Classification
 Nomenclature
Scientific Name:
Cheilanthes Sw., Syn. Fil. 126 (1806), nom. cons.
Type Taxon:
Cheilanthes micropteris Sw.
Etymology:
From the Greek cheilos (lip) and anthos (flower), a reference to the sori, which occur on the margin of the pinnae.
 Description

Terrestrial or rupestral ferns. Rhizomes short- to long-creeping, scaly. Rhizome scales non-clathrate, narrowly ovate, attached at base, concolorous or bicolorous, light to dark brown. Fronds monomorphic (NZ) or rarely dimorphic (not NZ). Stipes and rachises red-brown. Laminae 2–3-pinnate (NZ) or 1-pinnate and 4–5-pinnate (not NZ), herbaceous or coriaceous, glabrous or scaly and/or hairy, sometimes farinose abaxially (not NZ). Pinnae and ultimate lamina segments not articulated to rachis and costae. Veins free (NZ) or rarely reticulate (not NZ). Sori terminating veins at lamina margin, or extending almost continuously along the margins; paraphyses present; sori ± protected by the unmodified inrolled lamina margin. Spores trilete, lacking chlorophyll; perispores verrucate, tuberculate or echinate, without an equatorial flange.

 Taxonomy

A genus of c. 100 species, included in the subfamily Cheilanthoideae along with Myriopteris and Pellaea (PPG 1 2016).

Cheilanthes, as currently construed (Eiserhardt et al. 2011; Zhang & Yatskievych 2013; PPG 1 2016), is not monophyletic, and is in need of a comprehensive phylogenetic study to determine generic boundaries within the group. One New Zealand species was previously included in the genus Notholaena (e.g. Cheeseman 1925, as Nothochlaena), which was first described from Australia by Brown (1810). However, the typification of Notholaena has been very controversial, and the genus is now considered to be typified by a New World species, and confined to the Americas (Yatskievych & Smith 2003; Rothfels et al. 2008; PPG 1 2016).

The Australian species of Cheilanthes, including both species indigenous to New Zealand, have been extensively reviewed by Quirk et al. (1983) and Chambers & Farrant (1991, 1998), and those in southern Africa by Anthony (1984). Allan (1961) followed earlier New Zealand Flora writers in accepting three species, but one of them, C. tenuifolia, is now thought to occur in coastal regions of Queensland and Northern Territory, extending to Nepal, India, Sri Lanka, South-East Asia and the Pacific islands (Chambers & Farrant 1991). Plants attributed to this species in New Zealand have been re-identified as large forms of C. sieberi. In addition, C. viridis (as Pellaea viridis) was recognised as a casual species in New Zealand by Webb et al. (1995).

 Key
1Primary pinnae bearing abundant scales as well as hairsdistans
Primary pinnae glabrous or bearing only hairs2
2Tertiary pinnae 15–25 mm long, 6–12 mm wide, never strongly curled inwardsviridis
Tertiary pinnae 1–10 mm long, 1–3 mm wide, often strongly inrolledsieberi
 Recognition

In New Zealand, species of Cheilanthes can be recognised by their small to medium-sized 2–3-pinnate fronds, sori extending around the lamina edges protected by the inrolled pinna margins, and scabrate to verrucate spores (Large & Braggins 1991). They occur most frequently in dry, rocky habitats.

 Distribution

Cheilanthes is distributed in Europe, Africa, Asia, Australasia, Oceania and South America, with centres of diversity in South Africa, Australia and South America; 17 species in China (Zhang & Yatskievych 2013), 24 in South Africa (Roux 2009), 15 in Australia (Chambers & Farrant 1998), and five in the south-west Pacific (Nakamura 2008). Three species in New Zealand; two indigenous and one naturalised, none endemic.

 Biostatus
Indigenous (Non-endemic)
Number of species in New Zealand within Cheilanthes Sw.
CategoryNumber
Indigenous (Non-endemic)2
Exotic: Casual1
Total3
 Cytology

Base chromosome numbers of x = 26, 27, 28, 29 and 30 have been recorded in Cheilanthes. There is also an extensive polyploid series from diploid to hexaploid, and several species are known to be apomictic (Tindale & Roy 2002).

 Bibliography
Anthony, N.C. 1984: A revision of the southern African species of Cheilanthes Swartz and Pellaea Link (Pteridaceae). Contributions from the Bolus Herbarium 11: 1–293.
Brown, R. 1810: Prodromus Florae Novae Hollandiae et Insulae Van-Diemen. Johnson, London.
Brownsey, P.J.; Perrie, L.R. 2021: Pteridaceae. In: Breitwieser, I. (ed.) Flora of New Zealand — Ferns and Lycophytes. Fascicle 30. Manaaki Whenua Press, Lincoln.
Chambers, T.C.; Farrant, P.A. 1991: A re-examination of the genus Cheilanthes (Adiantaceae) in Australia. Telopea 4: 509–557.
Chambers, T.C.; Farrant, P.A. 1998: Cheilanthes. In: Flora of Australia. Vol. 48. 271–289.
Cheeseman, T.F. 1925: Manual of the New Zealand Flora. Edition 2. Government Printer, Wellington.
Eiserhardt, W.L.; Rohwer, J.G.; Russell, S.J.; Yesilyurt, J.C.; Schneider, H. 2011: Evidence for radiations of cheilanthoid ferns in the Greater Cape Floristic Region. Taxon 60: 1269–1283.
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.
Large, M.F.; Braggins, J.E. 1991: Spore atlas of New Zealand ferns and fern allies. SIR Publishing, Wellington.
Nakamura, M. (ed.) 2008: Illustrated flora of ferns and fern allies of South Pacific Islands. National Museum of Nature and Science Book Series No. 8. Tokai University Press, Tokyo.
PPG 1 2016: A community-derived classification for extant lycophytes. Journal of Systematics and Evolution 54(6): 563–603.
Quirk, H.; Chambers, T.C.; Regan, M. 1983: The fern genus Cheilanthes in Australia. Australian Journal of Botany 31: 501–553.
Rothfels, C.J.; Windham, M.D.; Grusz, A.L.; Gastony, G.J.; Pryer, K.M. 2008: Towards a monophyletic Notholaena (Pteridaceae): resolving patterns of evolutionary convergence in xeric-adapted ferns. Taxon 57: 712–724.
Roux, J.P. 2009: Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23: 1–296.
Swartz, O.P. 1806: Synopsis Filicum, earum genera et species systematicae complectens. Kiel.
Tindale, M.D.; Roy, S.K. 2002: A cytotaxonomic survey of the Pteridophyta of Australia. Australian Systematic Botany 15: 839–937.
Webb, C.J.; Sykes, W.R.; Garnock-Jones, P.J.; Brownsey, P.J. 1995: Checklist of dicotyledons, gymnosperms, and pteridophytes naturalised or casual in New Zealand: additional records 1988–1993. New Zealand Journal of Botany 33: 151–182.
Yatskievych, G.; Smith, A.R. 2003: Typification of Notholaena R.Br. (Pteridaceae). Taxon 52: 331–336.
Zhang, G.; Yatskievych, G. 2013: Cheilanthes. In: Zhengyi, W.; Raven, P.H.; Deyuan, H. (ed.) Flora of China. Lycopodiaceae through Polypodiaceae. Vol. 2–3. Science Press, Beijing.