Entomological investigations for onchocerciasis in South West Ghana (1996-1997): final report

ENTOAAOLO GTC AL INVESTIoATIONS FOR ONCHOCERCIA5IS IN 5OUTH WEST 6HANA (ree6-ree7) FINAL REPORT A WHOIONCHOCERCIASIS CONTROL PROGRAAAAAE AND NOGUCHI AAEAAORTAL INSTITUTE FOR AAEDICAL RE5EARCH COILABORATIVE RESEAR,CH PROJECT December 1999 CONTENTS PROJECT SUMMARY RECOMMENDATIONS CHAPTER I. INTRODUCTION 1.1. Objectives CHAPTER 2. BACKGROUND INFORMATION 2.1. Study Area 2.2. Current Status of Onchocerciasis in Southwest Ghana 2.3. Description of Breeding Sites 2.4. Catching Points CHAPTER 3. VECTOR STUDIES 3.1. Cytotaxonomy 3.2. Cytospecies ldentificationandKaryotyping Page 7 9 10 t2 t3 13 t4 16 t7 18 18 18 J.J 3.4. 3.5. Geographical and Seasonal Distribution of Cytospecies 19 3.3.1. Dry season 19 3.3.2. Wet season 20 3.3.3. Conclusions 2l Vector Abundance and Transmission Levels 22 3.4.1. Fly capture, identifications and dissections 22 3.4.2. Summary of results 22 Seasonal Distribution and Vectorial Role of ,Sizultum damnosum subcomplex (Savanna species) 25 3.5.1. Seasonal distribution 25 3.5.1.1. Dry season 25 3.5.1.2. Wet season 26 3.5.1.3. Summary of results 26 3.5.2 Vectorial role 27 3.5.2.1. Summary and discussion 27 CHAPTER 4. HYDROLOGY AND INVASION STUDIES 4.1. Hydrology Studies 4.2. Invasion Studies 4.3. Summary 29 29 30 Figures 1- I I Tables 1- l0 Appendices List of Figures Figure 1. Figure 2. Figure 3. Figure 4. Figure 5. Figure 6. Figure 7. Figure 8. Figure 9. Figure 10. Figure 1 1. Contents of Figures, Tables and Appendices Map of southem Ghana showing rivers and vegetation zones. An illustration of the complexity of breeding site downstream of Sekyere-Heman (R. Pra). Map of southern Ghana showing catching sites. Mean weekly river heights of R.Tano at Gyomuro bridge (1994 - ree6). Mean weekly river heights of R. Ofin at Adiembra bridge (1994 - te96). Plot comparing the periodicities of fly density at Akanyakrom-Bui, and water level of R. Tano rn 1994. Plot comparing the periodicities of fly density at Akanyakrom-Bui, and water level of R. Tano in 1995. Plot comparing the periodicities of fly density at Akanyakrom-Bui, and water level of R. Tano in 1996. Plot comparing the periodicities of fly density at Asubende and water level of R. Ofin in 1994. Plot comparing the periodicities of fly density at Asubende and water level of R. Ofin in 1995. Plot comparing the periodicities of fly density at Asubende and water level of R. Ofin in 1996. Results of river prospections for Simulium damnosum s.l. Dry season distribution of cytospecies of Sizulium damnosum s.l. on the major rivers in SW Ghana Wet season distribution of cytospecies of ,Sinulium damnosum s.l. on the major rivers in SW Ghana Summary of entomological results for the 1996 and 1997 dry SEASONS Summary of entomological results for the 1996 and 1997 wet SCASONS List of Tables Table I Table 2 Table 3 Table 4. Table 5. PROJECT SUMMARY The Como6-Bia, Tano, Ankobra and Pra-Ofin river systems of southwest Ghana were investigated for onchocerciasis during 1996-1997 with the ultimate aim of assessing whether they can be sources of recrudescence into the OCP area. Prospections for the pre-adult stages of the vector Simulium damnosum species complex were carried out at all the known breeding sites on the rivers. Fly catches were organized at a total of 19 sentinel villages during-six missions in the dry and wet seasons. Simulium damnosum s.l.larvae were collected at about 36 sites on all the major river basins in the region and identified to the species by cytotaxonomy. A total of 28,625 adult female flies were caught and 21,225 were morphologically identified before being dissected for Onchocerca volvulus infections. The infective L3 parasites that were found during dissections were analyzed by PCR-DNA probe method to determine the strain of O. volvulus. The results from cytotaxonomic studies showed that, the savanna vectors of the S. damnosum subcomplex were present on both river