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PLANTAGINACEAE 1. PLANTAGO Linnaeus, Sp. Pl. 1: 112. 1753

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PLANTAGINACEAE 1. PLANTAGO Linnaeus, Sp. Pl. 1: 112. 1753 PLANTAGINACEAE 车前科 che qian ke Li Zhenyu (李振宇)1, Wei Lai (魏来)1; Ronald K. Hoggard2 Herbs or small shrubs, rarely arborescent. Stipules absent. Leaves spirally alternate, usually in a basal rosette, rarely opposite or whorled, simple; leaf blade veins (1 or)3–11, arcuate. Inflorescences spikes, rarely racemes or 1-flowered, pedunculate, each flower with 1 bract. Flowers small, protogynous, bisexual, rarely polygamous or unisexual. Calyx usually 4-parted, persistent, segments imbricate. Corolla scarious or membranous, gamopetalous, actinomorphic or rarely zygomorphic, (3 or)4-lobed; lobes imbricate. Stamens (1 or 2 or)4, inserted on corolla tube, alternate with lobes; anthers free, versatile, with an apical appendage, thecae 2 and parallel. Pistil consisting of 2 carpels; ovary superior, (1 or)2-loculed, axile or rarely basal; style 1, filiform, mostly longitudinally stigmatic. Fruit a pyxis, rarely an indehiscent capsule or nutlet, with 1 to numerous seeds. Seeds minute; cotyledons narrow; endo- sperm fleshy. Two genera and more than 210 species: cosmopolitan; one genus and 22 species (three endemic, four introduced) in China. Li Zhenyu. 2002. Plantaginaceae. In: Hu Chiachi, ed., Fl. Reipubl. Popularis Sin. 70: 318–345. 1. PLANTAGO Linnaeus, Sp. Pl. 1: 112. 1753. 车前属 che qian shu Bougueria Decaisne; Psyllium Miller. Herbs, or rarely small shrubs or arborescent, annual, biennial, or perennial, acaulescent or infrequently stemmed, with a taproot, caudex, or only numerous fibrous roots. Leaves simple; petiole vaginate at base; leaf blade ovate, elliptic, oblong, lanceolate, linear, or subulate, margin entire, repand, erose, or toothed, rarely pinnately or palmately cleft. Inflorescences arising from rosette or leaf axil of stem, spikes, narrowly cylindric, capitate, or rarely 1-flowered. Corolla salverform or tubular, persistent; tube cylindric or constricted at throat, as long as or slightly longer than calyx; limb 4-parted. Stamens (1 or 2 or)4; anthers ovoid, suborbicular, ellip- soid, or oblong. Ovary 2-loculed or rarely 1-locular but sometimes with an apical third compartment, placentation axile or rarely basal, with 1 to numerous ovules per locule. Fruit a pyxis, rarely an indehiscent capsule or nutlet, with 1 to numerous seeds. Seeds peltately inserted; testa mucilaginous when wet; embryo straight; cotyledons parallel or vertical to ventral (hilum) side. About 200 species: cosmopolitan; 22 species (three endemic, four introduced) in China. 1a. Stem erect, with elongated internodes; leaves opposite, rarely in whorls of 3; corolla brownish .............................. 22. P. arenaria 1b. Stem absent or very short; leaves usually in basal rosette, rarely densely alternate; corolla white or yellowish. 2a. Plant with only fibrous roots. 3a. Leaf blade lanceolate to linear-lanceolate, 3–4 × as long as wide; corolla yellowish; anthers 1.8–2.2 mm ................................................................................................................................................. 7. P. fengdouensis 3b. Leaf blade broadly ovate to broadly elliptic, less than 2 × as long as wide; corolla white; anthers 0.8–1.5 mm. 4a. Flower sessile; pyxis circumscissile at or just below middle, with (8–)12–14(–34) seeds; seeds 0.8–1.2 mm ...................................................................................................................................................... 3. P. major 4b. Flower shortly pedicellate; pyxis circumscissile near base, with (2–)5–15 seeds; seeds (1.2–)1.3–2 mm. 5a. Sepals acuminate at apex, keel projecting beyond apex; anthers ca. 0.8 mm .................................. 5. P. cavaleriei 5b. Sepals obtuse or acute at apex, keel not projected at apex; anthers 1–1.5 mm. 6a. Petiole not winged at middle; corolla lobes narrowly triangular; anthers 1–1.2 mm .................... 4. P. asiatica 6b. Petiole broadly winged; corolla lobes ovate to broadly ovate; anthers ca. 1.5 mm ............... 6. P. gentianoides 2b. Plant with a taproot. 7a. Leaf blade broadly ovate to elliptic, veins 7–11; petiole retrorsely pubescent; stamens adnate to near base of corolla tube. 8a. Plants turning black when dry; petiole longer than or equaling blade, not winged; bracts glabrous ......... 1. P. maxima 8b. Plants not turning black when dry; petiole much shorter than blade, winged; bracts pubescent ................. 2. P. media 7b. Leaf blade obovate, elliptic, lanceolate, or linear, veins 1–5(or 7); petiole with patent, appressed upward, or rarely entangled trichomes; stamens adnate to middle of corolla tube or more apically. 9a. Corolla tube pubescent or lobes hirsute-villous outside. 