Peganum harmala


Peganum harmala, commonly called wild rue, Syrian rue, African rue, esfand or espand, or harmel, is a perennial, herbaceous plant, with a woody underground rootstock, of the family Nitrariaceae, usually growing in saline soils in temperate desert and Mediterranean regions. Its common English-language name came about because of a resemblance to rue. Its seeds contain a high concentration of diverse beta-carboline alkaloids.
It has deep roots and a strong smell, finely divided leaves, white flowers rich in alkaloids, and small seed capsules containing numerous dark, oily seeds. It is native to a vast region across North Africa, southern and eastern Europe, the Middle East, Central Asia, and parts of South and East Asia, and has been introduced to countries like South Africa, Mexico, France. It grows in dry, often saline or disturbed habitats, thriving from sea level to high elevations, is pollinated mainly by insects, disperses seeds mostly by dispersal vectors or human activity, and hosts a specialized beetle proposed for its biological control.
Some scholars have associated it with the sacred plant called soma or haoma in ancient Indo-Iranian texts and it has been described under various names by classical and medieval sources, with archaeological evidence suggesting its ritual use dating back to at least the 2nd century BCE. It was first described and illustrated in the 16th century by Rembert Dodoens and later classified by botanists such as Gaspard Bauhin and Carl Linnaeus. It has several recognized varieties distinguished by morphological traits and geographic distribution, with lectotype designations refined over time to clarify its taxonomy.
In the United States, it is banned or regulated as a noxious weed in several states requiring eradication, while internationally, possession and sale of the plant or its psychoactive alkaloids are illegal or controlled in several countries, including France, Finland, Canada, and Australia. It is used as a dye, incense, and in traditional medicine. It is also toxic to livestock and difficult to eradicate.

Etymology

Rue for the perennial evergreen shrub was first used in 14th century English, deriving from Old French rue and the Latin ruta.
Espand is derived from Middle Persian, which is thought, along with the English word spinach, to be ultimately derived from Proto-Iranian *spanta-, 'holy', itself thought to be ultimately derived from Proto-Indo-European *ḱwen-.

Common names

It is known by many names across regions and languages—including "African rue" in North America, "harmel" in India and North Africa, "espand" in Persian, and other local names in Pashto, Urdu, Turkish, Chinese, Spanish, French, and ancient Aramaic, reflecting its wide cultural and geographic significance.
It is known as اسپند in Persian, which is transliterated as, or but may also be pronounced or transliterated as,,,,,, or depending on source or dialect. The Persian word اسپند is also the name of the last month of the year, approximately March, in the traditional Persian calendar.
African rue is a common name.
Harmel is a name used in India, Algeria, and Morocco.
It is known as in Pashto. In Urdu it is known as,, or. In Turkish it is known as üzerlik. In Chinese it is 驼驼蒿,, or 骆驼蓬,.
In Spain, it is called hármala, alharma or gamarza, amongst dozens of other local names. In French, it is known as harmal.
In classical antiquity, it was known in Aramaic as šabbārā. In later Eastern Aramaic languages, it was also borrowed from the Middle Persian as spendā.

Description

Habitus

It is a perennial, herbaceous, suffrutescent, hemicryptophyte plant, which dies off in the winter, but regrows from the rootstock the following spring. It can grow to about tall, but normally it is about tall. The entire plant is hairless. Plants are bad tasting and smell foul when crushed.

Stems

Numerous erect to spreading stems grow from the crown of the root-stock in the spring, these branch in a corymbose fashion.

Roots

The roots of the plant can reach a depth of up to, if the soil where it is growing is very dry. The roots can grow to thick.

Leaves

The leaves are alternate, sessile, and have bristly, long stipules at the base. The leaf blade is dissected/forked twice or more into three to five thin, linear to lanceolate-linear, greyish lobes. The forks are irregular. The lobes have smooth margins, are long and broad, and end in points.

Flowers

It blooms with solitary flowers opposite to the leaves on the apical parts of branches. It flowers between March and October in India, between April and October in Pakistan, between May and June in China, between March and April in Israel, and between May and July in Morocco. The flowers are white or yellowish white, and are about 2–3 cm in diameter. Greenish veins are visible in the petals. They have a threadlike, 1.2 cm long pedicel. The flowers have five 12–15mm long, linear, pointy-ended, glabrous sepals, often divided into lobes, although sometimes entire and only divided at the end. There are five petals which are oblong-elliptic, obovate to oblong in shape, 14–15mm long, 6–8mm broad, and ending with an obtuse apex. The flowers are hermaphroditic, having both male and female organs. The flowers usually have 15 stamens ; these have a 4-5mm long filament with an enlarged base. The dorsifixed, 6mm long anthers are longer than the filaments. The ovary is superior, and has 3 locules and ends in an 8-10mm long style, the ending 6mm of which are triangular or 3-keeled in cross-section. The ovary is surrounded by a nectary which is glabrous and has five lobes in a regular pattern.
The flowers produce only a tiny amount of nectar. The nectar is rich in hexose sugars. It contains a relatively small concentration of amino acids among which there is an especially high amount of the glutamic acid, tyrosine and proline, the last of which can be tasted by, and is favoured by, many insects. It also contains alkaloids, in relatively high concentration compared to the flowers of other species, among them the toxins harmalol and harmine. The proportions and concentrations of the alkaloids in the nectar are different than in the other organs of the plant, indicating an adaptive reason for their presence.