basins, but they were limited in their distribution to within the 6" - 7oN latitudes. There, they formed the dominant species of the larval populations in the dry season but not in the wet season at most points. Morphological identifications of adult female flies caught at human bait supported the cytotaxonomic data in terms of geographic and seasonal distribution. In terms of vectorial role, just over l0o/o of the biting flies were found to be savanna flies, but they made up only 3o/o of the infective biting flies. The infected and infective rates of the total savanna flies dissected were 0.7% and 0.42% respectively. The numbers of infective larvae per thousand biting flies were much higher for the forest vectors than for the savanna vectors. It was therefore concluded that the threat of recrudescence posed by SW Ghana being the source of infective migrant savanna flies was minimal whose effect could only be felt over a long period of time. DNA identification of infective larvae found approximately equal numbers of savanna and forest parasite strains (among those that could be identified), and there was no evidence that either the savanna vectors or the forest vectors showed any preference for one strain or the other. The results also indicated that the savanna vectors were less efficient vectors of the savanna parasite strain than the forest vectors. In addition to these unexpected results, the DNA revealed that the savanna strains of the parasite were being transmitted as far south as the furthest catching points of Jema and Sekyere-Heman (lat. 5" I I 'N) which are situated deep in the forest belt. This lack of co-adapted vector- parasite complexes could be due to the pSF I diagnostic DNA sequences not 7 23 RECOMMENDATIONS 1 For this project, @_qq. infective larva/slide was analyzed for strain identity. This means that vital information was lost if there happened to be cases of mixed infections of forest and savanna O. volvulzs strains, and non- Onchocerca species in flies. For special studies such as the present one, attempts should be made to identify a significant proportion of all infective larvae that are found so that their relative proportions can be established in the parasite population occurring in the study area. This information is very relevant and can contribute to our understanding of the epidemiology of onchocerciasis. The DNA-PCR probe based on the pSFl sequences might have worked well so far in clearly defined biogeographic zones, but the lack of the classical parasite-vector complexes observed in SW Ghana seems to indicate that it probably breaks down in other areas. It is recommended that the method be re-appraised/evaluated especially in the forest, forest- savanna interphase zones. It is common knowledge that fly populations responsible for the re- invasion phenomenon observed in the OCP are made up of mainly old flies with infections. Notwithstanding that the infection rates of the savanna flies found in SW Ghana are very low, it could be that flies migrating from this area pick up infections en route. The hydrology and invasion studies convincingly implicate the Tano and the Ofin rivers as sources of two axis of fly invasion into the OCP. The studies also give an indication as to the timing and arrival of migrant flies in the invaded areas. This information can be exploited to plan the schedule of mass treatment with Ivermectin in the two OCP communities i.e. Asubende and Bui-Agbolokame. The Ivermectin ffeatment can be made to coincide with the onset of invasion. However, to be certain, the transmission rates obtained by the OCP during 1994 and 1995 can be revisited and re- analyzed in the light of the results of this study. 