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Asplenium scolopendrium

_Asplenium scolopendrium, commonly known as hart's-tongue fern, is an evergreen perennial fern in the family Aspleniaceae, characterized by a short that produces rosettes of arching, strap-shaped, leathery fronds up to 75 cm long with undulate margins and bright green coloration. The fronds are undivided, glossy, and bear sori in linear rows on their undersides, ripening in summer. Native primarily to the temperate regions of the , it thrives in shaded, moist environments on base-rich, soils. The species exhibits a wide but disjunct , occurring across central and southern (including the ), northwest , and parts of western extending to , with isolated populations in represented by the variety A. scolopendrium var. americanum. In , it is particularly common in the UK, , and , serving as an indicator of ancient woodlands due to its preference for stable, undisturbed habitats. Globally, it is classified as a rhizomatous geophyte adapted to temperate biomes, with notable biogeographic disjunctions, such as the isolated North American populations of var. americanum, linked to historical climate patterns. Ecologically, Asplenium scolopendrium favors damp, shady sites such as woodland floors, hedgerows, moist rock crevices, and walls, often in ashwoods or on humus-rich, well-drained soils with neutral to alkaline . It avoids acidic substrates and is frequently associated with bryophytes in cool, humid microhabitats, contributing to diversity in forests. The reproduces via spores and can spread vegetatively through rhizomes, though it is sensitive to , excessive shade removal, and fungal pathogens like (Milesina scolopendrii) in damp conditions. While the species is generally secure in its core range and not assessed as threatened by the IUCN, the North variety is federally listed as threatened due to habitat loss and limited populations. Historically, it has been used in for respiratory ailments and as an , with numerous cultivars selected for variations like crisped or crested margins.

Biology

Description

Asplenium scolopendrium is a rhizomatous that grows from a short, erect , forming dense clumps with erect to arching fronds. The fronds are simple and undivided, strap-shaped to lanceolate with a heart-shaped or auriculate base, leathery in texture, and glossy bright green in color, remaining year-round. They typically measure 30-45 cm (12-18 inches) in length and 2.5-5 cm (1-2 inches) in width, with entire margins that may become wavy or undulate with age, and a short stipe that is purplish-brown at the base, transitioning to green and covered in pale brown scales. The common name hart's-tongue fern refers to the distinctive tongue-like shape of the fronds, which resemble the tongue of a deer (hart being an archaic term for a red deer). On the underside of mature fronds, sori—linear clusters of sporangia—are arranged in pairs along the veins, primarily in the upper half of the blade, nearly perpendicular to the midrib, and covered by thin, whitish, flap-like indusia; these brown-striped sori evoke the segmented legs of a , inspiring the specific scolopendrium from for centipede. Cultivated and wild forms exhibit variations in frond texture and margins, such as more pronounced ruffled or crisped edges, while the subspecies A. scolopendrium subsp. americanum tends to have smaller fronds overall.

Taxonomy

Asplenium scolopendrium belongs to the family Aspleniaceae, within the order , and is placed in the genus , specifically the as resolved by global plastid phylogenies. The binomial name Asplenium scolopendrium was established by in his in 1753. The genus name derives from words a- (without) and splen (spleen), reflecting ancient medicinal uses believed to treat spleen ailments based on the . The specific epithet scolopendrium comes from the Greek skolopendra (), alluding to the linear rows of sori on the undersides that resemble the creature's segments. Three subspecies are currently recognized: A. scolopendrium subsp. scolopendrium, native to and (diploid, 2n=72); subsp. americanum, occurring in and distinguished by narrower fronds and scales (tetraploid, 2n=144); and subsp. japonicum, found in , which incorporates the former species A. komarovii based on phylogenetic evidence (tetraploid, 2n=144). Historically, A. scolopendrium was classified in the segregate genus Phyllitis as Phyllitis scolopendrium (L.) Newman from 1844 until the late 20th century, but molecular phylogenetic studies, including analyses of genomes and nuclear markers, have confirmed its within and supported the current placement. Notable synonyms include Scolopendrium vulgare Sm. and Scolopendrium officinale Sw.

