Hypericum perforatum
(St John's wort)

Pictures

Picture	Title	Caption	Copyright
Title Flowers Caption Hypericum perforatum (perforate St John's-wort); cluster of flowers. Copyright ©Trevor James/Hamilton, New Zealand-2015	Flowers	Hypericum perforatum (perforate St John's-wort); cluster of flowers.	©Trevor James/Hamilton, New Zealand-2015
Title Flower Caption Hypericum perforatum (St John's wort); individual flower. New Zealand. Copyright ©Trevor James/Hamilton, New Zealand-2015	Flower	Hypericum perforatum (St John's wort); individual flower. New Zealand.	©Trevor James/Hamilton, New Zealand-2015
Title Habit Caption Hypericum perforatum (St John's wort); vegetative prostrate plant. New Zealand. Copyright ©Trevor James/Hamilton, New Zealand-2015	Habit	Hypericum perforatum (St John's wort); vegetative prostrate plant. New Zealand.	©Trevor James/Hamilton, New Zealand-2015
Title Flowering habit Caption Hypericum perforatum (St John's wort); habit, as erect, flowering shoots. New Zealand. Copyright ©Trevor James/Hamilton, New Zealand-2015	Flowering habit	Hypericum perforatum (St John's wort); habit, as erect, flowering shoots. New Zealand.	©Trevor James/Hamilton, New Zealand-2015
Title Flowering habit Caption Hypericum perforatum (St John's wort); habit, flowering shoots. New Zealand. Copyright ©Trevor James/Hamilton, New Zealand-2015	Flowering habit	Hypericum perforatum (St John's wort); habit, flowering shoots. New Zealand.	©Trevor James/Hamilton, New Zealand-2015
Title Habit Caption Hypericum perforatum (St John's wort); tall, erect, flowering shoot. New Zealand. Copyright ©Trevor James/Hamilton, New Zealand-2015	Habit	Hypericum perforatum (St John's wort); tall, erect, flowering shoot. New Zealand.	©Trevor James/Hamilton, New Zealand-2015
Title Fruiting capsules Caption Hypericum perforatum (St John's wort); fruiting capsules. New Zealand. Copyright ©Trevor James/Hamilton, New Zealand-2015	Fruiting capsules	Hypericum perforatum (St John's wort); fruiting capsules. New Zealand.	©Trevor James/Hamilton, New Zealand-2015
Title Leaf Caption Hypericum perforatum (St John's wort); leaf, showing dark glands on underside. New Zealand. Copyright ©Trevor James/Hamilton, New Zealand-2015	Leaf	Hypericum perforatum (St John's wort); leaf, showing dark glands on underside. New Zealand.	©Trevor James/Hamilton, New Zealand-2015
Title Shoot Caption Hypericum perforatum (St John's wort); shoot. New Zealand. Copyright ©Trevor James/Hamilton, New Zealand-2015	Shoot	Hypericum perforatum (St John's wort); shoot. New Zealand.	©Trevor James/Hamilton, New Zealand-2015
Title Rhizomes and roots Caption Hypericum perforatum (St John's wort); rhizomes and rooting systems. New Zealand. March, 2012. Copyright ©Trevor James/Hamilton, New Zealand-2015	Rhizomes and roots	Hypericum perforatum (St John's wort); rhizomes and rooting systems. New Zealand. March, 2012.	©Trevor James/Hamilton, New Zealand-2015

Identity

Preferred Scientific Name

• Hypericum perforatum L.

Preferred Common Name

• St John's wort

Other Scientific Names

• Hypericum assurgens Peterm. ex Rouy
• Hypericum deidesheimense Sch.Bip. ex Trevir.
• Hypericum lineolatum Jord.
• Hypericum marylandicum Biroli ex Colla
• Hypericum perforatum subsp. angustifolium (DC.) GAUDIN.

International Common Names

• English: common goatweed; common St John's wort; common St Johnswort; goatweed; klamath weed; perforate St John's wort; perforate St Johnswort; racecourse weed; St Johnswort; tipton weed
• Spanish: hierba de San Juan; hipericon
• French: herbe a mille trous; millepertuis perfore
• Chinese: guan ye lian qiao
• Portuguese: hipericao; milfurada

Local Common Names

• Germany: Tuepfel- Hartheu; Tuepfel- Johanniskraut
• Italy: erba di San Giovanni; iperico perforato
• Netherlands: Sint-janskruid
• Russian Federation: zveroboj obyknovenny
• South Africa: Johanneskruid
• Sweden: äkta johannesört; Johannesoert

EPPO code

• HYPPE (Hypericum perforatum)

Summary of Invasiveness

H. perforatum is a medicinal plant which is poisonous to livestock, and which is considered a weed even in some of the countries where it is native. Such countries include Turkey, Italy, France, Hungary and Sweden, where it is a weed mostly in poorer pastures, neglected areas and occasionally in crops (Parsons and Cuthbertson, 1992). It has been widely introduced, and has become a serious invasive problem in parts of Canada, USA, Mexico, Argentina, Chile, South Africa, Reunion, Japan, Australia, New Zealand and Hawaii, often because of the damaging photosensitivity it can cause in livestock. Once established in an area it spreads vegetatively and is hard to control without damaging other pasture species. It is invasive in natural ecosystems as well as in agricultural land. This species is considered to have a number of herbal uses and has therefore been spread to many places as a useful medicinal or ornamental plant as well as an accidental hitchhiker on agricultural produce.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Theales
•                         Family: Clusiaceae
•                             Genus: Hypericum
•                                 Species: Hypericum perforatum

Notes on Taxonomy and Nomenclature

The family Clusiaceae comprises about 50 genera and 1200 species. The genus Hypericum has about 450 species, ranging from annual or perennial herbs to shrubs and trees in all temperate areas of the world. A number of species in the genus have been cultivated worldwide, several of them subsequently becoming invasive. Prohibited weeds and invasive species, as well as H. perforatum, include H. androsaemum, H. triquetifolium, H. calycinum and H. canariense.

