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Norwegian cinquefoil, Pennsylvania cinquefoil, pennsylvanian cinquefoil, potentille de pennsylvanie, prairie cinquefoil

Habit Perennials, ± tufted, not stoloniferous; taproots not fleshy-thickened; vestiture of long, short, crisped, and/or cottony hairs, glands sparse to abundant, sometimes absent, not red.
Stems

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.

ascending to erect, (0.3–)1–4(–6) dm.

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.

Basal leaves

pinnate, (2–)4–20(–25) cm;

petiole 1–8 cm, long hairs sparse to abundant, rarely absent, spreading to ascending, 0.5–3 mm, weak to ± stiff, short hairs abundant to dense, rarely absent, cottony or crisped hairs absent, glands sparse to abundant, often obscured;

leaflets (2–)3–5(–7) per side, on distal 1/2–3/5 of leaf axis, separate to scarcely overlapping, terminal ones narrowly to broadly oblanceolate, (1–)1.5–5(–6) × 0.5–1.5(–2) cm, margins strongly revolute, incised ± 3/4 to midvein, sometimes more or only ± 1/2, undivided medial blade 2–4(–8) mm wide, teeth 4–8(–10) per side, triangular-ovate to linear-elliptic, surfaces usually ± dissimilar, abaxial ± grayish to nearly white, rarely green, long hairs sparse to abundant (mostly on veins), rarely absent, 0.5–1.5 mm, ± weak, short hairs usually abundant to dense, rarely absent, crisped and cottony hairs absent, glands ± abundant, sometimes obscured, adaxial green to gray-green, long hairs sparse to abundant, rarely absent, spreading to ascending, ± 1 mm, short hairs ± abundant, sometimes obscured, rarely absent, cottony and crisped hairs absent, glands ± abundant.

Cauline leaves

1–4.

Inflorescences

(1–)2–40(–100)-flowered, cymose, congested to very open.

(1–)3–40-flowered, congested or elongating in fruit.

Pedicels

± straight in fruit, (0.1–)0.2–3(–5) cm, proximal often significantly longer than distal.

0.2–1 cm (proximal to 3.5 cm).

Flowers

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.

epicalyx bractlets usually ± lanceolate-elliptic, rarely ± ovate-elliptic, 3–6(–8) × (1–)1.5–2(–2.5) mm, lengths ± equal to sepals, margins ± revolute;

hypanthium 3–6 mm diam.;

sepals (3–)4–6(–8) mm, apex acute to obtuse, abaxial surfaces: venation indistinct, glands abundant, ± evident;

petals pale to bright yellow, (3–)4–7 × (3–)4–7 mm, lengths ± equal to sepals;

filaments (0.5–)1–2 mm, anthers 0.5–0.8(–1) mm;

carpels 50–100, styles papillate-swollen in proximal ± 1/2, ± 1 mm.

Achenes

smooth to rugose.

1.2 mm, ± to strongly rugose.

2n

= 28.

Potentilla sect. Pensylvanicae

Potentilla pensylvanica

Phenology Flowering summer.
Habitat Open prairies, steppe bluffs, shallow gravelly soil, rocky ridges and outcrops, dry stream channels, similar disturbed sites
Elevation 100–4000 m (300–13100 ft)
Distribution
North America; ne Mexico; Eurasia; n Africa
from FNA
AK; AZ; CA; CO; IA; ID; MN; MT; ND; NE; NM; NV; SD; TX; UT; WY; AB; BC; MB; NT; ON; QC; SK; YT; Mexico (Tamaulipas); Eurasia
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Discussion

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.)

Even in the reduced circumscription used here, Potentilla pensylvanica is exceptionally widespread, occurring in North America from Alaska to Hudson Bay, in the Great Plains and the North American Cordillera to northern Mexico. The species also grows across Siberia (where it has sometimes been called P. sibirica) and as isolated occurrences in the southwestern Alps and Spain (J. Soják 1987c; A. Kurtto et al. in J. Jalas et al. 1972+, vol. 13). One place P. pensylvanica does not occur is in Pennsylvania; the epithet was applied by Linnaeus to cultivated plants originating from Canada. Collections from Quebec are probably introductions (J. Cayouette, pers. comm.). The Oregon citation of P. pensylvanica (D. Mansfield 2000) refers to P. jepsonii.

