Gnaphalieae

Belongs within: Asteroideae.
Contains: Raoulia, Anaphalis, Ozothamnus, Rhodanthe, Podolepis, Angianthus, Craspedia, Calocephalus, Gnaphalium, Cassinia, Helipterum, Helichrysum, Haastia.

The Gnaphalieae are the tribe of composite-flowered plants including the everlastings and related taxa. Heads are mostly heterogamous in members of the subtribes Athrixinae and Gnaphaliinae but homogamous-discoid in Angianthinae.

Members of the Athrixinae have fewer outer female flowers in the heads than inner bisexual flowers with the former being mostly longer than the latter. Athrixia is an Australian and southern African genus with subulate, ciliate involucral bracts (Black & Robertson 1965).

Gnaphaliinae may have more or less outer female than inner bisexual flowers, or may have flowers all bisexual (Black & Robertson 1965). Everlastings such as species of Helichrysum and Helipterum have the flower-head surrounded by an involucre of several rows of papery bracts.

A brief flourish of nest straw

Published 17 March 2025

The desert is a punishing place in which to make a living. Harsh sun, high temperatures, searing winds, all challenge any attempt to construct a viable lifestyle. Desert plants have evolved all sorts of ways to meet these challenges: reduced leaves, succulent stems, deep roots. But some, the desert annuals, have found another way to deal with the harshest of climes: by not being there when they happen.

Stylocline is one genus of such desert annuals. Found in the southwest of North America, Stylocline species, commonly known as nest straws, are small, generally greyish composite-flowered plants with a cobwebby to tometose covering of small hairs (Hickman 1993). One to several stalks arise from the base of the plant, varying in habit from erect to spreading. In the largest species, stalks may grow to a bit over twenty centimetres; others reach little more than ten. Small leaves, about a centimetre to a centimetre and a half in length, are evenly distributed along the stalk. Flower heads are borne in clusters of two to ten and are likewise small, often only a few millimetres in diameter and at most less than a centimetre. Florets within the flower heads are each subtended by erect bracts bearing variously formed scarious wings around at least part of the margin.

Desert annuals such as Stylocline species make their living by sprouting from seed after the winter rains fall. Growth to maturity is rapid. When the season turns and the hot, dry summer weather comes back, the plant will die, but its seeds will remain in the ground to grow into the next generation. The winged bracts may play their part in quick seed production: as they grow, they push the styles of the outer male florets towards the central, functionally female florets, encouraging self-fertilisation (Morefield 1992). This does have the potential disadvantage of making cross-fertilisation apparently rare, but it is presumably better not to risk the possibility that no fertilisation occurs at all.

Seven species of Stylocline were distinguished by Morefield (1992). At least some of these appear likely to have hybrid origins; because of the rarity of cross-pollination events, those that do occur may be more likely to give rise to persistently discrete populations. Because of their small size and short life span, Stylocline species may be rarely collected. One species, S. sonorensis, was described from a single collection in 1950 and not seen again until 1988, at which point it turned out to be widespread in adjoining regions of Nevada and California. However, at least one species of Stylocline appears to be genuinely endangered. Stylocline citroleum is almost entirely restricted to petroleum-rich soils in the southern San Joaquin Valley. Needless to say, this area has been heavily impacted by human activity. In his initial description, Morefield (1992) speculated that this species might already be extinct; it thankfully appears that he was wrong, but the species does remain localised and uncommon.

