Hypericum Aegypticum

''Hypericum aegypticum'' is a species of flowering plant of the St. John's wort family (Hypericaceae) which is native to the Eastern Mediterranean. It was described by Carl Linnaeus in the second volume of his ''Species Plantarum'' in 1753, who named it after Egypt despite it not being distributed there. The plant is commonly known as shrubby St. John's wort or Egyptian St. John's wort in English. Like other members of section '' Adenotrias'', it is found among limestone rocks in coastal areas. While it has been evaluated as threatened on the island of Malta, the species has no legal protections.

The species is a shrub or shrublet that grows up to 2 meters (6.5 feet) tall with many branches, each having a single small flower with five yellow petals and many stamens. Its leaves lack hairs, and have densely patterned glands and large primary veins. The species is notable for its heterostyly, a trait which within the ''Hypericum'' genus is unique to section ''Adenotrias'', and exhibits one of two heterostylous flower types called "pins" and "thrums". It has three subspecies, which vary in their distribution and phytochemical makeup.

Description

''Hypericum aegypticum'' is a shrub or shrublet that varies in height from 0.05 to 2 meters (0.16 to 6.5 feet) tall. It grows in a crowded, spreading pattern, with its stems stretching out from the center of the plant. The branches grow roughly perpendicular from these stems, but can also spread out in different directions. It has a chromosome number of 2n=20.

Stems

The stems of ''Hypericum aegypticum'' usually have two visible lines running along their length, but can sometimes have four. When the plant is young, the stems are a flatter shape with two edges ( ancipitous), but quickly become more cylindrical as the plant matures. In a cross-section, the profile of the stems is circular and lacks wing-like extensions. They have a vessel density of roughly two vessels per square millimeter of stem, with each of these vessels measuring 15 micrometers in diameter. In between the nodes where the leaves branch from (in the internodes), there are small leaf-like structures which are shorter than the true leaves.

The xylem in the stems, a type of transport tissue, differs from other similar species of ''Hypericum.'' The xylem of ''H. aegypticum'' is diffuse; it is spread evenly throughout the stem. In other similar species, the xylem is ring-porous and more dense in a series of concentric rings. There are two secretory structures in the stems: small secretory canals in the phloem and many large secretory canals in the cortex.

Leaves

The leaves of ''Hypericum aegypticum'' are free and range from sessile to shortly petiolate. They persist for around two seasons. The blades are long and wide. Their shape is elliptic or narrowly oblong. The leaves are the same color as the rest of the plant and have a texture similar to that of leather. The shape of their tip can range from acute to obtuse and the base of the leaf is wedge-shaped. The leaves lack hairs (trichomes) on both the top and bottom.

They have one pair of lateral veins, with the midrib sometimes branched and visible beneath them. Their primary veins are very large and straight and are highly rigid. The secondary veins branch off at acute angles of wide divergence, and some tertiary veins with a branched pattern are also present.

The laminar and marginal glands are dense. Unlike some species of ''Hypericum'', such as '' H. perforatum'', ''H. aegypticum'' lacks black nodules on its leaves. Instead, it has small translucent glands which measure 54 by 45 micrometers and are found in the mesophyll.

The upper sides of the leaves have an outer layer (epidermis) made of cells with linear walls and rounded corners. This upper side also has stomata that are surrounded by smaller cells. The lower side of the leaves have a similar composition to the upper side, but with a higher density of stomata. On both sides, there is a cuticle that extends over the entire surface, even over the guard cells, which makes them very hard to distinguish. This thick cuticle leads to increased impermeability in areas where there are encrusted deposits.

Inflorescence

''Hypericum aegypticum'' is one-flowered, and its inflorescence is terminal and grows from around four axils below. It has short flowering branches past a "sterile" region which lacks them. These branches grow from around eleven different nodes. However, some plants lack the sterile region and can have flowers or flowering branches from up to twenty-two nodes. The inflorescence is similar in appearance to a spike, with longer branches below the flowering region. The bracts are often absent, but when they are present they are leaf-like in appearance, with a clasping calyx and several types of glands. The pedicels are very short or absent altogether.

The flowers of ''H. aegypticum'' range from in diameter. The shape of the buds is a narrow form between an oval and an ellipse which tapers to a rounded point. The sepals are green and are by in size. They overlap one another and are not quite the same size. Their shape ranges from oblong to elliptic, but they can also be lance-like in form. Their ends can be rounded, pointed, or obtuse. The sepals are hood-like and stand stiffly erect, and have roughly nine unbranched veins with prominent midribs and linear laminar glands. The petals of the flower range from bright yellow to a rather pale shade of yellow. They are by in size. They are persistent in fruit and there are roughly twice as many of them as there are sepals.

