Monograph
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Monograph
Monograph of wild and cultivated chili peppers (Capsicum L., Solanaceae)
expand article infoGloria E. Barboza, Carolina Carrizo García, Luciano de Bem Bianchetti§, María V. Romero, Marisel Scaldaferro|
‡ Instituto Multidisciplinario de Biología Vegetal, Córdoba, Argentina
§ Centro Nacional de Pesquisa de Recursos Genéticos e Biotecnologia, Brasília, Brazil
| Universidad Nacional de Córdoba, Córdoba, Argentina
Open Access

Abstract

Capsicum L. (tribe Capsiceae, Solanaceae) is an American genus distributed ranging from the southern United States of America to central Argentina and Brazil. The genus includes chili peppers, bell peppers, ajíes, habaneros, jalapeños, ulupicas and pimientos, well known for their economic importance around the globe. Within the Solanaceae, the genus can be recognised by its shrubby habit, actinomorphic flowers, distinctive truncate calyx with or without appendages, anthers opening by longitudinal slits, nectaries at the base of the ovary and the variously coloured and usually pungent fruits. The highest diversity of this genus is located along the northern and central Andes. Although Capsicum has been extensively studied and great advances have been made in the understanding of its taxonomy and the relationships amongst species, there is no monographic treatment of the genus as a whole. Based on morphological and molecular evidence studied from field and herbarium specimens, we present here a comprehensive taxonomic treatment for the genus, including updated information about morphology, anatomy, karyology, phylogeny and distribution. We recognise 43 species and five varieties, including C. mirum Barboza, sp. nov. from São Paulo State, Brazil and a new combination C. muticum (Sendtn.) Barboza, comb. nov.; five of these taxa are cultivated worldwide (C. annuum L. var. annuum, C. baccatum L. var. pendulum (Willd.) Eshbaugh, C. baccatum L. var. umbilicatum (Vell.) Hunz. & Barboza, C. chinense Jacq. and C. frutescens L.). Nomenclatural revision of the 265 names attributed to chili peppers resulted in 89 new lectotypifications and five new neotypifications. Identification keys and detailed descriptions, maps and illustrations for all taxa are provided.

Keywords

America, Capsicum, chili peppers, cytogenetics, morphology, phylogeny, taxonomy

Introduction

Capsicum L., with 43 species, is placed in the tribe Capsiceae (subfam. Solanoideae, Solanaceae) together with Lycianthes (Dunal) Hassl. It is native to temperate, subtropical and tropical regions of the Americas, growing from the southern United States of America to central Argentina and Brazil, with the primary centre of diversity in the Andes.

Capsicum is an important crop genus, comprising the chili peppers, bell peppers, ajíes, habaneros, jalapeños, ulupicas or pimientos, with five main domesticated species: C. annuum L., C. chinense Jacq. and C. frutescens L., now widely cultivated throughout the world and C. baccatum L. and C. pubescens Ruiz & Pav., cultivated predominantly in South America. The genus comprises a diverse group of sweet and hot chili peppers, which have been used as spices since 6000–6500 BCE (Perry et al. 2007; Kraft et al. 2014). Today, they are of great commercial interest and are consumed by one fourth of the global population (Masud Parvez 2017).

Chili peppers are known for their high nutritional value, health benefits and medicinal properties (Saleh et al. 2018). The fruits are highly appreciated not only for their taste and colour (Thampi 2003; Jarret et al. 2019), but also because of their essential oils and the presence of capsaicinoids (Pruthi 2003; Meckelmann et al. 2013). Capsaicinoids are the pungent principles of hot chili peppers, of which capsaicin is the most abundant (Mazourek et al. 2009). These bioactive compounds have various pharmacological effects, such as the treatment of chronic neuropathic pains and many other ailments (Papoiu and Yosipovitch 2010; Aza-González et al. 2011; Al-Snafi 2015; Saleh et al. 2018). In addition, the ornamental use of chili peppers as potted or bedding plants, particularly in the case of the most widely-cultivated species C. annuum, is gaining popularity in the ornamental plant market in many countries (Stommel and Griesbach 2005; Costa et al. 2019).

The genus is characterised by a distinctive truncate calyx, without or with appendages (vein prolongations) borne below the entire margin and by variously coloured and usually pungent berries (Hunziker 2001). Traditionally, Capsicum was placed in tribe Solaneae Miers (Wettstein 1891; Hunziker 1979, 2001), based on morphological grounds (valvate aestivation, straight filaments, dorsifixed to basifixed anthers, juicy fruits and annular to coiled embryos). Emphasising calyx vasculature and structure (D’Arcy 1986), D’Arcy and Averett (1996) proposed a new tribe Capsiceae D’Arcy, with Capsicum and Lycianthes as the core group of this tribe, but they also suggested the inclusion of another eight genera (Acnistus Schott, Dunalia Kunth, Aureliana Sendtn., Iochroma Benth., Saracha Ruiz & Pav., Tubocapsicum (Wettst.) Makino, Vassobia Rusby and Witheringia L’Hér.), based on possession of calyx features at least superficially similar to Capsicum (D’Arcy 1991). Hunziker (2001) maintained Capsicum within tribe Solaneae, which he divided into five subtribes, placing Capsicum within subtribe Capsicinae T.Yamaz. and Lycianthes within subtribe Solaninae Wetts., a view that has not been accepted by later workers.

In the molecular phylogenetic reconstructions of Solanaceae (Olmstead et al. 2008; Särkinen et al. 2013), Capsicum and Lycianthes are resolved as sister taxa and are now the only members of tribe Capsiceae (subfam. Solanoideae). Although acceptance of Capsicum as a monophyletic group has not been questioned, its relationship with Lycianthes is still unresolved (Carrizo García et al. 2016). However, a recent genomic study analysing transcriptome sequences in members of both genera suggests possible paraphyly of Lycianthes with respect to Capsicum (Spalink et al. 2018). If upheld with further study, this result has important implications for the taxonomy of tribe Capsiceae; either circumscription of Capsicum should be enlarged to include Lycianthes or Lycianthes should be split into different genera that are monophyletic (Spalink et al. 2018).

Although Capsicum has been extensively studied and great advances have been made in the understanding of its taxonomy and the relationships amongst the species, there is no taxonomic monograph of the genus as a whole. As part of ongoing projects to revise the genera Capsicum and Lycianthes, we present here a comprehensive taxonomic treatment of Capsicum, including updated information about morphology, anatomy, karyology, phylogeny and distribution and a revision of the nomenclature and typification of the 265 names in the genus. An identification key and descriptions of wild and domesticated taxa, together with distribution maps and illustrations for each, are provided.

Circumscription and infrageneric classification of Capsicum

Since Linnaeus (1753), there has been no agreement about the circumscription of Capsicum. Capsicum species have been moved back and forth between several genera: Acnistus, Athenaea Sendtn., Brachistus Miers, Bassovia Aubl., Withania Pauq., Witheringia and others. Dunal was the first botanist to confuse generic concepts of Capsicum by describing some species under Witheringia (Dunal 1816), an error perpetuated by Kunth (1818) and Walpers (1844) or under Bassovia (Dunal 1852). Hiern (1877), Johnston (1905) and Rusby (1927) also described Capsicum species under Bassovia. Miers (1849) was mistaken in interpreting the deeply 5-lobed calyx of Brachistus or the slightly 3–5-lobed calyx of Witheringia (Bohs 2015) as homologous with the entire calyx with appendages of Capsicum (Hunziker 2001) and he named some species now recognised as Capsicum under Brachistus (newly-described species or transferred from Witheringia). Other authors also added species recognised today as Capsicum under Brachistus (Watson 1890; Greenman 1903; Stewart 1911; Rusby 1912; Svenson 1946), Acnistus (Dammer 1905), Fregirardia Dunal ex Delile (Delile 1849; Dunal 1852) or Solanum L. (Van Heurck and Müller Argoviensis 1870). Conversely, a number of present-day species, placed in a variety of Solanaceae genera (i.e. Vassobia, Witheringia, Athenaea) and even in other families (Pombalia Vand., Violaceae), were originally described as members of Capsicum (see Excluded names). The circumscription of the genus began to stabilise with the publication of complete treatments for morphologically similar genera, such as Lycianthes (Bitter 1919; Dean 1995), Witheringia (Hunziker 1969a; Sousa-Peña 2001), Brachistus (D’Arcy et al. 1981), Acnistus (Hunziker 1982), Cuatresia Hunz. (Hunziker 1987), Aureliana (Hunziker and Barboza 1990) and Tubocapsicum (Zhang et al. 1994; Zhang and Lu 1999). The information provided in these taxonomic works allowed Hunziker (2001) to propose a convincing circumscription of Capsicum, based on the diagnostic characters of an entire calyx with or without appendices, staminal plaque (= stapet) with auricles, anthers with longitudinal dehiscence, annular nectary at the base of the ovary, usually piquant (pungent) berries, a pericarp usually possessing giant cells and berries lacking stone cells.

There has been little agreement on an infrageneric classification of Capsicum. Kuntze (1891) proposed three sections: 1. Bassovia (Aubl.) Kuntze, 2. Eucapsicum Kuntze and 3. Poecilochroma (Miers) Kuntze, creating a heterogeneous group where Aureliana (= Athenaea), Witheringia, Saracha, Brachistus, Lycianthes and Capsicum species were mixed. This classification was based on flower size and the ratio of anther to filament length. At the same time, Wettstein (1891) divided Capsicum into two sections according to corolla shape: Eucapsicum Wettst. (30 American species) and the monospecific section Tubocapsicum Wettst. that included only C. anomalum Franch. & Sav., a species native to Japan (= Tubocapsicum anomalum (Franch. & Sav.) Makino).

Bitter (1919, 1921, 1922) placed species of Capsicum with 10 linear calyx appendages in his section Decameris Bitter (C. dusenii Bitter, C. brachypodum (Dunal) Kuntze, C. eggersii Bitter), but he never proposed an infrageneric classification across the whole of Capsicum.

The taxonomic division of Capsicum was re-examined in Hunziker’s Capsicum synopsis (Hunziker 1956), presented in the International Botanical Congress held at Paris in 1954. He recognised three sections, comprising sect. Tubocapsicum (today, a valid genus with two species, D’Arcy et al. 2001), the monotypic sect. Pseudocapsicum Hunz. for C. breviflorum (Sendtn.) Hunz. (presently Vassobia breviflora (Sendtn.) Hunz.) and sect. Capsicum including 24 wild species, one variety and three cultivated species. Eshbaugh (1980) accepted this sectional classification suggesting other members in sect. Capsicum (22 wild species and five domesticated, with their varieties), in the light of new discoveries.

The amount of new evidence produced in recent years has allowed considerable progress in the characterisation of infrageneric groups in Capsicum. Some attempts to group the species were made, based on cytogenetic studies (Moscone et al. 2007 and references therein), as well as a combination of data from enzyme, crossing and molecular studies (e.g. Walsh and Hoot 2001; Carrizo García et al. 2016).

Species relationships in Capsicum have been analysed following a phylogenetic approach using a range of molecular data. Several early phylogenetic hypotheses involved and primarily concerned domesticated species, although they also included a small number of wild ones (Walsh and Hoot 2001; Jarret and Dang 2004; Guzmán et al. 2009). A comprehensive phylogenetic analysis that included 34 of the 35 species recognised at that time was published in 2016 (Carrizo García et al. 2016). Based on that study and subsequent additions (Barboza et al. 2019, 2020b, eleven major clades are resolved within Capsicum and given informal clade names: Andean, Caatinga, Flexuosum, Bolivian, Longidentatum, Atlantic Forest, Purple Corolla, Pubescens, Tovarii, Baccatum and Annuum (Fig. 1). Two main lineages can be distinguished in the genus: one formed by the early diverging Andean clade that includes species from Central America and north-western South America and the other including the remaining species of the genus (Table 1). The domesticated species and their closest relatives are resolved in this second lineage and contained within the Annuum, Baccatum and Pubescens clades, which form the most derived branches (Fig. 1). Data on phylogenetic affinities and clades to which each species have been assigned are detailed in the species descriptions and in Table 1.

Table 1.

Capsicum clades and species composition (after Carrizo García et al. 2016, Barboza et al. 2019, 2020b; *placement unassigned, **placement suggested in this work, based on morphology).

Clade Species
Unassigned C. benoistii Hunz. ex Barboza *
Andean C. dimorphum (Miers) Kuntze
C. geminifolium (Dammer) Hunz.
C. hookerianum (Miers) Kuntze
C. lanceolatum (Greenm.) C.V.Morton & Standl.
C. longifolium Barboza & S.Leiva
C. lycianthoides Bitter
C. piuranum Barboza & S.Leiva
C. regale Barboza & Bohs
C. rhomboideum (Dunal) Kuntze
Atlantic Forest C. campylopodium Sendtn.
C. carassense Barboza & Bianch.
C. cornutum (Hiern) Hunz.
C. friburgense Bianch. & Barboza
C. hunzikerianum Barboza & Bianch. **
C. mirabile Mart.
C. mirum Barboza
C. muticum (Sendtn.) Barboza
C. pereirae Barboza & Bianch.
C. recurvatum Witasek
C. schottianum Sendtn.
C. villosum Sendtn.
Flexuosum C. flexuosum Sendtn.
Caatinga C. caatingae Barboza & Agra
C. parvifolium Sendtn.
Longidentatum C. longidentatum Agra & Barboza
Bolivian C. caballeroi M.Nee
C. ceratocalyx M.Nee
C. coccineum (Rusby) Hunz.
C. minutiflorum (Rusby) Hunz.
C. neei Barboza & X.Reyes
Purple Corolla C. cardenasii Heiser & P.G.Sm.
C. eshbaughii Barboza
C. eximium Hunz.
Pubescens C. pubescens Ruiz & Pav.
Tovarii C. tovarii Eshbaugh, P.G.Sm. & Nickrent
Baccatum C. baccatum L.
C. chacoense Hunz.
C. rabenii Sendtn.
Annuum C. annuum L.
C. chinense Jacq.
C. frutescens L.
C. galapagoense Hunz.
Figure 1. 

Capsicum phylogeny. Cladogram summarising findings from Carrizo García et al. (2016) and Barboza et al. (2019, 2020b). Only one sample per species has been included; species names have been updated. Clades distinguished are labelled. Asterisks indicate branches with moderate/low support, letters ‘d’ clades with domesticated taxa.

Taxonomic history of Capsicum

The quest of Europeans for the “Indies” (namely the Americas) was accompanied by the discovery of new aromatic plants that extensively enriched cuisines around the world; amongst these were the chili peppers (reviewed in Andrews 1984). The first written documentation of these spices is that of Diego Alvárez Chanca, physician on Columbus’ second voyage to the West Indies. In a letter he wrote in 1494 to the Town Council of Seville, Spain, describing the principal events that occurred during the voyage, he referred to the ‘agi’ (word in Spanish) as a spice that the native Indians of Hispaniola employed to season their foods (American Journeys Collection 2003).

Fifty years after Columbus’ first voyage to the West Indies, Leonhard Fuchs, a German physician and botanist, published the first three scientific illustrations of chili peppers (Fuchs 1542). His illustrations (Calechutifcher Pfeffer, Langer Indianifcher Pfeffer and Greyter Indianifcher Pfeffer) were all given Pliny’s name of ‘Siliquastrum’ (large, elongate fruit), because he did not know the geographic origin of the plants (Bostock and Riley 1856) and he associated them with the spices native to India that were known at that time (Piperitis and Siliquastrum).

The word ‘capsicum’ was coined in the pre-Linnaean literature for the first time by Matthias de L’Obel (1576: 173) for the “Piper indicum longioribus siliquis”. L’Obel also provided two illustrations of fertile plants (C. annuum in its wild and domesticated forms). Pre-Linnaean botanists (e.g. Dodoens 1554, 1583; Clusius 1611; Bauhin 1623; Parkinson 1640; Morison 1669) proposed many polynomials for the peppers emphasising the variable fruit characters (colour, shape, size, position or pungency). Joseph Pitton de Tournefort (1719), in his influential work Institutiones, used the name Capsicum, gave an original description for the genus and listed 27 polynomial species that corresponded to his concept of the genus. He also mentioned the etymology of the word Capsicum, either from the Greek word δάγκωμα (= to bite), on account of the burning strength of the seeds or from the Latin voice capsa (= box), on account of the boxy shape of the cultivated fruits.

Linnaeus (1753) took Tournefort’s Capsicum as the generic name of peppers and reduced the number of species to two, the herbaceous annual C. annuum and the shrubby perennial C. frutescens. In subsequent works, Linnaeus (1767) added two more species, C. baccatum and C. grossum L., Jacquin (1776) described C. chinense, Ruiz and Pavón (1799) C. pubescens and Willdenow (1809) proposed 10 species, amongst them C. pendulum Willd. By the beginning of the 19th century, the five currently accepted domesticated Capsicum species had already been described.

Subsequently the taxonomy of the genus has been complicated by both generic circumscription (see above) and by differing species concepts. Opinions as to the number of taxa that belong to Capsicum range from as many as 61 species (plus infraspecific taxa) in the genus (e.g. Dunal 1852) to as few as two (Irish 1898) or one (Dierbach 1829; Bailey 1923; Shinners 1956). Understandably, there has been underlying disagreement amongst taxonomists about whether to accept emerging names as well as the different intricate infraspecific classifications proposed for the domesticated species (e.g. see Dierbach 1829; Terpó 1966). Further complications included: 1) the majority of the new taxa were described, based on the very wide range of variation observed in the fruit morphology of cultivated plants, which is highly influenced by human selection; 2) most herbarium specimens of Capsicum were inadequate, many consisting only of fruits and lacking flowers, the most critical and useful organ for identification purposes; and 3) type specimens for many names were difficult to locate or did not exist.

Taxonomy of the domesticated species

After Linnaeus, the British gardener Philip Miller (1768) recognised 10 domesticated species, five of them new species described by him. The descriptions of all these species were almost certainly made from living plants cultivated in the Chelsea Physic Garden in London.

The German botanist, Johann Heinrich Dierbach (1829), proposed a particularly confusing infraspecific classification. He accepted a single species and validated the pre-Linnaean name “Capsicum indicum” of L’Obel (1576), as C. indicum Dierb. He recognised infraspecific unranked taxa designated by numbers (1. macrocarpon, 2. pachycarb[p]on, 3. cerasocarpon, 4. elaeocarpon, 5. microcarpon), based on fruit shape; none of these numbered names had a direct reference to the author or a specimen. Within each numbered category, the different taxa were designated with a letter (a to e; e.g. C. indicum Lobel. 1. macrocarpon b. longum) and, in addition, before the designation of letters, the taxa were grouped according to the pericarp colour (e.g. pericarpiis rubris, pericarpiis flavis). Heiser and Pickersgill (1975) attempted to interpret Dierbach’s classification system, concluding that it “… should be ignored for nomenclatural purposes”. However, Dierbach based the lettered infraspecific taxa on the names of the chili peppers known at that time, with the result that many of them are now synonyms (especially under C. annuum) or as doubtful names (see Doubtful names).

The first illustrated monograph of Capsicum was published by the German botanist, Karl Anton Fingerhuth (1832), in which he included 32 species. He recognised the taxa already described by previous authors and described only three new species (C. cumanense Fingerh., C. strictum Fingerh. and C. ceratocarpum Fingerh.). He also recognised 29 varieties, most of which were newly described. Although he did not propose a formal infrageneric classification, he grouped the species into two groups (A and B) depending on the position of the fruits (A. Fruits erect; B. Fruits pendent). Within each group, he further separated the species into two subgroups (a. Fruits oblong; b. Fruits subglobose). The distinctions of his many varieties were based on the broad range of different fruit shapes. None of the taxa proposed by Fingerhuth is an accepted name in this treatment (see synonymy under the domesticated species).

In his monumental treatment of Solanaceae, Michel-Felix Dunal (1852) recognised 61 Capsicum species (16 newly described by him), together with 51 varieties (16 new), 11 species requiring further study and three doubtful names. From these, 44 species and 40 varieties referred to the domesticated species. Dunal did not establish an infrageneric classification, but grouped the species in a similar way as had Fingerhuth (1832), first by the position of the fruits (erect or pendent) and then by shape (oblong or globose-ovate).

In extreme contrast, other authors tended to reduce the number of accepted species to two (Irish 1898) or only one (Bailey 1923; Erwin 1929; Shinners 1956; and others), with very complex infraspecific hierarchical systems, based on the extreme variation in fruit characters of the domesticated taxa and their cultivars. Terpó (1966) summarised some of these elaborate systems, while Eshbaugh (1980) expressed the difficulties inherent in developing a useful and rational system that satisfied everyone. Capsicum annuum and C. frutescens and, to a lesser extent for C. chinense, have been the taxa most debated by taxonomists and horticulturists in classifications below the species level. From an evolutionary perspective, the relationships between these three species have been the most debated (Pickersgill 1988), whereas there was a general agreement that C. pubescens and C. baccatum var. pendulum were two well-defined domesticated taxa (Eshbaugh 1980; Pickersgill 1988).

As a crop genus, Capsicum has inspired researchers to follow a number of in-depth approaches since the mid-1900s, such as classical and molecular cytogenetic analyses, crossing experiments, biochemical and protein electrophoretic studies, molecular characterisation through genotypic markers (restriction fragment length polymorphism, RFLP, amplified fragment length polymorphism, AFLP, random amplified polymorphic DNA, RAPD, microsatellite or simple sequence repeat, SSR, random amplified microsatellite polymorphism, RAMPO and direct amplification of minisatellite DNA, DAMDPCR), phytogeographic and phylogeographic analyses, chloroplast and nuclear DNA and whole-genome sequencing studies (see Moscone et al. 2007; Aguilar-Meléndez et al. 2009; Carvalho et al. 2014; Carrizo García et al. 2016; Raveendar et al. 2017; Scaldaferro et al. 2018; Tripodi et al. 2019; and references in each). This evidence has contributed to a better understanding of the complex taxonomy of the domesticated (and wild) species, as well as to determine the origin of the crop species. Most current taxonomic works and breeding programmes recognise five domesticated species of Capsicum (Pickersgill 2016).

Taxonomy of the wild species

Taxonomic work on wild Capsicum species began in the 19th century. Initially, some authors (Cavanilles 1802; Willdenow 1809; Dunal 1816; Kunth 1818; Roemer and Schultes 1819; and others) described isolated species. The Capsicum treatment by Sendtner (1846) in “Flora Brasiliensis” is the first significant work from an extended geographical area; he accepted seven wild species and five infraspecific taxa in addition to several cultivated species. Later, Dunal (1852) made an important contribution with the addition of seven other novel wild species and three varieties.

During the early 20th century, sporadic descriptions of new Capsicum taxa continued (Chodat 1902; Chodat and Hassler 1903; Witasek 1910; Bitter 1919, 1921, 1922, 1924). Later, a new generation of taxonomists interested in this genus, led by Paul Smith and Charles Heiser, Jr. (in the United States) and Armando T. Hunziker (in Argentina), made great advances in the understanding of the generic limits of Capsicum (see above). These authors not only provided new names, but also new evidence to establish the relationships between species. Hunziker, especially, contributed greatly to a comprehensive understanding of the taxonomy of the genus (Hunziker 1950, 1956, 1961, 1969b, 1971, 1998, 2001). In his book “Genera Solanacearum”, Hunziker (2001) proposed Capsicum as a natural group with “ca. 20 species and a few varieties”.

In the last two decades, field explorations across South America, mainly in the central Andean countries and Brazil, have enabled us to gain a better understanding of the genus as a whole. Thirteen new wild species have been described and partial keys for the identification of the species for particular areas have been provided (Barboza and Bianchetti 2005; Nee et al. 2006; Barboza et al. 2011, 2019; Barboza et al. 2020a, Barboza et al. 2020b). We recognise here 43 species of Capsicum (including the domesticates), with new species still awaiting discovery in herbaria and in the field (especially in Peru and Bolivia).

Morphology

Habit and stems

Members of Capsicum plants are erect (e.g. C. schottianum, C. geminifolium), compact (e.g. C. chacoense, C. annuum var. annuum) or somewhat prostrate (e.g. C. annuum var. glabriusculum). They are subshrubs or shrubs, rarely trees (e.g. C. rhomboideum), short-lived perennials (e.g. C. chinense) or annual herbs (e.g. C. annuum var. annuum). Capsicum coccineum is unusual in being sprawling vines or scrambling shrubs. Stems are woody at the base (1.5–2.5 cm in diameter, rarely more) and some species have fissured bark and lenticels (e.g. C. rhomboideum, C. hookerianum); young stems are angular, herbaceous, usually hollow and weak and, occasionally, somewhat scrambling, range from glabrous to densely pubescent and may have anthocyanin along their length. The nodes are inflated and commonly green or purple.

Capsicum plants have typical solanaceous sympodial growth, giving the stems a “zig-zag” appearance. Initially, the vegetative growth is monopodial and the first stem to emerge has 8–39 leaves (C. annuum cultivars, Dorland and Went 1947; Patel and Shah 1977; Bosland and Votava 2000; C. chacoense, Barboza, pers. obs.) before the onset of sympodial ramification and flowering (Fig. 2A); the alternate leaves are arranged in a 2/5 phyllotaxic spiral. The main stem ends in a solitary flower or in a flower and one or two leaves (Fig. 2B, C). The number of leaves in a sympodial unit can be unifoliate or difoliate. In difoliate sympodial units, the leaves are geminate with the leaf pair slightly dissimilar in shape or size (e.g. C. cardenasii) or markedly dissimilar (e.g. C. dimorphum, C. lycianthoides) in size and/or shape.

Figure 2. 

Plant development in C. chacoense A monopodial vegetative growth B first dichotomy of the main stem and start of sympodial growth C initial branching with three branches. Photos by G.E. Barboza.

Leaves

Species of Capsicum have simple leaves that are generally membranous or less frequently coriaceous (e.g. C. hunzikerianum, C. longifolium, C. pereirae), concolourous to discolourous, ovate to elliptical, rarely lanceolate or narrowly elliptical (C. longifolium, C. carassense) in outline; in taxa with geminate leaves, the minor leaves can be orbicular and sessile (C. dimorphum, C. lycianthoides). Leaf margins are always entire, rarely slightly revolute (C. caballeroi, C. ceratocalyx, C. hunzikerianum) and the leaf base is asymmetric, attenuate or truncate and sometimes decurrent on to the petiole (C. piuranum, C. rabenii). Leaf apices are obtuse or acute to acuminate or long-acuminate in few species (e.g. C. benoistii, C. piuranum, C. hunzikerianum). Petioles are longer in the domesticated species and in the major leaves of geminate leaf pairs in wild species.

Leaf and petiole anatomy of Capsicum members were investigated in domesticated species (Idu and Ogbe 1997; Weryszko-Chmielewsk and Michałojć 2011; Dias et al. 2013; Wahua et al. 2014) and, more extensively, in 13 wild species from different biogeographic environments (Palchetti et al. 2014). Most of the studied species have amphistomatic leaves, but leaves are hypostomatic in some C. frutescens and C. annuum cultivars, C. caatingae, C. cornutum and C. geminifolium (Idu and Ogbe 1997; Palchetti et al. 2014). Anisocytic, anomocytic and hemiparacytic stomatal types were reported; this variation was observed within a same leaf, a common feature in Solanaceae (Metcalfe and Chalk 1957; Bessis and Guyot 1979; Karatela and Gill 1986). Palchetti et al. (2014) described leaf blades with dorsiventral mesophyll, sparse to abundant calcium oxalate crystals (druses, solitary crystals and/or crystal sands), bicollateral vascular bundles surrounded by parenchyma, fibres or collenchyma and petioles with arch- or U-shaped vascular bundles. Leaf epidermal variables and the internal structural differences of leaf and petiole were not significantly different in the 13 wild species from different environments (Palchetti et al. 2014), but some anatomical characters were useful for the identification (stomata position and types, type of trichomes, type of cells surrounding the vascular bundle in the leaf, presence/absence and abundance and types of crystals and others).

Pubescence

Trichomes in Capsicum are mostly eglandular and simple, although branched trichomes can also occur (e.g. C. longidentatum, C. rhomboideum and C. parvifolium). Simple trichomes are uniseriate and usually 1–11-celled (Fig. 3A–D, F, K, O), with slightly (Fig. 3F–H) to strongly (Fig. 3A–D, O) minutely warty cuticle (Dias et al. 2013; Palchetti et al. 2014). Trichomes can be straight (Fig. 3A, B, O) or curved (Fig. 3C, D), cylindrical (Fig. 3B, O) or conical (Fig. 3A, M), flexible or somewhat rigid, antrorse or spreading. The distal cell of the trichomes can be acute (Fig. 3A, D, F), obtuse (Fig. 3H) or strongly curved resulting in a hook-shaped trichome (Fig. 3K). Branched trichomes can be furcate (forked) (Fig. 3G, P) to many times branched (Fig. 3J, L, N, see also Barboza et al. 2011). Papillae (Fig. 3E, I) are common at the margin and the apex of the corolla lobes in most species.

Figure 3. 

Eglandular trichomes of Capsicum species A–D, F, H, K, M, O simple trichomes E, I papillae G, J, L, N, P branched trichomes. Scale bar: 50 μm (A, B, E, H–J); 100 μm (C, D, F, G, K–P).

Glandular trichomes are common in Capsicum species. In most species, glandular trichomes are simple, with short uni- or bicellular stalks and globose to ellipsoid multicellular heads (Fig. 4D, E, G, I). These are usually distributed on stems, leaves, pedicels and the outer calyx surface of some species (e.g. C. caatingae, C. cardenasii, C. galapagoense, C. tovarii) and on the inner calyx surface in all Capsicum species. Glandular trichomes with (1–) 2–3-celled stalks and globose or globose-peltate unicellular heads (Fig. 4B, C, F) are found on the interior surface of the corolla in many species. The most unusual glandular trichomes are found in C. eshbaughii (Fig. 4A, H, J) which has a distinctive branched glandular indument (Barboza 2011).

Figure 4. 

Glandular trichomes of Capsicum species. Scale bar: 50 μm (A–G, I); 100 μm (H, J).

Density of pubescence is highly variable both within and between species. Sometimes, this variability has been considered diagnostic at the infraspecific level. For example, the densely pubescent plants of C. chacoense, C. baccatum, C. eshbaughii or C. annuum have been recognised at the infraspecific level (see descriptions and synonymy of those species).

Inflorescences

Capsicum species have axillary flowers; they are solitary only in C. chacoense (Fig. 2C) or may be in 1–2 (–3)-flowered inflorescences in some species (e.g. C. baccatum, C. friburgense, C. piuranum) or in 4–13-flowered inflorescences in most of the species. Capsicum ceratocalyx and C. caatingae may have more than 18 flowers per inflorescence. The inflorescences are either epedunculate and unbranched or borne on a short (e.g. C. ceratocalyx, C. coccineum) or somewhat elongate rachis that is occasionally forked (e.g. C. regale). Flowers are usually deciduous, falling off the plant gradually and leaving conspicuous or obscure pedicel scars on the rachis.

All Capsicum flowers have distinct, usually pubescent, pedicels that may be terete (Fig. 5B), angled (Fig. 5A) or, rarely, almost winged (C. ceratocalyx). Pedicel/flower position is a useful identification character. Pedicels (and flowers) can be deflexed (termed pendent in this treatment; for example, C. pereirae, C. lanceolatum, Fig. 5C) or erect to spreading. In some cases, the pedicels are distally abruptly twisted forming a right or acute angle with the longitudinal axis of the flower (termed geniculate in this treatment). Flowers with geniculate pedicels may be orientated with their longitudinal axis in horizontal position (e.g. C. rabenii, Fig. 5A) or nearly parallel to the pedicel axis (e.g. C. annuum var. glabriusculum, Fig. 5B). In fruits, pedicels are usually green (Fig. 6A, B, G) or green with purple stripes (e.g. C. eximium, C. longifolium) or completely purple (C. regale, Fig. 6F). They are normally widened distally and can be pendent (e.g. C. recurvatum, C. flexuosum) or erect (e.g., C. baccatum var. baccatum, C. chacoense, C. frutescens).

