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Abundance and Diversity of Woody Undergrowth Reservoir as Indicator of Suitable Vegetation Patch for Natural Regeneration

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Open Journal of Ecology, 2022, 12, 113-132
https://www.scirp.org/journal/oje
ISSN Online: 2162-1993
ISSN Print: 2162-1985
DOI:
10.4236/oje.2022.122007 Jan. 30, 2022 113 Open Journal of Ecology
Abundance and Diversity of Woody
Undergrowth Reservoir as Indicator of Suitable
Vegetation Patch for Natural Regeneration
Saran Traoré1,2,3* , Issiaka Keïta1,2, Sébastien Ange Habih Nombré1,2,
Hassan Bismarck Nacro2, Brice Sinsin4
1UFR/de la Vie et de la Terre, Université Nazi Boni, Bobo-Dioulasso, Burkina Faso
2Laboratoire dEtude et de Recherche en Fertilité de Sols, IDR, Université Nazi Boni, Bobo-Dioulasso, Burkina Faso
3Laboratoire de Biologie et Ecologie Végétales, UFR-Sciences de la Vie et de la Terre, Université Joseph Ki-Zerbo, Ouagadougou,
Burkina Faso
4Laboratoire dEcologie Appliquée, Faculté des Sciences Agronomiques, Université d’Abomey-Calavi, Cotonou,
République du Bénin
Abstract
The diversity, abundance and height structure of naturally regenerated woody
species were analyzed and compared for undergrowth reservoir associated
with vegetation patches as bowal, shrubland and woodland for understanding
the effect of vegetation conditions
on spatial distribution of woody plants in
Sudanian ecosystem. All undergrowth individuals (height < 1.5 m) were re-
corded by species and the total height and collar diameter were measured in
50 subplots of 5 m × 5 m (or 25 m2) laid out in each vegetation p
atch during
rainy season. In total 52 undergrowth species and 2224 stems arranged in 38
genera and 21 families out of which 25, 36 and 38 species were found in bo-
wal, shrubland and woodland respectively and 15 shared species. Combreta-
ceae, Caesalpiniaceae and Mimosaceae families were the most abundant and
species density was significantly higher (P < 0.05) in shrubland and woodland
than bowal, and increase diversity in only shrubland (P = 0.002) comparing
with bowal. Undergrowth was abundantly concentrated in small height class
as canopy benefit and indicate a specific-
site distribution. Results also showed
that bowal had a particular vegetation which vary in function of tree commu-
nity characters in place (P < 0.05) while woodland and shrubland are
similar
vegetation patches. As conclusion,
the study vegetation patches form suitable
regeneration niches for the undergrowth reservoir of limited number of spe-
cies which in return can be predictor of species ric
hness and abundance and
thus the structure of tree community. Woody undergrowth community could
be considered in biodiversity management and degraded land restoration.
How to cite this paper:
Traoré, S.,
Keïta,
I
., Nombré, S.A.H., Nacro, H.B. and Sinsin
,
B
. (2022)
Abundance and Diversity of
Woody
Undergrowth Reservoir as Indica-
tor of Suitable Vegetation Patch for Natural
Regeneration
.
Open Journal of Ecology
,
12,
113
-132.
https:
//doi.org/10.4236/oje.2022.122007
Received:
December 19, 2021
Accepted:
January 27, 2022
Published:
January 30, 2022
Copyright © 20
22 by author(s) and
Scientific
Research Publishing Inc.
This work is licensed under the Creative
Commons Attribution International
License (CC BY
4.0).
http://creativecommons.org/licenses/by/4.0/
Open Access
S. Traoré et al.
DOI:
10.4236/oje.2022.122007 114 Open Journal of Ecology
Supplementary works could be undertaken to understand the characteristics
of seed bank for bowal, shrubland and woodland regarding the
undergrowth
and tree communities.
Keywords
Biodiversity Conservation, Burkina Faso, Regeneration, Spatial Heterogeneity,
Sudanian Ecosystems
1. Introduction
Restoring woody community for conservation goal with its species diversity based-
naturally occurrence in savanna and woodland ecosystems in arid and semi-arid
regions have always been of broad concerns for vegetation management by plant
ecologists, biologists, evolutionists and environmentalists. One reason is the fluc-
tuation observed in woody species composition and diversity, discontinuous struc-
ture and pattern of woody stratum within or between landscape which reflect the
magnitude and instability of species establishment and recruitment [1] [2]. Al-
though many efforts have been conceded and are increasingly underway [3] [4]
[5] [6] to overcome these concerns, the mechanisms or way in which plant spe-
cies colonize or regenerate and recruit in a given area and vegetation condition
vary largely in the literature [7] [8] [9] [10] [11]. It is reported in West Africa, in
Sudano-Sahelian ecosystems that many woody species asexually regenerate natu-
rally in the landscapes of
Daniellia oliveri
,
Detarium microcarpum
or
Isoberlinia
doka
[9] while species as
Crossopteryx febrifuga
is able to recruit and resist in a
wider range of fire conditions
than
Piliostigma
sp [8]. According to Do
et al
. [12]
undergrowth species decline in disturbed vegetation. These authors have recorded
54 woody species in high human impacted forest, 66 in low impacted and 56 in
intact forest at seedling stage against 55, 73 and 77 at sapling stage and 90, 102
and 98 at tree stage respectively and significant increase in sapling density for
high impacted forest and similar distribution in seedling density.
