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Alstonia
Alstonia scholaris, habit (above), details (below)
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Gentianales
Family: Apocynaceae
Subfamily: Rauvolfioideae
Tribe: Alstonieae
Genus: Alstonia
R.Br.[1]
Type species
Alstonia scholaris typ. cons.
R.Br.[2]
Synonyms[1]

Alstonia is a widespread genus of evergreen trees and shrubs, of the family Apocynaceae. It was named by Robert Brown in 1811, after Charles Alston (1685–1760), professor of botany at Edinburgh from 1716 to 1760.

The type species Alstonia scholaris (L.) R.Br. was originally named Echites scholaris by Linnaeus in 1767.

Description

[edit]

Alstonia consists of about 40–60 species (according to different authors) native to tropical and subtropical Africa, Central America, Southeast Asia, Polynesia and Australia, with most species in the Malesian region.[1]

These trees can grow very large, such as Alstonia pneumatophora, recorded with a height of 60 m and a diameter of more than 2 m.[3] Alstonia longifolia is the only species growing in Central America (mainly shrubs, but also trees 20 m high).[4]

The leathery, sessile, simple leaves are elliptical, ovate, linear or lanceolate and wedge-shaped at the base. The leaf blade is dorsiventral, medium-sized to large and disposed oppositely or in a whorl and with entire margin. The leaf venation is pinnate, with numerous veins ending in a marginal vein. Phyllotaxy is whorled i.e. two or more leaves arises at a node and form a whorl .

The inflorescence is terminal or axillary, consisting of thyrsiform cymes or compound umbels. The small, more or less fragrant flowers are white, yellow, pink or green and funnel-shaped, growing on a pedicel and subtended by bracts. They consist of 5 petals and 5 sepals, arranged in four whorls. The fertile flowers are hermaphrodite. The gamosepalous green sepals consist of ovate lobes, and are distributed in one whorl. The annular disk is hypogynous. The five gamesepalous petals have oblong or ovate lobes and are disposed in one whorl. The corolla lobes overlapping to the left (such as A. rostrata) or to the right (such as A. macrophylla) in the bud. The ovary has 2 separate follicles with glabrous or ciliate, oblong seeds that develop into deep blue podlike, schizocarp fruit, between 7–40 cm long. The plants contain a milky latex, rich in poisonous alkaloids. Fijians use the latex of A. costata (saurua, sorua) as a form of chewing gum.[5] The Alstonia macrophylla is commonly known in Sri Lanka as 'Havari nuga' or the 'wig banyan' because of its distinct flower that looks like a woman's long wig.

Alstonia trees are used in traditional medicine.[6]

Many Alstonia species are harvested for timber, called pule or pulai in Indonesia and Malaysia. Trees from the section Alstonia produce lightweight timber, while those from the sections Monuraspermum and Dissuraspermum produce heavy timber.

Alstonia trees are widespread and mostly not endangered. However a few species are very rare, such as A. annamensis, A. beatricis, A. breviloba, A. stenophylla and A. guangxiensis.

Species

[edit]

Alstonia has five distinct sections, each a monophyletic group; Alstonia, Blaberopus, Tonduzia, Monuraspermum, Dissuraspermum.[citation needed]

As of April 2025, Plants of the World Online accepts the following 44 species:[1]

[edit]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Alstonia

View on Grokipedia
from Grokipedia
Alstonia is a of flowering plants in the family , comprising approximately 43 species of trees and shrubs distributed across the paleotropics, from through tropical and (including and the ) to and the western Pacific islands. The genus was established by Robert Brown in 1810, with Alstonia scholaris designated as the , and is characterized by its production of milky , whorled leaves, and fragrant, salver-shaped flowers. Species typically inhabit diverse environments such as lowland rainforests, secondary regrowth, montane forests, and savannah grasslands, from sea level to elevations of 2100 meters. Morphologically, plants in the Alstonia are often large trees reaching up to 60 meters in height, featuring buttressed trunks and distinctive pagoda-like branching patterns in some . Leaves are arranged in whorls of 3 to 9, elliptic to lanceolate in shape, petiolate or sessile, and contain colleters in their axils for . Flowers are five-merous, with a corolla tube longer than the lobes and pubescence on the inner surface, producing paired linear follicles as fruits that contain numerous seeds. Ecologically, the is adapted to a range of tropical and subtropical conditions, though it rarely forms pure stands, and its abundant in young twigs serves as a defense mechanism. Many Alstonia species hold significant ethnobotanical value, particularly in across their native regions, where bark, leaves, and are used to treat ailments such as fever, , , , and conditions. The genus is rich in bioactive compounds, including over 100 alkaloids (such as echitamine and vallesamine), triterpenoids (e.g., and ), flavonoids, and , which contribute to its pharmacological potential. Scientific studies have validated several traditional uses, demonstrating antiplasmodial, , , antidiabetic, , and hepatoprotective activities in extracts of species like A. scholaris, A. boonei, and A. macrophylla. Additionally, the wood of certain species is valued for timber in furniture and construction, while from A. scholaris has been used in production.

