Convolvulus is a genus of approximately 190 species of flowering plants in the family Convolvulaceae, comprising mainly herbaceous vines or undershrubs that are typically twining or trailing, with simple spiral leaves, funnel-shaped corollas in various colors such as white, pink, or blue, and axillary inflorescences of cymes.[1] These plants are characterized morphologically by overlapping, unequal sepals, filiform stigmas, and petiolate leaves that are often cordate or sagittate at the base, though some species have sessile leaves with narrowed bases.[1] Native to a wide global distribution, Convolvulus species exhibit the greatest diversity in the Irano-Turanian region, including Mediterranean and semi-desert areas, and are adapted to dry, stony, or sandy habitats from sea level to elevations of 3000 meters.[1] Ecologically, they often occupy disturbed sites and can behave as aggressive weeds, such as Convolvulus arvensis (field bindweed), which is a persistent invader in agricultural fields due to its deep root system and perennial habit.[1] Several species hold ornamental value for their attractive foliage and flowers, while others, like Convolvulus pluricaulis, have traditional medicinal uses for central nervous system disorders, and Convolvulus scammonia as a purgative.[2] The genus is distinguished from related groups like Calystegia based on molecular and morphological evidence, with ongoing taxonomic revisions emphasizing its filiform stigmas and inflorescence structure.[1]
Taxonomy
Etymology and history
The genus name Convolvulus derives from the Latin verb convolvere, meaning "to roll together" or "to entwine," a reference to the characteristic twining habit of the plants' stems.[3] This etymology reflects the morphological feature that distinguishes many species as climbing or sprawling vines that wrap around supports.[4]The genus was formally established by Carl Linnaeus in his Species Plantarum (1753), where he provided the first binomial descriptions and included 31 species under a broad concept that encompassed various twining plants now classified elsewhere.[1] Although the name Convolvulus had been used informally for over a century prior in European herbals to describe bindweeds, Linnaeus's work marked its taxonomic foundation, emphasizing floral and vegetative traits like funnel-shaped corollas and alternate leaves.[1] Early recognition by Linnaeus built on observations from predecessors, but his inclusion of diverse taxa, such as those later segregated into Ipomoea, highlighted the initial vagueness in generic boundaries.[1]Through the 19th century, botanists like Robert Brown (1810), who refined distinctions based on stigma morphology, and Jean-Daniel Boissier (1875), who cataloged 66 species in Flora Orientalis, advanced the genus's delimitation by separating it from tropical elements like Ipomoea, characterized by larger flowers and different seed structures.[1] Jacques Denis Choisy (1845) further contributed by emphasizing ovary and fruit features in his revisions, excluding genera such as Evolvulus and early segregates of Calystegia. In the 20th century, monographers like Georg Peter (1891) and later molecular-informed studies consolidated the core concept, reducing the genus to around 200 species worldwide while splitting off woody or herbaceous outliers to related taxa.[1] These revisions progressively narrowed Convolvulus to primarily temperate, herbaceous climbers with filiform stigmas and a bilocular ovary.[1]
Classification and phylogeny
Convolvulus is a genus of flowering plants in the family Convolvulaceae, which belongs to the order Solanales, and comprises approximately 200 accepted species worldwide.[5] The family Convolvulaceae itself is monophyletic, as established by phylogenetic analyses of chloroplast DNA sequences including rbcL, atpB, psbE-J, and trnL-F regions, positioning it as sister to Solanaceae within Solanales.Within Convolvulaceae, Convolvulus resides in the tribe Convolvuleae, which molecular studies confirm as monophyletic based on sequences from ITS, matK, and rbcL loci.[6] Phylogenetic analyses demonstrate that Convolvulus is paraphyletic, with the genus Calystegia nested within it, rendering the traditional boundaries between these taxa indistinct.[7] The tribe Convolvuleae includes three genera: Convolvulus (the largest), Calystegia, and Polymeria, with Polymeria serving as sister to the Convolvulus-Calystegiaclade, which originated around 18 million years ago.[6]A comprehensive 2015 monograph utilized ITS, matK, and rbcL sequences from 140 Convolvulus species and 19 Calystegia species to resolve major clades, identifying two primary lineages: one comprising mostly herbaceous, trailing or climbing herbs with petiolate leaves (Clade X), and another of erect shrubs with indistinct petioles (Clade Y), further subdivided into 11 subclades (A-K) reflecting geographic and morphological patterns.[7] These clades highlight the genus's diversification primarily in the Irano-Turanian region, with subsequent radiations into other temperate areas. Evidence of hybridization, including between Convolvulus species and with Calystegia, has been documented through morphological intermediates and molecular markers, contributing to taxonomic complexity in the group.
