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Research Article
Mavoungou et al., Vector Biol J 2017, 2:1
DOI: 10.4172/2473-4810.1000115
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Vector Biology Journal
International Publisher of Science,
Technology and Medicine
Breeding Sites of Stomoxys
spp (Diptera: Muscidae),
a Preliminary Study in the
Makokou Region (North-East-
Gabon)
Jacques François Mavoungou1,2*, Rodrigue Mintsa Nguema1,
Geneviève Lydie Acapovi3, Roland Zinga Koumba1, Franck
Mounioko2, Sevidzem silas Lendzele4, Ingrid Kindzi Bakakas2,
Jérémie Gilles5, Gérard Duvallet6, Bertrand M’Batchi2 and
Nicolas Picard7
Abstract
In order to circumscribe the breeding sites of stomoxes in the
Makokou region, eight potentially favorable environments were
sampled from January to June 2007 in three habitats: primary
and secondary forest (natural environments) and the anthropic
environment (articial or disturbed areas). In the primary forest,
three types of substrates were collected: litter, elephant dung
and Cephalophus droppings. In the secondary forest, two types
of substrates: litter and monkey droppings. In villages or cities,
three substrates constituting of: decaying plant material, organic
waste from garbage in villages and cow dung were also collected.
These materials were placed in emergence cages while awaiting
the emergence of adult arthropods. In total, 14517 arthropods
emerged, including 11726 stomoxes and 2791 other arthropod
specimens. Seven Stomoxys spp were observed and included: S.
transvittatus with 8221 (70%) emerged individuals, S. niger niger
with 1679 (14%) observed individuals. However, S. omega (8%); S.
inornatus (4%); S. calcitrans (2%); S. niger bilineatus (1.9%); and
nally S. xanthomelas (0.1%) emerged in minor proportions. The
results obtained in this study indicate that stomoxes can lay their
eggs on several types of substrates hence colonizing many areas.
Keywords
Stomoxes; Breeding media; Development; Abundance; Gabon
*Corresponding author: Jacques François Mavoungou, Institut de Recherche
en Ecologie Tropicale (IRET), BP 13354, Libreville, Gabon, Tel: +241 733089/35;
E-mail: mavoungoujacques@yahoo.fr
Received: April 28, 2016 Accepted: May 05, 2017 Published: May 15, 2017
Introduction
Stable ies (Stomoxys spp.) are insects that feed on blood and are
potential vectors of several pathogens [1,2]. Indeed, because of their
blood feeding habits, they are a plague due to their direct nuisance
(harassment and blood predation), but also by the role they play as
potential vectors of various pathogenic agents [3-8]. Moreover, stable
ies cause considerable economic losses in stock farms by their direct
pathogenic eects. In the USA, economic losses inicted by stable ies
on the livestock industry were evaluated to be between 100 and 400
million dollars [9,10]. Some species of stable ies are described as a
limiting factor to milk and meat production, they equally cause severe
irritation, blood loss and at times the death of animals in extreme
cases [11,12]. ese insects are also potential mechanical vectors of
pathogens of animal origin such as viruses, equine infectious anemia,
Trypanosoma evansi and Besnoitia besnoiti [2]. Stable ies in the
Afrotropical region remain poorly understood. Presently, there is
little information about the biology and ecology of this group, despite
the existence of reports in some areas [13-20]. In addition, majority of
available information is based on the cosmopolitan species Stomoxys
calcitrans and more recently from S. niger niger Macquart, 1851 [13].
