Open access peer-reviewed chapter

Biodiversity Studies in Key Species from the African Mopane and Miombo Woodlands

Written By

Isabel Moura, Ivete Maquia, Alfan A. Rija, Natasha Ribeiro and Ana Isabel Ribeiro-Barros

Reviewed: 11 November 2016 Published: 01 March 2017

DOI: 10.5772/66845

From the Edited Volume

Genetic Diversity

Edited by Lidija Bitz

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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
  • Miombo
  • Mopane
  • Pterocarpus angolensis
  • woodlands

1. Introduction

Africa has a vast array of indigenous legumes (Fam. Leguminosae or Fabaceae), ranging from small annual herbs to large trees [1]. The potential of native legumes for multipurposes is high but poorly exploited [2]. 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 [3]. 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 [4]. 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: Brachystegia (Miombo in Swahili), Julbernardia, and Isoberlinia. On the contrary, Mopane woodlands (Figure 1B) are characterized by monospecific stands of Colophospermum mopane (Benth.) Léonard (also belonging to the Caesalpinoideae subfamily of legumes) [6, 8]. From the environmental point of view, these plants are determinant for energy, carbon, and water balance [9, 10]. At the socioeconomic level, Miombo-Mopane legumes are key providers of goods and services [11]. The woodlands are also very important to the national economies as they provide timber for exportation [5].

Figure 1.

Images of Miombo and Mopane woodlands. (A) Miombo regrowth in burned area; (B) Shrub-like C. mopane specimens.

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 [1416]. Additionally, the region is experiencing several major investments in mining, commercial agriculture, and infrastructures, which have further increased the pressure on the woodlands [17].

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 [1822]. For example, [19] predicts 5–15% reduction in precipitation for Southern Africa, while Green and coauthors [23] 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 [21]. 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: Brachystegia Benth., Julbernardia globiflora (Benth.) Troupin, Pterocarpus angolensis DC. (Miombo), and C. mopane (Mopane).


2. Brief description and major applications of typical Miombo-Mopane species

2.1. Brachystegia spp. (subfamily Caesalpinioideae)

Brachystegia (Miombo) is a large genus of trees confined to tropical Africa, representing major components of deciduous (Miombo) woodlands of central and Southern Africa. This is a taxonomically difficult genus, since some species hybridize freely with others, making classification difficult [24]. The most frequent species in dry Miombo woodland are Brachystegia boehmii and Brachystegia spiciformis. Brachystegia species are generally slender and graceful with a clean bole, a crown of handsome delicate leaves (Figure 2A) and a fibrous stem bark. The genus as a whole is noted for the great range of red colors of the young foliage, later turning into various shades of green [24]. Brachystegia species play an important role in formal and informal economies in the region. For example, barks, which are particularly fibrous, are commonly used for construction, being especially favored for weaving, fishnet, beds, and rope making. Bark is easily peeled from a number of species and is used for fabricating beehives. The ash is used as fertilizer in shifting cultivation, and the wood is used for domestic tools, canoes, firewood, and charcoal; timber is usually heavily susceptible to borers but when treated is suitable for several purposes [6, 25]. Brachystegia is also an important nectar-producing genus for apiculture [26]. Additionally, several species of Brachystegia are used in traditional medicine [2729], with antimicrobial, anti-inflammatory, and antidiabetic activities (Table 1) [3032].

SpeciesPhytochemical and pharmacological studiesGenetic diversity studies
Brachystegia boehmii Taub.Leaves: antibacterial [30]ISSR markers [70]
Leaves: anti-inflammatory [32]
Julbernardia globiflora (Benth.) TroupinNo studies reported
Pterocarpus angolensis DCAntischistosomal [90]RAPD [79, 80]
Stem, stem bark, leaves: anthelmintic [40]
Seeds: antibacterial [91]
Stem bark: antibacterial and anti-inflammatory; lack of mutagenicity [92]
Stem bark, leaves: anthelmintic, antibacterial, and cytotoxic [41]
Antibacterial [93]
Stem bark: antibacterial; epicatechin and derivatives identified [37]
Stem bark: antibacterial; leaves, stem bark: antifungal, HIV-1 reverse transcriptase inhibitory [38]
Leaves, stem bark: anti-inflammatory [94]
Stem bark, roots: antibacterial; tannins and saponins identified [39]
Colophospermum mopane (Benth.)Bark and seeds: significant cytotoxicity against a human breast cancer cell line [48]
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 [48]
Allozyme markers [83]
Seeds: anti-inflammatory and antioxidant activities [95]
Bark and seeds: significant cytotoxic activity against a human breast cancer cell line (aldehyde) [96]
Bark and seeds: three new diterpenes isolated [96]
Oligomeric flavonoids [97]
Leaves: antimicrobial activity [49]
Seed husks and leaves: five labdane (1–5), an isolabdane (6), and five clerodane diterpenoids (7–11) [49]

Table 1.

Phytochemical, pharmacological, and genetic diversity studies on the tree species Brachystegia boehmii Taub., Julbernardia globiflora (Benth) Troupin, Pterocarpus angolensis DC., and Colophospermum mopane (Benth).

Figure 2.

Details of the leaves from (A) Brachystegia boehmii, (B) Julbernardia globiflora (Barbosa and Carvalho, 2825, 1949 LISC Herbarium), (C) Pterocarpus angolensis, and (D) Colophospermum mopane (B and D credits to Maria Cristina).

