Scientific Name:
Lycopodiella Holub, Preslia 36: 20, 22 (1964)
  • = Lepidotis P.Beauv. ex Mirbel in Lamarck & Mirbel, Hist. Nat. Vég. 3, 477 (1802) nom. rej.
  • = Palhinhaea Franco & Vasc. ex Vasc. & Franco, Bol. Soc. Brot. Ser. 2 41: 24 (1967) nom. cons.
  • = Lateristachys Holub, Folia Geobot. Phytotax. 18: 441 (1983)
  • = Pseudolycopodiella Holub, Folia Geobot. Phytotax. 18: 441 (1983)
  • = Brownseya Li Bing Zhang, L.D.Sheph., D.K.Chen, X.M.Zhou & H.HE in Chen et al., Taxon 00: 17 (online) (2021)
Type Taxon:
Lycopodiella inundata (L.) Holub
From the related genus Lycopodium and the Latin diminutive suffix -ellus.

Terrestrial plants. Main stems indeterminate, horizontal and creeping or looping, giving rise to determinate, erect, simple or branched, aerial stems, often of different thickness to the horizontal stems. Roots arising at intervals along the length of horizontal stems. Leaves spirally arranged, usually all similar; leaf margins entire or minutely ciliate. Bulbils absent. Sporophylls modified and aggregated into distinct strobili, ephemeral. Strobili terminal on ultimate branches or lateral on overtopping branches, erect or pendent, sessile or stalked. Spores rugulate.


A genus of c. 54 species (PPG 1 2016, including all species in the subfamily Lycopodielloideae).

Following Øllgaard (1987, 1989, 1990) the lycopodioid clade within Lycopodiaceae is interpreted here to include just two genera, Lycopodiella and Lycopodium. However, more recent classifications further subdivide these taxa into four and nine genera, respectively (Field et al. 2016).

The lycopodioid clade is distinguished by anisotomous branching, with dichotomies resulting in branches of unequal thickness; horizontal, indeterminate main stems that are creeping, looping or climbing, and give rise to aerial determinate branches; roots that arise at intervals along the length of the main stem; strongly modified sporophylls aggregated into strobili (Øllgaard 1987, 1990); and reticulate, baculate or rugulate spores (Tryon & Lugardon 1991, figs 229.1–12, 230.1–12; Large & Braggins 1991).

Lycopodiella is distinguished from Lycopodium by its horizontal and subterranean, prostrate or looping stems, strobili that are terminal and either pendent and sessile, or erect on undivided stalks. In some species the strobili are lateral on overtopping branches. Spores are rugulate (Tryon & Lugardon 1991, figs 230.1–12; Large & Braggins 1991). In New Zealand, all species have spirally arranged leaves of one size.

Four species of Lycopodiella (as circumscribed here) are present in New Zealand. In the classification of Field et al. (2016), L. diffusa and L. lateralis would be placed in Lateristachys, L. cernua in Palhinhaea, and L. serpentina in Pseudolycopodiella. New Zealand species of Lycopodiella were included in Lycopodium by Allan (1961).

1Strobili mostly borne laterally on aerial stems, tan-brown, red-brown or chocolate-brown at maturity2
Strobili borne terminally on aerial stems and branches, yellow-brown at maturity3
2Aerial stems prostrate, 25–160 mm long; strobili red-brown or chocolate-brown at maturitydiffusa
Aerial stems erect or scrambling upwards, 50–700 mm long; strobili tan-brown at maturitylateralis
3Horizontal stems looping across the ground; aerial stems branched many times, the ultimate branches curved downwards; strobili pendent or inclined downwards cernua
Horizontal stems prostrate, not looping; aerial stems unbranched, erect; strobili erectserpentina

Lycopodiella is a genus of over 50 species occurring in moist temperate and tropical regions. The greatest diversity is in South America; four species in South Africa (Roux 2009), five in Australia (Chinnock 1998). Four indigenous, non-endemic species in New Zealand.

Indigenous (Non-endemic)
Number of species in New Zealand within Lycopodiella Holub
Indigenous (Non-endemic)4

Chromosome numbers in Lycopodiella are very variable. In sect. Lycopodiella (= Lycopodiella s.str.) most counts are diploid with n = 78; in sect. Caroliniana (= Pseudolycopodiella) n = 34, 35 and c. 69; and in sect. Campylostachys (= Palhinhaea) counts of n = 104, 108, 110, 136, c. 165 have been reported (Øllgaard 1987, 1990).


