Hypericum

Family 53. GUTTIFERAE

N.K.B. ROBSON

Herbs, shrubs or trees. Leaves opposite, simple, often glandular with translucent dots and streaks and sometimes with black gland-dots. Stipules absent. Flowers bisexual, actinomorphic, solitary or in terminal cymes or panicles. Sepals 5, free, imbricate,

often glandular. Petals 5, free, contorted in bud. Stamens numerous in 3 or 5 bundles.

Ovary superior, 1-5-celled; ovules numerous; styles 3 or 5, free or fused. Fruit a many­

seeded, 3- or 5-valved capsule. Robson, N.K.B. (1985, 1993). Studies in the genus Hypericum L. Bull. Brit. Mus.

(Nat. Hist.) Bot. 12: 163-325 (1985); 23 (2): 67-70 (1993).

HYPERICUM L.

Description as for the family.

1. Stems and leaves pubescent or puberulous

+ Stems and leaves glabrous2

4

2. Low shrub; leaves glabrous above; black glands absent 6. H. fieriense

+ Erect to prostrate perennial herbs; leaves with hairs on both sides; blackglands present 3

331

GUTTIFERAE

3. Stems erect, not rooting; leaves triangular-lanceolate to narrowly oblong;flowers c.25mm diam. 10. H. collenettiae

+ Stems ascending to prostrate, sometimes rooting; leaves narrowly oblong toelliptic or oblanceolate; flowers 9-13mm diam. 11. H. sinaicum

4. Erect perennial herbs 5+ Small trees or shrubs 6

5. Sepals and bracts fringed with black, glandular cilia; bracts auriculate9. H. annulatum

+ Sepals and bracts without black glands; bracts not auriculate12. H. perforatum

6. Styles 3, free, outcurving or spreading 7+ Styles 5, wholly or partly coherent 9

7. Leaves 25-75mm long, lanceolate to ovate, with densely reticulate venationbetween the main lateral veins; stamen bundles 5, deciduous 5. H. hircinum

+ Leaves 8-25mm long, oblong-elliptic to obovate, with obscure venationbetween the main lateral veins; stamen bundles 3, persistent 8

8. Flowers 1(2), terminal and in the axils ofolder leaves; sepals ensiform; capsulevalves vittate; habit erect 8. H. scopulorum

+ Flowers 7-13 in terminal subumbellate inflorescence; sepals elliptic or oblong-oblanceolate to oblong; capsule valves verrucose; habit spreading or straggling 7. H. tortuosum

9. Leaves with black submarginal gland-dots; styles completely coherent4. H. quartinianum

+ Leaves with pale (pellucid) submarginal gland-dots; styles partly coherentbelow 10

10. Sepals with gland-dots; flowers solitary; main lateral veins joining to forman uninterrupted submarginal vein 1. H. revolutum

+ Sepals without gland-dots; flowers in l-5(-10)-flowered terminal cymes; mainlateral veins not joining to form an uninterrupted submarginal vein 11

11. Leaves narrowly elliptic or narrowly lanceolate, acute; leaf venation parallel(with 3-7 main veins) towards the base, becoming pinnate along the midribtowards the tip 2. H. balfourii

+ Leaves elliptic to obovate or subcircular, acute to rounded; leaf venationmainly pinnate with 1 or 2 pairs of basal lateral veins extending around themargin towards the tip 3. H. socotranum

1. H. revolutum Vahl (1790 p.66). Syn.: H. lanceolatum auctt. non Lam. (1797). Illustr.:Webbia 22: 239 (1967); Collenette (1985 p.261). Type: Yemen (N), Forsskal (C).

