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Convolvulaceae

Convolvulaceae, commonly known as the , is a of flowering in the order that includes approximately 1,650 classified into about 60 genera. These are predominantly twining vines and lianas, though they also encompass erect herbs, shrubs, small trees, and leafless parasitic such as those in the genus . The is characterized by alternate, simple leaves that are often cordate or arrow-shaped, funnel- or tubular-shaped flowers with a sympetalous typically in , , , or blue, and fruits that are usually capsules or berries containing one to four seeds. Flowers are actinomorphic and five-merous, with five sepals, five stamens, and a superior , often featuring a milky latex in stems and leaves. Distributed primarily in tropical and subtropical regions but extending into temperate zones worldwide, Convolvulaceae species thrive in diverse habitats from moist forests and savannas to disturbed areas and aquatic environments. The largest genus, , accounts for about 600 species and includes economically significant plants like the sweet potato (I. batatas), a major global crop for its edible tubers, as well as water spinach (I. aquatica), a leafy vegetable widely cultivated in . Other notable genera include (bindweeds), known for their weedy habits in agricultural fields, and , which features large, showy flowers. Many species are ornamental, prized for their vibrant, funnel-shaped blooms that open in the morning, while some, like certain Ipomoea species, contain alkaloids with hallucinogenic properties used in traditional contexts. The family's ecological role is significant, with species serving as pollinator attractants for , , and bats, and contributing to through their climbing habits. However, several members are invasive weeds, such as field bindweed (), which can reduce crop yields in temperate . Taxonomically, Convolvulaceae has been revised in recent decades, incorporating former segregate families like Cuscutaceae based on , and it continues to be studied for its diverse breeding systems and evolutionary adaptations.

Taxonomy

Classification

Convolvulaceae is a family of flowering plants within the angiosperms, specifically placed in the core , , lamiids , and the order according to the APG IV classification system. This placement reflects molecular phylogenetic evidence supporting its close relationship to and Montiniaceae within . The family comprises approximately 60 genera and 1,977 species, highlighting its significant diversity within the order. The family was first established by in 1789, with Convolvulus L. designated as the . Historical classifications evolved from early morphological systems, such as those by Hallier f. in 1898, which divided the family based on pollen characteristics, to modern phylogenetic approaches incorporating DNA sequence data. The APG IV update in 2016 reaffirmed the family's and its position in without recognizing subfamilies, emphasizing tribal-level organization instead. Convolvulaceae is currently divided into 12 tribes based on phylogenetic analyses: Aniseieae, Cardiochlamyeae, Convolvuleae, Cresseae, Cuscuteae, Dichondreae, Erycibeae, Humbertieae, Ipomoeae, Maripeae, Poraneae, and taxa . This tribal framework, largely derived from Stefanović et al.'s study using and markers, has seen refinements, such as the dissolution of the non-monophyletic Merremieae in 2017, with its genera redistributed among existing tribes like Ipomoeae. These tribes encompass a range of habits from herbaceous vines to woody lianas, unified by shared reproductive and traits.

Etymology

The name Convolvulaceae is derived from the type genus , which originates from the Latin verb convolvere, meaning "to roll together" or "to entwine," a reference to the characteristic twining stems of many plants in this family. This family name was formally established by the French botanist in his seminal work Genera Plantarum, published in 1789. A common vernacular name for numerous species within Convolvulaceae, especially those in the genus Ipomoea, is "morning glory," so called because their funnel-shaped flowers typically open in the early morning and wilt by afternoon, showcasing a diurnal blooming pattern.

