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Azolla pinnata

Azolla pinnata, commonly known as mosquito fern or feathered mosquitofern, is a small, free-floating aquatic fern belonging to the family Salviniaceae.^[1] It features overlapping pairs of tiny, ovate to broadly elliptic leaves, each less than 1 mm long and often purplish to reddish in color, arranged along branched stems that form triangular fronds measuring 1–3 cm long and 1–2.5 cm wide, with roots up to 5 cm long dangling into the water.^[1] Native to tropical and subtropical regions across Africa, Madagascar, India, Southeast Asia, China, Japan, and Australia, it thrives in wind-protected, slow-moving freshwater bodies such as ponds, lakes, ditches, and rice paddies, tolerating low salinity up to 30 mM and optimal temperatures of 29–33°C.^[1]^[2] This species is renowned for its rapid vegetative reproduction through fragmentation, doubling its biomass every 2–5 days under ideal conditions, which allows it to form dense surface mats that can cover entire water bodies.^[2]^[1] It also reproduces sexually via heterosporous spores produced in specialized sporocarps, though this mode is less common.^[1] A defining feature is its obligate symbiosis with the cyanobacterium Anabaena azollae, hosted in leaf cavity hairs, which fixes atmospheric nitrogen at rates supporting 40–60 kg per hectare annually, enhancing its role as a natural biofertilizer.^[2] Ecologically, A. pinnata contributes to nutrient cycling, water purification by absorbing heavy metals and pollutants, and mosquito control by smothering breeding sites, while its dense growth can suppress weeds and algae in aquatic systems.^[1]^[2] In agriculture, Azolla pinnata has been utilized for centuries, particularly in Asia, as green manure in rice fields to boost soil fertility and yields without synthetic inputs, and as a high-protein forage for livestock, containing up to 25–35% crude protein.^[2]^[3] Its potential extends to sustainable practices like bioremediation, climate-resilient farming, and even as a model organism for genomic studies due to its unique symbiosis and fast growth.^[2] However, in non-native regions like North America, it is considered invasive, potentially disrupting native aquatic ecosystems by outcompeting other plants and altering water quality.^[1]

Description

Morphology

Azolla pinnata is a diminutive aquatic fern featuring a slender, triangular stem that measures up to 2.5 cm in length and floats on the water surface. The stem displays pinnate branching, with side branches arranged alternately and becoming progressively longer toward the base, facilitating vegetative fragmentation for propagation as the main axis decomposes.^[4]^[5] The leaves are small, typically 1-2 mm long, and overlap in two ranks along the stem, consisting of a bilobed structure with an upper aerial lobe and a lower submerged lobe. The upper lobes exhibit colors ranging from green to blue-green or dark red, influenced by environmental factors such as light intensity, while the lower lobes are translucent and brownish. These upper lobes are covered with multicellular hairs that confer a velvety texture and water-repellent properties; the dorsal leaf cavities formed by these structures house symbiotic cyanobacteria.^[6]^[7]^[8] Hairlike roots, often with a feathery appearance, extend downward from the fronds into the water, primarily for nutrient uptake. The overall fronds form dense, floating mats on water surfaces, with individual plants typically 1-2.5 cm across, though colonies can expand more broadly. Morphological variations occur under differing conditions, such as reduced frond size and growth in low light environments, where the plant adapts by forming tighter mats.^[4]^[5]^[7]

Symbiotic relationship

_Azolla pinnata forms a mutualistic symbiosis with the endophytic cyanobacterium Anabaena azollae (also referred to as Nostoc azollae), which resides within specialized cavities on the dorsal surface of its leaves. These leaf cavities are lined with multicellular hairs that facilitate the enclosure and protection of the cyanobacterial filaments, creating a microhabitat where A. azollae colonizes the periphery of the cavity. This association is obligate for the cyanobacterium, as it has lost the ability to survive independently outside the host.^[9]^[10] The primary function of this symbiosis is biological nitrogen fixation, where A. azollae employs nitrogenase enzymes in specialized heterocyst cells to convert atmospheric dinitrogen (N₂) into ammonia (NH₃), which is then assimilated into amino acids. This process supplies the majority of A. pinnata's nitrogen requirements, often meeting nearly the entire demand of the host under optimal conditions, enabling the fern to thrive in nitrogen-poor aquatic environments. In exchange, the plant provides fixed carbon to the cyanobiont in the form of carbohydrates, such as sucrose, glucose, and fructose, derived from host photosynthesis; these sugars support the energy needs of A. azollae for nitrogenase activity and growth. The exchange occurs via the cavity hairs, which feature labyrinthine wall ingrowths that enhance nutrient transfer between partners.^[11]^[9]^[12] The symbiosis exhibits high specificity, with distinct strains of A. azollae adapted to particular Azolla species, showing genetic divergence that correlates with host taxonomy, such as between the Azolla and Rhizosperma sections. This host-species specificity ensures compatibility and efficient mutualism. The association is maintained through vertical inheritance, where cyanobionts are transmitted maternally via the gametophytes enclosed within spores; during sporocarp development, akinetes (dormant cyanobacterial cells) are incorporated into megasporocarps and subsequently transferred to developing fronds upon germination.^[13]^[14] Evolutionary evidence indicates that the Azolla-Anabaena symbiosis originated approximately 80 million years ago during the Late Cretaceous in North America, coinciding with a whole-genome duplication event in the Azolla lineage that facilitated the stable, intergenerational transmission of the cyanobiont. Fossil records from formations like the Claggett and Judith River support this timeline, highlighting the ancient co-evolution of the partners into a highly integrated superorganism.^[15]