However, stable ies can exist in multiple environments including
urban [21-23], rural areas near stables, slaughter houses, cattle
markets, dumped waste and occasionally in natural areas such as
forests and national parks [24]. Moreover, factors such as temperature,
precipitation, physical barriers, habitat and presence of vertebrate hosts
are likely to inuence their distribution, development and abundance;
and consequently the distribution of diseases they transmit to humans
and animals [25]. e study of these factors in natural environments
is very important in the ecology and epidemiology as they are spatial
risk factors for vector-borne diseases [26]. Knowing these factors will
guide in the implementation of vector control strategies and also help
to locate areas of high transmission potential [1,18,19,26]. In Gabon,
much work has already been grounded on the population genetics of
stable ies [27], origin of blood meal [24] and detection of pathogenic
agents transmitted by these insects (Mounioko [27],). Other studies
have revealed the presence of seven species of stable ies in various
ecological zones [16-19,24]. ese studies seek to know the role of
landscape structure on the ecodistribution of populations of stable
ies [16-19]. However, few studies have focused on breeding and the
identication of the various breeding sites of these insects. e study
of habitat of pre-imaginal forms can provide information about the
life cycle, population dynamics and the physico-chemical composition
of a given area [26]. According to the World Health Organization
[28], sustainable management of breeding sites of insect vectors is of
paramount importance in vector control and preservation of human
health. To collect data on breeding of stable ies, an entomological
survey was conducted in three dierent habitats represented by
primary/secondary forests and neighborhoods of the city of Makokou
in Gabon. e objective of this study was to identify and characterize
potential breeding sites of stomoxines in order to set up strategies for
their ght and subsequently the pathogens they transmit.
Materials and Methods
Location and description of study area
e study was conducted in Makokou region in the province of
Ogooue Ivindo, North East of Gabon (Figure 1). e climate of this
region is hot, humid and equatorial and characterized by a double
alternation of rainy and dry seasons [29]. Four seasons exist in this
province and can be distinguished as such-short rainy season (from
mid-September to mid-December, characterized by fairly frequent
thunderstorms sometimes accompanied by tornadoes), short dry
season (from mid-December to mid-March which is oen marked
by clear skies and a few sporadic rains), long rainy season (from
mid-March to mid-June, this period is characterized by an increase
in precipitation which are most oen stormy) and long dry season
Citation: Mavoungou JF, Nguema RM, Acapovi GL, Koumba RZ, Mounioko F, et al. (2017) Breeding Sites of Stomoxys spp (Diptera: Muscidae), a Preliminary
Study in the Makokou Region (North-East-Gabon). Vector Biol J 2:1.
• Page 2 of 7 •
doi: 10.4172/2473-4810.1000115
Volume 2 • Issue 1 • 1000115
(from July to August, the negative eect about this period is the
sharp decrease in rainfall on vegetation which is also characterized
by an almost permanent high cloudiness throughout the season), it
was during this period that the lowest temperatures were observed
during the year. e annual rainfall is around 1700 mm on average;
while the average annual temperature is 24° C, with a minimum of
21.7°C in July and a maximum of 25 °C in April. e annual and daily
thermal amplitudes are low (about 3.3 °C). Vegetation consists mainly of
primary forest land and oodplain forests. ere are also some secondary
forest patches resulting from the impact of human activity in the region
and can be observed in various places such as roads or borders around
the formerly cultivated plots. ey are mostly canopy-trees (Musanga
cecropioides) and various Anthocleista spp of the family Loganiacae [1].
e fauna is rich and diverse; there are nearly 128 species of mammals,
424 species of birds, 65 species of reptiles and 47 species of amphibians.
Some of these species posses nocturnal activity and others diurnal, like
most bird species. Nocturnal forest mammals consist of 11 species.
Collection of substrates suitable for the development of
stable y’s larvae
Eight types of suitable substrates for the reproduction of stable
ies were collected along a transect from the primary forest in an
anthropic environment (Makokou districts) through secondary forest.
e primary and secondary forests represented natural environments
while the neighborhoods such as Makokou stood for articial or man-
made environments. In the primary forest, three types of substrates
were collected: litter (PFL), elephant dropping (ELD) and small
African antelope (Cephalophus) dropping (CED). In secondary forest,
two types of substrates were collected which consisted of litter (SFL)
and Chimpanzee dropping (MOD), while in the neighborhoods of
Makokou, three substrates were collected namely: decaying plant
matter (VMD), organic waste from garbage in villages (VIG) and
cow dropping (CAD). e substrates were collected with a shovel
and with a minimum of ground to avoid the presence of soil-dwelling
larvae. Collected substrates were placed in plastic bags (polyethylene)
labeled with information like place of collection and type of waste.
