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Piper (genus)

Piper (genus)

Overview
Piper, the pepper plants or pepper vines (a term used for certain Clematis
Clematis
Clematis is a genus of about 300 species within the buttercup family Ranunculaceae. Their garden hybrids have been popular among gardeners beginning with Clematis × jackmanii, a garden standby since 1862; more hybrid cultivars are being produced constantly. They are mainly of Chinese and Japanese...

in older times), are an economically and ecologically important genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 in the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 Piperaceae
Piperaceae
The Piperaceae, also known as the pepper family, is a large family of flowering plants. The group contains roughly 3,610 currently accepted species in five genera. The vast majority of peppers can be found within the two main genera: Piper and Peperomia .Members of the Piperaceae may be small...

. It contains about 1,000-2,000 species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 of shrubs, herbs, and liana
Liana
A liana is any of various long-stemmed, woody vines that are rooted in the soil at ground level and use trees, as well as other means of vertical support, to climb up to the canopy to get access to well-lit areas of the forest. Lianas are especially characteristic of tropical moist deciduous...

s, many of which are keystone species
Keystone species
A keystone species is a species that has a disproportionately large effect on its environment relative to its abundance. Such species play a critical role in maintaining the structure of an ecological community, affecting many other organisms in an ecosystem and helping to determine the types and...

 in their native habitat The diversification of this taxon is of interest to understanding the evolution of plants.
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Encyclopedia
Piper, the pepper plants or pepper vines (a term used for certain Clematis
Clematis
Clematis is a genus of about 300 species within the buttercup family Ranunculaceae. Their garden hybrids have been popular among gardeners beginning with Clematis × jackmanii, a garden standby since 1862; more hybrid cultivars are being produced constantly. They are mainly of Chinese and Japanese...

in older times), are an economically and ecologically important genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 in the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 Piperaceae
Piperaceae
The Piperaceae, also known as the pepper family, is a large family of flowering plants. The group contains roughly 3,610 currently accepted species in five genera. The vast majority of peppers can be found within the two main genera: Piper and Peperomia .Members of the Piperaceae may be small...

. It contains about 1,000-2,000 species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 of shrubs, herbs, and liana
Liana
A liana is any of various long-stemmed, woody vines that are rooted in the soil at ground level and use trees, as well as other means of vertical support, to climb up to the canopy to get access to well-lit areas of the forest. Lianas are especially characteristic of tropical moist deciduous...

s, many of which are keystone species
Keystone species
A keystone species is a species that has a disproportionately large effect on its environment relative to its abundance. Such species play a critical role in maintaining the structure of an ecological community, affecting many other organisms in an ecosystem and helping to determine the types and...

 in their native habitat The diversification of this taxon is of interest to understanding the evolution of plants.

Pepper plants belong to the magnoliids, which are angiosperms but neither monocots nor eudicots. Their family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

, Piperaceae
Piperaceae
The Piperaceae, also known as the pepper family, is a large family of flowering plants. The group contains roughly 3,610 currently accepted species in five genera. The vast majority of peppers can be found within the two main genera: Piper and Peperomia .Members of the Piperaceae may be small...

, is most closely related to the lizardtail family (Saururaceae
Saururaceae
Saururaceae is a plant family comprising four genera and seven species of herbaceous flowering plants native to eastern and southern Asia and North America. The family has been recognised by most taxonomists, and is sometimes known as the "lizard's-tail family"...

), which in fact generally look like smaller, more delicate and amphibious
Amphibious
Amphibious means able to use either land or water. In particular it may refer to:*Amphibious warfare, warfare carried out on both land and water*Amphibians, vertebrate animals of the Class Amphibia...

 pepper plants. Both families have characteristic tail-shaped inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...

s covered in tiny flowers. A somewhat less close relative is the pipevine family (Aristolochiaceae
Aristolochiaceae
The Aristolochiaceae, or the Birthwort family, are a family of flowering plants with 7 genera and about 400 species belonging to the order Piperales...

). A well-known and very close relative – being also part of the Piperaceae – are the radiator plants of the genus Peperomia.

The scientific name Piper and the common name "pepper" are derived from the Sanskrit
Sanskrit
Sanskrit , is a historical Indo-Aryan language and the primary liturgical language of Hinduism, Jainism and Buddhism.Buddhism: besides Pali, see Buddhist Hybrid Sanskrit Today, it is listed as one of the 22 scheduled languages of India and is an official language of the state of Uttarakhand...

 term pippali, denoting the Long Pepper
Long pepper
Long pepper , , sometimes called Indian long pepper, is a flowering vine in the family Piperaceae, cultivated for its fruit, which is usually dried and used as a spice and seasoning. Long pepper has a similar, but hotter, taste to its close relative Piper nigrum - from which black, green and white...