Phylogeny

The Convolvulaceae family occupies a well-defined position within the order , forming a monophyletic that is to the family. This relationship has been consistently supported by molecular phylogenetic analyses since the early 1990s, including the (APG) classifications, with the most recent update in APG IV confirming the placement of both families within alongside Montiniaceae as a basal . Early studies utilizing genes such as rbcL and ndhF established the monophyly of Convolvulaceae and its close affinity to , resolving long-standing uncertainties about its ordinal placement. Key insights into intrafamilial relationships derive from cladistic analyses integrating molecular and morphological data, notably the comprehensive study by Stefanović et al. (2003), which employed sequences from chloroplast loci (rbcL, ndhF, rpl16) and the nuclear waxy gene to reconstruct a robust phylogeny. This work delineated five subfamilies and 12 tribes, including the monophyletic tribe Ipomoeeae, which encompasses diverse genera like Ipomoea and Argyreia. Subsequent molecular evidence from chloroplast DNA and nuclear genes has reinforced these tribal divisions, with studies on the Ipomoeeae clade highlighting its internal structure and biogeographic patterns through expanded sampling of plastid and low-copy nuclear markers. The evolutionary origins of Convolvulaceae trace back to the late in East , as evidenced by leaves of an early -like taxon from dated to approximately 56 million years ago, predating previous North American records and aligning with molecular divergence estimates for the family. Diversification likely accelerated during the , coinciding with continental fragmentation and the of key adaptations such as the twining for exploitation and holoparasitism in the genus , which represents a derived lineage within the family based on phylogenetic reconstructions. These developments underscore the family's across tropical and temperate regions.

Characteristics

Morphology

Members of the Convolvulaceae family exhibit a diverse range of growth habits, predominantly as herbaceous twining vines that climb using their stems, though they also include prostrate herbs, subshrubs, shrubs, and rarely trees such as Humbertia madagascariensis. The stems are typically slender, herbaceous, and either winding or prostrate, often becoming woody with age in lianas up to 35 cm in diameter; they twine clockwise without stipules or tendrils and may contain milky sap or articulated laticifers. Leaves are simple and alternate, frequently cordate or ovate with pinnate venation, though shapes vary from lanceolate to trilobed and may be coriaceous in woody species; in parasitic genera like Cuscuta, leaves are reduced to minute scales. Flowers are actinomorphic and bisexual, featuring a sympetalous that is funnel-shaped or campanulate, formed from five fused petals often with colorful mid-petaline bands in , , , or purple. The consists of five free or basally fused sepals, while the androecium includes five epipetalous stamens that are equal or unequal in length and inserted at the base or throat. The features a superior, bicarpellate (rarely tricarpellate) with two locules, each containing two ovules. Inflorescences are typically axillary cymes that may be simple, compound, or rarely racemose, though flowers can also occur solitary on long peduncles. Fruits are diverse, most commonly loculicidal or septicidal capsules that are globose or ovoid and contain 1–4 per locule, though some are baccate berries or indehiscent nuts; are often pubescent or hairy and may be winged for dispersal. Parasitic species, particularly in the genus , deviate significantly by lacking , featuring yellow-orange stems with haustoria for attachment, reduced or absent leaves, and minute flowers and fruits. Anatomically, non-parasitic members often possess intraxylary , successive cambia producing anomalous , and extrafloral nectaries.

Reproduction

The Convolvulaceae family primarily reproduces sexually through perfect, bisexual flowers that are actinomorphic and typically 5-merous, featuring both stamens and pistils within the same floral structure. These flowers are often self-compatible, enabling autogamous pollination in many species, though the degree of reliance on external pollinators varies across genera such as . Pollen grains in Convolvulaceae are characteristically trinucleate at dispersal, a feature linked to their gametophytic development and compatibility systems that promote efficient fertilization. Following , seed development involves formation, which provides nourishment to the and is often starchy or oily in composition, supporting the large, straight or curved embryos typical of the family. In species like those in the genus , seeds exhibit physical mechanisms, primarily due to impermeable hard seed coats that prevent water uptake and until occurs, enhancing long-term viability in the . This is widespread in the subfamily Convolvuloideae and contributes to the persistence of species in diverse environments. Asexual reproduction occurs in several Convolvulaceae species through vegetative propagation, bypassing sexual processes to facilitate rapid clonal spread. For instance, the sweet potato (Ipomoea batatas) is commonly propagated via its tuberous storage roots, which develop adventitious buds that sprout into new plants, a method central to its . Similarly, species like field bindweed (Convolvulus arvensis) employ extensive s for underground vegetative growth, allowing the formation of new shoots and roots from fragmented rhizome segments, which bolsters invasiveness in disturbed habitats. Fruit and seed dispersal in Convolvulaceae is adapted to various mechanisms, with most species producing dry, dehiscent capsules that split open to release seeds. These capsules often exhibit explosive dehiscence, propelling seeds ballistically over short distances to reduce competition and promote establishment, as observed in genera like Ipomoea. In contrast, some taxa, such as Argyreia, develop fleshy berries that attract animals for endozoochorous dispersal, where seeds pass through digestive tracts unharmed, aiding long-distance spread.

