Cytotaxonomy of S.damnosum complex: January- December 1993

to/ { WORLD HEALTH ORGAI'IIZATION ONCHOCERCISIAS CONTROL PROGRAI.TME EASTERN OPERATTON AREA (EOA) I(ARA, TOGO. suBirlrr Cytotaxonomy of S.damnosum complex PERSOIII{BL ceorge K. FIASORGBOR (Technical Officer) and Messrs : Boukari K. BOUGSERE and Sedou NAI{IOGUE. PERIOD COVERBT} : January December 993. SICilATURE OT NESPOTTSI4A OFFICER : DAIE Of SUEIltSliIOf, z 27 January Leel 1. 2. 3. 4. E 6. 7. 8. 9. t_0. l-l-. @rlEEmS INTRODUCTION. CYTOLOGICAL IDENTTFICATIONS . I.IATERIALS AI.ID METHODS. RESULTS. THE ROLE OF LARVAL IDENTIFICATION IN VECTOR CONTROL OPERATIONS. SEASONAL INFILTRATTONS BY SIMULIUM SQUAI.IOSU{ INTO BENIN. GENERAL DISTRIBUTION OF THE CYTOTYPES. TRATNING. PUBLICATIONS. MEETTNGS. ACKNOWLEDMENTS. t a o o Po,l.- 3"6- 21. nmRoDucrron The present report is a summary of aII activities carried out bv the cvtotaxonomi:c Laboratory at Kara (Togo) in the year 1993' air-ia."diiication results pr"slnted here have already been commu- nicated to those who contrif,utea matrerials for cytoloqical identi-fication and to those invotvJit op"".tional research, Iarviciding and surveiltance activities. In the course of the year the cytotaxonomic Laboralory received the visit of Dr D.A;-noaiye iglHO Consultant)' Discussions focussed nainly on the analysis of Cytotaxonomic data and verifica-tion of r".uiir ; cytotaionirnic ianuscripts in preparation and introduction to the Uitrosoft Exce} and Maplnfo Conputer progrErmmesf;-p;.rit the manipulation of data and making of maps. 2. C{rorocrclr, rDEllfrprcf,rror]ll A total of Five Thousand Two Hundred and Twenty -Three (52231Iarvae hrere processed (stained) for cytological an-alysis in -the course of tne--y-eii. out'of this nurnber Four Thousand Nine Hundred irra Uit" (4909i ""pr"=""ting 94 * gf.the,total sample received were successfully iaentifiea. fh-e renaining 6 ? of the tanrae not iden-tiii"fa proiuced Poor chromosomes and therefore unreadable' 3' rnEBn'IjILs / rrEnroos lITI[tsRIlI.S : Speci.mens on which this report is based came frorn three main sources : Specinens from Routine Checks by.the Sectors'iirnpfes from Helicopter prospections and san[Ies from Susceptibility Tests. lGltfioDs : Larvae of the S.damnosum species conplex s'e!e collected from aII the main Rivers and thefFtfiUutaries in the Eastern Operation ar"" (EOA). The lanrae tere-preserved in the field in carnoy'sfixative (3 parts absolute aliohol : I patt glacial.acetic acid.ffie samples - *Lr" stained and chromosoTg. pr-e.par-a.tions made as aescriueh nV ou-nnar-(tgzz) and Boakye (1988)- rdentifications wereila; using- the chromosome maps ot Vajime and Dunbar (L9751 ,Meredith et ii.(1983), Post (fgee) and Surtees et al'(1988)' (i)(ii)(iii) a tsl. lmla IC l5la to EloI-l tPlolzlo l< lot_Io lrnlzl{lrr o -{ l6lz Lal- l<ol(o lG) (o tulu sI * €o m @(o(,l tu$() t\) o,{ o,o,o,ool No N{ o) o) 5 G) @J I @ !n I * . a \T x €x € CN crt @ N{ N5 45 ITIE ROLE OF IARVAL CYTO|rAXOrIOHIC IDENTIFICATIONIN VECIOR CONIROL OPERATIONS D The identlfication of vectors has proved a serious problemfor medical entomoloqists in recent years: th:-1ifll":tty aris:es from the inability to easily separate vector species on morpholo- sicar srounds;^;;a ;;"y-;aia.;i;i;sicar measures, e-e' vectoriar capacity, infectivity ltc- i-requir6 accurate species identifica-tions. Added to this probleml-fri"v