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Which one comes first, the tamarind or the Macrotermes termitarium?

Authors:
  • Université Nationale d'Agriculture, Bénin
  • Faculty of Agronomic Sciences | University of Abomey-Calavi

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The relationship between termitaria and their vegetation is being increasingly studied. Nonetheless, our understanding of the order of establishment of termitaria and their associated vegetations which may be relevant for developing conservation plans is limited. This study focuses on order of establishment of Macrotermes termitaria and associated plant species with a special focus on Tamarindus indica and was to answer whether tamarind trees establish before termitaria or reversely? A comparative analysis of T. indica-dominated vegetations on termitaria and adjacent vegetations was undertaken using a matrix involving 80 relevés across four phytogeographical districts (phytodistricts), and an Indicator Species Analysis. We discussed how informative vegetation data could be on termitaria and tamarind trees establishment order. Whatever the phytodistrict, vegetations associated with termitaria were found to be much similar to those of their adjacent areas. Overall, only seven species (not including T. indica) out of a total of 63 recorded were found to be confined to termitaria. These results would suggest termitaria to not be a factor controlling establishment of T. indica and most of the species their host. Comparative vegetation analysis was thus found not to be enough to stand on termitaria and T. indica establishment order. However, termitaria-tamarind associations may be profitable to both tamarind trees and termites: termitaria may help mitigate drought on tamarind trees under increasing drought conditions while tamarind trees may offer food to termites. Integration of information on termite species’ feeding preferences and ecology was proposed in order to improve current state of knowledge on tamarind-termite relationship.
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Which one comes first, the tamarind or the
Macrotermes termitarium?
B. Fandohan a b , A.E. Assogbadjo a , V.K. Salako a , P. van Damme c & B. Sinsin a
a Laboratoire d’Ecologie Appliquée, Faculté des Sciences Agronomiques, Université
d’Abomey-Calavi, 01BP 526, Cotonou, Bénin
b International Ecosystem Management Partnership (IEMP), United Nations Environment
Programme, c/o Institute of Geography and Natural Resources Research, Chinese
Academy of Sciences, No. 11A Datun Rd. Beijing 100101, China
c Laboratory of Tropical and Subtropical Agriculture and Ethnobotany, Faculty of
Bioscience Engineering, Ghent University, Coupure links 653, B- 9000, Ghent, Belgium
Version of record first published: 26 Nov 2012.
To cite this article: B. Fandohan , A.E. Assogbadjo , V.K. Salako , P. van Damme & B. Sinsin (2012): Which one comes first,
the tamarind or the Macrotermes termitarium?, Acta Botanica Gallica, 159:3, 345-355
To link to this article: http://dx.doi.org/10.1080/12538078.2012.721252
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Which one comes rst, the tamarind or the Macrotermes termitarium?
Lequel vient en premier, le tamarin ou le Macrotermes termitarium?
B. Fandohan
a,b
*, A.E. Assogbadjo
a
, V.K. Salako
a
, P. van Damme
c
and B. Sinsin
a
a
Laboratoire dEcologie Appliquée, Faculté des Sciences Agronomiques, Université dAbomey-Calavi, 01BP 526, Cotonou, Bénin;
b
International Ecosystem Management Partnership (IEMP), United Nations Environment Programme, c/o Institute of Geography and
Natural Resources Research, Chinese Academy of Sciences, No. 11A Datun Rd. Beijing 100101, China;
c
Laboratory of Tropical and
Subtropical Agriculture and Ethnobotany, Faculty of Bioscience Engineering, Ghent University, Coupure links 653, B- 9000, Ghent,
Belgium
Abstract: The relationship between termitaria and their vegetation is being increasingly studied. Nonetheless, our
understanding of the order of establishment of termitaria and their associated vegetations which may be relevant for
developing conservation plans is limited. This study focuses on order of establishment of Macrotermes termitaria and
associated plant species with a special focus on Tamarindus indica and was to answer whether tamarind trees establish
before termitaria or reversely? A comparative analysis of T. indica-dominated vegetations on termitaria and adjacent
vegetations was undertaken using a matrix involving 80 relevés across four phytogeographical districts (phytodistricts),
and an Indicator Species Analysis. We discussed how informative vegetation data could be on termitaria and tamarind
trees establishment order. Whatever the phytodistrict, vegetations associated with termitaria were found to be much
similar to those of their adjacent areas. Overall, only seven species (not including T. indica) out of a total of 63 recorded
were found to be conned to termitaria. These results would suggest termitaria to not be a factor controlling establishment
of T. indica and most of the species their host. Comparative vegetation analysis was thus found not to be enough to stand
on termitaria and T. indica establishment order. However, termitaria-tamarind associations may be protable to both
tamarind trees and termites: termitaria may help mitigate drought on tamarind trees under increasing drought conditions
while tamarind trees may offer food to termites. Integration of information on termite speciesfeeding preferences and
ecology was proposed in order to improve current state of knowledge on tamarind-termite relationship.
Keywords: plants-insect relationship; termitaria; indicator species; phytogeograpical districts
Résumé: La relation entre les termitières et leurs végétations a été assez documenté. Toutefois, notre compréhension de
lordre détablissement des termitières et des végétations associées qui peut être importante pour lélaboration de plan de
conservation est limitée. Cette étude sest focalisée sur lordre détablissement des termitières des Macrotermes et des
espèces de plantes associées en nous focalisant sur Tamarindus indica (le tamarinier sétablit-il avant la termitière ou le
contraire ?). Une comparaison des végétations des termitières à T. indica avec celles des zones adjacentes a été effectuée
en utilisant une matrice de 80 placeaux dans quatre districts phytogéographiques (phytodistricts) et une Analyse des
Espèces Indicatrices. La végétation des termitières sest révélée assez similaire à celle des zones adjacentes quelque soit
le phytodistricts. Seulement sept espèces sur un total de 63 inventoriées, T. indica non incluse, sétaient révélées
connées aux termitières. Ces résultats suggèrent que les termitières ne sont pas un facteur contrôlant létablissement des
de T. indica et de la plupart des espèces quelles abritent. Lanalyse comparative de la végétation sest donc révélée non
sufsante pour trancher sur lordre détablissement des termitières et T. indica. Indépendamment de lordre
détablissement, lassociation tamarinier-termitière peut être bénéque pour les tamariniers et les termitières: les
termitières pourraient contribuer à atténuer le stress hydrique sur les tamariniers dans un environnement de plus en plus
aride alors que les tamariniers pourraient procurer de la nourriture aux termites. La prise en compte dinformations
relatives aux préférences alimentaires et à lécologie des espèces de termites concernées pourraient aider à améliorer
létat des connaissances sur cette relation plante-insecte.
