Superregnum: Eukaryota
Regnum: Plantae
Divisio: Tracheophyta
Divisio: Lycopodiophyta
Classis: Lycopodiopsida
Subclassis: Lycopodiidae
Ordo: Lycopodiales
Familia: Lycopodiaceae
Subfamiliae: Lycopodielloideae – Lycopodiastroideae – Lycopodioideae – Huperzioideae
Genera: Austrolycopodium – Brownseya – Dendrolycopodium – Diphasiastrum – Diphasium – Huperzia – Lateristachys – Lycopodiastrum – Lycopodiella – Lycopodium – Palhinhaea – Phlegmariurus – Phylloglossum – Pseudodiphasium – Pseudolycopodiella – Pseudolycopodium – Spinulum
Paleogenera: †Lycopodicaulis
Name
Lycopodiaceae P.Beauv. ex Mirb. in J.B.A.P.M. de Lamarck & C.F.B. de Mirbel, Hist. Nat. Vég. 4: 293 (1802).
Type genus: Lycopodium L., (1753)
Synonyms
Heterotypic
Huperziaceae Rothm., Feddes Repert. 66: 236 (1962), nom. nov.
Urostachyaceae Rothm., Feddes Repert. 54: 58 (1944), nom. illeg.
Phylloglossaceae Kunze, Bot. Zeit. (Berlin) 1: 722 (1843).
Lycopodiellaceae Val.N.Tikhom., Novosti Sist. Vyssh. Rast. 49: 149 (2018).
References
Primary references
Lamarck & Mirbel, 1802. Hist. Nat. Veg. 4: 293
Additional references
Øllgaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Opera Botanica 92: 153–178. Reference page.
Christenhusz, M.J.M., Zhang, X.-C. & Schneider, H. 2011. A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19: 7–54. DOI: 10.11646/phytotaxa.19.1.2 Open access. Reference page
Pteridophyte Phylogeny Group (PPG I) 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54(6): 563–603. DOI: 10.1111/jse.12229 Reference page.
Greuter, W. & Troìa, A. 2014. (2292) Proposal to conserve the name Palhinhaea against Lepidotis (Lycopodiaceae). Taxon 63(3): 680–682. DOI: 10.12705/633.23 Reference page. (authorship of Palisot de Beauvois)
Links
Govaerts, R. et al. 2022. Lycopodiaceae in Kew Science Plants of the World Online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2022 Jun 24. Reference page.
Hassler, M. 2022. World Ferns. Checklist of Ferns and Lycophytes of the World. . Lycopodiaceae Accessed: 24 Jun 2022.
Vernacular names
العربية: رجل ذئبية
беларуская: Дзеразовыя
català: Licopodiàcies
čeština: Plavuňovité
dansk: Ulvefod-familien
Deutsch: Bärlappgewächse
English: Club-moss family
español: Lycopodiáceas
eesti: Kollalised
فارسی: پنجهگرگیان
suomi: Liekokasvit
français: Lycopodiacées
עברית: רגלי זאב
hrvatski: Crvotočine
magyar: Korpafűfélék
íslenska: Jafnaætt
日本語: ヒカゲノカズラ科
한국어: 석송과
lietuvių: Pataisiniai
latviešu: Staipekņu dzimta
македонски: Црвоточини
Nederlands: Wolfsklauwfamilie
norsk: Kråkefotfamilien
polski: Widłakowate
română: Licopodiacee
русский: Плауновые
slovenščina: Lisičjakovke
svenska: Lummerväxter
українська: Плаунові
oʻzbekcha / ўзбекча: Plaundoshlar
Tiếng Việt: Họ Thạch tùng
中文: 石松科
The Lycopodiaceae (class Lycopodiopsida, order Lycopodiales) are an old family of vascular plants, including all of the core clubmosses and firmosses, comprising 16 accepted genera[2] and about 400 known species.[3] This family originated about 380 million years ago in the early Devonian, though the diversity within the family has been much more recent.[4] "Wolf foot" is another common name for this family due to the resemblance of either the roots or branch tips to a wolf's paw.[5]
Description
Members of Lycopodiaceae are not spermatophytes and so do not produce seeds. Instead they produce spores, which are oily and flammable, and are the most economically important aspects of these plants. The spores are of one size (i.e. the plants are isosporous) and are borne on a specialized structure at the apex of a shoot called a strobilus (plural: strobili), which resembles a tiny battle club, from which the common name derives. Members of the family share the common feature of having a microphyll, which is a "small leaf with a single vein, and not associated with a leaf gap in the central vascular system."[4] In Lycopodiaceae, the microphylls often densely cover the stem in a linear, scale-like, or appressed fashion to the stem, and the leaves are either opposite or spirally arranged. The club mosses commonly grow to be 5–20 cm tall.[4] The gametophytes in most species are non-photosynthetic and myco-heterotrophic, but the subfamily Lycopodielloideae and a few species in the subfamily Huperzioideae have gametophytes with an upper green and photosynthetic part, and a colorless lower part in contact with fungal hyphae.[6][7] In Lycopodioideae monoplastidic meiosis is common, whereas polyplastidic meiosis is found in Lycopodielloideae and Huperzioideae.[8]
Taxonomy
The family Lycopodiaceae is considered to be basal within the Lycopodiopsida (lycophytes). One hypothesis for the evolutionary relationships involved is shown in the cladogram below.[2]
Lycopodiopsida
Lycopodiaceae
Isoetaceae
Selaginellaceae
Within the family, there is support for three subgroups. In 2016, Field et al. proposed that the primary division is between Lycopodielloideae plus Lycopodioideae and the Huperzioideae (names sensu PPG I).[9]
Lycopodiaceae
Lycopodielloideae (Lycopodiella s.l.)
