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FOC | Family List | FOC Vol. 18 | Scrophulariaceae

38. Veronica Linnaeus, Sp. Pl. 1: 9. 1753.

婆婆纳属 po po na shu

Hong De-yuan; Manfred A. Fischer

Herbs, perennial with rhizomes or annuals. Roots glabrous. Stems erect, procumbent, or creeping, base sometimes woody. Leaves mostly opposite, rarely whorled or upper ones alternate. Inflorescences terminal or axillary racemes, sometimes spicate, long or short and capitate. Calyx 4-lobed, if 5-lobed upper lobe much smaller than other lobes. Corolla 4-sect or 4-lobed, rarely slightly 2-lipped; tube short and limb subrotate, rarely tube conspicuous, 1/3-1/2 of corolla length, usually hairy inside; lobes usually patent, unequal in width, upper lobe widest, lower lobe narrowest. Stamens 2; filaments adnate to posterior part of corolla tube; anther locules parallel or divergent, apically confluent. Style persistent; stigma capitate. Capsule diverse in shape, slightly to strongly compressed laterally, 1-grooved, loculicidal. Seeds 2-50 per capsule, flattened and convex on both sides, navicular or cupular.

About 250 species: cosmopolitan, mainly in Asia and Europe; 53 species in China.

1 Inflorescences terminal, sometimes bracts leaflike and flowers appearing axillary.   (2)
+ Inflorescences axillary, usually in pairs, sometimes arrested development of stem apex causes inflorescences to appear terminal.   (20)
2 (1) Perennials with rhizomes or woody roots.   (3)
+ Annuals without rhizomes.   (9)
3 (2) Calyx lobes 4; corolla subrotate, tube inconspicuous; capsule strongly compressed.   (4)
+ Calyx lobes (4 or)5; corolla tube mostly conspicuous, 1.5-2 mm, rarely shorter; capsule slightly compressed.   (5)
4 (3) Capsule obovoid, longer than wide; inflorescences few flowered; stems erect or ascending, never creeping   6 Veronica stelleri var. longistyla
+ Capsule subreniform, shorter than wide; inflorescences many flowered; stems often creeping at base   7 Veronica serpyllifolia
5 (3) Plants woolly, white   5 Veronica lanuginosa
+ Plants tomentose or villous, with multicellular glandular hairs, green or gray-green.   (6)
6 (5) Inflorescences elongated, lax; infructescences 10-20 cm; leaf blade lanceolate to ovate-lanceolate, margin deeply cut   2 Veronica lanosa
+ Inflorescences capitate; infructescences less than 5 cm; leaf blade orbicular to ovate, margin shallowly cut or subentire.   (7)
7 (6) Style ca. 0.5 mm   3 Veronica alpina subsp. pumila
+ Style 6-7 mm.   (8)
8 (7) Stamens exserted; plants tomentose with white multicellular hairs, gray-green; infructescences slightly elongated   1 Veronica densiflora
+ Stamens included; plants densely with brown or brown-yellow multicellular glandular hairs; infructescences elongated to 5 cm   4 Veronica capitata
9 (2) Seeds convex on both sides; pedicel several X shorter than bract (as long as or slightly longer in V. pusilla).   (10)
+ Seeds navicular, convex on 1 side, with a deep groove on other; pedicel as long as or longer than bract, rarely slightly shorter.   (13)
10 (9) Leaves pinnatifid; stems simple or fastigiate; capsule pubescent   9 Veronica verna
+ Leaves entire, crenate, or dentate; stems branched; capsule glabrous or ciliate and otherwise glabrous.   (11)
11 (10) Pedicel 1.5-2 X longer than calyx   11 Veronica pusilla
+ Pedicel shorter than calyx.   (12)
