Commelina is a genus of approximately 170–215 herbaceous flowering plants in the family Commelinaceae, known as the dayflower or spiderwort family, characterized by their often succulent stems that range from prostrate and rooting at nodes to erect, ascending, or climbing, with alternate leaves that are linear to broadly ovate and clasping the stem.[1][2] These plants produce zygomorphic flowers typically in shades of blue (occasionally white or lilac), featuring three unequal petals—two larger and clawed, one smaller—and fertile stamens alongside staminodes, borne in leaf-opposed spathes that may be solitary or fascicled.[2][1] Fruits are loculicidal capsules, usually 2- or 3-locular and 2- or 3-valved, containing 1–5 seeds per locule with a distinctive hilum.[2]As the largest genus in Commelinaceae, which comprises about 40 genera and 600–650 species worldwide, Commelina exhibits a cosmopolitan distribution but is predominantly tropical and subtropical, extending into temperate regions in both hemispheres, with native ranges spanning Africa, Asia, Australia, the Americas, and oceanic islands.[1][3] In North America, for instance, nine species are recognized, several of which are introduced and can behave as aggressive weeds in disturbed habitats, agricultural fields, and along waterways.[2] Species vary from annuals to perennials, some forming tubers or rhizomes for vegetative propagation, and they thrive in moist to wet environments such as forests, grasslands, and margins of water bodies, often displaying C3 photosynthesis.[4][2]Notable for their ephemeral flowers that open in the morning and close by afternoon—hence the common name "dayflowers"—Commelina species play roles in ecosystems as ground cover and pollinator attractors, primarily by bees and flies, while some exhibit self-pollination or cleistogamy in closed flowers.[2] Several species, such as C. benghalensis (tropical spiderwort) and C. diffusa (climbing dayflower), are economically significant as invasive weeds that reduce crop yields through competition and allelopathy, prompting extensive management research.[4] Additionally, various Commelina taxa have traditional medicinal uses in indigenous cultures for treating ailments like inflammation, infections, and digestive issues, attributed to phytochemicals including flavonoids, alkaloids, and phenolics, though toxic potential exists in some.[1] The genus's taxonomic complexity, with ongoing revisions due to morphological variability and hybridization, underscores its botanical importance.[1]
Taxonomy
Etymology and history
The genus name Commelina was established by Carl Linnaeus in honor of the Dutch botanists Jan Commelijn (1629–1692) and his nephew Caspar Commelijn (1667–1731), with the two prominent showy petals of species like Commelina communis symbolizing the brothers who published significant botanical works, while the third smaller, often inconspicuous petal alluded to their less accomplished relative, Caspar Commelijn Jr. (1700–1734).[5][6] Linnaeus proposed the name as early as 1737 in Critica Botanica, drawing from earlier descriptions by Charles Plumier, but formally validated it in his seminal Species Plantarum (1753), where he described nine species primarily based on specimens from tropical and subtropical regions of the Americas and Asia.[6]The establishment of Commelina marked a key step in Linnaean binomial nomenclature for the Commelinaceae family, though the genus concept evolved amid challenges in distinguishing it from morphologically similar genera. Early classifications often confused Commelina with Tradescantia due to overlapping vegetative and floral traits, such as parallel-veined leaves and ephemeral flowers, leading to misplacements of species across Old World and New World taxa in pre-20th-century floras. Over time, refined morphological and later molecular analyses clarified boundaries, expanding recognition to approximately 170–215 species today, predominantly in tropical and subtropical habitats.[2]
Classification
Commelina is classified within the phylum Tracheophyta, class Liliopsida, order Commelinales, and family Commelinaceae, where it represents the largest genus with approximately 170–215 species.[1][2][7]The genus has been traditionally subdivided into subgenera based on floral and seed characteristics, particularly the number of ovules in the ventral locule of the capsule. Subgenus Didymoon C.B. Clarke includes species with two ovules per ventral locule, while subgenus Monoon C.B. Clarke features one ovule per ventral locule; these divisions were proposed by Clarke in 1881 and rely on capsule dehiscence patterns and seedmorphology.[8]Phylogenetic analyses have confirmed the monophyly of Commelina within Commelinaceae, supported by molecular evidence from nuclear ribosomal DNA (specifically the 5S non-transcribed spacer region) and chloroplast DNA (trnL-trnF intergenic spacer). A study sampling 22 species across the genus resolved two major clades— one comprising New World taxa and the other Old World taxa—with bootstrap support of 60% in combined analyses, though traditional subgenera like Didymoon and Monoon were not monophyletic.[9] More recent phylogenomic research using transcriptomic data across Commelinales further reinforces the monophyly of Commelinaceae and its genera, including Commelina, with robust support from hundreds of nuclear loci.[10]
