*Elsie Franklin Guimarães and **Daniele Monteiro
*Jardim Botânico do Rio de Janeiro, Brazil.** Universidade Federal de Juiz de Fora, Brazil.
Herbs (terrestrial, epiphytic or living on rocks) annual or perennial subshrubs or shrubs, less often lianas; aromatic, glabrous or with varied indumentum, frequently gland -dotted, with nodose stems. Leaves mostly alternate, sometimes opposite or whorled, simple, entire, sessile or petiolate, sometimes peltate, with variable size, shape and venation; prophyll (in Piper) persistent or deciduous and commonly united with petiole. Inflorescence terminal, leaf-opposed or axillary, usually fleshy, pedunculate, racemose or more commonly spike-like, solitary, umbellate or paniculate, erect, pendent or recurved, variable in size, white, cream, green or sometimes rose or reddish; rachis smooth, papillate or villous. Flowers very small, bisexual (all New World species), withouth perianth, numerous, each subtended by a single peltate or subpeltate bract, variable in shape, glabrous or with an indumentum; sessile or pedicellate; stamens 1-10, usually 2-6, arising near base of ovary with filaments free and generally short, anthers with 1, 2 or 4 thecae, laterally or apically dehiscent, deciduous after polinization; gynoecium with superior ovary, sessile or stipitate, with 1, 3 or 5 fused carpels, 1-locular with a single basal ovule, stigmas 1-5 sessile or stylose. Fruit a small berry or drupe, variously shaped, with a thin pericarp and sometimes hardened endocarp; seed small, solitary, endosperm scanty, perisperm abundant; embryo very small.
Notes on delimitation
- Piperaceae is placed in order the Piperales, one of the most species-rich and heterogeneous clades in the magnoliids, together with 4 families which also occur in the Neotropics: Saururaceae Rich. ex T. Lestib., Lactoridaceae Engl., Aristolochiaceae Juss. and Hydnoraceae C. Agardh.
- The family, considered sister of Saururaceae, includes about 3,000 species and has four genera distributed in two groups: Zippeliae consisting of Zippelia Blume and Manekia Trel., Pipereae comprising Piper L. and Peperomia Ruiz & Pav.
- The most detailed study of Peperomia (Dahlstedt 1900) divided the genus in nine subgenera and seven sections; however recent phylogenetic analyses only support the monophyly of two subgenera Micropiper and Shaerocarpidium (Samain et al. 2009).
- Some authors consider Peperomia and Manekia as a separate family - the Peperomiaceae, which also includes Verhuellia Miq. and Piperanthera DC. However, recent analyses show that Verhuellia is sister to all other Piperaceae (Wanke et al. 2007), that Manekia clearly is a separate genus (Arias et al. 2006) and that Piperanthera is synonymous with Peperomia (Wanke et al. 2006).
- Eight subgenenera of Piper are recognised in the Neotropics: Enckea Kunth (including Callianira Miq. and Arctottonia Trel.), Macrostachys Miq. (including Hemipodium Miq.), Ottonia Spreng., Peltobryon Klotzsch (including Trianaeopiper Trel.), Piper s.s., Pothomorphe Miq., Radula Miq. (including Isophyllon Miq.) and Schilleria Kunth (Jaramillo et al. 2008).
- The species circumscription of the neotropical Piper species is still unclear. Many taxa were described without reference to their position within the genus; many were incompletely described and some of their characteristics not evaluated; others resemble each other morphologically and may turn out to be synonymous once a considerable number of binomials have been evaluated.
- Finally, phylogenetic studies based on morphological and molecular data are also needed.
Distribution in the Neotropics
Piperaceae is pantropical, with major concentrations and centres of diversity in Central and South America and in Malaysia. The members of the family most frequently inhabit rain forests:
- Manekia: Haiti, Lesser Antilles (Guadalupe, Dominica, Martinique and St. Vincent), Southern Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, northern Peru and Brazil (Atlantic forest), growing in primary and secondary forest, from sea level to 2,000m.
- Peperomia: pantropical with the greatest diversity occurring in the Neotropics; the species are mostly found in humid forested regions and few in dry areas.
- Piper: pantropical with the greatest diversity in the Neotropics; the species occur in various habitats but are best represented in lowland tropical rainforests, frequently in formations below 1,500m elevation and also commonly found in disturbed areas.
Distinguishing characters (always present)
- Herbs, shrubs, subshrubs.
- Vegetative parts with spicy-aromatic odour when crushed.
- Leaves usually alternate, always simple and entire.
- Inflorescences in spikes or racemes.
- Flowers minute, without perianth, subtended by a small, variable bract.
- Ovary unilocular, with a single basal ovule.
Other important characters
- Small trees or lianas.
- Nodes prominently swollen (Piper).
- Prophyll present (Piper).
- Leaves opposite or whorled (Peperomia).
- Carpels 1 (Peperomia), 3 or 4 (5) fused (Piper and Manekia).
- Fruit a small berry or drupe.
