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'''''Hypericum hircinum''''' is a species of [[Perennial plant|perennial]] [[flowering plant]] in the St John's wort family, [[Hypericaceae]]. It is known as '''goat St John's wort''' and '''stinking tutsan'''; both names refer to the plant's distinctive odor. The species is a bushy shrub that can grow up to 1.5 meters tall, is many-stemmed, and has golden yellow flowers with conspicuous [[Stamen|stamens]]. The plant has been well-documented in botanical literature, with mentions dating back to at least 1627. [[Carl Linnaeus|Carl Linneaus]] described ''H. hircinum'' several times, including in his 1753 ''[[Species Plantarum]]'' which established its [[Binomial nomenclature|binomial]]. At one point the plant was placed into the defunct genus ''[[Hypericum sect. Androsaemum|Androsaemum]]'', but was returned to ''Hypericum'' by Norman Robson in 1985.
'''''Hypericum hircinum''''', also known as '''stinking tutsan''',<ref name=BSBI07>{{BSBI 2007 |access-date=2014-10-17 }}</ref> is a shrubby [[flowering plant]] in the St. John's wort family [[Hypericaceae]].

The distribution of ''Hypericum hircinum'' is continuous across the Mediterranean and parts of the Middle East. It has also been cultivated throughout Europe, and has become naturalized in several places where it escaped captivity. The species is highly variable in appearance, but its lack of geographic separation means that five subspecies have been established to account for its diversity. ''H. hircinum'' has been [[Hybrid (biology)|hybridized]] with the closely related ''[[Hypericum androsaemum|H. androsaemum]]'' to produce the [[Fertility|fertile]] [[Hybrid name|nothospecies]] [[Hypericum × inodorum|''H''. × ''inodorum'']], which lacks the goat-like smell of ''H. hircinum''.

While ''Hypericum hircinum'' lacks the high concentrations of several [[Phytochemical|phytochemicals]] found in other members of its genus, it still has a highly active and useful chemical profile. Extracts from the species contain the highest levels of chemicals when taken from the flowering structures during the [[Fruit|fruiting]] period of the plant. They have been used in folk medicine to treat respiratory diseases by ingestion, and it is also applied topically to treat burns and muscle ailments. Modern evaluations have demonstrated the plant's effectiveness as an [[antioxidant]], anti-[[collagenase]] (for cosmetic care), and [[antimicrobial]] agent.


==Description==
==Description==
''Hypericum hircinum'' is a perennial shrub that usually grows 0.6–0.9 meters tall, but can reach heights of 1.5 meters. It is bushy in shape, with many stems.{{Sfn|Robson|1985|pp=308-309}}
The species grows to be 2 meters tall. It has a branching base with grey-brown bark. Its leaves give off a goat-like scent when crushed ([[caproic acid]]).<ref>Studies in the genus Hypericum L. (Guttiferae): 3. Sections: 1. Campylosporus to: 6a. Umbraculoides.</ref>


=== Vegetative structures ===
==Distribution==
The base of the plant does not have exposed roots, but does branch directly from the ground. The stems vary in their arrangement, and can grow straight up, reach outwards, or droop to the ground. They are flat when the plant is young, but become more cylindrical as it matures. Their bark has long grooves and is grey-brown in color. The distance of the stem between each leaf is 0.2–0.9 centimeters long.{{Sfn|Robson|1985|pp=308-309}}
''Hypericum hircinum'' is found around the [[Mediterranean]], specifically in [[France]], [[Spain]], [[Italy]], [[Greece]], [[Turkey]], and [[Saudi Arabia]]. The species was naturalized in [[Great Britain|Britain]] as early as 1620. The species does not spread rapidly into natural vegetation, and so has very little spread.<ref>[https://www.brc.ac.uk/plantatlas/index.php?q=plant/hypericum-hircinum Online Atlas of British and Irish Plants]</ref>


The leaves are directly connected to the stem without a stalk, and sometimes envelop the stem. Their blades are a wide triangular lance-like shape, with a rounded point. They are paler in color on their undersides, and have a leathery or papery texture. There are several pairs of main lateral veins on the blade, and the many small tertiary veins are visible from either side of the leaf. There are small glands on the leaf, which are most dense near the edges of the blade.{{Sfn|Robson|1985|pp=308-309}}
==Subspecies==

