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Equisetales

Equisetales is an order of vascular plants within the class Polypodiopsida, subclass Equisetidae, consisting solely of the family Equisetaceae and the genus Equisetum, which includes 18 extant species worldwide, commonly known as horsetails or scouring rushes.^[1]^[2] These perennial, herbaceous plants are distinguished by their jointed, hollow stems reinforced with silica deposits in the cell walls, reduced scale-like leaves arranged in whorls at the nodes, and a life cycle involving alternation of generations with homosporous reproduction via spores produced in terminal strobili (cones).^[3]^[2] Representing the only surviving lineage of a once-dominant group from the Carboniferous period around 325 million years ago, Equisetales are considered living fossils with a rich evolutionary history, having contributed significantly to ancient coal and oil deposits through their fossilized remains.^[3]^[2] The morphology of Equisetales is highly specialized for survival in challenging environments, featuring extensive underground rhizomes that facilitate vegetative propagation and radial expansion, often reaching depths of over 4 feet and growing up to 20 inches per season.^[2] Many species exhibit dimorphism, with separate fertile shoots bearing spore-producing cones in spring and sterile, photosynthetic shoots that resemble miniature conifers or cattails, growing to heights of 0.25 to 8 meters (10 inches to 26 feet) depending on the species, though most are under 2 meters.^[3]^[2] The stems' ribbed, rough texture, due to silica content, historically led to their use as natural abrasives, while their resistance to herbivory stems from this same mineralization, which is a unique nutritional requirement among plants.^[3] Ecologically, species of Equisetales thrive in moist, disturbed habitats such as wetlands, roadsides, drainage ditches, and forest edges, preferring poorly drained soils for spore germination but capable of encroaching into drier areas via rhizomes.^[2]^[3] Distributed globally across temperate and boreal regions, they are often colonizers of open, high-light areas and play roles in soil stabilization, though they can become weedy in agricultural settings due to their persistent rhizomatous growth.^[3] Reproduction primarily occurs vegetatively through rhizomes, with spore-based sexual reproduction involving bisexual gametophytes producing eggs and antherozoids in moist conditions, though spores play a minor role in contemporary spread.^[3]^[2] From an evolutionary perspective, Equisetales diverged early in fern history, with fossil records showing ancestors that reached heights of up to 18 meters during the Paleozoic era, dwarfing modern forms that rarely exceed 2 meters.^[3]^[4] Their monophyletic lineage is marked by distinct nodal and internodal stem regions, a trait conserved since the Devonian period, underscoring their status as relics of a diverse ancient flora now reduced to a single genus.^[3]

Overview

Definition and Characteristics

Equisetales is an order of vascular plants within the subclass Equisetidae of the class Polypodiopsida, commonly known as ferns, distinguished by their unique articulate growth form and ancient lineage tracing back to the Devonian period.^[5] The order comprises the single extant family Equisetaceae, represented solely by the genus Equisetum, which includes approximately 15 living species of herbaceous perennials typically inhabiting wet environments worldwide.^[6] These plants are homosporous, reproducing via spores rather than seeds, and exhibit a clonal growth strategy through extensive rhizomes that produce upright, jointed stems.^[7] The defining morphological traits of Equisetales include hollow, articulate stems segmented into nodes and internodes, featuring prominent longitudinal ribs and furrows that provide structural support. Leaves are greatly reduced to whorled, scale-like phylloids that fuse into a sheath at each node, serving minimal photosynthetic roles while the photosynthetic function is primarily handled by the green stems or whorled branches. A hallmark feature is the deposition of silica in epidermal cells, which encrusts the ribs and enhances stem rigidity, making them abrasive and historically useful for scouring. The vascular system is notable for its siphonostelic arrangement, with vascular bundles arranged in a ring; each bundle includes carinal canals—water-conducting channels formed by the disruption of protoxylem during internode elongation—and vallecular canals in the cortex.^[8] Reproduction occurs through terminal strobili bearing sporangia that release green, spherical spores equipped with four ribbon-like elaters, which facilitate dispersal by hygroscopic movement.^[7] While modern Equisetales are small herbaceous plants rarely exceeding 1-2 meters in height, the order's extinct diversity was far more varied, including arborescent forms such as Calamites from the family Calamitaceae, which reached up to 20 meters tall with secondary xylem for added support.^[6] In comparison to other fern orders like Polypodiales, Equisetales stand out due to their jointed stems, silica impregnation, and whorled microphylls, reflecting an early divergence within the monilophyte clade despite shared vascular plant ancestry. These traits underscore their "living fossil" status, with minimal morphological change over millions of years.^[6]

Historical Recognition

Early European botanists in the 16th to 18th centuries provided the initial scientific descriptions of horsetails, noting their distinctive jointed stems and bushy appearance that evoked the tail of a horse, from which the genus name Equisetum (Latin for "horse bristle") derives. John Ray, a prominent English naturalist, detailed several Equisetum species in his Historia Plantarum (1686–1704), emphasizing their hollow, ridged stems and whorled branches while classifying them among British flora without flowers, distinguishing them from typical seed plants.^[9] Ray also observed fossilized stems resembling modern horsetails during his travels in Italy, marking an early link between living and extinct forms.^[10] In 1789, Antoine Laurent de Jussieu advanced the taxonomic framework in his Genera Plantarum, placing Equisetum within the class Acotyledones and order Filices (ferns), based on shared vegetative and reproductive traits like spore-bearing structures, thereby recognizing its non-seed-producing nature and affinity to other cryptogams.^[11] The family name Equisetaceae was formalized shortly after by Augustin Pyramus de Candolle in 1804, encompassing the genus Equisetum. By the early 19th century, the order Equisetales was established by Friedrich von Berchtold and Jan Presl in 1820, integrating paleobotanical evidence of ancient relatives and expanding the group to include both extant herbaceous forms and fossil arborescent sphenopsids from the Paleozoic era.^[12] Paleobotanists in the 1820s, such as Adolphe-Théodore Brongniart, further solidified the recognition of fossil Equisetales through detailed comparisons of Carboniferous calamites and other sphenophytes to living horsetails, revealing shared features like nodal ribs and whorled appendages despite vast size differences.^[13] Initially, horsetails were sometimes misclassified as related to rushes (family Juncaceae) or even primitive grasses due to their rush-like, non-woody stems and lack of obvious flowers, leading to confusion in early herbals. This ambiguity was resolved in the mid-19th century through advancements in microscopy, which revealed homosporous spores and prothallia; Wilhelm Hofmeister's seminal 1851 study on Vergleichende Untersuchungen demonstrated the alternation of generations in Equisetum, confirming its pteridophyte affinities akin to ferns rather than monocots.^[14] By the 20th century, integrative studies of morphology, anatomy, and fossils positioned living Equisetales as "living fossils," with Equisetum representing the sole surviving lineage of a once-dominant group that peaked in diversity during the Carboniferous period, over 300 million years ago.^[15]