9 ( CHAPTER ONE INTRODUCTION The Onchocerciasis Control Programme in West Africa (OCP) was established in 1974 with the main aim of eliminating blinding onchocerciasis as a public health problem. It therefore targeted for control, only the savanna vectoi species and parasite strain(s), using larvicides which until recently was supplemented with mass distribution of the drug Ivermectin. Therefore, only the northem savanna half of Ghana was included in its areas of operation but this was later extended to cover parts of Volta Region. In 1995, the OCP considered that since its operations in the sub-region will end in 2002, there was the need to reappraise its current activities and identiff priority areas for future action. It identified southwest Ghana among others, as potential trouble spots that needed fuither action and research studies in order to maintain achievements and avoid recrudescence of the disease in controlled areas (Report of Technical Meeting, OCP Document: IRU 95.05). Furthermore, the 1995 Expert Advisory Committee reviewed these identified spots and also recofirmended fuither studies and action (EAC Meeting, June 1995, Ouagadougou). The first indication that southwest Ghana could be important in this regard came from a 1994 OCP survey, which indicated that some villages close to the Tain/Black Volta river basin still had high disease prevalence (OCP/EPI unpublished data). Subsequent survey in the area further south but in the same Tain river basin, showed that in five villages 28oh of children of age 5-15 years were positive for skin biopsies. In addition, DNA-based method for detecting Onchocerca volvulus in skin and skin flakes revealed that almost all children 10 1.1. Objectives The specif,rc activities set for the present study are; 1. To cover 19 catching points on the Como6-Bia, Tano, Ankobra and R. Pra- Ofin river basins and tributaries in southwest Ghana. 2. To capture and dissect for infection biting female Simulium damnosum s.l. flies of the points 3. To prospect breeding sites and identify by cytotaxonomy the species of larval populations 4. To collate any available base-line epidemiological data such as from skin- snip, RAPD etc. for the area. 5. Since, scanty information was available to us at the beginning, it was thought necessary that the present study also resulted in the acquisition and compilation of accurate and detailed topographical, climatological, and biogeographic maps, census and demographic data. L2 CHAPTER TWO BACKGROUND INFORMATION 2.1. Study Area The study area is basically the large area of forest in the southwest Ghana. It is drained southwards by a series of sub-parallel river systems directly into the Atlantic Ocean from mountainouslhilly watershed running northwest to southwest in a line north of Sunyani - Kumasi - Koforidua. The uplands along the watershed consist of montane forest vegetation (Antiaris/Chlorophora association). North of this is guinea savanna, and the rivers drain into the Volta River system. South of the highlands is moist semi-deciduous forest (Celtis/Triplochiton association) which grades into tropical rainforest in the extreme southwest of the country (Figure 1). In the south of the study area are found coastal plains, swamps and lagoons and these limit the distribution of S. damnosum s.I. The Bia, Tano, Ankobra and Pra-Ofin and tributaries, the four significant river systems in the area, can be divided roughly into three sections; the upper, middle and lower sections to coincide with the different vegetation zones. The upper northern parts flow from and are near the uplands with the montane fype of vegetation. The middle sections flow through moist deciduous forest and the lower sections run either through tropical rain forest in various stages of deforestation as with the rivers Bia, Tano and Ankobra. Or as with the R. Pra, through moist deciduous forest and the transition between moist deciduous forest and tropical rain forest. t3 Research unpublished report, 1999), as well as this report provide supporting evidence of the role of S. sanctipauli subcomplex. However, along the upper reaches, the savanna species especially S. damnosum s.str. occur and may play a major role in disease transmission there. In fact, from casual observations made