Many subspecies and varieties of H. perforatum have been named but ITIS (2015) accepts none of these. The Plant List (2013), however, accepts three subspecies: - subsp. chinense N.Robson; subsp. songaricum (Ledeb. ex Rchb.) N.Robson; subsp. veronense (Schrank) H.Lindb. Others are still to be resolved.

H. perforatum subsp. chinense is native to China and has been introduced to Japan (GBIF, 2015). H. perforatum subsp. songaricum is present in the mountains of north-eastern Kazakhstan and adjacent Kyrghizstan and Xinjiang (China); then, after a gap of over 2500 km, it recurs in southern Russia (Astrakhan, Ciscaucasus) and Crimea (Robson, 2002). According to Robson (2003), H. perforatum subsp. veronense has a natural range extending from north of the Caucusus region eastwards to Tajikstan and north-west India, westward to Turkey, the Mediterranean region and Macaronesia as far as the Azores and southward to western Saudi Arabia.

Several suggestions have been put forward to explain the name common name ‘St. John’s wort’, among them: the use of the flowers as a source of red pigment known as St. John’s blood; in Europe the plant flowers on about 24 June, the anniversary of the birth of St. John; an association with the nights of St. John and the Crusades; it used to be hung around houses on St. John’s Eve to ward off evil spirits (Parsons and Cuthbertson, 1992). ‘Wort’ is from the old English word meaning plant or herb, especially one with medicinal properties.

Description

Erect perennial herb 30-90 cm high reproducing by seeds or from rhizomes at the base of the stem. Stems much branched, 2-sided or ridged with black glands along the ridges, hairless, a distinctly dark ring at the lower nodes. Leaves simple, opposite, pinnately netted, with no stalk, usually without teeth, hairless with translucent dots, apex rounded. Flowers showy, yellow, in fairly flat-topped clusters at the top of the plant. Sepals 5, green, lance-shaped, 5 mm long, 1 mm wide. Petals 5, yellow with black glandular dots along the margins, 14-15 mm long, 8 mm wide. Stamens many; one pistil with style 3-parted and widely divergent; fruit a capsule; seeds l.l x 0.5 x 0.5 mm, black, reticulate and with short, sharp points at the ends.

Plant Type

Distribution

H. perforatum was originally distributed in Europe, Western Asia and North Africa, with a broad Eurasian native range extending from the Atlantic seaboard of Europe to China and India. It has now been introduced, accidentally or deliberately, to many temperate parts of the world, most notably North America, Australia, South Africa and New Zealand, in all of which it has become a serious problem as a pasture weed poisonous to livestock. 

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

H. perforatum has been carried to new countries either by accident as a contaminant of crop seed, hay, straw and packing materials, or deliberately as a medicinal plant or garden ornamental. Its introduction to Australia in the 1800s appears to have been as a curiosity or ornamental, first grown in Botanical Gardens (it appeared in a catalogue of plants growing in the Melourne Botanic Gardens in 1857) but later sold in nurseries as a garden plant (Parsons and Cuthbertson, 1992). The first recorded occurrence of H. perforatum in North America was in Pennsylvania in in 1793 (Sampson and Parker, 1930). In 1893 it was reported on western United States grazing lands, and Gillett and Robson (1981, cited in Crompton et al., 1988) indicated its presence as a weed in western Canada by 1940. It was introduced into South Africa in 1942 from a contaminated batch of vetch seed (Gordon and Kluge, 1991).

Introductions

Risk of Introduction

The risk of H. perforatum being introduced to yet more temperate countries is clearly high, especially as its small seeds are easily hidden amongst other seed samples or in hay or straw taken from one country to another. In addition this species is also likely to be carried legally or illegally to other countries as a useful medicinal plant. 

Habitat

H. perforatum grows in humid and sub-humid temperate regions, usually on drier sites. It becomes invasive (weedy) when it invades poorly managed grazing land, sparse bushland, roadsides and neglected areas. In Australia it establishes well on loose soil in areas with a rainfall of above 750 mm (Parsons and Cuthbertson, 1992). Improved pastures are rarely invaded except where they are over-grazed or otherwise mistreated.

In Britain, Stace (2010) describes its habitat as ‘dryish grassland, banks and open woodland’. In Pakistan its habitat is described as ‘In wet or dryish, shaded or open grassy places often near streams or sometimes a weed of cultivation, 1200-3000m’ (Flora of Pakistan, 2015). Flora of China Editorial Committee (2015) says ‘Open woodlands, meadows, grasslands, and steppes, riverbanks, stony and grassy slopes, roadsides, in dry or well-drained habitats; 100-2800m.’