Diversity and species
The genus Convolvulus encompasses approximately 200 species, primarily consisting of herbaceous vines and shrubs, though some are subshrubs.[5] The currently accepted number stands at 203 species worldwide, reflecting ongoing taxonomic refinements.[5] These plants exhibit a range of growth habits, from trailing or climbing forms to more upright structures, adapted to various temperate and subtropical environments.Diversity is highest in the Mediterranean basin and adjacent regions, including Macaronesia and western Asia, where more than half of the species—over 100—are concentrated.[8] Notable examples include Convolvulus arvensis (field bindweed), a perennialvine notorious as a widespread agricultural weed due to its persistent rhizomes and ability to smother crops; C. cneorum (silverbush), an evergreenshrub prized for its silvery foliage and ornamental value in dry landscapes; and C. tricolor (dwarf morning glory), a short-lived perennial or annual with vibrant, multicolored flowers popular in horticulture.[9][10]Patterns of diversity within Convolvulus highlight significant endemism, particularly in the Canary Islands, where around 10 species are restricted to these volcanic archipelagos, and in North African regions like Morocco, which harbor unique adaptations to arid habitats.[11] The genus includes both annual and perennial life forms, with perennials dominating in stable environments and annuals more common in disturbed or seasonal settings, contributing to its ecological versatility.[4]A key recent taxonomic revision is the 2015 foundation monograph by Wood et al., which recognizes 190 species and provides detailed descriptions, illustrations, and keys while addressing morphological variability and phylogenetic relationships.[9] This work has served as a benchmark for subsequent studies, though updates like those in Plants of the World Online have slightly expanded the count based on new collections and molecular data.[5]
Description
Morphology
Convolvulus species display diverse growth habits, primarily as herbaceous annuals or perennials that are twining, trailing, prostrate, or occasionally erect, with some forming cushion-like or spiny undershrubs. Stems are typically slender, herbaceous to slightly woody, often producing milky latex, and can extend up to 3 meters in length, often climbing or sprawling over supports.[1][12]Leaves are arranged alternately along the stems, simple, and usually petiolate, though sessile in some subshrubs; blades vary from linear to ovate, cordate, sagittate, or hastate, with lengths of 1-5 cm, and margins that are entire, undulate, crenate, dentate, or sinuate-lobed. Stipules are absent, and the leaves are often glabrous to pubescent, with basal sinus present in sagittate or hastate forms.[1][12]Flowers are borne in axillary cymes or solitary on peduncles up to 5 cm long, with two small bracteoles subtending the pedicel. The corolla is funnel-shaped and 5-lobed, measuring 0.5-4 cm in diameter, typically white, pink, or blue, though yellow in some species like C. scammonia; sepals are five, imbricate, and ovate to lanceolate, while the androecium consists of five stamens and a pistil with a superior ovary and filiform stigmas.[1][12]Fruits are dry, dehiscent capsules that are ovoid to subglobose, bilocular, and 4-valved, containing 1-4 dark brown seeds, often reduced to fewer by abortion. Seeds are 2-6 mm long, smooth to reticulate or tuberculate in texture, and range from glabrous to densely pubescent or hirsute, with some species featuring marginal trichomes that form wing-like appendages aiding dispersal.[1][12]The root system in perennialspecies is extensive and persistent, with some, such as C. arvensis, featuring a primary taproot that can reach depths of 2-6 meters or more, supplemented by numerous adventitious lateral roots that spread horizontally up to 3-5 meters, allowing survival in arid or disturbed soils. Annual species have shallower fibrous roots, while some desert perennials develop thickened woody rootstocks.[10][1]
Reproduction and life cycle