Taxonomy

Classification

Azolla pinnata is classified within the kingdom Plantae, phylum Tracheophyta, class Polypodiopsida, order Salviniales, family Salviniaceae, genus Azolla, and species A. pinnata.^[16] The species was first described by Robert Brown in 1810 in his Prodromus Florae Novae Hollandiae et Insulae Van-Diemen.^[17] The genus Azolla comprises 6–7 species of small, floating aquatic ferns, traditionally divided into two sections—section Azolla and section Rhizosperma—distinguished primarily by differences in frond shape and the position of sporocarps.^[18] Within the family Salviniaceae, Azolla is the sister genus to Salvinia, with both genera characterized as heterosporous ferns adapted to aquatic environments.^[19]

Varieties

The taxonomy of Azolla pinnata and related forms is debated. Some classifications recognize three subspecies exhibiting subtle variations in morphology and distribution: A. p. subsp. pinnata, the typical form with green to reddish fronds and elliptical dorsal leaf lobes, native to Australia; A. p. subsp. asiatica, featuring lax fronds with short, wide submerged lobes and more prevalent in East Asia; and A. p. subsp. africana, primarily restricted to African regions.^[20]^[7]^[5] However, other authorities, such as Plants of the World Online, treat the Asian and African forms as distinct species: Azolla imbricata (corresponding to subsp. asiatica) and Azolla africana (corresponding to subsp. africana), with A. pinnata limited to its Australasian range. Identification to this level is often challenging without molecular tools due to morphological similarities.^[16]^[7] Historical synonyms include Azolla imbricata (applied to the Asian form due to misidentification of overlapping leaves).^[21]^[7] Genetic analyses reveal minor molecular differences among these forms, such as variations in DNA amplification patterns, alongside the noted morphological traits, yet they remain interfertile and capable of hybridization.^[22]^[23] The species A. pinnata holds an IUCN Red List status of Least Concern (as of 2018), reflecting its occurrence in its native range, though individual subspecies or related species have not been assessed separately.^[24]

Distribution and habitat

Native range

Azolla pinnata is native to tropical and subtropical regions across Africa, including areas such as the Nile Valley and Madagascar, and extends through South and Southeast Asia, New Guinea, and northern Australia, encompassing countries like India, China, Japan, Malaysia, and the Philippines.^[25]^[7]^[26] In these native areas, the species primarily occupies stagnant or slow-moving freshwater habitats, such as ponds, ditches, rice paddies, and protected backwaters of streams and rivers.^[7]^[1] Historical documentation of its presence and use dates back to ancient Asian agricultural texts, including the Er Ya from approximately 2000 years ago and Jia Ssu Hsieh's Chih Min Tao Shu in 540 A.D., where it is described in the context of rice cultivation practices.^[2]^[27] The plant prefers warm climates with temperatures ranging from 18°C to 28°C and neutral to slightly acidic water conditions with a pH of 4.5 to 7, though it is notably absent from high-altitude zones and saline waters.^[7]^[28]

Introduced ranges

Azolla pinnata, native to regions in Africa, Asia, New Guinea, and northern Australia, has been introduced to various areas outside its indigenous range through human-mediated pathways.^[29] The species has been intentionally introduced for use as a green manure and fertilizer in agriculture, as well as through the aquarium and ornamental plant trade. Accidental introductions have occurred via attachment to waterfowl or contaminated trade materials, facilitating its dispersal to new water bodies.^[30]^[31]^[30] Introduced populations are established in New Zealand, Pacific Islands such as the Cook Islands and Marshall Islands, and parts of North America including Florida and Hawaii. In these regions, it has colonized tropical to temperate wetlands, forming dense floating mats on still or slow-moving waters.^[5]^[1] In the United States, Azolla pinnata is classified as a federal noxious weed and is specifically listed as such in states including Alabama and North Carolina, reflecting its potential to disrupt aquatic ecosystems where established. It has been reported in parts of Europe, including historical records in the United Kingdom.^[7]^[32]^[33]^[34]^[35]

