Cytotaxonomy of S.damnosum complex: January- December 2000

aWORLD HEALTH ORGANIZATION Afllc^t{ lEcloN OROANISATION MONDIALE DE LA SANTE REG()N DE L'^fTIQTJE oMs/oNCHO, B.P.36 KARA, REPUBLIQUE TOGOLAISE T6l. 60.60.75 160.06.26 (Fax.60.06.26) STIBJECT : Cytotaxonomy of S. damnosum complex PERSONNEL : - George K. Fiasorgbor (Technical Officer) Coulibaly Yaya (Technician) Sedou Naniogud (Technician) PERIOD COVERED : January - December 2000 SIGNATT]RE OF' RESPONSIBLE OFFICER DATE OF SUBMISSION : 06 FEBRUARY 2001 CONTENTS I. INTRODUCTION 2. CYTOLOGICAL IDENTIFIATIONS 3. MATERIALS / METHODS 4. RESULTS 5. TT{E DISTRIBUTION OF THE CYTOPECIES OF THE SIMULTUM DAMNOSUM COMPLEX IN THE YEAR 2OOO. 6. RIVERS WITH MALE POPULATIONS OF S. SIRBANUM WITH OR WITHOUT TT{E CHARACTERISTIC SEX.LINKED IS.3 INVERSION 7. THE "NILE" FORM OF S. DAMNOSUM (S.S.) AND ITS DISTRIBUTION IN TFM OCP AREA IN TFM YEAR 2OOO. 8. LABORATORY REARING OF MEMBERS OF TI{E SIMULIUM DAMNOSUM COMPLEX tN TTIE YEAR 2OOO. t ! 9. TRAINING alO.MEETINGS 1 1 .OBSERVATIONS / COMMENTS l2.ACKNOWLEDGEMENTS 2 t 3 1. INTRODUCTION The period under review covers January to December 2000. In the course of this one year, the work load in the two cytotaxonomic laboratories of Odiennd in C6te d'Ivoire and Kara in Togo had more than doubled. Laboratory rearing of all members of the S.damnosun complex in the different seilsons of the year (montde des eaux, hautes eaux, ddcrue "p6riode d'harmattan et 6tiage") was introduced in March 2000 and was supposed to end in March or April2001. Activities related to the rearing process, coupled with the routine cytotaxonomic identifications, susceptibility tests and the frequent call on the reasearch team members to assist in other domains, were to the extent that, every member of the team was obliged to sacrifice most of the week-ends (saturdays and sundays) throughout the year. The calender for our "Annual leave" days was also not fully respected in the year. 2. CYTOLOGICAL IDENTIF'ICATIONS Routine identifications were carried out on the larvae of S. damnosum (s.1.) from all the river basins still under larviciding control in the Onchocerciasis Control Programme Area. In the East, these include the five major river basins ( Volta, Mono, Asukawkaw, Oueme, Niger) and their tributaries. In the West, the rivers include the Kaba, Niger, Tinkisso, Niandan, Milo, Sankarani, Bani, and the Sassandra in Ivory Coast. In the year 2000, a total of Eight Thousand Three Hundred and Ten ( 8310 ) larvae of S. damnosum ( s.l. ) were cytologically identified by the two cytota(onomy tearns of the OCP. 3. MATERIALS / METHODS MATERIALS Specimens on which this report is based came from there (3) main sources: (a) Specimens from routine checks by the Operational Bases. (b) Specimens from susceptibility tests. (c) Specimens originating from the laboratory rearing of members of the S. damnosum (s.1.) complex. METHODS Larrtae of the S. damnosum species complex were preserved in carnoy's fixative (3 parts absolute alcohol I part glacial acetic acid). The samples were stained by feulgen and chromosome preparations made as described by Dunbar (1972) and Boakye (1988). Identifications were made using the chromosome maps of Vajime and Dunbar (1975), Meredith et al. (1986) and Surtees et al. (1e88). 