Alchemilla
Sp. Pl. 1: 123. 1753.
Gen. Pl. ed. 5, 58. 1754.
Herbs, perennial, 0.2–4(–8) dm; rhizomatous, often forming woody stock. Stems 1–6(–10), decumbent to ascending or erect, usually pubescent or puberulent, rarely glabrous. Leaves not persistent, basal in rosette, alternate, usually simple, sometimes palmately compound; stipules persistent, adnate to petiole, proximally connate opposite petiole (abpetiolar fusion), semiorbiculate to ovate, margins lobed or dentate; petiole present; blade (of simple leaves) reniform to orbiculate, sometimes reniform-orbiculate, 1–15 × 1.5–15 cm, usually palmately 7–9(–11)-lobed, sometimes palmately compound with 5–7 leaflets, margins flat or undulate, crenate to dentate, venation palmate, that of lobes or leaflets pinnate, surfaces usually pubescent or glabrous, sometimes sericeous; cauline: stipules relatively large, variously colored, becoming brownish at maturity or on drying, toothed or lobed, petiole shorter, lobes 3–7(–11), often shallow. Inflorescences terminal, multi-flowered, cymes, much-branched; flowers ± aggregated into clusters of 5–15, glabrous or densely pubescent; bracts usually absent; bracteoles absent. Pedicels present. Flowers 2–4 mm diam.; epicalyx bractlets 4; hypanthium urceolate, 1.5–2.5 mm, glabrous or sparsely to densely pubescent; sepals 4, usually erect to spreading, rarely reflexed (A. glomerulans), ovate, glabrous or pubescent; petals 0; stamens 4, inserted outside nectar-secreting receptacular disc almost closing hypanthium, alternating with sepals, anthers often poorly developed; torus absent; carpel 1 (rarely, A. filicaulis, lowermost flower in a monochasium with 2), styles basal, stigmas capitate, ± exserted from hypanthia; ovules 1(–2). Fruits achenes, 1, ovoid, 2–2.5 mm, smooth; hypanthium persistent; sepals persistent, spreading or erect; styles persistent. x = 8.
Distribution
North America, Eurasia, Africa.
Discussion
Species 800–1000 (12 in the flora).
The occurrence of an additional species, not included in the account below, has recently been recognized. This is Alchemilla acutiloba Opiz naturalized near St. John's, Newfoundland. In the key below, it would most likely be identified as A. monticola. It is most readily distinguished by its almost triangular leaf lobes with straight sides and acute triangular teeth. It has yellowish green to grass green leaves. Unlike A. xanthochlora, the adaxial leaf surface is hairy but much less so than that of A. micans or A. monticola. In addition, it differs from A. micans in the rounded hypanthium base and in the stipules being only slightly wine red suffused, and from A. monticola by the hypanthium being glabrous or rarely sparsely hairy proximally.
The majority of Alchemilla species, including all those in the flora area, are agamospermic, high polyploids (to 28-ploid). Most occurrences in North America are as naturalized introductions. Only four species, restricted to Greenland, St. Pierre and Miquelon, and Atlantic coastal regions of Canada, may be native. Some of the introduced species that occur as scattered meadow and hedgerow plants in their native range grow in eastern Canada in large patches as aggressive weeds.
The name Alchemilla vulgaris Linnaeus, used in some North American accounts, was applied in a broad sense to include all the species of sect. Alchemilla, those being all the species occurring in the flora area except A. alpina (sect. Alpinae Camus) and A. mollis and A. venosa (sect. Erectae Fröhner). The type of A. vulgaris represents a mixture of A. glaucescens and A. wichurae (both with very limited occurrences in North America) and the name now is either informally rejected as confused or applied, also informally, in an aggregate sense.
The commonly cultivated species of Alchemilla all fall within sect. Erectae, distinguished from sect. Alchemilla (A. vulgaris of authors) by the epicalyx bractlets being as long as the sepals and almost as wide. In addition to A. venosa, well established on Cape Breton Island and sporadically elsewhere in Atlantic Canada, A. mollis and A. speciosa Buser are in cultivation, the former widely so and recently reported as naturalized. These species are distinguished from A. venosa by their flat or only slightly undulate, densely pubescent leaves. Alchemilla speciosa differs from A. mollis by its pubescent pedicels and its epicalyx bractlets narrower than the sepals.
Aphanes, the members of which are readily distinguished by their annual habit, has sometimes been included in Alchemilla in North American floras and manuals. Over the past half century, the two genera have generally been maintained as distinct, a pattern set even earlier in North America by P. A. Rydberg (1908). As a result of molecular studies, B. Gehrke et al. (2008) have suggested that Alchemilla should be circumscribed more widely to include Aphanes and also Lachemilla (Focke) Rydberg, which occurs in Mexico, Central America, and South America. Both of these genera are shown to be monophyletic groups, as is Alchemilla apart from the African species currently placed in it that form a clade separate from the genus as represented in Eurasia and North America. The African species deserve more thorough investigation before incorporating within Alchemilla such a morphologically distinct genus as Aphanes.
M. J. M. Christenhusz and S. M. Väre (2012) have proposed 35 new names and combinations in Potentilla for those species of Alchemilla and Aphanes native or cultivated in the Nordic countries. This is said to be based on the molecular findings of C. Dobeš and J. Paule (2010), which support previous results by T. Eriksson et al. (1998). These studies note that when Potentilla is traditionally circumscribed (as for example by P. W. Ball et al. 1968), it is paraphyletic and Alchemilla is nested within it, leading Chistenhusz and Väre to adopt an inclusive Potentilla that Dobeš and Paule consider an impractical and unconventional generic circumscription, a view that the authors share. As laid out by B. Ertter (2007), in this flora the authors follow the alternate option taken by Dobeš and Paule and other authors (A. Kurtto and Eriksson 2003; J. Soják 2008) of regcognizing within the monophyletic subtribe Fragariinae not only Alchemilla, Chamaerhodos, and Fragaria, but also other genera previously segregated from Potentilla on morphological grounds, such as Comarum, Dasiphora, Drymocallis, Farinopsis Chrtek & Soják, Sibbaldia, and Sibbaldiopsis.
Selected References
Lower Taxa
Key
| 1 | Leaves palmately compound. | Alchemilla alpina |
| 1 | Leaves palmately lobed | > 2 |
| 2 | Epicalyx bractlet lengths equal to or longer than sepals | > 3 |
| 2 | Epicalyx bractlet lengths shorter than sepals | > 4 |
| 3 | Leaf margins undulate, basal sinuses appearing closed, lobes overlapping, adaxial surfaces glabrous or hairy only on folds; hypanthia glabrous; leaves: middle lobes as long as or longer than their half-widths. | Alchemilla venosa |
| 3 | Leaf margins flat or slightly undulate, basal sinuses narrow or almost closed, lobes sometimes overlapping, adaxial surfaces hairy throughout; hypanthia usually densely, occasionally sparsely, hairy (in proximal 1/2); leaves: middle lobes shorter to as long as their half-widths. | Alchemilla mollis |
| 4 | Stems glabrous or appressed-hairy | > 5 |
| 4 | Stems spreading-, ascending-, or reflexed-pubescent or sericeous, sometimes glabrous (A. micans) | > 7 |
| 5 | Adaxial leaf surfaces sparsely to densely appressed-hairy throughout or only on folds, (nerves glabrous); primary inflorescence branches densely appressed- to ascending-hairy. | Alchemilla glomerulans |
| 5 | Adaxial leaf surfaces glabrous or slightly hairy on margins only; primary inflorescence branches glabrous | > 6 |
| 6 | Incisions absent from leaf sinuses; leaf teeth subacute or obtuse, strongly asymmetric (slightly concave near apex). | Alchemilla glabra |
| 6 | Incisions present in leaf sinuses; leaf teeth acute, symmetric to slightly asymmetric. | Alchemilla wichurae |
| 7 | Primary inflorescence branches glabrous. | Alchemilla filicaulis |
| 7 | Primary inflorescence branches sparsely to densely hairy | > 8 |
| 8 | Pedicels (all or most) hairy | > 9 |
| 8 | Pedicels usually glabrous or some of the proximal sparsely hairy | > 10 |
| 9 | Stipules strongly wine red-tinged proximally; leaf blades usually reniform, sometimes orbiculate, basal sinuses usually wide, sometimes narrow. | Alchemilla filicaulis |
| 9 | Stipules translucent, colorless, sometimes flushed pale wine red proximally; leaf blades orbiculate, basal sinuses closed. | Alchemilla glaucescens |
| 10 | Leaves: basal sinuses closed, basal lobes overlapping. | Alchemilla subcrenata |
| 10 | Leaves: basal sinuses wide or narrow, basal lobes not overlapping | > 11 |
| 11 | Adaxial leaf surfaces usually glabrous, rarely sparsely hairy on teeth, margins, and folds or on some folds only. | Alchemilla xanthochlora |
| 11 | Adaxial leaf surfaces densely appressed-hairy or pubescent throughout | > 12 |
| 12 | Hypanthia attenuate at base, usually glabrous; stipules translucent, strongly wine red-tinged proximally; stems usually sparsely to densely spreading- to slightly ascending-hairy, sometimes glabrous in distal 1/2. | Alchemilla micans |
| 12 | Hypanthia rounded at base, usually sparsely to densely spreading-hairy; stipules translucent to pale green proximally, (apices green); stems usually densely spreading-hairy in distal 1/2. | Alchemilla monticola |