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D e v e l o p m e n t b y in j e c t io n in S im u l iu m d a m n o s u m s . l .

o f t w o O n c h o c e r c a s p e c ie s f r o m t h e w a r t h o g

t o in f e c t iv e l a r v a e r e s e m b l in g t y p e d l a r v a e ( D u k e , 1 9 6 7 )

W A H L G .* & B A IN O . **

S u m m a r y :

Four wart hogs (Phacochoerus aethiopicus) examined in the Sudan savanna of North-Cameroon were all found infected with two types of skin microfilariae. One was O. ramachandrini Bain, W a h l and Renz, 1993, the adult worms of which live in the sub­

cutaneous tissues of the feet. The other, smaller type belongs to a new Onchocerca species, the adult worms of which were not yet found. O. ramachandrini-microfilariae were evenly distributed across the whole body surface, those of Onchocerca sp. were concentrated on the back. The two species of microfilariae were isolated from an infected hide, separated under the dissecting microscope and injected into the thorax of pupae-hatched S. squa­

mosum and S. damnosum s.str. females. Both filariae developed in both flies at high rates (33 - 47% of injected microfilariae) and without pathological forms to infective larvae (L3). Both L3-species had a caudal tip, were long, slender and very motile and had a conspicuous glandular oesophagus. L3 from O. ramachandrini- microfilariae had a long glandular oesophagus (55 % of total L3 length), a round head and measured an average of 9 5 5 μm long and 19.2 μm wide. L3 from the other microfilaria-species were shorter (845 μm, P< 0 .0 0 1) and thinner ( 1 6 .7 μm, P< 0 .0 0 1 ) and had a shorter glandular oesophagus (36%, P<0.001), a shor­

ter tail (P<0.01) and a conical head.

Both L3-species, by their caudal tip, their long and slender si­

lhouette, their great motility and their conspicuous glandular oeso­

phagus resemble non-O. volvulus filarial L3 known, since many years, to occur in "wild" S. damnosum s.l. in Cameroon (Type D larvae, Duke, 1967) and in Liberia (Agamofilaria Type VI, Voelker and Garms, 1972). During our study, 13 such larvae were found in 12 wild S. damnosum s.l. from two geographically different areas of North-Cameroon and all identified as O. ramachandrini.

The excellent development of the two Onchocerca species from the wart hog in S. damnosum s.l. after artificial infection, and the identification of all recently examined wild Type D larvae as O.

ramachandrini suggest that S. damnosum s.l. is a natural vector of O. ramachandrini and that most (if not all) of the Type D larvae in onchocerciasis vectors in North-Cameroon originate from wart hogs.

KEY WORDS : onchocerciasis. O.

ramachandrini. Onchocerca


Simulium damnosum

s.l. Type D larvae.


Type VI. suids. Cameroon, intra- thoracical injection.

* I n s titu t fü r T r o p e n m e d iz i n , E b e r h a r d - K a r l s - U n i v e r s i t ä t , W ilhelm strasse 27, D -72074 T ü b in g en , Germ any.

** Laboratoire de B io lo g ie Parasitaire, Protistologie, H elm inthologie CNRS LIRA 114, M uséum n ational d ’H istoire n aturelle, 61 rue de Bu ffon , 75231 Paris C éd ex 05, France.

Résumé : Ob t e n t i o nd el a r v e si n f e c t a n t e ss e m b l a b l e sa u x l a r v e sd e Ty p e D p a rin je c t io nc h e z Sim u u u md a m n o s u ms. l. d e s m i c r o f i l a i r e sd e sd e u xe s p è c e sd’ On c h o c e r q u e sd é c o u v e r t e s c h e zl e s Ph a c o c h è r e s

Quatre Phacochères (Phacochoerus aethiopicus) de la savane sou­

danaise au Cameroun sont tous quatre infestés par deux types de microfilaires dermiques. L'une correspond à O. ramachandrini Bain, W ahl et Renz, 1993, dont les adultes se trouvent dans les tissus sous-cutanés des pieds. L'autre, plus petite, appartient à une nou­

velle espèce d 'Onchocerque dont l'adulte n'a pas été découvert.