Taxonomy

Etymology and History

The genus Alstonia was named by Scottish botanist Robert Brown in 1810, in recognition of (1683–1760), a fellow Scottish botanist who held the position of Professor of Botany at the from 1738 until his death and served as the first keeper of the Edinburgh Botanic Garden, where he advanced botanical education and plant classification. Alston's contributions to and his critiques of early systematic botany, including Linnaean methods, influenced the naming choice, though he predeceased the genus's description by over half a century. In its early taxonomic history, Alstonia faced considerable confusion with other genera, including Ochrosia, Rauvolfia, Tabernaemontana, Tonduzia, Winchia, and Echites, owing to overlapping vegetative features like whorled leaves and production; distinctions were ultimately drawn based on the genus's paired, slender follicle fruits containing comose . This ambiguity persisted during 19th-century tropical explorations, which yielded initial species descriptions such as A. venenata from southern (based on collections from Indian subcontinental surveys) and A. spectabilis from (drawn from Southeast Asian voyages), contributing to the genus's recognition as a element within the family. Key historical revisions shaped the understanding of Alstonia, with Friedrich Markgraf's 1974 revision offering the first comprehensive global treatment, recognizing approximately 43 species across five sections and resolving many nomenclatural issues from prior scattered descriptions. Subsequent works built on this foundation, and modern phylogenetic analyses using morphological and molecular data have affirmed the genus's and its placement within the subtribe Alstoniinae of , underscoring its evolutionary coherence despite its wide distribution.

Classification and Phylogeny

Alstonia belongs to the tribe Alstonieae within the subfamily Rauvolfioideae of the family . This placement reflects modern classifications based on , which have restructured the family from earlier schemes where Alstonia was included in the subtribe Alstoniinae of the tribe Plumerieae (previously under Vinceae in some 19th- and early 20th-century systems). The genus is one of the largest in the tribe, comprising trees and shrubs adapted to tropical environments. Phylogenetic analyses using molecular data, including plastid genes such as rbcL and matK, confirm Alstonia as monophyletic and position the Alstonieae as an early-diverging lineage in the Rauvolfioideae, sister to the Aspidospermateae. These studies, incorporating extensive plastid supermatrices from over 1,000 species, highlight the genus's basal role in the family's evolutionary history, with estimates around 79–85 million years ago. While broader relations within Rauvolfioideae link Alstonieae to other tribes like Hunterieae (including genera such as Picralima), specific close affinities for Alstonia are primarily with congenera like Dyera based on shared morphological and anatomical traits. Approximately 44 are currently accepted in Alstonia, though taxonomic revisions continue due to morphological variability and new collections from understudied regions. Infrageneric divisions recognize five sections—Tonduzia, Monuraspermum, Alstonia, Blaberopus, and Dissuraspermum—supported by cladistic analyses of morphological characters, , and , which delineate groups based on , morphology, and adaptations. These sections provide a framework for understanding evolutionary diversification within the , with ongoing molecular work refining boundaries.

