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1. Caryophyllaceae A. L. Jussieu

石竹科 shi zhu ke

Authors: Dequan Lu, Zhengyi Wu, Lihua Zhou, Shilong Chen, Michael G. Gilbert, Magnus Lidén, John McNeill, John K. Morton, Bengt Oxelman, Richard K. Rabeler, Mats Thulin, Nicholas J. Turland & Warren L. Wagner

Dianthus chinensis
Image

Credit: Tatyana Shulkina

Herbs annual or perennial, rarely subshrubs or shrubs. Stems and branches usually swollen at nodes. Leaves opposite, decussate, rarely alternate or verticillate, simple, entire, usually connate at base; stipules scarious, bristly, or often absent. Inflorescence of cymes or cymose panicles, rarely flowers solitary or few in racemes, capitula, pseudoverticillasters, or umbels. Flowers actinomorphic, bisexual, rarely unisexual, occasionally cleistogamous. Sepals (4 or)5, free, imbricate, or connate into a tube, leaflike or scarious, persistent, sometimes bracteate below calyx. Petals (4 or)5, rarely absent, free, often comprising claw and limb; limb entire or split, usually with coronal scales at juncture of claw and limb. Stamens (2--)5--10, in 1 or 2 series. Pistil 1; carpels 2--5, united into a compound ovary. Ovary superior, 1-loculed or basally imperfectly 2--5-loculed. Gynophore present or absent. Placentation free, central, rarely basal; ovules (1 or) few or numerous, campylotropous. Styles (1 or)2--5, sometimes united at base. Fruit usually a capsule, with pericarp crustaceous, scarious, or papery, dehiscing by teeth or valves 1 or 2 × as many as styles, rarely berrylike with irregular dehiscence or an achene. Seeds 1 to numerous, reniform, ovoid, or rarely dorsiventrally compressed, abaxially grooved, blunt, or sharply pointed, rarely fimbriate-pectinate; testa granular, striate or tuberculate, rarely smooth or spongy; embryo strongly curved and surrounding perisperm or straight but eccentric; perisperm mealy.

Between 75 and 80 genera and ca. 2000 species: widespread but mainly of temperate or warm-temperate occurrence in the N hemisphere, with principal centers of distribution in the Mediterranean region and W Asia to W China and the Himalayas, fewer species in Africa S of the Sahara, America, and Oceania; 30 genera (two endemic) and 390 species (193 endemic) in China.

Arenaria, Silene, and Stellaria contain over half the species in the family in China. They are mostly concentrated in the Qinghai-Xizang plateau, and are especially rich from the Hengduan Mountains to the Himalayas. The main uses of this family are medicinal and ornamental. Dianthus superbus, Pseudostellaria heterophylla, Stellaria dichotoma var. lanceolata, and Vaccaria hispanica are commonly used in traditional Chinese medicine. Some species of Arenaria, Dianthus, Gypsophila, Psammosilene, and Silene are used as medicinal herbs among the people or are habitually used in local Chinese medicine. Many species of Dianthus, Gypsophila, Lychnis, Saponaria, and Silene are grown as ornamentals. Atocion armeria (Linnaeus) Rafinesque ( Silene armeria Linnaeus), native to Russia and Europe, is also cultivated in China. It differs from Silene in having a corymbose inflorescence and obscure calyx veins. Wu Cheng-yih, Ke Ping, Zhou Li-hua, Tang Chang-lin & Lu De-quan. 1996. Caryophyllaceae. In: Tang Chang-lin, ed., Fl. Reipubl. Popularis Sin. 26: 47–449.


1 Stipules present, rarely obscure (subfam. Paronychioideae).   (2)
+ Stipules absent.   (8)
       
2 (1) Fruit an achene; petals absent.   (3)
+ Fruit a capsule; petals present.   (4)
       
3 (2) Sepal apex aristate; pistil 3-carpeled; style apex 3-fid; leaves subulate-linear; shrublets   1 Gymnocarpos
+ Sepal apex not aristate; pistil 2-carpeled; style apex 2-fid; leaves oblong, elliptic, or subcordate; herbs perennial   2 Herniaria
       
4 (2) Styles free.   (5)
+ Styles connate at base or throughout.   (6)
       
5 (4) Styles 5; capsule 5-valved; leaves usually pseudoverticillate, stipules not connate   3 Spergula
+ Styles 3; capsule 3-valved; leaves decussate, stipules connate   4 Spergularia
       
6 (4) Sepals green, leaflike; petals 2--6-parted   5 Drymaria
+ Sepals white, scarious; petals entire or 2-lobed.   (7)
       
7 (6) Sepals centrally ridged abaxially, hyaline at margin; style apex 3-lobed; leaves obovate or spatulate   6 Polycarpon
+ Sepals not ridged, hyaline throughout; style apex entire; leaves linear or oblong   7 Polycarpaea
       