''Hypericum hircinum'' has five accepted subspecies:<ref>[http://hypericum.myspecies.info/taxonomy/term/615 Hypericum Myspecies Taxonomy]</ref>
=== Flowering structures ===
*[[Hypericum hircinum subsp. albimontanum|''H. hircinum'' subsp. ''albimontanum'']]
[[File:Bokkenkruid_(Hypericum_hircinum_L._Ascyrum_Cret._Vulgo)_Hortus_Botanicus_Leiden_(NL)2.jpg|left|thumb|''Hypericum hircinum'' in fruit]]
*[[Hypericum hircinum subsp. cambessedesii|''H. hircinum'' subsp. ''cambessedesii'']]
There are usually around three clusters of flowers on each plant with roughly twenty flowers each. The structure of the flower cluster is a wide pyramid. The flowers themselves are 2–4 cm in diameter, and are more or less globe-shaped when they are [[Flower bud|budding]]. The [[Sepal|sepals]] around the flowers vary in size, and are lance-like in shape. They grow larger during the flowering period, but fall off before the fruit ripens. They have glands both on their surface and more densely along their edges.{{Sfn|Robson|1985|pp=308-309}}
*[[Hypericum hircinum subsp. hircinum|''H. hircinum'' subsp. ''hircinum'']]

*[[Hypericum hircinum subsp. majus|''H. hircinum'' subsp. ''majus'']]
The flower [[Petal|petals]] are golden yellow, without any tint of red. They are 1.1–2.1 cm long and 0.4–0.9 cm wide, and they are roughly 3–4 times as large as the sepals. The stamens are bundled together in groups of around twenty, the longest of which are 1.2–2.2 cm long; this is conspicuously longer than the petals. The ovary is ellipse-shaped, and has styles 3–5 times its length that are upright. The seed capsule changes from green to a dull brown as the plant matures, and the seeds are an orange to reddish brown color.{{Sfn|Robson|1985|pp=308-309}}
*[[Hypericum hircinum subsp. metroi|''H. hircinum'' subsp. ''metroi'']]

== Chemistry ==
While most species of ''Hypericum'' contain high concentrations of the phytochemicals [[hypericin]] and [[pseudohypericin]], ''Hypericum hircinum'' may contain only trace amounts of them, and only in its flowers. However, it does contain other compounds that are uncommon within the genus, such as [[caffeic acid]] and [[amentoflavone]]. Other isolated compounds include [[Chlorogenic acid|chlorogenic]] and [[neochlorogenic acid]], small amounts of [[hyperforin]] and [[adhyperforin]], and [[2,4-Dihydroxybenzoic acid|dihydroxybenzoic acid]].{{Sfn|Odabas|Radusiene|Ivanauskas|Jakstas|2016|p=195-196}}

The concentration and composition of [[Essential oil|essential oils]] varies by the part of the plant. For example, the major [[Hydrocarbon|hydrocarbons]] in the leaves and flowers are [[Sesquiterpene|sesquiterpenes]], while those of the flowers are [[Monoterpene|monoterpenes]]. Some dominant compounds of the oil include [[guaiene]], [[selinene]], [[limonene]], and [[pinene]]. The most active and useful oil components have the highest concentrations when harvested while the plant is fruiting.{{Sfn|Maggi|Cecchini|Cresci|Coman|2010|p=3}}

== Taxonomy ==
[[File:Hortus_Eystettensis,_1640_(BHL_45339_265)_-_Classis_Aestiva_113.jpg|left|thumb|253x253px|The species in ''[[Hortus Eystettensis]]''. The inscriptions read (from left to right): Androsaemum foeditum seu Tragôdes, Siciliana, Hÿpericon.]]
The plant today known as ''Hypericum hircinum'' was well-studied and documented before the modern system of [[botanical nomenclature]] was established. For example, the plant may have been mentioned under the name "Ascyroides" as early as 1627 in [[Prospero Alpini|Prospero Alpini's]] ''[[De plantis exoticis]]''.{{Sfn|Turland|1995|pp=127, 146}} Additionally, it was depicted in the 1640 volume of ''[[Hortus Eystettensis]]'', a revolutionary book of [[Botanical illustration|botanical illustrations]]. In the codex, the plant was noted to be from [[Sicily]], of the grouping Hÿpericon (an early version of ''Hypericum''), and called "Androsaemum foeditum" or "Tragôdes".{{sfn|Besler|1640|p=265|ps=(folio 113)}} In [[Carl Linnaeus]]' early work ''Hortus Cliffortianus'', the name "Ascyroides" was [[Synonym (taxonomy)|synonymized]] with several others,{{Sfn|Turland|1995|p=146}} and Linnaeus gave the species the following polynomial [[Species description|description]]:{{Sfn|Linnaeus|1708|p=380}}{{Text and translation|Hypericum floribus trigynis, staminibus petalo longioribus caule fruticoso.|St John's wort with trigynous flowers on a bushy stem, and petals longer than the stamens.}}In his later work ''[[Species Plantarum]]'', which established the modern system of [[binomial nomenclature]] for plants, Linnaeus again described the species. This time, he did so using the currently accepted two-part name ''Hypericum hircinum''. He refined its short description, and noted several previous names for the plant, including variations of ''Androseamum foetidum'', ''Hypericum foetidum'', and ''Tragium''.{{Sfn|Linnaeus|1753|p=784}} After its original Linnaean treatment, there was only one time when a synonym for ''Hypericum hircinum'' arose. In 1836, [[Édouard Spach]] created the new genus ''Androsaemum'' (Androsème) out of several species from ''Hypericum''. The specific epithet ''hircinum'' was retained, creating the new combination ''Androsaemum hircinum'' for the species.{{Sfn|Spach|1836|p=419}}