Mots-clefs: relation plantes-insectes; termitières; espèces indicatrices; districts phytogéographiques
Introduction
Termite mounds (termitaria) are among the most
conspicuous elements of many tropical ecosystems,
especially in African savannah landscapes (Konaté et al.
1999). These particular social insect-made microhabitats
have been mentioned to be safe refuges for numerous
plant species (Harris 1966; Arshad 1982; Timberlake and
Childes 2004). How termitaria affect spatial distribution
*Corresponding author. Email: bfandohan@gmail.com
Acta Botanica Gallica: Botany Letters
Vol. 159, No. 3, September 2012, 345355
Société botanique de France
ISSN 1253-8078 print/ISSN 2166-3408 online
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and spatio-temporal dynamics of vegetation has been an
important question in plant ecology (Fanshawe 1968;
Malaisse and Anastassiou-Socquet 1977; Belsky et al.
1983; Konaté et al. 1999). Termitaria offer good living
conditions to plant species as they provide protection
from re, have well-drained and often deep soils, higher
soil moisture (Konaté et al. 1999) and higher soil
fertility than in the surrounding soils (Salick, Herrera,
and Jordan 1983; Tano 1993; Wood 1996). Recent
studies have evidenced positive inuence of termitaria
on plant community composition, structure, diversity and
recruitment (Traoré et al. 2008a, 2008b). Termitaria have
also been reported to be browsing hotspots for
mega-herbivorous such as large ungulates (Loveridge
and Moe 2004; Mobæk, Narmo, and Moe 2005).
Tamarindus indica L. (Fabaceae) is a monotypic
genus tree species that originated in Africa and Asia
which now occurs pantropically (Diallo et al. 2007). In
sub-Sahara Africa, this wind-resistant and long-living
tree thrives in arid and semi-arid ecosystems and is
associated with termitaria (Fandohan et al. 2010). Since
the previous century, T. indica has attracted increasing
attention as a plant associated with Macrotermes
termitaria in savannah ecosystems (Ouédraogo 1997;
Traoré et al. 2008a). It has been suggested that the
speciesafnity with termitaria is linked to its
preference for slightly loamy and well-aerated soils
(El-Siddig et al. 2006). However, this hypothesis has
never been directly tested.
On the other hand, an ethno-ecological study
reported that according to local communities, the
occurrence of tamarind individuals precedes that of
termitaria (Fandohan 2007). According to the latter
author, the afnity between tamarind and termitaria may
be explained by two reasons: rst, a tamarind tree
shadow creates a specic hygrometry (especially during
drought periods), which is appreciated by termites;
second, termites feed on tamarind fruits pulp and shells,
and may thus want to live near a sure food source.
The importance of thermoregulation systems for
social insects has been discussed in several studies
(Banschbach, Levit, and Herbers 1997; Korb and
Linsenmair 1999, 2000; Starks et al. 2004). It has also
been reported that the fungus-cultivating Macrotermitinea
(the subfamily that Macrotermes allies to) have
elaborated a thermoregulation mechanism within their
nests to yield a constant nest temperature of 30 °C and
humidity near saturation year round (Wood and Thomas
1989). In the absence of such thermoregulation nest
temperature would exponentially increase with ambient
temperature (Korb and Linsenmair 1999). Hence, the
association of termites with tamarind or another tree
species in dry ecosystems may also be seen as
adaptation strategy of termites, which have to maintain
an optimum thermoregulation system in their nests.
Despite all this research, it is still not clear which
comes rst, tamarind or termitarium? Information on the
specic plant communities associated with T. indica and
Macrotermes termitaria and how the latter communities
change with phytogeographical districts (phytodistricts)
may yield insight into (1) the relationship between plant
species associated with Macrotermes termitaria, (2) the
driving forces linking plants to these termitaria and, (3)
contribute to discuss uncertainties related to the
establishment order of these termitaria and their
associated plant communities.
A common research pathway in community analysis
consists to detect and describe the value of different
species for indicating environmental conditions. If
differences in environmental conditions are divided in
discrete groups of sample units, then Dufrene and
Legendres (1997) method of calculating species indicator
values provides a simple intuitive solution to detect
communities. This latter method combines information on
the concentration of species abundance and the
faithfulness of presence of a species in a particular
vegetation type. It produces indicator values for each
species in each vegetation type (McCune, Grace, and
Urban 2002).
The main objectives of the present study were to (1)
assess species richness and oristic composition of the
vegetation associated with Tamarindus indica on termitaria;
and (2) examine whether the latter information can provide
any information on the establishment order of Macrotermes
termitaria and the plant species they host. Understanding
the relationship between T. indica and Macrotermes
termitaria (thus answering, whether tamarind tree establish
before the termitarium or reversely) may provide relevant
information as far as conservation actions are concerned.
For instance, elucidating whether the association of
termitaria with T. indica underlies Macrotermes
termitaria-dependent establishment of T. indica or rather a
T. indica-dependent establishment of Macrotermes may be
critical to set conservation actions for both species.
The following research questions were addressed:
Are the species found on tamarind termitaria conned to
these habitats? Does the oristic composition of
termitaria associated with tamarind change according to
chorological zones? Do termitaria allow some species
that are typical for wetter chorological zone to establish
in a drier chorological zone? On the basis of previous
studies (see Fleming and Loveridge 2003) that stated
that termitaria vegetations vary with chorological
regions, we hypothesised that woody and liana plant
species associated with tamarind on termitaria are rather
dependent upon phytodistricts than termitaria.
Material and methods
Study area
The study was conducted in Benin (West Africa) in two
contrasting chorological regions where T. indica was
previously identied to occur naturally (Adomou, Sinsin,
and van der Maesen 2006): the Sudanian region
(9°45-12°25N) and the Sudano-Guinean transition
region (7°30-9°45N).
346 B. Fandohan et al.
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Study design and data collection
The study was put into place in four phytodistricts
which according to Adomou, Sinsin, and van der
Maesen (2006) differ for ecological conditions (Figure 1,
Table 1): in the Sudano-Guinean transition region, we
sampled the Zou and the Bassila phytodistrict; in the
Sudanian region, the two sub-branches of the
phytodistrict of Pendjari-Mekrou were separately
considered because of marked differences in annual
rainfall and temperature range (see Table 1).
Ten tamarind trees standing near a mound, with
diameter at 1.30 m above ground (D
130
) greater than
30 cm were randomly selected per phytodistrict. The
surface area of each Macrotermes mound associated with
each selected tamarind tree was considered as the
termitarium to be investigated. For each selected
termitarium, a circular plot of 5 m radius was laid
out in the adjacent area, at least 10 m away from the
base of the termitarium and where no perceptible
termitarium-building activity was observed. All woody
and liana plant species observed on each termitarium and
within its adjacent plot were recorded separately by
means of oristic relevés. The relevés were carried out
focusing on presence-absence and abundance (number of
individuals) data of recorded species.
