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Combretum

Combretum is a genus of approximately 300 species of flowering plants in the family Combretaceae, comprising trees, shrubs, and woody climbers with a pantropical distribution and the center of diversity in Africa. These plants typically feature opposite or whorled, petiolate leaves that are often hairy or scaly, bisexual flowers arranged in spikes or panicles with 4–5-merous parts and small or showy petals, and distinctive 4–5-winged, ridged, or angled fruits that are frequently papery in texture. Native primarily to low-altitude habitats such as riverine forests, coastal areas, and seasonally dry tropical biomes, species of Combretum play significant ecological roles in stabilizing soils and supporting biodiversity in tropical ecosystems. The genus, first described by Peter Loefling in 1758,^[1] is the largest in Combretaceae, a family of about 500 species across 10 genera distributed worldwide in tropical and subtropical regions.^[2] While most species occur in Africa and Asia, some extend to the Americas, Australia, and the Pacific, including five species in [New Guinea](/page/New Guinea). Combretum species are easily distinguished from similar genera, such as those in Malpighiaceae, by their inferior ovary and characteristic winged fruits. Beyond their botanical significance, many Combretum species have been utilized in traditional medicine across Africa, Asia, and the Americas for treating a range of conditions, including infections, inflammation, malaria, diarrhea, and diabetes.^[3] Phytochemical analyses reveal the presence of bioactive compounds such as triterpenes, flavonoids, and ellagitannins, which contribute to reported antibacterial, antifungal, antiviral, antimalarial, and antidiabetic properties.^[3] Notable examples include C. indicum, used for its showy flowers, and C. molle, valued for its hypotensive effects.^[3]

Taxonomy

Classification

Combretum Loefl. is the type genus of the family Combretaceae, which belongs to the order Myrtales within the rosids clade of angiosperms.^[4] This placement reflects the genus's position in the eudicot lineage, characterized by features such as opposite leaves and four- to five-winged fruits typical of the family.^[5] The taxonomic history of Combretum includes significant revisions, particularly the incorporation of species previously classified under the genus Quisqualis. Based on morphological similarities in inflorescence structure and fruit characteristics, as well as molecular phylogenetic analyses confirming their nested position within Combretum, species like Quisqualis indica were transferred to Combretum around 1998–1999, with further combinations validated by recent molecular evidence in 2020.^[6]^[7] These updates resolved long-standing debates over generic boundaries in Combretaceae subtribe Combretinae.^[8] Within Combretum, subgeneric divisions recognize three main subgenera: Combretum, Cacoucia (Aubl.) Exell & Stace, and Apetalanthum Engl. & Diels, distinguished primarily by petal presence and filament morphology.^[9] These are further organized into numerous sections, such as section Striata Engl. & Diels (characterized by striate twigs and specific leaf indumentum) and section Calescens Engl. & Diels (defined by calyx tube features), with circumscriptions refined through combined morphological traits like scale patterns on twigs and molecular markers from nuclear and plastid DNA sequences.^[10]^[11] For instance, sectional boundaries in subgenus Combretum have been adjusted based on phylogenetic studies showing polyphyly in earlier groupings.^[12] The current accepted taxonomy, as documented in Plants of the World Online (POWO) in 2025, recognizes Combretum as a pantropical genus with 287 accepted species, integrating these subgeneric and sectional frameworks from integrated morphological and molecular datasets.^[4]

Etymology and Synonyms

The genus name Combretum derives from the Latin term used by the Roman naturalist Pliny the Elder (23–79 CE) to describe a climbing plant of uncertain identity, later adopted by the Swedish botanist Peter Loefling in 1758 for this group of plants.^[13] The flexible branches of many Combretum species, often utilized in traditional basketry, align with characteristics that may echo the original descriptive intent behind Pliny's usage, though the precise plant he referenced remains unidentified.^[14] Historically, Combretum has undergone several nomenclatural revisions, with key synonyms including Cacoucia Aubl., Calopyxis Tul., Meiostemon Exell & Stace, Quisqualis L., and Thiloa Eichler.^[13] The genus Quisqualis, previously recognized as distinct and comprising about six species of climbers primarily from Asia and Africa, was merged into Combretum following phylogenetic analyses in the early 2000s that demonstrated its nested position within the clade, particularly in subgenus Cacoucia sect. Poivrea.^[15] This merger, supported by molecular data from plastid and nuclear sequences showing low divergence (around 9.6%) between Quisqualis and core Combretum, resolved longstanding debates on generic boundaries and led to transfers such as Quisqualis indica L. becoming Combretum indicum (L.) DeFilipps.^[16] Other obsolete names, like those in Calopyxis from Madagascar, were subsumed under Combretum sect. Calopyxis (Tul.) Jongkind in 1995 based on morphological and distributional evidence.^[17] The type species of Combretum is C. fruticosum (L.) Loefl., originally described by Linnaeus in 1753 under Gomphia fruticosa and lectotypified to stabilize nomenclature amid early taxonomic confusion with related climbers.^[18] These changes reflect broader post-2000 efforts to refine the genus using integrated molecular and morphological approaches, reducing polyphyly in the Combretaceae family while preserving monophyletic circumscriptions.^[16]