 (P. longum).

Distribution and ecology


Piper species have a pantropical distribution, and are most commonly found in the understory of lowland tropical rainforest
Tropical rainforest
A tropical rainforest is an ecosystem type that occurs roughly within the latitudes 28 degrees north or south of the equator . This ecosystem experiences high average temperatures and a significant amount of rainfall...

s, but can also occur in clearings and in higher elevation life zones such as cloud forest
Cloud forest
A cloud forest, also called a fog forest, is a generally tropical or subtropical evergreen montane moist forest characterized by a persistent, frequent or seasonal low-level cloud cover, usually at the canopy level. Cloud forests often exhibit an abundance of mosses covering the ground and...

s; one species (Japanese Pepper, P. kadsura, from southern Japan
Japan
Japan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...

 and southernmost Korea
Korea
Korea ) is an East Asian geographic region that is currently divided into two separate sovereign states — North Korea and South Korea. Located on the Korean Peninsula, Korea is bordered by the People's Republic of China to the northwest, Russia to the northeast, and is separated from Japan to the...

) is subtropical and can tolerate light winter frost
Frost
Frost is the solid deposition of water vapor from saturated air. It is formed when solid surfaces are cooled to below the dew point of the adjacent air as well as below the freezing point of water. Frost crystals' size differ depending on time and water vapour available. Frost is also usually...

. Peppers are often dominant vegetation
Vegetation
Vegetation is a general term for the plant life of a region; it refers to the ground cover provided by plants. It is a general term, without specific reference to particular taxa, life forms, structure, spatial extent, or any other specific botanical or geographic characteristics. It is broader...

 where they are found.

Most Piper species are either herbaceous
Herbaceous
A herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...

 or vine
Vine
A vine in the narrowest sense is the grapevine , but more generally it can refer to any plant with a growth habit of trailing or scandent, that is to say climbing, stems or runners...

s; some grow as shrub
Shrub
A shrub or bush is distinguished from a tree by its multiple stems and shorter height, usually under 5–6 m tall. A large number of plants may become either shrubs or trees, depending on the growing conditions they experience...

s or almost as small tree
Tree
A tree is a perennial woody plant. It is most often defined as a woody plant that has many secondary branches supported clear of the ground on a single main stem or trunk with clear apical dominance. A minimum height specification at maturity is cited by some authors, varying from 3 m to...

s. A few species, commonly called "ant pipers" (e.g. Piper cenocladum
Piper cenocladum
The tropical rainforest understory shrub Piper cenocladum is a member of the same genus as kava, betel, and black pepper. It is a myrmecophyte, a plant that lives in ecological mutualism with ants. This plant and three or four other closely related species are known collectively as ant plants or...

), live in a mutualism with ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s. The fruit of the Piper plant, called a peppercorn when it is round and pea-sized, as is usual, is distributed in the wild mainly by bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s, but small fruit-eating mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s – e.g. bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...

s of the genus Carollia
Carollia
Carollia is a genus of bats often referred to as the short-tailed fruit bats. Along with the genus, Rhinophylla, Carollia makes up the subfamily Carolliinae of family Phyllostomidae, the leaf-nosed bats. Currently, nine species of Carollia are recognized, with a number having been described since...

– are also important. Despite the high content of chemicals that are noxious to herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

s, some have evolved the ability to withstand the chemical defences of pepper plants, for example the sematurine
Sematurinae
Sematurinae is a subfamily of moths in the family Sematuridae, represented by at least 29 species in the Neotropics.-Taxonomy, classification and identification:Sematurinae has three principal genera, and possibly another two...

 moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...

 Homidiana subpicta or some flea beetle
Flea beetle
Flea beetles is a general name applied to the small, jumping beetles of the leaf beetle family . They make up the tribe Alticini, which is a part of the subfamily Galerucinae, though they were historically classified as a subfamily in their own right...

s of the genus Lanka. The latter can be significant pests to pepper growers.