Description
Vegetative characteristics
Commelina species are primarily annual or perennialherbs, exhibiting a range of growth habits from prostrate or decumbent forms to erect or ascending stems, occasionally becoming scrambling or twining in shaded or crowded conditions.[5][11] These plants typically reach heights of 10-100 cm, though some can extend up to 3 m in length under favorable conditions, forming dense mats or tufts adapted to various terrestrial environments.[11][12]The root systems of Commelina are generally fibrous and densely branched, facilitating efficient uptake in moist soils, though tuberous roots occur in certain species such as C. maculata and C. suffruticosa, providing storage for nutrients and water.[12][5] Rhizomatous growth is rare but present in some taxa, contributing to vegetative propagation in humid habitats.[5] Stems are herbaceous, cylindrical, and often succulent, measuring 0.5-1 cm in diameter, with elongated internodes and conspicuously swollen or brittle nodes that enhance structural support and water retention.[11][12] Branching is common, and in prostrate species like C. diffusa, stems root at lower nodes, producing adventitious aerial roots to anchor and spread vegetatively.[13]Leaves in the genus are simple, cauline, and arranged in a distichous (two-ranked) or spiral manner along the stems, typically lanceolate to ovate in shape and measuring 1-11 cm long by 1.5-4 cm wide.[5][11] Blades are sessile or pseudopetiolate, with closed, tubular sheathing bases that often bear reddish-brown hairs at the apex in species such as C. benghalensis, aiding in protection against herbivores or desiccation.[12] The plants produce a mucilaginous sap throughout their vegetative tissues, which contributes to their succulence and resilience in tropical or subtropical settings.
Flowers and fruits
The flowers of Commelina are zygomorphic, featuring three petals that are typically blue or lilac in color, with two larger showy petals and one smaller reduced petal.[11][14] These ephemeral blooms last only one day, contributing to the common name "dayflower" for the genus.[15]The inflorescence consists of flowers arranged in boat-shaped spathes that arise from vegetative nodes, often solitary or in fascicles, enclosing one to several flowers per spathe.[11][15] Each flower has six stamens, comprising three fertile ones—two lateral with introrse dehiscence and one central with abaxial dehiscence—and three staminodes that are bright yellow and serve as visual attractants mimicking pollen.[14][15]The fruits are dehiscent loculicidal capsules that are usually 2- or 3-locular with 1–5 seeds per locule, with the seeds featuring a reticulate or smooth testa and a starchy endosperm.[11][5][15]Pollination occurs primarily through insects attracted by the flower's visual cues, as the family lacks nectaries.[14][15]Most Commelina species exhibit a hermaphroditic sexual system in their chasmogamous flowers, which are open and promote cross-pollination, while some species produce cleistogamous flowers that remain closed for self-pollination, often subterranean in amphicarpic taxa.[15] This dimorphic floral strategy enhances reproductive assurance in varying environments.[15]
Distribution and habitat
Geographic range
The genus Commelina has a cosmopolitan distribution but is predominantly pantropical and subtropical, with native species occurring across tropical, subtropical, and extending into temperate regions of Africa, Asia, Australia, the Americas, and oceanic islands.[1] This widespread range reflects the family's adaptation to warm climates, encompassing diverse ecosystems from moist forests to open grasslands.[16] While the genus includes approximately 170–200 species globally, its highest diversity is concentrated in tropical Africa, where over 100 species are estimated to occur, representing a significant portion of the world's Commelinaflora.[17] This African hotspot underscores the continent's role as a primary center of origin and endemism for the genus, particularly in central and eastern regions like the Democratic Republic of the Congo and Ethiopia.[18]Notable centers of endemism extend to Southeast Asia, where species diversity is prominent in areas such as southern China, India, and the Indo-Malayan region, supporting unique adaptations to monsoon-influenced environments.[19] Recent taxonomic discoveries highlight ongoing revelations in this distribution; for instance, Commelina danxiaensis was described in 2023 from Mount Danxia in Guangdong Province, China, expanding knowledge of Asian endemics with its distinctive morphological traits.[17] Such findings emphasize the genus's understudied variability in eastern Asia, where new species continue to emerge from karst and mountainous habitats.Human-mediated introductions have broadened Commelina's presence beyond its native tropics, particularly into temperate zones. Commelina communis, originally from East Asia, was introduced to eastern North America in the early 19th century via botanical gardens near Philadelphia and has since naturalized widely as a weed.[20] Similarly, it reached central and southeastern Europe around the same period, often escaping cultivation to establish persistent populations in disturbed areas.[21] These patterns of dispersal illustrate the genus's opportunistic spread, now encompassing naturalized occurrences across much of the globe.[13]