Glabrous shrub or small tree up to 3m. Leaves sessile, lanceolate to narrowlyoblong-elliptic, 8-15 x2.5-4mm, acute, reflexed-auriculate at the base, black glandsabsent; main lateral veins joining to form an uninterrupted submarginal vein. Flowers

332

Hypericum

3.5-5.5cm across, solitary. Sepals ovate to broadly ovate, 7-9x4-8mm, acuteto obtuse, with marginal black gland-dots. Petals bright yellow to yellow-orange(15-)20-30x5-25mm, persistent. Stamen bundles 5. Styles 5, 6-10mm, unitedbelow. Capsule 5-valved, ovoid to subglobose, 9-12 x 8-1 Omm. Map 446, Fig. 63.

Montane shrubland and grassland; (2200-)2500-3500m; flowering in early summer.

Saudi Arabia, Yemen (N). Tropical and South AfricaH. revolutum comprises two subspecies which differ mainly in leaf venation and

style length. Subsp. keniense (Schweinf.) N. Robson is restricted to the mountains ofE Africa whereas subsp. revolutum is more widespread, occurring in SW Arabia andthroughout the mountains of tropical and southern Africa.

2. H. balfourii N. Robson, op. cit.: 191 (1985). Syn.: H. mysorense sensu Balf. f. (1888)non Heyne. Illustr.: Webbia 22: 263 (1967) as H. mysorense. Type: Socotra, Balfour,Cockburn & Scott 606 (A, BM, E, K).

Glabrous shrub or small tree up to 3m. Leaves sessile, narrowly elliptic to narrowlylanceolate, 10-45 x 4—12mm, acute, cuneate to subauriculate at the base, black glandsabsent; venation parallel (with 3-7 main veins) towards the base, becoming pinnatealong the midrib towards the tip. Flowers 4-7cm across, in l-3(-10)-flowered terminal cymes. Sepals ovate to oblong, 5-9 x 3-5.5mm, acute or obtuse, black glandsabsent. Petals golden yellow, 20-35 x 10-20mm, deciduous. Stamen bundles 5.Styles 5, 5-8mm, united below. Capsule 5-valved, ovoid, 13-15 x 7-9mm. Map 447,Fig. 63.

Low montane shrubland and dwarf Hypericum-Crotonopsis shrubland; (120-)600-1100m.

Socotra. Endemic.

Closely related to H. mysorense, a species from southern India and Sri Lanka.Normally found above about 600m, the lower altitudes are atypical.

3. H. socotranum Good in J. Bot. 65: 334 (1927).Glabrous shrub or small tree up to 4.5m. Leaves sessile, elliptic to obovate or

subcircular, 12-30 x 5-22mm, acute to rounded, cuneate at the base, black glandsabsent; venation mainly pinnate with 1-2 pairs of basal lateral veins extending aroundthe margin towards the tip. Flowers (2.5-)4-7.5cm across, in 1-5-flowered terminalcymes. Sepals ovate to oblong, 4.5-6.5 x 2.5-3mm, apiculate or obtuse to rounded,black glands absent. Petals golden yellow, 15-40x9-24mm, deciduous. Stamenbundles 5. Styles 5, 5-8mm, united below. Capsule 5-valved, ovoid, 12-13 x 6-7mm.Map 448, Fig. 63.

Socotra. Endemic.

333

GUTTIFERAE

Fig. 63. Hypericaceae. A, Hypericum revolutum: Aa flowering branch (x l-); Ab, sepal (x 3); Ac, leaf (x 1^);Ad, capsule (x 1±). B, H. balfourii: Ba, leaf (x 1±); Bb sepal (x 3). C,H. socotranum subsp. socotranum: Ca,leaf (x V-). D, H. socotranum subsp. smithii: Da, leaf (x I1-). E, H. quartinianum: Ea, leaf (x 1±); Eb, sepal(x 3); Ec, capsule (x 3). F, H. hircinum: Fa, leaf(x \1-); Fb, sepal (x 3); Fc, capsule (x 3) G, H.fierense:Ga,flowering branch (x ±); Gb,sepal (x 3); Gc, leaf (x U); Gd,capsule (x 3).