Obviously, in many arid and semi-arid regions, the natural vegetation is the
main source that supplies with natural resources and environment benefits for
local resident well-being [11] [13] [14]. Several species are overharvested for
household needs and others are ecologically used for land protection. The recur-
rent tree pruning of many species maintain these trees in juvenile stage while the
loss of many natural habitats generated by the constant change or transformation
of arable lands for agriculture extension and/or urbanization purposes lead to
modify the vegetation structure and conditions [15] [16] and thereafter hind their
natural regeneration by suppressing seed production. These factors also act to
reduce the size of plant population. Whenever the conditions of vegetation shape
the woody tree strata [17], the dynamic and quality of the natural undergrowth
successfully established and recruited remains the main responsible of the future
composition and structure in particular woody community. It is suggested that
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10.4236/oje.2022.122007 115 Open Journal of Ecology
the undergrowth ability to escape disturbance effects and changing environment
and therewith their performance to grow can be very considerable [7]. There-
fore, the characteristics of natural woody vegetation widely varied from site to
site in the same landscape as the clue of limited establishment and recruitment
in woody community. Subsequently, some species that escape changing environ-
ment manage to reproduce abundantly, persist and recruit in the conditions of
the new habitat and the others either decline or perish and hereafter jeopardize
the stability of local biodiversity inside the same phytogeographical edge. During
their investigations Attua
et al
. [15] have observed among study zones, the pre-
ponderance of the species of Fabaceae family in both least and medium human
activities while the species of Fabaceae, Malvaceae and Moraceae families were
most represented under higher activities.
In Sudanian Africa, strong soil compaction and moisture deficit are also fac-
tors that lead natural population regression in savanna. Therefore, quantifying
the regeneration status of woody species in these ecosystems become indisput-
able subject to understand the distribution and assemblage of species in a given
area despites the effects of ecological interactions [4]. Previous research have
highlighted that the success or fail of species to establish is guided by the prob-
ability and quality of undergrowth to establish while its recruitment is influ-
enced by habitat type or microsite characteristics and/or their interactions [10]
[18]. When scarce woody vegetation may be benefit for savanna native species
dense vegetation patch as woodland may facilitate shade tolerant and/or forest
native undergrowth species to establish [19] [12] [20] [21]. For [19] the under-
growth number of
Vitellaria paradoxa
was significantly higher under the tree
crowns of
Parkia biglobosa
and
V
.
paradoxa
than outside crown cover. Accord-
ing to [21] seedling density was higher
Eucalyptus camaldulensis
plantation for-
est (2567 stems/ha) in comparison with the native woodland (2267 stems/ha)
almost similar whereas unprotected site better promote seedling emergence with
the highest density than in protected one across phytogeographical regions [22].
Comparatively few or no attention has been addressed at understanding the
characteristics of the woody undergrowth reservoir along vegetation patches as
shrubland and woodland in the same landscape. Therefore, current investigation
was undertaken to assess the regeneration potential of woody species in secon-
dary disturbed ecosystems that could be candidate for conservation and/or res-
toration for particular woody species populations, or might be equivalent to the
same area at different times. The aim of this study was to characterize the com-
munity of woody undergrowth and the relationship between undergrowth and
trees of the same species across vegetation patches as habitats or regeneration
niches. We specifically assessed firstly, 1) the species composition and diversity
of undergrowth; 2) the density and structure of undergrowth stem; 3) whether
the species richness and stem density of undergrowth community vary in func-
tion of those of tree community within area and secondly, 4) the regeneration
status of the most shared species within area relating to the conspecific tree
community to find out the preferred regeneration niche. As research question
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“How important is undergrowth community in secondary disturbed vegetation
patch?”
2. Material and Methods
2.1. Study Site Description
The field work was carried out in a sub-urban forest in West part of Burkina
Faso at an altitude of 300 m a.s.l. This sub-urban forest is located in the north-
west of Bobo-Dioulasso the second main city of the country (11˚N - 12˚N and
10’W - 30W). The local climate is characterized by the alternance of a dry
and a rainy season. Data recorded by local meteorology department from 1999
to 2019 showed that the rain season lasts from April to October (7 months) with
an average rainfall of 1038.68 ± 174.31 mm per annum while a total rainfall of
1320 mm and 1349.8 mm were recorded in 2018 and 2019 respectively. The dry
season lasts from November to March (4 months). January is the coldest month
with 15.89˚C and April the hottest with 40.42˚C.
Soils based on sedimentary rocks are considered as Lixisols according to the FAO
classification system [23]. These soils are slightly deep (>85 cm) and dominated by
a loamy-sandy texture (57.5%) with the presence of sandy structures (17.5%), sandy
loamy (15%) and other structures (10%) (BUNASOLs, 1985) in [24].
The local vegetation is savannah of South-Sudanian domain included in the
Sudanian regional centre of endemism [25]. This vegetation is a mosaic patch
constituted of opened tree and bush savannah, wooded and opened woodlands
[26] with annual and/or perennial grasses and bare lands.
2.2. Study Plant Community
We considered vegetation patches on bowal outcrop (opened canopy), shrub-
land (semi-opened) and woodland (closed canopy). This is because the vegeta-
tion patches or canopy conditions can modify the regeneration degree through
the distribution, floristic composition, diversity and structure of naturally oc-
curred woody undergrowth reservoir. According to [27], woodland is defined by
the abundance of individuals of species
Anogeissus leiocarpus
(DC.) Guill. &
Perr.,
Combretum
spp,
Detarium microcarpum
Guill. & Perr.,
Isoberlinia doka
Craib & Stapf.,
Pterocarpus erinaceus
Pair.,
Terminalia
spp and
Vitellaria para-
doxa
var. paradoxa Gaertn f. The mean density was 497.00 ± 180.62 stemsha−1
and height 5.43 ± 0.73 m. Shrubland shelter
Acacia macrostachya
Reichenb. ex
DC,
Combretum
spp,
Lannea
spp,
Gardenia
spp,
Maranthes polyandra
(Benth.)