Description

Morphological Characteristics

Combretum species exhibit a woody habit, manifesting as trees, shrubs, or scrambling lianas, with stems that can reach diameters of up to 10 cm and lengths of 3–25 m in some cases.^[19] The leaves are simple, typically arranged in opposite or subopposite pairs, occasionally whorled, and lack stipules, featuring entire margins and pinnate venation; they often resemble those of willows in their lanceolate to elliptic shape and overall form, with short petioles that may persist as spines after leaf abscission.^[19]^[20] The inflorescences are axillary racemes, spikes, or terminal panicles, bearing small, actinomorphic, bisexual flowers that are 4–5-merous, with a calyx forming a tubular to bell-shaped hypanthium and free petals or none present.^[19] Flower colors vary across species, including white, cream, yellow, or red hues, with 8–10 exserted stamens in one or two whorls contributing to their often showy appearance.^[19]^[21]^[22] Fruits are characteristically dry, indehiscent or dehiscent samaras or capsules, typically 4–5-winged or ribbed, which facilitate wind dispersal through their lightweight, winged structure containing a single seed.^[19] Bark varies from smooth and gray in younger stems to rough or snakeskin-like in mature trees, often producing mucilaginous exudate when damaged.^[19]^[21] Wood anatomy is regular, featuring abundant paratracheal parenchyma and uniseriate rays; for instance, in C. imberbe (leadwood), the wood is notably hard, dense, and durable, with thick bark providing fire resistance.^[19]^[23]

Growth and Reproduction

Combretum species are typically long-lived perennials exhibiting diverse growth habits, including shrubs, small trees, and lianas, with heights ranging from 3 to 16 meters depending on the species and environmental conditions.^[24] Many exhibit a scrambling or scandent form with multi-stemmed structures, and growth rates vary from slow in arid-adapted species to fast in tropical climbers like C. indicum under favorable conditions.^[25]^[26] In seasonal environments, several species, such as C. erythrophyllum, are deciduous, shedding leaves in winter after autumnal reddening to conserve water during dry periods.^[27] Reproduction in Combretum occurs primarily through hermaphroditic flowers arranged in inflorescences, with breeding systems showing variability across species; many display partial self-compatibility, as evidenced by incomplete self-incompatibility indices (e.g., 0.53 in C. constrictum), leading to reduced but viable fruit set from self-pollination compared to cross-pollination.^[28] Fruit development typically spans several months post-anthesis, with maturation timelines of 1–4 months observed in species like C. constrictum, where fruiting peaks follow flowering by 1–3 months.^[28] Seeds of Combretum generally exhibit orthodox storage behavior, tolerating desiccation to low moisture levels (e.g., 1.65% in C. indicum) and remaining viable after cryogenic storage at -18°C for at least 60 days.^[29] Germination lacks physical dormancy in many species, achieving high rates (>80% in C. lanceolatum) across broad temperature ranges and light regimes, with cryptocotylar or hemi-cryptocotylar hypogeal patterns where cotyledons remain belowground for protection.^[30]^[29] In savanna species such as C. glutinosum and C. nigricans, germination is stimulated by fire cues like heat and smoke, enhancing seedling establishment in post-burn environments.^[31] Germination times vary from 5–44 days, often favoring moist substrates without specific light dependency.^[29]^[30] Phenological patterns in Combretum are adapted to seasonal climates, with flowering predominantly occurring during dry periods to align reproduction with favorable conditions; for instance, trees like C. farinosum peak in early dry season (e.g., March in tropical dry forests), while lianas may flower later in response to shortening daylength.^[32] In African savannas, species such as C. constrictum flower year-round but intensify from August to November, coinciding with dry season onset, followed by fruiting through December.^[28] This timing supports seed dispersal before wet seasons, promoting establishment in variable habitats.^[32]