Hypericum

1. Leaves elliptic to oblanceolate, 5-1 Omm wide, acute to obtuse; pedicels 1-6mm; flowers solitary subsp. socotranum

+ Leaves broadly elliptic to subcircular, 8-22mm wide, apiculate to rounded;pedicels 5-25mm; flowers 1-5 subsp. smithii

subsp. socotranum. Syn.: H. lanceolatum sensu Balf. f. (1888) non Lam. Illustr.:Webbia 22: 259 (1967); Robson, op. cit.: 193 (1985). Type: Socotra, Balfour, Cockburn& Scott 246 (BM, E, K, LE, P).

Limestone escarpments and rock crevices in the western part of the island on; 210m.

subsp. smithii N. Robson, op. cit.: 194 (1985). Illustr.: Robson, op. cit.: 194. Type:Socotra, Smith & Lavranos 272 (BM, K).

Limestone escarpments and rock crevices in the central and eastern part of the island;(120-)400-650m.

4. H. quartinianum A. Rich. (1847 p.97). Illustr.: Robson, op. cit.: 195 (1985); Webbia22: 255 (1967).

Glabrous shrub up to lm. Leaves sessile, lanceolate to oblong-elliptic, 20-100 x 5-35mm, acute, cuneate to subcordate at the base, with marginal black gland-dots;venation pinnate with only 2-3 pairs of lateral veins towards the base. Flowers 3.5-7cm across, in few-flowered terminal cymes. Sepals ovate to lanceolate, 5-15x2-5mm, acute, with submarginal black gland-dots. Petals bright yellow, 20-40 x 10-25mm, tardily deciduous. Stamen bundles 5. Styles 5, 8-13mm, completely united.Capsule 5-valved, ovoid, 10-16 x 5-1 Omm. Map 449, Fig. 63.

Humid gullies; 2000m.

Yemen (N). Eastern tropical Africa from Ethiopia and Somalia south to Zambia andMo9ambique.

A rare plant of the outer escarpment mountains in Yemen.

5. H. hircinum L., Sp. pi.: 784 (1753). Illustr.: Robson, op. cit.: 308 (1985); Fl. Palaest.1: t.324 (1966); Collenette (1985 p.261).

Glabrous shrub up to 1.5m. Leaves sessile, lanceolate to ovate, 25-60(-75)x 10-25(-35)mm, acute or obtuse, rounded or cordate at the base, black glands absent;venation pinnate with a prominent reticulum between the main lateral veins. Flowers3-4cm across, in few-flowered terminal cymes. Sepals lanceolate, 4-9x1.5-3mm,acute, black glands absent. Petals bright yellow, (11—)15—20 x (4-)8-9mm, deciduous.Stamen bundles 5. Styles 3, (10-)15-25mm, free. Capsule 3-valved, ellipsoid, 8-14 x 4-7mm. Map 450, Fig. 63.

In damp sand by streams; 1800-2250m.

Saudi Arabia. Mediterranean region; widely cultivated and naturalized elsewhere.The Arabian plants are all referable to subsp. majus (Aiton) N. Robson.

335

GUTTIFERAE

6. H. fieriense N. Robson, loc. cit.: 68 (1993). Type: Socotra, Smith & Lavranos475 (K).Low shrub up to lm with stems densely to sparsely fawn-puberulous and leaves

densely whitish-puberulous beneath. Leaves with petiole 4-6mm, triangular-ovate tooblong-ovate, 10-17 x7-12mm, subobtuse to rounded, broadly cuneate to truncateat the base, black glands absent; venation pinnate, obscure between the main lateralveins. Flowers clem across, in subumbellate 3-5-flowered terminal cymes. Sepalslinear-lanceolate, 4-5 x c. 1.5mm, acute, black glands absent. Petals not seen. Stamenbundles 3. Styles 3, free, not seen complete. Capsule 3-valved, pyramidal-ovoid, 6-7.5 x 4.5-5mm. Map 451, Fig. 63.

Low Hypericum-Crotonopsis shrubland among trees of Dracaena cinnabari; 1350m.