Significant variation occurs within Potentilla pensylvanica, especially regarding vestiture and plant stature. J. Soják (1987c) has described several varieties, all Eurasian except var. brevipila (long hairs 0.6–1 mm); diminutive specimens from the Brooks Range of northern Alaska, isolated from other known populations of P. pensylvanica, have been described as P. brooksensis. Although much of the ecogeographic variation may merit recognition, at least at varietal level, doing so is postponed pending further analysis and clarification of nomenclature. Among the extremes are plants from the Great Plains, which have abundant long silky hairs overlying and obscuring the velvety layer of short hairs (resembling sympatric P. litoralis), and glabrate (except for glands) populations in Alaska. E. Hultén (1968) erroneously referred to the latter as P. pensylvanica var. glabrata (Lehmann) S. Watson; that name is a synonym of P. ovina var. ovina (sect. Multijugae). Most populations in Alaska also have larger than average flowers (petals to 7 × 7 mm, anthers ca. 1 mm).

Although Potentilla pensylvanica var. strigosa Pursh has been one of the most commonly used varieties in North America, the name has been effectively lectotypified by J. Soják (1987c) on a collection from Siberia. Soják identified this collection as the hybrid between P. pensylvanica and the Asian P. sanguisorba D. F. K. Schlechtendal; var. strigosa accordingly does not occur in North America.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
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
Source FNA vol. 9, p. 211. Authors: Barbara Ertter, Reidar Elven. FNA vol. 9, p. 214.
Parent taxa Rosaceae > subfam. Rosoideae > tribe Potentilleae > Potentilla Rosaceae > subfam. Rosoideae > tribe Potentilleae > Potentilla > sect. Pensylvanicae
Sibling taxa
P. albiflora, P. ambigens, P. anachoretica, P. angelliae, P. anglica, P. anserina, P. arenosa, P. argentea, P. arizonica, P. basaltica, P. bicrenata, P. biennis, P. biflora, P. bimundorum, P. bipinnatifida, P. brevifolia, P. breweri, P. bruceae, P. brunnescens, P. canadensis, P. concinna, P. cottamii, P. crantzii, P. crebridens, P. crinita, P. cristae, P. demotica, P. drummondii, P. effusa, P. elegans, P. erecta, P. flabellifolia, P. fragiformis, P. furcata, P. glaucophylla, P. gracilis, P. grayi, P. hickmanii, P. hippiana, P. holmgrenii, P. hookeriana, P. hyparctica, P. inclinata, P. intermedia, P. jepsonii, P. johnstonii, P. lasiodonta, P. litoralis, P. macounii, P. millefolia, P. modesta, P. morefieldii, P. multijuga, P. multisecta, P. nana, P. newberryi, P. nivea, P. norvegica, P. ovina, P. paucijuga, P. pedersenii, P. plattensis, P. pseudosericea, P. pulchella, P. pulcherrima, P. recta, P. reptans, P. rhyolitica, P. rimicola, P. rivalis, P. robbinsiana, P. rubella, P. rubricaulis, P. sanguinea, P. saximontana, P. sierrae-blancae, P. simplex, P. sterilis, P. stipularis, P. subgorodkovii, P. subjuga, P. subvahliana, P. subviscosa, P. supina, P. thurberi, P. thuringiaca, P. tikhomirovii, P. townsendii, P. uliginosa, P. uschakovii, P. vahliana, P. verna, P. versicolor, P. villosa, P. villosula, P. vulcanicola, P. wheeleri
Subordinate taxa
P. anachoretica, P. bimundorum, P. bipinnatifida, P. jepsonii, P. lasiodonta, P. litoralis, P. pensylvanica, P. pulchella
Synonyms P. unranked Multifidae, P. section Multifidae, P. ser. P. atrovirens, P. brooksensis, P. pensylvanica var. arida, P. pensylvanica var. brevipila, P. sibirica
Name authority Poeverlein: in P. F. A. Ascherson et al., Syn. Mitteleur. Fl. 6(1): 669. (1904) Linnaeus: Mant. Pl. 1: 76. (1767)
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