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Drynaria quercifolia

Drynaria quercifolia is a species of epiphytic fern in the family Polypodiaceae, characterized by its dimorphic fronds and thick, fleshy rhizome covered in dark brown scales.^[1]^[2] It features two types of fronds: sterile nest fronds that are ovate, up to 40 cm long and 30 cm wide, and lobed to collect humus and debris; and fertile foliage fronds that are pinnatifid, up to 130 cm long and 40 cm wide, with coriaceous, pale to mid-green laminae and round sori arranged in two regular rows.^[2]^[3] The rhizome is approximately 2 cm thick, woolly, and bears scales that are 2–25 mm long and ginger to dark brown in color, allowing the plant to form large clumps on hosts.^[3] Native to tropical regions, D. quercifolia is widely distributed from Hainan in China through tropical Asia, including India, Sri Lanka, Nepal, Bangladesh, Myanmar, Thailand, Laos, Cambodia, Vietnam, Peninsular Malaysia, Singapore, Borneo, Indonesia (Jawa, Lesser Sunda Islands, Maluku, Sulawesi, Sumatera), the Philippines, New Guinea, and extending to northern and western Australia (such as Cape York in Queensland and Arnhem Land in the Northern Territory), as well as the Andaman and Nicobar Islands and Fiji.^[1]^[2] It thrives as an epiphyte on tree trunks, lithophyte on rocks and boulder crevices, or occasionally as a terrestrial plant in primary and secondary rainforests, freshwater swamp forests, tropical vine forests, beach scrubs, disturbed areas, and along creek rock faces, primarily in wet tropical biomes.^[1]^[2]^[3] This perennial, autotrophic fern grows up to 1.5 m in height, with wind-dispersed spores from exindusiate sori measuring 1–2 mm in diameter, and its fertile fronds are deciduous.^[2]^[3] Notable for its ornamental value and traditional medicinal applications, D. quercifolia is cultivated in parks and along roadsides for its attractive foliage, often propagated by spores.^[2] The whole plant, particularly the rhizome, which is bitter and astringent, is used in traditional medicine to treat conditions such as fever, swellings, bone fractures, cough, headache, and typhoid fever.^[2]^[4] Commonly known as the oakleaf fern or oak-leaf basket-fern, it was first described as Polypodium quercifolium by Carl Linnaeus and later transferred to Drynaria by John Smith in 1841.^[1]^[4]

Taxonomy

Classification

Drynaria quercifolia is classified within the kingdom Plantae, phylum Tracheophyta, class Polypodiopsida, order Polypodiales, family Polypodiaceae, genus Drynaria, and species D. quercifolia.^[1] The binomial name is Drynaria quercifolia (L.) J.Sm., with the authority attributed to John Smith in 1841; it is based on the basionym Polypodium quercifolium, originally described by Carl Linnaeus in Species Plantarum in 1753.^[1]^[5] In the Pteridophyte Phylogeny Group I classification published in 2016 (PPG I), the species was transferred to the genus Aglaomorpha as A. quercifolia due to phylogenetic evidence indicating that Drynaria was paraphyletic with respect to Aglaomorpha and related genera; however, a 2012 proposal to conserve the name Drynaria against Aglaomorpha for nomenclatural stability was advanced and accepted in 2016, and post-2016 taxonomic treatments, including updates in Plants of the World Online as of 2025, have reverted to recognizing it under Drynaria.^[1]^[6]^[7]^[8] The genus Drynaria comprises approximately 32 accepted species of primarily epiphytic ferns, characterized by dimorphic fronds—where sterile fronds are broad and persistent for debris capture, forming nest-like baskets, while fertile fronds are narrower and bear sporangia.^[9]

Etymology and synonyms

The genus name Drynaria derives from the Greek drys (oak tree) or dryas (oak nymph or oak-shaped), referring to the oak-like, pinnatifid fronds characteristic of the genus.^[10]^[2] The species epithet quercifolia originates from the Latin quercus (oak) and folium (leaf), emphasizing the fronds' resemblance to oak foliage.^[2] The currently accepted name is Drynaria quercifolia (L.) J. Sm., based on the basionym Polypodium quercifolium L. from Species Plantarum (1753).^[11] This species has accumulated numerous synonyms over time due to taxonomic revisions, with numerous heterotypic synonyms recognized, reflecting historical placements in genera such as Phymatodes, Aglaomorpha, and Drynariopsis. Key synonyms include:

Aglaomorpha quercifolia (L.) Hovenkamp & S. Linds. (2017)
Polypodium quercifolium L. (1753)
Phymatodes quercifolia (L.) C. Presl (1836)
Drynaria brancifolia (C. Presl) T. Moore (1859)
Drynaria morbillosa (C. Presl) T. Moore (1859)
Drynariopsis quercifolia (L.) C. Presl (1836)
Phymatodes brancifolia (C. Presl) C. Chr. (1906)
Phymatodes sylvatica (Schkuhr) C. Presl (1836)

These synonyms arise from varying interpretations of frond dimorphism and phylogenetic relationships within Polypodiaceae.^[1]^[11] Nomenclaturally, Drynaria quercifolia was temporarily classified under Aglaomorpha in the Pteridophyte Phylogeny Group I (PPG I) system of 2016, which recognized Aglaomorpha as a segregate genus based on molecular data.^[12] However, the name Drynaria was conserved against Aglaomorpha via a proposal published in 2012 and accepted in 2016 to promote nomenclatural stability, given Drynaria's widespread prior usage and the paraphyly concerns with Aglaomorpha.^[13]^[8] As of 2025, authoritative databases like Plants of the World Online and World Flora Online uphold Drynaria as the valid genus.^[1]^[11]

Description

Morphology

Drynaria quercifolia is a robust epiphytic fern that typically forms large, clumping masses up to 1 m tall, supported by long-creeping rhizomes that are thick, fleshy, woody, and 1-2 cm in diameter. These rhizomes are densely covered in imbricate scales that are pale to dark brown, 2-25 mm long, and 0.7-2.5 mm wide, often with a woolly or fuzzy appearance due to their soft, ginger-colored to reddish-brown texture and fine marginal teeth. The plant's growth habit is adapted for epiphytic life, occasionally lithophytic or terrestrial, with the rhizomes branching and providing structural support on tree trunks or rocks. In some populations, it exhibits deciduous tendencies, particularly with the shedding of fertile fronds seasonally.^[14]^[3]^[11]^[15] A key feature of D. quercifolia is its pronounced frond dimorphism, distinguishing sterile (nesting) fronds from fertile (foliar) fronds. Sterile fronds are shorter and broader, measuring 15-40 cm long and 9-25.5 cm wide, with a simple, broadly ovate to rounded shape, obtuse apex, and oblique base; they are leathery, stiff, and rough-textured, initially green but maturing to a shiny brown. These fronds overlap in a basket-like arrangement at the rhizome tips, functioning to trap falling debris and organic matter for nutrient uptake, and they remain persistent on the plant. Fertile fronds emerge from the center of this nest structure, reaching 50-100 cm or more in length, with deeply pinnatifid laminae 50-60 cm long and 5-6 cm wide, featuring strap-like, oak-resembling lobes that are 1.5-32 cm long and 1.1-4.9 cm wide, leathery, pale to mid-green, with a caudate apex and attenuate base. Stipes of both frond types are 1-20 cm long, green, adaxially grooved, and sparsely covered in hairs and paleae.^[3]^[15]^[16] The fertile fronds bear sori arranged in two regular rows between the main lateral veins, appearing as round, discrete structures 1-2 mm in diameter, sometimes slightly oblong and impressed, without significant indusia. Roots are numerous, aerial, and adventitious, fibrous and dark brown in color, emerging from the rhizome to facilitate attachment and anchorage to host substrates while absorbing moisture and nutrients from the air and surface films.^[3]^[15]

