The Contribution of Termitaria to Plant Species Conservation in the Pendjari Biosphere Reserve in Benin

Abstract

The role of termitaria in plant species diversity and conservation was investigated in Pendjari National Park and surroundings. The study objectives were to (i) compare termitaria vegetation similarity, life form and chorological composition between areas, (ii) investigate the difference of species richness and diversity on termitaria between living versus abandoned termitaria, and (iii) predict the species richness and diversity on termitaria as a function of termiaria size. Results show that species from the combretaceae family were the most abundant on termitaria; plant communities on termitaria were different among areas and between living and abandoned termitaria. Phanerophytes were abundant on mounds, therophytes were abundant on termitaria within the protected areas and farms outside of the park; however hemicryptophytes were as abundant as phanerophytes in fallows. From a phytogeographic perspective, species typical of Sudanian ecological region were the most abundant on termitaria. There was no difference in the richness and diversity of termitaria woody species, between termitaria state (dead or live) or between management type (protected or unprotected). Termitaria areas rather than termitaria height significantly correlated with the species richness and diversity. Larger termitaria had the highest diversity. Termitaria vegetation reflects the influence of local climate, moreover termitaria size can be used to predict the species richness and diversity related to them. The termitaria size has no influence on the richness and diversity of woody species related to them.

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Environment and Ecology Research 4(4): 200-206, 2016 http://www.hrpub.org
DOI: 10.13189/eer.2016.040403
The Contribution of Termitaria to Plant Species
Conservation in the Pendjari Biosphere Reserve in Benin
H. O. Dossou-Yovo*, A. E. Assogbadjo, B. Sinsin
Laboratory of Applied Ecology, Faculty of Agronomic Sciences, University of Abomey-Calavi, Benin
Copyright©2016 by authors, all rights reserved. Authors agree that this article remains permanently open access under the
terms of the Creative Commons Attribution License 4.0 International License
Abstract The role of termitaria in plant species diversity
and conservation was investigated in Pendjari National Park
and surroundings. The study objectives were to (i) compare
termitaria vegetation similarity, life form and chorological
composition between areas, (ii) investigate the difference of
species richness and diversity on termitaria between living
versus abandoned termitaria, and (iii) predict the species
richness and diversity on termitaria as a function of termiaria
size. Results show that species from the combretaceae family
were the most abundant on termitaria; plant communities on
termitaria were different among areas and between living
and abandoned termitaria. Phanerophytes were abundant on
mounds, therophytes were abundant on termitaria within the
protected areas and farms outside of the park; however
hemicryptophytes were as abundant as phanerophytes in
fallows. From a phytogeographic perspective, species typical
of Sudanian ecological region were the most abundant on
termitaria. There was no difference in the richness and
diversity of termitaria woody species, between termitaria
state (dead or live) or between management type (protected
or unprotected). Termitaria areas rather than termitaria
height significantly correlated with the species richness and
diversity. Larger termitaria had the highest diversity.
Termitaria vegetation reflects the influence of local climate,
moreover termitaria size can be used to predict the species
richness and diversity related to them. The termitaria size has
no influence on the richness and diversity of woody species
related to them.
Keywords Termitaria, Plant Diversity, Ecology,
Conservation, Pendjari National Park
1. Introduction
Termitaria play a key role in the maintenance of the tree
diversity in savannah across West Africa [1-3], however our
understanding of their contribution to the conservation of
natural resources is still limited. Termite mounds occur in
woodlands as well as agricultural land [4-6] in the tropical
and subtropical regions where they constitute a typical
feature [7-8]. Termitaria, colonised by various species of the
fungus-growing termites (Isoptera; Macrotermitinae) occur
at densities ranging between 2 to 7 mounds per ha [8]. The
physical and chemical processes driven by termite regulate
soil fertility around termite mounds [9]. During mound
building, termites intensely modify soil properties, enriching
the clay content and increasing soil infiltration capacities
thus promoting microbial activity an improving nutrient
composition and availability to plants [10-14]. As a
consequence of this positive effect of termitaria on soils
properties, there is often a high proportion of rapidly
growing annual grasses and functionally-annual herbs
around them, whereas slower growing perennial grasses and
sedges are predominant far from them. For example, recent
study in the sudanian region of Burkina-Faso [15] revealed
that termitaria maintains higher tree species density
compared to the surroundings. Furthermore, the diversity of
species around termitaria was significant on sites subjected
to fire and grazing. There is also evidence of termtiria effect
on plant water economy strategy. For instance, Konaté et al
[11] reported a lower dry season loss of leaves (17% of
leaves lost) by Crossopteryx febrifuga individuals occurring
on mounds than those occuring far from it (78% of leaves
lost).
