diversity
Article
Impacts of Diffuse Land-Use on Plant Diversity Patterns in the
Miombo Woodlands of Western Zambia
Priscilla Sichone 1,2, * , Jens Oldeland 3 , Patrick Phiri 4 , Norbert Jürgens 1
1
2
3
4
*
Citation: Sichone, P.; Oldeland, J.;
Phiri, P.; Jürgens, N.; Schmiedel, U.
Impacts of Diffuse Land-Use on Plant
Diversity Patterns in the Miombo
Woodlands of Western Zambia.
Diversity 2023, 15, 739. https://
and Ute Schmiedel 1
Institute of Plant Science and Microbiology, Universität Hamburg, Ohnhorststr. 18, 22609 Hamburg, Germany
Compassionate Carbon Zambia Project, 3 Nangwenya Road, Rhodes Park, Lusaka 10101, Zambia
Institute for Globally Distributed Open Research and Education (IGDORE), Burgunderweg 9d,
22453 Hamburg, Germany
School of Medicine, Cavendish University, 47 Senanga Road, Handsworth Park,
Lusaka P.O. Box 33145, Zambia
Correspondence: priscilla.sichone@compassionatecarbon.com
Abstract: Land use is known to influence the diversity of vascular plants in the Miombo woodlands.
However, little is known about the interaction between soil and land use in herbaceous and woody
species. We compared the diversity of vascular plants at the plot level (20 m × 50 m) and site level
for three sites in the Miombo woodlands of western Zambia subject to different levels of intensity
classes of diffuse land use (e.g., livestock herbivory and selective timber harvesting). For each of
the sites, twenty plots were randomly selected for assessment of species composition of vascular
plant species, indicators of land-use intensity, and soil chemistry per plot. We hypothesized that the
site with the lowest human impact would have the highest richness and diversity of woody and
herbaceous species. At the site level, we found that richness and diversity of woody species were
unaffected by land-use intensity, whereas herbaceous species richness was higher for the protected
site (28 species on average per 1000 m2 ) than the two other sites (23 and 21 species on average per
1000 m2 ). At the plot level, herbaceous species richness was positively associated with woodcutting
and soil pH. We interpret the positive effect of woodcutting on herbaceous species richness as the
effect of lower competition by the woody component for resources such as water, nutrients, and light.
With regard to the absence of any effect of land-use intensity on the richness of woody species, we
conclude that in our study areas selective timber harvesting may be at a sustainable level and might
even have a positive effect on the diversity of the herbaceous layer.
Keywords: species richness; Shannon diversity; herbaceous species; woody species; soil variables;
Miombo woodlands
doi.org/10.3390/d15060739
Academic Editors: Shengwei Wang
and Thomas Fickert
1. Introduction
Received: 26 March 2023
The Miombo woodlands form a widespread dry woodland belt covering large parts
of southern and eastern Africa, encompassing Angola, Botswana, the Democratic Republic
of Congo, Malawi, Mozambique, Tanzania, Zambia, and Zimbabwe [1]. While the flora of
the Miombo woodlands is characterised by high species richness, the diversity of canopy
trees is low [2]. The natural resources of the Miombo woodlands strongly contribute
to the wealth of the social and economic systems in their distribution area [3]. This
ecosystem and its resources, however, are prone to habitat transformation and biodiversity
loss [4]. Anthropogenic activities such as fuel combustion, commercial selective harvesting
of valuable timber, grazing, crop production, and complete deforestation threaten the
woodland system [5–7]. With a deforestation rate of 1.5% per year, Zambia is even classified
as one of the countries with the highest deforestation rates in the world [8]. Complete
habitat conversion from forest to cropland inevitably decreases both herbaceous and woody
species richness [9,10].
Revised: 29 May 2023
Accepted: 29 May 2023
Published: 3 June 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Diversity 2023, 15, 739. https://doi.org/10.3390/d15060739
https://www.mdpi.com/journal/diversity
Diversity 2023, 15, 739
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In contrast to deforestation, with its complete transformation of land cover, diffuse
disturbance is characterised by relatively small patches of change distributed over a large
area [11]. In woodlands, scattered, small-scale land use through grazing, selective timber
harvesting, or small-scale cropping has a diffuse but continuous negative impact on the
vegetation and woody and herbaceous forest biodiversity, as has been shown by studies
of (sub)tropical forest ecosystems in southern Africa [12], Brazil [13], the mountains of
Mexico [14], and the tropical rainforests of Madagascar [15]. Yet, the extent of the impact
appears to vary from low to high [16,17]. Revermann et al. [12] showed how plant species
richness and evenness respond to the diverging land-use patterns of spatially diffuse versus
intense land use in the dry woodlands along the Kunene in Angola and Namibia. They
showed that the spatially diffuse land use on unfenced communal land in Namibia has
measurable negative effects on the richness of mainly woody plant species. The authors
attributed this pattern to selective timber logging. In Zambia, Chidumayo [18] expressed
a growing concern countrywide about the negative effects of diffuse disturbance due to
the selective harvesting of trees for charcoal production and other uses. Irrespective of
the potential impact of diffuse land use on botanical diversity, the effects of spatiallydiffuse land use on the vegetation of the Miombo woodland have received little scientific
attention in the available literature. Most of the researchers who have investigated the
impact of land use on the Miombo woodlands have mainly focused on the effects on the
woody component of the vegetation [3,19,20]. However, as has been shown by Revermann
et al. [12], the response of forbs and grass species to diffuse disturbance of woodland
systems may differ from that of woody species. The herbaceous layer of the Miombo
woodlands shows great spatial variation in species composition, and several herbaceous
genera contribute to the species richness and local endemism in the system [2]. Irrespective
of the reported response of the herbaceous vegetation to diffuse land use in other vegetation
types [12] and the strong contribution of herbaceous flora to the overall species richness
and endemism, we are not aware of any study on the effect of land use on both the woody
and herbaceous vegetation of the Miombo woodlands.
