Silene

Herbs, annual, biennial, or perennial, often decumbent at base or sometimes cespitose. Taproots slender or often stout, deep, branched caudex often present, some species stoloniferous or rhizomatous. Stems simple or branched, terete or sometimes angular. Leaves opposite or occasionally whorled, connate proximally, petiolate (basal leaves) or sessile (most cauline leaves); blade 1–5-veined, linear to obovate or spatulate, herbaceous, apex acute to obtuse. Inflorescences terminal or sometimes axillary, simple or branched, sometimes condensed cymes, frequently flowers few or solitary, frequently glandular-pubescent and viscid; bracts paired, herbaceous or scarious, or absent; involucel bracteoles absent. Pedicels erect, rarely flowers sessile or subsessile. Flowers bisexual, sometimes unisexual (rarely so on separate plants); sepals connate proximally into tube, (4–)10–28(–40) mm; tube green, whitish, and/or purplish, 10–30-veined, cylindric to campanulate, urceolate, or clavate, terete, frequently inflated, membranous or more rarely herbaceous, commissures between sepals 1-veined, herbaceous; lobes green or purplish, 1–5-veined, broadly triangular to lance-oblong or linear, usually shorter than tube, margins whitish, scarious, apex acute to obtuse; petals 5, white, pink, scarlet, dusky purple, or off-white tinged with purple, clawed, claw usually conspicuous, sometimes small, rarely absent, auricles 2, coronal appendages 2, variously shaped or dissected; limb usually exserted and conspicuous, oblanceolate to obovate, apex 2-lobed, sometimes dissected into 1–4 linear lobes or irregular teeth, or fimbriate, rarely entire; nectaries at filament bases; stamens 10, rarely fewer or absent, frequently dimorphic with longer opposite petals, arising with petals from carpophore; filaments distinct nearly to base; staminodes absent (rarely to 10 in pistillate flowers, arising with petals from carpophore, filiform); ovary 1- or 3–5-locular; styles 3 or 5, occasionally 4 (absent in staminate flowers), filiform, 1.5–20 mm, glabrous proximally; stigmas 3 or 5, occasionally 4, linear along adaxial surface of styles, papillate (30×). Capsules ovoid to globose, opening along sutures into 3–5 valves, frequently splitting into 6–10 equal teeth; carpophore usually present. Seeds ca. (5–)15–100(–500+), reddish to gray or black, reniform to globose, usually tuberculate or papillate, papillae around margins sometimes larger and inflated, marginal wing sometimes present, appendage absent; embryo peripheral, curved. x = (10) 12.

Distribution

Mainly Northern Hemisphere.

Discussion

Species ca. 700 (70 in the flora).

Silene includes several important weeds and some very beautiful horticultural plants. In addition to the species described in this account, several others have occurred in the flora area as chance introductions or garden escapes, but they have not become established and most have not been seen recently. They include S. coeli-rosa (Linnaeus) Godron, S. cretica Linnaeus, S. (Lychnis) fulgens (Fischer) E. H. L. Krause, S. italica Persoon, and S. nutans Linnaeus.

In this account, Lychnis, Melandrium, and Viscaria have been included in Silene, their previous recognition as distinct genera having resulted in a great deal of confusion in both nomenclature and taxonomy. I have not presented an infrageneric classification of Silene because existing systems either do not include those other genera (e.g., P. K. Chowdhuri 1957) or do not deal with most of our native North American taxa [e.g., W. Greuter (1995) and the molecular studies by Oxelman and coworkers (e.g., B. Oxelman et al. 1997, 2000]. The recent molecular study by J. G. Burleigh and T. P. Holtsford (2003) provides little support for existing morphologically based sectional classifications within Silene insofar as they relate to endemic North American taxa. However, it does indicate the distinctness of our arctic alpine species (S. involucrata—as S. furcata, and S. acaulis) that are circumpolar in their distribution.

Selected References

Bocquet, G. 1969. Revisio Physolychnidum (Silene Sect. Physolychnis).... Lehre.

