Horkelia sect. Horkelia
Rosaceae tribe Potentilleae
decumbent or ascending to erect, (0.5–)1–10(–12) dm.
alternate, rarely opposite, pinnately (palmately) compound (simple in Alchemilla, Aphanes, and Chamaerhodos);
stipules persistent (absent in Chamaerhodos), adnate to petiole;
venation pinnate or palmate.
usually planar, sometimes ± cylindric;
stipules usually entire, sometimes basally lobed;
leaflets (1–)3–16(–20) per side, separate to overlapping, divided ± 1/6–3/4+ to midrib into 3–30(–60) teeth or lobes not restricted to apex.
open to congested, flowers arranged individually, in usually non-capitate glomerules, and/or in corymbiform clusters.
remaining straight, 1–30(–40) mm.
epicalyx bractlets narrowly elliptic-lanceolate to broadly ovate, 0.5–3 mm wide, usually entire, sometimes toothed;
hypanthium interior pilose or glabrous;
sepals acute;
petals white, oblong-oblanceolate to round, apex obtuse to truncate to emarginate;
filaments white, glabrous, anthers longer than wide;
carpels 10–200(–220).
perianth and androecium perigynous;
epicalyx bractlets present, sometimes absent;
hypanthium usually patelliform, cupulate, or campanulate, sometimes turbinate, saucer-shaped, flat-bottomed, or subglobose to ellipsoid or ovoid;
torus flat to conic or turbinate, enlarged (absent or reduced in Alchemilla, Aphanes, and Chamaerhodos);
carpels 1–260, styles basal or lateral to subterminal, distinct;
ovules 1(or 2), basal.
aggregated achenes (achenes in Alchemilla and Aphanes);
torus sometimes fleshy;
styles deciduous or persistent, not elongate.
0.8–2 mm, usually smooth or slightly rugose, sometimes merely roughened.
Horkelia sect. Horkelia
Rosaceae tribe Potentilleae
Species 9 (9 in the flora).
Section Horkelia encompasses the species that are most commonly encountered in heavily populated areas of California. Plants are notably glandular-viscid (unless obscured by dense vestiture) and have a distinctive resinous odor.
Previous revisions (for example, P. A. Rydberg 1908c; D. D. Keck 1938) have placed Horkelia frondosa (here treated as H. californica var. frondosa) at the beginning, implying that this is the least derived expression within the genus. Such an assumption is based on its gross resemblance to sympatric members of Drymocallis; molecular evidence (T. Eriksson et al. 1998; M. Lundberg et al. 2009; C. Dobeš and J. Paule 2010) confirms that this similarity is superficial. If, as speculated above, species composing sect. Hispidulae are relicts of the original radiation, then H. californica var. frondosa is actually one of the more derived members of the genus. Species within sect. Horkelia have been ordered here according to that interpretation.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
Genera 14–22, species ca. 860 (14 genera, 189 species, including 1 hybrid, in the flora area).
The base chromosome number for Potentilleae is mostly x = 7 (8 in Alchemilla and Aphanes; 14 in Comarum).
Variation in the number of genera recognized in Potentilleae is due to differences in generic delimitation between D. Potter et al. (2007) and the authors of Potentilla and segregates here (see 9. Ivesia and 8. Potentilla for discussion). In the former, Duchesnea, Horkelia, Horkeliella, and Ivesia are included within Potentilla. Likewise, Aphanes is included within Alchemilla by Potter et al. while it is kept distinct here.
Potentilla and its segregates and Fragaria are host to Phragmidium rusts, but not the other genera of the tribe.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
1. Leaflets 1–3 per side, sparsely short-pilose to glabrate, (15–)20–30-toothed 1/6–1/4 to midrib; petals obovate to round, (3–)4–7 mm wide. | H. truncata |
1. Leaflets (3–)5–16(–20) per side, hairy, 3–15(–30)-toothed or -lobed (collectively 10–60-toothed in H. californica) 1/6–3/4+ to midrib; petals narrowly obovate or oblanceolate to elliptic or oblong, 1–3(–4) mm wide | → 2 |
2. Hypanthia 3–5.5 mm, 1/2 to nearly as deep as wide; epicalyx bractlets 4–6(–8) mm, ± equal to sepals, entire or toothed. | H. californica |
2. Hypanthia 1–2(–3) mm, less than 1/2 as deep as wide (except H. clevelandii); epicalyx bractlets 1–4(–5) mm, shorter than sepals, entire | → 3 |
3. Leaflets ± elliptic to obovate, divided less than 1/3 to midrib into (5–)10–15 teeth; epicalyx bractlets 1.5–3 mm wide. | H. cuneata |
3. Leaflets cuneate or flabellate to ovate or nearly round, divided (1/5–)1/3–3/4+ to midrib into 3–10(–30 in H. yadonii) teeth or lobes; epicalyx bractlets 0.5–2(–2.5) mm wide | → 4 |
4. Stems ± decumbent to ascending, 1–3.5(–4.5) dm; leaflets 5–10(–12) per side, cuneate, ± overlapping; inflorescences usually congested; plants matted; coastal. | H. marinensis |