Socotra. Endemic

7. H. tortuosum Balf. f., (1882 p.502). Illustr.: Balfour (1888 t. 4B); Webbia 22: 271(1967). Type: Socotra, Schweinfurth 757 (E)

Spreading or straggling glabrous shrub, up to 50cm. Leaves shortly petiolate,becoming sessile above, obovate to oblong-elliptic, 8-25 x 4-20mm, obtuse, cuneateat the base, sometimes auriculate, black glands absent; venation pinnate, obscurebetween the main lateral veins. Flowers 1-1.5cm across, in subumbellate, many-flowered, terminal cymes. Sepals elliptic or oblong-oblanceolate to oblong, 3.5-6 x 1-2mm, acute or truncate, black glands absent. Petals yellow, 5-7.5 x 2-2.5mm, persistent. Stamen bundles 3. Styles 3, free, 2.5-3mm. Capsule 3-valved, subglobose, c.4.5 x 4mm, the valves verrucose. Map 452, Fig. 64.

Cliffs on granite and limestone and in cushion vegetation on granite pinnacles; 600-1150m.

Socotra. Endemic.

8. H. scopulorum Balf. f. (1882 p.502). Illustr.: Balfour (1888 t. 4A); Webbia 22: 270(1967). Type: Socotra, Schweinfurth 756 (E).

Erect, glabrous shrub up to 120cm. Leaves sessile, obovate to oblong-elliptic. 10-24 x 4-11mm, obtuse, subauriculate at the base, black glands absent; venation pinnate,obscure between the main lateral veins. Flowers 1-1.5cm across, solitary in the axilsof the uppermost leaves. Sepals ensiform, 5-7.5x0.5-1.2mm, acute, black glandsabsent. Petals golden-yellow to golden-orange, 5-11 x 1.5-2mm, persistent. Stamenbundles 3. Styles 3, free, c. 1.5mm. Capsule 3-valved, subglobose, c. 5 x4mm, thevalves vittate. Map 453, Fig. 64.

Common in dwarf Hypericum-Crotonopsis shrubland on the higher slopes of thegranite mountains and less commonly on limestone cliffs; 750-1000m.

Socotra. Endemic.

9. H. annulatum Moris, Stirp. sard, elench. 1:9 (1827). Illustr.: Webbia 22: 275 (1967);Collenette(1985p.261).

Erect, glabrous, perennial herb up to 40cm. Leaves sessile, oblong-ovate to oblong-elliptic, 15-40 x 0.5-15mm, acute or obtuse, rounded at the base, with marginal black

336

Hypericum

H.TtSM

Fig. 64. Hypericaceae. A, Hypericum tortuosum: Aa, flowering branch; Ab, sepal; Ac, leaf; Ad, capsule. B,H. scopulorum: Ba, flowering branch; Bb, leaf; Be, sepal; Bd, capsule. C, H. annulatum: Ca, floweringbranch; Cb, bracts; Cc, sepal. D, H. perforatum: Da, flowering branch; Db, sepal. E, H. sinaicum: Ea,flowering branch; Eb, leaf; Ec, sepal. F, H. collenettiae: Fa, flowering branch; Fb, leaf; Fc, sepal. Floweringbranches all x^, sepals and capsules all x 3, leaves all x2.

GUTTIFERAE

gland-dots. Flowers 2-3cm across, in many-flowered pyramidal cymes. Bracts withdense black glandular-ciliate auricles. Sepals lanceolate, 3.5-4.5 x 0.8-1.5mm, acute,with black gland-dots and black glandular-ciliate margins. Petals yellow, c. 15x3-4mm, persistent. Stamen bundles 3. Styles 3, 6-7mm, free. Capsule 3-valved, ovoid-pyramidal, c. 4 x 2.5mm. Map 454, Fig. 64.

Shady banks, amongst grass and shrubs; 1050-1750m.