Burleigh, J. G. and T. P. Holtsford. 2003. Molecular systematics of the eastern North American Silene (Caryophyllaceae): Evidence from nuclear ITS and chloroplast trnL intron sequences. Rhodora 105: 76–90.

Hitchcock, C. L. and B. Maguire. 1947. A Revision of the North American Species of Silene. Seattle. [Univ. Wash. Publ. Biol. 13.]

Kruckeberg, A. R. 1962. Intergeneric hybrids in the Lychnideae (Caryophyllaceae). Brittonia 14: 311–321.

McNeill, J. 1978. Silene alba and S. dioica in North America and the generic delimitation of Lychnis, Melandrium and Silene (Caryophyllaceae). Canad. J. Bot. 56: 297–308.

Oxelman, B. and M. Lidén. 1995. Generic boundaries in the tribe Sileneae (Caryophyllaceae) as inferred from nuclear rDNA sequences. Taxon 44: 525–542.

Oxelman, B., M. Lidén, and D. Berglund. 1997. Chloroplast rps16 intron phylogeny of the tribe Sileneae (Caryophyllaceae). Pl. Syst. Evol. 206: 411–420.

Oxelman, B., M. Lidén, R. K. Rabeler, and M. Popp. 2000. A revised generic classification of the tribe Sileneae (Caryophyllaceae). Nordic J. Bot. 20: 743–748.

Williams, F. N. 1896b. A revision of the genus Silene Linn. J. Linn. Soc., Bot. 32: 1–196.

Lower Taxa

Silene acaulis, Silene antirrhina, Silene aperta, Silene armeria, Silene bernardina, Silene bridgesii, Silene caroliniana, Silene chalcedonica, Silene conica, Silene coniflora, Silene conoidea, Silene coronaria, Silene csereii, Silene dichotoma, Silene dioica, Silene douglasii, Silene drummondii, Silene flos-cuculi, Silene gallica, Silene grayi, Silene greenei, Silene hitchguirei, Silene hookeri, Silene invisa, Silene involucrata, Silene kingii, Silene laciniata, Silene latifolia, Silene lemmonii, Silene marmorensis, Silene menziesii, Silene nachlingerae, Silene nivea, Silene noctiflora, Silene nuda, Silene occidentalis, Silene oregana, Silene ostenfeldii, Silene ovata, Silene parishii, Silene parryi, Silene pendula, Silene petersonii, Silene plankii, Silene polypetala, Silene pseudatocion, Silene rectiramea, Silene regia, Silene repens, Silene rotundifolia, Silene sargentii, Silene scaposa, Silene scouleri, Silene seelyi, Silene serpentinicola, Silene sibirica, Silene sorensenis, Silene spaldingii, Silene stellata, Silene subciliata, Silene suecica, Silene suksdorfii, Silene thurberi, Silene uralensis, Silene verecunda, Silene virginica, Silene viscaria, Silene vulgaris, Silene williamsii, Silene wrightii