4. Stems ascending to erect, (0.5–)1–7 dm; leaflets 6–16(–20) per side, cuneate to flabellate or nearly round, separate to ± overlapping; inflorescences open to ± congested; plants tufted to matted; coastal or interior | → 5 |
5. Epicalyx bractlets 1–2(–2.5) mm wide, lanceolate to ovate; anthers 0.8–1.2 mm; hypanthia 3–6.5 mm diam. | H. yadonii |
5. Epicalyx bractlets 0.5–1(–1.5) mm wide, narrowly elliptic to broadly lanceolate; anthers 0.4–1 mm; hypanthia 2–4.5(–6) mm diam | → 6 |
6. Stems: hairs ascending to appressed; plants usually grayish to grayish green; hypanthium interior pilose | → 7 |
6. Stems: hairs ± spreading; plants green to grayish; hypanthium interior glabrous or pilose | → 8 |
7. Carpels 20–50(–120); styles 2–4 mm; basal leaves (4–)8–20(–30) cm; Transverse Ranges, s California. | H. rydbergii |
7. Carpels 10–20(–27); styles (1–)1.5–2 mm; basal leaves 3–8(–9) cm; Coast Ranges, n California. | H. bolanderi |
8. Leaflets divided 1/2–3/4+ to midrib into linear to narrowly oblanceolate or narrowly elliptic lobes; hypanthium interior pilose; n Coast Ranges, California. | H. tenuiloba |
8. Leaflets divided 1/3 to midrib into acute to obtuse teeth; hypanthium interior glabrous; Peninsular Ranges, s California. | H. clevelandii |
1. Shrubs; leaf lobe margins entire; achenes hirsute. | Dasiphora |
1. Herbs, perennial, sometimes annual or biennial, or subshrubs; leaf lobe margins or apices ± toothed, sometimes entire; achenes glabrous (sometimes ± hairy) | → 2 |
2. Petals 0, sepals 4; achenes 1, enclosed in dry, urceolate or subglobose to ellipsoid or ovoid hypanthia | → 3 |
2. Petals and sepals usually 5; achenes 1–260, usually aggregated (sometimes on elongating tori), usually in (± open) patelliform, cupulate, campanulate, or turbinate hypanthia (not enclosed in dry hypanthium) | → 4 |
3. Herbs perennial; leaves basal, blades reniform to orbiculate, palmately lobed, sometimes palmately compound; stamens 4. | Alchemilla |
3. Herbs annual; leaves cauline, blades cuneate, deeply divided into segments, each lobed; stamen 1(or 2). | Aphanes |
4. Leaves all or mostly basal or proximal (if cauline, deeply pinnatifid), ternate or 2–4-ternate (sometimes simple and coarsely toothed apically in Sibbaldia) | → 5 |
4. Leaves basal or cauline, the latter usually reduced distally, odd-pinnate to palmate, rarely ternate or ± bipinnate | → 9 |
5. Tori becoming red and fleshy in fruit; leaf margins serrate to crenate | → 6 |
5. Tori hemispheric (not enlarged or fleshy) in fruit or absent; leaf margins entire or (2–)3(–5)-toothed apically | → 7 |
6. Leaves ± doubly serrate or crenate; stolons leafy; inflorescences: flowers solitary, axillary at stolon nodes; petals yellow. | Duchesnea |
6. Leaves serrate to crenate; stolons not leafy; inflorescences 1–10-flowered, cymes, axillary from leaf rosettes; petals usually white. | Fragaria |
7. Leaves pinnately compound or simple and deeply pinnatifid, margins entire, stipules absent. | Chamaerhodos |
7. Leaves ternate, margins toothed apically, stipules persistent | → 8 |
8. Petals ± yellow; stamens 5. | Sibbaldia |
8. Petals usually white; stamens 20(–30). | Sibbaldiopsis |
9. Petals deep red to purple, rarely pink, shorter than sepals; tori enlarged and spongy at maturity; horizontal stems sometimes floating, wetland habitats. | Comarum |
9. Petals yellow to white, rarely pink or red (then equal to or longer than sepals); tori not enlarged and spongy at maturity; stems erect to decumbent, not horizontal or floating even if in wetlands | → 10 |
10. Anthers dehiscing by continuous marginal slit (with a single theca); styles sub-basal. | Drymocallis |
10. Anthers dehiscing longitudinally; styles subterminal to lateral | → 11 |
11. Hypanthium patelliform to campanulate or cupulate to turbinate (not flat-bottomed); filaments not forming tube; petals white to yellow, sometimes reddish or pink tinged | → 12 |
11. Hypanthium ± cupulate or bluntly campanulate and flat-bottomed; filaments forming tube; petals usually white, sometimes pink-tinged, rose-veined, or cream | → 13 |
12. Plants not aromatic; leaves ± cordate or reniform to narrowly elliptic in outline, leaflets 3–15(–41); petals oblanceolate or obovate to obcordate to nearly round, rarely elliptic; carpels 3–260. | Potentilla |
12. Plants often aromatic; leaves planar to cylindric, leaflets (3–)7–161; petals linear or narrowly oblanceolate to obovate, sometimes obcordate; carpels 1–20(–40). | Ivesia |
13. Stamens 10; leaflets (3–)5–41. | Horkelia |
13. Stamens 20; leaflets 30–70. | Horkeliella |
