Medical and Veterinary Entomology (1998) 12, 267–275

Distribution of the Simulium damnosum complex on

Bioko island, Equatorial Guinea, and the potential for
onchocerciasis elimination by vector eradication
P. J . M c C A L L , 1 R . A . C H E K E , 2 M . D . W I L S O N , 3 R . J . P O S T , 4
P. K . F L O O K , 1 R . M A N K , 5 A . S I M A 6 and J . M A S 7
1Liverpool School of Tropical Medicine, U.K., 2Natural Resources Institute, University of Greenwich, U.K., 3Noguchi Memorial
Institute for Medical Research, University of Ghana, Accra, Ghana, 4Natural History Museum, London, U.K., 5Department of
Entomology, Wageningen Agricultural University, The Netherlands, 6Ministerio de Sanidad, Malabo, and 7Programa Control
Onchocercosis, University of Barcelona/Cooperación Española, Malabo, Bioko, Equatorial Guinea

Abstract. Onchocerciasis is endemic on the island of Bioko, Equatorial Guinea,

where it is transmitted by the ‘Bioko form’ of the Simulium damnosum complex, a
cytospecies unique to the island. To determine the distribution of vector breeding,
three dry season and two wet season expeditions were made in 1989, 1996 and 1997,
and 226 of the island’s 247 rivers (91.5%) were visited. Of these 226 rivers, 130
(58%) were flowing during the dry season, forty-five (20%) supported aquatic stages
of Simuliidae of any species and twenty-five (11%) contained larvae or pupae of the
S. damnosum complex. The twenty-one rivers not prospected were in the mountainous
south of the island, where an additional seventeen rivers were reached but not
satisfactorily prospected. Of these thirty-eight rivers, twenty-nine were considered
highly likely to support vector breeding, bringing the total number of rivers which
could harbour the vector during the dry season to fifty-four (21.9% of the island’s
total). Breeding was believed to be limited to river stretches below 1000 m altitude,
and during the dry season the total length of those stretches which could support
breeding on Bioko was estimated to be 1020 km. A combination of factors, including
low river discharges during the dry season, the relatively low water temperature on
Bioko, the suitability of limited stretches of most rivers as vector breeding sites and
the close proximity of many rivers within a small geographical area, render the vector
vulnerable to eradication by aerial treatment of rivers with insecticide. The isolation
of the Bioko form of the S. damnosum complex suggests that reinvasion following
treatment would be unlikely, and eradication of the vector might be achieved by a
dry season larviciding programme in one or two years.
Key words. Simulium damnosum, Simuliidae, onchocerciasis, vector control, Bioko,
Equatorial Guinea.

Introduction in the Bight of Bonny µ 40 km south-west of Mount Cameroon

The island of Bioko (formerly Fernando Póo) is 72 km long
and 35 km wide, covering an area of about 2000 km2, situated
Correspondence: Dr P. J. McCall, Division of Parasite and Vector
Biology, Liverpool School of Tropical Medicine, Pembroke Place,
Liverpool L3 5QA, U.K. E-mail: mccall@liv.ac.uk
(Fig. 1). Together with the mainland region of Rı́o Muni and
the small island of Annobón to the south, it forms the Republic
of Equatorial Guinea. Onchocerciasis occurs throughout the
island with a prevalence of 75% (85% of those older than
15 years), almost the entire human population of 62 000
inhabitants being at risk of infection (Mas et al., 1995).
Bioko is dominated by three extinct volcanic peaks: Basilé
(3011 m) in the north, Caldera de Luba (2261 m) and Biao

© 1998 Blackwell Science Ltd 267

268 P. J. McCall et al.

Fig. 1. Map of Bioko with the major geographical features and human habitations, showing the location of rivers known, or considered highly
likely, to support breeding of the Bioko form of the Simulium damnosum complex. Ninety of the largest rivers of the total of 226 prospected are
shown. Rivers where aquatic stages of the vector were found during the dry season are marked with arabic numerals, whereas rivers where vector
breeding was seen only in the wet season are marked with roman numerals: see Table 1 for details. Known positive rivers are numbered within a
circle, and unprospected rivers and rivers considered likely to support vector breeding but not yet fully investigated, are numbered within boxes.
The wet-season-positive rivers marked (iii – v) and (viii) are outlined in dotted circles, as their precise identity or location have not yet been
confirmed. Points labelled A, B, and C in the south of the island mark regions referred to in the text.