Figure 5. 

Flower morphology in Capsicum species A C. rabenii B C. annuum var. glabriusculum C C. lanceolatum D C. schottianum E C. frutescens F C. eximium G C. eshbaughii H C. cornutum I C. galapagoense J C. recurvatum K C. cardenasii L C. lycianthoides M C. chacoense N C. baccatum var. baccatum O C. caballeroi. Abbreviations. im interpetalar membrane sp staminal plaque.

Figure 6. 

A–G Fruit morphology in Capsicum species H, J epicarp structure A C. chinense B C. baccatum var. baccatum C C. hookerianum D C. lanceolatum E C. coccineum F C. regale G C. schottianum H epicarp with regular epidermal cells I epicarp with some sclereids amongst the regular epidermal cells J epicarp exclusively with sclereids. Abbreviation. sc sclereids. Scale bar: 10 μm (H, I, J).

Calyces

The calyx in Capsicum species is usually 5-merous (4–8-merous in domesticated taxa) and entirely synsepalous (Fig. 5E). The calyx tube is generally cup-shaped or campanulate and 5–10 nerved, with the margin always entire (Fig. 5E, L). Below the margin, up to10 calyx appendages may emerge (Fig. 5A, F–H, L), a character only shared with Lycianthes (Dunal 1852; Bitter 1919; D’Arcy 1986; Hunziker 2001). In calyces without appendages, the calyx outline can be circular with the five main veins completely immersed in the calyx tube and ending very near its margin (e.g. C. campylopodium, C. galapagoense) (Fig. 5E). In species such as C. schottianum, the calyx outline may be pentagonal (Fig. 5D) with the main veins ending in five tiny lateral umbos or lumps that barely emerge over the calyx margin. Calyx appendages are lateral elongate expansions of the calyx tube (Fig. 5C, F–H, L) with vasculature formed by each main or secondary vein arching outwards into the appendages and returning to the calyx tube to end in the margin of the calyx (see figs 1–4 in D’Arcy 1986). The calyx sleeve, which is a ring of tissue immediately below the margin, is rudimentary (e.g. C. recurvatum, C. hookerianum, C. coccineum, C. lycianthoides, Fig. 5L) or non-existent in Capsicum (Fig. 5F, H). This sleeve is well developed in many Lycianthes species (D’Arcy 1986; Dean et al. 2017, 2020) and, thus, serves as a trait to differentiate the two genera. The number, shape, size and orientation of the appendages are important identification characters within Capsicum. The number of appendages is consistent (5 or 10) in most species, but in others, it can vary (2–10) within a species and, sometimes, even within an individual (e.g. C. recurvatum). Appendages shape ranges from subulate (wider at the base and pointed at the tip, for example, C. lycianthoides, Fig. 5L), to cylindrical (e.g. C. eximium, Fig. 5C, F) and linear (e.g. C. eshbaughii, C. cornutum, Fig. 5G, H). In some species, the appendages are thick and strongly laterally flattened like triangular-compressed wings (e.g. C. longifolium, C. regale). Appendages are usually equal or subequal (e.g. C. eximium, C. piuranum), but when there are more than five in number, the main appendages (those with vasculature formed by the main veins) are longer than the secondary appendages (those with vasculature formed by secondary veins) (e.g. C. hookerianum, C. longidentatum). The longest main appendages (ca. 8.5 mm) are found in C. cornutum (Fig. 5H). Calyx appendages can be erect and appressed to the corolla (e.g. C. eximium, Fig. 5F), spreading (e.g. C. dimorphum, C. lycianthoides, Fig. 5L), recurved (e.g. C. recurvatum), reflexed (e.g. C. lanceolatum, Fig. 5C) or incurved and horn-like (C. ceratocalyx); orientation may change in the transition between flowering and fruiting (e.g. C. coccineum). An annular constriction at the junction between the fruiting calyx and pedicel is present in a few wild species (e.g. C. caatingae, C. regale) and it is a key character used in the identification of C. chinense (Fig. 6A). Fruiting calyces are persistent and not accrescent or only slightly accrescent (domesticated species). They are discoid in most species (Fig. 6G), but can be cup-shaped (e.g. C. frutescens) and are strongly reflexed in C. coccineum (Fig. 6E).

Corollas

Capsicum species have 5-merous (6–8-merous in domesticated taxa) sympetalous corollas. Corollas are usually of intermediate size (6–14 mm long), the smallest measuring 4–5 mm long (e.g. C. galapagoense) and the largest ones reaching 17–18 mm long (e.g. C. caballeroi, C. piuranum). Most species have stellate corollas (Fig. 5D–G, I, J), but campanulate (e.g. C. lanceolatum, Fig. 5C, C. cardenasii, Fig. 5K), broadly campanulate (C. lycianthoides, Fig. 5L, C. rhomboideum), campanulate-urceolate (C. friburgense), rotate-stellate (C. flexuosum, C. baccatum var. baccatum, Fig. 5N) or rotate (e.g. C. pubescens) corollas are also present. The corolla tube can be very short (C. benoistii), short (e.g. C. eximium, Fig. 5F) or long (e.g. C. caballeroi, C. cardenasii, Fig. 5K, C. piuranum). The lobes are connected by a thinner interpetalar membrane nearly completely (e.g. C. lycianthoides, Fig. 5K, L), partly (e.g. C. baccatum var. baccatum Fig. 5J) or just near the base (C. annuum var. glabriusculum, C. galapagoense, Fig. 5I); in very few species, interpetalar membrane is absent (e.g. C. caballeroi).

Corolla colour is highly variable in Capsicum species. Corollas can be entirely white (e.g. C. chacoense, C. galapagoense, C. annuum var. annuum), dull white or greenish-white (C. chinense, C. frutescens), light yellow (C. neei), yellow (e.g. C. piuranum, C. caballeroi) or violet or fuchsia (C. friburgense). Other species have corollas with a predominant primary colour (white, yellow or purple), as well as markings (spots) with diverse pigmentation. The abaxial surface (outer surface) of the corollas may have the same colouration as the adaxial surface (inner surface) (e.g. C. geminifolium, C. regale) or it may have a faded or a different colouration (e.g. C. tovarii, C. pereirae). In many species, the co-occurrence of different pigments results in multi-coloured corollas, for example, white corollas with purple (or variations) spots at the base of the lobes and limb and green or greenish-yellow centre (e.g. C. villosum, C. schottianum, C. pereirae). Descriptions in literature or on specimen labels usually refer to the colour of the inner (adaxial) corolla surface. In this monograph, description of the corolla colour of both surfaces is provided for each species; corolla colour can be very difficult to see on herbarium specimens.

Corolla pigmentation is due to anthocyanins which produce violet or purple shades (e.g. C. lycianthoides, C. lanceolatum, Fig. 5C, F, K, L) and chlorophyll which produces green colouring (C. baccatum, C. flexuosum, Fig. 5D, J).

The adaxial surfaces of the corollas are glabrous (many Andean species) or may be covered by sparse glandular trichomes (e.g. C. ceratocalyx, C. tovarii) or have a continuous ring of glandular trichomes (Fig. 4C, F) at the base of the lobes and in the throat (the Bolivian and Brazilian species). The abaxial surfaces of corolla lobes are mostly papillate, with minutely warty papillae (Fig. 3E, I) or with short simple trichomes on the margins and tips. The lobes are spreading in most species (Fig. 5D, F, G, I, J), erect in C. tovarii or slightly (e.g. C. cardenasii, Fig. 5K) to strongly recurved in some species (C. caballeroi and C. friburgense).

Androecium

Capsicum species have usually five (sometimes 6–8 in domesticated taxa) equal stamens; unequal stamens have only been observed in three species: C. campylopodium (and sometimes also C. lycianthoides), which has two stamens longer than the other three (Hunziker 2001) and C. regale, which has one longer than the other four (Barboza et al. 2020b). The free portion of the filaments is always distinct and glabrous and it is usually longer than the anthers (Fig. 5N). Each filament broadens at its base forming a staminal plaque (called a stapet, Hunziker 2001), with two short lateral auricles fused to the corolla base. In C. chacoense, these lateral auricles are long and are not fused to the corolla. The anthers are longitudinally dehiscent and are usually ellipsoid and yellow or cream in colour (Fig. 5M); in domesticated taxa and some wild species (e.g. C. dimorphum, C. carassense, C. regale), the anthers are blue (Fig. 5B), bluish-grey or purple. In pre-anthesis and early anthesis, the anthers are connivent (Fig. 5M), but they become separated during anthesis (Fig. 5N), remaining somewhat connivent in some species (Fig. 5B). Anther size is variable, ranging from 0.9–1.3 mm long in C. galapagoense, to 1–3 mm long in most species, to up to 4 mm in C. hunzikerianum.

Pollen is yellow or white, trizonocolporate, spheroidal, prolate, prolate-spheroidal or oblate-spheroidal, with a triangular to circular outline in polar view. It is usually small, from 15 µm in C. rhomboideum (Bo and Carrizo García 2015) to 18–24.73 µm in some domesticated species (Dharamadhaj and Prakash 1978; Martins et al. 2013; Adedeji and Akinniyi 2015; Bo and Carrizo García 2015; Kayani et al. 2018; Yang et al. 2018; Song et al. 2019), but in other domesticated and wild species, sizes range from 25.75 to 38 µm (Murray and Eshbaugh 1971; Quagliotti 1979; Martins et al. 2013). Pollen grains are shed in monads and at the binucleate stage (Cochran 1938; Yaqub and Smith 1971; Dharamadhaj and Prakash 1978; Kim et al. 2004; Bo and Carrizo García 2015) or trinucleate stage (e.g. a C. frutescens cultivar, Lengel 1960). Information on the ornamentation of the exine is variable in literature depending on whether observations were made with a light microscope (LM) or a scanning electron microscope (SEM). The exine appears to be clearly microechinate and perforate under SEM in C. annuum (Halbritter 2016a; Song et al. 2019) and C. pubescens (Bo and Carrizo García 2015; Halbritter 2016b). The exine has been described as reticulate (Murry and Eshbaugh 1971; Roubik and Moreno 1991), foveolate (Martins et al. 2013), psilate or faintly granulate (Kayani et al. 2018) and scabrate (Barth and Duarte 2008), based on LM observations for some Capsicum species, which should be corroborated with SEM. Pollen morphology is lacking for most wild Capsicum species and more information is needed to evaluate the usefulness of pollen to distinguish species.

Gynoecium

The gynoecium in Capsicum species is usually bicarpellate (2–5-carpellate in domesticated species). The ovary is superior with axile placentation, glabrous and usually subglobose to ovoid, rarely ellipsoid (e.g. C. frutescens). The style is simple, straight or slightly curved, cylindrical (the same width from the proximal to distal end, Fig. 5N) or clavate (broadened gradually from its base to the apex), glabrous, white, cream, lilac or purple and commonly exserted beyond the anthers. Most species have homostylous flowers, but some species (C. annuum, C. baccatum, C. benoistii, C. tovarii, C. pubescens) have heterostylous flowers, with different flowers on the same plant bearing short, medium or long styles. In domesticated species, style length variations have been observed amongst cultivars (Bosland and Votava 2000; Peña-Yam et al. 2019). The stigma is pale green or cream, globose or discoid, sometimes slightly bilobed (e.g. C. longifolium, C. lanceolatum) and finely papillate. The ovules are usually numerous, anatropous, tenuinucellate, with a single integument (Cochran 1938; Munting 1974; Dharamadhaj and Prakash 1978; Pérez-Pastrana et al. 2018). The nectary is located at the base of the ovary and is easily observed in fresh material, but sometimes can be very difficult to see in herbarium specimens; it is an inconspicuous annular disc, paler in colour than the rest of the ovary, variable in thickness and produces copious nectar. The nectar is colourless and is exposed in five nectar droplets on the corolla limb (see more detail in Floral biology and pollination).

Fruits

The fruit is usually a bicarpellate berry (Fig. 7A) that can also be 3–5-carpellate in domesticated species (e.g. C. annuum var. annuum, C. chinense, C. pubescens) or some wild species (e.g. C. tovarii). The berries are extremely diverse in shape, size and colour in domesticated species and more homogeneous in wild (and semi-domesticated) species. The fruits are juicy, fleshy, opaque (most species) or translucent (some Brazilian species). Wild species have globose or subglobose berries < 15 mm in diameter (Fig. 6A, C, E–G) or short ellipsoid or ovoid (e.g. C. baccatum var. baccatum, C. chacoense) berries 0.6–14 mm long (Fig. 6B). The fruits in domesticated species are much longer and wider, with the most elongate fruits (300 mm long or more) in C. annuum var. annuum (Bosland and Votava 2000). The great variation of the fruit shapes has resulted in the proposal of more than 100 names applied to the domesticated species (here assigned to any of the five recognised domesticated taxa). Within each of the domesticated taxon, a system of different fruit types (i.e. characterised by a defined fruit shape and colour, pungency level, aroma and/or flavour and uses) is used by horticulturists or plant breeders (Bosland et al. 1988); nearly 100 fruit types are known, the most frequent of which are described in Bosland and Votava (2000) and DeWitt and Bosland (1997, 2009). Red is the most frequent fruit colour in wild Capsicum species (Fig. 6B–E), with colours ranging from orange-red (e.g. C. geminifolium), dark burgundy (C. rhomboideum) to dark blue or purple-blue (C. regale, Fig. 6F). Some Brazilian species have greenish-golden yellow, translucent fruits (e.g. C. parvifolium, C. schottianum, Fig. 6G), a character state considered to be derived within Capsicum (Carrizo García et al. 2016).

Figure 7. 

Fruit anatomy in Capsicum species A, D–F C. baccatum var. pendulum B C. baccatum var. umbilicatum C, G, H C. pubescens A fruit, in cross section (note giant cells in the pericarp) B one locule of a fruit, in cross section (note the absence of giant cells in the pericarp) C, D epicarp and some layers of mesocarp (in D, observe cuticular wedges) E sector of pericarp (the arrow indicates the increase of the cell size ending in the giant cells) F detail of two adjacent giant cells G sector of homogeneous endocarp H sclereids of the endocarp. Abbreviations. c cuticle, cw cuticular wedge, epc epidermal cells, gc giant cells, p pericarp. Scale bars: 1 mm (A, B, E); 10 μm (C, D, H); 100 μm (F, G).

Fruiting pedicels are usually green (Fig. 6A, B, G) or green with purple stripes (e.g. C. eximium, C. longifolium) or completely purple (C. regale, Fig. 6F). They are normally widened distally and can be pendent (e.g. C. recurvatum, C. flexuosum) or erect (e.g., C. baccatum var. baccatum, C. chacoense, C. frutescens). Pedicels and ripe fruits are persistent in domesticated species, staying attached to the plant; in most wild species, the mature berries are usually deciduous and are easily detached from the calyx, leaving only pedicels and fruiting calyces on the plant (Carvalho et al. 2014, GEB, pers. obs.). In just a few species (e.g. C. muticum, C. coccineum), mature berries fall from the plant with pedicels attached, leaving conspicuous scars.

Pericarp structure

The development of the pericarp in Capsicum species is the typical of a true berry (Roth 1977), with the epicarp and endocarp originating respectively from the outer and inner epidermis of the ovary wall and the mesocarp derived from the middle layers between the two epidermal layers (Roth 1977; Filippa and Bernardello 1992); this structure is typical of most Solanaceae (Roth 1977; Bernardello 1983; Filippa and Bernardello 1992). The taxonomic significance of the pericarp structure in Capsicum has been rarely considered with some observations made on varietals or cultivars of the domesticated species (Tschirch and Oesterle 1900; Augustin 1907; Tschirch 1925; Winton and Winton 1939; András 1966; Fridvalszky and Nagy 1966; Konecsni 1971; Munting 1974; Dave et al. 1979; Dave 1986; Weryszko-Chmielewska and Michałojć 2011; Dias et al. 2013) and a few observations made on wild species (Filippa and Bernardello 1992). In this monograph, we include the most relevant taxonomic characters observed in nearly all the species of Capsicum (see Suppl. material 1: Appendix 1, for full details of each species).

The epicarp consists of a uniseriate epidermis covered by a smooth (e.g. C. annuum, C. chacoense, C. pubescens, Fig. 7C) or striate cuticle (e.g. C. campylopodium, C. mirabile). The cuticle can be thin (4.5–11 µm, for example, C. flexuosum, C. campylopodium) or thick (11.5–19 µm, for example, C. eximium, C. galapagoense) and usually projects towards the anticlinal walls of two adjacent epidermal cells forming conspicuous cuticular wedges (Fig. 7D). The cuticular wedges can be shallow to very deep, reaching the first layer of the mesocarp, with the cuticle extending below the inner periclinal wall of the epidermal cells (e.g. C. caatingae); these cuticular wedges vary from 7.5 to 54 µm thick. The epicarp is formed by regular epidermal cells (Fig. 6H, I), scarce stomata and sometimes sclereids (Fig. 6I, J). In cross section, epidermal cells are elongate tangentially (rectangular, Fig. 7C, D), isodiametric or flask-shaped (e.g. C. hookerianum) and in superficial view, they are polygonal, with straight (Fig. 6H) or slightly sinuous cell walls; pits appear frequently in thick- and thin-walled cells (Fig. 6I). The epicarp may have scattered sclereids amongst the epidermal cells (e.g. C. baccatum var. pendulum, Fig. 6I) or it can consist exclusively of sclereids (C. baccatum var. baccatum, Fig. 6J). Ventilating clefts are found in the epicarp of many species (Dave et al. 1979; Filippa and Bernardello 1982; this work), but their presence is not a constant feature within a species (Dave 1986).

The mesocarp consists of (5–) 7–26 layers, with up to 30 layers found in the thick pericarp of a sweet C. annuum var. annuum cultivar known as “calahorra”. The mesocarp can be homogeneous or heterogeneous; a homogeneous mesocarp is formed exclusively by parenchyma (thin-walled cells) or collenchyma (thick-walled cells) (e.g. C. rhomboideum, C. hookerianum), whilst a heterogeneous mesocarp consists of both collenchyma and parenchyma with a variable number of layers for each tissue, depending on the species (see Suppl. material 1: Appendix 1). The collenchymatous layers are always placed underneath the epicarp and both together have been referred as the “exocarp” (Roth 1977). The parenchymatous layers are underlying the collenchyma and the number of layers is usually greater than those of the collenchyma. Crystal sand and vascular bundles are found in the collenchyma or parenchyma (e.g. C. rhomboideum, C. dimorphum). A gradual increase of cell size occurs from the periphery inwards (Fig. 7E). In a few Capsicum species (e.g. the Andean clade species), the cells of the innermost layer of the mesocarp are equal in size and similar in shape to the adjacent mesocarp cells (Fig. 7B); thus, the pericarp surface facing the locule (endocarp) is smooth. In contrast, in most species, the inner surface of the pericarp is blistered due to the extensive development of the inner hypodermis (Fig. 7A, E, F); the inner hypodermis develops as a single layer of ‘giant cells’ that are initially large in the ovary wall (Augustin 1907; Munting 1974) and then increase tremendously in size towards fruit maturity (Fig. 7A, E). These giant cells are occupied by scanty cytoplasm, a large watery vacuole and a large nucleus pushed to the periphery by the vacuole. The size of the giant cells varies amongst the species; they can be nearly as long as wide, ranging from 390–1380 µm long and 300–1050 µm wide (most species) or 2–6 times longer than wide, varying from 540–2500 µm long and 150–490 µm wide (e.g. some Brazilian species). Each giant cell is attached to its neighbouring cell and the endocarp and they are surrounded by a triangular multiseriate wedge-shaped cluster of parenchymatous cells between their anticlinal cell walls (“bridge”, fide Roth 1977). The development of giant cells in the mesocarp is a derived anatomical trait in Capsicum, absent in the Andean clade and with a single reversion to the ancestral state in C. baccatum var. umbilicatum (Fig. 7B) (Carrizo García et al. 2016).

Hard inclusions of sclereids (stone cells) are developed in the mesocarp of some species (5 spp., see Suppl. material 1: Appendix 1); the stone cells can be completely immersed in the mesocarp and then only visible under microscope (C. piuranum) or they are easily seen in the pericarp surface in dried fruits (e.g. C. geminifolium). There are never more than six per fruit. The presence, number and position of the stone cells are not consistent within a single species; thus, these are not useful traits in the identification of Capsicum species, as they may be in other Solanaceous genera, such as Lycianthes (Bitter 1919) or Solanum (Särkinen et al. 2018; Knapp et al. 2019).

The endocarp develops from the inner epidermis of the ovary wall; it consists of one layer of polygonal or irregular cells (surface view) with straight or sinuate cell walls. The endocarp can be homogeneous, that is entirely with thin (e.g. C. recurvatum) or pitted thick-walled cells (sclereids) (e.g. C. rhomboideum, C. pubescens, Fig. 7G, H) or it may consist of a mixture of cells with thin and thick walls forming a heterogeneous tissue (e.g. C. annuum); in this latter endocarp type, groups of pitted sclereids lie on the inner periclinal walls of the giant cells that alternate with groups of thin-walled cells placed on the multiseriate wedge (“bridge”) of parenchymatous cells (Konecsni 1971; Dave 1986).

Some structural features of the pericarp useful in species-level taxonomy are detailed in Suppl. material 1: Appendix 1.

Septum and placenta

The presence of the pungent principles (capsaicinoids) of Capsicum fruits within cells of the interlocular septum and placenta has been demonstrated by histochemical (Ohta 1962d) and radioisotopic (Iwai et al. 1979) evidence, structural and ultrastructural microscopic analyses (Suzuki et al. 1980; Zamski et al. 1987; Stewart et al. 2007) and analytical chemical procedures (Manirakiza et al. 2003; Stewart et al. 2007; Nugroho 2016; Guillen et al. 2018). The histological characteristics of the septum during fruit development have been investigated extensively in some cultivars of C. annuum and C. frutescens (Tschirch and Oesterle 1900; Ohta 1962d; Munting 1974; Dave et al. 1979; Suzuki et al. 1980), as well as some wild Capsicum species (Filippa and Bernardello 1992). In general, the septum consists of epidermal cells and a loose parenchymatous tissue with intercellular spaces. The epidermal cells are responsible for the synthesis, accumulation (in pockets or blisters) and secretion of capsaicinoids. In early stages of fruit development (about 10 days after flowering), some of these cells become secretory (Ohta 1962d; Suzuki et al. 1980; Filippa and Bernardello 1992), but they fully develop the characteristic traits of a glandular tissue (i.e. large elongate size, abundant cytoplasm, a large vacuole, many small vesicles containing electron-dense granules and a large nucleus) nearly 30 days after flowering (Suzuki et al. 1980).

An ultrastructural study demonstrated that the major reservoir of capsaicinoids is in the capsisome, a specific capsaicinoid biosynthesising and accumulating vacuole, different from the vacuoles regarded as reservoirs of organic acids (Fujiwake et al. 1980, 1982).

Capsaicinoids have also been found in the pericarp and seeds in significant or low amounts in some species and cultivars (e.g. C. chinense, C. baccatum) (Pandhair and Sharma 2008; Bosland et al. 2015; Guillen et al. 2018).

Seeds

The gross morphology of the seeds and details of the sculpturing of the seed coat for all Capsicum species are summarised in Suppl. material 2: Appendix 2. The attributes used in species descriptions are illustrated in Fig. 8.

Figure 8. 

Seed morphology and seed coat structure A Capsicum chinense B C. chacoense, longitudinal section C, D C. schottianum, cross section (D detail of the seed coat structure) E C. annuum var. annuum, detail of seed coat structure (the rectangle indicates a cell of the seed coat) F, G C. eximium, cross sections at the seed margin and seed body, respectively H C. dimorphum, cross section at the seed body. Abbreviations. aw anticlinal cell wall, bp beak prominence, em embryo, en endosperm, h hilum, im inferior seed margin, ipw inner periclinal wall, opw outer periclinal wall, sb seed body, sc seed coat, sm superior seed margin, w seed width, l seed length. Scale bars: 200 μm (A–C); 100 μm (D, F, H); 20 μm (E, G).

Seed shape (and size) is influenced by the position in the berry. The seeds are flattened to slightly angled, mostly C- or D-shaped (Gunn and Gaffney 1974), subglobose or ellipsoid, rarely reniform or teardrop-shaped. Seed shape is sometimes difficult to define in the domesticated species due to the presence of a protrusion on the seed coat (a beak-like prominence) in a vertical or nearly vertical direction above the hilum (Fig. 8A, B). This protrusion can also occur in a lateral position in some wild species with the hilum placed on the protrusion edge (e.g. C. cornutum). Seed size varies from small (1.5–2.5 mm long) and medium (2.6–3.9 mm long) to large (4–7 mm long). Capsicum lycianthoides has the smallest seeds (1.5–1.8 mm long), whereas the largest seeds are found in the domesticated C. pubescens (5.5–7 mm long). Seed colour is usually uniform and is slightly shiny to shiny when observed dry under a stereomicroscope. Seeds can be grouped in three categories based on their colour: (1) pale yellow or nearly white (C. neei) to yellow (mostly domesticated species and their close relatives); (2) brownish-yellow to brown (many Bolivian species); and (3) brownish-black to black (many Andean and Brazilian species). Seed number per berry in Capsicum species is usually 10–45, with only four seeds found in C. campylopodium and more than 50 in C. lanceolatum, C. piuranum and some domesticated taxa.

The hilum is always marginal (on the inferior margin, Fig. 8A) and its position may be medial (Fig. 9Q), subterminal (Figs 9N, 10N) or terminal (Fig. 9I). The hilum area may be inconspicuous (minute to small) or conspicuous (medium to large). The hilum area shape may be elliptical, linear, ovoid or more rarely triangular (C. minutiflorum, C. pubescens) or circular (C. tovarii); some species may have two or three different hilar area shapes.

Figure 9. 

Seeds and seed coat morphology in species of the Annuum Clade A–D C. annuum var. annuum E–H C. annuum var. glabriusculum I–L C. frutescens M–P C. chinense Q–T C. galapagoense A, I seeds with testa partly digested B seed coat with the external periclinal cell wall partly removed C, D cross section of the seed at the seed margin and seed body, respectively; E, M, N, Q untreated seeds showing subterminal hilum (E, N) and medial hilum (Q) F, J, O, P, S, T detail of a non-digested portion of the seed coat G, K testa pattern with the external periclinal cell wall removed H, L detail of testa cells R hilum. Abbreviation. opw outer periclinal cell wall. Scale bars: 200 μm (A, E, I, M, N, Q); 20 μm (B, F–H, K, L, P, S, T); 50 μm (C, J, O, R); 10 μm (D).

Figure 10. 

Seeds and seed coat morphology in species of the Baccatum Clade A–D C. baccatum var. baccatum E C. baccatum var. pendulum F–H C. baccatum var. umbilicatum I–L C. chacoense M–O C. rabenii. A, I, M seeds with testa partly digested B, G detail of a non-digested portion of the seed coat C, H, J, K, O testa pattern of treated seeds showing anticlinal cell walls with fibrils (C, K), papillae (H) and ridge (O) D detail of a testa cell E, F untreated seeds L cross section of the seed at the seed body N seed showing the subterminal elliptical hilum. Scale bars: 10 μm (H); 20 μm (B–D, G, K, L, O); 100 μm (J); 200 μm (A, E, F, I, M, N).

Capsicum annuum (and its varieties) has been the most frequently studied species (Lohde 1875; Moeller 1886, 1905; Hanausek 1888; Hartwich 1894; Wojciechowska 1972; Krstić et al. 2001, Kong et al. 2011) before our studies reported here. Some other authors focused on the development and the structure and sculpture of the seed coat of many Solanaceae species, including some of the domesticated capsicums (Souèges 1907; Gunn and Gaffney 1974; Al-Nowaihi and Mourad 1999; Dias et al. 2013). More recently, Chiou and Hastorf (2014) provided a morphometric approach, based on 27 qualitative and quantitative seed attributes for the five domesticated species. They recovered some useful traits that can be used to compare features of modern Capsicum seeds to archaeological seeds. The gross morphology and the sculpture of the seed coat provided here for each Capsicum species (Suppl. material 2: Appendix 2) highlight the importance of some characters for delimiting species, species groups or clades. Carrizo García et al. (2016) found that seed colour is useful for identifying species or small clades (e.g. brownish-black or black seeds in the Atlantic Forest and in Andean clades or the pale yellow seeds in the Annuum and Baccatum clades). The seeds of species within the same clade are also characterised by similar sculpture and the structure of the outer layer. For example, species of the Atlantic Forest clade all have a seed coat that is reticulate-tuberculate/reticulate with pillar-like structures, has deep cells in the margin and seed body and has a thin cellulosic outer periclinal cell wall in the outer layer that is easily removed (naturally or by enzyme etching).

Seed ornamentation refers to the appearance of the seed coat (testa) (Figs 918). The testa appearance varies in the majority of the species, depending on the magnification (naked eye, stereomicroscope or SEM) and whether or not the seeds are untreated or treated with a technique, such as enzyme etching. By removing the outer periclinal wall with enzyme etching, the ornamentation of the seed coat (sculpture) can be observed in detail under SEM. From the SEM data obtained of all treated seeds, the sculpture of the seed coat and variations in the anticlinal (lateral) walls of the outer epidermal layer vary between species and provide useful diagnostic features.

Five major types of seed coat sculpture were observed after the outer periclinal wall was removed by enzymatic digestion: (1) reticulate, with straight to wavy cell walls (Fig. 14B, C, M, O, for example, C. dimorphum); (2) cerebelloid, with sinuate to strongly sinuate cell walls (Fig. 15B, E, F, K, for example, C. hookerianum); (3) reticulate-cerebelloid, reticulate at the seed margins and cerebelloid in the seed body (Fig. 16A, B, for example, C. caballeroi); (4) reticulate with pillar-like outgrowths at margins (Fig. 11M, R, S, for example, C. cornutum, C. mirabile), these outgrowths due to unequal height of the anticlinal walls thickenings (Fig. 8C, D); and (5) cerebelloid with pillar-like outgrowths at margins (Fig. 11B, C, for example, C. regale and some seeds of C. campylopodium and C. schottianum). The most common combinations in the ornamentation of untreated/treated seeds in each species are: smooth to obscurely reticulate/cerebelloid (Figs 9I, K, 10A, C); reticulate, marginally tuberculate/reticulate with pillar-like outgrowths at margins (Figs 11I–K, Q–S, 12L–O, 13E–H); reticulate/reticulate (Fig. 14A–C); smooth and reticulate at margins or completely reticulate/reticulate-cerebelloid (Fig. 15I–K) (see Suppl. material 2: Appendix 2 for each species).

Figure 11. 

Seeds and seed coat morphology in species of the Atlantic Forest Clade A–D C. campylopodium E–H C. carassense I–L C. cornutum M–P C. friburgense Q–T C. mirabile A untreated seed B, I, Q seeds with testa partly digested C, F, K, O, S marginal testa pattern of treated seeds (note pillar-like outgrowth in K, O, S) D, G, H, L, P, T testa pattern at the seed body of treated seeds showing anticlinal cell walls papillate and punctate E, J, M, R treated seeds (in J showing lateral prominence) N hilar zone with a linear hilum. Abbreviation. lp lateral prominence. Scale bars: 200 μm (A, B, E, I, J, M, Q, R); 100 μm (C, G, K, N, O, S); 20 μm (D, F, H, L, P, T).

