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Bidens

Bidens is a of flowering plants in the family , comprising approximately 230 of annual and perennial herbs, and occasionally shrubs, commonly known as beggar-ticks, bur marigolds, or sticktights. These plants are characterized by their opposite leaves and fruits bearing two to several barbed awns that facilitate dispersal by clinging to animal fur or human clothing. The name Bidens, derived from Latin for "two teeth," refers to the typical two-pronged awns on the cypselas (achenes), though some have more. These plants exhibit diverse habits, with stems ranging from prostrate to erect, often reaching heights of up to 36 inches (90 cm), and leaves that are or pinnate, arranged oppositely and either sessile or petiolate. Inflorescences consist of radiate, discoid, or disciform heads, typically few per in regions like , featuring an involucre with two series of phyllaries—the outer leaf-like and the inner thinner with scarious margins—and paleate receptacles. Flowers include ray florets (when present) and numerous disk florets, with some like Bidens pilosa occasionally lacking rays; fruits are narrowly club-shaped or compressed, topped by 0 to several barbed awns. Found worldwide, occurring on all continents except , Bidens species are particularly diverse in the , with about eight taxa in alone, including natives like B. alba and introduced ones like B. pilosa. They thrive in disturbed habitats such as roadsides, wetlands, and agricultural areas, preferring full sun and moist to dry s, with a deep system aiding . Ecologically, they serve as attractors and are self-compatible, producing 3,000 to 6,000 per plant that germinate readily on the soil surface in warm conditions above 60°F (15.6°C). Many Bidens species are considered weeds in ornamental and crop production due to their rapid growth and seed dispersal, but others hold cultural and medicinal value; for instance, B. pilosa is utilized traditionally worldwide for its anti-inflammatory and antimicrobial properties and as a food source. Some, like B. laevis, are ornamental with bright yellow flowers blooming until frost, while rare endemics such as B. wiebkei are protected due to habitat threats.

Taxonomy and Classification

Phylogeny

Bidens belongs to the tribe within the subfamily of the family, a placement supported by molecular phylogenetic analyses using ribosomal (ITS) sequences and plastid DNA markers such as matK and trnL-trnF. These studies resolve as part of the alliance, with Bidens species distributed across multiple clades in the tribal phylogeny, reflecting complex evolutionary relationships. Molecular evidence from ITS sequences indicates that Bidens is not monophyletic, with its species dispersed into at least three distinct clades that intermingle with those of closely related genera like and Thelesperma. Earlier plastid and combined analyses also suggested polyphyly, but a 2025 phylogenomic study using whole plastome sequences from 31 species supported Bidens monophyly with high bootstrap values, potentially indicating cytonuclear discordance and highlighting ongoing debate in resolving generic boundaries based on such as awn characteristics. No studies using external transcribed spacer () sequences have been reported specifically for Bidens, though ITS remains the primary nuclear marker for resolving intra-tribal relationships. Key evolutionary divergences within Coreopsideae, including the separation of Bidens lineages from , occurred during the late Eocene to early , with the crown age of the alliance tribes estimated at approximately 40–33 million years ago based on fossil-calibrated phylogenomic analyses. This timing aligns with broader diversification following the family's origin in the , where rapid radiations in the Asteroideae facilitated geographic expansions and trait lability in reproductive structures. Within Bidens, phylogenetic trees reveal major subclades corresponding to geographic distributions, including a North temperate lineage (B-3), a subtropical to tropical lineage (B-1), and an lineage (B-2), each supported by high bootstrap values in ITS and plastid analyses. The lineages form a well-supported to certain clades originating in , indicating a Mesoamerican center of diversification with subsequent dispersals, while the clade represents a distinct Old World radiation. These subclades highlight Bidens' distribution driven by long-distance dispersal events, as evidenced by the nested positions of insular endemics like Hawaiian species within the lineage.

Etymology and History

The genus name Bidens derives from the Latin words bi- (meaning "two") and dens (meaning "tooth"), alluding to the characteristic two awns or teeth on the fruits (achenes) of its species. This etymology highlights a defining morphological feature that aids in seed dispersal by catching onto animal fur or clothing. The name was first formally established by Carl Linnaeus in his seminal work Species Plantarum (1753), where he described the genus and included several initial species, such as B. tripartita as the type, based on specimens from European and North American collections. Subsequent taxonomic expansions began with Carl Ludwig Willdenow's fourth edition of Species Plantarum (1803), which incorporated new combinations and descriptions, such as B. leucantha and B. connata, drawing on herbarium materials to broaden the genus's scope beyond Linnaeus's original framework. In the 19th century, botanists like Edward Lee Greene further advanced the classification by describing numerous new species, including B. vulgata (1899) and B. riparia (1901), while resolving extensive synonymies arising from variable forms within the genus, such as segregating ecological variants of B. cernua and B. laevis. The marked a period of significant splits and monographic synthesis, exemplified by Earl Edward Sherff's comprehensive two-volume treatment The Genus Bidens (1937), published by the Field Museum of Natural History, which recognized over 230 species across 14 sections worldwide and refined distinctions from related genera like . Sherff's work built on prior revisions, incorporating thousands of specimens to address nomenclatural ambiguities and emphasize achene characteristics, thereby stabilizing the genus's amid its cosmopolitan diversity.

