Cytotaxonomy of S.damnosum complex: January-December 1994

aWORLD HEALTH ORGANIZATION ArRlc.4I{ X,EGlOt{ SUBJECT PERSONNEL PERIOD COVERED : January - SIGNATURE OF RESPONSTBLE OFFICER : DATE OF SUBMISSTON : ORGANISATION MONDIALE DE LA SANTE IIEOX)N OE UATRIQW Cytotaxonomy of S. damnosum complex George K. Fiasorgbor (TechnicalOfficer) and S6dou P. Naniogue (Assistant) ous / oNcHo, B.P. 36 KARA, REPUBLTQT E DU rrOGOTEL.60.60.75 ,/ Dec 28J ua 1,994 1995 CONTENTS 1. INTRODUCTION 2 . CYTOLOGICAL TDENTTFICATIONS 3.I,IATERIALS AND METHODS 4.RESULTS 5.SEASONAL DISTRIBUTION AND MOVEMENT OF THE CYTOSPECTES 6.REINVASION AND TNSECTICIDE RESISTANCE 7.THE FOREST CYTOSPECIES S.POSTTTVE GITES WITHOUT CARNOYS FOR CYTOLOGICAL ANALYSIS 9. TRAINING lO.MEETINGS r- r_ . oBSERVATIONS/COMI,IENTS ]- 2 . ACKNOWLEDGEMENTS haP s.6 , L'r't ) j\\ + arI 1. II{TRODUCTTON This report is'a summmary of all the main activities carried out by the Cytotaxonomy Laboratory at Kara (Togo) in L994. Results of some samples collected in L993 but which were not immediatelyidentified were also embodied in this report. It is important to note here that aIl identification results contained in this report have already been communicated to those who contributed materialsfor cytological analysis and to those involved in operational researth, larviciding and surveiltance activities- fn the course of the year,the two technicians of the Kara Cytotaxonomy Laboratory were invited to participate in a Workshop in Bamako. We joined our two counterparts of the Bamako Cytotaxonomy Laboiatory to do the course under the direction of Oi.Oaniel A: Boakye (WHO Consultant). Discussions focussed nainly on the following : Cytotaxonomic Results for the period L993/94. - Analysis of Cytotaxonomic Data for Publications. - Criteria for diagnosing different members of the S.damnosum complex. - Fixation Methods and Molecular Techniques. - Cytotaxonomy and Insecticide Research Studies. - Problems encountered during the past year (l-993). We also participated fulty in the Larval Rearing Work began by Dr. D.A.Boakye a few days prior to our arrival in Bamako. Mr.Sedou P.Naniogue, the Assistant in the Clrtotaxonomy Laratory Kara received training in dissection and morphology of the s.damnoiun comptex whitst the Senior Author was initiated in the identification- of S.damnosum (s.1.) by morphometry using the criteria estabtished by Dr. U. Wilson. 2. CYTOLOGICAL TDENTTFICATIONS Routine identifications were carried out on the larvae of S. damnosum (s.1.) from all the river basins making up the Eastern Operation Area of the OCP.These include the Bandama, Como6, Volta, Asukawkaw,Mono, Sio,Ou6m6 and the Niger. In Lgg1 a total of Six Thousand Three Hundred and Forty-Six(6346) Iarvae of S.damnosum(s.I.) were processed for cytological identification. Out of this number Five Thousand Seven Hundred and Seventy-One(5?7L) representing 90.92 of the total sample received were successfully identified.The remaining 9.Lt of the larvae not identified produced poor chromosomes and therefore unreadable. ih*l*r#:tr ."' ;' 'i;' e3. I,IATERTALS/METHODS I-IATERIALSS Specimens on which this report is based came from three sources: (i) Specimens from Routine Checks by the Sectors and Sub- Sectors. (ii)Specimens from Susceptibility Tests.