populations of species are diverging to n""o*" aistincC'forn3 irnicn exhibit varing degrees of intergraaation. This ".pfr"=i"es the dynamic nature of specia- tio,r which is a continuo,r= fio""ss, and. ioni"n must be taken into consideration in the aevero[rn""i-"i tecnnitues for vector identi-fication. Increasingly, the-idvances made fn analytical bioche- rnisrry ana moie""rii 6iorogy-;;;-ili"f . apprieg to the problem of vector identiai;;ai;" with-Lwo aims : to -provide a simple method of vectop iaentiiication which is accuratl,.which reflects the current status of the =p""i"= / font and which can be used byworkersinthefield.secondty.,toprovidearesearchtoolto investigate poputations, monilor chinge and thereby rnaintain the accuracy ana u!"i,rfr,"="'"f -I[e-le"nni{ue' To date' ro technique has fulfilled-th;;; i-quirements with each having their ownp"iti""far advantages and disadvantages' cytotaxonomy has been defined as a classification by means of the ceII ciiiiict"rs. s.dimnosum -(9.f -. ) Iarvae for. cytotaxo-I".i"-ia"rrtifi".tion are us-ually collected from all known breeding sites in the ocP. rne inetnod of collection' dissection' sraining .r,a Ipl"iJ=-ia"niiiication are basSd-91-:l:.procedures and criteria elianfished bt ;idely-known cytotaxonomists such as Dunbar (Lg?2),Vajime and Ounbar (igZS) , Quif f6vrirri (L975'), Meredith (1980) Heredith et aI.(igeg)' post (1986)' Boakye (1988) and. Surtees et aI.(1988)' As compared to other methods of identification (Morphometry' Electrophoresis, Adult cytotaxonomi, DNA Probes Cuticular Hvdro-carbon Analysis etc. i-iirvaf cytotaxonony still remains thefi;;;".;ii;;i"';;.i,=-"r-ident,ifyiTs simuliun damnosum (s' r' )' vectors of human Onchocerciasi3 in West Africa' Larval cytotaxo- nomic identification results are therefore very useful to the Onchocerciasis Control p."gi".ro" (OCP) for both operational and research Purposes' oPERAIIIIOI{AL PT RPOSE The main objective of oCP is to interrupt the- tt11=1i::i:' of River Blindne6s so that it ceases to be a health and socro- economic problem in the programme-area. The idea of the control of the adult females was abindoned at an early stage 3s a result of their large dispersion and for lack of sutlicient knot'rledge of 5their resting place. This, coupled with the absence of any effec- tive macrofrtaiiciae for the tieatment of the adult worm (oncho- cerca Volvulus) Ieaves larviciding as the only alternative method of control of the vector. In the OCp, the severe form of Onchocerciasis (the savanna strain) is caused by the savanna cytospecies. S.damnosum (s'S') ir.ra S.ii.n..,.r^. Lariicidinq operations are therefore mostly aimed ;I-"Iirinating these speci6s iather than the forest cytospecies(S.squamosum, -S.yahense, S.soubrense and S .san:tipauli) which ;ffi-$;=$re ior the transnission of the ress severe form (the forest strain) of the disease. In the tight of the fore-going explanations and the fact that OCp-""*-efrbarks on selgctive ierilt treatmelrF of the diffe- rent river basins, a good kn-owledge of the distribution of the various members oi th6 S.damnosum-(s.I.) complex is very impor- tant to ensqre effective vector control strategy' This is because the different "1.to=p"cies have varying vectorial capacities in iii" ii""smission f human onchocerciasis (river-blindness). rf,for exanPle, a river basin is found to be inhabited by the lessd.rrg.totri cytospecies, aerial treatments could be suspended and thi; is enoirgh Lo save costs in terms of fliqht hours, insecticide, kerosene, perdiern etc... RBSEARCR PT'RPOSES For reasons of research, cytotaxonomic identification could be used to carry out several investigations. For example, tle inversio.rs recoided on the banding pattern of the.giant saliveryqiana polytene chromosomes could be-utilised at time to detennine iesistint-cytospecies as was the case in 1980 and 1981 when S.