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"Schousboea" redirects here; this is also synonym of Alchornea.

Combretum
Combretum collinum with fruit
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Myrtales
Family: Combretaceae
Genus: Combretum
Loefl.
Species

About 3272 see text

Synonyms[1]
  • Aetia Adans.
  • Bucholzia Stadtm. ex Willemet
  • Bureava Baill.
  • Cacoucia Aubl.
  • Calopyxis Tul.
  • Campylochiton Welw. ex Hiern
  • Campylogyne Welw. ex Hemsl.
  • Chrysostachys Pohl
  • Codonocroton E.Mey. ex Engl. & Diels
  • Cristaria Sonn.
  • Embryogonia Blume
  • Forsgardia Vell.
  • Gonocarpus Ham.
  • Grislea L.
  • Hambergera Scop.
  • Kleinia Crantz
  • Meiostemon Exell & Stace
  • Mekistus Lour. ex Gomes Mach.
  • Physopodium Desv.
  • Poivrea Comm. ex DC.
  • Quisqualis L.
  • Schousboea Willd.
  • Seguiera Rchb. ex Oliv.
  • Sheadendron G.Bertol.
  • Sphalanthus Jack
  • Thiloa Eichler
  • Udani Adans.

Combretum, the bushwillows or combretums, make up the type genus of the family Combretaceae. The genus comprises about 272 species of trees and shrubs, most of which are native to tropical and southern Africa, about 5 to Madagascar, but there are others that are native to tropical Asia, New Guinea and the Bismarck Archipelago, Australia, and tropical America.

Around 17 species in the genus Quisqualis are very similar to Combretum and are now classified as species of the genus.[2]

Though somewhat reminiscent of willows (Salix) in their habitus, they are not particularly close relatives of these.

Ecology

[edit]
Combretum albidum in Kinnerasani Wildlife Sanctuary, Andhra Pradesh, India

Bushwillow trees often are important plants in their habitat. Savannahs in Africa, in particular those growing on granitic soils, are often dominated by Combretum and its close relative Terminalia. For example, C. apiculatum is a notable tree in the Angolan mopane woodlands ecoregion in the Kunene River basin in southern Africa.[3]

Other species of this genus are a major component of Southwestern Amazonian moist forests. This genus contains several species that are pollinated by mammals other than bats, which is quite rare indeed.[4] But most species are more conventionally pollinated by insects or birds.[citation needed]

Typhlodromus combretum, a mite of the family Phytoseiidae, was discovered on a bushwillow plant and is named after this genus. Other herbivores that eat Combretum foliage include the caterpillars of the Brown Awl (Badamia exclamationis) which is found on C. albidum, C. latifolium and C. ovalifolium; those of the Orange-tailed Awl (Bibasis sena) are recorded from C. extensum and C. latifolium.[citation needed]

Use by humans

[edit]
Combretum aculeatum inflorescence
Combretum paniculatum
Combretum acutum – MHNT

Several species are used in African or Indian herbal medicine.[additional citation(s) needed] Several species of this genus in Madagascar known collectively as voantamenaka or voatamenaka (from voa 'fruit' and menaka 'oil') are used in traditional Malagasy medicine as deworming remedies.[5] The class of chemical compounds known as combretastatins were first isolated from South African bushwillow (Combretum afrum), from which they get their name. One synthetic derivative, fosbretabulin disodium (combretastatin A4 phosphate), underwent preliminary study for the treatment of anaplastic thyroid cancer,[6] but it was not effective enough to progress to more advanced trials. C. molle is also recorded to contain antioxidants such as punicalagin,[citation needed] which is also found in pomegranates (Punica granatum), a somewhat related plant.

The botanist George Don studied this genus extensively. The Luvuvhu River in southern Africa was named after the river's bushwillows (C. erythrophyllum), locally known as muvuvhu.