of the inhabitants living along the upper reaches of the Tano River for example, it was clear that the blinding savanna type of the disease was being transmitted Moreover, the picture of distinct forest and savanna distributions is not the case on the ground, because of deforestation and also of human migration. Deforestation activities are creating derived savanna enclaves within the forest that might result in the southward extension of the savanna vectors and parasite (Baker et. al., 1990), thus extending the need for controlling blinding onchocerciasis. The reasons given are that the derived savanna will be suitable for the perennial breeding of savanna vector species and that human migration along the north-south axis of West Africa is also leading to the introduction of the savanna parasite strains into forested areas. Prior to this study, previous investigations by the Principal Investigator have all revealed that the savanna flies, especially S. damno,srlru sensu stricto occur in some river basins in SW Ghana. Intense human activities such as logging, farming, surface mining etc. in the area is leading to deforestation, which is widespread. The region is also characterizedby intense human migration, which is mostly in the north-south direction. Therefore all the elements that can contribute to the establishment of blinding onchocerciasis in SW Ghana are in place. Although the extent of blindness or visual impairment resulting from infections with onchocerciasis has not really been determined for this area. l5 2.4 Catching Points Catching sites on the Bia, Tano, Ankobra, Pra-Ofin and tributaries were chosen based on their location (nearby significant breeding site) and, accessibility (by road/vehicle) and to infrastructural needs such as availability of utilities etc. Most of the sites are basically small villages with populations of hundreds, and Nineteen sentinel villages, an initial five on the R. Tano and nine on the Pra- Ofin basin were chosen for adult fly collections in the first year. One site, Ntobroso (R. Ofin) was later dropped earlier in the study after preliminary catches yielded no flies and that at Akim-Oda (R. Birim) was substituted for Kusi Camp (R. Birim) at the end of the first year. Five additional sites three on the Bia and two on R. Ankobra were studied only in the second year. For the locations of these sites see Figure 3. To obtain detailed topographical maps of the area, survey maps (1: 50,000) and remote sensing images (Landsat TM) were used to obtain digitized image data initially for the whole country and then for the area of interest. A global positioning system (GPS) was then used to obtain the exact coordinates of the big breeding sites and catching points. l7 CHAPTER THREE VECTOR STUDIES 3.1 Cytotaxonomy Regular prospections of all the breeding sites were carried out and specimens of S. damnosum s.l.larvae found were identified by cytotaxonomy and the results analyzed to reveal their seasonal distribution in the area of study. For the list of breeding sites visited and the results of prospections that were carried out during the period of this study see Table I 3.2 Cytospecies Identification and Karyotyping During new cytotaxonomic prospections for this project full karyofypes have been regularly recorded to enable an assessment of whether new (previously uffecognized) cytoforms or cytospecies occur in the study area. New data from the present studies were also compared with those of previous studies carried out in 1995. There is no evidence for a new species in the study area. Some cytotaxonomic variation in inversion frequencies were recorded, but it is considered that this is not sufficient evidence for new cytoforms within S. damnosum s.str., S. sirbanum, S. squamosum, S. yahense and S. sanctipauli. No S. soubrense was found in the samples, but this is not surprising because it was always rare in the study area. A previous report (Boakye et al, in manuscript, The cytotaxonomy of S. sanctipauli