Capsicum baccatum
(pepper)

Pictures

Picture	Title	Caption	Copyright
Title Fruiting habit Caption Capsicum baccatum (pepper, bishop's hat, orchid pepper); fruiting habit. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014 Copyright ©Forest Starr & Kim Starr - CC BY 4.0	Fruiting habit	Capsicum baccatum (pepper, bishop's hat, orchid pepper); fruiting habit. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014	©Forest Starr & Kim Starr - CC BY 4.0
Title Fruiting habit Caption Capsicum baccatum (pepper, bishop's hat, orchid pepper); fruiting habit. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014. Copyright ©Forest Starr & Kim Starr - CC BY 4.0	Fruiting habit	Capsicum baccatum (pepper, bishop's hat, orchid pepper); fruiting habit. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014.	©Forest Starr & Kim Starr - CC BY 4.0
Title Fruits Caption Capsicum baccatum (pepper, bishop's hat, orchid pepper); close-up of fruits. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014. Copyright ©Forest Starr & Kim Starr - CC BY 4.0	Fruits	Capsicum baccatum (pepper, bishop's hat, orchid pepper); close-up of fruits. Pali o Waipio Huelo, Maui, Hawaii, USA. September, 2014.	©Forest Starr & Kim Starr - CC BY 4.0

Identity

Preferred Scientific Name

• Capsicum baccatum L.

Preferred Common Name

• pepper

Other Scientific Names

• Capsicum cerasiflorum Link
• Capsicum baccatum L. var. pendulum (Willd.) Eshbaugh
• Capsicum chamaecerasus Nees
• Capsicum ciliare Willd.
• Capsicum conicum Vell.
• Capsicum microcarpum Cav.
• Capsicum microphyllum Dunal
• Capsicum pulchellum Salisb.
• Capsicum umbilicatum Vell.

International Common Names

• English: aji; bishop's-hat; Christmas bell; Peruvian pepper
• French: piment chien

Local Common Names

• Bolivia: ají; arivivi
• Dominican Republic: ají bobito; ají bonito; ají caribe; ají montesino; ají tití; ajicito montesino
• Germany: peruanischer Pfeffer
• Haiti: piment zouézeau; piment zouézo; pimento z'oiseux
• Puerto Rico: ají caballero; ají pico de paloma
• Sweden: barpeppar
• United States Virgin Islands: wild pepper

Summary of Invasiveness

C. baccatum is a perennial plant listed as a ‘weed’ in the Global Compendium of Weeds (Randall, 2012), is reported to be invasive to Cuba (Oviedo-Prieto et al., 2012), and is reportedly a weed in Trinidad and Brazil (Holm et al., 1979; Randall, 2012). The species has been cultivated since pre-Columbian times in its native South America and later around the world, for use as a highly popular spice, vegetable, ornamental, and ingredient in such commodities as self defense pepper spray (Basu and De, 2003). The species reproduces by seeds encased in its famous chilli fruits, which are spread primarily by intentional and accidental biotic dispersal agents.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Solanales
•                         Family: Solanaceae
•                             Genus: Capsicum
•                                 Species: Capsicum baccatum

Notes on Taxonomy and Nomenclature

The genus Capsicum consists of all the ‘chilli pepper plants’ with 3-5 wild species and over 2000 cultivars (DeWitt and Bosland, 1996; Tewkesbury et al., 2006), and the confusing terminology ‘chilli’ is often used frequently and interchangeably with other names including ‘chile’, ‘aji’, and ‘paprika’ referring to multiple species (Basu and De, 2003). The genus name Capsicum derives from a Greek-based derivative of the latin word ‘kapto’, meaning ‘to bite’, in reference to the heat or pungency of the species’ fruit (Basu and De, 2003), although it has also been speculated to derive from the Latin word ‘capsa’, a box, referring to the shape of the fruit in forms of the typical species (Britton, 1918). The common name ‘chile’ is a variation of ‘chil’, derived from the Nahutal (Aztec) dialect (Basu and De, 2003).

The number of global species within the Capsicum genus has long been subject to debate, but there are presently considered to be five domesticated species of Capsicum, the primary distinguishing characteristics being flower and seed colour, shape of the calyx, number of flowers per node and their orientation; these five species are C. annuum, C. baccatum var. pendulum, C. frutescens, C. chinense, and C. pubescens (DeWitt and Bosland, 1996; Hawkes et al., 1979; Basu and De, 2003).