Lycopodioideae (Lycopodium s.l.)
Huperzioideae (Huperzia s.l.)
There are about 400 known species in the family Lycopodiaceae.[3] Sources differ in how they group these into genera. Field et al. (2016) say "Most Lycopodiaceae species have been re-classified into different genera several times, leading to uncertainty about their most appropriate generic identification."[9] In the PPG I system, the family has 16 accepted genera, grouped into three subfamilies, Lycopodielloideae, Lycopodioideae and Huperzioideae, based in part on molecular phylogenetic studies. The Huperzioideae differ in producing spores in small lateral structures in the leaf axils,[9] and it has been suggested that they be recognized as a separate family.[citation needed] Other sources use fewer genera; for example, the three genera placed in the subfamily Huperzioideae in PPG I, Huperzia, Phlegmariurus and Phylloglossum, have also all been treated within a broadly defined Huperzia.[9]
The species within this family generally have chromosome counts of n=34. A notable exception are the species in Diphasiastrum, which have counts of n=23.[10]
Genera
As of June 2024, the Checklist of Ferns and Lycophytes of the World recognized the following genera as members of Lycopodiaceae.[11] All of these are recognized by the Pteridophyte Phylogeny Group classification of 2016 (PPG I), except for the genus Brownseya, described in 2021.[2] Other classifications circumscribe the genera in the family more broadly, recognizing the subfamilies Lycopodielloideae, Lycopodioideae, and Huperzioideae as the genera Lycopodiella, Lycopodium, and Huperzia.
Phylogeny of Lycopodiaceae[12][13]
Lycopodiaceae
Huperzioideae
Huperzia s.s. Bernhardi
Phylloglossum Kunze
Phlegmariurus (Herter) Holub
Wagner & Beitel ex Øllgaard
Lycopodielloideae
Brownseya Zhang et al.
Palhinhaea Franco & Vasconcellos
Lateristachys Holub
Pseudolycopodiella Holub
Lycopodiella Holub
Wagner & Beitel ex Øllgaard
Lycopodiastroideae
Lycopodiastrum Holub ex Dixit
Zhang & Zhou
Lycopodioideae
Diphasiastrum Holub
Lycopodium s.s. von Linné
Spinulum Haines
Pseudolycopodium Preslia ex Holub
Pseudodiphasium Holub
Austrolycopodium Holub
Dendrolycopodium Haines
Diphasium Presl ex Rothmaler
Eaton ex Øllgaard
Distribution and habitat
The members of Lycopodiaceae are terrestrial or epiphytic in habit and are most prevalent in tropical mountain and alpine environments.[4] Though Lycopodiaceae are most abundant in these regions, they are cosmopolitan, excluding arid environments.[14]
Evolution
Lycopodiaceae (homosporous lycophytes) split off from the branch leading to Selaginella and Isoetes (heterosporous lycophytes) about ~400 million years ago, during the early Devonian. The two subfamilies Lycopodioideae and Huperzioideae diverged ~350 million years ago, but has evolved so slowly that about 30% of their genes are still in syntenic blocks (remaining in the same arrangement). They have also gone through independent whole genome duplications. In most plants the majority of duplicate genes are lost relatively quickly through diploidization, but in this group both sets of genes tends to be retained with relatively few alterations, even after hundreds of millions of years after the duplication event.[15][16] Spores indicate that the crown group of Lycopodiaceae had emerged by the Triassic-Jurassic boundary, around 200 million years ago,[17] with a member of the crown group of Lycopodioideae known from the Early Cretaceous of China.[18]
Uses
The running clubmosses (Diphasiastrum) have long been used as greenery for Christmas decoration.