All the collected substrates were transported to the laboratory and
distributed in cages.
Breeding method
e emergence cage traps are generally pyramidal in shape (40 cm
length x 30 cm width x 40 cm height). eir basal support is tightly
surrounded by a wooden frame (50 x 50 cm). is basal support is
covered by a black cotton fabric to ensure darkness. At the top of this
trap, a funnel-shaped cage is placed and this Roubaud cage is covered
with a bag made up of white mosquito net. is device was used to
trap the imagos aer emergence (Figure 2). An aliquot of 700 g of
each substrate was placed in each cage prior to emergence for a total
of 12 emergence cages. ese cages were installed in a room where the
temperature was maintained at 25°C throughout the study, but for
the Chimpanzee dropping (from the primary and secondary forest), 6
cages containing 400g each of the substrate were used (Tabl e 1).
e emergence cages were monitored daily for a period of 25
days. e appearance of adult ies (imago) indicated the completion
of their developmental cycle and they are attracted to light in the
Roubaud cages where they are trapped. ey were harvested and
preserved in 95 % alcohol prior identication as stomoxes or others.
Stomoxys spp were separated from other insects, counted, date noted,
species and capture time recorded. e identication of stable ies
was made under a dissecting microscope based on the identication
key issued by Zumpt [4] and Garros [30].
Data analysis
A correspondence analysis (CA) was conducted to see the
structuring of data on the aggregate table containing 84 rows and 7
Figure 1: Location of study area.
Figure 2 : Emergence cage trap.
Month (2006) Source of substrate Nature of substrate Number of cages used Mass of substrate in cages (grams)
Jan. Primary forest Litter (PFL) 12 700
Feb. Secondary forest Litter (SFL) 12 700
Mar. Anthropized milieu Cow dropping (CAD) 12 700
Apr. Primary forest Elephant dropping (ELD) 12 700
May Anthropized milieu Decomposing grass (VMD) 12 700
Jun. Primary forest Cephalophus dropping (CED) 12 700
Jul. Secondary forest Chimpanzee dropping (MOD) 6 400
Aug. Anthropized milieu organic waste from garbage (VIG) 12 700
Table 1: Data on the implementation of protocol.
Citation: Mavoungou JF, Nguema RM, Acapovi GL, Koumba RZ, Mounioko F, et al. (2017) Breeding Sites of Stomoxys spp (Diptera: Muscidae), a Preliminary
Study in the Makokou Region (North-East-Gabon). Vector Biol J 2:1.
• Page 3 of 7 •
doi: 10.4172/2473-4810.1000115
Volume 2 • Issue 1 • 1000115
increasing gradient readings from primary forest litter upto trash and
debris from the village, through Cephalophus dropping and elephant
dropping (Figure 6).
Analysis of variance (ANOVA)
To clarify the results of this rst CA, ANOVA abundance (log
data) of each species was conducted with a single factor, known as
medium (Figure 7). e results obtained for each species were as
follows:
S. calcitrans: log-abundance is signicantly (F6, 77 = 6.85, p-value
<0.001) dierent between environments. Peak abundance was
observed in the trash and debris from village.
S. niger niger: log-abundance was signicantly (F6, 77 = 7.41,
columns giving plenty of each of the seven species identied in each
of the 84 bred. is analysis took into account qualitative parameters
such as: species and dierent surveys. Information relating to media
(litter primary forest, cow dung, secondary forest litter, etc.) was not
used in this CA. It was rather used aer with CA, to classify records.