There are 18–48 stamens per inflorescence, with single fascicles containing 5–10 stamens and double fascicles containing 5–29 stamens. There are united filaments in each fascicle, the longest of which are in long-styled individuals and in short-styled individuals. Some stamen fascicles are sterile and are long and flat-topped. The ovaries are by in size and are narrow or very narrow and acute to truncate.

The seed capsule is by in size. Its shape is that of a cylindric ellipsoid to a rather broad ellipsoid. It is longer than the sepals, and its valves run longitudinally and have oil tubes. The seeds are dark brown and roughly long and lack a longitudinal ridge. The testa have small linear pits or wrinkles, and the elaiosome is often lobed.

Heterostyly

Individual plants of ''Hypericum aegypticum'' exhibit one of two different sets of characteristics, called distyly. This polymorphism is characterized by two different types of flowers: one has long styles and short stamens (such specimens are called "pins"), the other has short styles and long stamens (such specimens are called "thrums"). In this species, pin flowers produce and receive more pollen grains than thrum flowers. This is unusual, as it does not fit with the expected course of natural selection; plants in the species which produce and receive more pollen would be expected to outcompete those that produce and receive less. While pin flowers are more common in the wild with a ratio of approximately 3 pins to 1 thrum, that difference is only because the gene which codes for the pin is dominant while the thrum is recessive, and the 3:1 ratio is consistent with traits that have no effect on reproduction.

''Hypericum'' sect. ''Adenotrias'' is the only section in ''Hypericum'' that exhibits heterostyly, and all of its species, including ''Hypericum aegypticum'', are distylous. This trait is thought to have evolved within the genus as opposed to being primitive and is one of the unique characteristics of sect. ''Adenotrias''. In studies of eleven angiosperm genera which exhibited distyly, all but ''Hypericum'' followed a genetic system called the ''Primula'' system, in which the "thrum" form was dominant and the "pin" form was recessive. However, for an unknown reason, the opposite appears to be the case for the species of ''Hypericum''.

Phytochemistry

The phytochemistry of ''Hypericum aegypticum'' varies a great deal depending on which subspecies is analyzed. Subspecies ''aegypticum'', ''maroccanum'', and ''webii'' are all highly different in their major essential oil components, with only three constituents being found in common in all of the subspecies: spathulenol, caryophyllene oxide, and ''E''-caryophellene. The major constituent of subsp. ''aegypticum'' is ishwarane (a sesquiterpene) the major constituent of subsp. ''maroccanum'' is caryophyllene, and the major constituent of subsp. ''webii'' is pinene. These differences in the compounds detected in the species reveal that morphologically different populations of ''H. aegypticum'' are also likely to differ phytochemically. ''S''mall amounts of other metabolites are also present in ''H. aegypticum'', including flavonoids (3.95 g/kg), biflavones (1.45 g/kg), acylphloroglucinols (1.15 g/kg), cinnamic acids (0.35 g/kg), and naphtodianthrones (0.05 g/kg).

Essential oil extracts from ''Hypericum aegypticum'' subsp. ''webbii'' were shown ''in vitro'' to have moderate antimicrobial effects against Gram-positive bateria such as ''Bacillus subtilis'', ''Enterococcus faecalis'', ''Staphylococcus epidermidis'', and ''Micrococcus luteus''.

Distribution and habitat

''Hypericum aegypticum'' can be found frequently in the wild and is indigenous to Malta, Greece, Morocco, Algeria, and Libya. However, its distribution across these countries is unusual because it is highly fragmented into several different population strands. The three subspecies are divided among these strands: ''H. aegypticum'' subsp. ''aegypticum'' is native to Libya; subsp. ''maroccanum'' is native to Morocco and Algeria; subsp. ''webbii'' is native to Sardinia, Sicily, Malta, and Greece.

''H. aegypticum'' is found among limestone rocks and scree in valleys of coastal areas. It is found at elevations from sea level to 1600 meters (5250 feet) above sea level. In some habitats, particularly more wind-swept or exposed areas, the species has been observed in much denser, carpet-like growth patterns, sometimes growing to only tall. However, the species grows best in open, sunny areas free of other plants which could compete with it.

While ''H. aegypticum'' has been identified by the Maltese Red Data Book as having a threatened status on Malta, no legal action has been taken for the protection of the species as of 2021.

Cultivation

While ''Hypericum aegypticum'' is not widely cultivated, it is present in botanical gardens, including in a greenhouse at the University of Oxford Botanic Garden.