Therefore, in this study we focus on the effect of spatially diffuse land use on the
composition and diversity of herbaceous and woody plant species in the western Zambian
Miombo woodlands. To this end, we selected three sites in the Miombo woodlands of western Zambia, each representing a different land use type (i.e., national park, national forest,
and community forest) related to a different level of spatially diffuse land-use intensity.
National Park (Kafue): no land-use activities are permissible, and only prescribed fires
are used as a management intervention tool.
National Forest (Dongwe): a medium level of disturbance; only minimal land-use
activities are allowed, and harvesting of timber is only allowed upon issuance of a permit.
Community Forest (Luampa): high level of disturbance with several land-use activities permitted, including the harvest of both timber and non-timber products as well as
occasional agricultural activities.
We compared two species diversity measures for woody and herbaceous species at
the three sites and analysed the spatially heterogeneous effects of indicators of diffuse
land-use intensity and soil chemical variables on the species diversity measures at plot
scale (1000 m2 , n = 60).
Our hypothesis was that the Kafue site in the National Park would have a higher
richness and diversity of woody and herbaceous species at both the plot and site scale than
the two sites with higher land-use intensity. We additionally expected that, as observed
by Revermann et al. [12], the diffuse disturbances would have a greater negative effect on
species richness and biodiversity of trees than on the herbaceous layer.
2. Material and Methods
2.1. Study Area
The study was conducted at three sites in the Miombo woodlands of western Zambia
(Figure 1), extending over latitudes S 14◦ –16◦ and longitudes E 24◦ –26◦ at an elevation
Diversity 2023, 15, 739
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ranging from 1068 m to 1210 m a.s.l. (Table 1). The Miombo woodlands are the most
extensive tropical seasonal woodland considered part of the African Savannah, covering
about 2.4 million km2 in Africa [21]. Dominant plant families of Fabaceae tree species of the
genera Brachystegia Benth., Isoberlina Craib and Stapf ex. Holland, and Julbernardia Pellegr.,
which are clustered in the subfamily Caesalpinioideae and the tribe Ahmerstieae [22–24],
characterise the Miombo woodlands [24]. The area has a tropical sub-humid climate with
alternating dry and wet seasons. Rainfall occurs for 5–7 months in summer [25]. The mean
annual temperature of the study area is 20.8 ◦ C, and the mean annual rainfall ranges from
875 to 990 mm (Table 1). The soils consist of Kalahari sands from the Tertiary to the recent
period which, according to Japan Association for International Collaboration of Agriculture
and Forestry [26], cover western and northwestern Zambia. The main soil type is Arenosols,
a formation of the parent Basement and Katanga rocks, with the accumulation of Karoo
deposits [27].
Figure 1. Locations of study sites (C) in western Zambia, showing the sites in relation to the continent
(A) and the country (B).
Floristically, the three sites fall within the Sudano-Zambezian Phytoregion [28], whereas
the vegetation is classified as Miombo woodlands sensu [29], which forms part of the tropical seasonal woodlands [30]. Miombo woodlands are characterized by co-dominance of
trees, shrubs, and herbaceous plants, with respective proportions determined by environmental, ecological, and human parameters and adapted to fire [31]. The dominant tree
species are those referred to by Phiri [24] as dominant taxa of the Miombo woodlands.
2.2. Study Design and Data Collection
We selected three study sites in Kafue National Park (called Kafue for the rest of the
text), Dongwe, and Luampa, approximately 100 km apart (Figure 1), based on literature
consultations [29,32] and site visits to the study area of western Zambia in 2014. Classification according to land-use type and intensity was guided by the State of the Environment
in Zambia [33]. Each site was characterised by different land-use intensities, from low
Diversity 2023, 15, 739
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to medium to high (Table 1). For each study site, we classified the vegetation into open
or closed woodland using the most recent Google Earth image [34] at the time of data
assessment. Based on habitat stratification, we randomly selected twenty vegetation plots
for each of the sites. The plots were 1000 m2 (20 m × 50 m) in size and were laid out in an
east–west extension.
Table 1. Descriptive attributes of the three study sites.
Site Name
Mean Annual
Rainfall
[mm/yr]
Mean Annual
Temperature
[◦ C]
Land-Use Type
Land-Use
Intensity
Luampa
875.5
22.22
Forest Reserve
High
Dongwe
990.6
22.34
Community
Forest
Medium
Kafue NP
897.4
21.98
National Park
Low
GPS Coordinates
(North West
Corner)
S15.13782,
E24.48778
S14.09577,
E24.01520
S14.89830,
E25.43676
Elevation
Min–Max
[masl]
1152–1158
1066–1145
1091–1210
Sources: [35] for mean annual rainfall and temperature; [36] for land-use type; and [27] for soil type.
For each of the plots, we recorded the presence and identity of all angiosperms as
herbaceous or woody species (as defined by Petruzzello [37], with the former being plants
that do not have a true woody stem and may be either perennial or annual), estimated cover
per species in percent, and counts of individuals per species (see Appendix A, Table A1
for all recorded species identified). Grass species had to be excluded from the analyses
because at the time of sampling the majority of grasses did not have inflorescence, which
compromised identification of the species. To assess the presence of exotic species and
field weeds in the study area, we reviewed the literature sources in Zambia [38–41]. Indicators for land-use intensity, such as signs of recent woodcutting, grazing, and browsing,
were recorded semi-quantitatively. The categories of woodcutting were 1–2 stumps per
1000 m2 -plot = 1, 3–5 stumps = 2, 6–8 stumps = 3, and >8 stumps = 4. Signs of browsing
from game animals (domestic livestock were not observed in any of the study sites) were
graded from 1 (no browsing) to 4 (high abundance of signs of browsing) depending on the
frequency of signs of browsing observed. Grazing was not observed on any of the plots.
The time from the last fire event on the plot was determined based on consultation with
local field assistants as well as our own observations of the age of visible signs. We did
not determine the cause of the fire. However, in areas where human activity is frequent,
fires caused by human activity are more common than natural causes [42]. The determined
time from the last fire was translated to the ordinal scale for the recent occurrence of fire:
long ago = 2 (10 or more years since last fire), recent = 1 (5 years), and very recent = 0 (1 or
2 years).