In New Zealand, Webb et al. (1988) say of its habitat ‘A particularly troublesome weed in tussock grasslands, also very common in pastures, roadsides, riverbeds, waste places and in many modified open communities.’ At one time this species was responsible for appreciable harm to livestock, especially sheep in South Island highland tussock grassland, though cattle too sometimes suffered in the same environment (Connor, 1992). In Australia, H. perforatum occurs in grasslands, open Eucalyptus and Callitris woodlands, along river banks, cleared pasture land and forest plantations (Csurhes and Zhou, 2008).

H. perforatum is often mentioned as being found in semi-shade, but this is usually along roads and at the margins of forests: it rarely persists in mature forest stands (Tisdale et al., 1959). Buckley et al. (2003) indicate that plants growing in shaded sites grow more slowly and produce less fruit but live longer. 

Habitat List

Hosts/Species Affected

In most countries H. perforatum is a problem in poorly managed pastures or rangelands because it can crowd out existing useful pasture species and inhibit the germination and growth of other species. The plant contains hypericin, a red pibment concentrated in the black oil glands, which is toxic to livestock when consumed in quantity.

Elsewhere it is found along roadsides, in waste places, forest clearings and other sites (Crompton et al., 1988). Crompton et al. said it was becoming an increasingly important weed in lowbush blueberry culture in eastern North America.  

Biology and Ecology

In Australia and presumably elsewhere, seeds germinate in autumn, winter or spring and plants rarely flower in their first year. Established plants have two life forms. In winter and spring the plants form low-growing stems 30 cm long and bearing dense foliage which gives a dense, smothering mat. In spring, erect woody stems develop and flowering occurs (in Australia) in late spring (November) and they continue flowering well into summer. These flowering stems die back in late summer but can remain standing for several months or sometimes years (Parsons and Cuthbertson, 1992).

Zohar (2004) explained, quoting a number of studies, that H. perforatum exhibits a high degree of phenotypic and genotypic variation, particularly among populations. Studies comparing the characteristics of populations in California, Oregon and Montana found strong regional differences in chemical (hypericin and pseudohypericin) content and morphological (gland density, leaf area, stem length, leaf length:width ratio) characteristics (Shiflet, 1994;  Walker et al., 2001; Sirvent et al., 2002). The species also shows genetic variation in morphological variation and herbivore resistance (Campbell et al., 1997; Mayo and Roush, 1997; Buckley et al., 2003).

Genetics

Gillett and Robson (1981) found that in Canada haploid chromosome numbers varied from 16 to 18 while diploid numbers were 32 with one report of 48. The majority of studies cited in Missouri Botanical Gardens (2015) give the chromosome count as 2n = 32. 

Reproductive Biology

H. perforatum is a facultative apomict and is also pseudogamous (Pritchard, 1960) meaning that fertilisation is necessary to promote development of the endosperm in the seed but not to fertilise the ovum.

Seed production is prolific and a single plant can produce 33,000 seeds in a season, with some of these seeds remaining viable in the soil for 10 years (Parsons and Cuthbertson, 1992). Tisdale et al. (1959) observed a high degree of variation in field studies in Idaho, with seed production per plant ranging from a few thousand to nearly 100,000. Andersen (1968) reported that seeds maintained a germination rate of 50% after 15 years of dry storage at laboratory temperature. Tisdale et al. (1959) found no loss in germination for 3 years when seeds were buried 1.2 cm deep in the soil.  

In addition to its seed production, H. perforatum reproduces vegetatively by rhizomes, root fragments or lateral root sprouts (Zouhar, 2004). Sometimes this results in several crowns sharing a common rooting system, and also makes the longevity of individual plants next to impossible to determine. Harris (1971) says new St Johnswort plants originate at intervals along underground roots that may extend 90 cm or more from the parent plant. Plants propagated vegetatively severed connections with the parent plant fairly soon after new plants had established (Zouhar, 2004).

In eastern Canada, H. perforatum flowers during summer (late June, July and early August), with seeds reaching maturity in autumn (late August).

Thomson (1922) says that in Europe its flowers are visited by Apis mellifera, Bombus hortorum, B. terrestris, Eristalis tenax and Calliphora erythrocephala, presumably for nectar and pollen.

Physiology and Phenology

Fresh seed does not appear to germinate readily and requires 4-6 months after-ripening; this temporary dormancy may be due to an inhibitor in the sticky material associated with the seed capsules. However this requirement for after-ripening has been disputed by other authors (Crompton et al., 1988). Germination can be enhanced by alternating temperatures, gibberellic acid or potassium nitrate.

Seedling establishment is favoured by bare soil and wet summers (Clark, 1953).

Pritchard (1960) compared populations of H. perforatum from British Columbia, Ontario, California, Britain and New Zealand. He found that populations from British Columbia and California were taller and more aggressive, produced greater biomass, and were, as a consequence, more weedy. Under favourable conditions, plants developed vertical roots, whilst under less favourable conditions rhizomes developed actively. However other authors (Vila et al., 2003; Marron et al., 2004) have disputed these findings, observing that plants from the introduced range were neither uniformly larger nor better interspecific competitors than plants from their native range.

Longevity

Seeds can remain viable in the soil for considerable periods if conditions are suitable for long-term survival (Zouhar, 2004).

Under at least some conditions, seedling mortality can be high and is probably exacerbated by competition from mature plants of the same species or by other plant species (Tisdale et al. 1959; Campbell, 1985). This is at least partly due to the slow growth of H. perforatum seedlings.