Diversity and Species

The genus Bidens comprises approximately 223 of predominantly or herbs, distributed worldwide in tropical and warm temperate regions (as of ). These exhibit a range of growth forms, from erect forbs to prostrate vines, adapted to diverse environments such as wetlands, disturbed areas, and roadsides. The highest diversity occurs in the , where over 170 are native, particularly in North and , representing the center of origin and radiation for the . Several species are notable for their ecological and human significance. , a fast-growing annual herb, is a cosmopolitan weed originating from tropical and Central America but now invasive in tropical and subtropical regions worldwide, often dominating agricultural fields and disturbed habitats due to its prolific seed production and barbed achenes that aid dispersal. , an annual or short-lived perennial, is widespread across temperate zones of Europe, Asia, and North America, commonly found in moist, nutrient-rich soils along rivers and ditches. In contrast, , a perennial subshrub native to Mexico and the southwestern United States, is valued as an ornamental plant for its bright yellow, daisy-like flowers and fern-like foliage, thriving in sunny, well-drained conditions. Taxonomically, Bidens is recognized in a broad sense (Bidens lato), but phylogenetic studies have highlighted , leading to the recognition of infrageneric groups and segregate genera. Bidens stricto encompasses core species with typically two- to four-awned achenes and opposite, often pinnately divided leaves, while segregate genera like Coreocarpus (with about three in and ) are distinguished by their persistent pappus scales, campanulate involucres, and achenes with broader wings or reduced awns, reflecting distinct evolutionary lineages within the Coreopsideae tribe.

Description

Morphology

Bidens species are primarily erect or prostrate or perennial herbs, typically ranging from 10 to 150 cm in height, though some tropical forms exhibit shrubby or climbing habits up to several meters. Stems are often terete or angled, branched, and pubescent to glabrous, with variations in color from green to purplish. The is generally fibrous with a deep in many . Leaves in the genus Bidens are opposite (occasionally alternate on upper stems), simple to pinnately compound, with serrate or lobed margins, and measure 2–24 cm in length. They are typically lanceolate to ovate, petiolate, and may be glabrous or sparsely hairy, providing for the upright growth form. The consists of solitary or clustered capitula (flower heads) that are radiate or discoid, 6–40 mm wide, borne on peduncles up to 18 cm long. Ray florets, when present, number 0–8 and are yellow to white with ligules 2–40 mm long, surrounding 5–240 yellow tubular disc florets. The cypselae (fruits) are linear to obovate, 2.5–22 mm long, dorsally compressed or angled, and topped with 2–4 (up to 8) barbed awns 0.5–9 mm long that aid in dispersal.

Reproduction and Life Cycle

Bidens primarily reproduce sexually through , with many exhibiting self-compatibility that facilitates within individual flowers, though a subset of , particularly in diverse habitats like , display mixed mating systems involving both autogamy and . In autogamous such as , pollen transfer occurs autonomously due to the structure of the disc florets, leading to high seed set rates without external pollinators, while allogamous taxa rely on cross-pollination for . This reproductive strategy contributes to the genus's widespread success, as selfing ensures prolific seed production even in isolated populations. Flowering periods vary by and environmental conditions, generally spanning from spring to autumn in temperate latitudes, with blooms initiating as early as May and extending through in regions like , influenced by photoperiod and temperature cues. The life cycle of Bidens encompasses both annual and perennial forms, adapting to a range of ecological niches. Annual species, such as Bidens frondosa and Bidens pilosa, germinate in spring, undergo vegetative growth, flower, set seed, and senesce within a single growing season, typically completing their cycle in 3-6 months under favorable conditions. Perennial species, including Bidens laevis, persist for multiple years, overwintering as basal rosettes that protect meristems from frost and enable regrowth the following season. Seed germination across the genus is stimulated by light exposure and adequate soil moisture, with many species showing optimal rates at alternating temperatures of 25/15°C to 30/20°C and reduced viability under water stress below -0.3 MPa; light requirements prevent deep burial from triggering premature sprouting, promoting surface-level establishment. Asexual reproduction occurs in select perennial Bidens species via underground rhizomes, enabling clonal expansion and persistence in stable habitats. For instance, Bidens laevis produces horizontal rhizomes that generate new shoots and roots, bypassing to rapidly colonize wet areas and maintain genetic uniformity in populations. This vegetative propagation supplements seed-based dispersal, enhancing resilience against environmental variability, though it is less common than sexual modes across the .