One composite soil sample comprising five subsamples per plot was collected from
the topsoil layer (0–10 cm). Soil samples were analysed at the Mt. Makulu Research Centre
Soil Laboratory, Chilanga, Zambia for the variables of pH, nitrogen (N), phosphorus (P),
organic carbon (Org C), calcium (Ca), magnesium (Mg), sodium (Na), potassium (K), zinc
(Zn), manganese (Mn), iron (Fe), and cation exchange capacity (CEC). For the soil analytical
methods, see Appendix A, Table A2.
Within the plots, vegetation data were collected during the rainy seasons (March–May)
of 2014 and 2015. The data obtained from the Zambia Meteorology Department (ZMD)
indicated that the annual rainfall in these years was at 50–70% of the mean annual rainfall,
which ranges between 875 and 990 mm for the study area (Table 1). Mean annual rainfall per
site was available for the period from 1950 to 2010 from the Global Land Data Assimilation
System site [35].
Plant species identification in the field was carried out using field guides [43–46]
and later confirmed at the herbarium of the University of Zambia. Plant nomenclature
followed Phiri [24] for the majority of the identifications; in cases of ambiguity, the Flora
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of Zambia [39], the JSTOR Global Plants database [47], and the Plant List [48] were used
for verification. Voucher specimens were lodged at the Herbarium Hamburgense of the
Universität Hamburg (HBG) and the herbarium of the University of Zambia (UZL).
2.3. Data Analysis
Species inventories were analysed for richness (S) and Shannon index per plot [49].
The sites differed in the degree of exposure to human activities (Table 1). We tested the
semi-quantitative land-use variables (browsing, grazing, woodcutting, and time from last
fire) for differences among the three land-use intensities by applying a Kruskal–Wallis test
and then running a pairwise comparison of the land-use variables.
We tested the normal distribution of the diversity data. For the non-normally distributed Shannon diversity values, which are already on a log scale, we used the exponential
of the Shannon, which is a common way to transform Shannon entropy into Shannon diversity values [50]. The diversity values were then expressed as the effective number of
species. We applied one-way ANOVA to test for differences between the species richness
values and the exponential of the Shannon under different land-use intensities followed by
a Tukey HSD post hoc test.
We were interested in how well land-use intensity and soil chemistry can explain
variation in richness and Shannon diversity of woody and herbaceous plants at the plot scale
across the three study sites. We screened all soil variables visually for skewed distributions,
which are common for these kinds of data. Of the twelve measured soil variables, we had to
log-transform eight (P, Ca, Mg, K, Na, Zn, Mn, Fe). We tested the environmental variables
for pairwise correlation to exclude multicollinearity, and only maintained the relevant
variables of fire, woodcutting, browsing, soil pH, and above-mentioned eight soil variables.
We employed generalised linear models (GLM) using the Poisson distribution, as the data
were discrete and had a lower boundary (zero) for richness. A Gaussian distribution was
used for the exponential of the Shannon diversity values. For the four response variables in
the GLM (i.e., the richness and the diversity of woody and herbaceous species, respectively),
we created a full model including soil and land-use variables as predictors. We used a
heuristic approach that compared all 2100 possible models (not using interactions) and
identified a final best model. All models were fitted using the R package glmulti [51].
Because we were using GLMs, we used a ‘pseudo’-R2 measure, which behaves like R2 and
measures the improvement of a fitted model compared to a null model calculated as the
ratio [52]. Here, we used Cragg and Uhler’s R2 available from the R package pscl [53].
3. Results
3.1. Indicators for Land-Use Intensities at the Three Sites
We compared the indicator values for diffuse land-use intensity among the three sites.
Table 2 shows that the two sites which were subject to diffuse anthropogenic disturbance,
Dongwe National Forest and Luampa Community Forest, showed more frequent signs of
woodcutting and shorter time from the last fire along with a lower frequency of signs of
browsing compared to the Kafue site.
3.2. Diversity Patterns at the Different Land-Use Intensities
Due to our pre-classification of the sites into three different land-use types (Table 1)
and the higher frequency of recorded signs of anthropogenic disturbances at the two sites
under human land use (Luampa and Dongwe) compared with the National Park site (Kafue,
Table 2), we expected strong differences between the diversity indicators in the sites. For
all three sites, a cumulative total of 624 vascular plant species from 51 plant families was
recorded. Of these species, 239 were woody and 385 were herbaceous. The mean species
richness of the twenty plots (1000 m2 ) per site for woody species was 27 (Luampa), 25
(Dongwe), and 27 (Kafue), while for herbaceous species it was 23 (Luampa), 21 (Dongwe),
and 28 (Kafue) (Figure 2). In contrast to our expectations, species richness and Shannon
diversity at the plot level only differed in terms of herbaceous species (Figure 2). Kafue
Diversity 2023, 15, 739
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had on average eight more herbaceous species per 1000 m2 plot than the other two sites.
The mean richness of woody species per plot was about 26 species, and the mean diversity
was at about 13 effective species at all three sites. Very little research has been published on
exotic and invasive plant species in woodland habitats [38–41]. Of the herbaceous species
observed in the semi-disturbed sites of our study (Luampa and Dongwe), the following
herbaceous species have been classified as weeds in the literature: Crassocephalum rubens,
Striga asiatica, Vernonia petersii, and Crotalaria spp. Dichrostachys cineria, a species reported
by Blaser-Hart et al. [40] to cause bush encroachment in Zambia, was observed in the
woody vegetation of all the sites, though with low density.
Table 2. Pairwise comparisons using the Wilcoxon Rank Sum Test. Median values for the levels of
the land-use variables. Figures in brackets = the range of the observed intensity of the respective
land-use variables; hyperscripts indicate differences between sites at a significance level of p < 0.05.