The spores have long been used as a flash powder. See Lycopodium powder.
The spores have been used by violin makers for centuries as a pore filler.
In Cornwall, club mosses gathered during certain lunar phases were historically used as a remedy for eye disease.
References
James L. Reveal, Indices Nominum Supragenericorum Plantarum Vascularium
PPG, I (2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 563–603. doi:10.1111/jse.12229. S2CID 39980610.
Christenhusz, M. J. M. & Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3). Magnolia Press: 201–217. doi:10.11646/phytotaxa.261.3.1.
Judd; et al. (2015). Plant Systematics: A Phylogenetic Approach. Sunderland, MA: Sinauer Associates.
"Lycopodiaceae". www.flora.dempstercountry.org. Retrieved 2017-12-20.
Mycoheterotrophy: The Biology of Plants Living on Fungi
Phylogeny of Phlegmariurus (Lycopodiaceae) focusing on Brazilian endemic species
Sporogenesis, sporoderm and mature spore ornamentation in Lycopodiaceae
Field; et al. (January 2016). "Molecular Phylogenetics and the Morphology of the Lycopodiaceae Subfamily Huperzioideae Supports Three Genera: Huperzia, Phlegmariurus and Phylloglossum". Molecular Phylogenetics and Evolution. 94, Part B (Pt B): 635–57. doi:10.1016/j.ympev.2015.09.024. PMID 26493224.
Flora of North America, Diphasiastrum
Hassler, Michael. "Lycopodiaceae". World Ferns. Retrieved 2024-06-02.
Chen, De-Kui; Zhou, Xin-Mao; Rothfels, Carl J.; Shepherd, Lara D.; Knapp, Ralf; Zhang, Liang; Lu, Ngan Thi; Fan, Xue-Ping; Wan, Xia; Gao, Xin-Fen; He, Hai; Zhang, Li-Bing (2021), "A global phylogeny of Lycopodiaceae (Lycopodiales; lycophytes) with the description of a new genus, Brownseya, from Oceania", Taxon, 71 (1): 25–51, doi:10.1002/tax.12597
Chen, De-Kui; Zhou, Xin-Mao; Rothfels, Carl J.; Shepherd, Lara D.; Knapp, Ralf; Zhang, Liang; Lu, Ngan Thi; Fan, Xue-Ping; Wan, Xia; Gao, Xin-Fen; He, Hai; Zhang, Li-Bing (2021), "Fig. S1 Maximum likelihood phylogeny of Lycopodiaceae based on seven plastid markers (atpA, psbA-trnH, rbcL, rps4 & rps4-trnS, trnL & trnL-F)" (PDF), Taxon, 71 (1): 25–51, doi:10.1002/tax.12597
Øllgaard, B. (1990). "Lycopodiaceae". Pteridophytes and Gymnosperms. pp. 31–39. doi:10.1007/978-3-662-02604-5_10. ISBN 978-3-642-08080-7. {{cite book}}: |journal= ignored (help)
A window into plant evolution: The unusual genetic journey of lycophytes
Extraordinary preservation of gene collinearity over three hundred million years revealed in homosporous lycophytes
Wikström, Niklas; Larsén, Eva; Khodabandeh, Anbar; Rydin, Catarina (January 2023). "No phylogenomic support for a Cenozoic origin of the "living fossil" Isoetes". American Journal of Botany. 110 (1): e16108. doi:10.1002/ajb2.16108. ISSN 0002-9122. PMC 10108322. PMID 36401556.
Herrera, Fabiany; Testo, Weston L.; Field, Ashley R.; Clark, Elizabeth G.; Herendeen, Patrick S.; Crane, Peter R.; Shi, Gongle (March 2022). "A permineralized Early Cretaceous lycopsid from China and the evolution of crown clubmosses". New Phytologist. 233 (5): 2310–2322. doi:10.1111/nph.17874. ISSN 0028-646X. PMID 34981832. S2CID 245670357.
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