(iii)Specimens from Helicopter Prospections. I,IETHODS Larvae of the S.damnosum species complex were collected from aII the main Rivers and their tributaries in the Eastern Operation Area. The larvae hrere preserved in the field in carnoy's fixative(3parts abolute alcohol 1 part glacial acetic acid).The samples were stained and chromosome preparations made as described by Dunbar(L972) and Boakye (1988).fdentifications were made using the chromosome maps of Vajime and Dunbar(L975), Meredith et al. (L983), Post( L986 ) and Surtees et aI. ( l-988 ) . 4. RESULTS CYTOLOGICAL IDENTIFICATTONS (].994) BASIN SA},IPLE SIZE TOTAT IDENTIFIED z BANDAII{A 346 265 76.6 COMOE L257 l_050 83.5 VOLTA 2200 2084 94.7 AST'KAWKAW 273 255 93 .4 MONO 5l_8 509 98.3 sro 470 418 88.9 OUEME 855 808 94.5 NIGER 427 382 89.5 TOTAL 6346 577L 90.9 7, 5. SEASONAL PTSTRTBUTTON AND MOVEI,IENT OF THE DTFFERENT CYTOSPECTES In the dry season vast areas between the river systems faIIdry in the savanna areas. The perennial rivers in the southern areas however remain as refuges for S.damnosum (s.1.). With the beginning of the rainy season numerous new breeding sites hrere born and formerly dry areas vrere newly invaded by various pecies of the S.damnosum complex. As in the previous years, the savanna cytospecies (S.dannosum s.s. and S.sirbanum) were more widelydistributed in 1994 than the forest species. S.damnosun (s.s. ) (MAP 3) S.damnosum (s.s. ) is found mainty in the transition zonebetween the forest and the savanna. In the dry season the distribution area borders South and East close to the mountainforest in the Lower Dayi River, the Sio and Amou. The species bypasses the forest and spreads alongside the Mono and the ou6m6. T[ra breeding sites on the tributaries of the otilPendj.fi. river belong to the Sudan savanna.This species was also idntified on river Sota. In fvory Coast, S.damnosum (s.s. ) was recorded on the Lower Bandama, the Kan, the Nzi, and the Como6 rivers. In the rainy season S.damnosum(s.s. ) extends its distribution range upstream. During this period the species can be found in many areas in the Eastern OCP. S.sirbanum (MAP 2) Two forms of S.sirbanum exist in the Eastern Zone of the OCP.The two are geographically separated.Members of the northerndistribution arei ale cytotaxonomically homogeneous in a large measure. They miss the inversion l-S-3. The southern form is characterized by the sex-linked inversion l-S-3 which occurs eterozygous in males.The southern members show more chromosomal polymorphisms. Beside the southern breeding sites, S.sirbanum with 1S-: appeared in the northern part but only in a small area in the otilPendjari River. During the dry season the distribution of S.sirbanum without 1S-3 hras limited to the rivers Pendjari, K6ran, Kara, Koumongou, Sansargou and the Bouale in the northern part of the Zone.S.sirbanum with l-S-3 appeared only in the - southern rivers and in low numbers on the Dayi, Chai, Sio, Mono, ou6md and the Sota.In Ivory Coastrthe southern form of S.sirbanum