2.2. Julbernardia globiflora (subfamily Caesalpinioideae)

J. globiflora, known as Mnondo [6], is a common species of sandy woodlands, often codominant with Brachystegia species. It is distributed from DR Congo, Burundi, and Tanzania southward to Botswana, Zimbabwe, and Mozambique [33]. J. globiflora is usually recorded as a small- to medium-sized deciduous tree with a rounded crown that can grow up to 18 m in height, with alternate and compound leaves (Figure 2B) and thick fire-resistant bark [6]. Like Brachystegia species, J. globiflora is a useful local source of timber and nontimber products. The wood is hard, with a heartwood resistant to borers, and is widely used as a general-purpose timber and also for the construction, tool handles, mortars, and canoes as well as fuelwood and for making charcoal [24, 25, 33]. Ropes, fishnets, and other items are made from the bark fiber, which is fairly good, although of poorer quality than that of the Brachystegia spp. [6]. The bark yields tannin used for dyeing [33]. The tree is an important fodder species for early-season browsers [34]; it is also a bee forage, yielding honey of very high quality. Together with B. boehmii, J. globiflora is among the most important hive trees and nectar sources [26]. The leaves of J. globiflora are important food for edible caterpillars [33]. Although roots, barks, and leaves of J. globiflora have been recorded to be toxic, various plant parts are used in traditional medicine, mainly externally to treat ailments such as snake bites, leprosy, and conjunctivitis [28, 33].

2.3. Pterocarpus angolensis (subfamily Papilionoideae)

P. angolensis (commonly known in English as bloodwood, kiaat, and African teak) [6] is a medium-sized to large-sized tree up to 16 m in height but reaching 20 m under ideal conditions with a spreading crown and a single trunk. P. angolensis is widespread in southern tropical Africa, from Angola, DR Congo, and Tanzania south to northeastern South Africa and Swaziland [35]. The bark is dark gray to brown, rough, and longitudinally fissured. The drooping leaves have 5 to 9 pairs of subopposite to alternate leaflets (Figure 2C). The flowers are orange-yellow in large, branched sprays. The fruit is a very distinctive, indehiscent, and circular pod holding 1–2 small seeds [24]; the leaves are browsed by elephant and kudu; and fruits are eaten by baboons, monkeys, and squirrels [6]. P. angolensis serves a number of purposes, both utilitarian (as multipurpose timber, dye, forage) and in African folk medicine. It is the main Miombo and one of the most valuable timber trees in Africa heavily sought after for export and local use. The heartwood is golden or reddish brown, makes high-quality furniture, shrinks very little in drying, and is very resistant to borers and termites [24]. The wood is also used for construction, carpentry, flooring, boats, and wood carving. It is occasionally used for firewood [35]. The sap from the wood makes a permanent red stain on clothing and can be used as a dye. The powdered red inner bark of the roots, mixed with fat, is used to anoint bodies and faces by some ethnic groups in northern South-West Africa [6]. Traditionally, all parts of the plant are used for human and animal health purposes. For example, the bark is used as a powerful astringent to treat diarrhea, heavy menstruation, nosebleeding, headache, stomachache, schistosomiasis, sores, and skin problems; the root is believed to cure malaria, blackwater fever and gonorrhea [6, 24, 28, 35, 36]. Research studies reveal promising results concerning their antibacterial [3739], antifungal [38], anthelmintic [40, 41], and HIV-1 reverse transcriptase inhibitory properties [38] (Table 1). At the environmental level P. angolensis is able to establish nitrogen-fixing symbiosis with rhizobium bacteria and therefore relevant for soil fertilization. It is also planted for soil conservation, for dune fixation, and as live fencing [35]. Flowering trees are good sources of pollen and nectar [35].

2.4. Colophospermum mopane (subfamily Caesalpinioideae)

C. mopane is the single species in the genus Colophospermum, which only occurs in Africa, being characteristic of hot and dry river valleys, where it is often dominant and may form almost pure stands over large areas—the “Mopane woodlands” [42]. Commonly known as mopane, it is widespread in Southern Africa, where it occurs in Zambia, Malawi, southern Angola, northern Namibia, northeastern Botswana, Zimbabwe, southern Mozambique, and northern South Africa [43]. Mopane is adapted to a wide variety of soils and temperatures, presenting different growth forms, from shrub-like to tall slender trees. It is deciduous or sometimes semideciduous, as water availability determines leaf drop [44], with beautiful autumn and spring colors. The highly sclerophyllous compound leaves (Figure 2D) have two leaflets, large and butterfly-shaped, which fold together in the hottest time of day [6]. Leaves and fruits are very glandular and smell strongly as turpentine. As a fodder species, mopane is vital in areas of low rainfall. Like most Miombo legume species, C. mopane is a true multipurpose tree, not only important for its wood but also as source of medicine, forage, and edible caterpillars [43]. Cattle and wild animals browse the foliage eagerly and sometimes eat the dry leaves and seeds from the ground [6]. Seeds are consumed by humans as famine food [43]. Mopane heartwood is very hard, and it is known to be very durable and resistant to insect damage. It is traditionally used for posts and poles in the construction of houses and palisade fences. Mopane wood accounts for more than 90% of the wood used for living and storing huts in large parts of Southern Africa [43]. It is suitable for flooring and for carving and is excellent for turned objects, and to a lesser extent, due to its weight and hardness, it is used for joinery and furniture. Mopane gives excellent firewood and makes high valuable charcoal [43]. The rough bark is used to make twine and for tanning [6, 43]. Larvae of mopane moth (Gonimbrasia belina) feed on the mopane leaves. Those large caterpillars are rich in protein and considered a local delicacy [6]. Mopane worms are also traded to generate income providing a good economic return [45]. The tree also acts as a food plant for a wild silk moth (Gonometa rufobrunnea). Cocoons of the moth are harvested as wild silk and processed to make cloth [43]. C. mopane is an important medicinal species, and different parts of the plant are used in the preparation of traditional remedies [43, 46, 47]. Research into Mopane-active compounds revealed biological activities with potential for human and animal health [48, 49] (Table 1).


