Vol.1, No.1, 1-7 (2013) Advances in Entomology
Lack of evidence for sufficiently isolated populations
of Glossina morsitans submorsitans on the
Adamawa Plateau of Cameroon following geometric
morphometric analysis
Mbunkah Daniel Achukwi1*, Jessica Gillingwater2, Alexandre Michel Njan Nloga3,
Gustave Simo4
1Institute of Agricultural Research for Development (IRAD), Wakwa Regional Center, Veterinary Research Laboratory, Ngaoundere,
Cameroun; *Corresponding Author: achukwi_md@yahoo.co.uk
2London School of Hygiene and Tropical Medicine, London, UK
3School of Medicine and Veterinary Sciences, University of Ngaoundere, Ngaoundere, Cameroun
4Department of Biochemistry, Faculty of Science, University of Dschang, Dschang, Cameroon
Received 24 April 2013; revised 21 June 2013; accepted 8 July 2013
Copyright © 2013 Mbunkah Daniel Achukwi et al. This is an open access article distributed under the Creative Commons Attribution
License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Trypanosomosis remains the number one killer
of livestock in spite of efforts made to eradicate
t set se flies in the Adamawa plate au of Cameroon.
Acetone-baited Laveissière type biconical traps
were pitched at 100 meter intervals in strategic
geo-referenced positions in various ecological
niches of the landscape for 5 consecutive days
in selected villages in Mayo Rey, Mbere, Vina
and Faro et Deo divisions. All 493 tsetse flies
captured in sites other than Mboula were G.
morsitans submositans. Measurement of dif-
ferent morphometric characters on the wings of
each individual fly was undertaken using the Du
Jardin package. The data was processed and
analysed by “Permutaciones, Analisis Discrimi-
nante (PAD)” and “Bootstraps, Analisis en Com-
ponentes principales”. The three major sam-
pling sites on the plateau yielded similar results
as demonstrated by the neighbour joining tree
of Mahalanobis distances but tests using PAD
showed th e differen ces bet ween group mea ns to
be significant (P < 0.05) even when the same
number of flies was used. Mixing of tsetse
populations from the northern lowlands and
those on the plate au and K outine plain c ould not
be ruled out . These prel i mina ry f indin gs sug ge st
that the flies are not from isolated populations
and should be considered as populations fre-
quently exchanging migrants. However, mo-
lecular genetics techniques are necessary in ad-
dition to morphometric analysis to reach more
definitive conclusions.
Keywords: Glossina morsitans submositans;
Fly Wing; Geometric Morphometry;
Adamawa Plateau Cameroon
Tsetse flies (Diptera: Glossinidae) are vectors of try-
panosomes that cause sleeping sickness in humans and
nagana in livestock across sub-Saharan Africa. Try-
panosomosis has a major negative impact on agriculture,
especially on livestock economics, in areas such as the
Adamawa Plateau region of Cameroon where livestock
breeding is the mainstay of the economy. In the south-
west of neighbouring Nigeria sedentary and semi seden-
tary cattle herds had trypanosomosis prevalence rates of
9.8% and 42.8% respectively and more mechanical vec-
tors were caught than Glossina spps: the G. papalis and
G. tachinoides caught did not carry any trypanosome
infections [1]. In Zambia, 75.1% of G. m. morsitans
blood meals were from cattle, even when other domestic
animals (mainly goats, pigs and dogs) were present [2].
Thus tsetse control strategies rely on a detailed under-
standing of the epidemiology and ecology of tsetse flies
and the genetic variation within and among the fly popu-
In sub-Saharan Africa, conventional disease control
methods applied have inadequately addressed the prob-
lem of trypanosomosis and large scale control operations
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M. D. Achukwi et al. / Advances in Entomology 1 (2013) 1-7
have virtually stopped. Large scale tsetse eradication
campaigns instituted earlier in some countries including
Cameroon required huge budgetary allocations that most
affected countries could not afford on a sustainable basis.