Habitat preferences
Commelina species predominantly inhabit moist, shaded environments, including forest understories, wetlands, riverbanks, and disturbed areas such as roadsides and agricultural fields. These plants are well-adapted to conditions with high humidity and partial shade, where they exhibit efficient photosynthesis even under low light levels due to their ability to tolerate shaded canopies.[16] This preference for damp, sheltered niches supports their proliferation in tropical and subtropical ecosystems, reflecting their native origins in these regions.[10]In terms of soil requirements, Commelina thrives in fertile, well-drained loamy soils with a pH ranging from neutral to acidic, typically between 5.8 and 7.0, though some species tolerate slightly higher values up to 8.0. Nutrient availability plays a key role, with significant responses to total nitrogen and available phosphorus levels that enhance growth and distribution. Additionally, these plants demonstrate notable tolerance to seasonal flooding, often occurring in waterlogged sites like rice paddies and stream margins without succumbing to prolonged saturation.[22][16]The altitudinal range of Commelina extends from sea level to approximately 2000 meters, allowing colonization of diverse elevations within their preferred moist habitats. Certain species, such as Commelina erecta, feature underground tubers that enable survival during seasonal dry periods by storing reserves and facilitating regrowth upon the return of moisture.[23]
Ecology
Life cycle and growth
Commelina species exhibit varied life cycles depending on environmental conditions and ploidy levels. In tropical regions, many species function as perennials, persisting through underground rhizomes that allow overwintering and regrowth from rootstocks during favorable seasons.[24] In contrast, temperate populations often behave as annuals, completing their life cycle within a single growing season from seed germination to seed production and senescence.[25] This annual habit is particularly evident in diploid forms, while hexaploid variants in warmer climates support perennial growth.[26][25]Germination of Commelina seeds is primarily triggered by warm temperatures and adequate soil moisture, with optimal ranges of 18–27°C (65–80°F) for aerial seeds and slightly higher, 21–28°C (70–82°F), for subterranean seeds.[24] Once germinated, seedlings exhibit rapid vegetative growth, especially during wet seasons when high soil moisture promotes stem elongation and node rooting, leading to expansive prostrate or ascending shoots.[27] Under non-stressed conditions, plants can increase in height by approximately 2.4 cm per week and width by up to 20-fold over several months, forming dense mats through adventitious roots at nodes.[27]The phenology of Commelina involves flowering that often aligns with seasonal rainfall rather than strict day-length responses, though some species show insensitivity to photoperiod.[28] In tropical and subtropical areas, flowering commences shortly after the onset of rains, producing both aerial chasmogamous flowers and subterranean cleistogamous ones, with peak reproduction in the wet season.[26]Seeds develop within capsules that facilitate dispersal through explosive dehiscence or adhesion to passing animals, enabling colonization of new areas.[26]Growth rates enable Commelina to rapidly colonize disturbed sites, such as roadsides and agricultural fields, where it forms sprawling mats up to 1 m in width via vegetative propagation and seed spread.[26] This aggressive expansion is enhanced in moist, fertile soils, allowing biomass accumulation that outpaces associated crops under adequate water availability.[27]
Ecological interactions
