulum. Cirrus-sac c o n t a i n i n g a w e l l de­veloped v e s i c u l a s e m i n a l i s . Receptaculum seminis d o r s a l l y from v e n t r a l sucker. V i t e l l a r i a composed of numerous minute f o l l i c -

F i g s . 17-20 17 = Maxbraunium b a e r i Rohde, 1964, 18, 19 and 20 - Parabascus g h a t i sp. n. ( F i g s . 17-20 o r i g . )

l e s l y i n g between pharynx and v e n t r a l sucker. Two v i t e l l i n e ducts j o i n i n g behind v e n t r a l sucker.

D i f f e r e n t i a l d i agnosis: The most c h a r a c t e r i s t i c a l f e a t u r e of Parabascus g h a t i sp. n. i s the i r r e g u l a r l y shaped, lobed ovary, s i t u a t e d p r e a c e t a b u l a r l y . This species d i f f e r s from a l l other congeners by i t s w e l l developed, rounded c i r r u s -sac, and r e l a t i v e l y short caeca. A f u r t h e r s p e c i f i c f e a t u r e i s the f a i r l y equal l e n g t h of the l o n g i t u d i n a l and t r a n s ­v e r s a l axes.

Holotype: s l i d e I / 2 1 3 a ( L o c a l i t y : E l l o r a Caves; Date of c o l l e c ­t i n g * 16 August, 1967; c o l l . : Gy. TOPÁL) deposited i n the Para-s i t o l o g i c a l C o l l e c t i o n of the Hungarian N a t u r a l H i s t o r y Museum. Paratypes: s l i d e I/213b deposited as above.

Suming up the r e s u l t s described i n the present paper,from 22 trematode species c o l l e c t e d by Gy. TOPÁL i n I n d i a , 13 species were found t o be new f o r the I n d i a n trematode fauna: P I a g i o r ­c h i s v e s p e r t i l i o n i s (Müller, 1784), Neoheterophyes br e v i c o r p u s Matskási, 1973, H. t o p a l i sp. n., Prosthodendrium chilostomum ( M e h l i s , 1831), P. a s c i d i a (van Beneden, 1873), P. parvouterus (Bhalerao, 1926), P. minor Groschaft et Tenora, 1971 , P. bra- chyurna Groschaft et Tenora, 1971, P. robustum sp. n., Prostho­ dendrium sp., Pycnoporus macrolaimus (Linstow, 1894), Rensche-trema malayi Rohde, 1964, Parabascus g h a t i sp. n.

Although comparatively few bats have on t h i s occasion been exa­mined p a r a s i t o l o g i c a l l y , i t can be e s t a b l i s h e d t h a t the trema­tode fauna p a r a s i t i z i n g bats i n the Himalaya d i f f e r s from t h a t of a l l other l o c a l i t i e s . The trematode species o c c u r r i n g i n the Himalaya - Maxbraunium b a e r j . Renschetrema m a l a y i , Neohetero-phyes brevicorpus - occur i n h i g h mountains i n Vietnam and i n Malaysia.

M A T S K Á S I , L : Indiai denevérek mételyei

TOPÁL György Elő-India területén 36 denevérfaj 278 példányát gyűjtötte be. Ezek helmintológiai vizsgálata során 22 métely­f a j t sikerült meghatározni. Közülük három - a Neoheterophyes t o p a l i , a Prosthodendrium robustum é s a Parabascus g h a t i - a tudományra nézve új fajoknak b i z o n y u l t a k . A dolgozat a gyűj­tött métely-fajok f a u n i s z t i k a i a d a t a i t és r e n d s z e r t a n i f e l d o l ­gozását t a r t a l m a z z a .

References

BHALERAO, G.D.: The i n t e s t i n a l p a r a s i t e s of the bat (Nyctinomus p l i c a t u s ) w i t h a l i s t of the trematodes h i t h e r t o r e ­corded from Burma. - J. of the Burma Res. Soc. 15_. 181-189. 1926a.

BHALERAO, G.D.: A new species o f Trematode from N y c t i c e j u s p a l -l i d u s , w i t h a key t o the species o f Lecithodendrium. - Ann. et Mag. Nat. H i s t . ( 9 . ) 18^ 299-304. 1926b.

BHALERAO, G. D.: Studies on the helminths of I n d i a . Trematoda I I I . - J. Helminth. 14. 207-228. 1936.

CHEN, H.T.: On the taxonomy o f Prosthodendrium D o l l f u s 1931 and Longitrema gen. nov., w i t h a d e s c r i p t i o n of two new species and a new v a r i e t y (Trematoda: L e c i t h o d e n d r i -i d a e ) . - Acta Zool. S i n i c a . 6. 147-182. 1954.

DUBOIS, G.: Les Trematodes de Chiroptères de l a c o l l e c t i o n V i l -

l y A e l l e n . Étude s u i v i e d'une r e v i s i o n du sous-genre

Prosthodendrium Dollfus, 1 9 3 7 ( L e c i t h o d e n d r i i n a e Lühe)

- Rev. suisse Z o o l . 62. 469-506. 1955.

DUBOIS, G.: C o n t r i b u t i o n a l'étude des Trematodes de Chiroptè­

r e s . Revision du sous-genre Prosthodendrium D o l l f u s

1931 et des genres Lecithodendrium l o o s s 1896 et Pyc­

noporus Looss 1899« - Rev. suisse Zool. 67« 1-80.

I960.