12 (11) Stems glabrous or short glandular pubescent; leaves oblanceolate to narrowly oblong, base cuneate, margin entire or dentate above middle; corolla usually white   8 Veronica peregrina
+ Stems densely villous along 2 lines; leaves ovate-orbicular, base rounded, margin conspicuously crenate; corolla blue   10 Veronica arvensis
13 (9) Plants diffuse; bracts similar in size to leaves.   (14)
+ Plants erect, simple or branched below middle; bracts smaller than leaves.   (15)
14 (13) Pedicel slightly shorter than bract; capsule 4-6 mm wide, with inconspicuous reticulate veins, apically notched nearly at an acute angle, lobes rounded; style ca. 1.5 mm, as long as notch or slightly exserted   12 Veronica polita
+ Pedicel longer than bract; capsule 6-9 mm wide, with conspicuous reticulate veins, apically notched at a right to an obtuse angle, lobes obtuse; style 2-3 mm, conspicuously exserted   13 Veronica persica
15 (13) Leaves 4, under inflorescences, nearly whorled.   (16)
+ Leaves more than 4, pairs distant.   (17)
16 (15) Pedicel slender, 2-5 X longer than bract; calyx equally parted to base, lobes lanceolate   18 Veronica tenuissima
+ Pedicel slightly longer than bract; calyx unequally parted, laterally parted to middle, lobes rhomboid-ovate   19 Veronica cardiocarpa
17 (15) Seeds smooth; calyx lobes obscurely 3-veined or only midrib visible, red ciliate; plants often reddish   14 Veronica rubrifolia
+ Seeds transversely rugose; calyx lobes conspicuously 3-veined, white ciliate; plants green.   (18)
18 (17) Calyx lobes in fruit usually to 1.6 mm wide; seeds usually 8-14 per capsule, 1.3-1.7 X 0.7-0.8 mm, deeply transversely rugose (with 5 folds); pedicel in fruit usually distinctly recurved, usually longer than bract   17 Veronica campylopoda
+ Calyx lobes in fruit usually 2-3.5 mm wide; seeds usually 4-8 per capsule, 1.7-2.5 X 0.9-1.5 mm, obscurely reticulate-rugose; pedicel in fruit usually patent, often shorter than bract.   (19)
19 (18) Pedicel slightly recurved in fruit, pedicel and calyx densely glandular pubescent with hairs 0.4-0.7(-1) mm; corolla 4-6 mm in diam.; capsule (4-)5-8 mm wide, notch angle 30-45(-60); style (0.8-)1-1.3(-1.7) mm   16 Veronica arguteserrata
+ Pedicel straight in fruit, pedicel and calyx short puberulent with hairs 0.1-0.3 mm; corolla 3-4 mm in diam.; capsule (3-)3.5-5(-6) mm wide, notch angle acute (narrow); style 0.4-0.8 mm   15 Veronica biloba
20 (1) Plants aquatic or helophytic; stems succulent.   (21)
+ Plants terrestrial; stems not succulent.   (25)
21 (20) Stems creeping; capsule wider than long   53 Veronica beccabunga subsp. muscosa
+ Stems erect; capsule as wide as or narrower than long.   (22)
22 (21) Style ca. 3 mm; capsule basally wider than apically, apically notched   49 Veronica oxycarpa
+ Style less than 2.5 mm; capsule basally and apically equal in width, apically rounded or emarginate.   (23)
23 (22) Capsule ellipsoid, length exceeding width, apically emarginate; calyx lobes appressed to capsule, outside densely glandular hairy   52 Veronica anagalloides
+ Capsule subglobose, length equal to width, apically rounded; calyx lobes not closely appressed to capsule in fruit, glabrous or sparsely glandular hairy.   (24)
24 (23) Pedicel curved, at an acute angle with inflorescence axis; racemes less than 1 cm wide, usually glabrous   50 Veronica anagallis-aquatica