Figure 12. 

Seeds and seed coat morphology in species of the Atlantic Forest Clade A–C C. muticum D–G C. pereirae H–K C. mirum L–P C. schottianum Q–T C. recurvatum A, E, I, N, Q treated seeds B, F, J, O, S marginal testa pattern of treated seeds with pillar-like outgrowths C, G, P, T testa pattern at the seed body of treated seeds showing anticlinal cell walls papillate and punctate D, H, L untreated seeds K detail of papillate anticlinal cell walls M detail of a non-digested portion of the seed coat R hilar zone with a linear hilum. Scale bars: 200 μm (A, D, E, H, I, L, N, Q); 100 μm (B, F, J, O, R); 20 μm (C, G, K, M, P, S, T).

Figure 13. 

Seeds and seed coat morphology in species of the Atlantic Forest Clade A–D C. hunzikerianum E–I C. villosum A, G seeds with testa digested B seed showing medial and linear hilum C, H marginal testa pattern of treated seeds D, I testa pattern at the seed body of treated seeds showing anticlinal cell walls papillate (D) and punctate (I) E untreated seed F detail of a non-digested portion of the seed coat. Scale bars: 200 μm (A, B, E, G); 20 μm (C, D, F, I); 100 μm (H).

Figure 14. 

Seeds and seed coat morphology in species of the Andean Clade A–D C. dimorphum E–H C. geminifolium I–L C. lanceolatum M–P C. longifolium Q–T C. lycianthoides A, Q, S seeds untreated B, E, I, J, M seeds with testa digested C, G, K, O testa pattern at the seed body of treated seeds showing anticlinal cell walls punctate (C, J, O) and with fibrils (G, K, O) D detail of ridge H, L, P detail of testa cells F, N hilar zone R, T detail of a non-digested portion of the seed coat. Scale bars: 200 μm (A, B, E, I, J, M, Q, S); 100 μm (C, K, N, O); 20 μm (D, F, G, H, L, P, R, T).

Figure 15. 

Seeds and seed coat morphology in species of the Andean Clade A–D C. hookerianum E–H C. piuranum I–K C. rhomboideum A untreated seed B, E treated seeds C, D, G, J, K testa pattern of treated seeds showing anticlinal cell walls punctate (C, D, J, K) and papillate (G) F hilar zone H detail of testa cells densely papillate I seed partly digested. Scale bars: 200 μm (A, B, E, I); 20 μm (C, D, F, G, H, J, K).

The anticlinal cell walls observed with SEM are either: (1) papillate, with papillae 2.5–4 µm in diameter on the cell walls (Figs 10H, 11D, H, T, 12C, G, K, T, 15H, 16D, H, L, P, T, 17H, L, O, 18D, H, L); and (2) punctate, with perforations (holes, ca. 35 µm in diameter) at the bottom (Figs 9L, 11G, H, L, T, 12C, G, 14L, 16S) or uniformly distributed on the cell walls (Figs 11O, P, 12P, T, 14P, 15D, K, 17G, N).

Figure 16. 

Seeds and seed coat morphology in species of the Bolivian Clade A–D C. caballeroi E–H C. ceratocalyx I–L C. minutiflorum M–P C. neei Q–T C. coccineum A, Q seeds with testa partly digested B, F, G, J, O, R marginal testa pattern of treated seeds showing anticlinal cell walls punctate and densely papillate (G, O), punctate (J) and with ridge (R) C, H, K, P, S detail of testa cells D, L, T detail of papillae E, I, M treated seeds N linear hilum. Scale bars: 300 μm (A, E, I, M, N, Q); 50 μm (B, F, G, H, O, P); 20 μm (C, J, K, L, R, S); 5 μm (D, T).

Figure 17. 

Seeds and seed coat morphology in species of the Caatinga, Longidentatum, Flexuosum and Tovarii Clades A–D C. caatingae E–H C. parvifolium I–L C. longidentatum M–O C. flexuosum P–R C. tovarii A, M, P seeds with testa partly digested (in A hilum in terminal position, P hilum subterminal) B detail of a non-digested portion of the seed coat C, F, J, N, Q marginal testa pattern of treated seeds showing anticlinal cell walls papillate (F), with fibrils (J), punctate (N) and with fringe (Q) D, G, K, O, R detail of testa cells E, I treated seeds H, L papillae on anticlinal cell walls Scale bars: 200 μm (A, E, I, M, P); 20 μm (B, D, G, H, J, K, L, N, O, Q, R); 100 μm (C, F).

Figure 18. 

Seeds and seed coat morphology in species of the Purple corolla and Pubescens Clades A–D C. cardenasii E–H C. eshbaughii I–L C. eximium M–P C. pubescens A, M untreated seeds (in A hilum terminal, M hilum medial) B treated seed C, G, K, O testa pattern of treated seeds showing anticlinal cell walls with fringe (C) and fibrils (G) D, H, L, M detail of testa cells E, I, N seeds with testa partly digested (in I hilum subterminal) F, J detail of a non-digested portion of the seed coat. Scale bars: 200 μm (A, B, E, I, M, N); 20 μm (C, D, G, H, J, K, L, P); 50 μm (F, O).

The distal end of the anticlinal cell walls may have three different types of appendages: (1) a thin ridge (Figs 9K, 10D, H, O, 11K, L, O, P, T, 12P, T, 14D, 15K), which is very common in the genus; (2) a more or less wide fringe (Figs 16K, L, R, S, 17O, Q, R, 18C); and (3) strands of thickenings differentiated as finger-like laciniations or fibrils (hair-like structures, Figs 9G, 10C, K, 14G, H, K, L, P, 17J, 18G, K).

Embryo

The embryo in Capsicum is usually imbricate, meaning that the cotyledon tips are parallel or almost parallel to the radicle (Gunn and Gaffney 1974) and, less frequently, annular (cotyledon tips point toward the radicle, for example, C. dimorphum) or coiled (cotyledons coiled twice, Fig. 8B; for example, C. chacoense). The endosperm is firm, whitish and relatively abundant (Fig. 8B, C). Germination in Capsicum is epigeal.

Floral biology and pollination

Most work on pollination and floral biology in Capsicum has been done with the domesticated species used for their fruits. Capsicum species are generally reported to be self-compatible, although the studied cases concerned mainly domesticated species and a few wild relatives (e.g. C. annuum and C. galapagoense from the Annuum clade or C. baccatum and C. chacoense from the Baccatum clade) (Onus and Pickersgill 2004; Cauich et al. 2006). Self-incompatibility has only been documented in C. cardenasii, as well as in some accessions of C. pubescens (Yaqub and Smith 1971; CCG, pers. obs). Successful self-pollinations have also been achieved in C. cardenasii (CCG pers. obs.), suggesting a variable degree of self-(in)compatibility.

Flowering phenology, with particular attention to the timing of gynoecium and androecium maturity, has been studied to improve pollination, fertilisation and, ultimately, the fruit set, as well as to analyse the chances of doing targeted crosses (e.g. through bud pollination). Anther dehiscence has been registered to occur after flower opening, whereas the stigma is receptive before anther dehiscence, even in the buds and receptivity is maintained throughout the lifespan of the flower (Alleemullah et al. 2000; Crispim et al. 2017; Peña-Yam et al. 2019). Anthers open by a longitudinal slit; the anther walls extend outwards completely, fully exposing the pollen grains, which are held together in clumps on the anther walls by means of pollenkitt. Pollen viability is highest at the time of anther dehiscence, gradually decreasing towards the end of the flower anthesis (Peña-Yam et al. 2019; CCG, pers. obs.). Temperature has proven to be a key factor for pollen and pollen tube development, as well as for fertilisation, with high temperature being unfavourable (Aloni et al. 2001; Erickson and Markhart 2002; Reddy and Kakani 2007; Kafizadeh et al. 2008).

Capsicum species have served as models to examine unilateral self-incompatibility using reciprocal interspecific crosses mainly between the domesticated species and their closest relatives, both within and between clades that include domesticated species (e.g. Tong and Bosland 2003). Interspecific hybridisations can be successful amongst closely-related species (e.g. C. annuum × C. frutescens, C. pubescens × C. eximium), but crosses are not always successful or are only so in one non-reciprocal sense (e.g. C. chinense-male × C. baccatum-female, C. annuum-male × C. pubescens-female) (Emboden 1961; Peter and McCollum 1984; Tong and Bosland 2003; Onus and Pickersgill 2004; Vilagelim Costa et al. 2009; da Silva Monteiro et al. 2011; Kamvorn et al. 2014; Martins et al. 2015).

Nectar production and its presentation in Capsicum is due to the formation of nectar ducts, structures also found in other Solanaceae genera, such as Jaltomata Schltdl., Physalis L. (Vogel 1998; Cocucci 1999) and Mellissia Hook.f. (Fay et al. 2007). The base of the ovary contains a nectary disc that is enveloped by the narrow basal tube formed by the corolla and the adnate filaments of the stamens. The nectar passes through individual ducts formed between adjacent filaments and is exposed on the corolla limb as isolated droplets that are radially arranged at the base of the limb and alternating with the filaments (Fig. 5I). Even though nectar ducts are elaborate systems, such mechanism of nectar presentation, as an easily accessible reward, would lead to generalist pollinators (Vogel 1998), such as bees. Bees have been documented as effective pollinators of Capsicum (Kristjansson and Rasmussen 1991; Rabinowitch et al. 1993; Cauich et al. 2006; Cruz and Olivera de Campos 2007; Palma et al. 2008).

Fruit and seed dispersal

Wild Capsicum species typically produce colourful, juicy, fleshy, conspicuous, many-seeded berries that are attractive to their consumers. The fruits in most Capsicum species contain capsaicinoids (mainly capsaicin), the chemical principles responsible for their pungency, which are highly concentrated in the placental and septum tissues. Some authors (Mason et al. 1991; Norman et al. 1992; Tewksbury and Nabhan 2001) have experimentally proved that capsaicin repels rodents (house mice: Mus musculus; deer mice: Peromyscus maniculatus; Norway rats: Rattus norvegicus; cactus mice: Peromyscus eremicus; packrats: Neotoma lepida). This contrasts to the insensitivity of birds to capsaicin (European starlings: Sturnus vulgaris; parrots: Amazona spp.; pigeons; red-winged blackbirds: Agelaius phoeniceus; cedar waxwings: Bombycilla cedrorum; house finches: Carpodacus mexicanus; curve-billed thrashers: Toxostoma curvirostre; northern cardinal: Cardinalis cardinalis, northern mockingbird: Mimus polyglottos), which may reflect basic differences in trigeminal chemoreception (Mason et al. 1991 and literature therein; Norman et al. 1992; Tewksbury et al. 1999; Levey et al. 2006). Tewksbury and Nabhan (2001) proposed the directed deterrence by capsaicin hypothesis which predicts that this compound in ripe peppers fruits functions to selectively discourage fruit consumption by non-seed dispersing vertebrates without deterring beneficial seed dispersers. Levey et al. (2006) tested this hypothesis in the field with C. annuum var. glabriusculum (North America: Arizona) and C. chacoense (South America: Bolivia), both of which have upright, pungent, red fruits. Their studies support the hypothesis that Capsicum fruit consumption is diurnally facilitated by bird communities (Passeriformes: Fam. Tyrannidae, Turdidae, Mimidae, Cardinalidae and others), whereas mammals avoid consumption of the spicy fruits because of aversion to capsaicin (Jordt and Julius 2002). In addition, the gut passage and retention time of pepper seeds in birds enhance seed germination rates (Tewksbury and Nabhan 2001; Levey et al. 2006; Tewksbury et al. 2008a; Fricke et al. 2013). Frugivorous birds would be the legitimate seed dispersers of chili peppers. Rodents appear to be only significant as Capsicum seed predators, first when they directly consume the non-pungent fruits of C. chacoense, resulting in the destruction of the seeds and 0% germination (Tewksbury and Nabhan 2001) and second, when they consume the post-dispersal seed defecated by birds (Noss and Levey 2014).

It is expected that the non-pungent red or orange fruits of the Andean Capsicum species are also dispersed by birds. It would be would be interesting to test in nature if the directed deterrence hypothesis functions in a similar way in the Brazilian Atlantic forest species whose greenish-golden yellow fruits are not as showy as the red-fruited species; they are pendent and are somewhat masked amongst the copious green foliage of the plant, perhaps attractive to fauna moving underneath the plants.

Cytogenetics

Capsicum species are mostly diploid, with two chromosome numbers: 2n = 24 (x = 12) and 2n = 26 (x = 13), the latter appearing only in wild species (Heiser and Smith 1958; Pickersgill 1971, 1977, 1991; Eshbaugh et al. 1983; Moscone 1990, 1992, 1993, 1999; Moscone et al. 1996a, 2007; Tong and Bosland 2003; Pozzobon and Schifino-Wittmann 2006; Pozzobon et al. 2006; Barboza et al. 2011, 2019, 2020b; Scaldaferro et al. 2013, 2016; Scaldaferro and Moscone 2019). The number of species belonging to the x = 13 group has increased as new taxa have been discovered and now almost equals those with x = 12 (Table 2). In addition, although polyploidy is very rare in Capsicum, Greenleaf (1947), Pickersgill (1977) and Jha et al. (2012) described what they believed to be natural polyploids, triploids and tetraploids with 2n = 3x = 36 and 2n = 4x = 48, respectively. Polyploid forms can also be induced as has been shown (Gyorffy 1939; Nishiyama 1939, 1940; Pal and Ramanujam 1939; Pal et al. 1941; Aleksic 1960; Palfi et al. 1961; Ohta 1962c; Siskovic 1962; Raghuvanshi and Joshi 1964; Indira and Susan 1977; Panda et al. 1984; Rao 1987; Ishikawa et al. 1997; Ishikawa 2001; Kumar and Raja Rao 2003; Kumar et al. 2004; Takizawa et al. 2008; Kulkarni and Borse 2010).

Table 2.

Cytogenetic information on Capsicum species.

Taxon and voucher number n 2n Haploid karyotype formula Chromosomes with active NORs Hc amount (HKL in µm) 1C DNA co ntent in pg References
C. annuum var. annuum
No voucher cited - 24 - - - - Pickersgill 1971, 1977, 1991
cytotype 1 EAM 193, 251, 203 - 24 10 m + 1 sm + 1 st 11 sm 1.80 (68.51) 3.41* Moscone et al. 1993, 1995, 1996a, 2003*, 2007
cytotype 2 EAM 204, 252; NMCA 10544, 10272 - 24 10 m + 1 sm + 1 st 11 sm, 12 st 2.88 (70.40) 3.32* Moscone et al. 1993, 1995, 1996a, 2003*, 2007
Cuneo w.no. Doux Long des Landes w.no. - - - - - 3.83† Belletti et al. 1998
C. annuum var. glabriusculum
No voucher cited - 24 - - - - Pickersgill 1971 (as C. annuum var. minimum), 1977, 1991
cytotype 1 NMCA 10955 - 24 10 m + 1 sm + 1 st 11 sm 2.26 (59.53) - Moscone et al. 2007
cytotype 2 NMCA 10983 - 24 11 m + 1 st 1 m, 5 m 3.54 (51.95) - Moscone et al. 2007
cytotype 3 LQ w. no - 24 11 m + 1 st 11 m 2.33 (55.13) - Scaldaferro et al. 2013
cytotype 4 YSG w. no - 24 11 m + 1 st 5 m, 12 st 6.33 (53.56) - Scaldaferro et al. 2013
cytotype 5 Neth 804750009 - 24 11 m + 1 sm 12 sm 3.37 (55.43) - Scaldaferro et al. 2013
cytotype 6 PI 511885 - 24 11 m + 1 st 1 m, 5 m, 6 m, 12 st 2.97 (80.38) - Scaldaferro et al. 2013
cytotype 7 PI 511886 - 24 11 m + 1 st 1 m, 2 m, 5 m, 8 m 3.83 (70.05) - Scaldaferro et al. 2013
C. baccatum var. baccatum
Vouchers not cited - 24 - - - - Pickersgill 1977
GEB 163 - 24 11 m + 1 st 1 m, 3 m, 10 m, 12 st 7.45 (66.84) 3.71* Moscone et al. 2003*, 2007
Tuscia University, Italy w. no - - - - - 4.22 Belletti et al. 1998
C. baccatum var. pendulum
No voucher cited - 24 - - - - Pickersgill 1971, 1977
cytotype 1 EAM 192, 209 - 24 11 m + 1 st 1 m, 3 m, 12 st 7.30 (75.53) 3.71* Moscone et al. 1993, 1995, 1996a, 2003*, 2007
cytotype 2 EAM 205, 206, 247; ATH 25382; EAM & RN 211 - - 11 m + 1 st 1 m, 3 m, 10 m, 12 st 7.56 (74.31) 3.68* Moscone et al. 1993, 1995, 1996a, 2003*, 2007
Tuscia University, Italy w. no, Sao Paulo University w. no 74.92 - - - - 4.20 Belletti et al. 1998
C. baccatum var. umbilicatum
EAM 197, 253 - 24 11 m + 1 st 1 m, 3 m, 10 m, 12 st 9.06 (74.27) 3.76* Moscone 1999; Moscone et al. 2003*
C. caatingae
LBB 1560 12 - - - - - Pozzobon and Schifino- Wittmann 2006
LBB 1560 12 24 - - - - Pozzobon et al. 2006 (as C. parvifolium)
cytotype 1 ATH 25233 - 24 11 m + 1 st 12 st 5.52 (82.40) - Moscone 1993; Moscone et al. 1993a, 1995; 2007
cytotype 2 ATH 25233 bis - 24 12 m 12 m 7.47 (77.60) 5.77* Moscone et al. 2003*
(as C. parvifolium)
C. caballeroi
GEB et al. 3655 - 24 - - - - this monograph
C. campylopodium
LBB 1566 13 - - - - - Pozzobon and Schifino- Wittmann 2006
LBB 1566 13 26 - - - - Pozzobon et al. 2006
cytotype 1 ATH 25116 - 26 5 m + 6 sm + 1 st + 1 t 7 sm 32.49 (88.30) 5.74* Moscone et al. 2003*, 2007
cytotype 2 ATH 25128, 25130, 25136 - 26 10 m + 2 sm + 1 st 11 sm 20.41 (87.95) 4.53* Moscone et al. 2003*, 2007
C. cardenasii
Heiser 4196 12 - - - - - Heiser and Smith 1958
No voucher cited - 24 - - - - Pickersgill 1977
CORD 135 - 24 - - - - Moscone et al. 2007
cytotype 1 Neth 904750136 - 24 11 m + 1 sm 7 m, 12 sm 6.91 (62.00) - Scaldaferro et al. 2016
cytotype 2 AAC w. no; GEB w. no - 24 11 m + 1 sm 7 m, 12 sm 10.41 (76.01) - Scaldaferro et al. 2013, 2016
Budapest, Hungary w. no - - - - - 4.49 Belletti et al. 1998
C. chacoense
Argentina (Córdoba), no voucher cited 12 - - - - - Schnack and Covas 1947 (as C. microcarpum)
No voucher cited - 24 - - - - Pickersgill 1977
LB et al. 498 12 - - 1 m, 12 st - - Moscone 1992
cytotype 1 EAM 104, 195, 207, 250; AAC et al. 973 - 24 11 m + 1 st 1 m, 12 st 2.94 (65.02) 3.34* Moscone 1990; Moscone et al. 1993, 1995, 2003*, 2007; Scaldaferro et al. 2006 Moscone 1990; Moscone et al. 1993, 1995, 2003*, 2007
cytotype 2 LB & LG 525 - 24 11 m + 1 st - 2.44 (71.25) 3.36*
Tuscia University, Italy w. no - - - - - 3.83 Belletti et al. 1998
C. chinense
C 323, C 324 12 - - - - - Pickersgill 1966
No voucher cited - 24 - - - - Pickersgill 1977
cytotype 1 GEB et al. 797; GEB 807 - 24 11 m + 1 st 12 st 3.91 (61.31) 3.43* Moscone et al. 2003*, 2007
cytotype 2 EAM 201 - 24 11 m + 1 st 12 st 5.52 (61.36) 3.41* Moscone et al. 2003*, 2007
LBB 1720 12 - - - - - Pozzobon et al. 2006
MVR 9 - 24 11 + 1 st 12 st - - Romero da Cruz and Forni Martins 2015
Sao Paulo University w. no, Reading University U. K. w. no, I.N.R.A. France w. no - - - - - 4.02 Belletti et al. 1998
C. cornutum
LBB 1542, 1546 13 - - - - - Pozzobon and Schifino-Wittmann 2006
LBB 1546 - 26 - - - - Pozzobon et al. 2006
C. eshbaughii
CCG 91 - 24 - - - - Carrizo García et al. 2020
C. eximium
No voucher cited 12 - - - - - Heiser and Smith 1958
No voucher cited - 24 - - - - Pickersgill 1977
cytotype 1 EAM 254 - 24 11 m + 1 sm 7 m, 12 sm 4.90 (68.89) 4.06* Moscone et al. 2003*, 2007
cytotype 2 EAM 255 - - 11 m + 1 sm 7 m, 12 sm 2.10 (69.65) - Moscone et al. 2007; Scaldaferro et al. 2013; Debat et al. 2017
University of Reading, U. K. w. no, Budapest, Hungary w. no - - - - - 4.35 Belletti et al. 1998
C. flexuosum
RSu, EAM 4133 12 - - - - - Moscone 1992
LBB 1552 12 - - - - - Pozzobon and Schifino-Wittmann 2006
LBB 1552 - 24 - - - - Pozzobon et al. 2006
GEB et al. 3631 - 24 - - - - this monograph
GEB et al. 1034; JD & AIH 599 - 24 11 m + 1 st 2 m, 5 m 16.82 (103.69) - Moscone et al. 2007; Scaldaferro et al. 2013; Debat et al. 2016
No voucher cited - - - - - 7.2¤ Jarret et al. 2019 ¤
C. friburgense
LBB 1565 13 - - - - - Pozzobon and Schifino-Wittmann 2006; Pozzobon et al. 2006
C. frutescens
No voucher cited - 24 - - - - Pickersgill 1977
GEB et al. 795; EAM 200 - 24 11 m + 1 st 1 m, 12 st 5.55 (66.63) 3.40* Moscone et al. 1996; 2003*, 2007
Tuscia University, Italy w. no, Sao Paulo University w. no - - - - - 3.97 Belletti et al. 1998
C. galapagoense
No voucher cited 12 - - - - - Heiser and Smith 1958
No voucher cited - 24 - - - - Pickersgill 1977
PI 639682 - 24 11 m + 1 st 12 st 2.24 (48.66) - Moscone et al. 2007; Scaldaferro et al. 2013
C. hunzikerianum
GEB et al. 5041 - 26 - - - - this monograph
C. lanceolatum
NMCA 90016 13 - - - - - Tong and Bosland 2003
C. longidentatum
MFA & GEB 7086 - 24 12 m 12 m - - Barboza et al. 2011
C. longifolium
GEB & SLG 4821 - 26 9 m + 3 sm + 1 st 10 sm 3.77 (23.86) - Barboza et al. 2019
C. lycianthoides
GDB 85 - 26 9 m + 3 sm + 1 st 10 sm - - Scaldaferro and Moscone 2019
C. mirabile
NMCA 50029 13 - - - - - Tong and Bosland 2003
LBB 1559, 1564, 1568 13 - - - - - Pozzobon and Schifino-Wittmann 2006 (as Capsicum sp 6)
LBB 1550, 1554 13 - - - - - Pozzobon and Schifino-Wittmann 2006 (as C. buforum)
cytotype 1 ATH 25238, 25251 - 26 9 m + 2 sm + 1 st + 1 t 7 m 29.64 (83.81) - Moscone et al. 2007
cytotype 2 ATH 25238, 25255 - 26 8 m + 3 sm + 1 st + 1 t 7 m 29.25 (93.72) - Moscone et al. 2007
cytotype 3 ATH 25238, 25267 - 26 9 m + 3 sm + 1 t 9 m 30.93 (103.4) - Moscone et al. 2007
C. parvifolium
MFA & GEB 7075 - 24 11 m + 1 sm 12 sm - - Barboza et al. 2011; Romero da Cruz et al. 2017
C. pereirae
LBB 1558 13 - - - - - Pozzobon and Schifino-Wittmann 2006
LBB 1558 - 26 - - - - Pozzobon et al. 2006 (as Capsicum sp 7)
cytotype 1 ATH 26137 - 26 9 m + 1 sm + 2 st + 1 t 4 m, 11 st 11.42 (74.52) - Moscone et al. 2007
cytotype 2 ATH 25249 - 26 10 m + 2 st + 1 t 6 m, 11 st 16.04 (75.85) - Moscone et al. 2007
C. piuranum
GEB & SLG 4841 - 26 9 m + 3 sm + 1 st 10 sm 2.84 (22.97) - Barboza et al. 2019
C. pubescens
No voucher cited - 24 - - - - Pickersgill 1977, 1991
GEB 79; EAM 198, 202, 208, 256, 257 - 24 11 m + 1 st 10 m, 12 st 18.95 (80.53) 4.47* Moscone et al. 1993, 1995, 1996a, 2003*, 2007
Budapest, Hungary w. no - - - - - 4.86 Belletti et al. 1998
C. rabenii
No voucher cited 12 - - - - - Heiser and Smith 1958
No voucher cited - 24 - - - - Pickersgill 1977
LBB 1553, 1555 12 - - - - - Pozzobon and Schifino-Wittmann 2006
LBB 1524, 1553, 1555 - 24 - - - - Pozzobon et al. 2006 (as C. baccatum var. praetermissum)
cytotype 1 PI 441654 - 24 11 m + 1 st 7 m, 12 st 10.96 (72.55) - Moscone et al. 2007
cytotype 2 EFM 05-17 - 24 11 m + 1 sm 6 m, 12 sm 14.92 (76.20) - Grabiele et al. 2014; Scaldaferro et al. 2016
Budapest, Hungary w. no, Gatersleben, Germany w. no - - - - - 4.57 Belletti et al. 1998
C. recurvatum
LBB 1523 13 - - - - - Pozzobon and Schifino- Wittmann 2006
LBB 1523 - 26 - - - - Pozzobon et al. 2006 (as Capsicum sp. 2)
GEB et al. 915; GEB et al. 1629, 1632 - 26 10 m + 2 sm + 1 st 12 sm 5.73 (68.55) - Moscone et al. 2007; Scaldaferro and Moscone 2019
C. regale
AOR et al. 3034 - 26 - - - - Barboza et al. 2020b
C. rhomboideum
No voucher cited - 26 - - - - Pickersgill 1977 (as C. ciliatum)
YSG 19, 20 - 26 10 m + 1 sm + 2 st 9 m 4.88 (42.13) - Moscone et al. 2007; Scaldaferro et al. 2013; Aguilera et al. 2014
No voucher cited - - - - - 2.08¤ Jarret et al. 2019 ¤
C. schottianum
LBB 1535, 1536, 1540, 1544, 1545 13 - - - - - Pozzobon and Schifino-Wittmann 2006
LBB 1535, 1540 - 26 - - - - Pozzobon et al. 2006
ATH 25160 - 26 9 m + 2 sm + 1 st + 1 t 11 sm 23.28 (93.71) - Moscone et al. 2007
C. tovarii
No voucher cited 12 - - - - - Eshbaugh et al. 1983
cytotype 1 ATH & GEB 25653 - 24 11 m + 1 sm 10 m, 12 sm 38.91 (70.32) - Moscone et al. 2007
cytotype 2 NMCA 90008 - 24 11 m + 1 sm 6 m, 7 m, 12 sm 4.89 (67.02) - Scaldaferro et al. 2016
The Netherlands w. no - - - - - 3.97 Belletti et al. 1998
C. villosum
LBB 1538, 1539, 1543, 1557 13 - - - - - Pozzobon and Schifino-Wittmann 2006
LBB 1538, 1543, 1557 - 26 - - - - Pozzobon et al. 2006
ATH 25169; GEB et al. 1653 - 26 9 m + 3 sm + 1 t 12 sm 9.74 (75.89) - Moscone et al. 2007; Scaldaferro and Moscone 2019

Cytogenetics provides a valuable and irreplaceable source of information for solving taxonomic, evolutionary and applied questions (Poggio 1996; Guerra 2012). To this end, a comprehensive cytogenetic characterisation has been carried out in a large number of species in the genus. Techniques have included classical staining to indicate chromosome number and karyotype formula, silver staining to expose chromosomes with active nucleolar organiser regions (NORs), fluorescent banding to show heterochromatin amount (Hc) and distribution, fluorescence in situ hybridisation (FISH) to denote number and distribution of rDNA sites and flow cytometry and Feulgen densitometry to estimate nuclear DNA content (Pickersgill 1971, 1977, 1991; Moscone 1990, 1993, 1999; Moscone et al. 1993, 1995, 1996a, b, 1999, 2003, 2007; Belletti et al. 1998; Park et al. 2000; Pozzobon et al. 2006; Scaldaferro et al. 2006, 2013, 2016; Barboza et al. 2011, 2019, 2020b; Romero-da Cruz et al. 2016; Jarret et al. 2019; Scaldaferro and Moscone 2019; Carrizo García et al. 2020).

Almost half of the taxa that have been cytogenetically studied exhibit intraspecific karyotype variation, differing in karyotype formulas, number and location of active NORs, heterochromatin content and banding patterns (Moscone et al. 2007; Scaldaferro et al. 2013, 2016; Scaldaferro and Moscone 2019). The chromosome number of 2n = 26 for one species of the Atlantic Forest clade, C. hunzikerianum, is reported in this monograph for the first time, as is the number of 2n = 24 for C. caballeroi of the Bolivian clade (Table 2).

Capsicum disploidy (the presence of two basic chromosome numbers) has been examined in relation to genome size evolution and species diversification. The chromosome number 2n = 24 is dominant across the recognised Capsicum clades, whereas the 2n = 26 taxa are restricted to Andean and Atlantic Forest clades only. These last two clades are the more species-rich and include almost one-half of wild species of the genus.

Species with 2n = 24 chromosomes show rather uniform and comparatively the most symmetrical karyotypes, with the 11 m + 1 st karyotype formula, although 11 m + 1 sm is also frequent. In contrast, species with 2n = 26 karyotype formulas have more asymmetric, with nine different karyotypes amongst ten taxa. Out of these, the species of the Atlantic Forest clade are the most asymmetric, with seven different karyotype formulas found amongst them (Table 2).

It has been suggested that species with 13 chromosome pairs are derived from species with 12 pairs, since the latter have more symmetrical karyotypes (Pickersgill 1991; Moscone et al. 1993, 1995, 1996b, 2007; Tong and Bosland 2003; Barboza et al. 2011, 2019; Scaldaferro et al. 2013, 2016; Scaldaferro and Moscone 2019). However, the opposite scenario has also been proposed by Pozzobon et al. (2006), who hypothesised that x = 13 is the ancestral chromosome number of the genus and that the reduction in chromosome number is the result of the loss of the small 13th chromosome pair. Even though the origin and fate of this chromosome pair is not known, the occurrence of x = 13 taxa in distinct clades from x = 12 taxa (Carrizo García et al. 2016) suggests that the extra chromosome(s) arose and/or was lost on more than one occasion. It is conceivable that two 2n = 26 subgroups with asymmetrical karyotypes have arisen via a centric fission in a metacentric chromosome (Moscone et al. 2007; Scaldaferro et al. 2013). One subgroup of 2n = 26 is composed by all species of the Andean clade, which have smaller genomes, single heterochromatic banding patterns and one nucleolar organiser region (NOR) per haploid complement (Scaldaferro pers.obs.). The other subgroup includes C. mirabile, C. campylopodium, C. cornutum, C. friburgense, C. pereirae, C. recurvatum, C. schottianum and C. villosum, all representatives of the Atlantic Forest clade. These taxa have larger genomes and rich heterochromatic regions with complex banding patterns. They also contain AT-rich, GC-rich and moderately GC-rich satellite DNA in addition to one or two NORs (Moscone et al. 2007; Scaldaferro et al. 2013; Scaldaferro and Moscone 2019). The 13th chromosome pair shows distinctive characteristics amongst the subgroups.

