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Silene

Silene is a large of flowering in the family , consisting of approximately 850 species of annual to perennial herbs commonly known as campions or catchflies. These are typically erect, arising from a , , or , and are rarely dioecious, with simple, opposite leaves that are linear to oblanceolate and one-veined. Their flowers are generally bisexual, featuring five fused sepals forming a tube (4–38 mm long) and five petals (6–62 mm), along with three to five styles; the fruit is a capsule containing many gray to black seeds. Native to regions across , , , and , with a concentration in the Northern Hemisphere, Silene species have been introduced worldwide and often thrive in disturbed . Many exhibit sticky glandular hairs on stems and calyces, which give rise to the "catchfly" common name by trapping small , though this serves more as a defensive mechanism than for . Ecologically, the genus includes both native wildflowers that support pollinators and invasive weeds that can outcompete local in disturbed areas, such as roadsides and agricultural fields. Several species hold conservation significance, with some listed as rare or endangered due to habitat loss. The name Silene derives from the Greek mythological figure , possibly alluding to the swollen calyces of some resembling his bloated form, and the genus has been taxonomically challenging due to its morphological diversity and extensive hybridization.

Description

Morphology

Silene exhibit a range of habits, primarily as , , or herbs, rarely forming subshrubs or suffrutescent growths; they often appear decumbent at the base or cespitose. Stems are typically erect or ascending, simple or branched, terete or angular, and frequently glandular-pubescent or viscid, with heights varying from 5 cm in dwarf to over 1 m in tall . Root systems differ by life cycle: generally possess fibrous , while develop taproots that are slender or stout and deep, often with a branched ; some are stoloniferous or rhizomatous. Leaves are opposite, occasionally whorled, simple, and entire, connate proximally; basal leaves are petiolate and cauline leaves sessile, with blades that are 1–5-veined, linear to ovate or lanceolate, and 1–10 cm long, featuring acute to obtuse apices. Reproductive structures are borne in inflorescences that are terminal or axillary cymose panicles, often dichasial cymes, or solitary flowers, typically glandular-pubescent and viscid. The is tubular to campanulate or inflated, 10–30-veined, (4–)10–28(–40) mm long, 5-toothed, and often persistent in fruit. Flowers are typically bisexual, but unisexual in the dioecious and gynodioecious that occur in several sections of the , with 5 petals that are clawed, , , , or , and deeply bifid or dissected at the limb ; in dioecious , minor may occur in floral structures. The androecium consists of 10 dimorphic stamens arising with the petals, while the features an ovoid to globose that is 1- or 3–5-locular, with 3–5 glabrous styles 1.5–20 mm long. The fruit is a capsule, ovoid to globose, dehiscing by 3–5 valves into 6–10 teeth, often with a present carpophore; it contains (5–)15–100(–500+) reniform, tuberculate or papillate seeds that are reddish to black.

Reproduction

Silene species primarily reproduce sexually through seed production, though asexual reproduction occurs rarely in certain perennials via vegetative propagation from rhizomes. For instance, Silene nivea forms long rhizomes that allow clonal spread, enabling the plant to colonize new areas without seed dispersal. Similarly, Silene vulgaris and Silene chalcedonica produce rhizomes that facilitate vegetative reproduction alongside seed-based propagation. Many species exhibit self-incompatibility, promoting outcrossing and influencing seed set in hermaphroditic populations. Apomixis, or asexual seed formation, remains undocumented in the genus. Seed production in Silene involves the of dry capsules that dehisce at maturity to release numerous small , typically measuring 0.4–1.9 mm in length. These capsules, often with six teeth, open via hygroscopic movements triggered by , allowing to be shed gradually. Dispersal occurs primarily through , with falling near the parent plant, augmented by shaking the stems or capsules in species like . In some cases, limited animal-mediated dispersal may occur when adhere briefly to fur or are dislodged by passing animals. Germination in Silene varies by species and environmental conditions, with many exhibiting physiological broken by cold . For example, seeds of Silene elisabethae show improved with increasing durations of cold stratification, reaching up to 90%. Some species, such as Silene acaulis, show -dependent , where exposure to light post-stratification enhances establishment. Without stratification, percentages remain low, typically below 40% even under optimal temperatures. Life cycle variations among Silene species include annuals that complete their cycle in one , biennials that overwinter as rosettes before flowering, and perennials that persist for multiple years. Annuals like Silene antirrhina germinate, flower, and set seed rapidly in favorable conditions. Biennials and short-lived perennials, such as Silene latifolia, form basal rosettes in the first year, bolting and reproducing in the second. Perennials maintain rosettes or clumps across seasons, supporting repeated seed production. Fertility rates in Silene differ based on sexual systems, with hermaphroditic individuals generally achieving high seed set through self- or cross-pollination, often exceeding 80 seeds per capsule in optimal conditions. In dioecious species like and , seed set in females is typically reduced due to spatial separation of sexes, leading to pollen limitation over distances greater than a few meters. This reduction can lower overall fertility by 20–50% in isolated populations compared to hermaphroditic counterparts.