The species was brought back to ''Hypericum'' by [[Norman Robson (botanist)|Norman Robson]] in his [[monograph]] of the genus. Robson did not recognize ''Androsaemum'' as a genus-level taxon and reduced it to one of 36 new [[Section (botany)|sections]] within ''Hypericum''. Thus, ''Hypericum hircinum'' was assigned to [[Hypericum sect. Androsaemum|''Hypericum'' sect. ''Androsaemum'']] in 1985.{{Sfn|Robson|1985|p=308}}

A 2013 study used [[Bayesian inference]] to establish the [[Phylogenetic tree|phylogeny]] and close relations of ''Hypericum'' species. Section ''Androsaemum'', including ''Hypericum hircinum,'' was placed into an "''Androsaemum''-group" with several other sections. The study also determined that ''H. hircinum'' was most closely related to ''[[Hypericum foliosum]]''.{{sfn|Meseguer|Aldasoro|Sanmartín|2013|p=386}}

=== Etymology ===
The [[genus]] name ''Hypericum'' is possibly derived from the [[Greek language|Greek]] words {{Lang|el|hyper}} (above) and {{Lang|el|eikon}} (picture), in reference to the tradition of hanging the plant over religious [[Icon|icons]] in the home.{{Sfn|Coombes|2012|p=172}} The specific epithet "hircinum" comes from the Latin word "[[wiktionary:hircinus|hircīnus]]" and refers to something that is like a male goat.{{Sfn|Charles|Marchant|1927|p=252}} Another term used by early botanists to name the species was "foetidum", which comes from the Latin word "[[wiktionary:foetidus|foetĭdus]]" and means a foul or [[fetid]] odor.{{Sfn|Charles|Marchant|1927|p=229}}

In the United Kingdom, the species is known as '''stinking tutsan'''.<ref>{{Cite web |title=UK Wildflowers - Hypericaceae - Hypericum Hircinum, Stinking Tutsan |url=https://www.uksouthwest.net/wildflowers/hypericaceae/hypericum-hircinum.html |access-date=2023-12-02 |website=www.uksouthwest.net}}</ref> Elsewhere, it is called '''goat St John's wort'''. Both are in reference to the distinctive goat-like and foul smell of the plant.''{{Sfn|Maggi|Cecchini|Cresci|Coman|2010|p=1}}''

===Subspecies===
While Carl Linnaeus did not denote any [[subspecies]] in his descriptions of ''Hypericum hircinum'', it was later determined that he described specimens of the subspecies ''obtusifolium'', which was later designated as the type subspecies ''hircinum''.{{Sfn|Robson|1985|p=310}} In addition to his specific description of ''Hypericum hircinum'', Norman Robson also streamlined its [[Infraspecific name|infraspecific]] organization. He determined that the ancestral form of the species could be found in the eastern Mediterranean and Saudi Arabia, while more modern subspecies are present on the Mediterranean islands, in Greece, and in Morocco. Because of the species' high variability in appearance but lack of geographic discontinuity, Robson acknowledged that there are at least five subspecies of ''H. hircinum''.{{Sfn|Robson|1985|p=176}}

''Hypericum hircinum'' subsp. ''hircinum'' is the type subspecies, and was previously named subsp. ''obstusifolium'' or described as vartiety ''minus'' or ''pumilum''. It is found on the islands of [[Sardinia]] and [[Corsica]], but has also been cultivated in Holland and Britain. Today, the taxon is considered to be rare. ''H. h.'' subsp. ''majus'' is the most common subspecies, found around the Mediterranean and [[Naturalisation (biology)|naturalized]] in Britain. It grows larger than other plants of the species, up to 1.5 meters tall. ''H. h.'' subsp. ''cambessedesii'' was once considered its own species in the defunct genus ''Androsaemum''. It has smaller leaves and flowers than the other subspecies. ''H. h.'' subsp. ''albimontanum'' is only found on Cyprus and several Greek islands, including Crete.<ref>{{Cite web |last=Bean |first=William Jackson |date=1916 |title=''Hypericum hircinum'' |url=https://www.treesandshrubsonline.org/articles/hypericum/hypericum-hircinum/ |access-date=2023-12-02 |website=Bean's Trees and Shrubs Online}}</ref> ''H. h.'' subsp. ''metroi'' is very similar in appearance to subsp. ''majus'', but lacks the distinct goat-like smell of the species.{{Sfn|Robson|1985|p=298}}