(2001 m) in the south (Fig. 1). Apart from Malabo, which is
the main city on the island and capital of Equatorial Guinea,
there are two regional towns, Luba and Riaba, and one large
village in the south, Ureca. About half the population inhabits
the small villages or cocoa plantations outside these towns,
mainly in the north of the island. Although there is some
cultivated land, the island is clothed in tropical rain forest up
to the higher montane areas. Much of this forest shelters the
cocoa plantations, many of which are now derelict. Extensive
areas of uninhabited natural primary forest remain in remote
areas, typically in the southern part of the island, which is
accessible only on foot. Rainfall is very heavy but seasonal,
with many rivers drying up soon after the end of the wet
season. Bioko is one of the wettest areas in Africa, and the
south, where annual rainfall can exceed 14 m (de Teran, 1962),
is one of the wettest areas in the world.
Onchocerciasis prevalence is high everywhere, with infection
rates of 76% occurring at sites only 3 km from the capital,
although the highest prevalence of 87.1% was recorded at
Ureca (Mas et al., 1995). Malabo serves as the base for the
Programa Control Onchocercosis, a collaboration between
Departament de Microbiologı́a y Parasitologı́a Sanitaires,
Universitat de Barcelona/Cooperación Española and the
Ministerio de Sanidad, Malabo, Guinea Ecuatorial, which
initiated studies in 1987 (Mas et al., 1995). In 1990, control of
onchocerciasis began using the microfilaricidal drug Mectizan®

© 1998 Blackwell Science Ltd, Medical and Veterinary Entomology 12, 267–275

(Merck, U.S.A.), and since 1996 it has been administered once
a year to the entire population of the island.
The only known vector of onchocerciasis on Bioko is an
endemic member of the Simulium damnosum Theobald species
complex (Diptera: Simuliidae). Evidence from cytotaxonomy,
morphology, isozymes and DNA studies of this ‘Bioko form
of S. damnosum sensu lato’ confirm that it is taxonomically and
genetically distinct from other populations of the S. damnosum
complex on the African mainland, classified within the
Simulium squamosum (Enderlein) sub-complex and closely
related to Simulium yahense (Vajime & Dunbar) (Wilson et al.,
1994; Post et al., 1995). The Bioko form of S. damnosum s.l.
is a serious biting nuisance, with vectorial capacity comparable
to the highest transmission rates found in other forest vectors.
The infection rate with Onchocerca volvulus has been estimated
at over 400 infective larvae per 1000 biting flies (Cheke et al.,
1997), which is comparable to the high rates of transmission
found in S. yahense in Ghana (Cheke et al., 1992) and Côte
d9Ivoire (Quillévéré et al., 1977).
As an island with endemic onchocerciasis, disease
elimination from Bioko by vector eradication is an attractive
prospect. The geographic isolation of the vector population
suggests that, if eradication were attempted, the likelihood of
reinvasion would be very low, because (a) the Bioko form has
not been found anywhere else, and (b) the prevailing winds
blow almost constantly from Bioko island towards the African
mainland, making it difficult for S. damnosum s.l. to reach the
island from elsewhere. Until the Programa Control
Onchocercosis was set up in 1987, the only previous blackfly
records from Bioko were those of Calvo Picó (1962), Lewis
& Duke (1966) and Duke et al. (1966), who collected and
identified adult female flies of S. damnosum s.l. landing on
human bait at a few sites in the northern part of the island.
Our preliminary surveys in 1989 (Post et al., 1995; Flook &
Post, 1997) suggested that S. damnosum s.l. was restricted
during the dry season to a limited number of rivers, raising
the possibility of its eradication from the island by insecticide
treatment of breeding sites at that time of the year. Here we
report on a comprehensive survey of Bioko’s rivers and confirm
that, during the dry season, distribution of the S. damnosum
