''Piper'', the pepper plants or pepper vines, is an economically and ecologically important

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

in the family

Piperaceae The Piperaceae (), also known as the pepper family, are a large family (biology), family of flowering plants. The group contains roughly 3,600 currently accepted species in five genera. The vast majority of species can be found within the two mai ...

. It contains about 1,000–2,000

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

of shrubs, herbs, and lianas, many of which are dominant species 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

angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...

s but neither monocots nor eudicots. Their family,

, is most closely related to the lizardtail family ( Saururaceae), which in fact generally look like smaller, more delicate and amphibious pepper plants. Both families have characteristic tail-shaped

inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...

s covered in tiny flowers. A somewhat less close relative is the pipevine family ( Aristolochiaceae). A well-known and very close relative – being also part of the Piperaceae – are the radiator plants of the genus '' Peperomia''. The

scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...

''Piper'' and the common name "pepper" are derived from the

Sanskrit Sanskrit (; stem form ; nominal singular , ,) is a classical language belonging to the Indo-Aryan languages, Indo-Aryan branch of the Indo-European languages. It arose in northwest South Asia after its predecessor languages had Trans-cultural ...

term ''pippali'', denoting the long pepper (''P. longum'').

Evolution

The earliest fossil of ''Piper'' is of †''Piper margaritae'' from the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...

(

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

) of

Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...

. ''P. margaritae'' appears to be nested within the clade ''Schilleria'', indicating extensive Cretaceous diversification of ''Piper'' into the multiple extant clades, coinciding with the final breakup of Gondwana. This contrasts with previous theories assuming a younger radiation of the genus. An earlier potential record is of †''Piper arcuatile'' from the Cenomanian to Santonian Kaltag Formation of Yukon, although this affinity to ''Piper'' is not entirely reliable.

Distribution and ecology

''Piper'' species have a pantropical distribution, and are most commonly found in the understory of lowland tropical forests, but can also occur in clearings and in higher elevation life zones such as cloud forests; one species – the Japanese Pepper ('' P. kadsura'') from southern

Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...

and southernmost

Korea Korea is a peninsular region in East Asia consisting of the Korean Peninsula, Jeju Island, and smaller islands. Since the end of World War II in 1945, it has been politically Division of Korea, divided at or near the 38th parallel north, 3 ...

– is subtropical and can tolerate light winter frost. Peppers are often dominant species where they are found. Most ''Piper'' species are either herbaceous or

vine A vine is any plant with a growth habit of trailing or scandent (that is, climbing) stems, lianas, or runners. The word ''vine'' can also refer to such stems or runners themselves, for instance, when used in wicker work.Jackson; Benjamin; Da ...

s; some grow as shrubs or almost as small

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...

s. A few species, commonly called "ant pipers" (e.g. '' Piper cenocladum''), live in a mutualism with

ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...

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 Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

s, but small fruit-eating

mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...

s – e.g. bats of the genus '' Carollia'' – are also important. Despite the high content of chemicals that are noxious to herbivores, some have evolved the ability to withstand the chemical defences of pepper plants, for example the sematurine

moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...

'' Homidiana subpicta'' or some flea beetles of the genus '' Lanka''. The latter can be significant pests to pepper growers.

Usages

Many pepper plants make good ornamentals for gardens in subtropical or warmer regions. Pepper vines can be used much as ivy in temperate climates, while other species, like lacquered pepper ('' P. magnificum'') grow as sizeable, compact and attractive shrubs with tough and shiny leaves. Smaller

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

logging Logging is the process of cutting, processing, and moving trees to a location for transport. It may include skidder, skidding, on-site processing, and loading of trees or trunk (botany), logs onto logging truck, trucksprimary forests is threatening a number of peppers. The extent of the effect of such wholesale habitat destruction on the genus is unknown, but in the forests of

Ecuador Ecuador, officially the Republic of Ecuador, is a country in northwestern South America, bordered by Colombia on the north, Peru on the east and south, and the Pacific Ocean on the west. It also includes the Galápagos Province which contain ...

– the only larger region for which comprehensive data exists – more than a dozen species are known to be on the brink of

extinction Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

. 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 An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...

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

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 artisans at Spirit Cave,

Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...

. It is likely that these plants were collected from the wild rather than deliberately grown. Use of peppercorns as pungent spice is significant on an international scale. By

classical antiquity Classical antiquity, also known as the classical era, classical period, classical age, or simply antiquity, is the period of cultural History of Europe, European history between the 8th century BC and the 5th century AD comprising the inter ...