Les microfilaires d'O. ramachandrini sont uniformément réparties, celles d'Onchocerca sp. sont concentrées dans la région dorsale.

Des microfilaires prélevées dans une peau infestée ont été séparées sous la loupe binoculaire selon leur appartenance spécifique et injectées en intra-thoracique à des femelles de S. squamosum et S.

damnosum s. str. prélevées à leur sortie de la pupe. Les deux espèces de Filaires se développent en larves infectantes (L3) dans les deux espèces de Simulies avec un rendement élevé (33 - 4 7 % des microfilaires injectées), et sans anomalies morphologiques. Les deux espèces de L3 sont longues, grêles et très mobiles; elles ont une pointe caudale et un oesophage glandulaire très développé.

Les L3 d'O. ramachandrini ont un oesophage atteignant 5 5 % de la longueur du corps et une tête arrondie; elles mesurent en moyenne 95 5 μm de long et 19,2 μm de large. Les L3 de l'autre espèce sont plus courtes (8 4 5 μm, P<0,001) et plus minces ( 16,7 μm,

P < 0 ,0 0 1). L'oesophage est plus court (36 %, P<0,001), la queue plus courte (P<0,01) et la tête est conique. Les caractères communs aux deux espèces de L3 ressemblent à ceux des L3 connues depuis de nombreuses années chez les S. damnosum s. l. sauvages et dési­

gnées comme « Larves de Type D » de Duke, 1967 au Cameroun, ou comme « Agamofilaria Type VI » de Voelker et Garms, 1972, au Liberia. Au cours de notre étude, 13 larves de ce type ont été trouvées chez 12 S. damnosum s. 1. sauvages dans deux régions différentes du Nord-Cameroun et ont été identifiées comme O.

ramachandrini. Le parfait développement des deux espèces d'Onchocerques parasites des Phacochères, après inoculation intra- thoracique de S. damnosum s. I. et la présence du « Type D » dans les régions où se trouvent les Phacochères suggèrent que cette Simulie est le vecteur naturel de O. ramachandrini et que la plupart, sinon toutes les L3 de « Type D » chez les vecteurs de l'onchocer- cose humaine au Nord-Cameroun proviennent des parasites des Phacochères.


onchocercose. O . ramachandrini. Onchocerca sp. Simulium damnosum s. l. larve de Type D. A g a m o fila ria Type VI. phacochère.

Cameroun, injection intrathoracique.





n som e areas o f Africa the S im u liu m vectors o f O n c h o c e r c a v o lv u lu s, th e ca u sa tiv e a g en t o f hum an onchocerciasis (riverblindness), also fre­

q u en tly tran sm it fila ria e o f an im als (N elso n and Pester, 1962; Duke, 1967; Garms and Voelker, 1969;

V oelker and Garms, 1972, Renz et al., 1989). These

"anim al filariae" in the o n ch o ce rcia sis vecto rs are important for the epidem iology o f the human disease, because they are indicators o f the degree o f zoophily o f the vector populations, because they can hamper the co rrect assessm en t o f O. v olv u lu s-transm ission an d b e c a u s e th e y co u ld hav e an im m u n o lo g ica l impact in man (W ahl et al., 1994). Almost all o f the animal filariae so far found in onchocerciasis vectors are unidentified and only know n by their infective larvae (L3), which m orphologically differ from O. vol­

vu lu s ("n o n -O. volvulu s larvae"). However, onchocer­

ciasis vectors may also carry animal filariae, w hich closely resem ble O. volvulus, as has b een discovered by natural transmission experim ents o f the cattle-fila- ria O. o c h e n g i to S. d a m n o s u m s.l. (D enke and Bain, 1978). O ne com m on type o f n o n -O. volvulus larva in S. d a m n o s u m s.l. is longer, thinner and more motile than O. volvulu s and has a caudal tip and a conspi­

cuous glandular oesophagus. It was first described in Cam eroon as Type D larva (D uke, 1967). A similar larva was later described in Liberia as A g a m o fila r ia VI (V o elk er and Garms, 1972) and thought by the authors to be the same species as Type D. The g eo ­ graphical and seasonal pattern o f Type D fly-infec- tio n s in C a m e ro o n s u g g e s te d th a t th e s e la rv a e stemmed from gam e animals (Renz et al., 1989).

