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Article

Contrasting Herbaceous Communities in South African Savannas: A Comparative Analysis of Density, Composition, and Diversity Across Three Bioregions

by
Armand Arthur Biko’o
1,*,
Willem Johannes Myburgh
1 and
Brian Kevin Reilly
2
1
Department of Nature Conservation, Tshwane University of Technology, Private Bag X680, Pretoria 0001, South Africa
2
Department of Genetics, University of Free State, P.O. Box 339, Bloemfontein 9300, South Africa
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(7), 475; https://doi.org/10.3390/d17070475
Submission received: 10 June 2025 / Revised: 5 July 2025 / Accepted: 8 July 2025 / Published: 10 July 2025
(This article belongs to the Special Issue Plant Communities in Transition: Succession, Diversity, and Ecosystem Dynamics)
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Abstract

This study provides novel insight into herbaceous-layer dynamics across three distinct South African savanna bioregions (Central Bushveld, Lowveld, and Mopane) using a Total Count Quadrat approach to investigate species densities, community composition, diversity, and spatial patterns. A total of 196 unique herbaceous species were recorded across all bioregions. Contrary to typical expectations, our findings reveal no statistically significant differences in overall herbaceous density (ranging from 24.3 ± 2.31 to 32.0 ± 1.28 individuals/m2; F2,6 = 1.89, p = 0.23), species richness (F2,6 = 1.91, p = 0.23), or Shannon diversity (F2,6 = 3.23, p = 0.11) among bioregions, suggesting a more complex interplay of environmental drivers beyond broad climatic gradients. However, there was significant within-bioregion spatial heterogeneity in density, notably in the Central Bushveld (F2,87 = 4.96, p = 0.009) and Mopane (F2,87 = 7.54, p < 0.001) regions, indicating important fine-scale variation, unlike in the Lowveld region (F2,87 = 1.25, p = 0.292). Growth form analysis revealed that forbs consistently dominated species richness across all three bioregions (Central Bushveld: ~64%; Lowveld: ~70%; and Mopane: ~67%) and were also the dominant growth form by density in the Lowveld (54.3%) and Mopane (63.8%) regions. While numerical differences in grass density were observed, no statistically significant difference was found across bioregions (F2,6 = 4.15, p = 0.07). Sedges consistently contributed a small proportion to both species richness and total density. Non-metric multidimensional scaling further revealed patterns of dispersion in herbaceous community compositions between the Lowveld and Mopane regions, with Central Bushveld communities exhibiting greater variability. These findings underscore the critical ecological importance of forbs in South African savannas, not only for biomass but also for driving herbaceous diversity and highlighting the necessity of considering fine-scale spatial variation in future ecological research and conservation strategies.

Graphical Abstract

1. Introduction

Savannas constitute one of the largest biomes of the world [1], the largest portion of which occurs in Africa. The South African savannas constitute the southernmost extension of the savanna and represent 32.8% of South Africa [2]. Renowned for their rich biodiversity and essential ecological roles, they are intricate ecosystems where the herbaceous layer forms a critical base [3,4,5]. This layer, encompassing grasses, forbs, sedges, and other non-woody plants, drives primary production, sustains a diverse herbivore community, and significantly impacts nutrient cycles and fire regimes [6,7]. Understanding not only the absolute densities of its constituent species but also the broader community composition and diversity of this layer is paramount for effective conservation and management in these dynamic environments [8]. Changes in vegetation dynamics can profoundly impact savanna biodiversity and ecosystem function, necessitating detailed investigations into these multifaceted aspects.
Contemporary savanna ecology increasingly focuses on understanding ecosystem responses to change and the importance of heterogeneity [9]. Recent research continues to highlight the sensitivity of savanna herbaceous communities to environmental shifts [10], including altered rainfall patterns and land management practices [11]. These factors can induce substantial changes in species abundance and distribution, ultimately affecting ecosystem stability and resilience. The ecological strategies and adaptations of different plant growth forms play a crucial role in mediating these responses to environmental conditions and disturbances [12]. Furthermore, the encroachment of woody vegetation, a prevalent issue in many savannas, directly influences herbaceous layer density and diversity through competition for vital resources [13]. Understanding the contribution of various plant life and growth forms is also critical for appreciating global patterns of vascular plant diversity [14]. Thus, a comprehensive understanding of herbaceous layer dynamics, encompassing species abundance patterns and diversity, is crucial for effective conservation, sustainable land use, and the maintenance of vital ecosystem services [3].
Given the critical role and vulnerability of the herbaceous layer, accurate quantification of its structural attributes is indispensable. While absolute densities provide fundamental information, analyzing community composition and diversity patterns offers deeper insights into ecosystem health and resilience. Multivariate analyses, such as non-metric multidimensional scaling (nMDS), can reveal overall community relationships, while diversity indices, such as the Shannon index, can quantify species evenness and richness.
This study addresses the need for such a comprehensive understanding by employing a total count quadrat approach to determine the species and their absolute densities in the herbaceous layer across three distinct South African savanna subtypes: the Central Bushveld, Mopane, and Lowveld bioregions [2]. This research is particularly important as detailed, comparative baseline data on the herbaceous layer across these specific and ecologically significant South African bioregions remain limited. Furthermore, this research extends beyond absolute density comparisons to analyzing herbaceous community compositions using nMDS and assessing diversity using the Shannon index. By comparing these ecologically significant regions, this study aims to provide a nuanced understanding of the variations in herbaceous layer structure, diversity, and community patterns that characterize these important savanna ecosystems. Building on the premise that savanna herbaceous communities respond to environmental gradients [12], we hypothesized that distinct differences in herbaceous density, species richness, and community composition would be observed among these bioregions. Specifically, this study investigates (1) the differences in absolute herbaceous species densities, (2) the variations in community composition, and (3) the disparities in species diversity across the Central Bushveld, Mopane, and Lowveld bioregions of South Africa.