Species

[edit]

As of April 2021, there are 272 accepted species of Combretum:[1]

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Combretum

View on Grokipedia
from Grokipedia
Combretum is a of approximately 300 of flowering in the family , comprising trees, shrubs, and woody climbers with a distribution and the center of diversity in . These 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, of Combretum play significant ecological roles in stabilizing soils and supporting in tropical ecosystems. The genus, first described by Peter Loefling in , is the largest in , a family of about 500 species across 10 genera distributed worldwide in tropical and subtropical regions. While most species occur in and , some extend to the , , and the Pacific, including five species in [New Guinea](/page/New Guinea). Combretum species are easily distinguished from similar genera, such as those in , by their inferior ovary and characteristic winged fruits. Beyond their botanical significance, many Combretum species have been utilized in across , , and the for treating a range of conditions, including infections, , , , and . analyses reveal the presence of bioactive compounds such as triterpenes, , and ellagitannins, which contribute to reported antibacterial, , antiviral, antimalarial, and antidiabetic properties. Notable examples include C. indicum, used for its showy flowers, and C. molle, valued for its hypotensive effects.

Taxonomy

Classification

Combretum Loefl. is the of the family , which belongs to the order within the of angiosperms. 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. The taxonomic history of Combretum includes significant revisions, particularly the incorporation of species previously classified under the genus Quisqualis. Based on morphological similarities in structure and 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. These updates resolved long-standing debates over generic boundaries in subtribe Combretinae. 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. 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 DNA sequences. For instance, sectional boundaries in subgenus Combretum have been adjusted based on phylogenetic studies showing in earlier groupings. The current accepted , as documented in (POWO) in 2025, recognizes as a with 287 accepted species, integrating these subgeneric and sectional frameworks from integrated morphological and molecular datasets.

Etymology and Synonyms

The name Combretum derives from the Latin term used by the Roman naturalist (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. 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. Historically, Combretum has undergone several nomenclatural revisions, with key synonyms including Cacoucia Aubl., Calopyxis Tul., Meiostemon Exell & Stace, Quisqualis L., and Thiloa Eichler. 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. 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. 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. The of Combretum is C. fruticosum (L.) Loefl., originally described by Linnaeus in 1753 under Gomphia fruticosa and lectotypified to stabilize amid early taxonomic confusion with related climbers. These changes reflect broader post-2000 efforts to refine the genus using integrated molecular and morphological approaches, reducing in the family while preserving monophyletic circumscriptions.

Description

Morphological Characteristics

Combretum species exhibit a woody , 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. 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 . 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. 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. 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. Bark varies from smooth and gray in younger stems to rough or snakeskin-like in mature trees, often producing mucilaginous exudate when damaged. 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.

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 and environmental conditions. Many exhibit a 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. In seasonal environments, several , such as C. erythrophyllum, are , shedding leaves in winter after autumnal reddening to conserve water during dry periods. Reproduction in Combretum occurs primarily through hermaphroditic flowers arranged in inflorescences, with breeding systems showing variability across ; many display partial self-compatibility, as evidenced by incomplete indices (e.g., 0.53 in C. constrictum), leading to reduced but viable set from compared to cross-pollination. development typically spans several months post-anthesis, with maturation timelines of 1–4 months observed in like C. constrictum, where fruiting peaks follow flowering by 1–3 months. Seeds of Combretum generally exhibit orthodox storage behavior, tolerating to low levels (e.g., 1.65% in C. indicum) and remaining viable after cryogenic storage at -18°C for at least 60 days. lacks physical in many species, achieving high rates (>80% in C. lanceolatum) across broad ranges and regimes, with cryptocotylar or hemi-cryptocotylar hypogeal patterns where cotyledons remain belowground for protection. In species such as C. glutinosum and C. nigricans, is stimulated by cues like and , enhancing establishment in post-burn environments. times vary from 5–44 days, often favoring moist substrates without specific dependency. 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 (e.g., March in tropical dry forests), while lianas may flower later in response to shortening daylength. In African savannas, species such as C. constrictum flower year-round but intensify from to , coinciding with onset, followed by fruiting through December. This timing supports before wet seasons, promoting establishment in variable habitats.