C. baccatum was first described by Linnaeus in 1753, though this name has also been misapplied to a number of different taxa. The type material of C. baccatum can be matched with material found in the wild today; however, in his original description in the Species Plantarum, Linnaeus failed to mention the presence of white flowers with yellow corolla markings that are apparent in the type material. This led a number of workers to rely entirely on plant habit and fruit shape for their association of living plant material with the original description (Eshbaugh, 1970).

Historically, C. baccatum has been separated into the two species, C. microcarpum and C. pendulum, a separation based primarily on differences in the fruit characters. Today, separation of the wild and cultivated taxa is weak and is maintained primarily through geographic isolation outside the range of wild C. baccatum and by agricultural isolation within the overlap zone of the two varieties (Eshbaugh, 1970).

The species C. baccatum has both a wild form, C. baccatum var. baccatum, and a cultivated subspecies, C. baccatum var. pendulum; the cultivated form is widespread throughout tropical regions in South America, while the wild form is more restricted but ranges from Peru to Brazil (Basu and De, 2003). Eshbaugh (1970) described the qualitative characters of fruit colour, fruit position, and fruit persistence as easily distinguishing the two varieties of C. baccatum. C. baccatum var. baccatum has red, erect, and non-persistent fruits, and C. baccatum var. pendulum has red, orange, yellow, green, or brown fruits which are pendent and persistent. Basu and De (2003) later reported that the two are morphologically indistinguishable, with identical flavonoid and isoenyzme profiles, except for the organ size differences in var. pendulum.

An AFLP study of C. baccatum accessions from South America did not support taxonomic distinction of C. baccatum var. umbilicum from C. baccatum var. pendulum (Albrecht et al., 2012). A clustering analysis of the same data suggested that C. baccatum likely originated in Paraguay. The Plant List (2016) listed only 2 accepted names for infraspecific taxa of C. baccatum: var. pendulum (Willd.) Eshbaugh and var. praetermissum (Heiser & P.G. Sm.) Hunz.

Description

Erect or scrambling, often much branched, perennial herb to sappy shrub, up to 4.5 m (but often less). Branches ± angular, slightly striate, densely pubescent when young to ± glabrous. Leaves usually solitary, rarely 2 appearing together; petiole 0.3–3 cm long; lamina membranous, 1.5–7.5(10) × 0.8–4(4.5) cm, ovate to ovate-lanceolate, base rounded to narrowly cuneate, and often unequal-sided, apex long-acuminate, entire, ± ciliate, with scattered hairs, sometimes only along the nerves, paler and with a few hairs in the axils of the nerves beneath. Flowers 2-whorled, rarely solitary; pedicels 8-15 mm long, angular, striate, thickened upwards, scarcely pubescent to ± glabrous, ± erect or curved; in fruit elongated to 30 mm and slender. Calyx 2.5-3 mm long, shortly cupular, 5-ribbed, 5-dentate, subglabrous; teeth 0.5-0.8 mm long, apically thickened and somewhat obtuse to ± subulate, spreading; in fruit enlarged and surrounding the base of it. Corolla greenish-white to dirty-white, rotate-campanulate; limb 8-9 mm across; lobes 1.5->2.5 mm long, ovate-oblong or ± triangular, obtuse or slightly acuminate, ciliolate. Filaments 1-1.5 mm long; anthers yellow, 1.7-1.9 mm long, oblong. Ovary c. 1 mm long, ± ovoid, rounded distally, glabrous; style 3.5 mm long, slightly thickened into a small stigma. Fruit scarlet, erect, glossy, (7)9-10 × (5)6-7 mm, globose-ovoid or broadly ellipsoid, rounded distally, smooth, glabrous. Seeds pale brownish, 3.5-4 × 2.5-3 mm, ovate in outline or ± reniform. [taken from Flora Zambesiaca (2014) for wild species, C. baccatum var. baccatum].

The domesticated form, var. pendulum, of this lowland South American species has cream-coloured flowers with paired gold or green markings. Typically, fruit are elongate with cream-coloured seeds.