was recorded on the Lower Bandama, tne Uzi, and the Como6 both in the dry and rainy seasons. The distribution range of S.sirbanum with l-S-3 is very muchlimited in the rainy season. On the contrary, the distribution ofS. sirbanum without 1S-3 extends largely upstream during the dry season by the harmattan winds. ',qaHflti{L.g8 4S.squamosum (MAP 4) In the dry season larvae of S.squmosum were found in the rivers of the mbuntain forests extending alongside the Volta Lake in Ghana and Togo. In the rainy seasonrthe species extends its distribution range to intermittent rivers northeastwards to the tributaries of R-.Mono (Na, Aou, Kolowar6) and the Zou river in Benin. The entire population of S.squamosum recorded west of Lake Volta in Ghana on niver Oyoko (a small tributary of the Black Volta) was collected in the dry season (L6/2/94). S.yahense (MAP 5) S.yahense breeds mainly in small rivers in the forest zone. It is rareiy found in the savanha. In l-994 S.yahense sras registered at six co11-ecting sites, all confined to the immediate periphery east of the Volta Lake in Ghana and Togo.These are Djodji(R. Gban- Hou) ,Ayagba(R.Kpelekoum) , Tomegbe(R.Domi) , Menuso(R.Menu) ,Xudiraln.bayi), -and Katiin.Sio) .There was no signif icant extension of the distribution in the rainy season. S.sanctipauli (s.s. ) (MAP 6) S.sanctipauli(s.s. ) was recorded on the Lower Bandama, Nzirand Como6 both in the dry and rainy seasons in the course of the year L994. S.soubrense(Beffa) (MAP7) The only case of S.soubrense(Beffa) registered in L994 was collected in the rainy season (28/6/94) from Agolin-Pahou (R.Ou6m6)in Benin. dc ,6. RETNVSASION AND INSECTICTDE RESISTANCE REINVASION Reinvasion may be defined as a situation whereby blackflies of the S.damnosum complex are registered in good numbers on certain river basins under larviciding control and which are not of local origin. In the oCP,two methods are used to study the Reinvasionphenomena. These are by Morphometry and Larval Cytotaxonomy.Morphometric Analysis is carried on the adult flies and involves the observation of the colour of the hairs on certainparts of the fly (wing-tufts, antennarcoxararculusrscutulun and the 9th tergite hairs). In addition to the observation of these charactersrcertain parts of the fly are also measured (the antenna, the thorax, the fore-Ieg and the hind-leg.The results of the observations and measurements are then introduced into the computer in order to obtain the identity of the reinvading flies.The study of reinvasion by morphometry is at times beset with difficulties as this depends largely on the accuracy and efficiency of the technicians involved in the observasion and measurement of the many characters mentioned above. Larval Cytotaxonomy,which stilI remains the most reliable means of identifying S.darnnosum (s.I. ),the vector of human onchocerciasis in West Africa, rdY be used in the study of reinvasion. By this method some of the reinvading flies caught can be induced to Iay eggs which could be reared up to aged larvae suitable enough for cytological analysis. The inversions recorded on the banding pattern of the giant salivary gland