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millepertuis, St. Andrew's cross, St. John's-wort

Habit Herbs, annual or perennial, subshrubs, or shrubs [trees], sometimes rhizomatous, glabrous or hairy, with glandular canals, lacunae, or dots containing resins or waxes (amber), essential oils (pale, translucent), and/or, sometimes, hypericin and pseudohypericin (black or red) in various parts. Herbs (perennial), subshrubs, or shrubs; black glands absent.
Stems

internodes terete (not lined) or 2-, 4-, or 6-lined at first (lines usually raised), then sometimes becoming angled, terete, or winged;

bark smooth or striate, sometimes corky, punctiform.

Leaves

deciduous (base articulated) or persistent (base not articulated).

Inflorescences

terminal, cymose, 2+-flowered, or flowers solitary, branching stellate [cupulate];

sepals persistent or deciduous, (3–)4–5, distinct or ± connate, margins sometimes glandular-ciliate;

petals persistent or deciduous, (3–)4–5[–6], contorted, yellow to orange, sometimes red-tinged;

stamens persistent or deciduous, (5–)10–300(–650), in continuous or interrupted ring or in (3–)4–5 fascicles, fascicles distinct or connate, each with 1–60+ stamens;

filaments distinct or basally connate;

anthers yellow to orange, oblong to ellipsoid, almost isodiametric, sometimes with amber or black gland on connective;

staminode fascicles 0 [3];

ovary 2–5-merous;

placentation axile to parietal;

ovules 2+ on each placenta;

styles distinct or ± connate basally, spreading to ± appressed.

Flowers

7–45 mm diam.;

sepals deciduous or persistent, (2–)4–5;

petals deciduous, (3–)4–5;

stamens usually persistent, sometimes deciduous, 30–650, in continuous ring or in 4–5 barely discernable fascicles, each of 1–2 stamens;

ovary 2–5-merous;

placentation incompletely axile to parietal;

styles ± appressed, bases distinct;

stigmas minute.

Capsules

2–5-valved, sometimes with glandular vittae or vesicles.

Seeds

narrowly cylindric to ellipsoid, sometimes carinate;

testa foveolate or reticulate to scalariform [papillose].

± carinate or not.

x

= 12, 9–7, 6 (dihaploid).

Hypericum

Hypericum sect. Myriandra

Distribution
from USDA
Nearly worldwide
[BONAP county map]
e North America; e Mexico; West Indies (Bahamas, Greater Antilles); Bermuda; Central America (Belize)
Discussion

Species ca. 490 (54 in the flora).

Shrubs with deciduous leaves, petals, and stamens belong to either Hypericum sect. Ascyreia Choisy (with five stamen fascicles and five styles) or sect. Androsaemum (Duhamel) Godron (with five stamen fascicles and three or four styles). These are all introductions, mostly garden escapes. Those in sect. Ascyreia include: Hypericum calycinum Linnaeus, a low shrub with creeping stolons and flowers 50–95 mm diam. that has been found in California, Oregon, and Washington; H. ×moserianum Luquet ex André, its hybrid with H. patulum Thunberg, a low (sterile?) branching shrub with red anthers; and H. hookerianum Wight & Arnott, a shrub to 2 m tall with narrow leaves and a dense ring of relatively short stamens, recorded from California (its identity requires confirmation). In sect. Androsaemum, H. androsaemum is a deciduous shrub with relatively small flowers and baccate fruits that ripen from cherry-red to black; it has been found in British Columbia and in California and Washington.

Introduced herbaceous species with three stamen fascicles and three styles include: Hypericum hirsutum, with hairy stems and leaves (Ontario); H. tetrapterum, with four-winged internodes and lanceolate sepals (British Columbia and Washington); H. pulchrum with cordate leaves and red-tinged petals (Newfoundland, St. Pierre and Miquelon); and H. humifusum Linnaeus, a procumbent herb with unequal sepals (British Columbia).

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Species 29 (28 in the flora).

