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Lamiaceae Lindley

唇形科 chun xing ke

Authors: Xi-wen Li & Ian C. Hedge

Dracocephalum bullatum

Credit: Harvard University Herbaria

Herbs, sometimes subshrubs or shrubs, annual or perennial, usually aromatic. Stems and branches usually 4-angled. Leaves opposite, rarely whorled or alternate, simple to pinnately dissected or compound, without stipules. Inflorescences generally compound, sometimes flowers solitary and axillary; verticillasters 2- to many flowered, subtended by leaves or bracts. Flowers bisexual, zygomorphic, rarely subactinomorphic, bracteolate or not. Calyx persistent, 5-toothed, 2-lipped; upper lip 3-toothed or entire (deciduous in Scutellaria); lower lip 2- or 4-toothed; tube sometimes hairy annulate inside. Corolla limb usually 2-lipped; upper lip 2-lobed and lower 3-lobed, rarely upper lip entire and lower 4-lobed, also rarely limb (4- or) 5-lobed; tube hairy annulate inside. Stamens epipetalous, 4 or 2, free, rarely filaments connate, sometimes one staminodial; anther 1- or 2-celled, usually dehiscing longitudinally; disc persistent. Ovary superior, 2-celled and each cell 2-ovuled and style subterminal, or ovary 4-parted and each lobe 1-ovuled and style gynobasic (from bases of ovary lobes) with 2-cleft apex. Fruit usually 4 dry nutlets. Seeds with or without endosperm.

Approximately 3500 species in 220 genera, distributed worldwide, but mostly in the Mediterranean region and SW Asia. China has 807 species in 96 genera.

Cantino (Taxon 40: 441. 1991) suggests that Cardioteucris cordifolia C. Y. Wu, originally placed in the Lamiaceae, is identical with Caryopteris siccanea W. Smith (Verbenaceae, q.v.). Although the original author agrees with the identity of Cardioteucris cordifolia and Cardiopteris siccaneae, he believes that the generic placement of Cardioteucris is in the Lamiaceae because of its 2-lipped calyx and deeply 4-divided ovary. The last feature is aberrant in Caryopteris and all other Verbenaceae.

Wu Cheng-yih & Li Hsi-wen, eds. 1977. Labiatae. Fl. Reipubl. Popularis Sin. 65(2): 1-649, 66: 1-647.