Distribution and Habitat

Geographic Range

The genus Combretum is predominantly distributed across tropical and southern Africa, where the majority of its approximately 287 species occur, with diversity concentrated in this region.^[33]^[34]^[4] This core range spans from Kenya and Tanzania in the east to Angola and Namibia in the west, extending southward into subtropical zones, supported by extensive occurrence data from global biodiversity databases.^[35] Beyond Africa, the genus extends to other tropical regions, including about five species in Madagascar, roughly 25 in tropical Asia (such as India and Southeast Asia), around 40 in the tropical Americas (from Mexico to South America), and a few species in northern Australia and New Guinea.^[36]^[37]^[4] These disjunct distributions reflect a pantropical pattern, with limited presence in the Pacific islands, as evidenced by georeferenced records totaling over 47,000 occurrences worldwide (as of 2025).^[35] Endemic hotspots include the Cape Floristic Region in southern Africa, where certain species like Combretum afrum exhibit localized endemism amid the broader regional diversity.^[33] The historical biogeography of Combretum points to a Gondwanan origin, with ancestral range reconstructions indicating diversification across southern continents following the breakup of Gondwana.^[38] Fossil records, including wood impressions attributed to Combretaceae from the Miocene (approximately 23–5 million years ago) in sites such as the Ituzaingó Formation in Argentina and Fort Ternan in Kenya, support this ancient tropical lineage and its subsequent radiation.^[39]^[40] Current distribution patterns, as mapped in 2025 biodiversity databases, align with these inferences, showing concentrated diversity in African savannas and woodlands while highlighting range extensions via long-distance dispersal.^[35]

Environmental Preferences

Combretum species thrive in a variety of open habitats, including savannas, woodlands, and riparian zones, primarily within semi-arid to tropical climates across tropical and southern Africa. These environments are characterized by seasonal rainfall patterns, with annual precipitation typically ranging from 500 to 1500 mm, supporting deciduous or semi-deciduous growth forms that align with distinct wet and dry periods. For instance, in miombo and related woodlands where Combretum is prominent, mean annual rainfall often falls between 600 and 1000 mm, with higher values in moister variants reaching up to 1521 mm. Such conditions favor the genus's adaptation to water-limited ecosystems, where species like Combretum imberbe occur in arid to moist savannas receiving up to 750 mm annually.^[41]^[42]^[43] Soil preferences for Combretum emphasize well-drained substrates, such as sandy loams, ferralitic soils, alluvial deposits, and heavy sandy or dolerite-derived types, which prevent waterlogging while allowing nutrient access. Drought resistance is a key trait, facilitated by extensive deep root systems that tap into groundwater during prolonged dry seasons, enabling survival in nutrient-poor, low-fertility soils common to these habitats. Species tolerate a range of edaphic conditions, from basalt-derived soils in riverine areas to clay-influenced sites, but perform best in aerated, non-compacted profiles that support root penetration.^[43]^[44]^[45] The genus occupies altitudinal gradients from sea level to approximately 2000 meters, with many species concentrated at low to medium elevations (e.g., 1000–1800 m in East African savannas). Fire-prone habitats are integral to Combretum-dominated communities, where frequent surface fires shape vegetation structure by promoting resprouting and preventing woody encroachment, thus maintaining open woodland dynamics. Recent studies from the 2020s indicate that climate change may alter these niches, with projected distributional shifts in Africa; for example, Combretum imberbe could see range expansions of up to 52.8% under high-emission scenarios (RCP8.5) in southern Africa, while Combretum glutinosum faces habitat contractions of 22–38% by 2100 in West African regions like Burkina Faso due to rising temperatures and altered precipitation.^[42]^[46]^[47]^[48]

Ecology

Ecological Interactions

Combretum species play a significant role in African savanna ecosystems as key woody components, often dominating woodland structures and serving as a primary food source for large herbivores. In semi-arid savannas, such as those in Kruger National Park, South Africa, Combretum trees and shrubs constitute a substantial portion of the diet for browsers like giraffes, which favor species such as Combretum apiculatum and Combretum zeyheri, comprising up to half of their browse intake alongside acacias. Elephants (Loxodonta africana) also heavily utilize Combretum foliage, bark, and branches, with studies in Burkina Faso's Nazinga Game Ranch documenting selective feeding that shapes woodland composition by promoting regrowth in "browsing lawns" accessible to smaller herbivores like antelope. This browsing pressure positions Combretum as a keystone provider of forage, influencing herbivore distribution and maintaining savanna heterogeneity.^[49]^[50]^[51] Symbiotic relationships enhance Combretum's resilience in nutrient-poor savanna soils, particularly through associations with arbuscular mycorrhizal fungi (AMF). Species like Combretum apiculatum and Combretum hereroense form vesicular-arbuscular mycorrhizae (VAM), which facilitate nutrient uptake, especially phosphorus, in exchange for plant-derived carbohydrates, as observed in tropical African woodlands. Flavonoids such as catechin produced by Combretum roots stimulate AMF spore germination and hyphal branching, strengthening this mutualism and aiding establishment in dry, disturbed habitats. While Combretum lacks direct nitrogen-fixing symbioses—unlike associated legumes—it benefits indirectly from mycorrhizal networks that improve soil fertility in mixed savanna communities.^[52]^[53]^[54] Combretum contributes to savanna biodiversity by hosting diverse fauna and acting as a pioneer in disturbed areas. As early colonizers in fire-prone or overgrazed landscapes, species like Combretum collinum rapidly regenerate via resprouting, stabilizing soils and facilitating succession in Miombo and other woodlands. They support insect communities, attracting pollinators and herbivores such as leaf-eating beetles and moths, while providing nectar and foliage for birds including sunbirds (Cinnyris spp.) that forage on dry-season flowers of Combretum glutinosum. In biodiversity hotspots like Loky Manambato, Madagascar, Combretum shrubs enhance arthropod diversity, underscoring their role in trophic interactions without dominating climax communities.^[55]^[56]^[57] Despite these benefits, Combretum faces threats from intense herbivory and pests that can limit population dynamics. Elephants cause structural damage by uprooting or debarking trees, as seen in Combretum-Terminalia woodlands where browsing reduces canopy cover and alters regeneration patterns. Giraffes inflict targeted damage on upper branches, prompting chemical defenses like increased tannin production in Combretum apiculatum following simulated browsing. Insect pests, including defoliating caterpillars and borers, further exacerbate vulnerability, particularly in young saplings, though some Combretum extracts show potential as natural insecticides against such threats.^[51]^[58]^[59]