The type of termitaria as well as the higher diversity and
density of plants around them may also affect the spatial
distribution of wildlife. For instance, for nesting, females of
Andros Iguana (Cyclura cyclura cyclura) select mounds
with more than 5 cm surrounding soil depth [16]. Fleming
and Loveridge also reported that small vertebrates heavily
utilized Macrotermes mounds more than the surrounding
miombo woodland in Zimbabwe [17]. Large ungulates often
graze preferentially on mounds rather than on adjacent
vegetation [18, 19]. In Benin, Attignon et al [20, 21]
investigated the role of termites in nutrient recycling in
natural and plantation forests and concluded that both
changes in termite resource base and food-web disturbances
affect termite assemblages.
In this study, we investigate the effect of land-use system
on the diversity and functional role of termitaria-related

Environment and Ecology Research 4(4): 200-206, 2016 201
vegetation. Specifically, we tested if species diversity,
dominant life form and chorological composition of
vegetation on termite mounds were different between the
protected areas, farms and fallows in the Pendjari Biosphere
reserve in Benin. We also tested if termitaria status (live or
dead and abandoned by termites) and size affect plant
species diversity.
2. Materials and Methods
2.1. Study Area
This study was conducted from July to October 2007 in
the Pendjari Biosphere Reserve in Benin (10°30’-11°30 N,
0°50’-2°00’ E, Fig. 1). The reserve covers 471,140 ha of
which the Pendjari National Park covers approximately
56.47% and the Pendjari hunting zone only 43.53%. The
climate is dry with a single rainy season from mid-May to
mid-October and an annual mean rainfall of 1000 mm. The
dry season starts from mid-October to mid-May. The annual
mean temperature ranges between 18.6 ℃ and 36.8 ℃ in the
northern Reserve and in the southern, between 20.5 ℃and
34.2 ℃ [22]. Temperatures are highest in March and April
and lowest from December to January. The vegetation
consists of savannas, dry forest, woodlands and gallery
forests dominated by Terminalia, Combretum and Acacia
genera [23]. The most abundant species in shrub savanna are
Combretum glutinosum, Crossopteryx febrifuga, Acacia
seyal, Acacia senegal, Acacia gourmaensis [24]. Tree
savannas are dominated by Acacia sieberiana,
Pseudocedrela kotschyi, Terminalia macroptera, Detarium
microcarpum, Burkea africana, Afzelia africana and
Vitellaria paradoxa. Lastly, gallery forests are mostly
characterized by Diospyros mespiliformis, Borassus
aethiopium, Ficus capensis, Khaya senegalensis, Parinari
congoensis and Syzygium guineense. Thirteen ethnic groups
in the Pendjari Biosphere Reserve were identified and their
main income generating activities are agriculture, animal
husbandry and trade.
2.2. Data Collection
We identified homogeneous sites using the soil,
vegetation and rainfall maps of the Pendjari Biosphere
Reserve. A total of 84 termitaria (28 per land-use type:
protected areas, farms and fallows) were surveyed in the
study. We installed 50 m×50 m plots to count and measure
termite mounds. Mounds surveyed in the Pendjari National
Park were mostly epigeal termitaria. Several authors [13, 25,
26] identified them as Macrotermes genera buildings in
tropical savannahs. The status of each mound (living or dead
and abandoned) was assessed by slicing the mound to check
for presence of termintes and by looking at its colour. Living
mounds are mostly red while the dead and abandoned ones
are black sometimes going to white. Geographical
coordinates and altitude of each termitaria were recorded.
Around each mound, a single plot of 900 m2 (30 m×30 m)
was established to record the existing woody species. The
height and four radii of each termitaria were measured and
the mean radius was used to calculate mound areas [19].
Plant species found on mound were identified and
individuals’ woody species counted. The plants species were
identified at the National Herbarium of Université
d’Abomey-Calavi (Benin).
Figure 1. Location map of the biosphere reserve of Pendjari on the figure with the Atakora chain as southern border of the reserve, the Pendjari River the
north-western border and the country boundaries marked in grey.