Ecology

Growth dynamics

Azolla pinnata exhibits rapid vegetative growth, with biomass doubling every 1.9 to 7 days under optimal conditions such as high light intensity and adequate phosphorus availability.^[36] This fast proliferation is facilitated by its floating habit, allowing efficient capture of sunlight and nutrients at the water surface.^[37] Growth rates can reach up to 0.321 day⁻¹, corresponding to a minimum doubling time of approximately 2.16 days in controlled environments with sufficient phosphorus supplementation.^[36] Nutritionally, A. pinnata has a high demand for phosphorus, with optimal growth requiring at least 0.06 ppm in the medium, while it is largely independent of external nitrogen sources due to its symbiosis with the cyanobacterium Anabaena azollae, which fixes atmospheric nitrogen.^[38] Phosphorus limitation can significantly reduce biomass accumulation and nitrogen fixation efficiency.^[39] Growth is limited by sensitivities to certain environmental factors, including copper toxicity, which causes root detachment and chlorosis even at low concentrations.^[40] The plant tolerates salinity up to approximately 2 ppt, beyond which growth declines sharply due to osmotic stress.^[7] Additionally, A. pinnata prefers stagnant or gently flowing water, as high water velocities can disrupt mat formation.^[41] Under favorable conditions, A. pinnata populations rapidly expand to form dense surface mats, achieving 100% coverage of water bodies within 2 to 4 weeks from initial inoculation.^[42] This mat development supports high biomass yields, often reaching 10–20 t ha⁻¹ at saturation.^[42]

Interactions with environment

Dense mats formed by Azolla pinnata significantly reduce dissolved oxygen concentrations in water bodies, often leading to hypoxic conditions that adversely affect fish populations and macroinvertebrate densities.^[31] This oxygen depletion arises from the plant's rapid surface coverage, which blocks atmospheric exchange and promotes anaerobic decomposition beneath the mats.^[43] In addition, these mats degrade overall water quality by increasing turbidity through sediment entrapment and altering chemical parameters, such as reducing pH and conductivity while sequestering phosphorus at rates up to 122 kg/ha/year.^[31]^[44] Although this nutrient uptake can temporarily mitigate excess phosphorus, the subsequent decomposition of accumulated biomass releases bound nutrients back into the water, potentially perpetuating eutrophication cycles in nutrient-rich environments.^[45] On biodiversity, A. pinnata outcompetes native aquatic plants by forming impenetrable surface layers that restrict light penetration to submerged vegetation, thereby decreasing native plant richness and abundance.^[31] This shading effect has been observed to suppress species like Vallisneria americana, limiting their photosynthesis and growth, while also reducing phytoplankton populations due to diminished light availability.^[43] Dense mats also suppress algal blooms and weeds through nutrient competition and shading.^[2] In invaded ecosystems, such as northern New Zealand waterways, A. pinnata has displaced native congeners like Azolla rubra, further eroding local aquatic biodiversity.^[4] In terms of climate interactions, A. pinnata serves as a potential carbon sink, sequestering up to 21,266 kg CO₂/ha/year through its biomass accumulation, supported by symbiotic nitrogen fixation rates of 0.3–0.6 kg N/ha/day.^[44] However, the anaerobic decomposition of its dense mats can produce methane emissions, contributing to greenhouse gas releases in stagnant water systems.^[46] This dual role underscores its influence on carbon and nutrient cycles in aquatic habitats. A. pinnata exhibits sensitivity to abiotic stressors, particularly herbicides; low doses of glyphosate effectively inhibit its growth by disrupting amino acid synthesis, though regrowth may necessitate repeated applications for control.^[31]^[47]

Reproduction

Asexual reproduction

Azolla pinnata primarily reproduces asexually through fragmentation, in which portions of the stem, including attached roots and fronds, detach and develop into independent plants. This process allows separated fragments to rapidly establish new colonies on water surfaces, contributing to the species' ability to form dense mats in aquatic environments. Fragmentation occurs naturally as the plant grows, with stem pieces breaking off due to mechanical forces such as water currents or plant density.^[2]^[48] The heavy reliance on this clonal propagation mechanism results in a clonal process, where populations expand primarily through identical copies of the parent plant, leading to low genetic diversity within many stands. Studies on Azolla species indicate that infrequent sexual reproduction exacerbates this uniformity, limiting adaptability in some populations but enabling rapid colonization in favorable habitats. This clonal dominance is evident in both native and introduced ranges, where genetic analyses reveal minimal variation attributable to vegetative spread.^[49]^[2] As the primary mode of reproduction, asexual fragmentation accounts for the majority of population growth and spread in stable aquatic environments, often enabling biomass to double every 3-5 days under optimal conditions such as moderate temperatures and adequate phosphorus availability. This high efficiency allows A. pinnata to outcompete other floating plants and cover large areas quickly.^[2]^[31] Fragmentation in A. pinnata is often triggered by environmental factors like plant crowding, which increases mechanical separation of stems, or nutrient stress, such as phosphorus limitation, that prompts vegetative dispersal to exploit new resources. These triggers enhance propagation rates during periods of high density or suboptimal conditions, facilitating survival and expansion without reliance on sexual phases.^[50]^[51]