The median longevity of individual plants from four sites in Australia with various climatic profiles was 3 to 6 years with an overall range of 1 to 8 years (Briese, 1997). Very few crowns survived longer than 8 years: death of crowns was attributed to drought, fire, defoliation by Chrysolina  beetles, combinations of these, and senescence.

Activity Patterns

Seedlings of H. perforatum are very small and grow slowly so that young seedlings are susceptible to competition for light, nutrients, space and moisture from mature plants of their own and other species (Briese, 1997). In autumn and winter, H. perforatum has a low-growing, very leafy growth form, effectively swamping growth by low-growing or emerging plants of other species. In spring, stems become much taller and woody and carry flowers. Plants produce underground rhizomes and long taproots which allow the plants to be drought-tolerant by comparison with nearby neighbours of other species.

Population Size and Structure

Crompton et al. (1988) describe H. perforatum as a particularly aggressive weed of rangelands characterised by dry summers. Sampson and Parker (1930) said its deep root system can support the plant when water for more desirable plants has been depleted. It forms a dense, spreading canopy up to 1 m tall, and large infestations in western North America covered over 1 million hectares in the 1950s, according to Holloway (1957). These authors also said that in California plants could produce 4300 kg ha^-1 and that vegetative reproduction seemed to be stimulated by grazing, fire and defoliation.

According to Zouhar (2004), crown and stem density of H. perforatum may be affected by many variables. Stem production can be affected by rainfall pattern, insect defoliation, and site factors (Briese, 1997), with St Johnswort population fluctuations and dynamics driven by complex interactions of several stress factors (e.g. shallow, rocky soils, shading, drought, herbivory), and/or positive factors such as good rainfall years. For example, deep soils favour development of vertical roots and long-term crown survival, while shallow soils tend to support H. perforatum populations with more lateral roots, root sprouts, and shorter-lived crowns.

Harris (1971) describes a ‘typical’ St Johnswort near Colville, Washington, as dense, ‘almost to the exclusion of other herbaceous vegetation.’

Nutrition

Several authors have commented that H. perforatum is less common on calcareous soils and seems to prefer soils with slightly acid to neutral pH (Sampson and Parker, 1930; Daubenmire, 1947, cited by Crompton et al, 1988). Sampson and Parker suggested that while H. perforatum is not restricted to acid or alkaline soils, unusually luxurious growth and dense stands are more typically found on soils with pH 5 to 56.5 while very vigorous growth was not observed on strongly alkaline soils.

Associations

Crompton et al. (1988) list the species associated with H. perforatum at two locations in Nova Scotia. Most of the associated species are common weeds found in most places. Sampson and Parker (1930) describe dense St Johnswort stands in California with winter annuals in the understory.

Harris and Clapperton (1997) noted that 31% of the roots of H. perforatum plants growing in Canadian field infestations were infected with vesicular-arbuscular mycorrhizal (AM) fungus. H. perforatum seedlings inoculated with AM fungi were better able to tolerate harsh environmental conditions (Moora et al., 1998).

Environmental Requirements

H. perforatum is a predominantly temperate species with a very wide native range and which can tolerate a broad range of environmental conditions. Its natural range extends from parts of Central Asia with very low winter temperatures, to subtropical environments such as the Canary Islands. The species can tolerate a wide variety of soils, from dry, rocky, shallow soils to deep, fertile soils, with its best performance coming in areas with at least 760 mm of rainfall a year. Soil pH tolerance ranges from 4.3 to 7.6 (Rutledge and McLendon, 1996). It can tolerate drought and disturbance by means of reserves stored in the root crown (Buckley et al., 2003). It does not tolerate saturated soils.

Climate

Latitude/Altitude Ranges

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil reaction

• acid
• alkaline
• neutral

Soil texture

• heavy
• light
• medium

Special soil tolerances

• infertile

Natural enemies

Notes on Natural Enemies

H. perforatum hosts at least 37 species of insects in its native environment (Holloway, 1964), suggesting that at least part of its success in new countries is due to the absence of natural pests. Two leaf-feeding beetles Chrysolina hyperici and C. quadrigemina have been introduced as biological control agents and these have been very successful in some places (New Zealand, British Columbia and California) but less successful in others, like parts of Australia (see Biological Control section, under Prevention and Control). 

Means of Movement and Dispersal

Seeds can apparently be carried by wind for 30 m or more (Tisdale et al., 1959), by water, and by birds and animals. Country to country dispersal as well as wider dispersal within countries are almost certainly assisted by people taking the plants with them for herbal or medicinal purposes.

Natural Dispersal

The seeds of H. perforatum are tiny and are readily carried by air currents, water, mud, soil and agricultural produce (Parsons and Cuthbertson, 1992). Sampson and Parker (1930) suggested that the gelatinous seed coat may help dispersal by animals and that initial infestations often followed animal movements. Young plants fresh to a new area can spread locally by means of their long rhizomes.  

Vector Transmission (Biotic)

The sticky seed capsules of H. perforatum readily attach to wool or fur of animals and survive passage through the alimentary canal (Parsons and Cuthbertson, 1991).

Accidental Introduction

H. perforatum, which is found growing in pastures and along roadsides, has probably been harvested in hay crops and thus carried from farm to farm. Historically, hay has been carried from country to country as food for animals in transit and it is very likely that this species would have been carried to new countries like Australia and New Zealand.