=== Hybridization ===
''Hypericum hircinum'' can be [[Hybrid (biology)|crossed]] with ''[[Hypericum androsaemum]]'' to produce the fertile hybrid ''[[Hypericum × inodorum]].'' This hybrid differs in appearance in that its sepals remain on the flower through its period of fruiting. It also lacks the goat-like smell of ''H. hircinum'', hence the specific epithet ''inodorum'' from the Latin word "[[wiktionary:inodorus|inodōrus]]" which indicates a lack of fragrance.<ref>{{Cite web |title=''Hypericum'' x ''inodorum'' (St. John's Wort, Tutsan) |url=https://plants.ces.ncsu.edu/plants/hypericum-x-inodorum/ |access-date=22 November 2023 |website=North Carolina Extension Gardener Plant Toolbox}}</ref>

== Ecology ==
Carl Linnaeus noted the presence ''Hypericum hircinum'' in Sicily, Calabria, and Crete;{{Sfn|Linnaeus|1753|p=784}} Édouard Spach stated in 1836 that it was found in "southern Europe and the East".{{Sfn|Spach|1836|p=419}} Today, the species is recorded as native in Saudi Arabia, Jordan, Syria, Cyprus, Turkey, Greece, Italy, France, Spain, and Morocco. It has also been introduced to the Balkans, Portugal, and the British Isles.<ref>{{Cite web |title=''Hypericum hircinum'' L. |url=http://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:433491-1 |access-date=2023-12-02 |website=Plants of the World Online |language=en}}</ref> Its habitat is in areas that are damp and shady along riversides, generally at elevations of 300–1200 meters.{{Sfn|Robson|1985|p=309}}

''Hypericum hircinum'' has been [[Horticulture|cultivated]] across much of Europe. The species is highly prone to escaping from this cultivation, and has become integrated into the native ecosystems of numerous areas, especially in Spain, France, and Sicily.{{Sfn|Robson|1985|p=309}}

== Uses ==
The use of ''Hypericum hircinum'' as an [[ornamental plant]] dates back to at least 1836.{{Sfn|Spach|1836|p=419}} The species is also frequently used in [[Traditional medicine|folk medicine]], especially for the treatment of respiratory diseases. For example, it is used in Italy to treat persistent colds, [[asthma]], and coughing. The plant is also used [[Topical medication|topically]], with its oil applied to skin burns and its extracts used to relieve [[rheumatism]] and other muscular ailments.{{Sfn|Mandrone|Lorenzi|Venditti|Guarcini|2015|p=403}}

Formal scientific studies have confirmed some of ''Hypericum hircinum''<nowiki/>'s medicinal properties. It has greater [[antioxidant]] capabilities than the well-studied ''[[Camellia sinensis]]'', due largely to the presence of [[flavonols]], [[Flavanone|flavanones]], and [[Caffeoylquinic acid|caffeoylquinic acids]]. It also contains anti-[[collagenase]] components, which could lead to future use in [[Cosmetics|cosmetic products]].{{Sfn|Mandrone|Lorenzi|Venditti|Guarcini|2015|p=406-408}} The [[antimicrobial]] properties of the species have also been validated, and it is particularly effective against various ''[[Candida (fungus)|Candida]]'' fungus species and the bacterium ''[[Streptococcus mutans]].{{Sfn|Maggi|Cecchini|Cresci|Coman|2010|p=3}}''{{Sfn|Tocci|Perenzoni|Iamonico|Fava|2018|p=5}} ''H. hircinum'' is also effective against other species of fungi, and lacks the [[Cytotoxicity|cytotoxic]] effects in humans that other [[antifungal]] treatments may cause.{{Sfn|Tocci|Perenzoni|Iamonico|Fava|2018|p=8}}


==References==
==References==
<references group="" responsive="1"></references>
{{Reflist}}


=== Bibliography ===
{{cladogram|{{Clade|style=font-size:70%;line-height:75%;width:300px;
|label1="''Androsaemum''-group"
|1={{clade
|1=[[Hypericum sect. Arthrophyllum|sect. ''Arthrophyllum'']]
|2=[[Hypericum sect. Triadenoides|sect. ''Triadenioides'']]
|3={{clade
|label1=sect. ''Webbia''
|1=''[[Hypericum canariense|H. canariense]]''
|2={{clade
|label1=sect. ''Bupleuroides''
|1=''[[Hypericum bupleuroides|H. bupleuroides]]''
|label2= [[Hypericum sect. Androsaemum|sect. ''Androsaemum'']]
|2={{clade
|1=''[[Hypericum androsaemum|H. androsaemum]]''
|2={{clade
|1={{clade
|1='''''H. hircinum'''''
|2=''[[Hypericum foliosum|H. foliosum]]''
}}
|2=''[[Hypericum grandifolium|H. grandifolium]]''
}}
}}
}}
}}
}}
}}|caption=[[Cladogram]] showing the [[phylogeny]] and relationships of ''H.&nbsp;hircinum'' within the "''Androsaemum''-group" based on a 2013 study{{sfn|Meseguer|Aldasoro|Sanmartín|2013|p=386}}|align=}}