Key

1	Stigmas 5 (rarely 4 in individual flowers); capsule with 5-10 teeth (rarely 4 or 8)	> 2
1	Stigmas 3 (rarely 4 or 5 in individual flowers, usually 4 in S. scaposa); capsules with 3 or 6 teeth (rarely 4, 5, 8, or 10)	> 16
2	Limbs of petals shorter than calyx, not differentiated from claw, ascending to spreading but rarely in horizontal plane; petals off-white, dingy pink, dusky purple, or purple tinged	> 3
2	Limbs of petals clearly differentiated from claw, spreading horizontally, equaling or longer than calyx (shorter in S. chalcedonica, S. flos-cuculi, and S. involucrata); petals showy, scarlet, bright purple or pink, or white	> 9
3	Seeds winged	> 4
3	Seeds not winged	> 6
4	Calyces inflated in flower or fruit, thin and papery; petals dusky purple-red; seeds 1.5-2.5 mm, with broad wing; flowers nodding (erect in subsp. porsildii but seeds 2-2.5 mm)	Silene uralensis
4	Calyces neither inflated nor very thin; petals white or pink; seeds 0.5-1.5 mm, wing often narrow; flowers erect	> 5
5	Seeds (0.5-)0.8-1(-1.3) mm, wing to ca. 4 diam. of seed; calyces elliptic, 7-10(-12) mm; stem 2-10(-12) cm; flowers 1(-3)	Silene hitchguirei
5	Seeds 1-1.5 mm, wing to 2 diam. of seed; calyces often campanulate or ovate, 8-20 mm; stems 10-45 cm; often with 1-3 flowers per peduncle	Silene involucrata
6	Fruiting calyces 2-3 times longer than broad, broadly tubular to narrowly ellipsoid	Silene drummondii
6	Fruiting calyces to 2 times as long as broad, campanulate, ovate, or elliptic	> 7
7	Fruiting calyces 10-12 mm, not or only slightly contracted at mouth	Silene ostenfeldii
7	Fruiting calyces 13-17 mm, contracted at mouth to 2-2/ 3 diam	> 8
8	Inflorescences 1(-3)-flowered; pedicels 2-3 times calyx, length of pubescence less than 1/ 2 pedicel diam.; veins of calyx prominent	Silene kingii
8	Inflorescences (1-)3-flowered; flowers sessile or pedicellate; pedicels usually much shorter than calyx, longest hairs ca. equaling pedicel or peduncle diam.; veins of calyx partially obscured by dense pubescence	Silene sorensenis
9	Inflorescences dense (interrupted in S. viscaria) cymes of (3-)5-50 flowers; flowers sessile or pedicels shorter than calyx	> 10
9	Inflorescences open cymes, 1-3 to many-flowered; most pedicels longer than calyx	> 12
10	Petals scarlet (rarely white or pink); coarse hispid herbs 50-100 cm; leaf blades broadly ovate or elliptic to lanceolate	Silene chalcedonica
10	Petals bright pink or purple (rarely white); glabrous or sparsely pubescent herbs 5-90 cm; leaf blades narrowly lanceolate	> 11
11	Flowers 14-22 mm diam.; limb of corolla obovate, ± entire; carpophore 3-5 mm; nodes of stem viscid	Silene viscaria
11	Flowers 5-10 mm diam.; limb of corolla 2-lobed to middle; carpophore ca. 1 mm; nodes of stem not viscid	Silene suecica
12	Open flowers 8-16 mm diam	Silene involucrata
12	Open flowers 20-35 mm diam	> 13
13	Petal limbs deeply divided with 4 linear segments	Silene flos-cuculi
13	Petal limbs unlobed, emarginate or shallowly 2-lobed	> 14
14	Leaf blades with dense, silky, grayish white tomentum; flowers bisexual	Silene coronaria
14	Pubescence hirsute or sparingly to densely pubescent; flowers unisexual	> 15
15	Petals white; capsule teeth slightly reflexed or spreading	Silene latifolia
15	Petals bright pink; capsule teeth revolute	Silene dioica
16	Petals scarlet	> 17
16	Petals white, cream, pink, or dark red, off-white tinged with purple	> 23
17	Limbs of petals unlobed or emarginate, shallowly dentate, or crenate, rarely shallowly 2-lobed	> 18
17	Limbs of petals deeply lobed or laciniate	> 20
18	Stems and leaf blades glabrous	Silene subciliata
18	Stems and leaf blades pubescent	> 19