Intentional Introduction

There is little doubt that the successful movement of H. perforatum from country to country and within countries has been partly due to transport by humans who have long regarded this species as a useful medicinal plant. 

Pathway Causes

Pathway Vectors

Impact Summary

Economic Impact

H. perforatum has long been known for its poisonous properties, especially to livestock. The pigment hypericin, concentrated in the oil glands, is toxic when consumed in quantity (Csurhes and Zhou, 2008). Cirillo (1787, cited in Mabberley, 2002) was the first to report that its toxicity affected white but not black sheep. Light-coloured animals become sensitive to sunlight (photosensitisation) after eating the plants, and the exposed unpigmented parts of the bodies of animals are most affected. For example, the faces of sheep and horses and white body markings on cattle are severely affected and the animals suffer intensely, rubbing and biting at infected areas until they become raw and sometimes pieces of skin become sloughed off. Tips of the ears of affected sheep harden and may snap off and their sore mouths make grazing difficult so that the animals lose condition (Parsons and Cuthbertson, 1992). Furthermore, affected animals cannot stand contact with water and if this happens they will struggle and may drown. Animals will recover if access to the plant is prevented (Connor, 1992). Plant material is poisonous at all times of the year and drying (as in hay) does not diminish the risk.

The species is strongly competitive to other species throughout the year and, when firmly established, can eliminate almost all other plants. Parsons and Cuthbertson (1992) say that this has led to the abandonment of formerly productive land in north-eastern Victoria and New South Wales in Australia.

Environmental Impact

H. perforatum often invades poor or badly managed pastures but also occurs in open woodlands where its growth probably impedes the growth of native plant species. In southeastern Australia, it infests an estimated 400,000 ha of which some 80% supports natural vegetation, mainly open woodland (Csurhes and Zhou, 2008). It can become a serious temporary problem in some habitats but will eventually be succeeded by other, longer-lived species if these are left undisturbed. 

Social Impact

The economic hardships caused by this species must have disrupted livestock farmers and farming communities in areas where this weed has become a widespread problem.

PFAF (2015) lists a number of hazards associated with the plant and products from it, quoting a number of authors. Among these are: skin contact with the sap, or ingestion of the plant, can cause photosensitivity in some people; common side-effects are gastrointestinal disturbances, allergic reactions and fatigue; if used with drugs classed as serotonin reuptake inhibitors (fluoxetine:Prozac, paroxetine:Paxil) symptoms of serotonin syndrome may occur - mental confusion, hallucinations, agitation, headache, coma, shivering, sweating, fever, hypertension, tachycardia, nausea, diarrheoa, tremors. St John's wort can also reduce the effectiveness of prescription medicines including the contraceptive pill, antidepressants, immune suppressants, HIV medications, warfarin, and digoxin.

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Pioneering in disturbed areas
• Tolerant of shade
• Highly mobile locally
• Benefits from human association (i.e. it is a human commensal)
• Fast growing
• Has high reproductive potential
• Has propagules that can remain viable for more than one year
• Reproduces asexually
• Has high genetic variability

• Ecosystem change/ habitat alteration
• Negatively impacts agriculture
• Negatively impacts animal health
• Negatively impacts aquaculture/fisheries

• Causes allergic responses
• Competition - monopolizing resources
• Competition - shading
• Induces hypersensitivity
• Poisoning
• Rapid growth

• Highly likely to be transported internationally accidentally
• Highly likely to be transported internationally deliberately
• Difficult to identify/detect as a commodity contaminant
• Difficult/costly to control

Uses

H. perforatum is a well-known and well-studied medicinal herb (Sheahan, 2012). The translucent dots on the leaves and peppered spots on the petals are sacs of essential plant oils. When crushed, the black glands on the petals exude dark red oil. These essential oils make common St. Johnswort a popular homeopathic supplement for a variety of conditions. The most important risk of taking St. Johnswort products medicinally are their interactions with other drugs. Homeopathic tinctures, pressed juices, or oils can be prepared from the fresh plant material. Dried plant material can also be used in teas.

Plants of H. perforatum have been used as an ingredient in the distillation of vodka, as a herbal tea, and as a source of red, yellow, purple and orange dyes (Parsons and Cuthbertson, 1992). An essential oil from the plant is used in the cosmetics industry and the plant is grown commercially in Poland and other places in southern Europe for this purpose.

This species is widely known as a herbal treatment for depression; in Germany and some other countries it is commonly prescribed for mild to moderate depression, especially in children and adolescents (Fegert et al., 2006).

Linde et al. (2008) reviewed 29 trials involving 5489 patients testing H. perforatum extracts against either placebo or standard antidepressants and concluded that H. perforatum extracts are superior to placebo in patients with severe depression; they give similar results to standard anti-depressants and have fewer side-effects than standard anti-depressants. The authors stressed caution, however, saying that ‘However, findings were more favourable to St. John’s wort extracts in studies from German-speaking countries where these products have a long tradition and are often prescribed by physicians, while in studies from other countries St. John’s wort extracts seemed less effective.’ Nonetheless these findings have repeatedly been mentioned in many books and reports on the benefits of using H. perforatum for this purpose.

Economic Value

As a herbal remedy for depression H. perforatum is readily available either as seeds or as a herbal preparation via the internet and therefore presumably has economic value to herbalists. Recent interest in and widespread publication of its medicinal virtues has probably increased such commercial activity.