Prance,
Crossopteryx febrifuga
(Afzel. ex G. Don) Benth,
Sarcostemma
viminale
(L.) R.Br
with a total height not over 2 m above soil surface and annual grass
layer as
Loudetia togoensis
(Pilg.) C. E. Hubb., and
Pennisetum pedicellatum
.
The shrubland is dominated by woody shrub species with an opened layer (stem
density 620.00 ± 175.94 stemsha−1 and mean height 5.24 ± 0.40 m) and annual
grass layer, with occasional tree saplings. Bowal (plural: bowé) also refers to a
shrubland but corresponds to anedaphico-climaticgrassy formations of the
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semi-desert plateaus fossilized by an outcropping or sub-outcropping breastplates,
and with hydromorphic silts [28]. Soil surface is interrupted by crusted soils and
mushroom-shaped mounds erected by
Cubitermes
spp (humivorous) at 99 ha−1
[29]. Annual grass as
Loudetia togoensis
was recurrent with some scattered shrub
species during rainy season on bowal. The average density was 436.00 ± 235.10
stems∙ha−1 and height, 4.52 ± 0.78 m. These few scattered shrubs disappear dur-
ing the dry season leaving to discover a bare soil.
2.3. Experimental Design and Undergrowth Inventory
We firstly delimited 30 contiguous plots of 50 m × 20 m [30] randomly in the
state forest of Dinderesso following transect lines that included the direction of
the rainwater flow. All tree species which stem dbh was equal to or greater than
5 cm were systematically recorded and measured in the 30 plots (or 3 ha) or
tree-plots as adult community. We identified 58 species belonging to 42 families
and 18 genera, with Combretaceae (168 treesha−1), Fabaceae (111.67 treesha−1),
Sapotaceae (64.67 treesha−1), Caesalpiniaceae (63 treesha−1) the most abundant
families in terms of tree density [31]. The density and height of trees in at plot
scale was affected by the type of vegetation assemblage.
In each major plot, 50 subplots of 5 m × 5 m (or 25 m2) were established on
bowal, shrubland and woodland. Among these subplots, 5 were laid with one
subplot at the centre and one subplot at each of the 4 corners (diagonally ar-
ranged) for collecting undergrowth data in tree-plots. In total, 150 subplots were
established and considered.
Data on undergrowth plant communities were collected in the rainy season
during which all the woody species as seedlings, saplings and trees are leafy and
identifiable. This period also corresponded to that of the abundant emergence of
the undergrowth of many species either by seed germination or resprouting of
the remaining stem or sucker [10]. Inventory was proceeded from August to
November 2018. Undergrowth species were identified using vegetation commu-
nity characteristics and botanical criteria of [27] and by comparing morphologi-
cally with trees of the same species. The sample of undetermined species were
taken to be later identified at laboratory with referring to available data on Afri-
can woody species. Scientific name were cross-checked according to [32] and
[33]. For all live individual of which the diameter at breast height (dbh) less than
5 cm and total height of stem equal to or less than and 150 cm were systemati-
cally counted as woody undergrowth contrary to adult tree (mature). The total
height of undergrowth individual was measured using a graduated perch of 7 m,
at each 50 cm and their collar diameter was measured with calliper (0.01 mm
precision). Only the tallest stem was considered in case of more than stem for
individuals.
2.4. Undergrowth Data Analysis
We used the number of 25 m2 subplots in which undergrowth species occurred
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in each tree-plot (1000 m2). We firstly explored the data on undergrowth plant
community alone to assess species richness (S) and floristic composition. Species
richness was determined based on the number of species identified (S) and as
species density which refers to the number of species identified in the total area
of each subplot [34]. Species density is thus expressed as the number of species
(S) at 100 m2 of area. Species name were written in full for the first time and, ge-
neric name was used thereafter. When more than one species were identified in
the same genus, the generic name was abbreviated and followed by specific epi-
thet.
The species diversity of undergrowth was calculated and the Simpsons index
of diversity
I
(Equation (1)) [35] was calculated for each subplot of 5 m × 5 m to
evaluate the species diversity as the probability that two random individuals be-
long to the same habitat:
( )
( )
1
1
11 1
is
i
ni ni
ID NN
=
=
=−=
(1)
where
N
is the total number of woody undergrowth recorded, s the number of
species and
ni
the total number of individuals of species
i
. The Simpsons index
of diversity (
I
) consider the number of species present in a community and the
abundance of each species as single statistic. Its values range from 0 (low diver-
sity) to 1 (high diversity).
For assessing the height distribution in undergrowth community, all the re-
corded individuals growing on bowal, shrubland and woodland were separated
into six different height classes as ]0 - 25], ]25 - 50], ]50 - 75], ]75 - 100], ]100 -
125], ]125 - 150] with their corresponding density in each community.
We noted species richness and abundance as total number of species and stem
per unit of area or density. Since the size of sampled areas for undergrowth and
tree communities were different, both species richness and abundance were bal-
anced by the sampled plots (trees) or subplots (undergrowth) and then stan-
dardized per 100 m2 of area. This method also has been used by [10] for facili-
tating the comparison of species between various areas.