(s'n: ) were detected to have devetop"d resisiance to temephos and chlorphoxim respectively on the Lower Bandama in C6te d'Ivoire. fn the end, Dr.. R.J. Post discovered a new inversion on chromosome 2 (IfL-A) !o separate sl".."tipi"ii (s.n. ) as the cytospecies implicated in the resis- tance phenomena and not S.soubrense (s.n. )' Chromosome inversion could also be used to follow up the movement of reinvading f1ies. In the OCP, particutarly in theparent that the inveEse reinvasion that occurs annually i-n the btiTnenajari, Kara, Kriran and.Koumongou iir". basins s6on after each yeir's rainy season, originates from ifie-lrinutaries of R. Niger. inis positive conclusion $ras drawn o"-tn" basis of chromosome IS-3 inversion recorded on S.sirbanumpopufations from the tributaries of R. Niger and those other rivers named above. 66 SEASONAL INFILTRATIONS BY SIHULIIJT{ SQUAHOSMI INTO BENIN I qtroductialr In Benin, the Sinulium dannosum Theobatd species complex is represented by S. damnosum Vajime and Dunbar' S' sirbanun Va jirne and Dunbar, the Beffa form of S' soubrense (Mer-edith et aJ.. 1983) and S' squamosum (Enderlein). Bef'ore the start of the tIHo onchocerciasis Control Programme's (OCP) tarviciding campaign' Vajime & Quillevere (Lg?8) recorded s. squamosum in Benin only in upland areas in the Natitingou region' Their map shows sites wrth s. squamosum on the upper stretctres of the Pendjari, Sina Kire, Perma, Yarpao, Koumongou and Terou rivers (the latter site, their number L54' was incorrectly listed in their Table 5 as the Kara river) ' AII the specimens were collected durinq october to March, in the dry season, dt sites which are lnostly perennial' Wet season samples, cotlected in JuIy' August and September, were recorded by Cheke & Garms (1983) who confirmed the presence of s. squamosum in Benin at sites along the length of the Terou and in trre w6w6 (see their Fig. 3). However, since insecticidal treatments began in the Natitingou area s. sguamosum has not re-occurred in other rivers there, such as the Pendjari, but -it has been found in rivers where it was previously unknown. In this paper $re present the ne$, finding'; and suggest that they represent evidence for a regular seasonal infiltration into Benin from perennial sites in Togo or Ghana' a 7Methods Larvae of the S . damn< sum species complex wer ) collected between 1984 and 1993 Ln the Ou6m6 river and its tributaries. The larvae were preserved in the field in Carnoy's fixative (3 parts absolute alcohol: 1 part glaciat acotic acid). The samples were stained and chromosome preparations made as described by Dunbar (t972) and Boakye (L9e8). rdentifications were made using the'chromosome maps of Vajime and Dunbar (L975), Meredith et a7. (1983), Post (1986) and Surtees et al ' (1988)' Results and Discussion S. squamosum larvae were found at l-0 localities where they had not been reccrded