Hypericum limosum Grisebach is endemic to western Cuba.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Herbs, subshrubs, or shrubs, black glands absent; stamens in continuous or interrupted ring or in 4 or 5 barely discernable fascicles, each of 1 or 2 stamens
→ 2
1. Herbs or shrubs, black and/or red glands usually present throughout, sometimes absent; stamens in 5 fascicles, each of 2+ stamens
→ 3
2. Herbs (perennial), subshrubs, or shrubs; petals deciduous; stamens usually persistent, sometimes deciduous, 30–650, in continuous ring or in 4–5 barely discernable fascicles; styles ± appressed, bases distinct.
sect. Myriandra
2. Herbs (annual or perennial); petals persistent; stamens persistent, (5–)10–80, usually in continuous or interrupted ring, sometimes in 5 barely discernable fascicles; styles ± spreading, bases distinct.
Hypericum informal sect. group Brathys
3. Shrubs; leaves deciduous (base articulated); style bases distinct.
sect. Webbia
3. Herbs (perennial); leaves persistent or tardily deciduous (base not articulated) or; style bases ± connate or distinct
→ 4
4. Herbs, black glands absent; flowers 40–70 mm diam.; stamens 150, in 5 fascicles, fascicles usually distinct, rarely 1 pair connate; styles ± appressed, bases ± connate or distinct.
sect. Roscyna
4. Herbs, black glands usually on leaves, sepals, and petals and, sometimes, on stems and anthers; flowers 6–35 mm diam.; stamens 20–109, in 5 fascicles, fascicles connate (as 2 + 2 + 1); styles spreading, bases distinct.
Hypericum informal sect. group Hypericum
1. Sepals enclosing capsule, (2–)4, unequal (outer pair larger) [subsect. Ascyrum]
→ 2
1. Sepals not enclosing capsule, usually 5, if 3 or 4, usually subequal
→ 6
2. Stamens 30–50; ovaries 2-merous
→ 3
2. Stamens 70–100; ovaries 3(4)-merous
→ 4
3. Leaf bases without glandlike auricles; inflorescences 1-flowered, pedicels mostly recurved to reflexed in fruit, bracteoles proximal.
H. suffruticosum
3. Leaf bases with glandlike auricles; inflorescences 1–12-flowered, pedicels erect in fruit, bracteoles distal.
H. hypericoides
4. Leaf bases cordate-amplexicaul; inflorescences terminal, branching (at apical node) pseudodichotomous
H. tetrapetalum
4. Leaf bases usually cuneate to subrounded, rarely rounded to slightly subcordate-amplexicaul; inflorescences from 1–4 nodes, branching dichasial or pseudodichotomous
→ 5
5. Leaves: bases without glandlike auricles, margins plane to subrecurved; inflorescences 1–3(–7)-flowered, branching dichasial; outer sepals: apices apiculate or obtuse to rounded
H. crux-andreae
5. Leaves: bases with glandlike auricles, margins subrecurved to subincrassate; inflorescences 1-flowered, branching pseudodichotomous; outer sepals: apices acute to subacuminate
H. edisonianum
6. Leaves: bases usually not articulated, if articulated (22. H. myrtifolium), subcordate-amplexicaul; stamens (all or some) usually persistent (deciduous in H. apocynifolium, H. myrtifolium, and H. dolabriforme)
→ 7
6. Leaves: base articulated, not amplexicaul; stamens deciduous [subsect. Centrosperma]
→ 16
7. Shrubs, decumbent, mat-forming, wiry; inflorescences 1(–5)-flowered [subsect. Pseudobrathydium].
H. buckleyi
7. Herbs, subshrubs, or shrubs, erect to ascending or decumbent, not mat-forming, not wiry; inflorescences (1–)3–70-flowered
→ 8
8. Inflorescences widely branched; stamens 120–200 [subsect. Brathydium]
→ 9
8. Inflorescences narrowly branched; stamens 30–95 [subsect. Suturosperma]
→ 10
9. Shrubs, erect, unbranched or branched distally, bark corky; leaf blades oblong-ovate to triangular-lanceolate, (5–)7–20 mm wide, bases articulated, subcordate-amplexicaul.
H. myrtifolium
9. Subshrubs, decumbent, ± branching, bark not corky (thin); leaf blades linear-elliptic or linear-oblong to linear, 3–5 mm wide, bases not articulated, narrowly cuneate to rounded.
H. dolabriforme
10. Leaf blades (7–)10–25 mm wide; sepals deciduous or tardily deciduous; placentation incompletely axile
→ 11
10. Leaf blades 1–10(–15) mm wide; sepals persistent; placentation parietal
→ 12
11. Inflorescences terminal, (1–)3–5(–8)-flowered; sepals tardily deciduous, 1.5–2.3 mm wide; capsules 6–15 × 4.5–8 mm.
H. apocynifolium
11. Inflorescences from 1–3 nodes, the terminal 7–45-flowered; sepals deciduous, 1–1.5 mm wide; capsules 3.5–7 × 3–5 mm.
H. nudiflorum
12. Shrubs; leaf blades 1–3 mm wide; inflorescences 1–3-flowered; sepals (3–)4(–5).
H. microsepalum
12. Herbs, subshrubs, or shrubs; leaf blades (1–)3–15 mm wide; inflorescences usually 7–70-flowered, if flowers 1–6, herbs; sepals usually 5