To study the species-repeated relationship, the 84 × contingency
table S (where S is the number of species) giving abundance of
each species in each repetition and was analyzed using analysis of
correspondence (CA) [31]. e type of reproduction media was not
included in the CA, but was used aer to check whether the orderly
nature of replication by CA was compatible with the media type.
Based on each species, one-way analysis of variance was performed
to test the dierences in the abundance of species between breeding
substrates. e statistical unit consisted of 84 observations for each
species. To address the heterogeneity of the data, abundances were
transformed logarithmically before the analysis of variance: the
number of individuals x species were replaced by log10 (x + 1). e
analysis of variance was completed by a pair-wise comparison test
of Newman-Keuls. To study the relationship of media-species, a 7 ×
contingency table S giving the cumulative abundance of each species
in each reproduction medium was analyzed using correspondence
analysis. A signicant dierence was found between the substrates (P
< 0, 05). e CA was completed by a χ2- test to show the relationship
between species and breeding substrates. All statistical analyses
were performed using the R statistical soware of version 2.15.0 (R
Development Core Team 2012).
Results
Overall abundance of the emerged insects
In the collection cages, the overall adult that emerged was 14517
and consisted of 11726 (81%) stomoxes and 2791 (19%) specimens
other than Stomoxys spp (Figure 3).
Stomoxys spp identied
Of the 11726 stomoxines that emerged, seven species were
identied with varying abundances. ese species consisted of
Stomoxys transvittatus, S. niger niger, S. omega, S. inornatus, S.
calcitrans, S. xanthomelas and S. niger bilineatus. S. transvittatus was
the most abundant species with 8221(70%) individuals. S. niger niger
(1679) and S. omega (947) were represented with average rates of 14%
and 8% respectively. S. inornatus 463 (4%), S. calcitrans 224 (4%) and
S. niger bilineatus 185 (1.9%) were poorly represented. S. xanthomelas
(0.1%) was least represented (Figure 4).
A large variation was observed between the number of emerged-
ies and dierent substrates. Indeed, the litter of secondary forest
(SFL) (5,268 (45%)) and cow dropping (CAD) (3,882 (33%)) were the
preferred substrates of stomoxes (Figure 5). Primary forest litter (PFL)
was the least productive substrate with only 138 (1%) individuals of
stomoxes identied aer emergence. Other substrates established a
less favorable breeding ground and development of stable ies as seen
with an emergence rate of 1%.
CA analysis
e projection of every statement and the illustrations on the
rst two axes of the CA is shown in Figure 6. Although medium is
not used as a variable in the CA, consistency was observed between
the structure readings and settings. Indeed, rstly, axis 1 of the CA
explains 28.0% of the total inertia, while the second axis explains
only 22.6%. Furthermore, axis 2 of the CA can be interpreted as
Other arthrpoda
Stomoxes
Figure 3: Insect-types emerging from emergence cage traps.
Figure 4: Stomoxys spp identied.
Figure 5: Distribution of ies based on collected substrates. PFL: Primary
Forest Litter; CAD: Cow dropping; SFL: Secondary Forest Litter; ELD:
Elephant dropping; HD: Decomposing plant matter; B CH: Chimpanzee
dropping; PD:Dust bin and Discharge.
Citation: Mavoungou JF, Nguema RM, Acapovi GL, Koumba RZ, Mounioko F, et al. (2017) Breeding Sites of Stomoxys spp (Diptera: Muscidae), a Preliminary
Study in the Makokou Region (North-East-Gabon). Vector Biol J 2:1.
• Page 4 of 7 •
doi: 10.4172/2473-4810.1000115
Volume 2 • Issue 1 • 1000115
p-value <0.001) dierent between environments; this species had
peak abundance in cow dropping.
S. niger bilineatus: log-abundance was signicantly (F6, 77 = 8.70,
p-value <0.001) dierent between environments; this species also had
peak abundance in cow dropping.
