The Piperaceae (), also known as the pepper family, are a large

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

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

s. The group contains roughly 3,600 currently accepted species in five genera. The vast majority of species can be found within the two main genera: '' Piper'' (2,171 species) and '' Peperomia'' (over 1,000 species). Members of the Piperaceae may be small trees, shrubs, or herbs. The distribution of this group is best described as

pantropical A pantropical ("all tropics") distribution is one which covers tropical regions of both the Eastern and Western hemispheres. Examples of species include caecilians, modern sirenians and the plant genera ''Acacia'' and ''Bacopa''. ''Neotropical' ...

. The best-known species, '' Piper nigrum'', yields most peppercorns that are used as spices, including

black pepper Black pepper (''Piper nigrum'') is a flowering vine in the family Piperaceae, cultivated for its fruit (the peppercorn), which is usually dried and used as a spice and seasoning. The fruit is a drupe (stonefruit) which is about in diameter ...

, although its relatives in the family include many other spices. The family Piperaceae is unrelated to the family

Solanaceae Solanaceae (), commonly known as the nightshades, is a family of flowering plants in the order Solanales. It contains approximately 2,700 species, several of which are used as agricultural crops, medicinal plants, and ornamental plants. Many me ...

, which includes

bell peppers The bell pepper (also known as sweet pepper, paprika, pepper, capsicum or, in some parts of the US midwest, mango) is the fruit of plants in the Grossum Group of the species ''Capsicum annuum''. Cultivars of the plant produce fruits in diff ...

and

chili peppers Chili peppers, also spelled chile or chilli ( ), are varieties of berry-fruit plants from the genus ''Capsicum'', which are members of the nightshade family Solanaceae, cultivated for their pungency. They are used as a spice to add pungency ( ...

, which are so named due to

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 ...

ans taking part in the

Columbian exchange The Columbian exchange, also known as the Columbian interchange, was the widespread transfer of plants, animals, and diseases between the New World (the Americas) in the Western Hemisphere, and the Old World (Afro-Eurasia) in the Eastern Hemis ...

mistakenly believing the spicy fruits were a variety of the black pepper plant.

Etymology

The name Piperaceae is 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'', . Piper-caninum-SF21234-01

Taxonomy

The

APG III system The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a f ...

of 2009 recognizes this family, and assigns it to the order Piperales in the unranked clade

magnoliids Magnoliids, Magnoliidae or Magnolianae are a clade of flowering plants. With more than 10,000 species, including magnolias, nutmeg, bay laurel, cinnamon, avocado, black pepper, tulip tree and many others, it is the third-largest group of angiospe ...

. The family consists of five genera: '' Piper'', '' Peperomia'', '' Zippelia'', '' Manekia'', and '' Verhuellia''. The previously recognised Pacific genus '' Macropiper'', was recently merged into ''Piper''.Wanke, S., Jaramillo, M. A., Borsch, T., Samain, M.-T., Quandt, D., and Neinhuis, C. (2007) "Evolution of Piperales—''matK'' gene and ''trnK'' intron sequence data reveal lineage specific resolution contrast". ''Molecular Phylogenetics and Evolution'' 42: 477–497. A tentative cladogram showing relationships based on Wanke et al. (2007)Wanke, S., Vanderschaeve, L., Mathieu, G., Neinhuis, C., Goetghebeur, P., and Samain, M.S. (2007) "From Forgotten Taxon to a Missing Link? The Position of the Genus ''Verhuellia'' (Piperaceae) Revealed by Molecules". ''Annals of Botany'', 99: 1231–1238. is shown below. This phylogeny was based on 6,000

base pairs A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...

chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...

DNA. Only recently has it become clear that ''Verhuellia'' is sister to the other four genera in the family.Samain et al. (2010) "''Verhuellia'' is a segregate lineage in Piperaceae: more evidence from flower, fruit and pollen morphology, anatomy and development". ''Annals of Botany'', 105.

Characteristics

Members of pepper family are small trees, shrubs, or

perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...

or annual herbs.

Roots and stems

Plants are often

rhizomatous In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...

, and can be terrestrial or

epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...

. The stems can be either simple or branched.

Leaves

Leaves are simple with entire margins, and are positioned at the base of the plant or along the stem, and can be alternate, opposite, or whorled in arrangement.

Stipules In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). They are primarily found among dicots and rare among monocots. Stipules are considered part of the anatomy ...

are usually present, as are petioles. The leaves are often noticeably

aroma An odor (American English) or odour (Commonwealth English; see spelling differences) is a smell or a scent caused by one or more volatilized chemical compounds generally found in low concentrations that humans and many animals can perceive v ...

tic when crushed.

Flowers

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 (in the form of spikes) are terminal, opposite the leaves, or located in the axils. Flowers are bisexual, with no

perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower. It is a structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepal ...

, each flower is subtended by a peltate

bract In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...

. Stamens are 2–6, and hypogynous, with 2-locular

anther The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...

s. There are usually 3-4 stigmas attached to a single

pistil Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl (botany), whorl of a flower; it consists ...

per flower, which is 1 or 3-4

carpel Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more ...

late. The ovary is 1 locular, and superior.

Fruits and seeds

Fruits are

drupe In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or ''pyrena'') of hardened endocarp with a seed ...

like, with a single

seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...

per fruit. The seeds have a minute

embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...

, and mealy

perisperm In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fema ...

.Boufford, D.E. (1997)). Flora of North America - Piperaceae. http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10692l

Traditional medicinal uses

Numerous members of the Piperaceae family are used in the traditional medicinal systems of indigenous population for a wide variety of illnesses. Many studies have been undertaken to investigate these uses, with a large number of them focusing especially on the active ingredient

Piperine Piperine, possibly along with its isomer chavicine, is the compound responsible for the pungency of black pepper and long pepper via activation of TRPV1. It has been used in some forms of traditional medicine. Preparation Extraction Due t ...

and related compounds found in many members of this family, especially

Black pepper Black pepper (''Piper nigrum'') is a flowering vine in the family Piperaceae, cultivated for its fruit (the peppercorn), which is usually dried and used as a spice and seasoning. The fruit is a drupe (stonefruit) which is about in diameter ...

Long pepper Long pepper (''Piper longum''), sometimes called Indian long pepper or ''pippali'', 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 taste sim ...

and

Betel Betel (''Piper betle'') is a species of flowering plant in the pepper family Piperaceae, native to Southeast Asia. It is an evergreen, dioecious vine, with glossy heart-shaped leaves and white catkins. Betel plants are cultivated for their lea ...

, as well as

kavalactones 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, analg ...

found in

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 ...

References

External links

Plants of the World Online, ''Piperaceae''

Angiosperm Phylogeny Website

Piperaceae at the online ''Flora of North America''

Piperaceae at the online ''Flora of China''

Piperaceae at the online ''Flora of Zimbabwe''

Piperaceae at the NCBI Taxonomy Browser

{{Authority control Magnoliid families Pantropical flora