Taxonomy

History

The genus Silene was formally established by Carl Linnaeus in his seminal 1753 publication Species Plantarum, where he described 27 species based on morphological characteristics such as flower structure and calyx features. This initial description laid the foundation for the genus within the Caryophyllaceae family, drawing on earlier herbal traditions but introducing binomial nomenclature for clarity in botanical classification. Linnaeus's work emphasized the variability in seed capsules and petals, grouping species primarily from European and Mediterranean regions. Early 19th-century contributions expanded the significantly. In 1824, Daniel Otth, in de Candolle's Prodromus Systematis Naturalis Regni Vegetabilis, classified 217 Silene species into eight sections, relying on traits like type and arrangement to organize the growing diversity. The Melandrium, proposed by Elias Fries in 1828 for dioecious species previously under Lychnis and Silene, represented a key segregation effort, highlighting as a distinguishing factor; it was later merged back into Silene as classifications evolved. Paul Rohrbach's 1868 monograph Monographie der Gattung Silene provided the first comprehensive systematic treatment, detailing over 300 species and emphasizing patterns and capsule morphology, though it retained some artificial groupings. Charles Darwin contributed to early understandings of Silene biology in the 1860s, examining dimorphism and reproductive strategies in species like Silene (then classified under Lychnis), noting failures in self-fertilization and the adaptive value of derived from hermaphroditism. Pre-molecular era taxonomy faced challenges from artificial classifications based on geographic distribution or growth , leading to an of names—over 2,000 species described by 1900—many of which proved synonymous upon closer scrutiny. In the early , Silene emerged as a model for inheritance studies, with researchers like and others applying Mendelian principles to explore sex-linked traits and cytoplasmic effects in species such as S. latifolia. These efforts, building on Rohrbach's framework, underscored the genus's utility in investigating before molecular tools revolutionized phylogeny.

Etymology

The genus name Silene originates from the Latin feminine form of Silenus, woodland deity and foster father of (Bacchus), who was often portrayed as intoxicated and covered in or froth, an to the sticky, viscid exudates produced by the stems and calyces of many in the genus. An alternative etymology proposes derivation from the Greek sialon, meaning "" or "spittle," in reference to the glandular secretions that give some a foamy or sticky appearance. Common names for plants in the genus include "campion," derived from the Latin campio (champion), reflecting the showy, prominent flowers that were historically used in Europe to crown victors in sporting events or battles. Another widespread name is "catchfly," stemming from the viscid, glandular stems that can trap or "catch" small insects attempting to crawl over them. Within the genus, sectional names often draw from morphological traits using Greek or Latin roots; for example, the type section Silene is named directly after the genus, while sect. Conoimorpha refers to species characterized by cone-shaped or inflated calyces, combining Greek kōnos (cone) and morphē (form). Nomenclatural changes have included the merger of the former genus Lychnis (from Greek lychnos, meaning "lamp," alluding to flame-like flowers or the use of woolly leaves as lamp wicks) into Silene, driven by shared morphological and phylogenetic traits rather than etymological considerations, resulting in numerous species transfers such as Silene coronaria (formerly Lychnis coronaria).