Even innovative options like the use of live bait pyre-
throid-impregnated cattle against tsetse flies have had
problems with efficiency, time input an d the high cost of
pyrethroid in rural areas. After about thirty years of gov-
ernment efforts to control tsetse on the Cameroon Ada-
mawa Plateau, trypanosomosis still stands out from
various surveys [3-5], as the number one killer of cattle.
Large scale tsetse clearing activities were stopped in this
region in the mid-nineties [6,7]. These tsetse control ef-
forts have had mixed success records; ultimately, they
have been unable to prevent re-infestation of tsetse
cleared areas [8]. Consequently, there is a need for new
sustainable technical approaches that are adapted to ex-
isting local farming systems, more efficient and inexpen-
A pan African approach called “Pan African Tsetse
and Trypanosomosis Eradication Campaign” (PATTEC)
has been initiated under the support of African heads of
states. The objective of this initiative is to eliminate
tsetse flies and trypanosomosis in Africa. A critical ex-
amination of alternative options to currently employed
control strategies which include the use of screens, tsetse
traps and area-wide techniques to exterminate pockets of
tsetse infestations is urgently needed [9]. The success of
these control strategies requires the identification of ge-
netically isolated tsetse populations where elimination
can be undertaken. The identification of genetically iso-
lated tsetse populations is the critical point that will di-
rect the choice between elimination and suppression of
tsetse flies in a specific area.
Over the past 30 years, several techniques have been
used to assess genetic variation of tsetse flies to identify
genetically isolated populations. These techniques in-
clude allozymes [10-13], microsatellites [14-16], and
mitochondrial DNA techniques [12,15,17-19]. These
genetic markers have provided useful information on
phenotypic and genetic polymorphisms in Glossina spe-
cies. They have indicated a degree of genetic differentia-
tion between geographically separated tsetse populations.
This was surprising given the mobility of the flies com-
bined with expectations derived from population genetics
theory [20-22]. The analysis of mitochondrial DNA re-
vealed relatively high levels of haplotype divergence
between randomly chosen individuals among four spe-
cies analysed, including G. morsitans [20]. However,
arthropod mitochondrial genomes are small in size (14 -
25 kilobases) and maternally inherited [23], hence, data
interpretation differs from that of variation in nuclear
The number of tsetse loci in the nuclear genome using
the techniques sampled for variation to date is relatively
limited for microsatellite markers. For example, in
Glossina morsitans morsitans a total of 45 allozymes and
five microsatellites were analysed by Krafsur [20]. Mor-
phometric analyses have become interesting tools for the
study of population genetics [24,25]. The first characters
to be affected by micro-evolutionary changes are con-
tinuous characters. Morphometrics measure these con-
tinuous changes and represent a low-cost tool adapted to
the study of phenotypic evolution. This tool has shown
considerable epidemiological significance for medically
important insects such as the vectors of leishmaniasis
and Chagas disease in Latin America. Geometric mor-
phometrics allow comparisons of many samples with
homologous landmarks, e.g., the vein-patterns on wings.
Morphometric analyses are generally simple to perform
and easily realizable in most developing countries where
trypanosomosis has a major impact on animals and/or
humans. This tool can be used to determine how close or
distant tsetse fly populations thought to be isolated truly
are by normalising other factors such as size and rotation
of the wings from the different populations. This tech-
nique allows a more sensitive appraisal of the relation-
ship between groups than genetic techniques involving
microsatellites [26,27]. Therefore, the technique can be
used to reveal the structure of vector populations. If a
vector population is found to be sufficiently geographi-
cally isolated to prevent casual mixing with other popu-
lations, then it may be of interest for future vector control
efforts. The present study specifically sought to compare
Glossina morsitans submorsitans populations of different
sites of the Adamawa region using morphometry of
2.1. Description of the Geographic Location
The elevations where sampling was undertaken ranged
from 487 m above sea level in the Koutine plain to over
1497 m on the plateau (Table 1). The Adamawa region
has wooded savannah vegetation, a seven months rainy
season with moderate temperatures (22˚C - 28˚C) and the
average annual rainfall ranges from 1500 mm to 1900
mm. More details about these aspects of the study sites
have been described [28].
