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I. FIVE NEW SPECIES OF THAPAROCLEIDUS JAIN, 1 9 5 2 (ANCYLODISCOIDINAE) FROM PANGASIUS PANGASIUS, P. KINABATANGANENSIS, P. RHEOPHILUS AND P. NIEUWENHUISII

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MONOGENEANS FROM PANGASIIDAE (SlLURIFORMES) IN SOUTHEAST ASIA:

I. FIVE NEW SPECIES OF THAPAROCLEIDUS JAIN, 1 9 5 2 (ANCYLODISCOIDINAE) FROM PANGASIUS PANGASIUS, P. KINABATANGANENSIS, P. RHEOPHILUS AND P. NIEUWENHUISII

PARISELLE А.*,**, LIM L.H.S.*** & LAMBERT A . * * * *

S u m m a r y :

The examination of gill parasites from Pangasius pangasius (Hamilton, 1822) ; P. kinabatanganensis Roberts & Vidthayanon,

1991 ; P. rheophilus Pouyaud & Teugels, 2 0 0 0 and P. nieuwenbuisii (Popta, 1904) (Siluriformes, Pangasiidae) in Southeast Asia revealed the presence of six species of

Thaparocleidus Jain, 1952 (Monogenea, Ancylodiscoidinae). One has been previously described from P. pangasius: T. pangasi (Tripathi, 1957). The other five are new species: T. chandpuri n.

sp. from P. pangasius; T. bahari n. sp. and T. sabanensis n. sp.

from P. kinabatanganensis; T. redebensis n. sp. from P. rheophilus and T. mahakamensis n. sp. from P. nieuwenhuisii.

KEY WORDS : Monogenea, Pangasiidae, Thaparocleidus pangasi, T. chandpuri n, sp., T. bahari n. sp., T. sabanensis n. sp., T. redebensis n.

sp., T. mahakamensis n. sp

Résumé : MONOGÈNES DE PANGASIIDAE (SILURIFORMES) EN ASIE DU SUD-EST : I. CINQ ESPÈCES NOUVELLES DE THAPAROCLEIDUS JAIN, 1952 (ANCYLODISCOIDINAE), CHEZ PANGASIUS PANGASIUS, P. KINABATAN- GANENSIS, P. RHEOPHILUS ET P. NIEUWENHUISII

l'examen des parasites branchiaux de Pangasius pangasius (Hamilton, 1822); P. kinabatanganensis Roberts & Vidthayanon,

1991 ; P. rheophilus Pouyaud & Teugels, 2000 et P.

nieuwenhuisii (Popta, 1904] (Siluriformes, Pangasiidae), en Asie du sud-est, a révélé la présence de six espèces du genre Thaparocleidus Jain, 1952 (Monogenea, Ancylodiscoidinae).

l'une, T . pangasi (Tripathi, 1957), a été retrouvée sur P. pangasius. Cinq autres espèces nouvelles, du même genre, sont décrites: T. chandpuri n. sp. chez P. pangasius; T. bahari n.

sp. et T. sabanensis n. sp. chez P. kinabatanganensis;

T. redebensis n. sp. chez P. rheophilus et T . mahakamensis n. sp.

chez P. nieuwenhuisii.

MOTS CLÉS : Monogenea, Pangasiidae, Thaparocleidus pangasi, T. chandpuri n. sp., T . bahari n. sp., T , sabanensis n, sp., T . redebensis n. sp., T. mahakamensis n. sp.

I N T R O D U C T I O N

W

ithin t h e framework o f an EC project o n the biodiversity a n d culture o f Southeast Asian catfishes, t h e gills from pangasiid fishes (Siluriformes, P a n g a s i i d a e ) w e r e e x a m i n e d for m o n o g e n e a n parasites. T w e n t y t w o o f t h e 2 3 k n o w n s p e c i e s o f fishes b e l o n g i n g t o this family h a v e b e e n e x a m i n e d . This p a p e r presents t h e descriptions o f t h e five n e w s p e c i e s o f Thaparocleidus J a i n , 1 9 5 2 ( M o n o - g e n e a , A n c y l o d i s c o i d i n a e ) found o n four Pangasius V a l e n c i e n n e s , 1 8 4 0 s p e c i e s , viz., P. kinabatanganensis Roberts & Vidthayanon, 1 9 9 1 ; P. nieuwenhuisii (Popta, 1 9 0 4 ) ; P. rheophilus P o u y a u d & T e u g e l s , 2 0 0 0 a n d

* IRD ( e x ORSTOM), Catfish Asia Project, Instalasi Penelitian Per- ikanan Air Tawar, J a l a n Ragunan, Pasar Minggu, P.O. B o x 7220/jkspm, Jakarta 12540, Indonesia.

** GAMET, B.P. 5095, 34033 Montpellier Cedex 1, France.

*** Institute of Biological Sciences, University of Malaya, 50603 Kuala Lumpur, Malaysia.

**** Laboratoire Génome, Populations, Interactions, UMR 5000, c c 105, Université Montpellier II, Sciences et Techniques du Languedoc, Place Eugène Bataillon, 34095 Montpellier Cedex 5, France.