Morphology and Anatomy

Vegetative Structures

The vegetative structures of Equisetales are characterized by highly specialized stems, reduced leaves, and adventitious roots, which collectively support the plants' adaptation to wetland environments through enhanced mechanical strength and efficient water conduction. In living representatives of the genus Equisetum, the primary axis consists of a perennial, extensively branched underground rhizome that serves as the main vegetative organ, producing both upright aerial shoots and additional rhizomes in a monopodial branching pattern.^[16] These rhizomes are jointed, featuring distinct nodes and elongated internodes marked by longitudinal ribs that correspond to underlying vascular strands, providing structural rigidity.^[17] The aerial stems, which are photosynthetic and often green, exhibit similar jointed morphology with 6–40 ribs depending on species, and can reach heights of up to 8 meters in robust forms like Equisetum giganteum.^[18] A key adaptation in Equisetales stems is the impregnation of epidermal and cortical tissues with silica (SiO₂), which constitutes up to 25% of the dry weight and imparts abrasive texture and mechanical support against herbivory and environmental stress.^[19] This silicification occurs primarily in the outer walls of epidermal cells, enhancing stem stiffness while allowing flexibility in wind-prone habitats.^[20] Internally, the stems possess a unique vascular system with a central pith surrounded by a ring of collateral vascular bundles at each node, where the xylem forms a solid nodal plate.^[21] These bundles alternate between nodes, featuring protoxylem elements that develop into carinal canals—water-filled lacunae formed by the lysis of protoxylem during growth—and metaxylem regions that contribute to water transport efficiency.^[22] Vallecular canals, located in the cortex between ridges, further facilitate aeration and gas exchange in waterlogged soils.^[7] Leaves in Equisetales are greatly reduced and non-photosynthetic, functioning primarily as protective sheaths rather than major sites of carbon fixation, with photosynthesis occurring mainly in the chlorenchymatous cortex of the stems.^[21] Arranged in whorls at the nodes, these scale-like phylloids are fused at their bases into toothed sheaths that encircle the stem, with free tips forming small, triangular lobes; in Equisetum, there are typically 6–48 leaves per whorl, varying by species and stem type.^[17] This reduction reflects an evolutionary shift toward stem-dominated resource allocation, minimizing transpiration while shielding nodal buds and vascular tissues.^[23] Roots in Equisetum are adventitious, arising in whorls from the rhizome nodes and lacking a primary root system, which enables extensive horizontal spread and colonization of moist substrates.^[16] These roots are slender, dichotomously branched, and often form mycorrhizal associations with arbuscular fungi (Glomales), enhancing nutrient uptake—particularly phosphorus—in nutrient-poor, acidic soils typical of horsetail habitats. Root anatomy mirrors that of stems, with a stele featuring endarch xylem maturation and similar canal systems for water and oxygen movement.^[24] Branching in Equisetales vegetative structures is predominantly whorled at nodes in extant Equisetum, where lateral branches emerge from the axils of leaf sheaths in a helical pattern, often with reduced internodes near the base for compact growth.^[17] In contrast, fossil Equisetales, such as those in the Carboniferous genus Calamites, display dichotomous branching patterns in higher-order laterals, reflecting greater architectural diversity before the group's decline.^[7] This vascular arrangement, with protoxylem and metaxylem canals aligned along ribs, supports efficient hydraulic conductivity and mechanical stability across both living and extinct taxa.^[22]

Reproductive Structures

The reproductive structures of Equisetales are specialized for spore production and are primarily organized into terminal strobili, which are compact, cone-like aggregations borne on fertile shoots. These strobili consist of a central axis with successive whorls of sporangiophores, exhibiting determinate growth and lacking the node-internode differentiation typical of vegetative stems.^[25] In extant Equisetum species, strobili are typically 1–5 cm long and develop from an apical meristem that transitions to reproductive mode, producing a stack of modified phytomers where sporangiophores arise from periclinal divisions across the full thickness of each level.^[26] Sporangiophores within the strobili are peltate, umbrella-shaped structures arranged in whorls, each with a short stalk and a hexagonal or rounded disc that bears 5–10 elongated sporangia pendent from its underside near the periphery.^[27] These sporangiophores mature acropetally, originating as bulges on the flanks of the strobilus apex, and function as modified, condensed branch systems adapted for sporangium support.^[25] The sporangia themselves are eusporangiate, cylindrical sacs with a two-layered wall; the outer layer features helical thickenings that facilitate longitudinal dehiscence upon maturity, releasing spores in a controlled manner.^[28] Spores produced in the sporangia are homosporous, green due to chloroplasts, and measure approximately 30–60 μm in diameter, featuring a multi-layered wall with an outer epispore that splits into four hygroscopic, ribbon-like elaters banded for enhanced dispersal.^[29] These elaters respond to humidity changes, coiling in moist conditions and extending in dry air to aid in spore liberation and movement. Many Equisetum species exhibit shoot dimorphism, with distinct fertile and sterile forms; fertile shoots are often unbranched, tan-colored, and non-photosynthetic, emerging early in the season to bear strobili at their apices, while sterile shoots are green, branched, and photosynthetic.^[30] This separation optimizes resource allocation for reproduction, as seen in species like Equisetum arvense.(https://milnepublishing.geneseo.edu/suny-geneseo-botany/chapter/02:_Organisms/2.25:_Horsetails_the_genus_Equisetum) In extinct Equisetales, such as the calamitacean genus Calamostachys from the Pennsylvanian period, strobili were larger and more complex, often bracteate with whorls of sporangiophores on arborescent axes, contrasting the compact, bractless form in modern herbaceous Equisetum. These fossil variations, including genera like Palaeostachya, suggest an evolutionary reduction in strobilus size and complexity over time.^[31]