The mean values of 1) species density, 2) Simpsons index of diversity, 3) the
mean density and collar diameter of undergrowth stem were calculated for each
vegetation patch and compare to find out the regeneration niche preferences of
woody species. The variation of undergrowth stem density among habitats or
vegetation patches was analyzed using the one-factor ANOVA of Minitab 15 and
then the test of multiple comparisons of mean values were performed when the
observed differences were significant at probability P = 0.05 based on Tukeys
(HSD) test. In addition, the test of linear regression was used to highlight the re-
lationships that could exist between undergrowth and tree communities in each
habitat regarding their characteristics. These analysis involved both species rich-
ness and stem density to evaluate which of bowal, shrubland and wood shows
good undergrowth reservoir as suitable niche of natural regeneration.
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3. Results
3.1. Effects of Vegetation Patch on Alpha Diversity of Undergrowth
We recorded overall 52 undergrowth species and 2224 stems that belonged to 38
genera and 21 families along the study period. Among these species, 25, 36 and
38 species were identified in bowal, shrubland and woodland respectively (Ap-
pendix). The families of Combretaceae (6 species), Caesalpiniaceae (4 species), and
Mimosaceae (3 species) were largely represented in bowal while Combretaceae (9
species), Caesalpiniaceae (6 species), Euphorbiaceae (3 species), Rubiaceae (3 spe-
cies) dominated in shrubland. The families of Combretaceae (7 species), Caes-
alpiniaceae (6 species), Rubiaceae (4 species), Euphorbiaceae (3 species) and Mi-
mosaceae (3 species) prevailed in woodland.
All the three vegetation patches shared 15 species with each other while 3 spe-
cies were common in bowal and woodland, and 13 species were found in both
shrubland and woodland (Appendix). Only 1 species as
Opilia celtidiflora
Endl.
S. Walp. was found in both bowal and shrubland. In fact, 6 species were solely
identified in bowal, and 7 species in each of shrubland and woodland respec-
tively. The occurrence of species
O
.
celtidiflora
in bowal seems to be accidentally
and linked to fruit (or seed) dispersal process.
The calculated values of species richness as species density at 100 m2 unit of
area and Simpsons index of diversity I of undergrowth community were lower
for bowal, higher for shrubland, and intermediate for woodland (Table 1). Sta-
tistically, vegetation patching significantly influenced the species density (Anova,
F = 38.08, P < 0.0001) and Simpsons index of diversity I (Anova, F = 7.15, P =
0.003). Importantly, the pairwise comparison of mean value revealed that species
density was strongly increased for shrubland (P < 0.0001) and woodland (P<
0.0001) in comparison with bowal while it did not differ significantly between
shrubland and woodland (P = 0.618), and between woodland and bowal (P =
0.078) (Table 1). In contrast, the difference observed in the Simpsons index of
diversity was only significant for shrubland (P = 0.002) comparing with bowal
whereas it was insignificant between shrubland and woodland (P = 0.311) and,
between woodland and bowal (P = 0.078). The test of Tukey discriminated two
homogeneous groups (a et b) relating to specific density and stem density (Table
1): bowal (a) is species-poor site and shrubland-woodland together form a rich-
species site (b). In return, undergrowth community in bowal (a) was less diversi-
fied and shrubland (b) more diversified vegetation regarding the undergrowth
characteristics.
3.2. Effects of Vegetation Patch on the Structure of Undergrowth
Undergrowth individuals were counted in 50 × 25 m2 (or 125 m2) per vegetation
patch. The overall density of undergrowth ranged from 35.28 ± 19.11 stems∙100
m−2 (or 441 individuals125 m−2) on bowal to 76.88 ± 21.02 stems100 m−2 (961
individuals. 125 m−2) in woodland. This density was lower for bowal and interme-
diate for shrubland (Table 1). The most abundant families were Combretaceae,
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Table 1. Species density, Simpson’s index of diversity and abundance of woody undergrowth at sub-
plot-level on bowal, shrubland and woodland in the study site (mean ± standard deviation).
Variables
Subplot location
Bowal
Shrubland
Woodland
F
Subplot no.
50
50
50
Species diversity of undersgrowth
Number of observed genus
23
28
33
Number of observed family
13
18
19
Number of observed species
25
36
38
Species density (no. species 100 m−2)
4.72 ± 2.83a
13.52 ± 1.83b
12.48 ± 2.62b
38.08
Simpsons index of diversity I (1-D)
0.63 ± 0.24a
0.88 ± 0.03b
0.78 ± 0.08ab
7.15
Undergrowth abundance
Number of observed stems
441
822
961
Mean stem density (no. stems∙100 m−2)
35.28 ± 19.11a
65.76 ± 11.82b
76.88 ± 21.02b
14.7
Mean stem collar diameter (cm)
1.21 ± 0.77a
1.18 ± 0.91a
1.07 ± 0.83b
5.55
Minimum diameter (cm)
0.10
0.10
0.10
Maximum diameter (cm)
6.70
7.50
10.00
Mean values with different letter(s) across the same line for each vegetation type are significantly differ-
ent (at P < 0.05) according to Tukey’s HSD test.
Mimosaceae, Caesalpiniaceae, Sapotaceae and Euphorbiaceae (Appendix) with
abundant individuals.
At the scale of vegetation patches, the families of Caesalpiniaceae and Com-
bretaceae were prevalent in bowal whereas Combretaceae, Mimosaceae were sub-
stantially represented in shrubland and woodland respectively.
Out of the shared species,
Detarium microcarpum
Guill. & Perr.,
Guiera sene-
galensis
Lam. and
Diospyros mespiliformis
Hochst. ex A. DC. were abundantly
represented in bowal. Depending on site characteristics, species as
G
.
senegalen-
sis
,
Dichrostachys cinerea
(L.) Wight & Arn. and
Vitellaria paradoxa
var.
para-
doxa
C.F. Gaertn were the most abundant species in shrubland whereas the un-
dergrowth of
D
.
cinerea
,
G
.
senegalensis
and
V
.
paradoxa
dominated in wood-
land.