Pollination and Seed Dispersal

Pollination in Combretum species is predominantly entomophilous and ornithophilous, with insects such as ants (Meranoplus bicolor, Camponotus consobrinus), flies (Musca domestica), bees, and butterflies, as well as birds like sunbirds (Cinnyris venustus), serving as primary pollinators.^[28] These pollinators are attracted to the tubular flowers, which offer nectar rewards with sugar concentrations ranging from 9% to 21% in species like C. constrictum.^[28] In some cases, such as C. lanceolatum, birds are drawn to a unique sweet jelly secretion rather than traditional nectar, facilitating effective pollen transfer.^[60] Breeding systems in Combretum emphasize outcrossing, promoted by floral traits like protandry and self-incompatibility in many species, though some are self-compatible with pollinator visits significantly enhancing fruit set—for instance, visitor-pollinated flowers in C. constrictum yield 8.75±2.65 fruits compared to 2.60±1.10 from self-pollination.^[28]^[61] Apomixis, an asexual seed production mechanism, occurs rarely in certain lineages, such as Combretum (section Poivrea), allowing reproduction without fertilization in isolated populations.^[62] Seed dispersal in Combretum occurs mainly via anemochory, with many species producing dry, winged fruits—often four-winged samaras—that enable wind transport over distances up to 20 meters or more, depending on fruit morphology and wind conditions.^[63] Animal-mediated dispersal, particularly endozoochory, also plays a role; frugivorous birds like hornbills and mammals such as baboons ingest the fruits of species like C. zeyheri, with gut passage providing scarification that breaks seed dormancy and improves germination rates.^[63] This dual strategy ensures effective propagation across varied habitats.^[43]

Human Uses

Medicinal and Traditional Uses

Various species of the Combretum genus have been integral to traditional herbal medicine in Africa, Asia, and the Americas, where they are employed to treat a range of ailments including diarrhea, wounds, and malaria. In African ethnomedicine, bark decoctions of C. molle are commonly used to alleviate symptoms of malaria and other infectious diseases, reflecting its widespread application in regions like Ethiopia and South Africa. Similarly, in Indian traditional practices, leaves, roots, and bark of C. albidum are prepared as remedies for fever, diarrhea, and dysentery, often in decoction form to harness their purported antimicrobial properties. In Brazil, C. leprosum is used in folk medicine to treat inflammation, pain, and wounds.^[64]^[65] These uses underscore the cultural significance of Combretum in indigenous healing systems, where plant parts are selected based on observed efficacy passed down through generations.^[66]^[67]^[68] Key bioactive compounds isolated from Combretum species contribute to these therapeutic effects, including combretastatins, ellagitannins, and flavonoids. Combretastatins, particularly combretastatin A-4 derived from C. caffrum, exhibit potent anti-cancer activity by disrupting microtubule polymerization and inhibiting tumor vascularization. Ellagitannins, such as punicalagin found in C. molle, provide antioxidant benefits through radical scavenging, supporting traditional wound healing applications. Flavonoids in various Combretum extracts further enhance anti-inflammatory and antimicrobial actions, bolstering their role in treating infections like diarrhea.^[69]^[21]^[68] Modern pharmacological research has validated and expanded on these traditional applications, with combretastatins advancing to clinical trials for cancer therapy. For instance, the phase II trial NCT00060242 evaluated combretastatin A4 phosphate in patients with advanced anaplastic thyroid cancer, demonstrating its potential to inhibit tumor blood supply, though results highlighted challenges in efficacy and toxicity. As of 2025, ongoing developments include combretastatin hybrids and antibody-drug conjugates, showing enhanced anti-tumor effects in preclinical models and renewed interest in combination therapies. Ethnopharmacological studies further document specific uses, such as the application of C. coccineum fruits in Madagascar for treating intestinal parasites as an anthelmintic, aligning with broader African practices for deworming.^[70]^[71]^[72]