Reproduction

_Drynaria quercifolia exhibits the typical fern life cycle of alternation of generations, with a dominant diploid sporophyte phase and a free-living haploid gametophyte phase. The sporophyte, which is the visible plant form, produces spores through meiosis in sporangia located in sori on specialized fertile fronds. These spores germinate to form prothalli, the gametophyte stage, which are small, heart-shaped, and hermaphroditic, bearing both antheridia and archegonia for gamete production. Fertilization occurs in moist conditions, as sperm must swim to the egg, leading to the development of a new sporophyte. Unlike seed plants, D. quercifolia relies entirely on spores for sexual reproduction and requires humidity for successful gamete fusion.^[14]^[2]^[17] Sexual reproduction involves spore production on taller, annual fertile fronds, which reach up to 1 meter in height and are deciduous. The spores are wind-dispersed, facilitating colonization of new epiphytic sites. Upon landing in suitable moist substrates, spores germinate into filamentous prothalli that mature into cordate gametophytes, typically within weeks under optimal in vitro conditions with nutrient media. These gametophytes produce flagellated sperm in antheridia and eggs in archegonia, enabling self-fertilization or cross-fertilization in humid environments. The resulting zygote develops into a young sporophyte that emerges from the gametophyte.^[14]^[2]^[18] Asexual reproduction occurs primarily through extension and division of the long, creeping rhizome, enabling clonal propagation and spread in epiphytic habitats. Rhizome segments, especially larger ones with established shoots, root readily when detached, forming new individuals that maintain genetic uniformity. This vegetative mode supports population persistence in stable tree canopy environments without reliance on spore dispersal. Smaller rhizome sections lacking shoots can also generate adventitious shoots, enhancing resilience.^[14] In cultivation, propagation mirrors natural methods but emphasizes sterility to prevent contamination. Spores are sown on sterile media, such as agar with nutrients, where they germinate and develop into gametophytes before sporophytes emerge, often requiring controlled humidity and light. Rhizome division remains the simplest approach for horticultural use, with divisions planted in epiphytic mixes to mimic tree bark conditions. No specific hybrids involving D. quercifolia are documented.^[14]^[19]

Distribution and habitat

Geographic range

Drynaria quercifolia is native to tropical regions from Hainan in China through South and Southeast Asia—including the Andaman and Nicobar Islands, Sri Lanka, Nepal, India (including Assam), Bangladesh, Myanmar, Thailand, Laos, Cambodia, Vietnam, Peninsular Malaysia, Singapore, Indonesia (including Sumatra, Borneo, Java, Lesser Sunda Islands, Maluku, Sulawesi), and the Philippines—to New Guinea and northern and western Australia (Queensland, Northern Territory, Western Australia), as well as Fiji.^[1]^[11]^[2] This distribution spans diverse island and mainland habitats across the Indo-Australian archipelago, where the species thrives as an epiphyte or lithophyte.^[1] The species has been introduced and naturalized outside its native range, notably in central and southern Florida, USA, where it escapes cultivation and establishes in subtropical environments.^[20]^[21] It is also cultivated widely as an ornamental in tropical gardens worldwide, with occasional naturalization in some Pacific islands.^[14] Its range expansion beyond native areas is largely attributed to the international trade in ornamental plants since the 1800s, facilitating its spread as a popular epiphytic fern in cultivation across tropical regions.^[21]^[14]

Habitat preferences

Drynaria quercifolia primarily inhabits lowland tropical rainforests, monsoon forests, secondary woodlands, and occasionally disturbed areas as an epiphyte on tree trunks and branches, or epilithically on rocks and cliff faces.^[14]^[1]^[3] It is also recorded in riverine zones, freshwater swamp forests, beach scrubs, and bouldery areas along creeks, often forming large clumps in the understory.^[2]^[11] The species thrives in humid tropical climates at elevations from sea level to approximately 1500 m, with optimal conditions in shaded understories where temperatures range between 20–30°C and annual rainfall exceeds 2000 mm.^[14]^[2]^[22] It prefers semi-shaded environments with high humidity but can tolerate areas with a distinct dry season, provided moisture levels remain adequate.^[14]^[3] As an epiphyte, D. quercifolia favors rough-barked host trees such as dipterocarps, mangroves, Ficus species, and Mangifera indica for attachment and stability, though it shows variability across 11 or more host species in some regions.^[14]^[23]^[24] Its sterile nest fronds accumulate leaf litter and humus in the plant's crown, providing a nutrient-rich substrate independent of the host.^[3]^[14] This fern exhibits adaptations to its epiphytic lifestyle, including tolerance to periodic drought through water storage in rhizomes and fronds, but it is sensitive to frost and prolonged full sun exposure.^[14]^[25]