subcomplex in Ghana and C6te d'Ivoire) describes significant cytotaxonomic variation between the major river systems of southern Ghana and C6te d'Ivoire. Populations on the river Pra are considered distinct from t8 yahense were recorded at sites, which were closer to the 6"N latitude. However, above the 6"N latitude, S. damnosum s.str. was dominant at almost all sites even in the mountainous regions of the principal tributaries, R. Ofin and R. Anum.. For example at Pokukrom (lat. 6"01'N) which is situated on the middle of R. Ofin, it formed approximately 88% of the specimens (total of 73) identified there. At Anniamoa (lat. 6o33'N), which is located near the upper reaches of the same river both ,S. damnosum s.str. and S. strbanum were the only two species identified from the 76 larvae collected in April 1997. Simulium yahense was recorded mostly at sites near the hilly areas of the middle and upper reaches of the Pra-Ofin river system. No ,S. squamosum was ever recorded among the samples that were studied. A small number of breeding sites were found on river Bia and all were located above latitude 6"N. ,Sizulium sanctipauli was dominant at two sites, but S. damnosun s.str. was recorded at all the three sites. A few S. sirbanum and S. squamosum were also recorded together with S. yahense. Only two species, S. yahense and S. sancttpauli were found at the two sites on R. Ankobra and its tributary river Ehyere. The two sites are located below latitude 60N. 3.3.2 Wet season The wet season distribution of members of the ^S. damnosum complex is shown in Table 3 and it revealed that S. sanctipauli was the only species recorded on the lower and middle section of the R. Tano. There were only two specimens of the savanna cytospecies, S. damnosum s.str. and these were recorded at Ntotoroso around latitude 7"N. In the upper Tano only S. yahense was found. 20 Although the number of identifications made during the wet season is relatively on the low side, the general impression is that of fewer savanna cytotypes occurring in the study area during the wet season. 3.4 Vector Abundance and Transmission Levels 3.4.1 Fly capture, identification and dissection A total of 19 catching sites were covered during the study period. Each site was visited during the four missions of three days catch per site that covered the dry and wet seasons. Every fly was dissected, except the third day's catch which was dissected to determine the physiological age only and then preserved in 70%o alcohol for further morphological and molecular studies. The standard OCP protocols for fly catching and for determining indices of biting and transmissions were used and each fly was morphologically identified before dissection. The infective larvae that were found were sent to the OCP's laboratory at Bouak6, C6te d'Ivoire for DNA identification to determine whether they were forest or savanna strain of O. volvulus. 3.4.2 Summary of results For ease of comparison both I 996 and 1997 data are presented together. A total of 28,625 adult female flies were caught and 21,225 were morphologically identified before being dissected for infections. The details of biting densities and transmission levels expressed as monthly biting rates (MBR) and monthly transmission potential (MTP) for all sites in both the dry and wet seasons are shown in Tables 4 and 5 . It was found that, the lowest fly densities on the R. Tano basin were recorded at Tanoso with MBR ranging from 19 (dry season) to 238 (wet season). The 22 No transmissions were recorded at Bonsa during the wet season, but in the dry season the MTP was 78. The dry and wet season MTP values for Hiawa were 225 and 487 respectively. On the Bia basin, the MBR values ranged from zero to a high of 3,364 at Dadieso during the wet and dry season. The MBR values (wet and-dry season) for the other two sites were; 2129 and 878 at Kwesi-Addaekrom, and 31 and 93 at Eluokrom. No transmissions were recorded at both Eluokrom and Dadieso, but at Kwesi- Addaekrom the MTP values were 52 and 435 in the wet and dry season respectively. 