Reproduction and Life Cycle

Alternation of Generations

Equisetales display a heteromorphic diplohaplontic life cycle, featuring a prominent, independent diploid sporophyte phase that dominates the plant's observable form and a reduced, short-lived haploid gametophyte phase.^[32] The sporophyte generation includes articulate stems, whorls of reduced scalelike leaves, extensive rhizomes, and branching roots, enabling perennial growth and vegetative propagation in moist environments.^[3] This phase produces haploid spores through meiosis in terminal strobili, marking the transition to the gametophyte.^[3] The gametophyte in Equisetum is green and photosynthetic, typically developing as small, thalloid structures composed of flat lamellae or, in some species, a more tuberous form adapted to subterranean conditions; they are usually bisexual but can develop unisexual forms depending on environmental conditions like soil moisture.^[21]^[33] These gametophytes are free-living but brief, often measuring just a few millimeters in diameter, and bear both antheridia that release multiflagellated sperm and archegonia that house eggs. Archegonia generally mature before antheridia on the same thallus, facilitating self-fertilization in moist soil.^[21] Equisetales are homosporous, releasing a single type of spore from the sporophyte that germinates directly into a bisexual gametophyte, unlike the heterospory seen in certain ferns where distinct microspores and megaspores produce unisexual gametophytes.^[32] Fertilization occurs when sperm from antheridia swim to nearby eggs within archegonia under damp conditions, forming a diploid zygote.^[3] The young sporophyte develops from this fertilized egg embedded on the gametophyte surface, with the embryo exhibiting a well-defined foot for nutrient uptake, an elongating seta, and the initial rhizome primordium that establishes the underground axis for further growth.^[34] As the sporophyte matures and becomes independent, the gametophyte typically withers, completing the alternation back to spore production.^[32]

Spore Production and Dispersal

Sporogenesis in Equisetales occurs within the sporangia of strobili, where diploid sporocytes directly undergo meiosis without an archesporial stage, producing tetrahedral tetrads of haploid spores through two sequential divisions.^[35] During meiosis I, organelles such as plastids and mitochondria form a peripheral band around the metaphase plate, redistributing to daughter cells in meiosis II to ensure balanced inheritance in the resulting spores.^[35] A plasmodial tapetum surrounds the developing tetrads, providing nutrients before disintegrating to liberate the mature spores, which measure approximately 50 μm in diameter and contain chloroplasts.^[35] Each spore develops four ribbon-like elaters attached at the proximal pole; these hygroscopic appendages consist of two fibrillar layers—an inner longitudinal and an outer transverse—that twist and untwist in response to humidity changes, facilitating spore separation from the tetrad.^[35] Spore dispersal is primarily wind-mediated, initiated by the dehiscence of the sporangium, where a ring of thickened cells analogous to an annulus contracts sharply upon drying, ejecting clusters of spores from the strobilus.^[36] The elaters play a crucial role post-ejection by preventing spores from clumping together through their twisting motions, which also enable localized dispersal via "walking" steps—small, random displacements driven by humidity cycles—or sudden "jumps" when the elaters thrust the spore after folding tightly in dry conditions. These movements enhance ejection efficiency and allow spores to reorient or escape grounded positions, increasing the likelihood of wind capture for broader distribution. Equisetales spores exhibit short viability, typically remaining capable of germination for only a few days after release under natural conditions, though some can persist up to two weeks in a desiccated state.^[37] Germination requires moist soil, where the spore wall ruptures, and the prothallial cell divides to initiate the bisexual gametophyte.^[38] Fossil evidence reveals that elater-bearing spores, similar to those in modern Equisetum, appeared in sphenophyte clades by the Upper Carboniferous, as seen in genera like Elaterites, where trilete spores possessed three-layered elaters attached via a distal truss, underscoring the ancient origin of this dispersal adaptation within Equisetales.^[39] Ultrastructural comparisons show shared features in elater layering and composition between these fossils and extant forms, indicating evolutionary continuity despite morphological variations in attachment and number.^[39]

Taxonomy and Phylogeny

Living Taxa

The living taxa of Equisetales are restricted to a single family, Equisetaceae, which contains only one genus, Equisetum, with no other extant genera recognized.^[40] This genus comprises 18 accepted species worldwide, all of which are perennial plants characterized by extensive rhizomatous growth and dimorphic or monomorphic aerial stems that arise from underground tubers or rhizomes.^[41] The species exhibit a range of morphologies, from small, branched forms to tall, unbranched ones, adapted to wetland and moist terrestrial habitats. Within Equisetum, two subgenera are traditionally recognized: subgenus Equisetum and subgenus Hippochaete. Subgenus Equisetum includes species with typically branched, green vegetative stems and rounded cone apices, such as E. arvense (field horsetail), a widespread species with dimorphic stems—the fertile ones tan and unbranched, reaching up to 40 cm, while sterile stems are green, branched, and up to 60 cm tall.^[42] Another notable member is E. telmateia (giant horsetail), one of the tallest species in temperate regions, with sterile stems up to 2.4 m high, persistent and branched, featuring whorls of needle-like leaves fused into sheaths. Subgenus Hippochaete consists of species with unbranched, evergreen stems and pointed cone apices, including E. myriochaetum (Mexican giant horsetail), the tallest species in the genus reaching up to 7 m, and E. hyemale (scouring rush), which has rough, silica-rich stems up to 1.5 m tall used historically for polishing. Infrageneric taxonomy remains subject to debate, particularly regarding species delimitation, with morphological variations often influenced by environmental factors and frequent hybridization complicating boundaries.^[40] Genetic studies using plastid loci have supported the distinction of certain taxa, such as E. praealtum from E. hyemale in subgenus Hippochaete, while questioning the status of others like E. bogotense, potentially warranting a third subgenus, Paramochaete.^[15] Despite these discussions, the core division into two subgenera and 18 species holds in most contemporary classifications as of 2025.^[41] The distribution of living Equisetum species is primarily Holarctic, spanning temperate and boreal regions of North America, Europe, and Asia, with some southern extensions into montane areas of South America (e.g., E. bogotense in the Andes) and Africa.^[40] All species are perennial, relying on persistent rhizomes for survival and propagation in moist environments.^[40]