Plant Type

Distribution

The species C. baccatum originated from Peru and southern Bolivia with varying distributions; the domesticated form var. pendulum spans across tropical South America, while the wild var. baccatum is less widespread and ranges from Peru to Brazil (Eshbaugh, 1970; Basu and De, 2003). According to Russo (2012), the wild taxon is common in Bolivia and northern Argentina with outlier populations in Peru and Paraguay.

The species is included in the Vascular Plants of Ecuador (2014) which, however, notes its presence in the country is "questionably reported from the Galapagos Islands on the basis of a single collection" (Wiggins and Porter, 1971).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

Among the first domesticated plants of Mesoamerica, Capsicum has been known since the beginning of civilization in the Western hemisphere and has been part of the human diet since 7500 BC (Basu and De, 2003). It was either Christopher Columbus or his accompanying physician Chanca who first reported the use of Capsicum in the Americas to Europe around 1493-1494 and certainly Columbus who introduced it across the Atlantic; by mid-17^th century Capsicum was being cultivated throughout southern and middle Europe as a spice and medicinal drug, with introductions of one species to Japan and five to India (for mass cultivation in the colonies, from the Portuguese) around this time (Basu and De, 2003). At the time of Cristopher Columbus’ discovery of the Americas, the cultivated form of C. baccatum var. pendulum was only found in areas east and west of the Andes (Russo, 2012). Today the cultivated form, C. baccatum var. pendulum, has worldwide distribution and is reportedly the most widely domesticated pepper of Peru, even over C. annuum (Russo, 2012). The cultivated species has also been described as the most consumed species in Brazil (Spiller et al., 2008).

In the West Indies, Capsicum had been introduced to Jamaica by 1871, as Macfadyen observed the use of Capsicum fruit by Caribbean people as a food and drink condiment, but the plant is not mentioned by species (Macfadyen, 1871). The species was reported by name in Puerto Rico in 1881, during which Bello observed many forms being cultivated for culinary uses (Bello Espinosa, 1881). The species was present in Bermuda in 1918, as Britton reported it to be an occasional native plant occurring in rocky woodlands, and occasionally in gardens (Britton, 1918). The species can now be found across the Cayman Islands, Hispaniola, Jamaica, Puerto Rico, and the Virgin Islands, and is now considered an invasive introduction to Cuba (Oviedo-Prieto et al., 2011; Acevedo-Rodriguez and Strong, 2012). 

Risk of Introduction

Risk of introduction of C. baccatum is currently low to moderate, but the species possesses both desirable and undesirable traits that could cause potential invasiveness if not monitored, and further research is needed. The species is listed as a weed in the Global Compendium of Weeds (Randall, 2012) and is reported to be invasive to Cuba (Oviedo-Prieto et al., 2011) and a weed in Trinidad and Brazil (Holm et al., 1979; Randall, 2012). Invasive traits include tolerance of a wide range of precipitation and soil types, the species’ production of seeds viable for more than one year, wide distribution outside of its native range, and its ability to tolerate both acidic and alkaline soils (Basu and De, 2003; Ravishankar et al., 2003). Considering that the species has been cultivated in South America since pre-Columbian times (Basu and De, 2003) and is now present across tropical and subtropical regions around the world, the risk of introduction for this species may rise and should be monitored, especially in places where the species is cultivated.

Habitat

C. baccatum is primarily cultivated as a food and spice crop in agricultural and garden settings. In Zambia, it grows in disturbed ground and old cultivations as well as on anthills (Flora Zambesiaca, 2014). In Bermuda the species has been reported to grow in rocky woodlands and occasionally in gardens (Britton, 1918). In Peru, the domesticated form C. baccatum var. pendulum reportedly occurs in disturbed areas of the Andes and Amazonian regions, at altitudes 0-1500 m (Peru Checklist, 2014).

Habitat List

Biology and Ecology

Genetics

Sporophytic chromosome count for the species is 24 (IPCN Chromosome Reports, 2014).