24 The savanna flies formed significant proportions of flies at all sites on R. Bia, where they formed 67oh of biting flies at Eluokrom,34o/o at Kwesi-Addaekrom and 22%o at Dadieso. 3.5.1.2 Wet season It was observed that during the wet season, the savanna flies were still present on all the rivers where they occurred during the dry seasons, but their abundance were much reduced (Table 7). For example at Kpoikrom (R. Tano) the proportion of savanna flies had reduced from 92'/o in February 1996 to t6Yo in July of the same year. At Nsawora (R. Tano), their proportion of the biting population was reduced from 75%o and87o/o dunng the dry seasons of 1996 and 1997 to L6% and 4% respectively during the wet seasons. However, at Gyankobaa (R. Anum) the proportion of savanna flies did not follow this trend in 1997 , but instead increased from 630/o in the dry season to 7 5o/o in the wet season. At higher latitudes, savanna flies persisted in significant proportions at most sites during the wet season (e.g. at Tanoso on the R. Tano and at Anniamoa and Bontomurso both on the R. Ofin). Generally, no savanna flies were recorded in the wet season at sites where none or less that l%o was identified in the dry season in the dry season. These sites include Jema (R. Tano basin), Bonsa (R. Ankobra basin) and Sekyere-Heman (R. Pra-Ofin basin) which are all located below the 6"N latitude. 3.5.1.3 Summary of results The seasonal and geographic distribution of biting savanna flies of the S. damnosum complex during 1996 and I 997 is consistent with the general pattern revealed by larval cytotaxonomic identifications (see section above 3 above). 26 These results are unexpected according to the classical model of co-adapted vector-parasite complexes originally described by Lewis and Duke, and Quillevere. That model would predict that in the forest bioclimatrc zone, savanna flies would be less efficient vectors than forest vectors and that is what these data show. What is unexpected is that the savanna flies are also less efficient vectors of savanna strain pgslt: tE1--?fe _lhg-Qrest v€ctors. If this interpretation were tme it would imply that any event such as deforestation, .lorg and severe drought etc is unlikely to have undesirable consequences for onchocerciasis transmission in SW Ghana. As long as the biting rates do not change, the occurrence of savanna flies in SW Ghana would not necessarily lead to the replacement of forest strains by the savanna forest strains which are believed to be associated with blinding onchocerciasis. What this means is that the threat posed by SW Ghana as a source of in the OCP is minimal and if this is the case, then its influence can onlY/\srgn1 ficant over a long period of time. However, there is an alternative interpretation of the data, and that is that the DNA identification of the parasites strains is not working for the parasites occurring in SW Ghana. Under this interpretation there would be no variation in the pFS 1 DNA sequence family, but that variation would not be linked to the epidemiological strain variation in the parasites. 28 CHAPTER FOUR HYDROLOGY AND INVASTON STUDIES 4.1 Hydrology Studies Hydrological data of river gauge readings from 1994 to 1996, of the rivers Tano and Pra were obtained and then analyzed to establish hydrological stability of the river systems in the area. The analysis revealed that the river levels were significantly variable in timing and height between 1994,1995 and 1996 (Figures 4 and 5). 