Heterochromatin amount (Hc), indicated as percentage of haploid karyotype length (HKL), is quite variable in the genus (from 1.80 to 38.91) and correlates positively with the HKL in most of the taxa. Capsicum annuum and C. tovarii have the lowest and highest Hc, respectively, but across clades, the Annuum clade has the lowest Hc, whereas the highest Hc is found in the Atlantic Forest clade (Table 2).

DNA content analysis and characterisation of the 5S and 18S-5.8S-26S (45S) rDNA by FISH has been completed for only a few species of Capsicum (Belletti et al. 1998; Park et al. 2000; Moscone et al. 2003; Scaldaferro et al. 2006, 2016; Romero-da Cruz et al. 2016; Jarret et al. 2019; Scaldaferro and Moscone 2019). Although information is still lacking for many species/clades (e.g. the complete Atlantic Forest clade), genome sizes are highly variable between Capsicum species. This range of variation may be caused by different components of the genome that can modify the structure and composition of DNA; for example, more than 81% of the genome is composed of transposable elements in C. annuum (Qin et al. 2014).

Domestication

Five Capsicum species, C. annuum, C. chinense, C. frutescens, C. baccatum and C. pubescens, were independently domesticated for their fruits in different areas of Central and South America (Eshbaugh 1970, 1979; Pickersgill and Heiser 1977; Aguilar-Meléndez et al. 2009; Albrecht et al. 2012a, Albrecht et al. 2012b; Kraft et al. 2014; Scaldaferro et al. 2018). Archaeological records from ca. 8000 BCE indicate the presence of Capsicum species remains associated with human settlements in northern Peru (Dillehay et al. 2017). At least three Capsicum species (C. frutescens, C. chinense and C. baccatum) would have been cultivated/consumed along the Andes from around 4000–4500 BCE (Perry et al. 2007; Clement et al. 2010; Kraft et al. 2014). Paleobiolinguistic data suggest that Mesoamerican cultures had names for chili peppers by ca. 4500 BCE, such as in the Proto-Otomanguean language (Brown et al. 2013; Kraft et al. 2014), evidence that chilies were recognised and possibly consumed by the local people at that time. Different sources of evidence reveal that Capsicum species were used by Meso- and South American peoples as spices due to their pungent fruits (Yacovleff and Herrera 1935; Powis et al. 2013). On the whole, archaeological remains show that Capsicum species, together with beans (Phaseolus spp.) and several pumpkins (Cucurbita spp.), were amongst the first cultivated plants in the Americas (Pickersgill 1969b) that were domesticated by humans. Domesticated chili peppers spread outside the Americas after Columbus’ expeditions in the 15th century, first introduced into Europe and subsequently into Africa and Asia (Powis et al. 2013). Since that time, thousands of landraces have been developed in different regions around the world, some of which are currently recognised as secondary diversification centres (Nicolaï et al. 2013).

Domestication processes typically modify a few genes that affect domestication traits (Guerra-García and Piñero 2017). Capsicum species are cultivated for their fruits and are likely to have low quantities of proteins, carbohydrates and fats, then it is the presence of capsaicinoids, the compounds responsible for fruit pungency, the valuable and desired feature that may have led to its domestication. In cases like Capsicum, in which the target of selection has been the dispersal unit, modifications directed to reduce dispersal efficiency would increase the yield. For Capsicum, selection was orientated towards traits that would diminish fruit natural dispersal, as well as their consumption by birds, such as non-deciduous fruits and larger sizes. In addition to the loss of natural dispersal mechanisms, plants subjected to domestication can present other modifications regarded as part of the domestication syndrome in Capsicum, that is: changes in the reproductive system, increased morphological variability, changes in habit, loss of seed dormancy, loss of chemical or mechanical protection, as well as a variable degree of fruit pungency (Gepts 2004; Pickersgill 2007; Luna-Ruiz et al. 2018). All or some of these traits can be observed in the different domesticated species/varieties that belong to the Annuum (e.g. C. annuum var. annuum), Baccatum (e.g. C. baccatum var. pendulum) and Pubescens (C. pubescens) clades.

Distribution and habitat

Capsicum species are widely distributed across the Americas, from central Argentina and southern Brazil to the southern extreme of the United States of America, although most of the clades recognised here correspond to a particular geographic region (Fig. 19). Approximately half of the species are country-level endemics in South America: 14 species in Brazil (C. caatingae, C. campylopodium, C. carassense, C. cornutum, C. friburgense, C. hunzikerianum, C. longidentatum, C. mirabile, C. mirum, C. muticum, C. pereirae, C. recurvatum, C. schottianum and C. villosum), six species in Bolivia (C. caballeroi, C. cardenasii, C. ceratocalyx, C. eshbaughii, C. minutiflorum and C. neei) and two species in each of Ecuador (C. benoistii and C. galapagoense) and Peru (C. piuranum and C. tovarii). Another five species are native to only two countries: C. longifolium and C. hookerianum to Ecuador and Peru, C. lycianthoides to Ecuador and Colombia, C. eximium to Argentina and Bolivia and C. rabenii to Brazil and Paraguay. The remaining species have wider distributions, the most widely distributed are C. annuum var. glabriusculum, C. baccatum var. baccatum, C. chacoense and C. rhomboideum. Species richness is highest in the Andean Region (22 species), concentrated in Bolivia and Peru, each with 14 species.

Figure 19. 

Capsicum geographic distribution. Georeferenced collection points of all wild Capsicum species/varieties. Circles are coloured by clades (Andean: orange; Atlantic Forest: bright green; Flexuosum: bright light blue; Caatinga: lilac; Longidentatum: dark blue; Bolivian: yellow; Purple corolla: red; Tovarii: fuchsia; Baccatum: dark green; Annuum: pale light blue).

Wild Capsicum species are found in a wide variety of habitats, from xeric shrublands to rainforests (Suppl. material 3: Appendix 3). These habitats represent around 120 ecoregions, which belong to 10 different biomes (Suppl. material 3: Appendix 3). The high number of ecoregions represented is mainly due to the broad distribution of C. annuum var. glabriusculum (Fig. 24), spanning a total of ca. 100 ecoregions (particularly diverse in the southern United States of America, Mexico and Central America, including both continental and insular ranges) from nine out of the 10 possible biomes registered in Capsicum (Suppl. material 3: Appendix 3). Most species are found in habitats that correspond to the ‘tropical and subtropical moist broadleaf forest’ biome, except for C. cardenasii and C. galapagoense, the first of which occupies ‘montane grasslands and shrublands’ and ‘tropical and subtropical dry broadleaf forests’ biomes and the second of which occupies ‘deserts and xeric shrublands’ (Suppl. material 3: Appendix 3). The ‘tropical and subtropical dry broadleaf forests’, as well as the ‘tropical and subtropical grasslands, savannahs and shrubland’ biomes are next in number of species and include representatives of most clades (except for C. tovarii from the Tovarii clade; Suppl. material 3: Appendix 3). The remaining biomes are less represented, which encompass forests, shrublands and/or grasslands, ranging from desert to flooded conditions, including mangroves, as well as both tropical and temperate climates. The richest and most diverse clades in terms of biomes (and ecoregions) are found amongst the Baccatum and Annuum clades that include the domesticated species and their allies.

Materials and methods

Trichomes

To analyse the trichomes, temporary preparations of the epidermis of leaves, stems, calyx and corolla were made by making direct peels of the epidermis or cross sections; observations were made under light microscope and drawings were done with the help of a camara lucida.

Fruit anatomy

Mature fruits were used to analyse the anatomy of the pericarp. Fresh fruits were collected in the wild, bought at various markets (domesticated species) or obtained from plants cultivated at the University of Cordoba (Argentina) (see Suppl. material 1: Appendix 1). Fruits were fixed in formalin-acetic acid-ethanol (FAA; 3.7% formaldehyde; 5% glacial acetic acid; 50% ethanol) for at least 48 hours and then small pieces of 3–5 × 3–5 mm from the middle part of the fruit were examined. For light microscopy, fixed material was dehydrated through an ascendant ethanol series (50°, 60°, 70°, 80°, 90°, 95°, 100°), clarified in pure xylene and embedded in Paramat(TM). The samples were sectioned at 12 μm thick with a Minot rotary microtome. Sections were double-stained using Astra Blue/Basic Fuchsin (Kraus et. al. 1998; Zarlavsky 2014). Cross-sections were also made by hand. These were stained with safranin that stains intensely lignified cell walls (sclereids), which can then be easily differentiated from the cellulose-walled parenchyma cells. The cuticle was detected with Sudan IV. Outer and inner epidermal peels were obtained from the upper, middle and lower part of the fruit and were stained with safranin. The stained sections and peels were mounted in a drop of 50% glycerine solution. Measurements of the thickness of the cuticle and cuticular wedges, as well as the giant cells’ size, were made using a stage micrometer. Camera lucida drawings of pericarp sections are shown in many of the figures. Only dried fruits from herbarium specimens could be analysed for some species. Here, fruits were hydrated and kept in water at 90 °C for the necessary time (1–2 hours) to soften the tissues to be able to perform sectional cuts or pericarp peels.

Seeds

Mature seed samples were taken from herbarium material or collected from wild or cultivated sources (see Suppl. material 2: Appendix 2). All seeds were selected and analysed in three ways for: (1) stereomicroscope (SM), (2) scanning electron microscopy (SEM) and (3) light microscope (LM) observations. The processing of the seeds for each case was as follows: (1) seeds were removed from fresh or hydrated fruits (untreated seeds) and gross morphology observations were made on dried seeds with a Zeiss Stemi 2000-C stereomicroscope (SM) at 32× magnification; (2) for scanning electron microscopy (SEM), seeds were prepared following the enzyme etching technique (Lester and Durrands 1984) to dissolve the outer cell walls, affixed to aluminium stubs with double-sided adhesive tape, then coated with gold and examined using either a JEOL JSM 35 CF SEM (LABMEM, National University of San Luis, Argentina) or a FE-SEM Sigma (LAMARX, National University of Córdoba, Argentina; (3) for sectioning for light microscopy, seeds were soaked in boiling water, washed with a diluted sodium hypochlorite solution (10%) to remove possible fungal hyphae and rinsed in distilled water; cross sections were free-hand cut with glass knives and then stained with Astra Blue and safranin (Johansen 1940). Terminology for seed (and embryo) morphology largely follows Gunn and Gaffney (1974), Zhang et al. (2005) and Chiou and Hastorf (2014).

Cytogenetics

Chromosome counts, cytogenetic information and DNA content are based on voucher specimens for which we were able to verify their correct identifications (vouchers cited in Table 2) or based on studies with taxonomic identity, verifiable from the descriptions material (vouchers not cited in Table 2). We do not cite all the extant cytogenetic literature in Table 2 for the domesticated species and some related wild species, since these works are not comparable with the karyotyping nomenclature systems (e.g. Levan et al. 1964; Moscone et al. 1996a) currently used; this means that some works have been excluded (Huskins and La-Cour 1930; Dixit 1931; Yamamoto and Sakai 1932; Tokunaga 1934; Raghavan and Venkatasubban 1940; Chennaveeraiah 1947; Sinha 1950; Vazart 1950, 1951; Eshbaugh 1964; Chennaveeraiah and Habib 1966; Shopova 1966; Datta 1968; Carluccio and Saccardo 1977; Limaye and Patil 1989; Alcorcés de Guerra 2001; Nascimento Sousa et al. 2015).

Taxonomy

The monograph is based on results from many years of herbarium study and field work to collect these taxa across South America. Fresh material was preserved in FAA (formaldehyde–acetic acid–ethanol) or ethanol (70°) to perform measurements of reproductive organs using a Zeiss Stemi 2000-C stereomicroscope at 6.5–50× magnification or trichomes using a Leitz light microscope at 10–40× magnification. Descriptions were based on living plants observed during fieldwork and examination of ca. 6,900 herbarium specimens loaned from or inspected at 213 herbaria (acronyms follow Thiers 2021): A, AAU, AC, ALCB, ANDES, AS, ASE, ASSAM, B, BA, BAA, BAB, BAF, BHCB, BHZB, BKL, BM, BOLV, BR, C, CAL, CAS, CDS, CEN, CEPEC, CESJ, CHEP, CM, CNMT, COAH, COL, COLO, CONC, CONN, CORD, CR, CRI, CTES, CUVC, CUZ, DAV, DUKE, E, EAC, EAP, EFC, ENCB, ESA, ESAL, F, FCQ, FLAS, FLOR, FMB, FPS, FSU, FUEL, FURB, G, GA, GB, GBH, GH, GL, GOET, GUA, GUAY, HAL, HAJB, HAMAB, HAO, HAS, HB, HBG, HBR, HCF, HEH, HEPH, HFSL, HJ, HOXA, HRB, HRCB, HSB, HST, HSTM, HUA, HUAZ, HUCP, HUCS, HUEFS, HUEM, HUFU, HUFSJ, HULE, HURB, HURB, HUSA, HUSU, HUT, HVC, IAC, IAN, IEB, IBN, ICN, IND, INPA, IPA, ITIC, JAU, JAUM, JPB, K, KFTA, L, LAGU, LE, LINN, LP, LPB, LIL, LOJA, LUSC, M, MA, MAC, MCNS, MBM, MBML, MEDEL, MEL, MERL, MEXU, MICH, MISS, MNES, MO, MOL, MPU, MU, MY, NA, NDG, NY, OUPR, OXF, P, PACA, PEL, PEUFR, PH, PMSP, PR, PSO, PUL, PY, PYO, Q, QAP, QCA, QCNE, R, RB, REG, RIOC, RSA, RUSU, S, SAMES, SEV-H, SF, SMDB, SI, SJRP, SP, SPF, SPSF, TEFH, TUB, TULV, U, UB, UC, UDBC, UEC, UFG, UFP, UFRN, UNA, UNAH, UNOP, UNR, UOJ, UPCB, UPS, US, USCG, USF, USM, USZ, UT, VEN, VIC, VIES, VT, W, WAG, WU, WIS, XAL, YU and Z. Digital images were also accessed via different repositories, such as Global Plants (https://plants.jstor.org/), INCT Herbário Virtual (http://inct.splink.org.br), Herbário Virtual Reflora (http://reflora.jbrj.gov.br/reflora/herbarioVirtual), Atlas of the Florida Plants (http://florida.plantatlas.usf.edu) or Botanical Collections Databases of the following herbaria: B (http://ww2.bgbm.org/herbarium/default.cfm), COL (http://www.biovirtual.unal.edu.co/es/colecciones/search/plants/), F (http://fieldmuseum.org/explore/department/botany/collections), G (http://www.ville-ge.ch/cjb/bd.php), GH (https://huh.harvard.edu/pages/digital-resources), MA (http://colecciones.rjb.csic.es/), MPU (https://collections.umontpellier.fr/collections/botanique/herbier-mpu/base-herbier-mpu), NY, P (https://science.mnhn.fr/institution/mnhn/collection/p/item/search/form), S (https://herbarium.nrm.se/), US (https://collections.nmnh.si.edu/search/botany/), W (http://herbarium.univie.ac.at/database/search.php) and Z (http://www.herbarien.uzh.ch/index.html).

Measurements of dried material were made from dissections of flowers or fruits rehydrated in hot water, supplemented by measurements from living materials. Information about flower, fruit and seed colour was taken mainly from our own field observations and, in a few cases, flower colour was described from herbarium label data (e.g. C. hookerianum). The terminology used in the mature fruit descriptions of the domesticated species is based on the list of descriptors for Capsicum (IPGRI et al. 1995); pungency of immature and mature fruits was tested by tasting them in the field.

Distribution maps were produced using QGIS 3.16.0-Hannover (QGIS Development Team 2019) and were based on georeferenced data of all the herbarium collections analysed. Maps with boundaries of countries were imported from Natural Earth (www.naturalearthdata.com); elevation maps were obtained from the DIVA-GIS (http://diva-gis.org/gdata) spatial data. Specimens with latitude and longitude data on the labels were mapped directly; collections without geographical coordinates were georeferenced using different georeferencing tools, such as the point-radius method (determination of latitude and longitude and the maximum error distance), search for localities in Google Maps, Google Earth and GraphHopper Maps or in available databases with localities already georeferenced.

Preliminary conservation status was assessed using IUCN (2022) criteria B (for the relatively widespread species) and C (for some species with very restricted occurrences). We have given more weight to the extent of occurrence (EOO) in the threat assessments than to the area of occupancy (AOO), since the EOO is very sensitive to georeferencing bias and collecting effort (Knapp et al. 2020). The extent of occurrence and area of occupancy were calculated using the Geospatial Conservation Assessment Tool GeoCAT (Bachman et al. 2011; GeoCAT 2020).

Typification of cultivated taxa has been a particularly difficult task. For many taxa, the authors did not cite specimens or locality of the type. We searched for original material in potential herbaria and when we succeeded or duplicates were found, we designated lectotypes. In other taxa, lectotypifications were based on an illustration cited by the author in the protologue (e.g. Fingerhuth’s illustrations). For taxa recognised only as synonyms, we have cited the taxa in synonymy and indicated that duplicates have not been found rather than neotypifying these taxa. In cases where taxa were described from collections of living material cultivated in botanical gardens from unknown origin or the original material was destroyed in World War II, we designated neotypes when probable original material could be found (e.g. in C. ovatum) or using a modern collection (C. flexuosum). However, for the species described by Philip Miller (1768) in his“ Gardener’s Dictionary”, we have postponed typifications. Specimens made from plants grown by Miller at Chelsea Physic Garden in London (UK) are found in several different herbaria, mostly at BM and its associated historical herbaria; since we were unable to visit these herbaria for searching Miller’s original material due to restrictions for the current pandemic caused by Covid-19, we do not typify them here, leaving that for a separate study when these materials, including any non-digitised specimens, can be studied in detail.

In cases where specific herbaria have not been cited in protologues, we designate lectotypes rather than assuming holotypes exist (McNeill 2014). For cases in which the collection type was based on syntypes, we have examined as many as possible, choosing the best-preserved specimen as the lectotype; in all cases, we give the reasons for our choice in the species discussions.

Type specimens are cited with their barcodes in square brackets after each herbarium acronym, according to the style used in each herbarium (e.g. GOET003420 or MO-562486); we also indicate the sheet number after the barcode when available (i.e. IND-0153285, acc. # 139721). When barcodes are missing, we indicate only the sheet number (i.e. LIL acc. # 173409). In a few cases, we do not cite barcode or accession number (e.g. some type material from LE).

Identities of all numbered collections seen are presented in the List of Exsiccatae. Numbered and un-numbered collections are presented in Suppl. material 4: Appendix 4 where full citations of all the specimens examined for this treatment are presented in PDF format. We have not cited specimens of any Capsicum collected outside the Americas, with the exception of types.

Common names were taken from herbarium label data and reliable literature if we could verify the identity of taxa, but the list of common names for the cultivars of C. annuum, C. frutescens and C. chinense is not complete since we did not comprehensively examine the vast amount of literature where this information appears. Indigenous names are given in a separate paragraph for clarity indicating in brackets the indigenous language (if given). We cite only one specimen by provenance of the common and indigenous names per administrative division of each country. Uses as foods, spices or in ritual practices are cited in the species treatment and folk medicinal uses are summarised in Table 3, where the organ used and the medicinal properties, as well as the source of information, are indicated (verified specimens or literature).

Table 3.

Medicinal uses attributed to Capsicum species.

Taxa Organ Use Country Voucher/Reference
C. annuum var. annuum Colombia
Leaf, fruit Medicinal (no specification) Amazonas Alvarado C. 198
Ecuador
Fruit For snake bite Morona-Santiago Evans 4384
Peru
Leaf (juice) For pregnant women to help birth easily Loreto Williams et al. 10922
C. annuum var. glabriusculum Brazil
Leaf To cure acne Amapá Pereira et al. 2011
Colombia
Fruit To increase body temperature and for the skin fungi Huila Buendía 2
Fruit To soothe haemorrhoid pains Valle del Cauca Duque Jaramillo 4083-A
Ecuador
Leaf To reduce body temperature Orellana Carrillo & Reyes 434
Fruit Stomach medicine (for sore belly) Napo Davis & Yost 994
Fruit To kill parasites Morona-Santiago Bedoya 2
Fruit For skin diseases (measles, pox) Zamora-Chinchipe Santín et al. 100
Fruit For conjunctivitis Guatemala Kufer 99
Mexico
Leaf To relieve rashes in children (warm bath) Quintana Roo Serralta P. 104
Fruit For infected wounds Querétaro Martínez Torres 82
Fruit To treat ulcers Tabasco Orozco-Segovia 368
Fruit For skin wounds Yucatán Ucan 4617
Fruit Medicinal Yucatán Simá 517
U.S.A.
Fruit Stimulant Texas Chávez Jr. s.n.
C. chacoense Fruit Anti-rheumatic Argentina Martínez Crovetto (1981); Del Vitto et al. (1997)
Fruit Digestive Scarpa (2002)
Fruit Anti-spasmodic, vermifuge, stomach pain Filipov (1994)
Fruit extract Anti-inflammatory activity (mice) López et al. (2012)
Fruit Anti-parasitic Paraguay Friesen Ratzlaff (2017)
C. chinense Colombia
Seedling For haemorrhoids Meta Quevedo et al. 1816
Ecuador
Leaf To treat joint pains Sucumbíos Reyes & Moya 234
- Anti-parasitic Napo Bolotin 21
Fruit For stomach ache Napo Davis & Yost 993
Fruit For eye infections and coughing Napo Miller et al. 2404
Fruit For dysentery Orellana Herrera & Guerrero 186
Fruit Medicinal: cardiotonic Sucumbíos Moya & Reyes 206
Mexico
Leaf To treat wounds Yucatán Ucan Ek 4652
C. coccineum Bolivia
Entire plant In baths to relieve stomach pain La Paz Vargas 1310
C. frutescens Brazil
Leaf (infusion) Used for dizziness Minas Gerais Pereira 3219
Immature fruits For flu Minas Gerais Pereira 3219
C. frutescens Colombia
Root (infusion) To facilitate childbirth Guaviare Garzón et al. 3214
Buds To cure hand infection Guaviare Garzón et al. 3214
Fruit Medicinal Cundinamarca García Barriga 20315A
Ecuador
Leaf For fungal diseases Esmerlada Kvist 40356
Leaf, fruit To facilitate the fall of the baby’s umbilical cord Napo Siquihua 4
Fruit For snake bites Morona-Santiago van Asdall 82-59
Fruit For snake bites Zamora-Chinchipe Ortega 51
Guatemala
Fruit To treat conjunctivitis Chiquimula Kufer 100
C. pubescens Ecuador
Leaf For bites of dogs Loja Ellemann 66689
Fruit For headache, weakness and cold Loja Ellemann 66689
Fruit? Veterinary: to treat “moquillo” (catarrhal disease in dogs and cats) and “tos de nermo” (cough in horses) Peru
Oxapampa Chuck 137
C. rhomboideum Ecuador
- To heal skin eruptions Pichincha Cerón 6953

All species are illustrated with line drawings, colour illustrations or both; photos were taken by the authors of this treatment or were provided by other colleagues (credits are cited in each case). For some species (C. caatingae, C. friburgense, C. hunzikerianum, C. mirum), photos were provided and taken in the field by members of the Associazione PepperFriends (Verona, Italy); these photos have no herbarium voucher, but the identification was verified by the senior author of this treatment (GEB).

Biomes and ecoregions

Ecoregions were determined according to Olson et al. (2001). To determine the ecoregions and biomes occupied by the different species, maps were generated using QGIS 3.16.0-Hannover (QGIS Development Team 2019) superimposing layers of georeferenced data (as for the distribution maps, see above) with a layer of ecoregions obtained from Ecoregions2017©Resolve (https://ecoregions.appspot.com/).

Taxonomic treatment

Capsicum L., Sp. pl. 1: 188. 1753.

Capsicum section Decameris Bitter, Abh. Naturwiss. Vereins Bremen 24(2): 293. 1919. Type: C. dusenii Bitter

Capsicum section Capsicum, Huitième Congr. Int. Bot. Paris. Comptes Rend. Séances Rapp. & Commun. 1954, sect.4: 73. 1956. Type: C. annuum L.

Type

C. annuum L. (lectotype, designated by Britton 1918, pg. 338).

Description

Shrubs, subshrubs, rarely trees, vines or short-lived perennials or annuals, occasionally with a thick lignified xylopodium, glabrous or glabrescent or sparsely to densely pubescent with simple, branched, eglandular or glandular, uniseriate trichomes. Stems woody at the base, sometimes with fissured bark and lenticels; young stems angled, herbaceous, usually weak and fragile and occasionally somewhat scrambling. Sympodial units difoliate or unifoliate, the leaves usually geminate, blades simple, entire, concolorous or discolorous, glabrous to densely pubescent with eglandular and/or glandular simple or branched uniseriate trichomes; petioles generally well-developed. Inflorescences axillary, usually unbranched (rarely branched), with few to many (up to 20 or more) flowers clustered or, more rarely, on short rachis or spaced along an elongate rachis, sometimes with flowers solitary or paired. Flowers 5-merous (4–8-merous in domesticated species), actinomorphic, all perfect. Pedicels erect, slightly spreading or pendent, geniculate at their distal end or non-geniculate. Calyx truncate, entire, circular or five-angled in outline, often with 3–10 appendages. Corolla stellate, rotate-stellate, campanulate or campanulate-urceolate, entirely white, yellow, violet or fuchsia or with greenish-yellow and/or maroons or purple spots within, rarely entirely greenish-white or mostly purple, the lobes spreading or reflexed at anthesis, usually with interpetalar membrane. Stamens five (up to eight in domesticated species), usually equal (rarely unequal), the filaments glabrous and broadened at the base to form a staminal plaque fused to the corolla base, each plaque with two short lateral auricles, the anthers dorsifixed, ellipsoid or ovoid, yellow, cream or blue to purple, connivent in pre-anthesis, usually free when mature, dehiscent by longitudinal slits. Gynoecium usually bicarpellate, rarely 3–4-carpellate; ovary superior, glabrous, subglobose to ovoid (rarely ellipsoid), with an annular nectary at the base; styles straight or slightly curved, cylindrical or clavate, glabrous, commonly exserted beyond the anthers, sometimes heteromorphic (long, medium and short styles); stigma globose or discoid, sometimes somewhat bilobed, finely papillate. Fruit glabrous berry, globose, subglobose or somewhat elongate, the mesocarp juicy, the pericarp red, orange-red, greenish-golden yellow or, rarely, dark burgundy or purple-blue at maturity (in domesticated species, fruits of various shapes and colours), pungent or not; fruiting pedicels erect or deflexed; fruiting calyx discoid or campanulate, not accrescent or slightly accrescent. Seeds flattened to slightly angled, mostly C- or D-shaped, subglobose or ellipsoid (rarely reniform or teardrop-shaped), pale yellow to yellow, brownish-yellow to brown or brownish-black to black, seed coat smooth, reticulate or reticulate marginally tuberculate. Stone cells absent or present, if present, not more than six. Embryo usually imbricate (less frequently annular or coiled); endosperm firm, whitish and relatively abundant. Chromosome number: 2n = 24, 26 (see Table 2).

Distribution

(Fig. 19). Species of Capsicum are native to temperate, subtropical and tropical regions in the Americas, growing from southern United States of America to central Argentina and Brazil. Five taxa are widely cultivated elsewhere.