Sexual reproduction

Azolla pinnata exhibits heterospory, producing microspores in microsporocarps and megaspores in megasporocarps on separate fronds, specifically arising from the ventral lobe of the first leaf on a branch. This process maintains the symbiosis with the cyanobacterium Anabaena azollae, as megaspores contain filaments of the symbiont that are transmitted to the next generation.^[2] Microsporocarps are larger and brownish, containing up to 30 microsporangia per cross-section, with each microsporangium forming 3-6 massulae that hold 32 to 64 microspores approximately 0.035 mm in diameter, featuring a triradiate mark on their surface.^[52]^[53] In contrast, megasporocarps are smaller and develop a single functional megaspore, about 1.5 mm by 1 mm, surrounded by nine float-corpuscles (six below and three above) that provide buoyancy, while eight initial megaspore mother-cells abort during development.^[52] Sporocarp development occurs after periods of vegetative growth and is triggered by environmental cues such as short photoperiods, high plant density, low temperatures in tropical regions, or stress from early summer heat in temperate areas, often observed post-rainy season in September-October with ripening by April.^[52]^[53] The sporocarps initially remain sunken within the fronds but eventually float to the surface for reproduction.^[52] Fertilization is water-mediated, with microspores germinating into male prothalli that release multiflagellated antherozoids, which swim to the female prothallus within the megaspore to reach archegonia (up to 30 or more per prothallus); however, natural success rates are low due to the rarity of synchronized sporulation and environmental challenges.^[52]^[53] Following fertilization, the zygote develops into an embryo within the megaspore.^[53] Germination of the fertilized megaspore occurs after subsequent rains, forming a prothallus that displaces the float-corpuscles, leading to the development of a new sporophyte gametophyte; this process takes 1-2 months to produce a plant comparable in size to one grown vegetatively.^[52]^[53] Fertilized megaspores exhibit high viability, remaining dormant and viable for over a year in dry conditions.^[53] Sexual reproduction plays a minor role in population maintenance compared to asexual fragmentation, but it facilitates long-distance dispersal through buoyant sporocarps carried by water currents, floods, or attached to birds and other animals.^[7]^[53]

Uses

Agricultural applications

Azolla pinnata has been utilized in Asian agriculture for centuries, with records indicating its application as a green manure in rice cultivation in Vietnam and China dating back over 1,500 years in China, with records from the 6th century AD in texts like the Qi-Min-Yao-Shu, and extensively used in Vietnam today.^[2] This symbiotic fern, hosting the nitrogen-fixing cyanobacterium Anabaena azollae, fixes atmospheric nitrogen, contributing to its value in traditional farming systems.^[42] As a green manure, A. pinnata is incorporated into rice paddies, typically providing 20–40 kg of nitrogen per hectare, which can increase rice yields by approximately 20%.^[54] This practice involves growing the fern in flooded fields before or alongside rice and plowing it under to release nutrients, thereby improving soil organic matter and reducing reliance on synthetic inputs.^[55] In biofertilizer integration, A. pinnata is employed in dual cropping with rice, where it forms a floating mat that suppresses weeds by up to 50–60% and reduces the need for chemical nitrogen fertilizers by 25–50%.^[56]^[57] This method not only enhances nitrogen availability but also minimizes ammonia volatilization from applied fertilizers, promoting more efficient nutrient use in paddy systems.^[58] A. pinnata serves as a high-protein animal fodder, containing 25–35% crude protein on a dry weight basis, suitable for poultry, livestock, and fish feed, where inclusion rates of 5–25% have been shown to improve feed conversion efficiency and animal growth performance.^[59]^[60] Its nutrient-rich profile, including essential amino acids and minerals, makes it a cost-effective supplement in mixed farming operations. In modern sustainable agriculture, A. pinnata is promoted for enhancing soil fertility in tropical regions, particularly in integrated rice-fish systems and low-input farming, supporting climate-resilient practices amid growing concerns over chemical fertilizer overuse.^[44] Its rapid biomass production and multifunctional benefits align with global efforts to foster eco-friendly crop intensification in Asia and beyond.^[42]