Systematics of Gnaphalieae

<==Gnaphalieae
    |--AthrixinaeBR65
    |    |--PodolepisM99
    |    `--AthrixiaBR65
    |         |--A. athrixioides (Sonder & von Mueller) Druce 1917 (see below for synonymy)BR65
    |         |    |--A. a. var. athrixioidesBR65
    |         |    `--A. a. var. horripes (Black) Robertson in Black & Robertson 1965 (see below for synonymy)BR65
    |         `--A. croninianaBR65
    |--AngianthinaeBR65
    |    |--Basedowia tenerrima [=Humea tenerrima von Mueller 1896; incl. B. helichrysoides Pritzel 1918]BR65
    |    |--Eriochlamys behriiBR65
    |    |--Gnaphalodes uliginosumBR65
    |    |--Cephalipterum drummondiiBR65
    |    |--AngianthusBR65
    |    |--CraspediaM99
    |    |--Polycalymma stuartiiM99 [=Myriocephalus stuartiiBR65]
    |    |--CalocephalusBR65
    |    |--ChthonocephalusBR65
    |    |    |--C. pseudevaxBR65
    |    |    `--C. tomentellusKM08
    |    |--MyriocephalusBR65
    |    |    |--M. guerinaeK90
    |    |    |--M. occidentalisGK00
    |    |    |--M. rhizocephalus [=Hyalolepis rhizocephala]BR65
    |    |    |    |--M. r. var. rhizocephalaBR65
    |    |    |    `--M. r. var. pluriflorusBR65
    |    |    `--M. rudalliiBR65
    |    `--GnephosisBR65
    |         |--G. arachnoideaBR65
    |         |--G. cyathopappaM99
    |         |--G. eriocarpaBR65
    |         |--G. gnephosioides (von Mueller) Druce 1917 (see below for synonymy)BR65
    |         |--G. macrocephalaK90
    |         |--G. pusillaK90
    |         |--G. skirrophora [incl. G. codonopappa]BR65
    |         `--G. tenuissimaKM08
    `--GnaphaliinaeBR65
         |--Stuartina muelleri Sond. 1852BR65, C06 [incl. Gnephosis rotundifolia Diels 1904BR65]
         |--GnaphaliumA61
         |--CassiniaBR65
         |--HelipterumBR65
         |--HelichrysumBR65
         |--Quinetia urvilleiBR65
         |--Ixodia achilleoidesBR65
         |--HaastiaB93
         |--Pterygopappus lawrenceiB93
         |--RutidosisBR65
         |    |--R. helichrysoidesM99
         |    `--R. multiflora (Nees von Esenbeck) Robinson 1911 (see below for synonymy)BR65
         |--ToxanthusBR65
         |    |--T. muelleriBR65
         |    `--T. perpusillusBR65
         |--Podosperma Labillardière 1806 [incl. Podotheca Cassini 1822]BR65
         |    |--P. angustifolium [=Podotheca angustifolia]BR65
         |    |--‘Podotheca’ chrysanthaRL05
         |    `--‘Podotheca’ gnaphalioidesK90
         |--HumeaBR65
         |    |--H. cassiniaeformis (see below for synonymy)BR65
         |    |--H. pholidota (von Mueller) Black 1919 (see below for synonymy)BR65
         |    `--H. punctulataBR65
         |--IxiolaenaBR65
         |    |--I. brevicomptaM99
         |    |--I. leptolepisBR65
         |    |--I. supinaBR65
         |    |--I. tomentosaM99
         |    `--I. viscosaBR65
         |--Waitzia Wendland 1808BR65
         |    |--W. acuminataM99
         |    |--W. citrinaK90
         |    |--W. corymbosaK90
         |    |--W. nitidaGK00
         |    |--W. podolepisK90
         |    `--W. suaveolensRL05
         |--LeptorhynchosBR65
         |    |--L. elongatusBR65
         |    |--L. medius [incl. L. elongatus var. peduncularis]BR65
         |    |--L. squamatusBR65
         |    |--L. tenuifoliusBR65
         |    |--L. tetrachaetus (Schlechtendal) Black 1921 (see below for synonymy)BR65
         |    `--L. waitziaBR65
         `--Millotia Cassini 1829BR65
              |--M. greevesiiBR65
              |--M. kempeiBR65
              |    |--M. k. var. kempeiBR65
              |    `--M. k. var. helmsii [incl. Toxanthus whitei Black 1915]BR65
              |--M. myosotidifoliaGK00
              `--M. tenuifoliaBR65