Description

Morphology

Alstonia species are primarily evergreen trees or shrubs, ranging from 5 to 60 meters in height, characterized by straight trunks often featuring buttresses and fluted bases, particularly in larger individuals, along with distinctive whorled branching patterns in groups of four to five. Young plants may exhibit pagoda-like branching, contributing to their tiered canopy structure. The leaves are simple, entire, and typically arranged in whorls of three to nine, though rarely , with petiolate or sessile attachment; they are elliptic, lanceolate, or oblanceolate in shape, leathery to subcoriaceous in texture, and measure 5 to 20 cm in length. Lateral veins are numerous and often terminate in an intramarginal vein, while margins are entire and sometimes slightly revolute; colleters are present in leaf axils, and the leaves may be glabrous or pubescent, drying flat. Flowers are arranged in terminal or axillary cymes, forming much-branched inflorescences up to 120 cm long in some , and are 5-merous, actinomorphic, and fragrant, with colors ranging from white to yellowish, pink, or red. The corolla is salverform or hypocrateriform, featuring a cylindrical tube that dilates distally and is pubescent inside, measuring 5 to 10 mm in length and longer than the lobes, which overlap and at ; stamens are inserted in the throat with included ovate anthers, and an annular disk is present at the base of the superior, 2-carpellate united by a common style. Fruits consist of paired, slender follicles that are terete, slightly woody, and smooth, free or connate at the base, ranging from 10 to 60 cm in length and containing numerous elliptic to oblong . The are ciliated, with tufts of hairs () longest at the ends, facilitating dispersal. Milky latex is a characteristic feature present throughout all plant parts, abundant in young twigs and branchlets but scant or absent in mature trunks, typical of the family.

Reproduction

Alstonia species generally display seasonal flowering phenology, with blooming often synchronized to environmental cues such as the transition to the dry season or post-monsoon periods in tropical regions. In Alstonia scholaris, for instance, flowering occurs from November to February in subtropical Asia, featuring creamy white to greenish-white flowers that open at dusk and produce small amounts of nectar (approximately 2.7 μl per flower) alongside a pungent scent to attract nocturnal visitors. Individual trees may show asynchronous blooming within populations, but the overall pattern aligns with cooler, drier months to optimize reproductive success. Pollination in the genus is predominantly entomophilous, facilitated by the salverform corollas with cylindrical tubes that are pubescent inside and dilated distally, suiting long-tongued insects such as moths and bees. In A. scholaris, documented pollinators include moths (22% of visits, e.g., Palpita vitrealis), butterflies (29%), bees (40%), and flies (9%), with the dusk anthesis promoting cross-pollination through inter-tree movements. Many species are self-compatible and capable of facultative autogamy or geitonogamy, though some, like A. berteroi, exhibit self-incompatibility, resulting in higher fruit set from cross-pollination between unrelated individuals. Pollen-ovule ratios in studied species suggest adaptation for xenogamy, with hand-pollination experiments confirming effective outcrossing. Seed production follows successful , yielding paired or solitary dehiscent follicles that mature in approximately 3 months, containing numerous oblong or linear . In A. scholaris, open-pollinated fruit set reaches about 58%, with follicles dehiscing longitudinally during the to release comose bearing long silky hairs at both ends for dispersal (anemochory). viability is influenced by environmental factors, including storage conditions and ; submersion treatments and dry storage can reduce rates significantly, while of the hard seed coat enhances viability by up to several months post-dispersal. is typically and cued by adequate and light exposure following dispersal, with rates varying from 50–80% under optimal tropical conditions. Asexual reproduction is rare in Alstonia but occurs via root suckers in certain species under disturbance, such as fire or mechanical damage. For example, A. muelleriana produces coppice shoots and suckers from the base after fires, aiding regeneration in fire-prone habitats, while A. scholaris can form suckers post-cutting to facilitate vegetative spread. This mode is limited compared to and primarily serves recovery rather than primary propagation.