before (Tabl-e 1 anJ Figure 1) . The species $ras never found in the ou6n6 itself, only in its tributaries. Most samples invol'red S. sguamosum being in the minority amongst other members of the S. damnosum complex, with the notable exception of the Bor-rgi river collection. AII the positive samples $rere collected in the wet season, during the July to October period, consistent with a gradual infiltration in a northeastwards direction as the rainy season progresses, followe,l by disappearance for November to June. In eastern Gnana and in Togo S. sguamosurn survives the dry season in perennial rivers in, or at the edges of, highland areas- Principal amongst 8these rivers are the Asuakawkaw and Dayi systems and tributarres of the Mono, such as the Amou and the Sio (vajime i,nd Dunbar Lg?8, Vaiime .rnd Quil]6v6r6 197u). i' is known that S. sguamosum regularly migrates not-th and northeastwards, further into Togo, from these mountainous strongholds each wet season (Garms et a7. L982, Cheke and Garms 1983), as did the Djodji form of s. sanctiL',au7i before its eradication (Garms et a7. 1989). Thus, it seems very tikely that the populations in Benin, reporl-ed her6, were the progeny of migrants from Togo or eastertt Ghana. It. is not clear whether s. squamosum has only rer:e'rtly begun to infiltrate into Benin or if similar populations have been overlooked in the past. However, it is possible that the species only occurs :n sufficient quantity to be noticed in those years when there has been good rainfall, permitting opportunities for :-mproved breeding successes. The epidemiologjcal significance of the occurrence of S. squamosum in south and central Benin is trot known. Studies in Togo have shown that it is inefficient as a vector, in comparison with other members of the S' damnosum complex present there, during the wet se.l.son (Garms and Cheke l-985, Cheke and Denke t-988) but becomes a serious threat in the dry season (cheke et a7. L992). _J 9 h2) *_\ er. lrall.l' 8 o lo 160 I Ar a a 6 oa T2 10 l 0 to G 7 6 B I {i ), I t 5(\( -r 6 Mop of Benin showing the sites wherc S.sguomosum lorvqe hove bee,n idantitied during 1984 to 1993. .flro a Fig. l - ( E IO Table 1: Collection sites from where S. squamosum has been identified among populations in the ou6m6 River Basin during the rainy seasion. Species composition : (sq = S.squamosum; so = S.soubrense(Beffa); da = S.damnosum (s.s.); si = sirbanum): a Species Composition NO River Site Date Coordinates Sc So Da Si 1 lKouffo 2 Fonkpodii 24.7.90 r39'N - 1"59'E 1 2 Bougui Bouguidii 30.9.88 759'N -1"4'.E 43 3 Azokan Pt. 17 4.10.91 756'N-1053'E 2 47 1 4 Zou Mount Tob6 6.09.88 8019'N -1051 ',E 6 2 a t 8.09.92 ( I 14 3 I a 21.10.92 I t 12 20 1 5 Adjiro Banon 02.08.84 8036',N - 1056',E 5 16 a a 27.09.91 I I 1 55 6 Odola Odola Confl. 07.09.88 9"50'N -to7'E 3 22 41 I I 01.10.88 I t 4 18 50 i t 06.09.89 ! a 3 36 42 I 7 T6rou I 16.09.86 9012'N-1"50'E 6 1 1 62 8 W6w6 W6wr6 24.07.90 go22'N-2"17'E 17 35 1 9 K6m6tou R.4 01.09.87 goog'N-1049'E 2 2 10 Oudora Tchakparou 03.10.93 9054'N -2"43',E 1 45 4 II ?. GEilERAL DTIIIIRIEUITIOII OF T'EB CTK}I]III'ES Cytotaxononic identification results of .985 to 1993 brere used from t-he cytodatabanl (computer) to prcduce distribution maps ofihe varioui nernbers of tha S.damnosun comple>' in the Eastern oP.