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., Brachystegia spp., J. globiflora, P. angolensis, and C. mopane.

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 (J. globiflora and Brachystegia spp.) by subdominant species (Combretum spp., Terminalia sericea Burch. ex DC, and Diplorhynchus condylocarpon (Muell. Arg.) Pichon [51]. In southwestern Tanzania, [50] reported a tree species diversity score (Shannon-Wiener diversity) of 3.44 versus 2.86 in the Miombo woodlands subjected to low and high rates of resource use, respectively. In areas of moderate resources utilization, tree diversity was maintained, but the structure of the vegetation showed a reduction of Class 1 (diameter at breast height (DBH) < 10 cm) trees, indicating low recruitment. Key Miombo species from the genera Brachystegia and Julbernardia were present in all sites, but the frequency of the former declined by 60% from low to high utilization sites. The resilience of Julbernardia spp. to disturbances might be due to vigorous resprouting after cutting as observed in J. globiflora [33]. On the other hand, P. angolensis adult trees were harvested throughout the study site, and only immature specimens were recorded, suggesting that it is commercially extinct for the foreseeable future [50]. P. angolensis is widespread in many parts of Southern Africa; however, overexploitation endangers natural populations in all countries. None of the four species considered in this paper is listed in the Convention on International Trade in Endangered Species (CITES) of Wild Fauna and Flora Appendices. Only P. angolensis is reported by the International Union for Conservation of Nature (IUCN) as being near threatened. Technically, it does not meet the Red List criteria of a vulnerable or endangered species, but is close to qualifying for vulnerable [53].

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 [19]. Species distribution models developed for P. angolensis under two different climate change scenarios suggest that potential and realized distributions are very similar in Southern Africa, except for Madagascar where the species could grow but does not occur, and that species distribution may be particularly threatened in Namibia and Botswana [56]. Accordingly, pollen records from Brachystegia spp. suggest a retraction of Miombo in ca. 450 km over the past 6000 years [57] and that such changes in population dynamics may be associated with shifts in temperature and moisture regimes [7]. Based on the historical range shifts in B. spiciformis, an ecological niche retraction of ca. 31–47% in the continuous Miombo woodlands from Zimbabwe and southern Mozambique was predicted by 2050 [58]. However, considering the resilience of B. spiciformis to extreme environments (e.g., low precipitation, high temperatures), such retraction may be less exaggerated than the model predictions, which no account with genetic adaptation [58].

Contrary to Miombo, Mopane woodlands are constituted by nearly monospecific stands of C. mopane [59], resulting in extensive areas with low compositional and structural diversity [60]. Due to the same factors mentioned for the Miombo woodlands, some parts of Southern African Mopane woodlands are experiencing a decline in natural stands [43]. For example, [61] analyzed the effects of heavy land utilization (mainly grazing) on vegetation structure communal woodlands of the Mopane Bioregion of South Africa. The authors observed prominent effects in plant structure, i.e., reduced canopy and height, transforming the woodland into a shrubland. A significant decrease in biomass in post–boom charcoal production in southern Mozambique was also reported [62]. Govender et al. [63] analyzed the effect of two fire management strategies in one of the most important areas of mopane, the Great Limpopo Transfrontier Park (GLTP), which crosses two countries: South African component Kruger National Park (KNP) and Mozambique Limpopo National Park (LNP). KNP has a very structured and intensive fire management (FM), while in LNP, the fire regime is unstructured and highly associated to the large number of people living within its limits. Even so, in LNP natural fire frequency did not affect the ecological weight of Mopane, whereas in the KNP, frequent managed fires reduced the ecological value in 100–200 [63]. These results indicate that natural fire regimes are important to maintain the ecosystem’s equilibrium. Although less studied, C. mopane biodiversity shifts may also be affected by the environmental conditions. Stevens et al. [60] used MAXENT modeling to investigate which environmental variables may determine the distribution limits of C. mopane within Southern Africa. According to the results, both nonclimatic (dryness or hours of light) and climatic (temperature) variables may limit the regional distribution of C. mopane, which is restricted to warmer sites.

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 [6473]. 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. [70] have used ISSR markers to assess the genetic diversity in B. boehmii and Burkea africana, another typical legume tree from Miombo, across a fire gradient in NNR wherein the northeastern side is affected by annual fires and the western by bi- to triannual fires. The authors observed that the levels of genetic diversity were lower in B. boehmii (average genetic diversity, He = 0.1965) than in B. africana (He = 0.2972). Such difference was attributed to fire tolerance and adaptation, as B. africana is a typical fire-tolerant species, while B. boehmii is more sensitive to fire, particularly at young stages [74, 75]. Interestingly, in B. boehmii high fire frequency resulted in higher variability (i.e., He = 0.2059 in eastern versus He = 0.1482 in western NNR), while in the case of B. africana, the opposite was observed (He = 0.2977 in the west and 0.2184 in the east). Based on this observation, Maquia et al. [70] proposed that the increase in B. boehmii variation driven by frequent fires may be part of its evolutionary response, a phenomena called pyrodiversity-like effect [76], associated to a higher proportion of seed-derived propagation in detriment of vegetative reproduction. Overall, the study concludes that the genetic variability within and among populations as well as the estimated gene flow between populations represents a strong genetic pool of the two species in NNR, agreeing with the fact that the reserve is one of least disturbed areas of Miombo. Similar results were obtained with other species of the Fabaceae family, e.g., Afzelia quanzensis Welw. [73], Astragalus rhizanthus Royle ex Benth. [77], and Glycyrrhiza uralensis Fisch. ex DC [78], from conservation areas.