2.2. Entomological Approach
This study was con duc ted in fo ur d iv isions (Mayo Rey,
Mbere, Vina, and Faro et Deo) of the Adamawa Plateau
of Cameroon (Figure 1). The sites comprised: one vil-
lage (Gamba/Benoue National Park) of the Mayo Rey
Division, one village (Mboula) of the Mbere Division,
two villages (Mangoli, Mbe) of the Vina Division and
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M. D. Achukwi et al. / Advances in Entomology 1 (2013) 1-7
Copyright © 2013 SciRes. http://www.scirp.org/journal/ae/
Table 1. Geo-referenced positions of main villages and number of traps pitched in each village.
Village Geo-referenced positions Height above sea level Number of traps
Mboula N06˚ 69,366
E013˚ 98,333 932m 11
Mangoli N07˚ 02,841
E014˚ 00,819 1497m 25
Guemnfiti N07˚ 67,680
E01˚ 18,986 511m 21
Kontcha N07˚ 92,583
E012˚ 28,640 487m 25
Benoue National Park N08˚ 12,833
E13˚ 39,529 538m 18
Figure 1. Map of Adamawa region illustrating geo-referenced main positions of bi-conical traps pitched for tsetse fly catching.
The intensity of darkness of the trap reflects the abundance (is proportional to the number) of traps placed in the locality.
Openly accessible at
M. D. Achukwi et al. / Advances in Entomology 1 (2013) 1-7
two villages (Kontcha, Nguemfiti) of the Faro et Deo
Division. Tsetse fly acetone-baited biconical traps [29]
were pitched in tsetse fly favourable biotopes in each
village. The herdsmen guid ed the research team in locat-
ing tsetse fly biotopes. Geographical coordinates (the
main sites in Table 1) for each trap were recorded using
a GPS set. Traps were spaced at 100 meter intervals.
Tsetse flies were collected twice per day for five con-
secutive days. The relative G. m. submorsitans popula-
tion densities (flies/trap/day) for Nguemfiti, Kontcha and
Gamba/Benoue National Park trap sites were also deter-
mined. Henceforth the site Gamba/Benoue National Park
will simply be referred to as the Benoue National Park
because they are too close to each other. All flies from
each trap were counted, sorted according to sex and spe-
cies, and were prepared for wing morphometry. With
regard to the latter process, the wings were removed,
transferred to a glass slide, and covered with a cover slip.
The cover slip was sello-tapped onto the glass slide.
Slides were labeled with the sex, fly species and trap site
co-ordinates. The mounted wing preparations were sub-
sequently scanned into a co mputer (Figure 2(a)).
2.3. Morphometrics Analysis
The size of the wing is known to be proportio nal to the
Figure 2. (a) Above is a scanned image of the dissected
wing of a G. morsitans sub morsitans collected from one
of the three sampling sites. Below the same image with
yellow dots indicating the chosen land-marks for mor-
phometric analysis; (b) Joining tree of Mahalanobis dis-
tances. This shows that the fly populations at sites one
and two (Kontcha and Nguemfiti) are more closely re-
lated to one another than to the population from site 3
(Benoue National Park).
size of the individual insect [30]. Here, the features of
“COO” (collection of coordinates) software version 37 of
Du Jardin package; (http://www.mpl.ird.fr/morphometrics/bac/)
were used. This software allows the measurement of dif-
ferent morphometric characters on the wings of each
individual. Briefly, digital images of each wing were
obtained and the COO software was used to record 9
Cartesian coordinates (homologous points on the wing
referred to as landmarks where veins intersect or end
(Figure 3(a)) on each wing. T he x and y co ordinates data
were then formatted using TET (“Tabla Espacios Tabu-
laciones”) software (version TET 0 45) to allow proc-
essing with the MOG (“Morfometria Geometrica”) soft-
ware (version MOG 0 79) which aligns the scanned im-
ages and allows comparison of their size and shape pro-
ducing both partial and relative warps. These were then
further examined using PAD (“Permutaciones, Analisis
Discriminante”) and BAC (“Bootstraps, Analisis en
Componentes Principales”) software. Principal compo-
nents analyses were performed on the covariance matrix
(total sample) or the consensus covariance matrix (sub-
divided sample) of either raw data or size free variables.
