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Phytolacca
Phytolacca
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Phytolacca
Phytolacca acinosa foliage and fruit
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Phytolaccaceae
Subfamily: Phytolaccoideae
Genus: Phytolacca
L.[1]
Species

About 25 to 35 species

Synonyms

Pircunia Bertero ex Ruschenb.[1]

Phytolacca is a genus of perennial plants native to North America, South America and East Asia. Some members of the genus are known as pokeweeds or similar names such as pokebush, pokeberry, pokeroot or poke sallet.[2][3] Other names for species of Phytolacca include inkberry and ombú. The generic name is derived from the Greek word φυτόν (phyton), meaning "plant," and the Latin word lacca, a red dye.[4] Phytolaccatoxin and phytolaccagenin are present (in the leaves, stems, roots, blossoms, berries etc.) in many species which are poisonous to mammals if not prepared properly. The berries are eaten by birds, which are not affected by the toxin. The small seeds with very hard outer shells remain intact in the digestive system and are eliminated whole.[citation needed]

The genus comprises about 25 to 35 species of perennial herbs, shrubs, and trees growing from 1 to 25 m (3 to 80 ft) tall. They have alternate simple leaves, pointed at the end, with entire or crinkled margins; the leaves can be either deciduous or evergreen. The stems are green, pink or red. The flowers are greenish-white to pink, produced in long racemes at the ends of the stems. They develop into globose berries 4 to 12 millimetres (0.2 to 0.5 in) diameter, green at first, ripening dark purple to black.[5][6][7]

Selected species

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Although many species are herbaceous, P. dioica forms a substantial tree
Nymboida National Park, NSW, Australia, August 2014.

The following species are accepted by one or more regional floras:[5][6][7][8][9][10]

Formerly placed here

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Ecology

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Members of the genus Phytolacca can be found on every continent except Antarctica, though it is introduced in Europe and Australasia.[11]

Phytolacca is known to be dispersed by birds and box turtles.[12][13]

The ombú (Phytolacca dioica) grows as a tree on the pampas of South America and is one of the few providers of shade on the open grassland. It is a symbol of Uruguay, Argentina and gaucho culture. P. weberbaueri from Peru also grows to tree size. Both species have massively buttressed bases to their trunks, and very soft wood with a high water storage capacity which makes them resistant to grass fires and drought.[14]

In the Pacific Northwest of North America, pokeweed is an invasive species.[15]

Uses

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Phytolacca americana (American pokeweed, pokeweed, poke) is used as a folk medicine and as food, although all parts of it must be considered toxic unless, as folk recipes claim, it is "properly prepared."[citation needed] The root is never eaten and cannot be made edible.[16] Poke salad ('poke salat') is considered part of traditional southern U.S. cuisine, where it is cooked three times in three changes of boiling water to remove some of the harmful components.[17] Toxic constituents which have been identified include the alkaloids phytolaccine and phytolaccotoxin, as well as a glycoprotein.[18]

Fossil record

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A Phytolacca-like fossil has been described from the Upper Cretaceous (late Campanian) Cerro del Pueblo Formation, Coahuila, Mexico, it is a permineralized multiple infructescence composed of berries with six locules, each containing a single seed with a curved embryo developed in a curved ovule with pendulous placentation, a berry anatomy that is similar to that of the genus Phytolacca. Though this new plant from Coahuila shares reproductive characters with Phytolacca, the constant number (six) of carpels per fruit and pendulous placentation support the recognition of a new genus, Coahuilacarpon phytolaccoides.[19]

Notes and references

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Phytolacca is a of flowering plants in the family Phytolaccaceae, within the order , consisting of approximately 25 species of perennial plants that exhibit diverse growth forms ranging from herbaceous perennials to shrubs and small trees. These plants are primarily native to the , including North and , and , where they thrive in a variety of habitats such as open woods, fields, roadsides, and disturbed areas. Morphologically, species in the genus are characterized by alternate, simple leaves, small white to pinkish flowers arranged in racemes or , and fruits that are typically berry-like and range from green to deep purple or black at maturity. The most well-known species is , commonly called American pokeweed or pokeberry, which is a tall herbaceous native to the and parts of and . This species features robust reddish-purple stems, large lanceolate leaves up to 10 inches long, and clusters of dark purple berries that attract birds but are toxic to humans and . Other notable species include , a tree-like form from with a massive trunk and pendulous berry clusters, and from , valued in . Members of the Phytolacca genus have been utilized in across various cultures for treating inflammatory conditions, with compounds such as esculentosides exhibiting properties. Additionally, some species demonstrate antiviral activity due to proteins like pokeweed antiviral protein (PAP), which inhibits , and they possess a conserved ability to accumulate rare earth elements from . However, many parts of these plants contain toxic alkaloids like phytolaccatoxin and phytolaccigenin, which can cause severe gastrointestinal distress, making proper preparation essential for any uses, such as the young shoots of P. americana. In some regions, Phytolacca species are considered invasive, spreading rapidly via bird-dispersed seeds.