Classification

Silene belongs to the family , within the subfamily Silenoideae, characterized by flowers with united petals and absence of stipules. The closest relatives to Silene within this subfamily include the genus Atocion, often treated as congeneric or closely allied based on shared morphological and molecular traits, and , which shares a broader tribal affinity in the Sileneae. These relationships have been clarified through phylogenetic analyses emphasizing nuclear and plastid DNA markers. Infrageneric classification of Silene recognizes three main subgenera—Behenantha (Otth.) Torr. & A. Gray, Lychnis (L.) Greuter, and Silene—supported by comprehensive molecular phylogenies spanning 2003 to 2020 that integrate multi-locus data such as ITS, trnL-F, and sequences. These subgenera encompass approximately 35 sections, with examples including sect. Silene (characterized by annual habits and inflated calyces) and sect. Psammophila (featuring psammophilous adaptations in sandy habitats), reflecting monophyletic groups defined by floral, fruit, and ecological traits corroborated by Bayesian and maximum likelihood analyses. This framework resolves many historical ambiguities in sectional boundaries, though ongoing refinements continue based on expanded sampling. The genus comprises about 908 accepted species according to the 2023 update of , though species delimitation remains challenging due to frequent hybridization events and , which blur morphological boundaries and complicate phylogenetic resolution in complexes like sect. Physolychnis. Hybrid zones often produce intermediate forms that evade clear taxonomic assignment, while allopolyploid origins—evident in taxa up to 10-ploid—further confound diploid-polyploid species clusters, necessitating integrative approaches combining and for accurate circumscription. Recent taxonomic revisions include the merger of the former genus Melandrium Röhl.—encompassing dioecious like S. latifolia—into during the 2010s, driven by phylogenetic evidence placing it within sect. Melandrium based on and markers that rejected its distinctness. Additionally, a 2020 study formally described sect. Arenosae as a new infrageneric unit, confirming nine annual primarily from the to southwestern Asia, distinguished by dehiscent capsules and psammophilous distributions, with delimitation supported by RAD-seq and morphological revisions.

Sexual Systems

The genus Silene displays remarkable diversity in sexual systems, reflecting its evolutionary lability and serving as a key feature in taxonomic and phylogenetic studies. Hermaphroditism predominates, occurring in approximately 58% of species, while and each account for about 14% and 13%, respectively, based on an extensive survey of 98 across subgenera Silene and Behenantha. These proportions highlight hermaphroditism as the ancestral state, with transitions to separate-sex systems occurring independently multiple times within the . Dioecy in Silene is primarily confined to specific lineages, such as section Melandrium (including model species like S. latifolia and S. dioica), where sex determination is governed by heteromorphic X and s. The harbors male-determining genes that suppress female function and promote male traits, leading to distinct ; for instance, males typically exhibit larger flowers and higher floral display compared to females, traits partially linked to . These genetic mechanisms facilitate the of from gynodioecious ancestors, with the showing degeneration over time due to suppressed recombination. Gynodioecy, characterized by co-occurring female and hermaphroditic individuals, is maintained through interactions between (CMS) factors in the mitochondria and nuclear restorer genes that counteract sterility. In species like S. nutans and S. vulgaris, multiple CMS mitotypes exist, with nuclear restorers exhibiting epistatic interactions that allow variable female frequencies across populations, often ranging from 10% to 50% depending on environmental and genetic contexts. This nuclear-cytoplasmic conflict underpins the stability of , enabling females to persist via fertility advantages despite pollen production limitations in hermaphrodites. Less common systems include , where females, males, and hermaphrodites coexist, as observed in certain populations of S. acaulis, potentially representing transitional states toward . Andromonoecy, featuring plants with both staminate and hermaphroditic flowers, is rare but documented in some Asian species, such as those in lineages, contributing to variation. Overall, evolutionary transitions among these systems are frequent and labile, with molecular phylogenies revealing at least three independent origins of from hermaphroditic or gynodioecious progenitors, driven by genetic conflicts and selection pressures.