S. transvittatus: log-abundance was signicantly (F6, 77 = 2.69,
p-value = 0.02) dierent between environments. Peak abundance was
noticed in secondary forest litter.
S. inornatus: log-abundance was signicantly (F6, 77 = 2.44,
p-value = 0.03) dierent between environments. e peak abundance
was recorded with cow dropping
S. omega: log-abundance was signicantly (F6, 77 = 4.68, p-value
<0.001) dierent between environments, peak abundance for this
species was found in secondary forest litter.
S. xanthomelas: log-abundance, unlike other species, revealed no
signicant (F6, 77 = 2.2, p-value = 0.052) dierence between substrate
environments.
CA and the χ2-test
In this second analysis, each medium was represented by 12
readings. Aer cumulative abundances of each medium were
considered, a new contingency table was generated. A CA latter and
Figure 6: Projection of the CAS. tran: S. transvittatus, S. ino: S. inornatus,
S. x: S. xanthomelas, S. cal: S. calcitrans, S. n nig. S. niger niger, S ome
S. omega, S. n bil. S. niger bilineatus. Each black dot represents a record,
while ellipses represent circles.
Figure 7: Emergence of Stomoxys Spp based on environments S. t. S. transvittatus, S. i: S. inornatus, S. x: S. xanthomelas, S. Cal: S. calcitrans, S. n. n:
S. niger niger, S. o: S. omega, S. n. b: S. niger bilineatus
Citation: Mavoungou JF, Nguema RM, Acapovi GL, Koumba RZ, Mounioko F, et al. (2017) Breeding Sites of Stomoxys spp (Diptera: Muscidae), a Preliminary
Study in the Makokou Region (North-East-Gabon). Vector Biol J 2:1.
• Page 5 of 7 •
doi: 10.4172/2473-4810.1000115
Volume 2 • Issue 1 • 1000115
cumulative abundance of each species in each medium was carried
out. e rst axis of the CA explains 50.4% of the total inertia, while
the second axis explains 40.7%. e projection of 7 environments and
7 species on the rst two axes of the CA is shown in Figure 8. is
Figure 8 shows associations between environments and species. CA
was supplemented by a χ2-test that revealed a signicant (χ2 = 4046.7,
p-value <0.001) binding between species and environment.
Discussion
e results obtained in this study is preliminary and focused
in dening the potential suitable development substrates for the
larvae of stable ies in the Makokou region of north east, Gabon.
Furthermore, the methodology developed in this work can be used,
notably in primary /secondary forests, savannahs and anthropogenic
habitats (villages). is present research work reveals that stable
ies can lay their eggs on several types of substrates including: litter,
elephant dropping, Cephalophus dropping, chimpanzee dropping,
decomposing plant material, cow dropping and organic waste from
dust bins in villages. ese results corroborated with those obtained
by Gamal [32], Grimaud [33] which showed that stable ies develop
in sites associated with unsanitary conditions, animal waste coupled
with organic waste from plant origins. e results obtained in this
study have also shown that stable ies can colonize several types of
environments such as in primary/secondary forests and man-made
environments (villages). ese observations are similar to that made
from the studies conducted by several authors who showed that
stable ies are present in many habitats including primary /secondary
forests and anthropic environments [1,13,16,17,34]. Knowledge of
biological diversity and risk assessment related to biting ies (stable
ies) in human and animal circles are of paramount importance to
human health. In this study, seven species of stomoxes previously
described in this region were signaled [1,27]. ese species were
represented by S. calcitrans, S. transvittatus, S. inornatus, S. omega,
S. niger niger, S. niger bilineatus and S. xanthomelas. Several types
of potentially favorable breeding grounds of these species have thus
been identied. Generally, substrates that favoured the emergence
of ies are from disturbed environments such as secondary forests
and man-made environments. ese substrates are represented by
secondary forest litter and cow dung from man-made environments.