, there was a vigorous trade of spices including black pepper ('' P. nigrum'') from

South Asia South Asia is the southern Subregion#Asia, subregion of Asia that is defined in both geographical and Ethnicity, ethnic-Culture, cultural terms. South Asia, with a population of 2.04 billion, contains a quarter (25%) of the world's populatio ...

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

. The '' Apicius'', a recipe collection complied about 400 AD, mentions "pepper" as a spice for most main dishes. In the late

Roman Empire The Roman Empire ruled the Mediterranean and much of Europe, Western Asia and North Africa. The Roman people, Romans conquered most of this during the Roman Republic, Republic, and it was ruled by emperors following Octavian's assumption of ...

, black pepper was expensive, but was available readily enough to be used more frequently than

salt In common usage, salt is a mineral composed primarily of sodium chloride (NaCl). When used in food, especially in granulated form, it is more formally called table salt. In the form of a natural crystalline mineral, salt is also known as r ...

sugar Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose Glucose is a sugar with the Chemical formula#Molecular formula, molecul ...

. As Europe moved into the

Early Middle Ages The Early Middle Ages (or early medieval period), sometimes controversially referred to as the Dark Ages (historiography), Dark Ages, is typically regarded by historians as lasting from the late 5th to the 10th century. They marked the start o ...

, trade routes deteriorated and the use of pepper declined somewhat, but peppercorns, storing easily and having a high mass per volume, never ceased to be a profitable trade item. In the

Middle Ages In the history of Europe, the Middle Ages or medieval period lasted approximately from the 5th to the late 15th centuries, similarly to the post-classical period of global history. It began with the fall of the Western Roman Empire and ...

, international traders were nicknamed ''Pfeffersäcke'' ("pepper-sacks") in German towns of the

Hanseatic League The Hanseatic League was a Middle Ages, medieval commercial and defensive network of merchant guilds and market towns in Central Europe, Central and Northern Europe, Northern Europe. Growing from a few Northern Germany, North German towns in the ...

and elsewhere. Later, wars were fought by European powers, between themselves and in complex alliances and enmities with

Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or approximately 20% of the water area of Earth#Surface, Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia (continent), ...

states, in part about control of the supply of spices, perhaps the most

archetypal The concept of an archetype ( ) appears in areas relating to behavior, History of psychology#Emergence of German experimental psychology, historical psychology, philosophy and literary analysis. An archetype can be any of the following: # a stat ...

being black pepper fruit. Today, peppercorns 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. Long pepper (''P. longum''), is possibly the second-most popular ''Piper'' spice internationally; it has a rather chili-like "heat" and the whole

is used as the fruits are tiny. Cubeb (''P. cubeba''), also known as tailed pepper, played a major role in the spice trade. Reputedly Philip IV of Spain suppressed trade in cubeb peppercorns at the end of the 1630s to capitalize on his share of the black pepper trade. It remains a significant spice around the

region today, however. West African pepper ('' P. guineense''), is commonly used in West African cuisine, and is sometimes used in the

East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...

n '' berbere'' spice mix. This species, despite being traded more extensively in earlier times, is less common outside Africa today. Not only the seeds of ''Piper'' are used in cooking. West African Pepper leaves, known locally as ''uziza'', are used as a flavoring vegetable in

Nigerian Nigerians or the Nigerian people are citizens of Nigeria or people with ancestry from Nigeria. The name Nigeria was derived from the Niger River running through the country. This name was allegedly coined in the late 19th century by British jo ...

stews. In Mexican-influenced cooking, ''hoja santa'' or Mexican pepperleaf ('' P. auritum'') has a variety of uses. In

Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...

, leaves of two species of ''Piper'' have major importance in cooking: lolot (''P. lolot'') is used to wrap meat for grilling in the Indochina region, while wild betel ('' P. sarmentosum'') is used raw or cooked as a vegetable in Malay and Thai cuisine; The stems and roots of '' Piper chaba'' are used as a spice in Bangladeshi cuisine.