Environmental remediation

Azolla pinnata plays a significant role in environmental remediation through its capacity for bioaccumulating heavy metals from contaminated water bodies. The plant effectively absorbs lead (Pb), zinc (Zn), and cadmium (Cd), with accumulation levels ranging from 100 to 500 mg/kg dry weight in its tissues, depending on exposure concentration and duration. For instance, studies on Azolla species have reported up to 416 mg/kg for Pb and 259 mg/kg for Cd in fronds after exposure to polluted solutions, with A. pinnata showing similar accumulation potential (e.g., 310–740 mg/kg for Pb).^[61] This bioaccumulation process helps mitigate heavy metal toxicity in aquatic ecosystems, preventing their entry into the food chain. In addition to heavy metals, A. pinnata excels in adsorbing organic pollutants such as textile dyes and pesticides from wastewater. It achieves removal efficiencies of 70-90% for dyes like methylene blue and methyl violet, primarily through surface adsorption on its biomass and enzymatic degradation. For pesticides, including neonicotinoids like imidacloprid and fungicides like difenoconazole, removal rates in constructed systems reach up to 92%, facilitated by the plant's rapid biomass production and high surface area. These capabilities make A. pinnata a cost-effective biosorbent for treating industrial effluents.^[62]^[63] A. pinnata contributes to water purification by absorbing excess nutrients, thereby reducing eutrophication in nutrient-enriched waters. In constructed wetlands, it removes 25-38% of ammonia-nitrogen (NH₃-N), phosphate (PO₄³⁻), and nitrate (NO₃⁻) from agricultural runoff, promoting clearer water and preventing algal blooms. Its symbiotic association with nitrogen-fixing cyanobacteria enhances nutrient uptake efficiency. Furthermore, the plant's biomass sequesters 10-20 tons of CO₂ per hectare per year, supporting climate mitigation efforts through carbon fixation during growth.^[64]^[65] Recent research in the 2020s has explored A. pinnata in bioreactors and constructed wetlands for treating industrial effluents, such as palm oil mill and paper mill wastewater. These systems achieve substantial pollutant reduction, with heavy metal removals up to 88% for nickel and improved overall water quality, highlighting its scalability for sustainable remediation. As of 2025, ongoing research highlights A. pinnata's application in constructed wetlands for treating agricultural runoff, achieving up to 80% reduction in pesticide residues in some systems.^[66]^[67]^[42]

Management

Cultivation techniques

Cultivation of Azolla pinnata typically begins with inoculum preparation using fresh, healthy biomass to establish dense mats in controlled environments. A common approach involves inoculating shallow ponds, trays, or pits with 200–500 g/m² of fresh Azolla biomass, often sourced from established cultures, to achieve rapid coverage.^[42]^[68] For larger setups, such as 5 × 4 × 0.3 m pits lined with impermeable sheets and a 10–15 cm layer of soft soil, an initial dose of 5 kg of pure inoculum is applied, followed by filling with water to a depth of 7–11 cm.^[69] This method ensures pest-free starting material and promotes vegetative propagation, with the biomass doubling every 2–5 days under favorable conditions.^[70] Optimal growth requires temperatures between 20–30°C, as higher levels above 37°C can harm the plants, and a pH range of 5.5–7.0, with neutral pH being ideal for maximal biomass accumulation.^[70]^[69] Light exposure of 6–8 hours of full sunlight per day, or equivalent intensities of 1413–1561 lux, supports robust development, while relative humidity of 65–80% and long photoperiods further enhance doubling rates.^[42]^[69] Phosphorus supplementation is critical, typically at 0.5–1 mg/L or 20 kg/ha of superphosphate, to boost nitrogen fixation by the symbiotic cyanobacterium Anabaena azollae and increase phosphorus availability by 20–30%; no additional nitrogen fertilizers are needed due to the plant's autotrophic capabilities.^[42]^[68] Organic amendments like 15 kg of fermented buffalo feces every 15 days can also sustain nutrient levels in pit systems.^[69] Harvesting occurs every 7–15 days once the mat reaches saturation, using manual methods to skim the surface biomass without disturbing the water column.^[42] Yields typically range from 10–20 t/ha fresh weight or 3–9 t/ha dry matter annually, with weekly harvests producing up to 3.73 kg fresh and 172 g dry per pit under optimal conditions; this equates to approximately 1–2 kg dry matter/m² per month in high-productivity setups.^[42]^[70] Half of the harvested material is often reinoculated to maintain coverage, while the remainder supports agricultural applications like green manure.^[42] For scaling, A. pinnata is integrated into rice fields at 0.5–1 t/ha post-transplanting or grown in separate ponds using the half-saturation method, where initial densities of 300–500 kg/ha double weekly to cover larger areas.^[42]^[71] This approach is adaptable to low-labor systems, with the half-saturation technique originating in Vietnam allowing expansion from small plots to 10–20 t/ha saturated densities.^[42] Pest management involves neem extracts to control insects, alongside integrated systems like rice-Azolla-duck-fish models where ducks naturally suppress pests; conventional options include furadan granules at 100 g/plot applied seven days post-inoculation if needed.^[42]^[72] Key challenges include controlling pests such as snails and weevils, which can damage mats, and preventing fragmentation from wind or turbulent water; pits should be cleared and reinoculated if infestations occur.^[42]^[69] Post-2020 sustainable protocols emphasize organic farming integrations, such as reducing synthetic fertilizers by 25% through Azolla biofertilization and incorporating it into carbon-sequestering systems for climate resilience.^[70]^[42]