4t. 4. RESULTS CYTOLOGICAL IDENTTFICATIONS ( 2000 ) ZONE EAST ZONE WEST BASINS TOTALIDENTIFIED BASSIN TOTAL IDENTIFIED VOLTA 2767 MILO 104 MONO t532 SANKARANI 208 ASUKAWKAW 338 MADAN 74 OUEME 2096 TINKISSO 318 MGER (Benin) 154 NIGER ( Mali ) l3r Kaba 6 SASSANDRA 219 NIGER ( Guinea 363 TOTAL 6887 1423 GRAND TOTAL 83r0 TIIE DISTRIBUTION OF'TIIE CYTOSPECIES OT' S. DAMNOSUM COMPLEX IN TIIE YEAR 2OOO. S. sirbanum : In the course of the year 2000, S.sirbanum was registered on all the river basins in the OCP with the exception of the Kaba river in Guinea and the tributaries of the Volta Lake in Ghana/Togo. It has a large distribution range in the northern half of the Programme and makes periodic movements to the south in the dry season, especially in the harmattan. S. damnosum (s.s.) was recorded in sympatry with S. sirbanum on almost all the rivers in the OCP except on the Kaba in Guine4 Niger in Guinea and the Niger in Mali. It is found mainly in the transition zone between the forest and the savanna. S.squamosum : This species was observed in the West on the Kaba, Niandan, Milo, Sankarani and Sassandra. In the East, it was registered in its natural habitat in the mountaneous forest regions east of the Volta Lake in Ghana and Togo. The main rivers are Asukawkaw and its tributaries, the Mono and its tributaries, and on the tributaries of the Kara and the Mo. Few cases of S.squamosum were also found on River Zou, a tributary of the Oueme River in Benin. S.yahense : The breeding site of Tsatsadu Falls on the Tsatsadu River in Ghana is the only point from where this species was identified in the course of the year 2000. S.soubrense : The "Beffa" form of S.soubrense was observed in the year 2000 on R. Mono at Landa-Mono in Togo. In Benin it was registered on the Oueme River at Agolin-Pahou, M'betekoukou, Atchakpa, and Gba. On the Okpara, it was found at Bassa-Barrage and Kaboua. 56. No case of S. souDrense has ever been recorded in Ghana since the inception of OCP operations in the country in 1975. RIVERS WITH THE POPT]LATION OF'S. SIRBANT]M WITH OR WITHOUT TIIE CHARATERISTIC SEX.LINKED IS.3 IIYVERSION Two forms of S. sirbanum are known to exist in the Onchocerciasis Control Programme Area. Cytologically, almost all the male populations of the first group, the northern population or in other words, the monomorphic form of S. sirbanum lack the characteristic sex-linked Is-3 inversion. It was first described as "Sirba" form in 1973 by Dr. C.G. Vajime. In the eastern part of the OCP, cases of this first group of S. sirbanum were recorded on the following rivers:Niger, Sirba, Goroubi, Tapoa, Dyamongou, Alibori, Mekrou, and oti/Pendjari. The second group of S. sirbanum, the southern population or the polymorphic form, was also described inl973 by Dr. Vajime. It was originally separated form S.sirbanum as a separate species and referred to as S. sudanense. Its specific status as a separate species was widely criticised by other cytoto(onomists who consider it as just a form of S. sirbanum species. The male populations of this form always possess the characteristic sex-linked IS-3 inversion. Rivers on which this form of S. sirbanum were registered in the OCP area include the Bandama Blanc, Bas Bandama, Sassandr4 Marahou6, Volta Noire, Volta Blanche, Comoe, Asukawkaw, Mono, and their tributaries. All cases of S. sirbanum recorded on the Tano and Pra rivers in South-west Ghana belong to this form. Studies on the adult populations of S. sirbanum by R.A. Cheke (pers. Communication) also confirmed the existence of two forms of the species in the OCP area. During morphometric studies, Dr. Cheke, working as a WHO Consultant for the OCP, observed that populations