Data processing
Species richness and dissimilarity between phytodistricts
A dataset based on 80 relevés (twenty per phytodistrict)
and 63 species was used to explore the degree of
distinctiveness of woody and liana plants associated with
termitaria and their adjacent areas, in each chorological
region. For this purpose, we used a presence-absence
matrix built on all species recorded according to
termitaria, adjacent plots and chorological regions.
Jaccards Index of similarity of was computed as follows:
Pj¼c
aþbcð1Þ
where P
j
is Jaccards community coefcient, ais number
of species present in community A, bis the number of
species in community B, and cis the number of species
shared by A and B. This index is a good measure of
similarity for presence-absence data (Adomou, Sinsin,
and van der Maesen 2006). In addition, all recorded
species were identied according to their chorological
status i.e. the endemism centre to which they belong after
White (1983). Botanical nomenclature followed Lebrun
and Stock (1991-1997). Presence/absence data of all
recorded species were assembled in a binary matrix and
submitted to multidimensional scaling to map relevés
according to their species composition. The
DECORANA procedure of PC-ORD software was used
to build a bidimensional plan based on the observed
dissimilarity among relevés.
Figure 1. Location of the study sites.
Figure 1. Localisation des sites détude.
Table 1. Characteristics of the study four chorological regions [climatic data source: Hijmans et al. (2004)].
Tableau 1. Caractéristiques des quatre régions chorologiques détude [source de données climatiques: Hijmans et al. (2004)].
Sudano-Guinean region Sudanian region
Phytodistrict of Zou Phytodistrict of Bassila Sub-Phytodistrict of Mekrou Sub-Phytodistrict of Pendjari
Location N8°2-E1°4N7°5-E2°2N12°06-E2°54N11°25- E1°30
Rainfall (mm) 1164 1280 675 875
Temperature (°C) 18.3-35.9 20.1-32.8 17.1-42.1 18.6-39.3
Relative humidity (%) 31-98 31-98 18-99 18-99
Climate type Tropical Tropical Tropical Tropical
Legend: phytodistrict = phytogeographical district.
Légende: phytodistrict = district phytogéographique.
Acta Botanica Gallica: Botany Letters 347
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Identication of indicator species according to
chorological regions
The approach used, assumes that two or more a priori groups
of sample units exist (here, phytodistricts), and that species
abundance values (i.e. overall number of individuals) were
recorded in each of the sample units. For each species, the
following steps were used in the implementation of
PC-ORD (McCune, Grace, and Urban 2002):
First step: the software rst computed the
proportional abundance of a particular species in a
particular group relative to the abundance of that species
in all groups;
say: A= sample unit xspecies matrix; a
ijk
=
abundance of species jin sample unit (SU) iof group k;
nk = number of sample units in group k;g= total number
of groups; mean abundance X
kj
of species jin group kis
calculated as:
Xkj ¼Pn1
i¼1aijk
nk
ð2Þ
whereas the relative abundance RA
kj
of species j in
group k is calculated as:
RAjk ¼Xkj
Pg
k¼1Xkj
ð3Þ
Second step: the proportional frequency of the
species in each group (i.e. the percentage of sample
units in each group that contain that species) was
computed:
A was rst transformed to a matrix of
presence-absence, B:
bij ¼a0
ij ð4Þ
then relative frequency RF
kj
of species j in group k was
calculated as:
RFkj ¼Pn1
i¼1bijk
nk
ð5Þ
Third step: the two proportions calculated in steps 1
and 2 were then combined by multiplying them. The
result was expressed as a percentage, yielding an
indicator value IV
kj
for each species jin each group k.
IVkj ¼RAkj RFkj 100:ð6Þ
Fourth step: the highest indicator value (IV
max
) for a
given species across
groups was saved as a summary of the overall
indicator value of that species.
Fifth step: statistical signicance of IV
max
was
determined using 4999 randomisations in a Monte Carlo
test. The probability of type I error was based on
the proportion of randomisations that the IV
max
from the
randomised data set equals or exceeds the IV
max
from the
actual data set. The null hypothesis was that IV
max
is no
larger than would be expected by chance (i.e. that the
species has no indicator value).
The indicator values range from zero (no indication)
to 100 (perfect indication). Perfect indication means that
presence of the species points to a particular group of
relevés without error at least with the data set in hand
(McCune, Grace, and Urban 2002).
Results
Species richness and oristic composition
Sudano-Guinean region
Twenty three (23) species, 21 genera and 14 families,
were recorded on termitaria in Zou phytodistrict,
whilst 22 species 20 genera and 13 families were
recorded on their adjacent plots (Figure 2). On the
other hand, 20 species, 19 genera and 13 families
Figure 2. Species, genus and family richness according to study sites.
Legend: + indicates relevés on termitaria.
Figure 2. Richesse en espèces, genres et familles par site détude.
Légende: + indique les relevés sur les termitières.
348 B. Fandohan et al.
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were recorded on termitaria in Bassila phytodistrict,
whereas 17 species, 16 genera and 10 families were
recorded on their adjacent plots (Figure 2). Overall,
Fabaceae (20%) Sterculiaceae (20%) Sapotaceae (10%)
and Combretaceae (10%) were the best-represented
families in this region. Ebenaceae and Capparaceae
were conned to termitaria.
Sudanian region
On the one hand, 30 species, 25 genera and 17 families
were recorded on termitaria in Mekrou sub-phytodistrict,
while 36 species, 29 genera and 17 families were recorded
on their adjacent plots (Figure 2). On the other hand, 35
species, 31 genera and 20 families were recorded on
termitaria in Pendjari sub-phytodistrict, whereas, 25
Table 2. Index of similarity (P
j
in percentage) among termite mounds associated with tamarind and adjacent areas, from the study
different chorological regions.
Tableau 2. Indice de similarité (P
j
en pourcentage) des termitières associées aux tamariniers et leurs zones adjacentes, dans les
différentes régions chorologiques détude.
Phyt-Zo+ Phyt-Zo Phyt-Ba+ Phyt-Ba Phyt-M+ Phyt-M Phy-P+ Phyt-P
Phyt-Zo+
Phyt-Zo 95.45
Phyt-Ba+ 41.38 37.93
Phyt-Ba 32.14 33.33 78.94
Phyt-M+ 15.91 13.64 17.07 15.79
Phyt-M 21.74 22.22 17.78 19.51 46.51
Phy-P+ 27.91 25.58 30.10 29.73 44.19 34.00
Phyt-P 24.32 25.00 26.47 30.00 29.27 38.88 67.65
Legend: phyt = phytogeographical district; Ba = Bassila, M = Mekrou, P = Pendjari, Zo = Zou; + indicates relevés on termite mounds.