2. Study Area

The Lowveld, Central Bushveld, and Mopane regions are three savanna bioregions that represent a moisture gradient from moist to arid savanna [2]. This study was conducted in three different study areas, each purposively selected as a representative site within these three major savanna bioregions.

2.1. Letlapa Pula Game Reserve: Central Bushveld

The Central Bushveld is a significant bioregion in the northern interior of South Africa, characterized by a warm, semi-arid climate [2]. Its diverse geology, stemming from the Bushveld Igneous Complex, results in a mosaic of soil types that support mixed savanna vegetation. This bioregion is typically dominated by a variety of woody species such as Vachellia and Combretum, with an herbaceous layer primarily composed of grasses such as Themeda and Eragrostis. Fire, rainfall, and herbivory are key ecological drivers, and woody encroachment is a notable management challenge [9,15].
The study within the Central Bushveld Bioregion was conducted at the Letlapa Pula Game Reserve (LPGR), situated in the Waterberg area (Limpopo Province). The LPGR experiences a Köppen Cwa climate (summer rainfall and hot summers) and falls within the Crocodile River catchment area, characterized by lowland sand beds or floodplain geomorphology [16,17]. The reserve encompasses two vegetation units of the Central Bushveld: Western Sandy Bushveld and Waterberg Mountain Bushveld, with a mean annual precipitation of 584 mm [2].

2.2. Selati Game Reserve: Mopaneveld

The Mopane bioregion, prominent in the northern parts of South Africa, is characterized by a generally hot and arid climate with summer rainfall, often with high inter-annual variability [2]. This region experiences hot summers and a prolonged dry season, with evaporation rates often exceeding precipitation. The dominant vegetation is typically Colophospermum mopane, which can form extensive woodlands, often on nutrient-poor and sometimes sodic soils derived from underlying gneiss and granite. The herbaceous layer in mopane veld tends to be less dense and diverse compared to more mesic savannas, with grasses adapted to drier conditions [18].
The study within the Mopane Bioregion was conducted at the Selati Game Reserve (SGR), situated south of the Murchison Mountain range and north of the Olifants River in the Limpopo Province [19]. According to the Köppen climate classification, SGR falls within the hot, arid steppe climate zone (BSh), characterized by hot summers, warm-to-cool winters, and a negative net water balance due to high evaporation [16]. The Selati Game Reserve lies largely within the Mopane Bioregion and is predominantly covered by the Phalaborwa-Timbavati Mopane veld vegetation unit (SVmp 7), with a mean annual precipitation of 522 mm [2]. The geology of the SGR is influenced by the Murchison Greenstone belt in the northwest and various granite and gneiss formations across the remainder of the reserve [19,20].

2.3. Kempiana Nature Reserve: Lowveld

The Lowveld Bioregion, situated in the eastern, lower-altitude parts of South Africa, exhibits a subtropical climate characterized by hot, wet summers and mild, dry winters [2]. The vegetation is diverse and adapted to the warmer conditions and often nutrient-poor soils. It typically comprises a mix of woody species such as Sclerocarya birrea and various Vachellia, Senegalia, and Combretum species, alongside an herbaceous layer often dominated by grasses such as Themeda triandra and Panicum maximum. Herbivory by a diverse range of ungulates and fire play significant roles in shaping the ecological dynamics of this region [21,22].
The study within the Lowveld Bioregion was conducted in the Kempiana Nature Reserve (KNR) and focused specifically on the Granite Lowveld (SVl 3) vegetation unit, with a mean annual precipitation of 633 mm [2]. The KNR forms part of the greater Kruger National Park, located approximately 50 km southeast of Hoedspruit. According to Kottek et al. [16], the area falls into the BSh climate classification, indicating an arid area with hot average annual temperatures exceeding 18 °C. Geologically, Kempiana is dominated by ancient granitoid rocks of the Basement Complex, primarily gneiss, granite, and migmatite.