Distribution and Habitat

Geographic Range

The genus Bidens is native to all continents except and exhibits a , with the highest concentrations of occurring in the tropical regions of the —particularly and —and throughout . This widespread native range spans tropical, subtropical, and warm-temperate zones globally, encompassing diverse ecosystems from the to , , and . Comprising approximately 230 species, Bidens demonstrates significant regional variation in species richness, with over 150 species documented across North, Central, and , including 25 species in north of alone. In , more than 50 species contribute to this diversity, while hosts around 30 species, many adapted to tropical and environments. Several species, such as B. , have become highly invasive cosmopolitan weeds, now established in over 40 countries across tropical and subtropical regions through human-mediated dispersal.

Ecological Preferences

Species of the genus Bidens predominantly favor disturbed habitats, including wetlands, roadsides, agricultural fields, and areas, where they exhibit a ruderal strategy that allows rapid colonization of successional environments following disturbances such as or . These thrive in moist, moderately fertile soils that retain water, often associating with nutrient-rich, eutrophic conditions in or semi-aquatic settings, reflecting their high demands particularly for . Bidens species demonstrate broad tolerance to ranging from 4 to 9, encompassing slightly acidic to neutral conditions, and they adapt to various substrates like , , and even saline soils up to 100 mM NaCl. They prefer full sun exposure for optimal growth and flowering, though some tolerate partial shade in semi-open habitats. In terms of climatic preferences, tropical forms of Bidens, such as B. pilosa, exhibit optima between 20°C and 30°C, with favored at 25–30°C, while the genus overall tolerates a wider range of 15–45°C and annual rainfall from 500 to 3,500 mm. Altitudinal distribution spans from to over 3,000 m, with Andean species like B. macroptera extending up to 3,700 m and some reaching 4,000 m in high-elevation successional sites.

Ecology and Interactions

Pollination and Dispersal

Pollination in the genus Bidens is predominantly entomophilous, with insects serving as the primary vectors through attraction to nectar rewards in the disc florets. Species such as B. pilosa produce sucrose-dominant nectar with a concentration of approximately 25%, supplemented by amino acids, which draws a diverse array of pollinators including butterflies (e.g., Eurema hecabe), bees (e.g., Apis cerana), wasps, and flies. Recent studies have identified specific floral volatiles in Bidens species that enhance attractiveness and selectivity for pollinating insects, varying by species and influencing visitation rates. In related species like B. eatonii, bees (e.g., bumblebees and halictids) and various flies (e.g., syrphids and tachinids) are key visitors, facilitating pollen transfer via secondary pollen presentation on the style branches. Wind plays a secondary role in some contexts, but entomophily remains essential, as evidenced by the absence of seed set in pollinator-excluded populations of species like B. campylotheca. Many Bidens species exhibit self-compatibility, enabling or when pollinators are scarce, though breeding systems are typically mixed with significant . For instance, in Hawaiian Bidens taxa, self-compatibility supports facultative selfing, but rates across 15 populations average 0.65 (ranging from 0.43 to 0.88), promoting through insect-mediated . In invasive species like B. pilosa and B. alba, self-compatibility is pronounced, with high fruit set (up to 84%) under open pollination, yet pollinator visits by hymenopterans and lepidopterans reduce by favoring xenogamy. This mixed strategy enhances reproductive assurance in variable environments, with protandry and herkogamy further modulating selfing levels. Seed dispersal in Bidens relies heavily on epizoochory, where barbed awns on the achenes adhere to animal , feathers, or , facilitating external transport away from the parent plant. The fruits, characterized by two to four rigid, retrorse awns, exemplify the "Bidens type" dispersal syndrome, combining epizoochory with limited autochory and occasional endozoochory for effective spread. This mechanism enables long-distance dispersal, with seeds potentially traveling several kilometers per event depending on vector mobility, as attachments persist until grooming or mechanical removal occurs. For example, in B. , the awned cypselas readily hitchhike on mammals or birds, contributing to rapid colonization of disturbed habitats. Anthropogenic dispersal has amplified Bidens invasions, particularly through contamination of seeds and agricultural trade. B. pilosa, native to tropical , was introduced to regions like in 1857 via imported and , subsequently spreading across as a in and other fields since the mid-1800s. This human-mediated pathway, combined with epizoochory, has enabled B. pilosa to establish in over 40 countries, forming dense stands that outcompete natives and reduce crop yields by up to 50%.

