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Boraginaceae A. L. Jussieu

紫草科 zi cao ke

Authors: Gelin Zhu, Harald Riedl & Rudolf V. Kamelin

Herbs perennial, biennial, or annual, less often lianas, shrubs, or trees, usually bristly or scabrous-pubescent. Leaves simple, exstipulate, alternate, rarely opposite, entire or serrate at margin. Inflorescences often double scorpioid cymes, rarely solitary; bracts present or absent. Flowers bisexual, actinomorphic, rarely zygomorphic. Calyx usually 5-parted or lobed, mostly persistent. Corolla tubular, campanulate, rotate, funnelform, or salverform; tube appendages 5, rarely more, mostly trapeziform, rarely absent, sometimes a ring of hairs present; limb usually 5-parted; lobes overlapping, rarely twisted in bud. Stamens 5, inserted on corolla tube or rarely at throat, included or rarely exserted; anthers introrse, 2-loculed, usually dorsifixed at base, less often medifixed, dehiscence longitudinal. Nectaries at base of corolla tube or on disc below ovary. Ovary superior, 2-carpellate; locules 2 and each with 2 ovules, or 4 and each with 1 ovule; ovules nearly atropous, semianatropous, or anatropous. Style terminal or gynobasic, branched or not. Gynobase flat, fastigiate, or subulate. Fruit 1-4-seeded drupes or nutlets (mericarps); nutlets mostly dry, often ornamented with wings, prickles and/or glochids (stiff bristles with barbed or anchorlike tips). Seeds vertical or oblique, coat membranous; embryo straight, less often curved; cotyledons flat, fleshy.

About 156 genera and 2500 species: temperate and tropical regions, centered in the Mediterranean region; 47 genera and 294 species in China, of which four genera and 156 species are endemic.

Anchusa italica Retzius, A. officinalis Linnaeus, and Borago officinalis Linnaeus are cultivated. Borago officinalis is uncommon and not naturalized, and because its seeds contain quality oil, it is probably more commonly grown than previously.

Wang Wen-tsai, Liu Yu-lan, Zhu Ge-ling, Lian Yong-shan, Wang Jing-quan & Wang Qing-rui in Kung Xian-wu & Wang Wen-tsai, eds. 1989. Boraginaceae. Fl. Reipubl. Popularis Sin. 64(2): 1-253.


1 Ovary undivided; style terminal (at ovary apex).   (2)
+ Ovary (2- or) 4-lobed; style gynobasic (between ovary lobes).   (8)
       
2 (1) Style twice 2-cleft; stigmas 4; fruit drupes with 1 pyrene; cotyledons plicate   1 Cordia
+ Style 2-cleft or entire; stigmas 1 or 2; fruit drupes or nutletlike, usually divided into 2 or 4 pyrenes, rarely undivided; cotyledons not plicate.   (3)
       
3 (2) Style divided into a basal sterile terete portion that may be strongly reduced, and an upper conical portion; stigma 1, often forming a ring around style.   (4)
+ Style usually 2-cleft, rarely entire; stigmas 2, capitate or elongated, rarely subpeltate, or 1, 2-cleft, lower part not inflated, without sterile portion.   (5)
       
4 (3) Mature fruit with a fleshy or corky mesocarp   7 Tournefortia
+ Mature fruit dry, with thin mesocarp and bony endocarp   6 Heliotropium
       
5 (3) Fruit dry, without evident mesocarp, divided at maturity into 4 1-seeded mericarps; annual herbs   5 Coldenia
+ Fruit berrylike or drupe with evident fleshy mesocarp, undivided or divided at maturity into 2-4 mericarps; trees or shrubs.   (6)
       
6 (5) Style undivided; stigma somewhat 2-cleft, subpeltate, endocarp dividing into 4 1-seeded pyrenes   3 Rotula
+ Style 2-cleft; stigmas 2.   (7)
       
7 (6) Style 2-cleft to below middle; endocarp undivided, ovoid; leaf blade densely white spotted adaxially   4 Carmona
+ Style 2-cleft to above middle; endocarp divided into 2 2-seeded, rarely 4 1-seeded pyrenes; leaf blade not white spotted adaxially   2 Ehretia
       
8 (1) Ovary 2-lobed; ovules 2, potentially maturing into 2 nutlets   39 Rochelia
+ Ovary 4-lobed; ovules 4, potentially maturing into 4 nutlets.   (9)
       
9 (8) Connective of anthers prominently subulate, spirally twisted   36 Trichodesma
+ Connective of anthers not subulate, not spirally twisted.   (10)
       
10 (9) Corolla throat unappendaged.   (11)
+ Corolla throat or tube with prominent appendages, longitudinal folds, or hair bands.   (17)
       
11 (10) Anthers coherent at least at base, sagittate; corolla lobes toothed.   (12)
+ Anthers free, not sagittate at base; corolla lobes mostly not toothed.   (13)
       
12 (11) Calyx parted to middle or slightly below, lobes triangular; corolla with inflated ribs below lobes and deep furrows between them   11 Maharanga
+ Calyx parted to base, lobes linear or linear-lanceolate; corolla without ribs or deep furrows   12 Onosma
       
13 (11) Stamens exserted; nutlet with a cupular emergence abaxially, attachment scar below middle adaxially; gynobase narrowly fastigiate   35 Craniospermum
+ Stamens included; nutlet without a cupular emergence, attachment scar basal; gynobase flat.   (14)
       
14 (13) Corolla zygomorphic, lobes unequal   13 Echium
+ Corolla actinomorphic, lobes subequal.   (15)
       
15 (14) Style undivided; nutlet smooth, attachment scar with umbilical tissue   14 Pulmonaria
+ Style 2- or 4-divided; nutlet tuberculate, attachment scar without umbilical tissue.   (16)
       