3. Methods

3.1. Sampling Design and Site Selection

This study focused on the herbaceous layer within the Savanna Biome [2]. In each of the three study areas, representing the Central Bushveld, Mopane, and Lowveld bioregions, a relatively homogenous study site, typical of the dominant vegetation unit, was selected. Only one representative study area was sampled per bioregion due to logistical constraints. Therefore, comparisons should be interpreted as between sites representative of the bioregions, rather than definitive bioregional differences, acknowledging the potential for site-specific factors to influence results.
At each of these study sites, three independent 40 × 40 m2 quadrats were established using tape measures. These 40 × 40 m2 quadrats served as the primary sampling sites within each bioregion. These quadrats were spatially separated by tens to hundreds of meters within each study site to capture local variability. A schematic diagram illustrating this multi-level sampling hierarchy is provided in Figure 1.
To ensure representative sampling within each 40 × 40 m2 quadrat, a grid system was overlaid, dividing each quadrat into 400 potential 2 × 2 m2 sampling plots. Subsequently, a random number generator was used to select thirty of these 2 × 2 m2 sampling plots within each 40 × 40 m2 quadrat (Figure 1). This random selection was implemented to minimize the potential for pseudo-replication. Each of the thirty selected 2 × 2 m2 sampling plots was then demarcated for data collection. This sampling design ensured that the herbaceous data were spatially distributed across each of the three 40 × 40 m2 quadrats per study site while maintaining a consistent sampling unit size of 2 × 2 m2 for detailed plant counts and identification.

3.2. Data Collection

Within each of the thirty randomly selected 2 × 2 m2 sampling plots, a thorough inventory of the herbaceous vegetation was conducted. This 2 × 2 m2 plot size was chosen to comprehensively capture the diversity and density of all herbaceous growth forms, including larger or more sparsely distributed forbs, through a total count approach. Starting from a predetermined reference corner within each 2 × 2 m2 sampling plot, all herbaceous plants rooted within its boundaries were meticulously identified and recorded. This detailed and systematic recording of species identity, growth form, and the number of individuals within each sampling plot formed the basis for the subsequent calculation of absolute densities and the analysis of herbaceous community composition across the three savanna bioregions.

3.3. Data Analysis

3.3.1. Calculation of Absolute Density and Species Richness per Quadrat

The absolute density and richness of herbaceous species were calculated at the sampling plot, quadrat, and bioregion levels. The absolute density was taken as the number of individuals per m2 (ind./m2). Species richness was recorded as the count of species per plot, and the total species richness per quadrat or bioregion was the cumulative count of all the unique species recorded within that quadrat or bioregion. Mean values for density and diversity metrics are reported with their associated standard error (SE). These methods for quantifying vegetation density and species richness through quadrat sampling align with standard ecological practices for population and community analysis [23,24].

3.3.2. Comparison of Absolute Density and Species Richness Across Bioregions

To compare the absolute density and species richness of the overall herbaceous layer among the Central Bushveld, Mopane, and Lowveld bioregions, a one-way analysis of variance (ANOVA) was performed. Assumptions of normality and homogeneity of variances were checked, and non-parametric alternatives (e.g., Kruskal–Wallis) were considered if violated. Post hoc tests (Tukey’s HSD) were only conducted to identify significant pairwise differences between bioregions when the ANOVA yielded a statistically significant main effect (p < 0.05).

3.3.3. Shannon Diversity Index (H)

For each of the nine 40 × 40 m2 quadrats, the Shannon diversity index (H) was calculated [24]. One-way ANOVA (or the Kruskal–Wallis test if assumptions were violated) was used to compare the mean Shannon diversity index values among the three bioregions. Post hoc tests were conducted as needed to identify pairwise differences only if the overall ANOVA was statistically significant.

3.3.4. Non-Metric Multidimensional Scaling

To visualize the relationships in herbaceous community composition, a non-metric multidimensional scaling (nMDS) ordination was performed. A species abundance matrix was constructed using the abundance of each species per quadrat and per plot. From this matrix, a Bray–Curtis dissimilarity matrix, accounting for species abundances, was calculated to quantify the ecological dissimilarity between quadrats and between plots. The ordination plot was used to examine quadrat and plot clustering and the separation of bioregions based on their herbaceous communities.