* {{Cite book |last=Besler |first=Basilius |url=https://www.biodiversitylibrary.org/item/98364#page/265/mode/1up |title=Hortus Eystettensis |date=1640 |publisher= |location=Nürnberg |doi=10.5962/bhl.title.45339}}
* {{Cite book |last=Charles |first=Joseph |url=https://catalog.hathitrust.org/Record/009163191 |title=Cassell's Latin dictionary |last2=Marchant |first2=J.R.V. |publisher=Funk & Wagnalls |year=1927}}
* {{cite book |last1=Coombes |first1=Allen J. |title=The A to Z of plant names : a quick reference guide to 4000 garden plants |date=2012 |publisher=Timber Press, Inc. |isbn=978-1-60469-196-2 |location=Portland, Oregon |page=}}
* {{Cite book |last=Linnaeus |first=Carl |url=https://www.biodiversitylibrary.org/item/13838 |title=Hortus Cliffortianus |year=1708 |location=Amsterdam |language=la |doi=10.5962/bhl.title.690}}
* {{Cite book |last=Linnaeus |first=Carl |url=https://www.biodiversitylibrary.org/item/13830 |title=Species Plantarum |year=1753 |edition=2 |location=Stockholm |language=la |doi=10.5962/bhl.title.669}}
* {{Cite journal |last=Maggi |first=Filippo |last2=Cecchini |first2=Cinzia |last3=Cresci |first3=Alberto |last4=Coman |first4=Maria |date=2010 |title=Chemical composition and antimicrobial activity of ''Hypericum hircinum'' L. subsp. ''majus'' essential oil |url=https://www.researchgate.net/publication/229070049_Chemical_composition_and_antimicrobial_activity_of_Hypericum_hircinum_L_Subsp_majus_essential_oil |journal=Chemistry of Natural Compounds |volume=46 |issue=1 |doi=10.1007/s10600-010-9545-1 |via=ResearchGate}}
* {{Cite journal |last=Mandrone |first=Manuela |last2=Beatrice |first2=Lorenzi |last3=Venditti |first3=Alessandro |last4=Guarcini |first4=Laura |date=2015 |title=Antioxidant and anti-collagenase activity of ''Hypericum hircinum'' L. |url=https://journals.scholarsportal.info/details/09266690/v76icomplete/402_aaaaohhl.xml |journal=Industrial Crops and Products |volume=76}}
* {{Cite journal |last=Meseguer |first=Andrea |last2=Aldasoro |first2=Juan |last3=Sanmartín |first3=Isabel |date=2013 |title=Bayesian inference of phylogeny, morphology and range evolution reveals a complex evolutionary history in St. John’s wort (''Hypericum'') |url=https://digital.csic.es/bitstream/10261/167045/1/Bayesian%20inference%20of%20phylogeny.pdf |journal=Molecular Phylogenetics and Evolution |volume=67 |pages=379-403 |via=Elsevier}}
* {{Cite journal |last=Odabas |first=Mehmet |last2=Radusiene |first2=Jolita |last3=Ivanauskas |first3=Liudas |last4=Jakstas |first4=Valdas |date=2016 |title=Secondary metabolites in ''Hypericum'' species and their distribution in different plant parts |journal=Zemdirbyste-Agriculture |volume=1-3 |issue=2 |doi=10.13080/z-a.2016.103.025}}
* {{Cite journal |last=Robson |first=Norman |date=1985 |title=Studies in the genus ''Hypericum'' {{small|L.}} (Guttiferae). '''3'''. Sections 1. ''Campylosporus'' to 6a. ''Umbraculoides'' |url=https://www.biodiversitylibrary.org/item/127083 |journal=Bulletin of the British Museum (Natural History) Botany |volume=12 |pages=163-325 |via=Biodiversity Heritage Library}}
* {{Cite book |last=Spach |first=Édouard |url=https://www.biodiversitylibrary.org/bibliography/44839 |title=Histoire naturelle des végétaux |publisher=Librairie encyclopédique de Roret |year=1836 |edition=5th |location=Paris |language=fr |doi=10.5962/bhl.title.44839}}
* {{Cite journal |last=Tocci |first=Noemi |last2=Perenzoni |first2=Daniele |last3=Iamonico |first3=Duilio |last4=Fava |first4=Francesca |date=2018 |title=Extracts From ''Hypericum hircinum'' subsp. ''majus'' Exert Antifungal Activity Against a Panel of Sensitive and Drug-Resistant Clinical Strains |url=https://www.frontiersin.org/articles/10.3389/fphar.2018.00382/full |journal=Frontiers in Ethnopharmacology |volume=9 |doi=10.3389/fphar.2018.00382}}
* {{Cite journal |last=Turland |first=Nicholas |date=1995 |title=Linnaeus's interpretation of Prospero Alpino's ''De plantis exoticis'', with special emphasis on the flora of Crete |url=https://ia802507.us.archive.org/22/items/biostor-265894/biostor-265894.pdf |journal=Bull. not. Hist. Mus. Land. (Bot.) |volume=25 |issue=2 |pages=127-159 |via=Internet Archive}}
{{Taxonbar|from=Q889234}}
{{Taxonbar|from=Q889234}}