202 The Contribution of Termitaria to Plant Species Conservation in the Pendjari Biosphere Reserve in Benin
2.3. Data Analysis
Plant communities established on the mounds were
compared between land use type and status of termitaria
using the Jaccard’s similarity index Ij on species
presence-absence data [27, 28]. Ij is defined as follows:
Where: Ij ＝Jaccard’s similarity index, a ＝number of
species found on mounds in land-use type A; b＝number of
species on mounds in land-use type B and c＝number of
species shared by A and B. Plant communities on mounds
were considered similar if Ij was more than 50%.
The dominant life form and phytogeographical spectra of
termitaria vegetation were compared between land-use type
and termitaria status. Woody and herbaceous species found
on termitaria were taken to determine the life form and
phytogeographical types of each species as indicated
respectively by Sinsin [29], Houinato and Sinsin [30] and
Adomou [31]. The main life forms used were: Th:
Therophytes, Ch: Chamephytes, G: Geophytes Ph:
Phanerophytes and Hc: Hemicryptophytes. About
chorotypes, the following symbols were used: At: Atacorian
species; PAL: Paleotropical species; Pt: Pantropical species;
S: Sudanian species; SG: Guineo-congolian / Sudanian
transition species and SZ: Sudano-zambesian species. The
proportions of various life form and phytogeographical types
were computed using the mound species checklists compiled
for each area.
We calculated Shannon index (H) and Evenness index (E)
as follows:
where Pi＝proportion of individuals of species i per mound
and S＝woody species richness for each mound.
We used Kruskal-Wallis H-test to compare mean species
richness, diversity parameters Shannon index (H) and
Evenness index (E) between land-use types and
Mann-Whitney to test for differences in the parameters
between live and dead abandoned termitaria. To test if
species richness and diversity on termitaria were height or
areas-dependent, we used linear regression. The dependent
variables were the species richness and the diversity indices;
mound height or areas were the independent variables. All
statistical analyses were performed using SPSS 16.0 for
Windows.
3. Results
3.1. Species Composition
The number of plant species recorded on termitaria varied
among land-use types. The proportion of species restricted to
termitaria as shown in Table 1 was higher in farms than in
the Park and fallows.
Tab l e 1. Number of plant species counted on termitaria per site categories
and proportion of restricted species per area
Land-use
type
Total
mounds
surveyed
Woody
species
richness
Total
species
richness
Number and
percentage of
restricted
species
Park 28 37 49 5 (13,5%)
Fields 28 42 74 11 (26,2%)
Fallows 28 33 50 4 (12,1%)
The lists and full names of plant species are available with
the author.
Among the woody species found on termitaria,
Combretaceae was overall dominant in all land-use types
(Fig. 2). Species from the Rubiaceae family were
co-dominant in the Park. In the farms, Mimosaceae ranked
second whereas Caesalpiniaceae was co-dominant on
mounds in fallows. The Shannon index and Evenness index
showed similar trend and low diversity of termitaria-related
woody species in our study areas (Table 2).
Tab l e 2. Summary of Shannon and Evenness indices of termitaria surveyed in different site categories.
Diversity
indices
Park Fields Fallows
Range Mean±SD Range Mean±SD Range Mean±SD
Shannon index
(Bits) 0-2.32 1.25±0.73 0-2.95 1.31±1 0-2 0.88±0.74
Evenness index 0-1 0.80±0.38 0-1 0.66±0.44 0-1 0.58±0.45

Environment and Ecology Research 4(4): 200-206, 2016 203
0 5 10 15 20
Anarcadiaceae
Annonaceae
Bombacaceae
Capparaceae
Caesalpiniaceae
Curcubitaceae
Euphorbiaceae
Liliaceae
Malvaceae
Mimosaceae
Rhamnaceae
Sterculiaceae
Vitaceae
family
% Species
Fallows
Fields
Park
Figure 2. Plant family diversity and ranking by site category showing combretaceae species as the most abundant on termitaria whatever the area.
3.2. Species Richness and Evenness Comparison
There was no significant difference in species diversity
between termitaria (Kruskal-Wallis, prichness ＝0.192) and
land-use types (pshannon＝0.07). However, the Evenness index
of termitaria was different between sites categories
(Kruskal-Wallis, p＝0.03). Furthermore, the Evenness mean
rank for the three land-use types revealed highest evenness
(best occupation of termitaria by species) in farms and the
Park compared to fallows (ZFallows＝-3.06; ZFields＝1.55 and
ZPark＝1.52). For all site categories combined, there was no
significant difference in species richness, as shown by both
the Shannon and Evenness indices between living and
abandoned termitaria (Mann Whitney, prichness＝0.66; pshannon
＝0.56 and pevenness＝0.82).
