Athanasiinae Pfeiff., Nomencl. Bot.

Oberprieler, Christoph, Töpfer, Alisha, Dorfner, Marco, Stock, Miriam & Vogt, Robert, 2022, An updated subtribal classification of Compositae tribe Anthemideae based on extended phylogenetic reconstructions, Willdenowia 52 (1), pp. 117-149 : 126

5. Athanasiinae Pfeiff., Nomencl. Bot. 1: 323. 1872. – Type: Athanasia L. ( Athanasia crithmifolia ( L.) L.) .

Description — Shrubs or shrublets, rarely perennial to annual herbs ( Adenoglossa , Lasiospermum ). Indumentum absent or of basifixed or stellate ( Athanasia , Hymenolepis ) hairs. Leaves alternate or opposite, entire or lobed to pinnatifid or 1- or 2-pinnatisect. Capitula solitary or in lax to dense corymbs, radiate, disciform or discoid. Involucre hemispheric, spheric to urceolate. Phyllaries in 2–5 rows, without or with scarious margins, with central resin canals or sacs (in Eriocephalus in 2 unequal rows, outer phyllaries with very wide scarious margins, inner ones connate and hairy). Receptacle flat, hemispheric to conic, paleate or epaleate ( Adenoglossa , Leucoptera ); paleae flat or canaliculate, rarely villous ( Eriocephalus ). Ray florets female; limb yellow, white or reddish. Disc florets hermaphrodite (male in Eriocephalus ); corolla 5-lobed; tube sometimes with long stalked hairs ( Athanasia ); anthers with polarized endothecial tissue, rarely non-polarized ( Eriocephalus ), and a slender filament collar. Achenes cylindric to obovate, either terete and with 5–12(–18) ribs or dorsiventrally flattened and laterally winged ( Adenoglossa , Leucoptera ); apex marginally rounded, with a short, thickened rim ( Athanasia ) or with a corona or scales ( Adenoglossa , Hymenolepis , Leucoptera ); pericarp glabrous or densely hairy ( Eriocephalus , Lasiospermum ), with or without myxogenic cells and/or resin sacs. Embryo sac development monosporic (only known in Lasiospermum ). Base chromosome number x = 8, 9.

Distribution — South Africa, Namibia, Botswana, Lesotho, Egypt (Sinai), Israel, Jordan.

Members — Adenoglossa B. Nord. (1), Athanasia L. (41), Eriocephalus L. (32), Hymenolepis Cass. (7), Lasiospermum Lag. (4), Leucoptera B. Nord. (3).

Notes — The subtribe Athanasiinae in its present circumscription is surely the most problematic generic assemblage in the subtribal classification of the tribe presented here. The present circumscription corresponds completely with the one proposed in our former treatment ( Oberprieler & al. 2007, 2009), where its non-monophyletic nature was discussed to some extent. Our present phylogenetic reconstructions do not support a monophyly of the Athanasiinae —neither the gene trees with representatives of all six mentioned genera (Fig. 1, 2) nor the species trees with the reduced set of those four genera, for which a complete data matrix was achieved ( Fig. 3, 4). This is paralleled by the quite diverse morphological circumscription of the subtribe in itself. Nevertheless, we presently refrain from a decomposition of this possibly paraphyletic subtribe until a more complete sampling and a more reliable phylogenetic reconstruction is available. Species numbers of the present account were updated following taxonomic work done by Powell & Magee (2013) in Athanasia and by Magoswana & Magee (2014) and Magoswana & al. (2015) in Hymenolepis .