Pests and Diseases

Bidens species are susceptible to several insect pests that can cause significant defoliation and transmit pathogens. , particularly , are common pests on Bidens plants, feeding on sap and weakening growth while serving as vectors for viruses. These aphids have been observed on native Hawaiian Bidens species, contributing to stress in natural populations. Leaf beetles in the genus Calligrapha, such as C. bidenticola and C. californica, also target Bidens foliage, with larvae and adults skeletonizing leaves and causing substantial defoliation in affected stands. These beetles are particularly associated with Bidens and related , leading to reduced photosynthetic capacity in infested plants. Fungal diseases pose a notable threat to Bidens, especially in humid environments. Rust fungi, including Uromyces bidentis (a in the Pucciniales order closely related to ), infect , producing pustules on leaves that impair and promote premature leaf drop. This is prevalent in tropical and subtropical regions, exacerbating damage under high moisture conditions. Viral infections, such as Bidens mottle virus (BiMoV), affect weedy Bidens populations by causing mosaic symptoms, stunting, and reduced seed production, with transmission primarily by like M. persicae. , caused by Ralstonia pseudosolanacearum, occurs in wet soils and leads to vascular blockage and plant collapse in like B. pilosa, particularly in tropical agricultural settings. Certain Bidens species exhibit invasive potential that impacts and native ecosystems. Bidens pilosa, a , infests over 40 types worldwide, competing for resources and reducing yields through rapid and prolific production. Its allelopathic effects, mediated by in root exudates and residues, suppress and of native and crops, altering community dynamics in invaded habitats. This invasiveness is amplified in disturbed soils, where B. pilosa can dominate and facilitate secondary pest establishment. Recent research as of 2025 indicates that invasive Bidens species, such as B. frondosa, may show greater resilience to compared to native congeners, potentially expanding their ecological impact through altered distribution and interactions.

Human Uses and Significance

Medicinal and Culinary Applications

Bidens pilosa has been employed in traditional folk medicine across and for treating , with roots and the whole prepared as juices or decoctions to alleviate symptoms. In regions such as and , extracts from B. pilosa are used to combat , supported by in vitro studies showing inhibition of growth by up to 90% with extracts. For , the whole is applied topically as crushed herb or fresh material in African and Asian practices, including decoctions documented since the late 19th century in ethnobotanical records from and . The properties of B. pilosa are primarily attributed to its , such as , centaureidin, and butein, which suppress inflammatory mediators like TNF-α and IL-6 in rodent models. These compounds exhibit activity comparable to standard agents, reducing and with IC₅₀ values of 12.5–25 µg/ml in methanolic extracts. Among Native groups in , such as Amazonian communities, flowers and young leaves are incorporated into edible preparations, including teas and raw additions to meals, as a staple wild green. Pharmacological studies from the 2010s confirm the antioxidant activity of B. pilosa, with extracts demonstrating strong radical scavenging and protection against in cellular models, often outperforming . These effects are linked to and phenolic acids like and , which elevate endogenous antioxidants such as and in assays. However, cautions are advised, though overall extracts show low (LD₅₀ > 2000 mg/kg in rats).

Ornamental and Agricultural Roles

Hybrids of Bidens ferulifolia, such as 'Goldilocks Rocks', are popular in ornamental for their compact growth and prolific bright yellow, daisy-like blooms that persist from spring through fall. These cultivars thrive in full sun and well-drained , making them ideal for hanging baskets, window boxes, and container plantings where their trailing habit adds vibrant color and texture. They exhibit strong once established, requiring minimal watering beyond occasional deep soaks during prolonged dry spells. Modern breeding efforts for B. ferulifolia hybrids began intensifying in the early , focusing on traits like reduced height for better container performance and extended bloom periods without deadheading. 'Goldilocks Rocks', developed through selective hybridization, exemplifies this progress with its mounded-trailing form reaching 12-18 inches tall and wide, suited to USDA hardiness zones 9a through 11b. In agricultural contexts, certain Bidens species, particularly B. pilosa, serve as spontaneous cover crops that help mitigate by stabilizing particles and reducing runoff in tropical and subtropical systems. For instance, in orchards, groundcover including B. pilosa has been shown to limit loss to as low as 0.55 Mg ha⁻¹ per period, comparable to planted cover crops like jack beans. Additionally, B. pilosa provides value in some regions, acting as a supplementary feed for and due to its nutritional content. However, Bidens species are often regarded as problematic weeds in row crops like soybeans, where B. pilosa can cause significant yield reductions through competition for resources. Reported losses in soybean production range from 9.4% to 43% depending on weed density and environmental factors, underscoring the need for integrated management in affected areas.