polytene chromosomes are then utilized to determine cytotypes of the various members of the S.damnosum (s.1.) implicated in the reinvasion.fn this $ray the movement of the reinvading flies could be easily monitored. In the Eastern oCP the results of larva1 cytotaxonomy have made it possible to distinguish two forms of reinvasion in the oti/Pendjari, Kara, K6ran and Koumongou river basins annually.Normal Reinvasion occurs during the rainy season (May- September) when almost aII the male populations of S.sirbanum trear the usual 1S-3 heterozygote inversion. Inverse Reinvasion replaces the Normal Reinvasion soon after the rainy season (October-ApriI) and it is presumed to be largely influenced by the North-Easterly Winds (The Harmattan).At this time of the year almost all the males of S. sirbanum populations lack the characteristic 1S-3 inversion.These populations are the original inhabitants of the tributaries of R Niger. ,rr ll:' 6INSECTICIDE RESISTANCE Insecticide Re'sistance in the OCP means non-susceptibility of S.damnosum (s.1.) Iarvae exposed to known and accepted diagnostic doses.The Programme has seven (7) different chemical products for its larviciding operations. These are Abate (Temephos), Chlorphoxim/Phoxim, Bacillus Thuringiensis (Bt), Carbosulfan, Pyrachlofos, Permithrine, and Vectron. Before any of these products are finally setected for operational use, several susceptibility tests lrere performed to determine their efficacityrcarry, efficiency, residual effect etc.. However,the frequent use of the products permit some of the target cytospecies to develop resistance. These may be due to underdosirgrage of the chemicalruse of same insecticide for several cycles without rotationrinterference with the use of agricultural products of the same class in the area etc.. The ability to detect resistance to larvicides in the OCP has been greatly improved, thresholds of resistance of populations ofS. damnosum complex have been reassessed, and a computerized cytotaxonomic data can be correlated with known resistancepitterns, thus facilitating redefinition of the range of susceptibility of the various members of the S.damnosum complex. In 1-980 and 198L when S.soubrense and S.sanctipauli were detected to have developed resistance to Temephos (Abate) and Chlophoxim respectively on the Lower Bandama in the Ivory Coast, Dr. R.J. Post discovered a new inversion recorded on Chromosome 2(IIL-A) on the banding pattern of the giant salivary gland chromosomes to separate S.sanctipauli as the cytospecies implicated in the resistance study and not S.soubrense (s.n. ) - Cytotaxonomic studies also reveals that of the two savanna cytotypes (S.damnosum s.s. and S.sirbanum) S. damnosum s.s. is more susceptible to insecticide resistance than S.sirbanum. 