Phylogenetic Relationships

Equisetales are classified within the Polypodiopsida, the class encompassing ferns and their allies, where they form the monophyletic subclass Equisetidae, consisting solely of the order Equisetales and the genus Equisetum.^[31] Phylogenetic analyses based on molecular data have variably positioned Equisetales as sister to all other monilophytes or as sister to Ophioglossales plus Psilotales, with Marattiales emerging as sister to the remaining monilophytes in multi-gene datasets.^[43]^[44] Molecular evidence from chloroplast genes, such as rbcL, trnL-F, and complete plastid genomes, alongside nuclear loci, robustly supports Equisetum as the sole extant representative of the ancient Sphenopsida clade, a lineage once diverse but now reduced to this genus.^[45]^[46] These datasets highlight the deep divergence of Equisetales within monilophytes, with no evidence of close affinities to lycophytes or seed plants.^[47] Key morphological synapomorphies defining Equisetales include the reduction of leaves to small, scale-like sheaths fused around the stem, the presence of elaters on spores that aid in dispersal, and secondary xylem tracheids featuring scalariform wall pitting.^[48] These traits distinguish Equisetales from other fern groups and align with their basal position in monilophyte phylogenies.^[43] Post-2010 studies, incorporating phylogenomic approaches with low-copy nuclear genes and expanded plastid data, have confirmed the Devonian origins of Equisetales through calibration with fossil records and underscore their isolation, with no close living relatives beyond the monilophyte ferns.^[46]^[31]

Evolutionary History

Origins in the Devonian

The origins of Equisetales, commonly known as sphenopsids, trace back to the Devonian period, with the earliest fossils appearing in the Late Devonian around 375 million years ago.^[49] These primitive forms characterized by dichotomous branching and the emergence of whorled appendages, representing transitional vascular plants between simpler rhyniophytes and more advanced lineages. Such fossils, preserved in formations like those of Germany and New York, indicate the initial diversification of upright, branching axes in early land plant communities.^[50] By the Late Devonian, more definitive Equisetales fossils emerged, with key genera such as Pseudobornia regarded as potential ancestors. Pseudobornia ursina, documented from sites in Spitsbergen and North America, exemplifies an early monopodial, arborescent form with clonal growth via rhizomes, marking the onset of sphenopsid morphology.^[16] Articulate stems, defined by distinct nodes and internodes, first appeared in these Late Devonian taxa, enabling modular growth and resilience in variable environments.^[31] Significant evolutionary innovations during this period included the initial development of stem articulation, which facilitated repeated branching and elongation. These adaptations coincided with a transition from leafless, isotomous axes to microphyllous forms, where small, scale-like leaves in whorls improved light capture without compromising vascular efficiency. Such changes positioned Equisetales as key contributors to Devonian terrestrialization.^[31] Paleoenvironments for these early Equisetales spanned aquatic to semi-terrestrial settings within expansive wetlands, as evidenced by fossils from floodplain and coastal plain deposits in Euramerica and Gondwana. These habitats, characterized by high humidity and periodic flooding, supported the proliferation of sphenopsids alongside lycophytes and cladoxylopsids, fostering the evolution of root-like rhizomes for anchorage in unstable substrates.^[50]

Diversity and Decline

Equisetales achieved their greatest diversity during the Carboniferous period, around 300 million years ago, when arborescent forms such as Calamites dominated wetland forests and grew to heights of up to 20 meters.^[51] These tree-like sphenopsids coexisted with smaller taxa like sphenophylls, and the group included diverse families such as Archaeocalamitaceae, which featured early-branching lineages with distinct stem and leaf anatomies preserved in Mississippian deposits.^[52] This peak reflects the order's adaptation to humid, tropical environments following their initial emergence from Devonian precursors.^[53] Diversity declined sharply during the Permian, with calamitacean lineages experiencing a drastic reduction, although some arborescent forms persisted locally into the Late Permian.^[7] The Permian-Triassic mass extinction exacerbated this trend, leading to the complete disappearance of tree-like species such as Calamites by the Early Permian, while herbaceous representatives like Neocalamites survived into the Mesozoic as common elements in Triassic floras of Gondwana and elsewhere.^[54]^[55] Throughout the Cenozoic, Equisetales underwent further reduction, with fossils attributable to the genus Equisetum appearing in Eocene strata and becoming more prevalent in Miocene deposits, where they closely resembled extant forms in stem structure and habitat preferences.^[56]^[57] This ongoing diminishment, driven by competition from diversifying seed plants in varied terrestrial habitats and climate shifts tied to global cooling and aridification, resulted in the extinction of nearly all lineages, leaving only Equisetum as the surviving genus.^[58]^[57]

Distribution and Ecology

Global Range

Equisetum, the sole surviving genus of Equisetales, comprises approximately 15 extant species with a subcosmopolitan distribution, predominantly in the Holarctic realm but with disjunct populations in the Southern Hemisphere. The genus is widespread across North America, Europe, and Asia, where it occupies temperate and boreal zones, extending northward to the Arctic Circle and southward into montane regions. In the Southern Hemisphere, occurrences are more fragmented, including southern South America, scattered sites in Africa, and isolated populations in Australasia and oceanic islands such as the Galápagos, Mascarenes, and Fiji, though it is absent as a native from Australia and New Zealand.^[59] Species richness peaks in the temperate zones of the Northern Hemisphere, reflecting historical patterns of expansion following the Pleistocene glaciations. For instance, North America hosts around 10 species, including widespread taxa like Equisetum arvense and E. hyemale, while Europe supports a similar number, such as E. palustre and E. telmateia. In contrast, tropical regions harbor fewer species, with only about four native to South America (E. bogotense, E. giganteum, E. myriochaetum, and E. ramosissimum), underscoring a bias toward cooler climates.^[40]^[59]^[60] The biogeographic patterns of Equisetum trace back to vicariance events associated with the breakup of Pangaea, with post-glacial recolonization driving recent expansions in the Northern Hemisphere during the Holocene. Ancient Gondwanan elements are evident in southern lineages, particularly subgenus Paramochaete, which includes high-elevation species in the Andes and disjunct tropical distributions suggestive of long-distance dispersal. Human-mediated introductions have further altered ranges, notably E. arvense as an invasive species in New Zealand and parts of South America.^[59]^[61]