complex is indeed very restricted, and that vector eradication
may be feasible.
Materials and Methods
Rivers were visited and surveyed for Simuliidae during the
dry season in May 1989 (R.J.P., P.K.F. and J.M.), April and
May 1996 (P.J.M., R.A.C., A.S. and J.M.), and April 1997
(P.J.M.). Additional expeditions were made during the wet
season in November 1989 (P.K.F. and J.M.) and in July 1996
(M.D.W. and R.M.).
Attempts were made to visit all rivers, especially during the
dry season expeditions, whether or not they were marked on
maps of the island (1:50 000, Instituto Geográfica Nacional de
España, 1980, 1981). For surveys in 1996 and 1997, the precise
location of each river visited was determined using a portable
Global Positioning System (GPS; Garmin GPS 40 Navigator,
© 1998 Blackwell Science Ltd, Medical and Veterinary Entomology 12, 267–275
Simulium damnosum s.l. on Bioko 269
Garmin International, Kansas, U.S.A.). Where the forest canopy
was too dense to use the GPS, and for all prospections
prior to 1996, the locations were determined from the maps.
Prospections were carried out in all accessible flowing rivers
and streams. Aquatic stages of Simuliidae were detected by
examining trailing vegetation or rocks and stones in the running
water. Late-instar larvae of the Simulium damnosum complex
were recognizable by their dark colour and (using a hand lens)
by the presence of short, broad, bluntly pointed spines on the
dorsal cuticle (Crosskey, 1969). Larvae or pupae were removed
and preserved in 100% alcohol (for DNA analysis) or Carnoy’s
solution (for cytotaxonomy). Voucher material was deposited
in the Natural History Museum, London.
Results
Results of prospections
The island of Bioko contains 247 rivers and streams of
which 226 (91.5%) were examined. The remaining twenty-one
rivers were inaccessible and could not be visited during any
of the expeditions. A total of thirty-seven (16.4%) rivers were
found to contain aquatic stages of the S. damnosum complex
at some time of the year. The most comprehensive surveys
were carried out during the dry season, in particular during
April and May 1996 when 224 rivers, including those in the
south of the island, were visited. During the dry season, only
130 of the 226 rivers (58%) were flowing and forty-five (20%)
of these supported populations of various Simuliidae (McCall
& Cheke, unpubl. data). Larvae and/or pupae of the Simulium
damnosum complex were found in twenty-five (11%) rivers
during the dry season and in an additional twelve rivers during
the wet season prospection in November 1989. Table 1 lists
the rivers where aquatic stages of the S. damnosum complex
were found, with the relevant hydrological data for each river
where known, and the length of river which is estimated to
support breeding of the vector. Figure 1 shows ninety of the
major rivers on Bioko, with the locations of the known and
potential breeding sites of the vector marked and numbered
according to Table 1.
Access to most rivers in the northern part of the island was
possible by road. It was sometimes possible to visit these rivers
at different reaches upstream and downstream of the main
road, although treks on foot of up to 4 h were necessary in
some cases. However, there are no roads in the southern part
of the island and prospections to all of the rivers south of
Riaba in the east to point C on the western coast of this part
of the island (Fig. 1) were made by landings from the sea
during April and May 1996. It was possible to examine these
rivers for no more than a few 100 m from their mouths.
Prospections in the south of the island were further limited by
the need to travel along the beach as the only means of gaining
access to rivers in the region. High cliffs to the north of Punta
Oscura as far as point C (Fig. 1) prohibit entry from the sea,
and the mountainous inland terrain renders them inaccessible
on foot.
Of interest was the proximity to the sea of aquatic stages of
270 P. J. McCall et al.