Other Applications

Species of the genus Combretum provide valuable timber, particularly in Africa, where hardwoods like leadwood (C. imberbe) are utilized for durable furniture, carvings, and turned objects due to their density and resistance to wear.^[73]^[74] The wood's slow-burning properties also make it ideal for fuelwood, with smaller trees commonly harvested for this purpose across southern and eastern Africa.^[73]^[75] In cultural contexts, flexible stems and roots of certain Combretum species, such as C. indicum and C. zeyheri, are employed in basketry and weaving traditions in tropical regions.^[76]^[77] Bark from species like C. collinum yields dyes and serves in rituals among local communities in southern Africa.^[78] Combretum species contribute to ornamental landscaping and agroforestry practices in drylands, where they are planted as live fences for boundary demarcation and livestock control, as seen with C. collinum and C. molle in Uganda and Zambia.^[79]^[80] Their root systems aid in erosion control and soil stabilization in semi-arid environments.^[79] Economically, Combretum wood supports significant trade in firewood and charcoal production throughout sub-Saharan Africa, with species like C. molle preferred for their high energy yield in miombo woodlands.^[74]^[81] However, intensified harvesting has raised sustainability concerns, as noted in 2021 FAO reports highlighting deforestation risks and the need for regulated practices to balance energy demands with forest conservation.^[82]

Diversity

Number and Variation

The genus Combretum encompasses 287 accepted species as recognized in 2025 by Plants of the World Online, representing an update from earlier estimates; including synonyms and unresolved taxa, the total approaches approximately 370.^[4]^[5] This diversity underscores the genus's prominence within the Combretaceae family, with species exhibiting a range of growth forms from shrubs and small trees to lianas. Patterns of variation in Combretum are pronounced, particularly in geographic distribution and morphology. Over 80% of species demonstrate high endemism in Africa, with more than 140 recorded in tropical regions and around 20 endemic to Madagascar alone.^[83] Morphological diversity manifests in leaf shapes—varying from ovate and elliptic to oblong-lanceolate—and in fruit structures, where wing development ranges from absent to prominently four-winged, adaptations often linked to dispersal strategies in diverse habitats.^[84] Infrageneric classification recognizes about 17 sections, primarily delineated by floral traits such as petal shape and presence of calyx appendages, alongside fruit morphology. Molecular phylogenetic analyses, incorporating nuclear and plastid DNA sequences, have largely confirmed these sections as monophyletic clades, revealing evolutionary relationships that support the current taxonomy while highlighting polyphyly in some broader groups.^[85]^[9] Regarding conservation, the majority of Combretum species are assessed as Least Concern on the IUCN Red List, reflecting their widespread distributions in resilient savanna and woodland ecosystems. However, a subset—such as C. esteriense (Endangered) and C. hartmannianum (Vulnerable)—face threats from habitat loss due to deforestation and agricultural expansion, particularly in fragmented African ranges.^[86]^[87]

Notable Species

Combretum imberbe, commonly known as the leadwood tree, is renowned for producing Africa's heaviest and densest wood, which is highly resistant to termites and decay. This slow-growing deciduous tree typically reaches 6-10 meters in height but can attain up to 20 meters under optimal conditions, with a broad spreading canopy and pale grey bark that fissures into rectangular flakes resembling snakeskin. It holds deep cultural importance in southern African communities, where it is often regarded as sacred, used for ancestral rituals, carvings, and as a symbol of longevity due to its exceptional durability.^[75]^[88]^[89] Combretum erythrophyllum, or river bushwillow, is a riparian specialist adapted to riverbanks and moist environments across southern Africa. This medium to large deciduous tree grows to 5-10 meters, featuring dense foliage that turns vibrant yellow to red in autumn and sweetly scented cream-colored flowers in spring. Its bark and roots are rich in tannins, traditionally harvested for tanning leather and producing dark brown dyes.^[90]^[91]^[92] Combretum molle, the velvet bushwillow, is a widespread species in East and southern Africa, valued for its extensive medicinal applications in traditional practices. This variable shrub or small tree, reaching 3-10 meters tall, has velvety leaves and is commonly used to treat ailments such as fever, wounds, constipation, headaches, and chest complaints through decoctions of roots, leaves, and stems. It thrives in a variety of habitats, from woodlands to rocky outcrops, contributing to its broad distribution.^[93]^[94]^[95] Combretum fruticosum serves as the type species of the genus, originally described based on specimens from the Americas by Peter Loefling, a student of Carl Linnaeus, in 1758 as Gaura fruticosa. This scrambling shrub or vine, native from Mexico to northern Argentina, can grow up to 10 meters with support, producing striking orange spike-like flowers that attract pollinators. Though now less common in the wild due to habitat changes, it remains notable for its ornamental value and role in defining the genus.^[96]^[97]^[98]