Habitat and Ecological Interactions

Equisetales, solely represented today by the genus Equisetum, predominantly occupy moist and disturbed terrestrial habitats across temperate, subarctic, and tropical regions. These plants favor wetlands, streambanks, lake shores, seepage areas, meadows, marshes, and wet forest edges, often in poorly drained or waterlogged soils that retain high moisture levels. Many species exhibit remarkable tolerance for acidic, nutrient-poor, and compacted substrates, enabling persistence in marginal environments like roadsides, ditches, and areas with low fertility or high metal content. For instance, Equisetum arvense thrives in acidic conditions with pH levels below 6, serving as a reliable bioindicator of such soils.^[40]^[62]^[63]^[64] Key adaptations facilitate their success in these dynamic settings. Extensive rhizomatous systems, extending up to 1.2 meters deep and spreading laterally over several meters, enable rapid vegetative colonization of disturbed or open ground, often outpacing competitors in early successional stages. The accumulation of silica (SiO₂) in epidermal cells, reaching concentrations of 10–25% dry weight, imparts mechanical rigidity to stems and acts as a physical deterrent to herbivores by increasing tissue abrasiveness and reducing palatability. Furthermore, aerenchyma tissues in roots and stems allow oxygen diffusion in anoxic wetland soils, while efficient ion uptake supports growth in low-nutrient or saline conditions, as seen in Equisetum giganteum across Andean wetlands with varying salinity.^[2]^[64]^[65]^[66] Ecologically, Equisetales function as pioneer species in habitat succession, rapidly invading post-disturbance sites such as burned areas or eroded slopes to stabilize soil and facilitate later colonization by other vegetation. Their interactions with biota are limited by inherent toxicities, including thiaminase enzymes that degrade vitamin B1 and alkaloids like nicotine in some species (e.g., Equisetum palustre), which restrict consumption by most livestock and many vertebrates despite occasional foraging by tolerant animals such as greater snow geese (Anser caerulescens) and black bears (Ursus americanus). Insect herbivory occurs on select species like E. arvense, involving specialized feeders including beetles, sawflies, and flies, but overall diversity of attackers remains low compared to angiosperms. Mycorrhizal associations with arbuscular and dark septate fungi enhance nutrient acquisition, though symbiotic nitrogen-fixing partnerships with bacteria, while documented in rhizomes of species like E. arvense, are infrequent and contribute minimally to nitrogen budgets relative to other wetland plants.^[64]^[67]^[68]^[69] Climate dynamics significantly influence their persistence, with populations declining in aridifying landscapes due to reduced soil moisture availability, as moist conditions are essential for spore germination and rhizome viability. In contrast, they proliferate in persistently wet, stable wetlands, where species like E. pratense indicate facultative wetland status and overall ecosystem health through sensitivity to hydrological alterations and acidification.^[64]^[70]

Human Uses and Conservation

Traditional and Modern Applications

Equisetales, particularly species in the genus Equisetum, have been utilized by various cultures for their abrasive properties and medicinal potential. Native American groups, such as the Gwich'in, have traditionally prepared teas from Equisetum to address kidney problems, bladder infections, and urinary tract issues.^[71] In ancient Roman and Greek medicine, as well as among Native American and Chinese practitioners, Equisetum arvense was employed to treat a range of ailments, including urinary disorders.^[62] European herbal traditions similarly valued its diuretic effects for managing edema and kidney conditions, often in the form of infusions from E. arvense.^[72] The stems' high silica content imparts an abrasive texture, leading to historical uses as scouring tools for cleaning pots, polishing metal, and smoothing wood across European and Native American practices.^[51]^[73] In modern contexts, Equisetum extracts are incorporated into supplements for their silica content, which supports bone and connective tissue health by enhancing collagen formation and mineralization.^[74]^[75] Studies indicate anti-inflammatory properties in E. arvense, potentially aiding conditions like arthritis and wound healing through flavonoid and phenolic compounds, though clinical evidence remains preliminary.^[76]^[77] These supplements are marketed for benefits to hair, nails, skin, and joint health, but consumption requires caution due to thiaminase enzyme in fresh plants, which degrades vitamin B1 (thiamine) and may cause deficiency in large amounts, particularly in animals like horses.^[78]^[79] Processed dried forms mitigate this risk, but long-term use in humans should be monitored, and no strong evidence links Equisetum to carcinogenicity in typical medicinal doses.^[79] Industrially, Equisetum stems serve as natural abrasives in polishing and cleaning applications, leveraging their siliceous ridges.^[80] Certain species, like E. hyemale, are woven into baskets, mats, and crafts due to their sturdy, jointed structure.^[81] Historical attempts to use Equisetum as fodder were limited by its toxicity, causing thiamine deficiency and weight loss in livestock when consumed in quantity.^[82] Ornamentally, E. hyemale (scouring rush) is cultivated in gardens for its evergreen, architectural stems, valued in wetland or modern landscape designs.^[83]