19	Stems erect, 50-160 cm; inflorescences many-flowered; leaf blades ovate to lanceolate, apex acute, not acuminate	Silene regia
19	Stems ascending, 10-20 cm; inflorescences with flowers usually solitary, terminal; leaf blades linear-lanceolate, apex sharply acuminate	Silene plankii
20	Petals deeply 4-6-lobed, small lateral teeth may be present	Silene laciniata
20	Petals deeply 2-lobed, lobes often with lateral teeth	> 21
21	Leaf blades subrotund to broadly lanceolate; petal lobes ciliate	Silene rotundifolia
21	Leaf blades lanceolate, oblanceolate, or obovate to narrowly elliptic; petal lobes glabrous or nearly so (claw may be ciliate)	> 22
22	Stems 4-10(-15) cm; inflorescences 1-3(-4)-flowered; leaves longest in mid stem, basal tufts absent; w United States	Silene serpentinicola
22	Stems 20-80 cm; inflorescences (3-)7-11(-20)-flowered; leaves longest at base of stem, in basal tufts; e United States	Silene virginica
23	Calyces with 20-30 parallel veins	> 24
23	Calyces with 10 or fewer veins or venation obscure	> 27
24	Calyces glabrous, veins obscure	Silene csereii
24	Calyces pubescent and usually glandular, veins prominent	> 25
25	Mature calyces 20-30 mm; seeds 1.3-1.8 mm broad	Silene conoidea
25	Calyces 8-15 mm; seeds 0.6-1 mm broad	> 26
26	Petals ± equaling calyx, limb 1-3 mm, inconspicuous; corollas cream to dull orange abaxially, often purple tinged adaxially; calyx lobes 2-3 mm	Silene coniflora
26	Petals much exceeding calyx, conspicuous, limb 3-6 mm; corollas pink (rarely white or dark red); calyx lobes ca. 5 mm	Silene conica
27	Inflorescences congested, capitate, or flowers clustered; flowers sessile or pedicels shorter than calyx	> 28
27	Inflorescences open; flowers pedicellate, if pedicels short, then flowers in several separate whorls, or solitary	> 30
28	Plants glabrous	Silene armeria
28	Plants pubescent	> 29
29	Calyx veins purple tinged or entirely purple; corollas rose pink, limb 2-lobed	Silene repens
29	Calyx veins green; corollas greenish yellow to white, limb deeply lacin- iate into 6 or more linear lobes	Silene parishii
30	Leaves whorled at each node, 4 to many	> 31
30	Leaves opposite, 2 at each node (smaller axillary shoots sometimes present), or all or mostly proximal	> 32
31	Leaf blades lanceolate to ovate-lanceolate, 4 per node	Silene stellata
31	Leaf blades linear to very narrowly elliptic-lanceolate, more than 4 pernode	Silene sibirica
32	Calyces herbaceous, conspicuous pale commissures between veins absent, veins obscure	> 33
32	Calyces with pale commissures between prominent parallel veins	> 43
33	Limbs of petals with several narrow, linear lobes	Silene greenei
33	Limbs of petals unlobed, sometimes emarginate, or 2-lobed	> 34
34	Calyces glabrous	> 35
34	Calyces pubescent	> 38
35	Petals bright pink, rarely white	Silene acaulis
35	Petals white	> 36
36	Fruiting calyces ovoid, contracted at mouth to ca. 2 its diam.; stamens ca. 2 times calyx; filaments purple	Silene csereii
36	Fruiting calyces clavate or campanulate, not contracted at mouth; stamens ca. 1-11/ 3 times calyx; filaments usually white	> 37
37	Fruiting calyces broadly clavate, strongly contracted at base around carpophore; carpophores5-6 mm; inflorescences leafy	Silene nivea
37	Fruiting calyces campanulate, not contracted at base; carpophores 2-3 mm; inflorescences withleaves reduced, resembling bracts	Silene vulgaris
38	Seeds winged	Silene spaldingii
38	Seeds not winged	> 39
39	Mature calyces contracted towards base around carpophore; carpophores 4-7 mm	> 40
39	Mature calyces not contracted at base; carpophores less than 2 mm	> 41
40	Corollas white; calyces green	Silene nivea
40	Corollas pink (rarely white); calyces purple, purpleveined, or purple tinged	Silene repens