Distribution and Habitat

Geographic Range

The genus Silene is primarily native to the , with its core range encompassing temperate and Mediterranean regions of , where the majority of its approximately 700–900 species are found. This area hosts the highest diversity, estimated at around 500 species, reflecting the genus's evolutionary origins in the mid-late within the . The distribution extends eastward to the mountains of , southward to and parts of , as well as westward to , where about 70 species occur, many of them polyploid endemics. Scattered populations also inhabit high-elevation habitats in tropical African mountains, underscoring the genus's preference for cooler, temperate-like conditions even in subtropical zones. Key centers of endemism for Silene include the , particularly in and , where nearly half of regional species are unique, including members of section Sclerocalycinae. The region and the , especially northwestern , represent additional hotspots, harboring high species richness and narrow endemics adapted to alpine and rocky terrains. These areas contribute significantly to the genus's overall variation, with biogeographic patterns dominated by Holarctic distributions and disjunct occurrences between and , likely facilitated by ancient migrations across the during glacial periods. Several Silene species have been introduced outside their native ranges, notably S. vulgaris and S. latifolia, which arrived in North America in the late 18th to early 19th centuries via ship ballast or contaminated seeds and have since become invasive in disturbed habitats across the continent. Similarly, both species were introduced to Australia in the 19th century as ornamental or fodder plants and now exhibit invasive tendencies in temperate grasslands and roadsides. A 2019 European study found significant phenotypic plasticity in S. vulgaris populations, enabling compensatory adaptations such as reduced flower number under warmer and drier conditions, suggesting potential resilience to climate change.

Habitat Preferences

Species in the genus Silene exhibit a wide range of habitat preferences, primarily favoring well-drained substrates such as sands, rocks, and chalk, with many showing a strong affinity for calcareous soils. For instance, Silene nutans thrives on limestone outcrops and steep grassy slopes, particularly in regions like the Peak District where such substrates predominate. Similarly, Silene vulgaris prefers moderately fertile, open, calcareous, sandy, or gravelly conditions with some humus content, reflecting the genus's general adaptation to neutral to basic pH environments. These preferences enable Silene species to colonize rocky slopes and open grasslands, where soil texture is often sandy and slightly alkaline. Climatically, Silene occupies temperate to alpine zones, with notable drought tolerance in Mediterranean habitats and cold hardiness in boreal and arctic regions. In Mediterranean climates, species like Silene gallica endure dry, non-saline conditions such as meadows and wastelands, supported by adaptations to hot, dry summers and mild, wet winters. Boreal representatives, including Silene dioica, persist in moister, cooler settings up to 500 m elevation, while arctic-alpine species such as Silene acaulis withstand temperatures down to -40°C in tundra environments. Altitude ranges broadly from sea level in coastal and lowland areas to 5000 m in high-elevation alpine meadows, as seen in Silene himalayensis across the Himalayas. Moisture levels vary from xeric in open, dry grasslands—where glandular hairs on species like Silene scouleri aid in protection—to mesic in riparian zones, exemplified by Silene dioica in rich meadows and streamside habitats. Many Silene species act as pioneers in disturbed habitats, readily establishing on roadsides, waste grounds, and other sites with loose, well-aerated soils. This is evident in Silene vulgaris, which frequently appears in such areas alongside natural settings like alluvial woodlands for Silene nivea. While global distribution spans the with hotspots around the Mediterranean, these abiotic preferences underpin the genus's versatility across diverse landscapes.

Ecology

Pollination

The genus Silene displays a range of strategies, with many relying on vectors that exploit floral scents, , and rewards. Nocturnal moths, particularly from the genus Hadena (), serve as primary pollinators for numerous Silene , often functioning as nursery pollinators that both transfer and oviposit in flowers or developing fruits. These moths are attracted to the nocturnal fragrance emitted by flowers, which peaks in the evening to align with their activity patterns. Diurnal or those with mixed regimes are frequently visited by bees (e.g., bumblebees like Bombus fervidus in S. spaldingii) and flies (including syrphids and tachinids), which collect and . Floral traits in Silene are adapted to facilitate effective while deterring inefficient visitors. Flowers are typically -producing with hexose-rich rewards that vary by time of day, sex phase, and age, though is abundant and serves as a key resource for many visitors. The often forms a tube that accommodates proboscides or mouthparts, and many species emit scents from osmophores to lure pollinators. Viscid glandular stems and calyces trap small crawling , reducing or theft by non-pollinating arthropods and favoring larger flying visitors. Selfing rates in hermaphroditic Silene vary widely, often reaching moderate to high levels (e.g., up to 41% in S. acutifolia), facilitated by the absence of strong and opportunities for or facilitated . Despite this, is promoted by temporal protandry, where anthers dehisce before stigmas become receptive, minimizing within flowers. This dichogamy, combined with spatial separation in some dioecious taxa, enhances flow between plants. Pollinator specialization differs across Silene sections. In section Silene, species tend toward pollination, attracting a broad array of diurnal and nocturnal without strict syndromes. For instance, S. otites features long-tubed flowers suited to hawkmoths and other crepuscular/nocturnal , though it also receives visits from diurnal flies and bees, indicating partial .