Convolvulus species exhibit both sexual and asexual reproduction, with strategies varying by species and environmental conditions. Sexual reproduction primarily occurs through insect-mediated pollination, where flowers, typically funnel-shaped and white to pink, attract a range of pollinators such as bees and butterflies. Breeding systems vary across the genus, with some species self-compatible and others, like C. arvensis, self-incompatible; outcrossing is generally favored due to the protandrous nature of the flowers and the activity of pollinators during the day. Flowering generally takes place from spring through summer, with individual flowers lasting only one day before wilting, and seed production following pollination leads to the formation of capsules containing 1–4 seeds each.[1][10][13]Asexual reproduction is prominent in perennial species, particularly through vegetative propagation via rhizomes or root fragments. For instance, in Convolvulus arvensis, root segments as small as 5 cm can regenerate into new plants, forming extensive underground networks that enable clonal spread and persistence in disturbed habitats. This mode of reproduction allows perennials to rapidly colonize areas without reliance on seed production, contributing to their invasiveness in agricultural settings.[10][14]The life cycle of Convolvulus differs between annual and perennial species. Annuals, such as C. tricolor, complete their cycle in a single growing season, germinating in spring, flowering in summer, and setting seed before senescence. Perennials, including C. arvensis, overwinter as dormant roots or rhizomes, resuming growth in spring from underground buds. Seeds across the genus often enter dormancy, remaining viable in soil seed banks for over 10 years—up to 50 years in some cases—due to a hard, impermeable seed coat that enforces physical dormancy.[1][10][15]Seed germination typically requires breaking dormancy through scarification—either mechanical abrasion or chemical treatment like sulfuric acid—or exposure to light, along with alternating temperatures (e.g., 20–30°C day/10°C night) and adequate moisture. In the absence of these cues, seeds can persist dormant in the soil, forming long-lived banks that replenish populations after disturbances. For example, in C. arvensis, scarification increases germination rates from under 10% to over 80%, highlighting the adaptive role of dormancy in survival.[15][10][16]
Distribution and habitat
Native range
The genus Convolvulus is primarily native to the Mediterranean Basin, with its range extending across temperate Eurasia, North Africa, and the Macaronesian islands.[5] This distribution encompasses diverse regions including southern Europe (e.g., Spain, Italy, Greece), the Middle East (e.g., Turkey, Iran), and northwest Africa (e.g., Morocco, Algeria), where the genus has evolved in response to seasonal climates.[1] In Macaronesia, particularly the Canary Islands, over 10 species are endemic, highlighting the archipelago's role as a hotspot for speciation within the genus.[17]Centers of diversity are concentrated in Anatolia (Turkey), which hosts approximately 35 species, and the Iberian Peninsula, where species richness is notably high due to varied microhabitats.[18] These areas feature a combination of geological and climatic factors that support elevated endemism and morphological variation. Biogeographically, the genus exhibits disjunct distributions in southern Africa and Australia, resulting from ancient long-distance dispersals rather than recent introductions.[1]Convolvulus species typically inhabit scrublands, grasslands, rocky slopes, and coastal dunes, showing strong adaptations to dry, calcareous soils and semi-arid conditions from sea level to elevations of around 3000 m.[1] These preferences align with Mediterranean-type ecosystems, where the plants thrive in well-drained, stony or sandy substrates that experience low rainfall and high solar exposure.[5]
Introduced ranges and invasiveness
Convolvulus arvensis, commonly known as field bindweed, was introduced to North America as early as 1739, likely as a contaminant in crop and garden seeds brought from Europe.[10] This species has since become widely distributed across the Americas, including both North and South America, as well as Australia and New Zealand, where it thrives in temperate and disturbed habitats far beyond its native Eurasian range.[19]In introduced regions, C. arvensis exhibits high invasiveness by forming dense, sprawling mats of twining vines that smother crops, native vegetation, and other plants, reducing biodiversity and agricultural productivity.[20] It is particularly problematic in farmlands, roadsides, and riparian areas, where its extensive root systems allow it to persist and regenerate even after disturbance.[21] Due to these impacts, C. arvensis is listed as a noxious weed in over 30 U.S. states and several Canadian provinces.[10]The primary pathways of introduction and spread for C. arvensis include contamination of agricultural seeds during trade, as well as intentional planting for ornamental purposes in gardens and landscapes, from which it escapes and establishes wild populations.[10] Seeds can remain viable in soil for decades, facilitating long-distance dispersal via machinery, hay, and human activities.[22]Management of C. arvensis in introduced ranges often involves integrated approaches, including biological controls introduced since the 1980s. Notable among these is the bindweed moth Tyta luctuosa, released in North America from Europe to target C. arvensis foliage and seeds, with establishments reported in several western U.S. states and Canada, though its overall impact remains variable.[23] Additional agents, such as the gall miteAceria malherbae, have been deployed alongside chemical and mechanical methods to suppress populations.[24]