Gnaphalieae incertae sedis:
  RaouliaBS97
  AnaphalisBS97
  AliellaBS97
  Apalochlamys spectabilis [=Cassinia spectabilis]BW97
  MniodesBW98
  RaouliopsisBW98
  Leucogenes Beauverd 1910BS97, A61
    |--L. grandiceps (Hooker) Beauverd 1910 (see below for synonymy)A61
    `--L. leontopodium (Hooker) Beauverd 1910 (see below for synonymy)A61
  ×Leucoraoulia Allan 1939A61
    |--‘Haastia’ × loganii Buchan. 1882 (see below for synonymy)A61
    `--‘Helichrysum’ × pauciflorum Kirk 1895 (see below for synonymy)A61
  PhagnalonBS97
    |--P. graecumPT98
    |--P. nitidumDH98
    |--P. purpurascensD17
    `--P. sinaicumDH98
  Pseudognaphalium (subg. Laphangium) luteoalbumBS97, B93 [=Gnaphalium luteo-album Linnaeus 1753B93, A61]
    |--P. l. var. luteoalbumA61
    |--‘Gnaphalium’ l. var. compactum Kirk 1899A61
    `--‘Gnaphalium’ l. var. incanum Rich. 1832A61
  Ewartia Beauverd 1910BS97, A61
    |--*E. catipesA61
    |--E. meredithiaeBS97
    |--E. nubigenaKC83
    |--E. planchoniiBS97
    `--E. sinclairii (Hooker) Cheeseman 1925 [=Gnaphalium (Helichrysum) sinclairii Hooker 1864]A61
  OzothamnusBS97
  RhodantheM99
  ChrysocephalumM99
    |--C. apiculatumB00
    `--C. semipapposumB00
  Lawrenalla davenportiiM99
  LeucochrysumM99
    |--L. fitzgibboniiG04
    `--L. stipitatumM99

Athrixia athrixioides (Sonder & von Mueller) Druce 1917 [=Panaetia athrixioides Sonder & von Mueller 1852; incl. A. tenella Bentham 1866]^BR65

Athrixia athrixioides var. horripes (Black) Robertson in Black & Robertson 1965 [=A. tenella var. horripes Black 1929]^BR65

Gnephosis gnephosioides (von Mueller) Druce 1917 [=Cyathopappus gnephosioides; incl. G. arachnoidea var. foliata]^BR65

‘Haastia’ × loganii Buchan. 1882 [=Helichrysum loganii (Buchan.) Kirk 1899, Raoulia loganii (Buchan.) Cheeseman 1925; Leucogenes leontopodium × Raoulia rubra]^A61

‘Helichrysum’ × pauciflorum Kirk 1895 [Leucogenes grandiceps × Raoulia bryoides; incl. R. gibbsii Cheeseman 1910, H. (Leucogenes) grahamii Petrie 1913]^A61

Humea cassiniaeformis [=Haeckeria cassiniaeformis von Mueller 1855; incl. Humea cassiniacea von Mueller 1858]^BR65

Humea pholidota (von Mueller) Black 1919 [=Ozothamnus pholidotus von Mueller 1861, Helichrysum pholidotum von Mueller 1866; incl. Humea squamata von Mueller 1880]^BR65

Leptorhynchos tetrachaetus (Schlechtendal) Black 1921 [=Doratolepis tetrachaeta Schlechtendal 1847; incl. L. pulchellus von Mueller 1858]^BR65

Leucogenes grandiceps (Hooker) Beauverd 1910 [=Gnaphalium (Helichrysum) grandiceps Hooker 1864, Helichrysum grandiceps Kirk 1899]^A61

Leucogenes leontopodium (Hooker) Beauverd 1910 [=Helichrysum leontopodium Hooker 1853; incl. Gnaphalium (Helichrysum) colensoi Hooker 1864]^A61

Rutidosis multiflora (Nees von Esenbeck) Robinson 1911 [=Styloncerus multiflorus Nees von Esenbeck 1846-1847; incl. Pumilo argyrolepis Schlechtendal 1848, Rutidosis pumilo Bentham 1866]^BR65

*Type species of generic name indicated

References

[A61] Allan, H. H. 1961. Flora of New Zealand vol. 1. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. R. E. Owen, Government Printer: Wellington (New Zealand).

