Piper (genus)

Piper
Lacquered Pepper (P. magnificum) inflorescence
Scientific classification
Kingdom:	Plantae
(unranked):	Angiosperms
(unranked):	Magnoliids
Order:	Piperales
Family:	Piperaceae
Genus:	Piper L.
Species
Some 1,000-2,000, see text
Synonyms
Anderssoniopiper Trel. Arctottonia Trel. Artanthe Miq. Chavica Miq. Discipiper Trel. & Stehlé Lepianthes Raf. Lindeniopiper Trel. Macropiper Miq. Ottonia Spreng. Pleiostachyopiper Trel. Pleistachyopiper Trel. Pothomorphe Miq. Trianaeopiper Trel.

Piper, the pepper plants or pepper vines (a term used for certain Clematis in older times), are an economically and ecologically important genus in the family Piperaceae. It contains about 1,000-2,000 species of shrubs, herbs, and lianas, many of which are keystone 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 angiosperms but neither monocots nor eudicots. Their family, Piperaceae, 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 inflorescences 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 Piper and the common name "pepper" are derived from the Sanskrit term pippali, denoting the Long Pepper (P. longum).

Distribution and ecology

Piper species have a pantropical distribution, and are most commonly found in the understory of lowland tropical rainforests, but can also occur in clearings and in higher elevation life zones such as cloud forests; one species (Japanese Pepper, P. kadsura, from southern Japan and southernmost Korea) is subtropical and can tolerate light winter frost. Peppers are often dominant vegetation where they are found.

Most Piper species are either herbaceous or vines; some grow as shrubs or almost as small trees. A few species, commonly called "ant pipers" (e.g. Piper cenocladum), live in a mutualism with ants. 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 birds, but small fruit-eating mammals – 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 Homidiana subpicta or some flea beetles of the genus Lanka. The latter can be significant pests to pepper growers.

Piper and humans

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

Unsustainable logging of tropical primary forests is threatening a number of peppers. The extent of the effect of such wholesale habitat destruction on the genus is unknown, but it is to be noted that in the forests of Ecuador – the only larger region for which comprehensive data exists^[1] – 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 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 artisans at Spirit Cave, Vietnam. It is not too likely that these plants were deliberately grown rather than collected from the wild.^[2]

Black Pepper (Piper nigrum) corns, from left to right:
Green (pickled ripe fruits)
White (dried ripe seeds)
Black (dried unripe fruits)

Use of peppercorns as pungent spice is significant on an international scale. In classical antiquity, there was a vigorous trade of spices including Black Pepper (P. nigrum) from South Asia to Europe already. The Apicius, a recipe collection complied about 400 AD, mentions "pepper" as a spice for most main dishes.^{[citation needed]} Judging from the Apicius, in the late Roman Empire, Black Pepper was probably still expensive, but nevertheless seems to have been available readily enough to be used more frequently than salt^[3] or sugar.^[4] As Europe turned into the Early 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, international traders were nicknamed Pfeffersäcke ("pepper-sacks") in German towns of the Hanseatic League 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 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 (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. But reputedly Philip IV of Spain at the end of the 1630s suppressed trade in Cubeb peppercorns to capitalize on his massive share of the Black Pepper trade.^[5] 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 region today, and popular as far as Morocco where it flavors spicy-sweet markout pastry and is sometimes included in the famous ras el hanout mix. Further east, it is well-loved across Indonesia, where it is the popular pepper to use in gulé (curried stews).

West African Pepper (P. guineense), also given a variety of "regionalized" names like "Benin pepper", "Ashanti pepper, or the rather ambiguous "Guinea pepper", is considered supreme for use in stews and other regional specialties. It is used in addition to Black Pepper to impart a particularly refined aroma with hints of nutmeg and saffron. Sometimes it is used in the East African berbere 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 (P. longum), commonly traded under its Indian name pipalli. This is possibly the secondmost popular Piper spice internationally; it has a rather chili-like "heat" and the whole inflorescence 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 Nigerian stews. In Mexican-influenced cooking, hoja santa or Mexican Pepperleaf (P. auritum) has a variety of uses, mainly to impart flavor too. In Southeast Asia, 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), often seen under the Thai name cha phlu, is used raw or cooked as a vegetable in its own right in Malay and Thai cuisine;^[6] it is also used to prepare the famous miang kham snacks of northwestern Thailand.