3.3.5. Analysis of Growth Forms

The mean absolute densities of the major growth forms (grasses, forbs, and sedges) per 40 × 40 m2 quadrat were compared across the bioregions using ANOVA (or non-parametric alternatives) to assess functional group differences.

4. Results

This study recorded a total of 196 unique herbaceous species across the three South African savanna bioregions. Specifically, 108 species were recorded in the Central Bushveld, 74 in the Lowveld, and 93 in the Mopane bioregions across all sampled quadrats. The detailed abundance, density, frequency, relative density, relative frequency, and modified importance value index (IVI) values for all observed species within each bioregion are presented in Supplementary Table S1.

4.1. Species Richness and Absolute Density

The mean total herbaceous density per square meter varied across the bioregions: 24.3 ± 2.31 individuals/m2 (ind./m2) in Central Bushveld, 32.0 ± 1.28 individuals/m2 in Lowveld, and 29.6 ± 4.19 individuals/m2 in Mopane (Table 1). The analysis of variance on the total herbaceous density per square meter revealed no statistically significant differences across the three bioregions (F2,6 = 1.89, p = 0.23). Similarly, there was no statistically significant difference in total species richness per m2 across the bioregions (F2,6 = 1.91, p = 0.23).
Analysis of variance at the plot level revealed significant spatial heterogeneity in total herbaceous density within the Central Bushveld (F2,87 = 4.962, p = 0.009) and the Mopane (F2,87 = 7.543, p < 0.001) bioregions. In contrast, there was no significant spatial variation in total herbaceous density within the Lowveld region (F2,87 = 1.25, p = 0.292). There was no significant variation in plot-level species richness within the Central Bushveld region (F2,87 = 1.19, p = 0.311). The Lowveld region showed a marginally significant trend towards spatial heterogeneity in plot-level species richness (F2,87 = 2.79, p = 0.067). In contrast, the Mopane bioregion exhibited significant spatial heterogeneity in plot-level species richness (F2,87 = 6.27, p = 0.003).

4.2. Shannon Diversity Index

The Shannon diversity index (H) per quadrat showed some variation across the bioregions (Table 1). The mean Shannon diversity was highest in the Mopane region (3.39 ± 0.09), followed by the Lowveld (3.32 ± 0.04) and the Central Bushveld (3.11 ± 0.09) regions. Analysis of variance, however, revealed no statistically significant difference in Shannon diversity across the three bioregions (F2,6 = 3.228, p = 0.112). Within-bioregion variation in Shannon diversity at the quadrat level was lowest in the Lowveld region with a standard deviation (SD) of 0.07, followed by the Central Bushveld and Mopane regions (SD = 0.16). At the plot level, the standard deviation of Shannon diversity within each bioregion was 0.32 in the Central Bushveld region, 0.27 in the Lowveld region, and 0.33 in the Mopane region.
Analysis of variance revealed significant differences in plot-level Shannon diversity within the Central Bushveld (F2,87 = 4.01, p = 0.022) and the Mopane (F2,87 = 3.67, p = 0.030) bioregions, indicating spatial heterogeneity in diversity at this finer scale. In contrast, there was no significant variation in plot-level Shannon diversity within the Lowveld region (F2,87 = 1.11, p = 0.335).

4.3. nMDS Ordination

The quadrat-level ordination resulted in a low stress value of approximately 0.045, indicating a good representation of the original dissimilarities in the two-dimensional ordination space. In contrast, the plot-level ordination yielded a considerably higher stress value of approximately 0.219, suggesting a poor fit and that the two-dimensional representation should be interpreted with caution.
The nMDS ordination at the quadrat level (Figure 2a) revealed patterns of dispersion in herbaceous community compositions. Quadrats from the Lowveld and Mopane bioregions formed relatively tight and separate clusters, indicating a degree of similarity in herbaceous community composition within each of these bioregions and clear differences between them. In contrast, the quadrats from the Central Bushveld region were more scattered in the ordination space, suggesting greater variability in community composition among the sampling units within this bioregion. Notably, one Central Bushveld quadrat appeared to be positioned intermediately, exhibiting a similar distance to both the Lowveld and Mopane clusters.
The nMDS ordination at the plot level (Figure 2b, stress = 0.219) showed a more complex pattern. The Central Bushveld plots formed two relatively distinct clusters, each with several outliers. These two clusters were positioned close to each other and did not overlap with the clusters of the Mopane and Lowveld regions. The Lowveld plots formed a relatively tight cluster, with some overlap with the Mopane cluster. The Mopane plots also formed a cluster, but with numerous outliers that exhibited considerable overlap with the Lowveld cluster.