Piper and humans


Many pepper plants make good ornamental
Ornamental plant
Ornamental plants are plants that are grown for decorative purposes in gardens and landscape design projects, as house plants, for cut flowers and specimen display...

s for gardens in subtropical or warmer regions. Pepper vines can be used much as ivy
Ivy
Ivy, plural ivies is a genus of 12–15 species of evergreen climbing or ground-creeping woody plants in the family Araliaceae, native to western, central and southern Europe, Macaronesia, northwestern Africa and across central-southern Asia east to Japan and Taiwan.-Description:On level ground they...

 in temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

 climates, while other species, like Lacquered Pepper (P. magnificum) grow as sizeable, compact and attractive shrubs with tough and shiny leaves. Smaller species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

, like Celebes Pepper (P. ornatum) with its finely patterned leaves, are also suitable as indoor pot plants.

Unsustainable logging
Logging
Logging is the cutting, skidding, on-site processing, and loading of trees or logs onto trucks.In forestry, the term logging is sometimes used in a narrow sense concerning the logistics of moving wood from the stump to somewhere outside the forest, usually a sawmill or a lumber yard...

 of tropical primary forests is threatening a number of peppers. The extent of the effect of such wholesale habitat destruction
Habitat destruction
Habitat destruction is the process in which natural habitat is rendered functionally unable to support the species present. In this process, the organisms that previously used the site are displaced or destroyed, reducing biodiversity. Habitat destruction by human activity mainly for the purpose of...

 on the genus is unknown, but it is to be noted that in the forests of Ecuador
Ecuador
Ecuador , officially the Republic of Ecuador is a representative democratic republic in South America, bordered by Colombia on the north, Peru on the east and south, and by the Pacific Ocean to the west. It is one of only two countries in South America, along with Chile, that do not have a border...

 – the only larger region for which comprehensive data exists – more than a dozen species are known to be on the brink of extinction. On the other hand, other Piper species (e.g. Spiked Pepper, P. aduncum) have been widely distributed as a result of human activity and are a major invasive species
Invasive species
"Invasive species", or invasive exotics, is a nomenclature term and categorization phrase used for flora and fauna, and for specific restoration-preservation processes in native habitats, with several definitions....

 in certain areas.

The most significant human use of Piper is not for its looks however, but ultimately for the wide range of powerful secondary compounds found particularly in the fruits.

As spice and vegetable

For other spices called "pepper", see Pepper.

Culinary use of pepper plants is attested perhaps as early as 9,000 years ago. Peppercorn remains were found among the food refuse left by Hoabinhian
Hoabinhian
The term Hoabinhian was first used by French archaeologists working in Northern Vietnam to describe Holocene period archaeological assemblages excavated from rock shelters. It has become a common term in the English based literature to describe stone artifact assemblages in Southeast Asia that...

 artisans at Spirit Cave, Vietnam
Vietnam
Vietnam – sometimes spelled Viet Nam , officially the Socialist Republic of Vietnam – is the easternmost country on the Indochina Peninsula in Southeast Asia. It is bordered by China to the north, Laos to the northwest, Cambodia to the southwest, and the South China Sea –...

. It is not too likely that these plants were deliberately grown rather than collected from the wild.
Use of peppercorns as pungent spice is significant on an international scale. In classical antiquity
Classical antiquity
Classical antiquity is a broad term for a long period of cultural history centered on the Mediterranean Sea, comprising the interlocking civilizations of ancient Greece and ancient Rome, collectively known as the Greco-Roman world...

, there was a vigorous trade of spices
Spice trade
Civilizations of Asia were involved in spice trade from the ancient times, and the Greco-Roman world soon followed by trading along the Incense route and the Roman-India routes...

 including Black Pepper
Black pepper
Black pepper is a flowering vine in the family Piperaceae, cultivated for its fruit, which is usually dried and used as a spice and seasoning. The fruit, known as a peppercorn when dried, is approximately in diameter, dark red when fully mature, and, like all drupes, contains a single seed...

 (P. nigrum) from South Asia
South Asia
South Asia, also known as Southern Asia, is the southern region of the Asian continent, which comprises the sub-Himalayan countries and, for some authorities , also includes the adjoining countries to the west and the east...

 to Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

 already. The Apicius
Apicius
Apicius is the title of a collection of Roman cookery recipes, usually thought to have been compiled in the late 4th or early 5th century AD and written in a language that is in many ways closer to Vulgar than to Classical Latin....

, a recipe collection complied about 400 AD, mentions "pepper" as a spice for most main dishes. Judging from the Apicius, in the late Roman Empire
Roman Empire
The Roman Empire was the post-Republican period of the ancient Roman civilization, characterised by an autocratic form of government and large territorial holdings in Europe and around the Mediterranean....

