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Flora of China 16: 329–427. 1995. BORAGINACEAE 紫草科 zi cao ke Zhu Ge-ling1; Harald Riedl2, Rudolf Kamelin3 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. 1a. Ovary undivided; style terminal (at ovary apex). 2a. Style twice 2-cleft; stigmas 4; fruit drupes with 1 pyrene; cotyledons plicate .................................................. 1. Cordia 2b. 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. 3a. 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. 4a. Mature fruit with a fleshy or corky mesocarp .............................................................................. 7. Tournefortia 4b. Mature fruit dry, with thin mesocarp and bony endocarp ............................................................ 6. Heliotropium 3b. Style usually 2-cleft, rarely entire; stigmas 2, capitate or elongated, rarely subpeltate, or 1, 2-cleft, lower part not inflated, without sterile portion. 5a. Fruit dry, without evident mesocarp, divided at maturity into 4 1-seeded mericarps; annual herbs ............................................................................................................................................................. 5. Coldenia 5b. Fruit berrylike or drupe with evident fleshy mesocarp, undivided or divided at maturity into 2–4 mericarps; trees or shrubs. 6a. Style undivided; stigma somewhat 2-cleft, subpeltate, endocarp dividing into 4 1-seeded pyrenes ........................................................................................................................................................... 3. Rotula 6b. Style 2-cleft; stigmas 2. 7a. Style 2-cleft to below middle; endocarp undivided, ovoid; leaf blade densely white spotted adaxially ................................................................................................................................ 4. Carmona 7b. 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 1b. Ovary (2- or) 4-lobed; style gynobasic (between ovary lobes). 8a. Ovary 2-lobed; ovules 2, potentially maturing into 2 nutlets ....................................................................... 39. Rochelia 8b. Ovary 4-lobed; ovules 4, potentially maturing into 4 nutlets. 9a. Connective of anthers prominently subulate, spirally twisted .......................................................... 36. Trichodesma 9b. Connective of anthers not subulate, not spirally twisted. 10a. Corolla throat unappendaged. 11a. Anthers coherent at least at base, sagittate; corolla lobes toothed. 12a. Calyx parted to middle or slightly below, lobes triangular; corolla with inflated ribs below lobes and deep furrows between them ...................................................... 11. Maharanga ______ 1 Herbarium, Institute of Botany, Northwest Normal University, Shilidian, Lanzhou, Gansu 730070, People’s Republic of China. 2 Herbarium, Botanische Abteilung, Naturhistorisches Museum, Burgring 7, A-1014 Wien, Austria. 3 Herbarium, V. L. Komarov Botanical Institute, Russian Academy of Sciences, Prof. Popov Street 2, 197376 St. Petersburg, Russia. 12b. Calyx parted to base, lobes linear or linear-lanceolate; corolla without ribs or deep furrows ............................................................................................................................ 12. Onosma 11b. Anthers free, not sagittate at base; corolla lobes mostly not toothed. 13a. Stamens exserted; nutlet with a cupular emergence abaxially, attachment scar below middle adaxially; gynobase narrowly fastigiate ................................ 35. Craniospermum 13b. Stamens included; nutlet without a cupular emergence, attachment scar basal; gynobase flat. 14a. Corolla zygomorphic, lobes unequal ........................................................................ 13. Echium 14b. Corolla actinomorphic, lobes subequal. 15a. Style undivided; nutlet smooth, attachment scar with umbilical tissue ....... 14. Pulmonaria 15b. Style 2- or 4-divided; nutlet tuberculate, attachment scar without umbilical tissue. 16a. Nutlet with short stipe at base; stamens spirally arranged; style shortly 2-cleft ................................................................................. 10. Stenosolenium 16b. Nutlet without stipe; stamens in single whorl, if spirally arranged (Arnebia decumbens) then flowers yellow; style twice 2-cleft ...................................................................................................................... 9. Arnebia 10b. Corolla throat or tube with prominent appendages, longitudinal folds, or hair bands. 17a. Calyx lobes unequal, strongly enlarged in fruit, bilaterally compressed, becoming some- what clamshell-shaped .......................................................................................................... 38. Asperugo 17b. Calyx lobes subequal, slightly enlarged in fruit, not clamshell-shaped. 18a. Attachment scar concave, with umbilical tissue, margin with ringlike emergence. 19a. Corolla appendages below throat ................................................................................ 16. Nonea 19b. Corolla appendages at throat. 20a. Stamens and corolla appendages at same level; corolla lobes dentate ........ 18. Symphytum 20b. Stamens below corolla appendages; corolla lobes not dentate. 21a. Corolla appendages short pubescent; plants sparsely strigose or hispid ................................................................................................................... 15. Anchusa 21b. Corolla appendages pilose; plants short strigose, bristly .................. 17. Gastrocotyle 18b. Attachment scar not concave, without umbilical tissue or ringlike emergence. 22a. Anthers mucronulate at apex; nutlets ovoid, smooth, white or gray ................. 8. Lithospermum 22b. Anthers not mucronulate at apex; nutlets not ovoid, often not smooth and/or not white or gray. 23a. Nutlets with glochids or wings. 24a. Gynobase flat; stem leaves subglabrous. 25a. Abaxial surface of nutlet with cupular or crownlike emergence formed by confluent bases of glochids .................... 31. Actinocarya 25b. Abaxial surface of nutlet without emergence; glochids on adaxial surface coherent into wings ..................................... 32. Metaeritrichium 24b. Gynobase fastigiate or subulate; stem leaves bristly, rarely glabrous. 26a. Corolla limb shorter than tube; stamens and style exserted. 27a. Stamens inserted above appendages; anthers much shorter than filaments, base not hastate .......................................... 45. Solenanthus 27b. Stamens inserted below appendages; anthers ca. as long as to longer than filaments, base hastate. 28a. Nutlets globose or ovoid, ca. 15 mm, without prickles, broadly winged .................................................................. 44. Rindera 28b. Nutlets compressed, ovate, ca. 6 mm, margins without wings, with dense glochids ........................................... 43. Lindelofia 26b. Corolla limb ca. as long as to longer than tube; stamens and style included. 29a. Attachment scar at apex of nutlet. 30a. Abaxial cupular emergence of nutlet broadly winged ........................................................................ 47. Mattiastrum 30b. Abaxial cupular emergence of nutlet not winged ..................................................................................
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