Reproduction

Asplenium scolopendrium displays the characteristic found in ferns, with a prominent diploid phase that dominates the visible form and produces haploid spores through in sporangia clustered in linear sori on the undersides of fertile fronds. The has a chromosome number of 2n = 72 in subsp. scolopendrium and 2n = 144 in subspp. americanum and japonicum. It can produce millions of spores annually, facilitating long-distance dispersal primarily via . These tiny, homosporous spores germinate under suitable conditions to form the free-living stage, a small, thalloid prothallus typically heart-shaped and measuring just a few millimeters in diameter. The gametophyte is bisexual, bearing both antheridia (male structures producing multiflagellated sperm) and archegonia (female structures containing eggs) on its ventral surface, enabling sexual reproduction. Fertilization requires a film of water to allow sperm motility from antheridia to archegonia, a process that results in a diploid zygote developing into a young sporophyte attached to the gametophyte. This species exhibits a mixed mating system, favoring outcrossing between different gametophytes for higher sporophyte production rates (up to 30%), though intragametophytic selfing is possible and occurs at lower success rates (around 10%) due to inbreeding depression. Gametophytes are short-lived, typically surviving less than four months in culture, underscoring the ephemeral nature of this phase compared to the long-lived sporophyte. In addition to , A. scolopendrium reproduces asexually through extension of its short , which allows the to form dense clumps via vegetative propagation. The American subspecies (subsp. americanum) further demonstrates potential in the stage, where specialized outgrowths develop into independent prothalli, enhancing persistence in isolated habitats without formation; this trait is absent or minimal in the subspecies (subsp. scolopendrium). Apogamy, the of directly from gametophytic tissue without fertilization, has not been documented in this species. Spore release from mature sori generally occurs in summer, with individual fronds capable of producing up to 18,000 spores adapted for wind dispersal over long distances. Spores maintain viability for several months under dry storage conditions, though viability declines with exposure to salt or desiccation. Reproductive success, particularly of the gametophyte, is heavily influenced by environmental factors; shaded, humid conditions with high moisture availability (e.g., 90% relative humidity) are essential for spore germination, prothallus growth, and fertilization, as desiccation beyond -8.4 MPa water potential halts development. The American subspecies shows heightened sensitivity to temperature fluctuations and low moisture, with optimal gametophyte development at 20°C rather than 25°C, contributing to its rarity in variable climates.

Geographic Range

Distribution

Asplenium scolopendrium is native to the temperate regions of the , with a disjunct distribution spanning , eastern , and temperate . The species occurs widely across , from the and in the north to the in the south, including countries such as the , , , , , , , , and . It is also present in , particularly in northwest regions like , and extends eastward to . In eastern , the species is rare and restricted primarily to the , with current populations known from , , and , , where it can be locally abundant along the and . Historical records document additional occurrences in southern states including and , but these populations have since become extirpated or extremely limited, with disjunct populations in and northeastern . The subspecies A. scolopendrium subsp. japonicum is native to temperate , ranging from eastern and the [Russian Far East](/page/Russian Far East) through , , and , often found in inland mountainous areas such as Gangwon-do in Korea. In North America, A. scolopendrium var. americanum represents the distinct eastern variant, while A. scolopendrium subsp. scolopendrium dominates in and . Outside its native range, A. scolopendrium has been introduced in , where it is naturalized in limited wooded areas of the North and South Islands, including sites near , , and , but remains not widely established. In , the species maintains a stable yet patchy distribution, common in woodlands of the but rarer in peripheral regions.

Habitat

Asplenium scolopendrium thrives in shaded, humid environments characterized by cool microclimates, often on mesic slopes with north to northeast aspects. It prefers calcium-rich substrates such as dolomitic , thin soils over , and open talus or slopes, where it can grow epipetrically in rock crevices or overhangs. These conditions provide the stable, base-rich medium essential for its growth, with populations frequently occurring in understories beneath canopies like sugar maple or northern hardwoods. The requires moist but well-drained soils that are to alkaline, maintaining moderate levels supported by high and annual of 50–150 cm. It is sensitive to summer and drying conditions, though established individuals show some tolerance in humid settings; adequate winter snow cover (200–300 cm) protects against freeze in temperate regions. Light preferences include filtered or indirect illumination under 25–50% herbaceous cover, allowing it to sustain its deep green fronds while avoiding full sun exposure that could cause scorching. In terms of climate and elevation, A. scolopendrium favors mild, climates with temperatures ranging from 10–30 °C and high , occurring from lowlands to submontane altitudes, including mid- to upper-slope positions above 800 ft in regions. It can also establish terrestrially in humus-rich soils within these niches, particularly in that offer consistent moisture and shade. Such habitats are prevalent across its disjunct distributions in , eastern , and .