Correspondence : Alain Lambert.

E-mail: lambert@crit.univ-montp2.fr

P. pangasius (Hamilton, 1 8 2 2 ) a n d a redescription o f Thaparocleidus pangasi. T h e first three fish s p e c i e s from B o r n e o ( I n d o n e s i a a n d Malaysia) have not b e e n e x a m i n e d for parasites; w h i l e t h e fourth s p e c i e s (from B a n g l a d e s h ) , w a s e x a m i n e d b y Tripathi ( 1 9 5 7 ) in India.

M o n o g e n e a from t h e o t h e r 18 pangasiid s p e c i e s will b e d e s c r i b e d e l s e w h e r e .

T o date a total o f three s p e c i e s o f Thaparocleidus have b e e n d e s c r i b e d from P. pangasius a n d P. hypoph- thalmus ( S a u v a g e , 1 8 7 8 ) from India a n d Peninsular Malaysia, respectively ( s e e Tripathi, 1 9 5 7 ; Lim, 1 9 9 0 ) .

MATERIALS A N D M E T H O D S

F

ish were bought in fish markets or directly from fishermen in Indonesia (East Kalimantan region, South part of Borneo Island), Malaysia (Sabah state, North part of Borneo Island) and Bangladesh.

Fish were caught in the rivers using lines. The fish were dissected as soon as possible; the left branchial arches, separated by dorsal and ventral section, were frozen in liquid nitrogen, until examination. To verify the spe- cific identity of host fishes, the carcasses were num- Article available athttp://www.parasite-journal.orgorhttp://dx.doi.org/10.1051/parasite/2001082127

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bered, fixed and preserved in formalin. In the labora­

tory, the gills w e r e t h a w e d and the m o n o g e n e a n s w e r e d e t a c h e d from the gill using a strong water cur­

rent. T h e w o r m s w e r e then transferred individually o n a slide with a m o u n t e d n e e d l e directly into a drop o f a m m o n i u m picrate-glycerine (mixture d e s c r i b e d b y Malmberg ( 1 9 5 7 ) ) . T h e preparation w a s then c o v e r e d with a round cover slip and sealed with Glyceel (GURR - B D H Chemicals Ltd.). From these preparations, dra­

wings w e r e m a d e o f the sclerotised p i e c e s o f the haptor and o f the copulatory c o m p l e x using a c a m e r a lucida. M e a s u r e m e n t s , m a d e with a digitiser, in micro­

meters as the m e a n ± the standard deviation followed b y the range in parentheses, are those p r o p o s e d b y G u s s e v ( 1 9 6 2 ) (Fig. 1 ) . T h e m e t h o d o f numbering o f the haptoral p i e c e s is that adopted at ICOPA IV (Euzet

& Prost, 1 9 8 1 ) . T h e terminologies p r o p o s e d b y Pari- selle and Euzet ( 1 9 9 5 ) and b y N ' D o u b a et al. ( 1 9 9 9 ) are used.

R E S U L T S

S

ix s p e c i e s o f M o n o g e n e a w e r e r e c o r d e d ; o n e , Thaparocleidus pangasi (Tripathi, 1 9 5 7 ) , had already b e e n described, while the other five are n e w s p e c i e s o f Thaparocleidus ( A n c y l o d i s c o i d i n a e , M o n o g e n e a ) as defined b y Lim ( 1 9 9 6 ) .

R E D E S C R I P T I O N

THAPAROCLEIDUS PANGASI (TRIPATHI, 1957) (Fig. 2)

Type-host: Pangasiuspangasius (Hamilton, 1 8 2 2 ) . Site: gills.

Type-locality: Rivers Malta, G a n g a ( B u x a r ) , Cauvery and Hoogly.

O t h e r l o c a l i t y : a l s o f o u n d o n t h e s a m e h o s t at Chandpur, G a n g e s Delta, B a n g l a d e s h .

Fig. 1. - Measurements used in this study.

C - cuneus: L = length; 1 = largest width;

e = extension length.

DB = dorsal transverse bar: x = total length;

w = width in the middle.

DG - dorsal gripus; a, b, c, d and e = stan­

dard measurements.

MA = male apparatus: Pe = total length of the penis; Ap = length of the accessory piece; He = length of the heel.

U = total length of the uncinulus.

VB = ventral transverse bar: x = length of one branch; w = largest width.

VG = ventral gripus: a, b, c, d and e = stan­

dard measurements; L and 1 = length and width of gripus aperture.

Vg = vagina.