, Black Pepper was probably still expensive, but nevertheless seems to have been available readily enough to be used more frequently than salt
Salt
In chemistry, salts are ionic compounds that result from the neutralization reaction of an acid and a base. They are composed of cations and anions so that the product is electrically neutral...

 or sugar
Sugar
Sugar is a class of edible crystalline carbohydrates, mainly sucrose, lactose, and fructose, characterized by a sweet flavor.Sucrose in its refined form primarily comes from sugar cane and sugar beet...

. As Europe turned into the Early Middle Ages
Early Middle Ages
The Early Middle Ages was the period of European history lasting from the 5th century to approximately 1000. The Early Middle Ages followed the decline of the Western Roman Empire and preceded the High Middle Ages...

, trade routes deteriorated and the use of pepper declined somewhat.

But this dearth was hardly ever absolute and altogether rather short-lived. Black peppercorns, storing easily and having a high mass per volume, never ceased to be a profitable trade item. In the Middle Ages
Middle Ages
The Middle Ages is a periodization of European history from the 5th century to the 15th century. The Middle Ages follows the fall of the Western Roman Empire in 476 and precedes the Early Modern Era. It is the middle period of a three-period division of Western history: Classic, Medieval and Modern...

, international traders were nicknamed Pfeffersäcke ("pepper-sacks") in German
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

 towns of the Hanseatic League
Hanseatic League
The Hanseatic League was an economic alliance of trading cities and their merchant guilds that dominated trade along the coast of Northern Europe...

 and elsewhere. As the Modern Era came into full swing, wars were fought by European powers, between themselves and in complex alliances and enmities with Indian Ocean
Indian Ocean
The Indian Ocean is the third largest of the world's oceanic divisions, covering approximately 20% of the water on the Earth's surface. It is bounded on the north by the Indian Subcontinent and Arabian Peninsula ; on the west by eastern Africa; on the east by Indochina, the Sunda Islands, and...

 states, about control of the supply of spices, perhaps the most archetypal being Black Pepper fruit. Today, Black Pepper corns of the three preparations (green, white and black) are one of the most widely used spices of plant origin worldwide.

Due to the wide distribution of Piper, the fruit of other species are also important spices, many of them internationally. Mecaxochitl (P. amalgo) was used by the Aztecs to spice up cocoa. Cubeb
Cubeb
Cubeb , or tailed pepper, is a plant in genus Piper, cultivated for its fruit and essential oil. It is mostly grown in Java and Sumatra, hence sometimes called Java pepper. The fruits are gathered before they are ripe, and carefully dried...

 (P. cubeba), also known as Tailed or Javan Pepper actually played a major role in Early Modern Era spice trade; ocet kubebowy, Cubeb-flavored vinegar, was a popular condiment in 14th century Poland
Poland
Poland , officially the Republic of Poland , is a country in Central Europe bordered by Germany to the west; the Czech Republic and Slovakia to the south; Ukraine, Belarus and Lithuania to the east; and the Baltic Sea and Kaliningrad Oblast, a Russian exclave, to the north...

. But reputedly Philip IV of Spain
Philip IV of Spain
Philip IV was King of Spain between 1621 and 1665, sovereign of the Spanish Netherlands, and King of Portugal until 1640...

 at the end of the 1630s suppressed trade in Cubeb peppercorns to capitalize on his massive share of the Black Pepper trade. After a brief comeback as a medical plant, Cubeb is nowadays fairly obscure in the West. It is however a most significant spice around the Indian Ocean
Indian Ocean
The Indian Ocean is the third largest of the world's oceanic divisions, covering approximately 20% of the water on the Earth's surface. It is bounded on the north by the Indian Subcontinent and Arabian Peninsula ; on the west by eastern Africa; on the east by Indochina, the Sunda Islands, and...

 region today, and popular as far as Morocco
Morocco
Morocco , officially the Kingdom of Morocco , is a country located in North Africa. It has a population of more than 32 million and an area of 710,850 km², and also primarily administers the disputed region of the Western Sahara...

 where it flavors spicy-sweet markout pastry
Pastry
Pastry is the name given to various kinds of baked products made from ingredients such as flour, sugar, milk, butter, shortening, baking powder and/or eggs. Small cakes, tarts and other sweet baked products are called "pastries."...

 and is sometimes included in the famous ras el hanout
Ras el hanout
Ras el hanout or Rass el hanout is a popular Moroccan blend of spices that is used across North Africa. The name means "top of the shop" in Arabic and refers to a mixture of the best spices a seller has to offer...

mix. Further east, it is well-loved across Indonesia, where it is the popular pepper to use in gulé (curried stews).