of S. sirbanum having males with type IV scutal pattems were associated with male larvae lacking the IS-3 inversions. In contrast, those populations with type III scutal patterns were found with larvae which possess the IS-3 in versions. On the basis of cytota(onomic identifications and morphology of the adult flies, it is quite apparent therefore that two forms of S. sirbanum ( northem and southern forms) exist in the OCP area. A publication on this subject appeared in 1992 in Medical and Veterinary Entomology (1992) 6,139-142: cytotaxonomic confirmation of two forms of Simulium sirbanum in the Eastem part of the Onchocerciasis Control Programme in West Africa by G.K Fiasorgbor and R.A Cheke. THE "NILE" ['ORM OF S. DAMNOSUM (S.S) AIYD ITS DISTRIBUTION IN TIIE OCP AREA IN THE YEAR 2OOO. As in the case of S. sirbanum two forms of S.damnosum (s.s.) are known to exist in west Africa (Dunbar and Vajimd ( unpublished). These two authors indicated that one form (the "Volta" form) occurs in the OCP area and the other form (the'Nile" form) in Nigeria and some places east of the oCP. The'Nile" form has an inversion on chromosome 2 which has never been recorded in the OCP until in 1988. This was right in the harmattan (29llll88) which was quite severe and very much prolonged in the year. Since 1988 to date, the "Nile" form continues to spread to many rivers in the eastern part of the OCP especially into Benin and Togo. In the year 2000 this form of S. damnosum (s.s.) was recorded at the following breeding points: 7, 6DATE BREEDING SITE RIVER 04.01.2000 04.0r.2000 13.01.2000 13.01.2000 18.01.2000 18.01.2000 18.01.2000 19.01.2000 19.01.2000 02.02.2000 09.02.2000 24.02.2000 17.05.2000 10.08.2000 17.08.2000 0s.09.2000 l1.09.2000 Atchakpa Kaboua Sarakawa Barrage Dokomey Bagan Landa-Pozanda Agninda M'bdtdkoukou Al6hdridd Kpani Banage W6ria Djabignon Barrage Bassa Tchakparou Bogbohonou Barrage Biro Barrage Atagara Oudm6 Okpara Kpdlou Doko M6 Kara Kara Oudm6 Boual6 6ti Toussi Kara Okpara Oudora Dra I Souansird soubourdkou CHROMOSOMAL CHARACTERISTICS Morphologically the chromosomes of the two forms look alike. The banding pattern of the two is just the same. The difference between the two forms is largely based on inversion differences and sex-determining systems as follows: CHROMOSOME INVERSION DIFF'ERENCES -On chromosome I the t'wo populations of the "Volta" form and the "Nile" form share three floating inversion. : IS-3; lS-2;lL-2. -On chromosome 2 the two populations share a common standard sequence IIL-3. Inversion IIL-8 is present in the "Volta" form and it is lacking in the'Nile" population. -On chromosome 3 the heterozygous inversion IIIL-7 is absent from the'Nile" population but is found in about 30-40% of the "Volta" population. DIF'F'ERENCES IN SEX DETERMINATION ln the "Volta" form sex determination is on chromosome IIL and associated with inversions IIL-3 and IIL-8 whilst in the "Nile" form this is found on chromosome IS and in associated with the sex linked inversion IS-3 and IS-2. LABORATORY REARING OF' MEMBERS OF' TIIE SIMULIUM DAMNOSUM COMPLEX IN THE YEAR 2OOO. Single egg-batches of three members of the S. damnosum (S.l) complex were successfully reared to adult flies in the laboratory in 1995 and 1996. These were the Beffa form of S. soubrense from M'b6tikoukou and Kaboua on the Ou6md and Okpara rivers in Benin; S.yatrense from Tstsadu Falls in Ghana and S. squamosum from Kouniohou on R. Ban-Hon in Togo. 8. 