Biotic Interactions

Silene species engage in a range of biotic interactions beyond pollination, including antagonistic relationships with herbivores and pathogens as well as mutualistic associations that influence plant fitness and distribution. Specialized herbivores, such as the noctuid moth Hadena bicruris, target reproductive structures by ovipositing eggs directly on the ovaries of host plants like Silene latifolia. Upon hatching, the larvae consume developing seeds, effectively manipulating seed set and reducing host reproductive output, though females preferentially avoid oviposition on pathogen-infected flowers to ensure larval survival. Other folivores, including aphids such as Brachycaudus lychnidis, feed on leaves and stems, inducing sex-specific changes in floral volatiles that alter interactions with other insects. These herbivory pressures can vary with plant population density, with higher damage observed in larger, less isolated stands. Pathogenic interactions further constrain Silene populations, particularly through fungal and viral agents that impair growth and reproduction. Rust fungi in the genus Uromyces, such as U. silenes-chloraefoliae on Silene chlorifolia, infect leaves and stems, producing pustules that disrupt and vascular function, though direct links to sterility are more commonly associated with related pathogens like the anther-smut fungus Microbotryum silenes-dioicae, which sterilizes infected by replacing with spores. Viral infections, including potexviruses like Silene virus X (SVX) on Silene pratensis and vein-clearing virus on Silene flos-cuculi, cause mosaic symptoms and stunting in wild populations, with prevalence potentially elevated in dense stands due to increased transmission efficiency. These pathogens exploit high host densities, contributing to episodic outbreaks that reduce population viability. Mutualistic relationships provide compensatory benefits, notably through mycorrhizal associations that enhance nutrient uptake in challenging environments. Many Silene species form arbuscular mycorrhizal symbioses, albeit sometimes weakly, with fungi improving acquisition in nutrient-poor soils and boosting overall growth, though this can inadvertently facilitate infection rates. In glandular species, viscid trichomes on stems and leaves act as passive traps for small , potentially benefiting by immobilizing herbivores and providing resources for predatory arthropods, thereby enhancing indirect protection. Invasive dynamics exemplify how altered biotic interactions drive ecological impacts, as seen with in . Introduced populations outcompete native flora partly by escaping specialist enemies from their native range while co-opting novel mutualists and antagonists, such as the native seed predator Hadena ectypa, which pollinates but also consumes seeds, potentially favoring invasive genotypes through relaxed predation pressure. Recent studies highlight how these shifted interactions, including changes in volatile emissions under herbivory, contribute to invasion success by disrupting native community structures. Such patterns underscore the role of biotic novelty in facilitating range expansion.

Species Diversity

Number and Variation

The genus Silene encompasses approximately 908 accepted species worldwide, according to the 2025 assessment by (POWO), though estimates vary between 700 and over 1,000 due to the prevalence of cryptic that are difficult to distinguish morphologically and genetically. This variation stems from ongoing taxonomic revisions, where molecular data often reveal hidden diversity within apparent morphospecies. Between 2020 and 2025, phylogenetic and ecological studies have led to the description of at least 20 new , primarily through species delimitation in understudied regions, including Silene isabellae from in 2023 and Silene vanchingshanensis from in 2022. Morphological diversity in Silene is extensive, with plants ranging from prostrate forms as short as 5 cm, such as cushion-like S. acaulis, to tall biennials reaching 1.5 m, like S. ovata. Key identification features include variation in shape and , where teeth are often heteromorphic, with three differing from the other two, aiding in sectional classification. At the genetic level, the genus exhibits high , with numbers ranging from diploid 2n=24 (the most common) to higher levels up to 2n=120 in decaploid forms, contributing to . Hybridization is frequent in contact zones between closely related species, such as S. latifolia and S. dioica, leading to and further complicating species boundaries. Species richness is highest in Eurasia, with over 600 species concentrated in temperate and montane habitats, compared to about 150 in the Americas (including roughly 70 in ) and around 100 in . Taxonomic challenges persist due to undersampling in Asian biodiversity hotspots, where remote montane areas remain poorly explored, potentially harboring additional cryptic taxa. Additionally, is accelerating in mountain ranges by promoting and adaptation in fragmented habitats, as evidenced by shifting distributions and novel genetic variants in alpine Silene populations.