RAPD markers were used to characterize accessions of P. angolensis from Zimbabwe, Zambia, and Tanzania [79, 80]. Chisha-Kasumu and colleagues [79] characterized the genetic structure of different populations from Zimbabwe and Zambia. According to their study, the variability within each population was high (Shannon’s information index, I = 81%). In Tanzania, [80] used the same strategy to analyze genetic diversity and population structure in six natural populations. In line with the results of [79], genetic diversity within populations was high (I = 77%), and in both cases, an effective gene flow was suggested. The results are within the range of those reported for other tree species [81, 82].

Villoen and collaborators [83] have used 13 allozyme markers to analyze five populations of C. mopane in South Africa: these included aspartate aminotransferase (AST), alcohol dehydrogenase (ADH), esterase (EST), isocitrate dehydrogenase (IDH), guanine deaminase (GDA), glucose-6-phosphate isomerase (GPI-1), leucine aminopeptidase (LAP), malate dehydrogenase (MDH), mannose-6-phosphate isomerase (MPI), peroxidase (PER), phosphoglucomutase (PGM), shikimate dehydrogenase (SKDH), and superoxide dismutase (SOD). As for B. boehmii [70] and P. angolensis [79, 80], most of the variation in C. mopane was observed within populations. Additionally, gene flow between populations was also effective. The results were also comparable to those obtained in the legume Virgilia oroboides (Berg.) [84]. The authors suggest that C. mopane developed an effective outcrossing mechanism that avoid inbreeding and maintain a considerable level of diversity necessary to ensure adaptation and survival under a context of ever-changing environments.

It should be highlighted that except for [70], 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, [85] examined the effect of three different environmental gradients on the genetic diversity of the semishrub legume Caragana microphylla Lam. The authors observed that higher levels of genetic diversity were correlated with optimal humidity, soil fertility, and to a less extent temperature. Dai et al. [86] examined the genetic variation of marginal Bombax ceiba L. (silk-cotton tree) populations in China and South Asia, based on the sequences of nuclear and chloroplast genes. The results revealed extremely low levels of genetic diversity without significant differences between cultivated and natural populations. This might reflect a small number of individuals (or other founder effects) during the establishment of populations or genetic drift. Addisalem et al. [87] examined the genetic structure of Boswellia papyrifera (Del.) Hochst across threaten tropical dry forest (Terminalia-Combretum) woodlands in Ethiopia, where populations retain high levels of genetic diversity despite the diverse environmental conditions (including harsh environments, i.e., high temperatures and low precipitation) as well as progressive deforestation and degradation.

Altogether, these studies [7087] 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 Hevea brasiliensis (Willd. ex A. Juss.) (Rubber tree) by de Souza and collaborators [88, 89], which have established a foundation of molecular markers (microsatellites and SNPs) highly valuable for breeding and conservation programs.


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.