16 (15) Nutlet with short stipe at base; stamens spirally arranged; style shortly 2-cleft   10 Stenosolenium
+ Nutlet without stipe; stamens in single whorl, if spirally arranged (Arnebia decumbens) then flowers yellow; style twice 2-cleft   9 Arnebia
       
17 (10) Calyx lobes unequal, strongly enlarged in fruit, bilaterally compressed, becoming somewhat clamshell-shaped   38 Asperugo
+ Calyx lobes subequal, slightly enlarged in fruit, not clamshell-shaped.   (18)
       
18 (17) Attachment scar concave, with umbilical tissue, margin with ringlike emergence.   (19)
+ Attachment scar not concave, without umbilical tissue or ringlike emergence.   (22)
       
19 (18) Corolla appendages below throat   16 Nonea
+ Corolla appendages at throat.   (20)
       
20 (19) Stamens and corolla appendages at same level; corolla lobes dentate   18 Symphytum
+ Stamens below corolla appendages; corolla lobes not dentate.   (21)
       
21 (20) Corolla appendages short pubescent; plants sparsely strigose or hispid   15 Anchusa
+ Corolla appendages pilose; plants short strigose, bristly   17 Gastrocotyle
       
22 (18) Anthers mucronulate at apex; nutlets ovoid, smooth, white or gray   8 Lithospermum
+ Anthers not mucronulate at apex; nutlets not ovoid, often not smooth and/or not white or gray.   (23)
       
23 (22) Nutlets with glochids or wings.   (24)
+ Nutlets without glochids or wings.   (34)
       
24 (23) Gynobase flat; stem leaves subglabrous.   (25)
+ Gynobase fastigiate or subulate; stem leaves bristly, rarely glabrous.   (26)
       
25 (24) Abaxial surface of nutlet with cupular or crownlike emergence formed by confluent bases of glochids   31 Actinocarya
+ Abaxial surface of nutlet without emergence; glochids on adaxial surface coherent into wings   32 Metaeritrichium
       
26 (24) Corolla limb shorter than tube; stamens and style exserted.   (27)
+ Corolla limb ca. as long as to longer than tube; stamens and style included.   (29)
       
27 (26) Stamens inserted above appendages; anthers much shorter than filaments, base not hastate   45 Solenanthus
+ Stamens inserted below appendages; anthers ca. as long as to longer than filaments, base hastate.   (28)
       
28 (27) Nutlets globose or ovoid, ca. 15 mm, without prickles, broadly winged   44 Rindera
+ Nutlets compressed, ovate, ca. 6 mm, margins without wings, with dense glochids   43 Lindelofia
       
29 (26) Attachment scar at apex of nutlet.   (30)
+ Attachment scar at or below middle on adaxial surface of nutlets.   (31)
       
30 (29) Abaxial cupular emergence of nutlet broadly winged   47 Mattiastrum
+ Abaxial cupular emergence of nutlet not winged   42 Cynoglossum
       
31 (29) Gynobase pyramidal, much shorter than nutlets or nearly flat.   (32)
+ Gynobase subulate, ca. as long as to longer than nutlets.   (33)
       
32 (31) Gynobase short pyramidal, to 2-3 mm; fruiting pedicel always deflexed   29 Hackelia
+ Gynobase nearly flat, to 1 mm; fruiting pedicel erect or deflexed   28 Eritrichium
       
33 (31) Adaxial surface of nutlet joined to gynobase (not easily stripped)   34 Heterocaryum
+ Adaxial surface of nutlet free from gynobase (easily stripped)   33 Lappula
       
34 (23) Nutlets tetrahedral or lenticular.   (35)
+ Nutlets neither tetrahedral nor lenticular.   (40)
       
35 (34) Nutlet with adaxial membranous cupular emergence.   (36)
+ Nutlet without membranous cupular emergence.   (37)
       
36 (35) Leaf blade ovate-cordate; fruiting calyx extremely enlarged, enclosing fruit   21 Sinojohnstonia
+ Leaf blade elliptic-ovate; fruiting calyx slightly enlarged, not enclosing fruit   22 Omphalotrigonotis
       
37 (35) Stamens included.   (38)
+ Stamens exserted.   (39)
       
38 (37) Corolla lobes convolute; nutlets lenticular   19 Myosotis
+ Corolla lobes overlapping; nutlets tetrahedral or 3-angled   20 Trigonotis
       
39 (37) Leaves nearly whorled at apex of stem   23 Brachybotrys
+ Leaves alternate along stem   24 Mertensia
       
40 (34) Stamens exserted   41 Antiotrema
+ Stamens included.   (41)
       
41 (40) Flowers solitary, terminal; perennial alpine cushion plant   27 Chionocharis
+ Flowers several to numerous, in cymes.   (42)
       
42 (41) Nutlet without an emergence on either surface.   (43)
+ Nutlets with a discoid or cupular emergence on abaxial or adaxial surface.   (45)
       
43 (42) Gynobase nearly flat; nutlets glabrous, shiny   25 Amblynotus
+ Gynobase subulate or terete.   (44)
       
44 (43) Nutlets short hispid, abaxial surface completely coherent to gynobase   37 Lasiocaryum
+ Nutlets glabrous, abaxial surface free except for basal attachment scar which is coherent to gynobase   26 Microcaryum
       
45 (42) Cupular emergence with 2 layers, margin of outer layer dentate   46 Thyrocarpus
+ Cupular emergence with 1 layer, if 2, margin of outer layer entire.   (46)
       
46 (45) Nutlet with cupular emergence adaxially   40 Bothriospermum
+ Nutlets with cupular emergence on abaxial surface, surrounding an aperture   30 Microula

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