West African Pepper
West African Pepper
West African Pepper, also known as Ashanti Pepper, Benin Pepper, False Cubeb, Guinea Cubeb, Uziza Pepper or "Guinea pepper", called locally kale, kukauabe, masoro, sasema and soro wisa, is a West African spice that corresponds to the dried fruit of Piper guineense...

 (P. guineense), also given a variety of "regionalized" names like "Benin
Benin
Benin , officially the Republic of Benin, is a country in West Africa. It borders Togo to the west, Nigeria to the east and Burkina Faso and Niger to the north. Its small southern coastline on the Bight of Benin is where a majority of the population is located...

 pepper", "Ashanti pepper, or the rather ambiguous "Guinea pepper
Guinea pepper
Guinea pepper is the name for several unrelated pepper-like spices traded from the general region of West Africa:* Aframomum melegueta from the ginger family...

", is considered supreme for use in stews and other regional specialties
West African cuisine
West African cuisine is a general term referring to many distinct regional and ethnic cuisines in West African nations, with climates ranging from desert to tropical...

. It is used in addition to Black Pepper to impart a particularly refined aroma with hints of nutmeg
Nutmeg
The nutmeg tree is any of several species of trees in genus Myristica. The most important commercial species is Myristica fragrans, an evergreen tree indigenous to the Banda Islands in the Moluccas of Indonesia...

 and saffron
Saffron
Saffron is a spice derived from the flower of Crocus sativus, commonly known as the saffron crocus. Crocus is a genus in the family Iridaceae. Each saffron crocus grows to and bears up to four flowers, each with three vivid crimson stigmas, which are each the distal end of a carpel...

. Sometimes it is used in the East Africa
East Africa
East Africa or Eastern Africa is the easterly region of the African continent, variably defined by geography or geopolitics. In the UN scheme of geographic regions, 19 territories constitute Eastern Africa:...

n berbere
Berbere
Berbere is a spice mixture whose ingredients usually include chile peppers, garlic, ginger, dried basil, korarima, rue, white and black pepper, and fenugreek...

spice mix. This species, despite being traded more extensively in earlier times, is very hard to get outside Africa today. More readily available in the West is Long Pepper
Long pepper
Long pepper , , sometimes called Indian long pepper, is a flowering vine in the family Piperaceae, cultivated for its fruit, which is usually dried and used as a spice and seasoning. Long pepper has a similar, but hotter, taste to its close relative Piper nigrum - from which black, green and white...

 (P. longum), commonly traded under its India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

n name pipalli. This is possibly the secondmost popular Piper spice internationally; it has a rather chili
Chili pepper
Chili pepper is the fruit of plants from the genus Capsicum, members of the nightshade family, Solanaceae. The term in British English and in Australia, New Zealand, India, Malaysia and other Asian countries is just chilli without pepper.Chili peppers originated in the Americas...

-like "heat" and the whole inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...

 is used as the fruits are tiny.

Not only the seeds of Piper are used in cooking. West African Pepper leaves, known locally as uziza, are used as flavoring vegetable in Nigeria
Nigeria
Nigeria , officially the Federal Republic of Nigeria, is a federal constitutional republic comprising 36 states and its Federal Capital Territory, Abuja. The country is located in West Africa and shares land borders with the Republic of Benin in the west, Chad and Cameroon in the east, and Niger in...

n stews. In Mexican
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...

-influenced cooking, hoja santa or Mexican Pepperleaf (P. auritum) has a variety of uses, mainly to impart flavor too. In Southeast Asia
Southeast Asia
Southeast Asia, South-East Asia, South East Asia or Southeastern Asia is a subregion of Asia, consisting of the countries that are geographically south of China, east of India, west of New Guinea and north of Australia. The region lies on the intersection of geological plates, with heavy seismic...

, leaves of two species of Piper have major importance in cooking: Lolot
Lolot
Lolot is a flowering vine in the family Piperaceae, cultivated for its leaf which is used in Lao and Vietnamese cuisine as a flavoring wrap for grilling meats, namely the bò lá lốt sausages of Vietnam. It is also known as lolot pepper...