In search for the host and adult stages o f Type D lar­

vae, w e screened som e com m on wild mammals o f North-Cameroon for filariae and their possible deve­

lopm ent in S. d a m n o s u m s.l. A first paper described a new O n c h o c e r c a sp ecies, O. r a m a c h a n d r in i Bain, Wahl and Renz, 1993, discovered in the subcutaneous tissues o f the feet o f wart hogs ( P h a c o c h o e r u s a e th io - p ic u s). This paper reports the attempt to experim en­

tally infect S. d a m n o s u m s.l. with O. r a m a c h a n d r in i and with a second O n c h o c e r c a microfilaria, coinci- dently found in the sam e wart hogs. T he p ossible relation o f their L3 to Type D larvae is examined.


Study site


he game animals w ere exam ined at the hun­

ting lodge “Campement de Vogzoum ” on the river Vina du Nord, 160 km to the North-East o f Ngaoundé ré , North-Cameroon.

Isolation o f microfilariae

Large num bers o f microfilariae for the experim ental infection o f flies w ere obtained by incubating superfi­

cial skin strips from the neck and the feet o f an infec­

ted wart hog in 25 ml cone-bottom ed plastic tubes with RPM I-1640 medium, penicillin (200 U/ml) and streptomycin ( 2 0 0 ug/ml) for several hours at ambient temperature (28 ±3 °C).

Collection and identification o f S. d a m n o s u m s.l.

S. d a m n o s u m s.l. pupae w ere collected in the nearby rapids o f the Vina du Nord and left to em erg e in cages o f black cloth. Eight larvae w ere cytotaxonom i- ca lly id e n tified as S. s q u a m o s u m ( 6 la rv a e) and S. d a m n o s u m s.str. (2). Adult flies w ere accordingly identified by the colour o f the basal segments o f the antennae as S. s q u a m o s u m (≤ 2.5 pale segm ents) and S. d a m n o s u m s.str. (≥ 3 pale segm ents).

Experimental infection o f flies

Microfilariae which had emerged from the skin were transferred to fresh RPMI-1640 medium, with penicillin and streptomycin (see above), am photericin B (0.25 ug/ml) and fetal calf serum (1 0%) and injected in por­

tions o f 15 microfilariae/fly through fine glass capilla­

ries into the thorax o f pupae-hatched female flies. The flies were then kept individually in 2 ml gauze-cove- red plastic tubes in a humid cham ber in total darkness and at ambient temperature (28 ± 3 °C), feeding on a 30% sugar solution with penicillin, streptomycin and amphotericin (same concentrations as above).

Control o f identification o f injected microfilaria

In each trial som e o f the injection capillaries with the m icrofilariae separated under the dissecting m icro­

scope (50x) w ere blow n out into a drop o f glycerol on a slide, covered with a slip and sealed with nail polish for later identification under the com pound m icroscope (100 - 400x magnification).

M orphology o f infective larvae

Mature third stage larvae from the head, thorax or abdom en o f flies having reached infective age, w ere consid ered infective larvae = L3. T heir length was m easured through an ocu lar scale on a dissecting m icro sco p e at 50x m ag n ification . T h e o th er m ain parameters (maximal diameter, muscular oesophagus + buccal cavity, glandular oesophagus, intestine and tail) w ere m easured through a scale on a com pound m icroscope at 4 00x m agnification. Arithmetic m ean m easurem ents w ere com pared by the t-test.

Hide mapping

T he hide (w ithout head and tail) o f o n e wart hog


Fig. 1. - Distribution of O. ramachandrini and Onchocerca sp.

microfilariae in the hide of a Pbacochoerus aetbiopicus. Number of O. ramachandrini / Onchocerca sp. microfilariae emerged per 100 mg skin biopsy.

infected with both m icrofilaria-species was sampled on four equidistant sites on the back, the sides and the b elly and on ea ch leg by tak in g a squ are o f approxim ately 2 c m x 2 cm su perficial skin with a scalpel (Fig. 1). T he biop sies w ere w eighed, finely cut w ith sc isso rs and in c u b a te d in m edium (s e e above) for 4 hours. The microfilariae w hich em erged from each skin sample w ere quantitatively transferred to a slide with glycerol for later identification under the com pound m icroscope (see above).