Taxonomy

''Hypericum aegypticum'' was first described by Carl Linnaeus in his second volume of ''Species Plantarum'' in 1753. He noted the species' Egyptian habitat, its often paired leaves, growing pattern, and the shape of its leaves. However, even though Linnaeus mentioned "Aegypto" and claimed that his type specimen for the species was from Egypt, ''H. aegypticum'' is not actually native to Egypt, and the specimen Linnaeus used most likely came from one of the Libyan populations. Today, ''H. aegypticum'' is commonly referred to in English as Shrubby St. John's Wort or Egyptian St. John's Wort, while in Malta it is known as Fexfiex tal-irdum.

The species' placement within ''Hypericum'' can be summarized as follows:

''Hypericum''
: ''Hypericum'' subg. ''Hypericum''
:: ''Hypericum'' sect. ''Adenotrias''
::: '' Hypericum russeggeri''
::: '' Hypericum aciferum''
::: ''Hypericum aegypticum''

History

After the original description by Linnaeus, various botanists identified the plant under different names which are now considered invalid synonyms of the species. Michel Adanson, in his 1763 work establishing the genus ''Elodes'', erroneously described the species '' Hypericum elodes'' by relying upon a supposed description by Linneaus of the species in his 1753 ''Species Plantarum''. However, the name ''H. elodes'' did not appear it was mentioned in the 1762 '' Flora Anglica'', and the name that Adanson actually referenced was ''Hypericum aegypticum''. Therefore, his description under ''H. elodes'' is considered a synonym of ''H. aegypticum''. Confusion around ''Elodes'' continued, but it was eventually restricted to only the species ''H. elodes'' as defined by Linneaus. In the early 19th century, the species ''Hypericum'' were being divided into numerous smaller, new genera by different authors. Antoine Laurent de Jussieu divided it into five genera, confusingly including ''Elodea.'' Édouard Spach contributed his own restrictive genera, and placed ''H. aegypticum'' first into ''Triadenia'' and then ''Adenotrias''. Finally, Robert Keller reduced all of these new genera to sections within ''Hypericum'' in 1925, bringing ''H. aegypticum'' back into ''Hypericum'' in section ''Adenotrias''.

In 1996, British taxonomist Norman Robson published the sixth volume of his comprehensive monograph on the genus ''Hypericum.'' In this volume, Robson recognized Linnaeus' original name as correct and maintained ''H. aegypticum'' in sect. ''Adenotrias'', incorporating it into his new framework of 36 sections. ''Adenotrias'' was established to comprise three species of shrubs found among limestone rocks which vary greatly in appearance. However, the species of the section were united because they shared the trait of heterostyly.

Phylogeny

Molecular phylogenetics has demonstrated that ''Hypericum aegypticum'' is most closely related to '' H. balfourii'' and '' H. socotranum'' of sect. ''Campylosporus''''.'' However, no conclusive evidence was found which would suggest that these similarities would warrant the inclusion of ''H. aegypticum'' in sect. ''Campylosporus.'' Phylogenetic studies have also found that '' H. heterophyllum'', despite lacking the floral specializations and heterostyly of ''H. aegypticum'', has similar characteristics to the species, especially ''H. aegypticum'' subsp. ''maroccanum''. This suggests that ''H. heterophyllum'' is a derivative of the species which has lost some of the specializations that are unique to ''H. aegypticum'' and sect. ''Adenotrias''.

Subdivision

The following subspecies are accepted by Plants of the World Online:

* ''Hypericum aegypticum'' subsp. ''aegypticum''
* ''Hypericum aegypticum'' subsp.'' maroccanum'' (Pau) N. Robson
* ''Hypericum aegypticum'' subsp.'' webbii'' (Spach) N. Robson
Beyond their separate distributions, the subspecies also vary slightly in appearance. Subsp. ''maroccanum'' grows more upright than its counterparts, subsp. ''webbii'' is typically taller and larger, and subsp. ''aegypticum'' has petals that are somewhat shorter. Additionally, the branches of ''maroccanum'' tend to reach upwards, while ''aegypticum'' usually forms a lower bush. Lastly, the leaves of ''maroccanum'' are markedly larger than ''aegypticum'' or ''webbii''.

Subspecies ''maroccanum'' was originally described as a variety of ''H. aegypticum'' by Carlos Pau in 1932, while subsp. ''webbii'' was originally considered to be its own species under the name ''Triadenia webbii'' by Édouard Spach in 1836. Norman Robson reclassified both as subspecies of ''H. aegypticum'' in 1993, and established subsp. ''aegypticum'' to denote those specimens which were separate from both ''maroccanum'' and ''webbii''.

References

{{Taxonbar, from=Q16653686

aegypticum
Flora of Malta
Flora of Libya
Flora of Greece
Flora of Morocco
Flora of Algeria
Taxa named by Carl Linnaeus
Plants described in 1753