Artificial key to Capsicum

1 Stem and mature leaf blades completely glabrous, if trichomes present, sparsely distributed on the veins and margins 2
Stem and mature leaf blades variously pubescent 10
2 Calyx appendages (3–) 5, strongly incurved; flowering pedicels slightly winged and conspicuously winged in fruit; leaves coriaceous; Bolivia C. ceratocalyx
Calyx appendages absent or up to 10, straight; flowering and fruiting pedicels not winged; leaves coriaceous or membranous 3
3 Corolla tubular-campanulate to broadly campanulate, lobed less than 1/3 of the way to the base 4
Corolla usually stellate or stellate-campanulate, lobed 1/3 up to nearly halfway to the base 6
4 Calyx appendages 10, unequal (5 long, 5 short); corolla lobes recurved; fruits pungent; seeds 3–4 (–5) mm long, pale yellow to nearly white; Bolivia C. caballeroi
Calyx appendages 2–5, equal or subequal; corolla lobes erect; fruits non-pungent; seeds 1.5–2.5 mm long, brown to black 5
5 Leaves coriaceous; calyx appendages (2–) 3–5, spreading or reflexed; filaments 1–2.5 mm long; corolla broadly campanulate; Colombia and Ecuador C. lycianthoides
Leaves membranous; calyx appendages 5, erect; filaments 3–5 mm long; corolla tubular-campanulate; Peru C. piuranum
6 Inflorescences with 5–13 flowers on an elongate rachis, sometimes rachis forked; fruiting calyx with a conspicuous annular constriction at the junction with the pedicel; fruiting pedicels erect, brilliant dark purple; fruits dark blue to purple; Colombia, Ecuador and Peru C. regale
Inflorescences with 3–7 (–9) axillary flowers, rarely on a very short unbranched rachis or flowers solitary; fruiting calyx without an annular constriction at the junction with the pedicel; fruiting pedicels pendent or rarely curved, green or greenish-purple; fruits of other colours 7
7 Calyx appendages absent or 5, minute (< 0.5 mm long); corolla tube and base of the lobes with a sparse but continuous ring of glandular trichomes adaxially; fruits greenish-golden yellow; Brazil 8
Calyx appendages 2–10, longer (1–5 mm long); corolla tube and base of the lobes glabrous adaxially; fruits orange or greenish-golden yellow 9
8 Leaves membranous, elliptic to ovate; flowering pedicels 9–14 mm long, erect, geniculate at anthesis; corolla small, 4.5–6.5 (–8) mm long; stamens unequal (3+2); ovules 2 per locule; fruits 4-seeded; Brazil (Rio de Janeiro, Minas Gerais, Espírito Santo) C. campylopodium
Leaves coriaceous, elliptic to narrowly elliptic, flowering pedicels 15–30 mm long, pendent, non-geniculate at anthesis; corolla larger, 9–10 mm long; stamens equal; ovules more than 2 per locule; fruits multi-seeded (up to 20 seeds); Brazil (Bahía, Espírito Santo, Minas Gerais, São Paulo) C. pereirae
9 Major leaves narrowly elliptic; calyx appendages 2–3, triangular-compressed wings; flowering pedicels 3–8 mm long, pendent, non-geniculate; corolla 6–8.5 mm long, 8–11 mm in diameter, entirely yellow or with red-brown pigmentation within; fruits orange, non-pungent; seeds 1.7–2.3 mm long; Peru and Ecuador C. longifolium
Major leaves ovate to elliptic; calyx appendages 5 (6–10), cylindrical; flowering pedicels (13–) 20–38 (–48) mm long, erect to spreading, geniculate; corolla 10–14 (–16) mm long, (10–) 15–23 mm in diameter, white with diffuse brown-purple spots and a greenish-yellow centre within; fruits greenish-golden yellow, pungent; seeds 2.5–3.2 mm long; Brazil C. hunzikerianum
10 Pubescence mostly of branched eglandular or glandular trichomes, few simple trichomes 11
Pubescence mostly of simple eglandular trichomes, rarely furcate eglandular trichomes or simple glandular trichomes 13
11 Dense pubescence of long furcate and simple glandular trichomes; calyx appendages usually 10 (rarely 5 or up to 12); filaments 2.5–3 mm long; flowering and fruiting pedicels erect; fruits pungent; Bolivia C. eshbaughii
Dense pubescence of furcate to dendritic eglandular trichomes mixed with few simple eglandular trichomes; calyx appendages usually 5 (rarely 3–4 or 6); filaments 1.2–2.3 mm long; flowering and fruiting pedicels pendent; fruits non-pungent 12
12 Calyx appendages (4.5–) 5–8.5 mm long; corolla stellate, lobed almost halfway to the base, white with dark greenish-yellow spots within, with small glandular trichomes adaxially; style exserted ca. 1 mm beyond the anthers; seeds < 20 per fruit, 3–3.7 mm long, 2.5–2.8 mm; inflorescences usually (1–) 2–5-flowered; Brazil C. longidentatum
Calyx appendages 0.9–3 mm long; corolla campanulate or campanulate-rotate, shallowly lobed, entirely yellow or sometimes tinged greenish within, glabrous adaxially; style barely exserted beyond the anthers; seeds > 20 per fruit, 2.4–2.8 mm long, 1.8–2.2 mm wide; inflorescences usually (1–) 3–8 (–13)-flowered; Mexico to Peru C. rhomboideum
13 Corolla nearly lobed to the base, the tube 4–4.5 times shorter than the lobes; Ecuador C. benoistii
Corolla shallowly lobed or lobed halfway or to 2/3 of the way to the base, the tube as long as the lobes or 1.5 times shorter than the lobes 14
14 Staminal plaques with conspicuous auricles not fused to the corolla at the point of insertion of the filaments; Bolivia, Argentina and Paraguay C. chacoense
Staminal plaques with inconspicuous auricles fused to the corolla at the point of insertion of the filaments 15
15 Major leaves narrowly elliptic to lanceolate, the length/width ratio 5–10 (–16) 16
Major leaves ovate or elliptic, if elliptic the length/width ratio (2–) 2.5–4 (–4.5) 17
16 Calyx appendages 5, erect, green; corolla stellate, white with large purple spots and a cream centre within, with glandular trichomes in the throat and lobes adaxially; flowering pedicels erect to spreading, geniculate at anthesis; anthers blue; berry 6–7 mm in diameter, greenish-golden yellow, pungent; seeds 3.5–4 mm long, 2.5–3 mm wide; Brazil C. carassense
Calyx appendages (2–) 3–5, spreading or erect, green, greenish-purple or purple; corolla campanulate, entirely yellow or yellow with smaller maroon or purple spots within, glabrous adaxially; flowering pedicels pendent, non-geniculate at anthesis; anthers cream, yellow or rarely white; berry 7–12 mm in diameter, pale orange or orange, non-pungent; seeds 1.8–2.3 mm long, 1.3–1.5 mm wide; Colombia, Ecuador and Peru C. geminifolium
17 Flowering pedicels pendent, non-geniculate at anthesis 18
Flowering pedicels erect to slightly spreading, geniculate at anthesis 29
18 Flowers 5–7-merous; calyx thick, strongly 5–10-nerved; style heteromorphic (included at the same level as the stamens or exserted); corolla usually entirely white, dull white or greenish-white, rarely entirely purple or pale yellow; seeds pale yellow or nearly white, the seed coat smooth; fruits of various shape, size and colour; widely cultivated in the Americas 19
Flowers 5-merous (rarely 4-merous); calyx usually thin, comparatively weakly 5–10-nerved; style usually homomorphic, rarely dimorphic (included and exserted); corolla entirely yellow, yellow with maroon or purple spots within, white with greenish-yellow spots within or primarily purple or lilac; seeds usually brown or brownish-black to black, rarely yellow or pale yellow, the seed coat uniformly reticulate or reticulate and tuberculate at margins; fruits usually globose or subglobose, not more than 16 mm in diameter, orange to red or greenish-golden yellow; wild species, mostly from South America 20
19 Flowers solitary, rarely in pairs or more; petioles up to 10 cm long; corolla 8–15 mm long, entirely white, rarely entirely purple or pale yellow; fruiting calyx without a prominent annular constriction at junction with the pedicel C. annuum var. annuum
Flowers 2–4 (–5); petioles up to 3.5 cm long; corolla (5–) 6.5–8 mm long, entirely dull white or greenish-white (occasionally with purple spots); fruiting calyx with a prominent annular constriction at junction with the pedicel C. chinense
20 Corolla tubular-campanulate, 14.5–17 mm long; calyx appendages 5; stone cells 2; northern Peru C. piuranum
Corolla campanulate, campanulate-stellate, rotate-stellate or stellate, 4.5–12 (–15) mm long; calyx appendages absent or up to 10; stone cells absent or up to 6 21
21 Calyx appendages absent or up to 5, equal and minute, < 1 mm long 22
Calyx appendages 2–10, subequal or unequal, > 1 mm long 25
22 Young stems, leaves and calyx with simple eglandular trichomes mixed with small dark glandular trichomes 23
Young stems, leaves and calyx only with simple eglandular trichomes, glandular trichomes absent 24
23 Corolla campanulate to campanulate-stellate, lobed less than ⅓ of the way to the base; style dimorphic, short style 1–1.6 mm, long style 4.1–4.2 mm long; flowering pedicels 3–10 mm long; inflorescences few-flowered (2–8 flowers); fruiting pedicels erect, green; central Peru C. tovarii
Corolla stellate, lobed nearly halfway to the base; style homomorphic, 4.3–4.8 mm long; flowering pedicels longer, 7–21 (–28) mm long; inflorescences multi-flowered (5–13 flowers or up to 20 or more); fruiting pedicels pendent, green or purple; north-eastern Brazil C. caatingae
24 Leaf pair strongly dissimilar in shape and size; flower buds ovoid, purple or yellowish; fruits non-pungent; seeds 1.9–2.7 mm long, 1.8–2.1 mm wide; corolla entirely yellow or with purple or maroon spots within; Colombia, Ecuador and Peru C. dimorphum
Leaf pair similar or dissimilar in size, similar in shape; flower buds globose, white with green spots; fruits pungent; seeds 2.8–3.4 mm long, 2.2–3 mm wide; corolla white with greenish-yellow spots, rarely also with purple spots; Argentina, Paraguay and Brazil C. flexuosum
25 Calyx appendages 8–10, unequal 26
Calyx appendages 2–5 (–7), subequal 27
26 Leaves coriaceous; inflorescences (1–) 2-flowered; flowering pedicels 20–40 (–50) mm long; corolla ≥ 10 mm long, 4–6 mm in diameter; filaments 4–6 mm long; style 7–9 mm long; fruits pungent; fruiting calyx appendages appressed to the berry; Bolivia C. caballeroi
Leaves membranous; inflorescences with (1–) 2–7 flowers; flowering pedicels shorter, 8–15 mm long; corolla 7.5–9 mm long, 8–10 mm in diameter, filaments (1.5–) 1.8–2 mm long; style ca. 4 mm long; fruits non-pungent; fruiting calyx appendages spreading or reflexed; Ecuador and Peru C. hookerianum
27 Leaf pair dissimilar in size, similar in shape; major leaves ovate; corolla stellate, lobed nearly halfway to the base; fruits greenish-golden yellow, pungent; seeds 3–3.8 mm long, 2.7–3 mm wide; Colombia, Venezuela and Brazil C. parvifolium
Leaf pair markedly dissimilar in size and shape, rarely similar in shape; major leaves elliptic to lanceolate; corolla campanulate, lobed ⅓ of the way to the base; fruits orange to orange-red, non-pungent; seeds 1.8–2.8 mm long, 0.8–1.8 mm wide 28
28 Mature leaves glabrous adaxially; flowers solitary, rarely 2 per node; calyx appendages (4–) 5, spreading or strongly reflexed; corolla purple with white interpetalar membrane; North America (Mexico) and Central America (Guatemala and Honduras) C. lanceolatum
Mature leaves sparse to densely pubescent adaxially; flowers 2–5 (–6) per node; calyx appendages 2–3 (–5), erect or spreading; corolla entirely yellow or with purple or maroon pigmentation within and yellow interpetalar membrane; South America: Colombia, Ecuador and Peru C. geminifolium
29 Corolla broadly campanulate, campanulate-urceolate, rotate or rotate-stellate, lobed 1/3 or less of the way to the base 30
Corolla stellate, lobed more than 1/3 up to 2/3 of the way to the base 36
30 Stems and mature leaves with minute dark simple glandular trichomes mixed with sparse eglandular trichomes; corolla broadly campanulate or campanulate-urceolate 31
Stems and mature leaves lacking minute dark glandular trichomes mixed with abundant or sparse eglandular trichomes; corolla rotate or rotate-stellate 32
31 Major leaves (5.5–) 8.5–13 (–21) cm long; corolla campanulate-urceolate, entirely fuchsia or violet, glabrous adaxially, the lobes spreading to strongly recurved; fruits greenish-golden yellow, slightly pungent; seeds brownish-black to black; Brazil C. friburgense
Major leaves 3–5 (–6) cm long; corolla campanulate, almost completely violet or lilac and a greenish-yellow to white centre within, with short glandular trichomes adaxially, the lobes erect or spreading; fruits red, pungent; seeds pale yellow to brownish-yellow; Bolivia C. cardenasii
32 Flowers 4–8-merous; corolla 8.5–15 mm long; fruiting pedicels pendent; fruits large, > 10 mm in diameter, persistent, variously coloured; cultivated in the Americas 33
Flowers 5-merous (rarely 4-merous); corolla 4–7 mm long; fruiting pedicels erect; fruits small, < 10 mm in diameter, deciduous, orange or red; wild or semi-domesticated in South America 35
33 Leaves densely pubescent, rarely glabrescent; flower buds dark purple on pendent pedicels; style clavate; seeds 5.5–7 mm long, 4.8–6 mm wide, brownish-black to black, the seed coat reticulate; corolla dark purple or violet with a white or yellowish-green centre within C. pubescens
Leaves glabrous to sparsely pubescent; flower buds greenish-white or purple on geniculate pedicels; style cylindrical; seeds 3–5.2 mm long, 3–4 mm wide, pale yellow to yellow, the seed coat smooth to slightly reticulate; corolla white with large greenish-yellow spots and white centre within 34
34 Fruits pungent, rarely non-pungent, usually elongate, endocarp alveolate, pericarp with giant cells C. baccatum var. pendulum
Fruits non-pungent, campanulate-umbilicate, endocarp smooth, pericarp without giant cells C. baccatum var. umbilicatum
35 Leaves with dense pubescence, especially abaxially; corolla marginally purple with greenish-yellow centre; Brazil and Paraguay C. rabenii
Leaves mostly glabrescent, more rarely moderately pubescent; corolla white with greenish-yellow spots within, purple pigmentation absent; widely distributed across South America C. baccatum var. baccatum
36 Flowers 4–8-merous; style usually heteromorphic (three different lengths), when homomorphic carpels 2; fruits of various size, shape and colours; mostly cultivated or semi-domesticated plants across the Americas 37
Flowers always 5-merous; style homomorphic; fruits small, not more than 15 mm in diameter, globose or subglobose, most rarely ellipsoid or ovoid, usually red or red-orange or greenish-golden yellow; wild plants from South America 41
37 Calyx appendages absent or if present, minute, ≤ 0.5 mm long 38
Calyx appendages > 0.5 mm long 40
38 Flowers solitary, rarely in pairs or more; petioles up to 10 cm long; corolla 8–15 mm long, entirely white, rarely entirely purple or pale yellow; fruits usually large, up to 300 mm long, pungent or non-pungent C. annuum var. annuum
Flowers (1–) 2–5; petioles up to 3.5 cm long; corolla 3.75–8 mm long, entirely dull white or greenish-white (occasionally with purple spots); fruits small to medium-sized, up to 100 mm long, pungent, rarely non-pungent 39
39 Corolla glabrous adaxially; style heteromorphic; fruits highly variable in shape (domesticated), with the base obtuse or truncate (fruits subglobose in wild populations); fruiting calyx discoid or shallowly cup-shaped, with a prominent annular constriction at junction with the pedicel; ovary subglobose, 2–2.5 mm long, 2.5–3.5 mm in diameter C. chinense
Corolla with small glandular trichomes adaxially; style homomorphic; fruits usually elongate and narrowly triangular, with the base abruptly narrowed; fruiting calyx deeply cup-shaped lacking a constriction at junction with the pedicel; ovary oblong-ovoid, 2.5–4 mm long, 1.3–1.8 mm in diameter C. frutescens
40 Young leaves rugose; flowers 4–8-merous; corolla 10–15 mm long, 15–22 (–25) mm in diameter; style heteromorphic; fruiting pedicels pendent; fruits > 10 mm in diameter, persistent; seeds 5.5–7 mm long, 4.8–6 mm wide, brownish-black to black, the seed coat reticulate; cultivated from Mexico to Bolivia C. pubescens
Young leaves plane; flowers 5-merous, rarely 4-merous; corolla 6–7 mm long, (9–) 12–15 mm in diameter; style homomorphic; fruiting pedicels erect; fruits < 10 mm in diameter, deciduous; seeds 3–4.2 mm long, 2.5–2.8 mm wide, yellow, the seed coat smooth. Wild or semi-domesticated; Brazil and Paraguay C. rabenii
41 Androecium heterodynamous (three long and two short filaments); ovules two per locule; fruits 4-seeded; corolla white or cream with golden yellow or ochraceous spots within; Brazil C. campylopodium
Androecium homodynamous (filaments equal or slightly subequal); ovules more than two per locule; fruits many-seeded; corolla variously coloured 42
42 Calyx appendages absent or five, minute, < 0.5 mm long 43
Calyx appendages 2–10, the main appendages ≥ 0.5 mm long 47
43 Filaments < 2 mm long; fruiting pedicels erect; seeds pale yellow, yellow or brownish-yellow; fruits yellow, red-orange or red 44
Filaments ≥ 2 mm long; fruiting pedicels pendent; seed brownish-black to black; fruits greenish-golden yellow 46
44 Flowers 4–13 (–18) on a short rachis; style clavate; seeds 4–4.6 mm long, (–2.8) 3.2–3.75 mm wide, yellow to brownish-yellow; sprawling vines or scrambling shrubs, with stems to 7 m long; Peru, Bolivia and Brazil C. coccineum
Flowers 1–2 per axil, rarely up to 3; style cylindrical; seeds 3–4 mm long, 2.5–3.2 mm wide, pale yellow to yellow; perennial herbs or low shrubs or subshrubs up to 2 m, rarely larger 45
45 Calyx circular in outline; corolla 4–5 mm long, ca. 6 mm in diameter, with glandular trichomes adaxially; style 2.25–2.5 mm long, exserted 0.5–0.8 mm beyond the anthers; seed coat smooth; plants densely pubescent, the trichomes spreading; Ecuador: Galapagos Islands C. galapagoense
Calyx pentagonal in outline; corolla (5–) 6–8 mm long, 8–10 (–12) mm in diameter, glabrous adaxially; style 4–4.8 mm long, exserted 1.5–2 mm beyond the anthers; seed coat reticulate to obscurely reticulate; plants glabrescent to densely pubescent, the trichomes appressed-antrorse, sometimes spreading; widespread in the Americas C. annuum var. glabriusculum
46 Stems, leaves, pedicels and calyx densely pubescent with long spreading eglandular trichomes 0.5–2 mm long; major leaves elliptic to narrowly elliptic; corolla white with large greenish-yellow spots and sparse diffuse purple or brown spots within, the lobes widely triangular; Brazil (Rio de Janeiro) C. muticum
Stems, leaves, pedicels and calyx glabrescent to moderately pubescent, with short antrorse eglandular trichomes 0.25–0.5 mm long; major leaves elliptic to ovate; corolla white with large or small purple or brownish spots and large greenish-yellow spots within (in some populations purple or brownish pigmentation completely absent); the lobes triangular or ovate; Brazil (Minas Gerais, Rio de Janeiro and São Paulo) C. schottianum
47 Leaves coriaceous; calyx appendages strongly incurved, flattened laterally; Bolivia C. ceratocalyx
Leaves membranous; calyx appendages straight or recurved, filiform or cylindrical 48
48 Fruiting pedicels usually erect, rarely pendent; fruits red; seeds nearly white or yellow to brown 49
Fruiting pedicels pendent; fruits greenish-golden yellow; seeds brownish-black to black 52
49 Calyx strongly 10-nerved with prominent venation; seeds pale yellow to nearly white, the seed coat smooth and reticulate at margins; Bolivia C. neei
Calyx slightly 5-nerved with inconspicuous venation; seeds yellow to brown, the seed coat faintly reticulate and slightly tuberculate at margins 50
50 Inflorescences many-flowered (4–18 flowers); pedicel scars prominent, corky; filaments < 2 mm long; sprawling vines or scrambling shrubs; Peru, Bolivia and Brazil C. coccineum
Inflorescences few-flowered (up to 5 flowers); pedicels scars inconspicuous; filaments ≥ 2 mm long; erect shrubs or subshrubs 51
51 Calyx appendages 5, 1–1.5 mm long; filaments 2–2.5 mm long; corolla yellow with small and faint greenish-yellow spots within; seeds 4–4.5 mm long, 3–3.5 mm wide, dark brown; Bolivia C. minutiflorum
Calyx appendages (4–) 5, 1.2–2.7 (–3) mm long; filaments 2.7–3.8 mm long; corolla lilac, purple or magenta with a continuous greenish-yellow or ochre tube within, sometimes the corolla white with greenish-yellow spots; seeds 2.5–4 (–4.2) mm long, 2.1–3 mm wide, brownish-yellow; Bolivia and Argentina C. eximium
52 Calyx appendages five 53
Calyx appendages 10 or 6–10, rarely five 54
53 Plants densely pubescent, the stem trichomes spreading; Brazil: Rio de Janeiro, Minas Gerais and São Paulo C. villosum
Plants glabrescent to moderately pubescent, the stem trichomes antrorse; Brazil: Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro and São Paulo C. mirabile
54 Calyx appendages 10, subequal; filaments 3–3.2 mm long; style barely exserted beyond the anthers; anthers lilac or pale blue; corolla almost entirely purple; Brazil (São Paulo) C. mirum
Calyx appendages ranging from 5 to 10, unequal; filaments 1.4–2.5 mm long; style exserted 0.8–1.3 mm beyond the anthers; anthers yellow, light green or grey; corolla white with greenish-yellow or purple spots 55
55 Flowering calyx appendages cylindrical or triangular-compressed, glabrous to moderately pubescent with antrorse trichomes, the longest appendages 1–2.5 mm; corolla 6–7 mm long, ca. 11 mm in diameter, white with greenish-yellow spots within; style 3.2–3.5 mm long; fruiting calyx appendages strongly recurved; fruiting pedicels 18–25 mm long; Brazil (Minas Gerais, Paraná, Rio de Janeiro, Santa Catarina, and São Paulo) C. recurvatum
Flowering calyx appendages linear or subulate, densely pubescent with spreading trichomes, the longest appendages 2.5–5 (–6) mm long; corolla (8–) 9–14 mm long, 18–22 mm in diameter, white with purple or reddish-brown spots within; style 4–6.8 mm; fruiting calyx appendages spreading; fruiting pedicels (25–) 30–38 mm long; Brazil (São Paulo and Rio de Janeiro) C. cornutum

Key to the wild Capsicum species from North America, Central America and the Caribbean

1 Leaf pair subequal in size and shape; calyx appendages absent or five, minute, < 0.5 mm long; corolla (5–) 6–8 mm long, stellate, lobed nearly halfway or up to 2/3 of the way to the base, entirely white or almost pale yellow, rarely greenish-white; style cylindrical; flowering and fruiting pedicels erect; fruits pungent; seeds pale yellow to yellow; perennial herbs or prostrate subshurb; southern United States of America, Mexico, Central America and the Caribbean islands C. annuum var. glabriusculum
Leaf pairs markedly dissimilar in size and shape; calyx appendages (3–4) 5, > 0.5 mm long; corolla 5–14 mm long; campanulate or campanulate-rotate, lobed not more than 1/3 of the way to the base, yellow or purple marginally white; style clavate; flowering and fruiting pedicels pendent; fruits non-pungent; seeds brown or brownish-black to black; erect shrubs or shrubs, rarely trees 2
2 Plants glabrescent to densely pubescent with simple, furcate or dendritic trichomes; major leaves ovate, elliptic or rhomboid-ovate; inflorescences of 3–8 (–13) flowers, rarely flowers solitary; calyx appendages erect or spreading, 0.9–3 mm long; corolla (5–) 6–10 mm long, entirely yellow or with diffuse greenish spots within; Mexico, Guatemala, Honduras, El Salvador, Nicaragua and Costa Rica (also South America) C. rhomboideum
Plants glabrous or glabrescent only with simple eglandular trichomes; major leaves elliptic to lanceolate; inflorescence of a solitary flower, rarely two; calyx appendages spreading or strongly reflexed, (2–) 3–5 mm long; corolla 9.8–14 mm long, purple with white margin within; Mexico, Guatemala and Honduras C. lanceolatum

Key to the wild Andean and adjacent Andean (Venezuela, Colombia, Ecuador, Peru, Bolivia and Argentina) Capsicum species

1 Stem and mature leaf blades completely glabrous, if trichomes present sparsely distributed on the veins and margins 2
Stem and mature leaf blades variously pubescent 7
2 Calyx appendages (3–) 5, strongly incurved; flowering pedicels slightly winged and conspicuously winged in fruit; leaves coriaceous; Bolivia C. ceratocalyx
Calyx appendages absent or up to 10, straight; flowering and fruiting pedicels not winged; leaves coriaceous or membranous 3
3 Corolla tubular-campanulate to broadly campanulate, lobed less than 1/3 of the way to the base 4
Corolla stellate or stellate-campanulate, lobed between 1/3–2/3 of the way to the base 6
4 Calyx appendages 10, unequal (five long, five short); corolla lobes recurved; fruits pungent; seeds 3–4 (–5) mm long, pale yellow to nearly white; Bolivia C. caballeroi
Calyx appendages 2–5, equal or subequal; corolla lobes erect; fruits non-pungent; seeds 1.5–2.5 mm long, brown to black 5
5 Leaves coriaceous; calyx appendages (2–) 3–5, spreading or reflexed; filaments 1–2.5 mm long; corolla broadly campanulate; Colombia and Ecuador C. lycianthoides
Leaves membranous; calyx appendages five, erect; filaments 3–5 mm long; corolla tubular-campanulate; Peru C. piuranum
6 Leaves coriaceous; major leaves narrowly elliptic (length/width ratio 6–10.8); calyx tube membranaceous; stamens equal, 2–2.6 mm long; fruits 8–13 mm in diameter, orange at maturity; fruiting pedicels 10–16 mm long, green, pendent; fruiting calyx green-purple or green; seeds 1.7–2.3 mm long, 1.7–2.2 mm wide, seeds D or teardrop-shaped, the surface reticulate; Peru and Ecuador C. longifolium
Leaves membranaceous; major leaves elliptic (length/width ratio 2.5–4); calyx tube fleshy; stamens subequal (one longer), (2–) 3–4.3 mm long; fruits 6–9 mm in diameter, dark blue to purple at maturity; fruiting pedicels ca. 18 mm long, brilliant dark purple, erect; fruiting calyx entirely brilliant purple; seeds 2.75–3.40 mm long, 2.25–2.70 mm wide, C-shaped, the surface smooth and tuberculate at margins; Colombia, Ecuador, and Peru C. regale
7 Pubescence mostly of branched eglandular or long forked glandular trichomes, few simple trichomes 8
Pubescence mostly of simple eglandular trichomes, rarely furcate eglandular trichomes or simple long glandular trichomes 9
8 Dense glandular pubescence of long furcate and simple trichomes; calyx appendages usually 10 (rarely 5 or up to 12); corolla stellate, lobed nearly halfway to the base, white with greenish-yellow spots within (sometimes nearly white or with purple spots in the lobes); flowering and fruiting pedicels erect; fruits pungent; inflorescences usually 2–3 (–4)-flowered; Bolivia C. eshbaughii
Dense eglandular pubescence of simple, furcate and dendritic trichomes; calyx appendages usually five (rarely 3–4); corolla campanulate or campanulate-rotate, shallowly lobed, entirely yellow or sometimes tinged greenish within; flowering and fruiting pedicels pendent; fruits non-pungent; inflorescences usually 3–8 (–13)-flowered; Venezuela to Peru (also in Mexico and Central America) C. rhomboideum
9 Corolla nearly lobed up to the base, tube 4–4.5 times shorter than the lobes; Ecuador C. benoistii
Corolla shallowly lobed or lobed halfway or up to 2/3 of the way to the base, tube as long as the lobes or 1.5 times shorter than the lobes 10
10 Calyx appendages (3–) 5, strongly incurved; flowering pedicels nearly winged and conspicuously winged in fruit; leaves coriaceous; Bolivia C. ceratocalyx
Calyx appendages absent or up to 10, straight; flowering and fruiting pedicels not winged; leaves coriaceous or membranous 11
11 Young stems, lower surface of the leaves and calyx with small dark glandular trichomes mixed with simple eglandular trichomes; corolla primarily purple, violet or lilac; fruits pungent 12
Young stems, leaves and calyx only with simple eglandular trichomes, glandular trichomes absent; corolla entirely yellow or nearly white or primarily purple and usually with maroon, purple or greenish-yellow pigmentation within; fruits pungent or non-pungent 13
12 Calyx appendages absent or five, minute ≤ 0.5 mm long; leaves moderately to densely pubescent; flowering pedicels usually pendent, non-geniculate at anthesis, 3–10 mm long; style dimorphic; fruiting pedicels erect; Peru C. tovarii
Calyx appendages five, 1–2 mm long; leaves glabrescent; flowering pedicels usually erect, geniculate at anthesis, 8–18 (–22) mm long; style homomorphic; fruiting pedicels pendent; Bolivia C. cardenasii
13 Corolla tubular-campanulate or campanulate, lobed less than 1/3 of the way to the base 14
Corolla stellate or rotate-stellate, lobed between 1/3–2/3 of the way to the base 17
14 Calyx appendages 8–10, unequal 15
Calyx appendages 2–5, equal or subequal 16
15 Leaves coriaceous; inflorescences 2-flowered or flowers solitary; flowering pedicels 20–40 (–50) mm long; corolla ≥ 10 mm long, 4–6 mm in diameter; filaments 4–6 mm long; style 7–9 mm long; fruits pungent; fruiting calyx appendages appressed to the berry; seeds pale yellow to nearly white; Bolivia C. caballeroi
Leaves membranous; inflorescences with 2–7 flowers, rarely flowers solitary; flowering pedicels shorter, 8–15 mm long; corolla 7.5–9 mm long, 8–10 mm in diameter, filaments (1.5–) 1.8–2 mm long; style ca. 4 mm long; fruits non-pungent; fruiting calyx appendages spreading or reflexed; seeds yellow or brown; Ecuador and Peru C. hookerianum
16 Calyx appendages five, erect; corolla tube narrow, ca. 6 mm in diameter; filaments 3–5 mm long; mature leaves glabrescent adaxially; stone cells two; northern Peru C. piuranum
Calyx appendages 2–3 (–5), erect or spreading; corolla tube broad, 8–10 mm in diameter; filaments 2–3 mm; mature leaves sparse to densely pubescent adaxially; stone cells 1–5 or absent; Colombia, Ecuador and Peru C. geminifolium
17 Flowers 4–13 (–18) on a short rachis; pedicel scars prominent, corky; fruiting calyx usually recurved; sprawling vines or scrambling shrubs; Peru, Bolivia (also Brazil) C. coccineum
Flowers 2–6 (–8) per axil, rarely flowers solitary; pedicel scars usually inconspicuous, rarely prominent; fruiting calyx not recurved; usually scandent or erect shrubs or subshrubs, rarely perennial herbs 18
18 Calyx appendages absent or minute, ≤ 1 mm long 19
Calyx appendages (4–) 5–10, > 1 mm long 21
19 Leaf pair strongly dissimilar in shape and size; flower buds ovoid, purple or yellowish; fruits non-pungent; seeds 1.9–2.7 mm long, 1.8–2.1 mm wide, brownish-black to black; corolla entirely yellow or with purple or maroon spots within; Colombia, Ecuador and Peru C. dimorphum
Leaf pair similar or dissimilar in size, similar in shape; flower buds globose or ovoid, white cream or greenish-white; fruits pungent; seeds 3.2–4 mm long, 2.5–4 mm wide, pale yellow to yellow; corolla entirely white or pale yellow, rarely greenish-white 20
20 Calyx circular in outline; corolla 4–5 mm long, ca. 6 mm in diameter, with glandular trichomes adaxially; style 2.25–2.5 mm long, exserted 0.5–0.8 mm beyond the anthers; seed coat smooth; plants densely pubescent, the trichomes spreading; endemic; Ecuador, Galapagos Islands C. galapagoense
Calyx pentagonal in outline; corolla (5–) 6–8 mm long, 8–10 (–12) mm in diameter, glabrous adaxially; style 4–4.8 mm long, exserted 1.5–2 mm beyond the anthers; seed coat reticulate to obscurely reticulate; plants glabrescent to densely pubescent, the trichomes appressed-antrorse, sometimes spreading; widespread; Colombia, Venezuela, Ecuador, Peru, Bolivia (also in North and Central America, the Caribbean Islands and Brazil C. annuum var. glabriusculum
21 Flowers solitary; corolla entirely white; filaments with conspicuous auricles not fused to the corolla at the point of insertion; Bolivia and Argentina (also Paraguay) C. chacoense
Flowers 2–8 per axil, rarely solitary; corolla white, purple or yellow with greenish-yellow spots or greenish-yellow centre within; filaments with inconspicuous auricles fused to the corolla at the point of the insertion 22
22 Flowering pedicels pendent, non-geniculate at anthesis; calyx appendages 5–10 23
Flowering pedicels erect, geniculate at anthesis; calyx appendages (4–) 5 24
23 Calyx appendages always 10; calyx tube strongly 10-nerved with prominent venation; pedicels scars inconspicuous; corolla entirely yellow or with small greenish-yellow spots within; fruits red; seeds 4–5 mm long, 3–4.25 mm wide, pale yellow to white; Bolivia C. neei
Calyx appendages 5 (–7); calyx tube strongly 5-nerved with prominent venation; pedicels scars prominent; corolla purple with a narrow white margin and yellowish-green centre; fruits greenish-golden yellow; seeds 3–3.8 mm long, 2.7–3 mm wide, brownish-black; Colombia and Venezuela (also Brazil) C. parvifolium
24 Corolla rotate-stellate; white with greenish-yellow spots within; seeds pale yellow to yellow; fruits globose, subglobose or ellipsoid; Colombia to Argentina (also Paraguay and Brazil) C. baccatum var. baccatum
Corolla stellate; primarily yellow or lilac, purple or magenta; seeds brownish-yellow or dark brown; fruits globose 25
25 Calyx appendages five, 1–1.5 mm long; filaments 2–2.5 mm long; corolla yellow with small and faint greenish-yellow spots within; seeds 4–4.5 mm long, 3–3.5 mm wide, dark brown; Bolivia C. minutiflorum
Calyx appendages (4–) 5, 1.2–2.7 (–3) mm long; filaments 2.7–3.8 mm long; corolla lilac, purple or magenta with a continuous greenish-yellow or ochre tube within, sometimes the corolla white with greenish-yellow spots; seeds 2.5–4 (–4.2) mm long, 2.1–3 mm wide, brownish-yellow; Bolivia and Argentina C. eximium