Reproduction

The flowers are protogynous, but readily self pollinate. In the field, high rates of outcrossing (up to 90%) can occur with insect pollination. Capsicum exhibits no inbreeding depression. The stigma is positioned slightly below the level of the anthers or exserted slightly beyond, in which case the chances for cross-pollination are greater. Pepper breeders and seed producers use caution when producing a seed crop to prevent uncontrolled cross-pollination. The flowers are normally solitary in the axils of the branches, with the occasional cluster type that causes multiple flowers to form at a node. Many of the wild species have multiple flowers per node. There are two to four or more locules within the fruit. The locules are separated by the placentae where the capsaicinoids are produced. The outer wall, or pericarp, is fleshy and varies in thickness. It consists of a very thin cuticle, five to eight compact layers of small collenchyma cells that are cutinized during maturation and provide a tough, colourless, epidermal layer or skin. The growth of the fruit in the early stage consists of rapid cell multiplication; in the later stages growth is chiefly by enlargement of the cells already formed.

Environmental Requirements

Capsicum plants grow best at low-to mid elevations with 7-8°C, annual rainfall of 300-4600 mm and well-drained, sandy or silt-loam soil with pH of 4.3-8.7 (Basu and De, 2003; Ravishankar et al., 2003). FAO reports optimal annual rainfall levels for C. baccatum var. pendulum to be 600-1250 (absolute 500-1500) and temperature absolutes of 15-32°C, with the ability to grow in climatic zones ranging from tropical wet or wet and dry, to subtropical humid, dry summer, or dry winter (FAO EcoCrop, 2014). In Bolivia, C. baccatum grows in dry to montane forest and dry valleys at altitudes of 0-2000 m (Bolivia Checklist, 2014), while in Paraguay, the species occurs in low and high-altitude forests (Paraguay Checklist, 2014).

Climate

Soil Tolerances

Soil drainage

• free

Soil reaction

• acid
• alkaline
• neutral

Soil texture

• light
• medium

Means of Movement and Dispersal

C. baccatum spreads by seeds. Dispersal is primarily through intentional and accidental introduction, as the species continues to be cultivated around the world for human consumption as a food, spice, and medicinal ingredient (Basu and De, 2003); the domesticated species is reportedly the most widely domesticated pepper of Peru (Russo, 2012) and has been described as the most consumed species in Brazil (Spiller et al., 2008). In Oceania the species reportedly escaped cultivation but has been grown as both a spice and ornamental plant; in Vanuatu it is used first and foremost as an ornamental shrub and as a boundary marker, which may contribute to spread of the species through accidental introduction and cultivation escape (Walter et al., 2007).

Pathway Causes

Pathway Vectors

Impact Summary

Environmental Impact

C. baccatum is classified as a weed in the Global Compendium of Weeds (Randall, 2012), indicating its potential threat to the environment. Little data is currently available on the impacts of C. baccatum becoming weedy or invasive, and this should be researched in order to prevent and control the problem proactively. 

Risk and Impact Factors

• Proved invasive outside its native range
• Abundant in its native range
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Pioneering in disturbed areas
• Tolerant of shade
• Has propagules that can remain viable for more than one year

• Highly likely to be transported internationally deliberately
• Difficult to identify/detect in the field

Uses

C. baccatum has been valued by cultures around the world for its culinary use as a spice and vegetable, as well as in medicine and as an ornamental. In Oceania the species reportedly escaped cultivation and has been grown as both a spice and ornamental plant; in Vanuatu it is used first and foremost as an ornamental shrub and as a boundary marker (Walter et al., 2007). C. baccatum is also used as the ‘pepper’ chemical ingredient in self-defense pepper sprays (Basu and De, 2003). Pharmacological studies have also shown the species possesses anti-inflammatory properties and potential for pharmaceutical development (Spiller et al., 2008).

Uses List

Environmental

• Boundary, barrier or support

Human food and beverage

• Food additive
• Spices and culinary herbs

Materials

• Chemicals

Medicinal, pharmaceutical

• Source of medicine/pharmaceutical
• Traditional/folklore

Gaps in Knowledge/Research Needs

Less is known about this species compared with other members of the Capsicum genus, and further research is recommended in the areas of the potential environmental, social and economic impact if the species becomes invasive. As it is only reported to be invasive to Cuba and Trinidad, additional data on the species’ invasive or weed status would help to steer future actions for monitoring and, if necessary, control.

References

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Links to Websites

Contributors

04/12/2014 Original text by:

Marianne Jennifer Datiles, Department of Botany-Smithsonian NMNH, Washington DC, USA

Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

Distribution Maps