128 Mémoire Parasite, 2001, 8, 127-135

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Adults 534 ± 103-5 ( 4 0 7 - 8 0 0 ) long, 109 ± 12.3 (76-129) w i d e at the level o f the penis. Pharynx 49 ± 6.1 (37- 63) wide. Dorsal gripus with blade bent in middle, very short guard: a = 61 ± 1.7 (55-64), b = 5 0 ± 1.6 (44-

53), c = 2 + 0.5 (1-4), d = 14 ± 1.2 (12-17), e = 3 0 ±

1.1 (26-32). Cuneus with long extension: L = 20 ± 1.1 (17-22), 1 = 6 ± 0.5 (5-8), e = 11 ± 1.4 ( 8 - 1 5 ) . Slightly curved dorsal transverse bar: x = 3 8 ± 1.5 (35-42), w

= 6 + 0.6 (5-7).

Ventral gripus with well marked aper­

ture, poorly m a r k e d outer root: a = 2 8 ± 0.8 ( 2 5 - 3 0 ) ,

b = 25 ± 0.8 (23-26), c = 2 ± 0.4 (1-3), d = 9 ± 0.7 (7- 10),

e = 18 ±

0.9

( 1 5 - 1 9 ) , L =

4 ± 0.5 (3-5), 1 = 2 ± 0.3 (1-2).

Thin V-shaped ventral transverse bar: x =

33 ± 1.4 (29-36),

w =

4 ± 0.5 (3-4).

Uncinuli II =

16 ± 1.3

(13-21), uncinuli I and III to VII = 16 ± 1.8 (9-19). Long and folded penis, with ovoid basal bulb and large heel:

Pe = 181 ±

4.7 (169-192),

He -

11 ± 1.5

( 8 - 1 5 ) . Curved a c c e s s o r y p i e c e linked to the basal bulb o f the penis:

Ap = 54 ± 2.1 ( 5 0 - 5 8 ) . No sclerotised vaginal system

Fig. 2. - Thaparocleidus pangasi (Tripathi. 1957) : C = cuneus; DB = dorsal transverse bar; DG - dorsal gripus; MA = male apparatus; VB - ventral transverse bar; VG = ventral gripus; U = uncinulus. Bar - 30 urn.

This species is identified as Thaparocleidus pangasi first described b y Tripathi ( 1 9 5 7 ) from Pangasius panga­

sius in India. T h e m e a s u r e m e n t s for the present spe­

c i m e n s are not different from those given by Tripathi ( 1 9 5 7 ) . T h e s h a p e o f the sclerotised parts are also similar. T h e parasites w e r e r e c o v e r e d from the s a m e fish species, P. pangasius w h i c h is the only Pangasius s p e c i e s occurring naturally in India and the present host s p e c i e s w e r e from o n e o f the type localities River ( G a n g a ) , as noted by Tripathi ( 1 9 5 7 ) .

D E S C R I P T I O N

THAPAROCLEIDUS CHANDPURI N. SP. (Fig. 3)

Type-host: Pangasius pangasius (Hamilton, 1822).

Site: gills.

Type-locality: Chandpur, G a n g e s Delta, Bangladesh.

Material studied: 2 9 individuals fixed and m o u n t e d in Malmberg solution.

MONOGENEANS FROM PANGASIIDAE

Material studied: 3 0 individuals stained and m o u n t e d C o m m e n t s in Malmberg solution.

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Fig. 3. - Thaparocleidus chandpuri n. sp. : C = cuneus ; DB = dorsal transverse bar; DG - dorsal gripus; MA

= male apparatus; VB = ventral transverse bar; VG

= ventral gripus; Vg - vagina; U = uncinulus. Bar

= 30 urn.

T y p e - m a t e r i a l : h o l o t y p e d e p o s i t e d at t h e M u s e u m National d'Histoire Naturelle (Paris): n° 3 HG, slide Tg 1 1 8 . Paratypes d e p o s i t e d at t h e M u s e u m National d'Histoire Naturelle (Paris): n° Tg 1 1 9 ; T h e Natural His­

tory Museum ( L o n d o n ) : Reg. n° 2 0 0 0 . 9 . 1 4 . 1 .

Adults 1 4 3 4 ± 227.1 ( 9 9 2 - 1 9 2 9 ) long, 180 ± 2 1 . 6 ( 1 4 8 - 2 2 0 ) wide at the level o f the penis. Pharynx 7 3 ± 9.4 ( 5 8 - 9 5 ) wide. Dorsal gripus with blade regularly bent and well m a r k e d guard: a = 82 ± 3-1 ( 6 7 - 8 6 ) , b = 7 4