  1. 1. Lock JM. Legumes of Africa: A Check List. Kew: Royal Botanic Gardens; 1989.
  2. 2. Sprent JI, Odee DW, Dakora FD. African legumes: a vital but under-utilized resource. Journal of Experimental Botany. 2010;61:1257–1265.
  3. 3. Mittermeier R, Mittermeier C, Brooks TM, Pilgrim JD, Konstant WR, da Fonseca GAB, Kormos C. Wilderness and biodiversity conservation. Proceedings of National Academy of Sciences. 2003;100:10309–10313.
  4. 4. Burgess ND, Hales JDA, Underwood E, Dinerstein E. Terrestrial Ecoregions of Africa and Madagascar: A Conservation Assessment. Washington DC: Island Press; 2004.
  5. 5. Dewees P, Campbell B, Katerere Y, Sitoe A, Cunningham AB, Angelsen A, Wunder S. Managing the Miombo Woodlands of Southern Africa: Policies, Incentives, and Options for the Rural Poor. Washington DC: Program on Forests (PROFOR); 2011.
  6. 6. Palmer E, Pitman N. Trees of Southern Africa, Vol. 2. A.A. Cape Town: Balkema AA; 1972.
  7. 7. Campbell B, Frost P, Byron N. Miombo woodlands and their use: overview and key issues. In: Campbell B, editor. The Miombo in Transition: Woodlands and Welfare in Africa. Bogor: CIFOR; 1996. pp. 1–5.
  8. 8. Frost P. The ecology of Miombo woodlands. In: Campbell B, editor. The Miombo in Transition: Woodlands and Welfare in Africa. Bogor: CIFOR; 1996. pp. 11–55.
  9. 9. Chidumayo E, Marunda C. Dry forests and woodlands in Sub-Saharan Africa: context and challenges. In: Chidumayo EN, Gumbo D, editors. The Dry Forests and Woodlands of Africa: Managing for Products and Services. London: Earthscan; 2010. pp. 1–10.
  10. 10. Ribeiro N, Cumbana M, Mamugy F, Chaúque A. Remote sensing of biomass in the Miombo woodlands of southern Africa: opportunities and limitations for research. In: Fatoyinbo L, editor. Remote Sensing of Biomass – Principles and Applications. Rijeka: InTech; 2012. pp. 7–98.
  11. 11. WWF. Miombo Eco-Region “Home of the Zambezi” Conservation Strategy 2011–2020. Harare: WWF-World Wide Fund for Nature; 2014.
  12. 12. Ribeiro NS. Interaction between Fires and Elephants in Relation to Vegetation Structure and Composition of Miombo Woodlands in Northern Mozambique [PhD Thesis]. Charlottesville: University of Virginia; 2007.
  13. 13. Gumbo D, Chidumayo E. Managing dry forests and woodlands for products and services: a prognostic synthesis. In: Chidumayo EN, Gumbo D, editors. The Dry Forests and Woodlands of Africa: Managing for Products and Services. London: Earthscan; 2010. pp. 261–279.
  14. 14. Stromgaard P. Early secondary succession on abandoned shifting cultivator’s plots in the miombo of south central Africa. Biotropica. 1987;18:97–110.
  15. 15. Chidumayo EN. Woody biomass structure and utilization for charcoal production in a Zambian miombo woodland. Bioresources Technology. 1991;37:43–52.
  16. 16. Malambo FM, Syampungani S. Opportunities and challenges for sustainable management of Miombo woodlands: the Zambian perspective. In: Varmola M, Valkonen S, Tapaninen, editors. Research and Development for Sustainable Management of Semi-Arid Miombo Woodlands in East Africa. Vantaa: Finnish Forest Research Institute; 2008. pp. 125–130.
  17. 17. Ribeiro NS, Syampungani S, Nangoma D, Ribeiro-Barros A. Miombo Woodlands Research Towards the Sustainable use of Ecosystem Services in Southern Africa [Internet]; 2015. Available from: [Accessed: 2016-09-14].
  18. 18. Hulme M, Doherty R, Ngara T, New M, Lister D. African climate change: 1900–2100. Climate Research. 2001;17:145–168.
  19. 19. Chidumayo EN. Effects of climate on the growth of exotic and indigenous trees in central Zambia. Journal of Biogeography. 2005;32:111–120.
  20. 20. IPCC 2007. Climate change 2007: The physical science basis: Contribution of working group I to the fourth assessment report of the intergovernmental panel on climate change. In: Solomon S, Qin D, Manning M, Chen Z, Marquis M, Averyt K, Tignor M, Miller H, editors. Cambridge: Cambridge University Press; 2007.
  21. 21. Gonzalez P, Tucker CJ, Sy H. Tree density and species decline in the African Sahel attributable to climate. Journal of Arid Environments. 2012;78:55–64.
  22. 22. IPCC 2014. Climate change 2014: Synthesis report. Contribution of working groups I, II and III to the fifth assessment report of the intergovernmental panel on climate change. In: Core writing team, Pachauri RK, Meyer LA, editors. Geneva: IPCC; 2014.
  23. 23. Green JMH, Larrosa C, Burgess ND, Balmford A, Johnston A, Mbilinyi BP, Platts PJ, Coad L. Deforestation in an African biodiversity hotspot: extent, variation and the effectiveness of protected areas. Biological Conservation. 2013;164:62–72.
  24. 24. Palgrave KC. Trees of Southern Africa. Cape Town: Struik Publishers; 1981.
  25. 25. Bruschi P, Mancini M, Mattioli M, Michela M, Signorini MA. Traditional uses of plants in a rural community of Mozambique and possible links with Miombo degradation and harvesting sustainability. Journal of Ethnobiology and Ethnomedicine. 2014;10:59. DOI: 10.1186/1746-4269-10-59.
  26. 26. Snook L, Alves T, Sousa C, Loo J, Gratzer G, Duguma L, Schrotter C, Ribeiro N, Mahanzule R, Mazuze F, Cuco E, Elias M. Relearning traditional knowledge to achieve sustainability: honey gathering in the miombo woodlands of northern Mozambique. In: Tropentag 2016; 18–21 September 2016, BOKU Vienna, Austria.