Ecology

Asplenium scolopendrium exhibits gregarious growth, forming dense colonies through the vegetative spread of its short, creeping rhizomes, which allow the plant to establish compact clumps of overlapping fronds in favorable sites. These colonies can expand into large drifts, particularly in shaded, moist environments, contributing to local population persistence by enabling resource sharing and protection from environmental extremes. In regions like the Niagara Escarpment, such clonal growth supports colony sizes ranging from small groups to larger aggregations, with over 90% of North American populations concentrated in discrete areas. The species frequently associates with mosses and lichens in rocky crevices, where these bryophytes and crustose lichens create a shared microhabitat that retains moisture and provides structural support for establishment. Such associations enhance suitability by stabilizing substrates and moderating humidity, though direct symbiotic nutrient exchange remains unconfirmed. While like A. scolopendrium show variable and often limited mycorrhizal associations compared to seed plants, potential fungal partnerships may aid nutrition in nutrient-poor rock environments, as observed in related taxa. Dispersal in A. scolopendrium primarily occurs through wind-borne spores, which facilitate long-distance transport and enable the species to colonize distant sites, as evidenced by its disjunct temperate distributions across , , and . However, successful establishment is constrained by the need for specific microclimates with consistent and , limiting to suitable crevices where spores can germinate into gametophytes. The mixed , including occasional intragametophytic selfing, further aids by allowing homozygous sporophytes to develop from single spores, promoting initial population founding in fragmented landscapes. Climate influences A. scolopendrium's significantly; milder winters reduce damage to fronds, potentially benefiting northern populations under projected warming scenarios. Conversely, the species is vulnerable to in drier conditions, as its leathery fronds and shallow roots offer limited , exacerbating risks from altered precipitation patterns. Herbivory on A. scolopendrium is occasional, with browsing by deer noted in some settings, though the tough fronds deter heavy consumption. Slugs and snails can young fronds in moist areas, necessitating management in but appearing minor in wild populations. Fungal diseases, including from pathogens like Phytophthora spp., arise in overly humid or poorly drained conditions, potentially stressing colonies during prolonged wet periods. In its ecosystem, A. scolopendrium contributes to by anchoring rock fissures with its rhizomatous roots, preventing in outcrops and maintaining integrity. The dense clumps also provide microhabitats for small , offering shelter and humidity in otherwise exposed rocky terrains, thereby supporting local .

Conservation

Europe

Asplenium scolopendrium is assessed as Least Concern on the European Red List of Lycopods and Ferns, with a 2017 assessment indicating a stable population across the continent. This status reflects its widespread occurrence in western and central Europe, where it remains common in suitable habitats. However, national assessments reveal regional vulnerabilities; for instance, it is classified as Vulnerable in Albania based on the 2014 national Red List due to limited populations and habitat pressures. Similarly, Norway's 2010 Red List rates it as Vulnerable under Criterion D1, highlighting small population sizes at its northern range edge. The primary subspecies in Europe is A. scolopendrium var. scolopendrium, which dominates these assessments. Population trends show stability and abundance in , particularly in the and , where it forms dense clusters in humid, shaded environments. In contrast, eastern and southern European ranges exhibit fragmentation and localized declines, attributed to habitat loss over the past several decades. Reports indicate contractions in distribution during the last 50 years, especially in southern regions, with abundance decreasing eastward from northwest hotspots like the . These trends underscore the species' sensitivity to range-edge effects, though overall European numbers remain robust. Key threats to A. scolopendrium in include from quarrying and , which disrupt its preferred damp, rocky sites. , particularly historical acid deposition, has impacted fern communities, while poses emerging risks through increased drying and shifts in humidity levels, potentially reducing habitat suitability in southern areas. Fragmentation exacerbates these issues by isolating populations and limiting . Protection efforts focus on national legislation rather than species-specific EU measures, as A. scolopendrium is not listed in Annex II or IV of the Habitats Directive. In the , it receives full protection under Schedule 8 of the , prohibiting intentional picking, uprooting, or destruction. Reintroduction initiatives address localized threats; a notable 1998 project in Germany's Mountains successfully transplanted 59 plants from a quarry-threatened site, achieving over 1700% after 10 years of monitoring. Such efforts, combined with habitat safeguards in sites, support recovery in degraded areas.