Conservation Challenges

Equisetales, commonly known as horsetails, face several anthropogenic threats that impact their wetland and riparian habitats. Habitat loss due to drainage for agriculture and urban development is a primary concern, as many species rely on moist environments that are increasingly converted for human use. For instance, wetland drainage in Europe has degraded suitable sites for Equisetum species, leading to population declines in affected areas. Additionally, competition from invasive species, such as Phragmites australis, can outcompete native horsetails in altered wetlands, particularly for rarer taxa like Equisetum scirpoides. Pollution, including heavy metal contamination, poses another risk; horsetails bioaccumulate metals like cadmium, lead, and zinc from polluted soils and water, potentially stressing populations in industrialized regions despite their tolerance. While most Equisetum species are assessed as Least Concern globally by the IUCN Red List, local declines highlight vulnerabilities for certain taxa. Equisetum variegatum, for example, is endangered in the Parisian region of France due to habitat fragmentation and small population sizes. Similarly, Equisetum scirpoides is listed as a species of special concern in Massachusetts, USA, and extremely rare in Connecticut, where local extirpations have occurred from habitat alteration. Equisetum pratense experiences regional rarity, such as in parts of New England, though it remains secure overall. These cases underscore that while the order is resilient, isolated populations are at higher risk of extinction without intervention. Conservation efforts for Equisetales emphasize wetland protection and restoration. Many horsetail habitats are safeguarded within Ramsar wetland sites, such as those in Austria's Tyrolean mires, where Equisetum species contribute to mire ecosystems protected under international conventions. Restoration projects in degraded wetlands, including rehydration and invasive species removal, have supported recovery of Equisetum populations in North American fens. Research on genetic diversity is ongoing, particularly for threatened species like Equisetum variegatum, where propagation from extant genotypes is recommended to preserve variability and aid reintroduction. Climate change exacerbates these challenges through potential range shifts driven by altered hydrology and temperature; rising temperatures may expand suitable areas for some species but threaten others via drying wetlands, with monitoring efforts incorporating citizen science platforms to track distributions.