Notable Species

Silene latifolia, commonly known as white campion, is a dioecious species native to but invasive in , where it has spread widely in disturbed habitats such as roadsides and fields. It serves as a key in studies of due to its , which has been extensively researched for insights into genetic mechanisms of sex differentiation. Silene vulgaris, also called bladder campion, is a widespread weed originating from and Asia, now naturalized globally, including in , where it thrives in agricultural fields and waste areas. The plant is notable for its edible young leaves used in traditional cuisines, particularly in Mediterranean regions, and it possesses a diploid number of 2n=24, facilitating genetic studies on hybridization and . Silene acaulis, known as moss campion, is a low-growing perennial adapted to and environments across the , forming dense cushion-like mats that protect against harsh winds and cold. This species acts as an indicator of , with populations showing shifts in flowering times and distribution in response to warming temperatures in polar regions. Silene undulata, referred to as the African dream root, is indigenous to the of and holds cultural significance among the , who use its tuberous roots in rituals to induce vivid, lucid dreams for divination and spiritual purposes. The plant's roots contain believed to influence sleep patterns, though scientific validation of these effects remains limited. Silene alexandri is an endangered endemic found only on the island of in , growing to heights of 30-60 cm in montane wet forests. It faces severe threats from non-native feral goats that degrade its habitat through browsing and , contributing to its status.

Human Uses

Ornamental Cultivation

Silene are valued in ornamental for their vibrant flowers and versatile growth habits, making them suitable for borders, rock gardens, and containers. Popular cultivars include Silene armeria, commonly known as none-so-pretty or sweet William catchfly, which forms compact mounds ideal for garden borders and cottage-style plantings with its clusters of bright to magenta blooms on 16- to 20-inch stems. Similarly, Silene pendula, or nodding catchfly, is favored for its trailing habit, cascading gracefully in rock gardens, hanging baskets, or as a groundcover with nodding flowers. Propagation of Silene is straightforward, primarily through sown directly in the garden in after the last frost or in fall for natural , or started indoors 6 to 8 weeks prior under cool conditions of 60-70°F to ensure even . species can also be divided in early or fall, with success depending on well-drained to minimize rot. Optimal growing conditions for most ornamental Silene include full sun to partial shade and moderately fertile, well-drained soils with neutral to slightly acidic , as poor can lead to root issues. They are generally hardy in USDA zones 3 to 9, tolerating drought once established but benefiting from consistent moisture during dry spells. Several Silene species have been cultivated in gardens since at least the , with modern breeding producing hybrids like 'Sibella Carmine' for enhanced color range in and . Challenges in include susceptibility to slugs and , particularly in damp conditions, which can damage foliage and flowers; cultural controls like reducing and evening watering help mitigate this. Recent trends toward native plant , prominent in 2023, have increased interest in indigenous Silene species for sustainable landscapes, promoting in ornamental settings.

Culinary and Medicinal Applications

Silene vulgaris, commonly known as bladder campion, has been traditionally utilized in , where its young leaves are boiled and served as a substitute, often incorporated into Greek horta—a dish of wild greens seasoned with and . This preparation method preserves the plant's nutritional profile, including its high content, which contributes to its antiscorbutic properties and supports immune health when consumed regularly. Recent analyses of wild edible greens confirm that S. vulgaris retains significant ascorbic acid levels even after cooking, making it a valuable source of this essential nutrient in traditional diets. In medicinal contexts, Silene undulata roots are employed by the of as ubulawu, a traditional preparation consumed to induce vivid and lucid dreams for spiritual communication with ancestors, attributed to the presence of triterpenoid that act as inhibitors. Various Silene , including those used in Asian folk medicine, serve as emollients applied topically to soothe wounds, cleanse , and promote healing through baths or direct washes. For instance, in Iranian traditional practices, S. conoidea seeds are utilized as a to alleviate urinary issues and reduce fluid retention. Modern pharmacological has validated some of these applications, suggesting potential for treating inflammatory conditions. Preparations typically involve infusions for internal use, such as brewing into teas for dream induction, or poultices made from crushed leaves and stems applied externally for care. However, caution is advised due to the hemolytic properties of in many Silene species, which can cause gastrointestinal upset or toxicity if consumed in excess, particularly in raw forms. Ethnobotanical surveys conducted in 2024 have expanded documentation of Silene uses, emphasizing its role in sustainable foraging practices amid urbanization, with S. vulgaris noted as a commonly gathered species in peri-urban areas for both nutritional and therapeutic purposes.