± 2.3 ( 6 9 - 7 9 ) , c = 4 ± 0.9 ( 2 - 6 ) , d = 19 ± 1.8 ( 1 5 - 2 3 ) , e = 3 3 ± 2.5 ( 2 3 - 3 7 ) . Large c u n e u s with very short extension: L = 39 ± 2.5 ( 3 2 - 4 6 ) , 1 = 16 ± 1.6 ( 1 1 - 1 9 ) , e - 3 ± 1 ( 1 - 7 ) . Nearly straight dorsal transverse bar:

x = 6 8 ± 2.5 ( 6 3 - 7 4 ) , w = 11 ± 1.1 (8-13). Ventral gripus with well marked aperture and well d e v e l o p e d guard:

a = 27 ± 0.8 ( 2 5 - 2 8 ) , b - 2 1 ± 0.9 ( 1 9 - 2 3 ) , c = 7 ± 0.8 ( 5 - 8 ) , d - 10 ± 1 ( 6 - 1 2 ) , e = 17 ± 0.9 ( 1 4 - 1 9 ) , L = 4 ± 0.7 ( 2 - 5 ) , 1 - 2 ± 0.5 ( 1 - 4 ) . Thin V-shaped ventral trans­

verse bar with b u b b l e d extremities: x = 41 ± 2 ( 3 7 - 4 7 ) , w = 8 ± 0.8 ( 6 - 1 0 ) . Uncinuli II - 17 ± 1.3 ( 1 3 - 1 9 ) , uncinuli I and III to VII - 15 ± 1.6 ( 1 1 - 2 1 ) . Long and

thin penis folded just after the o v o i d basal bulb, with a spirally c o i l e d thickening (5-7 turns) at its extremity, small heel: Pe = 106 ± 3.1 ( 9 8 - 1 1 3 ) , He = 5 ± 1.4 ( 4 - 9 ) . Long, straight and thin slightly sclerotised a c c e s s o r y p i e c e linked to the basal b u l b o f the penis: Ap = 4 5

± 3 ( 3 8 - 5 0 ) . Slightly sclerotised vagina with round well sclerotised part: diameter = 11 ± 1.3 ( 9 - 1 6 ) .

C o m m e n t s

B e s i d e s T. pangasi ( s e e a b o v e ) only two o t h e r s p e c i e s b e l o n g i n g to Thaparocleidus have already b e e n des­

c r i b e d from pangasiid hosts: T. caecus (Mizelle &

Kristsky, 1 9 6 9 ) and T. siamensis (Lim, 1 9 9 0 ) from P. hypophthalmus (syn. P. sutchi) ( s e e Lim, 1 9 9 0 ) .

This species is the only species o f Thaparocleidus with the dorsal gripus having a regularly a r c h e d blade, a spirally coiled thickening at the extremity o f the c o p u - latory tube and a sclerotised part o f the vagina.

Thaparocleidus chandpuri n. sp. is n a m e d after the type locality, Chandpur Riverine Station ( B a n g l a d e s h ) .

130. Mémoire Parasite, 2001, 8, 127-135

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MONOGENEANS FROM PANGASIIDAE

THAPAROCLEIDUS BAHARI N. SP. (Fig. 4 )

Type-host: Pangasius kinabatanganensis Roberts &

Vidthayanon, 1 9 9 1 . Site: gills.

Type-locality: Kinabatangan River, Sukau, Sabah State (North part o f B o r n e o Island), Malaysia.

Material studied: 3 0 individuals stained and m o u n t e d in Malmberg solution.

T y p e - m a t e r i a l : h o l o t y p e d e p o s i t e d at t h e M u s e u m National d'Histoire Naturelle (Paris): n° 2 HG, slide Tg 116. P a r a t y p e s d e p o s i t e d at the M u s e u m National d'Histoire Naturelle (Paris): n° Tg 1 1 7 ; T h e Natural His­

tory M u s e u m ( L o n d o n ) : Reg. n° 2 0 0 0 . 9 . 1 4 . 2 .

Adults 6 5 9 ± 107 ( 4 5 6 - 1 0 6 0 ) long, 124 ± 13.5 ( 8 8 - 1 5 7 ) wide at the level o f the penis. Pharynx 4 5 ± 4 . 8 ( 3 4 - 5 7 ) wide. Dorsal gripus with b l a d e b e n t in middle and very short guard: a - 53 ± 1.3 ( 5 0 - 5 6 ) , b = 4 3 ± 1.3 ( 4 0 - 4 5 ) , c - 1 ± 0.3 ( 0 . 4 - 2 ) , d = 14 ± 0.8 ( 1 2 - 1 7 ) , e = 28 ± 1.1 ( 2 5 - 3 1 ) . Cuneus with long extension: L = 1 6

± 1.1 ( 1 3 - 1 9 ) , 1 = 6 ± 0.6 ( 4 - 7 ) , e = 12 ± 1.6 ( 9 - 1 7 ) . Slightly curved dorsal transverse bar: x = 4 0 ± 1.3 ( 3 7 - 4 2 ) , w = 5 ± 0.4 ( 4 - 6 ) . Ventral gripus with well marked