C om m on o c c u r r e n c e o f t w o d e rm a l m i c r o f il a ­ r ia e ,




w o types o f dermal microfilariae were found in the wart hog : a larger one with a cylindrical head and lashing m ovem ents and a smaller one with an attenuated anterior end and sluggish, coi­

ling movements (Fig. 2, Table 1). The bimodal distri­

b u tio n o f th eir len g th s (Fig. 3 ), th eir d ifferen tial distribution in the hide (Fig. 1) and their development into different types o f infective larvae show that they are two distinct species. The larger one was identified as O. r a m a c h a n d r in i Bain, Wahl and Renz, 1993. The small one, the adult stages o f w hich w ere not yet found, is a new species. By the morphology of its L3 (see below ), it was assigned to the genus O n ch ocerca.

The two species w ere associated in all four wart hogs exam ined. W hereas O. r a m a c h a n d r in i- m icrofilariae were more or less evenly distributed across the whole hide, with a local maximum at the flanks, those o f

O n c h o c e r c a sp. w ere concentrated at the back and neck and were rare in the feet (Fig. 1).

D e v e lo p m e n t t o t h e in fe c tiv e s t a g e in


s q u a ­ m o su m AND



Experimental set-up

Microfilariae w ere isolated from the skin o f an infec­

ted wart hog and injected into nulliparous (pupae- h a tc h e d ) fe m a le flie s . T h e d e v e lo p m e n t o f th e microfilariae was monitored in flies dying until day 6

post infection (p.i.). All surviving flies w ere killed and checked for filarial larvae on day 7 p.i. Three inocula­

tion trials were conducted (Table 2). In Trial 1 and 2 microfilariae isolated from the neck w ere separated under the dissecting m icroscope into O. r a m a c h a n ­ d r in i and O n c h o c e r c a sp. prior to injection. In Trial 3 undifferentiated microfilariae from the feet w ere ino­

culated. This site was known, from hide-mapping, to contain almost exclusively O. r a m a c h a n d r in i. In all trials the “purity” o f the inoculum was checked by randomly leaving aside som e o f the injection capilla­

ries and identifying the microfilariae in them under the com pound m icroscope (Table 2).

Filarial developm ent in the three trials

In all trials the m icrofilariae d eveloped to infective larv ae at a h ig h rate (3 3 - 47 % ) an d p ra ctica lly without pathological stages (Table 2). The time span o f developm ent was slightly faster than for O. volvu ­ lu s and O. o c h e n g i under the same conditions (Wahl unpubl. data), with first sausage stages (L1 ) appearing already on day 1 p.i., second stage larvae (L2) on day 3 p.i., preinfective larvae on day 4 p.i. and mature third stage larvae (L3) in the head on day 5 p.i. In the three trials six S. s q u a m o s u m and two S. d a m n o s u m s. str. survived to infective age. Developm ent in both fly sp ecies w as equally good for both m icrofilaria species (Table 2). The total num ber o f L3 (both spe­

cies) recovered from the head, the thorax and abdo­

m en was 24, 17, and 6 respectively.

D escription o f O. r a m a c h a n d r in i and O n c h o c e r c a sp. infective larvae

Among the experim entally obtained infective larvae, two morphological types w ere found (Fig. 4, Tables 1 and 3) : a large one had a round head, a long tail and a conspicuously long glandular oesophagus, which made up about 1/2 o f the total L3-length. This type was found for all L3 in Trial 3 (injection o f almost pure samples of “big microfilariae”) and was identified as O. r a m a c h a n d r in i . T h e other type was shorter (P < 0.001) and thinner (P < 0.001) and had a conical



F ig . 2. - T h e tw o s p e c i e s o f d e r m a l m i c r o f ila r ia e fo u n d in

Phacochoerus aethiopicus : O. ramachandrini an d Onchocerca sp.

A-C : O. ramachandrini; A : silh o u ette, c e p h a lic h o o k in lateral view ; B : cep h alic region o f an o th er sp ecim en , h o o k in front view ; C : sam e sp ecim en , posterior region.