7The main objective of the rearing exercise in 1995 and 1996 at M'bdtdkoukou and Okpara was to provide enough samples of dark-wing tuft adult females and larvae of the S.damnosum (theobald) complex for DNA, ta<onomic and cytotaxonomic determinations. Before this date, these dark-wing tuft adult females from the Oudmd and Okpara rivers in Benin were scored by the DNA technique as savanna cytospecies (S.damnosum (s.s) and S. sirbanum). The DNA determination was contrary to all previous cytotaxonomic indentification of the same populations as the Beffa form of S. soubrense, which infiltrates into the OCP area from Nigeria each year during the rainy season. lt was therefore an attempt to investigate this discrepancy between the DNA and cytota(onomic identificationsthat VCU hierarchy called for the rearing of wild-caught females of the S. damnosum complex in the Oudm6 Basin to provide more samples to confirm once and for all the actual identity of these biting adult females. Rearing work was also undertaken on the Comoe river in Zone West between 1995 andl996 with S.sanctipauli(s.s) as the target cytospecies. In the year 2000, the same exercise was carried out in the two zones (East and West) with the same objective. In addition, the rearing was supposed to cover all the various members of the S. damnosum complex in the OCP area. This was to provide enough reference samples as the Programme draws to an end in2002. We report below the full results of the study in the different river basins ( see cytotaxonomic identification results in table 2 ) of this report. 9. TRAINNING - The following four Laboratory Assistants in Zone East who were initiated in 1998 in VCU technical activities (dissection , prospection, ground larrriciding treatrnents etc.) have completed their training and issued with dissection codes. They are: Atakpam6: Allassani Abdulaye: code N" 625 Parakpou: Boukari Gibrilou Arimouyaou: code N" 626 Bohicon: Houessou Ahognon : code N' 627 Kara : Batascome Tchilabalo: code N'628 The issue of dissection codes qualifies the four assistants to take effective part in the weekly routine dissection activities in their respective bases. In addition, they can now officially run the bases at least temporarily in the absence of the chiefs when they are either on " annual Leave" for some weeks only or on missions. Henceforth, there is no need for sending a technician from the Kara Base to do the interim for the base chiefs when they are away. [t is now time for the laboratory assistants to play more roles in the technical activities of the Vector Control Unit of the OCP. - As the Programme draws to an end in 2002, it is very doubtful if aerial treatments of potential breeding sites would continue beyond this date. For this reason, VCU has found it necessary to train more boatmen to reinforce ground treatments of river basins where entomological and epidemiological results still remain poor. The intention in to select a candidate each for Benin, Ghana and Togo, the remaining there (3) countries of the Zone East still under aerial larviciding control. 