Taxonomy

Etymology and history

The genus name Phytolacca derives from the phytón (φυτόν), meaning "plant," combined with lakke or lac, referring to a red dye, alluding to the pigment extracted from the berries of species like P. americana. The genus was initially proposed by in his Institutiones Rei Herbariae around 1700 and formally validated by in the first edition of in 1753, where he established Phytolacca americana as the based on specimens from and . Linnaeus's description emphasized the plant's herbaceous habit, racemose inflorescences, and berry-like fruits, distinguishing it within the nascent system of . Taxonomic understanding of Phytolacca advanced through the as European botanists incorporated collections, with Alfred Moquin-Tandon providing the first monographic treatment of the Phytolaccaceae in Augustin Pyramus de Candolle's Prodromus Systematis Naturalis Regni Vegetabilis (volume 13, part 2) in 1849; this work expanded the genus by describing over a new and varieties, such as P. octandra and P. icosandra var. fraseri, while clarifying sectional divisions based on and carpel numbers. Earlier confusions with genera like Rivina—described separately by Linnaeus in 1753—were resolved by emphasizing differences in fruit morphology, with Rivina featuring single-seeded berries versus the multi-seeded ones in Phytolacca. By the early , further revisions, including those by Heinrich Gustav Adolf Engler and Käthe Krause in Das Pflanzenreich (1934), consolidated the genus around 25 , incorporating palynological and anatomical data to affirm its core position within Phytolaccaceae.

Classification and phylogeny

Phytolacca belongs to the family Phytolaccaceae, which is placed in the order according to the IV (APG IV) classification system. This family, in its strict sense (Phytolaccaceae s.s.), comprises approximately 5 genera and 35 , primarily distributed in the , with Phytolacca as the largest genus containing 20–35 . The APG IV framework, based on extensive molecular data, recognizes Phytolaccaceae as distinct from previously broader circumscriptions that included polyphyletic elements, emphasizing its position within the non-core subclade. Phylogenetic analyses have confirmed the monophyly of , supported by high bootstrap values (BS = 100, posterior probability PP = 1) across multiple datasets, including whole genomes, nuclear ribosomal DNA (nrDNA), and genes such as rbcL and matK. The genus forms a well-supported within Phytolaccaceae, closely related to genera like Ercilla and Anisomeria, with Ercilla as its nearest sister. Broader relationships place Phytolaccaceae sister to Petiveriaceae (including ) and near Nyctaginaceae within , diverging from the core that includes . This divergence is evidenced by early 2000s molecular studies using nuclear 18S rDNA, rbcL, atpB, and matK sequences, which demonstrated the of the traditional Phytolaccaceae s.l. and its separation from through cladistic analyses of 127 taxa. Phylogenomic studies estimate the genus diversified around 20 million years ago in the early . Infrageneric classification of Phytolacca formerly recognized three subgenera based on the degree of carpel connation (Nowicke ). However, recent phylogenomic reconstructions indicate that these subgenera do not correspond to monophyletic groups, with species like P. dioica diverging early and morphological traits failing to align with molecular clades. Key studies, such as those employing (ITS) sequences in the early 2010s, further supported the narrowed family boundaries and of Phytolacca by resolving its position amid the family's historical taxonomic revisions.