Table 1. List of known breeding sites of the S. damnosum complex on Bioko (see Fig. 1), with the dates on which the material was collected,
and hydrological data for each river. *Length refers only to the estimated length of each river which could support the aquatic stages of
the S. damnosum complex. n.r. 5 not recorded. Hydrological data were not recorded during the wet season visits of 1989 and 1996.
Map numbers correspond to Fig. 1.

Temp. Discharge Length*

Map No. River Grid Ref. Date Material Collected (°C) pH (m3 sec–1) (km)

1 Ruma 3°239180N 8°449180E 23.05.89 lv 15

14.11.89 lv
23.04.96 36 lv 1 1 p 23.8 7.98 2.41
19.07.96 lv
Ruma – upstream 3°329500N 8°429550E 23.04.96 lv
2 Grande 3°239180N 8°449180E 23.04.96 11 lv 1 p 23.1 7.98 1.86 30
19.07.96 lv
3 Muedede 3°179100N 8°449030E 29.04.96 lv 1 p 26.2 7.40 0.57 22
4 Iladyi 3°159180N 8°429380E 30.04.96 4 lv 1 4 p 24.6 8.12 4.46 30
5 Leke 3°149050N 8°369450E 02.05.96 1 lv 23 7.45 0.28 10
6 Ole 3°159510N 8°289280E 03.05.96 1 lv 26.1 6.15 2.79 30
7 Cibitá (Tope-Bue) 3°269140N 8°269210E 09.05.96 10 lv 24.6 7.50 0.12 18
8 Booló 3°279050N 8°359280E 24.04.96 lv 1 p 22.6 8.08 0.22 10
Booló – irrigation canal 3°279050N 8°359280E 24.04.96 lv 1 p 4
9 Musola 3°269360N 8°379180E 24.04.96 2 lv 1 1 p 24.3 7.66 0.51 25
30.04.97 4 lv 1 1 p
Musola – dam 3°279140N 8°369150E 24.04.96 2 lv 1 6 p
Musola – mouth 3°289200N 8°359000E 17.07.96 lv
10 Ebá 3°269460N 8°379200E 13.05.89 lv 10
11.11.89 lv
30.04.97 7 lv 1 2 p 22.5 n.r. 0.65
11 Bisoco 3°289250N 8°359050E 24.04.96 3 lv 8
10.05.96 lv 24 7.35 0.22
12 Lada 3°309170N 8°369520E 13.05.89 lv 9
13.04.96 11 lv
10.05.96 5 lv 24.2 7.30 0.52
13 Matogi/Edda tributary 3°309310N 8°379040E 13.05.89 lv 10
13.04.96 8 lv 1 1 p
10.05.96 2 lv 1 2 p 23.5 7.62 0.70
17.07.96 lv
14 Edda 3°309420N 8°369560E 13.04.96 2 lv 25
10.05.96 lv 25 6.13 0.11
15 Maloho 3°319390N 8°359560E 10.04.96 lv 1 p 95
08.05.96 lv 1 p 25.4 7.20 0.88
16 Puputu 3°349190N 8°379070E 13.04.96 2 lv 8
08.05.96 4 lv 1 2 p 24.8 6.81 0.32
17 Rupe 3°359180N 8°369560E 12.05.89 lv 24
13.04.96 15 lv 1 2 p
08.05.96 6 lv 24.5 7.06 0.47
16.07.96 lv 1 p
Rupe – upstream 3°359180N 8°379210E 20.04.96 39 lv 1 12 p
18 Moramope 3°359240N 8°369550E 13.04.96 1 lv 28
08.05.96 24.8 7.28 0.31
19 Ope 3°379180N 8°379250E 12.04.96 10 lv 1 3 p 23
08.05.96 lv 1 p 23.9 6.94 1.05
20 Ope tributary 3°359180N 8°369560E 12.04.96 3 lv 5
08.05.96 24.6 6.06 0.19
21 Biala 3°379380N 8°379430E 18.05.89 lv 20
08.11.89 lv
12.04.96 1 lv 1 1 p
08.05.96 lv 23.8 7.09 1.24
16.07.96 lv 1 p
Biala – upstream 3°379380N 8°389360E 20.04.96 3 lv

© 1998 Blackwell Science Ltd, Medical and Veterinary Entomology 12, 267–275
 Simulium damnosum s.l. on Bioko 271

Table 1. Continued.

Temp. Discharge Length*

Map No. River Grid Ref. Date Material Collected (°C) pH (m3 sec–1) (km)

22 Apu 3°409530N 8°399090E 18.05.89 lv 30

08.11.89 lv
16.11.89 lv
12.04.96 25 lv 1 23 p
08.05.96 3 lv 23.5 7.37 1.06
16.07.96 lv 1 p
23 Tiburones 3°439170N 8°429110E 16.05.89 lv 40
12.04.96 22 lv 1 4 p
08.05.96 lv 23.2 7.54 1.15
16.07.96 lv
24 Biolo-Timbabé 3°439180N 8°459150E 08.04.96 2 lv 50
22.07.96 lv 1 p
Biolo/Timbabé – upstream 3°419520N 8°459280E 08.05.96 6 lv 24.3 7.18 0.25
25 Togecha 3°449410N 8°489420E 17.05.89 lv 12
i Ko (Borabuope?) 3°489N 8°529E 18.11.89 lv 8
ii unnamed stream ‘a’ 3°439N 8°449E 15.11.89 lv 10
iii unnamed stream ‘b’ 3°439N 8°449E 15.11.89 lv ?
iv unnamed stream ‘c’ 3°439N 8°449E 15.11.89 lv ?
v Bosao 3°319N 8°399E 15.11.89 lv 17
vi Bioco 3°369N 8°379E 09.11.89 lv 5
vii (16?) unnamed (Puputu?) 3°339N 8°379E 09.11.89 lv 8
viii (18?) unnamed (Moramope?) 3°349N 8°379E 11.11.89 lv 28
ix Maloho tributary (?) 3°329N 8°379E 11.11.89 lv 8
x Sabe 3°289N 8°379E 11.11.89 lv 12
xi Ria-ria (?) 3°339N 8°509E 14.11.89 lv 7
xii Toho (?) 3°349N 8°519E 14.11.89 lv 8