  27. 27. Agostinho AB, Cuinica DF, Chelene I, José AE. Medicinal plants and traditional medical practices in the treatment of mental disorders. Registo n6009rlind/2009. Maputo: UDEBA-LAB; 2009. (in Portuguese).
  28. 28. Augustino S, Hall JB, Makonda FBS, Ishengoma RC. Medicinal resources of the Miombo woodlands of Urumwa, Tanzania: Plants and its uses. Journal of Medicinal Plants Research. 2011;5:6352–6372.
  29. 29. Amri E, Kisangau DP. Ethnomedicinal study of plants used in villages around Kimboza forest reserve in Morogoro, Tanzania. Journal of Ethnobiology and Ethnomedicine 2012;8:1.
  30. 30. Chitemerere TA, Mukanganyama S. In vitro antibacterial activity of selected medicinal plants from Zimbabwe. The African Journal of Plant Science and Biotechology. 2011; 5(1):1–7.
  31. 31. Irondi EA, Oboh G, Akindahunsi AA. Methanol extracts of Brachystegia eurycoma and Detarium microcarpum seeds flours inhibit some key enzymes linked to the pathology and complications of type 2 diabetes in vitro. Food Science and Human Wellness. 2015;4:162–168. DOI: 10.1016/j.fshw.2015.08.002
  32. 32. Chirisa E, Mukanganyama S. Evaluation of in vitro anti-inflammatory and antioxidant activity of selected Zimbabwean plant extracts. Journal of Herbs, Spices & Medicinal Plants. 2016;22(2):157. DOI: 10.1080/10496475.2015.1134745
  33. 33. Jimu L. Julbernardia globiflora (Benth.) Troupin. [Internet] Record from PROTA4U. In: Brink M, Achigan-Dako ED, editors. Wageningen: PROTA; 2010. [Accessed: 2016-09-19].
  34. 34. Campbell BM. The Miombo in Transition: Woodlands and Welfare in Africa. Bogor: Centre for International Forestry Research; 1996.
  35. 35. Takawira-Nyenya R. Pterocarpus angolensis DC. [Internet] Record from PROTA4U. In: Louppe D, Oteng-Amoako AA, Brink M, editors. Wageningen: PROTA; 2005. [Accessed: 2016-09-19]
  36. 36. Cheikhyoussef A, Shapi M, Matengu K, Ashekele HM. Ethnobotanical study of indigenous knowledge on medicinal plant use by traditional healers in Oshikoto region, Namibia. Journal of Ethnobiology and Ethnomedicine 2011; 7:10.
  37. 37. Samie A, Housein A, Lall N, Meyer JJ. Crude extracts of, and purified compounds from Pterocarpus angolensis and the essential oil of Lippia javanica: their in-vitro cytotoxicities and activities against selected bacteria and Entamoeba histolytica. Annals of Tropical Medicine and Parasitology. 2009;103(5):427–439.
  38. 38. Mulaudzi RB, Ndhlala AR, Kulkarni MG, Finnie JF, Van Staden J. Antimicrobial properties and phenolic contents of medicinal plants used by the Venda people for conditions related to venereal diseases. Journal of Ethnopharmacology. 2011;135(2):330–337. DOI: 10.1016/j.jep.2011.03.022
  39. 39. Munodawafa T, Chagonda LS, Moyo SR. Antimicrobial and phytochemical screening of some Zimbabwean medicinal plants. Journal of Biologically Active Products from Nature. 2013; 3(5/6):323–330.
  40. 40. Molgaard P, Nielsen SB, Rasmussen DE, Drummond RB, Makaza N, Andreassen J. Anthelmintic screening of Zimbabwean plants traditionally used against schistosomiasis. Journal of Ethnopharmacology. 2001;74(3):257–264.
  41. 41. McGaw L, Van der Merwe D, Eloff JN. In vitro anthelmintic, antibacterial and cytotoxic effects of extracts from plants used in South African ethnoveterinary medicine. The Veterinary Journal. 2007;173:366–372.
  42. 42. Brummitt RK, Chikuni AC, Lock JM, Polhill RM. Leguminosae, subfamily Caesalpinioideae. In: Timberlake JR, Pope GV, Polhill RM, Martins ES, editors. Flora Zambesiaca. Vol. 3, part 2. Richmond: Royal Botanic Gardens, Kew; 2007.
  43. 43. Melusi R, Mojeremane W. Colophospermum mopane (Benth.) J.Léonard. [Internet] Record from PROTA4U. In: Lemmens RHMJ, Louppe D, Oteng-Amoako AA, editors. Wageningen: PROTA; 2012. [Accessed: 2016-09-19].
  44. 44. Stevens N, Archibald SA, Nickless A, Swemmer A, Scholes RJ. Evidence for facultative deciduousness in Colophospermum mopane in semi-arid African savannas. Austral Ecology. 2016;41(1):87–96.
  45. 45. Makhado R, Potgieter M, Timberlake J, Gumbo D. A review of the significance of mopane products to rural people's livelihoods in southern Africa. Transactions of The Royal Society of South Africa. 2014;69(2):117–122. DOI: 10.1080/0035919X.2014.922512
  46. 46. Ribeiro A, Romeiras MM, Tavares J, Faria T. Ethnobotanical survey in Canhane village, district of Massingir, Mozambique: medicinal plants and traditional knowledge. Journal of Ethnobiology and Ethnomedicine. 2010;6:33. DOI: 10.1186/1746-4269-6-33
  47. 47. Lukwa N, Mutambu SL, Makaza N, Molgaard P, Furu P. Perceptions about malaria transmission and control using anti-malaria plants in Mola, Kariba, Zimbabwe. Nigerian Journal of Natural Products and Medicine. 2001;5:4–7
  48. 48. Ferreira D, Marais JP, Slade D. Phytochemistry of the mopane, Colophospermum mopane, Phytochemistry. 2003;64:31–51.
  49. 49. Du K, Marston A, Van Der Westhuizen J, De Mieri M, Hamburger M, Neuburger M, Ferreira D. Labdane and clerodane diterpenoids from Colophospermum mopane. Journal of Natural Products. 2015;78(10):2494–2504. DOI: 10.1021/acs.jnatprod.5b00729
  50. 50. Jew EKK, Dougill AJ, Sallu SM, O'Connell J, Benton TG. Miombo woodland under threat: Consequences for tree diversity and carbon storage. Forest Ecology and Management. 2016;361:144–153. DOI: 10.1016/j.foreco.2015.11.011