aperture and short guard: a = 27 ± 0.6 ( 2 6 - 2 8 ) , b = 23

± 0.8 ( 2 1 - 2 4 ) , c = 1 ± 0.5 ( 1 - 3 ) , d = 9 ± 0.7 ( 7 - 1 1 ) , e

= 17 ± 0.6 ( 1 6 - 1 8 ) , L = 4 ± 0.6 ( 3 - 6 ) , 1 = 2 ± 0.4 ( 1 - 3 ) . Thin V-shaped ventral transverse bar: x - 39 ± 1.6 ( 3 5 - 4 3 ) , w = 3 ± 0.4 ( 3 - 4 ) . Thin uncinuli II = 15 ± 1 ( 1 3 - 17), uncinuli I and III to VII - 15 ± 1.7 ( 1 2 - 1 9 ) . 3- shaped, thin w a l l e d and large penis with ovoid basal bulb and large heel: Pe = 8 9 ± 5.9 ( 7 5 - 9 9 ) , He = 9 ± 1 ( 6 - 1 2 ) . Curved a c c e s s o r y p i e c e with slightly sclero- tised vast e x t e n s i o n linked to the basal b u l b o f the penis: Ap = 3 5 ± 1.5 ( 3 1 - 3 9 ) . No sclerotised vaginal system.

C o m m e n t s

This s p e c i e s is easily distinguished from T. caecus, T. chandpuri n. sp. and T. pangasi by having a large penis s h a p e d like a 3 {vs. T. caecus with thin and curved o n e and T. siamensis with a sinuous o n e ) and in having a long extension on the cuneus {vs. n o exten­

sion). Thaparocleidus babari n. sp. is p r o p o s e d for M. Basri B a h a r responsible for the Sukau River Lodge (Wildlife Expedition SDN. B H D . ) w h o kindly found, bought and provided the rare Pangasius kinabatan­

ganensis s p e c i m e n s .

Fig. 4. - Thaparocleidus bahari n. sp. C - cuneus;

DB = dorsal transverse bar; DG = dorsal gripus; MA

= male apparatus; VB = ventral transverse bar; VG

= ventral gripus; U = uncinulus. Bar = 30 um.

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THAPAROCLEIDUS SABANENSIS N. SP. (Fig. 5 )

Type-host: Pangasius kinabatanganensis Roberts &

Vidthayanon, 1 9 9 1 . Site: gills.

Type-locality: Kinabatangan River, Sukau, S a b a h State (North part o f B o r n e o Island), Malaysia.

Material studied: 6 individuals stained and m o u n t e d in Malmberg solution.

T y p e - m a t e r i a l : h o l o t y p e d e p o s i t e d at the M u s e u m National d'Histoire Naturelle (Paris): n° 5 HG, slide Tg 122. P a r a t y p e s d e p o s i t e d at the M u s e u m National d'Histoire Naturelle (Paris): T g 1 2 3 ; the Natural History Museum ( L o n d o n ) n° 2 0 0 0 . 9 . 1 4 . 3 .

Adults 1 2 8 9 ± 1 5 1 7 ( 1 0 1 7 - 1 4 6 2 ) long, 165 ± 3 7 . 3 ( 1 2 8 - 2 3 1 ) w i d e at the level o f the penis. Large and well m a r k e d pharynx: 9 4 ± 9.8 ( 7 9 - 1 0 7 ) at its width. Dorsal gripus with regularly a r c h e d blade and short guard: a

= 77 ± 3.1 ( 7 2 - 8 2 ) , b = 6 6 ± 2.6 (61-70), c = 3 ± 0.5 ( 2 - 4 ) , d = 15 ± 1.1 ( 1 4 - 1 8 ) , e = 34 ± 1 ( 3 2 - 3 6 ) . Large c u n e u s with very short extension: L = 3 5 ± 0.8 ( 3 3 -

3 6 ) , 1 = 10 ± 0.8 ( 9 - 1 1 ) , e - 3 ± 1.3 (2-6). Slightly curved dorsal transverse bar: x = 56 ± 1.6 ( 5 4 - 5 8 ) , w = 9 ± 0.7 ( 8 - 9 ) . Ventral gripus with small aperture and short guard: a = 3 3 ± 0.7 ( 3 2 - 3 4 ) , b = 27 ± 0.8 (26-28), c = 2 ± 0.7 ( 1 - 3 ) , d = 10 ± 0.7 ( 8 - 1 1 ) , e - 19 ± 1 ( 1 8 - 2 1 ) , L = 4 ± 0.5 ( 3 - 4 ) , 1 = 2 ± 0.3 ( 1 - 2 ) . Thin V - s h a p e d ven­

tral transverse bar: x - 3 6 ± 1.7 ( 3 4 - 3 9 ) , w = 4 ± 0.6 ( 4 - 6 ) . Thin uncinuli II = 18 ± 1 ( 1 6 - 1 9 ) , uncinuli I and III to VII = 17 ± 1.9 ( 1 2 - 2 0 ) . Curved and short penis with ovoid basal b u l b and m e d i u m heel: Pe = 56 ± 3.8 (52-62), He = 8 ± 1.9 ( 5 - 9 ) . Slightly curved and short a c c e s s o r y p i e c e with gutter-like d e p r e s s i o n , linked to the basal b u l b o f the penis: Ap = 31 ± 1 ( 3 0 - 3 3 ) . No sclerotised vaginal system.