e dierences in adult Stomoxys spp emergence in various
substrates isolated could be explained by the dierent micro-climatic
conditions prevailing in these substrates which oer a conducive
milieu for the development of stable y larvae. Furthermore,
temperature, relative humidity and rate of drying substrates may
vary depending on the type of environment. In addition, the litter
of secondary forest and cow dropping seem to oer ideal conditions
for stable y larval survival. is nding is similar to that obtained
by Nibaruta [35] who showed that diptera are likely to grow in large
numbers in cow dropping. ese results also support the work of
Vaillant [36] who reported that some species of blood-sucking ies
prefer to deposit their eggs or larvae on organic material of animal
origin. According to the WHO [37], an increment in the number of
disease cases transmitted by certain species of biting ies could be
due to organic material of animal origin as well as faeces from cattle,
providing conditions for the development of eggs and larvae of these
insects. ese substrates could provide an ideal environment for the
development of stable y larvae and thus a better reason for their
blooms in the surveyed environments. Furthermore, data analysis
showed that some Stomoxys species were substrate specic. Similarly,
the correspondence analysis conrmed this assertion, based on the
association between environment and species. Indeed, S. omega, is
associated with primary forest litter, Cephalophus dropping and
elephant dropping. Considering S. transvittatus and S. inornatus,
they are only associated with secondary forest litter. S. niger niger is
more frequent in cow dropping and rotting weeds. S. xanthomelas,
S. calcitrans and S. niger bilineatus are associated with garbage and
village debris (man-made environments). ese results could be
explained by the fact that each species has its own bio-ecological
requirements for instance S. niger niger is a species that is most
prevalent in tropical rain forest areas of Africa and in South Sudan
farmer-settlement areas [1,4]. Considering S. calcitrans, it is known as
a cosmopolitan species oen associated with human activities related
to livestock [4,13]. S. xanthomelas is present in forest areas [4,24]. S.
transvittatus, S. omega and S. inornatus seem to be more subservient
to the forest where human density is low and where the oor of this
forest is covered with organic material. Microclimate presented by
dierent substrates could therefore play a role in the distribution and
abundance of dierent species of stable ies. Gilles [13,14] showed
that temperature had an eect on the rate of development of some
species including Stomoxys calcitrans and S. n. niger. Comparing the
abundances using ANOVA, it was noted that there were signicant
dierences in the distribution of certain Stomoxys species depending
on the substrate. S. xanthomelas is the least abundant species (seven
occurrences only) while S. transvittatus was most abundant (with
8,189 occurrences). Similarly, substrates such as secondary forest
litter and cow dropping were those that manifested high abundance
of stable ies with 5,268 and 3,883 proportions respectively. ese
environments appear to be richer in minerals that favor the larval
development of stable ies than other substrates like primary forest
litter. It is not surprising with the unusual peaks of abundance of
many species. S. calcitrans peak was highest in trash and debris from
village. Considering S. niger niger, S. niger bilineatus and S. inornatus,
their peak abundances occurred in cow dropping. S. transvittatus and
S. omega, presented their peak abundances in secondary forest litter.
S. xanthomelas did not present signicant dierences between the
sampled environments.
Figure 8: Projection of the association between environments and species.
S. t: S. transvittatus, S. i: S. inornatus, S. x: S. xanthomelas, S. Cal. S.
calcitrans, S. n. n: S. niger niger, S. o: S. omega, S. n. b: S. niger bilineatus.
Citation: Mavoungou JF, Nguema RM, Acapovi GL, Koumba RZ, Mounioko F, et al. (2017) Breeding Sites of Stomoxys spp (Diptera: Muscidae), a Preliminary
Study in the Makokou Region (North-East-Gabon). Vector Biol J 2:1.