Social Benefit

Parsons and Cuthbertson (1992) list a number of traditional virtues such as its magical properties against storms, thunder, and witches. In medieval times it was apparently used a cure for bed wetting, insomnia and other nervous conditions as well as dysentery, rheumatism, gout, internal parasites, lung disorders, urinary problems, healing burns, other wounds and skin disorders. A tincture of flowers mixed with wine is a cure for melancholy and madness.

To that long list of benefits, Crompton et al. (1988) quote several sources in adding that Hypericum extracts are well known as antibiotics and may also be used for their anti-viral benefits.

Uses List

Drugs, stimulants, social uses

• Psychoactive

General

• Ritual uses
• Sociocultural value

Materials

• Chemicals
• Essential oils
• Lipids
• Poisonous to mammals

Medicinal, pharmaceutical

• Source of medicine/pharmaceutical
• Traditional/folklore

Similarities to Other Species/Conditions

Other species of Hypericum have similar flowers but the only ones likely to be confused with H. perforatum are those with similar life-forms. Thus H. canariense, native to the Canary Islands but introduced to several other countries as an ornamental, is described by GISD (2015) as ‘similar’ but is a shrub. Dwarf St. John's wort (Hypericum mutilum), also described by GISD (2015) as ‘similar’ is a smaller plant, half the height of H. perforatum, with a one-chambered capsule unlike H. perforatum which has a three-chambered capsule. This species is native to North America but has been spread to other countries. In New Zealand Popay et al. (2010) describe H. pulchrum as similar to H. perforatum, and Webb et al. (1988) report that ‘H. pulchrum may grow in the same locality as H. perforatum, but can be distinguished by the ± terete [round] stems, the even fringe of black glands on the sepals, and the reddish petals, anthers and stigmas.’

In Canada, Crompton et al. (1988) claim that H. scouleri and H. punctatum are similar but point out the important differences. In H. punctatum the black glandular dots are scattered throughout the petals whereas in H. perforatum they are restricted to the margins of the petals. H. scouleri has ovate-elliptical sepals and the black dots on them are elongated more than those sometimes found in H. perforatum.

In Australia, the indigenous species H. gramineum and H. japonicum can co-occur with H. perforatum and be confused with it (Csurhes and Zhou, 2008). Both native species can be distinguished by the absence of oil glands on their leaves and petals, the presence of four longitudinal ridges on the stem (two for St John’s wort) and the stamens not being fused into bundles.

Prevention and Control

Eradication

Zouhar (2004) stresses that control of H. perforatum infestations requires several strategies designed to affect different plant parts and different stages of the plant’s life history. These strategies will depend be site-specific and long in term as populations can quickly build up again.

Physical/Mechanical Control

Isolated plants should be grubbed before they produce seed, but large areas are best ploughed in the autumn, cultivated whenever regrowth shows, and sown the following autumn in an appropriate pasture mixture (Parsons and Cuthbertson, 1992). Crompton et al. (1988) said that H. perforatum is readily controlled by tillage and is therefore seldom a problem in cultivated crops. Such techniques are obviously limited to small infestations or to areas where tillage is possible.

Biological Control

The introduction of weevils of the genus Chrysolina has led to effective control of H. perforatum in several countries where the weed was a serious problem earlier. Thus it has been very effective in New Zealand, South Africa, the USA and Canada. In South Africa H. perforatum is now considered to be under satisfactory control in the Southwestern Cape (Gordon and Kluge, 1991), thanks to the introduction of Chrysolina quadrigemina in 1960 and of the gall midge Zeuxidiplosis giardi in 1972. In Australia, C. hyperici and C. quadrigemina adequately control the weed under open but not under shaded conditions (Shepherd, 1985). At shaded sites, plants appeared less aggressive than elsewhere and their growth habits differed from those of plants growing at open or semi-shaded sites. C. quadrigemina is now the dominant species and the main insect exerting some control on the plant. In New Zealand C. hyperici was introduced in 1943 and was followed two years later by releases of C. quadrigemina. Syrett et al. (1997) concluded that ‘There are no longer reports of areas in New Zealand where St. John's wort is a problem weed, and we conclude that successful biological control with insects is as least partially responsible for this change.’

In the western USA, where the species was once a problem, the success of the biocontrol agents has led to its replacement with annual grasses or with mixtures of forbs and grasses (Zouhar, 2004).

Chemical Control

Herbicides such as glyphosate can be used to get effective control of small infestations but are rarely the complete or long-term answer to the problem unless competitive species of plants are encouraged by oversowing and by appropriate grazing management. Herbicides are too expensive for use over very large areas of infestations, especially since consideration must also be given to oversowing of desirable species to replace the Hypericum and to prevent its resurgence. Campbell and Nicol (1987) tested a range of herbicides for the control of H. perforatum in Australia but concluded that the species is hard to selectively control with herbicides and that every effort should be made to promote biological control and/or effective grazing management.

IPM

Seedlings of H. perforatum are very susceptible to competition and offer the best stage at which to tackle infestations. This can be done by encouraging competitive species either by using careful grazing management or oversowing with pasture species. Any strategy must consider the presence of long-term soil seed banks. In Australia a programme of seeding pasture species (T. subterraneum and Phalaris spp.) combined with cultivation and fertilization on arable land has been proposed (Campbell and Nicol, 1997; Groves, 1997).

According to these same authors, in New South Wales and Victoria, change in land use from pastures badly invaded with H. perforatum to pine plantations has been used by farmers as an evasion strategy.