the Bioko form of the S. damnosum complex at two sites.
Larvae and pupae were found in the Iladyi (river 4, Fig. 1) at
only 50 m from the sea, and in the Musola (river 9, Fig. 1) at
c. 750 m from the sea. Aquatic stages of the vector were found
in a range of breeding site conditions, occurring in rivers with
temperatures from 22.5°C (Ebá, river 10) to 26.2°C (Muedede,
3), pH from 6.06 (Ope tributary, 20) to 8.08 (Booló, 8), and
in a wide range of river types (Fig. 2). Although many of the
rivers found with aquatic stages conformed to typical ‘forest-
type’ habitat, as exemplified by the Tiburones (Fig. 2a,b),
larvae were also found in rivers ranging from large shallow
rivers, such as the Ole in the south (Fig. 2c), to small streams
like the Lada (Fig. 2d), and in man-made sites such as the
irrigation canal deriving from the R. Booló (no. 8, Table 1).
Inaccessible rivers
found in two of these rivers (Table 1), the R. Leke (5) and the
R. Ole (6). Listed anti-clockwise from Punta Oscura (Fig. 1),
and marked with numerals in boxes the others are:
Punta Oscura to Ureca (map nos. 23–12): Tope Tomu
(3°1594099 N 8°2894599 E), Bacha (3°1593199 N 8°3092099 E),
Djique (3°1591999 N 8°3191499 E), Motalelo (3°1592299 N
8°3095899 E), Maila (Sangre) (3°1591299 N 8°3192799 E),
Zio/Sodje (3°1590899 N 8°3194099 E), Mosejo (3°1495899 N
8°3195999 E), Lobibo (3°1494899 N 8°3292899 E), Vatalalo
(3°1493799 N 8°3295699 E), Poto (3°1493499 N 8°3193499 E),
Osa (3°1494599 N 8°3490599 E), Rute/Mbaali/Salabe (3°1494099
N 8°3592299 E);
Ureca to ‘point B’ (map nos. 11–7): Olabaita (3°1494099 N
8°3592799 E), Baca (3°1490099 N 8°3692099 E), Mococobe
(3°1490799 N 8°3695099 E), Moaba (3°1490099 N 8°3792599 E),
Loara (3°1490899 N 8°3793099 E).

Seventeen of the 130 rivers flowing were inaccessible, but
considered highly likely to be breeding sites. These rivers were
all in the south of the island, flowing into the sea on the
beaches between the Rı́o Moaba (point B, Fig. 1) and Punta
Oscura. Little is known of these rivers and some are not
marked on the available maps of Bioko or have names known
only to the local inhabitants in Ureca. Many are large, and
flow over cliffs (µ 10 m high) onto the beach, or have
deepwater mouths which are inaccessible for long stretches
upstream. Aquatic stages of the S. damnosum complex were
Unprospected rivers
A total of twenty-one (9%) rivers were not visited because
they are difficult to reach, with few existing paths or routes
available, and the difficulty of access is increased by cliffs
(lower reaches of the rivers of the south and south-west) or
mountainous terrain (upper reaches of the same rivers). All are
in the southern part of the island, and are shown in Fig. 1
marked with numerals in boxes. They can be conveniently
classed in three groups:

© 1998 Blackwell Science Ltd, Medical and Veterinary Entomology 12, 267–275
272 P. J. McCall et al.

Fig. 2. Examples of the range of breeding sites of the Bioko form of the S. damnosum complex. R. Tiburones (23), typical of the majority of
breeding sites seen in the northern part of the island, at two locations: (a) roadside at an altitude of 90 m, (b) upstream at altitude of 400 m. (c)
An example of the large Southern rivers, the R. Ole (6) close to its mouth on the south coast, and (d) an example of the smaller streams in which
the Bioko form of S. damnosum also breeds, R. Lada (12) at an altitude of 50 m.