As medicine and stimulant

At a Kava club in Tonga

Cubeb (P. cubeba) has been used in folk medicine, herbalism and related fields (e.g. alchemy, folk magic or exorcism), as well as, particularly in the early 20th century, as a cigarette flavoring. P. darienense is used medically by the Kuna people of the Panama-Colombia border region, and elsewhere it is used to intoxicate fish which then can be easily caught. Spiked Pepper, often called "matico",^[7] appears to have strong disinfectant and antibiotic 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.

See also Betel leaf, Kava culture.

Two Piper species have gained large-scale use as a stimulant. 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. In the Pacific region, where it has been widely spread as a canoe plant, Kava (P. methysticum) is used to produce a calming drink somewhat similar to alcohol but without many of the negative side effects and addiction 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.^[8]

In science

The genus contains species suitable for studying natural history, molecular biology, natural products chemistry, community ecology, and evolutionary biology.^[9]

Crystallized piperine, extracted from Black Pepper (Piper nigrum)

Piper is a model genus for research in ecology and evolutionary biology. 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), P. methysticum (Kava), and P. betle (Betel).

The obligate and facultative ant 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. 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 kavalactones and flavokawains. Pipermethystine 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.^[10]

Some species are sometimes segregated into the genera Pothomorphe, Macropiper, Ottonia, Arctottonia, Sarcorhachis, Trianaeopiper, and Zippelia, but other sources keep them in Piper.^[10]

Piper curtipedunculum

Piper dilatatum

Piper friedrichsthalii

Piper grandifolium

Piper guanacostense

Piper pseudofuligineum

Piper rivioides

Piper achupallasense Piper aduncum – Spiked Pepper, "matico" (South America except Ecuador) Piper aequilaterum C.DC. Piper amalgo – mecaxochitl (Nahuatl) Piper angamarcanum Piper auritum – Mexican Pepperleaf, Sacred Pepper, root beer plant, hoja santa or "yerba santa" (Mexican Spanish) Piper azuaiense Piper baezanum Piper baezense Piper begoniiforme Piper betle – Betel, vilya or veeleya (Kannada), vettila (Malayalam), vidyache pan (Marathi), nagavalli or taambuul (Sanskrit), vetrilai (Tamil) Piper borbonense – Voatsiperifery Pepper, Madagascan wild pepper Piper borneense Piper brachipilum Piper brachystylum Piper bullatifolium Piper caldense C.DC. Piper campii Piper caninum – Common Pepper, Native Pepper (Australia) Piper candollei Piper cenocladum – "ant piper" Piper chaba Piper chimborazoense Piper chimonanthifolium Kunth Piper clathratum Piper claussenianum Piper coeloneurum Piper cordulatum Piper cubeba – Cubeb, Java Pepper, Tailed Pepper Piper curtipedunculum Piper cutucuense Piper darienense – duermeboca (Panama), kana (Kuna) Piper decurrens Piper densiciliatum Piper diffundum Piper dilatatum Piper disparipilum Piper distigmatum Piper dodsonii Piper entradense Piper eriocladum Piper eustylum Piper fallenii Piper fimbriulatum Piper friedrichsthalii Piper gaudichaudianum Kunth Piper glabratum Kunth Piper grandifolium Piper gualeanum Piper guanacostense Piper guayasanum Piper guineense – West African Pepper, Ashanti Pepper, Benin Pepper, False Cubeb, Guinea Pepper, Guinea Cubeb, Uziza Pepper, kale, kukauabe, masoro, sasema, soro wisa Piper hillianum C.DC. Piper hispidinervum Piper huigranum Piper hydrolapathum Piper hylebates Piper hylophilum Piper kadsura – Japanese Pepper, fūtōkazura (Japanese) Piper imperiale Piper laevigatum Piper lhotzkyanum Kunth Piper lineatipilosum Piper lolot – Lolot, lá lốt (Vietnamese) Piper longicaudatum Piper longum – Long Pepper, Indian Long Pepper, pipalli (Hindi), pippali (Sanskrit)