Ecology
Interactions with pollinators and herbivores
Convolvulus species are primarily pollinated by a range of insects, including bees and butterflies, which are attracted to the nectar rewards produced at the base of the corolla tube. Flowers of Convolvulus arvensis, for example, open for only one day and are visited by halictid bees, honeybees, bumblebees, and various butterflies and moths, facilitating cross-pollination in natural settings.[10] These generalist pollinators benefit from the floral structure, where nectar guides, including potential ultraviolet patterns common in the Convolvulaceae family, direct foragingbehavior.[25]Herbivory on Convolvulus plants involves specialized insects, notably serving as a larval host for Lepidoptera species such as the convolvulus hawkmoth (Agrius convolvuli), whose caterpillars feed on the foliage.[26] This moth's larvae can defoliate plants significantly in native Mediterranean habitats, representing a key biotic interaction.[27] To counter generalist herbivores, Convolvulus species produce alkaloids, including those derived from fungal endophytes, which act as feeding deterrents and protect seeds and tissues from bruchid beetles and other pests.[28] These chemical defenses, such as shankhapushpine, enhance resistance without broadly inhibiting specialized herbivores like Agrius convolvuli.[2]Beyond pollinators and herbivores, Convolvulus forms mutualistic associations with arbuscular mycorrhizal fungi (AMF), which colonize roots to improve nutrient uptake, particularly phosphorus, in nutrient-poor soils common to their habitats.[29] Species like Convolvulus arvensis exhibit 21-70% AMF colonization, aiding establishment in arid or disturbed environments by enhancing water and mineral acquisition in exchange for plant carbohydrates.[30] These symbioses are widespread in the Convolvulaceae, supporting plant growth without reliance on high soil fertility.[31]Seed dispersal in Convolvulus varies by species but generally lacks specialized mechanisms, with lightweight seeds primarily dispersed short distances by gravity or wind (anemochory).[32] In Mediterranean species like Convolvulus arvensis, longer-distance dispersal occurs via wind, water, or attachment to animals and birds, allowing colonization of new areas while maintaining proximity to parent plants for most seeds.[33] This passive strategy aligns with the genus's adaptation to open, windy habitats, though myrmecochory is not documented.[34]
Role as a weed and pest
Convolvulus arvensis, commonly known as field bindweed, poses significant economic challenges in agriculture due to its competitive nature and twining growth habit, which allows it to smother crops and reduce yields substantially. In cereal fields and vineyards, infestations can lead to yield reductions of 20-80%, with specific studies reporting losses of 33-64% in wheat and 17-19% in sunflowers when densities exceed 14 plants per square meter. This competition for water, nutrients, and light, combined with the plant's ability to form dense mats, exacerbates the issue in perennial systems like orchards and vegetable production.[35][36]As a pest, C. arvensis serves as a reservoir for plant pathogens, including cucumber mosaic virus (CMV), which it transmits to nearby crops, further compounding agricultural losses. Control efforts are complicated by the weed's extensive regenerative root system, capable of producing new shoots from fragments as small as 5 cm, making mechanical and chemical methods often ineffective without repeated applications over multiple years.[37][38]Environmentally, C. arvensis outcompetes native flora in grasslands and disturbed habitats, leading to decreased biodiversity and altered ecosystem dynamics in invaded areas. By dominating resources, it reduces cover of native grasses and forbs, impacting forage quality for wildlife and overall plant community structure.[39][10]The global extent of C. arvensis as a weed is vast, infesting millions of hectares in North America, particularly in the Great Plains agricultural regions, where it has become widespread since its introduction from Eurasia. Additionally, populations have shown developing resistance to certain herbicides, such as Group 22 compounds, reported since the early 2010s, complicating long-term management strategies.[40][41]