1 Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 2 Robert Bebb Herbarium, 770 Van Vleet Oval, University of Oklahoma, Norman, Oklahoma 73019-0245, U.S.A. 495 496 PLANTAGINACEAE 10a. Leaf blade papery, sparsely villous; corolla lobes densely hirsute-villous outside; stamens adnate to near apex of corolla tube ........................................................................................... 20. P. lagocephala 10b. Leaf blade leathery, sparsely strigillose or glabrous; corolla tube pubescent; stamens adnate to middle of corolla tube ..................................................................................................... 21. P. maritima 9b. Corolla glabrous. 11a. Lower sepals connate to near apex; stamens adnate to middle of corolla tube .......................... 16. P. lanceolata 11b. All sepals distinct; stamens adnate to or near apex of corolla tube. 12a. Bracts with linear to subulate apex, much longer than flower ................................................ 17. P. aristata 12b. Bracts with obtuse or acute apex, shorter, equaling, or slightly longer than flower. 13a. Leaf blade glabrous or abaxially pubescent, vein 1; seeds 7–30 per pyxis, fusiform. 14a. Bracts broadly ovate to suborbicular; sepals broadly elliptic to orbicular; pyxis with 12–30 seeds .................................................................................. 14. P. polysperma 14b. Bracts ovate; sepals obovate; pyxis with 7–15 seeds ....................................... 15. P. tenuiflora 13b. Leaf blade hairy on both surfaces, veins (1 or)3–7; seeds 1–4 per pyxis, ellipsoid to oblong. 15a. Leaf blade elliptic, ovate-lanceolate, obovate-lanceolate, or spatulate, usually more than 1 cm wide. 16a. Leaf blade spatulate to obovate-lanceolate, veins (3 or)5; flowers heteromorphic; corolla yellowish, lobes erect when flowers fertile; seeds 2 per pyxis, ventral surface grooved .................................................. 8. P. virginica 16b. Leaf blade elliptic to ovate-lanceolate, rarely obovate-elliptic, veins 5 or 7; flowers homomorphic; corolla white, lobes patent to reflexed; seeds 4 per pyxis, ventral surface flat or prominent but not grooved. 17a. Bracts triangular-ovate; corolla lobes 0.5–1 mm; anthers white; pyxis 4–5 mm ..................................................................................... 9. P. depressa 17b. Bracts ovate-elliptic; corolla lobes 1–1.5 mm; anthers reddish brown; pyxis 2.5–3 mm ............................................................. 10. P. camtschatica 15b. Leaf blade ± linear to narrowly lanceolate or narrowly elliptic, usually less than 1 cm wide. 18a. Leaf blade arachnoid or silky villous; spikes cylindric to narrowly cylindric. 19a. Leaf blade arachnoid, veins 1 or 3, margin entire or repand- crenate; spikes 1–2.5(–5) cm; flowers dense; anthers ca. 1.2 mm .................................................................................. 13. P. arachnoidea 19b. Leaf blade sparsely silky villous, veins 3 or 5, margin remotely and retrorsely denticulate or triangular dentate; spikes 4–10 cm; flowers usually interrupted at base; anthers ca. 2 mm ........................................................................................... 12. P. perssonii 18b. Leaf blade villous or lanate; spikes capitate or ovoid at first but later shortly cylindric. 20a. Keel not extending to apex of sepals; seeds 3 or 4 per pyxis, black, (1–)1.5–2 mm, ventral face flat; cotyledons parallel to ventral side .................................................................................. 11. P. komarovii 20b. Keel extending to apex of sepals; seeds 1 or 2 per pyxis, yellowish brown to brown, (2–)3–4 mm, ventral face with a groove; cotyledons vertical to ventral side. 21a. Plants with short stems, usually branched at base; corolla lobes orbicular-ovate .................................................................... 18. P. ovata 21b. Plants stemless; corolla lobes elliptic to narrowly ovate ........... 19. P. minuta 1. Plantago maxima Jussieu ex Jacquin, Collectanea 1: 82. 7–11, base broadly cuneate to decurrent onto petiole, margin 1787. entire or remotely repand-crenate, apex acute to rounded. Spikes cylindric, 6–20 cm, densely flowered; peduncle 20–50 巨车前 ju che qian cm, pubescent; bracts narrowly ovate, slightly shorter than ca- Herbs, perennial, robust. Taproot cylindric. Leaves basal; lyx, glabrous, keel apparent. Sepals narrowly elliptic to ovate- petiole 8–20 cm, densely retrorsely pubescent; blade broadly elliptic, 2.2–2.5 mm, glabrous, keel broad, extending to apex. elliptic, broadly ovate, or broadly obovate, 8–20 × 5–14 cm, Corolla white, glabrous; lobes ovate-lanceolate, 1.5–2 mm, thickly papery, turning black when dry, white pubescent, veins patent to reflexed, apex acute. Stamens lilac turning black when PLANTAGINACEAE
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