Piper lucigaudens Piper macrophyllum Piper magnificum – Lacquered Pepper Piper malacophyllum (C.Presl) C.DC. Piper manabinum Piper marginatum Piper mekongense Piper mendezense Piper mestonii Piper metallicum Piper methysticum – Kava, yaqona (Fijian), ʻawa (Hawaiian), sakau (Pohnpeian), 'ava (Samoan) Piper mexiae Piper molliusculum Piper mullesua Piper nanegalense Piper napo-pastazanum Piper nebuligaudens Piper nigrum – Black Pepper, white pepper, green pepper, pamintâ (Philippines) Piper novae-hollandiae – Giant Pepper Vine Piper ornatum – Celebes Pepper Piper pachyphyllum – Mahalatsaka Piper peltatum Piper permucronatum Yunck. Piper perstrigosum Piper platylobum Piper porphyrophyllum Piper poscitum Piper prietoi Piper productispicum Piper pseudofuligineum Piper pseudonigrum Piper pseudopothifolium C.DC. Piper puyoense Piper regale Piper retrofractum – Balinese Long Pepper, Javanese Long Pepper, Indonesian Long Pepper Piper richardiifolium Kunth Piper rivioides Piper rubronodosum Piper rubrovenosum Piper sagittifolium Piper saigonense – Saigon Pepper Piper saloyanum Piper san-miguelense Piper sarmentosum – Wild Betel, phak i leut (Lao), pokok kadok (Malay), cha phlu (Thai) Piper schuppii Piper seychellarum – Seychelles Pepper Piper skutchii Piper sodiroi Piper stipulosum Piper subaduncum Piper subnitidifolium Piper supernum Piper sylvaticum – Mountain Long Pepper Piper tectonifolium Kunth Piper trachyphyllum Piper translucens Yunck. Piper triandrum F.Muell. Piper trioicum Piper truman-yunckeri Piper truncatum Vell. Piper umbellatum Piper unguiculatum Piper valladolidense Piper verrucosum Piper warakahoura Piper wibomii Piper wightii Piper zarumanum

Piper aggregatum and P. fasciculatum are actually Lacistema aggregatum, an entirely unrelated angiosperm from the family Lacistemataceae.

Footnotes

1. ^ IUCN (2007)
2. ^ German (1969, 1971)
3. ^ Apicius generally uses garum fish sauce instead; raw brine and large quantities of herbs were also employed by many.
4. ^ Crystal sugar was about as expensive as precious metals and even to the wealthiest people less available than diamonds are today. Apicius uses honey and caroenum (grape must concentrate).
5. ^ Parkinson (1640)
6. ^ Salomon (1998), Ecocyber (2008)
7. ^ Except in Ecuador, where this name refers to the unrelated Aristeguietia glutinosa.
8. ^ Sorrentino et al. (2006), Lim et al. (2007)
9. ^ Dyer & Palmer (2004)
10. ^ ^{a b} Jaramillo & Manos (2001)

References

• Dyer, L.A. & Palmer, A.N. (2004): Piper: A model genus for studies of evolution, chemical ecology, and trophic interactions. Kluwer Academic Publishers, Boston. ISBN 0-306-48498-6 PDF fulltext
• Ecocyber (2008): Tropical Root Crops. Retrieved 2008-JUL-15.
• Gorman, Chester F. (1969): Hoabinhian: A pebble tool complex with early plant associations in Southeast Asia. Science 163(3868): 671-673. doi:10.1126/science.163.3868.671 (HTML abstract)
• Gorman, Chester F. (1971): The Hoabinhian and After: Subsistence Patterns in Southeast Asia during the Late Pleistocene and Early Recent Periods. World Archaeology 2(3): 300-320. HTML abstract and first page image
• International Union for the Conservation of Nature and Natural Resources (IUCN) (2007): 2007 IUCN Red List of Threatened Species. IUCN, Gland.
• Jaramillo, M. Alejandra & Manos, Paul S. (2001): Phylogeny and Patterns of Floral Diversity in the Genus Piper (Piperaceae). Am. J. Bot. 88(4): 706-716. doi:http://www.amjbot.org/cgi/content/full/88/4/706 (Abstract and first page image)
• Lim, S.T.; Dragull, K.; Tang, C.S.; Bittenbender, H.C.; Efird, J.T. & Nerurkar P.V. (2007): Effects of Kava Alkaloid, Pipermethystine, and Kavalactones on Oxidative Stress and Cytochrome P450 in F-344 Rats. Toxicological Science 97(1): 214-221. doi:10.1093/toxsci/kfm035 PMID 17329236 PDF fulltext
• Parkinson, John (1640): Theatrum Botanicum: the Theater of Plants, etc. T. Cotes, London.
• Solomon, Charmaine (1998): Cha Plu. In: Encyclopedia of Asian Food. Periplus Editions. HTML fulltext
• Sorrentin, L.; Capasso, A. & Schmidt, M. (2006): Safety of ethanolic kava extract: Results of a study of chronic toxicity in rats. Phytomedicine 13(8): 542-549. doi:10.1016/j.phymed.2006.01.006 PMID 16904878 (HTML abstract)

External links

• The families of flowering plants: descriptions, illustrations, identification, information retrieval.
• Piperaceae in the Flora of North America
• Piperaceae in the Flora of China
• NCBI Taxonomy Browser
• Piper species in Thailand