Genus Hypericum L.
Род 317. ЗВЪНИКА, ЖЪЛТ КАНТАРИОН, ПОРЕЗНИЧЕ —HYPERICUM L.²
L., Sp. Pl. ed. 1(1753)783; Gen. Pl. ed. 5(1754)341Fam: Guttiferae Juss.
Genus: Hypericum L.
English Name: St. John's wort, goatweedDescription:
Semi-shrubs or perennial herbaceous, bare or fibrous plants. Stems round, quadrangular, double-edged or winged. Leaves opposite, rarely in vertebrae, from linear to almost round, sat down or with short petioles, often with punctate, elliptical or linear, black, brown or transparent glands.² Inflorescences thyroid or racemose, rarely single flowers. Sepals 5, rarely 4, from almost round to wedge-shaped, entire or glandular, or simply ciliated. Petals 5, rarely 4, slightly or several times longer than the sepals, entire or in its upper half on the edge glandular ciliated, yellow to orange, on the surface or on the edge often with black or brown punctate, elliptical ate linear glands. Stamens numerous, fused at the base in 3. or 5 groups, rarely free; anthers yellow or reddish, often with a black gland. Carpel from 3 to 5 fully or semi-fused carpels; the upper ovary; the styles 3 - 5, free or fused at the base. The ovules are numerous. The fruit 3 - 5-nested ovoid to conical, cracked, smooth or on the surface with glandular bubbles or canal box, or not cracked with a fleshy shell - strawberry. Seeds 1 - 3 mm long, usually smooth on the surface, reticulate wrinkled or warty. Probably self-pollinating. Propagated by seeds and vegetatively.
Fossil remains of the genus have been found in the Sarmatian deposits near the village of Slavotin, Mihaylovgrad region - Hypericum cf. ponticum Lip.
Table for determination of the species
1 The leaves on both sides, as well as the stems, at least in their lower half densely bristly or short bristly fibrous .................. .. 2
1* The whole plant bare or at most the leaves on its underside along the veins scattered short bristly or fibrous ......................... 4
2 Sepals entire, without intramarginal glands ……..................................................…..... 7. - H. cerastioides (Spach) Robson
2* Sepals glandular ciliated or with 5 - 8 intramarginal glands ................................................................................................... 3
3 Inflorescences glabrous. The leaves with a heart-shaped stem-covering base ………...........……… 8. - H. degenii Vognm.
3* Inflorescences, as well as stems, bristly fibrous. The leaves are broadly wedge-shaped at the base ........... 3. - H. hirstitum L.
4 The sepals entire or shallowly unevenly toothed, without marginal glands or very rarely with several located at the apex ....…. 5
4* The sepals are glandular or glandular and simply serrated or ciliated, often with marginal glands .................………………. 11
5 The styles 5. Flowers 5.0 - 9.0 cm in diameter …………...............................................……...…… ... 1. - H, calycinum L.
5* The styles 3. Flowers 1.5 - 4.0 cm in diameter ……………………………...............................................……………… 6
6 Semi-shrub. Leaves without transparent glands. The fruits are not cracked - strawberries .............. . 2. - H. androsaemum L.
6* Herbaceous plant. The leaves with transparent glands. The fruits are cracked in 3 - 5 part boxes .......................................... 7
7 Sepals 5 - 10 (14) mm long, strongly unequal to each other, ovate or broadly ovate, paniculate. Stem leaves without shortened
twigs in their axils ..................................................................................................................................... 6, - H. olympicum L.
7* Sepals up to 5 (6) mm long, equal to each other, lance, ovate-lance, wedge-shaped, elliptical or ovate. Stem leaves in the axils
with shortened, often shorter, twigs .......................................................................................................................................... 8
8 Stems in cross section round or double-edged .......................................................................................................... ......... 9
8* Stems in their cross section quadrangular ...................................................................................................... ................... 10
9 Surface boxes with longitudinal resin grooves. Leaves without intramarginal glands. Inflorescences narrowly clustered-vertebrate …………..……....................................................................................................................……… 5. - H. linarioides Bosse
9* Boxes with short linear or renal glandular bubbles parallel to the vertical axis. Leaves with intramarginal glands. Inflorescences
semi-thyroid…. …………..........................................................................................................……… 21. - H. perforatum L.