Distribution and Habitat

Geographic Range

The genus Alstonia exhibits a distribution, with native species occurring primarily in tropical and subtropical regions of , , , and the Pacific islands, while representation in the is limited to and . In , species are concentrated in tropical West and Central regions, extending eastward to and , where they inhabit lowland rainforests. Asian distribution spans from the and through to southern , encompassing diverse tropical forests. In , the genus reaches (particularly ), , and extends across numerous Pacific islands as far east as the Marquesas, forming a broad palaeotropical belt. Centers of diversity for Alstonia are most pronounced in the region, including , , the , and , where approximately 19–20 species occur, representing a significant portion of the genus's estimated 44 accepted species worldwide. For instance, African species such as A. boonei are widespread across rainforest zones in West and , contributing to regional timber and medicinal resources. This concentration in Malesia underscores the genus's evolutionary hotspot in Southeast Asian archipelagos. Several Alstonia species have been introduced beyond their native ranges for ornamental and medicinal purposes. Notably, A. scholaris is cultivated in subtropical areas such as , where it has naturalized in parts of the state, and , including islands like Oahu and Kauai, often planted in urban and coastal landscapes. These introductions highlight the genus's adaptability to non-native tropical environments. Historical biogeography suggests that Alstonia likely originated in , supported by the oldest known fossils from the Eocene Palana Formation in , indicating early diversification in the region. Subsequent dispersal to , , and the Pacific is inferred to have occurred via long-distance mechanisms such as ocean currents and avian vectors, facilitating the genus's expansion across isolated landmasses.

Habitat Preferences

Alstonia species primarily inhabit tropical biomes such as rainforests, , and secondary woodlands, often occurring as canopy or emergent trees in lowland to mid-elevation settings ranging from to approximately 2,100 m. These environments provide the humid, shaded conditions essential for their growth, with the showing a particular affinity for disturbed or regrowth areas like edges and lands where light penetration supports establishment. The prefers well-drained, fertile soils, including alluvial, basaltic red earths, and lateritic types with a range of 5.5 to 7.5, thriving in climates characterized by high annual rainfall of 1,000–3,800 mm and mean temperatures between 12–32°C. Some exhibit tolerance to seasonal , enabling persistence in monsoon forests with periodic dry periods. These conditions support rapid growth, particularly in moist, nutrient-rich substrates that facilitate root development. Alstonia trees are frequently associated with mixed dipterocarp forests, where they contribute to canopy diversity, and are commonly found along riverbanks in damp, floodplain habitats that retain moisture during dry seasons. As , they colonize disturbed sites due to their fast growth rates and ability to regenerate in canopy gaps or cleared areas, aiding in tropical ecosystems. Adaptations such as prominent buttress roots provide stability in wet, shallow soils prone to waterlogging and erosion.

Ecology

Pollination and Seed Dispersal

Alstonia species exhibit a mixed pollination strategy involving both nocturnal and diurnal pollinators, which contributes to their reproductive flexibility in tropical environments. Nocturnal moths, particularly from the family Sphingidae such as Daphnis nerii, are key pollinators for species like Alstonia scholaris, drawn to the strongly fragrant, white flowers that bloom at night. Diurnal pollinators include bees (e.g., honey bees) and butterflies, which visit flowers during the day to collect nectar or pollen, facilitating both self- and cross-pollination in species such as Alstonia venenata and A. scholaris. These plants are self-compatible and capable of facultative autogamy, allowing reproduction even in low pollinator density, though cross-pollination enhances genetic diversity. Seed dispersal in Alstonia primarily occurs through anemochory, facilitated by , with seeds featuring hairy or appendages that enable them to float over moderate distances. Fruits are paired follicles that dehisce when dry, releasing numerous flat, winged seeds—typically several to hundreds per follicle—typically during the for optimal conditions, as observed in A. scholaris and A. venenata. Autochory, involving dehiscence for short-range dispersal, supplements this mechanism, while hydrochory may play a minor role in riparian species and animal-mediated dispersal remains rare across the genus. Fresh seeds generally show high germination rates, often approaching 100% without pretreatment, though viability declines rapidly post-dispersal. The reproductive efficiency of Alstonia is bolstered by high seed output per follicle, compensating for variable success and dispersal losses, with enabling colonization of new areas in disturbed habitats. poses emerging threats by altering and abundance, potentially disrupting moth and bee visitation patterns, and shifting dry-season dispersal windows through irregular rainfall. These factors could reduce and establishment success, particularly for specialist moth-pollinated populations.