Control as invasive species

Mechanical removal is a primary strategy for managing small infestations of Azolla pinnata, involving raking, seining, or scraping the floating mats from water surfaces to physically extract the plant.^[73] This method is effective for localized outbreaks in ponds or ditches, as it directly reduces biomass without chemicals, but it is labor-intensive and requires frequent repetition due to the plant's rapid regrowth, potentially doubling coverage every 4-5 days under favorable conditions. Water level drawdown can complement raking by exposing and desiccating the fern during dry periods, though it may disrupt aquatic ecosystems if not carefully managed.^[28] Chemical control targets A. pinnata with aquatic herbicides applied at low concentrations to minimize environmental impact. Diquat, a contact herbicide, provides good control when applied at rates of 0.1-0.37 kg active ingredient per hectare, achieving up to 80% reduction in coverage within weeks by disrupting photosynthesis.^[73] Fluridone, a systemic herbicide, is highly effective at 10-30 parts per billion (0.01-0.03 ppm) in whole-water treatments from spring to mid-summer, inhibiting carotenoid synthesis and leading to plant death over 30-90 days, with efficacy exceeding 90% in enclosed systems. These applications must account for water flow and dilution to ensure sustained exposure, and follow-up treatments are often needed for regrowth.^[47] Biological control employs the weevil Stenopelmus rufinasus, a native North American herbivore specialized on Azolla species. This weevil has been introduced to manage invasive A. filiculoides in non-native regions such as South Africa, where releases since the 1990s have reduced Azolla biomass by up to 90% through larval and adult feeding on fronds, establishing self-sustaining populations that provide long-term suppression. It has also adopted A. pinnata as a host plant, with potential for use against it in invaded areas like North America, though establishment success varies with climate and initial densities. This agent is host-specific, minimizing risks to non-target species.^[74]^[75] Integrated management combines multiple approaches for larger-scale infestations, enhancing efficacy while reducing reliance on any single method. Strategies include shading to limit photosynthesis, nutrient reduction to slow growth, and physical barriers like booms to contain spread, often paired with targeted herbicide or biological releases; these programs emphasize early detection to prevent dense mat formation, which exacerbates control challenges due to the plant's rapid vegetative spread. As of 2025, A. pinnata is part of Early Detection Rapid Response (EDRR) efforts in Florida, including manual removal in waterways like Lake Okeechobee.^[76]^[77] Regulations play a key role in preventing A. pinnata proliferation, with the species listed as a Federal Noxious Weed in the United States, prohibiting its possession, transport, or sale across states like Alabama, North Carolina, and Vermont where it is classified as a Class A noxious weed.^[5] Similar restrictions apply internationally, including bans in parts of Canada under Ontario's Invasive Species Act effective January 1, 2024.^[30] Monitoring efforts utilize remote sensing, such as satellite imagery and normalized difference vegetation index (NDVI) analysis, to detect and track infestations at landscape scales, enabling timely interventions in wetlands and rivers.^[7]