Land-Use
Luampa (High
Land-Use Intensity)
[Median Values and
(in Brackets)
Min–Max Values]
Dongwe (Medium
Land-Use Intensity)
[Median Values and
(in Brackets)
Min–Max Values]
Kafue (No Land-Use)
[Median Values and
(in Brackets)
Min–Max Values]
Woodcutting
1a
(0–3)
1a
(0–3)
0b
(0–1)
Browsing
0a
(0–1)
0a
(0–1)
1b
(0–2)
Time since last fire
0a
(0–2)
1a
(0–2)
2b
(1–2)
Figure 2. Bar plots of the diversity indices of the 1000 m2 plots for species richness and Shannon index
as follows (a) Richness for the woody species; (b) Richness for the herbaceous species; (c) Shannon
index for the woody species and; (d) Shannon index for the herbaceous species. Different groups
ff
(based on Tukey’s HSD test) are indicated by superscripts.
ff
tt
ff
Diversity 2023, 15, 739
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3.3. Environmental Drivers of Diversity at Plot Level
Although the recorded disturbance variables of woodcutting, browsing, and fire events
differed between Kafue and the other two sites, their distribution varied within each of the
sites as well (Table 2). We were therefore interested in the effects of the plot-based land-use
intensities on the diversity measures per plot. We related the observed signs of disturbance
per plot to the species richness and Shannon diversity of woody and herbaceous plant
species for the respective plots. To discriminate between the disturbance effects and those
of other abiotic habitat variables, we included plot-based soil variables in the analyses as
well. For each of the four response variables (i.e., richness and diversity of woody and
herbaceous species), we did not find any significant variation in richness or diversity of
the woody component in response to any of the disturbance or soil variables (Table 3).
We did, however, find a positive response of the species richness in the herbaceous layer
to an increase in soil pH and to woodcutting at the plot level. The Shannon diversity of
herbaceous species solely responded to organic carbon (positively) and iron (negatively).
Overall, the models were rather weak, with low pseudo-R2 values except for the richness
of the herbaceous layer.
Table 3. Generalised linear models for species richness (S) and the exponential of the Shannon
diversity (H’) of woody and herbaceous species after multi-model inference. The values for the
logarithm of iron (logFe) and woodcutting are chi-square statistics, indicating an overall significant
effect of the parameters. pR2 is Cragg and Uhler’s pseudo r-squared [51] measure based on the
differences between best model and null model. SOC = soil organic carbon.
Diversity
Model
Distribution
and Link
n
Intercept
Swoody
Sherbs
H’woody
H’herbs
Poisson (log)
Poisson (log)
Gaussian (id)
Gaussian (id)
60
60
60
60
3.275 ***
2.593 ***
11.880 ***
2.773 ***
log Mg
SOC
pH
log Fe
‡
0.151 **
0.935
2.506
−5.970
−12.952 *
Wood
Cutting ‡
9.432 *
−2.141 *
pR2
0.00
0.36
0.07
0.18
* p < 0.05; ** p < 0.01; *** p < 0.001 ‡ Chi-square statistic.
4. Discussion
4.1. Effect of Land Use on Plant Species Diversity
Our study showed that the two sites under medium and high land-use-intensity,
namely, Dongwe Forest Reserve and Luampa Community Forest, indeed had higher
frequency of woodcutting and a shorter time from the last fire than Kafue (Table 2). Selective
woodcutting in the Miombo woodlands in Zambia targets valuable timber species. The
most harvested species according to the Forest Department are Baikiaea plurijuga and
Pterocarpus angolensis [54], which are used for both domestic and commercial purposes.
Selective harvesting of valuable timber species could lead to overharvesting and even local
extinction, as has been shown by De Cauwer et al. [55] for Pterocarpus angolensis in Namibia.
Therefore, we expected woodcutting for selective timber harvesting to have a negative
effect on woody species richness and diversity. Even though intensity of woodcutting
intensity differed between Kafue and the other two sites, woody species richness and
diversity were not significantly different among the three sites. One possible reason for this
might be that the observed woodcutting intensity at Dongwe Forest Reserve and Luampa
Community Forest, with 0–3 tree stumps per 1000 m2 , while significantly higher than in
Kafue (Table 2), was too low to have a measurable negative effect on the tested diversity
measures. In previous studies, very moderate land-use intensity did not show any negative
effect on species richness and abundance of woody species in the Miombo woodlands of
Tanzania [56] and in the West African savannah of Burkina Faso [57], and even showed an
increase in the tree species diversity of the Miombo woodlands in Angola [12]. It appears
that in the present study the impact of land use on woody species diversity was below
the threshold, in contrast to that observed in other studies [56,57]. This observation shows
Diversity 2023, 15, 739
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that the offtake of woody species through woodcutting and destruction by fire can be
considered sustainable if the impact remains on a small spatial scale.
In contrast to the woody species richness and diversity, we found the richness of
herbaceous species to be lower at Luampa and Dongwe than at the Kafue National Park
site, suggesting a negative impact of land use on the herbaceous layer. Nacoulma et al. [57]
compared the herbaceous species richness of sites under land use and in protected areas
for savannah woodlands in Burkina Faso. They found that sites under higher land-use
intensity to have lower herbaceous species richness, which they explained by the effect
of higher grazing intensity in the unprotected area. Unsustainable grazing pressure from
livestock reduces the number of palatable herbaceous species in woodlands [58]. The
impact of grazing livestock (i.e., livestock that predominantly consume the herbaceous
layer of the vegetation) on biodiversity has been addressed by previous studies on savannah
ecosystems [59–61], showing that grazing may, depending on its intensity, have either a
positive or negative effect on species richness. In our study, however, we did not find any
signs of grazing in either Kafue, where one might expect game to graze, or at either of the
other two sites, where grazing domestic cattle could be expected. The observed absence of
grazing signs at the Miombo woodland site in Kafue supports reports that the grazers of
the national park prefer the open grasslands (called dambos) [62]. Similarly, at Luampa and
Dongwe, where grazing is also absent, the livestock farmers prefer to graze their cattle in
the open grasslands (dambos), where the quality of forage is better.