41	Leaf blades oblanceolate to elliptic-lanceolate, rarely obovate or elliptic, broadest distally or atmiddle	Silene menziesii
41	Leaf blades narrowly lanceolate to elliptic- or ovate-lanceolate, broadest proximally	> 42
42	Petals dark red (rarely white); calyces 6-9 ×3-4 mm	Silene seelyi
42	Petals white; calyces 9-11 × 4-6 mm	Silene williamsii
43	Plants annual or biennial, often with slender taproot	> 44
43	Plants perennial, rhizomatous or with stout, often woody taproot and branched (or sometimes in S. oregana), woody caudex	> 49
44	Mature calyces clavate, urceolate, or fusiform, contracted proximally around carpophore	> 45
44	Mature calyces campanulate, elliptic, or ovoid, not contracted at base around carpophore	> 47
45	Petals white, often pink tinged	Silene noctiflora
45	Petals bright pink	> 46
46	Petals unlobed	Silene pseudatocion
46	Petals 2-lobed	Silene pendula
47	Stems subglabrous to puberulent, often glutinous on distal internodes	Silene antirrhina
47	Stems coarsely hispid	> 48
48	Mature calyces (7-)10-15 mm; seeds grayish brown to black, ca. 1 mm diam.; stems (20-)50-100 cm	Silene dichotoma
48	Mature calyces 7-10 mm; seeds dark reddish brown, ca. 0.5 mm diam.; stems 15-45 cm	Silene gallica
49	Calyx lobes equaling or exceeding tube	Silene aperta
49	Calyx lobes shorter than tube. [50. Shifted to left margin.—Ed.]	> 50
50	Mature calyces clavate, funnelform, turbinate, urceolate, tubular, or fusiform, narrowed proximally around carpophore	> 51
50	Mature calyces campanulate, elliptic, or ovate, not contracted proximally around carpophore, or if so, broadly turbinate or obconic, at least 1/ 2 as broad as long	> 61
51	Corolla limbs unlobed or nearly so	Silene caroliniana
51	Corolla limbs lobed or laciniate	> 52
52	Calyx lobes 4-8 mm, lanceolate or narrowly triangular	> 53
52	Calyx lobes 2-4 mm, lanceolate to ovate or triangular-acute, or 2-5 mm with broad, obtuse, membranous margins	> 56
53	Corollas coral pink, pink, or white, limb 7-25 mm; calyces umbilicate or turbinate in fruit	> 54
53	Corollas pale greenish yellow or pale yellow to white, sometimes purple tinged, limb 5-8 mm; calyces clavate in fruit	> 55
54	Corolla limbs laciniate into many narrow segments; calyces thinly pilose,funnelform in flower, umbilicate in fruit	Silene polypetala
54	Corolla limbs 4-lobed, usually deeply so, rarely 2-lobed with 2 smaller lateral teeth; calyces canescent, eglandular or glandular, broadly tubular inflower, turbinate in fruit	Silene hookeri
55	Corolla limbs deeply laciniate into 6 or more linear lobes; calyces (20-)25-30 mm, densely puberulent and viscid-glandular, tubular in flower	Silene parishii
55	Corolla limbs cleft ca. to middle into (2-)4-8 lanceolate to oblong lobes; calyces 16-20 mm, densely and coarsely pubescent and viscid-glandular,tubular to narrowly obconic in flower	Silene wrightii
56	Leaf blades ovate-acuminate, sessile; mature calyces turbinate	Silene ovata
56	Leaf blades linear or spatulate to ovate-lanceolate or oblanceolate, at least proximal ones petiolate; mature calyces clavate to tubular or fusiform	> 57
57	Corolla limbs pink or rose red (rarely white), (7-)10-20 mm, with 4 (rarely 2 with 2 teeth) divergent, lanceolate lobes; mature calyces 15-38 mm	Silene occidentalis
57	Corolla limbs white, greenish, pink, or dingy red, 3-8 mm, with 2-6(-10) linear lobes; mature calyces 8-17(-20) mm	> 58
58	Corolla limbs 2-lobed (rarely with small lateral teeth)	> 59
58	Corolla limbs 4-6-lobed	> 60
59	Inflorescences open cymes with many elongate, ascending branchesand pedicels longer than calyx	Silene verecunda
59	Inflorescences pseudoracemose and nodding with flowers usually paired at nodes and pedicels ± equaling calyx, or erect with dense, many-flowered whorls and most flowers sessile or shortlypedicellate	Silene scouleri