Distribution and Ecology

Geographic Distribution

The Convolvulaceae family displays a predominantly distribution, encompassing tropical, subtropical, and extending into temperate regions worldwide, with the greatest in the and . In the , approximately 500 species occur, with notable concentrations in hosting over 300 species, including diverse assemblages that exemplify regional hotspots around 20–25° N . Africa features high diversity in its tropical zones, with about 115 species documented in southern regions such as the and , where dominates as the most speciose genus across the continent. supports roughly 300 species, distributed across tropical and subtropical areas, as seen in with 129 species and with 119. In contrast, and harbor fewer species, with recording about 155 (134 native) primarily in warmer coastal and inland areas, and limited to around 20–30 native taxa mainly in Mediterranean climates. Endemism is pronounced in , home to the monotypic woody genus Humbertia, and on oceanic islands, where isolated evolution has produced numerous endemic species. The family's global spread likely originated through long-distance dispersal, including buoyant seeds transported by ocean currents, enabling colonization over millions of years, with some later introduced anthropogenically as weeds.

Habitat and Adaptations

Members of the Convolvulaceae family inhabit a broad spectrum of environments, including tropical and subtropical forests, open grasslands, arid deserts, and wetlands, reflecting their versatility across moisture gradients and soil types. Many prefer disturbed habitats such as roadsides, agricultural fields, and riverbanks, where they can establish quickly in nutrient-poor or compacted soils. For example, , a semi-aquatic , thrives in freshwater wetlands, canals, ditches, and shallow ponds, growing as a floating or trailing with hollow stems that root at nodes to exploit shallow bodies. A prominent in the family is the twining or climbing habit, which enables many herbaceous to ascend supporting and compete effectively for in shaded or dense understories. This allows like to physically overtop competitors, reducing shading stress and enhancing . is another key trait, facilitated by deep, extensive root systems that access subsurface moisture; for instance, field bindweed persists in dry, sandy, or moderately moist soils during extended arid periods by storing carbohydrates in its perennial roots. The genus exemplifies extreme specialization within Convolvulaceae through obligate holoparasitism, where plants lack roots and , instead forming haustoria—specialized appendages that penetrate host tissues to extract water, minerals, and organic nutrients via direct vascular connections. These haustoria establish symplastic continuity with the host's and , supporting the parasite's leafless, twining habit across various habitats. Many Convolvulaceae respond to environmental cues with diurnal flowering patterns, where corollas open at dawn in response to light signals, aligning with peak activity and minimizing nocturnal exposure. Coastal species, such as , exhibit salt tolerance through osmotic adjustments and ion compartmentalization, enabling survival in saline dune and beach habitats subjected to spray and inundation.

Ecological Interactions

Members of the Convolvulaceae family exhibit diverse strategies, primarily relying on animal pollinators such as bees, s, and hummingbirds, though some species are wind-pollinated. Day-opening flowers, characteristic of many genera like , attract diurnal pollinators including bees and hummingbirds through rewards and colorful corollas, facilitating cross- in tropical and subtropical environments. Nocturnal species, such as certain Calonyction vines, are adapted for moth , with white flowers emitting scents to draw in sphingid moths during evening hours. In rare cases, bat pollination has been documented in genera like Merremia, where large, durable flowers produce copious to support these mammalian visitors. Herbivory poses a significant threat to Convolvulaceae, prompting the evolution of chemical defenses centered on production, particularly pyrrolizidine (PAs) and , which deter generalist by disrupting their . These , biosynthesized via pathways like homospermidine synthase, accumulate in leaves and seeds, reducing and to and mammals. A key mutualistic interaction enhances this defense: many species form with clavicipitaceous fungi, such as those in the Periglandula, which colonize seeds and vegetative tissues to produce vertically transmitted to . This fungal not only boosts levels for resistance but also illustrates a co-evolutionary where the fungi gain nutrients from the host plant. Within the family, the genus represents a parasitic extreme, functioning as holoparasites that lack and roots, instead attaching via haustoria to extract water, nutrients, and carbohydrates from host plants, including like and . These twining parasites can severely impact agriculture by reducing crop yields—up to 20% in infested fields—and increasing management costs through their rapid spread and broad host range exceeding 150 species. 's alters host physiology, suppressing growth and while promoting its own proliferation, often leading to weakened or dead hosts in unmanaged systems. Beyond , Convolvulaceae engage in mutualisms that support stability, with secretion providing essential rewards to , thereby enhancing plant reproductive success and pollinator foraging efficiency. Vining growth habits in genera like and contribute to , particularly in coastal dunes and riparian zones, where sprawling roots and stems bind sandy substrates, preventing and aiding habitat restoration. For instance, , known as railroad vine, effectively anchors beach sands against wave action and storm surges, underscoring the family's role in geomorphic processes.