Figure 3. Projection of the relevés in system axes 1 and 2. Stress = 0.085; R-square (%) = 95.38. V1: relevés in adjacent areas
from Mekrou sub-phytodistrict; V2: relevés from Mekrou (only termitaria relevés) and Pendjari sub-phytodistricts; V3: relevés from
Bassila and Zou phytodistricts.
Figure 3. Projection des relevés dans le système axes 1 et 2. Stress = 0,085; R-carré (%) = 95,38 V1: relevés dans les zones
adjacentes dans le sous-phytodistrict de la Mékrou; V2: relevés des sous-phytodistrict de la Mékrou (uniquement les relevés sur
termitières) et de la Pendjari; V3: relevés des phytodistricts de Bassila et du Zou.
Acta Botanica Gallica: Botany Letters 349
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species, 24 genera and 16 families where recorded in their
adjacent plots (Figure 2). Overall, Fabaceae (22%)
Combretaceae (14%) and Rubiaceae (10%) were the
best-represented families in this region. Ebenaceae,
Opiliaceae and Capparaceae were conned to termitaria.
Pattern of dissimilarity
Within each phytodistrict, vegetations on termitaria
weakly differed from those in adjacent plots (46 < P
j
<
96; Table 2). By contrast, there were signicant
dissimilarities (15 < P
j
< 30) between species associated
with tamarind termitaria for all pairs of chorological
regions. However, oristic composition of vegetations on
termitaria in geographically close phytodistricts were
found to be similar (41 < P
j
< 45).
Multidimensional scaling (Figure 3) was congruent
with similarity analysis and discriminated relevés
according to their phytodistrict of origin. Only relevés
from the Mekrou sub-phytodistrict showed a clear
distance between termitaria and adjacent areas.
In Zou phytodistrict (Sudano-Guinean transition
region), four species belonging to the wetter
Guineo-Congolian endemism centre were recorded on
termitaria and adjacent areas (Cola gigantea, Cola
millenii, Flacourtia avescens and Pouteria alnifolia). In
Bassila phytodistrict (Sudano-Guinean transition region),
only one species from the Guineo-Congolian endemism
centre (Capparis viminea) was found to occur only on
termitaria whereas two other species from the same
endemism centre (Flacourtia avescens and Mimusops
kummel) were recorded on both termitaria and adjacent
areas. By contrast, in Mekrou and Pendjari
sub-phytodistricts, no species from the wetter
Guineo-Congolian endemism centre were recorded neither
on termitaria nor on adjacent plots.
Indicator species of termitaria associated with
T. indica with respect to chorological regions
Indicator values and results of the randomisation test are
shown in Table 3. Zanthoxylum zanthoxyloides,Diospyros
mespiliformis and Lannea barteri were the best signicant
indicator species of the vegetation on termitaria associated
with T. indica in Zou phytodistrict (25 < IV < 59; 0.0002
p< 0.02) whereas Acacia sieberiana,Ceiba pentandra,
Cissus populnea and Pterocarpus erinaceus were found to
be characteristic of adjacent vegetations (19 < IV < 24;
0.002 p< 0.02). Only D. mespiliformis was found to be
conned to termitaria.
Capparis viminea and Mimusops kummel were
indicators of the vegetation on termitaria in Bassila
phytodistrict (27 < IV 100; p < 0.0003) while
Gymnosporia senegalensis, Pseudocedrela kotschyi,
Berlinia grandiora, Terminalia macroptera and Albizia
zygia were seen to be characteristic of adjacent
vegetations (26 < IV 50; 0.0001 < p < 0.02). C. viminea
and D. mespiliformis were exclusively recorded on
termitaria.
Capparis sepiaria, Cissus quadrangularis, Allophylus
africanus, Opilia amentacea were indicators of the
vegetation of termitaria in the Mekrou sub-phytodistrict
(IV = 50; p = 0.0002) whereas Bombax costatum,
Combretum glutinosum, Combretum nigricans, Guiera
senegalensis, Mimosa pigra and Albizia chevalieri were
found to be characteristic of adjacent vegetations (49
IV < 72; 0.0002 p0.0004). A. africanus,
C. sepiaria, C. quadrangularis, D. mespiliformis, and O.
amentacea were exclusively recorded on termitaria.
Feretia apodanthera and Grewia bicolor were
indicators of the vegetation of termitaria in the Pendjari
sub-phytodistrict (58 < IV < 68; p= 0.0002) while
Borassus aethiopum,Cassia sieberiana,Paullinia
pinnata,Daniellia oliveri and Anogeissus leiocarpa were
the most signicant indicator species of the adjacent
vegetation (20 < IV < 50; 0.001 p0.0002). A.
africanus,C. sepiaria,C. quadrangularis,G. bicolor,
and D. mespiliformis were conned to termitaria.
Except for T. indica itself, D. mespiliformis was the
only one species recorded on all termitaria, regardless of
phytodistricts. A vicariance was also observed between C.
sepiaria recorded on termitaria in the sub-phytodistricts
of Pendjari and Mekrou (Sudanian region) and C. viminea
recorded on termitaria in the Bassila phytodistrict
(Sudano-Guinean region).
Discussion
From this study, it appears that whatever the
phytodistrict, vegetations associated with termitaria are
much similar to those of their adjacent areas. Overall,
only seven species (not including T. indica) out of a
total of 63 recorded were found to be conned to
termitaria. These ndings would suggest termitaria to not
be a factor controlling establishment of T. indica and
most the species their host.
However, results also show that the specic
composition of the vegetation associated with T. indica on
termitaria varies between phytodistricts, which is
consistent with our hypothesis. Observed variations reect
differences between original native chorological types.
Such chorological variation is consistent with results from
previous studies on termitaria vegetations (Glovers,
Trump, and Wateridge 1964; Fanshawe 1968; Ouédraogo
1997; Fleming and Loveridge 2003). From our results, the
number of plant families conned to termitaria seemed to
increase with drier conditions. This is congruent with the
hypothesis that under a given climatic condition,
termitaria would appear as factors inuencing species
diversity and oristic composition (Traoré et al. 2008a).
The association between tree species and termitaria
may vary with environmental conditions. In the present
study, T. indica was recorded on 5% to 50% of plots in
adjacent areas, irrespective of chorological regions. A
previous survey by Fandohan (2007), reported that up to
90% of tamarind trees in savannahs were found on
350 B. Fandohan et al.
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Table 3. Monte Carlo test of signicance of observed maximum indicator values (IV) for each species, based on 4999 randomisations.
Tableau 3. Test de signication de Monte Carlo des valeurs indicatrices maximales observées (IV) pour chaque espèce, basées sur 4999 randomisations.
