Potentilla sect. Pensylvanicae
Rosaceae tribe Potentilleae
decumbent to erect, not flagelliform, not rooting at nodes, lateral to persistent basal rosettes, (0.1–)0.5–4(–6) dm, lengths (1–)1.5–3(–4) times basal leaves.
basal not in ranks;
cauline 1–4;
primary leaves pinnate to subpinnate or subpalmate (with distal leaflets distinct), 1.5–25(–30) cm;
petiole: long hairs spreading to appressed, soft to stiff, glands absent or sparse to abundant;
leaflets (3–)5–13(–19), on distal 1/6–1/2+, rarely less, of leaf axis, separate to ± overlapping, narrowly oblanceolate or elliptic to obovate or broadly oblong, margins ± to strongly revolute, ± whole length evenly incised ± 1/2 to nearly completely to midvein, teeth (2–)3–8(–12) per side, surfaces similar to strongly dissimilar, abaxial green to white, cottony hairs absent or sparse to dense, adaxial green to grayish, not glaucous, long hairs soft to ± stiff.
alternate, rarely opposite, pinnately (palmately) compound (simple in Alchemilla, Aphanes, and Chamaerhodos);
stipules persistent (absent in Chamaerhodos), adnate to petiole;
venation pinnate or palmate.
(1–)2–40(–100)-flowered, cymose, congested to very open.
± straight in fruit, (0.1–)0.2–3(–5) cm, proximal often significantly longer than distal.
5-merous;
hypanthium 2.5–10 mm diam.;
petals pale to bright yellow, obovate to obcordate, (2–)3–5(–10.5) mm, +\- equal to or longer, sometimes shorter, than sepals, apex rounded or truncate to retuse;
stamens (15–)20;
styles subapical, conic to columnar, papillate-swollen in proximal 1/4–3/4+ or only at base, if at all, 0.8–1.2(–1.5) mm.
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.
smooth to rugose.
Potentilla sect. Pensylvanicae
Rosaceae tribe Potentilleae
Species ca. 20 (8 in the flora).
Section Pensylvanicae, as here presented, accommodates most perennial species with pinnate to subpinnate leaves and styles ca. 1 mm that are frequently papillate-swollen proximally. Plants are often notably glandular, leaflet margins are usually revolute, and petals are most commonly about the same length as the sepals and often somewhat erect at anthesis. Section Multifidae was commonly used for this group of species until J. Soják (1987b) noted that sect. Pensylvanicae was an older, validly published name.
About half of the North American members of sect. Pensylvanicae have been previously included in a broadly defined Potentilla pensylvanica, with or without infraspecific divisions (for example, C. L. Hitchcock and A. Cronquist 1961b; S. L. Welsh 1974; R. L. McGregor 1986b; H. A. Gleason and Cronquist 1991). In support of a more narrowly circumscribed P. pensylvanica, B. L. Kohli and J. G. Packer (1976) found that major morphologic types coincide with three different ploidy levels. Most plants with fully pinnate leaves (five to nine leaflets in remote pairs) and other characters of typical P. pensylvanica in the narrow sense are tetraploid; plants with subpinnate leaves (mostly five leaflets in approximate pairs) are octoploid (or sometimes tetraploid; Packer, pers. comm.). Diploid populations from the northern Great Plains, morphologically closest to P. pensylvanica in the narrow sense but with epicalyx bractlets usually longer than the sepals, were named P. finitima Kohli & Packer (a later synonym of P. lasiodonta).
The ploidy differences noted by B. L. Kohli and J. G. Packer (1976), in combination with morphology (vestiture, glandularity, leaf dissection, and features of the epicalyx) and ecogeographic correlates, serve collectively to delimit the species that are recognized here. The most distinctive vestiture type is a velvety covering of short (0.2 mm), straight, spreading hairs that generally characterizes Potentilla lasiodonta and P. pensylvanica in the narrow sense, although this feature is sometimes obscured by longer ascending hairs or is missing altogether. Leaves of P. lasiodonta and P. pensylvanica in the narrow sense are also distinctly pinnate, with the blade comprising at least half the total leaf length and being no more than about half as wide as long, usually bearing at least three pairs of lateral leaflets gradually reduced in size from the distalmost, and the leaflets themselves being pinnately toothed with 4–12 blunt teeth per side. Epicalyx bractlets in both species, and also in P. litoralis, tend to be nearly as long as the sepals (to twice as long in P. lasiodonta) and concave, at least in fruit, with revolute margins and prominent midveins. These diagnostic features are not consistently present and/or evident, especially in depauperate and immature specimens, but collectively they serve to identify most specimens with a reasonable level of confidence. Occasional hybrids are to be expected.