3.3. Correlation between Mound Size and Species
Diversity
There was no significant correlation between termitaria
height and species richness and diversity whereas mound
surface area was positively correlated with the
above-mentioned parameters [Linear regression of surface
area with (i) species richness: p＝0.03 and R2＝10.6%; (ii)
Shannon index: p＝0.03 and R2＝10.4%; Evenness: p＝0.20
and R2＝6.5%].
3.4. Comparison of Plant Communities
Jaccard’s values were lower than 50% for all land-use
types but the comparison between farms and fallows
revealed the highest values of Jaccard’s index for both
woody communities and the total termitaria flora. So,
termitaria in farms and fallows had more species in common
compared with the mounds vegetation in the Park. For all
category sites, woody species on abandoned termitaria (46
species) were significantly different (Ij ＝43.07%) from
living termitaria (47 species).
3.5. Life form Spectrum
Phanerophytes were the most abundant on mounds.
Among herbaceous species, therophytes were the most
abundant on mounds in the Park, farms and fallows.
Hemicryptophytes were co-abundant with therophytes on
termitaria in fallows.
3.6. Chorological Spectrum
Sudano-zambesian species were the most abundant on
mounds in the Park whereas in farms and fallows, sudanian
species were the most abundant. In total, species of sudanian
chorological group consisted of Sudanian and

204 The Contribution of Termitaria to Plant Species Conservation in the Pendjari Biosphere Reserve in Benin
Sudano-Zambesian were the most abundant on mounds in
our study areas.
4. Discussion
4.1. Species Composition and Diversity
With respect to species restricted to termitaria, Cadaba
farinosa (Capparaceae) was also reported on mounds by
Traoré et al [15]. The latter had mentioned the genera
Capparis and Maerua (both Capparaceae) as solitary on
mounds. In addition, Arbonnier [32] mentioned termitaria as
Capparaceae’s habitat. Results from our investigation and
mentioned studies show termitaria to be hotspots for
Capparaceae species [15, 32]. Indeed, Arbonnier [32] also
mentioned dry stations and rocky soils and mountains among
Capparaceae’s habitat. As a result, we assume that there are
some similar patterns between types of termitaria on which
these species grow and the other habitat types where the
species are found. An investigation can be undertaken to
state this similarity.
The abundance of Combretaceae species on termitaria
(Fig. 2) is consistent with the results of Thiombiano et al [33]
who reported an increased number of Combretaceae species
in Burkina Faso. Similarly, Traoré et al [15] identified
Combretaceae, Mimosaceae, Rubiaceae and Caesalpiniaceae
as predominant families on termitaria. So Combretaceae
species seem dominant on termitaria in sudanian regions and
this finding could be explained by the compliance of
vegetation with climatic conditions. The implication of this
result for conservation and vegetation sampling is the
conservation of the concerned species through the
sustainable management of termitaria. In addition, termite
mounds can serve as hotspots for Combretaceae sampling in
sudanian regions and further researches can be conducted to
highlight the ecological patterns responsible of
Combretaceae distribution on termitaria in sudanian
savannahs.
The low values of Shannon index (Table 2) may be
explained by the fact that termitaria are selective habitats. In
other words, they may be some specific plant species having
termitaria as habitats and these species are poorly diversified
and this fact leads to low termitaria species diversity.
4.2. Species Diversity and Termitaria Size
Our study reveals no significant effect of termitaria status
on plant species richness and diversity. This is consistent
with findings by Traoré et al [15] in similar ecological
settings. However, termitaria status tends to influence their
floristic composition. Specifically, we found that species
such as Maerua oblongifolia, Flueggea virosa and Feretia
apodanthera were more abundant on abandoned mounds
while Detarium microcarpum, Combretum molle and
Diospyros mespiliformis were observed on live termitaria
(complete list of species rank-abundance available with
author). With regards to the habitat of the three species found
on abandoned mounds according to Arbonnier [32], F.
virosa abundant on abandoned mounds is characteristic of
disturbed habitats while F. apodanthera is said to colonize
compact and battleship soils. M. oblongifolia is found on all
types of soil. We assume in this study that these species have
in common the disturbed and poor habitat since compact and
battleship soils mentioned above are mostly poor in terms of
fertility. Abandoned and dead mounds are consequently seen
as disturbed habitats in this paper but this has not been stated
yet.