{{Hypericum species Navbox}}
{{Hypericum species Navbox}}
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[[Category:Hypericum|hircinum]]
[[Category:Hypericum|hircinum]]
[[Category:Plants described in 1753]]
[[Category:Plants described in 1753]]
[[Category:Taxa named by Carl Linnaeus]]
[[Category:Taxa named by Carl Linnaeus]]
{{Hypericum-stub}}

Revision as of 16:36, 2 December 2023

Hypericum hircinum
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Malpighiales
Family: Hypericaceae
Genus: Hypericum
Section: Hypericum sect. Androsaemum
Species:
H. hircinum
Binomial name
Hypericum hircinum
Synonyms
  • Androsaemum hircinum

Hypericum hircinum is a species of perennial flowering plant in the St John's wort family, Hypericaceae. It is known as goat St John's wort and stinking tutsan; both names refer to the plant's distinctive odor. The species is a bushy shrub that can grow up to 1.5 meters tall, is many-stemmed, and has golden yellow flowers with conspicuous stamens. The plant has been well-documented in botanical literature, with mentions dating back to at least 1627. Carl Linneaus described H. hircinum several times, including in his 1753 Species Plantarum which established its binomial. At one point the plant was placed into the defunct genus Androsaemum, but was returned to Hypericum by Norman Robson in 1985.

The distribution of Hypericum hircinum is continuous across the Mediterranean and parts of the Middle East. It has also been cultivated throughout Europe, and has become naturalized in several places where it escaped captivity. The species is highly variable in appearance, but its lack of geographic separation means that five subspecies have been established to account for its diversity. H. hircinum has been hybridized with the closely related H. androsaemum to produce the fertile nothospecies H. × inodorum, which lacks the goat-like smell of H. hircinum.

While Hypericum hircinum lacks the high concentrations of several phytochemicals found in other members of its genus, it still has a highly active and useful chemical profile. Extracts from the species contain the highest levels of chemicals when taken from the flowering structures during the fruiting period of the plant. They have been used in folk medicine to treat respiratory diseases by ingestion, and it is also applied topically to treat burns and muscle ailments. Modern evaluations have demonstrated the plant's effectiveness as an antioxidant, anti-collagenase (for cosmetic care), and antimicrobial agent.

Description

Hypericum hircinum is a perennial shrub that usually grows 0.6–0.9 meters tall, but can reach heights of 1.5 meters. It is bushy in shape, with many stems.[1]

Vegetative structures

The base of the plant does not have exposed roots, but does branch directly from the ground. The stems vary in their arrangement, and can grow straight up, reach outwards, or droop to the ground. They are flat when the plant is young, but become more cylindrical as it matures. Their bark has long grooves and is grey-brown in color. The distance of the stem between each leaf is 0.2–0.9 centimeters long.[1]

The leaves are directly connected to the stem without a stalk, and sometimes envelop the stem. Their blades are a wide triangular lance-like shape, with a rounded point. They are paler in color on their undersides, and have a leathery or papery texture. There are several pairs of main lateral veins on the blade, and the many small tertiary veins are visible from either side of the leaf. There are small glands on the leaf, which are most dense near the edges of the blade.[1]

Flowering structures

Hypericum hircinum in fruit

There are usually around three clusters of flowers on each plant with roughly twenty flowers each. The structure of the flower cluster is a wide pyramid. The flowers themselves are 2–4 cm in diameter, and are more or less globe-shaped when they are budding. The sepals around the flowers vary in size, and are lance-like in shape. They grow larger during the flowering period, but fall off before the fruit ripens. They have glands both on their surface and more densely along their edges.[1]