Key to the wild Capsicum species occurring in Brazil, Paraguay and Argentina

1 Stem and mature leaf blades completely glabrous, if trichomes present, sparsely distributed on the veins and margins 2
Stem and mature leaf blades variously pubescent 4
2 Calyx appendages five (6–10), unequal, 1–5 mm long; corolla 10–14 (–16) mm long; Brazil (São Paulo) C. hunzikerianum
Calyx appendages absent or five, minute, < 0.5 mm long; corolla 4.5–10 mm long 3
3 Leaves membranous, elliptic to ovate; flowering pedicels 9–14 mm long, erect, geniculate at anthesis; corolla small, 4.5–6.5 (–8) mm long; stamens unequal (3+2); ovules 2 per locule; fruits 4-seeded; Brazil (Rio de Janeiro, Minas Gerais, Espírito Santo) C. campylopodium
Leaves coriaceous, elliptic to narrowly elliptic, flowering pedicels 15–30 mm long, pendent, non-geniculate at anthesis; corolla larger, 9–10 mm long; stamens equal; ovules more than 2 per locule; fruits multi-seeded (up to 20 seeds); Brazil (Bahía, Espírito Santo, Minas Gerais, São Paulo) C. pereirae
4 Pubescence of furcate to dendritic eglandular trichomes mixed with few simple eglandular trichomes; fruits non-pungent; calyx appendages long, (4.5–) 5–8.5 mm long; Brazil (Bahía, Minas Gerais, Pernambuco) C. longidentatum
Pubescence of simple eglandular trichomes, rarely furcate trichomes; fruits usually pungent; calyx appendages absent or, if present, up to 6 mm long 5
5 Corolla campanulate-urceolate, rotate or rotate-stellate, lobed less than 1/3 of the way to the base 6
Corolla stellate, lobed more than 1/3 up to 2/3 of the way to the base 8
6 Corolla campanulate-urceolate, entirely fuchsia or violet, glabrous adaxially, the lobes spreading to strongly recurved; fruiting pedicels pendent; fruits greenish-golden yellow, slightly pungent; seeds brownish-black to black; Brazil (Rio de Janeiro) C. friburgense
Corolla rotate or rotate-stellate, white or primarily purple or lilac with greenish-yellow pigmentation within, with glandular trichomes adaxially; the lobes spreading; fruiting pedicels erect; fruits usually red, pungent; seeds pale yellow to yellow 7
7 Leaves with dense pubescence, especially underneath; corolla marginally purple or lilac with greenish-yellow centre; Brazil and Paraguay C. rabenii
Leaves mostly glabrescent, more rarely moderately pubescent; corolla white with greenish-yellow spots within, purple pigmentation absent; widely distributed across South America C. baccatum var. baccatum
8 Calyx appendages absent or up to five, minute, < 0.5 mm long 9
Calyx appendages 2–10, > 0.5 mm long 16
9 Flowering pedicels pendent, non-geniculate at anthesis 10
Flowering pedicels erect, geniculate at anthesis 11
10 Young stems, leaves and calyx with simple eglandular trichomes mixed with small dark glandular trichomes; inflorescences multi-flowered (5–13 flowers or up to 20 or more); corolla purple with a white margin within; fruiting pedicels green or purple, with a constriction at the junction with the calyx; seeds pale yellow; Brazil C. caatingae
Young stems, leaves and calyx only with simple eglandular trichomes, glandular trichomes absent; inflorescences few-flowered (2–6 flowers), rarely solitary flowers; corolla white with greenish-yellow spots, rarely also with purple spots; fruiting pedicels green, without a constriction at the junction with the calyx; seeds brownish-black; Argentina, Paraguay and Brazil C. flexuosum
11 Flowers 4–13 (–18) on a short rachis; pedicels scars prominent, corky; fruiting calyx usually recurved; seeds 4–4.6 mm long, (–2.8) 3.2–3.75 mm wide, yellow to brownish-yellow; sprawling vines or scrambling shrubs, with stems to 7 m long; Brazil (also Peru and Bolivia) C. coccineum
Flowers 2–5 (–7) per axil, rarely solitary; pedicels scars inconspicuous; fruiting calyx not recurved; seeds 2–4 mm long, 2.5–3.5 mm wide, pale yellow or brownish-black to black; erect shrubs or subshrubs up to 2.5 m tall, rarely low perennial herbs or small trees 12
12 Stamens unequal (3+2); ovules two per locule; fruits 4-seeded; Brazil (Rio de Janeiro, Minas Gerais, Espírito Santo) C. campylopodium
Stamens equal; ovules more than two per locule; fruits multi-seeded (up to 20 seeds) 13
13 Filaments short, < 2 mm long; fruiting pedicels erect; seeds pale yellow to yellow; fruits yellow, red-orange or red 14
Filaments longer, 2.4–4 mm long; fruiting pedicels pendent; seeds brownish-black to black; fruits greenish-golden yellow 15
14 Fruiting calyx without a prominent annular constriction at junction with the pedicel C. annuum var. glabriusculum
Fruiting calyx with a prominent annular constriction at junction with the pedicel C. chinense
15 Stems, leaves, pedicels and calyx densely pubescent with long spreading eglandular trichomes (0.5–2 mm long); major leaves elliptic to narrowly elliptic; corolla white with large greenish-yellow spots and sparse diffuse purple or brown spots within, the lobes widely triangular; Brazil (Rio de Janeiro) C. muticum
Stems, leaves pedicels, and calyx glabrescent to moderately pubescent, with short antrorse eglandular trichomes (0.25–0.5 mm long); major leaves elliptic to ovate; corolla white with large or small purple or brownish spots and large greenish-yellow spots within (in some populations, purple or brownish pigmentation absent entirely); the lobes triangular or ovate; Brazil (Minas Gerais, Rio de Janeiro and São Paulo) C. schottianum
16 Flowers solitary; corolla entirely white; staminal plaques with auricles not fused to the corolla at the point of insertion; Argentina and Paraguay (also Bolivia) C. chacoense
Flowers 2–18, rarely flowers solitary; corolla yellow, purple or white tinged of different colours within; staminal plaques with auricles fused to the corolla at the point of insertion 17
17 Corolla usually yellow with yellowish-green or purple-brown spots within; fruiting calyx usually recurved; seeds 4–4.6 mm long, (–2.8) 3.2–3.75 mm wide, yellow to brownish yellow; sprawling vines or scrambling shrubs, with stems to 7 m long; western Brazil (also Peru and Bolivia) C. coccineum
Corolla primarily purple or white; fruiting calyx not recurved; seeds (2–) 2.2–3.5 (–4) mm long, (–1.8) 2–3 (–3.5) mm wide, yellow or brownish-black to black; erect shrubs or subshrubs, up to 5 m tall 18
18 Calyx appendages 10 or 6–10, rarely five 19
Calyx appendages five (very rarely up to seven) 21
19 Calyx appendages 10, subequal; filaments 3–3.2 mm long; style barely exserted beyond the anthers; anthers lilac or pale blue; corolla almost entirely purple; Brazil (São Paulo) C. mirum
Calyx appendages ranging from 6 to 10, rarely five, unequal; filaments 1.4–2.5 mm long; style exserted 0.8–1.3 mm beyond the anthers; anthers yellow, light green or grey; corolla white with greenish-yellow or purple spots within 20
20 Flowering calyx appendages cylindrical or triangular-compressed, glabrous to moderately pubescent with antrorse trichomes, the longest appendages 1–2.5 mm; corolla 6–7 mm long, ca. 11 mm in diameter, white with greenish-yellow spots within; fruiting calyx appendages strongly recurved; fruiting pedicels 18–25 mm long; Brazil (Minas Gerais, Paraná, Rio de Janeiro, Santa Catarina and São Paulo) C. recurvatum
Flowering calyx appendages linear or subulate, densely pubescent with spreading trichomes, the longest appendages 2.5–5 (–6) mm long; corolla (8–) 9–14 mm long, 18–22 mm in diameter, white with purple or reddish-brown spots within; fruiting calyx appendages spreading; fruiting pedicels (25–) 30–38 mm long; Brazil (São Paulo and Rio de Janeiro) C. cornutum
21 Major leaves ovate; corolla primarily purple with greenish-yellow or cream centre within 22
Major leaves elliptic or narrowly elliptic, more rarely ovate 23
22 Pedicels scars prominent; flowering and fruiting pedicels pendent; fruits greenish-golden yellow; seeds brownish-black, the seed coat reticulate and tuberculate at margins; Brazil (also Colombia and Venezuela) C. parvifolium
Pedicels scars inconspicuous; flowering and fruiting pedicels erect; fruits orange or red; seeds pale yellow or yellow, the seed coat smooth; Paraguay and Brazil C. rabenii
23 Plants densely pubescent on stems, petioles, pedicels and sometimes also on the leaf nerves beneath, the trichomes spreading; Brazil (Rio de Janeiro, Minas Gerais, São Paulo and Espírito Santo) C. villosum
Plants glabrescent to densely pubescent on stems, leaves and pedicels, the trichomes appressed-antrorse; calyx appendages up to 5 mm long 24
24 Plants glabrous to sparsely pubescent; major leaves elliptic to ovate, rarely narrowly elliptic (length/width ratio: (2–) 2.5–4 (–4.9), apex acuminate to long acuminate; calyx appendages (0.4–) 0.5–1.5 (–3) mm; flower buds purple; corolla (6–) 7.5–12 mm long, (9–) 10–13 mm in diameter; Brazil (Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro and São Paulo) C. mirabile
Plants moderately to densely pubescent; major leaves narrowly elliptic to lanceolate (length/width ratio: (4–) 5–10 (–16), apex acute to obtuse; calyx appendages (2.8–) 3–4 (–5) mm; flower buds cream with greenish and purple spots; corolla (8–) 10–12 mm long, 13–20 mm in diameter; Brazil (Minas Gerais) C. carassense

Key to the domesticated Capsicum species

1 Calyx appendages absent or minute, ≤ 0.5 mm long 2
Calyx appendages > 0.5 mm long 4
2 Flowers solitary, rarely in pairs or more; petioles up to 10 cm long; corolla 8–15 mm long, entirely white, rarely entirely purple or pale yellow; fruits usually large, up to 300 mm long, pungent or non-pungent C. annuum var. annuum
Flowers 2–5, rarely solitary; petioles up to 3.5 cm long; corolla 3.75–8 mm long, entirely dull white or greenish-white (occasionally with purple spots); fruits small to medium-sized, up to 100 mm long, pungent, rarely non-pungent 3
3 Corolla glabrous adaxially; style heteromorphic; fruits subglobose to highly variable in shape, with the base obtuse or truncate; fruiting calyx discoid or shallowly cup-shaped, with a prominent annular constriction at junction with the pedicel; ovary subglobose, 2–2.5 mm long, 2.5–3.5 mm in diameter C. chinense
Corolla with small glandular trichomes adaxially; style homomorphic; fruits usually elongate, narrowly triangular, with the base abruptly narrowed; fruiting calyx deeply cup-shaped lacking a constriction at junction with the pedicel; ovary oblong-ovoid, 2.5–4 mm long, 1.3–1.8 mm in diameter C. frutescens
4 Leaves densely pubescent, rarely glabrescent, the youngest leaves rugose; flower buds dark purple on pendent pedicels; corolla dark purple or violet with a white or yellowish-green centre within; style clavate; seeds 5.5–7 mm long, 4.8–6 mm wide, brownish-black to black, the seed coat reticulate C. pubescens
Leaves glabrous to sparsely pubescent; the youngest leaves even; flower buds greenish-white (rarely purple) on geniculate pedicels; corolla white with large greenish-yellow spots and white centre within, style cylindrical; seeds 3–5.2 mm long, 3–4 mm wide, pale yellow to yellow, the seed coat smooth to slightly reticulate 5
5 Fruits pungent, rarely non-pungent, usually elongate, endocarp alveolate, pericarp with giant cells C. baccatum var. pendulum
Fruits non-pungent, campanulate-umbilicate, endocarp smooth, pericarp without giant cells C. baccatum var. umbilicatum

Species descriptions

Capsicum annuum L., Sp. Pl. 1: 188. 1753.

Type

“Habitat in America meridionali” Herb. Clifford: 59, Capsicum 1 (lectotype, designated by D’Arcy 1974 [’1973’], pg. 591: BM [BM000558022]).

Capsicum annuum L. var. annuum

Figs 20, 21

Capsicum grossum L., Mant. Pl.: 47. 1767. Type. “Habitat in India … H.U.” HU [Horto Upsaliensis]: Fructu vario crasso. Caulis biennis, Herb. Linn. N° 249.5 (lectotype, designated here: LINN [LINN-HL249-5]).

Capsicum cordiforme Mill., Gard. Dict. ed. 8, no. 2. 1768. Type. Cultivated at the Chelsea Physic Garden (no specimens cited; no original material located).

Capsicum tetragonum Mill., Gard. Dict. ed. 8, no. 3. 1768. Type. Cultivated at the Chelsea Physic Garden (no specimens cited; no original material located).

Capsicum angulosum Mill., Gard. Dict. ed. 8, no. 4. 1768. Type. Cultivated at the Chelsea Physic Garden (no specimens cited; no original material located).

Capsicum olivaeforme Mill., Gard. Dict. ed. 8, no. 6. 1768. Type. Cultivated at the Chelsea Physic Garden, seeds from “Barbadoes” (no specimens cited; no original material located).

Capsicum pyramidale Mill., Gard. Dict. ed. 8, no. 7. 1768. Type. Cultivated at the Chelsea Physic Garden, seeds from Egypt (no specimens cited; no original material located).

Capsicum conicum Lam., Tabl. Encycl. 2: 26. 1794. Type. “Ex Indiis” Herb. Lamarck s.n. (lectotype, designated here: P-LAM [P00357734]).

Capsicum bicolor Jacq., Fragm. Bot. 66, tab 99, fig. 1. 1809. Type. “Patriam ignoro” (no specimens cited; lectotype, designated here: Jacquin, Fragm. Bot.: 66, tab 99, fig. 1. 1809).

Capsicum grossum Willd., Enum. Pl. [Willdenow] 1: 241. 1809, nom. illeg., not Capsicum grossum L. (1767). Type. “Habitat in India orientaliCapsicum grossum [sheet] 2, Herb. Willdenow (lectotype, designated here: B [B-W04425-02-0]).

Capsicum sphaericum Willd., Enum. Pl. [Willdenow] 1: 241. 1809. Type. “Habitat…. ” (lectotype, designated here: B [B-W04426-01-0, F neg. 2886]).

Capsicum nigrum Willd., Enum. Pl. [Willdenow] 1: 242. 1809, nom. illeg. superfl. Type. Based on Capsicum bicolor Jacq. (cited in synonymy).

Capsicum purpureum Vahl ex Hornem., Hort. Bot. Hafn. 1: 224. 1813. Type. [Denmark]. Hort. Haf., 1802, Herb. Vahl s.n. (lectotype, designated here: C [C10019148]).

Capsicum ovatum DC., Cat. Pl. Horti Monsp.: 86. 1813. Type. “Habitat….” (no specimens cited; no original material located; Capsicum ovatum, Anonymous s.n. (neotype, designated here: G-DC [G00200072]).

Capsicum longum DC., Cat. Pl. Horti Monsp.: 86. 1813. Type. “Hab... in hortis frequens” (no specimens cited; lectotype, designated here [illustration]: “Piper Calecuticum sive Capsicum oblongius, Bauhin et al., Hist. pl. 2: 943, f. I. 1651).

Capsicum globiferum G.Mey., Prim. Fl. Esseq.: 113. 1818. Type. “In plantationibus”, no specimens cited; [Guyana]. Río Essequibo, E.K. Rodschied 29 (lectotype, designated here: GOET [GOET003420]).

Capsicum purpureum Roxb., Fl. Ind., ed. Carey & Wall. 2: 259. 1824, nom. illeg., not Capsicum purpureum Vahl ex Hornem. (1813). Type. “Most likely from the Molucca Islands” (no specimens cited; neotype, designated here: “C. purpureum, H.B.C.” [Horto Botanici Calcutta]: K [K001132446]).

Capsicum indicum Dierb. var. vulgatum Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 22. 1829, nom. illeg. superfl. Type. Based on Capsicum annuum L. (cited in synonymy).

Capsicum indicum Dierb. var. longum (DC.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 23. 1829. Type. Based on Capsicum longum DC.

Capsicum indicum Dierb. var. tetragonum (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 23. 1829. Type. Based on Capsicum tetragonum Mill.

Capsicum indicum Dierb. var. angulosum (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 25. 1829. Type. Based on Capsicum angulosum Mill.

Capsicum indicum Dierb. var. cordiforme (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 25. 1829. Type. Based on Capsicum cordiforme Mill.

Capsicum indicum Dierb. var. grossum (L.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 26. 1829. Type. Based on Capsicum grossum L.

Capsicum indicum Dierb. var. sphaericum (Willd.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 27. 1829. Type. Based on Capsicum sphaericum Willd.

Capsicum indicum Dierb. var. ovatum (DC.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 27. 1829. Type. Based on Capsicum ovatum DC.

Capsicum indicum Dierb. var. pyramidale (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 28. 1829. Type. Based on Capsicum pyramidale Mill.

Capsicum indicum Dierb. var. olivaeforme (Mill.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 28. 1829. Type. Based on Capsicum olivaeforme Mill.

Capsicum indicum Dierb. var. nigrum (Willd.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 29. 1829. Type. Based on Capsicum nigrum Willd.

Capsicum axi Vell., Fl. Flumin.: 61. 1829 (“1825”); Fl. Flumin. Icon. 2: t. 6. 1831 (“1827”). Type. Brazil. [Rio de Janeiro]: “Colitur hortis” (lectotype, designated by Knapp et al. 2015, pg. 824: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_009] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 6. 1831).

Capsicum silvestre Vell., Fl. Flumin. 60. 1829 (“1825”); Fl. Flumin. Icon. 2: t. 1. 1831 (“1827”). Type. Brazil. [Rio de Janeiro]: “Ad declivium Alpium Fluminensium” (lectotype, designated by Knapp et al. 2015, pg. 824: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_004] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 1. 1831).

Capsicum annuum L. var. rugosulum Fingerh., Monogr. Capsic.: 13. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II b. 1832).

Capsicum annuum L. var. acuminatum Fingerh., Monogr. Capsic.: 13. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II c. 1832).

Capsicum annuum L. var. subangulosum Fingerh., Monogr. Capsic. 13. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II d. 1832).

Capsicum annuum L. var. ovoideum Fingerh., Monogr. Capsic.: 14. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II e. 1832).

Capsicum annuum L. var. abbreviatum Fingerh., Monogr. Capsic.: 14. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II f. 1832).

Capsicum annuum L. var. olivaeforme Fingerh., Monogr. Capsic.: 14. 1832. Type. “Crecit in America meridionali et India oriental” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. II g. 1832).

Capsicum bicolor Jacq. var. purpureum (Vahl ex Hornem.) Fingerh., Monogr. Capsic.: 16. 1832. Type. Based on Capsicum purpureum Vahl ex Hornem.

Capsicum strictum Fingerh., Monogr. Capsic.: 21. 1832. Type. “Patria…..” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. V a. 1832).

Capsicum grossum Willd. var. pomiforme Fingerh., Monogr. Capsic.: 22. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. V c. 1832).

Capsicum grossum Willd. var. ovatum Fingerh., Monogr. Capsic.: 22. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. V d. 1832).

Capsicum grossum Willd. var. cordatum Fingerh., Monogr. Capsic.: 22. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VI a. 1832).

Capsicum grossum Willd. var. angulosum Fingerh., Monogr. Capsic.: 22. 1832. Type: “Patria India orientalis (Herb. Wight et Herb. Hamilt.)” (no specimens found; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VI d. 1832).

Capsicum ceratocarpum Fingerh., Monogr. Capsic.: 22. 1832. Type. “Patria….” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VI c. 1832).

Capsicum longum DC. var. incrassatum Fingerh., Monogr. Capsic.: 24. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VII a. 1832).

Capsicum longum DC. var. latum Fingerh., Monogr. Capsic.: 25. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VII b (as - - [Capsicum longum] luteum). 1832).

Capsicum longum DC. var. rectum Fingerh., Monogr. Capsic.: 25. 1832. Type. “Cresit in Indiis et America meridionali” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VII c. 1832).

Capsicum pendulum Willd. var. torulosum Fingerh., Monogr. Capsic.: 26. 1832. Type. [Indonesia] “in Amboina” (no specimens cited; lectotype, designated here [illustration]: Capsicum rubrum minus Rumphius, Herbarium Amboinense 5, Tab. LXXXVIII, fig. 1, 1747, cited in synonymy).

Capsicum angulosum Mill. var. conicum Fingerh., Monogr. Capsic.: 28. 1832. Type. No locality cited (no specimens cited, no original material located).

Capsicum angulosum Mill. var. ovale Fingerh., Monogr. Capsic.: 28. 1832. Type. “Patria….?” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VIII b. 1832).

Capsicum hamiltonii G.Don, Gen. Hist. 4: 447. 1838. Type. [Caribbean Islands] “Native of the Island of Nevis, in gardens” (no specimens cited, no original material located).

Capsicum annuum L. var. longum (DC.) Sendtn., Fl. Bras. (Martius) 10(6): 144. 1846. Type. Based on Capsicum longum DC.

Capsicum annuum L. var. grossum (Willd.) Sendtn., Fl. Bras. (Martius) 10(6): 147. 1846. Type. Based on Capsicum grossum Willd.

Capsicum annuum L. var. cordiforme (Mill.) Sendtn., Fl. Bras. (Martius) 10(6): 148. 1846. Type. Based on Capsicum cordiforme Mill.

Capsicum abyssinicum A.Rich., Tent. Fl. Abyss 2: 96. 1850. Type. [Ethiopia] “Abyssinia, Ouedjerate”, R. Quartin Dillon s.n. (lectotype, designated here: P [P00329903]; isolectotypes: P [P00329904, P00329905]).

Capsicum annuum L. var. oblongum Dunal, Prodr. [A. P. de Candolle] 13(1): 412. 1852. Type. “Capsicum annuum α oblongum fructibus rubris”, 1844, Herb. Dunal (lectotype, designated here: G-DC [G00131768]).

Capsicum pyramidale Mill. var. longicorne Dunal, Prodr. [A. P. de Candolle] 13(1): 414. 1852. Type. [Indonesia] Java, 1843, H. Zollinger 489 (lectotype, designated here: G-DC [G00131841]; isolectotypes: G [G00390281], LE).

Capsicum bicolor Jacq. var. purpureum (Vahl ex Hornem.) Dunal, Prodr. [A. P. de Candolle] 13(1): 414. 1852. Type. Based on Capsicum purpureum Vahl ex Hornem.

Capsicum testiculatum Vis. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 424. 1852. Type. In Hort. Montpellier [seeds sent by R. de Visiani], 1837, Anonymous s.n. (lectotype, designated here: G-DC [G00200067]; isolectotype: MPU [MPU023039]).

Capsicum angulosum Mill. var. macrocarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 426. 1852. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. VIII a (as Capsicum angulosum M.). 1832).

Capsicum leucocarpon Dunal, Prodr. [A. P. de Candolle] 13(1): 429. 1852. Type. Cultivated in England “Capsicum americanum latifolium, fructu oblongo erecto candido” (Miller 1752) (no specimens cited, no original material located).

Capsicum dulce Dunal, Prodr. [A. P. de Candolle] 13(1): 428. 1852. Type. Cultivated in Montpellier, France, “In hortis botanicis cultum” (no specimens cited; no original material located).

Capsicum annuum L. var. cordiforme (Mill.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum cordiforme Mill.

Capsicum annuum L. var. angulosum (Mill.) Alef., Landw. Fl.: 132. 1866, as ‘angulatum’. Type. Based on Capsicum angulosum Mill.

Capsicum annuum L. var. pyramidale (Mill.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum pyramidale Mill.

Capsicum annuum L. var. globiferum (G.Mey.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum globiferum G.Mey.

Capsicum annuum L. var. longum (DC.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum longum DC.

Capsicum annuum L. var. tetragonum (Mill.) Alef., Landw. Fl.: 133. 1866. Type. Based on Capsicum tetragonum Mill.

Capsicum annuum L. var. tetragonum (Mill.) Alef., Landw. Fl.: 133. 1866. Type. Based on Capsicum tetragonum Mill.

Capsicum annuum L. var. purpureum (Roxb.) Alef., Landw. Fl.: 134. 1866. Type. Based on Capsicum purpureum Roxb.

Capsicum annuum L. var. ceratocarpum (Fingerh.) Alef., Landw. Fl.: 134. 1866. Type. Capsicum ceratocarpum Fingerh.

Capsicum annuum L. var. bicolor (Jacq.) Alef., Landw. Fl.: 134. 1866. Type. Capsicum bicolor Jacq.

Capsicum fasciculatum Sturtev., Bull. Torrey Bot. Club 15(5): 133. 1888. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: “Tenjikumamori, Capsicum annuum L. (Solaneae)”, Tanaka & Motoyoshi, Sô-Mokou-Zoussets, vol. 3, Tab. 38. 1874).

Capsicum annuum L. var. longum (DC.) Kuntze, Revis. Gen. Pl. 2: 449. 1891. Type. Based on Capsicum longum DC.

Capsicum annuum L. var. erectum Kuntze, Revis. Gen. Pl. 2: 449. 1891. Type. “Java, cult.” (no specimens cited, no original material located).

Capsicum annuum L. var. grossum (L.) Kuntze, Revis. Gen. Pl. 2: 449. 1891. Type. Based on Capsicum grossum L.

Capsicum annuum L. var. fasciculatum (Sturtev.) Irish, Rep. (Annual) Missouri Bot. Gard. 9: 68, pl. 9, f. 4. 1898. Type. Based on Capsicum fasciculatum Sturtev.

Capsicum frutescens L. var. lanicaule Greenm., Proc. Amer. Acad. Arts 39: 88. 1903. Type. Mexico. Jalisco: along Ave. Vallarta in Ciudad Granja, on the western outskirts of Guadalajara, 31 Dec 1886, E. Palmer 639 (lectotype, designated here: US [00816554, acc. # 92534], isolectotype: BM [BM000775827]).

Capsicum velutinum De Wild., Pl. Bequaert. 1: 413. 1922. Type. [Democratic Republic of the Congo]. Basankusu, Mar 1913, O. Lamboray 22 (lectotype, designated here: BR [BR000000649909]).

Capsicum frutescens L. var. fasciculatum (Sturtev.) L.H.Bailey, Gentes Herbarum 1: 129. 1923. Type. Based on Capsicum fasciculatum Sturtev.

Capsicum frutescens L. var. grossum (Willd.) L.H.Bailey, Gentes Herbarum 1: 129. 1923. Type. Based on Capsicum grossum Willd.

Capsicum annuum L. forma erectum Makino, J. Jap. Bot. 3(8): 29. 1926, as “Capsicum annuum var. fasciculatum f. erectum”. Type. “Hab. JAPAN, cultivated” (no specimens cited, no original material located).

Capsicum annuum L. forma pendulum Makino, J. Jap. Bot. 3(8): 29. 1926, as “Capsicum annuum var. fasciculatum f. pendulum”. Type. “Hab. JAPAN, cultivated, rare” (no specimens cited, no original material located).

Capsicum petenense Standl., Publ. Carnegie Inst. Wash. 461(4): 84. 1935. Type. Guatemala. Distr. Peten, La Libertad, Jun 1933, C. L. Lundell 3754 (holotype: F [v0072800F, acc. # 685329]; isotypes: CORD [CORD00101764 fragment ex MICH], MICH [1109873]).

Capsicum sonitpurense J.Sarma & G.Dutta, Bangladesh J. Pl. Taxon. 24(2): 215. 2017. Type. India. Assam, Sonitpur, Tezpur, 49 m, 22 Oct 2016, J. Sarma & G. Dutta 394 (holotype: ASSAM [acc. # 95893, sheet 394A]; isotypes: TUH [Tezpur University Herbarium, 3 sheets 394 B, C, D]).

Description

Annual herbs or short-lived, compact, low subshrubs, 1–1.5 m tall, the main stem 0.5–1 cm in diameter at base, branched from near the base. Young stems 3–4-angled, fragile, green to brownish-green, sometimes with purple lines, glabrous, glabrescent to moderately pubescent, rarely densely pubescent, with appressed-antrorse, simple, uniseriate, (5–) 8–13)-celled, eglandular trichomes 0.5–1 (–2) mm long; nodes green or with purple spots; bark of older stems light brown or brown, glabrous to sparsely pubescent; lenticels absent or few. Sympodial units difoliate, the leaves geminate; leaf pair similar in size and shape. Leaves membranous, concolorous, pale to dark green, glabrous to moderately pubescent on both sides, especially on the main veins abaxially, the trichomes similar to those of the stems; blades of all leaves 3–7 (–15.5) cm long, 2.5–5 (–8) cm wide, ovate to elliptic, the major veins (3–) 5–8 on each side of mid-vein, the base truncate to cordate or cuneate to attenuate, the margins entire, the apex acuminate or long-acuminate; petioles (0.5–) 4–7 (–10) cm, with the same pubescence as the stems. Inflorescences axillary, 1 (– 2) flowers per axil, rarely more; flowering pedicels (6–) 10–40 mm long, angled, erect and geniculate at anthesis or pendent and non-geniculate, green or purple, glabrous to moderately pubescent, the eglandular trichomes usually short, antrorse; pedicels scars inconspicuous. Buds globose, white or purple. Flowers 5–7-merous. Calyx 1–4 mm long, 3–5 mm wide, cup-shaped, green, strongly 5–10-nerved, glabrous to moderately pubescent with similar short or long eglandular trichomes as the stems, the calyx appendages usually 5 (–7), minute, 0.3–0.5 mm long. Corolla 8–15 mm long, (8–) 10–22 mm in diameter, entirely white, rarely entirely pale yellow or purple, stellate with narrow interpetalar membrane, lobed ca. halfway or 2/3 of the way to the base, glabrous adaxially and abaxially, the tube 3–8 mm long, the lobes 5–7 mm long, 3.5–5.5 mm wide, ovate, spreading, the margins finely ciliate, the tips acute, papillate. Stamens 5–7, equal; filaments 1–3 mm long, white or cream, sometimes purple, inserted on the corolla 1–1.5 mm from the base, with auricles fused to the corolla tube at the point of insertion; anthers 2–3 mm, ellipsoid or ovoid, pale blue to purplish, very rarely yellow, connivent or not connivent at anthesis. Gynoecium with ovary 1.5–3 mm long, 1.2–2.5 mm in diameter, ovoid or globose, green; nectary ca. 0.5 mm tall, pale green; style heteromorphic, short style 2.2–2.5 mm, not exceeding the anthers, medium style nearly the same height as the anthers, long style 3–5.1 mm, exserted 1.3–2.3 mm beyond the anthers, cylindrical, white or purple; stigma 0.1–0.2 mm long, ca. 0.4 mm wide, discoid or capitate, pale green or yellow. Berry highly variable in shape, size and colour, usually blocky or elongate, less commonly globose, up to 300 mm long, 6–65 mm in diameter, green, yellow or purple when immature, yellow, red, brown, purple or purple-black at maturity, persistent, pungent or non-pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 25–50 (–70) mm, erect or pendent, rigid, angled, uniformly widened, green; fruiting calyx 15–25 mm in diameter, slightly accrescent, discoid or rather cup-shaped, green. Seeds more than 50 per fruit, 3.8–4.4 mm long, 3.2–3.6 mm wide, C-shaped, pale yellow, the seed coat smooth to slightly reticulate (SM), cerebelloid (SEM), the cells irregular in shape, the lateral walls sinuate; embryo imbricate.