 (P. lolot) is used to wrap meat
Meat
Meat is animal flesh that is used as food. Most often, this means the skeletal muscle and associated fat and other tissues, but it may also describe other edible tissues such as organs and offal...

 for grilling in the Indochina
Indochina
The Indochinese peninsula, is a region in Southeast Asia. It lies roughly southwest of China, and east of India. The name has its origins in the French, Indochine, as a combination of the names of "China" and "India", and was adopted when French colonizers in Vietnam began expanding their territory...

 region, while Wild Betel (P. sarmentosum), often seen under the Thai name cha phlu, is used raw or cooked as a vegetable in its own right in Malay
Malay cuisine
Malay cusine is the cuisine of Malay people of Malaysia, Indonesia, Singapore, Brunei and Southern Thailand. Different Malay regions are all known for their unique or signature dishes - Terengganu and Kelantan for their Nasi dagang, Nasi kerabu and Keropok lekor, Negeri Sembilan for its...

 and Thai cuisine; it is also used to prepare the famous miang kham snacks of northwestern Thailand.

As medicine and stimulant



Cubeb
Cubeb
Cubeb , or tailed pepper, is a plant in genus Piper, cultivated for its fruit and essential oil. It is mostly grown in Java and Sumatra, hence sometimes called Java pepper. The fruits are gathered before they are ripe, and carefully dried...

 (P. cubeba) has been used in folk medicine
Folk medicine
-Description:Refers to healing practices and ideas of body physiology and health preservation known to a limited segment of the population in a culture, transmitted informally as general knowledge, and practiced or applied by anyone in the culture having prior experience.All cultures and societies...

, herbalism
Herbalism
Herbalism is a traditional medicinal or folk medicine practice based on the use of plants and plant extracts. Herbalism is also known as botanical medicine, medical herbalism, herbal medicine, herbology, herblore, and phytotherapy...

 and related fields (e.g. alchemy
Alchemy
Alchemy is an influential philosophical tradition whose early practitioners’ claims to profound powers were known from antiquity. The defining objectives of alchemy are varied; these include the creation of the fabled philosopher's stone possessing powers including the capability of turning base...

, folk magic or exorcism
Exorcism
Exorcism is the religious practice of evicting demons or other spiritual entities from a person or place which they are believed to have possessed...

), as well as, particularly in the early 20th century, as a cigarette
Cigarette
A cigarette is a small roll of finely cut tobacco leaves wrapped in a cylinder of thin paper for smoking. The cigarette is ignited at one end and allowed to smoulder; its smoke is inhaled from the other end, which is held in or to the mouth and in some cases a cigarette holder may be used as well...

 flavoring. P. darienense
Piper darienense
Piper darienense is a flowering plant in the family Piperaceae. It is called Duermeboca in Panama. In El Chocó, the plant is used as a fish poison. Cuna, an indigenous people of Panama and Colombia, call it Kana, and they use it in a bath, for snakebites and colds....

is used medically by the Kuna people of the Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

-Colombia
Colombia
Colombia, officially the Republic of Colombia , is a unitary constitutional republic comprising thirty-two departments. The country is located in northwestern South America, bordered to the east by Venezuela and Brazil; to the south by Ecuador and Peru; to the north by the Caribbean Sea; to the...

 border region, and elsewhere it is used to intoxicate fish
Fish
Fish are a paraphyletic group of organisms that consist of all gill-bearing aquatic vertebrate animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish, as well as various extinct related groups...

 which then can be easily caught. Spiked Pepper, often called "matico", appears to have strong disinfectant and antibiotic
Antibiotic
An antibacterial is a compound or substance that kills or slows down the growth of bacteria.The term is often used synonymously with the term antibiotic; today, however, with increased knowledge of the causative agents of various infectious diseases, antibiotic has come to denote a broader range of...

 properties. Black Pepper
Black pepper
Black pepper is a flowering vine in the family Piperaceae, cultivated for its fruit, which is usually dried and used as a spice and seasoning. The fruit, known as a peppercorn when dried, is approximately in diameter, dark red when fully mature, and, like all drupes, contains a single seed...

 (P. nigrum) essential oil
Essential oil
An essential oil is a concentrated hydrophobic liquid containing volatile aroma compounds from plants. Essential oils are also known as volatile oils, ethereal oils or aetherolea, or simply as the "oil of" the plant from which they were extracted, such as oil of clove...

 is sometimes used in herbalism, and Long Pepper
Long pepper
Long pepper , , sometimes called Indian long pepper, is a flowering vine in the family Piperaceae, cultivated for its fruit, which is usually dried and used as a spice and seasoning. Long pepper has a similar, but hotter, taste to its close relative Piper nigrum - from which black, green and white...