8 Two candidates from Benin and Togo were already selected and trained for this purpose by the boatman from Odienn6, Mr Bindd Kissi in the year 2000. The search for a suitable candidate from Ghana continues. - Two technician entomologists from Equatorial Guinea (Malabo) received two weeks(wks 48 and 4912000) training in larval prospections an ground larviciding treatment at Kara in the course ofthe year 2000. IO. MEETINGS - The senior author participated in the Annual Entomological and Insecticide Research Meeting in Ouagadougou from 24 to25 March 2000. -Mr Yaya Coulibaly and the senior author participated in the Annual Zonal VCU Meeting at Bobo- Dioulasso from 23 to 28 October 2000 II. OBSERVATIONS/COMMMENTS Five cytospecies of the S. damnosum complex (S squamosum, S. yahense, .S. damnosum(s.s.) S. sirbanum and the "Beffa" form of S.soubrense were recorded in the year 2000. - As already reported in the 1998 and 1999 reports, more cases of the backcross (hybrid) between S. damnosum(s.s) and S. sirbanum were again recorded in the year 2000. We present here below a table of the cases registered so far in the year 2000. DATE BREEDING SITES RIVER 04.01.2000 04.01.2000 10.01.2000 11.01.2000 13.01.2000 19.01.2000 20.01.2000 20.01.2000 20.01.2000 26.01.2000 01.02.2000 02.02.2000 03.02.2000 15.02.2000 15.02.2000 17.02.2000 17.02.2000 01.03.2000 30.03.2000 20.04.2000 03.05.2000 Atchakpa Agolin-Pahou Naboulgou Toutionga Blitta M'bdt6koukou Angaraddbou B6tdrou Igbomakro Bagan Barrage Datcha Titira Aguigadji Koutoukp6 Nuja Djabignon Sikan Pont-Routier Ekpla Kati ConIl. Katcha Ouimd Oudmd Koumongou Oti Anid Ou6md T6rou Oudm6 Tirou M6 Amoutchou Kdran Oudmd Pru Pru Kara Kara Oti Sio Sio M6 9 08.06.2000 15.06.2000 27.06.2000 27.06.2000 r3.07.2000 18.07.2000 20.07.2000 21.08.2000 24.08.2000 07.09.2000 07.09.2000 13.09.2000 20.09.2000 Kama-Pont Asubende Gba Sinahou Barrage Tchakparou Dassa-gledji Konfokpa Barrage Biro Barrage Parakou Kelan Sarakawa Djougou Banage Logozoho6 Kama Pru Ou6m6 Ou6md Oudora Azokan Kouffo 1 Souansird Sokounon Mabo Kp6lou T6rou Klou - The "Beffa" form of S.soubrense was identified among samples from the following breeding sites -Six cases of backcross (hybrid) between S. damnosum (s.s) and the "Beffa" form of S. soubrense were recorded at the following breeding sites: DATE BREEDING SITES RTVER 05.01.2000 01.02.2000 27.04.2000 17.05.2000 06.06.2000 12.07.2000 1 1.08.2000 23.08.2000 25.08.2000 05.09.2000 Agolin-Patrou Kaboua Gba Barrage Bassa Atchakpa Barrage Tob6 M'bdtikoukou Gounso6 Landa-mono Barrage Nikki Oudmd Okpara Ou6m6 Okpara Oudm6 Zott Ou6md T6wd Mono Sora DATE BREEDING SITES RIVER 15.06.2000 I 1.07.2000 1 1.08.2000 16.08.2000 23.08.2000 20.09.2000 Atchakpa Kaboua M'bdt6koukou Barrage Ayekotonia Barrage Manigri W6ria Ouim6 Okpara Oudmd Issanhoun Odola Toussi l0 12. ACKNOWLEDMENTS We are sincerely grateful to all members of the Vector Control Unit (VCU) and in particular to Dr L. YAMEOGO, (Chief VCU), Dr L. K. B. AKPOBOUA (ATO), Dr H. AGOUA (CAT Zone East), Mr M. SARR (CAT Zone West), Dr Y. BISSAN (CRE), Mr AKE Assi ( Coordinator/Aerops); the Chiefs of Operational Bases and their Assistants as well as the entire staff of the two aerobases at Kara (Togo) and Odiennd (C6te d'Ivoire) including all the Evergreen personnel who had provided materials or assisted in one way or the other to make this work a success. wr oozz m ma @7c{ m CN o ma -o mz{ .Tl -o { oza o {o{ xoz o = -oC ma ! n TD aIz t: ooT c, m C. z m n o mo m =@ n m Nooo F N =zT @ CN o a o @ z ov a zx v z z 6 m7 =g z Dzo z =t-o x @D - C{ z 6 mv o =. =O.o @ z Iz 6 m7 r o ol-{ ma{ o 1-{ or{ o 1-{ oC m =m oC m =m oC m =m ot-{ ol-{D ol-{ ot-{ z 6 mv =oz o =oz o @ a c2 z =z =aa o a ao z ov 3n Dzx 7 z l! m z z o z = t-o =ozo o z 6 mv oC =mm @ oCt- m D1l 'nt-C mz{a t-Do or{D mo{ o x !vC E =m ot-{ ox! v EDo oC nt =m -DC{ oc m =m o =!4z o ! t- x 7D =o x m7 z z 6 mv ma{ @ @ =oz o - C{ =oz o v s mv EI o,j cr