As medicine

Cubeb (''P. cubeba'') has been used in folk medicine and herbalism as well as, particularly in the early 20th century, as a

cigarette A cigarette is a narrow cylinder containing a combustible material, typically tobacco, that is rolled into Rolling paper, thin paper for smoking. The cigarette is ignited at one end, causing it to smolder; the resulting smoke is orally inhale ...

flavoring. '' P. darienense'' is used medically by the Guna people of the

Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...

border region, and elsewhere it is used to intoxicate fish which then can be easily caught. Spiked pepper, often called ''matico'' appears to have strong disinfectant and

antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting pathogenic bacteria, bacterial infections, and antibiotic medications are widely used in the therapy ...

properties. Black pepper (''P. nigrum'') essential oil is sometimes used in herbalism, and long pepper (''P. longum'') is similarly employed in Ayurveda, where it was an ingredient of ''Triphala Guggulu'' and (together with black pepper) of ''Trikatu'' pills, used for '' rasayana'' (rejuvenating and detoxifying) purposes. One ''Piper'' species has gained large-scale use as a

stimulant Stimulants (also known as central nervous system stimulants, or psychostimulants, or colloquially as uppers) are a class of drugs that increase alertness. They are used for various purposes, such as enhancing attention, motivation, cognition, ...

. Betel (''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, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...

. Conversely, another ''Piper'' species,

kava Kava or kava kava (''Piper methysticum'': Latin 'pepper' and Latinized Ancient Greek, Greek 'intoxicating') is a plant in the Piperaceae, pepper family, native to the Pacific Islands. The name ''kava'' is from Tongan language, Tongan and Marqu ...

(''P. methysticum''), is used for its depressant and euphoriant effects. In the Pacific region, where it has been widely spread as a canoe plant, kava is used to produce a calming and socializing drink somewhat similar to alcohol and benzodiazepines but without many of the negative side effects and less of an addiction risk. It has also become popular elsewhere in recent decades, and is used as a medical plant. However, pills that contain parts of the whole plant have occasionally shown a strong hepatotoxic effect, which has led to the banning of kava 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,

molecular biology Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...

, natural products

chemistry Chemistry is the scientific study of the properties and behavior of matter. It is a physical science within the natural sciences that studies the chemical elements that make up matter and chemical compound, compounds made of atoms, molecules a ...

, community ecology, and

evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes such as natural selection, common descent, and speciation that produced the diversity of life on Earth. In the 1930s, the discipline of evolutionary biolo ...

''Piper'' is a model genus for research in ecology and evolutionary biology. The diversity and ecological importance of the genus makes it a strong candidate for ecological and evolutionary studies. Most research has focused on the economically important species ''P. nigrum'' (black pepper), ''P. methysticum'' (kava), and ''P. betle'' (betel). A recent study based on DNA sequence analysis suggest that ''P. nigrum'' originated in the Western Ghats hot spot in India. The obligate and facultative

mutualists found in some ''Piper'' species have a strong influence on their biology, making them ideal systems for research on the evolution of symbioses and the effect of mutualisms on biotic communities. Important secondary metabolites found in pepper plants are piperine and chavicine, which were first isolated from Black Pepper, and reported to have antibiotic activities. Preliminary research reports has shown that piperine has an antibacterial activity against various bacteria such as ''S. aureus'', ''Streptococcus mutans'', and gastric cancer pathogen ''Helicobacter pylori'' and decreased ''H. pylori'' toxin entry to gastric epithelial cells. The piperidine functional group is named after the former, and piperazine (which is not found in ''P. nigrum'' in noticeable quantities) was in turn named after piperidine. The significant secondary metabolites of kava are

kavalactone Kavalactones are a class of lactone compounds found in kava roots and ''Alpinia zerumbet'' (shell ginger) and in several Gymnopilus, Phellinus and Inonotus fungi. Some kavalactones are bioactive. They are responsible for the psychoactive, analges ...

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

Repelling insects

Studies have been done to determine the effectiveness of piper leaves to repel different types of insects. Capuchin monkeys have been recorded by BBC Earth rubbing the piper leaves on them to repel insects.

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 In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

; the affinity of the African species is unclear. Some species are sometimes segregated into the genera ''Pothomorphe'', ''Macropiper'', ''Ottonia'', ''Arctottonia'', ''Sarcorhachis'', ''Trianaeopiper'', and ''Zippelia'', but other sources keep them in ''Piper''. The species called "''Piper aggregatum''" and "''P. fasciculatum''" are actually '' Lacistema aggregatum'', a plant from the family Lacistemataceae.

Notes

References

External links

The families of flowering plants
descriptions, illustrations, identification, information retrieval.

{{Authority control Pantropical flora Piperales genera