D -F : Onchocerca sp.; D : silhou ette; E : attenuated an terior region o f an o th er sp ecim en , h o o k in front view (o n th e left) o r lateral view (o n th e right); F : posterior region o f tw o sp ecim en s. (O n ly th e first c ep h alic n uclei, R1 cell and last caudal n uclei w ere draw n;

A-F : sca le = 30 μm).

Fig. 3. - B im o d a l le n g th d istrib u tio n o f derm al m icrofilariae from a Phacochoerus aethiopicus in fected w ith b o th sp e cies, O.

ramachandrini w ith lon g m icrofilariae (ra) and Onchocerca sp. w ith sh orter m icrofila­

riae (sp ).

A bscisse : classes o f length : 220 to 229 μm , 230 to 239 μm, etc.


1. MICROFILARIAE (n ) 35 29

LENGTH (μm) ± SD (RANGE) 301 ± 1 2 (272-333) 251 + 10 (227-272)

DIAMETER (|am) ± SD (RANGE) 7.4 ± 0.3 (6.7-8.4) 6.9 ± 0.5 (5.7-7.4)

ANTERIOR END cylindrical attenuated

BEHAVIOUR lashing front end sluggish coiling

LOCALISATION IN THE HIDE evenly distributed neck and back

2. INFECTIVE LARVAE (n ) 17 1 2

LENGTH (μm) ± SD (RANGE) 955 ± 55 (850-1040) 845 ± 65 (750-1000)

DIAMETER (μm) ± SD (RANGE) 19.2 ± 0.8 (17.8-21.2) 16.7 ± 0.8 (16.0-18.5) GLAND. OES (% L3-length) ± SD (RANGE) 55.5 + 2.3 (52.3-61.5) 36.3 ± 1.9 (32.6-39.2) TAIL (% of L3-length) ± SD (RANGE) 5.7 ± 0 .4 (5.1-6.3) 5.2 ± 0.4 (4.7-6.0)

HEAD rounded conical

T a b le 1. - D ifferential characters o f O. ram achan drin i and O nchocerca sp.

(resp ectiv ely left and right colu m ns)


number of capillaries tested 1 2 4

number of O. ramachandrini identified 1 2 0 56

number of Onchocerca sp. identified 0 25 7


number of flies injected (15 mff/fly) 14 18 2 2

number of SQUAMXDAM surviving 1 / 0 2 / 1 3/1

number of SQUAM\DAM with L3 1/- 2 / 1 3/1

total number of L3 in SQUAM\DAM 6/- 10/5 19/7

% development in SQUAM\DAM 40/- 33/33 42/47

T ab le 2. - D evelo p m en t o f O. ram achan drin i (o r) and O nchocerca sp. (o sp ) in S. squam osum and S. dam nosum s.str., follow ing intrathoracical injection

head, a shorter tail (P<0.01) and a shorter glandular oesophagus (P<0.001), which made up only about 1/3 o f the total L3-length. All L3 from Trial 2 (injection of selected “small m icrofilariae”) corresponded to this second type and w ere identified as O n c h o c erc a sp., with the ex cep tio n o f o n e 0 . r a m a c h a n d r in i - L3, w hich ap p aren tly d ev elo p ed from an accid en tally injected “big microfilariae” (Table 3). Both L3-species w ere long and slender in general appearance and very motile, when recovered from fresh flies, with quick, cu rlin g m o v em en ts. T h ey all had a co n sp icu o u s, clearly visible glandular oesophagus (Fig. 4 A,F), a laterally flattened buccal cavity and a caudal tip, which was most com m only ben t ventrally or dorsally and only rarely straight (Fig. 4 D, E, H, I).


T w o

On c h o c e r c a s p e c i e s f r o m t h e w a r t h o g i n C a m e r o o n


his paper describes the microfilaria and the in fective larvae (L3) o f tw o filarial sp ecies from the wart hog, P h a c o c h o e r u s aeth iop icu s, in Cameroon. O ne o f them is O n c h o c erc a r a m a c h a n ­ d rin i., the L3 o f which were unknown sofar. The other is a new species, the adult worms of which were not yet found. Due to the morphology o f its infective lar­

vae the new species can be assigned to the genus O n c h o c erc a ( in Bain and Chabaud, 1986). In view o f the high infection rate o f P. a e th io p ic u s (4 o f 4 exam i­

ned animals infected with both species), the wart hog seems to be a major host o f the two filariae.