Interactions and Conservation Status

Alstonia species exhibit various biotic interactions that contribute to their ecological roles and survival strategies. The milky produced by these trees acts as a primary defense against herbivory, with its sticky and toxic alkaloids entangling and poisoning mouthparts or bodies of feeding , thereby reducing damage from pests like caterpillars and beetles. Additionally, the fruits of several Alstonia species are consumed by birds and bats, which facilitate by carrying and depositing seeds away from the parent tree, enhancing colonization in fragmented habitats. These trees form symbiotic relationships with arbuscular mycorrhizal fungi, which colonize their roots to improve nutrient uptake, particularly phosphorus, in nutrient-poor tropical soils, promoting growth and resilience in forest understories. As emergent or canopy trees in tropical rainforests and secondary forests, Alstonia species provide essential ecosystem services, including shade that moderates microclimates for understory plants and habitat structure for epiphytes, birds, and arboreal mammals, supporting overall biodiversity in these dynamic environments. Conservation challenges for Alstonia are significant, driven primarily by habitat loss through for and , as well as overharvesting of bark and for timber and traditional medicines. Species-specific threats vary; for instance, Alstonia beatricis is classified as Vulnerable due to restricted range and logging pressures in Indonesian New Guinea, while is assessed as Least Concern owing to its wide distribution and adaptability. Efforts to conserve Alstonia focus on integrating and strategies to mitigate these threats. Protected areas within hotspots, such as the Heart of Borneo initiative spanning , , and , safeguard habitats for multiple Alstonia species against . includes seed banking and cultivation programs to support sustainable harvesting, particularly for medicinal species like Alstonia boonei, reducing pressure on wild populations while preserving genetic diversity.

Chemical Composition and Uses

Bioactive Compounds

Alstonia species are particularly noted for their abundance of monoterpenoid alkaloids (MIAs), which constitute the primary class of bioactive compounds isolated from various plant parts. These alkaloids are structurally diverse, encompassing subtypes such as corynantheine-strychnine, sarpagine, ajmaline, macroline, eburnamine/kopsifoline, and bisindole alkaloids, with over 300 reported variants across the genus. Prominent indole alkaloids include alstonine, echitamine, and akuammicine, all derived from the strictosidine biosynthetic pathway, where strictosidine serves as the key intermediate formed from and secologanin precursors. Alstonine is a characteristic MIA featuring an core with a complex polycyclic structure, while echitamine, first isolated in 1875, is a crystalline with characteristics, and akuammicine belongs to the corynantheine-strychnine group, exhibiting a pentacyclic . These structures arise through enzymatic cyclizations and rearrangements in the MIA pathway, contributing to the chemical diversity observed in Alstonia. In addition to alkaloids, Alstonia contains such as , a pentacyclic triterpenoid found in bark and leaves; , including glycosides like isookanin-7-O-α-L-rhamnopyranoside; and steroids such as β-sitosterol and , primarily in root and stem bark. Iridoids and triterpenes, such as and β-amyrin, further diversify the terpenoid profile. Biosynthesis of these compounds predominantly occurs in the bark and leaves, where MIA production is regulated by enzymes catalyzing oxidative steps from geissoschizine intermediates. Species variation influences compound abundance; for instance, A. scholaris exhibits higher levels of alstonine compared to other species like A. boonei, which is richer in echitamine and . Such variations are linked to genetic differences in biosynthetic clusters, as revealed in studies of A. scholaris. Extraction of bioactive compounds from Alstonia traditionally involves aqueous decoctions of bark or leaves to yield crude mixtures, as practiced in ethnomedicinal preparations. Modern methods employ solvent extraction with or , followed by chromatographic techniques such as , (HPLC), and HPLC-ESI-MS/MS for isolation and purification of specific alkaloids and terpenoids. These approaches enable precise standardization, as demonstrated in validated protocols for fractions rich in MIAs.