When we calculated the value of the ratio of shared undergrowth species over
tree communities among patch, we noticed a preponderance of undergrowth in-
dividuals compared with adult tree within species (Appendix). There was much
more undergrowth than adult trees as limited regeneration of these species. Un-
dergrowth density was significantly different across vegetation type (Anova, F =
14.7, P < 0.001 in Table 1). According to Tukeys test the mean density of un-
dergrowth was higher for shrubland (P = 0.002) and woodland (P < 0.0001) than
bowal but this observed difference between shrubland and woodland was not
significant (P = 0.355). Regarding the abundance distribution of undergrowth
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individuals, bowal (a) refers to a particular vegetation set and pool community
of shrubland-woodland (b) form the other homogeneous vegetation.
The collar diameter of undergrowth varied from 1.07 ± 0.83 cm in woodland
to 1.21 ± 0.77 cm in bowal as the highest value (Table 1). The minimum collar
diameter recorded in each area was the same while the maximum ranged from
6.70 cm in bowal to 10 cm in woodland indicating the development level of their
underground systems.
The mean values of stem collar diameter remarkably varied from one site to
another (Anova, F = 5.55, P = 0.004). Difference observed in these mean values
was strongly higher for bowal (P = 0.006) and shrubland (P = 0.032) compared
with woodland while it was not significant between bowal and shrubland (P =
0.849) showing similar community (Table 1). The multiple comparison of mean
values indicated two homogeneous groups as bowal-shrubland (a) and wood-
land (b) in term of stem collar diameter.
3.3. Effects of Vegetation Patch on Height Class Distribution
The abundance distribution of undergrowth varied according to the height classes
(Figure 1). This distribution was in bell-shaped and represented a gauss-type
structure reflecting poor regeneration and recruitment on bowal regarding the
tendency curve shape (Figure 1(a)). Most individuals in this community or 92%
occurred as aggregated in the midst height classes between 50 and 100cm tall. In
contrast, the tendency curve showed that the distribution of undergrowth stems
was in reverse J shape for the shrubland and woodland communities (Figure
1(b) and Figure 1(c)). There was a decrease in undergrowth abundance from ]25
- 50], ]50 - 75], ]75 - 100], ]100 - 125] to ]125 - 150] in shrubland and woodland
towards higher height class. The incidence of undergrowth individuals in height
class ]0 - 25] and those with height > 100 were scarce. Such distribution seems to
be linked to canopy degree and indicate distribution as specific-site (bowal) vs.
similar homogeneous distribution model in both shrubland and woodland com-
munities.
3.4. Relationship between Undergrowth and Tree Communities
The linear regression analyses were performed to explore the relationship be-
tween tree community as explanatory variable and undergrowth community as
response variable (specie and stem densities). The expected results highlighted
positive relationship between undergrowth and tree communities (Anova, P <
0.05) in bowal while it was statistically good between undergrowth and tree stem
densities only in woodland (Anova, P < 0.05) (Table 2). The observed species
richness of undergrowth was 53% explained by that of tree species and their
stem densities at 42% by that of tree density for bowal. In shrubland, under-
growth and tree communities were almost independent (P > 0.05) with weak re-
lationship between them. Only few variations of undergrowth species richness
(26.50%) and stem density (16.20%) was explained respectively by species richness
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Figure 1. Pattern of the height class pattern of woody undergrowth recorded in 2018 in bowal (a), shrubland (b)
and woodland (c).
Table 2. Linear relationship between the species richness and stem density of woody un-
dergrowth and tree communities for each vegetation patch.
Subplot location
Bowal
Shrubland
Woodland
Species richness (no. species∙100 m−2)
Regression equation
6.34x − 0.79
3.09x + 8.30
1.53x + 10.9
R2
52.50%
26.50%
3.00%
ANOVA, P
0.018
0.128
0.630
Density (stems∙100 m−2)
Regression equation
5.78x’+14.7
2.40x’+52.5
7.51x’+ 42.4
R2
42.20%
16.20%
55.80%
ANOVA, P
0.042
0.249
0.013
Regression equation corresponded to species richness (or stem density) = a x (with x =
species number∙100 m−2 or x = stem density (stems∙100 m−2) + b; P: probability.
S. Traoré et al.
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10.4236/oje.2022.122007 123 Open Journal of Ecology
Table 3. Ratio of woody undergrowth over tree densities (stems∙100 m−2) for the most 15 shared species in the study vegetation
patches.
Species name
Bowal
Shrubland
Woodland
Undergrowth
Tree
Ratio
undergrowth
Tree
Ratio
Undergrowth
Tree
Ratio
Annona senegalensis
0.40
0.02
20.00
1.44
0.06
24.00
2.16
0.08
27.00
Burkea africana
0.72
0.03
24.00
1.20
0.37
3.24
0.48
0.07
6.86
Cassia sieberiana
0.88
0.44
2.00
0.48
0.02
24.00
0.08
0.01
8.00
Combretum nigricans
0.08
0.03
2.67
0.24
0.16
1.50
0.48
0.08
6.00
Daniellia oliveri
0.08
0.03
2.67
0.56
0.15
3.73
0.56
0.53
1.06
Detarium microcarpum
10.08
1.1
9.16
1.36
0.30
4.53
0.80
0.01
80.00
Dichrostachys cinerea
0.40
-
-
10.56
-
-
28.48
0.01
2848
Diospyros mespiliformis
1.52
0.04
38.00
0.16
0.01
16.00
0.72
0.09
8.00
Flueggea virosa
0.48
-
-
1.52
-
-
1.12
-
-
Gardenia ternifolia
0.64
-
-
1.20
-
-
2.08
-
-
Guiera senegalensis
4.72
0.11
42.91
12.48
0.05
249.60
17.44
0.03
581.33
Holarrhena floribunda
0.80
-
-
0.08
-
-
0.72
-
-
Pteleopsis suberosa
0.72
-
-
3.12
0.49
6.37
2.08
0.04
52.00
Terminalia macroptera
0.64
0.22
2.91
4.00
0.39
10.26
2.64
0.19
13.89
Vitellaria paradoxa
0.72
0.08
9.00
5.76
0.9
6.40
4.40
0.96
4.58
and density of woody tree. Among the shared undergrowth species (Table 3),
G
.