North America

The American variety of hart's-tongue fern, Asplenium scolopendrium var. americanum, is a distinct subspecies characterized by smaller fronds measuring 9 to 60 cm in length, compared to the larger fronds of the European variety, and it is tetraploid with 144 chromosomes. In North America, this subspecies faces significant conservation challenges, primarily due to its restricted distribution and vulnerability to habitat alteration. It is federally listed as Threatened under the U.S. Endangered Species Act, a status it has held since its original listing on July 14, 1989. At the state level, it is classified as Endangered in New York, Michigan, and Tennessee, reflecting its precarious situation in these core regions. In Canada, it is designated as Special Concern under the Species at Risk Act (SARA) Schedule 1, with Ontario listing it similarly under provincial law since 2017. The population of A. scolopendrium var. americanum is rare and fragmented across the , with an estimated total of approximately 122,000 individuals globally for this variety, the majority concentrated in , , and . In alone, there are about 74 extant populations totaling a conservative estimate of 30,000 plants, while supports 16 to 20 extant populations. Historically, its range extended to southern states including , , and , but it has been extirpated from most southern locales, leaving only disjunct, small populations in areas like and . This contraction underscores the subspecies' reliance on northern habitats, where it persists in low numbers vulnerable to local extirpations. Key threats to A. scolopendrium var. americanum include from , which thins the canopy and exposes to ; and quarrying operations that fragment outcrops; and urban or residential development that encroaches on sites. , such as pale swallow-wort () and common buckthorn (), compete directly with the fern for space and resources, exacerbating declines in invaded areas. poses an additional risk through rising summer temperatures, reduced snow cover that limits winter insulation, and altered precipitation patterns, which may shift suitable microclimates beyond current ranges. Protections for the subspecies are anchored in the U.S. Endangered Species Act, which prohibits take and mandates , alongside state-level safeguards in and that mirror federal prohibitions. In , the Species at Risk Act provides for habitat protection on and encourages voluntary stewardship on private properties, while Ontario's Endangered Species Act restricts activities that could harm populations. efforts include comprehensive plans emphasizing habitat restoration, such as invasive species removal and canopy maintenance, with goals to secure at least 15 self-sustaining populations in the U.S. These initiatives have supported reintroduction and augmentation in protected areas. The 2025 draft Species Status Assessment confirms ongoing population declines, with four sites extirpated and two others showing reduced numbers due to invasives and changes, though it highlights stability or increases at 13 sites bolstered by in state parks. Monitoring programs track nine populations annually and others on a five-year cycle, with similar efforts in Michigan's national forests and Ontario's reserves to inform adaptive . These updates indicate that while protections have stabilized some core sites, broader threats continue to challenge recovery across , in contrast to the more secure populations.

Asia

In , the subspecies Asplenium scolopendrium subsp. japonicum, adapted to the region's temperate climates with higher humidity and seasonal monsoons, is not separately assessed at the global level by the , which has no entry for the post-2016. This subspecies occurs patchily in temperate forests across , , and northeastern , where it is generally considered rare due to fragmented habitats, though populations are relatively more abundant on several Japanese islands compared to sites. In , it is recognized as a rare and based on national assessments. Regional evaluations suggest a potential Vulnerable status under IUCN criteria, driven primarily by ongoing habitat loss, though formal rankings vary by country. Population trends indicate declines across its East Asian range, with patchy distributions in shaded, moist forest understories making it vulnerable to fragmentation; studies on regional assemblages as of 2025 infer reductions from and , particularly in low-elevation areas where suitable microhabitats are diminishing. In and , small, isolated stands persist in mountainous regions, but overall numbers are low and decreasing, while populations show some stability in protected highlands but face similar pressures in developed zones. Quantitative data remain sparse, with no comprehensive available, highlighting the need for updated surveys to quantify extent of occurrence and population sizes. Key threats include encroaching on forest edges, industrial degrading soil and air quality in fern habitats, and competition from that alter dynamics. shifts, such as reduced humidity from altered precipitation patterns, further stress this humidity-dependent , potentially exacerbating declines in southern parts of its range. These pressures are compounded by limited monitoring, as fern often prioritizes vascular with higher economic value. Protection efforts are modest and habitat-focused rather than species-specific; the subspecies appears in national red lists in as rare, warranting general safeguards, and is indirectly protected in Japan's mountainous reserves, though not explicitly listed as threatened nationally. In , assessments under the 2017 red list for lycophytes and ferns recommend , but implementation is uneven due to competing land uses. Overall, efforts emphasize broader protection in temperate s, with calls for targeted surveys to address significant data gaps in distribution, abundance, and threat impacts across .