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[34]
Combretum | Flora of the Guianas
The genus is divided in 3 subgenera and 52 sections, of which 2 and 11 respectively occur in America, and 2 and 5 respectively in the Guianas: Subgenus 1.
[35]
Combretum | Flora of Australia
A genus of some 250+ species throughout the tropics, with greatest diversity in Africa; three species in Australia (one of these on Christmas Island), two ...
[36]
Combretum indicum (L.) DeFilipps | Plants of the World Online
It is a climber and grows primarily in the wet tropical biome. It is used to treat unspecified medicinal disorders, has environmental uses, as a poison and a ...
[37]
Drift in the tropics: Phylogenetics and biogeographical patterns in ...
Jul 19, 2023 · The absence of fossils of non-winged fruits before the Miocene is coherent with our inferences which suggest that other fruit types evolved ...
[38]
Combretaceous fossil wood from Ituzaingó Formation (Late Miocene ...
Ituzaingó Formation is the first record of fossil wood that indicates a coastal marine environment, which is further south of the tropics.
[39]
Middle Miocene Fossil Plants from Fort Ternan (Kenya) and ... - jstor
as Cassia, Combretum, Terminalia, and Commi- phora. The fossil stumps and logs at Fort Ter- nan are larger than would be expected in des- ert vegetation ...
[40]
Dry Miombo Woodlands | One Earth
Mean annual rainfall ranges between 600 and 1,000 millimeters (24-39 in). Mean maximum temperatures range between 18°C and 30°C (64-86°F) depending on elevation ...Missing: conditions | Show results with:conditions
[41]
None
Summary of each segment:
[42]
[PDF] combretum imberbe warwa (combretaceae) and - DFFE
Aug 6, 2003 · Details on the density of specific stage classes of trees are not available, but rather for all growth stages combined. Clarke (1997) ...<|separator|>
[43]
2. Background to miombo woodlands in Mozambique
It is in the presence of red clay soils of low fertility and low retention of water that high forest and dense miombo woodland take place. 2.4 Vegetation.2.1 Distribution And... · 2.4 Vegetation · 2.7 Land Tenure And Forest...Missing: conditions | Show results with:conditions
[44]
Miombo - an overview | ScienceDirect Topics
The deep roots of woodland trees recycle nutrients that would otherwise be lost in deep soil layers. The transpiration of forest trees helps regulate rainfall.
[45]
(PDF) The ecology of miombo woodlands - ResearchGate
Aboveground woody biomass (B, Mg ha-1 ) of oldgrowth, mixed-age stands of miombo woodland increases with mean annual rainfall (P, mm): B = 0.14 P-55.48 (r = ...
[46]
Climate change threatens the distribution of major woody species and ecosystem services provision in southern Africa
### Summary: Combretum Species and Climate Change Impacts in Africa
[47]
Modelling the current and future geographical distribution of ...
Combretum glutinosum is a keystone species prevalent in the West Africa savannas, particularly thriving in arid ecosystems. It plays a vital role in local ...
[48]
Grazers and Browsers Kruger National Park Wildlife
They are adaptable browsers which feed on 70% of the tree species in Kruger, but favour acacias and combretum species which make up half their diet. Giraffe ...
[49]
Elephant impact on shoot distribution on trees and on rebrowsing by ...
Elephants, therefore, seem to facilitate browsing by mesoherbivores by generating 'browsing lawns'. Such benefits need to be considered when making decisions on ...
[50]
Effects of browsing by elephants on the Combretum/Terminalia ...
Effects of browsing by elephants on the Combretum/Terminalia woodland at the Nazinga Game Ranch, Burkina Faso, West Africa ... Elephant-woodland interactions at ...
[51]
Arbuscular mycorrhizal fungi communities from tropical Africa reveal ...
Aug 25, 2016 · robusta Burch., Albizia spp., Combretum spp. and Milletia ... Arbuscular mycorrhiza and nitrogen: implications for individual plants through to ...
[52]
Rhizosphere Microbiome Modulators: Contributions of Nitrogen ...
The flavonoid naringenin produced by legumes activates germination of mycorrhizal spore and hyphal branching while the flavonoid catechin produced by Combretum ...
[53]
(PDF) Investigating the impact of herbivory and nitrogen-fixation on ...
Jun 7, 2017 · Selecting individuals of the N2-fixing Acacia tortilis and the non-N2-fixing Combretum hereroense both inside an exclosure and on the ...
[54]