+ Pedicel straight, at a right angle with inflorescence axis; racemes 1-1.5 cm wide, sparsely glandular pubescent   51 Veronica undulata
25 (20) Rhizomes less than 1 cm, nodes and internodes inconspicuous; calyx 5-lobed, upper lobe much smaller (sometimes 4-lobed in V. rockii and V. filipes); corolla tube conspicuous; capsule slightly compressed (strongly compressed in V. filipes), length exceeding width.   (26)
+ Rhizomes more than 1 cm, with obvious nodes and internodes; calyx 4-lobed (5-lobed in V. teucrium); corolla tube inconspicous; capsule usually strongly compressed (not strongly in V. teucrium and V. morrisonicola), width exceeding length (length exceeding width in V. teucrium and V. morrisonicola.   (31)
26 (25) Capsule oblong to ovoid-globose, strongly compressed; corolla tube inside with a ring of villous hairs; stamens longer than or as long as corolla   25 Veronica filipes
+ Capsule ovoid to ovoid-conical, slightly compressed; corolla tube inside often glabrous (villous in V. himalensis); stamens shorter than corolla.   (27)
27 (26) Corolla to 10 mm, outside with multicellular glandular hairs; capsule more than 5 mm wide; leaves amplexicaul, margin conspicuously dentate   24 Veronica himalensis
+ Corolla 0.3-0.6 mm, outside glabrous; capsule less than 5 mm wide; leaves never amplexicaul, margin not conspicuously dentate.   (28)
28 (27) Ovary and capsule glabrous or sparsely pubescent   22 Veronica rockii
+ Ovary and capsule hirsute with multicellular hairs.   (29)
29 (28) Inflorescences long and lax, not capitate at anthesis   20 Veronica eriogyne
+ Inflorescences usually capitate at anthesis (elongated in V. ciliata subsp. zhongdianensis).   (30)
30 (29) Capsule ovoid, 4-5 mm wide; calyx lobes narrowly elliptic to oblanceolate; plants less than 10-flowered; leaves densely villous   23 Veronica chinoalpina
+ Capsule narrowly ovoid to narrowly ovoid-conical, less than 4 mm wide; calyx lobes linear-lanceolate; plants usually more than 10-flowered; leaves glabrescent   21 Veronica ciliata
31 (25) Stems densely white villous; lower leaves withered at anthesis   48 Veronica tibetica
+ Above combination of characters absent.   (32)
32 (31) Inflorescences less than 3 cm, 1-5-flowered.   (33)
+ Inflorescences usually more than 3 cm, lax, more than 5-flowered.   (34)
33 (32) Flowers 2-5, in corymbs; capsule obcordate-deltoid   30 Veronica szechuanica
+ Flowers 1-3, in fascicles; capsule reniform   31 Veronica chayuensis
34 (32) Capsule obcordate to obovoid-cordate.   (35)
+ Capsule mostly triangular or pliciform-rhomboid, rarely obcordate-reniform or reniform.   (39)
35 (34) Calyx 5-lobed; pedicel as long as or longer than bract; capsule glabrous   26 Veronica teucrium subsp. altaica
+ Calyx 4-lobed; pedicel shorter or rarely longer than bract; capsule hairy or ciliate.   (36)
36 (35) Stems creeping; capsule longer than wide   32 Veronica morrisonicola
+ Stems erect or ascending; capsule as long as or shorter than wide.   (37)
37 (36) Stems branched at base; style 0.3-0.5 mm; inflorescences usually less than 5 cm, rarely to 10 cm   29 Veronica javanica
+ Stems simple; style 3-6 mm; inflorescences to 30 cm.   (38)
38 (37) Stems densely villous with multicellular hairs along 2 lines; pedicel longer than or as long as bract; style 5-6 mm   27 Veronica chamaedrys