Saudi Arabia. External distribution of subsp. intermedium: Sudan and northernEthiopia.

A species with an interesting disjunct distribution between southern Europe andtropical E Africa with the Arabian population, which is somewhat intermediategeographically and morphologically, referable to subsp. intermedium (Steud. ex A.Rich.) N. Robson, op. cit.: 69 (1993).

10. H. collenettiae N. Robson, op. cit.: 69 (1993). Illustr.: Collenette (1985 p.262) asH. sp. aff. sinaicum. Type: Saudi Arabia, Collenette 3752 (BM, K).

Erect woody-based herb up to 20cm; stem and leaves white-puberulous. Leavessessile, markedly tetrastichous, narrowly oblong to triangular-lanceolate, 7.5-13 x 1.5-4mm, obtuse, rounded to subcordate at the base, with submarginal blackgland-dots only. Flowers c.2.5cm across, in (2—)5—11-flowered terminal cymes. Sepalsnarrowly lanceolate, 3.5-4 x 0.8-1.2mm, acute to subaristate, with black gland-dots,the margins with irregular black glandular cilia and prominent sessile black glands.Petals golden yellow, 9-10 x c.4.5mm, persistent. Stamen bundles 3. Styles 3, c.4mm,free. Capsule 3-valved, ovoid-cylindric, 4.5-5 x 3-4mm. Map 455, Fig. 64.

In shady rock crevices; 1800m.

Saudi Arabia. Endemic.

Known only from two localities in the mountains of the Asir. It is intermediate,both morphologically and geographically, between H. somaliense N. Robson (NSomalia) and H. sinaicum.

11. H. sinaicum Hochst. & Steud. ex Boiss., Fl. orient. 1: 808 (1867). Illustr.: Tackholm(1974U2B).

Ascending woody-based herb up to 15cm; stems and leaves white-tomentose. Leavessessile, ovate to elliptic, 5-17x3-8mm, obtuse to acute, subcordate at the base,with scattered black gland-dots. Flowers 1-1.5cm across, in few to several-floweredterminal cymes. Sepals ovate, 3-4.5 x 1-1.25mm, acute, with black gland-dots, themargins with stalked black glands. Petals yellow, 6-8 x 3-4mm, persistent. Stamenbundles 3. Styles 3, 4-6mm, free. Capsule 3-valved, ovoid, c. 5 x 3mm. Map 456, Fig.64.

In rock crevices by water; 1200-1500m.

Saudi Arabia. Sinai.

Outside the Sinai Peninsula known only from around J. Lawz in the extreme NWof Saudi Arabia.

338

Hypericum

12. H. perforatum L., Sp. pl.: 785 (1753).Erect, glabrous, perennial herb up to 50cm. Leaves sessile, ± linear to narrowly

oblong or narrowly lanceolate, 5-30 x 2-7mm, acute or obtuse, cuneate or± rounded at the base, with marginal black gland-dots. Flowers l-2cm across, in many-flowered pyramidal cymes. Bracts non-auriculate, with scattered black gland-dots. Sepals linear to lanceolate, 5-9 x l - l .25mm, entire or with a few non-glandular cilia on the margins. Petals yellow, c. 12 x 5mm, persistent. Stamen bundles 3. Styles 3, 4-5mm, free. Capsule 3-valved, ovoid to pyramidal, 5-8 x 3-4.5mm. Map 457, Fig. 64.

Banks bordering fields; 2070m.

Saudi Arabia. Europe, N Africa, SW and C Asia, Siberia and W China.

339

Map 446. Hypericum revolutum

Distribution Maps

Map 447. H. balfourii

545

Distribution Maps

Map 448. H. socotranum Map 449. H. quartinianum Map 450. H. hircinum

Map 451. H. fieriense Map 452. H. tortuosum Map 453. H. scopulorum

Map 454. H. annulatum Map 455. H. collenettiae Map 456. H. sinaicum

546

Map 457. H. perforatum

Distribution Maps

547