Diversity and selected species

The genus Phytolacca comprises 25 accepted species according to POWO (2024), though estimates vary from 20 to over 35 depending on taxonomic treatments, reflecting ongoing refinements in classification. The highest diversity occurs in the Neotropics, particularly including , , and , with additional centers in tropical and temperate to subtropical . Among the most notable species is Phytolacca americana, the North American pokeweed, a perennial herbaceous erect herb that reaches up to 3 m in height, characterized by its robust stems, large ovate leaves, and clusters of dark purple berries. In tropical regions, P. dioica, known as the ombú tree, stands out as a deciduous tree growing 10–25 m tall with a distinctive swollen trunk base up to 4 m in diameter and an expansive umbrella-like canopy. Representing African diversity, P. dodecandra is a scrambling shrub or perennial climber up to 6 m long, featuring tuberous roots, elliptic leaves, and bright red berries used traditionally as a soap substitute due to their saponin content. Some genera previously included in broader Phytolaccaceae s.l. have been segregated based on phylogenetic analyses, such as Lophiocarpus now placed in Lophiocarpaceae, highlighting morphological and anatomical divergences within the order . Infrageneric variation in Phytolacca is pronounced in growth habits, ranging from annual or herbs in temperate zones to woody shrubs and trees in tropical environments, with stems often exhibiting green to red coloration and semi-succulent textures in arid-adapted species. This diversity in habit correlates with ecological adaptations, such as climbing forms in P. dodecandra versus upright trees like P. dioica.

Description

Morphology and growth habit

Phytolacca species display diverse growth habits, ranging from perennial herbs and shrubs to small trees, with approximately 25 in the . These are characterized by succulent, often branched stems that are typically green but frequently exhibit reddish-purple coloration due to the accumulation of pigments, which are water-soluble nitrogenous compounds unique to the order. The stems can be herbaceous and erect in smaller species or more woody at the base in larger forms, supporting upright or spreading growth. Leaves in the are simple, alternate, and petiolate, usually lanceolate to ovate in shape with entire or slightly undulate margins, measuring 10–30 cm long and 5–15 cm wide. They are bright green, smooth or slightly pubescent, and exude a disagreeable when crushed, a trait common across . In forms like certain shrubs and trees, the leaves persist year-round, contributing to the plant's ornamental appeal. Herbaceous possess deep, fleshy systems that store nutrients and water, often displaying a vibrant red interior from pigmentation, which aids in identification and may deter herbivores. Growth patterns vary by ; most have bisexual flowers with simultaneous organs on the same flowers, though dioecious exceptions exist, such as P. dioica. Heights range from 0.5–3 m in herbaceous perennials like P. americana to 10–25 m in arborescent like P. dioica, the ombú tree, which features a broad, domed canopy and rapid juvenile growth from a tuberous .

Flowers, fruits, and reproduction

The inflorescences of Phytolacca species are typically axillary, terminal, or leaf-opposed racemes or spikes bearing 5–100 small flowers, with proximal pedicels occasionally supporting multiple blooms; these structures are often pendulous in species such as P. americana. The flowers are apetalous, featuring 5–8 petaloid sepals that range from white to pinkish or purplish, along with 8–25 stamens arranged in one or two whorls and 6–12 carpels that may be distinct or connate at the base. Most species produce bisexual flowers, though unisexual flowers occur in some, such as P. dioica, where plants are dioecious with separate individuals. Pollination in Phytolacca is primarily entomophilous, mediated by insects such as hymenopterans including bees (Apis cerana indica) and wasps (Crabronidae), which visit the nectar-producing flowers; wind assistance contributes in an ambophilous system observed in species like P. acinosa. The sexual system varies across the genus, with dioecy in certain species necessitating cross-pollination between male and female plants, while bisexual flowers in others allow for self-compatibility and geitonogamy alongside xenogamy. The fruits of Phytolacca are berries, typically oblate with 6–12 locules containing one each (thus 6–12 per ), maturing to a glossy purple-black color that persists through styles at the apex; these berries contain toxic . occurs mainly via endozoochory, with frugivorous birds consuming the attractive fruits and excreting viable , facilitating spread over distances while promoting after passage through digestive tracts. exhibit high longevity, remaining viable in for up to 40 years in like P. americana.

Distribution and ecology

Geographic range

The genus Phytolacca is primarily native to the , with its range extending from southeastern southward through , the , and into as far as . Some species are also indigenous to other continents, including a few in and . Centers of highest species diversity occur in , particularly in , , and , where multiple endemic taxa contribute to the genus's richness; secondary hotspots include tropical and eastern . Patterns of endemism are pronounced, with numerous confined to localized regions—for instance, P. heterotepala is restricted to the state of in northeastern . Several Phytolacca species have been introduced beyond their native ranges, often through human-mediated trade and ornamental planting. Notably, P. americana, originating from eastern , spread to in the 17th century and to by the 19th century, establishing invasive populations in parts of both continents where it disperses rapidly via seeds.