1. Riaba to ‘Point A’. These nine rivers occur in the south- to dry out during the December–January period (R. Castelo,
eastern coastal region, between Riaba and the R. Obába (Point personal communication). In the southern part of this coast,
A, Fig. 1). Although access to the northern rivers in this section between the R. Iladyi (no. 4, Fig. 1) and Punta Santiago, a
is possible on foot, the rivers were not visited as all are known number of small rivers and streams exist, but are of a size

© 1998 Blackwell Science Ltd, Medical and Veterinary Entomology 12, 267–275

suggesting they are unlikely to be perennial, and therefore
unlikely to support breeding of the S. damnosum complex
during the dry season.
2. Punta Santiago to ‘point B’ (rivers 1–6). These six rivers
flow to the south coast between Punta Santiago and point B
(Fig. 1). No information exists regarding their condition during
the dry season. The rivers are the Nkari/Sodyo (3°1194299 N
8°4091599 E), Maama (3°1390599 N 8°3995899 E), Lede
(3°1391199 N 8°3994199 E), Biadyi (3°1393599 N 8°3895099 E),
Mombebaca (3°1393899 N 8°3894299 E), unnamed (3°1395299
N 8°3795899 E). All rivers are of a nature which suggest they
are likely to support breeding of the S. damnosum complex.
3. Punta Oscura to Rı́o Boó (‘Point C’) (map nos. 24–29).
These six rivers flow to the west coast between Punta Oscura
and the Rı́o Boó (river C, Fig. 1). No information exists
regarding their condition during the dry season. The rivers are
the Eburi (3°1892599 N 8°2691099 E), Baña (3°1991499 N
8°2594599 E), unnamed (3°2090299 N 8°2592099 E), unnamed
(3°2190999 N 8°2591099 E), Bisambu (3°2290699 N 8°2591299
E) and Etepo (3°2294899 N 8°2591299 E). All were considered
likely to support the S. damnosum complex.
The total number of rivers where aquatic stages of the
S. damnosum complex were found was twenty-five. The total
number of inaccessible rivers which were considered likely to
support breeding was seventeen, and the total number of
unprospected rivers which are of a nature suggesting they
could also support the vector was twelve. The maximum
number of rivers therefore, which might support vector breeding
and require treatment in a vector elimination programme is
fifty-four (21.9% of all rivers on Bioko).
Altitude limit of breeding
Most river prospections in the northern part of Bioko were
made with access from the main road which circles this part
of the island below 100 m, occasionally rising to 450 m on the
east coast and to a maximum of 750 m on the pass between
Riaba and Luba. Extra prospections in this region were made
at higher altitudes in a number of selected rivers where the
S. damnosum complex had been found further downstream.
The rivers were selected on the basis of their relatively large
size and accessibility.
At 250 m, larvae were found in the Biala (river 21, 20 April
1996) but none were found in the Edda (14) or the Maloho
(15) at 200 m (29 April 1997). At 300 m, no larvae were found
in the Ruma (1) or its two tributaries on 23 April 1996 despite
the apparent suitability of these sites. A series of prospections
of the Tiburones (23) on 23–24 April 1997 found that larvae
were present at all points up to an altitude of 400 m, but it
was not possible to sample higher up. In contrast, the Ope (19)
was dry (24 April 1997) at the same altitude. The Musola (9)
was prospected at numerous points along its length: larvae
were found up to 400 m (24 April 1997), but no larvae were
found at 600 m (30 April 1997). The Musola tributary, the Ebá
(10), had larvae at 300 m (13 May 1989 and 30 April 1997)
but not at 550 m (24 April 1997). No larvae were found at
altitudes above 400 m. Temperature may not have been the
© 1998 Blackwell Science Ltd, Medical and Veterinary Entomology 12, 267–275
Simulium damnosum s.l. on Bioko 273
limiting factor at these altitudes as, for example, larvae were
found in the Ebá at 22.5°C (300 m), but not in the Musola at
22°C (600 m). In most cases, because of the low discharges
during the dry season, the rivers were flowing as narrow and
shallow streams even at relatively low altitudes, and so were
unsuitable for the S. damnosum complex. It was clear that few
rivers would provide suitable vector breeding sites at altitudes
above 500 m. The Grande (2), Iladyi (4), Musola (9), Maloho
(15), Tiburones (23), and probably some of the unprospected
rivers in the south, were large enough to support breeding
above this altitude, although in these cases temperature could
be the limiting factor at higher altitudes. Temperatures in the
Iladyi at 1100 m (3°1991899 N 8°4091299 E) ranged from 18.5
(12 May 1996) to 19.5°C (1 May 1997). Few members of the
S. damnosum complex are known to occur at these low
temperatures (see Discussion below). It seems likely therefore
that there is a maximum altitudinal limit to breeding in even
the largest rivers at some point below 1000 m.