IV from de Randomisation groups
Indicator species Districts Chorological types Observed IV Mean Std p
Zanthoxylum zanthoxyloides (Lam.) Zepernick & Timber Phyt-Zo+ SG 58.2 9.0 4.23 0.0002
Diospyros mespiliformis Hochst. ex A.DC. Phyt-Zo+ AT 25.0 13.2 2.89 0.0030
Lannea barteri (Oliv.) Engl. Phyt-Zo+ SG 24.6 11.5 3.54 0.0106
Acacia sieberiana DC. Phyt-Zo Pl-Af 23.3 13.5 2.76 0.0020
Ceiba pentandra (L.) Gaertn Phyt-Zo Pan 26.7 7.9 4.30 0.0130
Cissus populnea Guill. & Perr. Phyt-Zo SG 22.7 13.6 2.72 0.0132
Pterocarpus erinaceus Poir Phyt-Zo A-Am 19.2 9.4 4.11 0.0178
Sterculia setigera Del. Phyt-Zo AT 20.8 9.3 4.24 0.0468
Piliostigma thonningii (Schumach.) Milne-Redh. Phyt-Zo AT 18.9 10.6 3.95 0.0814
Hildegardia barteri (Mast.) Kosterm. Phyt-Zo SG 18.0 7.7 4.62 0.1008
Cola gigantea A. Chev. Phyt-Zo GC 18.0 7.8 4.60 0.1022
Cola millenii K. Schum. Phyt-Zo GC 18.0 7.7 4.64 0.1046
Pouteria alnifolia (Baker) Roberty Phyt-Zo AT 18.0 7.7 4.64 0.1046
Lannea acida A. Rich. Phyt-Zo SG 17.1 10.7 3.67 0.1324
Mitragyna inermis (Willd.) O. Kuntze Phyt-Zo SG 11.4 9.5 3.95 0.4665
Capparis viminea Hook. f. & Thoms. ex Oliv. Phyt-Ba+ GC 100.0 8.7 4.22 0.0002
Mimusops kummel Bruce ex A.DC. Phyt-Ba+ GC 27.8 8.3 4.32 0.0024
Flacourtia avescens Willd. Phyt-Ba+ AT 16.0 8.6 4.16 0.1138
Gymnosporia senegalensis (Lam.) Loes Phyt-Ba Pal 29.4 12.6 3.22 0.0002
Pseudocedrela kotschyi (Schweinf.) Harms. Phyt-Ba SG 32.3 12.2 3.34 0.0002
Berlinia grandiora (Vahl) Hutch. & Dalziel Phyt-Ba SG 50.0 10.6 3.76 0.0004
Terminalia macroptera Guill. & Perr. Phyt-Ba SZ 29.1 10.9 3.79 0.0026
Albizia zygia (DC.) J.F. Macbr. Phyt-Ba AT 26.7 7.9 4.36 0.0158
Bridelia ferruginea Benth. Phyt-Ba SG 15.0 7.9 4.34 0.1900
Combretum collinum Fresen Phyt-Ba SZ 12.3 9.4 4.09 0.3643
Vitellaria paradoxa C.F.Gaertn. Phyt-Ba S 12.4 11.9 3.43 0.6279
Stereospermum kunthianum Cham. Phyt-Ba SZ 6.0 9.7 4.13 0.9684
Allophylus africanus P. Beauv. Phyt-M+ AT 50.0 10.6 3.74 0.0002
Capparis sepiaria L. Phyt-M+ SZ 50.0 10.6 3.74 0.0002
Cissus quadrangularis L. Phyt-M+ Pl-Af 50.0 10.6 3.74 0.0002
Opilia amentacea Roxb. Phyt-M+ SZ 50.0 10.6 3.74 0.0002
Balanites aegyptiaca (L.) Del. Phyt-M+ Pl-Af 24.5 10.5 3.73 0.0124
Sclerocarya birrea (A.Rich.) Hochst.) Phyt-M+ S 10.0 10.0 0.14 1.0000
Bombax costatum Pellegr. & Vuill. Phyt-M S 51.4 8.0 4.35 0.0002
Combretum glutinosum Perr. ex DC. Phyt-M S 47.6 10.8 3.77 0.0002
Combretum nigricans Lepr. ex Guill. & Perr. Phyt-M S 71.4 9.5 4.11 0.0002
Guiera senegalensis J.F.Gmel. Phyt-M S 60.0 7.9 4.31 0.0002
Mimosa pigra L. Phyt-M Pal 50.0 10.6 3.82 0.0002
Albizia chevalieri Harms Phyt-M S 49.0 8.7 4.40 0.0004
Hannoa undulata (Guill. & Perr.) Planch. Phyt-M S 40.0 7.2 4.97 0.0010
Lannea microcarpa Engl. & K. Krause Phyt-M SZ 30.0 6.7 5.30 0.0132
(Continued)
Acta Botanica Gallica: Botany Letters 351
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Table 3. (Continued)
IV from de Randomisation groups
Indicator species Districts Chorological types Observed IV Mean Std p
Acacia seyal Del. Phyt-M Pl-Af 26.7 8.0 4.50 0.0162
Detarium microcarpum Harms Phyt-M S 22.5 7.1 4.95 0.0422
Celtis toka (Forssk.) Hepper & Wood Phyt-M SZ 20.0 6.7 4.77 0.1142
Crossopteryx febrifuga (G. Don) Benth. Phyt-M Pl-Afr 14.5 8.9 4.16 0.1772
Afzelia africana Smith ex Pers. Phyt-M AT 13.3 6.7 5.27 0.3189
Terminalia laxiora Engl. & Diels Phyt-M Pl-Af 13.3 6.8 5.34 0.3215
Combretum molle R.Br. ex G. Don Phyt-M Pl-Af 13.3 6.7 5.20 0.3233
Adansonia digitata L. Phyt-M Pl-Af 12.3 9.4 4.11 0.3575
Ziziphus mauritiana Lam. Phyt-M Pal 10.0 7.2 5.02 0.5477
Acacia macrostachya Reichenb. ex DC. Phyt-M S 10.0 7.2 5.12 0.5483
Khaya senegalensis (Desr.) A.Juss. Phyt-M S 9.0 8.6 4.19 0.5971
Feretia apodanthera Del. Phyt-P+ SZ 67.5 9.2 4.09 0.0002
Grewia bicolor Juss. Phyt-P+ SZ 58.8 10.1 3.95 0.0002
Borassus aethiopum Mart. Phyt-P Pl-Af 44.5 8.8 4.09 0.0002
Cassia sieberiana DC. Phyt-P S 50.0 10.5 3.77 0.0002
Paullinia pinnata L. Phyt-P Pan 50.0 10.5 3.77 0.0002
Daniellia oliveri (Rolfe) Hutch. & Dalziel Phyt-P SZ 32.0 7.7 4.55 0.0044
Anogeissus leiocarpa (DC.) Guill. & Perr. Phyt-P Pl-Af 20.8 13.8 2.51 0.0092
Ximenia americana L. Phyt-P Pan 14.5 8.8 4.12 0.1782
Grewia avescens Juss. Phyt-P SZ 15.0 7.9 4.25 0.1806
Vitex doniana Sweet Phyt-P SG 15.0 7.9 4.19 0.1846
Legend: The mean and standard deviation (m; sdt) of IV from the randomisations are given with p-values for the hypothesis of no differences between termitaria and adjacent areas and between phytodistricts; IV
= Indicator Value; phyt = phytogeographical district; Ba = Bassila; M = Mekrou; P = Pendjari; Zo = Zou; + indicates relevés on termite mounds; AT = Afro-Tropical; A-AM = Afro-American; GC = Guineo-
Congolian; S = Sudanian; SG = Sudano-Guinean; SZ = Sudano-Zambezian; Pal = Paleotropical; Pan = Pantropical; Pl-Af = Pluri-regional African.
