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Dodonaea
Dodonaea
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Dodonaea
Dodonaea viscosa foliage and flowers
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Sapindales
Family: Sapindaceae
Subfamily: Dodonaeoideae
Genus: Dodonaea
Mill.[1]
Species

See text

Dodonaea, commonly known as hop-bushes,[2][3] is a genus of about 70 species of flowering plants in the soapberry family, Sapindaceae. The genus has a cosmopolitan distribution in tropical, subtropical and warm temperate regions of Africa, the Americas, southern Asia and Australasia, but 59 species are endemic to Australia.[4][5]

Plants in the genus Dodonaea are shrubs or small trees and often have sticky foliage, with simple or pinnate leaves arranged alternately along the stems. The flowers are male, female or bisexual and are borne in leaf axils or on the ends of branchlets and lack petals. The fruit is an angled or winged capsule.

Description

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Plants in the genus Dodonaea are shrubs or small trees that typically grow to a height of 0.1–4 m (3.9 in – 13 ft 1.5 in) and are dioecious, monoecious or polygamous and often have sticky foliage. The leaves are arranged alternately along the stems and are simple or pinnate. The flowers have three to seven sepals but that fall of as the flowers mature, but no petals. There are six to sixteen stamens (except in female flowers), the filaments shorter than the anthers, the style is threadlike and the ovary in female flowers has two to six carpels. The fruit is a two to six angled or winged capsule.[2][4][5]

Taxonomy

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The genus Dodonaea was first formally described in 1754 by Philip Miller in The Gardeners Dictionary and the first species he described was Dodonaea viscosa.[6] The genus name (Dodonaea) honours Dodonaeus, (the latinised form of Dodoens), a Flemish physician and botanist.[7]

Systematics

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Dodonaea is one of the largest genera in the Sapindaceae, and includes 70 species widely distributed in continental Australia.[8] The only other species of the Dodonaea widely spread beyond mainland Australia, Dodonaea viscosa, is believed to be one of the world's most greatly disseminated transoceanic plants.[8]

The first attempts to distinguish infrageneric categories within genus Dodonaea were based on leaf morphology, specifically, two sections - Eu-Dodonaea (simple leaves) and Remberta (pinnate leaves) were differentiated.[8] Later this sectional classification was expanded by Bentham, who included 39 species in five series - four simple-leaved series further divided on capsule-appendage morphology (series Cyclopterae, Platypterae, Cornutae and Apterae) and one pinnate-leaved species (series Pinnatae).

Later the genus was reviewed extensively two times. Radlkofer identified Dodonaea as a part of the tribe Dodonaeeae, within Dyssapindaceae, together with Loxodiscus, Diplopeltis and Distichostemon. Dodonaea and Distichostemon share similar morphological characteristics which include plants having regular flowers without petals and an intrastaminal disc. Therefore, these two genera are considered to be closely related.[9]

54 Dodonaea species identified by Radlkofer were divided into three series (Cyclopterae, Platypterae and Aphanopterae) and six subseries.[8]

Another revision of the genus was proposed by West, where Dodonaea were divided into six species groups by using a combination of characters.[10] Species with the most primitive characters were classified in Group 1 and Group 6 included plants with the most derived states. For instance, the character of an aril possession was recognized as a derived trait.

The most recent molecular study of phylogenetic relationships within the genus revealed some discrepancy with the previously stated hypotheses of morphological evolution within Dodonaea which classified taxa by the combination of leaf, capsule and seed characters. As in preceding morphological research,[10] species with compound leaves were identified in several clades, interspersed among species with simple leaves (e.g. D. humilis is the only species in Clade I with imparipinnate leaves). The breeding system has great variation across the phylogeny, and although most species are dioecious, sometimes some species may differ from this state being monoecious. Most genera in Sapindaceae are dioecious, however, most closely related to Dodonaea in the phylogeny (Diplopeltis, Diplopeltis stuartii and Cossinia) are monoecious. It has also been reported that whereas normally breeding system in Harpullia is dioecism, a few species have also been recognized as monoecious.[11] It was stated that during evolution a general breeding-system across the phylogeny was dioecism, however, the polygamous state was intermediate or, might be partially reversible.