4.4. Growth Form Analysis

The percentage contribution of different growth forms to the total herbaceous density per square meter varied across the bioregions (Table 1). In the Central Bushveld region, grasses (G) contributed 48.3% to the total density, followed by forbs (F) at 47.4% and sedges (Se) at 4.3%. In the Lowveld region, forbs (F) were the dominant growth form, contributing 54.3%, with grasses (G) at 43.2% and sedges (Se) at 2.5%. The Mopane region showed a strong dominance of forbs (F) at 63.8%, followed by grasses (G) at 33.4% and sedges (Se) at 0.7%. Statistical comparisons using ANOVA on the mean absolute densities of each growth form per quadrat revealed no statistically significant differences across the bioregions for grasses (F2,6 = 4.15, p = 0.074), forbs (F2,6 = 2.72, p = 0.145), or sedges (F2,6 = 0.1, p = 0.904). The ten most abundant herbaceous species in each bioregion (Table 2) comprised a mix of growth forms. In the Central Bushveld region, the dominant species were distributed among grasses (four species), forbs (four species), and sedges (two species). The Lowveld region was characterized by a higher proportion of grasses among the most abundant species (six grass species and four forb species), while the Mopane region had a higher representation of forbs (eight forb species and two grass species).
The percentage contribution of different growth forms to the total species richness also varied across the bioregions (Table 1). In the Central Bushveld region, forbs (F) constituted the highest proportion of species richness (63.9%), followed by grasses (G) at 33.3% and sedges (Se) at 2.8%. In the Lowveld region, forbs (F) again represented the largest proportion of species richness (70.3%), with grasses (G) at 27.0% and sedges (Se) at 2.7%. The Mopane bioregion showed a similar pattern, with forbs (F) contributing 66.7% to the total species richness, followed by grasses (G) at 30.1% and sedges (Se) at 3.2%.

5. Discussion

5.1. Herbaceous Density and Species Richness Across Bioregions

A central tenet in savanna ecology is the influence of resource availability, particularly rainfall, on primary productivity and subsequently on vegetation structure and diversity [1]. Our initial expectation was to observe a gradient in herbaceous density and species richness corresponding to the known rainfall patterns across the Lowveld, Central Bushveld, and Mopane bioregions [16,17]. However, the absence of statistically significant differences in both total herbaceous density and species richness per m2 suggests a more nuanced relationship. This finding, while potentially influenced by the limited statistical power inherent in our sampling design (three quadrats per bioregion), aligns with the understanding that savanna ecosystems are shaped by a complex interplay of factors, where soil nutrient availability [3], fire regimes [9,22], and herbivory [7,25] can mediate or even override the direct effects of rainfall. Meta-analyses have also demonstrated that the relationship between productivity and species richness is often unimodal, with diversity peaking at intermediate productivity levels due to trade-offs in competition and resource use [26]. The three bioregions may represent different points along such a complex curve, leading to similar overall density and richness despite variations in rainfall.

5.2. Shannon Diversity Patterns

The lack of significant differences in Shannon diversity across the three bioregions, despite potential variations in environmental conditions and species pools, highlights the multifaceted nature of ecological diversity [24]. Shannon diversity is sensitive to both the number of species (richness) and their relative abundances (evenness) [27]. It is possible that while species richness may exhibit subtle trends related to environmental gradients, differences in species evenness across the bioregions could compensate for these, resulting in statistically similar overall diversity indices. Notably, the Mopane bioregion, despite being the driest site, exhibited the numerically highest mean Shannon diversity (3.39 ± 0.09). This apparent paradox can be ecologically interpreted through several mechanisms. Arid environments can foster high species diversity via strong selection for specialized traits related to water acquisition and survival, promoting niche partitioning among species, particularly within functionally diverse groups such as forbs [28]. Furthermore, pulsed resource availability in arid savannas can lead to rapid growth and reproduction during short favourable periods, allowing more species to coexist [29]. The specific disturbance regimes (e.g., fire and selective herbivory) in arid savannas can also prevent competitive exclusion, thereby promoting higher evenness and diversity. This suggests that even if overall species richness is not dramatically higher, a more equitable distribution of abundances among species might contribute to the higher Shannon diversity. This pattern underscores the complex interplay of aridity, functional traits, and disturbance in shaping community structures [28]. Furthermore, the scale of our sampling (quadrats of a specific size) may have influenced our ability to detect finer-scale differences in diversity that could be apparent at different spatial scales [30,31].