Key 4

1 Calyx with very dissimilar teeth, throat closed in fruit by 2 obliquely upwardly directed teeth of lower lip; upper corolla lip galeate   25 Prunella
+ Calyx with ± unlike teeth, throat open in fruit.   (2)
2 (1) Calyx with 5, obtuse, triangular equal teeth or with 3 or 4 lobes; corolla broad at base or gradually dilated ascending, with a wider, somewhat convex upper lip   26 Chelonopsis
+ Calyx with acute, subulate, or rarely spinescent lanceolate teeth; corolla upper lip convex, always helmet-shaped, rarely flat.   (3)
3 (2) Upper lip of corolla mostly short, ± flat (except in Colquhounia), glabrous or sparsely hairy.   (4)
+ Upper lip of corolla convex or galeate, rarely nearly flat, always densely hairy.   (10)
4 (3) Anther cells confluent at apex; nutlets winged or scaly; shrubs or herbs.   (5)
+ Anther cells separate (except in Microtoena); nutlets not winged (except in Chamaesphacos) or scaly; herbs.   (6)
5 (4) Shrubs; nutlets membranous winged at apex; corolla large, scarlet   49 Colquhounia
+ Dwarf herbs; nutlets densely linear scaly at apex and adaxial side   50 Achyrospermum
6 (4) Anthers 1-celled, longitudinally dehiscent; corolla tube saccate in front   51 Eurysolen
+ Anthers 2-celled or anterior anthers 2-celled and posterior anthers 1-celled; corolla tube not saccate in front.   (7)
7 (6) Anterior anthers transversely 2-celled, cells parallel, posterior anthers 1-celled; calyx 5-dentate, only slightly dilated in fruit; verticillasters arranged in continuous or interrupted, long terminal spikes   52 Anisomeles
+ Anther cells of both pairs equal, divaricate; calyx deformed or dilated in fruit; inflorescences of terminal loose panicles, rarely verticillasters axillary.   (8)
8 (7) Upper corolla lip falcate, sometimes very short; nutlets wingless; inflorescences of loose terminal panicles.   (9)
+ Upper corolla lip flat; nutlets narrowly winged at apex and on 2 sides; leaves spiny on margin; verticillasters axillary, 2-6-flowered   55 Chamaesphacos
9 (8) Upper corolla lip very short; anther cells separate, divaricate; flowers very small; calyx equally 5-toothed   53 Craniotome
+ Upper corolla lip galeate, long; filaments sometimes bearded near base; anther cells divaricate, confluent at apex; flowers large; one posterior tooth of calyx sometimes dilated   54 Microtoena
10 (3) Calyx teeth 5, spinescent, hooked at apex   27 Notochaete
+ Calyx teeth 5-10, not spinescent and hooked at apex.   (11)
11 (10) Style lobes unequal in length (sometimes equal in Phlomis), posterior much shorter than anterior.   (12)
+ Style lobes subequal or equal in length.   (15)
12 (11) Nutlets prominently hairy at apex; filaments with comblike-fimbriate appendages at base; calyx teeth truncate, apiculate at apex; desert or desert-meadow plants   28 Eremostachys
+ Nutlets slightly hairy or glabrous; other characters not as above.   (13)
13 (12) Calyx 10-toothed   29 Leucas
+ Calyx 5-toothed.   (14)
14 (13) Upper corolla lip always hairy or fringed-incised; posterior stamens mostly appendiculate at base; mostly erect herbs or subshrubs; leaf venation not fan-shaped; verticillasters axillary, lax or dense   30 Phlomis
+ Upper corolla lip not fringed or incised; posterior stamens not appendiculate at base; stemless herbs; leaf venation fan-shaped; verticillasters in a dense scapose capitulum or short spike   31 Lamiophlomis
15 (11) Anther cells transversely 2-valved; lower lip of corolla with an apophysis on connective between middle and lateral lobes   32 Galeopsis
+ Anther cells parallel or spreading in vertical or oblique position; lower lip of corolla without apophyses.   (16)
16 (15) Nutlets ovoid, rounded at apex.   (17)
+ Nutlets ± acutely 3-angled, not truncate at apex (except in Loxocalyx and partly in Lagochilus).   (22)
17 (16) Upper corolla lip galeate, longer than lower lip, bearded; calyx obconic-campanulate, 10-veined, equally 5-toothed   44 Paraphlomis
+ Upper corolla lip shorter than or equal to, rarely longer than lower lip, if equal or longer then calyx 5-8(-10)- or 11-veined.   (18)
18 (17) Calyx 2-lipped, 5-8(-10)- or 11-veined.   (19)
+ Calyx 5-toothed, 5-10-veined; teeth equal in length, rarely calyx 2-lipped.   (20)
19 (18) Calyx 5-8(-10)-veined, upper lip 3-toothed, lower 2-toothed; plants without woody underground stem; verticillasters axillary; stamens much exserted   42 Loxocalyx
+ Calyx 11-veined, 2-lipped, upper lip 2-toothed, lower lip 3-toothed; plants with woody underground stem; verticillasters in terminal racemes; stamens slightly exserted   45 Ombrocharis
20 (18) Calyx teeth needlelike; shrubs with many spines arising from base of verticillasters and leaf axils; leaf margin spiny   40 Lagochilus
+ Calyx teeth not needlelike; spineless herbs.   (21)
21 (20) Verticillasters many flowered, densely arranged in short cylindric or subcapitate spikes; anther cells parallel; corolla tube exserted, not villous annulate inside; anther cells parallel   46 Betonica
+ Verticillasters 2- to many flowered, not in dense spikes; anther cells divaricate; corolla tube included or exserted, villous annulate or not at middle inside   47 Stachys
22 (16) Calyx 2-lipped, unequally 5-toothed, upper lip 3-toothed and lower 2-toothed, or upper lip entire and lower 4-toothed.   (23)
+ Calyx inconspicuously 2-lipped, equally or subequally 5-toothed.   (24)
23 (22) Calyx tubular, upper lip 3-toothed, lower 2-toothed; verticillasters axillary   42 Loxocalyx
+ Calyx broadly campanulate, upper lip entire, lower lip 4-toothed; verticillasters in dense terminal racemose panicles   43 Paralamium
24 (22) Posterior filaments appendiculate or thickened at base; desert or alpine plants.   (25)
+ Posterior filaments not appendiculate at base; plants of various habitats.   (26)
25 (24) Calyx tubular-campanulate, teeth subulate or spinescent; nutlets hairy at apex; tall desert plants   28 Eremostachys
+ Calyx broadly campanulate, teeth triangular; nutlets smooth; dwarf herbs of strongly weathered stony alluvial fans   41 Eriophyton
26 (24) Throat of corolla dilated, tube elongated; calyx teeth not spinescent.   (27)
+ Throat of corolla barely dilated, tube slightly exserted or included; calyx teeth ± needlelike or spinescent.   (29)
27 (26) Middle lobe of lower corolla lip largest; non-alpine plants; leaves various.   (28)
+ Lateral lobes of lower corolla lip larger, ovate; alpine plants; leaves broadly rhombic or cuneate fan-shaped, tomentose   35 Alajja
28 (27) Lateral lobes of lower corolla lip not developed, with a small, acute tooth on margin; anther cells divaricate, hairy   33 Lamium
+ Lateral lobes of lower corolla lip well developed, with acute teeth on margin; anther cells divergent, glabrous   34 Galeobdolon
29 (26) Calyx teeth needlelike; shrubs with spines at base of verticillasters and in leaf axils; leaf margin spiny   40 Lagochilus
+ Calyx teeth spinescent; spineless plants.   (30)
30 (29) Anther cells divaricate; leaves ovate, entire.   (31)
+ Anther cells parallel; leaves other than ovate, dissected.   (32)
31 (30) Corolla 6-7 mm, slightly longer than calyx; nutlets pubescent at apex   36 Chaiturus
+ Corolla more than 10 mm, much longer than calyx; nutlets glabrous   37 Stachyopsis
32 (30) Calyx funnelform, 5-veined; corolla purple-red, reddish, or white, tube puberulent or hairy annulate inside   38 Leonurus
+ Calyx tubular-campanulate, 10-veined; corolla yellowish white, tube not hairy annulate inside   39 Panzerina

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