Molecular data supports an evidence that monophyly of Dodonaea includes all species of Distichostemon.[8] It is also supported by the morphological characters as synapomorphies of flowers with reduced petal number and with a highly reduced intrastaminal disk, the trait which is absent in staminate flowers. Both West and Radlkofer used an aril presence or absence as a character to define species groups. All the main clades of Dodonaea and also two species of Diplopeltis have small funicular arils.[8] Seeds of D. viscosa have very small funicular aril, and are harvested by Pheidole ants and deposited in middens outside the nest after the elaiosome has been consumed.[12]

Bayesian MCMC estimation of Dodonaea phylogeny supported the hypothesis that two species of Cossinia are sisters to Diplopeltis and Dodonaea.[8] Nevertheless, Diplopeltis is identified as a paraphyletic group. The monophyly of Dodonaea is well supported by Bayesian MCMC estimation (1.00 posterior probability, PP).[8]

Dodonaea viscosa is placed within the Clade IV, being closely related to D. biloba, D. procumbens and D. camfieldii. It is known that D. viscosa and D. camfieldii evolved in Australia from their most recent common ancestor.[8]

Species

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As of April 2024, the following species are accepted by Plants of the World Online in the genus Dodonaea:[13]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Dodonaea is a genus of approximately 70 species of flowering shrubs and small trees in the soapberry family, Sapindaceae, commonly known as hop-bushes. The genus exhibits a primarily Australian distribution, with more than 60 species endemic to the continent and found across all states in diverse habitats such as rocky ridges, woodlands, and shrublands, excluding alpine, heathland, and rainforest communities. It also has a broader cosmopolitan range in tropical, subtropical, and warm temperate regions of Africa, southern Asia, the Americas, and the Pacific islands. Most species are small shrubs reaching 1–2 meters in height, though some, like , can form trees up to 8 meters; they are typically dioecious with alternate, simple or compound leaves that are often glabrous or variably hairy. Flowers are inconspicuous and greenish, borne in raceme-like or paniculate inflorescences, while the fruits are distinctive papery capsules with three or four wings that facilitate wind dispersal. Dodonaea species hold ecological value as plants for and birds, and they are widely cultivated for ornamental purposes due to their , colorful winged fruits, and attractive foliage—particularly cultivars like D. viscosa 'Purpurea' with its purple leaves. Traditionally, various species have been used in for treating ailments such as , wounds, fevers, and gastrointestinal issues, attributed to their content including and terpenoids, while some show potential in agricultural .

Description

Morphology

Dodonaea species are typically shrubs or small trees, ranging from 0.1 to 8 m in , with erect to spreading growth forms that may include prostrate habits in certain taxa. The stems feature branchlets that are terete to angular or winged, often viscous due to sessile glands, and covered in resinous exudates that impart a sticky texture to the foliage. This resinous quality arises from glandular hairs, contributing to the plant's characteristic glossy or viscid appearance. Leaves in the genus are simple or pinnately compound, arranged alternately along the stems, and measure 0.2–15 cm in length, with widths of 0.05–4.7 cm. They exhibit coriaceous texture, eucamptodromous venation, and margins that are entire or serrulate to toothed, often with acute to obtuse apices and attenuate bases; many species display viscid surfaces from glandular secretions. Compound leaves, when present, are imparipinnate with 2–31 leaflets that are oblong to lanceolate. Flowers are small and inconspicuous, typically under 5 mm in diameter, and lack petals entirely. They consist of 3–7 free or basally fused sepals that are ovate to lanceolate, valvate or imbricate, and often viscous or caducous, surrounding 6–16 stamens with filaments 0.1–3.5 mm long and oblong to linear anthers. Plants are dioecious or polygamo-dioecious, with unisexual flowers occasionally showing aborted organs indicative of bisexual origins. The fruit is a dry, dehiscent capsule, 2–6 angled or winged, measuring 2–38 mm long and up to 30 mm wide, with a membranous to crustaceous texture. It splits septicidally or septifragally into three valves, releasing 1–5 lenticular, black seeds per locule that are 1.4–3.8 mm in size and may be arillate or exarillate. Wings, when present, are broad (0.5–11.5 mm) and extend from apex to base, often turning red to at maturity. For instance, D. viscosa features narrow, wavy, viscid leaves 2.5–8 cm long and three-winged capsules 10–15 mm across. Dodonaea belongs to the family.