 (P. longum) is similarly employed in Ayurveda
Ayurveda
Ayurveda or ayurvedic medicine is a system of traditional medicine native to India and a form of alternative medicine. In Sanskrit, words , meaning "longevity", and , meaning "knowledge" or "science". The earliest literature on Indian medical practice appeared during the Vedic period in India,...

, where it was an ingredient of Triphala Guggulu and (together with Black Pepper) of Trikatu pill
Pill (pharmacy)
A pill is a small, round, solid pharmaceutical oral dosage form that was in use before the advent of tablets and capsules. Pills were made by mixing the active ingredients with an excipient such as glucose syrup in a mortar and pestle to form a paste, then rolling the mass into a long cylindrical...

s, used for rasayana
Rasayana
Rasayan, a Sanskrit word , is used to describe chemistry and alchemy, and chemistry is generally called Rasayan Shastra in Sanskrit, Nepali, Marathi, Hindi, Kannada and several other languages.Ancient rasayan texts center around the use of prepared forms of mercury or cinnabar , as do...

(rejuvenating and detoxifying
Body cleansing
Detoxification is an alternative medicine approach that proponents claim rids the body of "toxins", accumulated harmful substances that are alleged to exert undesirable effects on individual health...

) purposes.
See also Betel leaf, Kava culture
Kava culture
Kava cultures are the religious and cultural traditions of western Oceania which consume kava. There are similarities in the use of kava between the different cultures, but each one also has its own traditions.-Hawaii:...

.

Two Piper species have gained large-scale use as a stimulant
Stimulant
Stimulants are psychoactive drugs which induce temporary improvements in either mental or physical function or both. Examples of these kinds of effects may include enhanced alertness, wakefulness, and locomotion, among others...

. Betel
Betel
The Betel is the leaf of a vine belonging to the Piperaceae family, which includes pepper and Kava. It is valued both as a mild stimulant and for its medicinal properties...

 (P. betle) leaves are used to wrap Betel palm nut slices; its sap helps release the stimulating effect of these "cookies" which are widely known as pan in India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

. In the Pacific region, where it has been widely spread as a canoe plant
Canoe plants
Canoe plants, or Polynesian introductions, are plants taken from ancient Polynesia and transplanted to other Pacific IslandsThe term is particularly used to refer to plants brought to Hawaii 1,700 years ago by Polynesian explorers....

, Kava
Kava
Kava or kava-kava is a crop of the western Pacific....

 (P. methysticum) is used to produce a calming drink somewhat similar to alcohol
Alcohol
In chemistry, an alcohol is an organic compound in which the hydroxy functional group is bound to a carbon atom. In particular, this carbon center should be saturated, having single bonds to three other atoms....

 but without many of the negative side effects and addiction
Substance use disorder
Substance use disorders include substance abuse and substance dependence. In DSM-IV, the conditions are formally diagnosed as one or the other, but it has been proposed that DSM-5 combine the two into a single condition called "Substance-use disorder"....

 risk. It has also become popular elsewhere in recent decades, and is used as a medical plant. However, Kava whole-plant "herbal supplement" pills have occasionally shown a strong hepatotoxic effect, which has led to their banning in many countries. On the other hand, the traditional preparation of the root as a calming drink appears to pose little if any such hazard.

In science


The genus contains species suitable for studying natural history
Natural history
Natural history is the scientific research of plants or animals, leaning more towards observational rather than experimental methods of study, and encompasses more research published in magazines than in academic journals. Grouped among the natural sciences, natural history is the systematic study...

, molecular biology
Molecular biology
Molecular biology is the branch of biology that deals with the molecular basis of biological activity. This field overlaps with other areas of biology and chemistry, particularly genetics and biochemistry...

, natural product
Natural product
A natural product is a chemical compound or substance produced by a living organism - found in nature that usually has a pharmacological or biological activity for use in pharmaceutical drug discovery and drug design...

s chemistry
Chemistry
Chemistry is the science of matter, especially its chemical reactions, but also its composition, structure and properties. Chemistry is concerned with atoms and their interactions with other atoms, and particularly with the properties of chemical bonds....