C o m m e n t s

Only six specimens o f this species w e r e recovered from two o f five fishes sampled, w h i l e 1 2 5 - 3 5 0 s p e c i m e n s o f T. bahari n. sp. w h e r e found o n the s a m e fishes.

Thaparocleidus sabanensis n. sp. is easily distingui­

shable from T. siamensis and T. bahari b y the s h a p e

Fig. 5- - Thaparocleidus sabanensis n. sp. C - cuneus;

DB = dorsal transverse bar; DG = dorsal gripus; MA

= male apparatus; VB - ventral transverse bar; VG - ventral gripus; U - uncinulus. Bar = 30 urn.

132- Mémoire Parasite, 2001, 8, 127-135

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M O N O G E N E A N S F R O M P A N G A S I I D A E

of the penis (sinuous or M-shaped vs. c u r v e d ) ; from T. caecus, and T. pangasi mainly by having a very short e x t e n s i o n o n the large c u n e u s (vs. long e x t e n s i o n and small cuneus) and from T. chandpuri b y the shape and size o f the penis (simple and 5 6 urn long vs. spirally coiled thickening at the extremity and 106 p m long).

Thaparocleidus sabanensis n. sp. is n a m e d after the type locality ( S a b a h State).

THAPAROCLEIDUS MAHAKAMENSIS N. SP. (Fig. 6 )

Type-host: Pangasius nieuwenhuisii (Popta, 1 9 0 4 ) . Site: gills.

Type-locality: M a h a k a m River, Samarinda, East Kali­

mantan (Southeast part o f B o r n e o Island), Indonesia.

Material studied: 3 0 individuals stained and m o u n t e d in Malmberg solution.

T y p e - m a t e r i a l : h o l o t y p e d e p o s i t e d at t h e M u s e u m National d'Histoire Naturelle (Paris): n° 1 HG, slide Tg 1 1 4 . P a r a t y p e s d e p o s i t e d at the M u s e u m National d'Histoire Naturelle (Paris): n° Tg 1 1 5 ; T h e Natural His­

tory M u s e u m ( L o n d o n ) : n° 2000.9-14.4.

Adults 1 0 0 4 ± 157.5 ( 7 1 6 - 1 3 2 0 ) long, 182 ± 24.3 ( 1 3 5 - 2 1 8 ) wide at the level o f the penis. Pharynx 61 ± 6 ( 5 0 - 7 5 ) wide. Dorsal gripus with blade bent in the middle and very short guard: a = 59 ± 2.5 ( 5 3 - 6 5 ) , b

= 4 9 ± 2.1 ( 4 5 - 5 4 ) , c - 1 ± 0.3 ( 0 . 5 - 2 ) , d - 14 ± 1.3 ( 1 1 - 1 7 ) , e = 32 ± 1.5 ( 2 9 - 3 6 ) . Cuneus with medium extension: L - 1 3 ± 1.4 ( 1 3 - 1 9 ) , 1 = 5 ± 0.5 ( 4 - 7 ) , e = 8 ± 1.3 ( 4 - 1 0 ) . Nearly straight dorsal transverse bar:

x = 3 3 ± 1.4 ( 3 1 - 3 9 ) , w = 6 ± 0.4 ( 5 - 7 ) . Ventral gripus with well marked aperture and very short guard: a = 27 ± 0.7 ( 2 6 - 2 9 ) , b = 25 ± 0.7 ( 2 3 - 2 7 ) , c - 1 ± 0.3 ( 1 - 2), d = 5 ± 0.6 ( 4 - 7 ) , e - 1 6 ± 0.7 ( 1 4 - 1 7 ) , L = 6 ± 0.6 ( 5 - 8 ) , 1 = 3 ± 0.3 ( 2 - 3 ) . Thin V-shaped ventral trans­

verse bar: x - 3 4 ± 1.5 ( 3 0 - 3 8 ) , w = 4 ± 0.4 ( 3 - 5 ) . Thin uncinuli II = 13 ± 1 ( 1 0 - 1 6 ) , uncinuli I and III to VII

= 13 ± 0.7 ( 1 1 - 1 5 ) . Long, thin and regularly folded penis with a small spine-like structure at its distal third, an ovoid basal b u l b and a large heel: Pe = 169

± 7 ( 1 5 5 - 1 8 1 ) , He - 12 ± 1.3 ( 1 0 - 1 6 ) . Curved a c c e s ­ sory p i e c e with a central gutter-like depression, linked to the basal bulb o f the penis: Ap = 8 0 ± 3 7 ( 7 2 - 8 7 ) . No sclerotised vaginal system.

Fig. 6. - Thaparocleidus mahakamensis n. sp. C

= cuneus; DB = dorsal transverse bar; DG = dorsal gripus; MA - male apparatus; VB = ventral transverse bar; VG = ventral gripus; U" uncinulus. Bar - 30 µm.