Although we did not find any signs of grazing (consumption of the vegetation of the
herbaceous layer), we did find signs of browsing (consumption of the leaves on twigs and
branches of shrubs and trees), which was significantly less at the two sites under land use
(Luampa and Dongwe) than at Kafue. At Luampa and Dongwe, browsing livestock such
as goats have been restricted to areas close to land users’ homesteads, and are kept away
from the distant woodlands, according to personal communication with local farmers. At
Kafue, we found signs of browsing, even though no presence or signs of large mammals
were observed. The browsing signs could be attributed to the presence of insects known to
browse on woody species [62]. However, increased density of browsing insect species in
the woodlands of Kafue National Park is unlikely to have a negative effect on the woody
component of the vegetation. This means that the low density of woody species at the
Kafue site cannot be explained by grazing. Reduced woody cover facilitates the abundance
of herbaceous species by reducing the competition with woody components for resources
such as light, water, and nutrients [30].
Field weeds and invasive species such as Crassocephalum rubens, Striga asiatica, Vernonia
petersii, and Crotalaria spp. were recorded only occasionally or even rarely in the plots of the
semi-disturbed study sites. Dichrostachys cineria was only rarely observed at all three sites.
Owing to the very limited information available in the literature on exotic and invasive
plant species in the woodlands of Zambia, very few species in our study were identified as
exotic or invasive, and these had very low densities. Therefore, our data suggest that the
influence of synanthropic plants on the diversity patterns is likely to be low.
4.2. Drivers of Diversity at Plot Level
The range of land-use variables per site (Table 2) revealed within-site variability
of land-use effects within the three study sites. Therefore, we tested for the effect of
land-use intensity on species richness and diversity per plot. Because soil characteristics
show interplay with biotic drivers and drive vascular plant diversity in the Miombo
woodlands [25], we tested the effect of both land-use indicators and soil variables on
diversity and richness at plot level irrespective of the land-use type they were exposed
to. Land-use (woodcutting) and soil variables (soil pH) showed effects on the diversity
patterns of herbaceous species, whereas woody species diversity and richness were not
affected. The absence of land-use effects on woody species diversity and richness is in
contrast to other studies showing that selective harvesting of valuable timber species can
lead to local extinction [55]. The patchy nature of diffuse disturbance of woodlands referred
Diversity 2023, 15, 739
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to by Asefa et al. [63] is defined by the absence of settlements and crop cultivation as well
as relatively low-level logging and grazing, and had positive effects on woody species
richness in Ethiopia. In turn, we found the herbaceous species richness to increase with
woodcutting. As discussed earlier, this positive effect might have been a result of better
light conditions for the herbaceous species coupled with reduced competition for space,
water, and nutrient resources [4,30]. Reduction of competition for resources in combination
with the release of organic nutrients through decaying tree stumps have previously been
shown to increase herbaceous biomass in the Miombo woodlands [64]. At the site level,
Kafue had the highest herbaceous species richness as well as the highest density in terms
of signs of browsing (Table 2). Browsing, as discussed earlier, may already have a positive
effect on herbaceous species and species diversity. Our study showed a positive effect of
diffuse disturbance caused by woodcutting (plot level) and browsing (site level in Kafue)
on herbaceous species richness and diversity.
We further found a positive relationship between richness of herbaceous species and
an increase in soil pH at the plot level. Soils in humid subtropical regions are typically
acidic [65]. In our study area, the dominant soil type (Arenosols) is leached under high rainfall conditions, resulting in low pH [66], with soil pH ranging from 3.7 to 5.4 (Appendix A,
Table A3). Acidity in soils slows down the rate of decomposition of soil organic material,
and thereby reduces the availability of nutrients for plant uptake [67]. Thus, highly acidic
soils may provide a low nutrient supply, which limits the range of plant species that can
cope with these conditions [68]. Therefore, very low soil pH has previously been found
to be negatively associated with species diversity of herbaceous species in the Miombo
woodlands [2,69,70] and the tropical montane forests of Cameroon [71].
We further found SOC at the plot level to be negatively associated with the Shannon
diversity of herbaceous species. Generally, SOC content increases with precipitation and
with optimal levels in humid and cold climates and decreases with soil pH; beyond that,
SOC storage links to biophysical factors and management practices [72]. A study in Ghana
by Quaye et al. [73] revealed unsuitably low SOC in strongly acidic soils in the western
African Savannah woodlands. In the Miombo Woodlands of our study area, which are
Savannah woodlands, low soil pH might have negatively affected SOC as well. The SOC
appeared to be negatively related to the biodiversity of herbaceous species, which could be
because herbaceous species have a lower root network than woody species [30].
At the plot level, our study showed that both land use (woodcutting) and soil acidity
were drivers of the diversity of herbaceous species, whereas woody species were unaffected.
The absence of variance in the richness and diversity of woody species, however, does
not exclude the fact that there are differences in other vegetation characteristics, such as
species composition. Variances in small-scale species composition in response to fire and
herbivory have previously been shown in the Miombo woodlands of Zimbabwe [74]. We
expect similar effects in our study area, which we will analyse in a subsequent study.
5. Conclusions
This study revealed that diffuse land use has no influence on woody species richness
and diversity in the Miombo woodlands in our study area. The absence of such effects on
woody species richness and diversity was consistent across all three of our sites. However,
the study showed influence of both diffuse land use and soil acidity on herbaceous species
richness at the plot level. Among the land-use parameters, woodcutting (at plot level)
and browsing (at site level) showed positive effects on herbaceous species richness; both
of these result in opening up of the tree canopy, providing water, nutrients, and sunlight
for herbaceous species. Other land-use effects, such as fire and grazing, which have been
shown in other studies to influence patterns of richness and diversity of vascular plant
species in the region, were not found in our study. We assume that the intensities of these
disturbances at all three sites and plots were too low to show any effects.
Diversity 2023, 15, 739
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Author Contributions: P.S. (Hamburg University)—Data sampling, data analysis, and drafting of
the manuscript; J.O. (Hamburg University)—Statistical support; N.J. (Hamburg University)—Study
design and conceptual guidance; P.P. (Cavendish University)—Identification of specimens and
conceptual guidance; U.S. (Hamburg University)—Study design, conceptual guidance, and support in
drafting the manuscript. All authors have read and agreed to the published version of the manuscript.