- ration Area of the ocP. In general, the savanna cytospecies(S.damnosum (s-s.; anci S.sirbantlm) seetn to be more wldely distri- buted in the East than the forest species' S.dannoeut ( s. s. ) This species is found nainl1 in the transition zone between the forest and the savanna. It has been recorded from a1l the major rivers and their tributaries. ft is the mosE predominant cytotype of the savanna group found in the forest areas of c6te d'Ivoire, Ghana, Togo and Benin (See l{aP 2). S.eLttanul Of the two savanna cytospecies, S.sirbanum is the most widelydistruted in the Eastern OCP. It has been recorded from all theiir". basins in the area although it occurs predoninantly in the savanna zones. The presence of this species in the forest areas is usually during the dry season (See l{ap 3). S.eeuanoann S.squamosum is found in the forest zone breeding. in small riverslE nail-ccasionally been identified from some nediun to big rivers in the savanna zone. In the East S.squanosum seems to bef.tq.fy confined to the Asukawkaw and Dayi ri19r systems in the noufrtairous forest on the Ghana/Togo border. fn the wet season the specles makes 6ome incursion into Benin (see Fig f and l{ap 4). S.yaDgngc S.yahense is found breeding in snall rivers in the forest zone. It itrarely found in the savanna. In the Eastern zone of the OCp its distribuf,ton has been found to be extendipg in southern Ghana due to the insecticidal control of the other cytospecies(Flasorgbor et aI L992). See ltlap 5. Sreanc;tltnulL ftro populations of S.sanctLpauli are referred to on l'iap 6. s.sanct-ipiuii (s.s.) or type forn has been recorded predorninantlyffid,Ivoirea-naSoutht{estofGhana.Thepopu1ation a IE found East of the Volta Lake is known as the OjodJt forn ofS.sanctipauli. This Djodji form has totally disappeared from larval samples as a result of the extension of larviciding to rivers in the south of Togo and Ghana in 1988. S.eoubrnn3g (Beffa). fn uhe Eastern OCP S.soubrense (Beffa) is found only in Benin and Togo (See Map 7). fn Benin the species is found on R. Ouene andits trlbutaries whllst R. Mono and its briutaries and the Slo are the known habitats in Togo. 8. 'IIRAIf,ING - AII the Sub-Sector Chiefs of the Eastern OCP received training in morphology in the course of tl:e year. - A11 Ehe Laboratory Assistants also received training in dissec-tion and norphologrl; with emphasis on dlssection. They are now to betested and those found to be competent be issued with dissection codes. - fhe staff of the cytotaxonomy team in Kara and oEher AREM staff were introduced to a computer progranme. This would pernit then toproduce species distribution maps from the cytodatabank. 9. Pt Brrcf,.rlds Four l,Ianuscripts are in preparation for publication. These are : (iii) Seasonal infiltrations by Simulir.un Squamosum into Benin. The cytotaxonomy of the Sinuliun dannosum (s.s) Theobald(Diptera : Simuliidae) . The cytotaxonomy of the Simuliurn Sanctipauli subconplexin the Eastern Operation Area of the Onchocerciasis Con-trol Programme in West Africa inctuding the C6ted'fvoLre. ( iv) Seasonal patterns of distribution of merubers of the Sinu-liun damnosum Theobald conplex in the OnchocerciasisControl Prograrnme Area in !{est Africa. 