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C o m m e n t s

This s p e c i e s is easily distinguished from:

- T. pangasi b y the size o f the a c c e s s o r y p i e c e o f the penis ( 8 0 urn vs. 54 p m ) , the c u n e u s ( 1 3 um vs. 2 0 pra in length), the p r e s e n c e o f a spine-like structure at the distal third o f the penis (vs. a b s e n c e ) and the s h a p e of ventral gripus (thick vs. thin) e v e n if the measure­

ments are similar;

- T. caecus by the sizes o f the sclerotised parts w h i c h are larger in the present species T. mahakamensis, even if the shapes are morphologically similar (i.e. s a m e spine-like structure at the distal third o f the penis), penis is 1 6 9 p m in length (vs. 62 p m ) , dorsal gripus 59 p m (vsAi p m ) , accessory p i e c e 8 0 p m (vs. 41 p m ) , - T. chandpuri b y the s h a p e o f the penis (spirally coiled thickening), the s h a p e and size o f dorsal gripus (regularly curved blade, total length 8 2 p m vs. b e n t in the middle, total length 59 p m ) and ventral gripus (well d e v e l o p e d guard 7 p m vs. very short outer root 1 p m ) ; - T. siamensis and T. bahari n. sp. b y the s h a p e o f the penis (regularly folded vs. sinuous or 3 - s h a p e d ) ;

- T. sabanensis n. sp. b y the size o f the sclerotised parts: i.e. penis ( 1 6 9 p m vs. 5 6 p m ) , cuneus (L = 13 p m vs. 3 5 p m or e x t e n s i o n = 8 p m vs. 3 p m ) , etc.

T h e n a m e Thaparocleidus mahakamensis n. sp. refers to the type locality.

THAPAROCLEIDUS REDEBENSIS N. SP. (Fig. 7 )

Type-host: Pangasius rheophilus Pouyaud & T e u g e l s , 2 0 0 0 .

Site: gills.

Type-locality: B e r a u River, Tanjung R e d e b , East-Kali­

mantan state (East part o f B o r n e o Island), Indonesia.

Material studied: 3 0 individuals fixed and m o u n t e d in Malmberg solution.

T y p e - m a t e r i a l : h o l o t y p e d e p o s i t e d at t h e M u s e u m National d'Histoire Naturelle (Paris):n° 4 HG, slide T g 120. Paratypes d e p o s i t e d at the M u s e u m National d'Histoire Naturelle (Paris):n° Tg 1 2 1 ; T h e Natural His­

tory Museum ( L o n d o n ) : n° 2 0 0 0 . 9 . 1 4 . 5 .

Adults 1 4 8 9 ± 180.6 ( 1 2 3 8 - 1 9 8 5 ) long, 2 3 3 ± 2 3 . 4 ( 1 7 5 -

Fig. 7. - Thaparocleidus redebensis n. sp. C = cuneus; DB = dorsal transverse bar; DG - dorsal gripus; MA = male apparatus; VB = ventral trans­

verse bar; VG = ventral gripus; U = uncinulus. Bar - 30 pm.

134 Mémoire Parasite, 2001, 8, 1 2 7 - 1 3 5

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M O N O G E N E A N S F R O M P Â N G A S I I D A E

2 9 5 ) wide at the level o f the penis. Pharynx 8 0 ± 6 ( 7 0 - 9 1 ) wide. Large dorsal gripus with blade bent in the middle and very short guard: a = 97 ± 2.9 ( 9 1 - 1 0 3 ) , b = 8 3 ± 2.9 ( 7 7 - 8 9 ) , c = 2 ± 0.6 (1-4), d = 21 ± 1.8 (16-25), e = 4 8 ± 2.3 ( 4 4 - 5 4 ) . Large c u n e u s with short e x t e n s i o n : L = 4 4 ± 2.1 ( 4 0 - 4 9 ) , 1 - 1 2 ± 0.9 (11-16), e = 8 ± 2.1 (5-12). Slightly curved and long dorsal trans­

verse bar: x = 6 6 ± 2.3 ( 6 2 - 7 1 ) , w = 9 ± 0.7 ( 8 - 1 2 ) . Ventral gripus with well m a r k e d aperture and short guard: a - 3 8 ± 1.3 ( 3 5 - 4 2 ) , b - 3 4 ± 1.1 ( 3 1 - 3 6 ) , c - 2 ± 0 . 6 (1-4), d - 11 ± 1 ( 9 - 1 3 ) , e = 25 ± 0.9 (22-26), L = 5 ± 0.6 ( 4 - 7 ) , 1 = 2 ± 0.4 (2-3). Thin and long V- s h a p e d ventral transverse bar: x = 4 7 ± 2 ( 4 2 - 5 3 ) , w

= 6 ± 0.5 ( 5 - 7 ) . Long uncinuli II - 1 9 ± 1.4 (14-22), uncinuli I and III to VII - 1 9 ± 3 ( 1 1 - 2 5 ) . Short large and 3-shaped penis with a long thin b r a n c h e d wall e x p a n s i o n at the first quarter, an ovoid basal bulb and a large heel: Pe = 6 8 ± 3-9 ( 6 0 - 7 8 ) , He - 9 ± 1.8 (7- 14). Curved a c c e s s o r y p i e c e with a central gutter-like depression, linked to the basal b u l b o f the penis: Ap

= 4 0 ± 1.6 ( 3 8 - 4 4 ) . No sclerotised vaginal system.