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During their ontogeny, both hair types undergo a succession of transfer cell differentiation and senescence which begins in their terminal cells and ends in ...<|control11|><|separator|>
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Classification ; Kingdom Plantae ; Phylum Streptophyta ; Class Equisetopsida ; Subclass Polypodiinae ; Order. Salviniales. View Order Tree opens in a new tab.Missing: current | Show results with:current<|control11|><|separator|>
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Kingdom: Plantae ; Phylum: Filicinophyta ; Class: Filicopsida ; Order: Salviniales ; Family: Salviniaceae.
[25]
The East discovers Azolla | The Azolla Foundation
The earliest known written record of this practice is in a book written by Jia Ssu Hsieh (Jia Si Xue) in 540 A.D on The Art of Feeding the People (Chih Min Tao ...
[26]
Azolla growth in farm dams - Water - Agriculture Victoria
May 31, 2021 · Azolla can survive within a water pH range of 3.5 to 10, but optimum growth occurs in the pH range of 4.5 to 7 and temperature range of 18°C to ...
[27]
Azolla pinnata
Azolla pinnata is locally distributed in its native range of Africa and Madagascar, India, Southeast Asia, China and Japan, Malaysia and the Philippines.
[28]
Water Ferns - Invasive Species Centre
Common Names: Waterferns, water ferns, waterfern, mosquito fern, mosquitofern, crested mosquitofern, Carolina mosquitofern, eastern mosquitofern, ...<|separator|>
[29]
[PDF] Azolla pinnata - Aquatic Plant - Wisconsin DNR
Native North American Azolla weevil, Stenopelmus rufinasus (Coleoptera: Curculionidae), uses the invasive old world Azolla pinnata as a host plant. The ...
[30]
Invasive Plants of the Thirteen Southern States
Currently, there are 7 States with noxious weed laws and 8 States with invasive plant lists. ... Azolla pinnata R. Brown, feathered mosquito-fern, a, a, a.
[31]
[PDF] 15a ncac 02g .0602 noxious aquatic weed list - NC.gov
plants as noxious aquatic weeds: (1). Species Listed on the Federal Noxious Weed List. Azolla pinnata R. Brown - Pinnate mosquitofern. Eichhornia azurea (Sw ...Missing: Alabama | Show results with:Alabama
[32]
Azolla; Fairy Fern, Invader, and World Changer – Portsmouth ...
Azolla filiculoides is a small aquatic fern that floats on water, where it can form thick mats on the surface, capable of doubling in size in just a few days in ...<|separator|>
[33]
Optimal Growth Conditions for Azolla pinnata R. Brown
Apr 12, 2022 · The symbiotic relationship that all Azolla species have with the nitrogen-fixing bacteria Anabaena azollae allows the plants to live in nitrogen ...
[34]
Evaluating growth-dependent enhanced carbon dioxide ...
Evaluating growth-dependent enhanced carbon dioxide sequestration potential of Azolla pinnata using cattle wastes (cow dung and cow urine) ... doubling time of ...
[35]
Growth and Survival of Azolla filiculoides in Britain. I. Vegetative ...
forming rhizomes and turions which escape freezing in bottom sediment ... of Azolla pinnata R. Brown. Acta Botanic Indica 15: 196-201. Kannaiyan S ...Missing: dormancy | Show results with:dormancy
[36]
Occurrence of cytokinin-like compounds in two aquatic ferns and ...
Aug 7, 2025 · ... formation and dormancy of turions in U. ... Enhancement of growth and endogenous phytohormones of azolla pinnata in response to tryptophan.
[37]
Differential phosphorus requirements of Azolla species and strains ...
Mar 29, 2012 · A continuous-flow device for determining the minimum phosphorus level was developed. Azolla pinnata from Bangkok grew normally at 0.06 ppm P.
[38]
Growth and nutrient composition of Azolla pinnata R. Brown and ...
Growth and nutrient composition of Azolla pinnata R. Brown and Azolla filiculoides Lamarck as affected by water temperature, nitrogen and phosphorus supply, ...
[39]
Interspecific Variability in Growth Characteristics and ... - MDPI
Detaching of roots and changes in color were more pronounced in Azolla pinnata compared to other species, which might be due to the strength of Cu toxicity for ...Missing: flow | Show results with:flow
[40]
Developing Guidelines for Azolla microphylla Production as ... - MDPI
... Azolla pinnata in fresh and powder form as organic fertilizer on soil chemical properties, growth and yield of rice plant. AIP Conf. Proc. 2018, 1927 ...
[41]
Role of Azolla in sustainable agriculture and climate resilience
Azolla pinnata, native to Asia, Africa, and Australia, has been introduced to the USA and South America. Azolla filiculoides, tolerant of cold climates, was ...<|control11|><|separator|>
[42]
https://www.frontiersin.org/journals/plant-science/articles/10.3389/fpls.2025.1661720/full
[43]
Role of Azolla in sustainable agriculture and climate resilience
Oct 14, 2025 · Azolla has long been used in agriculture mostly for water conservation, weed control, and soil fertility enhancement. Its use as a biofertilizer ...
[44]
https://pmc.ncbi.nlm.nih.gov/articles/PMC12569520/
[45]
https://doi.org/10.1007/s11356-021-16986-6
[46]
Background on the Aquatic Herbicides Registered for Use in Florida
May 15, 2025 · ... (Azolla pinnata). Diquat is also applied alone or in combination with ... plants may be dormant. Cooler water slows microbial ...
[47]
Azolla pinnata Growth Performance in Different Water Sources
Azolla increased its population size (the number of plants) by fragmentation (similar to observations of Wagner, 1997; Lumpkin, 1987) within 3-4 days. The ...Missing: asexual apomixis
[48]
Ecological distribution and genetic diversity of Azolla in Uganda
Mar 8, 2023 · According to [29], Azolla. pinnata and Azolla nilotica were morphologically identified in Uganda in the regions of Kigezi, West Nile, Lake ...
[49]
(PDF) Effects of physical space and nutrients on the growth and ...