10 Sepals broadly ovate, to broadly elliptical, obtuse or shortly pointed, 3 - 4 times shorter than the petals ............................... ……………………………......................................................................………………………. 20. - H. maculatum Crantz
10* The sepals are narrowly lance or linearly wedge-shaped, long pointed, 1.5 - 2 shish shorter than the petals ......................... …………………………….....................................................................……………………...… 19. - H. tetrapterum Fries
10* The sepals are narrowly lance or linearly wedge-shaped, long pointed, 1.5 - 2 shish shorter than the petals .......................... ………………………………………………......................................................................…..… 19. - H. tetrapterum Fries
11 (4*) The sepals over their entire surface with black punctate or linear glands or only with well-developed intramarginal glands ………………………..…………........................................................................................................…………………… 12
11* Sepals pure green or with reddish-brown or transparent superficial punctate or linear glands, rarely at most with 1 - 3 (4 - 8)
large black punctate or almost marginal or superior glands; their central part is always pure green ............................................. 16
12 The sepals on the edge are simply ciliate; lashes but longer than the width of the sepals, curly and whitish at the end ……..…. ……….........................................................................................................................................…... 15, - H. barbatum Jacq.
12* The sepals on the edge are glandular ciliated; cilia almost to the glands green and protruding, at most equal to the width of the
sepals .................................................................................................................................................................................... 43
13 Petals over their entire surface covered with black punctate or linear glands ….....................................................…....... 14
13* Petals only with intramarginal black dot glands ………….................................................……… 9, - H. montbretii Spach
14 Leaves on its lower surface mostly with intramarginal glands. Boxes with black hemispherical glandular bubbles or almost smooth ………………...……...................................................................................................................................................……. 15
14* Leaves on its lower surface with black or red and less often with transparent superficial and marginal glands. Boxes with
strongly protruding orange elliptical and linear glandular bubbles ………….……...…………............ 11. - H. umbellatum Kern.
15 The boxes are smooth or only on the edge of the carpel with small reddish or transparent glandular bubbles. Glandular cilia
on the edge of the sepals protruding and at least equal to half the width of the sepals. Bracts at base with glandular ears ................. ............................................................................................................................................................ 16. - H. rumelimm Boiss.
15* Boxes with black violet round glandular bubbles. The glandular cilia on the edge of the sepals abut the edge, less than half the
width of the sepals. Bracts at base without glandular ears .............................................................................. 12. - H. richerii Vill.
16 The sepals are just long ciliated. The styles 5 ……………........................................................…… 17. - H. thasium Griseb.
16* The sepals are glandular ciliated, glandularly toothed, or glandular and simply ciliated. Columns 3 .............……………...... 17
17 Leaves without intramarginal glands, rarely with several apical ones. Bracts entire ……..........…. 4. H. hyssopifolium Chaix
17* Leaves along the entire edge with intramarginal glands. Bracts glandular toothed or glandular ciliated .................................. 18
18 Leaves on the underside of the veins adjacent scattered short bristly ………..............…………….. 10. - H. setiferum Stef.
18* Leaves on the underside bare ……………………………….................................................…..………………………. 10
19 Glandular cilia equal to or longer than the width of the sepals, whitish below the gland. Boxes with longitudinal parallel resin
grooves ………............................................................................................................……. 18. - H. aucherii Jaub. and Spach
19* Glandular cilia shorter than the width of the sepals and up to the gland green. Boxes with ovoid, oval or elliptical glandular
vesicles or vesicles linear but then oblique to the edges of the carpels ......................................................................................... 20
20 The sepals are linearly lanceolate or wedge-shaped. The box most often with short oval glandular bubbles ……...............……. …........................................................................................................................……… 9. Moitbret's bell - H, montbretii Spach
20* Sepals ovate, ovate-lanceolate, broadly oval or broadly lanceolate. Boxes with ovoid, long oval or short linear, but in this case
oblique bubbles ........................................................................................................................................................................ 21
21 Leaves in the axils with shortened twigs or tufts of leaves. Sepals ovate or broadly lanceolate, obtuse or shortly pointed. The
boxes with linear, short, to the edges of the fruiting resinous channels. Petals with marginal glands only ............................................. ……...................................................................................................................................................... 22. - H. elegans Stephen
21* Leaves in the axils without shortened twigs. Sepals ovate or broadly lance, more or less pointed. The boxes with ovoid or
elliptical glandular bubbles. Petals at least to the upper half with superficial black glands ............................................................. 22
22 Leaves on the underside except with intramarginal and black superficial glands; the upper leaves most often around the midrib
and with transparent glands …....................................................................................................................14. - H. boissieri Petr.