5.3. Community Composition Differentiation

The distinct clustering of herbaceous communities in the Lowveld and Mopane bioregions, as revealed by nMDS, provides strong support for the bioregional classifications proposed by Mucina and Rutherford [2]. These bioregions are characterized by unique combinations of climatic conditions, geology, and associated vegetation types [2]. Our findings suggest that these broad environmental differences have resulted in the development of distinct herbaceous species assemblages, indicating significant beta diversity [31] between these regions. The greater dispersion of Central Bushveld quadrats in the nMDS ordination suggests a higher degree of within-bioregion variability in community composition. This could be attributed to the Central Bushveld region being a more heterogeneous landscape, potentially encompassing transitional zones between other savanna types or exhibiting greater local-scale environmental gradients [32]. Although quantitative support for these specific drivers was not included in the current study, factors such as soil heterogeneity [33] are known to drive such patterns.

5.4. Growth Form Dynamics and the Ecological Importance of Forbs

Our analysis revealed significant patterns in the contribution of different growth forms to the herbaceous layer across the three bioregions. In terms of density, grasses and forbs showed relatively even contributions in the Central Bushveld region, while forbs dominated in the Lowveld region and were particularly prominent in the Mopane bioregion. Sedges consistently contributed a small proportion to the overall density. The statistical analysis indicated that no statistically significant differences were found across the bioregions for grasses (F2,6 = 4.153, p = 0.0738), forbs (F2,6 = 2.716, p = 0.145), or sedges (F2,6 = 0.103, p = 0.904).
Beyond density, forbs consistently emerged as the most speciose group, comprising the largest proportion of total species richness across all three bioregions. Grasses contributed the second highest proportion to species richness, while sedges consistently accounted for a small fraction.
This dual dominance of forbs, contributing the highest percentage to both density (in the Lowveld and Mopane regions) and species richness (across all bioregions), underscores their potentially significant ecological role in South African savannas. As a diverse group encompassing various functional traits, forbs contribute to multiple ecosystem processes [34]. They can enhance nutrient cycling through their often deeper root systems and differing decomposition rates compared to grasses [5]. Furthermore, forbs are crucial for supporting a wide array of herbivores, including insects and small mammals, thus contributing to overall food web complexity [35,36]. The high species richness of forbs suggests a wide array of ecological niches they occupy and the diverse functions they perform within these savanna ecosystems. The numerically higher contribution of forb density in the Mopane bioregion (63.8%) compared to Central Bushveld (47.4%), while not statistically significant across all bioregions, warrants specific discussion as it appears to contradict general expectations of lower overall herbaceous biomass in more arid zones [15]. This pattern might be explained by specific adaptive strategies of forbs in arid environments, such as deeper root systems for accessing water, greater phenological plasticity for rapid growth during resource pulses, and efficient nutrient acquisition in challenging soils [29]. These adaptations could confer a competitive advantage in the Mopane region’s hot and arid conditions, allowing forbs to maintain higher densities or relative contributions compared to grasses. The consistently low contribution of sedges to both density and richness suggests that they might play a more specialized or localized role in these herbaceous communities. Future research should continue to explore the functional roles of different growth forms and the environmental drivers of their abundance and diversity in these savannas [37].