Reproduction

Dodonaea species exhibit through flowers that typically bloom from spring to summer, depending on regional and . The are predominantly dioecious, bearing separate (staminate) and (pistillate) flowers on different individuals, though some species or populations produce bisexual flowers, indicating monoecious tendencies in certain cases. flowers consist of exposed stamens that release , while flowers feature a pistil adapted for development. Pollination in Dodonaea is primarily anemophilous, relying on wind to transfer pollen between flowers, facilitated by the lack of petals and nectar to avoid attracting unnecessary visitors. However, opportunistic insect pollination occurs, with bees observed collecting pollen from stamens despite the absence of rewards like nectar. This dual mechanism ensures reproductive success in diverse environments, though wind remains the dominant vector. Following successful , female flowers develop into dehiscent capsules that split open to release . These capsules bear papery wings, enabling wind-mediated dispersal of the small , which measure approximately 1.4–3.8 mm in size. The are black, compressed-ovoid or lenticular, and are dispersed by the winged capsules over short to moderate distances. Seed germination in Dodonaea is constrained by physical imposed by a hard, water-impermeable seed , necessitating to break dormancy and allow . Mechanical or chemical methods, such as acid treatment or hot water immersion, promote rates up to 91% under optimal temperatures around 25°C. Scarified seeds exhibit , with viability persisting for several years in soil seed banks due to the impermeable . In some regions, secondary seed dispersal in D. viscosa involves ants of the genus Pheidole, which harvest seeds attracted to a small funicular aril, transporting them to nests where the aril is removed, potentially enhancing survival by burying seeds. This myrmecochory supplements wind dispersal, particularly in semi-arid grasslands.

Taxonomy

Etymology and History

The genus Dodonaea is named after Rembert Dodoens (1517–1585), a prominent Flemish physician and botanist known in Latin as Dodonaeus, in honor of his influential work on and plant descriptions. formally established the genus in 1754 within the fourth edition of his The Gardeners Dictionary, adhering to the Linnaean system of that emphasized precise, standardized naming. Prior to this formal description, referenced the —now classified as —in his foundational (1753) under the name Ptelea viscosa, reflecting early European botanical interest in the plant's distinctive features but placing it within a different . This pre-establishment mention underscores the gradual recognition of Dodonaea species amid the era's expanding exploration of global . The common name "hop-bush" originated from historical confusions in European herbals, where the 's papery, three-winged seed capsules were likened to those of the unrelated hop plant (), leading to misidentifications and shared vernacular uses in brewing substitutes. The today encompasses approximately 70 species, predominantly native to and surrounding regions.

Systematics

Dodonaea is the largest within the Dodonaeoideae of , comprising approximately 70 , the majority of which are endemic to . The is positioned in Dodonaeoideae, one of four subfamilies in , and its is strongly supported by molecular phylogenetic analyses using nuclear ribosomal ITS and partial ETS sequences, with Bayesian posterior probabilities of 1.00. This includes the former Distichostemon, whose have been transferred to Dodonaea based on shared morphological synapomorphies such as petal-less flowers and the absence of a reduced intrastaminal disk in staminate flowers, corroborated by unequivocal molecular evidence. Internally, Dodonaea is classified into sections such as Eu-Dodonaea, characterized by simple leaves and winged capsules, and Remberta, distinguished by pinnate leaves, with divisions primarily based on capsule wing structure and leaf morphology. Cladistic analyses reveal that Australian endemics form distinct clades, often linked to arid and temperate biomes, reflecting recent s from the onward. The closest relatives to Dodonaea include genera such as Diplopeltis, Cossinia, and Loxodiscus within Dodonaeoideae, with Distichostemon now subsumed, highlighting a dry-adapted in the Australian flora. Hybridization within Dodonaea is rare but documented, particularly between D. viscosa and D. procumbens, with putative hybrids observed in populations from and Victoria exhibiting intermediate morphological features. Recent molecular studies since 2000 have refined the genus's , increasing the recognized count from 68 (as per earlier revisions) to around 70, incorporating new descriptions and phylogenetic insights that resolve previous uncertainties in circumscription.