Figure 20. 

Capsicum annuum var. annum. Lectotype (BM). Copyright The Trustees of the Natural History Museum, London. Reproduced with permission.

Figure 21. 

Capsicum annuum var. annuum A flower bud on pendent pedicel B flower with connivent anthers C flower with hexamerous corolla (note nectar droplets on the limb) and style near the same length as the anthers D flower with heptamerous purple corolla E flower with pentamerous corolla and style exceeding the anthers F, H mature fruits on pendent pedicels G mature fruits on upright pedicels A–H no specimen vouchers, photos by G.E. Barboza and C. Carrizo García taken at different greenhouses.

Distribution

Capsicum annuum var. annuum is the most extensively cultivated pepper worldwide.

Ecology

Capsicum annuum var. annuum is found in diverse habitats throughout its wide distribution and is well adapted to the highlands environments (0–2,600 m elevation).

Phenology

Flowering and fruiting all year.

Chromosome number

2n = 2x = 24 (Pickersgill 1971, 1977, 1991; Moscone et al. 2007).

Common names

Argentina: Ají balita (Jujuy, Moscone 204), Pimiento (Corrientes, Anzótegui & Benitez 237; Salta, Hunziker 25498), Serrano (Salta, Hunziker 25492), Pimiento Calahorra (Córdoba, Hunziker 29428); Bolivia: Ají (Beni, Balderrama 10; Santa Cruz, Saldías P. 759), Urubibi (Beni, Ticona & Saravia May 10), Pimentón colorado (Santa Cruz, Libreros et al. 2014), Brazil: Pimentão (Roraima, Barbosa et al. 2006; São Paulo, Duth s.n.), Pimenta-americana, Pimentão-vermelho, Pimentão-indigena, Pimenta-de-mesa, Pimenta-ornamental, Pimentão-bola, Pimenta-doce (Roraima, Barbosa et al. 2006), Pimenta-jalapeño, Pimenta-cayenne, Pimenta-serrano (Carvalho et al. 2006); Chile: Ají (Concepción, Junge 5755); Colombia: Ají (Amazonas, Cordero P. 683; Chocó, La Rotta & Martínez 737; Guainía, Espina et al. 189; Vichada, Rodríguez 165), Pimentón (Amazonas, Torres & Morales 139; Caquetá, Cárdenas et al. 9310; Cundinamarca, Correa N. 006), Ají amarillo (Caquetá, Cárdenas et al. 9375, Vichada, Rodríguez 1), Ají bravo (Amazonas, Posada 2580), Ají dulce (Amazonas, Cárdenas et al. 9438; Cundinamarca, Duque-Jaramillo 3555), Ají grande (Amazonas, Cárdenas et al. 9432), Ají pajaa (Vaupes, Rodríguez 86), Ají picante (Amazonas, Posada 2581), Ají pimentón (Amazonas, Torres et al. 4011), Ají yunga (Nariño, de Benavides 4673), Ají yuquitania (Vaupes, Plowman 11986), Pimentón dulce (Caquetá, Cárdenas et al. 9360), Pimentón quisquis (Meta, Álvarez & Montañez 1), Ají de agua (Vaupes, Rodríguez 102), Ají de blanco (Amazonas, Henao & Kuiru 172), Ají de curripaco (Vaupés, Rodríguez 105), Ají de gente (Amazonas, Cárdenas et al. 9424), Ají mas picante (Vaupes, Rodríguez 83), Diente de chucha (Caquetá, Cárdenas et al. 9319), Pipí de perro (Caquetá, Cárdenas et al. 9306), Ají largo de blanco (Amazonas, Henao 316), Ecuador: Ají (Chimborazo, Lara s.n.; Pichincha, Mejía 001), Pimiento (Guayas, Bonifaz & Cornejo 4158; Pichincha, Narváez 018), Ají colorado (Tungurahua, Acosta Solís 8848), Ají manzana (Guayas, Valverde 392), Ají patateño (Chimborazo, Ganchozo 007), Manzanita de Eva (Guayas, Valverde 35); El Salvador: Chile de relleno (San Salvador, Calderón 523); Guatemala: Chile (Quezaltenango, Steyermark 34462); Honduras: Chile fuerte (Morazán, Molina R. & Molina 34535), Chile picante (Copán, Molina R. & Molina 33572; Morazán, Molina R. 34008), Chile de gallina (Cortes, Chevez 40); Mexico: Chile (Chiapas, Calzada et al. 3774; Michoacán, Miranda et al. 1519B; Tamaulipas, Rodríguez & Lira 63; Veracruz, Baizabal & Zola B. 12), Picante (Veracruz, Baizabal & Zola B. 11), Pimentón (Michoacán, Bye et al. QD 246), Chile ancho (Guanajuato, Benítez 689; Tamaulipas, Rodríguez & Lira 63b), Chile bolita (Zacatecas, Benítez 738), Chile camote (Tamaulipas, Hernández 1944), Chile cimarrón (México, Hinton 4336), Chile chaua (Yucatán, Simá 610), Chile cora (Zacatecas, Benítez 734), Chile costeño (Campeche, Ramírez A. 56), Chile chilaca (Guanajuato, Benítez et al. 386), Chile delgado (Quintana Roo, Gutiérrez 85-27), Chile dulce (Michoacán, Bye et al. 90; Tabasco, Ortíz 01; Yucatán, Simá 606), Chile gordo (Oaxaca, Hernández Ortega 484; Veracruz, Vázquez 673), Chile guajon (Zacatecas, Benítez 724), Chile kat (Yucatán, Ucan et al. 3519), Chile largo (Quintana Roo, Gutiérrez 85-41), Chile pableño (Guanajuato, Vieyra s.n.), Chile pasilla (Tamaulipas, Rodríguez & Lira 63ª), Chile pimienta (Mexico, Rodin 40), Chile pimiento (Mexico, Linares 846), Chile serrano (Hidalgo, Villa 71; Oaxaca, Martínez Calderón 1643; Veracruz, Zola & Baizabal 1439), Chile uñepicho (Veracruz, Díaz Rico 48), Chile verde, chile de huerta (Michoacán, Soto Núñez et al. 5442), Chile xalapeño, Calzada 2367), Fruto azul (Guerrero, Díaz Rico 221), Pimiento grande (Oaxaca, Bamonte 77), Chile de agua (Oaxaca, Acosta Castellanos 9417), Chile de árbol (Michoacán, Soto N. 14302), Chile de vida o chilar (Oaxaca, Moreno 29), Chile mira parba (Tabasco, Ortíz 07), Chile pico paloma (Tabasco, Ortíz 20), Chile de árbol de bola (Michoacán, Soto Núñez 14047), Chile de uña de perro (Veracruz, Vázquez 998); Nicaragua: Chile (León, Guzmán et al. 1011); Panamá: Ají (Canal Zone, Standley 28523), Sweet pepper, pimiento morrón, ají (Canal Zone, Standley 29880); Peru: Encarnado (Lima, Velarde Nuñez 20), Pimiento (Lima, Vilcapoma S. 84), Aji amarillo (Lima, Vilcapoma S. 85), Ají cerezo (Lambayeque, Libreros et al. 2013), Ají dulce (Loreto, Hormia 2228), Ají limón (Trujillo, Plowman 14541), Ají tomate (Junín, Ridoutt 11714), Bobo panca (Lima, Velarde Nuñez 19), Cerezo triangular (Lambayeque, Libreros et al. 2013), Conico amarillo (Lima, Velarde Nuñez 26), Cónico panca (Lima, Velarde Nuñez 18), Tambo Tacna (Lima, Velarde Nuñez 22), Aji Acarí Moquegua (Lima, Velarde Nuñez 7); United States of America: Black Chile or Chile negro (New Mexico, Wooton 48), Red Chile or Chile rojo (New Mexico, Wooton 49). Venezuela: Ají caribe (Portuguesa, Aymard 5108).

Indigenous names

Bolivia: Ta (Beni, Ticona & Saravia May 10); Colombia: Aati (Curripaco, Guainía, Espina et al. 189), Aii (Cauca, Plowman & Vaughan 5370), Asi (Piapoco, Vichada, Rodríguez 177), Azi (Piapocos, Vichada, Rodríguez 165), Biaá (Tanimuka, Amazonas, Cárdenas et al. 9406), Coc (Puinabe, Vichada, Rodríguez 169), Curripaati (Tucano, Guainía, Marín & Rodríguez 502), Fecogɨ (Bora, Amazonas, Torres et al. 4020), Fekorai (Huitoto-Mɨnɨka, Amazonas, Henao 167), Jipujou (Caquetá, Cárdenas et al. 9330), Jumerien (Sukuare, Vichada, Rodríguez 1), Mèe (Colona, Amazonas, Torres & Rodríguez 2021), Munɨ (Huitoto, Amazonas, Posada 2577), Nubata (Andoque, Amazonas, Torres et al. 4047), Pidá (Emberá, Chocó, La Rotta & Martínez 737), Rɨairai (Huitoto-Mɨnɨka, Amazonas, Henao & Kuiru 172), Arera rɨairai (Huitoto-Mɨnɨka, Amazonas, Henao 316), Yicane (Miraña, Caquetá, Cárdenas et al. 9375), Jeba gayebá (Mui, Castro et al. 238), Masan via (Amazonas, Cárdenas et al. 9424), Pipita deé (Mui, Amazonas, Castro 305), Viahoracá carunoje (Tanimuka, Amazonas, Cárdenas et al. 9432); Ecuador: Aatyu (Chapalaachi, Yañez et al. 1485), Uchu (Quichua, Napo, Kohn 1225), Ahí bia (Siona & Secoya Indians, Napo, Vickers 211), Suara pia (Siona & Secoya Indians, Napo, Vickers 227), Soa horo bia (Siona & Secoya Indians, Napo, Vickers 200); Mexico: Cants (Huave, Oaxaca, Zizumbo & Colunga 145), Chaunik (Yucatán, Vargas 66), Guiin-cànár (Zapateco, Oaxaca, Hunn OAX-1345), Guiin-ló-yág (Zapateco, Oaxaca, Hunn OAX-1341), Guiin-ló-ngÚbidz (Zapateco, Oaxaca, Hunn OAX-1343), Guiin-nàl-zhàb (Zapateco, Oaxaca, Hunn OAX-1342), Guiin-txxtlé (Zapoteco, Oaxaca, Hunn OAX-1344), Moo-o-re (Oaxaca, Hernández Ortega 482), Moo-o-qui (Oaxaca, Hernández Ortega 481), Niiy (Oaxaca, Antonio B. GUI 201), Xcatic (Maya, Quintana Roo, Villanueva 591), X-mash ik (Quintana Roo, Gutiérrez 85-26), X-mehen (Quintana Roo, Gutiérrez 85-27), Xkat-ik (Maya, Quintana Roo, Gutiérrez 26), Ya Jimia (Morona-Santiago, Evans 4384), Ya’axik (Maya, Quintana Roo, Gutiérrez 109), Chaua ik (Yucatán, May 39), Ixa nadun (Guerrero, Wagenbreth 130), Kat ik (Yucatán, Ucan et al. 3529), Nadam kanc (Huave, Oaxaca, Bamonte 77), Namis kanc (Huave, Oaxaca, Bamonte 79), Yaá dia (Mixteco, Guerrero, Díaz Rico 221), Yak ik (Quintana Roo, Gutiérrez 85-71), Ixe dun xkuiya smidi (Guerrero, Wagenbreth 687); Peru: Iwiá (Mayna Jívaro, Loreto, Lewis et al. 10922), Kistian jima (Amazonas, Ancuash 297), Mun hima (Amazonas, Berlín 1572), Tsitikana ogat-santsakarioni (Machiguenga, Cuzco, Johnson 70).

Uses

Capsicum annuum var. annuum is the economically most important member of the genus. The fruits are widely used in international cuisine in a broad spectrum of meals and preparations, because of their aroma, flavour, texture and level of pungency. Some cultivars have good acceptance as ornamental plants due to the colour of the leaves and the brightness of the colourful and usually erect fruits (e.g. Christmas peppers, Bolivian rainbow, Fig. 21). There are few instances where medicinal uses have been recorded on herbarium labels (Table 3), but the medical and nutritional importance, as well as the pharmacological properties and therapeutic effects of the active compounds, of C. annuum fruits have been extensively highlighted (Al-Snafi 2015; Srinivasan 2016; Masud Parvez 2017; Sricharoen et al. 2017; Saleh et al. 2018; Sanati et al. 2018; Roman et al. 2020).

Preliminary conservation assessment

Capsicum annuum var. annuum is not under threat.

Discussion

The domesticated taxon C. annuum var. annuum belongs to the Annuum clade, together with C. chinense, C. frutescens and C. galapagoense (Carrizo García et al. 2016). The three domesticated species and their conspecific wild populations constitute the Capsicum annuum primary gene pool (van Zonneveld et al. 2015).

The vast majority of the modern landraces, varietals and hybrids of chili peppers belong to this variety (Bosland and Votava 2000) and it is consequently the most intensively studied species of Capsicum with regard to diversity, domestication and genetics (OECD 2006; Pickersgill 2016; Acquadro et al. 2020 and references therein). Conversely, a full comprehension of its taxonomy has not been achieved in the last 50 years. Research indicates that its domestication could have been initiated in central-east Mexico over 6,500 years ago (Kraft et al. 2014).

Due to the selective pressure for domestication and diversification, defining a characteristic group of traits for var. annuum is difficult; however, the most distinctive features are its herbaceous to shrubby, annual or perennial habit, the solitary axillary flowers (rarely two or more), the strongly 5–10-nerved calyx, the large white (or purple) corollas (up to nearly 22 mm in diameter) and the usually persistent and pendent fruits, which are highly variable in size, form, colour and pungency. Some of these traits contrast with those of var. glabriusculum which has a shrubby habit, 5-nerved calyx, smaller corollas (≤ 12 mm in diameter) and small (< 10 mm in diameter), globose, ellipsoid or ovoid, erect, red or red-orange, deciduous fruits.

Philip Miller was the curator of the Chelsea Physic Garden in London in the late 18th century. Many of the plants he grew there were new taxa in his “Gardener’s Dictionary” (1768). He described several Capsicum species (C. cordiforme, C. tetragonum, C. angulosum, C. olivaeforme and C. pyramidale), based on cultivated specimens obtained from seeds of different provenance. As was the practice at the time, he did not cite specimens and is likely to have based his descriptions on living plants. Most of these plants were described as annuals with white flowers and a variety of fruit sizes, shapes (heart-shaped, angular-obtuse, oval-shaped, pyramidal), colours (yellow, scarlet, red), textures and positions (pendent or upright), characters that are highly variable due to human selection in these domesticated species. Specimens made from plants grown by Miller are found in several different places, mostly at BM and its associated historical herbaria. As these names are almost certainly described from living plants and, thus, will need neotypification, we do not typify them here, but leave that for a separate study when these materials, including any non-digitised specimens, can be studied in detail.

We found a collection in the Lamarck Herbarium with a label indicating that it belongs to C. conicum (P00357734) which we designate here as the lectotype.

Capsicum bicolor was probably described only from living material cultivated in the gardens of Schönbrunn Palace near Vienna (Austria). Jacquin (1809) cited no specimens for C. bicolor and gave no place of origin for his species. The description is quite complete and includes a colourful illustration (Tab. 99, fig. 1); both fall within our circumscription of C. annuum. Since no specimens have been found, we designate the illustration here as the lectotype.

There are two sheets of original material labelled C. grossum in Willdenow’s Herbarium held at Berlin. Both contain reproductive branches; one of these (B-W 04425 -02 0) consists of two fruiting branches that exactly match Willdenow’s description (Willdenow 1809) and is, therefore, designated the lectotype here.

Willdenow (1809) cited neither specimen nor locality when describing C. sphaericum. We found at B original material labelled “C. sphaericum, Hort. Bot. Berol. W” (B-W04426-01-0) and select it here as the lectotype.

In the protologue of C. purpureum, Hornemann (1813) stated “Herb. Vahlii … Hab. - -”, referring to material cultivated at Horto Hafniensis (Hafnia = Haunia = Copenhagen). A specimen held in C [C10019148], with good flowering material and bearing a label (on the verso of the sheet) with data matching the protologue, is here selected as the lectotype. Additional original material is found at C (C10019147, upper stem only), which corresponds to a Herb. Hornemann specimen cultivated in Copenhagen; this specimen is sterile.

De Candolle (1813) described C. ovatum, based on a living specimen of unknown origin cultivated at the Montpellier Botanical Garden (hort Bot. Monspeliensis), but he cited no herbarium material. We found no original material at MPU; Dunal (1852) expanded the description in the Prodromus with specimens seen in the De Candolle Herbarium, now held in G-DC. At G-DC, there are two elements: a small fruiting fragment possibly from Montpellier (“h.m.” [my herbarium] on the label) and a more complete specimen with a label stating “Capsicum ovatum D.C.” in Dunal’s hand; we designate here this latter specimen (G00200072) as the neotype.

When coining the name C. longum, De Candolle (1813) cited in synonymy pre-Linnean works, some with illustrations. We examined the illustrations and the one from J. Bauhin et al. (1651) best illustrates and corresponds to the original diagnosis and, therefore, is selected here as the lectotype.

Meyer (1818) based his description of C. globiferum on collections made by Ernst Karl Rodschied in what is now Guyana. In Göttingen, where Meyer worked, we found two Rodschied specimens labelled as C. globiferum, both from Rio Essequibo, that are certainly original material. One (GOET003420) has what appears to be a collecting number, Rodschied 29, while the other lacks any indication of a number (GOET003419). We designate here the former and most complete specimen (Rodschied 29) as the lectotype.

Capsicum purpureum is based on a single plant found in the Botanic Garden of Calcutta (India), whose exact origin is unknown, but Roxburgh (1824) suggested the seeds came from the “Molucca Islands”. A specimen at Kew (K001132446), labelled as “Capsicum purpureum” and with a faint annotation of “H.B.C.” at the bottom of the sheet, is possible original material, but we cannot be sure it was used by Roxburgh or when it was prepared. We, therefore, designate this sheet as the neotype of Capsicum purpureum.

Dierbach (1829) referred to his C. indicum var. vulgatum as “Capsicum annuum Auctor. plurimor.” making this name superfluous. His species was characterised by its red, oblong and straight fruits, one of the most common forms of the domesticated C. annuum.

Fingerhuth (1832) described many domesticated chili pepper taxa (see above); he did not mention specimens and, if he made specimens, the fate of his herbarium is unknown (Stafleu and Cowan 1976). Fingerhuth provided a set of ten plates, each one with figures in colour that illustrated almost all of the taxa included in his monograph. These figures are of good quality and match the protologues, which allowed us to assign the species to which they belong and to use them as lectotypes for Fingerhuth’s names. The names Capsicum strictum, C. ceratocarpum, C. angulosum (var. macrocarpum and ovale), some varieties of C. annuum (var. rugosulum, acuminatum, subangulosum, ovoideum, abbreviatum, olivaeforme), C. grossum (var. pomiforme, ovatum, cordatum, angulosum) and C. longum (var. incrassatum, latum, rectum) are here lectotypified, based on the Fingherhuth figures indicated above.

In the protologue of C. abyssinicum, Richard (1850) cited two collections from “Abyssinia” (Ethiopia), one made by León Richard Quartin Dillon and the other by Antoine Petit, botanists on the Lefebvre expedition to the mountains of Africa. Quartin Dillon’s collection is housed at P and consists of three sheets (P00329903, P00329904, P00329905), all of them with complete flowering and fruiting branches. We were unable to find the second collection at P, but a duplicate of the original Petit collection is at MEL (MEL 2442182) and is also a well-preserved specimen. We designate here the best-preserved collection and that which Richard is likely to have seen and used (P00329903) as the lectotype.

Dunal (1852) coined C. annuum var. oblongum with a direct citation to a polynomial and illustration in “Fingerh. l.c. t. 2 f.a” (Fingerhuth 1832); he also stated “v.s. in h. DC”. A sheet in G-DC (G00131768) is labelled “Capsicum annuum α oblongum fructibus rubris Fingerh.” from “Herb. Dunal 1844”, both in Dunal’s hand; we select this specimen as the lectotype.

In his description of C. pyramidale var. longicorne, Dunal (1852) cited the collection Zollinger 489 that he had seen in “h. Boiss. et h. DC.” (now G and G-DC). Both specimens consist of fertile branches, but that in G-DC (G00131841) has an immature fruit which confirms the identity of this name and is selected here as the lectotype.

In the protologue of Capsicum testiculatum, Dunal (1852) stated “v.s. in h. Dc. et herb. meo”. The original material came from plants grown in the Botanical Garden in Montpellier from seeds sent by R. de Visiani. The sheet in G-DC (G00200067) is a more complete fruiting branch which we select here as the lectotype.

In the protologue of C. angulosum var. macrocarpum, Dunal (1852) cited no specimens, but referred his variety to the illustration “tab. 8, fig. a” provided by Fingerhuth (1832) which is selected as the lectotype.

Dunal based C. leucocarpum on Miller’s (1752) polynomial “Capsicum americanum latifolium, fructu oblongo erecto candido”, referring to a sort of white-coloured C. annuum fruit; Fingerhuth (1832) also transcribed Miller’s polynomial exactly, but he did not provide a formal name, as appears in Index Kewensis (Capsicum leucocarpon Fingerh., accessed on 20 April 2020) or IPNI (Capsicum leucocarpon Mill. ex Fingerh. (accessed on 20 April 2020).

Sturtevant (1888b) described C. fasciculatum, based on his own cultivated living material known as Bouquet rouge (French garden name) or Red Cluster (American name). He distinguished this species from C. annuum primarily by its peculiarly clustered leaves and fruits at the summit of the plant. We found no herbarium material corresponding to this name, but Sturtevant stated in the protologue that C. fasciculatum is “well figured under the name Tenjikumamori in a Japanese botanical work…” referring to Tanaka and Ono (1874). The figure cited by Sturtevant (1888b) is very accurate and represents very well one of the many variations of the domesticated C. annuum and is here selected as the lectotype.

Greenman (1903) based C. frutescens var. lanicaule on four syntypes and characterised it as a more pubescent variant of C. frutescens. We studied all the syntypes (Palmer 639, 640, 642 and González 975). Palmer’s collections are flowering and fruiting branches, while González’s specimen has flowers and very young fruits. The fruiting calyx is critical in assigning the correct placement of this name and, amongst the fruiting Palmer specimens, the most informative one is Palmer 639 which has mature fruits typical of C. annuum var. annuum and is designated here as the lectotype.

Capsicum velutinum (De Wildeman 1922) was described based on five syntypes from different localities and collectors in what is now the Democratic Republic of the Congo, all of them well preserved at BR (Blommaert s.n., Jespersen s.n., Lamboray 22 and Lescrauwaet 315). All the specimens consist of strikingly pubescent flowering or fruiting branches or both. We designate Lamboray 22 (BR 000000649909) as the lectotype for this name, since it matches the protologue most closely.

Capsicum annuum L. var. glabriusculum (Dunal) Heiser & Pickersgill, Baileya 19 (4): 156. 1975.

Figs 22, 23

Capsicum hispidum Dunal var. glabriusculum Dunal, Prodr. [A. P. de Candolle] 13(1): 420. 1852. Type. [United States of America. Texas: Bexar Co., San Antonio]: “Mexico, circa Bejar”, Sep 1828, J.L. Berlandier 1863 (lectotype, designated by Barboza 2011, pg. 27: P [P00410138]; isolectotypes, BM [BM000775839], F [F0072795F, acc. # 680282], G [G00390278], NY [000138591], P [P00409852], YU [YU.065273]).

Capsicum minimum Mill., Gard. Dict. ed. 8, no. 10. 1768. Type. “Cultivated in England” (no specimens cited; no original material located).

Capsicum havanense Kunth, Nov. Gen. Sp. [H.B.K.] 3: 38. 1818. Type. [Cuba]. “in arenosis maritimis, prope Havanam (Insulae Cubae)” [Havana] s.d., F.W.H.A. von Humboldt & A.J.A. Bonpland 4518 (lectotype, designated here: P [P00670653]).

Capsicum indicum Dierb. var. aviculare Dierb., Arch. Apotheker-Vereins Nördl. Teutschl. 30(1): 30. 1829. Type. Based on Capsicum minimum Mill. and C. microcarpon DC. (cited in synonymy), PANAMA: Coclé, 10 mi. E of Nata at Rio Grande, 4 Jan 1969, E.L. Tyson 5222 (neotype, designated here: MO [MO-562584, acc. # 1980106]; isoneotype, FSU [000064909, acc. # 119808]).

Capsicum frutescens L. var. minus Fingerh., Monogr. Capsic.: 17. 1832. Type. “Crecit in India orientali et America meridionali” (no specimens cited; lectotype, designated here [illustration]: “Capsicum rubrum minimum” Rumphius, Herbarium Amboinense 5, Tab. 88, fig. 2, 1747]).

Capsicum pendulum Willd. var. minus Fingerh., Monogr. Capsic.: 25. 1832. Type. Based on Capsicum havanense Kunth.

Capsicum chlorocladum Dunal, Prodr. [A. P. de Candolle] 13(1): 415. 1852. Type. Mexico. Tamaulipas: “Tampico da Tamaulipas”, 1827, J.L. Berlandier 97 (lectotype, designated here: G-DC [G00131884]; isolectotypes: BM [BM000775807, BM000775821], G [G00342805], F [v0072794F, acc. # 680277], LE [LE01072484], MPU [MPU023049], P [P00410031, P00410147, P00409849]).

Capsicum laurifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 418. 1852. Type. Brazil. Bahia: “partie mérid. de la prov. de Bahia”, 1840, J.S. Blanchet 3098 A (lectotype, designated here: G-DC [G00131882]; isolectotypes: MPU [MPU023046, MPU023047).

Capsicum hispidum Dunal, Prodr. [A. P. de Candolle] 13(1): 419. 1852. Type. Mexico. Tamaulipas: circa Tupan et Tampico de Tamaulipas, 1827, J.L. Berlandier 152 (lectotype, designated here: G-DC [G00131880]; isolectotypes: BM [BM000775838], G [G00390276], G [G00390277], MO [MO-562486, acc. # 1690380], MPU [MPU013437], P [P00409850, P00410137]).

Capsicum angustifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 420. 1852. Type. “In Indiâ utrâque colitur”. Capsicum baccatum hort. Geneve, 1836, Anonymous 1414/6 (lectotype, designated here: G-DC [G00131878]).

Capsicum microphyllum Dunal, Prodr. [A. P. de Candolle] 13(1): 421. 1852. Type. [Cuba]: La Havanna, 1828, R. de la Sagra 3 (lectotype, designated by D’Arcy and Eshbaugh 1974, pg. 99: G-DC [G00131975]; isolectotype: MPU [MPU023042]).

Capsicum pendulum Willd. var. minus Dunal, Prodr. [A. P. de Candolle] 13(1): 425. 1852, nom. illeg., not C. pendulum var. minus Fingerh. (1832). Type. Based on Capsicum havanense Kunth.

Capsicum annuum L. var. minus (Fingerh.) Shinners, Baileya 4: 82. 1956. Type. Based on Capsicum frutescens L. var. minus Fingerh.

Capsicum annuum L. var. minimum (Mill.) Heiser, Ci. & Nat. 7: 52. 1964. Type. Based on Capsicum minimum Mill.

Capsicum annuum L. var. aviculare (Dierb.) D’Arcy & Eshbaugh, Phytologia 25(6): 350. 1973. Type. Based on Capsicum indicum Dierb. var. aviculare Dierb.

Capsicum frutescens L. var. glabriusculum (Dunal) M.R.Almeida, Fl. Maharashtra 3B: 356. 2001. Type. Based on Capsicum hispidum Dunal var. glabriusculum Dunal.

Type

Based on Capsicum hispidum Dunal var. glabriusculum Dunal.

Description

Perennial low herbs or somewhat prostrate subshrubs, (1–) 1.5–2 (–3) tall, the main stem woody, 0.5–1 cm in diameter at base, much branched from near the base, the branches dichotomously spreading in a typical “zig-zag” appearance above. Young stems angled, fragile, green to greenish-grey or purple-striped, glabrescent to densely pubescent, with appressed-antrorse to spreading, simple, uniseriate, (2–) 3–8 (–12)-celled, eglandular trichomes 0.3–0.9 (–2) mm long, rarely furcate trichomes; nodes solid, green or purple; bark of older stems light brown or brown, glabrous to sparsely pubescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair subequal in size and shape. Leaves membranous, discolorous, dark green above, light green beneath, glabrescent to densely pubescent on both sides, if glabrescent with an evident tuft of trichomes in the vein axils beneath, the trichomes similar to those of the stems; blades of all leaves 2.5–6 (–8.5) cm long, 1.15–2.5 (–3.4) cm wide, ovate to elliptic, the major veins 4–5 on each side of mid-vein, the base attenuate or truncate and rather unequal, the margins entire, the apex acuminate; petioles (0.5–) 1.5–2.5 (–3) cm, glabrous to moderately pubescent. Inflorescences axillary, 1–2 flowers per axil, more rarely 3 flowers; flowering pedicels 7.5–27.8 mm long, angled, erect, geniculate at anthesis, green, glabrous to moderately pubescent, the eglandular trichomes short, antrorse; pedicels scars inconspicuous. Buds globose, white, cream or greenish-white. Flowers 5-merous. Calyx 1.5–2.5 (–3) mm long, 2–3.8 mm wide, cup-shaped, green, pentagonal in outline, glabrous to moderately pubescent with similar short or long eglandular trichomes as the stems, without appendages or with five minute appendages less than 0.5 mm long. Corolla (5–) 6–8 mm long, 8–10 (–12) mm in diameter, entirely white or almost pale yellow, rarely greenish-white, stellate with narrow interpetalar membrane, lobed ca. halfway or 2/3 of the way to the base, glabrous adaxially and abaxially, the tube (2–) 3–3.5 mm long, the lobes 3–4.5 mm long, 2–2.5 mm wide, triangular, spreading, the margins slightly involute and finely ciliate, the tips acute to long-cucullate, densely papillate. Stamens five, equal; filaments 1–1.25 mm long, white, cream or purple, sometimes lilac at the apex, inserted on the corolla 1–1.3 mm from the base, with auricles fused to the corolla tube at the point of insertion; anthers 0.95–2.55 mm, broadly ellipsoid or ellipsoid, blue, bluish-grey or purple, very rarely yellow, connivent at anthesis. Gynoecium with ovary 1.2–1.5 (–2.5) mm long, 1–2 mm in diameter, green or cream, ovoid or globose; nectary ca. 0.3 mm tall, pale yellow; style homomorphic, 4–4.8 mm, exserted 1.5–2 mm beyond the anthers, cylindrical, white or pale lilac; stigma 0.1–0.2 mm long, ca. 0.3 mm wide, discoid or bilobed, pale bright green or white. Berry 6–8.5 mm in diameter, globose (larger in semi-domesticated specimens, 9–11 mm in diameter) or ellipsoid or ovoid with acute to slightly obtuse apex, 9–13 mm long, 5–6.5 mm in diameter (larger in semi-domesticated specimens, 15–25 mm long, 7–12 mm in diameter), green or green and partly dark purple or purple when immature, bright lemon-yellow, bright red-orange or red at maturity, deciduous, very pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 16–28 (–35) mm, erect, rigid, angled, widened distally, green; fruiting calyx 4–4.5 mm in diameter, persistent, not accrescent, discoid or rather cup-shaped, green. Seeds (6–) 8–26 per fruit, 3.2–4 mm long, 2.5–3.2 mm wide, C- or D-shaped, pale yellow to yellow, the seed coat reticulate to obscurely reticulate (SM), cerebelloid (SEM), the cells irregular in shape, the lateral walls strongly sinuate; embryo imbricate.

