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FNA | Family List | FNA Vol. 5 | Caryophyllaceae

36. Silene Linnaeus, Sp. Pl. 1: 416. 1753. Gen. Pl. ed. 5, 193. 1754.
[name conserved]

Campion, catchfly [Greek seilenos, probably derived from Silenus, the intoxicated foster father of the Greek god Bacchus, who was described as covered with foam; perhaps allud-ing to the viscid secretion covering many species]

John K. Morton

Anotites Greene; Atocion Adanson; Coronaria Guettard; Gastrolychnis (Fenzl) Reichenbach; Lychnis Linnaeus; Melandrium J. C. Röhling; Physolychnis (Bentham) Ruprecht; Viscaria Bernhardi; Wahlbergella Fries

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.

Species ca. 700 (70 in the flora): mainly Northern Hemisphere.

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.


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

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