NJ @ Noo {5 {5 Io5 ot or or or C'I{ o.to No, Jo ot 5o Ot C'IC' JNo C.l of5g, {o5 6o,J or{5 NclC) IOt5 or@(o oorgr {o{ t- 6) I I { ct o CD 5o) o N No IC'I (o -l Crl or Io 56ro Io(.t(o NNo N(,l J ctN @N (,t N J or 5{ Iocl o =z ol c, @ NI No {5 oOr 5Ctr o o,o (r,{ ol I5 N{ glo (,rc) {o N(rl {C'I{ C'ta,(o NNo NNql ol6 Nol gr(t) 2,7 @ o o{ A o C'I Aol (rt o o C'tC'I@ o o o o g,or o o {N orC'I o ct NC'IOl o{{ aoc (rr o o o o o o c, o gro, o c, o C' o o C) o o o C) J o - m o o o o c, o o o o o o o o g,5 OlC,' o c, o o o C) o J5 a o @ o o c, o o o o o o o o Io N!,o .l o, l.to 5 C) !.r C' I(o or o o I{ o gr C) I ol (,t o @ !,to (D I o 5 o Ol o PI I o o o o o o C' o o o o o ct o o N o N o o o o o o o C' o a o \o @ o 2 o 5 a! m a ma oo =!oa +ul= :z1r,f;Iv:)Ef B920 "88 IF o m{ mv =z { 6zo trl Co Cc N ac c CI tt c €c N '€ €c C.l ca Cc .l € cc zo mo c cCCccC CI GG CIGG € CI CI CIC' C! €!o c Gc €ccC o o CIcc6 CI 5o E,oo m 0 fi E7 m o!'t D oI D oI D oI D oI D oID oI D oI D o D o , D oI D o , D oI D oI D o D oI D ot D oI D oI D oI D oI D o D oI D oI D o D o D oI D o D oI D o D oI D cI! oI D oI D o E D o D o D E U' !22 o o a.l €c c ogl G CI o o(d la CI c c c o o Cl o o € o c ce o o c € I cG o o GG o o oc c c G tC o o la €l o o G c c ,3l c otclGo ot ol € CI o ,glcG o ,N otN CI € o ,NG c o ,N r=c o ,Ncc oIN €G o ,NG c o ,Nce oTNc € o ^N€ € o ,N €tc c ,NCC oTN € € o ,NG c otNo €! o N^ € € c ,NG e c ,Nac E mn =o =o =o tr oC D m tr o C D lll E ocD lll tr oc D m tr oC D m E ocD m E oC D lll tr oc DI ln tr oc D m tr o C D m tr oc D m , oc =oz o oc , oc =o2 6'l oc , o C 3 o z o cc I oC =oz a oC Ez D - ,lIIz I ma Dz Illlv Dz , mIz ,lllt z I ma Dz IlIIz t m a Dz , lnIz InIz t m E2 T nta 2 , ln a D z I ma D2 m^Iz I mv 2 IlllIz ,llt 42 4, s mv m =o 4d =o Dm - m v tr nl D D DIfl . ln , tr tn DlF nl -lI 4olll D mTn 4trlll D lI - ln a trlll DiF nl - nta o m D m -lllv trlll Dl mIlll 4tr m D m - m 4trlI DI nl! nl a Elll I oc! ov 6t oc z Dtr oC 5 oC z Dtr oc 5 oc n D 7 m =o E z D - z o - = a D D D a D =oz = a D n I! s = =aD o ttlt s = a D o tr t? 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G CI c el c CC N c CI IG 6 € € ao € @ G € gl € G6 'c c N N c c c C €e Cla € GG r= Gt, cC N C C c N € 6 GG N G c c N € 'o 6 €t a.l € cG gr c C4 cc o m ; 3 ! I ll3 2o mo € €cccc C: € € €c €l C! € c c €ccc €i € 4co c Gccccc c cc Cicce c € €c €cc G €cc € €c € a €t CI € € Cl c €G €c €tc c GccCCC CI € € €6 €6 ooooU' m 6 ll =tr 6) ll Ev m oI D oI D c! D oI D oI D o D o D o? D c =D oI D oI D o E D oI D o D oI D oI D oI D o D oI D oI D oI D o D oI D o! D oI D o D oI D oI D o =D oI D o D o D oI ; o E D o D oI D cI D o D oI D o! D o , D o D o D @ o 9,z o dcc c d CIc o c!c6 o acc c dcC o a6G o o € € c d CC o dt6 c o atG € o 6GG c oCc o atG € o Nc c c N e c o N66 o NGG c NG c I Ncc ! 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NC c an a) CT4 C 9,7E C C C c C Cl C G C c G ooC c G c C C G C G c CI CI m C a € C C G c a C 6 6 o o @D @Iz o E qt ! m a m U' oo =! oo =ur1Z 4, rt<Y neof >.9 20 L8IF o m{ mv =z {6za a CI a, G c c,l co C'c 6 cd cG NG c4 G c CIE € C.l cc c c €i c Gc Nc 'cq r=c 6l € cl cc Ga G .,! €C N ol ca GC 6N Ggl GG GN G c Nd Ga cc Nc CI cc E'D m 2 U' mo € Gccccc € cccc €i c 6 €6666 c G GGGGG € € tO € € €G € CI c CIcccc CI €Gcccc CI a € CI e C! c G G €G €cc o € C! c CI CI € CI CIec €t €c CI €6666 CI c GGG cG € CI CI c €t CI CIG 6 €6 ai c Gcc CIc C: € 6 € €G €c Cl €G c CIGc CI cccccc c 6 €ccc € c €6 €Gc € c C! €l c c CI c oooo aD m 3tr oll Ev m o D oI D oI D o D o D o D o D o