The flower petals are golden yellow, without any tint of red. They are 1.1–2.1 cm long and 0.4–0.9 cm wide, and they are roughly 3–4 times as large as the sepals. The stamens are bundled together in groups of around twenty, the longest of which are 1.2–2.2 cm long; this is conspicuously longer than the petals. The ovary is ellipse-shaped, and has styles 3–5 times its length that are upright. The seed capsule changes from green to a dull brown as the plant matures, and the seeds are an orange to reddish brown color.[1]

Chemistry

While most species of Hypericum contain high concentrations of the phytochemicals hypericin and pseudohypericin, Hypericum hircinum may contain only trace amounts of them, and only in its flowers. However, it does contain other compounds that are uncommon within the genus, such as caffeic acid and amentoflavone. Other isolated compounds include chlorogenic and neochlorogenic acid, small amounts of hyperforin and adhyperforin, and dihydroxybenzoic acid.[2]

The concentration and composition of essential oils varies by the part of the plant. For example, the major hydrocarbons in the leaves and flowers are sesquiterpenes, while those of the flowers are monoterpenes. Some dominant compounds of the oil include guaiene, selinene, limonene, and pinene. The most active and useful oil components have the highest concentrations when harvested while the plant is fruiting.[3]

Taxonomy

The species in Hortus Eystettensis. The inscriptions read (from left to right): Androsaemum foeditum seu Tragôdes, Siciliana, Hÿpericon.

The plant today known as Hypericum hircinum was well-studied and documented before the modern system of botanical nomenclature was established. For example, the plant may have been mentioned under the name "Ascyroides" as early as 1627 in Prospero Alpini's De plantis exoticis.[4] Additionally, it was depicted in the 1640 volume of Hortus Eystettensis, a revolutionary book of botanical illustrations. In the codex, the plant was noted to be from Sicily, of the grouping Hÿpericon (an early version of Hypericum), and called "Androsaemum foeditum" or "Tragôdes".[5] In Carl Linnaeus' early work Hortus Cliffortianus, the name "Ascyroides" was synonymized with several others,[6] and Linnaeus gave the species the following polynomial description:[7]

Hypericum floribus trigynis, staminibus petalo longioribus caule fruticoso.
St John's wort with trigynous flowers on a bushy stem, and petals longer than the stamens.

In his later work Species Plantarum, which established the modern system of binomial nomenclature for plants, Linnaeus again described the species. This time, he did so using the currently accepted two-part name Hypericum hircinum. He refined its short description, and noted several previous names for the plant, including variations of Androseamum foetidum, Hypericum foetidum, and Tragium.[8] After its original Linnaean treatment, there was only one time when a synonym for Hypericum hircinum arose. In 1836, Édouard Spach created the new genus Androsaemum (Androsème) out of several species from Hypericum. The specific epithet hircinum was retained, creating the new combination Androsaemum hircinum for the species.[9]

The species was brought back to Hypericum by Norman Robson in his monograph of the genus. Robson did not recognize Androsaemum as a genus-level taxon and reduced it to one of 36 new sections within Hypericum. Thus, Hypericum hircinum was assigned to Hypericum sect. Androsaemum in 1985.[10]

A 2013 study used Bayesian inference to establish the phylogeny and close relations of Hypericum species. Section Androsaemum, including Hypericum hircinum, was placed into an "Androsaemum-group" with several other sections. The study also determined that H. hircinum was most closely related to Hypericum foliosum.[11]

Etymology

The genus name Hypericum is possibly derived from the Greek words hyper (above) and eikon (picture), in reference to the tradition of hanging the plant over religious icons in the home.[12] The specific epithet "hircinum" comes from the Latin word "hircīnus" and refers to something that is like a male goat.[13] Another term used by early botanists to name the species was "foetidum", which comes from the Latin word "foetĭdus" and means a foul or fetid odor.[14]

In the United Kingdom, the species is known as stinking tutsan.[15] Elsewhere, it is called goat St John's wort. Both are in reference to the distinctive goat-like and foul smell of the plant.[16]

Subspecies

While Carl Linnaeus did not denote any subspecies in his descriptions of Hypericum hircinum, it was later determined that he described specimens of the subspecies obtusifolium, which was later designated as the type subspecies hircinum.[17] In addition to his specific description of Hypericum hircinum, Norman Robson also streamlined its infraspecific organization. He determined that the ancestral form of the species could be found in the eastern Mediterranean and Saudi Arabia, while more modern subspecies are present on the Mediterranean islands, in Greece, and in Morocco. Because of the species' high variability in appearance but lack of geographic discontinuity, Robson acknowledged that there are at least five subspecies of H. hircinum.[18]