[BR65] Black, J. M., & E. L. Robertson. 1965. Flora of South Australia. Part IV. Oleaceae–Compositae. W. L. Hawes, Government Printer: Adelaide.

[B00] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 2. CSIRO Publishing: Collingwood (Victoria).

[B93] Breitwieser, I. 1993. Comparative leaf anatomy of New Zealand and Tasmanian Inuleae (Compositae). Botanical Journal of the Linnean Society 111: 183–209.

[BS97] Breitwieser, I., & F. B. Sampson. 1997. Pollen characteristics of New Zealand Gnaphalieae (Compositae) and their taxonomic significance. Grana 36: 80–95.

[BW97] Breitwieser, I., & J. M. Ward. 1997. Transfer of Cassinia leptophylla (Compositae) to Ozothamnus. New Zealand Journal of Botany 35: 125–128.

[BW98] Breitwieser, I., & J. M. Ward. 1998. Leaf anatomy of Raoulia Hook. f. (Compositae, Gnaphalieae). Botanical Journal of the Linnean Society 126: 217–235.

[C06] Cheeseman, T. F. 1906. Manual of the New Zealand Flora. John Mackay, Government Printer: Wellington.

[DH98] Danin, A., & I. C. Hedge. 1998. Contributions to the flora of Jordan 2. A new species of Satureja (Labiatae) and some new records. Willdenowia 28: 135–142.

[D17] Diels, L. 1917. Beiträge zur Flora der Zentral-Sahara und ihrer Pflanzengeographie. Nach der Sammelausbeute des Freiherrn Hans Geyr von Schweppenburg. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 54 (Beiblatt 120): 51–155.

[G04] Gibson, N. 2004. Flora and vegetation of the Eastern Goldfields Ranges: part 6. Mt Manning Range. Journal of the Royal Society of Western Australia 87 (2): 35–47.

[GK00] Gibson, N., & G. J. Keighery. 2000. Flora and vegetation of the Byenup-Muir reserve system, south-west Western Australia. CALMScience 3 (3): 323–402.

Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[K90] Keighery, G. J. 1990. Vegetation and flora of Shark Bay, Western Australia. In: Berry, P. F., S. D. Bradshaw & B. R. Wilson (eds) Research in Shark Bay: Report of the France-Australe Bicentenary Expedition Committee pp. 61–87. Western Australian Museum.

[KM08] Keighery, G. J., & W. Muir. 2008. Vegetation and vascular flora of Faure Island, Shark Bay, Western Australia. Records of the Western Australian Museum Supplement 75: 11–19.

[KC83] Körner, C., & P. Cochrane. 1983. Influence of plant physiognomy on leaf temperature on clear midsummer days in the Snowy Mountains, south-eastern Australia. Acta Œcologica Œcologiae Plantae 4 (2): 117–124.

[M99] Matthews, M. 1999. Heliothine Moths of Australia: A guide to bollworms and related noctuid groups. CSIRO Publishing.

Morefield, J. D. 1992. Three new species of Stylocline (Asteraceae: Inuleae) from California and the Mojave Desert. Madroño 39 (2): 114–130.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

[RL05] Rafferty, C., & B. B. Lamont. 2005. Selective feeding by macropods on vegetation regenerating following fire. Journal of the Royal Society of Western Australia 88 (4): 155–165.