Phytochemical, pharmacological, and genetic diversity studies on the tree species
The Southern African Miombo-Mopane woodlands are globally considered as ecosystems with irreplaceable species endemism, being the most important type of vegetation in the region. Among the approximately 8500 plant species, legume trees play a crucial role in biodiversity dynamics, being also key socioeconomic and environmental players. From the ecological point of view, they contribute significantly to ecosystem’s stability as well as to water, carbon, and energy balance. Additionally, legume species represent an immensurable source of timber and nontimber products. Research in Miombo-Mopane biodiversity has been mainly focused on the analysis of ecosystem drivers by means of ecological parameters and models, lacking interdisciplinary with relevant cross-cutting tools, such as the application of molecular markers to assess genetic diversity within the region. In this chapter, the applications and biodiversity dynamics of typical legume species from Miombo (Brachystegia spp., Julbernardia globiflora, and Pterocarpus angolensis) and Mopane (Colophospermum mopane) are reviewed. Gaps and challenges are also brought forward in the context of the lack of genetic diversity assessments and the need of an effective and coordinated network of interdisciplinary research.
- Brachystegia spp.
- Colophospermum mopane
- Julbernardia globiflora
- Pterocarpus angolensis
Africa has a vast array of indigenous legumes (Fam. Leguminosae or Fabaceae), ranging from small annual herbs to large trees . The potential of native legumes for multipurposes is high but poorly exploited . This is particularly the case of the woody species from the Miombo-Mopane woodlands, one of the five ecozones (together with Amazonia, Congo, New Guinea, and the North American deserts) with irreplaceable species endemism . Miombo is the most widespread and important type of vegetation in Southern Africa covering approximately 2.4 million km2 across seven countries: Angola, Democratic Republic of Congo (DR Congo), Malawi, Mozambique, Tanzania, Zambia, and Zimbabwe [4, 5]. Mopane extends over 0.48 million km2 in Botswana, Malawi, Mozambique, Namibia, South Africa, Swaziland, Zambia, and Zimbabwe, constituting the second most important type of vegetation in the region . The woodlands are the main source of woody species and a wealth of resources to the livelihood systems of millions of rural and urban dwellers that depend on these ecosystems to meet their food, health, energy, and housing needs [6, 7]. Environmentally, Miombo and Mopane add to biodiversity and have a global impact in energy, water, and carbon balances.
Miombo woodlands (Figure 1A) are dominated by legume trees belonging to three Caesalpinoideae genera:
The ecological dynamics of Miombo-Mopane is strongly influenced by a combination of climate, disturbances (e.g., drought, fire, grazing, and herbivory primarily by elephants), and human activities [12, 13]. The growing population in the region over the last 20–25 years has resulted in increased woodlands degradation and deforestation. Slash-and-burn agriculture and charcoal production are the major causes of forest loss and degradation [14–16]. Additionally, the region is experiencing several major investments in mining, commercial agriculture, and infrastructures, which have further increased the pressure on the woodlands .
Changes in the global climatic pattern constitute another major threat for these ecosystems. They are mainly associated with more extreme wet and dry seasons as well as with extreme temperatures, which may change disturbance regimes (e.g., fire, shifting cultivation) and thus the prevailing biodiversity status [18–22]. For example,  predicts 5–15% reduction in precipitation for Southern Africa, while Green and coauthors  hypothesize that the combined effect of climate changes and disturbances may cause the loss of ca. 40% of the woodlands by the middle of the century. In line with these predictions, field studies combined with remote sensing and Geographic information system (GIS) methodologies indicated a decline in vegetation richness of 10–30% across Sahel and a southward shift of Sahel, Sudan, and Guinea zones due to shifts in temperature and precipitation regimes . This may impose changes in biodiversity and biomass with associated modifications on the pattern of goods and services offered by the ecosystems. Under this scenario, several researchers are currently investigating (i) the impacts of the different ecosystem drivers on the woodland biodiversity, (ii) the capacity of biodiversity to supply and underpin goods and services, (iii) the patterns of genetic diversity of important species across environmental gradients, (iv) how different land cover types affect the existing patterns of biodiversity, and (v) how the changes in biodiversity will affect the availability and accessibility of resources to rural and urban dwellers.