Similarly to our findings, many other authors reported the
independence between land-use type on mean species
richness and diversity [34-36].
Moreover, termitaria areas can be used to predict a part of
plant species richness and diversity on mounds. This positive
relation is consistent with the findings of Harner and Harper
[37]. Chapin III et al [38] reported that several experiments
indicate linear and asymptotic relationship of ecosystem
process rates with species richness. So as long as termitaria
are conserved, termite species will be conserved.
Consequently the building activities may lead to increasing
termitaria areas. The implication of this latter for
conservation is that the probability of species occurrence
with strong ecosystem effects will increase. In contrast to the
positive relation between termitaria areas and species
evenness found in our study areas, Hurlbert [39] found that
species evenness tends to decrease with sample size. The low
values obtained for the coefficient of determination (R2)
suggest that a little part of the variation of plant species
richness and diversity was explained by surface variation.
Therefore, other additional ecological factors such as
chemical characteristics of mounds and the annual rainfall
received by termitaria, may have contributed to plant species
richness and diversity on termitaria. Regardless of all other
factors susceptible of influencing plant species richness and
diversity in relation with termitaria, results highlighted in
this paper allow us to conclude that termitaria conservation
will lead to increasing species richness and diversity since
they increase with termitaria surface.
Finally, we suggest that further studies focusing on the
spatial distribution of termitaria and variation in their size be
done. Studies on the variation in species diversity related to
the distribution of termitaria could serve to predict the spatial
heterogeneity of microhabitats and its impact on plant
species diversity in semi-arid ecosystems.
4.3. Comparison of Plant Communities
Plant communities on mounds were different among
land-use types (Ij＜50%). Termitaria in the various areas
offer various floristic compositions. The assumption is that
the ecological factors which determine species distribution
on termitaria seem to be different between management
areas. For example, the dissimilarity in floristic composition
between land-use types may reflect the difference in
dispersal agents between areas. Farms and fallows are easily

Environment and Ecology Research 4(4): 200-206, 2016 205
accessible to people and domestic animals contrary to the
Park wherein wild animals are predominant. As a result, wild
and domestic animals as well as humans will transport
different diaspores cross termitaria in investigated areas. The
fact that termitaria in fields and fallows shared more species
(Ij ＝47.1%) may probably be due to the effect of the
common diaspores transport agents in these areas and other
factors such as wildland fire. Traoré et al [15] stated the
positive effect of fire and livestock grazing on tree density
found on termitaria but no investigation has been directed to
measure the impact of these factors on termitaria floristic
composition.
4.4. Termitaria Life Forms and Phytogeography
Life form spectra reflect not only the sum of ecological
factors, including climate and human influences, but the
historical factors of transport and introduction of new species
from abroad [40] Here we do not focus discussion on
climatic factors since they are not measured in this study.
However, we assumed that as termitaria are located in the
same bioclimatic zone, climate influence may be less
important. The importance of therophytes on termitaria in
fields might be due to the fact that after ploughing or
weeding farms, farmers put heaps on termite mounds. The
herbs mostly consist of therophytes of seed origin.
Elsewhere, animals and humans contribute also to the
abundance of therophytes [40-41]. In savannah ecosystems,
termitaria are browsing spots for megaherbivores [14-19].
Some animal species such as lions (Panthera leo) eat their
prey on termitaria whereas other animal species get rest on
mounds (example hyena: Crocuta crocuta; unpublished
data). Despite the likely low disturbance on termitaria in the
National Park compared to farms, all the actions highlighted
above could contribute to the abundance of therophytes on
termitaria within the National parks.
The importance of sudanian species on mound reflects the
effect of climate [28-33]. This confirms the fact that the
specific plant composition of termitaria and their
physiognomy vary between phytogeographical regions
[15-17, 42, 43, 44]. The abundance of sudanian species
confirms the compliance of termitaria flora with the study
area which is a sudanian zone. The sustainable management
of termitaria will consequently contribute to sudanian
species conservation in the Pendjari Biosphere Reserve.
Acknowledgements
This work was financially supported by BIOTA. We thank
this institution and its donors. Our acknowledgements also
go to local communities around the National Park of Pendjari,
Dr. Aristide Adomou (National Herbarium, University of
Abomey-Calavi Benin) and Aristide Tehou (Pendjari
National Park). We are also grateful to Orou Gaoué (PhD)
and Cuthbert Katsvanga for editing the draft of this paper.
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