→ 13
13. Shrubs; capsules ovoid-cylindric to broadly ovoid.
H. cistifolium
13. Herbs or subshrubs; capsules usually broadly ovoid to depressed-globose, if ovoid-ellipsoid to ellipsoid or globose, rhizomatous herbs
→ 14
14. Herbs or subshrubs, not or rarely rhizomatous; capsules broadly ovoid to depressed-globose; seeds 2–2.7 mm.
H. sphaerocarpum
14. Herbs, rhizomatous; capsules ellipsoid to ovoid-ellipsoid or globose; seeds 0.6–0.7 mm
→ 15
15. Herbs, 2–8 dm, relatively stout; leaf blades narrowly oblong or linear to lanceolate or narrowly elliptic.
H. adpressum
15. Herbs, 1–3(–5) dm, relatively slender; leaf blades elliptic or oblanceolate to oblong-elliptic.
H. ellipticum
16. Leaf blades usually elliptic, lanceolate, linear, oblanceolate, or oblong, (1–)3–22 mm wide; sepals usually elliptic, linear, oblanceolate, oblong, obovate, ovate, or spatulate
→ 17
17. Inflorescences usually 1–3(–7+)-flowered from apical node, sometimes with single flowers at 1(–2) proximal nodes
→ 18
17. Inflorescences usually (1–)3–25-flowered from apical node, sometimes with triads, dichasia, or flowering branches from 1–4 proximal nodes
→ 19
18. Shrubs (6–)10–30 dm; leaf blades 8–22 mm wide, margins plane or subrecurved; sepals 4–10 mm wide; ovaries 3-merous.
H. frondosum
18. Shrubs (1.4–)2–6(–10) dm; leaf blades 3–7(–10) mm wide, margins subrecurved to revolute; sepals 1.5–5 mm wide; ovaries (3–)5(–6)-merous.
H. kalmianum
19. Stem internodes 4- or 6-lined at first; inflorescences cylindric or broadly to narrowly elongate-cylindric with 1–5(–7)-flowered lateral dichasia proximally; internodes 4-lined at first and capsules 7–13 mm or internodes 6-lined at first and capsules 4.5–6 mm
→ 20
19. Stem internodes 4-lined at first; inflorescences shortly and broadly pyramidal or broadly cylindric to globose-cylindric, or obpyramidal with (2–)5–15-flowered lateral dichasia proximally; capsules 5–7 mm
→ 21
20. Leaf blades 30–70 mm, narrowly oblong to narrowly elliptic-oblanceolate; flowers 15–30 mm diam.; placentation incompletely axile; capsules 7–13 mm; seeds 1.5–2 mm.
H. prolificum
20. Leaf blades 15–32(–37) mm, narrowly oblong-elliptic or oblanceolate to linear; flowers 9–14 mm diam.; placentation parietal; capsules 4.5–6 mm; seeds 0.7–0.8 mm.
H. galioides
21. Sepals: basal veins 3–7; ovaries (3–)4–5-merous; capsules notably lobed.
H. lobocarpum
21. Sepals: basal veins 1–3; ovaries 3–4(–5)-merous; capsules not or scarcely lobed.
H. densiflorum
22. Shrubs, erect, 3–45 dm
→ 24
23. Leaf blades 13–25 mm; sepals (3–)4.5–7 mm.
H. lloydii
23. Leaf blades 4–11 mm; sepals 2–4 mm.
H. tenuifolium
24. Capsules usually cylindric, rarely narrowly conic or ovoid-conic; bark smooth, not metallic-silvery
→ 25
24. Capsules ± narrowly ovoid-conic or pyramidal-ovoid to ovoid or ellipsoid; bark smooth and metallic-silvery or corky to spongy
→ 26
25. Leaves 9–26 mm; styles shorter than ovaries; capsules (4.5–)5–7 mm.
H. nitidum
25. Leaves 6–12 mm; styles longer than ovaries; capsules 3.5–5 mm.
H. brachyphyllum
26. Bark smooth and metallic-silvery, exfoliating in thin, curled plates; styles 5 mm; seeds 1–1.6 mm; young stems, leaves, and sepals glaucous.
H. lissophloeus
26. Bark corky to spongy, exfoliating in thin, papery sheets or plates; styles 2.5–4 mm; seeds 0.4–0.8 mm; young stems, leaves, and sepals not glaucous
→ 27
27. Stems: internodes 6-lined at first, soon 2-winged, then terete; bark smooth; terminal inflorescence (3–)7–32-flowered.
H. fasciculatum
27. Stems: internodes 4-lined at first, soon 4-angled, then terete; bark striate; terminal inflorescence 1–3-flowered.
H. chapmanii
Source FNA vol. 6, p. 72. Treatment author: Norman K. B. Robson. FNA vol. 6, p. 74.
Parent taxa Hypericaceae Hypericaceae > Hypericum
Subordinate taxa
H. sect. Myriandra, H. sect. Roscyna, H. sect. Webbia
H. adpressum, H. apocynifolium, H. brachyphyllum, H. buckleyi, H. chapmanii, H. cistifolium, H. crux-andreae, H. densiflorum, H. dolabriforme, H. edisonianum, H. ellipticum, H. fasciculatum, H. frondosum, H. galioides, H. hypericoides, H. kalmianum, H. lissophloeus, H. lloydii, H. lobocarpum, H. microsepalum, H. myrtifolium, H. nitidum, H. nudiflorum, H. prolificum, H. sphaerocarpum, H. suffruticosum, H. tenuifolium, H. tetrapetalum
Synonyms Ascyrum, Crookea, Sanidophyllum, Sarothra section Myriandra
Name authority Linnaeus: Sp. Pl. 2: 783. (1753): Gen. Pl. ed. 5, 341. (1754) (Spach) R. Keller: in H. G. A. Engler and K. Prantl, Nat. Pflanzenfam. 95[III,6]: 214. (1893)
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