Figure 22. 

Capsicum annuum var. glabriusculum A reproductive branch B eglandular trichome of the leaf C flower D section of the calyx showing the venation E opened corolla F gynoecium G fruit H anatomical detail of the pericarp (note the giant cell in the mesocarp) I seed J seed, in cross section K structure of seed coat at the seed margin L structure of seed coat at the seed body M embryo. A–H from Singleton 195 I–M from Scolnik 19An329. Drawn by L. Ochoa. Published in Hunziker (2001), reproduced with permission.

Figure 23. 

Capsicum annuum var. glabriusculum A apex of a reproductive branch B flower bud on geniculate pedicel C–F flowers in anthesis showing variations in corolla, stamens and style colouration G immature fruit H mature and immature fruits A, B, D, F from Barboza et al. 5049, photos by G.E. Barboza C, E, H Carrizo García 102, photos by C. Carrizo García G Leiva González et al. 2105, photo by S. Leiva González.

Distribution

Capsicum annuum var. glabriusculum is the most widely distributed member of the genus, from southern United States of America to northern Bolivia and northern Brazil (Fig. 24). It is more common in Mexico, Central America, the Caribbean, Colombia and Venezuela. In eastern Australia, it is reported as a weed (Symon 1981; Purdie et al. 1982).

Figure 24. 

Distribution of C. annuum var. glabriusculum.

Ecology

Capsicum annuum var. glabriusculum occupies a wide variety of habitats throughout its wide distribution, including tropical deciduous, semi-deciduous and evergreen forests, less frequently in dry tropical or subtropical forests or in thorny scrub, from sea level to ca. 2,500 m elevation. It is found in shade along roadsides, stream banks, meadows near shores or as a weed in pastures or on the edges of cultivated lands. Indigenous communities and rural people cultivate C. annuum var. glabriusculum for self-consumption and it is often found escaped from cultivation.

Phenology

Flowering and fruiting all year.

Chromosome number

2n = 2x = 24 (Pickersgill 1971, 1977, 1991; Moscone et al. 2007; Scaldaferro et al. 2013).

Common names

Bahamas: Bird pepper (Long Island, Richey 98-355), Pepper bush (Bimini, Howard & Howard 10055); Boliva: Ají (Pando, Beck et al. 19150); Brazil: Pimenta-de-mesa, Pimenta-peito-de-moça, Pimenta-ova-de-tamuata (Pereira et al. 2011), Pimenta açaí (Amapá, Pereira & Severino 1853), Pimenta chumbinho (Amapá, Pereira et al. 1807), Pimenta de cheiro vermelha (Amapá, Pereira et al. 1747), Pimenta ova de aruanã (Amapá, Pereira et al. 1867); Colombia: Ají (Amazonas, Henao & Padd 168; Guainía, Espina et al. 191; Huila, Llanos & Camacho 1827; Valle del Cauca, Cuatrecasas 22800), Ajicito (Valle del Cauca, Cuatrecasas 22800), Ajijito (Valle del Cauca, Lehmann 4730), Cimarrón (Bolívar, Killip & Smith 14251), Ají chilca (Guainía, Augusto 4847), Ají chiquito (Norte de Santander, Carvajalino & Díaz 11; Santander, Tochoy & Garzón 675), Ají chirel (Antioquia, Santa María 733), Ají chivato (Meta, García Barriga 5056; Valle del Cauca, Duque Jaramillo 4083-A), Ají guagua (Bolívar, Espina 577), Ají pajarito (Antioquia, Barkley & Gutiérrez V. 1776; Caldas, Galán & Cárdenas 12; Norte de Santander, Garganta F. s.n.), Ají perfumado (Amazonas, Henao 169), Ají picante (Bolívar, Killip & Smith 14534; Norte de Santander, Garganta F. 818), Ahipique (Valle del Cauca, Dryander 2178), Ají pimienta (Bolívar, Romero Castañeda 9255; Santander, Betancur et al. 10158), Ají pique (Cundinamarca, Dumont et al. 43; Nariño, de Benavides 4693, Valle del Cauca, Soukup 1851), Ají piquicho (Huila, Buendía S. 2), Ají del monte (Atlántico, Bro. Elías 1461), Aji amarillo de culebra (Amazonas, Henao & Zɨuec 247), Ají ojo de charapa (Amazonas, Henao & Kuiru 173), Ají ojo de sapo (Amazonas, Henao 170), Chicha e gato (Santander, Tochoy & Garzón 675), Pimiento (Norte de Santander, Carvajalino & Díaz 11); Costa Rica: Chile, Chile congo, Chiltepe (Bohs 2015); Cuba: Ají guaguao (La Habana, García Cañizares 127; Villa Clara, Luna 356); Dominican Republic: Ají (Altagracia, Zanoni & Mejía 17077 A), Ajicito montecino (Azua, Zanoni et al. 22115; Monte Cristi, Valeur 476; Santiago, Valeur 275); Ecuador: Ají (Napo, Alarcón 102), Veneno de perro (Esmeraldas, Freire & Ruales 2931); El Salvador: Chiltepe (Ahuachapán, Standley 19872; Sonsonate, Calderón 1658), Chile de zope (Ahuachapán, Standley 19872; La Unión, Standley 20689), Chile chocolate pequeño (San Salvador, Calderón 1197); Guadeloupe: Piment Moka (Basse-Terre, Duss 3575), Piment grives (Basse-Terre, Duss 3681); Guatemala: Chiltepe, chile chiltepe (Chimaltenango, Porter 1299; Chuiquimula, Kufer 99; Huehuetenango, Steyermark 51277), Chiltep, chile de monte (Huehuetenango, Steyermark 51364), Chile de montaña (Huehuetenango, Steyermark 51277); Honduras: Chilpepe (Atlantida, Standley 54485), Chiltepe (Morazán, Standley 26254), Chiltepin (Atlantida, Standley 53628), Chile bravo (Atlantida, Standley 53392); Jamaica: Bird pepper (Kingston, Harris 10051; Saint Mary, Yuncker 18460); Leeward Islands: Piment café (Duss 149); Martinica: Piment rond, Piment sauvage (Duss 349); Mexico: Chile (Colima, Eyerdanm & Beetle 8720; Hidalgo, Blanco-Macías 1559; Puebla, Villalobos C. & Guerrero 182; Veracruz, Olivares Hernández 1), Chigundo (Oaxaca, Torres et al. 199), Chilegole (Oaxaca, Salas M. et al. 1851), Chilepepina (Baja California, Carter 4921), Chiletepin (Puebla, Sarukhán et al. 3581), Chilillo (Oaxaca, González Olivares 494), Chilpalla (Veracruz, Martínez C. 1287), Chilpitín (Baja California, León de Luz 2019; Coahuila, Wendt & Riskind 1611), Chiltapin (Chihuahua, Gentry 949 & 1541), Chiltepín (Puebla, Bye et al. 16386; Sonora, Reina G. 1012; Veracruz, Cortés 552), Chiltipen (Chihuahua, Bye Jr. 1875, Sonora, Joyal 1815), Chiltipin (Tamaulipas, Nee 32699; Veracruz, Acosta et al. 39), Chipilin (Chiapas, Calzada et al. 3990), Chirripitín (Michoacán, Madrigal Sánchez 4875), Pinchitle (Veracruz, Nee 1986), Piquín (Querétaro, Martínez Torres 57; San Luis Potosí, Edwards 592; Veracruz, Vázquez 611), Quipín (Querétaro, Martínez Torres 57), Tempenchile (Chiapas, Arcos Vernet 40), Tempinchile (Chiapas, Trujillo Eslava 75), Tepechile (Chiapas, Calzada et al. 9679), Chile bola (Veracruz, Nee 1986), Chile congo (Chiapas, Castillo C. et al. 4136), Chile chichalaco (Guerrero, Guízar Nolazco & Pimentel B. 2884), Chile chiguado (Oaxaca, Nava Zafra et al. 1812), Chile chilpalla (Veracruz, Ibarra Manríquez 3616), Chile Gole (Oaxaca, Vásquez & Ortega 865), Chiltepin grande (Puebla, Villalobos C. & Guerrero 205), Chile maxito (Campeche, Ramírez A. 57), Chile piquín (Chihuahua, Torres C. & Tenorio L. 3751; Veracruz, Nee 1986), Chile de árbol (Veracruz, Ortega T. 88), Chile de bolita (Veracruz, Martínez C. 2011), Chile Amash, chile mashito (Tabasco, Orozco-Segovia 368), Chile amachito (Tabasco, Guadarrama et al. 865), Chile chigol (Oaxaca, Gopar Vásquez 122), Chile garbanzo (Tabasco, Escolastico 165), Chile gordo (Veracruz, Castillo C. et al. 214), Chile machito (Tabasco, Ortega O. 870), Chile Pekin (Veracruz, Calzada 5577), Chili pequin (Sinaloa, Ferris & Mexia 5130), Chile piquín (Campeche, Alvaro M. 397; Nueva León, Cano s.n.; S. L. Potosí, Gómez-Lorence 877; Sinaloa, Vega A. 1297; Tamaulipas, Rodríguez 104; Veracruz, Robles G. 886), Chile silvestre (Campeche, Álvarez 89), Chile de monte (Jalisco, Pérez J. 521; Oaxaca, Gopar Vásquez 122; Yucatán, Ordonez 239), Chilote de monte (Tamaulipas, Martínez Ojeda 263), Jonnihui, tempen-chile (Chiapas, Palacios E. 636), Max, Chile max (Campeche, Álvarez 89; Quintana Roo, Serralta P. 104; Yucatán, Estrada 43), Max-hic (Campeche, Bacab W. 115), Siete Cardo (Chiapas, Matuda 17590), Chilitos de monte (Guerrero, Kruse 1906); Nicaragua: Chile (Carazo, Aranda et al. 94; Managua, Guzmán et al. 396; Masaya, Guzmán et al. 1307), Chile congo (Chontales, Nee 28288; Estelí, Nee 27745; Managua, Araquistain 5; Rivas, Araquistain 278), Chile montero (Managua, Grijalva 706); Peru: Ají (Loreto, Lewis et al. 11196), Ají del monte o charapilla (San Martín, Woytkowski 35161), Ají charapilla (Loreto, de Jong 53), Ají charapita (Ucayali, Graham & Schunke V. 460), Ají del trueno (Cajamarca, Campos & Díaz 2293), Pipi de mono (Huánuco, Becerra González & Perea 1152); Puerto Rico: Ahi caballero (Adjuntas, Stimson 3922), Ají caballos (Ponce, Britton & Britton 7344); Surinam: Spaanse pepper (Marowijne/Sipaliwini, Rombouts 735), Busi peper (Commewijne, Heilbron & Sanredjo 6); United States of America: Chillipiquin (Florida, Cory 51407), Bird pepper (Florida, Kral 1884), Cayenne Pepper (Florida, Bishop & B. Holst CC0048); Venezuela: Pajarito (Lara, Tamayo 2618), Ají corito (Mérida, Trujillo 6351), Ají pajarito (Carabobo, Hunziker 9032; Mérida, Pittier 12837), Ají de mono (Bolívar, Liesner & González 5417), Chirel del Mono (Bolívar, Knab-Vispo et al. 1288).

Indigenous names

Colombia: Aati (Curripaco, Guainía, Espina et al. 191), Aiyo borarede jairai (Huitoto-Nɨpode, Amazonas, Henao & Zɨueche 247), Beeakxtú (And, Amazonas, Castro & Andoke 607), Jɨgɨngo uijɨ (Huitoto-Mɨnɨka, Amazonas, Henao 170), Jimorai (Huitoto-Mɨnɨka, Amazonas, Henao & Padd 168), Kupirapa’ajiné (Amazonas, Castro & Matapí 523), Meniño uijɨ (Huitoto-Mɨnɨka, Amazonas, Henao 173), Wainpiraicha (Guajira, Betancur et al. 11258), Ziorai (Huitoto-Mɨnɨka, Amazonas, Henao 169); Ecuador: Bula uchu (Napo, Irvine 773), Giimo (Oncaye, Napo, Davis & Yost 994), Jimiea (Achuar Jívaro, Pastaza, Lewis et al. 14010), Jimia (Shuar, Zamora-Chinchipe, Van den Eynden et al. 700), Sampíajimia (Shuar, Zamora-Chinchipe, Santín et al. 100), Uchu (Quichua, Napo, Alarcón 102), Uchumuyu (Quichua/Spanish, Pastaza, Lewis et al. 14010); Guatemala: Chi-ik (Alto Verapaz, Standley 90936), Tamut ich (Ch’orti’, Chuiquimula, Kufer 99); Mexico: Guiiña (Zapateco, Oaxaca, Sánchez L. et al. 1190), Skapin (Totonaco, Veracruz, Cortés-Vásquez 552 & 143), Guiiña dxuladi (Zapateco, Oaxaca, Sánchez L. & Trujillo V. 874), Guien guiix (Oaxaca, Ruiz Núñez 7), Guiinya xigundu (Zapateco, Oaxaca, Sánchez L. 317), Kulum its (Huastec, S. Luis Potosí, Alcorn 2369), Max ik (Campeche, Álvarez 89; Yucatán, Ucan et al. 3527 & 3893), Lak’su pin (Tot, Puebla, Villalobos C. & Guerrero 205), Tsakam its (Huastec, S. Luis Potosí, Alcorn 1406; Veracruz, Alcorn 1903), Xmax ik (Yucatán, Ucan 4617), Aj max iik (Yucatán, Ucan 5058); Peru: Ají (Quichua, Loreto, Lewis et al. 12906), Cusharu’ nu’ca” (Chayahuita, Loreto, Odonne 561), Imiá (Achual Jívaro, Loreto, Lewis et al. 11196), Nuca (Loreto, Odonne 25), Nu’ca (Loreto, Odonne 626), Uchu (Loreto, Lewis et al. 12552), Yaa Jimia (Amazonas, Salaün 185), Yampit jima (Amazonas, Berlin 2016), Yanco nu’ca” (Chayahuita, Loreto, Odonne 563); Surinam: Lombo riwit (Ja, Commewijne, Heilbron & Sanredjo 6), Lombo kusti ‘pepper of god’ (Commewijne, Heilbron & Sanredjo 6).

Uses

This taxon is used as an ornamental, for food and for medicine. The fruits are harvested by local people and are widely used and much prized throughout its distribution as a hot seasoning; they are also eaten fresh, dry or in vinegar, raw or toasted. Some medicinal properties have been attributed to the leaves and fruits in different countries (see Table 3). In some communities, fruits are used against evil spirits in ritual practices of the Day of the Dead (Mexico).

Preliminary conservation assessment

EOO (37,301,728.615 km2); AOO (4,496 km2). Capsicum annuum var. glabriusculum is not under threat for the time being.

Discussion

Capsicum annuum var. glabriusculum, better known as ‘chiltepin’ or ‘chilipequin’ (with some variations of these names) in Mexico and Central America (see common names) or ‘bird pepper’ in the United States and the Caribbean, belongs to the Annuum clade (Carrizo García et al. 2016). It is considered the wild progenitor of the cultivated C. annuum var. annuum from which it can easily be differentiated by its fragile-stemmed somewhat prostate habit, very small flowers, calyx with 0–5 inconspicuous appendages, short filaments and small globose, ellipsoid or ovoid red-orange or red fruits.

In herbaria or in literature, many names have been misapplied to the specimens of this variety, such as C. baccatum, C. frutescens, C. conoides, C. annuum var. conoides, C. annuum var. baccatum, C. frutescens var. baccatum and so on. Based only on the morphology of the fruits, this variety is sometimes confused with wild C. baccatum var. baccatum. While the fruiting calyx of C. annuum var. glabriusculum has 0–5 inconspicuous appendages and the fruits are generally more ovoid with an acute to slightly obtuse apex (rarely truncate), in C. baccatum var. baccatum, the calyx has five appendages up to 2 mm in length and the fruits are generally more globose or subglobose to ellipsoid with a truncate or flattened apex (very rarely acute to slightly obtuse). In addition to the differences in the fruits, C. annuum var. glabriusculum has solitary or paired flowers (rarely three flowers), stellate corollas that are entirely white to greenish-white without spots within and connivent blue, bluish-grey or purple anthers at anthesis (Fig. 23C–F), whereas C. baccatum var. baccatum has usually 2–3 flowers (rarely one), white rotate or rotate-stellate corollas with greenish-yellow spots within and not connivent, usually white or pale yellow anthers at anthesis (Fig. 26D–F).

Kunth (1818) did not cite a specific specimen from Cuba in the protologue of C. havanense. At P, we found a sheet labelled C. havanense with the number ‘4518’ (P00670653) in the Herbarium of Humboldt and Bonpland; this sheet is selected as the lectotype.

Dierbach (1829) based Capsicum indicum var. aviculare on C. minimum Mill. and C. microcarpum DC. Capsicum minimum applies to the wild specimens of C. annuum while C. microcarpon refers to the wild C. baccatum form which is, in fact, very similar to the wild C. annuum in the fruiting stage. D’Arcy and Eshbaugh (1973) proposed C. annuum var. aviculare, based on C. indicum var. aviculare, a name that was frequently used in literature to refer to the widespread spontaneous variety of C. annuum. In the interest of fixing the application of the basionym, we are designating a neotype using the modern collection E.L. Tyson 5222 (MO-562584, acc. # 1980106) that D’Arcy used to illustrate this taxon (figure 4 in D’Arcy 1973) which shows its diagnostic characters.

Fingerhuth (1832) gave a very brief description for C. frutescens var. minus (“fructu ovato obtuso minori”) and based this name on an illustration in the pre-Linnean Rumphius’ Herbarium Amboinense (1747: Tab. 88, fig. 2, as Capsicum rubrum minimum). This figure is of a complete flowering and fruiting branch that unequivocally matches with our concept of C. annuum var. glabriusculum. As no collections were cited and no specimens with internal evidence of being part of the original material were found, Tab. 88, fig. 2 of Rumphius’ work is selected as the lectotype.

In describing C. chlorocarpum, Dunal (1852) cited in the protologue “Berland. n. 97, in h. DC. et Boiss”. We did not find the duplicate that Dunal saw in the “h. Boiss” (now part of the general herbarium at G). Therefore, we select here the duplicate at G-DC (G00131884) as the lectotype.

Capsicum laurifolium was described, based on two different specimens in G-DC, both mounted on the same sheet. The right hand specimen (Anonymous 67, G00131902) comes from the Island of Guadeloupe in the Leeward Islands, part of the Lesser Antilles in the Caribbean. The left hand specimen is that of Blanchet from Bahia (Brazil). Of these two collections, we have selected the most complete specimen that most closely matches the data in the protologue (Blanchet 3098 A) as the lectotype (G00131882).

Dunal (1852) based the description of C. hispidum on three specimens of Berlandier 152 today housed at G. The one in G-DC (G00131880) is the best-preserved and is here selected as the lectotype.

Dunal (1852) described C. angustifolium, based on specimens from plants cultivated in the Botanic Gardens in Geneva (“hort. Geneve 1836”). We have selected as the lectotype a sheet in G-DC (G00131878) with the same data as the protologue (“Capsicum baccatum hort. Geneve, 1836”) and a handwritten label with “Capsicum angustifolium Dun., janvier 1845” in Dunal’s hand.

Barboza’s (2011) lectotypification for C. microphyllum using the collection Berlandier 1907 from “Béjar” (Texas, United States of America) must be set aside, since D’Arcy and Eshbaugh (1974) explicitly lectotypified this name earlier with the collection of Sagra from Cuba at G-DC.

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum baccatum L., Syst. Nat., ed. 12, 2: 174. 1767; Mant. Pl.: 47. 1767.

Type

“Habitat in Indiis” Herb. Linn. N° 249.3 (lectotype, designated by D’Arcy and Eshbaugh 1974, pg. 95: LINN [LINN-HL249-3]).

Capsicum baccatum L. var. baccatum

Figs 25, 26

Capsicum pulchellum Salisb., Prodr. Stirp. Chap. Allerton: 134. 1796, nom. illeg. superfl. Type. Based on Capsicum baccatum L. (cited in synonymy).

Capsicum microcarpum Cav., Descr. Pl. (Cavanilles): 371. 1802. Type. Cultivated in the Royal Botanical Garden in Madrid, Spain “H.R.M. [Hortus Regis Matritensis]. Se cría en la Havana... y se cultiva en el Jardín botánico” (lectotype, designated here: MA [MA-307276]).

Capsicum ciliare Willd., Enum. Pl. [Willdenow] 1: 243. 1809. Type. Cultivated in Berlin, Germany, of unknown origin “Cult. in Hort. Bot. Berol.”, C.L. Willdenow s.n. (lectotype, designated here: B [B-W04430-01-0]).

Capsicum indicum Dierb. var. ribesium Dierb., Arch. Apotheker-Vereins Nördl. Teutschl. 30 (1): 29. 1829. Type. Based on C. baccatum L.

Capsicum comarim Vell., Fl. Flumin.: 60. 1829 (“1825”); Fl. Flumin. Icon. 2: t. 2. 1831 (“1827”). Type. Brazil. [Rio de Janeiro]: “Colitur hortis, et sponte undequaque crescit” (lectotype, designated by Knapp et al. 2015, pag. 284: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_005] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 2. 1831).

Capsicum cumanense Fingerh., Monogr. Capsic.: 17. 1832, nom. illeg. superfl. Type. Based on (renaming of) “Capsicum baccatum Kunth” [= C. baccatum L.] (cited in synonymy).

Capsicum microcarpum DC. forma fruticosum Sendtn., Fl. Bras. (Martius) 10(6): 146. 1846. Type. Brazil “In Brasilia”, Pohl s.n. (lectotype, designated here: M [M-0171544]).

Capsicum microcarpum DC. forma herbaceum Sendtn., Fl. Bras. (Martius) 10(6): 146. 1846. Type. Brazil. “Martius Mss. in Itinerario n. 132”, Prope Polafoco, Sept., C.F.P. Martius 132 (lectotype, designated here: M [M-0171543]; isolectotype, CORD [CORD00101765]).

Capsicum annuum L. var. microcarpum (DC.) Alef., Landw. Fl.: 133. 1866. Type. Based on Capsicum microcarpum DC.

Capsicum annuum L. var. baccatum (L.) Kuntze, Revis. Gen. Pl. 2: 449. 1891. Type. Based on Capsicum baccatum L.

Capsicum annuum L. var. microcarpum (Cav.) Voss, in Vilm. Blumengärtn., ed. 3. 1: 723. 1894. Type. Based on Capsicum microcarpum Cav.

Capsicum frutescens L. var. baccatum (L.) Irish, Rep. (Annual) Missouri Bot. Gard. 9: 99. 1898. Type. Based on Capsicum baccatum L.

Capsicum microcarpum Cav. var. glabrescens Hassl., Repert. Spec. Nov. Regni Veg. 15: 244. 1918. Type. Paraguay. Canindeyú: “Iter ad Yerbales montium Sierra de Maracayu, in regione cursus superioris fluminis Jejui guazú”, Dec. 1898-99, É Hassler 5703 (lectotype, designated by Barboza 2011, pg. 28, second step designated here: G [G00390268]; isolectotypes: BM [BM000074084, acc. # 5447772; BM000074084a, acc. # 4575837; G [G00390266 two sheets with same barcode, G00390267], GH [00936720], K [K000585896], MO [MO-503802, acc. # 1574551], NY [00138600], P [P00410160, P00410161, P00482076], UC [UC944854], W [acc. # 1902-0002869]).

Capsicum annuum L. subsp. baccatum (L.) Terpó, Feddes Repert. 72: 173. 1966. Type. Based on Capsicum baccatum L.

Description

Erect shrubs or perennial herbs 0.50–3 (–3.5) m tall, rarely small trees, the main stem 2-2.5 cm in diameter at base, much branched from near the base and above, the branches spreading in a typical “zig-zag” appearance. Young stems 3–4-angled, fragile, green, sometimes the ridges purple, mostly glabrous to sparsely or moderately pubescent with appressed-antrorse, simple, uniseriate, 4–7-celled, eglandular trichomes 0.2–1.2 mm long; nodes usually purple; bark of older stems fissured, dark brown, glabrous; lenticels abundant. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar in shape. Leaves membranous, slightly discolorous, dark green above, light green beneath, glabrescent to moderately pubescent with appressed-antrorse trichomes like those of the stems on both surfaces and margins; blades of major leaves 4.5–10 cm long, 2.5–6 cm wide, ovate, the major veins 5–8 on each side of mid-vein, the base somewhat asymmetric and attenuate, the margins entire, the apex acute; petioles (1.5–) 2–4 cm long, moderately to densely pubescent; blades of minor leaves 3.5–4.5 cm long, 2–3 cm wide, ovate, the major veins 4–5 on each side of mid-vein, the base rounded or truncate, the apex acute; petioles 0.5–1 cm long, moderately to densely pubescent. Inflorescences axillary, 2–3 flowers per axil, rarely flowers solitary; flowering pedicels (17–) 20–35 mm long, angled, erect or slightly spreading, geniculate at anthesis, glabrescent to moderately pubescent, the eglandular trichomes short, spreading or antrorse; pedicel scars inconspicuous. Buds globose, white with greenish-yellow spots, occasionally purple. Flowers 5-merous. Calyx 1.5–2 (–2.5) mm long, ca. 2–2.5 mm wide, cup-shaped, thick, green, pubescent with the same trichomes as pedicels and some glandular trichomes, the calyx appendages 5, (0.3–) 0.5–2 mm long, 0.2 mm wide, subequal, thick, erect, cylindrical, inserted close to the margin, pubescent with the same trichomes as calyx tube. Corolla 4.5–7.5 mm long, 10–13 mm in diameter, thick, white with greenish-yellow spots and white centre outside and within, rotate or rotate-stellate, with interpetalar membrane, lobed 1/3 or less of the way to the base, pubescent adaxially with short glandular trichomes (stalk 1–3-celled; head globose, peltate, unicellular) in the throat and base of the lobes, glabrous abaxially, the tube 4–5 mm long, the lobes 2.5–2.7 mm long, 3.4–3.5 mm wide, broadly triangular, spreading, the margins with very short eglandular trichomes, the tips acute, papillate. Stamens five, equal; filaments 2.5–3.5 mm long, white, inserted on the corolla 1–1.1 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.5–1.8 mm long, ellipsoid, white or pale yellow, more rarely greyish, not connivent at anthesis. Gynoecium with ovary 2.5–2.7 mm long, ca. 2 mm in diameter, ovoid, green; nectary 0.3–0.5 mm tall; styles dimorphic, short style 2–2.5 mm long, not exceeding the anthers length, long style ca. 3.5 mm long, exserted 1.4–1.7 mm beyond the anthers, cylindrical, white; stigma 0.3 mm in diameter, globose or discoid, pale green. Berry 6–8 (–10) mm in diameter, globose or subglobose, less frequently ellipsoid with truncate or flattened apex, 10–20 mm long, 4–7 mm in diameter, green when immature turning to greenish-black and bright red at maturity, deciduous, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 20–35 mm long, erect, strongly angled, widened distally, green; fruiting calyx 3–4.5 mm in diameter, persistent, not accrescent, cup-shaped or discoid, green, the appendages 1.5–2.3 mm long, appressed to the berry or spreading. Seeds 12–15 per fruit, 2.5–4 mm long, 2.3–3 mm wide, ovoid, subglobose or C-shaped, pale yellow to yellow, the seed coat smooth or slightly reticulate (SM), cerebelloid (SEM), the cells irregular in shape, the lateral walls sinuate strongly sinuate; embryo imbricate.

Figure 25. 

Capsicum baccatum var. baccatum A flowering branch B calyx C glandular trichome of the calyx D eglandular trichome of the calyx E flower, upper view F sector of opened corolla G gynoecium H fruit I seed J seed, in cross section K embryo A–G from Hunziker 7350 H–K from Hunziker 1579. Drawn by N. de Flury. Published in Barboza (2013), courtesy of the Board of the Instituto Darwinion (San Isidro, Buenos Aires, Argentina), reproduced with permission.

Figure 26. 

Capsicum baccatum var. baccatum A plant B flower bud C purple flower bud D, E, F flowers at anthesis, different views G young fruiting branch showing ovary with long and short styles H immature fruits I mature fruit A, C from Barboza 4913 B, D–F, H from Barboza 5038 G from Barboza 2431 bis, I from Barboza et al. 3419. Photos by G.E. Barboza.

Distribution

Capsicum baccatum var. baccatum is widely distributed in South America from northern Venezuela and Colombia through Peru, Bolivia, Paraguay, northern and north-eastern Argentina to south and eastern Brazil (Fig. 27).

Figure 27. 

Distribution of C. baccatum var. baccatum, C. baccatum var. pendulum and C. baccatum var. umbilicatum.

Ecology

Capsicum baccatum var. baccatum occurs in dry or humid subtropical or tropical forests with semi-deciduous or deciduous vegetation, between 150 and 1,900 m elevation; it is quite common in Chaco scrub forests, in gallery forests and in the margins or interior of secondary forests. It is frequently a ruderal in disturbed areas.

Phenology

Flowering from October to May. Fruiting from late November to September.

Chromosome number

2n = 2x = 24 (Pickersgill 1977; Moscone et al. 2003, 2007).

Common names

Argentina: Coincho (Jujuy, Fabris 3454), Cumbarí (Misiones, Montes 15164), Quitucho (Salta, Hunziker 1985), Puta-parió (Corrientes, Martínez Crovetto 11125), Ají quitucho (Salta, Hunziker 1579), Ají del campo, Ajitucho (Salta, West 8389), Ají del monte (Salta, Rial Alberti s.n.), Pimenta del monte (Misiones, Montes 15164), Pimentón del monte, ají cumbarí (Misiones, Montes 15202); Bolivia: Arabibi (Santa Cruz, Zenteno-R 12798), Aribibe (Santa Cruz, Hurtado 296), Aribibi (Chuquisaca, Debouck 3019; Santa Cruz, de Michel 159), Arivivi (Chuquisaca, Serrano 1903; Santa Cruz, Cárdenas 4702), Cobincho (Tarija, Krapovickas & Schinini 39010), Ají aribibi (Cochabamba, Thomas 705), Aribibi silvestre (Beni, Rivero 218), Ají del campo (Tarija, Krapovickas & Schinini 39010), Arivivi grande o cumbarito (Fundación PROINPA 2007), Arivivi last’a o miska (Fundación PROINPA 2007), Arivivi morado (Fundación PROINPA 2007), Arivivi tuna árbol grande (Fundación PROINPA 2007), Arivivi tuna con flor blanca (Fundación PROINPA 2007); Brazil: Cumari (Espírito Santo, Crepaldi 59), Pimenta-cumari (Goiás, Mendonça et al. 5971), Pimenta de passarinho (Bahia, Mori 11603); Colombia: Ají (Vichada, Rodríguez 164), