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Comarum palustre

comaret des marais, marsh potentilla, marsh-cinquefoil, purple cinquefoil, purple marsh-locks

rose family

Habit Herbs (annual or perennial), shrubs, or trees.
Stems

1–10(–18) dm.

simple or branched.

Leaves

petiole 5–10(–15) cm;

leaflets usually abaxially glaucous, adaxially green, sometimes silvery-silky, often aging reddish, 1.5–10(–12) × (0.8–)1–5 cm.

persistent or deciduous, basal and/or cauline, usually alternate, rarely opposite, simple or compound (palmate or imparipinnate);

stipules usually present, sometimes absent;

petiole present or absent;

blade thin to coriaceous, margins ± lobed or unlobed, usually toothed.

Inflorescences

terminal, sometimes axillary, panicles with terminal flower (that is, determinate) or reductions of this: 1-flowered, glomerules, fascicles, spikes, racemes, corymbs, umbels, or cymes.

Pedicels

5–25(–30) mm.

Flowers

epicalyx bractlets narrowly lanceolate to lanceolate, 2–7(–9) mm, usually equal to or shorter than sepals;

sepals greenish purple adaxially, (5–)7–12(–14) mm, apex acuminate;

petals 2.5–6(–8) mm;

anthers dark purple, 0.8–1.2 mm, connectives thickened, greenish purple to purple;

styles 0.9–2 mm.

usually bisexual, rarely unisexual, perianth and androecium perigynous or epigynous;

epicalyx bractlet sometimes present;

hypanthium flat to hemispheric, or cylindric to funnelform or urceolate;

sepals (0–)4 or 5(–10), distinct, free;

petals (0–)4 or 5(–12, rarely more in double ornamentals), distinct, free;

nectar disc sometimes absent;

stamens 0–130(–220), distinct, free, anthers usually longitudinally dehiscent;

torus well developed, inconspicuous, or absent;

pistils 1–250(–450), distinct or ± connate, free or ± adnate to hypanthium, ovary superior or inferior (then 2–5-carpellate and -locular and ± connate with axile placentation), styles terminal, subterminal, lateral, or ± basal, sometimes basally connate, stigmas usually capitate;

ovules 1 or 2(–5+), basal, marginal, or apical, collateral, superposed, biseriate, or clustered, integuments 2, crassinucellate, with or without obturator.

Fruits

achenes aggregated or not, follicles aggregated or not, drupes aggregated or not, aggregated nutlets, pomes, aggregated drupelets, or capsules;

sometimes involving accessory organs, for example, hypanthium, torus.

Seeds

1 or 2(–12+), not arillate.

2n

= 28, (35)36, 42, (60, 62, 63)64.

Comarum palustre

Rosaceae

Phenology Flowering summer.
Habitat Shallow, often sphagnum-dominated ponds, bogs, wet meadows, marshes, stream banks, and lake margins, mainly in boreal and low arctic habitats in the north, subalpine to alpine sites in the south
Elevation 0–3100 m (0–10200 ft)
Distribution
from FNA
AK; CA; CO; CT; IA; ID; IL; IN; MA; ME; MI; MN; MT; ND; NH; NJ; NY; OH; OR; PA; UT; VT; WA; WI; WY; AB; BC; MB; NB; NL; NS; NT; NU; ON; PE; QC; SK; YT; SPM; Greenland; Eurasia
[WildflowerSearch map]
[BONAP county map]
North America; Mexico; Central America; South America; West Indies; Bermuda; Eurasia; Africa; Atlantic Islands; Indian Ocean Islands; Pacific Islands (Hawaii, New Zealand); Australia
[BONAP county map]
Discussion

Comarum palustre is circumboreal; it occurs in a broad band across the northernmost United States and Canada as far north as the low arctic. Disjunct populations occur well to the north, as on Baffin and Victoria islands, and in the south, as in Gunnison County, Colorado, and the Uinta Mountains of Utah. Appropriate habitats are uncommon and geographically restricted in the southern part of the range, for example, in Idaho, Montana, and North Dakota.

Comarum palustre is sometimes divided into subspecies, varieties, and/or forms on the basis of vestiture, leaflet dimensions, and flower number (for example, M. L. Fernald and B. H. Long 1914; N. N. Tzvelev 2007). Because there is no existing consensus, and a global evaluation of the situation is beyond the scope of this work, no infraspecific taxa are recognized here. Within North America, northern populations tend to have broader leaflets and inflorescences composed of one to relatively few flowers with relatively short sepals and petals, but these differences are not consistent; similar plants occur in subalpine settings in the mountains well to the south. Chromosome numbers also vary, apparently not correlated with morphology or geography; some populations occasionally include individuals of different disploid and polyploid levels.

Horizontal stems were used occasionally by Native Americans as a medicinal aid in the treatment of dysentery and stomach cramps (D. E. Moerman 1998). Aquatic birds in particular eat the leaves and achenes.

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

Genera 88, species ca. 3000 (68 genera, 680 species, including 22 hybrids, in the flora).

Three subfamilies and 16 tribes are recognized for the family with representatives of all tribes found in the flora area.

Rosaceae grow most commonly in north-temperate regions and are more or less absent from hot deserts and high-rainfall, low-altitude tropics. The family is large and diverse, characterized by radially symmetric flowers with a fundamentally saucer-shaped hypanthium and peripheral calyx, corolla, androecium, and, usually, superior gynoecium. Considerable variation occurs in important details of flower and fruit, this enhanced by a very adaptable hypanthium, which is discussed in more detail below.

Rosaceous inflorescences vary in the number of flowers from one to about 500. A great variation occurs also in inflorescence form, though a pattern is perceived when they are viewed as reduction series from a terminal, determinate panicle that is fundamentally bracteate, thus generating a range of panicles, racemes, corymbs, cymes, solitary flowers, or other forms. In this treatment, appendages on the inflorescence are distinguished by the terms bract and bracteole. Bract is used for the larger, laminate, usually chlorophyllous leaf homologues that subtend axes and may be indistinguishable from foliage leaves. Bracteoles are scalelike, often membranous, often caducous, and of uncertain homology, in part due to not being restricted to axis-subtending positions.

The floral architecture in Rosaceae is radially symmetric around a disc-shaped to urceolate hypanthium. Flowers normally have a four- or five-merous corolla and calyx; great variation is found in the numbers of stamens and carpels. Pollination is usually entomophilous, the flowers having normally green sepals and showy, often more or less clawed, usually white, yellow, or pink, less commonly red or green, petals. The flowers in some genera are relatively small and anemophilous and may lack one or two of the principal whorls. An unusual feature of some rosaceous flowers is the torus, a pad of receptacular, usually spongy, tissue, sometimes relatively large, in the center of the hypanthium that, when present, bears the gynoecium. Another unusual feature of some genera is an epicalyx that comprises a ring of sepaloid bractlets, usually of the same number as sepals, which is located on the hypanthium proximal to the calyx.

Fruit types are particularly significant both in rosaceous identification, taxonomy, and diversity, as well as for successful dispersal. Fruit in Rosaceae may be either dry, then dehiscent or not, or succulent and indehiscent; it sometimes involves accessory organs such as the torus or a persistent hypanthium.

Dry indehiscent fruits are achenaceous, in certain cases involving modifications to the style. In anemochorous (adapted for dispersal by wind) situations, this may involve a plumose style, for example, Cercocarpus; in epizoochorous (distributed on the outside of animals) cases, the styles may bear stiff hairs or barbs, for example, Geum. Alternatively, achenaceous fruits may lack styles (sometimes due to abscission) or, more commonly, have a relatively short one, for example, Potentilla. In situations where there appears to be no significant post-flowering function for the style, dispersal may be myrmecochorous (by ants), anemochorous, or in the rare torus-borne cases (for example, Fragaria), the fruit may be endozoochorous (eaten by animals and passed through the gut). Sometimes, fruits with such styles are aggregated in acheneta that rely on barbed, persistent hypanthia for epizoochorous dispersion, for example, Acaena and Agrimonia. Dry dehiscent fruits are follicular with seeds normally distributed by air after splitting of the ripe follicle, for example, Gillenia.

Succulent, endozoochorous fruits exhibit a similar range of variation. The most common types are: multiple drupelets on the surface of a more or less conic torus (for example, Rubus); individual and sometimes large drupes on a flat receptacular apex (Prunus); pomes, which are berrylike fruits in which a fleshy hypanthium more or less completely surrounds and is generally more or less fully adnate to the carpels (for example, Amelanchier, Crataegus). The result is that a terminal orifice may remain open, often bearing floral remnants around its rim. Pomes with hard pyrenes are sometimes distinguished as polypyrenous drupes, a term not used in this treatment due to antonymic confusion caused by combining the roots for pyrene (hardened mesocarp) and drupe (fleshy mesocarp), although the relevant distinction is well recognized (J. Rohrer et al. 1991).

Some important temperate fruits are members of the Rosaceae: apples (Malus), pears (Pyrus), almonds, apricots, cherries, peaches, and plums (Prunus), blackberries and raspberries (Rubus), strawberries (Fragaria), loquat (Eriobotrya); minor fruits include those of Amelanchier, Crataegus, Cydonia, Mespilus, and others. Some genera are popular in ornamental horticulture in North America, for example, most of the above as well as, especially, Chaenomeles, Cotoneaster, Pyracantha, Rhaphiolepis Lindley, Sorbus, Photinia, Physocarpus, Rosa, and Spiraea among trees and shrubs; and Filipendula, Geum, Potentilla, and Spiraea among plants suitable for flower borders.

Apomixis is a feature of some rosaceous genera and may make taxonomic decisions difficult or equivocal in genera such as Alchemilla, Amelanchier, Crataegus, Rubus, and others. Apomixis always seems to be associated with polyploidy and often with hybridization; in some apomictic genera, sexual species and apomicts are facultatively sexual.

Rosaceae lack alkaloids and blue anthocyanic pigments. Some have foliage or seed that becomes toxic due to hydrolysis of benzaldehyde cyanohydrins.

The great diversity of Rosaceae reflects the age of the family (since at least the mid Eocene) and its evolutionary success. The family has needed no fundamental change in circumscription since Jussieu first recognized it, the sometimes included outgroups, for example, Chrysobalanaceae, Neuradaceae, already being confidently excluded in more recent pre-molecular classifications, while segregate families (for example, Malaceae) have received little currency. This reflects a lack both of close neighbors and of large internal discontinuities. Species limits are often debatable, especially where apomixis is present; generic limits are much more fixed, apart from the familiar historic trend to recognizing smaller, more discrete units. What has principally been fluid in rosaceous taxonomy until modern molecular research are the phylogenetic relationships, both to neighboring families and within the family. Rosales once contained families such as Crassulaceae and Saxifragaceae, which have many morphological similarities to Rosaceae, but Angiosperm Phylogeny Group (2003) indicated instead that in Rosales, Rosaceae is sister to a group of families including Moraceae, Rhamnaceae, Ulmaceae, and Urticaceae.

The internal relationships of Rosaceae have received much study, including morphological, anatomical, cytological, phytochemical, breeding system, fungal pathogenicity (especially rusts), and molecular. Among these studies, that of D. Potter et al. (2007) resulted in the first comprehensive molecular phylogeny of Rosaceae, and the treatment in this volume reflects it. Potter et al. recognized three subfamilies with 15 tribes worldwide. Their classification used the ranks supertribe and subtribe, not used here. The subfam. Dryadoideae, with one tribe and four genera, diverged early and is unique in Rosaceae for its actinorhizal symbiosis; it has achenaceous fruits. Subfamily Rosoideae has six tribes and 29 genera and is only slightly altered from its traditional circumscription by shedding Dryadoideae and three small genera. Most Rosoideae have achenaceous fruit; some have multiple drupelets or achenes borne on a torus. The remainders of Rosaceae are found in the large and heterogeneous subfam. Amygdaloideae of nine tribes and 54 genera. This subfamily has the largest diversity of fruit types. Amygdaloideae now contains the traditional spiraeoid genera with follicular fruit, Maleae with pome fruit, and Amygdaleae with drupaceous fruit.

Key to Subfamilies and Tribes of Rosaceae

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

Key

Key to Subfamilies and Tribes of Rosaceae (Luc Brouillet)

1. Leaves simple, sometimes pinnately, rarely palmately compound, stipules deciduous or absent, rarely persistent, if pinnately compound and stipules persistent, then fruits follicles (rarely achenes); flowers: perianth and androecium perigynous or epigynous, tori absent or minute; ovules 2+ (rarely 1, sometimes by abortion); fruits usually aggregated follicles or drupes or nutlets, pomes, ± woody capsules, follicles, or drupes, rarely achenes [c. Rosaceae subfam. Amygdaloideae, p. xxx]
→ 2
1. Leaves pinnately, sometimes palmately, compound, sometimes simple, stipules persistent and adnate to petioles, sometimes deciduous and free, rarely absent; flowers: perianth and androecium perigynous, tori enlarged, sometimes small or absent; ovules 1(or 2, then fruits achenes or drupes); fruits achenes, or aggregated drupelets or achenes (within fleshy, urn-shaped hypanthia [hips] in Roseae)
→ 10
2. Leaves opposite; carpels 2(or 3), distinct, free; fruits aggregated follicles
c1 Lyonothamneae
2. Leaves alternate, rarely opposite; carpels 1–8, distinct or connate, free or ± adnate to hypanthium; fruits achenes, drupes, aggregated achenes, drupes, nutlets, or follicles, or capsules or pomes
→ 3
3. Shrubs or herbs; stipules present; leaves pinnately compound, rarely simple
→ 4
3. Trees or shrubs, rarely herbs (then stipules absent); stipules present or absent; leaves simple, rarely pinnately compound (then herbs or trees)
→ 5
4. Shrubs; carpels 1–5, basally connate or distinct
c6 Sorbarieae
4. Herbs; carpels 5, basally connate
c8 Gillenieae
5. Carpels 1; fruits drupes
c3 Amygdaleae
5. Carpels 1–5; fruits achenes, aggregated drupes, aggregated nutlets, aggregated follicles, capsules, or pomes
→ 6
6. Flowers: perianth and androecium epigynous (perigynous in Vauquelinia); carpels ± connate or distinct, adnate to hypanthium (free in Vauquelinia); fruits pomes or woody capsules (surrounded by persistent hypanthia and splitting into 5 follicles Vauquelinia)
Maleae
6. Flowers: perianth and androecium perigynous; carpels distinct, sometimes ± connate, free or rarely adnate to hypanthium base; fruits achenes, aggregated follicles, aggregated nutlets, aggregated drupes, or capsules splitting into 5 follicles (not surrounded by hypanthia)
→ 7
7. Shrubs; leaf venation palmate (stipules present); carpels basally connate
c2 Neillieae
7. Shrubs, sometimes subshrubs, trees, or herbs; leaf venation pinnate (sometimes palmate in Spiraeeae, then stipules absent); carpels distinct (sometimes connate in Exochordeae)
→ 8
8. Fruits capsules or aggregated drupes
c4 Exochordeae
8. Fruits aggregated follicles or nutlets, or achenes
→ 9
9. Shrubs; stipules persistent or deciduous; leaves alternate or opposite; carpels 2–8 or 1 (in Coleogyne); ovules 1 or 2, marginal; fruits achenes or aggregated nutlets
c5 Kerrieae
9. Shrubs, subshrubs, or herbs; stipules absent; leaves alternate; carpels 3–5(or 6); ovules 2–5, apical; fruits aggregated follicles or achenes (in Holodiscus)
c7 Spiraeeae
10. Shrubs or trees, unarmed; leaves simple, sometimes pinnately compound; styles persistent, elongate in fruit [b. Rosaceae subfam. Dryadoideae]
b1 Dryadeae
10. Herbs, shrubs, or subshrubs, these sometimes armed; leaves pinnately or palmately compound, rarely simple; styles deciduous, if persistent in fruit, then either not elongate, or rarely elongate (in Colurieae) [a. Rosaceae subfam. Rosoideae]
→ 11
11. Shrubs or subshrubs
→ 12
11. Herbs
→ 15
12. Hypanthia ovoid, oblong, globose to cupulate or urceolate, rarely obovoid or hemispheric, tori absent (conic if present); fruits aggregated achenes within hypanthium (hips)
a4 Roseae
12. Hypanthia saucer-, cup-, or funnel-shaped, hemsipheric to flat, or obconic to obcampanulate, tori enlarged; fruits achenes, or aggregated achenes or drupelets
→ 13
13. Plants armed, sometimes unarmed; stipules free or adnate to petiole base; fruits aggregated drupelets…
Rubeae
13. Plants unarmed; stipules absent or ± free or ± adnate to petiole; fruits aggregated achenes, sometimes achenes
→ 14
14. Epicalyx bractlets absent, sometimes present; styles terminal, sometimes subterminal, elongate in fruit
Colurieae
14. Epicalyx bractlets present, sometimes absent; styles basal or lateral to subterminal, not elongate in fruit
Potentilleae
15. Leaves pinnately compound, sometimes simple with palmate venation; stipules free; fruits aggregated drupelets; ovules 2, collateral
Rubeae
15. Leaves pinnately or palmately compound, rarely simple with pinnate or palmate venation; stipules adnate to petiole, sometimes free; fruits achenes or aggregated achenes; ovules 1, if 2 then superposed
→ 16
16. Tori inconspicuous or absent, sometimes enlarged in fruit; carpels 1–12
→ 17
16. Tori usually enlarged; carpels 1–260(–450)
→ 18
17. Ovules 2; fruits aggregated achenes, not enveloped by hypanthia; tori enlarged in fruit
a1 Ulmarieae
17. Ovules 1; fruits achenes, enclosed within enlarged, often hardened, hypanthia; tori not enlarged in fruit
a6 Agrimonieae
18. Epicalyx bractlets absent, sometimes present; styles terminal, sometimes subterminal, elongate in fruit
Colurieae
18. Epicalyx bractlets present, sometimes absent; styles basal or lateral to subterminal, not elongate in fruit
Potentilleae
Source FNA vol. 9, p. 301. FNA vol. 9, p. 18. Authors: James B. Phipps, Luc Brouillet.
Parent taxa Rosaceae > subfam. Rosoideae > tribe Potentilleae > Comarum
Subordinate taxa
a1 Ulmarieae, a4 Roseae, a6 Agrimonieae, b1 Dryadeae, c1 Lyonothamneae, c2 Neillieae, c3 Amygdaleae, c4 Exochordeae, c5 Kerrieae, c6 Sorbarieae, c7 Spiraeeae, c8 Gillenieae
Synonyms C. palustre subsp. angustifolium, C. palustre subsp. arcticum, Fragaria palustris, Potentilla palustris, P. palustris var. parvifolia, P. palustris var. villosa
Name authority Linnaeus: Sp. Pl. 1: 502. (1753) Jussieu
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