Distribution and Habitat

Geographic Range

The genus Combretum is predominantly distributed across tropical and , where the majority of its approximately 287 occur, with diversity concentrated in this region. This core range spans from and in the east to and in the west, extending southward into subtropical zones, supported by extensive occurrence data from databases. Beyond Africa, the genus extends to other tropical regions, including about five species in , roughly 25 in (such as and ), around 40 in the tropical (from to ), and a few species in and . These disjunct distributions reflect a pattern, with limited presence in the Pacific islands, as evidenced by georeferenced records totaling over 47,000 occurrences worldwide (as of 2025). Endemic hotspots include the in , where certain species like Combretum afrum exhibit localized amid the broader regional diversity. The historical biogeography of Combretum points to a origin, with ancestral range reconstructions indicating diversification across southern continents following the breakup of . Fossil records, including wood impressions attributed to from the (approximately 23–5 million years ago) in sites such as the Ituzaingó Formation in and Fort Ternan in , support this ancient tropical lineage and its subsequent . 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.

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 . These environments are characterized by seasonal rainfall patterns, with annual 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 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 occur in arid to moist savannas receiving up to 750 mm annually. Soil preferences for emphasize well-drained substrates, such as sandy loams, ferralitic , alluvial deposits, and heavy sandy or dolerite-derived types, which prevent waterlogging while allowing nutrient access. resistance is a key trait, facilitated by extensive deep root systems that tap into during prolonged dry seasons, enabling survival in nutrient-poor, low-fertility common to these habitats. Species tolerate a range of edaphic conditions, from basalt-derived in riverine areas to clay-influenced sites, but perform best in aerated, non-compacted profiles that support root penetration. The genus occupies altitudinal gradients from to approximately 2000 meters, with many species concentrated at low to medium elevations (e.g., 1000– m in East African savannas). Fire-prone habitats are integral to Combretum-dominated communities, where frequent surface fires shape structure by promoting resprouting and preventing woody encroachment, thus maintaining open dynamics. Recent studies from the 2020s indicate that may alter these niches, with projected distributional shifts in ; for example, could see range expansions of up to 52.8% under high-emission scenarios (RCP8.5) in southern , while Combretum glutinosum faces habitat contractions of 22–38% by 2100 in West African regions like due to rising temperatures and altered precipitation.

Ecology

Ecological Interactions

Combretum species play a significant role in African ecosystems as key woody components, often dominating woodland structures and serving as a primary food source for large s. In semi-arid s, such as those in , , 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. (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 s like . This browsing pressure positions Combretum as a keystone provider of , influencing herbivore distribution and maintaining savanna heterogeneity. Symbiotic relationships enhance Combretum's resilience in nutrient-poor 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 , in exchange for plant-derived carbohydrates, as observed in tropical African woodlands. such as produced by Combretum roots stimulate AMF and hyphal branching, strengthening this mutualism and aiding establishment in dry, disturbed habitats. While Combretum lacks direct nitrogen-fixing symbioses—unlike associated —it benefits indirectly from mycorrhizal networks that improve in mixed communities. Combretum contributes to biodiversity by hosting diverse 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 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, , Combretum shrubs enhance diversity, underscoring their role in trophic interactions without dominating climax communities. Despite these benefits, Combretum faces threats from intense herbivory and pests that can limit . 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 production in Combretum apiculatum following simulated browsing. 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.