DUBOIS, G.: C o n t r i b u t i o n a l'étude des Trematodes de Chiroptè­

r e s . Revision du genre' Allassogonoporus O l i v i e r 1938

et note a d d i t i o n e i l e sur l e sous-genre Prosthoden-

drium D o l l f u s 1931. - Rev. suisse Z o o l . 70. 103 -125 . 1963.

GUPTA, N, K. - BHARDWAJ, 0. N.: On two new Trematodes of b a t s , Lyroderma l y r a ( G e o f f r o y ) i n Gurdaspur, I n d i a . - Res. B u l l . Punjab Univ. 141. 75-79. 1958.

GR0S0HAPT, J. - TENORA, P.: Trematodes of the genus Prostho­dendrium D o l l f u s 1931 ( L e c i t h o d e n d r i i d a e ) - p a r a s i t e s of bats i n Afghani s t a n . - P o l i a Par. (Praga) 18. 127-138. 1971.

KHOTENOVSKY, I . A.: N o v i j r od i v i d t r e m a t o d i Neoheterophyes bychowskyi gen. et sp. n. i z l e t u t s c h e j myshi Myotis c o p a c c i n i i . - P a r a s i t o l o g i j a 4. 122-124. 1970.

KHOTENOVSKY, I . A.: Semejstvo Pleurogenidae Looss, 1899- I n : SKRJABIN, K.: Trematodi z h i v o t n y h i cheloveka 23 .spec. p. 137 - 2 9 7 . 1970.

LOOSS, A.: Notizen zur Helminthologie Aegyptens. V I I I . Über ei n i g e neue Trematoden der ägyptischen Fauna. - Z b l . Bakt. P a r a s i t e n k . I n f e k t i o n s k r a n k h . Abt. I I . 43_. 478-490. 1907.

MACY, R.W.: A new species o f Trematode, Pycnoporus ramsesi (Le­c i t h o d e n d r i i d a e ) , from Egypt and notes on P. acetabu­l a r s Looss, 1899. - Proc. of the Helm. Soc. 31. 292-297. 1964.

MACY, R.W. - HEYNEMANN, D. - KUNTZ, R.E.: Records of Trematodes of the f a m i l i e s L e c h i t h o d e n d r i i d a e , D i c r o c o e l i i d a e from C h i r o p t e r a c o l l e c t e d i n Egypt and Yemen, S. W. Arabia. - Proc. Helm. Soc. Wash. 28. 13-17. 1961.

MATSKÁSI, I . : Flukes from bats i n Vietnam. - Acta Zool. Hung, ( i n p r i n t ) .

MEHRA, H. R.: New Trematodes of the f a m i l y L e c i t h o d e n d r i i d a e Odhner, 1911 w i t h a d i s c u s s i o n on the c l a s s i f i c a t i o n of the f a m i l y . - Proc. Acad. S e i . U. P. I n d i a 4. 99-121. 1934-35.

ODENING, K.: Exkretionssystem und systematische S t e l l u n g e i n i ­ger Pledermaustrematoden aus B e r l i n und Umgebung nebst Bemerkungen zum l e c i t h o d e n d r i i d e n Komplex. - Z.f. Pa­r a s i t e n k . 24. 453-483. 1964-

ODENING, K.: Trematoden aus vietnameaiachen C h i r o p t e r e n . - Zool. Abhandl. 29. 119-157. 1968a.

ODENING, K.: Zur Kenntnis der einheimischen Fledermaustremato-den. Zweiter B e i t r a g . - Zool . Jb. Syst. 95_. 265-296. 1968b.

OGATA, T.: Sur un nouveau trematode d'une chauve-souris du Pa-l a o s , I I . Lecithodendrium l e p t o c o e l i u m n. sp. - Zool. Mag. Tokyo 289-291. 1942.

OZAKI, Y.: Three new Trematodes from b a t . - Annot. Zool. Jap. 12. 91-98. 1929.

PANDE, B. P.: C o n t r i b u t i o n s t o the d i g e n e t i c trematodes of the M i c r o c h i r o p t e r a of Northern I n d i a . Studies on the ge­nus Lecithodendrium Looss. - Proc. Acad. S e i . U. P. I n d i a 4. 86-97. 1934-35a.

PANDE, B. P.: C o n t r i b u t i o n to the d i g e n e t i c trematodes of the M i c r o c h i r o p t e r a of Northern I n d i a . New Distomes of the genus Mesodendrium Paust (1919). - Proc. Acad. Se i . U. P. I n d i a 4. 243 -249 . 1934-35b.

PANDE, B. P.: C o n t r i b u t i o n s t o the d i g e n e t i c trematodes of the M i c r o c h i r o p t e r a of Northern I n d i a . New species of the genus Pycnoporus Looss w i t h a note on Anchitrema Looss. - Proc. Acad. Sei. U. P. I n d i a 4. 371-380. 1934-35C.

ROHDE, K.: Trematoden malayischer Fledermäuse. - Z. f . P a r a s i -tenk. 23_. 324-339 . 1963.

ROHDE, K.: Eine neue Trematoden-Gattung (Renschetrema, Micro-p h a l l i d a e ) m i t d r e i Arten aus dem Darm malayischer Fledermäuse. - Z. f . P a r a s i t e n k . 24. 13-22. I 9 6 4 .

SOGANDARES-BERNAL, P.: Pour trematodes from Korean bats w i t h d e s c r i p t i o n s of th r e e new species. - J. Par. 42. 200-206. 1956.

Received: 14. 2. 1973. Dr. MATSKÁSI, I . Z o o l o g i c a l Department of the

Hungarian H a t u r a l H i s t o r y Museum 1088 Budapest, Baross u. 13.