Ecology

Interactions with other organisms

_Drynaria quercifolia, as an epiphytic fern, forms symbiotic associations with root fungi that enhance nutrient uptake and drought tolerance in its canopy habitat. Root symbiotic fungi (RSF) isolated from this species, including genera like Trichoderma and Fusarium, promote plant growth by improving water retention and stress adaptation mechanisms, with debris accumulated in the nest fronds supporting fungal communities that aid in organic matter decomposition for nutrient cycling.^[26] These nest fronds also provide microhabitats for invertebrates, including ants, which may nest within the protective basket-like structures formed by the sterile fronds. In the genus Drynaria, extrafloral nectaries on fronds attract ant bodyguards, offering indirect defense against herbivores through mutualistic interactions that originated convergently with those in angiosperms during the Cretaceous.^[27] Spore dispersal in D. quercifolia primarily occurs via wind, enabling colonization of new phorophytes in humid tropical environments, though occasional bird-mediated transport has been noted in similar epiphytic ferns. Fronds are subject to browsing by insects, but chemical defenses such as tannins in leaves and rhizomes deter herbivory by binding to proteins and reducing palatability.^[2]^[28] Reproduction involves alternation of generations, with haploid gametophytes bearing antheridia and archegonia that require moist microenvironments for sperm to swim to eggs, without reliance on animal vectors typical of seed plants.^[29] In rainforest communities, D. quercifolia contributes to epiphyte diversity by trapping litter that retains canopy humidity and fosters microbial activity, supporting broader ecosystem stability in humid tropical forests.^[30]

Conservation status

_Drynaria quercifolia has not been globally assessed by the IUCN Red List and is categorized as Not Evaluated.^[11] Local conservation statuses vary across its range; it is rated as Least Concern in Singapore^[2] and Sri Lanka,^[31] Not Evaluated in India,^[32] Not Threatened in Western Australia,^[33] and Vulnerable in the Philippines.^[24] The species faces threats from deforestation and logging, which fragment habitats in regions like India and Australia.^[34] ^[35] Overcollection for traditional medicinal uses contributes to population declines in areas with high demand, such as parts of South Asia.^[36] Populations appear stable in core tropical ranges with intact forests but are declining in fragmented lowland areas due to these pressures.^[37] Conservation measures include protection within national parks and reserves, such as Key Biodiversity Areas in the Philippines^[38] and various Australian protected areas where it occurs naturally.^[35] The species is not listed under CITES, though international trade is monitored in some contexts due to medicinal demand. Efforts focus on habitat preservation to mitigate ongoing threats from land-use changes.

Human uses

Medicinal applications

Drynaria quercifolia has been utilized in traditional medicine across various cultures, particularly for its rhizome, which is employed in Ayurvedic formulations to promote bone healing, alleviate pain, reduce fever, treat diarrhea, address skin diseases, and manage infertility.^[39] Indigenous groups, including the Kadars in India's Anamalai Hills, use the plant as an antifertility agent, with ethnobotanical records documenting its application by women to prevent conception.^[40] Additionally, traditional practices incorporate the rhizome for treating rheumatoid arthritis, worm infections, and as an adjunct therapy for diabetes, often in decoction or paste forms to relieve joint pain and gastrointestinal issues.^[41] In the Philippines, folklore associates the fern, locally known as pakpak lawin, with wound healing, where frond poultices are applied to swellings and injuries.^[42] Phytochemical analysis reveals the presence of bioactive compounds in D. quercifolia, including flavonoids such as quercetin, triterpenoids like β-amyrin and friedelin, and polysaccharides including glucans and mannans, primarily concentrated in the rhizome and fronds.^[43] These constituents contribute to the plant's pharmacological properties; rhizome extracts exhibit anti-inflammatory effects by reducing paw edema in animal models, antioxidant activity through DPPH scavenging (IC<sub>50</sub> = 52.4 μg/mL), and antibacterial efficacy against pathogens like Escherichia coli (MIC 125-250 μg/mL).^[43]^[44] Modern research in the 2020s has validated these traditional applications, particularly the rhizome's role in managing rheumatic pain. A 2025 study using a Freund's Complete Adjuvant-induced arthritis model in Wistar rats demonstrated that methanolic rhizome extract (DME) at doses of 250-500 mg/kg orally for 28 days significantly reduced paw swelling, arthritis scores, and pro-inflammatory markers (TNF-α, IL-6, COX-2), while normalizing oxidative stress parameters (SOD, GSH, MDA, CAT), suggesting immunomodulatory mechanisms via AKT and PPARG pathways.^[45] Earlier investigations, including a 2021 in vivo study, confirmed the extract's anti-rheumatoid potential by inhibiting paw edema and inflammatory cytokines, comparable to standard treatments like indomethacin.^[39] Preparations such as pastes or decoctions remain common in contemporary herbal practices for these indications.