Aug 9, 2025 · We grew either one or four clonal fragments of a floating clonal fern, Azolla ... These fragmented or intact clones were grown under two nutrient ...
[50]
[PDF] A review of some ecological factors affecting the growth of Azolla spp.
(2006) Protection against salt toxicity in Azolla pinnata-. Anabaena azollae symbiotic association. Page 10. A review of some ecological factors affecting… 74.
[51]
[PDF] the structure and life-history of azolla - pinnata r. brown with remarks ...
They are green at first, but acquire a brownish colour in the cold season on account of a pigment formed in the leaves. The plants produced from the sporocarps ...
[52]
[PDF] Azolla-Anabaena symbiosis : its physiology and use in tropical ...
The growth rate is retarded as the plant density increases [2] . The maximum biomass or nitrogen accumulation reported by researchers is summarized in Table 2.
[53]
Utilization Azollapinnata as substitution of manure to improve ...
Nitrogen fixation potential of the Azolla anabaenasystem has been estimated to be 1.1 kg N ha-1 day-1 and one crop of Azollaprovided 20–40 kg N ha-1 to the rice.
[54]
An overview of underutilized benefits derived from Azolla as a ...
The use of Azolla pinnata as green manure significantly increases grain yields by 7.7% more compared to control [31,32]. The study carried out by Ref. [22] in ...Missing: velocity | Show results with:velocity
[55]
[PDF] Control of Weeds by Azolla in Rice.
Azolla dual cropping alone reduced the weed quantity by 50%, whereas green manuring plus dual cropping reduced weeds by 60% over uninoculated and un-.
[56]
[PDF] AN OVERVIEW OF AZOLLA IN RICE PRODUCTION
Dec 4, 2020 · Azolla dual cropping also helps in suppressing the weeds in ... and reduces 30-35 kg of nitrogenous fertilizer requirement of rice crop.
[57]
Rice production | The Azolla Foundation
[9] When grown in a rice field, azolla reduces the ammonia volatilization that occurs following the application of inorganic nitrogen fertilizers by 20% to 50%.Missing: dual | Show results with:dual
[58]
Livestock feed | The Azolla Foundation
Suitability of Azolla as a livestock feed On a dry weight basis, azolla has 25-35% protein content, 10-15% mineral content, and 7-10% comprising a combination ...
[59]
Azolla as a beneficial macrophyte for livestock feed: a review
Jun 19, 2024 · Azolla is a cheap and alternative source of protein that can improve feed conversion ratio (FCR), energy efficiency, and animal performance ...
[60]
Phytoremediation Potential of Aquatic Macrophyte, Azolla - PMC
The removal of heavy metal contaminants by aquatic macrophytes presents the problem of plant biomass disposal so as to prevent recycling of accumulated metals ( ...
[61]
(PDF) Azolla pinnata: An Efficient Low Cost Material for Removal of ...
Aug 7, 2025 · ... In our investigation, 88% removal was seen with absorbent dosages of 2, 4, and 8 g after a 60-minute contact period. Kooh et ...Missing: herbicides | Show results with:herbicides
[62]
Experiments on Pilot-Scale Constructed Floating Wetlands ...
Dec 15, 2022 · Four experimental CFWs were designed and constructed; three of them were planted with the aquatic plant species Lemna minor, Azolla pinnata and ...
[63]
Phytoremediation potential of Azolla pinnata on water quality and ...
In addition, turbidity and pH value diminished to 15.65% and 18.6%, respectively. However, it indicated that Azolla pinnata has the potential as an agent of ...
[64]
CO2 sequestration by propagation of the fast-growing Azolla spp - NIH
This is equivalent to 50,633 kg of Azolla (dry weight) per year, or 21,266 kg of carbon per year, per 1 ha. This means that to sequester the carbon dioxide ...
[65]
[PDF] Green Remediation Strategy Using Azolla pinnata: Heavy Metals ...
Oct 14, 2025 · Phytoextraction potential of water fern (Azolla pinnata) in the removal of a hazardous dye, methyl violet 2B: artificial neural network ...
[66]
Paper Mill Pollutants Reduction-Azolla pinnata - TSI Journals
In this present study, Azolla pinnata an aquatic plant was used to remove pollutants from effluent of paper mill. The constructed wetland study was conduct.
[67]
https://www.tsijournals.com/articles/paper-mill-pollutants-reductionazolla-pinnata.html
[68]
Azolla pinnata as unconventional feeds for ruminant feeding
Mar 14, 2022 · The optimum temperature for growing Azolla is 18–28 °C with a pH of 4.5–7 (Kathirvelan et al. 2015). Azolla (Azolla pinnata) is a good ...<|control11|><|separator|>
[69]
Azolla Plant Production and Their Potential Applications - Korsa - 2024
Jan 17, 2024 · In return, Anabaena azollae fixes atmospheric nitrogen and supplies Azolla with ammonia. So, lowland rice fields are a cost-effective ...
[70]
https://onlinelibrary.wiley.com/doi/10.1155/2024/1716440
[71]
https://doi.org/10.1007/s13593-017-0440-z
[72]
How to Control Mosquito Fern (Azolla) - AquaPlant
1. Physical Management Options. Mosquito fern can be removed by raking or seining it from the pond's surface. 2. Biological Control Options. Tilapia will ...
[73]
Azolla control - CABI.org
Azolla is one of the UK's most invasive plants. CABI is working on a biocontrol agent to help control the weed naturally.Missing: pinnata | Show results with:pinnata
[74]
Life history and laboratory host range of Stenopelmus rufinasus, a ...
The frond-feeding weevil, Stenopelmus rufinasus Gyllenhal, was imported into quarantine for testing as a potential natural enemy for the invasive fern Azol.
[75]
Feathered Mosquitofern - FWC
Feathered mosquitofern (Azolla pinnata) is a free-floating small aquatic fern that is distinguished from the native Azolla filiculoides by its triangular ...Missing: var. | Show results with:var.
[76]
feathered mosquito-fern (Azolla pinnata R. Brown) - Invasive.Org
Taxonomic Rank ; Kingdom: Plantae ; Phylum: Pteridophyta ; Class: Filicopsida ; Order: Hydropteridales ; Family: Salviniaceae.