Human uses
Ornamental cultivation
Several species of Convolvulus are valued in ornamental gardening for their attractive foliage, funnel-shaped flowers, and versatile growth habits, particularly in dry, sunny landscapes. Convolvulus cneorum, known as silverbush, is a popular evergreenshrub reaching up to 60 cm tall and wide, featuring narrowly oblong silvery leaves and white flowers tinged pink in bud from late spring to summer.[42] This species excels in rock gardens and borders, where its compact, mounding form provides year-round interest and softens hardscape elements like retaining walls.[43] Similarly, Convolvulus sabatius, or blue rock bindweed, serves as an effective groundcover in Mediterranean-style landscapes, forming trailing mats up to 20 cm high with small oval leaves and purplish-blue flowers throughout summer.[44] Its sprawling habit makes it ideal for cascading over edges or filling gaps in xeriscapes, enhancing drought-tolerant designs.[45]Cultivation of these ornamentals requires full sun and well-drained, gritty soil to prevent root rot, with C. cneorum thriving in poor, alkaline conditions once established.[46] Both species are hardy in USDA zones 8-10, tolerating light frost but needing protection in colder areas through mulching or container growth.[47]Propagation is straightforward via seeds sown in spring or softwood cuttings taken in summer, allowing gardeners to easily expand plantings.[44] They demand minimal maintenance, with occasional pruning to maintain shape and encourage blooming, and are drought-tolerant after the first year, making them suitable for low-water gardens.[48]In modern horticulture, hybrids like C. sabatius 'Two Moons' offer enhanced ornamental appeal with a mix of white and lilac flowers on deep green foliage, blooming prolifically from spring to autumn.[49] This cultivar performs well in containers or over embankments, providing colorful, low-maintenance coverage in contemporary landscapes.[50]While generally non-invasive, some Convolvuluscultivars can spread vigorously if not contained, necessitating use in pots, raised beds, or bordered areas to prevent unwanted encroachment in gardens.[45]
Medicinal and other applications
Various species of Convolvulus have been employed in traditional medicine across Europe and Asia, particularly for their diuretic and laxative properties. In traditional European phytotherapy, decoctions of the roots of Convolvulus arvensis (field bindweed) are used as cholagogues, diuretics, laxatives, and purgatives to promote bile flow and alleviate constipation.[51] Similarly, in Asian folk medicine, including practices in Turkey and Pakistan, C. arvensis decoctions treat cough, flu, jaundice, skin diseases, and urinary complaints, with leaves applied as diuretics and laxatives.[52] In Ayurvedic medicine, Convolvulus pluricaulis (known as Shankhpushpi) serves as a nervinetonic for cognitive enhancement, improving memory, reducing mental tension, and treating anxiety, epilepsy, hallucinations, chronic cough, sleeplessness, and hypertension; preclinical studies in animal models demonstrate its efficacy in promoting calm and learning at doses of 100-200 mg/kg of root extract, while limited human studies suggest benefits at lower doses such as 500 mg twice daily, though more research is needed.[53][54]Active compounds in Convolvulus species include tropane alkaloids such as convolvine and convolamine, which exhibit anticholinergic properties by blocking muscarinic receptors (M2 and M4), contributing to their pharmacological effects.[55] Modern pharmacological research has identified additional bioactive constituents, including flavonoids like kaempferol and steroids such as β-sitosterol, supporting anti-inflammatory, antioxidant, antibacterial, and neuroprotective activities; for instance, ethanolic extracts of C. arvensis show strong free radical scavenging (ABTS: 1.62 mmol TE/g) and inhibit lipid oxidation in food models.[56][51]Beyond medicinal applications, stems of Convolvulus arvensis have been historically used for cordage production, yielding flexible and strong fibers suitable for pack ropes and twine, though not durable for long-term use.[57] In arid regions, C. arvensis serves as fodder for cattle, sheep, and goats, providing nutritional value in low-forage environments despite its widespread invasiveness.[58]However, Convolvulus species pose toxicity risks, particularly to livestock, due to their tropane alkaloids; documented poisoning cases in cattle, horses, swine, and sheep have occurred since the 1800s, causing symptoms like photosensitization, digestive upset, and neurological effects, with field bindweed implicated in North American incidents leading to economic losses.[59][60]