7. THE FOREST CYTOSPECIES (MAP 8) In the Eastern OCPrs5-x cytospecies are known to be the vectors of human onchoceriasis. These are S.Squmosufi, S.yahense, S.damnosum s.S. r S.sirbanum, S.sanctipauli,and S.soubrense (Beffa). Of these six S.damnosum s.s. and S.sirbanum are known as the savanna cytospecies and are the cause of the severe form of human onchocerciasis(River Blindness) in the entire Programme Area. The remaining four (4) cytospecies (S.squamosum, S-yahense,S.sanctipauli and S.soubrense (Beffa) are known as the forest cytospecies though some of them make incursions into the savanna aieas-during the rainy season. In L994 the forest cytospecies are recorded mainly from the mountainous forest region east of theVolta Lake in Ghana and Togo with S.squamosum as the dominantforest cytotype in the Eastern Zone. S.squamosum was registered in Benin on R.Zou a tributary of the Ou6rn6 in the rainy season. TS.soubrense (Beffa) was recorded on the Ou6m6 River at Agolin-Pahou also in the wet season. fn the West S.sanctipauli (s.s. ) was registered on the Lower Bandama, Nzi, and the Como6 in lvory Coast whilst S.squamosum sras recorded at Yabraso on R. Oyoko a srnall tributary of the Black Vo1ta in Ghana. It is very irnportant to note here that S.squamosum population of R.Oyoko, West of the Volta Lake differs from S.squamosum populations East of the Lake. This observation once again confirms earlier identifications of S.squamosum populations from South West Ghana on rivers Tano, Pra, Afram, ChirimfarSumampa and Oda. 88. POSITIVE GITES WITHOUT CARNOYS FOR CYTOLOGICAL ANALYSIS (MAP e) ft is observed with much regret that most of the larval samples of S.damnosum (s.1.) collected in the Sub-Sectors were not sent to the Cytotaxonomy Laboratory in Kara for identification. From January to October L994, that is just before the East Zonal Meeting (24 -28 /LO / 94), larvae from 57 points (see Map 9) were still not yet received. Some of these samples vrere collected as early as in January L994. This unfortunate situation was seriously discussed at the Zonal Meeting especially during the the Workshop on Friday 28 /LO/94. Together with the Sub/Sector Chiefs thefollowing arrangements for the expedition of larval samples to the Cytotaxonomy Laboratory in Kara was concluded: BOUAKE SECTOR Subsectors of Bondoukou, Bouak6, Bobo-Dioulassou. The Sub/Sector Chief of Bouak6 assumes the responsibility of collecting aII samples and despatching them to Kara. TA},IALE SECTOR The Sub/Sector Chief of Hohoe is to send his samples to Atakpam6 for onward transmission to Kara by the Sub/Sector Chief of Atakpam6. The Sub/Sector Chief of Kintanpo is to send his sanples to Tamald where they are to be added to those from Tamal6 and be forwarded to Kara. The Sub/Sector Chief of Tamal6 is to assume this responsibility. PARAKOU SECTOR The Sub/Sectors of Kandi and Bohicon are to send their collections to Parakou. The Sub/Sector Chief of Parakou then adds the samples from Parakou to those from Kandi and Bohicon and forwards everything to Kara through the courrier of saturday. KARA SECTOR The Subsector of Kara has direct contact with the Cytotaxonomy Team. Larval samples for cytological analysis are therefore to be handed over to the Cyto Team immediately they return from theprospection trips. N. B. Apart from the arrangements made above, aII theSub/Sector Chiefs can benefit from the presence of any pilotrthe hydrology team, the insecticide team or the big trucks of theAerial Operation Team for the distribution of insecticides and aviation fuel,to send their larval samples directly to the Cyto Team in Kara. .. 