Diversity

Genera

The Convolvulaceae family encompasses approximately 57 genera, contributing to its overall diversity of about 1,955 worldwide. These genera exhibit a range of growth forms, from herbaceous vines and shrubs to woody climbers and parasites, reflecting the family's adaptability across tropical and temperate regions. The classification into tribes highlights phylogenetic relationships, with most genera aligned within monophyletic groups based on molecular and morphological evidence. The largest genus, , comprises 500–650 species of primarily twining vines and herbs, accounting for a significant portion of the family's and distributed pantropically. It dominates the tribe Ipomoeeae, which includes over 800 species and represents nearly half of Convolvulaceae's total diversity, underscoring the tribe's central role in the family's . Other prominent genera in this tribe and beyond include Convolvulus, with around 250 species of perennial bindweeds known for their trailing or climbing habits in temperate zones; , encompassing about 200 species of obligate parasitic, leafless vines that lack and attach via haustoria to host plants; and , featuring approximately 25 species of hedge bindweeds characterized by large, funnel-shaped flowers and robust stems. These genera collectively illustrate the family's morphological variation, from free-living climbers to specialized parasites. Notable genera outside the core diversity hubs include Erycibe, a group of about 70 woody climbers or scandent shrubs native to , distinguished by their lenticellate stems and often reddish tomentose indumentum. In contrast, Humbertia stands out as a monotypic , represented solely by H. madagascariensis, an endemic in Madagascar's dry forests, notable for its primitive traits and basal position in the family's phylogeny. Endemic genera further enhance regional biodiversity, such as Astripomoea with 12 species restricted to , including the , where they occur as geoxylic suffrutices or perennial herbs adapted to arid environments. Together, these genera highlight Convolvulaceae's ecological versatility and biogeographic patterns.

Species Diversity

The Convolvulaceae family encompasses approximately 1,955 accepted species distributed across 57 genera, predominantly in tropical and subtropical regions worldwide. As of 2024 (per ), this reflects 295 new species described since 2000, with ongoing molecular studies suggesting further generic reclassifications. This diversity is unevenly distributed, with the largest concentrations in the Neotropics, where the genus has experienced a major evolutionary radiation, accounting for around 425 species in the , many endemic to biodiversity hotspots like the and Andean regions. The tropics also represent a key center of diversity, particularly for the Ipomoeeae tribe, which includes numerous climbing and herbaceous forms adapted to varied ecosystems. Speciation patterns within Convolvulaceae are notably high among vining species, which dominate the family and facilitate rapid adaptation and diversification in tropical forests and open habitats through climbing habits that enhance access to light and dispersal. The parasitic genus Cuscuta, comprising about 200 species, exemplifies specialized evolution, with many taxa exhibiting narrow host ranges that restrict their distribution to specific plant communities and contribute to localized endemism. Conservation challenges are acute for Convolvulaceae, driven primarily by habitat loss from agricultural expansion, urbanization, and deforestation, which fragment populations and reduce suitable environments for these often habitat-specific plants. A significant number of species have been assessed as threatened by the IUCN Red List, with examples including the critically endangered Convolvulus argyrothamnos in Europe, a shrubby endemic restricted to cliff habitats in Crete, Greece, where ongoing habitat degradation exacerbates its vulnerability.