In addition to the species addressed here, Potentilla lyngei Jurtzev & Soják subsp. lyngei has been reported from northeastern Greenland. At least one purported voucher (annotated by Jurtzev in ALA) has been reidentified to P. pedersenii (sect. Rubricaules), leaving the North American presence of P. lyngei subsp. lyngei in question. This is not the case for P. lyngei subsp. spissa Soják, described from Greenland; as discussed elsewhere (B. Ertter et al. 2013), this name is probably based on hybrids between P. pulchella and P. hyparctica (sect. Aureae), for which P. ×safronoviae Jurtzev & Soják is the priority name.
Additional nomenclatural details of species in sect. Pensylvanicae were provided by M. L. Fernald (1935), J. Soják (1987c), and B. Ertter (2008). For species of presumed hybrid origin with sect. Pensylvanicae as one parent, see sect. Rubricaules and Ertter et al. (2013).
(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. Styles papillate-swollen at very base, if at all; inflorescences ± congested to very open | → 2 |
1. Styles papillate-swollen in proximal 1/4–3/4+; inflorescences usually congested, sometimes elongating in fruit (± open in P. pulchella) | → 3 |
2. Sepal venation prominent at least proximally; inflorescences (5–)10–40+-flowered, ± congested to open; pedicels 0.3–1 cm (proximal to 3 cm). | P. bimundorum |
2. Sepal venation indistinct; inflorescences 2–5-flowered, very open; pedicels 2–4 cm. | P. anachoretica |
3. Basal leaves pinnate, short hairs usually abundant to dense, rarely absent, sometimes obscured (especially in Great Plains); leaflets (2–)3–6(–9) per side, on distal (1/3–)1/2–3/5 of leaf axis; cottony and crisped hairs absent abaxially; epicalyx bractlets: lengths 1–2 times sepals, margins ± revolute | → 4 |
3. Basal leaves subpinnate to subpalmate, short hairs absent, sparse, or obscured; leaflets (1–)2–3 per side, on distal 1/6–1/3(–1/2) of leaf axis; cottony and/or crisped hairs absent or sparse to dense abaxially; epicalyx bractlets: lengths 1/2–1 times sepals, margins revolute or not | → 5 |
4. Leaflets: teeth 4–8(–10) per side, margins incised ± 3/4 to midvein, leaving undivided medial blade 2–4(–8) mm wide; epicalyx bractlets equal to length of sepals; widespread. | P. pensylvanica |
4. Leaflets: teeth 8–12 per side, margins incised ± 1/2 to midvein, leaving undivided medial blade 5–9 mm wide; epicalyx bractlets 1–2 times length of sepals; sandy sites, n Great Plains. | P. lasiodonta |
5. Inflorescences ± open, (1–)2–5(–9)-flowered; pedicels 0.5–5 cm; leaflet teeth 2–5 per side; Alaska, n Canada, Greenland. | P. pulchella |
5. Inflorescences congested, sometimes elongating in fruit, (1–)3–50(–100)-flowered; pedicels 0.2–1(–3) cm; leaflet teeth (2–)3–8 per side; Alaska to Newfoundland, s to California, Colorado, and New Hampshire | → 6 |
6. Petals (4–)5–6 × 4.5–5.5 mm, longer than sepals; styles 1.2–1.5 mm; leaflets incised 2/3–3/4 to midvein; stems 0.5–2(–2.5) dm; La Sal Mountains, Utah. | P. paucijuga (sect. Rubricaules) |
6. Petals 2–5 × 2–4 mm, ± equal to sepals; styles 0.8–1.2(–1.5) mm; leaflets incised 3/4+ to midvein; stems (0.2–)0.5–5(–6) dm; widespread | → 7 |
7. Sepals: glands absent, sparse, or obscured; leaflets usually white abaxially, cottony (and sometimes crisped) hairs ± dense. | P. bipinnatifida |
7. Sepals: glands abundant, evident at least distally; leaflets usually green to grayish abaxially, cottony hairs absent, crisped hairs absent or sparse to abundant | → 8 |
8. Epicalyx bractlets: lengths 2/3–1 times sepals, margins ± revolute (at least in fruit); stems (0.4–)1–4.5(–6) dm; n Atlantic Coast and Hudson Bay across n prairies to n Rocky Mountains; 0–2200 m. | P. litoralis |
8. Epicalyx bractlets: lengths 1/2–3/4 times sepals, margins flat; stems (0.2–)0.5–2.6 dm; Great Basin and surrounding mountains; 2500–3700 m. | P. jepsonii |
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 |
