Named for the type genus Lycopodium, from Greek λύκος (lykos) meaning wolf, and πόδι (pod) meaning foot.

Club Moss, Fir Moss, Ground Pine, Tassel Fern.

Lycopodium L.

Terrestrial, epilithic or epiphytic. Sporophytes emergent, photosynthetic, annually deciduous or perennially evergreen and long-lived. Protostelic (mostly), plectostelic (in lateral shoots of some species) or solenstelic (rarely). Habit varied, including: (i) solitary unbranched determinate shoots arising from a subterranean tuber bearing the roots; (ii) tufted isotomous indeterminate shoots arising from the axils of basal leaves, with roots arising from the narrow stem stele and descending through the broad cortex to the stem base to emerge as a basal tuft in Huperzioideae; and (iii) non-tufted, anisotomous shoots with indeterminate horizontal stems and determinate aerial stems usually of differing thickness and branching pattern, with roots arising from a broad stem stele and emerging directly through the stem wall in Lycopodioideae and Lycopodielloideae. Shoots may be erect, pendulous, scrambling or climbing. Shoots homophyllous with undifferentiated sterile leaves and sporophylls, or heterophyllous with markedly differentiated sterile leaves and sporophylls. Sterile leaves simple and single-veined, isophyllous or anisophyllous, lacking a ligule, arranged in alternating subspiral whorls of 2–11, appearing subopposite decussate, longitudinally ranked, verticillate or spiralling. Sporophylls intermingled with undifferentiated sterile leaves or arranged into sessile or leafless pedunculate terminal or lateral spikes. Sporophylls simple and single-veined, isophyllous, lacking a ligule, persisting after sporangia dehiscence or deciduous after sporangia dehiscence. Sporophyll arrangement similar to sterile leaves or with a reduced number of parts per subspiral whorl in heterophyllous species. Sporangia axillary, reniform to globular, isovalvate or anisovalvate. Spores trilete tetrahedral (sometimes appearing round) with bacculate, fovelate, fossulate, reticulate, rugate or scabrate surface ornamentation. Gametophytes subterranean and mycoheterotrophic or emergent and photosynthetic, tuberous of various forms, with gametangia on the upper surface.

Readily identified by the combination of being microphyllous (having scale-like leaves with a single vein) and being homosporous (having a single type of sporangium) with sporangia born in the axils of leaves.

Evergreen, except Phylloglossum drummondii Kunze which is annually deciduous.

Native. Cultivated. None known to be naturalised.

Globally distributed. In Australia, occurring in the coastal and subcoastal regions of all states and territories and most offshore islands.

The habit and habitat of Lycopodiaceae is extremely diverse, including a myriad of terrestrial, epilithic and epiphytic forms ranging from colonial terrestrial plants in oligotrophic wetlands to hanging epiphytes in rainforest canopies. They occur in a variety of high light humid habitats and are generally absent from deep shade or arid zones. Most species diversity globally is associated with montane areas, especially in the tropics.

Reproduce and disperse as spores, which give rise to short-lived surface-living photosynthetic gametophytes (Lycopodielloideae) or long-lived subterranean mycoheterotrophic gametophytes (Huperzioideae and Lycopodioideae). Gametrophytes produce both antheridia and archaegonia and may self or cross fertilise. The period between spore germination and sporophyte emergence is usually more than several years.

Many species also reproduce vegetatively by layering of shoot apices or by detachable bulbils in Huperzia. Lycopodielloideae and Lycopodioideae form colonies and spread by means of lateral or layering stems. Huperzioideae are localised plants that form a dense local tuft of roots and shoots. Mature Huperzia colonies may form 'fairy rings' as the living part grows out and the centre senesces. Mature hanging Phlegmariurus may revert multiple times between sterile and fertile shoots. Phylloglossum is annually deciduous, forming a subterranean tuber in the hot and dry time of the year, re-emerging with a tiny leafy shoot the following cool and wet season.

Many species of the genus Phlegmariurus are Vulnerable or Endangered and one species of Huperzia  is presumed Extinct in the Queensland Nature Conservation Act (Anon. 1992) and the Australian Federal Environment Protection and Biodiversity Conservation Act (Anon. 1999). Trade in Phlegmariurus has been regulated in Queensland to prevent poaching of wild plants (Anon. 2010).