Habitat preferences and interactions

Species of the genus Phytolacca predominantly occupy disturbed habitats such as edges, roadsides, clearings, pastures, borders, and thickets, where they benefit from reduced competition and increased availability. These demonstrate broad tolerance, thriving in well-drained substrates across a wide spectrum from mildly acidic to mildly alkaline conditions, and they prefer medium moisture levels while accommodating short periods of drought. The genus exhibits climatic adaptability, with species distributed across temperate zones in and to tropical and subtropical regions in Central and , the , and parts of . Phytolacca species engage in symbiotic relationships that enhance their ecological fitness, including associations with arbuscular mycorrhizal fungi that facilitate nutrient uptake, particularly , in nutrient-limited soils. Additionally, certain species like P. americana exert allelopathic effects through root exudates containing phytotoxic compounds, which suppress the and growth of neighboring and contribute to competitive dominance in invaded areas. Regarding biotic interactions, Phytolacca display partial resistance to herbivory owing to toxic secondary metabolites that deter mammalian grazers and many herbivores, though generalist insects such as commonly infest foliage and can vector diseases. Fruits, despite their toxicity to mammals, are readily consumed by birds, which play a key role in through endozoochory. Some species, such as P. dioica, further demonstrate , enabling persistence in arid or semi-arid environments.

Human uses and toxicity

Culinary and medicinal applications

In cuisine, the young shoots and leaves of are traditionally prepared as "poke sallet," a dish involving multiple boilings to leach out toxic compounds and render them edible. The process typically includes the greens for about 5 minutes, discarding the water, rinsing, and reboiling until tender, often followed by frying with ingredients like bacon or eggs for flavor. Berries of the plant have been used in limited culinary applications, such as cooked pie fillings, after thorough preparation to mitigate risks. Historically, Native American communities utilized pokeweed for treating , skin ailments like and ringworm, and as an emetic or purgative, often applying root salves mixed with lard for topical relief. In , dating back over 2,000 years, the plant (known as Shang Lu) has been employed for conditions such as , abscesses, and water retention, with root extracts serving as diuretics and anti-inflammatory agents. During the , European settlers and American pharmacopeias incorporated pokeweed extracts as emetics and purgatives for gastrointestinal issues and respiratory infections. In modern herbal practices, poke root is cautiously used for supporting lymphatic drainage in conditions like , , and swollen glands, though professional supervision is advised due to potential side effects. has focused on pokeweed antiviral protein (PAP), a ribosome-inactivating protein isolated from the plant, which exhibits broad-spectrum antiviral activity against viruses including , , and by inhibiting synthesis in host cells. This has led to biotechnological applications, such as transgenic for enhanced resistance and exploring PAP for therapeutic antiviral treatments. Ethnobotanical records emphasize preparation techniques like repeated boiling for leaves (at least twice, changing water each time) and low-dose tinctures for roots to minimize while preserving .

Phytochemicals and toxicity risks

Phytolacca species contain a variety of bioactive phytochemicals, including triterpene saponins such as phytolaccosides A, B, D, E, and G, derived from the aglycone phytolaccigenin, which are present throughout the plant. Lectins, notably pokeweed mitogen (PWM), and the pokeweed antiviral protein (PAP) are key proteinaceous toxins, while betalains contribute to the red-violet pigmentation of fruits and other tissues. These compounds are concentrated in roots, leaves, stems, and seeds, with lower levels in ripe berries. The toxicity of Phytolacca arises primarily from these phytochemicals' disruptive effects on cellular and physiological processes. Triterpene saponins like phytolaccigenin and phytolaccosides induce gastrointestinal distress by disrupting cell membranes, leading to , , , and . such as PWM cause hemagglutination by binding to carbohydrates on surfaces, potentially resulting in systemic effects if absorbed. PAP exerts by depurinating a specific residue in the sarcin/ loop of , thereby inhibiting protein synthesis and contributing to cell death. All parts of Phytolacca are toxic when consumed raw, with roots and seeds posing the highest risk due to elevated concentrations of and PAP. Documented human poisonings often involve ingestion by children, causing acute symptoms like , persistent , , and , while larger exposures can lead to seizures or respiratory distress. In , ingestion results in similar gastrointestinal and neurological effects, underscoring the plant's broad hazard profile. Detection of these toxins relies on bioassays, such as hemagglutination tests for and ribosomal inhibition assays for PAP, while content can be quantified via . Mitigation involves thorough processing, such as repeated boiling and water changes, which reduces and levels to safer thresholds for limited culinary use of young shoots, though complete detoxification is not guaranteed. Professional medical intervention, including activated charcoal and supportive care, is essential for cases.