Fig. 4. - T h e infective larvae o f O. ram achan drin i an d O nchocerca sp.

A-E : O. ram achan drin i; A : silhou ette, lateral view ; B : an terior region, left lateral view ; C : h ead , m edian view ; D , E : tail, ventral and left lateral view respectively.

F-I : O nchocerca sp.; F: silhou ette, lateral view ; G : head, lateral view ; H, I : tail, ventral and right lateral view respectively;

(A, F: sca le = 2 00 μm; B,D ,H ,I: scale = 3 0 μm; C, G: scale = 10 μm ).


FLY/LOC LENGTH DIAMETER M. OES G. OES INTEST TAIL L3 IDENT (μ m) ( μim ) (% o f len gth ) (% o f length) (% o f length) (% o f length)


F1H1 840 16.0 17.1 37.8 40.0 5.0 O SP

F1H2 800 16.0 18.8 35.4 40.3 5.5 OSP

F1T1 820 17.0 17.4 36.7 41.2 4.7 OSP

F1T 2 800 / 15.9 33.7 45.1 5.3 OSP

F2H1 860 16.0 17.3 39.2 38.3 5.0 OSP

F2H 2 800 16.5 19.3 37.0 38.8 5.0 OSP

F2H 3 750 17.3 15.4 35.9 43.6 5.1 OSP

F2H 4 900 17.3 16.1 32.6 46.3 4.9 OSP

F2H5 850 16.1 13.2 38.0 42.7 6.0 OSP

F2H 6 900 18.5 18.2 36.5 39.6 5.6 OSP

F2H 7 1000 16.1 / / 42.0 4.9 OSP

F2T1 820 16.8 16.1 36.7 41.9 5.3 OSP

F3T1 8 50 19.8 16.0 57.5 21.4 5.1 OR


F1H1 1010 19.5 14.8 56.8 22.2 6.2 OR

F1H2 1000 21.2 16.5 54.5 23.6 5.3 OR

F1H3 9 40 18.5 16.4 57.5 20.9 5.2 OR

F1H4 9 60 20.0 14.5 53.8 26.2 5.5 OR

F2T1 9 40 19.0 16.4 57.0 20.7 5.8 OR

F2T2 9 50 18.8 16.0 52.8 25.3 5.9 OR

F3H1 9 90 19.8 16.8 61.5 15.4 6.3 OR

F3A1 1040 18.5 16.6 53.9 24.0 5.5 OR

F3A2 990 18.5 17.1 55.0 22.2 5.8 OR

F3A3 8 80 18.5 16.2 53.3 24.8 5.7 OR

F3A4 9 90 18.5 15.4 56.6 22.6 5.4 OR

F4H1 1040 19.5 16.8 55.7 21.5 6.0 OR

F4H 2 9 40 19.8 15.6 52.9 25.2 6.3 OR

F4H 3 9 00 17.8 14.7 55.2 24.2 5.9 OR

F4H 4 900 18.5 16.5 52.3 25.8 5.4 OR

F4T1 920 19.8 16.0 57.2 20.9 5.9 OR

T ab le 3. M orphology o f infective larvae from tw o injection trials

FLY/LOC = number o f fly and localisation w here L3 were found (F=fly number, H=head, T=thorax, A=abdomen); M. OES = muscular oeso­

phagus + buccal cavity; G. OES=glandular oesophagus.

D e v e lo p m e n t in Sim uliu m v e c t o r s

Com plete extrinsic developm ent o f the two species was obtained by intrathoracical injection o f microfila­

riae into pu pae-hatched S. d a m n o s u m s.l. fem ales.

D evelop m ent w as excep tio n ally g ood and w ithout pathological forms, so that, despite the artificial route o f infection used, it can be concluded with near cer­

tainty that both species are naturally transmitted by S im u liu m sp ecies. This is the first acco u n t o f life cycles o f O n c h o c e r c a species o f suids.