22* Leaves on the underside with only marginal glands …...................................................... 13. - H. rochelii Griseb . et Schenk² The location and shape of the glands on the leaves, sepals and petals as well as the glandular bubbles on the surface of the fruit box are of great importance for the taxonomy of the genus. Glands in shape are: punctate, elliptical or linear, and in location: marginal, if their position breaks the line at the edge of the leaf {sepals, petals), intramarginal, if they are located very close to the edge line, but do not break it and superficial, if scattered on the surface of the organ concerned.
The glandular cells in the peripheral layer of the fruit box are: hemispherical, oblong or linear, in the latter case most often parallel to each other, with a tubular or peripheral location on the carpels, parallel or inclined with respect to the seams of the latter.From: „Флора на Н. Р. България”, том IV, Издателство на БАН, София, (1970) * * * * * Hypericum /ˌhaɪˈpiːrɪkəm/ is a genus of flowering plants in the family Hypericaceae (formerly considered a subfamily of Clusiaceae).[2][3] The genus has a nearly worldwide distribution, missing only from tropical lowlands, deserts and polar regions.[4] Many Hypericum species are regarded as invasive species and noxious weeds. All members of the genus may be referred to as St. John's wort, and some are known as goatweed.[2] The white or pink flowered marsh St. John's worts of North American and eastern Asia are now separated into the genus Triadenum.[5]
Hypericum is unusual for a genus of its size because a worldwide taxonomic monograph[6] was produced for it by Norman Robson (working at the Natural History Museum, London). Robson recognizes 36 sections within Hypericum.Description
Hypericum species are quite variable in habit, occurring as trees, shrubs, annuals, and perennials. Trees in the sense of single stemmed woody plants are rare, as most woody species have multiple stems arising from a single base. Shrubs have erect or spreading stems but never root from nodes that touch the ground. However, perennial herbs tend to root from these horizontal nodes, especially those that occur in wet habitats. Annual herbs tend to have taproots with a developed system of secondary hair roots. Many species of Hypericum are completely glabrous, others have simple uniseriate hairs, and some species have long, fine hairs.[7]
Two types of glands form the characteristic punctiform patterns of Hypericum, "dark glands" and "pale glands". Dark glands consist of clusters of cells with a distinct black to reddish color. Their hue is indicative of a presence of naphthodianthrone, either hypericin or pseudohypericin, or both. These glands occur in about two-thirds of Hypericum sections and are usually restricted to certain organs. When these glands are crushed, the naphthodianthrones give a red stain. Paracelsus called the red secretions "Johannes-blut" in the 16th century, linking the plant to the martyr St. John and giving rise to the English and German common names of "St. John's wort". The pale glands, forming the pellucid dots, are each a schizogenous intracellular space lined with flattened cells that secrete oils and phloroglucinol derivates, including hyperforin. The distribution of these hypericin glands dissuades generalist herbivores from feeding on the plants. When generalist insects feed on Hypericum perforatum, 30-100% more naphthodianthrones are produced, repelling the insects.[7]
The four thin ridges of tissue along the stems are closely to the opposite-decussate leaves of Hypericum. The ridges can be minor, just being called "ridges", or prominent, being called "wings". Terete, two-lined, and six-lined stems can occur occasionally. When a species has a tree or shrub habit, the internodes become mostly terete with age, though some trace of lines can still be detected in mature plants. The number of lines is an important distinguishing characteristic; for example, H. perforatum and Hypericum maculatum are easily confused save for H. perforatum having two lines and H. maculatum having four. The pale and dark glands are present on stems of various species, and other various species have stems without any glands. In section Hypericum, the glands are only present on stem lines, and in other sections, including Origanifolia and Hirtella, the glands are distributed across the stems.[7]