• Page 6 of 7 •
doi: 10.4172/2473-4810.1000115
Volume 2 • Issue 1 • 1000115
e second CA, associated with the χ2-test enabled us to show
clearly that axis 1 thereof include exactly the same gradient circles on
axis 2 of rst CA, with one end standing for un-disturbed environment
(primary forest litter, elephant dropping and Cephalophus dropping
and the other end anthropogenic environments (garbage and debris
village and decaying grass) and between the two secondary areas
(secondary forest litter and cow dropping). Moreover, relying on
the contributions of the χ2-test, we can identify some denitive
associations. Indeed, S. omega is specically associated with primary
forest litter, the duiker (genus: Cephalophus) dropping and elephant
dropping. is is consistent with the rst CA. However, this same
species has peak abundance in litter of secondary forest. e peak
abundance could simply result from the fact that the abundance is
globally (all species) higher in secondary forest litter, which does not
reect a specic association. S. niger niger is specically associated
with decaying plant material and is consistent with the rst CA.
is species showed peak abundance in cow dropping and could be
explained by the fact that the abundance of stable ies is generally
very high in cow dung. S. calcitrans is specically associated with
village trash and debris. Moreover it is a cosmopolitan species that
can grow on various substrates and environments. Considering S.
inornatus, S. transvittatus, S. xanthomelas and S. niger bilineatus, they
do not show a clearly dened media preference.
Conclusion
is study is a valuable contribution to identifying potentially
suitable substrates for the development of stable y larvae. e
identied substrates were represented by litter, elephant dropping,
Cephalophus dropping, chimpanzee dropping, decaying plant
material, organic waste from garbage in villages and cow dropping.
Furthermore, there is substrate preference of these species and
this depends on the type of substrate for example S. omega is
specically associated with the litter of primary and secondary forest,
Cephalophus and elephant droppings. S. niger niger is associated
with decaying plant material and cow dropping. S. calcitrans is
associated with village trash and debris. S. inornatus, S. transvittatus,
S. xanthomelas and S. niger bilineatus do not have a clearly dened
preferred environment. It is interesting to know that each species
presented a maximum abundance in a particular environment. In
cages, S. calcitrans peak abundance was noticed in trash and debris
from village. S. niger niger, S. niger bilineatus and S inornatus were
most abundant in cow dropping. Based on S. transvittatus and S.
omega, they presented the highest and eective development in
litter from secondary forest. S. xanthomelas presented no particular
abundance with substrate. Biostatistical analyses revealed a net
liation between the dierent species described and selected potential
breeding site. e distribution and abundance of these blood-sucking
insects varied with environment. is reects the openness, ora and
fauna composition of the environment.
A thorough study of each substrate including factors such
as: humidity, percentage of organic matter (plant and animal),
particle size of the ground and monitoring of annual variations of
these substrates is essential in the determination of the ecological
requirements for larval development of each kind of stable y.
Acknowledgements
This work received funding from the European Union through the
European Development Fund Project 8 ACP GA 009 (PSVAP-Component II
Makokou, CNPN-MESRIT-U.E./IRETCIFOR). We would like to thank Philippe
Hecketsweiller, Paul Posso, Robert Nasi and Charles Doumenge for their advice
and assistance. Thanks to Nicolas Yao Emba, Joseph Okouyi.
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Author Afliations Top
1Institut de Recherche en Ecologie Tropicale (IRET-CENAREST), Libreville,
Gabon
2Université des Sciences et Techniques de Masuku (USTM), Franceville,
Gabon
3Laboratoire de Zoologie & Biologie Animale, UFR Biosciences, université Felix
Houphouet Boiny , Abidjan-Côte d’Ivoire
4Laboratory of vector-borne-infectious Diseases, Unit of Laboratory of Biology
and applied Ecology, Department of animal Biology, Faculty of Science,
University of Dschang, Cameroon
5Institut für Vergleichende Tropenmedizin und Parasitologie, München,
Germany
6Université Paul-Valéry Montpellier 3, Centre d’Ecologie Fonctionnelle et
Evolutive, UMR 5175, Montpellier, France
7Chercheur associé du CIRAD au CENAREST Cirad, unité de recherche
B&SEF, BP 4035, Libreville, Gabon
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