R e l a t i o n o f t h e t w o On c h o c e r c a sp ecies fr o m THE WART HOG WITH NON-O . VOLVULUS LARVAE IN WILD ONCHOCERCIASIS VECTORS

The infective larvae o f both species, by their long and slender silhouette, their caudal tip, their great motility an d th e ir c o n s p ic u o u s g la n d u la r o e s o p h a g u s , resem ble a certain form o f “wild” filarial larva com ­ monly found in onchocerciasis vectors in Africa : it was first described in Cameroon and named Type D (Duke, 1967). This form o f larva has since been fre­

quently found by other authors in Cameroon, as well

in S. d a m n o s u m s.l. (Franz and Renz, 1980; Renz et a l ., 1989; Schibel and Renz, 1990; Wahl et a l., 1994) as in S. h o v is (R en z et a l., 1989; W ahl and Renz, 1991). A similar type o f larva was found in S. d a m n o ­ su m s.l. in Liberia and named A g a m o fila r ia Type VI (V oelker and Garms, 1972). The authors thought it to b e “[...] m ost p ro b a b ly id e n tica l w ith T y p e D in Cameroon [...]”. Long and slender n o n -O. volvulus L3 with a caudal tip w ere also found in other parts o f the O nchocerciasis Control Programme area (B ak er and Bain, unpubl. data).

The (retrospective) synonym ization o f our tw o L3- species from the w an hog with the historical accounts o f T y p e D- and/or A g a m o f i l a r i a T y p e VI larvae seem s, how ever, im possible: although m ost o f the m o rp h o lo g ic a l c h a r a c te rs g iv en by V o e lk e r and Garms (1972) for A g a m o fila r ia Type VI larvae corres­

pond to O. r a m a c h a n d r i n i (leng th : 9 15-985 μm, glandular o eso p h ag u s : 49% o f total length, tail : 5.8% ), its inferior diam eter (m ean : 17.5 μm ) pre­

cludes a formal synonymization. Many o f the charac­

ters o f Type D larvae given by Duke (1967) show a wide range (length : 750 - 925 μm, diameter : 15 - 20



μm, glandular oesophagus 270 - 475 μ m) and thus cover the m orphology o f both O. r a m a c h a n d r in i and O n c h o c e r c a sp . M o re o v e r , th e a b u n d a n c e o f O n c h o c e r c a species in game animals in Africa (Bain et al., 1978; D enke and Bain, 1981), all o f them with unknow n life cycles, cautions to consider that other L3-species with similar morphology may occur.

H o w e v e r, s in c e th e d e s c r ip tio n o f th e tw o O n c h o c e r c a L3 from wart hogs, w e w ere able to exa­

m ine 13 Type D-like larvae from 11 wild S. d a m n o ­ s u m s .s tr . an d o n e S. s q u a m o s u m in tw o geographically different regions in North-Cameroon.

Significantly, they w ere all m orphologically indistin­

guishable from O. r a m a c h a n d r in i.

On c h o c e r c a sp ecies f ro m w a r t h o g s in t h e EPIDEMIOLOGY OF HUMAN ONCHOCERCIASIS

In view o f the ex cellen t developm ent o f O. r a m a ­ c h a n d r i n i in je c te d in to S. d a m n o s u m s.str. and S. s q u a m o s u m and its L3 being indistinguishable from

“wild” Type D-like larvae found in these fly species, it seem s alm ost certain that S. d a m n o s u m s.str. and S. s q u a m o s u m are natural vectors o f O. r a m a c h a n ­ d rin i. W e can n o t sp ecu late at this stage o f know ­ le d g e , w h e th e r O n c h o c e r c a sp. is also n atu rally transmitted by S. d a m n o s u m s.l. or is insusceptible, nor w hether S. d a m n o s u m s.l. is the only natural vec­

tor o f O. r a m a c h a n d r in i (and O n c h o c e r c a sp. ?), nor can w e exclude that there are other unknow n infec­

tive larvae with Type D m orphology. However, the coincidence o f all recently exam ined Type D larvae in North-Cameroon being identified as O. r a m a c h a n ­ d r in i and the abundance o f P. a e th io p ic u s in this area, strongly suggests that many (if not all) o f the Type D larvae in S. d a m n o s u m s.l. in Cam eroon (and o th er parts o f Africa ?) stem from w art hogs. The annual prop ortion o f Type D larvae in the Sudan savanna o f Cam eroon is as high as one fifth o f all fila­

rial infections (W ahl et al., 1994). This underlines the high d egree o f zoophily o f the vecto r populations and raises questions as to a possible immunological impact o f these animal filariae in man.