't llt**1* t TRAIMXG PROGRAMME DE FORMATION POUR I.ES TECHNICIENS EN ZONE EST 1994 AGBOSSOU AMADOU BOUKARI KARA Morphologie PARAKOU 9 b u 37 tr 21-27181s4 & 1 1 - 1Blele4 2818/94 o3l9le4 4-1019194 36 tr 35 GBAGUIDI GUEGUEDEGBO Morphologie & Dissection Morphologie DATES SEMAINES S/SECTEUR PERSONNEL SWET UEU DE FORMATION AWADI BIRREGAHs/s KARA S/S KARA GOUNI AREM/KARA BILEOU FIASORGBOR AREM/KARA AREM/I(ARA GOMINA GONE S/S ATAKPAME S/S KARA KOUMOUVI NANIOGUE S/S ATAKPAME AREM/KARA S/S KANDI S/S PARAKOU S/S PARAKOU S/S BOHICON S/S PARAKOU Morphologie & Dissection Morphologie Morphologie & Morphom6trie Morphologie Morphologie & Dissection Morphologie Morphologie & Dissection S/S KINTAMPO S/S HOHOE S/S TAMALE S/S KINTAMPO S/S TAMALE S/S HOHOE ADAMS ADZAH AGUNGA GYAMERA OSEI TETTEH Morphologie Morphologie Morphologie Morphologie & Dissection Morphologie Morphologie & Dissection KINTAMPO AREM / KARA FIASORGBOR NANIOGUE35-37 Atelier CYTO BAMAKO1 - 1ol9l9q n I I I ro 10. MEETINGS The Senior Author of this report participated in the following meetings in the course of the year L9942 (i) Meeting with Chief VcU in connection with the PIan of Action and Budget for l-995 Kara ,6-7 April L994. (ii) Bi-Annual Sector Meeting, Kara L4-L5 Avril L994. (iii) Research Meeting, Bouakd 2_6 ylay L994. (iv) Workshop in Bamako 1-l-o September L994. (v) East Zonal VCU Meeting, Kara 25-28 October L994. 1l-. oBSERVATIONS/COMMENTS -AIt the six main cytospecies of the S.damnosum complex (S. squamosum, S.yahense, S.damnosum (s.s. ), S.sirbanum,S.sanctipaul(s.s.)rand S.soubrense have been registered in the Eastern Operation Area. The NiIe form of S.damnosum (s.s. ) samples from Bassa (R. okpara), Bagan and(R.Sora). was identified Mo (R.Mo),and amongNikki - The Northern or Monomorphic form of S.sirbanum was recordedfrom Porga R.Pendjari, Borgou (R.Sansargou) rNaboulgou(R.Koumongou),Titira and Tapound6 (R.K6ran), Sara-Kawa and Landa- Pozanda (R.Kara) and A16h6rid6 (R.Boua16). One case of backcross between S.damnosum (s.s. ) andS.sanctipauli(s.s. ) was identified from Koutoukro (R.Como6) on7.I.94. Chromosome I and IfI have inversions typical of S.damnosum(s.s.). Chromosome II vras fixed for the inversion TIL-4.6 A /4 6A and was therefore considered as S.sanctipauli (s.s. ). Backcross on the basis of chromosome II. One case of backcross between S.damnosum (s.s. ) andS.soubrense (Beffa) was recorded from Barrage Tob6 on R. Zou in Benin on 28.9.94. Chromosome I and fff have inversions typicat of S.damnosum (s.s. ) whilst chromosome II wears inversions ofS.soubrense (Beffa). Backcross on the basis of chromosome ff. During the preparation of chromosomes frorn Koutoukro (R. Como6 in Ivory Coast) and Bagan (R. Mo in Togo) some larvae wearing female gonads turned out to be males in the final identification. This newdiscovery is to be seriously followed in L995. .t 1,rrlffif1.;.6g1 II L2. ACKNOWLEDGEMENTS we are sincerely grateful to Dr. o.Qui116v6r6 (Ex-chief VCU),Dr. J.Hougard (Chief VCU), Dr.H. Agoua (CAT-Zone East), Dr. B.K. Akpoboua (CAT-Zone West),Dr. C.Back (Chief AREM/IRU),Dr.M. Kassambara (Sector Chief/Kara) rDr. A.K.DiaIIo (Sec torChiefr/Parakou),Mr. S.A. Sowah (Sector ChiefrzTamald),Mr.M.Sarr(Sector Chief/Bouak6),The Assistant Entomologists, Laboratory Research Assistantsrand the entire staff of the two Aerobases at Kara and Odi6nn6 including all the Evergreen personnels,who have contributed materials or assisted in one hray or the other to make this work a success. 