Pollen

P. harmala has smallish, tricolpate pollen grains with a rugulate-reticulate surface. The exine has a sexine which is thicker than the nexine. These grains are well distinguishable from pollen of related plants in Pakistan.

Fruit

The plant fruits between July and November in China. The fruit is a dry, round seed capsule which measures about 6–10 mm in diameter, These seed capsules have three chambers and carry more than 50 seeds. The end of the fruit is usually somewhat sunken inwards and retains a persistent style.

Seeds

The seeds are colored dark brown to blackish-brown, slightly curved, triangular, about long with an irregular surface. The endosperm is oily.

Cytology

The cells have 24 chromosomes, although sometimes 22 have been found.

Distribution

Native

Peganum harmala is native to a wide area stretching from Morocco in north Africa and Spain and Italy in Europe, north to Serbia, Romania, Dagestan, Kazakhstan, south to Mauritania, Yemen, Saudi Arabia, Kuwait and Pakistan In Zhob district, and east to western Mongolia, northern China and possibly Bangladesh. It is a common weed in Afghanistan, Iran, parts of Israel, eastern and central Anatolia, and Morocco.
In Africa it is known from Morocco, Algeria, Tunisia, Libya, and Egypt. It likely does not occur in Mauritania. In Morocco it is quite common and occurs throughout the country, excepting Western Sahara. In Algeria it is found mostly in the north bordering Morocco and Tunisia, being absent in the south and central regions. It is reasonably commonly found throughout Tunisia. In Libya it is found in the maritime zone, especially around Bengazi, and is not abundant. In Egypt it grows in the Sinai, has been recorded from the east of the Eastern Desert, and been rarely collected on the mid-west of the Mediterranean coast.
In Europe it is native to Spain, Corsica, much of Russia, Serbia, Moldova, Ukraine, Romania, Bulgaria, Greece, Cyprus, Turkey and southern Italy. It also is native to the Caucasus: Armenia, Azerbaijan, and Georgia. On the Iberian Peninsula it is absent from Portugal and Andorra, but it is not uncommon in Spain, especially in the southeast, the Ebro depression, and the inland valleys of the Duero and Tajo, but it is rare in Andalusia and it does not occur on the Balearic Islands and the Canary Islands, and in the west along the Portuguese border, Galicia, the northern coast, and the northern mountain ranges.
In Turkey it is found both in Thrace and across most of Anatolia, but is absent from the northern Black Sea coast. It is abundant in some regions of south and central Anatolia.
In Israel it is most commonly found around the Dead Sea, in the Judean mountains and desert, in the Negev and its surrounding areas, including areas in Jordan and Saudi Arabia, being rare or very rare in the northern mountains, Galilee, coastal areas and the Arava valley.
It grows in drier parts of the northern half of India but is possibly only native to the Kashmir and Ladakh regions. It also occurs in, and is possibly native to, Bangladesh.
The distribution in China is in dispute. The 2008 Flora of China considers it to be native to northern China in the provinces of Gansu, western Hebei, western Inner Mongolia, Ningxia, Qinghai, northern Shanxi, Tibet and Xinjiang. The 2017 Species Catalogue of China considers it to be restricted to Inner Mongolia, Ningxia and Gansu.

Adventive distribution

It has been added to the lists of the Global Register of Introduced and Invasive Species for the countries of South Africa, Mexico, France and Ukraine, although it is not reported as having a negative impact in any of these countries. Most Ukrainian and other references consider the plant native to Ukraine. Sources are in disagreement regarding rare collections in coastal Romania, but many consider it introduced. At least 7 occurrences have been registered in South Africa, and none in Mexico. As of 2020 it is included in South African National Biodiversity Institute's Plants of Southern Africa website as an introduced plant to South Africa. One database has it occurring as a non-native in Hungary.
In France it is considered a former accidental introduction once uncommonly found on the Côte d'Azur along the Mediterranean coast. It has very rarely been found elsewhere in France in the past. According to the Flora Europaea there is a native population on Corsica, however, according to Tela Botanica it does not occur on the island, either as a native or not.
It was first planted in the United States in 1928 in New Mexico by a farmer wanting to manufacture a dye called "Turkish red" from its seeds. From here the plant spread over most of southern New Mexico and the Big Bend region of Texas. An additional spread has occurred from east of Los Angeles in California to the tip of southernmost Nevada. Outside of these regions the distribution in the US is not continuous and localised. As of 2019 it has been reported in southern Arizona, northeastern Montana, northern Nevada, Oregon and possibly Washington. "Because it is so drought tolerant, African rue can displace the native saltbushes and grasses growing in the salt-desert shrub lands of the Western U.S."
Although the distribution in New Mexico and Texas would suggest it has spread to parts of northern Mexico, the species has not been included in the 2004 list of introduced plants of Mexico.