Key differences from similar families
- Saururaceae differs from Piperaceae by having (2-)3-4 distinct or fused carpels (vs. 1), if distinct then each carpel with 2-4 ovules, if fused then ovary 1-locular and each carpel with 6-13 ovules, placentation parietal (vs. basal), and fruit dry, schizocarp or apically dehiscent capsule (vs. berry or drupe).
Number of genera
- 3 genera: Manekia (ca. 4 species), Piper and Peperomia (the two largest of the family in number of representatives; both with about 1,500 species).
Useful tips for generic identification
Key to genera of Neotropical Piperaceae
1. Herbs with alternate, opposite or whorled leaves; stamens 2; stigma 1 … Peperomia1. Shrubs or small trees, sometimes lianas or climbing shrubs with alternate leaves; stamens 2-6; stigmas 3-5 ... 2.
2. Shrubs or small trees, glabrous or indumented; inflorescence a solitary spike or raceme, sometimes a compound umbel of spikes, leaf-opposed or terminal … Piper2. Lianas, glabrous; inflorescence a solitary or paired spike, axillary or occasionally terminal … Manekia
Notable genera and distinguishing features
- Manekia: lianas or climbing shrubs, glabrous, sometimes with succulent stems. Leaves alternate, venation palmate; petioles vaginate for their entire length. Inflorescences spikes usually axillary, occasionally terminal, solitary or paired. Stamens 2-4. Ovary sessile with 3-4(-5) fused carpels and 3-4(-5) sessile stigmas. Fruit laterally compressed, immersed and coalescent with the somewhat fleshy rachis at maturity.
- Peperomia: herbs erect, prostate, pendent or assurgent, usually succulent, terrestrial, rock-living or epiphytic. Leaves alternate, opposite or whorled, sometimes clustered near the apex, usually petiolate and often yellowish or black gland-dotted; venation palmate, pinnate or plinerved. Inflorescence axillary, terminal or leaf-opposed, in spikes or racemes, solitary or paniculate. Stamens 2. Ovary sessile or stipitate, with 1 carpel and 1 apical or subapical stigma. Fruit in depression of the rachis or stipitate, commonly verrucose and viscid.
- Piper: shrubs, subshrubs, occasionally subherbaceous or small trees with nodose stems. Leaves alternate, sometimes scabrous and with the upper surface smooth, rugose or bullate and lacunose beneath, gland-dotted, petiole usually present; prophyll persisted or deciduous and commonly united with petiole; venation palmate or pinnate. Inflorescence a solitary spike or raceme (subg. Ottonia) leaf-opposed or terminal, erect, pendent or recurved, occasionally compound with simple spikes borne in an umbellate cluster on an axillary stalk (sug. Pothomorphe). Stamens (1) 2-6, usually 4. Ovary sessile or stipitate, with 3-4 fused carpels and 3-4 sessile or stylose stigmas. Fruit in depression of the rachis or stipitate sometimes laterally compressed, glabrous or with trichomes.
- Some species are widespread but several are restricted to specific centres of diversity and each country has a large number of endemics.
- Cultivated: Piper nigrum L.
- Many species of Piperaceae are rich in secondary metabolites, including alkaloids, amides, flavonoids and terpenes, having economic and medicinal importance: Piper nigrum L. (black pepper), Piper umbellatum L. (pariparoba) and Piper methysticum L. (kava-kava). Species of Peperomia are commonly used as ornamental house plants (Yuncker 1958; Milliken 1997).
- Generalist insects usually act as pollinators, being attracted by color and/or perfume, nectar or pollen (Figueiredo & Sazima 2000). Birds, bats and other animals help to spread the fruits (Yuncker 1958). Some species have interaction with ants (Tepe et al. 2007).
- Zippelia is a monotypic genus of Asian distribution, with floral morphology considered intermediate between Piperaceae and Saururaceae. This genus does not occur in the Neotropics.
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Tepe, E. J., Vincent, M.A. and Watson, L.E. 2007. Stem diversity, cauline domatia, and the evolution of ant-plant associations in Piper sect. Macrostachys (Piperaceae). American Journal of Botany 94: 1-11.
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Wanke, S., Vanderschaeva, 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.
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How to cite
Guimarães, E.F. & Monteiro, D. (2010). Neotropical Piperaceae. In: Milliken, W., Klitgård, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Piperaceae.htm.
Click images to enlarge
Leaves of Manekia obtusa © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Peperomia arifolia © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Peperomia corcovadensis © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Peperomia hispidulla © Marcelo Vianna-Filho, Museu Nacional-Rio de Janeiro.
Whorled leaves and terminal inflorescence of Peperomia megapotamica © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Inflorescences of Peperomia urocarpa © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Habit of Peperomia velloziana © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Leaf-opposed inflorescences of Piper abutiloides © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Piper belloi © Marcelo Vianna-Filho, Museu Nacional-Rio de Janeiro.
Axillary inflorescence of Piper duartei © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Leaf-opposed inflorescences of Piper hispidum © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Leaf-opposed inflorescences of Piper lhotzkyanum © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Piper solmsianum © Daniele Monteiro, Universidade Federal de Juiz de Fora.
Habit of Piper translucens © Daniele Monteiro, Universidade Federal de Juiz de Fora.