+ Stems evenly villous with multicellular hairs; pedicel much shorter than bract; style 3-4 mm   28 Veronica laxa
39 (34) Capsule obcordate-reniform or reniform, base truncate-rounded.   (40)
+ Capsule pliciform-rhomboid or deltoid-flabellate, base truncate or cuneate-truncate.   (42)
40 (39) Stems creeping and rooting at least below middle; bracts and calyces glabrous; style 5-6 mm   34 Veronica laxissima
+ Stems ascending; bracts and calyces ciliate; style less than 5 mm.   (41)
41 (40) Inflorescences several flowered; leaf base truncate   33 Veronica sutchuenensis
+ Inflorescences 1- or 2-flowered; leaf base shallowly cordate   35 Veronica vandellioides
42 (39) Stems creeping; leaf blade less than 2 cm.   (43)
+ Stems erect or ascending (sometimes creeping basally or below middle in V. tsinglingensis. V. riae, and V. henryi); leaf blade mostly more than 2 cm (less than 2 cm in V. taiwanica).   (44)
43 (42) Lower leaves petiole ca. 7 mm; leaf blade triangular-orbicular, margin with 2 or 3 broadly crenate lobes on each side, base truncate; inflorescences 1-4-flowered   47 Veronica oligosperma
+ Lower leaves subsessile; leaf blade elliptic, margin serrate, base cuneate; inflorescences ca. 10-flowered   46 Veronica forrestii
44 (42) Stems 1-5 cm; leaves often rosulate; inflorescences much longer than stems   45 Veronica piroliformis
+ Stems (8-)10-30 cm; leaves never rosulate; inflorescences shorter than stems.   (45)
45 (44) Leaves sessile, amplexicaul   44 Veronica deltigera
+ Leaves conspicuously petiolate.   (46)
46 (45) Stamens longer than or as long as corolla; style more than 5.5 mm; pedicel more than 5 mm.   (47)
+ Stamens shorter than corolla; style less than 4 mm (6-7 mm in V. longipetiolata); pedicel less than 3 mm (5-7 mm in V. riae).   (48)
47 (46) Longest petioles 1.5-3 cm; leaves papery, glabrous; capsule almost entirely glabrous   36 Veronica fargesii
+ Longest petioles 0.5-0.7 cm; leaves membranous, adaxially sparsely puberulent; capsule apically sparsely glandular ciliate   37 Veronica tsinglingensis
48 (46) Style 6-7 mm; petiole 3-4 cm on lower leaves   38 Veronica longipetiolata
+ Style less than 4 mm; petiole 0.3-1.5 cm.   (49)
49 (48) Leaves membranous; inflorescences to 11 cm; peduncle more than 3 cm; pedicel to 5 mm at anthesis, to 7 mm in fruit; style 3-4 mm   40 Veronica riae
+ Leaves papery; inflorescences less than 8 cm; peduncle less than 2 cm; pedicel less than 3 mm; style less than 3 mm.   (50)
50 (49) Calyx lobes narrowly elliptic to oblanceolate, sparsely glandular ciliate, apex obtuse; inflorescences axis and pedicels glandular hairy   43 Veronica yunnanensis
+ Calyx lobes linear-lanceolate, glabrous or sparsely ciliate, apex acute or acuminate; inflorescences axis and pedicels villous.   (51)
51 (50) Inflorescences 1.5-2 cm; corolla 4-5 mm in diam.; capsule 7-8 mm wide; style 1.5-1.8 mm; leaf blade 1-1.5 X 0.8-1.2 cm   42 Veronica taiwanica
+ Inflorescences 3-7 cm; corolla 10-12 mm in diam.; capsule 9-11 mm wide; style 2-3 mm; leaf blade 2-5 X 1.2-3 cm.   (52)
52 (51) Stems basally densely pubescent; lower leaves deltoid-orbicular to shallowly cordate, upper leaves ovate to ovate-orbicular, apex often obtuse; petiole 1-1.5 cm   39 Veronica cana
+ Stems basally subglabrous; leaves ovate to narrowly ovate, apex often acute; petiole less than 1 cm   41 Veronica henryi

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