4.2 Invasion Studies The hydrology data (see section 4.1) were used in analyses to determine if there was any correlation between these variables (i.e. timing and height) and the onset and periodicity of invasion at certain points in the OCP area of Ghana. Two sites in the OCP area which are seasonally invaded by flies and lying upwind of the SW monsoon were selected for analysis, to determine if the periodicity of river levels was correlated to the onset and invasion. Akanyakro-Bui (R. Black Volta) and Asubende (R. Pru) were the sites chosen in the OCP area for analysis. The weekly biting rates (WBR) of Akanyakro-Bui and the river level periodicity of R. Tano were compared, and Asubende and R.Ofin were similarly treated. The results obtained revealed that during the first 15 weeks of 1994 and 1995, the two parameters were not correlated, but there were strong correlation from then on between WBRs and river levels (Figures 6-11). The L996 data however did not show the same degree of positive correlation observed for the two previous years (Figures 8 and 11). This might 29 CHAPTER FIVE COMPILATION OF EPIDEMTOLOGY, DEMOGRAPHY, CLTMATE AND TOPOGRAPHY DATA 5.1 Epidemiology Data The data of reported onchocerciasis cases at hospitals, clinics and Health centres in southern Ghana for the last ten years has already been compiled and are deposited in an electronic database created at NMIMR. 5.2 Demography Data The last population census in Ghana was held in 1984 and as a result any current information of the demographic and economic characteristics of the study area will have to be extrapolated from this data. The data have been published as series for each administrative region and the relevant ones for Western, Brong Ahafo, Ashanti and Central Regions (Boateng et al., I987a; 1987b; 1987c; 1987d) were acquired during the period of this project. However, the contents are but summaries and the format of the publications is such that it will be difficult to analyzed the data specifically for the populations and villages at risk. It is doubtful if such information can be had from the existing government agencies. Ideally, a comprehensive survey of villages at risk should be carried out and the data managed using geographical information system (GIS). 3t ACKNOWLEDGMENTS We are very grateful to the Director of OCP, Dr Yankum Dadzie and Prof. David Ofori-Adjei, Director of Noguchi Memorial Institute for Medical Research, the present Chief VCU, Laurent Yameogo and his predecessor, Drs. Jean Marc Hougard and H. Agoua for their support and encouragement. Mr. George Fiasorgbor, OCP Kara and Mr. Michael Osei-Atweneboana carried out the cytological analyses. Dr. Laurent Toe and staff carried out the DNA identification of the parasite. We also appreciate the support given to us by the Ghana Sector of the OCP, especially, Mr. S. Sowah and Mr. Naps Fusseini. Mr. Anthony Tetteh-Kumah, Messieurs Christoff Tettegah, Mr. Boadu, Asaah- Asare, Sampson Otoo provided technical and logistic support. 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UJoo -oz o E. o lJ- U)z o tr o tr U) uJ z J o o oJ o oFz LU G 'a c o o o_ c .9o 3EEoroc. E'-o *>, >r: !o z illlo-EFG] oOrOo$(f)OOtit (Y) cO(fr_ @_ (o_ r-@ NO@tr)co(o o{ r-- o_(ocoN 9o.o .n-$ 3 EPBNO(O@oos (o l'- oco(f) (f)(f)(f) NCo (f) oo o oo o (oN o(f)(o -$ os3 O-@ OO O Ese 3-. EB€ -.oN (a (o o) r tf) (O(r)sd) 33H Or O rr)N F.- rt o(f)o(f)@s --@ lrrcoS8E^- EE L -. Soa Q <o r'-BEE ::;.(trX(E ><*> EEE L..>.(6=(U > a> f.- o) O) (I, E in o E(, o 'a =Y E in oD o I E = E fC (o(u _o ovc(o o (! o_ c(o E o -I c) (I) -Yoa & co a o-cEO ,{(I,ilo .z- o-F (O J 9E1a I o (o E. o)c in o(,tr co o o) C o o G) c oo o o) C a foL o o- :o o o oa .9o E o)L =o-(g o o o)(! E o -- U) tr u) oe E(Ub u) o (l) -o E =z (l) (o o o) E. (l) =U) ocoo(, oa -E f.* O) o, oc(u (o o, O) r o .c. Lo a = =ooL (o .9(,) o o E o c o o z-(I, E E :fa ii o)lc c o(,) s o _o(t,F z I o F U) tU 3 IF :) o U) z 9. 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'- o o o) E C. o c.o .E f -o .E .a! o .c.F iiq) :l c c oo(o o) _o C'F o E. lJ- F U) lU = -F :f o @ z U) a (J E, lljoo -o z. o x. o lJ- U) z. o tr o tr @ TU =J I ooJ o oF z. tU 6 .F C, o o o- c o 'U)a -ta iE cLbqc.E(g= .!= -o2.>E -C; LL oO ll ll oGFCO oo o oo o F-toBs6*ooo ooo ooo ooo ooo oo o ooo ooo oo o ooo ooo oo o ooo oo@ oo o NNN NNN C{N O SEE Esg o(ao(o@s rr@ oo o l'*$ r F*S r ^^a;ro -o o\o-o\m sRgl=od.r3 (O o f.