The type of Lepidotis was first designated by Rothmaler (1944), who chose L. cernua (L.) P.Beauv. That choice was challenged by Pichi Sermolli (1971) who instead designated Lycopodium clavatum L., thereby making Lepidotis a synonym of Lycopodium, a view shared, among others, by Øllgaard (1989, 2012). However, Greuter & Troia (2014) dismissed Pichi Sermolli’s arguments and asserted that Lepidotis is a legitimate name typified by L. cernua, stating that even if Rothmaler’s typification was disregarded, a subsequent typification by Fuchs (1955) could not be rejected. As a consequence, the name Palhinhaea (L.) Franco & Vasc. was made illegitimate because it was based on the previously designated type of Lepidotis. Greuter & Troia therefore proposed the conservation of Palhinhaea against Lepidotis to preserve the use of Palhinhaea and remove the threat to Lycopodiella, a proposal now accepted, leaving Lepidotis a rejected name under Lycopodiella.  

Brownsey, P.J.; Perrie, L.R. 2020: Lycopodiaceae. In: Breitwieser, I.; Wilton, A.D. (ed.) Flora of New Zealand — Ferns and Lycophytes. Fascicle 27. Manaaki Whenua Press, Lincoln.
Chen, D.-K.; Zhou, X.-M.; Rothfels, C.J.; Shepherd,L.D.; Knapp, R.; Zhang, L.; Thi Lu, N.; Fan, X.-P.; Wan, X.; Gao, X.-F.; He, H.; Zhang, L.-B. 2021: A global phylogeny of Lycopodiaceae (Lycopodiales; lycophytes) with the description of a new genus, Brownseya, from Oceania. Taxon 00(00): 1–27 (online).
Chinnock, R.J. 1998: Lycopodiaceae. In: Flora of Australia. Vol. 48. 66–85, 705.
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.
Field, A.R.; Testo, W.; Bostock, P.D.; Holtum, J.A.M.; Waycott, M. 2016: Molecular phylogenetics and the morphology of the Lycopodiaceae subfamily Huperzioideae supports three genera: Huperzia, Phlegmariurus and Phylloglossum. Molecular Phylogenetics and Evolution 94: 635–657.
Fuchs, H.P. 1955: Urostachys (Herter 1909, 5/29) Herter 1922, 249 nomen genericum conservandum? Verhandlungen der Naturforschenden Gesellschaft in Basel 66: 33–48.
Greuter, W.; Troia, A. 2014: (2292) Proposal to conserve the name Palhinhaea against Lepidotis (Lycopodiaceae). Taxon 63: 680–682.
Holub, J. 1964: Lycopodiella, novy rod radu Lycopodiales. Preslia 36: 16–22.
Holub, J. 1983: Validation of generic names in Lycopodiaceae: with a description of a new genus Pseudolycopodiella. Folia Geobotanica et Phytotaxonomica 18: 439–442.
Kramer, K.U.; Green, P.S. 1990: Pteridophytes and gymnosperms. Kubitzki, K. (ed.) The Families and Genera of Vascular Plants. Vol. 1. Springer-Verlag, Berlin.
Lamarck, J.B.A.P.M de; Mirbel, C.F.B. de 1802: Histoire naturelle des végétaux, classés par familles. Vol. 3. Crapelet, Paris.
Large, M.F.; Braggins, J.E. 1991: Spore atlas of New Zealand ferns and fern allies. SIR Publishing, Wellington.
Øllgaard, B. 1987: A revised classification of Lycopodiaceae s. lat. Opera Botanica 92: 153–178.
Øllgaard, B. 1989: Index of the Lycopodiaceae. Biologiske Skrifter 34: 1–135.
Øllgaard, B. 1990: Lycopodiaceae. In: Kramer, K.U.; Green, P.S. (ed.) The families and genera of vascular plants. 1 Pteridophytes and gymnosperms. Springer-Verlag, Berlin.
Øllgaard, B. 2012: New combinations in Neotropical Lycopodiaceae. Phytotaxa 57: 10–22.
Pichi Sermolli, R.E.G. 1971: Names and types of the genera of fern-allies: Lycopodiaceae, Selaginellaceae, Isoetaceae, Equisetaceae, Psilotaceae, Tmesipteridaceae. Webbia 26: 129–194.
PPG 1 2016: A community-derived classification for extant lycophytes. Journal of Systematics and Evolution 54(6): 563–603.
Rothmaler, W. 1944: Pteridophyten-Studien, I. Feddes Repertorium Specierum Novarum Regni Vegetabilis 54: 55–82.
Roux, J.P. 2009: Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23: 1–296.
Tryon, A.F.; Lugardon, B. 1991: Spores of the Pteridophyta. Springer-Verlag, New York.
Vasconcellos, J. de Carvalho e; Franco, J. do Amaral 1967: Breves notas sobre Licopodiáceas. Boletim da Sociedade Broteriana (Series 2) 41: 23–25.