D oID o D oI D o E D o D o D oI D oI D o D oI D o D oI D oI D or D oI D o =D oI D oI D oI D o D oID o , D oI D ot D oI D oI D o D o D oI D o D oI D o =D oI D o D o D @ lh 9,z cN I CN t cN I cN tr(a6 € tr @GG trta GC GG CI C! o G c c o d c e o ctc € o dGG o dGc o o,cc o 6l c c o o)GG c C.C c o a CI CI o atGG o 6t c c I dcc I ar, €c o qt €c o ql €c o dcc 9 ecc o 6i66 o 6: € € o dGG o dcc c ec € o 6l € € o 6l c c o dcc o gt €6 o 6tGG o dcc c ecc o 6l €6 o o)cc o 6! c6 o ('lG c E mn t7c !vC ! ,C !vC =I =m o D = =n or D o tocI oc3tr oc I oCt oc =tr oC o = o o = o = oI o = o = o = o o o = o = o = o o = o = o = o o = o = o o = o = o o o = o = o o = o 4, s mv m Ductr m z tr ln a ln r D t,uctrtltztr nt gl m m D Ductr m2tr m N lllIfi D Ductr n z tr ln llI!l D oc oc ol lI lI P U'c oc GI mIlI P uc 6)c N ,n lll P oc oC IT ,n F o c6 CI 0 m o^C o^c 3 tr oC IdCI oc =troc oc =oz 6t D ol m m D oc o z 6) D cl ln ll P oc =o2 o D I llIll ! oCI oz o D cl m lll D oc =oz o Dgl m E P oC =oz 6) D N ll t? 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G G C G u N G c C G G C c (, G C N N e c CI c c c u c c C G o =z € € C € € C C c l. c alc G 4 € 6 I 6 NN q4 G Ne N GI€ GIc N N gl t N an NE d 6 gIc N a 9,v @ c € c € G C € € G C € € C C CI c c c € € c c c c € C Cl €t ooCD c G C € G a c c C c c c € 6 G G C C CI € G C c c € C Cl CI D - m € G C c c C G G C G G G C G G G C a G G G C a G c C € G o otr = DE ts LE)= >= ts @Iz o q, ! m a m cn o o =! ot2{96:zlJrtf;I v:JtE] B920 'o B6 TF o m{ mv =z =oza o c C aC c CIb4 C Ci Ctc €(! €c c4 €(! c CI CI c C(a Ci c 6t 6 € gl G GG Nt c CC rO CIc € G c c € ci € € € G CC u € CIG Na c €G G bd CC al CI G c Na ctc G4 € Cc Ngl 6o oc Ng, €4b €t N ol Ge € € N el 'c 4 C c c,l CIN G € a) €c Nc C CC Nc 'c CC N c C CC a' G cl c CI ol G(, cC 6 cd CC 6t G CI N CI 'c GG g cd CiC o Ce cc E'D{lll m =m =omo € cCcCCC c € € cG CI c CI G c c €c Ci c cCcc €C € €G CI Cl CI € G Cl CI € € CI c G G c cc Ci c 6 c6 € € €C' € € € ClGG c c G €c CIcG CI G eCCC r= c c €G € €c € gt @ tttG c) Gccc Ci tC c Ctc € €6 € € €G CI CIc € € €cGc €c € GGG G €c Cl G c cc €c c CCC ccc c €ccocc C €C CiCCC CI CI CI eccc c cGG €cc G GG c c eC c 6G cGc € G G C: c Ci cC 3 2) 't, ll m = e m Ev m c!I o D o D oID o D o D c E D o D o D o! 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C CI N N c C c C ooC C G G c G c c € G € G c C c D - m C CI 6 G G a c 6 CI € CI G G G U' o @ EEEEEEEEEEEEEEE llllillllilltE lrililrilllllH W rfi Hffi m ffi gP ffi [*,i ilil c e C C c N cgl CC CNt CT Cc (o c6l GG Ngt G cl 6c u ca C c € Nt CT Cc € N c GI G c cN cgI cc CN cq bc 6N 6 elb Cl Ngl € clbc c Ce cc gl ct a cc cN ec 6 6N c c tr 6N cG cc GI cc C: cl €a cc oD m zo mo c G c ooooo!I E7 m I Dc =6) m v o EI :I! a n 7 6 E f! s 2 a n 7 G, Et - DC =o m v oc I 6: E = =Dc 26ll 7 oE I a,! =2 G, n 7 G,c I -Dc =6 m v oc ! =D =z all 7 6 E3 :! =2 all a 6 E I DC z 6lI v o E =Dc =olll v o E =Dc =Gll 7 6 E @ oo2 3 'ti € € 3 Ncc =N Cl CI =gl €c =ngl cc =gl cc :2occ -z orcc =Ncc f2 Ncc : aN c c trt Ncc =aC c c =ze CI CI =zacc =aocc : zq c € E m .n 3I .t c c 3 Da cc =! c c z 6 m v 6)C z mfi z 6lll u o s zlll m =6ll 7 G s zttl! =! c C =D oC , otr D 2 6 tn u , c E! z G.ll 7 =6rln v 6)c zI! m =6n 7 c C 2It l? t s mv m =D oc =6n , 6 s zlltt e n , G, C z n n =D1l oc eIt 7 6: s z tll tll z -ci ll 7 6 E allll tr D m -z ! 5 -1! 