Lengths of rivers supporting breeding during the dry season
Based on the data presented, and assuming that the largest
rivers could support breeding up to 1000 m, the extent of
breeding of the S. damnosum complex on Bioko was estimated.
The total length of rivers which are known to support this
vector during the dry season is 591 km (Table 1). The total
length of inaccessible rivers which were thought likely to
support development (Fig. 1, boxed numbers 7–23) is 256 km,
whereas the total length of unprospected rivers also thought to
be likely to harbour the vector (Fig. 1, boxed numbers 1–6 and
24–28) is 173 km. The total length of the stretches of rivers
which might support breeding of the S. damnosum complex
is 1020 km.
Discussion
The major prerequisites listed by Walsh (1990a) for successful
blackfly eradication are isolation of the vector population and
accessibility of its breeding sites for larviciding. A number of
discrete foci of onchocerciasis have previously been eradicated
successfully by vector control: S. neavei in Kenya (Garnham
& McMahon, 1954; McMahon et al., 1958); S. neavei Roubaud
from one focus (Budongo forest) and a member of the
S. damnosum complex from two other foci (Victoria Nile and
Ruwenzori) in Uganda (Prentice, 1974; Walsh, 1990a; Davies,
1994); the Djodji form of S. sanctipauli V. & D. from parts of
Ghana and Togo by the UNDP/World Bank/WHO
Onchocerciasis Control Programme (OCP) (Walsh, 1990b).
Bioko is the only island with autochthonous onchocerciasis
transmission, and as such represents a uniquely isolated focus
of the disease, potentially vulnerable to eradication. Taxonomic
studies have now demonstrated that the Bioko form is unique
to the island (Post et al., unpubl. data), remote from mainland
populations of other sibling species and forms of the
S. damnosum complex. The possibility of reinvasion following
a vector eradication programme on Bioko would therefore
274 P. J. McCall et al.
be minimal, satisfying Walsh’s first requirement in the case for
vector eradication on Bioko. The surveys reported here have
determined the extent of breeding on the island, and the
accessibility of the breeding sites to treatment; Walsh’s second
requirement for the planning of an eradication programme.
Although Bioko has 247 rivers and streams, only fifty-four
of these are capable of supporting vector breeding during the
dry season. Assuming that all of these fifty-four rivers do
support vector breeding and would require treatment, the
prospect of controlling so many rivers might initially appear
to be a formidable task. This may not be the case. A maximum
distance of only 1020 km of river would need to be treated
with insecticide to destroy all larvae of the vector. Achieving
vector eradication on Bioko may require a significantly lower
degree of technological input and logistic support than that
required to achieve similar results in mainland West Africa by
the OCP, for a number of reasons.
Firstly, during the dry season most of the positive rivers
have relatively low discharges, ranging from 0.11 (Edda) to
4.46 (Iladyi) m3 sec–1, although an exceptionally high rate of
8.18 m3 sec–1 was recorded at the mouth of the R. Moaba – a
probable breeding site. Data for the rivers Ruma and Musola
(Castelo, 1994) show that monthly discharges are at their
lowest during the period from December to June.
Secondly, the water temperatures at the breeding sites on
Bioko are relatively low for the S. damnosum complex, ranging
from 22.5 to 26.2°C during the dry season (Table 1). Larval
development could be expected to require a minimum of about
12 days at 25–26°C (Davis et al., 1992) to 16 or more days at
temperatures less than 24°C (Elsen, 1979). Insecticide treatment
could therefore be applied only fortnightly, instead of weekly,
significantly reducing the cost of treatment and increasing the
chances of effectively reaching all breeding sites at each
treatment cycle.
Thirdly, the steep togography of the island results in shorter
river stretches suitable for vector breeding as, above 500 m,
discharges in smaller rivers become too low for S. damnosum
s.l. Low temperatures at higher altitudes limit S. damnosum
s.l. breeding to stretches below 1000 m in the largest rivers.
Grunewald (1976) found breeding of the ‘Sanje’ form of the
S. damnosum complex at water temperatures down to
a minimum of 16.8°C, but recorded higher minima of 22.7
and 23.4°C for S. yahense and S. squamosum, respectively.
These latter species are the most closely related to the Bioko
form, for which a minimum temperature of 22.5°C was recorded
in this study. The temperature above 1000 m on Bioko (18.5–
19.5°C recorded in the Iladyi at 1100 m in the present study)
would prohibit breeding of all S. damnosum complex species,
with the exception of the East African ‘Sanje’, ‘Nkusi’ and
‘Nyamagasani’ forms, which are found at temperatures between