Many species of the epiphytic genus Phlegmariurus (tassel ferns) are cultivated as ornamental hanging basket plants, especially in tropical and subtropical regions (see Jones 1987; Elliot & Jones 1993; Spencer 1995; McAuliffe 2012 under the generic names Lycopodium L. and Huperzia Bernh.). The trade in tassel ferns has been regulated in Queensland to prevent poaching of wild plants, with native species being required to be produced artificially and tagged by approved propagators (Anon. 2010).

Huperzine extracted from Huperzia serrata (Toothed Clubmoss) has been used as a traditional medicine in China and has been reported to have efficacy in preventing Alzheimer's disease (Ma & Gang 2008). Several Australian species of the genera Huperzia and Phlegmariurus contain Huperzine, but are not presently used as a commercial source of Huperzine (Goodger et al. 2008; Lim et al. 2010).

Several species of Phlegmariurus (tassel ferns) are named by, and have cultural significance for, indigenous people of the Wet Tropics, Queensland.

A monophyletic family with three subfamilies and 16 genera and 388 species globally with 12 genera and 24 species occurring in Australia (Du Puy & Orchard 1993; Green 1994; Chinnock 1998; Wikström & Kenrick 1997, 2000, 2001; Field et al. 2016; PPG I 2016; Testo et al. 2018a, 2018; Bauret et al. 2018; Field 2020). The classification at generic rank differs considerably from Chinnock (1998) and several new species are recognised for Australia. The reinstatement of Holub’s (1964, 1975, 1983) changes to the generic classification of Lycopodiaceae have resulted from molecular data revealing the non-monophyly of the genus Huperzia Bernh. and also recognition of the deep-time divergence and morphological specialisation within the genera recognised by Chinnock (1998) as Lycopodium L. and Lycopodiella Holub (Wikström & Kenrick 1997, 2000, 2001; Øllgaard 2015; Field et al. 2016; Bauret et al. 2018; Testo et al. 2018a, b; Field 2020).

Many species referred to Lycodium and Lycopodiella in the Australian literature are now placed in different genera, including Austrolycopodium, Diphasium, Lateristachys, Palhinhaea, Pseudodiphasium, Pseudolycopodiella and Pseudolycopodium (PPG I 2020; Field 2020). Similarly, the non-bulbiferous epiphytic species referred to Huperzia species are now placed in Phlegmariurus. The monotypic genus Phylloglossum remains unchanged.

Lycopodiaceae is an ancient family with a Carboniferous crown group age (Testo et al. 2018b). The genera as adopted by PPG I (2016) and Field (2020) diversified during the Mesozoic, and most species diversity is Cenozoic (Bauret et al. 2018; Testo et al. 2018b).

The generic system adopted by Pteridophyte Phylogeny Group I (2016) and for Australian Lycopodiaceae by Field (2020) has not been fully processed by the Australian Plant Census (accessed February 2020). As such, the generic classification adopted by Chinnock (1998) is used for Lycopodium and Lycopodiella as an interim, with the correct current name listed in Taxonomic Notes of the interim profiles.

Key to subfamilies of Lycopodiaceae

1a. Shoots a tuft of isodichotomous branches with new shoots arising from the axils of basal leaves and roots emerging as a basal tuft; or shoots a solitary tuft of leaves arising from a subterranean tuber = subfamily Huperzioideae (see profiles for Huperzia, Phlegmariurus, Phylloglossum).

1b. Shoots non-tufted, anisodichotomously branched, usually with differentiated lateral and upright shoots and with roots arising adventitiously from the lateral stems = 2

2a. Strobili pendulous and sessile; erect terminating unbranched aerial stems; or lateral on branched aerial stems; spores rugate = subfamily Lycopodielloideae (see profile 'Lycopodiella').

2b. Strobili erect, terminating branched aerial stems; or strobili pendulous, terminating hanging peduncles; spores baculate, reticulate, rugate or scabrate = subfamily Lycopodioideae (see profile 'Lycopodium').

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Submitted 2 February 2020. Published 14 February 2020.