Légende: La moyenne et lécart type (m; sdt) des IV issues des randomisations sont données avec les valeurs de ppour lhypothèse nulle dabsence de différence entre les termitières et leurs phytodistricts; IV =
Valeur Indicatrice; phyt = district phytogéographique; Ba = Bassila; M = Mekrou; P = Pendjari; Zo = Zou; + indique les relevés sur les termitières; AT = Afro-Tropicale; A-AM = Afro-Americaine; GC = Guinéo-
Congolaise; S = Sudanienne; SG = Sudano-Guinéenne; SZ = Sudano-Zambézienne; Pal = Paléotropicale; Pan = Pantropicale; Pl-Af = Pluri-régionale Africaine.
352 B. Fandohan et al.
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termitaria while only 24% of tamarind trees in riparian
forests were found on termitaria. This observation would
suggest that T. indica tends to be associated with
termitaria under more severe drought conditions and
contrasts with ndings of Traoré et al. (2008a) who
argued that T. indica is always found on termitaria.
Possibly, wetter species habitats (i.e. riparian forest) were
not included in the sampling design of the latter study.
Furthermore, typical Guineo-Congolian species such as
C. viminea and M. kummel may need a critical reduction
in drought stress (soil) in order to be able to live under
the Sudano-Guinean climatic conditions where they were
recorded during our survey. This is consistent with the
hypothesis that termitaria contribute to mitigate drought
stress in tree species (Konaté et al. 1999).
Despite the similarity between termitaria and the
vegetation in their immediate vicinity (adjacent areas),
results from the indicator species analysis suggest that
important differences exist between species characterising
each habitat type (i.e. termitaria versus adjacent areas)
within each phytodistrict. The connement of seven
species i.e., Allophylus africanus,Capparis sepiaria,
Capparis viminea,Cissus quadrangularis,Opilia
amentacea,Grewia bicolor and Diospyros mespiliformis
to termitaria as well as the vicariance observed between
C. sepiaria and C. viminea suggests that termitaria may
provide suitable habitats for these species (i.e., specic
soil conditions requirements), as previously suggested by
Traoré et al. (2008a). Indeed, except for D. mespiliformis,
all these species are shrubs or lianas and typical of
tropical Africa thickets, known as vegetation type that
depends on soil conditions (White 1983).
The consistency of presence of D. mespiliformis on
all sampled termitaria may be either because D.
mespiliformis and T. indica share the same dispersal
agents (i.e. primates) or because the dispersal agents of
one species also use the other species as a feeding roost.
In both cases, probable dispersal agents are Papio anubis,
Erythrocebus patas and Cercopithecus aethiops that are
known to feed on T. indica and D. mespiliformis in
savannah-forest mosaics of West Africa (Kunz and
Linsenmair 2008a, 2008b; B. Fandohan, pers. obs.). In
addition, a species such as Grewia bicolor has been
reported to grow under T. indicas shadow in the
Sudano-Zambezian chorological region (Nyadoi 2004).
Such consistency of association across phytodistricts or
chorological regions may also suggest that rather than the
termitaria, the dispersal agents may be the driving force
bringing them together. This is congruent with previous
studiesresults that reported T. indica and many Grewia
species (e.g. G. saligna, G. calvata, G. humbertii, G.
grevei and G. leucophylla) to be fed on by primate
species such as Lemur catta and Eulemur fulvus in
Madagascar (Simmen, Hladik, and Ramasiarisoa 2003).
According to Hubbell (2001), organisation of
species assemblages in communities may be viewed as
resulting from two main kinds of processes:
niche-assembly rules and dispersal assembly rules. By
niche assembly, it is meant that speciesadaptation to
particular ecological niches determines the particular
species composition and organisation of a community,
following deterministic processes (Hardy and Sonké
2004). On the contrary, the organisation of a
dispersal-assembled community results from localised
dispersal events and local demographic stochasticity,
following unpredictable processes (Hubbell 2001; Hardy
and Sonké 2004). However, in cases two or more plant
species share the same dispersal agents, their
assemblage would rather be deterministic than
stochastic.
From our study, it is still not clear whether tamarind
comes before the termitarium or reversely. Available
literatures on establishment order of termitaria and plant
species have yielded various ndings. Some studies
highlighted that some woody/liana plants are only found
on termitaria (i.e. Tamarindus indica,Boscia
senegalensis,Cadaba farinosa,Capparis sepiaria,
Maerua angolensis and Diospyros mespiliformis),
suggesting that termitaria came rst and favoured the
establishment of the plant species (Ouédraogo 1997;
Traoré et al. 2008a). It has also been suggested that the
presence of some woody plants on termitaria (i.e.
tamarind trees) may result from some viable seeds carried
and left inside termitarium cavities by rodents (Alexandre
2002). Results in this study suggest that T. indica is not
always found on termitaria but conrm the
above-mentioned assumptions for species such as
C. sepiaria and O. amentacea.
However, before exclusively associating the
establishment of a species with termitaria, its seedlings
should be observed on termitaria with no adult
individuals nearby (at least, for barochore and
root-sucking species such as T. indica). Up to this date,
no study has recorded such observation for T. indica
despite Traoré et al. (2008b) having argued for the role
of termitaria in a number of tree speciesregeneration.
For species such as D. mespiliformis,G. bicolor and
C. sepiaria, it is common to observe seedlings and
saplings on termitaria with no mature individuals nearby
(B. Fandohan, pers. obs.). Contrastingly, for T. indica,if
on the one hand adult individuals are rarely seen out of
termitaria in savannah habitats, on the other hand,
tamarind seedlings are also hardly seen standing on
termitaria with no mature tamarind nearby (B. Fandohan,
pers. obs.).