Human Uses and Significance

Economic Importance

The Convolvulaceae family holds substantial economic value through its food crops, with the (Ipomoea batatas) serving as a primary example of a global staple . Annual worldwide production was approximately 86 million metric tons as of 2022, with over 95% occurring in developing countries, underscoring its role in and in tropical and subtropical regions. Ranked as the seventh most important crop globally, sweet potato contributes significantly to caloric intake and export revenues in major producers like and . Another key food crop is water spinach (), a semi-aquatic prized in Asian markets for its nutrient-rich shoots and leaves, which are harvested as a fast-growing leafy green. Its high yield potential and adaptability to conditions make it a profitable specialty in , supporting local economies through fresh sales and culinary applications. In , species such as the () are widely cultivated as ornamentals for their striking, trumpet-shaped flowers in shades of blue, purple, and white, enhancing gardens, trellises, and containers. These vines provide aesthetic value in and are commercially propagated for the nursery trade, contributing to the global industry. Conversely, certain Convolvulaceae species pose economic challenges as invasive weeds in ; field bindweed (), for instance, infests crops like and reduces yields by 50-60% in dense stands due to its aggressive rhizomatous and for resources. This perennial weed necessitates costly control measures, including herbicides and , amplifying production expenses for farmers in temperate regions. Beyond food and ornamentals, some Convolvulaceae species offer industrial potential through fiber extraction; for example, beach morning glory () yields natural cellulosic fibers suitable for textiles, ropes, and composite materials after chemical treatments to improve tensile strength. Additionally, tubers exhibit promise as a biofuel feedstock, with varieties yielding up to 82 grams of per kilogram of fresh root through processes, positioning them as a sustainable alternative in production. Recent breeding efforts focus on biofortified varieties to combat in and .

Medicinal and Cultural Uses

The roots of Ipomoea purga, commonly known as jalap root, have been utilized historically as a potent purgative in and colonial medicine, originating from trade routes where they were valued for inducing peristaltic movements to balance bodily humors. Resin glycosides extracted from these roots contribute to their effects, making them a key ingredient in traditional remedies for digestive disorders. In Ayurvedic medicine, Argyreia speciosa (known as Vriddhadaru) is employed as a rejuvenator for conditions involving inflammation, such as and , with root extracts demonstrating significant activity in animal models by inhibiting paw and formation. Psychoactive properties in the family stem from alkaloids, such as those in the seeds of Ipomoea tricolor (ololiuqui), which were used by in divinatory rituals to induce hallucinations, as documented in 16th-century accounts. These alkaloids hold pharmaceutical potential, serving as precursors for drugs treating migraines, , and endocrinological disorders, with biosynthetic pathways in Convolvulaceae enabling their accumulation in plant tissues. Culturally, morning glories symbolize rebirth in various folklores, representing renewal through their daily cycle of blooming at dawn and withering by dusk, evoking themes of life's transience and resurgence. Ornamentally, they feature prominently in festivals, such as Japan's Iriya Festival, an Edo-period tradition revived post-World War II, where over 1,000 varieties are displayed to honor summer aesthetics and deities like Kishimojin for protection and fertility. Historically, species like Calystegia soldanella have been incorporated into as an edible and therapeutic herb to alleviate rheumatic arthritis, , dropsy, and .

Toxicity and Risks

Members of the Convolvulaceae family produce several toxic compounds, including ergoline alkaloids such as found in species, which can induce hallucinations and upon ingestion. These alkaloids, structurally similar to , originate from symbiotic fungi in the plant tissues and pose risks to both humans and animals. Additionally, some genera contain tropane alkaloids like and pseudotropine, particularly in species, contributing to their poisonous effects. Livestock poisoning from (field bindweed) manifests as vomiting, , weight loss, , and digestive disturbances due to the alkaloids disrupting normal gastrointestinal function. In humans, ingestion of seeds, often for their psychoactive properties, leads to symptoms including , , disorientation, tremors, rapid heartbeat, and hallucinations, with effects varying by dose and individual sensitivity. Severe cases may require medical intervention for or cardiovascular complications. Invasive weeds within the family, such as , exacerbate toxicity risks by contaminating pastures and spreading alkaloids to grazing animals, potentially causing chronic health issues in herds. The genus (dodder), a holoparasitic member of Convolvulaceae, acts as a vector for plant pathogens including viruses, fungi, , and phytoplasmas, transmitting them between host plants and amplifying disease spread in agricultural settings. Management of these risks includes detoxification methods like applying fungicides to asarifolia, which eliminates alkaloids by targeting the associated fungal symbionts in leaf glands, thereby reducing plant toxicity. For psychoactive species, regulatory measures in various jurisdictions ban or restrict the sale and distribution of seeds containing high levels of , such as those from , to prevent abuse and associated health hazards, with LSA itself classified as a under Schedule III in the United States. Preventive strategies for livestock involve pasture management to control and prompt veterinary care for affected .

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