References

[1]
Equisetales - Explore the Taxonomic Tree | FWS.gov
Location in Taxonomic Tree ; Division, Tracheophyta ; Subdivision, Polypodiophytina ; Class, Polypodiopsida ; Subclass, Equisetidae ; Order, Equisetales ...
[2]
Equisetum: Biology and Management
Horsetails are members of the genus Equisetum, the only genus in the family Equisetaceae. There are 15 species of equisetum found worldwide; field horsetail ...Missing: Equisetales | Show results with:Equisetales
[3]
Horsetails, the genus Equisetum – Inanimate Life - Milne Publishing
Horsetails, the genus Equisetum, are a very easily recognized group of plants that are commonly found throughout the world. They represent a very small remnant ...
[4]
https://www.britannica.com/plant/Pseudobornia-ursina
[5]
https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&anchorLocation=SubordinateTaxa&credibilitySort=Subordinate%20Taxa&rankName=Species&search_value=846529&print_version=SCR&source=from_print
[6]
https://www.sciencedirect.com/science/article/pii/S0034666721001275
[7]
Equisetales - an overview | ScienceDirect Topics
The Equisetales are characterized by the presence of distinct nodes, internodes, and longitudinal internodal ridges on their stems.
[8]
https://doi.org/10.1093/aob/mcr161
[9]
[PDF] The Historia Plantarum Generalis of John Ray: Book I
Ray's final classification system as given in the Met hodus Plantarum emendata et aucta is given in the introduction to the notes to Chapter 27 of this.
[10]
John Ray - Rocky Road - Strange Science
In a few cases, Ray found fossil identification straightforward. When he saw the fossil stems of Equisetum (giant, extinct relatives of horsetails) in Italy, he ...
[11]
Antonii Laurentii de Jussieu Genera plantarum
The Biodiversity Heritage Library works collaboratively to make biodiversity literature openly available to the world as part of a global biodiversity ...
[12]
The first fossil record of a giant horsetail (Equisetum, Equisetaceae ...
Order Equisetales de Candolle ex Bercht. & Presl., 1820. Family Equisetaceae ... Phylogenetics, classification and typification of extant horsetails (Equisetum, ...
[13]
[PDF] sketch of paleobotany. - USGS Publications Warehouse
Paleobotany is a science of the nineteenth century. Nevertheless its dawn at the beginning of this century was preceded by a long fading twilight extending ...
[14]
Trends and concepts in fern classification | Annals of Botany
Feb 13, 2014 · Ferns are vascular plants that produce spores and undergo an alternation of generations (with separate gametophyte and sporophyte generations ...
[15]
None
Nothing is retrieved...<|control11|><|separator|>
[16]
Biology and Functional Ecology of Equisetum with Emphasis on the ...
Jan 15, 2013 · without mycorrhizal associations (Read et al., 2000). For examp found no mycorrhizal colonization of Equisetum roots in the they studied ...
[17]
Equisetum - an overview | ScienceDirect Topics
The Equisetales are characterized by the presence of distinct nodes, internodes, and longitudinal internodal ridges on their stems. Branches and leaves are ...
[18]
Roles of silica and lignin in horsetail (Equisetum hyemale), with ...
Feb 29, 2012 · The silica content in horsetail amounts to 25% of the dry weight and is derived from silicic acid from the soil,8 and a part of deposition ...
[19]
New insight into silica deposition in horsetail (Equisetum arvense)
Jul 29, 2011 · The horsetails (Equisetum sp) are known biosilicifiers though the mechanism underlying silica deposition in these plants remains largely unknown.
[20]
[PDF] VIII. Equisetophytes: The Horsetails - A. Equisetum
Horsetails, in the genus Equisetum, are the only living Equisetophytes, with fused leaves in whorls, and sometimes branches. They have nodes with leaf sheaths.
[21]
An extensin-rich matrix lines the carinal canals in Equisetum ...
Jul 12, 2011 · Previous studies on the carinal canals in several Equisetum species suggest that they convey water from one node to another.
[22]
Horsetails get the wind up - Raven - 2009 - Wiley Online Library
Sep 2, 2009 · They function in the movement of gases in photosynthetic structures of land plants and in the movement of oxygen to below-ground structures ...
[23]
Anatomy of Equisetum (With Diagram) | Pteridophyta
In this article we will discuss about the anatomy of equisetum. Also learn about its spore-producing organs.
[24]
Developmental programmes in the evolution of Equisetum ... - NIH
Equisetales are characterized by whorled, usually single-veined leaves that may be basally fused forming a sheath, and fertile regions consisting of one or more ...
[25]
https://pmc.ncbi.nlm.nih.gov/articles/PMC5458719/
[26]
Spore size In the genus Equisetum | Request PDF - ResearchGate
Aug 9, 2025 · Cones consist of whorls of peltate sporangiophores bearing approximately 10 sporangia each. Spores have a perispore and four elaters with ...
[27]
Equisetum: Habitat, Structure and Reproduction - Biology Discussion
In Equisetum, silica is deposited on the outer wall of the ... The sporophytic plant body of Equisetum is differentiated into stem, roots and leaves (Fig.<|control11|><|separator|>
[28]
The walk and jump of Equisetum spores - PMC - PubMed Central
Equisetum plants (horsetails) reproduce by producing tiny spherical spores that are typically 50 µm in diameter. The spores have four elaters, which are ...
[29]
Equisetum arvense - USDA Forest Service
GENERAL BOTANICAL CHARACTERISTICS : Field horsetail is a native, perennial, rhizomatous cryptogam. The sporophyte is dimorphic with unbranched, fertile ( ...
[30]
Origin of Equisetum: Evolution of horsetails (Equisetales) within the ...
Jul 19, 2018 · We provide a comprehensive phylogenetic analysis of Equisetales, with special emphasis on the origin of genus Equisetum.
[31]
Life History and Ecology of the Sphenophyta
Sphenophytes show the same pattern of alternation of generations found in all plants: asexually produced spores give rise to gametophytes, which produce eggs ...
[32]
[PDF] The Early Tracheophytes - PLB Lab Websites
The resulting zygote develops into an embryo that possesses a well-developed foot, a short primary root, leaf primordia, and a shoot apex (Fig. 23.9i). The ...<|control11|><|separator|>
[33]
Esporogénesis y esporas de Equisetum bogotense (Equisetaceae) de las áreas montañosas de Colombia
### Summary of Sporogenesis in *Equisetum bogotense*
[34]
[PDF] Fabulous Ferns - UNCW
Equisetophyta: Equisetum. Equisetophyta. Equisetopsida. Equisetales ... dehiscence mechanism. – Ring of thickened cells. – Contract sharply when dry.
[35]
Desiccation-time limits of photosynthetic recovery in Equisetum ...
The chlorophyllous spores of Equisetum survive desiccation, yet cannot tolerate this quiescent state for more than ~2 wk. The hypothesis that spore ...
[36]
Comparative ultrastructure of the sphenophyte spores Elaterites and ...
Aug 8, 2025 · Elaters possess three distinct layers. A comparison of the spores of Elaterites and Equisetum reveals a number of morphological and ...
[37]
Equisetum - FNA - Flora of North America
Sep 23, 2025 · Equisetum, also known as horsetail, has hollow stems with whorled leaves, found in moist places. Its name refers to its coarse black roots.
[38]
Equisetum arvense - FNA - Flora of North America
Mar 1, 2024 · Equisetum arvense var. boreale (Bongard) Ruprecht has been most generally accepted and has been applied to plants with tall, erect stems with 3-ridged branches.
[39]
Horsetails are the sister group to all other monilophytes and ...
A nine-gene data set and an indel in rpl2 place horsetails (Equisetales) are sister to all other monilophytes (ferns). Marattiales are the sister order to the ...
[40]
Complete plastid genomes from Ophioglossum californicum ...
Jan 11, 2013 · We sequenced the plastid genomes from three early diverging species: Equisetum hyemale (Equisetales), Ophioglossum californicum (Ophioglossales), and Psilotum ...Missing: microscopy | Show results with:microscopy
[41]
Molecular phylogeny of horsetails (Equisetum) including chloroplast ...
Equisetum is a genus of 15 extant species that are the sole surviving representatives of the class Sphenopsida. The generally accepted taxonomy of Equisetum ...Missing: survivor evidence
[42]
The evolutionary history of ferns inferred from 25 low‐copy nuclear ...