Nearly all leaves of Hypericum species are arranged opposite and decussate, an exception being section Coridium in which whorls of three to four leaves occur. The leaves lack stipules and can be sessile or shortly petiolar, though long petioles exist in sections Adenosepalum and Hypericum. Basal articulation can be present, in which case leaves are deciduous above the articulation, or absent, in which case the leaves are persistent. Some species in sections Campylosporus and Brathys have an auricle-like, reflexed leaf base, whereas true auricles only exist in sections Drosocarpium, Thasia, and Crossophyllum. Laminar venation is highly variable, being dichotomous to pinnate to densely reticulate. Leaves are typically ovoid to elongate to linear in shape. Leaves are typically shorter than the internodes. Pale or dark glands can be present on or near the leaf margin and on the main leaf surface.[7]
Typically there are four or five sepals, though in section Myriandra there are rarely three. When five sepals are present they are quincuncial, and when four sepals are present they are opposite and decussate. Sepals can be equal or unequal. Sepals can be united at their base, as seen in sections Hirtella, Taeniocarpium, and Arthrophyllum. The margins are variable, having marginal glands, teeth, or hairs. The presence or absence of dark glands on the sepals is a useful distinguishing characteristic.[7]
Almost all Hypericum petals are yellow, though a range of color exists from a pale lemony hue to a deep orangish-yellow. Exceptions include the white or pinkish petals of Hypericum albiflorum var. albiflorum and H. geminiflorum. Many species have petals that are lined or tinged with red, including the deep crimson petals of H. capitatum var. capitatum. Petal lengths can be equal or unequal. The petals are mostly asymmetrical except those of sections Adenotrias and Elodes. In those two sections, sterile bodies have developed between the stamen fascicles, working as lodicules to spread the petals of the pseudotubular flower, a specialized pollination mechanism. Nearly all species have glands on their petals; only section Adenotrias has completely eglandular petals. It has been hypothesized that the intensity of red on the petals is correlated with the hypericin content of the glands, but other pigments including skyrin derivatives can create a red color.[7]
Hypericum flowers have four or five fascicles that have, in total, five to two hundred stamens. The fascicles can be free or fused in various ways, often into three apparent fascicles. In sections Myriandra, Brathys, and some of Trigynobrathys, the stamens form a ring. Though stamens are usually persistent, some are deciduous. The stamens have an anther gland on the connective tissue, varying in color from amber to black.[7]
The ovaries are three or five-merous, occasionally two-merous, with a corresponding number of free or united styles. Developing seeds are borne on axile or parietal placentae, with at least two ovules per placenta. Hypericum fruits are dissimilar to most of Hypericaceae, being capsular and dehisce from the apex. The capsule can be dry or remain fleshy when mature. The capsules have elongate or punctate glands on their surface that create various shapes and patterns. These glands are typically pale amber, though in section Drosocarpium the glands are reddish-black. Extractions of these glands in certain species yielded phloroglucinol and terpenoid derivatives, suggesting a connection between these glands and the pale glands of vegetative tissue. Seeds of Hypericum species are small and range in color from a yellowish brown to dark purplish brown. The seeds are cylindric to ellipsoid and may have narrow wings. In some seeds, a basal ridge may be present, and rarely in section Adenotrias an apical caruncle is present which attracts ants to disperse seeds. Some species have highly specific germination and survival condition requirements. For example, H. lloydii is susceptible to a fungal infection as a seedling if conditions are too moist, whereas other species including H. chapmanii can grow underwater.[7]
From Wikipedia, the free encyclopedia
* * * * *
Distribution in Bulgaria: (Conspectus of the Bulgarian Vascular Flora) = conspectus&gs_l= Zlc.
Distribution:
References: „Флора на Н. Р. България”, том IV, Издателство на БАН, София, (1970), Wikipedia, the free encyclopedia
SPECIES:
Hypericum hirsutum L. - Hairy St John's-wort
Hypericum montbretii Spach - not found common name
Hypericum perforatum L. - Perforate St. John’s-wort
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