Handeliinae K. Bremer & Humphries

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 : 128

8. Handeliinae K. Bremer & Humphries in Bull. Nat. Hist. Mus. London, Bot. 23: 108. 1993. – Type: Handelia Heimerl ( Handelia trichophylla (Schrenk) Heimerl ). = Cancriniinae K. Bremer & Humphries in Bull. Nat.

Hist. Mus. London, Bot. 23: 96. 1993. – Type: Cancrinia Kar. & Kir. ( Cancrinia chrysocephala Kar. &

Kir.).

Description — Subshrubs or annual, biennial (hapaxanthic) or perennial (pollacanthic) herbs. Indumentum absent or of basifixed, rarely of medifixed hairs (some species of Tanacetopsis and Xylanthemum ). Leaves alternate, lobed or apically trifid, 1- or 2-pinnatipartite to 3- or 4-pinnatisect. Capitula solitary, in lax to dense corymbs or in a long panicle ( Lepidolopsis ), discoid or radiate. Involucre hemispheric to spheric or obconic ( Trichanthemis ), sometimes umbonate ( Sclerorhachis ). Phyllaries in 2–4(or 5) rows, with narrow to broad scarious margins. Receptacle hemispheric, epaleate or paleate ( Handelia , Sclerorhachis ); paleae subulate to linear, flat to canaliculate, sometimes readily deciduous ( Sclerorhachis ). Ray florets female or neuter; limb white, yellow, pink or violet. Disc florets hermaphrodite; corolla 5-lobed, sometimes 4–6-lobed ( Lepidolopsis ), sometimes hairy ( Tanacetopsis , Trichanthemis ); anthers with non-polarized endothecial tissue and a balusterform filament collar (slender in Allardia and some Tanacetopsis species). Achenes cylindric to obconic, circular to elliptic in cross-section, with 4–10(–15) ribs, sometimes with 3–5 adaxially arranged ribs ( Microcephala ); apex with a corona formed by 5–20 ( Cancrinia , Ugamia ) or 25–50 bristle-like scales ( Allardia ), a short rim or of laciniate scales of various shapes, rarely ecoronate ( Pseudohandelia , Sclerorhachis ); pericarp with or without myxogenic cells, without resin sacs, glabrous or rarely sparsely to densely hairy ( Allardia , Cancrinia , Trichanthemis , Ugamia ). Embryo sac development unknown. Base chromosome number x = 7 ( Microcephala ), 9.

Distribution — Iran, Afghanistan, Pakistan, C Asia, Mongolia, China.

Members — Allardia Decne. (4), Cancrinia Kar. & Kir. (4), Cancriniella Tzvelev (1), Handelia Heimerl (1), Lepidolopsis Poljakov (1), Microcephala Pobed. (5), Polychrysum (Tzvelev) Kovalevsk. (1), Pseudohandelia Tzvelev (1), Richteria Kar. & Kir. (6), Sclerorhachis

(Rech. f.) Rech. f. (4), Tanacetopsis (Tzvelev) Kovalevsk. (21), Trichanthemis Regel & Schmalh. (9), Ugamia Pavlov (1), Waldheimia Kar. & Kir. (4).

Notes — The phylogeny of the subtribe was studied in detail by Oberprieler & al. (2019) in a multi-locus approach based on both plastid and nuclear markers. Incongruence among the marker-wise gene-tree reconstructions was interpreted there as a result of ample hybridization having occurred in the evolutionary history of the members of Handeliinae . While generic circumscriptions were unaddressed by the mentioned study due to the limited sampling of representatives from the species-rich genera, the exclusion of Xylanthemum from the subtribe and its transfer to Anthemidinae as proposed earlier by Sonboli & al. (2012) and the polyphyly of Allardia and its classification as two independent genera ( Allardia and Waldheimia ) received good support from these phylogenetic reconstructions. There is still a lack of information for the subtribal placement of the unispecific genus Cancriniella Tzvelev. However , the close relationship to Cancrinia and Trichanthemis karataviensis (the type of Trichanthemis ), as proposed by Poljakov (1959) by uniting them under a more comprehensive genus Cancrinia , argues for the classification of Cancriniella in Handeliinae and awaits support from a molecular phylogenetic analysis of this C Asian endemic.