Funding: This research was part of the BMBF-funded SASSCAL initiative (Task no. 159) promotion
number 01LG1201M.
Institutional Review Board Statement: Not applicable.
Data Availability Statement: The data in this publication is deposited in BIOTABase and can be
accessed on request from pmwinji@yahoo.com.
Acknowledgments: This study formed a part of the research portfolio of the BMBF-funded SASSCAL
initiative (Task no. 159) promotion number 01LG1201M. We are very grateful to the field assistants
who participated in the collection of the data, namely, Alex Liseli, Andrew Munshukulumbwe,
and Charles Sitali. Gerhard Muche and Rasmus Revermann provided guidance for the use of the
database software BIOTABase and R statistics package. We are grateful for the support on climate
data provided by Verena Baumberg. Zambia Wildlife Authority granted us permission to undertake
this study in its protected areas.
Conflicts of Interest: The authors declare no conflict of interest.
Appendix A
Table A1. List of woody and herbaceous plant species with their life form, collection number,
and location, arranged according to family: Abundance was assigned according to the frequency
of observations of each woody plant species: rare (1 or 2 recordings), occasional (3–5 recordings),
frequent (6–10 recordings), and common (>11 recordings). The locations where these species occurred
in the study sites are abbreviated as D = Dongwe, K = Kafue National Park, and L = Luampa. The different uses were coded as TI = timber production, PO = posts, pole, and roundwood, WO = fuelwood
and charcoal, PU = pulp and paper production, FD = fodder, FO = food, NW = other non-wood
products (gums, medicines, dyes, tanning, etc.), AE = aesthetic and ethical values, and TX = toxic
to livestock.
Species Name
Acanthaceae
Duosperma quadrangulare (Klotzsch) Brummitt
Hypoestes forskaolii (Vahl) R.Br.
Amaryllidaceae
Crinum macowanii Baker
Anacardiaceae
Searsia quartiniana (A. Rich.) A.J. Mill.
Sclerocarya birrea (A.Rich.) Hochst.
Annonaceae
Friesodielsia obovata (Benth.) Verdc.
Uvariastrum hexaloboides (R.E.Fr.)
Xylopia odoratissima Welw. ex Oiv.
Apocynaceae
Diplorhynchus condylocarpon (Müll.
Arg.) Pichon
Landolphia parvifolia K. Schum.
Strophanthus welwitschii (Baill.) K. Schum.
Life Form
Abundance
Uses
Voucher
Number
Location
Herbaceous
Herbaceous
rare
FO
TX
138076
132148
K
K
Herbaceous
rare
rare
NW
132141
L, K
Woody
Woody
Frequent
rare
FO
FO
131071, 142196
132128
L, D
K
Woody
Woody
Woody
occasional
occasional
common
FO, FD
FO
NW
142610
132107
131181
K
L, K
L, D
Woody
common
NW
140121
D, L, K
Woody
Woody
frequent
occasional
FO
142505
142649
D, L
D
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Table A1. Cont.
Voucher
Number
Location
142788
K
NW
131355, 132214
D, K
occasional
rare
occasional
frequent
frequent
rare
rare
NW
NW
132207
131301
138099
131119
142503
142698
142660
Herbaceous
occasional
NW
142665
Herbaceous
Herbaceous
Herbaceous
occasional
frequent
rare
FD
131401
142678
131157
K
D
K
D, L, K
L, K
D
K
L, K
L, K
L
D, K
L
Capparis tomentosa Lam.
Herbaceous
frequent
NW,
FO, AE
138093
K
Cleome hirta (Klotzsch) Oliv.
Maerua triphylla ssp. pubescens A. Rich.
(Klotzsch) DeWolf
Chrysobalanaceae
Parinari capensis Harv.
Herbaceous
frequent
138076
K
Herbaceous
frequent
131358
K
Woody
frequent
Parinari curatellifolia Planch. ex Benth.
Woody
frequent
Combretaceae
Combretum collinum Fresen.
Combretum elaeagnoides Klotzsch
Combretum molle R.Br. ex G.Don
Combretum psidioides Welw.
Woody
Woody
Woody
Woody
occasional
occasional
frequent
occasional
Combretum zeyheri Sond.
Woody
common
Pteleopsis anisoptera (Welw. ex M.A.Lawson)
Engl. & Diels
Woody
occasional
Terminalia brachystemma Welw. ex Hiern
Woody
occasional
Herbaceous
Species Name
Asparagaceae
Asparagus racemosus Willd.
Asphodelaceae
Bulbine abyssinica A.Rich.
Asteraceae
Conyza gouanii (L.) Willd.
Crassocephalum rubens (Jacq.) S. Moore
Dicoma anomala Sond.
Elephantopus scaber L.
Erythrocephalum zambesianum Oliv. & Hiern
Felicia welwitschii (Hiern) Grau
Macledium poggei (O.Hoffm.) S.Ortiz
Pleiotaxis eximia O. Hoffm.
Vernonia glabra ssp. laxa (Seetz) Vatke
Vernonia melleri Oliv. & Hiern
Vernonia petersii Oliv. & Hiern ex Oliv.
Vernonia poskeana Vatke & Hildebr.
Capparaceae
Life Form
Abundance
Herbaceous
occasional
Herbaceous
rare
Herbaceous
Herbaceous
Herbaceous
Herbaceous
Herbaceous
Herbaceous
Herbaceous
Uses
NW
NW, FO
NW,
FO, WO
L
131213
L, K
131113
131403
142578
131402
D, L, K
D, L
D, L, K
D
131183
L, K
1320102
D
131165
L, K
frequent
131016
D, L, K
Woody
Woody
occasional
occasional
140121
132222
K
K
Woody
occasional
142639, 141189
L
Diospyros batocana Hiern
Woody
common
140159
D, L, K
Diospyros mespiliformis Hochst. ex A.DC.
Woody
frequent
Commelinaceae
Cyanotis longifolia Benth.