senegalensis
and
D
.
mespiliformis
had high undergrowth density compared with
their corresponding trees in bowal while
D
.
cinerea
and
G
.
senegalensis
had rela-
tively abundant undergrowth over tree ratio in shrubland and woodland. From
the overall 15 shared undergrowth species,
Flueggea virosa
(Roxb. ex Willd.)
Voigt,
Gardenia ternifolia
Schumach. & Thonn. and
Holarrhena floribunda
T.
Durand & Schinz did not exist as tree in the study areas.
4. Discussion
The present research analyzed the characteristics of woody undergrowth com-
munity in natural vegetation patches as habitats or regeneration niche in a state
forest. Our results revealed that the type of vegetation or habitats importantly
affects woody undergrowth community through establishment and recruitment.
In accordance with our first specific objective, woody undergrowth associated
with woodland sheltered higher number of species corresponding to 73.08% of
the total counted species versus bowal with 48.08% (the lowest rate) and shrub-
land 69.23% (intermediate rate). At the scale of occupied surface, the species
density (species.100 m−2) of undergrowth community was lower in bowal but it
was similar between both shrubland and woodland. Ecological undergrowth
species were also represented and but differed between the three types of vegeta-
S. Traoré et al.
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10.4236/oje.2022.122007 124 Open Journal of Ecology
tion. The species of Caesalpiniaceae (
Burkea
sp) and Mimosaceae (
Acacia
sp)
families were the most abundant and frequent in each of the vegetation patches.
Species richness observed in study areas may indicate that of viable seeds and
propagules (mainly seed stock) that were stored on the floor or underground
relatively to their dispersal system and have emerged during the favourable con-
ditions of humidity and temperature. Nonetheless this significant difference may
certainly result from the chance of species undergrowth to abundantly establish,
grow and co-exist during their early stage [36] and hereafter influence the spe-
cies diversity as response to heterogeneous areas described in community plant
material (see Section 2.2). Current results on species richness are consistent with
previous studies carried out in different climate zones [37] [38] [39] [40] [41].
By using a theoretical model during their investigation, [37] have demonstrated
a positive effect of canopy composition (adult tree composition) on the under-
canopy composition and shrub diversity through the modification of the soil
conditions in where they grow. Therefore, in function of their ecological adap-
tive strategy, a many species may persist under particular canopy conditions as
bowal (plenty opened), shrubland (light opened) and woodland (full cover) as
area specific-species because vegetation patch likely act as filter or refuge for
them to the detriment of area sensitive species [42]. This is the case of shared
undergrowth species that can abundantly colonize and grow simultaneously in
these three vegetation patches but their persistence is driven by their plasticity
profile.
Regarding the second specific objective, species diversity in each vegetation
patch was the consequence of capability to occur on bowal, shrubland and wood-
land leading to the co-existence of few species richness and abundant with un-
even stem density even if the observed difference in the value of Simpsons di-
versity index for shrubland and woodland was insignificant. In shrubland, the
most common families were Caesalpiniaceae, Combretaceae and Mimosaceae
and highly contributed to increase the species diversity of undergrowth reservoir
through the species number representing each of them in study sites during
sampling period (see Appendix). Although the community associated with bo-
wal sheltered lower species richness and diversity relating to shrubland, its di-
versity was similar with that of woodland undergrowth community as the proven
of preference regeneration niche for each species. Current observations cor-
roborate that obtained by [42] [43]. By comparing plant diversity between bowé
(plural of bowal) and adjacent woodlands, these authors have pointed out lower
woody species occurring on bowé with the predominance of
A
.
macrostachya
,
C
.
glutinosum
,
C. nigricans
,
D. microcarpum
and
L. microcarpa
individuals as re-
sistant species to drought and with high regeneration potentials whereas Fa-
baceae family was well represented along the Njoro and Kamweti rivers by
Aca-
cia
sp [44]. Therefore the low species diversity of woody undergrowth on bowal
may result from the stressful soil conditions (edaphic factors), such as incrusted
soil surface, absence of litter (bare soil) leading to water deficiency. Such in-
S. Traoré et al.
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10.4236/oje.2022.122007 125 Open Journal of Ecology
crusted soil certainly reduce significantly space available for tree species to estab-
lish or prevent undergrowth to take root.
While vegetation patches significantly differ in species diversity, the overall
abundance of undergrowth individuals evolved across sites and so did collar di-
ameter and height class distribution. Our results showed that the density of un-
dergrowth was significantly lower on bowal only in where the density value was
twice that observed in shrubland and woodland (see Table 1). Such a reduced
stem density of undergrowth community on bowal may depict limit in under-
growth establishment with scattered individuals [43] while shrubland and wood-
land together appeared with aggregated individuals as positive effect of canopy
extend for undergrowth establishment contrary to [45]. Another explanation of
unbalanced distribution in the different habitats may be that some species better
fit a given environment than others at both undergrowth and adult tree stages.
