Cotulinae Kitt., Taschenb. Fl. Deutschl.

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 : 124-125

2. Cotulinae Kitt., Taschenb. Fl. Deutschl. , ed. 2, 2: 609. 1844. – Type: Cotula L. ( Cotula coronopifolia L.) .

= Thaminophyllinae K. Bremer & Humphries in Bull. Nat. Hist. Mus. London, Bot. 23: 144. 1993. – Type: Thaminophyllum Harv. ( Thaminophyllum mundtii Harv. ).

Description — Shrubs, subshrubs, perennial or annual herbs ( Cotula , Leptinella , Soliva ). Indumentum of basifixed hairs or absent. Leaves alternate or opposite, entire, lobed, pinnatifid to 1- or 2-pinnatisect. Capitula solitary or in lax to dense corymbs, radiate, disciform or discoid. Involucre broadly campanulate, hemispheric to cylindric or obconic, sometimes ( Leptinella ) umbonate. Phyllaries in 2–4 rows, without ( Lidbeckia ) or with narrow to broad scarious margins, sometimes ( Cotula ) with central resin ducts. Receptacle flat to hemispheric or conic, glabrous or hairy ( Lidbeckia , Thaminophyllum ), epaleate or with few marginal paleae ( Schistostephium ). Ray florets and outer disc florets (when present) female, rarely sterile or neuter ( Lidbeckia ), sometimes stalked ( Cotula ); limb white or yellow, rarely pilose ( Inezia ), sometimes confluent with achene and tube short or absent ( Adenanthellum , Inezia , Thaminophyllum ). Disc florets hermaphrodite or functionally male ( Hippia , Leptinella , Schistostephium , Soliva ); corolla 3-, 4- or sometimes 5-lobed ( Adenanthellum , Hippia ); anthers with non-polarized endothecial tissue and a slender filament collar; stylopodium sometimes large and persistent in fruit ( Lidbeckia , Thaminophyllum ). Achenes oblong to obovoid, terete with 2 or 3 adaxial or 3–10 ribs, sometimes 3- or 4-angled, often dorsiventrally compressed with 2 lateral, wing-like ribs; apex truncate or marginally rounded, ecoronate, rarely with minute scales ( Inezia ); pericarp with or without myxogenic cells and/or resin canals in ribs, sometimes papillose or hairy ( Cotula , Hippia ). Embryo sac development monosporic (only known in Cotula ). Base chromosome numbers x = 8, 9, 10, 13.

Distribution — S and E Africa, Australia, New Guinea, New Zealand, South America, S oceanic islands; some species widespread and naturalized as weeds.

Members — Adenanthellum B. Nord. (1), Cotula L. (58), Hilliardia B. Nord. (1), Hippia L. (8), Inezia E. Phillips (2), Leptinella Cass. (33), Lidbeckia P. J. Bergius (3), Schistostephium Less. (12), Soliva Ruiz & Pav. (8), Thaminophyllum Harv. (3).

Notes — The circumscription of this subtribe with its ten genera proposed by Oberprieler & al. (2007) is corroborated by all present phylogenetic reconstructions except the coalescent-based species-tree reconstruction ( Fig. 4), where Hippia is associated with members of subtribe Pentziinae . Since this position and the reciprocal monophyly of each of both subtribes ( Cotulinae and Pentziinae ) is not supported by posterior probabilities, we think that any hypothesis of a non-monophyly of the subtribe is unsubstantiated. Species numbers given for Cotula and Leptinella are only approximations due to the lack of recent and/or comprehensive revisions of the two genera ( Leptinella : Lloyd & Webb 1987; Himmelreich & al. 2012, 2014; Cotula : Powell & al. 2014; Jakoet & al. 2019).