Statistical significance of the test was estimated accord-
ing to the angles between first principal components and
associated permutation tests. All of the software used for
our analyses is contained in Du Jardin package at
Only the three largest groups of samples were exam-
ined since morphometric analysis requires at least twice
the number of samples as there are landmarks; thus the
smaller collections were ruled out. The biggest differ-
ences in wing shape were between the samples collected
in the Northern lowlands (Benoue National Park) and
those collected from the other two sites. The fly catch in
Mbe was very scanty and no flies were caught near
Mangoli village. A total of 495 Glossina sps were caught
and identified. Of these, 493 were of the G. morsitans
sub morsitans group trapped at the three main sites men-
tioned above. The remaining two flies caught in Mboula
were not included in the analysis. The highest catch was
in the neighbourhood of Nguemfiti village while the
lowest catch was in the jungle ne ighbourhood of Mboula.
Trapped flies were mainly G. morsitans submorsitans.
The relative densities o f G. m. submorsitans for the three
main trap sites (Nguemfiti, Kontcha and Benoue Na-
tional Park) were estimated at 9.6, 7.4 and 3.7 flies/
trap/day, respectively. These densities were superior to
the index of apparent abundance reported by Mamoudou
et al. [31] for other villages in the same region where the
present study was undertaken. The differences may sim-
ply be due to differences in biotopes as our study sites
were more remote and inaccessible than those of Ma-
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M. D. Achukwi et al. / Advances in Entomology 1 (2013) 1-7 5
Figure 3. (a) Results of a principle component analysis run
in BAC. The three sampled populations show overlap be-
tween the data. The numbers in the key refer to the sample
sizes which were 96 for Kontcha (yello), 126 for Nguemfenti
(green), and 163 for Gamba (red) at the Benoue National
Park. This diagram also clearly shows the more dissimilar
northern (Gamba) population overlapping least with the
other two; (b) Principle component analysis run in BAC.
This time the sample sizes are equal to ensure that it is not
the difference in sample size causing the observed differ-
ences between the populations. The first 96 samples in each
of three groups Kontcha (K), Nguemfeti (N) and Gamba (G)
were arbitrarily selected for this analysis.
moudou et al. [31]. During a longitudinal study in the
Adamawa region it was found that the incidence of try-
panosomosis was high in the valley (3.7% - 20%) and the
buffer zone (1.8% - 13.4%), and was significantly lower
(0% - 2.1%) on the plateau where tsetse clearing had
previously been undertaken [31]. They also caught main-
ly Glossina morsitans submorsitans and a few G. tach in-
oides in the valley and the buffer zone, but none on the
plateau and concluded that the distribution of tsetse, in a
large part of the valley, undergoes substantial seasonal
changes depending on either the presence or absence of
cattle. Other studies undertaken to the north of Kontcha
(also one of the sites in the present study) in villages
around Poli near the Faro National Park [3] showed a
parasite detectable trypanosomosis prevalence rate of
14.3% in cattle. A consequence of the negative impact of
the trypanosomosis in the Adamawa and northern re-
gions of Cameroon is that huge quantities of trypano-
cides are sold to farmers by vet pharmacies in these re-
gions [32].
Results of the joining tree of Mahalanobis distances
(Figure 2(b)) suggest a subdivision of G. m. submorsis-
tans on the Adamawa Plateau according to the sampling
sites. Looking at the permutation analysis, it appears that
the two major sampling sites on the plateau yielded much
more similar results as demonstrated by the neighbour
joining tree of Mahalanobis distances (Figure 2(b)).