The co-existence of undergrowth individuals of the same and different species in
each community is linked to the type of growing niche and their relationships
with the existing ecological factors.
Besides, density distribution, undergrowth community on both bowal and
shrubland had larger stump trunk in view of the collar diameter value. These
communities constitute similar homogeneous group for stem collar diameter but
differed from woodland. One may question whether the size of stem (big strunk)
respond to an edaphic heterogeneity or to particular stress (competition with
established species) in place or ontogenic factor? This is because high densities
of undergrowth are likely to be correlated with small undergrowth sizes on each
area. The higher collar diameter found on bowal mainly give an insight on the
fact that woody undergrowth develop priority their underground systems as en-
vironment-feedback for a good anchoring, storing much nutrients and water to
face unfavourable conditions for survival. Report indicated that undergrowth
with larger stem diameter tend to survive better than small stem diameter and,
are strong competitors [46]. Interestingly, most undergrowth individuals were
under 100 cm tall and the proportion of those in the upper height class decreased
on bowal (bell-shaped or gauss distribution). This fact may indicate that the high
densities of undergrowth are likely to be correlated with small undergrowth sizes
on each area. As for the distribution of undergrowth observed in shrubland and
woodland, we can state that woody species roughly emerged jointly from propagules
and/or soil seed stock (germination by seed) as response to favourable condi-
tions (study period) and only the most successful species may be recruited in
taller groups. This variation may also result from higher mortality rate or a slow
growth that individuals experience along their life cycle and hereafter inducing a
decreased density in taller groups (e.g. density in shrubland) corresponding to
an instable communities and extreme vulnerability at early stage. Broadly, the
three vegetation patches have a poor recruitment status regarding a gauss type
structure observed, although some species abundantly established successfully in
detail contributing to increase stem density. This is a case of D.
cinerea
,
D
.
mi-
crocarpum
in scrubland and woodland, and
D
.
mespiliformis
in bowal. Similar
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10.4236/oje.2022.122007 126 Open Journal of Ecology
result was reported for
Boswellia papyrifera
population by [47] in Ethiopia where
species exhibited a poor regeneration status.
Globally, the recruitment of woody species open to criticism because of the
preponderance of undergrowth as regards adult trees through shared species as
undergrowth > trees. Nevertheless the variation of undergrowth species and stem
density was significantly related to adult tree on bowal solely whereas 55.80%
variations in woodland resulted from adult stem density only as the regression
analysis results. This strong relationship suggest that woody undergrowth spe-
cies successfully regenerated abundantly beneath adult trees but decline during
undergrowth to tree transition in both bowal and woodland. Establishment pat-
tern was likely promoted by the way adult tree may act on habitat conditions as
facilitator or inhibitor during growth process.
5. Conclusion
Our results showed that undergrowth reservoirs varied across vegetation patches
as response to the degree of heterogeneity in the study sites. In the Dinderesso
state forest. These undergrowth reservoirs and may be an indicator of the species
richness of soil propagules and/or seed stock. The three vegetation patches
ranged into two homogeneous groups as the undergrowth community linked to
bowal and that of shrubland-woodland. A particular undergrowth vegetation set
is growing in bowal as area specific-species and stem density which positively
vary in function of the associated tree community characters. Bowal as well as
shrubland and woodland acts as suitable regeneration niche with reservoir for
limited number of species that favourably co-exist. We may state that the co-exi-
stence of undergrowth of the same and different woody species and afterwards
tree individuals in a particular community is in function of woody species
themselves (intrinsic factors as biotic), habitat-specific biotic (protection from
stand canopy) and the type of growing niche and their different interrelated
mechanisms. Nonetheless, the main driver of observed distinction is the edaphic
factors at small spatial scale (bowal) but undergrowth reservoir remain con-
nected with family existing in its surroundings. Furthermore, this reservoir con-
stitutes an indicator of species richness and abundance and hereafter a predictor
of the physiognomy of woody tree community in place. The woody undergrowth
community should be considered in the program of diversity management and
degraded land restoration or other area similar to bowal. Based on the results of
this study, field works could be undertaken to understand the characteristics of
seed bank for bowal, shrubland and woodland regarding the undergrowth and
tree communities and various conditions as the infiltration of soil water and bi-
otic factor effects.
Acknowledgements
Current study did not obtain any external financial support. Authors would like
to thank the staff of Direction provincial des Eaux et Forêts, Houetfor author-
S. Traoré et al.
DOI:
10.4236/oje.2022.122007 127 Open Journal of Ecology
izing us to work and ensuring our security in Dindéresso forest. Authors are
grateful to the staff of IRD-Bobo for allowing us to access the institute e-library
and to the anonymous reviewers for their valuables comments and suggestions.
Data Availability Statement
All data supporting the results of this study are available from the corresponding
author (Traoré Saran) on decent demand.
Conflicts of Interest
The authors declare no conflicts of interest regarding the publication of this pa-
per.
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S. Traoré et al.
DOI:
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Appendix. Complete List of Undergrowth Species and Stem Number (Density Stems100
m2) in 50 Subplots Laid in Each Vegetation Patch
Species-name
Family
Number of undergrowth (N)
Density (stems∙100m2)
Bowal
Shrubland
Woodland
Bowal
Shrubland
Woodland
Acacia dudgeoni
Craib ex Hall.
Mimosaceae
-
-
1
-
-
0.08
Acacia erythrocalyx
Brenan
Mimosaceae
-
9
-
-
0.72
-
Acacia macrostachya
Reichenb. ex DC
Mimosaceae
2
-
-
0.16
-
-
Annona senegalensis
Pers.