Conservation and Threats

Status of Key Species

The genus Bidens encompasses approximately 230 worldwide, with the majority assessed as () on the due to their widespread distributions and stable populations, though several cosmopolitan such as B. pilosa are actively monitored for their invasive tendencies in non-native regions. However, a subset of endemic faces significant conservation challenges, particularly those restricted to oceanic islands and isolated continental habitats, where and competition from introduced exacerbate vulnerability. In the Hawaiian Islands, 19 species of Bidens are endemic, representing an from a single ancestral lineage, and many are highly threatened by non-native competitors that displace them through resource competition and alteration. For instance, Bidens wailele, a recently described endemic to , is classified as (CR) under IUCN criteria, with an estimated population of only 700–800 individuals across three remote locations, primarily due to ongoing loss from and feral ungulates. Similarly, Bidens hillebrandiana subsp. hillebrandiana, restricted to the windward eastern coast of Kohala on the island of Hawai'i in coastal dry lands on bluffs and sea cliffs, holds a CR D status ( due to very small population), with threats intensified by competition from alien plants in its specialized dry land . Among continental endemics, Bidens meyeri from Rapa, , exemplifies acute risk, assessed as (CR) based on its tiny population of fewer than 50 mature individuals confined to a single remote site on volcanic cliffs.

Conservation Efforts

Conservation efforts for within the Bidens primarily target those classified as threatened or endangered, focusing on protection, , and population . Many Bidens are widespread and resilient, but a subset, particularly endemics in ecosystems and coastal wetlands, face risks from loss, , and climate impacts. Efforts are often coordinated through national and state agencies, botanical gardens, and conservation networks, emphasizing ex situ preservation and management to prevent extinctions. One prominent example is Bidens wiebkei, an endangered Hawaiian species endemic to Moloka'i, where federal recovery plans guide actions under the U.S. Endangered Species Act. The 1995 Recovery Plan for the Moloka'i Plant Cluster outlines priorities such as fencing to exclude ungulates like deer and goats, control of invasive plants (e.g., Melinus minutiflora and Schinus terebinthifolius), and fire prevention in its . Propagation efforts include seed collection and cultivation by the National Tropical Botanical Garden and Waimea Arboretum, supporting genetic stock maintenance and potential reintroductions. In 2016, the U.S. Fish and Wildlife Service designated 7,515 acres of critical habitat across coastal, lowland wet, and montane wet ecosystems on Moloka'i to safeguard recovery areas. Population augmentation and monitoring continue, with interim goals aiming for three stable populations of at least 50 mature individuals each. For Bidens eatonii (Eaton's beggar-ticks), a globally vulnerable (G3) annual herb restricted to brackish marshes along the U.S. Atlantic coast, conservation integrates state-level protections and national stewardship. Listed as endangered in Massachusetts, New Jersey, Connecticut, and New York, it benefits from legal safeguards under state endangered species acts, prohibiting collection or habitat disturbance without permits. The Center for Plant Conservation maintains B. eatonii in its National Collection, providing long-term seed banking and propagation to preserve genetic diversity. In Massachusetts, efforts include habitat monitoring through the Natural Heritage Program and prevention of degradation from shoreline development, pollution (e.g., oil spills), and hydrological alterations. Field surveys in the Hudson River estuary have documented and mapped populations to inform management, emphasizing restoration of tidal wetlands. Similar actions in Pennsylvania and Maine focus on maintaining estuary integrity against adjacent land-use pressures. Other threatened Bidens species receive targeted interventions, often through regional programs. Bidens meyeri, a (CR) species from Rapa in known from fewer than 50 individuals on volcanic cliffs, requires immediate monitoring and habitat assessment, though specific actions remain limited due to its remote location; recommendations include ex situ propagation to mitigate risks from stochastic events. In the Galápagos and other Pacific islands, broader ecosystem restoration indirectly supports Bidens endemics by controlling invasives and protecting cliff and habitats. For mainland species like Bidens bidentoides (endangered in ), efforts center on preservation and removal to sustain small populations. Across the genus, collaborative networks like the Plant Conservation Alliance promote seed banking and research to enhance resilience against , prioritizing high-risk island taxa.

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