60	Corolla appendages 2, laciniate, apex rounded, 2-3 mm; claw andfilaments ciliate; corolla lobes 4, linear	Silene bernardina
60	Corolla appendages 4-6, linear, apex acute, 1-1.5 mm; claw and filaments glabrous; corolla lobes 4-6, often forked, producing to 10 linear segments	Silene oregana
61	Styles exceeding 2 times calyx; stamens exserted from calyx	> 62
61	Styles and stamens ± equaling calyx and/or corolla	> 63
62	Leaves mostly in dense basal tufts	Silene lemmonii
62	Leaves mostly cauline	Silene bridgesii
63	Sepals and pedicels without glandular hairs	> 64
63	Glandular hairs present at least on pedicels and calyces. [65. Shifted to left margin.—Ed.]	> 65
64	Petals to 2 times calyx	Silene douglasii
64	Petals slightly longer than calyx	Silene nachlingerae
65	Leaves mostly basal, densely tufted, cauline leaves usually in 1-3 pairs and/or reduced	> 66
65	Basal leaves not densely tufted; cauline leaves in 3-12 pairs	> 72
66	Corollas slightly longer than sepals, limbs 1-2 mm, unlobed, or apex notched	Silene invisa
66	Corollas exceeding or sometimes equaling calyces; limbs 2-10 mm, 2(-4)-lobed	> 67
67	Plants small alpine species; stems 3-20(-30) cm; leaf blades fleshy, 0.5-5(-6) cm	> 68
67	Plants lowland and montane species; stems usually exceeding 30 cm; leaf blades not fleshy, 2-15 cm	> 70
68	Pubescence on calyces and pedicels with purple septa	Silene suksdorfii
68	Pubescence on calyces and pedicels with colorless septa	> 69
69	Seeds 2-3 mm, margins thickened and winglike; proximal leaf blades oblanceolate-spatulate, 2-7 mm wide	Silene grayi
69	Seeds ca. 1.5 mm, margins with large inflated papillae; proximal leaf blades linear-oblanceolate, 0.5-3 mm wide	Silene sargentii
70	Calyx lobes narrowly lanceolate with narrow membranous margins proximally	Silene nuda
70	Calyx lobes ovate to broadly triangular with broad membranous margins	> 71
71	Claws of corollas exceeding calyx, ciliate proximally; veins to calyx lobes conspicuously broadened distally and lance-shaped, commissural veins slender, not forked distally	Silene scaposa
71	Claws of corollas equaling calyx, glabrous; veins of calyx lobes not conspicuously broadened distally, commissural veins slender and forked distally, connate with those to lobes	Silene parryi
72	Cymes sessile (proximal ones may be pedunculate); pedicels usually shorter than calyxand becoming deflexed at base of calyx	Silene scouleri
72	Cymes pedunculate or flowers solitary; pedicels shorter than to exceeding calyx, ascending (may be partially deflexed in S. thurberi and S. petersonii)	> 73
73	Petals bright pink, limb 5-15 mm; plants rhizomatous from thick rootstock; stems5-15 cm	Silene petersonii
73	Petals white, pale pink, or yellowish, often tinged green or dusky purple, limb 3-8 mm; plants not rhizomatous; stems erect, usually exceeding 15 cm	> 74
74	Pubescence on calyces and pedicels short, deflexed, grayish white, eglandular(glandular hairs often present in var. rupinae)	Silene douglasii
74	Pubescence scabrid-puberulent or often viscid glandular-pubescent with septate hairs, glandular hairs present at least distally	> 75
75	Mature calyces ca. as broad as long; flowers nocturnal	Silene marmorensis
75	Mature calyces ca. 2 times as long as broad, flowers diurnal	> 76
76	Corolla limbs 5-7 mm, mature calyces 12-16 mm; stems softly viscid-puberulent distally	Silene parryi
76	Corolla limbs 2-4 mm, mature calyces 7-12 mm; stems scabridpuberulent	> 77
77	Calyx lobes 1-2 mm, ovate, as long as broad, corolla appendagesentire	Silene rectiramea
77	Calyx lobes 3-4 mm, narrowly lanceolate, much longer than broad, corolla appendages erose	Silene thurberi