Pollination and Seed Dispersal

Pollination in Combretum species is predominantly entomophilous and ornithophilous, with such as (Meranoplus bicolor, Camponotus consobrinus), flies (Musca domestica), bees, and , as well as birds like sunbirds (Cinnyris venustus), serving as primary pollinators. These pollinators are attracted to the tubular flowers, which offer rewards with sugar concentrations ranging from 9% to 21% in species like C. constrictum. In some cases, such as C. lanceolatum, birds are drawn to a unique sweet jelly secretion rather than traditional , facilitating effective transfer. Breeding systems in Combretum emphasize , promoted by floral traits like protandry and in many species, though some are self-compatible with 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 . , an asexual seed production mechanism, occurs rarely in certain lineages, such as Combretum (section Poivrea), allowing without fertilization in isolated populations. 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. 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. This dual strategy ensures effective propagation across varied habitats.

Human Uses

Medicinal and Traditional Uses

Various species of the Combretum genus have been integral to traditional in , , and the , where they are employed to treat a range of ailments including , wounds, and . In African ethnomedicine, bark decoctions of C. molle are commonly used to alleviate symptoms of and other infectious diseases, reflecting its widespread application in regions like and . Similarly, in Indian traditional practices, leaves, roots, and bark of C. albidum are prepared as remedies for fever, , and , often in form to harness their purported properties. In , C. leprosum is used in folk medicine to treat , pain, and wounds. These uses underscore the cultural significance of Combretum in indigenous systems, where parts are selected based on observed efficacy passed down through generations. Key bioactive compounds isolated from Combretum species contribute to these therapeutic effects, including combretastatins, ellagitannins, and . Combretastatins, particularly combretastatin A-4 derived from C. caffrum, exhibit potent anti-cancer activity by disrupting polymerization and inhibiting tumor vascularization. Ellagitannins, such as punicalagin found in C. molle, provide benefits through radical scavenging, supporting traditional applications. Flavonoids in various Combretum extracts further enhance and actions, bolstering their role in treating infections like . 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 , 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 for treating intestinal parasites as an , aligning with broader African practices for .

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. 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. 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. Bark from species like C. collinum yields dyes and serves in rituals among local communities in . Combretum species contribute to ornamental and practices in , where they are planted as live fences for boundary demarcation and livestock control, as seen with C. collinum and C. molle in and . Their root systems aid in and in semi-arid environments. Economically, Combretum wood supports significant trade in and production throughout , with species like C. molle preferred for their high energy yield in woodlands. However, intensified harvesting has raised concerns, as noted in 2021 FAO reports highlighting risks and the need for regulated practices to balance energy demands with forest conservation.

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. 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 in , with more than 140 recorded in tropical regions and around 20 endemic to alone. Morphological diversity manifests in 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. Infrageneric classification recognizes about 17 sections, primarily delineated by floral traits such as shape and presence of calyx appendages, alongside morphology. Molecular phylogenetic analyses, incorporating nuclear and DNA sequences, have largely confirmed these sections as monophyletic clades, revealing evolutionary relationships that support the current while highlighting in some broader groups. Regarding conservation, the majority of Combretum species are assessed as Least Concern on the , reflecting their widespread distributions in resilient and ecosystems. However, a subset—such as C. esteriense (Endangered) and C. hartmannianum (Vulnerable)—face threats from habitat loss due to and agricultural expansion, particularly in fragmented African ranges.

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 and decay. This slow-growing 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 . It holds deep cultural importance in southern African communities, where it is often regarded as sacred, used for ancestral rituals, carvings, and as a of due to its exceptional durability. Combretum erythrophyllum, or river bushwillow, is a riparian specialist adapted to riverbanks and moist environments across . This medium to large 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 , traditionally harvested for tanning and producing dark brown dyes. Combretum molle, the velvet bushwillow, is a widespread species in East and , valued for its extensive medicinal applications in traditional practices. This variable or small , reaching 3-10 meters tall, has velvety leaves and is commonly used to treat ailments such as fever, wounds, , 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. Combretum fruticosum serves as the of the , originally described based on specimens from the by Peter Loefling, a student of , in 1758 as Gaura fruticosa. This scrambling shrub or vine, native from to northern , 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 changes, it remains notable for its ornamental value and role in defining the .

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