16.8 and 22.9°C (Grunewald, 1976). As a consequence of
these altitude restrictions, the longest of the fifty-four rivers
which would require treatment is the Maloho, where up to
95 km could support vector breeding. The other large rivers
with long stretches requiring treatment are the rivers Moaba
(75 km), Tiburones (40 km) and Biolo/Timbabé (50 km). In
the remaining major rivers, including those with high discharges
© 1998 Blackwell Science Ltd, Medical and Veterinary Entomology 12, 267–275
(1.5–5.0 m3 sec–1; Ole, Iladyi and Grande), the stretches
requiring treatment are less than 30 km each.
Finally, the majority of the rivers requiring treatment are
grouped extremely closely together in two discrete areas.
Between Malabo and Luba, sixteen positive rivers (circles 8–
23, Fig. 1) occur in a 35 km flight path, with some rivers only
500 m (Rupe and Moramope, 17 and 18) or 1 km (Ope, Ope
tributary and Biala, 19, 20 and 21) apart. Two of these rivers,
the Biolo/Timbabé (24) and the Tiburones (23) are within 5 km
of the airport, with the lower reaches of the latter passing
within 1500 m of the airport runway. In the southern part of
the island, thirty-three definite or probable positive rivers occur
in a coastal flight path of less than 50 km (boxes 1–29, circles
3–6, Fig. 1), twenty-five of these in a straight line of less than
25 km, between Punta Oscura and Punta Santiago.
The ideal period for larvicide treatment on Bioko would be
during the first 3–4 months of the year, from January to April.
Comparison of both 1930s and 1990s rainfall data (unpublished
data, ASECNA, Malabo) showed that the distribution pattern
has been generally stable, with minimum precipitation occurring
during the period from December to May. River discharge
rates are also at their lowest during this period, optimizing
insecticide usage (Castelo, 1994). The levels of vector biting
are lowest during January to April (unpublished data),
coinciding with the dry season reduction in suitable breeding
sites to the maximum of fifty-four rivers described above. At
this time the crucial parameters of hydrology and abundance
of the vector population are at their most favourable to enable
a cost-effective eradication campaign. The inaccessibility by
road of many of the rivers, especially those in the southern
part of Bioko, precludes the possibility of eradication measures
being conducted from the ground and aerial application would
be essential. For the aerial applications a helicopter would be
needed, as the topography of the island is unsuitable for
treatment with fixed-wing aircraft. There are a few sites which
will, however, remain inaccessible to treatment by helicopter
because of dense canopy. These could be reached and easily
treated by a ground-based operation.
It is unlikely that the unique ecosystem of this island would
be greatly affected by such a programme. In the dry season,
seven other species of Simuliidae have been recorded on Bioko,
from many rivers in addition to those with the S. damnosum
complex (McCall & Cheke, unpubl. data). The short duration
of required treatment with a suitable insecticide means that
the environmental and non-target impact would be minimal.
Although the use of temephos in mainland Africa by the OCP
is believed to have caused little or no long-term effect to the
fauna or human population/food chain over a 20-year period
(Lévêque et al., 1988), an assessment of the environmental
impact resulting from such treatments on Bioko would be a
pre-requisite of any such control programme.
In conclusion, the results of this study suggest that a control
campaign using larvicides against the vectors of onchocerciasis
on the island of Bioko is feasible. If treatments were conducted
during the dry season (December–March), a maximum of fifty-
four rivers would require treatment and could be dealt with
effectively by a helicopter-based programme supplemented by
ground-based delivery. A vector control programme could

conceivably achieve vector eradication in 1 or 2 years,
with minimal (if any) risk of reinvasion of vectors from
elsewhere, and could provide a quick and cost-effective solution
to the onchocerciasis problem on Bioko.
Acknowledgements
We are grateful to the staff of the Programa Control
Onchocercosis, in particular Sres. Enrique Mba Oudo, Francisco
Sidibola, Rufino Nguema Andeme and Sra Maria del Mar
Ardevol. We are also grateful to the Ministerio de Sanidad and
Ministerio de Medio Ambiente, Malabo, Guinea Ecuatorial,
for their support. We thank the Servicio de Meteorologı́a/
ASECNA Malabo for providing meteorological data. This
investigation received financial support from the UNDP/World
Bank/WHO Special Programme for Research and Training
in Tropical Diseases (WHO/TDR Research Task Force on
Onchocerciasis Operational Research, ref. no. 151/181/34).
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Accepted 1 March 1998