Annonaceae
5
18
27
0.4
1.44
2.16
Anogeissus leiocarpus
(DC.) Guill. & Perr.
Combretaceae
-
1
37
-
0.08
2.96
Baissea multiflora
A. DC.
Apocynaceae
-
-
1
-
-
0.08
Brideliaferruginea
Benth
Euphorbiaceae
-
2
-
-
0.16
-
Bridelia micrantha
(Hochst.) Baill.
Euphorbiaceae
-
28
28
-
2.24
2.24
Burkea africana
Hook.
Caesalpiniaceae
9
15
6
0.72
1.2
0.48
Cassia arereh
Delile
Caesalpiniaceae
-
3
7
-
0.24
0.56
Cassia sieberiana
DC.
Caesalpiniaceae
11
6
1
0.88
0.48
0.08
Celtis integrifolia
Lam.
Ulmaceae
4
-
1
0.32
-
0.08
Combretum glutinosum
Perr. ex DC.
Combretaceae
23
-
-
1.84
-
-
Combretum micranthum
G. Don
Combretaceae
6
-
-
0.48
-
-
Combretum molle
R. Br ex G. Don
Combretaceae
-
6
-
-
0.48
-
Combretumnigricans
Lepr. exGuill & Perr.
Combretaceae
1
3
6
0.08
0.24
0.48
Crossopteryx febrifuga
Benth.
Rubiaceae
-
-
2
-
-
0.16
Daniellia oliveri
(Rolfe) Hutch. & Dalz.
Caesalpiniaceae
1
7
7
0.08
0.56
0.56
Detarium microcarpum
Guill. & Perr.
Caesalpiniaceae
126
17
10
10.08
1.36
0.8
Dichrostachys cinerea
(L.) Wight & Arn.
Mimosaceae
5
132
356
0.4
10.56
28.48
Diospyros mespiliformis
Hochst. Ex A. DC
Ebenaceae
19
2
9
1.52
0.16
0.72
Entada africana
Guill. & Perr.
Mimosaceae
86
-
3
6.88
-
0.24
Flueggea virosa
(Roxb. ex Willd.) Voigt
Euphorbiaceae
6
19
14
0.48
1.52
1.12
Gardenia aqualla
Stapf & Hutch.
Rubiaceae
-
11
2
-
0.88
0.16
Gardenia erubescens
Stapf & Hutch.
Rubiaceae
-
8
6
-
0.64
0.48
Gardenia ternifolia
Schumach. & Thonn.
Rubiaceae
8
15
26
0.64
1.2
2.08
Grewia bicolor
Juss.
Tiliaceae
-
-
4
-
-
0.32
Guiera senegalensis
Lam.
Combretaceae
59
156
218
4.72
12.48
17.44
Holarrhenafloribunda
T. Durand & Schinz
Apocynaceae
10
1
9
0.8
0.08
0.72
Hymenocardia acida
Tul.
Euphorbiaceae
-
-
4
-
-
0.32
Isoberlinia doka
Craib & Stapf
Fabaceae
-
-
1
-
-
0.08
Jatropha curcas
L.
Euphorbiaceae
2
-
-
0.16
-
-
S. Traoré et al.
DOI:
10.4236/oje.2022.122007 132 Open Journal of Ecology
Continued
Khaya senegalensis
A. Juss.
Meliaceae,
-
18
11
-
1.44
0.88
Lannea acida
A. Rich., Guill. & Perr.
Anacardiaceae
-
1
2
-
0.08
0.16
Lannea velutina
A. Rich., Guill. & Perr.
Anacardiaceae
6
-
-
0.48
-
-
Maranthes polyandra
(Benth.) Prance
Chrysobalanaceae
-
12
-
0
0.96
-
Opilia celtidifolia
(Guill. & Perr.) Endl.
ex Walp.
Opiliaceae 1 7 - 0.08 0.56 -
Parinari curatellifolia
Planch. ex Benth.
Chrysobalanaceae
-
2
4
-
0.16
0.32
Piliostigma thonningii
(Schumach.)
Milne Redh.
Caesalpiniaceae - 43 2 - 3.44 0.16
Prosopis africana
Taub.
Fabaceae.-
-
45
4
-
3.6
0.32
Pteleopsis suberosa
Engl. & Diels
Combretaceae
9
39
26
0.72
3.12
2.08
Pterocarpus erinaceus
Poir.
Fabaceae
-
9
1
-
0.72
0.08
Saba senegalensis
(A. DC.) Pichon
Apocynaceae
20
-
-
1.6
-
-
Securidaca longepedunculata
Fresen
Polygalaceae
-
21
-
-
1.68
-
Strychnos spinosa
Lam.
Loganiaceae
5
-
1
0.4
-
0.08
Terminalia avicennioides
Guill. & Perr.
Combretaceae
-
9
24
-
0.72
1.92
Terminalia laxiflora
Engl.
Combretaceae
-
24
11
-
1.92
0.88
Terminalia macroptera
Guill. & Perr.
Combretaceae
8
50
33
0.64
4
2.64
Terminalia mollis
M.A. Lawson
Combretaceae
-
6
-
-
0.48
-
Vitellaria paradoxa
var.
Paradoxa
C. F.
Gaertn.
Sapotaceae 9 72 55 0.72 5.76 4.4
Vitex doniana
Sweet
Lamiaceae
-
-
1
-
-
0.08
Vitex simplicifolia
Oliv.
Lamiaceae
-
5
-
-
0.4
-
441
822
961
35.28
65.76
76.88
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