Uses

Cultivation

Asplenium scolopendrium thrives in partial to full shade, where it can maintain its vibrant green fronds without scorching. It prefers moist, well-drained, fertile, humus-rich soils that are neutral to alkaline, often performing best in limestone-based conditions to mimic its natural . The is hardy in USDA zones 5 to 9, tolerating a range of temperate climates but requiring protection from intense summer heat and winter . Propagation of A. scolopendrium can be achieved through several methods, including sowing spores in mid to late summer at around 15°C or dividing established clumps in spring to separate or offsets. This is notably slow-growing, often taking several years to reach maturity from spores, which makes a more practical option for gardeners seeking quicker results. Peeled bases can also be removed from the rhizome for vegetative , though success rates vary. Several cultivars of A. scolopendrium are popular in for their distinctive variations, enhancing ornamental appeal through diverse textures and forms. Notable selections include 'Angustatum', featuring narrow, strap-like fronds and holding the Royal Horticultural Society's for its reliability; 'Cristata', with crested tips adding a ruffled appearance; and 'Kaye's Lacerated', characterized by deeply and irregularly lobed margins for a frilled effect. Other variants, such as those with undulating or furcate edges, provide options for textured contrasts in shaded plantings. In garden settings, A. scolopendrium excels in rock gardens, walls, and borders, where its rosettes offer year-round structure in shaded areas. It is also well-suited to container cultivation in placed in protected, shaded spots, provided the medium remains consistently moist. Mulching around the base helps retain and suppress weeds, contributing to low-maintenance growth. While generally pest-resistant, A. scolopendrium can be susceptible to damage from slugs, snails, and grubs, which feed on the fronds and roots; controls or barriers are recommended to manage these threats. Routine involves removing any damaged or browned fronds to promote vigor, along with ensuring consistent without waterlogging to prevent . Historically, A. scolopendrium gained prominence during the Victorian era's , or fern fever, when it was widely cultivated in shaded conservatories and gardens for its elegant form, reflecting the period's fascination with exotic and native ferns.

Herbal Medicine

Asplenium scolopendrium, commonly known as hart's-tongue fern, has been employed in traditional primarily for its soothing and properties, with historical records dating back to 17th-century European herbals. In Nicholas Culpeper's Complete Herbal (1653), the fern is commended against the hardness and stoppings of the and liver, the stone in the kidneys, all inward swellings especially in the veins and liver, pains in the joints, and the bloody flux; the distilled water drunk twice a day is singular for those troubled with the stone in the reins or kidneys, while the juice dropped into the ears takes away pain, and heals wounds, ulcers, and burns. Culpeper also noted its application in decoctions or ointments from fronds and rhizomes for treating earaches, conditions like burns and ulcers, and as a vulnerary for . The fern's traditional uses extend to folk remedies in , where infusions were prepared for coughs and relief, leveraging its mucilaginous content to soothe irritated mucous membranes in the . These applications were often based on the , linking the fern's tongue-like fronds to oral and throat ailments. Key active compounds in A. scolopendrium include , which provides and expectorant effects for respiratory soothing; , contributing properties for skin and wound applications; and such as derivatives, offering and benefits. These constituents are concentrated in the fronds, supporting the fern's historical use in teas, decoctions, and topical preparations. In modern contexts, scientific validation for these traditional uses remains limited, with no robust clinical supporting for respiratory conditions, earaches, or skin issues. Due to the presence of potential carcinogens common in and the enzyme , which can deplete , is not recommended, and caution is advised, especially for pregnant or breastfeeding individuals. Professional consultation is essential to avoid risks associated with related species.

Other Uses

Asplenium scolopendrium holds cultural significance in , particularly within medieval Welsh traditions, where it is referenced in texts like the Meddygon Myddfai as 'dauot yr hydd' for use as an and general wellness. Its distinctive strap-shaped fronds have also inspired representations in and botanical art, symbolizing endurance and elegance in temperate landscapes. Beyond cultivation, the fern contributes to efforts, where it is incorporated into native plantings for rehabilitation in shaded woodlands and areas. Studies on relocated populations demonstrate effective establishment, with spore-based propagation yielding viable communities over decadal monitoring in European projects. Similar initiatives in utilize the species to enhance in conservation sites, supporting recovery in threatened ecosystems. Commercially, Asplenium scolopendrium sees limited use, mainly as spores in propagation kits for hobbyists and educators, or as live in eco-tourism exhibits highlighting native . Ornamental cultivars like 'Undulatum' are briefly noted in these markets for their wavy fronds.

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