Diet DNA reveals novel African Forest elephant ecology on the ...
Mar 23, 2022 · In this study, the elephant diet frequently contained known pioneer woody species, such as Combretum and Terminalia, which were represented in ...
[55]
(PDF) Diversity and abundance of insect visitors to flowers of trees ...
Aug 7, 2025 · Tiliaceae (Combretaceae: Combretum hereroense Schinz, Combretum zeyheri ... americana ), was a potential hosting environment for such insects(Hill ...
[56]
Selection by Birds of Shrub and Tree Species in the Sahel - BioOne
Jul 5, 2023 · The grid cells span a range of climate zones from Sahara to Guinean forests, with an annual rainfall between 30 and 1800 mm (see Figure 4 in ...
[57]
Giraffe browsing in response to plant traits - ScienceDirect.com
We used Masai giraffe (Giraffa ... Bergström et al. Plant responses and herbivory following simulated browsing and stem cutting of Combretum apiculatum ...
[58]
Growth, chemical responses and herbivory after simulated leaf ...
Jul 25, 2006 · Growth, chemical responses and herbivory after simulated leaf browsing in Combretum apiculatum · Abstract · Article PDF · References.Missing: damage | Show results with:damage
[59]
Birds accept sweets from strange flowers - Nature
Jun 25, 2001 · Combretum lanceolatum: a sweet shop for birds. Most plants offer visitors a soft drink, rewarding pollinators with liquid nectar. Researchers ...
[60]
Reproductive biology, variability of nectar features and pollination of ...
Aug 7, 2025 · Previous studies on the breeding biology of Combretum ... The pollination biology of Combretum farinosum, a hummingbird-pollinated, outcrossing ...
[61]
Apomixis in Neotropical Vegetation - IntechOpen
Mar 14, 2018 · In Combretaceae, apomixis was observed in Combretum ( Poivrea ). In Myrtaceae, adventitious embryony was recorded in Callistemon , Eugenia and ...
[62]
Combretum zeyheri – Eco Restoration
Combretum zeyheri. Combretum zeyheri Sond. Large-fruited Bushwillow, Zeyher's ... High Browsing Value, Host plant Butterfly, Host plant Moth, Nesting and Shelter, ...
[63]
Anti-plasmodial activities of Combretum molle (Combretaceae ...
May 18, 2018 · Combretum molle is a popular medicinal plant widely used in Africa for treatment of different aliments including malaria and HIV [5,6,7].
[64]
Combretum albidum G. Don: a medicinal liana of India - ResearchGate
Aug 15, 2024 · Traditionally, different parts of the plant, such as leaves, roots, and bark, are used in the treatment of ailments like fever, diarrhoea, ...
[65]
Bioactivities of the Genus Combretum (Combretaceae): A Review
The present work is a literature survey of Combretum species that have been evaluated for their ability to exert biological activities. A total number of 36 ...Missing: habits | Show results with:habits
[66]
In vitro evaluation of the antineoplastic activity of combretastatin A-4 ...
Combretastatin A-4 is a natural product which was isolated from the South African tree Combretum caffrum. In this study, the cytotoxic activity of ...Missing: cafre | Show results with:cafre
[67]
Combretastatin A4 Phosphate in Treating Patients With Advanced ...
This phase II trial is studying how well combretastatin A4 phosphate works in treating patients with advanced recurrent or metastatic anaplastic thyroid cancer.Missing: 2025 | Show results with:2025
[68]
A novel life for antitumor combretastatins: Recent developments of ...
Jan 5, 2025 · The combination of CA4 and antitumor agents enhances the anticancer efficacy. •. Combretastatin A4 as a payload in novel antibody-drug ...Missing: cafre | Show results with:cafre
[69]
https://pubmed.ncbi.nlm.nih.gov/8457709/
[70]
Leadwood | The Wood Database (Hardwood)
Common Uses: Carving, furniture, turned objects, and other small specialty items. Comments: Smaller trees are used as fuelwood, as the wood burns slowly at ...
[71]
Combretum imberbe - PROTA4U
It is popular for sculpture and lathe work, whereas it is also used to make heavy, extremely durable furniture. The wood is favoured for use as firewood and for ...
[72]
Combretum imberbe - PlantZAfrica |
Leadwood ash is used as a toothpaste. The wood is very hard and tough, and burns very slowly with intense heat. Africans used this wood to make hoes before ...
[73]
Combretum indicum (PROTA) - Pl@ntUse - PlantNet
Apr 20, 2020 · After a revision all 6 African Quisqualis species were transferred to Combretum, although this transfer is disputed by some taxonomists.