Conservation and fossil record

Conservation status

Most Phytolacca species are widespread and adaptable, with few formally assessed by the , where evaluated species such as P. dioica are categorized as Least Concern due to their stable populations across South American and savannas. According to the Angiosperm Extinction Risk Predictions developed by the Royal Botanic Gardens, , common species like P. americana and P. dioica are predicted to face no significant risk, with high confidence based on their broad geographic ranges and tolerance. However, rarer taxa show vulnerability; for instance, the recently described P. exiensis from is proposed as Critically Endangered (CR B1ab(iii)) under IUCN criteria, owing to its extremely restricted distribution limited to three localities with fewer than 50 mature individuals and ongoing degradation from agricultural encroachment. Primary threats to Phytolacca species stem from through and agricultural expansion, particularly in tropical and subtropical regions of and where many species are endemic to forest edges and disturbed areas. In the and Himalayan regions, exacerbates these pressures by altering suitable habitats for species like P. acinosa, potentially leading to range contractions and population declines. For P. acinosa in , overharvesting for exacerbates these risks, contributing to declining populations as of 2025. Furthermore, the aggressive invasive behavior of P. americana outside its native North American range can outcompete native vegetation in introduced ecosystems, particularly in parts of , by dominating resources in open woodlands and roadsides, with recent expansion noted in southern as of 2025. Conservation efforts for Phytolacca emphasize in situ protection within established reserves; for example, native populations in southern Brazil's remnants and Uruguay's are safeguarded in areas like the Campos do Planalto Protected Landscape, while P. americana occurs naturally in U.S. national forests and parks such as the . Ex situ strategies include seed banking and living collections in botanic gardens worldwide, such as those at the , to preserve amid . Ongoing research focuses on and propagation techniques to support restoration initiatives in degraded native habitats, though no Phytolacca species are currently listed under for international trade regulation.

Paleobotanical history

The fossil record of the genus Phytolacca and its Phytolaccaceae is sparse, reflecting the limited preservation of these herbaceous to shrubby , but available evidence points to an ancient origin within the order . The earliest definitive attributable to Phytolaccaceae is Coahuilacarpus phytolaccoides, an infructescence consisting of multiple berry-like fruits, each with six locules containing a single and curved , discovered in the Upper (late , approximately 72 million years ago) Cerro del Pueblo Formation in , . This specimen closely resembles reproductive structures in modern Phytolacca species, supporting its assignment to the , though it exhibits fixed carpel number and pendulous not seen in all extant members. Pollen grains resembling those of modern , the broader order containing Phytolaccaceae, have been recorded from Eocene deposits (approximately 50 million years ago) in , indicating early presence of related lineages in the region. Key fossil discoveries further illuminate the family's early diversification. The Coahuilacarpus infructescence represents the oldest reliable record for Phytolaccaceae, predating previously known occurrences and highlighting the family's diversity in low-latitude during the . Additional evidence includes permineralized wood assigned to Petiveria (a in Phytolaccaceae) from early sediments (approximately 20 million years ago) of the El Cien Formation in , , suggesting persistence and possible expansion in western during the . These finds, combined with sparse records, underscore a restricted but significant macro- and history. Phylogenomic analyses estimate the divergence of Phytolaccaceae within around 40–60 million years ago, during the late to early Eocene, with the family's stem age at approximately 59.7 million years ago and crown diversification of Phytolacca sensu stricto beginning in the early around 20.3 million years ago. Fossil evidence, particularly from Laurasian deposits like the Mexican site, supports an origin in northern Gondwana-Laurasia, contrasting with the family's current predominantly distribution. These paleobotanical records imply that Phytolaccaceae adapted to open, marginal habitats—such as fluvial-lacustrine environments—well before their modern pantropical ranges, with early forms likely exploiting disturbed or successional ecosystems in subtropical settings. This prehistoric niche occupation predates the radiation of the genus, highlighting long-term ecological flexibility within the family.

References

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