References

Andersen RN, 1968. Germination and establishment of weeds for experimental purposes: (A Weed Science Society of America Handbook). Illinois, USA: Weed Science Society of America, 236 pp.

Briese DT, 1997. Population dynamics of St. John's wort in south-eastern Australia. In: Plant Protection Quarterly, 12(2) [ed. by Jupp, P. W.\Briese, D. T.\Groves, R. H.]. 59-63.

Buckley YM; Briese DT; Rees M, 2003. Demography and management of the invasive plant species Hypericum perforatum. I. Using multi-level mixed-effects models for characterizing growth, survival and fecundity in a long-term data set. Journal of Applied Ecology, 40(3):481-493.

Campbell MH, 1985. Germination, emergence and seedling growth of Hypericum perforatum L. Weed Research, UK, 25(4):259-266.

Campbell MH; May CE; Southwell IA; Tomlinson JD; Michael PW, 1997. Variation in Hypericum perforatum L. (St. John's wort) in New South Wales. In: Plant Protection Quarterly, 12(2) [ed. by Jupp, P. W.\Briese, D. T.\Groves, R. H.]. 64-66.

Campbell MH; Nicol HI, 1997. Improving herbicide effectiveness on Hypericum perforatum L. (St. John's wort) and replacing it with pastures sown on non-arable land. In: Plant Protection Quarterly, 12(2) [ed. by Jupp, P. W.\Briese, D. T.\Groves, R. H.]. 93-96.

Cirillo DML, 1787. Fundamenta Botanicae.

CLARK N, 1953. The biology of Hypericum perforatum L. var. angustifolium DC (St. John's wort) in the Ovens Valley, Victoria, with particular reference to entomological control. Australian Journal of Botany, 1:95-120.

Connor HE, 1992. The poisonous plants in New Zealand. Wellington, New Zealand: Government Printer, 247 pp.

Council of Heads of Australasian Herbaria, 2015. Australia's virtual herbarium. Australia: Council of Heads of Australasian Herbaria. http://avh.ala.org.au/#tab_simpleSearch

Crompton CW; Hall IV; Jensen KIN; Hildebrand PD, 1988. The biology of Canadian weeds. 83. Hypericum perforatum L. Canadian Journal of Plant Science, 68(1):149-162.

Csurhes S; Zhou Y, 2008. Pest plant risk assessment: St John's wort (Hypericum perforatum). Brisbane, Australia: Biosecurity Queensland, Department of Primary Industries and Fisheries, 12 pp. https://www.daf.qld.gov.au/__data/assets/pdf_file/0011/63857/IPA-Hypericum-Perforatum-Risk-Assessment.pdf

Daubenmire RF, 1947. Plant and environment. A text book of plant autecology. New York, USA: John Wiley and Sons.

Fegert JM; Kölch M; Zito JM; Glaeske G; Janhsen K, 2006. Antidepressant use in children and adolescents in Germany. Journal of Child and Adolescent Psychopharmacology, 16(1-2):197-206.

Flora of China Editorial Committee, 2015. Flora of China. St. Louis, Missouri and Cambridge, Massachusetts, USA: Missouri Botanical Garden and Harvard University Herbaria. http://www.efloras.org/flora_page.aspx?flora_id=2

Flora of Pakistan, 2015. Flora of Pakistan/Pakistan Plant Database (PPD). Tropicos website. USA: St. Louis, Missouri and Cambridge, Massachusetts. http://www.tropicos.org/Project/Pakistan

GBIF, 2015. Global Biodiversity Information Facility. http://www.gbif.org/species

Gillett JM; Robson NKB, 1981. The St. John's-worts of Canada (Guttiferae). Publications in Botany, National Museum of Natural Sciences, No. 11. 40 pp.

GISD, 2015. Global Invasive Species Database (GISD). IUCN. http://www.issg.org/database/welcome/

Gordon AJ; Kluge RL, 1991. Biological control of St. John's wort, Hypericum perforatum (Clusiaceae), in South Africa. Agriculture, Ecosystems and Environment, 37(1-3):77-90.

Groves RH, 1997. St. John's wort: introduction. In: Plant Protection Quarterly, 12(2) [ed. by Jupp, P. W.\Briese, D. T.\Groves, R. H.]. 51-52.

Harris GA, 1971. Johnswort on western ranges: a review of the literature and bibliography. U.S. Department of Agriculture, Forest Service, Northern Rocky Mountain Forest and Range Experiment Station, 18 pp. [Station Paper No. 26. [Missoula, MT].] https://archive.org/stream/stjohnswortonwes26harr/stjohnswortonwes26harr_djvu.txt

Harris P; Clapperton MJ, 1997. An exploratory study on the influence of vesicular-arbuscular mycorrhizal fungi on the success of weed biological control with insects. Biocontrol Science and Technology, 7(2):193-201; 46 ref.

Holloway JK, 1957. Weed control by insects. Scientific American, 197:56-62.

Holloway JK, 1964. Projects in biological control of weeds. In: Biological control of insect pests and weeds [ed. by DeBach, P.]. New York, USA: Reinhold Publishing Corp.

ITIS, 2015. Integrated Taxonomic Information System online database. http://www.itis.gov

Linde K; Berner MM; Kriston L, 2008. St John's wort for major depression. Cochrane Database of Systematic Reviews, 2008(4):106 pp.