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C'T !ao(,r !oo C'I !o cr C'I a.l a, C, C, Ftt c! o irrql ro o ro o ro ql @ o @ o oL ot oL C'I o 5 C'I o 5 C'I c! l\t C't Pht C'I Pl\lgr PNgr PN(,t \ { :.1 r I _{o Io(eo 5 @o F(o 6 Io(o 6 o C.l cl o C.l ct o C.t cl o(.t cl N(,l Nor it NI !.rN i.lN l\lN !.,N IC' NN Io NN IPNN IPNN !.. N P N !.t N l.l N !.r N N .r C'I N 5gr N 5gr NL C'T NL C'I t- oz o = = = = = m m m m m m m m m m m = E = = m m m m m m m m m m m = o o o o o cr cr (o 5 o o (o o o o o C., 5 q! o cl o C' o o or o o { o{ o o o o o o o o o o o o qr N o o 5 o o (n o o o o l\t q! o o =z o o o o o o o o o o o .o o o 5 N o c! o o o o o o o o o o o n(E o o o o o o o o o o o o o o o o o o o o o o CI o o o o o o o oC o o o o o cl ot @ o o o o o o o o o o o o o o o o o o o o o D - m o o o o o o o ct 5 6 ct cl cl Ct o c o o o o o o o o o o o o o o o @ o o o o oo o CI Ct cl o o o o N NIo gr or a, o CA qr o C.f(.' o o o o o o o o o C.l 6lo CI c, o U' :s tT rno ! aCl-L =tt o rTq oo =Efi NC/)OhfrFmrnqs oorn rn FOgu !g =c1- m U, rll =o rnoIq rn a I =Dail3 ul ol oo dr oo o) CIC' (ri oo o o oo o o oo o o oo o o oo qr o oo o oo { CI CIo (., o oo ql o oo (.t o oo C'I o oC' oF Poo ol Poo o5 o oo oF o oo o { m or{ ot.{ oF{ o{ or{ D or{ or{ or{ or{D ot-{D acxD Ex = DoCx =I = DoCx =I = =oz o =oz o =oz o =oz o =oz o @ oIz t- oN t- oN r ol\t t- oN oINoo oxNoo oxNoo oxNoo oxI\'oo o{(Doo o{ oroo t- qr N t- C.lN t- (.t N 3z oo =z oo =z €l €l =z oo =z oo @ m 'n !vc !vc !vC !vC x mv z x mv Dz x mv Dz x mv 2 x mv z o = o = Docx =x E ocx =xD = ocxD =x = =oz o =oz o =oz o =oz o =oz o v mv m oc @ mzo m C'I oc @ mzo m 5 oC @ mzo m Ot oc @ mzo m = =v C'I = _{ v 5 =Iv qt j Jv N j _{v { oC =oz o C'I { oC =oz o q, Iz =vo or Jz 5v o 5 =zIv ogl x! o @ o I! o @ 5 x! o @ gr x! o @ l\l x! o @ D tr mc f.l @ N f.l rON :.t(o N :.t(o N !o(oo !o rOo .o @o @ @q, @ ioo o C., C'I o(.,(,r Poo o oo cr oq! or q!{ (r) cl or ql{ Ic!{ Po{ I I I { I N N N N N I C'I I or P or o C'I o\ ol C'Iql C'Io C'Io C'Ic C'Io oz Q = E = = = m m m m m m m m m m m m m m m m = o ht o o 4., 6 (O C'I o o o (o o o {o{ o CI o o o or o o o o o o (o o o c, =z 5 o o al o aO C'I o o o o o o o o Iv @ o o o o o c, o o o o o CI o o o o o o o oc o o o o o o o o o o o o o o o o o o o -m o o o o o o C' o o o o o o o o o o o o o o TD C,l N oo cl t\, N o o 5 o o o CI o CI ct 6 o o o o Pa s or{ Bll{l ol 38 COMPOSITION DES ESPECES BASSINS / RIVIERES CYTOTYPES KABA NIGER (GUINEE) TINKISSO NIANDAN MILO SANKARANI NIGER (MALI) BANI SASSANDRA PRU DAKA WHITE VOLTA OTI PRTNCTPAL KERAN KARA MO NTGER EST (BENIN) LAC VOLTA HT. MONO BAS MONO HT. OUEME BAS OUEME OKPARA S. squamosum S.sirbanum S.sirbanum, S.damnosum (s.s.) S.sirbanum, S.damnosum (s.s), S.squamosunr S.sirbanum, S.damnosum (s.s), S.squamosum S.sirbanum, S.damnosum (s.s), S.squamosum S.sirbanum, S.sirbanum, S.damnusum (s.s.) S.damnosum (s.s.), S.squamosum, S.sirbanum, S.sirbanum, S.damnosum (s.s.) S.sirbanum, S.damnosum (s.s.) S.sirbanum, S.damnosum (s.s.) S.sirbanum, S.damnosum (s.s.), S.sirbanum, S.damnosum (s.s.), S.damnosum (s.s.), S.sirbanum, S.squamosunr S.sirbanum, S.damnosum (s.s.), S.squanrosun'r S.squamosum, S.yahense, S.damnosum (s.s) S.squamosum, S.damnosum (s.s.), S.sirbanurn, S.soubrense S.damnosum (s.s), S.squamosum, S.sirbanum, S.damnosum (s.s.), S.sirbanum, S.damnosum (s.s.), S.sirbanum, S.soubrense, S.squamosum S.damnosum (s.s.), S.sirbanum, S.soubrense S.damnosum (s.s.), S.sirbanum Itr m ={I .It I o D -lI o =o o -{ o { D x o =o = I o mo D =z C mFF mo Nooo !lr Fa o o! l) \ \ -f- z t_1 '\i 'i)i \i il l,\r \ .Tl .Tl !- o ol-{ m U,{ .r.\ bc ox! v \-Ir =oz a o =m =oz o "1r-/ J( bol- *$3IECT XE Ed TTITE a<aao EHBA= tJt\o \ \ )-1 ),.- r I l n I rn