e are grateful to Mr. D. Ekale, Programme O n c h o c e r c o s e s H u m aine et A n im ales, Ngaoundé ré , Cameroon for the cytotaxo- nom ic identification o f the S. d a m n o s u m s.l. larvae and for technical assistance. W e are much indepted to Mr. and Mrs. Koufou, ow ners o f the Campement de V ogzoum , for their kind perm ission to exam ine the hunted anim als and to the M inistry o f H igher Education, Scientific Research and Computer Sciences

o f Cam eroon for granting a research permit to one o f us (G .W .). This investigation received financial su p - port from the Commission o f European Communities (TS2/0184-D(AM) and TS3/CT92-006).


Bain O., Ch a b a u d A.G. & La n d a u I. Trois nouvelles oncho- cerques chez des céphalophes au Gabon. A nnales d e Parasitologie H um aine et Comparée, 1978, 53, 403-419.

Bain O. & Ch a b a u d A. G . Atlas des larves infestantes de Filaires. Tropical M edicine a n d Parasitology, 1986, 37, 301-340.

Bain O., Wahl G . & Ren z A. O nchocerca ram achan drin i n.sp. from the wart hog in Cameroon. A n n a les d e Parasitologie H um aine et Comparée, 1993, 68, 139-143.

Denke A.M. & Bain O. Données sur le cycle d’Onchocerca ochengi chez Simulium dam nosum s.l. au Togo. Annales de Parasitologie H um aine et Comparée, 1978, 53, 757-760.

Den k eA.M. & Bain O. Deux nouvelles onchocerques nodu- laires chez des Bovidés sauvages en Haute-Volta.

A nnales d e P arasitologie H u m ain e et Com parée, 1981, 56, 339-347.

Du k e B.O.L. Infective larvae, other than O nchocerca volvu­

lus in Simulium dam nosum . Annals o f Tropical M edicine a n d Parasitology, 1967, 61, 200-205.

Franz M. & Ren z A. Scanning electron microscope study of infective filaria larvae of Type D and O nchocerca volvu­

lus. Tropical M edicine a n d Parasitology, 1980, 3 1, 31-33.

Garm s R. & Vo e l k e rJ. Unknown filarial larvae and zoophily in Simulium dam nosum in Liberia. Transactions o f the Royal Society o f Tropical M edicine a n d Hygiene, 1969, 63, 676-677.

Nelson G.S. & Pe ster F .R . The identification of infective filarial larvae in Simuliidae. B u lletin o f th e W orld H ealth Organization, 1962, 27, 473.

Renz A., Sc h ib e lJ.. Eic h n er M. & En y o n g P . Animal filariae in anthropophilic Sim ilium spp. in North Cameroon.

Tropical M edicine a n d Parasitology, 1989, 40, 386-387.

Sc h ib e l J.M. & Re n z A. Differentiation of non-O nchocerca volvulus filarial larvae in anthropophilic Simulium spp.

in North Cameroon. Acta Leidensia, 1990, 59, 460-461.

Vo e l k e r J. & Ga r m s R . Zur Morphologie unbekannter Filarienlarven aus dem O nchozerkoseuebertraeger Simulium dam nosum und aus S. kenyae in Liberia und zur Frage der moeglichen Endwirte. Tropical M edicine a n d Parasitology, 1972, 23, 285-301.

Wahl G. & Ren z A. Transmission of O nchocerca du kei by Simulium bovis in North-Cameroon. Tropical M edicine a n d Parasitology, 1991, 42, 368-370.

Wahl G., En y o n g P ., Sc h ib e lJ.M., Ng o s s o A., Tu b b e s in g II.,

Mo y o u R. & Ren z A. Transmission of O nchocerca volvu­

lus, O. ochengi and Type D filariae by Simulium d am n o­

sum s.l. in North-Cameroon and its influence on the epidemiology of human onchocerciasis. P arasite, 1, 1994, Suppl. 7-10.

Accepté le 13 novembre 1994


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