  51. 51. Ribeiro NS, Saatchi SS, Shugart HH, Washington-Alen RA. Aboveground biomass and leaf area index (LAI) mapping for Niassa Reserve, northern Mozambique. Journal of Geophysical Research. 2008;113:G02S02. DOI: 10.1029/2007JG000550.
  52. 52. Ribeiro NS, Shugart HH, Washington-Allen R. The effects of fire and elephants on species composition and structure of the Niassa Reserve, northern Mozambique. Journal of Forest Ecology and Management. 2008;255:1626–1636.
  53. 53. World Conservation Monitoring Centre 1998. Pterocarpus angolensis. The IUCN Red List of Threatened Species 1998: e.T33190A9759374. [Internet]; 1998. Available from: [Accessed: 2016-09-28].
  54. 54. Shirima DD, Munishi PKT, Lewis SL, Burgess ND, Marshall AR, Balmford A, Swetnam RD, Zahabu EM. Carbon storage, structure and composition of miombo woodlands in Tanzania’s Eastern Arc Mountains. African Journal of Ecology. 2011;49:332–342.
  55. 55. Madoffe SS, Rija AA, Midtgaard F, Katani JZ, Mbeyale G, Zahabu E, Liwenga E, Christopher B. Preliminary assessment of forest structure, management and carbon stocking in Tanzania Miombo woodland. In: Proceedings of the First Climate Change Impacts, Mitigation and Adaptation Programme Scientific Conference. 2012; pp. 106–117.
  56. 56. De Cauwer V, Muys B, Revermann R, Trabucco A. Potential, realised, future distribution and environmental suitability for Pterocarpus angolensis DC. in southern Africa. Forest Ecology and Management. 2014;315:211–226. DOI: 10.1016/j.foreco.2013.12.032
  57. 57. Scott L. A late quaternary pollen record from the Transvaal bushveld, South Africa. Quaternary Research. 1982;17:339–370.
  58. 58. Pienaar B, Thompson DI, Erasmus BN, Hill T R, Witkowski EF. Evidence for climate-induced range shift in Brachystegia (miombo) woodland. South African Journal of Science. 2015:111(7/8):1–9. DOI: 10.17159/sajs.2015/20140280
  59. 59. White F. The Vegetation of Africa: A Descriptive Memoir to Accompany the UNESCO/AETFAT/UNSO, Vegetation Map of Africa (3 plates), 1:5,000,000. Paris: UNESCO; 1983.
  60. 60. Stevens N, Swemmer AM, Ezzy L, Erasmus BFN. Investigating potential determinants of the distribution limits of a savanna woody plant: Colophospermum mopane. Journal of Vegetation Science. 2014;25(2):363–373. DOI: 10.1111/jvs.12098
  61. 61. Rutherford MC, Powrie LW, Thompson, DI. Impacts of high utilisation pressure on biodiversity components in Colophospermum mopane savanna. African Journal of Range and Forage Science. 2012; 29(1):1–11. DOI: 10.2989/10220119.2012.687039
  62. 62. Woollen E, Ryan CM, Baumert S, Vollmer F, Grundy I, Fisher J, Fernando J, Luz A, Ribeiro N, Lisboa SN. Charcoal production in the Mopane woodlands of Mozambique: What are the trade-offs with other ecosystem services? Philosophical Transaction of the Royal Society B. 2016;371:20150315.
  63. 63. Govender N, Ribeiro N, Macanza V, Ruecker G. Comparing Two Fire Management Strategies in the GLTP: Creating the Foundations for a Fire Management System in the LNP. Project Report, GIZ; 2015.
  64. 64. Vos P, Hogers R, Bleeker M, Reijans M, Van de Lee T, Hornes M, Frijters A, Pot J, Peleman J, Kuiper M,Zabeau M. AFLP: a new technique for DNA fingerprinting. Nucleic Acids Research. 1995; 23(21):4407–4414.
  65. 65. Appleby N, Edwards D, Batley J. New technologies for ultra-high throughput genotyping in plants. Methods in Molecular Biology. 2009;513:19–39. DOI: 10.1007/978-1-59745-427-8_2.
  66. 66. Mondini L, Noorani A, Pagnotta MA. Assessing plant genetic diversity by molecular tools. Diversity. 2009;1:19–35.
  67. 67. Abdul Kareem VK, Rajasekharan PE, Ravish BS, Mini S, Sane A, Vasantha Kumar T. Analysis of genetic diversity in Acorus calamus populations in South and North East India using ISSR markers. Biochemical Systematics and Ecology. 2012;40:156–161.
  68. 68. Pillai PP, Sajan JS, Menon KM, Jayakumar KSP, Subramoniam A. SSR analysis reveals high intraspecific variation in Rauvolfia serpentina L. – a high value medicinal plant. Biochemical Systematics and Ecology. 2012;40:192–197.
  69. 69. Yao QL, Chen FB, Fang P, Zhou GF, Fan YH, Zhang ZR. Genetic diversity of Chinese vegetable mustard (Brassica juncea Coss) landraces based on SSR data. Biochemical Systematics and Ecology 2012;45:41–48.
  70. 70. Maquia I, Ribeiro NS, Silva V, Bessa F, Goulao LF, Ribeiro A. Genetic diversity of Brachystegia boehmii Taub. and Burkea africana Hook. f. across a fire gradient in Niassa National Reserve, northern Mozambique. Biochemical Systematics and Ecology. 2013;48:238–247.
  71. 71. Sarwat M, Yamdagni MM. DNA barcoding, microarrays and next generation sequencing: recent tools for genetic diversity estimation and authentication of medicinal plants. Critical Reviews in Biotechnology. 2014;36(2):191–203.
  72. 72. Grover A, Sharma PC. Development and use of molecular markers: past and present. Critical Reviews in Biotechnology. 2016;36 (2): 290–302.
  73. 73. Ribeiro NS, Jetimane JL, Militão E, Maquia I, Chirizane C, de Sousa C, Alves T, Veloso MM, Goulao LF, Ribeiro-Barros AI. Ecological characterization of an ex situ conservation plantation in southeastern Mozambique. 10 p. African Journal of Ecology [serial online]. 2016. Available from: Scopus® DOI: 10.1111/aje.12320. [Accessed: 2016-09-28].
  74. 74. Trapnell CG. Ecological results of woodland burning experiments in northern Rhodesia. Journal of Ecology. 1959;47:129–168.