Jul 1, 2015 · Over the past two decades, molecular phylogenetic approaches have fundamentally altered our understanding of the evolution of ferns. These ...Discussion · The Fern Tree Of Life · Appendix 1
[43]
Chloroplast genome structure analysis of Equisetum unveils ... - NIH
Apr 11, 2024 · The phylogeny position of Equisetum has traditionally been determined using morphological characters, as well as various chloroplast segments ...
[44]
Phylogenetic diversification of Equisetum (Equisetales) as inferred ...
Jul 1, 2009 · Likewise, there are five carinal canals, five vallecular canals, and five leaves that make up each stem segment (Figs. 3, 4). Stems measure 1.2– ...
[45]
Early Fossil Record of Sphenopsids and Ferns
However, the early fossil record of these groups is marked by a number of taxa with dubious combinations of characters which could tie them to either group, ...Missing: Equisetales origins
[46]
Sphenophyta: Fossil Record
The first fossils that can be unambiguously placed in the Sphenophyta are Late Devonian in age. The Sphenophyta appear to have evolved from ancestors in the ...Missing: Pseudobornia Taeniocrada
[47]
Common Horsetail - University of Puget Sound
Our familiar horsetails are relatively primitive plants, first detected in the fossil record in the Carboniferous period (>300 million years ago), when they ...
[48]
Archaeocalamites from the upper Mississippian of Arkansas
Petrified Archaeocalamites fragments are described from the Mississippian of Arkansas. Anatomical features of the wood of stems and rhizomes are detailed.Missing: Equisetales | Show results with:Equisetales
[49]
Evidence of the oldest extant vascular plant (horsetails) from the ...
Horsetails, the unique survivors of a very ancient group of vascular plants, the Sphenophyta, have a history reaching back to the Upper Devonian and may ...
[50]
Evidence of the oldest extant vascular plant (horsetails) from the ...
Horsetails, the unique survivors of a very ancient group of vascular plants, the Sphenophyta, have a history reaching back to the Upper Devonian and may ...
[51]
Systematic and organ relationships of Neocalamites (Halle ...
Neocalamites carrerei is the best represented equisetalean species in the Triassic of Gondwana. Nododendron suberosum is widespread in the Middle–Upper Triassic ...
[52]
[PDF] U.S. GEOLOGICAL SURVEY BULLETIN 2085-B
Species richness of ferns, conifers, and Equisetum in. 20 plant-fossil localities from the Eocene Puget Group, west-cen- tral Washington. Localities are grouped ...<|control11|><|separator|>
[53]
[PDF] The first Cenozoic Equisetum from New Zealand - ScienceDirect.com
Here we describe Equisetum remains from two early to earliest middle Miocene fossil assemblages of New Zealand. We also assess the potential causes of ...<|separator|>
[54]
[PDF] Ecological Sorting of Vascular Plant Classes During the Paleozoic ...
Terra firma habitats, the favored territory of seed plants, were the most physically diverse and were thus capable of supporting the most species and the most ...Missing: Equisetales | Show results with:Equisetales<|control11|><|separator|>
[55]
Biogeography and genome size evolution of the oldest extant ... - PMC
Feb 18, 2021 · Phylogenetic diversification of Equisetum (Equisetales) as inferred from Lower Cretaceous species of British Columbia, Canada. American ...
[56]
Equisetum L. - GBIF
variegatum). Only four (E. bogotense, E. giganteum, E. myriochaetum, and E. ramosissimum) of the fifteen species are known to be native south of the Equator.<|control11|><|separator|>
[57]
Equisetum arvense (field horsetail) | CABI Compendium
Nov 16, 2021 · This datasheet on Equisetum arvense covers Identity, Overview, Distribution, Hosts/Species Affected, Diagnosis, Biology & Ecology, Natural Enemies, Impacts, ...
[58]
Common Horsetail (Equisetum arvense). - Forest Service
Equisetum arvense is distributed throughout temperate and arctic areas of the northern hemisphere, growing typically in moist soils.
[59]
[PDF] Equisetum arvense L. AS A BIOINDICATOR OF ACID SOILS
In agricultural practice Equisetum arvense is known as a bio-indicator species for acid soils with excess moisture. Diminution of acid soils can be achieved by ...
[60]
Biology and Functional Ecology of Equisetum with Emphasis on the ...
Biology and Functional Ecology of Equisetum with Emphasis on the Giant Horsetails. Published: 15 January 2013. Volume 79, pages 147–177, (2013); Cite this ...
[61]
Silicon and Plants: Current Knowledge and Technological ... - NIH
Mar 23, 2017 · Biogenic silica is also a deterrent against herbivores. Additionally ... New insight into silica deposition in horsetail (Equisetum arvense).
[62]
Salinity tolerance ecophysiology of Equisetum giganteum in South ...
Equisetum giganteum is a widespread species associated with wetlands and other moist habitats at elevations typically above 500 m in the American tropics ...
[63]
The History of Herbivory on Sphenophytes: A New Calamitalean ...
Equisetum (Equisetales: Equisetaceae) extends back to the earliest Jurassic and is approximately 200 million years old (Elgorriaga et al. 2018). The species ...<|separator|>
[64]
Variation of the Main Alkaloid Content in Equisetum palustre L. in ...
Nov 9, 2020 · We tested the hypothesis that the ontogenetic stage of plant development may explain a significant part of the variations in the main Equisetum-type alkaloids.
[65]
https://pmc.ncbi.nlm.nih.gov/articles/PMC5362598/
[66]
[PDF] Equisetum pratense - NJ.gov
Apr 13, 2022 · Equisetum pratense sporophytes have perennial rhizomes that produce annual stems. Two kinds of stems are produced by Meadow Horsetail: The ...
[67]
Horsetail | Gwich'in Social & Cultural Institute
The horsetail plant is used to make a medicinal tea to treat kidney problems, bladder infections or urinary track problems. To treat a bladder infection, ...
[68]
Horsetail - LiverTox - NCBI Bookshelf - NIH
Jul 25, 2022 · Horsetail is an extract of the plant Equisetum arvense which has been used in traditional medicine for bladder and kidney conditions and to promote wound ...
[69]
Horsetails and Scouring Rushes | The Outside Story
May 12, 2008 · The scouring rushes, on the other hand, mostly have single erect stems whose high silica content makes them suitably abrasive for scrubbing pots ...
[70]
SILICON AND BONE HEALTH - PMC - NIH
Accumulating evidence over the last 30 years strongly suggest that dietary silicon is beneficial to bone and connective tissue health.
[71]
Botanicals in Postmenopausal Osteoporosis - PMC - NIH
It contains abundant constituents that may exert beneficial effects on bone health, e.g., silica, flavonoids, and triterpenoids. ... horsetail extract enhanced ...
[72]
Equisetum arvense (common horsetail) modulates the function of ...
Aug 4, 2014 · In Europe, extracts of Equisetum arvense (common horsetail) have a long tradition in the treatment of inflammatory disorders.Missing: silica | Show results with:silica
[73]
Phytochemistry and Pharmacology of the Genus Equisetum ...
Mar 5, 2021 · The Equisetum genus, Equisetaceae family, is widely distributed worldwide and may be the oldest nonextinct genus on Earth.
[74]
Horsetail - Uses, Side Effects, and More - WebMD
When taken by mouth: It is possibly unsafe to take horsetail products that contain thiaminase. Thiaminase breaks down the vitamin thiamine. This could lead to ...Missing: carcinogens | Show results with:carcinogens
[75]
[PDF] Assessment report on Equisetum arvense L., herba
Feb 2, 2016 · Equisetum is widely distributed throughout the temperate zones of the northern hemisphere, Canada, the USA, Europe and Asia south to Turkey, ...
[76]
Scouring Rush - Green Timbers Heritage Society
Nov 29, 2016 · The stems are very rich in silica. They are used for scouring and polishing metal and as a fine sandpaper. The stems are first bleached by ...
[77]
https://pmc.ncbi.nlm.nih.gov/articles/PMC7954623/
[78]
Toxicity of equisetum to horses - Ontario.ca
May 19, 2022 · Equisetum is toxic to horses in large quantities, especially in contaminated hay. Symptoms include weight loss, diarrhea, and loss of muscle ...
[79]
[PDF] RESTORING RARE NATIVE HABITATS IN THE WILLAMETTE ...
introduced to North America as a garden ornamental in the late 1800s. It ... Equisetum hyemale. X. X. X ee,. Sword Fern. Polystichum munitum. X. X m, ee, ii.