Evolutionary History

Fossil Record

The earliest known fossils attributable to the genus Silene consist of seeds of S. microsperma Mai recovered from the Oberleichtersbach Formation in the Rhön Mountains of central Germany, dating to the Chattian stage of the Late Oligocene (approximately 26–23 million years ago). These small seeds, measuring about 0.5–0.7 mm in length, exhibit a reniform shape and finely reticulate surface ornamentation closely resembling those of extant small-seeded Silene species, suggesting morphological conservatism within the genus over this period. The fossils were preserved in blue-black clayey sediments within a doline (sinkhole) deposit, indicative of an open, mesophytic forest environment. Additional fossil evidence includes dispersed pollen grains of , to which Silene belongs, from sediments (23–5 million years ago) across and . In , such pollen first appears in the Early , often as psilate or faintly ornamented tricolpate grains typical of the family, recorded in various lacustrine and fluvial deposits. In , comparable pollen has been identified in the Middle Shanwang flora of Shandong Province, , where it represents a minor component of diverse palynofloras from subtropical mixed forests. The fossil record of Silene and related is notably sparse, primarily due to the predominantly herbaceous growth of the group, which favors rapid over mineralization or preservation in most depositional environments. The fossil record of Silene and related remains sparse, with only a handful of formally described taxa, mostly as seeds, fruits, or , with macrofossils being particularly rare outside of exceptional lagerstätten.

Phylogenetic Insights

Phylogenetic analyses of the genus Silene have revealed a complex evolutionary history, with major clades organized into three subgenera: S. subg. Behenantha (encompassing 18 sections), S. subg. Lychnis (divided into three main clades), and S. subg. Silene (including 11 sections). A comprehensive multi-locus phylogeny, based on nuclear ribosomal DNA ITS sequences from 415 and chloroplast rps16 from 397 , resolved relationships for approximately 80% of the 33 recognized sections, highlighting monophyletic groups that differ from traditional morphology-based classifications. However, basal nodes among subgenera remain poorly supported, with S. subg. Lychnis weakly positioned as to the others and limited resolution within S. subg. Behenantha. The exhibits a pattern of early primarily in Eurasian lineages, from which multiple independent dispersals led to American clades, including polyploid groups in . For instance, North American taxa in S. sect. Physolychnis s.l. derive from Eurasian ancestors via at least three separate migrations, with poor resolution attributed to rapid and in and nuclear markers. Hybridization has driven across these lineages, as demonstrated by successful between species diverged for about 6 million years, contributing to and in regions like the Mediterranean and . A pivotal evolutionary event is the origin of in S. sect. Melandrium (now often treated within S. subg. Behenantha), estimated at approximately 11 million years ago based on estimates from resequencing of S. latifolia. This shift from hermaphroditism involved the of young , with the accumulating mutations over this timeframe. Recent 2025 genomic sequencing of S. latifolia has further elucidated the structure and of its giant , supporting the estimated origin of around 11 million years ago. further illuminate these dynamics, as Silene harbors the largest known mitochondrial genomes, exceeding 11 Mb in like S. conica, characterized by multichromosomal structures, high (up to 50 times the angiosperm average), and extensive recombination. (CMS), linked to in such as S. vulgaris, is associated with accelerated substitution rates in genes that promote diversity. Despite advances, unresolved phylogenetic issues persist, including the of traditional S. . Silene s.l., where molecular data place constituent across multiple subgenera, necessitating revised circumscriptions. Ongoing 2025 studies employing restriction-site associated (RAD-seq) are targeting cryptic complexes, such as those in contact zones of closely related taxa, to clarify boundaries obscured by hybridization and incomplete lineage sorting. Evolutionary trends indicate that transitions to correlate with colonization of higher northern latitudes, potentially driven by climatic pressures favoring separate sexes for , as seen in S. . Melandrium distributions.