Species

The genus Dodonaea comprises approximately 72 accepted species worldwide, with the vast majority—around 59—endemic to . This high level of underscores Australia's role as the primary center of diversity for the , where exhibit adaptations to a range of environmental conditions. Diversity within Dodonaea is particularly pronounced in Australia's arid and semi-arid zones, with over 50 occurring in these regions, often on rocky outcrops or in mallee communities. In contrast, fewer are found outside ; and host only a handful, predominantly the widespread D. viscosa and its , with limited endemics in highland or coastal habitats. Notable species include , a cosmopolitan known as sticky hop-bush for its resinous foliage, which grows up to 5 meters tall and features winged capsules; it is the most widespread member of the . Dodonaea biloba is a from , reaching 0.5 meters high and up to 3.5 meters wide, distinguished by its bilobed leaves and occurrence in dry shrublands. Dodonaea procumbens, or trailing hop-bush, forms low-growing mats up to 1 meter across and 10-20 centimeters tall, with wedge-shaped leaves and characteristic winged fruits, endemic to southeastern . Dodonaea camfieldii, a rare endemic to , is a spreading to 1 meter tall with narrow leaves and solitary or paired flowers, confined to specific coastal habitats. Several Dodonaea species face conservation challenges due to loss from , , and grazing; for instance, D. stenophylla is listed as vulnerable in , with populations restricted to fragmented arid woodlands.

Distribution and Ecology

Geographic Distribution

The genus displays a , though it is predominantly concentrated in tropical, subtropical, and warm temperate zones worldwide. Native occurrences span southern regions of , southern , parts of the including and , and , with the highest in these areas. This broad range reflects the genus's adaptability to diverse climates within these zones, though it remains absent from colder temperate or boreal regions. Australia stands out as the epicenter of Dodonaea diversity, accommodating the majority (approximately 60) of the genus's 72 accepted species across coastal habitats to inland arid interiors. In contrast, African species are largely restricted to southern locales, with notable endemism and concentrations in Madagascar and South Africa. Southern Asian representatives occur in scattered subtropical pockets, while in the Americas, the genus is native to select Pacific islands like Hawaii and southern continental sites such as Chile. The species D. viscosa exemplifies the genus's expansive reach, exhibiting a distribution facilitated by transoceanic dispersal through ocean currents and, more recently, human-mediated introductions. This has led to established populations far beyond its Australian origins, including in regions like the and where it was not originally native. No Dodonaea species are native to , but D. viscosa has been introduced and naturalized in Mediterranean countries as an .