5.5. Within-Bioregion Spatial Heterogeneity

Greater spatial heterogeneity in plot density and Shannon diversity in the Central Bushveld region compared to the Lowveld and Mopane regions suggests differing fine-scale ecological processes. Patchiness in resource availability, localized disturbances, and microtopography can drive this heterogeneity [38,39], with the Central Bushveld region potentially exhibiting a more dynamic local environment. Understanding these fine-scale patterns is increasingly recognized as important for maintaining biodiversity and ecosystem functioning [40,41,42,43].
Our analysis revealed significant spatial heterogeneity in the herbaceous layer at both bioregional and finer scales. We found significant variation in total herbaceous density across quadrats within the Central Bushveld and Mopane regions, suggesting patchiness in biomass, possibly due to localized differences in soil properties [44], topography [32], or disturbances such as grazing and fire [21,45]. In contrast, the Lowveld region showed a more uniform density distribution. Subplot species richness also varied significantly across Mopane quadrats, indicating patchiness, which is potentially linked to the same factors influencing density. The Central Bushveld region showed uniform richness, and the Lowveld region showed a marginal trend toward heterogeneity.
These findings highlight the importance of considering spatial scale. While bioregional differences provide a broad overview, finer-scale heterogeneity reveals ecological processes and informs management. The Mopane region, with heterogeneity in both density and richness, might have a more dynamic environment. The uniform Lowveld region might reflect more homogenous conditions. Within-bioregion patterns may interact with growth form variation; for example, forb dominance in Mopane, coupled with high heterogeneity, could indicate forb responsiveness to localized resource or disturbance variation.
While broad environmental contexts, including general rainfall patterns, differ among the chosen bioregions, it is important to acknowledge that our study sampled only one representative site per bioregion. This design, while providing valuable comparative data, means that our inter-bioregional findings should be interpreted with caution, as site-specific factors (e.g., local management, microclimates, and unique disturbance histories) could potentially confound broader bioregional signals. A comprehensive multivariate analysis explicitly modeling the direct effects of dynamic environmental drivers, such as precise rainfall amounts during the study period, fire frequency, and quantified herbivore biomass or utilization, would require a dedicated study design focused on capturing these variables contemporaneously with vegetation sampling across a broader range of environmental variation [3,46]. Such an analysis, while providing valuable mechanistic insights into the drivers of herbaceous community structure, was beyond the scope and data collection focus of this initial comparative assessment across distinct bioregional classifications.
Furthermore, collecting spatially explicit data on dynamic processes such as fire history and herbivore movement or the localized impact at the resolution of our sampling quadrats is logistically challenging and often relies on long-term ecological monitoring infrastructures or targeted experimental setups [47,48]. Therefore, while we acknowledge that these factors undoubtedly contribute to the observed patterns, including within-bioregion heterogeneity, robust statistical modeling of their individual and interactive effects would necessitate a dataset specifically designed for that purpose, which differs from the objective of characterizing herbaceous layer attributes within defined bioregional units as presented here. Future research building upon these observed patterns could quantify these environmental variables to unravel their specific roles in shaping savanna herbaceous communities.

6. Conclusions

In conclusion, our analysis of the herbaceous layer across three South African savanna bioregions revealed a complex pattern of similarities and differences in density, species richness, and community composition. Notably, the growth form analysis highlighted that forbs consistently dominated the species richness across all bioregions, representing the largest proportion of the total number of herbaceous species. Furthermore, forbs were also the dominant growth form in terms of density in the Lowveld and Mopane bioregions, contributing a substantial proportion to the overall herbaceous biomass. While numerical differences in grass density were observed across the bioregions, no statistically significant difference was found, and their contribution to species richness was consistently lower than that of forbs. Sedges consistently played a minor role in both density and species richness. These findings underscore the potentially critical ecological importance of forbs in these savanna ecosystems, not only in terms of biomass contribution but also in driving the diversity of the herbaceous layer [5,36]. Understanding the factors that influence the distribution and abundance of different growth forms, particularly the diverse forb community, is crucial for the effective conservation and management of these valuable savanna ecosystems. Future research should continue to investigate the functional roles of these growth forms and the drivers of community assembly to inform strategies that maintain their biodiversity and ecological integrity.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17070475/s1, Table S1: Detailed abundance, density, frequency, relative density, relative frequency, and modified importance value index (IVI) values for all observed species within each bioregion..

Author Contributions

A.A.B.: Conceptualization; methodology; data collection; sample analysis; data analysis; validation; data curation; writing—the initial draft; writing—revisions. W.J.M.: Conceptualization; methodology; validation; student supervision; project leadership; project management; writing—revisions. B.K.R.: Conceptualization; methodology; data analysis; data curation; student supervision; project leadership; project management; writing—revisions. All authors have read and agreed to the published version of the manuscript.

Funding

This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

Data used in the preparation of this manuscript are available as Supplementary Table S1.

Conflicts of Interest

The authors declare no conflicts of interest.