Commelina species, particularly C. diffusa and C. benghalensis, often function as invasive weeds in agricultural systems, where they compete aggressively with crops such as rice, cotton, and soybeans for light, water, and nutrients.[29] These plants can also serve as alternative hosts for pests, including nematodes like Rotylenchulus reniformis and Meloidogyne incognita, as well as fungal pathogens and viruses such as Groundnut rosette virus, thereby exacerbating crop damage in affected regions.[16][26]Flowers of Commelina attract pollinators primarily from the orders Hymenoptera and Diptera, including honey bees, stingless bees, syrphid flies, and bee flies, which facilitate cross-pollination through visits to the pollen-rewarding blooms.[30] Seed dispersal occurs mainly via water in species adapted to moist environments, with some evidence of ant involvement in certain taxa, aiding propagation in wetland and riparian ecosystems.[31][32]Herbivory on Commelina primarily involves insect damage to leaves, with leaf miners from families Agromyzidae and Chironomidae creating tunnels, and chrysomelid beetles like Lema quadrivittata feeding on foliage.[33][34] Some species employ chemical defenses, such as cyanogenic glycosides in C. diffusa, which release hydrogen cyanide upon tissue damage to deter further insect consumption.[35]Most Commelina species are classified as Least Concern by the IUCN, reflecting their widespread distribution and adaptability, though habitat loss in tropical regions threatens narrow endemics like C. catharinensis (Endangered) and C. mwatayamvoana (Critically Endangered).[36][37] Certain species, including C. benghalensis and C. erecta, demonstrate potential in bioremediation by accumulating heavy metals such as cadmium, chromium, and lead from contaminated sediments and soils.[38][39]
Human uses
Medicinal and culinary
Various species of Commelina, such as C. benghalensis and C. diffusa, have been employed in traditional African and Asian medicine for treating inflammation, wounds, and infections, with leaves and roots commonly used in poultices or decoctions to reduce swelling and promote healing.[40] In African traditions, extracts address urinary tract infections, respiratory issues, diarrhea, and rheumatoid arthritis, while in Asian contexts, they manage dermatitis, snake bites, and fever.[40] These effects are attributed to phytochemicals including flavonoids (e.g., quercetin and luteolin derivatives) and alkaloids, which exhibit anti-inflammatory activity by inhibiting pathways like NF-κB and reducing edema in models such as carrageenan-induced paw inflammation.[41][40][42]Historical records document Commelina species in Ayurvedic medicine since ancient times for anti-inflammatory and diuretic purposes, as noted in early texts like Nadkarni's Indian Materia Medica (1926), and in Traditional Chinese Medicine for cooling the blood, clearing heat, and reducing swelling.[43] Modern studies corroborate these uses, demonstrating antioxidant activity through phenolic compounds that scavenge free radicals and protect against oxidative stress, with ethanolic extracts showing significant DPPH radical inhibition.[44][45] However, some species, such as C. benghalensis, have traditional abortifacient uses and may pose toxicity risks, particularly for pregnant individuals; caution is advised, and further safety research is needed.[46]Culinary applications of Commelina focus on young leaves and shoots of C. benghalensis, which are consumed as vegetables in India, Africa, and Southeast Asia, often steamed, stir-fried, boiled in soups, or added to stews and salads.[47] These parts provide nutritional value, including high levels of vitamin C (up to 72.46 mg/100 g dry weight) and carotene (a vitamin A precursor, up to 335.41 mg/100 g dry weight), contributing to dietary intake of antioxidants and essential micronutrients.[48]
Ornamental and other
Several species of Commelina are cultivated ornamentally for their vivid blue flowers, which bloom prominently in summer and autumn, adding aesthetic value to gardens and naturalized landscapes. For instance, Commelina tuberosa forms spreading clumps of erect stems with striking blue petals, making it suitable for herbaceous borders in warm, sheltered sites with fertile, well-drained soil and partial shade.[49]Commelina erecta is similarly valued for its medium blue flowers, often appearing in spring or fall, and thrives in naturalized areas across North America.[23] In tropical regions, C. tuberosa serves as an effective groundcover due to its low-growing, tuberous habit and ability to spread in moist environments.[50]Beyond ornamentals, Commelina species have utilitarian applications, including as fodder for livestock in regions with limited feed resources. Commelina benghalensis offers high nutritional value, with crude protein levels reaching approximately 163 g/kg dry matter in samples from coastal lowlands.[51][52] and is commonly supplemented in ruminant diets in East Africa and Mauritius to enhance intake, digestibility, and nitrogen balance. In southern Ethiopia, C. benghalensis and C. imberbis improve average daily weight gain in lambs by up to 50.63 g when added to low-quality basal diets like teff straw, providing economic returns through better nutrient utilization.[53] The flowers of Commelina communis yield a traditional blue dye used in Japanese crafts, extracted from petals to color woodblock prints (ukiyo-e) and kimonos since the 11th century, producing purplish-to-slate-blue shades on specialized aobanagamipaper.