[74]
Species information: Combretum zeyheri - Flora of Zambia
Combretum zeyheri is a small to medium tree with distinctive banded bark. Its roots are used for basketry, and it grows in Miombo and munga woodland.
[75]
Cultural, forage, medicinal and potential applications of Combretum ...
Aug 26, 2025 · Minor uses include edible seeds, edible cooked tubers, edible flower nectar, ritual purposes, edible gum, edible root tubers, edible ...
[76]
Uganda | Interactive Suitable Tree Species Selection and ...
Combretum collinum, Bimwa, Combretaceae, Native, Firewood, Charcoal, Timber for ... Ornamental, Live fence, Dead fence, Shade, Soil erosion control, Soil ...
[77]
[PDF] Adoption of Agroforestry Systems by Farmers in Masaka District of ...
Mar 29, 2012 · Combretum molle R.Br. ex G. Don. Ndagi. 0.00 Timber, fuel wood. T. I ... 0.16 Fruit jam, live fence, ornamental. SOLANACEAE. S. I. Cyphomandra ...
[78]
[PDF] SOCIO-ECONOMICS OF CHARCOAL EXTRA- TION IN TANZANIA
There is a lucrative charcoal trade between producers and urban traders rendering the often re- garded minor forest product a source of livelihood and income to ...
[79]
In the spotlight: how to make Africa's reliance on charcoal and ...
Nov 23, 2021 · In the spotlight: how to make Africa's reliance on charcoal and firewood more sustainable. Format: News and Press Release; Source. FAO. Posted ...Missing: Combretum | Show results with:Combretum<|control11|><|separator|>
[80]
A new species of Combretum sect. Ciliatipetala (Combretaceae ...
Oct 10, 2025 · About 140 species have been recorded in tropical Africa and approximately 20 species are endemic to Madagascar (Schmelzer & Gurib-Fakim 2013). .
[81]
systematic significance of morphology and leaf epidermal ...
All Combretum species possessed amphistomatic and hypostomatic leaves. Systematically, the leaf epidermal ultrastructure revealed remarkable variation. All ...
[82]
[PDF] Combretaceae - UJ Content - University of Johannesburg
Sampling includes 14 genera of the family Combretaceae, 73 taxa of two subgenera (Combretum and Cacoucia), 17 sections of genus. Combretum, 71 species of the ...
[83]
Combretum esteriense THE IUCN RED LIST OF THREATENED ...
Dec 4, 2017 · This species is assessed as Category of Endangered: EN B1ab(iii)+2ab(iii). It occurs in the estuary area of Gabon and is currently only known ...<|control11|><|separator|>
[84]
Combretum Facts - Photos - Earth's Endangered Creatures
Status/Date(s) Listed as Endangered ; 1. Combretum hartmannianum, VU-IUCN, 1998, Eritrea, Ethiopia, Sudan ; 2. Combretum rochetianum, VU-IUCN, 1998, Eritrea, ...
[85]
https://ujcontent.uj.ac.za/view/pdfCoverPage?instCode=27UOJ_INST&filePid=135894720007691&download=true
[86]
Fascinating Flora: The Leadwood Tree - Singita
The leadwood is one of the largest trees in Africa, and is so called because of the wood which is extremely dense and heavy. As such, it is impermeable to ...Missing: characteristics | Show results with:characteristics
[87]
https://www.earthsendangered.com/profile.asp?gr=P&sp=16470
[88]
Combretum erythrophyllum - CJM Tree Growers
Usually a smooth, pale grey or yellow to orange-brown on young trees, becoming darker as the tree matures. · With age, the bark tends to flake in patches to ...
[89]
Combretum erythrophyllum - Tree SA
Bush Willow, Bushveld Willow, River Combretum, River Bushwillow, River Combretum, Vaal River Yellowood. ... leaves in a flower cluster or associated with flowers) ...
[90]
https://tropical.theferns.info/viewtropical.php?id=Combretum+erythrophyllum
[91]
Combretum molle | PlantZAfrica
Velvet bush willow tree is important in traditional medicine. Most African people use boiled root decoction to treat constipation, headaches, stomachs, fever, ...
[92]
Phytochemical Composition of Combretum molle (R. Br. ex G ... - NIH
Apr 19, 2023 · Combretum molle has also been reported to possess cardioprotective properties [45], possibly due to the presence of mollic acid glucoside ( ...
[93]
Combretum - Atlas of Florida Plants
Contains 1 accepted taxa overall. Characteristics: Genus: Combretum Loefl. Family: COMBRETACEAE. Common Name: -. Specimen: View USF Herbarium specimens of this ...
[94]
https://pza.sanbi.org/combretum-molle
[95]
FPS138/FP138: Combretum fruticosum Orange Flame Vine
Combretum fruticosum Orange Flame Vine ; General Information. Scientific name: Combretum fruticosum ; Description. Height: depends upon supporting structure.