, community ecology, and evolutionary biology.
Piper is a model genus
Model organism
A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the organism model will provide insight into the workings of other organisms. Model organisms are in vivo models and are widely used to...

 for research in ecology and evolutionary biology
Ecology and evolutionary biology
Some U.S. universities are home to degree programs titled Ecology and Evolutionary Biology, offering integrated studies in the disciplines of ecology and evolutionary biology. The wording is intended as representing the alternative approach from the frequently used pairing of Cell and Molecular...

. The diversity and ecological importance of the genus makes it an obvious candidate for ecological and evolutionary studies, though not surprisingly, most research has focused on the economically important species P. nigrum (Black Pepper
Black pepper
Black pepper is a flowering vine in the family Piperaceae, cultivated for its fruit, which is usually dried and used as a spice and seasoning. The fruit, known as a peppercorn when dried, is approximately in diameter, dark red when fully mature, and, like all drupes, contains a single seed...

), P. methysticum (Kava
Kava
Kava or kava-kava is a crop of the western Pacific....

), and P. betle (Betel
Betel
The Betel is the leaf of a vine belonging to the Piperaceae family, which includes pepper and Kava. It is valued both as a mild stimulant and for its medicinal properties...

).

The obligate and facultative ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

 mutualists found in some Piper species have a strong influence on their biology, making them ideal systems for research on the evolution of symbioses
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

 and the effect of mutualisms on biotic communities.

Important secondary metabolite
Secondary metabolite
Secondary metabolites are organic compounds that are not directly involved in the normal growth, development, or reproduction of an organism. Unlike primary metabolites, absence of secondary metabolities does not result in immediate death, but rather in long-term impairment of the organism's...

s found in pepper plants are piperine
Piperine
Piperine is the alkaloid responsible for the pungency of black pepper and long pepper, along with chavicine . It has also been used in some forms of traditional medicine and as an insecticide. Piperine forms monoclinic needles, is slightly soluble in water and more so in alcohol, ether or...

 and chavicine
Chavicine
Chavicine is an alkaloid found in pepper. It is one of the four diastereomeric geometric isomers of piperine....

, which were first isolated from Black Pepper. The piperidine
Piperidine
Piperidine is an organic compound with the molecular formula 5NH. This heterocyclic amine consists of a six-membered ring containing five methylene units and one nitrogen atom...

 functional group
Functional group
In organic chemistry, functional groups are specific groups of atoms within molecules that are responsible for the characteristic chemical reactions of those molecules. The same functional group will undergo the same or similar chemical reaction regardless of the size of the molecule it is a part of...

 is named after the former, and piperazine
Piperazine
Piperazine is an organic compound that consists of a six-membered ring containing two opposing nitrogen atoms. Piperazine exists as small alkaline deliquescent crystals with a saline taste....

 (which is not found in P. nigrum in noticeable quantities) was in turn named after piperidine.

The significant secondary metabolites of Kava are kavalactone
Kavalactone
Kavalactones are a class of lactone compounds found in the kava shrub.-Compounds:At least 18 different kavalactones have been identified to date, with methysticin being the first identified.Analogues, such as ethysticin, have also been isolated....

s and flavokawains. Pipermethystine
Pipermethystine
Pipermethystine is an alkaloid present in the aerial portions of the kava plant.-Toxicity:Pipermethystine is known to induce hepatoxicity in humans, and is believed to be the cause of liver failure in individuals consuming kava supplements....

 is suspected to be the main hepatotoxic compound in this plant's stems and leaves.

Selected species


The largest number of Piper species are found in the Americas (about 700 species), with about 300 species from Southern Asia. There are smaller groups of species from the South Pacific (about 40 species) and Africa (about 15 species). The American, Asian, and South Pacific groups each appear to be monophyletic; the affinity of the African species is unclear.

Some species are sometimes segregated
Segregate (taxonomy)
In taxonomy, a segregate, or a segregate taxon is created when a taxon is split off, from another taxon. This other taxon will be better known, usually bigger, and will continue to exist, even after the segregate taxon has been split off...

 into the genera Pothomorphe, Macropiper, Ottonia, Arctottonia, Sarcorhachis, Trianaeopiper, and Zippelia, but other sources keep them in Piper.
Piper aggregatum and P. fasciculatum are actually Lacistema aggregatum, an entirely unrelated angiosperm from the family Lacistemataceae
Lacistemataceae
Lacistemataceae is a family of flowering plants, consisting 2 genera, Lacistema Sw. and Lozania Mutis ex Caldas .- Common name :Waits Numi...

.

External links