  75. 75. Cauldwell AE, Ziegler U. A reassessment of the fire-tolerance of some miombo woody species in the Central Province, Zambia. African Journal of Ecology. 2000;38:138–146.
  76. 76. Parr CL, Andersen AN. Patch mosaic burning for biodiversity conservation: a critique of the pyrodiversity paradigm. Conservation Biology. 2006;20:1610–1619.
  77. 77. Anand KK, Srivastava RK, Chaudhary LB, Singh AR. Delimitation of species of the Astragalus rhizanthus complex (Fabaceae) using molecular markers RAPD, ISSR and DAMD. Taiwania. 2010;55:197–207.
  78. 78. Yao H, Zhao Y, Chen DF, Chen JK, Zhou TS. ISSR primer screening and preliminary evaluation of genetic diversity in wild populations of Glycyrrhiza uralensis. Biologia Plantarum. 2008;52:117–120.
  79. 79. Chisha-Kasumu E, Woodward S, Price A. RAPD markers demonstrate genetic diversity in Pterocarpus angolensis from Zimbabwe and Zambia Southern Forests. A Journal of Forest Science. 2009:71(1):63. DOI: 10.2989/SF.2009.
  80. 80. Amri E, Mmoya F. Genetic diversity in Pterocarpus angolensis populations detected by Random Amplified Polymorphic DNA markers. International Journal of Plant Breeding and Genetics. 2012; 6(2):105–114.
  81. 81. Newton AC, Allnutt TR, Dvorak WS, del Castillo RF, Ennos RA. Patterns of genetic variation in Pinus chiapensis, a threatened Mexican pine, detected by RAPD and mitochondrial DNA RFLP markers. Heredity 2002;89:191–198.
  82. 82. Bouvet JM, Fontaine C, Sanou H, Cardi C. An analysis of the pattern of genetic variation in Vitellaria paradoxa using RAPD markers. Agroforestry Systems. 2004;60:61–69.
  83. 83. Villoen L, Van der Bank FH, Van der Bank M, Grobler JP, Wessels D. Allozyme variation in five populations of mopane, Colophospermum mopane (Fabaceae). South African Journal of Botany. 2003;69(3):282–286. DOI: 10.1016/S0254-6299(15)30314-8
  84. 84. Van Der Bank H, Van Der Bank M, Van Wyk B. Allozyme variation in Virgilia oroboides (Tribe Podalyrieae: Fabaceae). Biochemical Systematics and Ecology. 1995;23:47–52.
  85. 85. Huang W, Zhao X, Zhao X, Li Y, Lian J. Effects of environmental factors on genetic diversity of Caragana microphylla in Horqin Sandy Land northeast China. Ecology and Evolution. 2016. DOI: 10.1002/ece3.2549.
  86. 86. Dai S, Wang S, Ruan L, Zhou Y, Wu W, Liu Y, Shi S, Zhou R. Extremely low genetic diversity and extensive genetic admixture at the northern range margins of Bombax ceiba. Biochemical Systematics and Ecology. 2015;60:177–185.
  87. 87. Addisalem A, Bongers F, Kassahun T, Smulders M. Genetic diversity and differentiation of the frankincense tree (Boswellia papyrifera (Del.) Hochst) across Ethiopia and implications for its conservation. Forest Ecology and Management. 2016;360:253–260.
  88. 88. de Souza L M, Le Guen V, Cerqueira-Silva CM, Silva CC, Mantello CC, Conson A O, Souza A. Genetic diversity strategy for the management and use of rubber genetic resources: More than 1,000 wild and cultivated accessions in a 100-Genotype Core Collection. Plos One. 2015;10(7):1–20. DOI: 10.1371/journal.pone.0134607.
  89. 89. de Souza LM, Toledo-Silva G, Cardoso-Silva CB, Silva CC, Andreotti I A, Conson AO, Mantello CC, Guen V, Souza AP. Development of single nucleotide polymorphism markers in the large and complex rubber tree genome using next-generation sequence data. Molecular Breeding. 2016;(8)1. DOI: 10.1007/s11032-016-0534-3.
  90. 90. Ndamba J, Nyazema N, Makaza N, Anderson C, Kaondera KC. Traditional herbal remedies used for the treatment of urinary schistosomiasis in Zimbabwe. Journal of Ethnopharmacology 1994;42:125–132.
  91. 91. Steenkamp V, Mathivhaa E, Gouwsb MC, Van Rensburga CEJ. Studies on antibacterial, antioxidant and fibroblast growth stimulation of wound healing remedies from South Africa. Journal of Ethnopharmacology. 2004; 95(2):353–357.
  92. 92. Luseba D, Elgorashi EE, Ntloedibe NT, Van Staden J. Antibacterial, anti-inflammatory and mutagenic effects of some medicinal plants used in South Africa for the treatment of wounds and retained placenta in livestock. South African Journal of Botany 2007;73:378–383.
  93. 93. Obi CLJ, Ramalivhana J, Samie A, Igumbor EO. Prevalence, pathogenesis, and antibiotic susceptibility profiles, and in vitro activity of selected medicinal plants against Aeromonas isolates from stool samples of patients in the Venda region of South Africa. Journal of Health, Population and Nutrition. 2007;25(4):428–435.
  94. 94. Mulaudzi RB, Ndhlala AR, Kulkarni MG, Finnie JF, Van Staden J. Anti-inflammatory and mutagenic evaluation of medicinal plants used by Venda people against venereal and related diseases. Journal of Ethnopharmacology. 2013;146(1): 173–179. DOI: 10.1016/j.jep.2012.12.026
  95. 95. Motlhanka DM. Free radical scavenging activity of selected medicinal plants of Eastern Botswana. Pakistan Journal of Biological Sciences. 2008;11(5):805–808.
  96. 96. Mebe PP. Diterpenes from the bark and seeds of Colophospermum mopane. Phytochemistry. 2001;57(4):537–541.
  97. 97. Steenkamp JA, Malan JCS, Roux DG, Ferreira D. Oligomeric flavanoids. Part 1. Novel dimeric profisetinidins from Colophospermum mopane. Journal of the Chemical Society, Perkin Transactions. 1988;1:1325–1330.

Written By

Isabel Moura, Ivete Maquia, Alfan A. Rija, Natasha Ribeiro and Ana Isabel Ribeiro-Barros

Reviewed: 11 November 2016 Published: 01 March 2017