Habitat and Adaptations

Dodonaea species primarily inhabit dry, rocky, or sandy soils within open s, scrublands, and coastal dunes, where they demonstrate remarkable tolerance to poor, saline, and drought-prone conditions. The genus, comprising 72 accepted , is commonly found in , , and communities across tropical and subtropical regions, often on well-drained reddish silty clay loams or basalt-derived soils. For instance, , one of the most widespread , thrives from sea level to elevations near 8,000 feet in diverse settings, including high mountains like those in south-western at 2,100 meters. These plants exhibit key adaptations suited to arid and disturbed environments, including deep or fibrous spreading root systems that enhance water access and on slopes and dunes. Resinous leaves, characteristic of species like D. viscosa, form a protective that minimizes and water loss while potentially deterring herbivores through and . Additionally, some species display fire-resistant bark and coats that weaken under heat, promoting post-fire and survival in fire-prone ecosystems. Dodonaea plants also tolerate levels up to 300 mM NaCl during germination, further enabling persistence in coastal or saline habitats. Ecologically, Dodonaea serves as a in disturbed areas, rapidly colonizing sites to facilitate succession by providing shade and suppressing competitors through allelopathic effects from leaf extracts that inhibit growth. It supports by offering and resources for and birds, while its root systems aid in and watershed protection. The genus thrives in semi-arid to Mediterranean climates, with D. viscosa enduring annual rainfall from 12 to 98 inches, temperatures ranging from 10–32 °C, and conditions like dry desert heat or ocean winds. However, species such as D. viscosa can become invasive in non-native grasslands, encroaching on communities due to their competitive growth and tolerance to disturbance.

Uses and Cultivation

Traditional and Medicinal Uses

Australian Aboriginal communities have traditionally utilized Dodonaea viscosa root juice for treating cuts and toothache. The plant's seeds contain saponins and have been used traditionally as fish poison in some cultures. The leaves of Dodonaea species, particularly D. viscosa, contain bioactive compounds such as saponins and flavonoids, which contribute to anti-inflammatory and antimicrobial effects by inhibiting microbial growth and reducing inflammation in affected tissues. Similar traditional medicinal uses, such as for inflammation and wounds, are reported for other Dodonaea species, attributed to shared phytochemicals like flavonoids and terpenoids. Pharmacological studies have demonstrated analgesic properties of these extracts in animal models, where they exhibited pain-relieving effects comparable to standard anti-inflammatory agents in induced pain assays. In Australian traditions, the bark of D. viscosa is employed for tanning leather, owing to its tannin content that binds to animal hides for preservation, though it is noted as lower quality compared to other sources. Across Pacific regions, including Hawaii, the plant serves cultural roles such as using seeds and papery capsules as beads in jewelry and leis, while D. viscosa is applied in traditional Hawaiian medicine to treat skin ailments like rashes and infections through topical leaf poultices that soothe irritation and promote healing. Early European settlers in Australia historically substituted the hop-like capsules of D. viscosa for true hops (Humulus lupulus) in beer brewing, utilizing their bitter flavor and aromatic qualities to ferment yeast and produce acceptable brews during shortages.

Ornamental and Economic Value

_Dodonaea species, particularly D. viscosa, are widely cultivated as drought-tolerant ornamental shrubs in xeriscaping and landscape designs, valued for their evergreen foliage, rapid growth, and adaptability to harsh conditions. The species serves as an effective hedge, screen, or specimen plant, with varieties such as D. viscosa 'Purpurea' prized for its striking purple-bronze leaves that provide year-round color contrast in gardens. Its tolerance to salt spray, wind, and poor soils makes it ideal for coastal or arid landscapes, where it enhances biodiversity by attracting birds and insects. Propagation of Dodonaea is straightforward, typically via seeds or semi-hardwood cuttings, with seeds requiring or hot water soaking for optimal rates of around 10 days. thrive in full sun and well-drained soils, ranging from sandy to loamy, and are hardy in USDA zones 9–11, where they can withstand temperatures down to about -7°C once established. In cultivation, they grow 3–5 meters tall, preferring minimal after rooting and occasional to maintain shape. Economically, holds potential in revegetation projects for , as its deep stabilize on slopes, gullies, and dunes, significantly improving erosion resistance indices compared to bare or cropped . Its fast growth also supports biomass production for research, with seed oil yields of up to 23% enabling that meets ASTM standards, positioning it as a viable non-edible feedstock for marginal lands. Additionally, the plays a minor role in extraction from leaves, yielding less than 5% oil with properties, though commercial production remains limited. Despite these benefits, D. viscosa can become invasive in certain regions, forming dense thickets in disturbed areas and outcompeting native vegetation; it is considered a weed in parts of and , necessitating careful management such as monitoring and removal in non-native or sensitive ecosystems.

References

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