In this review, we present the current efforts, gaps, and challenges toward biodiversity conservation of key legume trees from the Miombo and Mopane woodlands, respectively:
2. Brief description and major applications of typical Miombo-Mopane species
Brachystegiaspp. (subfamily Caesalpinioideae)
|Species||Phytochemical and pharmacological studies||Genetic diversity studies|
|Leaves: antibacterial ||ISSR markers |
|Leaves: anti-inflammatory |
|No studies reported|
|RAPD [79, 80]|
|Stem, stem bark, leaves: anthelmintic |
|Seeds: antibacterial |
|Stem bark: antibacterial and anti-inflammatory; lack of mutagenicity |
|Stem bark, leaves: anthelmintic, antibacterial, and cytotoxic |
|Stem bark: antibacterial; epicatechin and derivatives identified |
|Stem bark: antibacterial; leaves, stem bark: antifungal, HIV-1 reverse transcriptase inhibitory |
|Leaves, stem bark: anti-inflammatory |
|Stem bark, roots: antibacterial; tannins and saponins identified |
|Bark and seeds: significant cytotoxicity against a human breast cancer cell line |
Heartwood: mono-, di-, and triflavonoids; leaves: beta-sitosterol and stigmasterol; aerial part: essential oils that comprise mainly alpha-pinene and limonene; bark and seeds: diterpenes, including dihydrogrindelaldehyde 
|Allozyme markers |
|Seeds: anti-inflammatory and antioxidant activities |
|Bark and seeds: significant cytotoxic activity against a human breast cancer cell line (aldehyde) |
Bark and seeds: three new diterpenes isolated 
|Oligomeric flavonoids |
|Leaves: antimicrobial activity |
Seed husks and leaves: five labdane (1–5), an isolabdane (6), and five clerodane diterpenoids (7–11) 
Julbernardia globiflora(subfamily Caesalpinioideae)
(subfamily Papilionoideae) Pterocarpus angolensis
(subfamily Caesalpinioideae) Colophospermum mopane
3. Diversity and population structure studies
Miombo and Mopane woodlands face major threats related to climate, human, and animal pressure which in the midterm may reduce tree species abundances and, thus, ecosystem services [17, 43, 50]. Thus, understanding the interaction between the main woodland’s drivers is crucial for the development of effective and sustainable management strategies for biodiversity conservation and resource use. In this section we focus on diversity and structure of the key legume trees referred above, i.e.,
In the Niassa National Reserve (NNR), one of the most pristine and least disturbed areas of Africa’s deciduous Miombo woodlands, notable alterations in vegetation structure and composition were reported in areas with high fire (mostly anthropogenic) frequency [51, 52]. These included a decrease in woody parameters and a replacement of typical Miombo species (
Despite the factors referred above, natural factors, i.e., soil fertility, topography, and local hydrology, are also important determinants of biodiversity variation, although these factors are less studied [54, 55]. The vulnerability of the woodlands to climate changes is particularly high . Species distribution models developed for
Contrary to Miombo, Mopane woodlands are constituted by nearly monospecific stands of
Genetic diversity is the basis for stability and survival under the ever-changing environments. Populations with high levels of genetic variation offer a diverse gene pool from which breeding and conservation programs can be designed. The over exploitation of the reported species may threaten their genetic diversity in the future and hence might limit their ecological and evolutionary development. Therefore, genetic diversity and structure studies are of utmost importance for designing appropriate conservation strategies.
The use of molecular markers constitutes an effective approach to evaluate genetic variation within and between species and populations, because they are expedited and precise and are not affected by the environmental processes. Polymerase chain reaction (PCR)-based markers, like random amplified polymorphic DNA (RAPD), Amplified fragment length polymorphism (AFLPs), inter-simple sequence repeats (ISSRs), simple sequence repeats (SSRs), or single-nucleotide polymorphisms (SNPs), are commonly used in plant science for a wide variety of purposes such as genome mapping, gene tagging, phylogenetic analysis, taxonomy, marked-assisted selection, and genetic diversity studies [64–73]. However, the analysis of the genetic variation and structure is an incipient issue in Miombo and Mopane research. Regarding the species selected for this review, up to our best knowledge, only four reports are available in the scientific literature (Table 1).
Maquia et al.  have used ISSR markers to assess the genetic diversity in
RAPD markers were used to characterize accessions of
Villoen and collaborators  have used 13 allozyme markers to analyze five populations of
It should be highlighted that except for , the studies of [79, 80, 83] did not address the impact of environmental and/or anthropogenic drivers, which are determinant to understand ecosystem’s dynamics. In fact, the use of molecular markers to assess tropical tree biodiversity dynamics across environmental gradients is an issue that deserves more attention. Using ISSR markers,  examined the effect of three different environmental gradients on the genetic diversity of the semishrub legume
Altogether, these studies [70–87] reveal that the response of tropical trees to environmental and anthropogenic pressure is highly variable and that, in general, most of the species are resilient to extreme soil and climate conditions, being able to retain high levels of genetic diversity. This in turn highlights the relevance of integrative molecular analyses at a regional scale, to understand the mechanisms of species adaptation and evolution within the context of climate changes. A good example of the potentialities of such studies is the work developed in
4. Concluding remarks
African woodlands support the livelihoods of millions of rural and urban people, providing valuable sources of wood, edible products, fibber and related products, insect products (honey and beeswax, edible insects), and medicinal plants, among others.
Based on the available scientific information, it is our understanding that major gaps and challenges need urgently to be addressed. The development of coordinated research throughout the region to assess genetic diversity and structure as well as to define common conservation strategies, adapted to each country needs and facilities, should be prioritized. For that, effective networking between Southern African institutions and their partners from Europe and the USA seems to be the most appropriate approach. However, such interactions are not reflected in a considerable number of scientific publications. It is our conviction that collaborative work is the best way to consolidate and/or promote partnerships, resulting in mutual benefits, e.g., scientific excellence, critical thinking, team playing, access to funding, broadening information sharing to prompt innovation, translation of knowledge from “local to global,” and from “global to local” contexts. The involvement of the Miombo Network for Southern Africa would be of utmost importance to incorporate key issues, such as genetic diversity and bioprospection in Miombo and also Mopane, in its research strategy plan. The inclusion of these two issues in Miombo and Mopane research will not only ensure the sustainable use and conservation of key species but also allow the establishment of modern biotechnology platforms toward the incorporation of the most valuable species into bio-based economy schemes.
The authors thank the Portuguese Cooperation through Camões, Instituto da Cooperação e da Língua, Fundação para a Ciência e Tecnologia through the contribution to IRRI/CGIAR and to Research unit LEAF (UID/AGR/04129/2013) and Maria Cristina Duarte for providing the photograph for Figures 2B and 2D.
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