Finally, we conclude that comparative vegetation
analysis is not enough to stand on termitaria and T.
indica establishment order. Termite-tamarind relationships
may be symbiotic. On the one hand whether they come
rst or not, Macrotermes may benet from tamarind trees
for their thermoregulation and feeding needs (Fandohan
2007). In addition, some observations in tiger bush in the
Sahel would suggest that termitaria become non-active
when the overhead vegetation disappears. This may
suggest that it is the presence of trees that favour
termitesactivities (B. Sinsin, pers. Comm.). On the other
Acta Botanica Gallica: Botany Letters 353
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hand, there are indications that alternating deterioration
and rebuilding of Macrotermes termitaria may bare roots
leading to coppicing of which additional stems could
grow from the base of tamarind trees (B. Fandohan, pers.
obs.). Coppicing can result in the longevity of a plant
greatly exceeding the age of individual original trunks
(Peakall et al. 2003). These termitaria may thus
contribute to the protection and long-term conservation of
tree species in the continuously degrading environment of
arid and semi-arid ecosystems. Besides, as previously
suggested, termite effect on soils may also offer to plants
well-drained and deep soils with higher moisture and
nutrients (Konaté et al. 1999; Traoré et al. 2008b). To
better elucidate the relationship between plant and termite
species in general and particularly T. indica and termite
species, it is necessary to move from vegetational/
phytosiciological perspective to insect-plant
co-evolutionary and/or insect-plant dependency or
co-dependency perspective. In view of these perspectives,
integration of information on termite speciesfeeding
preferences and ecology may help to improve current
state of knowledge on this plant-insect relationship.
Acknowledgements
We are very grateful to Fulani herders and Gourmantché
women for their help during eld surveys. We also thank Aida
Cuni Sanchez and the reviewers for their comments on a
previous version of the manuscript.
Notes on contributors
B. Fandohan holds a PhD in natural resources
management / conservation biology and is currently a
post doctoral research fellow at the International
Ecosystem Management Partnership of the United
Nation Environment Programme (IEMP-UNEP). He
conceptualised, designed and performed the eld work,
analysed the data and wrote the manuscript.
A.E. Assogbadjo, holds a PhD in applied biological
sciences and is a senior lecturer at the University of
Abomey Calavi (Benin Republic). He read and improved
the manuscript.
V.K. Salako, holds a MSc in natural resources
management and is a research assistant in the laboratory
of applied ecology (Benin Republic). He read and
improved the manuscript.
P. van Damme, Chair of tropical, subtropical agriculture
and ethnobotany at Ghent University (Belgium). He
co-supervised the work, read and improved the
manuscript.
B. Sinsin, Chair of tropical ecology and is currently the
rector of the University of Abomey calavi (Benin
Republic). He supervised the work, read and improved the
manuscript.
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... In the course of their foraging and building activities, workers of Macrotermes subhyalinus Rambur (Termitidae, subfamily Macrotermitinae) directly or indirectly rearrange huge amounts of soil, in which they concentrate nutrient content deeply improved and stock in form of mounds above soil surface commonly considered as "heterogeneity drivers" (Holt & Lepage, 2000;Moe et al., 2009;Jouquet et al., 2011;Muller & Ward, 2013;Davies et al., 2016). Such fertile microhabitats scattered and available in savanna contribute to enhance safe ecological niches for plant species and for other organisms, and as rooting niche in where varied woody species occur, establish, develop and coexist for creating termite mound savanna despite species biogeographical origin as Sudanian, Guinea or Sahelian species and ecophysiological characteristics as fire sensitive species (Mobaek et al., 2005;Moe et al., 2009;Fandohan et al., 2012;Okullo & Moe, 2012). Such microhabitats likely seem to provide fundamental attribute to seedlings of these woody species and then facilitate their establishment and recruitment into mature trees as specific response. ...
... Plant communities on termitaria were different from agricultural lands (field and fallows) to the National Park (Dossou-Yovo et al., 2016). Fandohan et al. (2012) suggested termitaria as a factor used in controlling the establishment of Tamarindus indica and most of the plant species they host, although the termitaria-tamarind associations may be profitable to both termites and tamarind trees. They added that, termitaria may help to mitigate drought on tamarind trees while the trees in turn may offer food to termites. ...
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Findings are available on termitaria and their vegetation in the Pendjari Biosphere Reserve and other Sudanian regions in West Africa, but research without dissemination and impacts on communities seems not to be useful. This work aims at providing non-governmental organization (NGOs) and forestry advisers with useful data for environmental education projects and taking termitaria and their vegetation into account for ecotourism in Pendjari Reserve. This article on termitaria and termitaria-related vegetation summarizes data useful for two purposes. Traditional knowledge on termitaria is useful for education; termitaria plants are used as medicine. Mushrooms growing on termitaria and small mammals living in dead and abandoned mounds are consumed in the reserve. There is a need to train kids and students on termitaria and their vegetation conservation. The panoramic view of mounds and their vegetation is really attractive to tourists. Vegetation on termitaria differs between management types of an area and is dominated by woody species belonging mostly to Combretaceae botanical group. Cappareae species seem restricted to termitaria. The three major ethnic groups in the Reserve hold a diversity of ethnological knowledge on termitaria and their vegetation. These can serve for ecotourism development towards termitaria to lower poverty probability of small households in the Reserve.
... Two of them, namely Mekrou-Pendjari and Plateau, are composed each of two sub-phytodistricts which are geographically separated, the Mekrou-Pendjari into Mekrou and Pendjari sub-phytodistricts and Plateau into East Plateau and West Plateau sub-phytodistricts ( fig. 1, Adomou et al. 2006). A recent study showed that at certain scale these subphytodistricts could differ as far as the vegetation pattern is concerned (Fandohan et al. 2012). Taking this into account, we considered them as separate entities i.e. eight phytodis- tricts and four sub-phytodistricts were considered here (for simplicity all named phytodistricts). ...
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Background and aims – Borassus aethiopum Mart. is a wild palm species with high subsistence importance in West Africa. Extensive agriculture and overharvesting of its stem and fruits for multiple uses have caused a decrease of its natural populations in its native range. For conservation purposes, the distribution, abundance and structural diversity of the species were investigated across ten phytodistricts in three biogeographical zones in Benin. Two hypotheses were tested (i) tree floristic composition of B. aethiopum natural habitat changes with phytodistricts and (ii) structural diversity of B. aethiopum changes with phytodistricts, both as potential adaptation strategies to changing ecological conditions. Methods – Geographical coordinates of the species occurrence were recorded. Abundance was assessed in 852 one-ha plots. Structural diversity was studied using structural indices on data from ecological inventories and neighbourhood survey in 70 one-ha plots. Key results – The two hypotheses proved true. B. aethiopum was found in all phytodistricts but with strong variations in abundance. Overall, floristic composition of its natural habitats showed dissimilarities among phytodistricts. Three main vegetation types sheltered B. aethiopum: mixed grass and shrub savannas, savanna woodlands and woodlands, all of which were found in gallery forest landscapes. The density of B. aethiopum was lower in grass savannas but larger, shorter and distant individuals were found there than in savanna woodlands and woodlands. In the latter vegetation types, its density was high with thin, tall and closely spaced individuals. B. aethiopum tolerates mingling with several other tree species but increased mingling tends to lead to positive differentiation in diameter and height. Conclusions – Borassus aethiopum is a sun-demanding species and establishes successful populations in various ecological conditions. It could be mixed with other tree species in tree plantations and modern agroforestry systems as long as water requirements are met. However, it would be preferable that the species is associated with shade tolerant or medium sun-demanding species.