[54] Additionally, species like C. benghalensis and C. communis demonstrate potential in phytoremediation, accumulating heavy metals such as copper (bioaccumulation factor >1) for extraction or stabilizing cadmium (>100 mg/kg in shoots) and lead in polluted soils to reduce mobility.[55][56]Cultivation of Commelina can present challenges due to its weedy tendencies, particularly in temperate zones where species like C. communis and C. benghalensis invade crop fields, reducing yields in soybeans and ornamentals through prolific seeding (up to 1,600 seeds per plant) and vegetative propagation.[16][24] In the southeastern United States, C. benghalensis is classified as a federal noxious weed, requiring integrated management with mulching, crop rotation, and herbicides like glyphosate for control in gardens and agriculture, as it forms dense stands in moist, fertile soils.[24][16]
Species
Diversity
The genus Commelina comprises approximately 207 accepted species, rendering it the largest within the family Commelinaceae, which encompasses around 40 genera and over 700 species total.[57][58] Estimates have varied historically from 170 to 215 species due to taxonomic revisions and ongoing field explorations, with recent discoveries including Commelina danxiaensis from Guangdong Province in China (Asia) in 2023 and two new gem-fruited species (C. almandina and C. sugariae) described in a 2025 revision, highlighting continued additions particularly from Asia and Africa.[58][17][59]Morphological diversity in Commelina spans a wide array of growth forms, from succulent aquatic or semi-aquatic herbs adapted to wetland environments to robust, drought-tolerant perennialspecies capable of surviving in arid or seasonally dry habitats.[19] Flowers exhibit variation in color, predominantly vivid blue but ranging to white or pale variants in certain lineages, often lasting only a single day (hence the common name "dayflowers").[19] Seed morphology shows significant ornamentation diversity, including differences in testa sculpture (smooth, rugose, or tuberculate) and hilum structure, which contribute to dispersal strategies and taxonomic delimitation across the 22 species examined in detailed studies.[60]Infragenerically, Commelina is divided into about 10 sections, primarily delineated by characteristics of the stamens (e.g., filament coloration, presence of staminodes) and capsules (e.g., locule dehiscence patterns, fruit wall texture).[61] Endemism patterns underscore continental specialization, with roughly 50% of species restricted to a single continent—most notably around 100 species endemic to tropical Africa, where diversity peaks, alongside notable endemics in Asia and the Americas.[17] The genus displays a pantropical distribution, with centers of diversity in the Old World tropics.[57]
Notable species
Commelina communis, commonly known as the Asiatic dayflower, is a widespread annual herb native to East Asia and introduced to the Americas and Europe, where it has become a significant invasive weed in agricultural fields, orchards, and disturbed moist areas.[62] It features prostrate to ascending stems up to 1 meter long, with blue flowers that last only a day, and reproduces primarily via seeds from two-valved capsules that aid in dispersal across wet habitats like ditches and roadsides.[62] Its persistence as a weed stems from prolific seed production and ability to root at nodes, making it challenging to control in crops.[62]Commelina benghalensis, or Benghal dayflower, originates from tropical Africa and Asia, serving as a native vegetable in those regions where its leaves and stems are chopped and cooked similarly to spinach during food shortages.[63] As an aggressive invader, it has spread to the Pacific islands, thriving in moist, disturbed sites like roadsides and grasslands, where it competes with crops such as cotton and soybeans through rapid vegetative spread and long-lived seeds.[64] Its creeping habit and drought tolerance contribute to its establishment in tropical environments.[26]Commelina diffusa, the spreading dayflower, is prevalent throughout the Americas, from Mexico to Brazil and the West Indies, often occurring in open swamps, marshes, and agricultural fields as a cosmopolitanweed.[13] It holds medicinal value in traditional practices, with its juice applied to treat wounds, inflamed eyes, and digestive issues, as well as serving as an external remedy for fractures.[13]Commelina tuberosa, a tuber-forming perennial from the Andean regions of Bolivia and Peru through Central America to Mexico, is notable for its edible, starch-rich underground tubers that are locally cooked and consumed.[65] Grown worldwide as an ornamental, it produces bright blue flowers on short stems about 10 cm tall, preferring light, well-drained soils in sunny, sheltered spots.[65]In contrast to the prostrate forms like C. communis and C. diffusa, Commelina erecta exhibits an erect to ascending habit with cespitose stems arising from tufted roots, distinguishing it as a perennial adapted to drier, sandy soils in prairies and open woods.[66]