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Diversity 17 00475 g001
Figure 1. Sampling framework showing the hierarchy between the study area, sampling sites, and sampling plots.
Figure 1. Sampling framework showing the hierarchy between the study area, sampling sites, and sampling plots.
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Figure 2. nMDS plots of quadrats (a) and plots (b) showing the level of differentiation between the Central Bushveld (CB), Lowveld (LV), and Mopane (MV) regions.
Figure 2. nMDS plots of quadrats (a) and plots (b) showing the level of differentiation between the Central Bushveld (CB), Lowveld (LV), and Mopane (MV) regions.
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Table 1. Results of the species richness, density, and Shannon diversity index in the Central Bushveld, Mopane, and Lowveld regions.
Table 1. Results of the species richness, density, and Shannon diversity index in the Central Bushveld, Mopane, and Lowveld regions.
Growth FormsCentral BushveldLowveldMopane
Species
Richness		Grasses362028
Forbs695262
Sedges323
Overall1087493
Density
(ind./m2)		Grasses11.7 ± 1.4113.8 ± 0.4410.1 ± 0.59
Forbs11.5 ± 0.4617.4 ± 2.0219.5 ± 3.82
Sedges1.5 ± 0.531.6 ± 0.571.86 ± 0.50
Overall24.3 ± 2.3132.0 ± 1.2829.6 ± 4.19
Shannon Diversity Index 3.11 ± 0.093.32 ± 0.043.39 ± 0.09
Table 2. Densities of the ten herbaceous species with the highest densities at the Letlapa Pula Game Reserve (LPGR; Central Bushveld), Selati Game Reserve (SGR; Mopane), and Kempiana Nature Reserve (KNR; Lowveld). In the table, Ab is the abundance, mean Ab is the mean abundance per sampling plot, SD is the standard deviation, CV is the coefficient of variation, CB is the Central Bushveld region, MV is the Mopane region, and LV is the Lowveld region.
Table 2. Densities of the ten herbaceous species with the highest densities at the Letlapa Pula Game Reserve (LPGR; Central Bushveld), Selati Game Reserve (SGR; Mopane), and Kempiana Nature Reserve (KNR; Lowveld). In the table, Ab is the abundance, mean Ab is the mean abundance per sampling plot, SD is the standard deviation, CV is the coefficient of variation, CB is the Central Bushveld region, MV is the Mopane region, and LV is the Lowveld region.
Study AreaSpeciesFamilyAbMean AbSDCVDensity (ind./m2)
LPGR
(CB)		Melinis repensPoaceae6897.6614.001.831.91
Heteropogon contortusPoaceae6557.288.541.171.82
Becium obovatumLamiaceae4364.847.151.481.21
Waltheria indicaMalvaceae3904.3312.852.961.08
Limeum viscosumLimeaceae3834.267.421.741.06
Tragus berteronianusPoaceae3704.117.871.911.03
Tephrosia longipesFabaceae3543.936.381.620.98
Setaria sphacelata var. sphacelataPoaceae3143.494.551.300.87
Coleochloa setiferaCyperaceae2973.3010.003.030.83
Cyperus obtusiflorusCyperaceae2893.216.912.150.80
SGR
(MV)		Hoffmannseggia burchelliiFabaceaea7838.708.6900.9992.175
Indigofera oxytropisFabaceaea7418.238.5211.0352.058
Zornia milneanaFabaceaea6076.7413.3831.9841.686
Digitaria erianthaPoaceae5446.046.0741.0051.511
Waltheria indicaMalvaceae5045.6012.3242.2011.400
Becium obovatumLamiaceae4765.2913.2092.4971.322
Evolvulus alsinoidesConvolvulaceae4745.275.6231.0681.317
Chamaecrista mimosoidesFabaceaea4585.097.4571.4651.272
Urochloa mosambicensisPoaceae4525.025.0771.0111.256
Ocimum americanumLamiaceae4254.7210.3752.1971.181
KNR
(LV)		Indigofera oxytropisFabaceae112512.5011.540.923.13
Digitaria erianthaPoaceae95610.626.670.632.66
Panicum maximumPoaceae7858.726.670.762.18
Urochloa mosambicensisPoaceae7268.075.240.652.02
Hibiscus calyphyllusMalvaceae6206.8912.871.871.72
Bidens bipinnataAsteraceae5125.6912.732.241.42
Oldenlandia herbaceaRubiaceae4705.229.751.871.31
Perotis patensPoaceae3644.047.181.771.01
Aristida congesta subsp. congestaPoaceae3373.745.821.560.94
Pogonarthria squarrosaPoaceae3243.605.181.440.90
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Biko’o, A.A.; Myburgh, W.J.; Reilly, B.K. Contrasting Herbaceous Communities in South African Savannas: A Comparative Analysis of Density, Composition, and Diversity Across Three Bioregions. Diversity 2025, 17, 475. https://doi.org/10.3390/d17070475

AMA Style

Biko’o AA, Myburgh WJ, Reilly BK. Contrasting Herbaceous Communities in South African Savannas: A Comparative Analysis of Density, Composition, and Diversity Across Three Bioregions. Diversity. 2025; 17(7):475. https://doi.org/10.3390/d17070475

Chicago/Turabian Style

Biko’o, Armand Arthur, Willem Johannes Myburgh, and Brian Kevin Reilly. 2025. "Contrasting Herbaceous Communities in South African Savannas: A Comparative Analysis of Density, Composition, and Diversity Across Three Bioregions" Diversity 17, no. 7: 475. https://doi.org/10.3390/d17070475

APA Style

Biko’o, A. A., Myburgh, W. J., & Reilly, B. K. (2025). Contrasting Herbaceous Communities in South African Savannas: A Comparative Analysis of Density, Composition, and Diversity Across Three Bioregions. Diversity, 17(7), 475. https://doi.org/10.3390/d17070475

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