- N@s oo(o tooB8E-- N o, O) o E in o_ E(I, o a :lY E in (o Eo I Ev E =c (U(I, _oo _v.C(, o (E L o_ C(U E o :tr I o)L o) -)< c) U) EE L .- Soa Q <o tt-833 ::;(o=(U HHi r-:, g F: (l) o G) C, lC(l) o o) c Eo() G)a .9.o aq) (I, E(t) o Lq) -o E f z. t m = o-F (O J9E1d) :tr ogE .(oooz (gC\o Eo(EOu)a .ro9.q o'E !q)2E E o) =ooo o .qo(l) o-a Lo o o (o cco G, U) U' o (I, E(l) j a Ef, .4 oq E(oB U) o oz G) (E o L(D t (l) =U) ac oa(, oo b ro F.- o) o, E c G,(o O) O) o)c .E =o xq) o- E oo -o)a E .t oc E(gb u) siLo oo oCc(I' oa o o og o .E o() q) ac(U Co .F f -o .9E oEF .U o = 'a c oo(o o -o(oF o d. tL F U) TU =-F =oa z U) a o E. IIJoo -o z. o & olr U)z o tr o tra LU z J o ooJ o oFz tU ooo)f.-lr)5$str) r.-@-Oo.l No5E(oC\lN -(Ol() oo ooo ooo ooo ooo oo o o o ooo ooo ooo ooo oo o o oooo ooo ooo ooo oo o o ooo ooo ooo ooo oo o o o oPSo N@(O OO f.*s oo rSt() ONC.) NN(O c\tN(f) NN(f' NC{(f) NN (.) (O (f) N N r.- o, o) f - .q co No F.- r C\lo F.-- F- O) o) = - (o L -oolzc F- o{ r.-ol F.- o, o) f - C,L -ooYc o (E L coo'ai6 .eLC eb -Lc =C, -oE E. -CCC.Oc) ll il Eo_coF EoL .Yof LU oaco co C' =.gI C' E o -) \oo\ -o$o\go3 (o lf, \O ^\oo\ \o o\(oo)@ c.i I a'j -o -N(O sslr)(o(r)rO OrS ss oQnr (r) c! €)EgB -$@ 3$s 3$s (o t-o) o)O, O, ff -- (O (O F-O) o, O)o, o, o) >>>(IT:: 2-- Esu s 5i EEE e= +>:< EEE e. q,>=< oc(! 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'a a cIE o E o -Yflzo o_ E o (o o E .gcc o <t) =E o C o co .g CD oaIE(U o .o co E o -v.o(o E ro U'o 3Y E G] o_F (o J9E1o) - a -sE .(oo(J z- C,C\o Eo-G, o-@a o ,F o o) c :o o o o) c Eq) oq) a .9.o a o) o E o) o L c) -o E =z. a .ooo o- @ o oo occ(U (o U) .Eoo .C IEo- 'F(J q) ro oL :l o-(E o oo(s E o) J a E q o E(!B u) o o z- o (o o (l) t o -_U) 2 oa!(tr .L{ o)d </)/:\ -r5\ t-\ o, o, EC o(o O) o, o)c =ro xo -o- E o(,) -o =a U)o (!B u) U; Lo oq) (E C c(I, (I, a o o oL (o .E o (.) o) oc(U c o f -o .9o (I) -cF iiq) f,C 'a c oo l'- o -oGF z -o F U) TU =IFf o U) =U) a ot LUo o -oz o E. oII U)z o tr(, tr U) tU z J 9 o oJ o oFz LU g C,L c o fiELC PE =o -oE -C. E =uLCOo llil Eo_(DF oS-B o_ P .e_ c!"(o ooo ooo ooo oo3 oo oo o oo ooo oo o ooo oo(o oo o ooo oo(f, oo o ooo ooF.- oo o $pE NNN Nc\IN NN (f) soN tr) ^^Gss=&i pS;; -Nr rf or-- 5 ,o;o_x o:F8ro--@*ir-O)(0lr)0)NF-cO EE >. o) 5< <o I r-- E9E >o> =--a= -<--) <o I r'.-o) )< o) o) uJ o) > 9> = --l :f -<-- @ @ E in o:o o I Ev E =c o(! -oolzc(o (, (g L o_ c(g E G)I I(D Lq) lz oa Srlt-(o(ooNr'-c.) tr) F- O) o) l - E in o_ E C'O .U' =Y & co o-F (f) J9ELo - agE .ooo z. (oC\oE;(I' 0- tDa o oo c io o) o .9 C Eo o c)o .9,o ao (\, E e o Lq) -o E :f z. a o '5 q) o-o o o o (g C C(I, (o U) .!oo) oo) o o)L f o-(U O a o o E o J a E U) o Eo 'o U) o oz o (g o q) t o =U) a C. oaooa a.E r.- o) O) E c(o (o o, O) o)c =E x c) o- Eoo -of a U) o (UE u) ui o o c) occ o oo o o oL o .E o oq) E c(U C o f _o .9.ro o _c.F ,o o) fc c oo f.- o) _o o F- z -o F. U) tIJ =IFf o U) =U) a oE tu() oIoz o x. o lJ- U)z o tro tr U) I,JJ z J I o o -Jo oFz ul Season No. Flies Caught No. Flies Identified No. Savanna Flies (7o of Total) No.Infective Flies No.Infective Savanna Flies (o/. of Total) Dry 10,841 7,954 1,395 (17.5o/o\ 149 6 (4.0%) Wet t7,784 13,271 845 6.40h\ 126 3 (2.4%) Total 28,625 21,225 2240 (10.5oh') 275 9 (3.3%\ Table 8.1 : Summary of dissection data from 19 sites in SW Ghana during 1996 and 1997 Table 8.2 : Numbers of infective larvae per 1,000 biting flies Vector species Wet Season Dry Season tr'orest 55 111 Savanna 7 6 Table 8.3 : Summary of the results of DNA identification of infective larvae from all flies Species Strain Numbers Identified f,'rom : All Flies Savanna Flies Onchocerca volvulus Savanna Forest Unknown 85 77 o5 I I 3 Ortchocerca ochengi I 0 Non-Onchocerca 2 I a .9 o, o x. c o o o il E. c .9 o) ou g o o(U trJ t D c .9 O) ou (E oO ll x O i .9(r, ou o (u _c cnc o c0 il m c .9 o, c)E c6 -co ll Ia oc! tr) oF-:6 c{ (o;r.ri(os Nr (oros c.ioo r$(O qqct(ooc') oo)c) (o.t(f) or+(arf) sl @ (f)(OF-3gg(f)N o)@@ -t ci c.i@rC.) 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