8 (1) Sepals free, rarely connate at base; petals nearly clawless, rarely absent; stamens often perigynous (subfam. Alsinoideae).   (9)
+ Sepals connate into a distinct calyx tube; petals usually clawed; stamens hypogynous (subfam. Caryophylloideae).   (23)
       
9 (8) Flowers of 2 types: chasmogamous at stem apex, with petals, usually sterile; cleistogamous at stem base, without petals, fertile; plants with fleshy root tubers   8 Pseudostellaria
+ Flowers not of 2 types: cleistogamous flowers absent; plants usually without fleshy root tubers.   (10)
       
10 (9) Capsule teeth as many as styles.   (11)
+ Capsule teeth 2 × as many as styles.   (14)
       
11 (10) Styles 4 or 5; petals shorter than or equaling sepals, rarely absent.   (12)
+ Styles 2 or 3; petals longer than sepals.   (13)
       
12 (11) Petals entire   9 Sagina
+ Petals 2-fid   10 Stellaria
       
13 (11) Styles 3; seeds numerous   11 Minuartia
+ Styles 2(or 3); seeds 1 or 2   12 Lepyrodiclis
       
14 (10) Styles (4 or)5.   (15)
+ Styles (2 or)3(or 4).   (17)
       
15 (14) Petals 2-fid for up to 1/3 their length, retuse, or rarely entire; capsule cylindric, usually greatly exceeding calyx, apex 8--10-toothed   13 Cerastium
+ Petals deeply 2-fid; capsule ovoid to shortly cylindric, shorter than or slightly exceeding calyx, apex 10-toothed, or 5-valved to middle with valves 2-fid at apex.   (16)
       
16 (15) Capsule shortly cylindric, shorter than calyx, apex 10-toothed; styles episepalous   14 Pseudocerastium
+ Capsule ovoid, slightly exceeding calyx, 5-valved to middle, valves 2-fid at apex; styles alternisepalous   15 Myosoton
       
17 (14) Petals 2-fid, rarely absent, lobes not toothed at apex.   (18)
+ Petals entire, rarely emarginate, toothed, or 2-fid with toothed lobes.   (19)
       
18 (17) Petals 2-fid for up to 1/3 their length; capsule cylindric, toothed at apex   13 Cerastium
+ Petals deeply 2-fid, rarely absent; capsule ovoid or globose, opening by valves   10 Stellaria
       
19 (17) Sepals connate below middle; seed testa spongy   16 Thylacospermum
+ Sepals free; seed testa not spongy.   (20)
       
20 (19) Seeds with membranous strophiole   17 Moehringia
+ Seeds without strophiole.   (21)
       
21 (20) Inflorescence umbellate; capsule cylindric; seeds dorsiventrally compressed   18 Holosteum
+ Inflorescence cymose, paniculate, or flowers solitary; capsule ovoid or oblong; seeds reniform or globose.   (22)
       
22 (21) Seeds few or numerous, rarely 1; sepals herbaceous; petals shorter or longer than sepals; herbs erect, diffuse, or cushionlike   19 Arenaria
+ Seed 1; sepals subscarious, semihyaline; petals much shorter than sepals; herbs climbing   20 Brachystemma
       
23 (8) Styles 3 or 5.   (24)
+ Styles 2(or 3).   (27)
       
24 (23) Fruit berrylike, drying when mature; irregularly dehiscent (Silene baccifera)   21 Silene
+ Fruit a capsule, with regular teeth.   (25)
       
25 (24) Calyx lobes leaflike, longer than tube; styles hairy   22 Agrostemma
+ Calyx teeth shorter than tube; styles glabrous.   (26)
       
26 (25) Capsule septicidal, 5-toothed; style base reflexed and persistent in fruit   23 Lychnis
+ Capsule loculicidal (usually also septicidal), 6- or 10-toothed; style base not persistent in fruit   21 Silene
       
27 (23) Calyx 5-winged   24 Vaccaria
+ Calyx wingless.   (28)
       
28 (27) Calyx with 1 to several pairs of bracts at base; seeds dorsiventrally compressed   25 Dianthus
+ Calyx without bracts at base; seeds reniform or ± so, rarely dorsiventrally compressed.   (29)
       
29 (28) Capsule membranous proximally, irregularly transversely dehiscent; seeds 1 or 2; leaves spiny   26 Acanthophyllum
+ Capsule dry and brittle, 4-toothed or valved; seeds numerous; leaves not spiny.   (30)
       
30 (29) Petals with coronal scales   27 Saponaria
+ Petals without coronal scales.   (31)
       
31 (30) Stamens 5; calyx 15-veined   28 Psammosilene
+ Stamens 10; calyx 5-veined.   (32)
       
32 (31) Seeds reniform   29 Gypsophila
+ Seeds dorsiventrally compressed   30 Petrorhagia

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