Mabberley, 2002. The Plant Book. Cambridge, UK: Cambridge University Press, 858pp.

Maron JL; Vilà M; Bommarco R; Elmendorf S; Beardsley P, 2004. Rapid evolution of an invasive plant. Ecological Monographs, 74(2):261-280. http://www.esajournals.org/perlserv/?request=get-document&doi=10.1890%2F03-4027

Mayo GM; Roush RT, 1997. Genetic variability of Hypericum perforatum L. (Clusiaceae) and the detection of resistance to the biological control agent Aculus hyperici Liro (Eriophyidae). In: Plant Protection Quarterly, 12(2) [ed. by Jupp, P. W.\Briese, D. T.\Groves, R. H.]. 70-72.

Moora M; Zobel M, 1998. Can arbuscular mycorrhiza change the effect of root competition between conspecific plants of different ages? Canadian Journal of Botany, 76(4):613-619.

Parsons WT; Cuthbertson EG, 2001. Noxious Weeds of Australia. Collingwood, Australia: CSIRO Publishing, 698 pp.

PFAF, 2015. Plants For A Future. http://www.pfaf.org/user/Default.aspx

PIER, 2015. Pacific Islands Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Popay I; James T; Champion P, 2010. An Illustrated Guide to Common Weeds of New Zealand. Christchurch, New Zealand: New Zealand Plant Protection Society, 448 pp.

PRITCHARD T, 1960. Race formation in weedy species with special reference to Euphorbia cyparissias L. and Hypericum perforatum L. In: Biology of Weeds, Symp. Brit. ecol. Soc. 61-6 pp.

Robson NKB, 2002. Studies in the genus Hypericum L. (Guttiferae) 4(2). Section 9. Hypericum sensu lato (part 2): subsection 1. Hypericum series 1. Hypericum.

Robson NKB, 2003. The genus Hypericum. In: Hypericum: the genus Hypericum. Medicinal and Aromatic Plants - Industrial Profiles [ed. by Ernst, E.]. London, UK: Taylor and Francis, 1-22.

Rutledge CR; McLendon T, 1996. An Assessment of Exotic Plant Species of Rocky Mountain National Park., USA: Department of Rangeland Ecosystem Science, Colorado State University, 97 pp. http://www.npwrc.usgs.gov/resource/plants/explant/index.htm

Salisbury E, 1964. Weeds and Aliens. 2nd Edition. London, UK: Collins.

Sampson AW; Parker KW, 1930. Johnswort on rangelands of California. California Agricultural Experiment Station Bulletin, 503:89 pp.

Sheahan CM, 2012. Plant guide for common St. Johnswort (Hypericum perforatum). Cape May, NJ, USA: USDA-NRCS, Cape May Plant Materials Centre. http://plants.usda.gov/plantguide/pdf/pg_hype.pdf

Shepherd RCH, 1985. The present status of St. John's wort (Hypericum perforatum L.) and its biological control agents in Victoria, Australia. Agriculture, Ecosystems and Environment, 12(2):141-149.

Shiflet TN, 1994. Rangeland cover types of the United States [ed. by Shiflet, T. N.]. Denver, CO, USA: Society for Range Management, 157 pp.

Sirvent TM; Walker L; Vance N; Gibson DM, 2002. Variation in hypericins from wild populations of Hypericum perforatum L. in the Pacific Northwest of the USA. Economic Botany, 56(1): 41-48

Stace CA, 2010. New flora of the British Isles. 3rd edition. Cambridge, UK: Cambridge University Press, 1232 pp.

Syrett P, 1997. Biological control of St. John's wort in New Zealand. In: Plant Protection Quarterly, 12(2) [ed. by Jupp, P. W.\Briese, D. T.\Groves, R. H.]. 88-90.

The Plant List, 2013. The Plant List: a working list of all plant species. Version 1.1. London, UK: Royal Botanic Gardens, Kew. http://www.theplantlist.org

Thomson GM, 1922. The naturalisation of animals & plants in New Zealand. Cambridge, UK: Cambridge University Press, 607 pp.

TISDALE EW; HIRONAKA M; PRINGLE WL, 1959. Observations on the autecology of Hypericum perforatum. Ecology, 40(1):54-62.

USDA-ARS, 2015. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

USDA-NRCS, 2015. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Vilà M; Gómez A; Maron JL, 2003. Are alien plants more competitive than their native conspecifics? A test using Hypericum perforatum L. Oecologia, 137(2):211-215.

Walker L; Sirvent T; Gibson D; Vance N, 2001. Regional differences in hypericin and pseudohypericin concentrations and five morphological traits among Hypericum perforatum plants in the northwestern United States. Canadian Journal of Botany, 79(10):1248-1255.

Webb CJ; Sykes WR; Garnock-Jones PJ, 1988. Flora of New Zealand. Vol. IV. Naturalised Pteridophytes, Gymnosperms, Dicotyledons. Christchurch, New Zealand: Botany Division, D.S.I.R. http://floraseries.landcareresearch.co.nz/pages/Taxon.aspx?id=_e58919e7-2438-4828-a928-3192b39645b4&fileName=Flora%204.xml

Zouhar K, 2004. Hypericum perforatum. Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory. http://www.fs.fed.us/database/feis/plants/forb/hypper/all.htm

Contributors

22/06/15 Original text by:

Dr. Ian Popay, Hamilton, New Zealand

Distribution Maps