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D LtJ J ITE! IJJiF B I! T? il e?IJJ ul tlJ *,w aaHi=====f=HE=sfr IJJE lrJ E -qr CE og tJ 'oo lr IJJ E c, c, c, o (J I cr 9 ct q, tr, tr ct EfIrrEooooooooo0000000 .D!l.1ltlFFFFF(\rrrFFOlFaocnOAAOOOOOOOOOOzzzzzzzzzzz22222 uJ !E{ IJJ -Ffioootr== t\G)O- 88Orso) o,(oFO'ooqci o fi ID = IIJ IJJ ==uJ tri LrJF o #tEsgs$g$iii$ IJJ IIJ IJJ IJJ IIJ IJJ IJJ IJJ UI T! U,I TIJ IJJ IU IIJ z5 ZONE EST CYTOTAXONOMY t.tilP r E I l I I a \L/ I,a Ja- -)- O tO aOOl. -- - ,} - - \ I \\ ( , \ I \ I I \ t \ est I! Il t I a 7 { \ Dlslrlbutloh of 'thc Cytotypcs of thc LEGENDE O. S. slrDonrm O S. rlonrranm (rsl O S. rquomorum a S yohonrl i E S. robnnsc (bcffol . A S. ronc?lpouli (r.s.t -,rJ- Llmltc drs phosct op&otltrncttcs S. domnosum complcx EASTERN :rlJjloN AREA ( EoA' ll, ,$ r^f i,,. 24 CYTOTAXONOI{Y LIST OT @I,LECTING SITES - 1994 l- B6riaboukro -R'Bas Bandama 2 Gat6goua -R'N' Zi 3 Kravassou -R'N'Zi 4 Ass6r6kro -R'Kan 5 MrBahiakro -R.N'Zi 6 F6t6kro -R'N' zi 7 Amouakoukro -R'como6 8 Arniankouassikro -R'Como6 9 Koutoukro -R'como6 1-O Yabraso -R'OYoko Ll- Porga -R'Pendjari L2 Panseni -R'oti 13 Borgou -R'Sansargou 14 Kpalgou -R'KPalgou l-5 Toutionga -R'Oti 16 Naboulgou -R'Koumongou 17 Titira -R'K6ran 18 Pouda -R'Binah 19 Landa-Pozanda -R'Kara 20 Atchangbad6 -R'KPaYa 2L Gu6rin-Kouka -R'DankPen 22 Kama -R'Kama 23 Bagan -R'I'{6 24 ALeberid6 -R'Boual6 25 Kass6na -R'Na 26 Landa-Mono -R'Mono 27 Aou-Losso -R'Aou 28 KPassa -R'Bassa 29 Kechebi -R'Chai 3O Tinkiro -R'Asukawkaw 3L Menuso -R.Nenu 32 Djodji -R'Gban-Hou 33 AYagba -R'Gban-Hou 34 DaYi-HoteI -R'DaYi 35 Kudzra -R'DaYi 36 Amou-Oblo -R'Amou 37 Kati -R'Sio 38 Wonougba -R'Sio 39 Konigbo -R'Ani6 40 Ad6dakoP6 -R'Amou 41 KPoba -R'Mono 42 Tobd -R' Zou 43 Angarad6bou -R'T6rou 44 Gba -R'Ou6m6 45 AgaIin-Pahou -R'Ou6n6 46 Y]'B6t6koukou -R'ou6m6 47 Bassa -R'OkPara 48 Nikki -R'sora(oli) 49 Gbass6 -R'Sota 50 zondji -R'Sota 2' DT$TRTBIITION E[ I{OUTEI{EII'IS DES DTFFBREITTES ESPECE.S c1-,:ro1r,N(oNoEIQt ES DAIIS L,A ZONE EST (1994) - Bas-Bandama et ses affruents : s.damnosum (s.s. ), s.sirbanum, etS.sanctipauli (s.s. ). - comori 3 s.damnosum (s.s. ), s.sirbanum et s.sanctipauri (s.s). - Volta Noire (R.Oyoko) : S.squamosum. - oti/Pendjari et ses afftuents : s.damnosum (s.s), s.sirbanum et S. squamosum. - Asukawkaw et ses affluents : S.squamosum et S.yahense - Dayi 3 s.sguamosum, s.darnnosum (s.s.), s.sirbanum, et s.yahense. - sio : s.damnosum (s.s. ), S.sirbanum, s.squamosum et s.yahense. - Mono et ses affluents: S.damnosum(s.s), s.sirbanum et s.squamosum - ouen6 et ses affruents: s.damnosum (s.s), S.sirbanum, s.squamosum et S.soubrense (Beffa). - Les affluents du Niger : S.damnosum (s.s) et S.sirbanum. En ce qui