Hypericum hircinum subsp. hircinum is the type subspecies, and was previously named subsp. obstusifolium or described as vartiety minus or pumilum. It is found on the islands of Sardinia and Corsica, but has also been cultivated in Holland and Britain. Today, the taxon is considered to be rare. H. h. subsp. majus is the most common subspecies, found around the Mediterranean and naturalized in Britain. It grows larger than other plants of the species, up to 1.5 meters tall. H. h. subsp. cambessedesii was once considered its own species in the defunct genus Androsaemum. It has smaller leaves and flowers than the other subspecies. H. h. subsp. albimontanum is only found on Cyprus and several Greek islands, including Crete.[19] H. h. subsp. metroi is very similar in appearance to subsp. majus, but lacks the distinct goat-like smell of the species.[20]

Hybridization

Hypericum hircinum can be crossed with Hypericum androsaemum to produce the fertile hybrid Hypericum × inodorum. This hybrid differs in appearance in that its sepals remain on the flower through its period of fruiting. It also lacks the goat-like smell of H. hircinum, hence the specific epithet inodorum from the Latin word "inodōrus" which indicates a lack of fragrance.[21]

Ecology

Carl Linnaeus noted the presence Hypericum hircinum in Sicily, Calabria, and Crete;[8] Édouard Spach stated in 1836 that it was found in "southern Europe and the East".[9] Today, the species is recorded as native in Saudi Arabia, Jordan, Syria, Cyprus, Turkey, Greece, Italy, France, Spain, and Morocco. It has also been introduced to the Balkans, Portugal, and the British Isles.[22] Its habitat is in areas that are damp and shady along riversides, generally at elevations of 300–1200 meters.[23]

Hypericum hircinum has been cultivated across much of Europe. The species is highly prone to escaping from this cultivation, and has become integrated into the native ecosystems of numerous areas, especially in Spain, France, and Sicily.[23]

Uses

The use of Hypericum hircinum as an ornamental plant dates back to at least 1836.[9] The species is also frequently used in folk medicine, especially for the treatment of respiratory diseases. For example, it is used in Italy to treat persistent colds, asthma, and coughing. The plant is also used topically, with its oil applied to skin burns and its extracts used to relieve rheumatism and other muscular ailments.[24]

Formal scientific studies have confirmed some of Hypericum hircinum's medicinal properties. It has greater antioxidant capabilities than the well-studied Camellia sinensis, due largely to the presence of flavonols, flavanones, and caffeoylquinic acids. It also contains anti-collagenase components, which could lead to future use in cosmetic products.[25] The antimicrobial properties of the species have also been validated, and it is particularly effective against various Candida fungus species and the bacterium Streptococcus mutans.[3][26] H. hircinum is also effective against other species of fungi, and lacks the cytotoxic effects in humans that other antifungal treatments may cause.[27]

References

  1. ^ a b c d e Robson 1985, pp. 308–309.
  2. ^ Odabas et al. 2016, p. 195-196.
  3. ^ a b Maggi et al. 2010, p. 3.
  4. ^ Turland 1995, pp. 127, 146.
  5. ^ Besler 1640, p. 265(folio 113)
  6. ^ Turland 1995, p. 146.
  7. ^ Linnaeus 1708, p. 380.
  8. ^ a b Linnaeus 1753, p. 784.
  9. ^ a b c Spach 1836, p. 419.
  10. ^ Robson 1985, p. 308.
  11. ^ Meseguer, Aldasoro & Sanmartín 2013, p. 386.
  12. ^ Coombes 2012, p. 172.
  13. ^ Charles & Marchant 1927, p. 252.
  14. ^ Charles & Marchant 1927, p. 229.
  15. ^ "UK Wildflowers - Hypericaceae - Hypericum Hircinum, Stinking Tutsan". www.uksouthwest.net. Retrieved 2023-12-02.
  16. ^ Maggi et al. 2010, p. 1.
  17. ^ Robson 1985, p. 310.
  18. ^ Robson 1985, p. 176.
  19. ^ Bean, William Jackson (1916). "Hypericum hircinum". Bean's Trees and Shrubs Online. Retrieved 2023-12-02.
  20. ^ Robson 1985, p. 298.
  21. ^ "Hypericum x inodorum (St. John's Wort, Tutsan)". North Carolina Extension Gardener Plant Toolbox. Retrieved 22 November 2023.
  22. ^ "Hypericum hircinum L." Plants of the World Online. Retrieved 2023-12-02.
  23. ^ a b Robson 1985, p. 309.
  24. ^ Mandrone et al. 2015, p. 403.
  25. ^ Mandrone et al. 2015, p. 406-408.
  26. ^ Tocci et al. 2018, p. 5.
  27. ^ Tocci et al. 2018, p. 8.

Bibliography

"Androsaemum‑group"
Cladogram showing the phylogeny and relationships of H. hircinum within the "Androsaemum-group" based on a 2013 study[1]
  1. ^ Meseguer, Aldasoro & Sanmartín 2013, p. 386.
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