These results were confirmed by a permutation test using
PAD. The PAD showed significant (P < 0.05) differ-
ences (Figure 3(a)) between tsetse populations from
different capture sites. To ensure that the observed dif-
ferences between the tsetse populations were not due to
differences in sample sizes alone, the same analysis was
performed using equal sample sizes for all three groups
(Figure 3(b)); the same result was obtained. Conse-
quently, morphometric analysis can be used to different-
tiate Glossina m. submorsistans from the Adamawa Pla-
teau of Cameroon and this technique can reveal differ-
ences between vectors of medical and veterinary impor-
tance. The geometric morphometrics technique allows a
more sensitive appraisal of the relationships between
groups than genetic techniques involving microsatellites
[26,33]. Geometric morphometrics techniques are useful
in evaluating the micro-evolutionary changes occurring
during phenotypic evolution in tsetse flies, especially in
developing countries where laboratories to perform ge-
netic analyses are lacking. These techniques are valuable
to the Pan African Tsetse and Trypanosomiasis Eradica-
tion Campaign which envisages eradicatio n of tsetse flies
from the African continent. Most countries infested by
tsetse flies are poor and have few laboratories capable of
genetic analyses. Several tsetse fly species are found in
some tsetse infested areas. In such areas, the geometric
morphometrics technique will be of great value because
differences between tsetse species can be easily evalu-
ated. In environments with different tsetse species, spe-
cific markers for each species must be developed if ge-
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M. D. Achukwi et al. / Advances in Entomology 1 (2013) 1-7
netic analysis techniques are required. The use of genetic
techniques can be time consuming and expensive.
Despite the morphometric differences observed be-
tween tsetse populations from different sites, the permu-
tation analysis test showed some identical phonetic
characters for some flies sampled at different capture
sites (Figures 3(a) and (b)). These results indicate that
mixtures of tsetse fly populations may exist, the sam-
pling site notwithstanding. The presence of mixed tsetse
fly populations at each capture site can be explained by
the exchange of tsetse flies between different sampling
sites. The probability of a tsetse fly to fly fro m one sam-
ple site to another is low because of the distance between
sites. However, during transhumance, it is likely that
livestock transport a few tsetse flies between sites.
The flies in the sampled areas on the Cameroon
Adamawa Plateau, although different, are not from suffi-
ciently isolated populations to allow treatment for the
purposes of elimination. These flies should be considered
as populations which are able to exchange migrants, es-
pecially through the movement of livestock. The neces-
sity for geographic isolation for eradication makes is-
lands ideal targets for such strategies. This was demon-
strated in 1979 using SIT with G. austeni from Unguja
Island of Zan zibar [34]. Mainland vector populatio ns are
considerably harder to define and delineate, especially
where several species are present as in the Southern part
of the Adamawa Plateau where both G. morsitans sub-
mositans and G. tachinoides are found. However, artifi-
cial barriers such as screens and traps impregnated with
insecticides may be usefu l during initial contro l interven-
tions in mainland vector populations.
The present study shows that the natural reserve parks
are a safe haven for tsetse flies and are major sources for
acquisition of trypano somosis by domesticated livesto ck.
The predominant livestock is cattle in the Adamawa Pla-
teau, which move in and out of these reserves in search
of fodder. Villages like Nguemfiti, Kontcha and Gamba
(Benoue National Park) which are near national parks in
this region have very high tsetse fly densities. Future
studies could be directed towards DNA sequence analy-
ses of tsetse flies, estimating the vectorial capacity of the
flies and relating this index to trypanosome infection
rates of inhabitants, both humans and animals. The pre-
sent findings may be valuable in establishing practical
guidelines for tsetse fly control strategies in the region in
partnership with neighbouring countries.
This work received financial support from Leverhulme Trust and
material assistance from Vestergaard and the Institute of Agricultural
research for development (IRAD) Cameroon. The field work received
technical assistance from Dr. Bouyer Jeremy (CIRAD-CIRDES Burk-
ina Faso), the rural field staff of MINEPIA Cameroon headed by Mr.
Samuel Abba and Ministry of health for which we are grateful. We are
also grateful to Dr. Manchang TK, Mr. Mforpit Y. Mrs. Wachong-Kum
HNF and Faustin Samba, all of IRAD Wakwa, who provided valuable
logistical assistance during some of the fi eld trips.
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