concerne Ia distribution et les mouvements desdiff6rentns especes cytotaxonomiques, il n,y a pas de grand change- ment en L994 par rapport a 1993. '#1 r I( I '-a ) (' ) / '._-- ( *9' ) \-/- -"'a- -' I ,, ')\ a.' \_-. _' t (-'= .t .. \\ \.1 \ q -<' I l\ o-I Qo E a c,6 .o rn Oj- uzoo trj o-Oe ffr lrJoEl o IL tuttr: 1 N 3{ s o-U o -J tUo Fa tU LUz oN qrI TIJ \oN .rt o) o) IF z UJ E =c({ .cIL a-tr U) tUo z o -F tr(tr {L tT ttr I I v.itl-.5. ''-.. '.:,; ( l ( )I / I,I r\). \ ...J ) q o E =at o E6.2+a= 3 -lXYfl; |.oF Fz tr.lo tr,f o 0- t F-N N\ A s TLU o IJ Lu r) F(/) tlJ I.IJz oN z UJ ET o) o) rF- z tU ,i a E =(n oE E(utt a U) I.IJo =o EFE { o- TIJ ttr ( \ *:' I I \ a,.--\. ,--/ ) (, €DN & * ET o, o) 'F z, Lu tro- =U8pEJ(U l,u +E .,tf F nfr HH =oONtrztr I.UE { TL LuE c =o o E6f CTulL aiui t -OY HE <.Y<I(o- , IUoEf o 0- I € ,??- / I ( \ Ir ) (, oc o 6 U'F wzIJO 9 uJ 0- si2o- lrJotrf o 0- ,/(t oN tft A s ET o) o) i- z tIJ O. nruPPLr(U -l .C TUSo .l-fi'fi I.U TU()1IJ zzooE!\l-i-1 Etu { o- TUE I ( ro \o ft E R P{U oi rq e F. V1 F4 14 z o N z rq q+ o\(}t F{ z 14 a v2 fi r- - tJfit& r- +)UAH(tr ?? I U1 FI1 R z o lE{ E-lH Efr4 P{ rqfr i r / I \1- ic- ( ) ( I 'i' r \2 I (--- ---{ \' J I ) =) al o. Ecdo [H E - Ug ! o<d z U c.) o o- ( I -( t t'--<- I -\- \ ,ffittr H N A. d o-U o -J tU E] F Cfi LtJ tU -'4- oI\ =Lu rf o) ol F z tU (d {r{ro -cl o El- ora- o U,tr oL .cI =o q, {fi [Uo z o -F tr TE {L lU TE r ( j (l (\ '.i ) LJ\\ ) o al,c o Ttt oO oy1 TE E . llq ! rv<(o- z tuotrl o o. ( ,Ih ( tvl I ( \ u <? ) --a'/ N N N F - N @ - - nt N N (.) - - = Gl - u) lO Ol Ol - u) - -:=:vv::vvg--9vv-o- riraggggagisrirr garggfasi =g8sx1egB3eI==9i;6::e:??99: a z llJ U,lrlE(L o 1! 0- Eo Fo h f trJ J oo ., o o o ral o F o lc 6 or OoN o c oo@ co ., rrl ffiffiffi+r* € A{ # o- L) o :J LlJ o F tJ) lrl lrJZ o r\I z tU F{ tL TL trJE UJ =cto LL o a o -o =oa tJ) I a a Jd o- E fi a U) I o(n o d >, cn I E a E H g a fJ) lrlo II(J Z. lrJ cn tU cr o_ -k- I fr lsP 9 LABO C}/TO I(ARA HEPAHTITION DEs GITES POSITIFS EN 1994 SANS EAHNOY POUH I-A DETEHMINATIBN DEs DI,TOTYPES PHESENTS RENTRANT DANS LE CADRE DES ACTIVITES OPERATIONNELLES .l .\1),: /'1 I /:) I /\ ii /1 t I i(, 2:, i.l t\ l. I I \ -\ \ a I 'l' n II rt /- -) GITES I,ARVAIRES OAGOUT+PA}|OU (r) oALEHERTDE (r) oAMOTIoBLO l2l OASSEREKnO (4) o ASUBENDE 121 arAst (Aw(Aw-BR (r) o AYAGBA (2) G)AYAGBAKPELE(UM (2) €)BARRAGE-U'OUGOU (r) Gl BARRAGEGOTTTEBE (r) c,BARRAGE-NI(K] (r) ct BARRAGE-PAMKOU (4) o BARRAGE-PB. T(ARA (r) o BAnRAGE-TAMAROU (r) o BASSAKPASSA (6) o BASSA-OKPARA (2) o BERTAEOTTKFO (8) o BETEROU (r) c) BTELTKPONG (r) G) BUTA (r) @ CHUTE-GAUTHTER (5) GtDATCHA-BAFRAGE (r) o DAY-HOTEL (r) o DTMBOKRO (21 o a,o},r (2) 6) EBEVA (l) Gt FETEKRO (r) o GBA (r) o GLE| (r) o GNAGAGNA (r) cTHAHAMANSU (Z) G)|(ATI (5) e KECHEBT (3) @ KEREMOU (2) c,KoD,lNA (l) e)KoNTGBO (r) o KouNroHou-PTGKA (2) e KFAVASSOU (3) o KUDZRA (5) a) r-ANDA-POZANOA (r) o MENUSo (6) o Mo (r) o N'DENOU PR.AV.AM (5) o N'zNouA (5) o NAS|A (r) o oussou-PoNT (') etPONr-r(AN (r) e)suBrNso (4) a) suGU (r) o TARE-PONT (t) a)TETETOU (r) OTTNKTRO (r) o TUTT,XGENE (3) a)wAwA (r) owoNouGBA (3) aIYAGABAGOR. (21 a)zoucouNou (r) ,'Jffitm, I a