Traditional and Modern Applications

Alstonia species have been utilized in across and , particularly through bark decoctions employed to treat , fever, and . In , the stem bark of A. boonei is widely prepared as a febrifuge and antimalarial remedy, often combined with other herbs for enhanced efficacy against febrile illnesses and gastrointestinal disorders like dysentery. In Asian indigenous practices, the bark of A. scholaris serves similar purposes, addressing , , and via oral decoctions that act as tonics and antiperiodics. Contemporary pharmacological studies have validated several of these traditional applications, focusing on bioactive alkaloids such as alstonine derived from Alstonia species. Alstonine demonstrates potent antimalarial activity against Plasmodium falciparum, with an IC50 of 0.17 μM in the 3D7 strain over 96 hours, showing high selectivity (>140-fold) over human cells and no cross-resistance to multidrug-resistant parasites. Extracts and alkaloid fractions from A. scholaris leaves exhibit anti-inflammatory effects by inhibiting COX-2, 5-LOX, and reducing pro-inflammatory mediators like NO and PGE2 in murine models of edema and air pouch inflammation. Additionally, the n-butanol fraction of A. scholaris stem bark displays antihypertensive potential through calcium channel blockade and activation of soluble guanylate cyclase, reducing blood pressure dose-dependently in rat models (e.g., 54.87 mmHg decrease at 100 mg/kg intravenously). Recent studies (as of 2025) have also explored anticancer activities, with extracts showing cytotoxic effects against various cancer cell lines, including breast and lung cancers. Beyond medicinal uses, Alstonia provides practical resources in various regions. The lightweight, workable timber of species like A. boonei and A. scholaris is employed for furniture, components, and light construction, valued for its durability in humid climates. Several species are cultivated as ornamental shade trees in tropical gardens due to their foliage and attractive form. The bark of A. constricta yields dyes for traditional coloring. Alkaloids in Alstonia, including alstonine, contribute to both therapeutic benefits and potential risks, as referenced in detailed chemical analyses. Safety profiles indicate primarily from high doses of these alkaloids, manifesting as emetic effects like and in animal models, observed acutely at 4 g/kg in dogs and transiently in sub-chronic studies up to 120 mg/kg over 13 weeks. Ongoing pharmacological research emphasizes and antimalarial efficacy, with extracts of A. boonei showing promise in murine models of P. berghei , supporting further development as affordable therapies in endemic areas.

Species

Diversity and Key Characteristics

The genus Alstonia comprises approximately 40–45 accepted species of trees and shrubs, with around 100 synonyms arising from historical taxonomic descriptions. The highest diversity occurs in the region, particularly , though the genus has a distribution extending from and tropical Africa to the southwestern Pacific. Infrageneric classification recognizes five sections—Tonduzia, Monuraspermum, Alstonia, Blaberopus, and Dissuraspermum—delineated primarily by leaf arrangement and morphology. For instance, section Alstonia features species with whorled leaves, while variations in structure, such as follicle shape and characteristics, distinguish other sections. These groupings are supported by macromorphological, , and wood anatomical data, reflecting evolutionary patterns within the genus. All Alstonia species are latex-bearing plants typical of the Apocynaceae family, producing digitoxigenin-type cardenolides as defensive compounds in their latex and tissues. Taxonomic revisions have significantly reduced the species count from over 150 proposed names through extensive synonymy; a key 1998 monograph by Sidiyasa recognized 43 species, including five newly described, by consolidating variants based on type specimens and regional floras. Earlier works, such as Monachino's 1949 revision of African taxa, laid groundwork but left broader inconsistencies that the later study resolved.

Notable Species

Alstonia scholaris, commonly known as the devil tree or blackboard tree, is a prominent in the , capable of reaching heights of up to 50 meters with a straight bole and whorled leaves measuring 10-20 cm in length. Native to tropical regions spanning southern , , , and , it thrives in rainforests and is valued for its lightweight timber used in furniture, blackboards, and construction. The bark has long been employed in across for treating , respiratory ailments, and , with phytochemical studies confirming its antimalarial and properties. Alstonia boonei, referred to as the African stinging or God's , is a large endemic to tropical West and , growing up to 45 meters tall with a cylindrical bole reaching 27 meters and a girth of 3 meters. Its bark is a key resource in traditional African medicine, particularly for combating fevers, , and rheumatic pain, and it is listed in the African pharmacopoeia for antimalarial applications. The species faces threats from overharvesting and habitat loss, rendering it endangered in certain regions like parts of and . Alstonia macrophylla, a restricted to including , , and the , features large, elliptic leaves up to 30 cm long and is recognized as a medium-heavy suitable for timber applications. It plays an important ecological role in swamp forests, contributing to the stability of these fragile ecosystems in areas like and . The wood is utilized locally for construction and crafting, supporting regional economies while highlighting the need for sustainable harvesting in these biodiverse habitats. Other notable species include Alstonia congensis from , where its stem bark is traditionally used to treat , , and , and Alstonia angustiloba, an endemic to valued for its medicinal properties against and wounds, as well as its potential in .

References

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