Ornamental cultivation

Drynaria quercifolia, commonly known as the oakleaf fern or basket fern, is cultivated as an ornamental plant for its distinctive dimorphic fronds and epiphytic growth habit, making it suitable for hanging baskets, mounted displays, and tropical garden settings.^[14] This fern is appreciated for its aesthetic appeal in indoor environments and outdoor landscapes in suitable climates, where it can form large clumps up to 1 meter in height.^[3] Propagation of Drynaria quercifolia is primarily achieved through spores or division of the rhizome. For spore propagation, spores are harvested from the undersides of mature fertile fronds and sown on a sterile mix of peat moss and vermiculite, maintained at 18-24°C with high humidity under a dome; germination typically occurs in weeks to months.^[46] Rhizome division is performed in early spring by separating sections of the mature plant, each with at least one growth point and frond, and planting them in a well-draining medium while keeping fronds above the surface; larger divisions establish more readily, though smaller ones may produce adventitious shoots.^[47]^[14] In cultivation, Drynaria quercifolia thrives as an epiphyte in pots filled with orchid bark, a mix of bark, coco coir, and perlite, or mounted on wood slabs or cork for optimal air circulation.^[47] It requires bright indirect light or partial shade, avoiding direct sunlight to prevent frond burn, and temperatures between 18-38°C, with protection from anything below 10°C.^[46]^[47] High humidity of at least 60% is essential, achieved through regular misting, humidifiers, or pebble trays, while watering should keep the medium consistently moist but not waterlogged—allowing the top inch to dry slightly between waterings.^[48]^[47] Fertilization with a diluted, high-nitrogen formula every two weeks during spring and summer supports growth, ceasing in fall and winter.^[47] The plant is hardy in USDA zones 10-11, where it can be grown outdoors in frost-free areas.^[47]^[48] Common cultivation issues include root rot from overwatering or poor drainage, which can be prevented by using well-aerated substrates and monitoring moisture levels.^[46] Low humidity or inconsistent watering may cause frond browning or slowed growth, while excessive direct light leads to scorching; regular inspection for pests such as scale insects is recommended, though specific infestations are rare in proper conditions.^[46] Pruning of damaged fronds in early spring with sterilized tools helps maintain plant health.^[47] Historically, Drynaria quercifolia has been traded and cultivated as an ornamental since at least the late 20th century in tropical regions, featuring in decorative displays and noted for its use in hanging containers or rockeries in warm climates.^[49] Today, it is popular in terrariums, indoor collections, and tropical gardens worldwide for its unique form.^[3]

References

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[45]
Unveiling the mechanism of amelioration of adjuvant-induced ...
Apr 8, 2025 · Drynaria quercifolia (L.) J. Sm, also known as Oak leaf fern or “Pankhiraj” locally in West Bengal (India), is a medicinal pteridophyte, found ...Missing: etymology | Show results with:etymology
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How to Propagate Drynaria quercifolia
### Summary of Drynaria quercifolia Propagation and Care
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Buy 1 Oakleaf Fern, Aglaomorpha Drynaria Quercifolia - Jiffy Plants
14-day returnsThe Oakleaf fern can be propagated by division. In early spring, carefully separate the rhizomes, ensuring each division has at least one growth point (frond or ...
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How to Grow and Care for Oakleaf fern - PictureThis
Key care points are ensuring consistently moist soil without overwatering, and maintaining high humidity, often requiring regular misting or a pebble tray. As ...Missing: Drynaria | Show results with:Drynaria
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Decorative Fern Fronds Can Be Cooling - The New York Times
Jul 29, 1979 · Oak‐leaf fern, Drynaria quercifolia, is another native of the South Pacific Islands. I recall seeing one in Fiji, high in a windswept palm tree ...