10. IEEEIilGg The Senlor Author of this report participated in the following ueetings 3 Research l.leeting, Bouake (C6te drfvoirel 2-6 l,tarch 1993.Discussion of PIan of Action and Budget for 1994 with Chlef VCU,Kara6-9April1993. Bi-Annual Sector Meeting, Kara 26 May 1993.Briefing of Chlef l{edical Officers in tne Oncho Zone of Togo : (i)(ii) T3 Kara, 20 21 JuIy Lgg3. The Cyto Team was invited to give a demonstration of our activities to this session on the last day 2L JuIy l-e93. tnfornil I'teeting with Chief VCU aE Kara, 6 October 1993. East Zonal Meeting, Kara 2 6 Norrember 1993. tr1. A(XrtOnriEDGEltBllrS hle are sincerely grateful to Dr. D. Quill6v6rri (Chief VCU) iDr. H. Agoua (Chief nnlomological Evaluation Officer) ; Mr. G.D. Zerbo (Ctiiet - EOA) i Dr. J.M. Hougard (Chief - WOA) i Dr. C. Back(Chief AREU/fRU) ; Ur. S.A. 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ZON E EST 3' 2' 0' 24 lr' r5' l. 7' 4 'vl- - \ 0 50 l00xn E:- t5' 11 t3' lt' l0' t.. 'r. -/\-)-i\ N,_tI c. /( ( I I I I \ LEGEND sirbonum domnosum ( s.s) squcmosum yohcnsc soubrensc ( Belfo) sonctipouli ( typs) l"v tt h- a* i 21 8 ( t- \ ) v -O\!.J 9' 9' 8' 7' 6 5' 1 oS oS 0s 0s OS rS i ) 7 5 I 5 l0 )\ ( .")(-i I i I.l Gr\ n'/ .., \1A 3 2 t ) I I .L ) )I .\t I I5 t: 6' DISTRIBUTION OF THE CYTOTYPES OF THE S. domnosum complcx FJSTERN OPERATION AREA ( EOA ) l9 93 5 2 Iti 'a 1 3' t' 0' 2' 3' l' I + I t' 25 a Ch . Gauthier Beriaboukro N'Zinoua Dimbokro M'Bahiakro r6t6rro Mafa-Confluencu - Karfiguela Kodjina Amiankouassikro - ?rmouakoukro l.outoukro Tagadi l'oumbiel rroud jolo Bielikpong Yagaba-Gor6ba Zongoiri Rpds Nasia Wawa Weni Sabari Bridge Bukasu Porga Pt. 1 Pt. L Titira LTST OF COI,LEETr}IG STTES R. Ilas-B.rndama R. Bas-Bandama R. N'Zi R. N' Zi R.N'Ni R. N' Zi R.Mafa R.Como6 R. Dior6 R.Comoe R. Como 5 R. Como 5 Vo1ta N:ireVolta {oire R..Siss LIi R. KuIPo.rn R. Kulpawn White Volta R. Nasia white Vo1ta White Volta R. Oti R. Oti R. Pendjari R.ll6krou R. Pendjari R. Keran 28. Sarakawa -R.Keran 29. Landa-Pozadan -R.Kara 30. Bagan -R.Mo3L. KPassa -R.Basa 32. Kechebi -R.Chai 33. Asukawkaw-Br. -R.Asukawkaw 34. D:iodji -R.Gban-Hou 35. Ayagba -R.Gban-Hou 36. rLyi-Hotet -R-DaYi 3?. hudzra -R-DaYi 38. Kati -R.Sio 39. Wonougba -R.Sio 40. Datcha -R.Amoutchou 4L. Amou-Oblo -R.Amou 42. ad6dakoPe -R.Amou 43. KPoba -R.Mono 44. Landa-Mono -R.Momo 45. Tobe -R-Zou 46. Pt 2(B6terou) -R.Orrem6 47. AtchakPa -R.Otteme 48 . M'Bet6ioukou -R - orremei 49. Agolin-Pahou -R.ou6me 50. Gba -R.Oueme 51-. Kaboua -R.OkPara 52. Bassa -R.OkPara 53. TchaParou -R.Oudora 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. L]-. L2. 13. L4. 15. 16. L?. 18. 19. 20. 2L. 22. 23. 24. 25. 26. 27. 26 DISTRIBUTION DES DIFFERENTES ESPECES CYTOTAXONOHIQUES DANS LA ZONE EST ( T99' ) a - Bas-Bandanra et ses affluents : S.damnosum (s.s. ) et S.sirbanurn - Colno6 : S.darnnosum (s.s. ), S.sanctipauli et S.sirbanum. - Volta Noire et ses affluents : S.sirbanum. - Volta Blanche et ses affluents : S.sirbanurn et S.darrrnosum (s.s. ). - 0ti/Pendjari et ses affluents : S.sirbanum et S.datrtnosum (s.s.). - Asukawkaw et ses affl uents : S. squamosunr et S. darnnosunr ( s. s . ) . - Dayi : S. squamosurn et S. damnosurn ( s . s . ) - Sio:: S. damnosum ( s. s. ) , S. s'irbanunr, S. squamosunr et S.yahense. - Mono et ses affluents : S.damnosum (s.s.), S.sirbanum et S.squamosum. - 0u6rne et ses aff luents : S.darnnosum (s.s. ), S.sirbanurrr, S.soubrense (Beffa) et S. squanrosum. - Les affluents du Niger : S.sirbanum. --+.- I\1 i T m BTm-{2-i B6z7m- EH JrnmctC> oJO@to(,l D (t! 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