C o m m e n t s

This s p e c i e s is different from the other Thaparocleidus s p e c i e s from pangasiids b y the p r e s e n c e o f a b r a n c h e d e x p a n s i o n at the first quarter o f the penis wall and in having large anchors and bars. Thaparocleidus rede- hensis n. sp. is n a m e d after the type locality

C O N C L U S I O N S

T

h e present five n e w s p e c i e s bring the n u m b e r o f Thaparocleidus s p e c i e s found o n five spe­

cies o f pangasiids (P. pangasius, P. hypoph- thalmus, P. rheophilus, P. nieuwenhuisii and P. kina- batanganensis) to eight. In India only o n e s p e c i e s , T. pangasi, has b e e n r e c o r d e d on P. pangasius, while the s a m e host species in Bangladesh harbours two spe­

cies o f Thaparocleidus. T h e diversity o f m o n o g e n e a n s p e c i e s o n the five host s p e c i e s is low, ranging from o n e to t w o m o n o g e n e a n s p e c i e s per host s p e c i e s .

ACKNOWLEDGEMENTS

T

his w o r k was part o f a project on the charac­

terisation utilisation and m a i n t e n a n c e o f b i o ­ logical diversity for the diversification and sus- tainability o f catfish culture in Southeast Asia supported b y the E u r o p e a n Commission (grant I C 1 8 - C T 9 6 - 0 0 4 3 ) . W e w o u l d like to thank Dr G.C. Haldar (Chief Scien­

tific Officer), Dr A.K. Y o u s u f Haroun (Principal Scien­

tific Officer) and MD Rabiul Awual Hossain (Scientific Officer) o f the Chandpur Station in B a n g l a d e s h for assistance in getting Pangasius pangasius.

D r a w i n g s b y : Dr. N a t h a l i e Le B r u n , l a b o r a t o i r e G é n o m e , Populations, Interactions, UMR 5000, c c l 0 5 , Université Montpellier II, p l a c e E. Bataillon, 3 4 0 9 5 Montpellier c e d e x 0 5 .

R E F E R E N C E S

E U Z E T L. & P R O S T M. Report of the meeting on Monogenea:

problems of systematics, biology and ecology. In: Slusarski, W. (Ed.) Review of advances in parasitology. Warsaw:

P.W.N. Polish Scientific Publishers, 1981, 1003-1004.

G U S S E V A.V. Monogenoidea. In: Key to parasites of freshwater fish of the USSR. Bychovskaya-Pavlovskaya I.E. et aided.) Moscow-Leningrad: Academiya Nauk SSSR, 1962, 919 pp.

(In Russian: English translation IPST, Ser. No. 1136, Jeru­

salem, 1964).

LIM L.H.S. Silurodiscoides Gussev, 1961 (Monogenea: Ancy- rocephalidae) from Pangasius sutchi Fowler, 1931 (Pan- gasiidae) cultured in Peninsular Malaysia. Raffles Bulletin of Zoology, 1990, 38, 55-63.

LIM L.H.S. Thaparocleidus Jain, 1952, the senior synonym of Silurodiscoides Gussev, 1976 (Monogenea: Ancyrodiscoi- dinae). Systematic Parasitology, 1996, 35, 207-215.

M A L M B E R G G. On the occurrence of Gyrodactylus on Swedish

fishes. Skrifterutgivna av Sodra Sveriges Fiskeriforening, 1957, 19-76. (In Swedish, with description of species and a summary in English).

N ' D O U B A V., L A M B E R T A. & E U Z E T L. Gill parasites of the genus Quadriacanthus Paperna, 1961 from Heterobranchus lon- gifilis and H. isopterus with description of seven new spe­

cies from Côte d'Ivoire, West Africa. Systematic Parasito­

logy, 1999, 44, 105-118.

P A R I S E L L E A. & E U Z E T L. Gill parasites of the genus Cicblido- gyrus Paperna, I960 (Monogenea, Ancyrocephalidae) from

Tilapia guineensis (Bleeker, 1862), with descriptions of six new species. Systematic Parasitology, 1995, 30, 187-198.

P O U Y A U D L. & T E U G E L S G. Description of a new pangasiid cat­

fish from East Kalimantan, Indonesia (Siluriformes, Pan- gasiidae). Ichtyol. Exptor. Fresbwaters, 2000, 11, 193-200.

T R I P A T H I Y.R. Monogenetic trematodes from fishes in India.

Indian Journal of Helminthology, 1957, 9, 1-149.

Reçu le 2 novembre 2000 Accepté le 5 février 2001

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