Dipterocarpaceae
Marquesia macroura Gilg
Gilg
Monotes glaber Sprague
NW, TI
NW, TI
NW
NW,
WO
NW, TI,
WO
NW,
FO, WO
NW,
WO
Ebenaceae
Diospyros virgata (Gürke) Brenan
Ericaceae
Cleistanthus polystachyus Hook. f. ex Planch.
Erythroxylaceae
Erythroxylum emarginatum Thonn.
NW,
FO, PU
NW, FO
NW
K
131054, 131191,
140143
D, L, K
Woody
common
Herbaceous
rare
132122
L
Woody
occasional
142649
K
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Table A1. Cont.
Species Name
Life Form
Abundance
Uses
Herbaceous
Woody
Woody
Woody
occasional
occasional
common
frequent
NW, FO
NW
NW
Woody
frequent
Woody
common
NW, FO
Herbaceous
rare
NW
Woody
Woody
occasional
occasional
NW, FO
NW, FO
Afzelia quanzensis Welw.
Woody
frequent
Albizia antunesiana Harms
Woody
frequent
Albizia versicolor Welw. ex Oliv.
Woody
common
Anisophyllea boehmii Engl.
Woody
occasional
Baphia massaiensis var. obovata Taub.
Woody
common
Bauhinia petersiana Bolle
Woody
frequent
Bobgunnia madagascariensis (Desv.) J.H.Kirkbr.
& Wiersema
Woody
common
Brachystegia boehmii Taub.
Woody
common
Brachystegia spiciformis Benth.
Woody
common
Burkea africana Hook.
Woody
common
Cassia abbreviata Oliv.
Chamaecrista mimosoides (L.) Greene
Copaifera baumiana Harms
Crotalaria alexandri Baker f.
Crotalaria anisophylla (Hiern) Welw. ex
Baker f.
Crotalaria caudata Welw. ex Baker
Crotalaria cephalotes Steud. ex A.Rich.
Crotalaria laburnifolia L.
Crotalaria microcarpa Hochst. ex Benth.
Cryptosepalum exfoliatum ssp. pseudotaxus
De Wild.
Woody
Herbaceous
Woody
Herbaceous
occasional
occasional
common
occasional
Herbaceous
occasional
Herbaceous
Herbaceous
Herbaceous
Herbaceous
occasional
occasional
common
rare
Woody
common
Dichrostachys cinerea (L.) Wight & Arn.
Woody
common
Erythrophleum africanum
Woody
common
Guibourtia coleosperma (Benth.) J.Leonard
Indigofera demissa Taub.
Indigofera flavicans Baker
Woody
Herbaceous
Herbaceous
common
occasional
common
Euphorbiaceae
Acalypha ornata Hochst. ex A. Rich.
Flueggea virosa (Roxb. ex Willd.) Voigt
Hymenocardia acida Tul.
Maprounea africana Müll.Arg.
Oldfieldia dactylophylla (Welw. ex
Oliv.) Léonard
Pseudolachnostylis maprouneifolia Pax
Sclerocroton oblongifolius (Müll. Arg.) Kruijt
& Roebers
Uapaca kirkiana Müll. Arg.
Uapaca nitida ssp. nitida
Fabaceae
NW, FO
TI, WO
NW,
WO
NW,
WO
NW, FO
NW,
FO, FD
NW,
FO, FD
NW,
FO, TI
TI, WO,
PO,
NW
TI, WO,
PO,
NW
WO,
PO,
NW
NW
NW
NW
NW
TI, NW,
PO,
NW,
WO
WO,
PO,
NW
TI, NW,
Voucher
Number
Location
132101
142566
142526
L
L, K
D, L, K
L
142693
D
140131, 131047
131056, 142172,
135679
142817
D, L, K
131163
L, K
142766
D, L, K
132156
D, K
132216
K
131007
D, L, K
142520
D, L, K
131191
L, K
L, K
D, K
L, K
D, L, K
131146
D, L, K
D, L, K
142622
131025, 142559
131085
142576
K
D, K
L, K
L, D
142774
L
131176
132180
142570
131099
D, L
K
D
D, L
142504
D
D, L, K
131124
D, L, K
140151
131174
131130
D, L
D, L
L, K
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Table A1. Cont.
Species Name
Life Form
Abundance
Isoberlinia angolensis (Benth.) Hoyle & Brenan
Woody
occasional
Julbernardia paniculata (Benth.) Troupin
Woody
common
Keetia venosa (Oliv.) Bridson
Lannea edulis (Sond.) Engl.
Mucuna poggei Taub.
Woody
Woody
Woody
occasional
frequent
occasional
Pericopsis angolensis (Baker) Meeuwen
Woody
occasional
Piliostigma thonningii (Schumach.)
Milne-Redh.
Woody
occasional
Pterocarpus angolensis DC.
Woody
common
Rhynchosia caribaea (Jacq.) DC.
Flacourtiaceae
Flacourtia indica (Burm.f.) Merr.
Hypericaceae
Psorospermum baumii Engl.
Lamiaceae
Ocimum africanum Lour.
Tinnea vestita Baker
Woody
Vitex doniana Sweet
Vitex madiensis Oliv. subsp. milanjiensis
(Britten) F. White
Lauraceae
Cassytha pondoensis ssp. Pondoensis Engl.
Malvaceae
Abutilon angulatum (Guill. & Perr.) Mast.
Pavonia senegalensis (Cav.) Leistner
Meliaceae
Bersama abyssinica Fresen.
Trichilia emetica Vahl
Myrtaceae
Syzygium guineense (Willd.) DC.
Ochna pulchra Hook.
Olacaceae
Olax obtusifolia De Wild.
Ximenia americana L.
Ximenia caffra Sond.
Oleaceae
Olea capensis L.
Schrebera trichoclada Welw.
Orobanchaceae
Striga asiatica (L.) Kuntze
Oxalidaceae
Biophytum abyssinicum Steud. Ex A. Rich.
Biophytum umbraculum Welw.
Passifloraceae
Paropsia brazzeana Baill.
Polygalaceae
Securidaca longepedunculata Fresen.