... Two of them, namely Mekrou-Pendjari and Plateau, are composed each of two sub-phytodistricts which are geographically separated, the Mekrou-Pendjari into Mekrou and Pendjari sub-phytodistricts and Plateau into East Plateau and West Plateau sub-phytodistricts ( fig. 1, Adomou et al. 2006). A recent study showed that at certain scale these subphytodistricts could differ as far as the vegetation pattern is concerned (Fandohan et al. 2012). Taking this into account, we considered them as separate entities i.e. eight phytodis-P r o o f s tricts and four sub-phytodistricts were considered here (for simplicity all named phytodistricts). ...
Article
Background and aims – Borassus aethiopum Mart. is a wild palm species with high subsistence importance in West Africa. Extensive agriculture and overharvesting of its stem and fruits for multiple uses have caused a decrease of its natural populations in its native range. For conservation purposes, the distribution, abundance and structural diversity of the species were investigated across ten phytodistricts in three biogeographical zones in Benin. Two hypotheses were tested (i) tree floristic composition of B. aethiopum natural habitat changes with phytodistricts and (ii) structural diversity of B. aethiopum changes with phytodistricts, both as potential adaptation strategies to changing ecological conditions. Methods – Geographical coordinates of the species occurrence were recorded. Abundance was assessed in 852 one-ha plots. Structural diversity was studied using structural indices on data from ecological inventories and neighbourhood survey in 70 one-ha plots. Key results – The two hypotheses proved true. B. aethiopum was found in all phytodistricts but with strong variations in abundance. Overall, floristic composition of its natural habitats showed dissimilarities among phytodistricts. Three main vegetation types sheltered B. aethiopum: mixed grass and shrub savannas, savanna woodlands and woodlands, all of which were found in gallery forest landscapes. The density of B. aethiopum was lower in grass savannas but larger, shorter and distant individuals were found there than in savanna woodlands and woodlands. In the latter vegetation types, its density was high with thin, tall and closely spaced individuals. B. aethiopum tolerates mingling with several other tree species but increased mingling tends to lead to positive differentiation in diameter and height. Conclusions – Borassus aethiopum is a sun-demanding species and establishes successful populations in various ecological conditions. It could be mixed with other tree species in tree plantations and modern agroforestry systems as long as water requirements are met. However, it would be preferable that the species is associated with shade tolerant or medium sun-demanding species.
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Termite mounds represent abundant microhabitats of high biodiversity in tropical savanna ecosystems and are an important source of landscape heterogeneity in Sub–Saharan West Africa. Floristic composition as well as density, structure and zonation of plant cover on the mounds were investigated in northern Benin and compared to the adjacent savanna vegetation. A total of 57 abandoned and densely vegetated termite mounds of comparable size and similarly affected by erosion located in different types of savannas inside and outside of the W National Park and in cotton fields were studied. This study revealed that termitaria are special habitats differing in density, composition and structure from surrounding savannas. The plant cover of termite mounds showed a distinctive zonation. Succulents, geophytes, and lianas were much more abundant on mounds, the family Capparaceae was found exclusively on mounds. The floristic composition and vegetation on termitaria proved to be rather homogeneous; although those mounds located in cotton fields differed by higher abundance of Poaceae and lower species richness.
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Heat shielding is a recently identified mechanism used by worker honey bees (Apis mellifera) to help maintain constant hive temperatures. Only workers perform this behavior; in our experiment, drones actively avoided heated hive regions. Observations of marked day-old cohorts within broodcomb regions indicate that heat shielding is performed by young bees to preferentially protect advanced stage larvae and pupae. As expected, the number of heat-shielders significantly increased with both the temperature of the heat source and the size of the colony. Of the young bees observed to perform the behavior, those aged 12--14 days were significantly more likely to heat-shield than expected. Combined, these data suggest that classifications of age-based tasks in honey bees should include heat shielding, and that the behavior is an adaptation designed to protect temperature-sensitive brood. Copyright 2005.
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Termite population parameters and termite nest-nutrient content are compared among 3 nutrient-deficient vegetation types along the Rio Negro. Termite population parameters are positively associated with forest productivity, biomass, stature, and soil fertility. A minimum of 3-5% of total litter is consumed by termites, but high spatial variance (patchiness) of termite consumption and nutrient concentration may be much more important to forest dynamics. Termitaria form rich nutrient patches that contrast significantly with the highly weathered soils of the region. Termitaria are abandoned at a rate of 165+ or -66 nests ha-1 yr-1, providing abundant and potentially important microsites for tree-seedling establishment.-from Authors
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1. To determine whether agroforestry and silvopastoralism might be introduced more successfully into xeric or into mesic environments, the effects of isolated, mature trees of Acacia tortilis (acacia) and Adansonia digitata (baobab) on herb-aceous-layer composition and productivity, soil properties, and microclimate in a moderately mesic savanna (c. 750 mm annual rainfall) were investigated and compared with an earlier study of the effects of the same two species in a more xeric savanna (c. 450 mm annual rainfall) in Tsavo National Park (West), Kenya. 2. Similar to the more xeric site, solar radiation was reduced by 45-65% and soil temperatures were reduced by 5-12 C⚬ under both tree species. Except for early in the growing season, soil-moisture values were similar under tree canopies and in open grasslands. 3. Compared to the more xeric site, where herbaceous-layer productivity was 95% higher under trees than in the open, productivity in the mesic site was 52% higher under acacia canopies (a mean of 808 g m-2) than in the open (533 g m-2), but only 18% higher under baobab canopies (569 g m-2) than in the open (484 g m-2). 4. Concentrations of organic matter, total N, 15N, P, K and Ca were significantly higher, and C:N ratios and soil bulk density significantly lower under tree canopies than in the open at both sites. Mg concentrations were significantly higher in the open than under tree canopies at the mesic, but not the xeric, site. 5. The contrast noted in 3 above between the herbaceous layer productivity below tree canopies and that in areas beyond them was attributed higher soil-N concentrations and to reduced evapotranspiration in these N- and water-limited systems. 6. The contrast in forage production under tree canopies between xeric and mesic sites may be due to the greater importance of shade in reducing temperatures and evapotranspiration in more arid environments.