Uses
TI, NW,
AE
TI, WO,
PO,
NW, FD
NW
NW
TI, WO,
NW, FD
Voucher
Number
Location
K
140155
D, L, K
142561
131178
142564
K
L, K
K
138072
L, K
K
occasional
TI, WO,
NW
NW
131195
L, K
Woody
frequent
NW, FO
131198
K
Woody
occasional
NW, FO
131162
D
Herbaceous
Herbaceous
rare
occasional
131021, 142739
K
L, K
Woody
occasional
NW, FO
TI, FO,
NW, FD
140154
L
Woody
rare
142638
L
Woody
occasional
NW
131134
D, L
Woody
Woody
occasional
rare
NW
142504
131268
D
D, L
Woody
Woody
rare
rare
NW, AE
NW, AE
142874
K
K
Woody
Woody
occasional
common
NW, FO
NW
142200
131059
L
D, L, K
Woody
Woody
Woody
occasional
frequent
frequent
NW, FO
NW, FO
142598
142523
131047
L
L, K
K
Woody
Woody
occasional
rare
138065
144524
K
L, K
Herbaceous
occasional
NW
138077
L, D
Herbaceous
Herbaceous
occasional
occasional
NW
131098
D
D, L
Woody
frequent
NW
131080
D, L, K
Woody
occasional
NW
131204
L
D, L, K
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Table A1. Cont.
Species Name
Life Form
Abundance
Uses
Proteaceae
Protea angolensis Welw.
Protea gaguedi J.F. Gmel.
Ranunculaceae
Woody
Woody
rare
frequent
NW
NW
Clematis chrysocarpa Welw. ex Oliv.
Herbaceous
occasional
Ziziphus mucronata Willd.
Woody
occasional
Rubiaceae
Agathisanthemum bojeri Klotzsch
Fadogia cienkowskii Schweinf.
Gardenia ternifolia Schumach. & Thonn.
Pavetta schumanniana F. Hoffm ex K. Schum
Rothmannia engleriana (K.Schum.) Keay
Spermacoce pusilla Wall.
Tricalysia longituba De Wild.
Woody
Woody
Woody
Woody
Woody
Woody
Woody
occasional
occasional
occasional
frequent
frequent
occasional
occasional
Vangueriopsis lanciflora (Hiern) Robyns
Woody
frequent
Voucher
Number
142795
131351, 142755,
142609
Location
K
K
L, K
Rhamnaceae
FO,
NW
NW
K
142781
142191
132132
142749
131196
131100
131255
L, K
L
D
D, K
D, L
D, L
L, K
NW,
WO
131180
D, L
NW, FO
NW
Sapindaceae
Zanha africana (Radlk.) Exell
Sapotaceae
Englerophytum magalismontanum (Sond.)
T.D.Penn.
Solanaceae
Solanum mauritianum Scop.
Strychnaceae
Woody
occasional
WO
142751
K
Herbaceous
occasional
NW
131079
D, L
Herbaceous
occasional
Strychnos cocculoides Baker
Woody
frequent
Strychnos pungens Soler.
Thelypteridaceae
Christella chaseana (Schelpe) Holttum
Tiliaceae
Grewia flavescens Juss.
Triumfetta annua L.
Verbenaceae
Endostemon obtusifolius (E. Mey. ex Benth.)
N.E.Br.
Woody
frequent
Herbaceous
occasional
Herbaceous
Herbaceous
occasional
occasional
Herbaceous
occasional
Herbaceous
occasional
Herbaceous
Herbaceous
frequent
frequent
occasional
Lantana angolensis Moldenke
Vitaceae
Cyphostemma junceum Wild & R.B. Drumm.
Cyphostemma princeae Wild & R.B. Drumm
Zingiberaceae
Aframomum alboviolaceum (Ridl.) K.Schum.
Herbaceous
K
NW,
FO, AE,
PO
NW, FO
131083
D, L, K
140149
D, L, K
D
NW, FO
131013
131109
D, L, K
L, K
D, L
NW,
FO, FD
142644
L, K
NW, FO
NW, FO
132169
131382
D
D, L, K
NW, FO
D
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Table A2. Methods of soil sample analysis.
Analysis Variable
Method and Reference
Unit
pH
using the CaCl2 mixture in H2 O [75]
Nitrogen (N)
Kjeldahl method [76]
percentage per total weight
Phosphorus (P)
Bray I extractant [77]
parts per million
Organic Carbon (Org C)
Walkley Black technique [78]
percentage per total weight
Calcium (Ca)
Ammonia acetate extraction [79]
parts per million
Magnesium (Mg)
Ammonia acetate extraction [79]
parts per million
Sodium (Na)
Ammonia acetate extraction [79]
parts per million
Potassium (K)
Ammonia acetate extraction [79]
parts per million
Zinc (Zn)
DPTA method [80]
parts per million
Manganese (Mn)
DPTA method [80]
parts per million
Iron (Fe)
DPTA method [80]
parts per million
Cation Electronic Exchange (CEC)
Conductivity method [81]
Milli-equivalents
Table A3. Soil variables from the observatory samples analysis.
Variables
pH
N
P
Org C
Ca
Mg
Na
K
Zn
Mn
Fe
CEC
Mean and Covariance
Luampa
Dongwe
Kafue National Park
x
4.4
3.9
4.9
CV
0.04
0.07
0.08
x
0.016
0.011
0.018
CV
0.5
0.64
0.66
x
5.0381
3.01
7.73
CV
0.42
0.39
0.83
x
0.25
0.21
0.26
CV
0.43
0.37
0.5
x
40
32
120
CV
0.71
1.61
0.49
x
10.23
11
24.31
CV
0.58
0.14
0.57
x
2.23
3.75
6.53
CV
1.49
0.64
0.65
x
10.91
10.35
42.05
CV
1.28
1.14
0.48
x
0.27
0.05
0.15
CV
2.19
1.43
1.4
x
14.42
12.1
82.05
CV
1.15
2.6
0.58
x
11.23
11.95
28.47
CV
0.49
0.84
0.51
x
3.45
2.4
3.83
CV
0.35
0.58
0.32
Soil variables with x for mean values and CV for covariance.
Diversity 2023, 15, 739
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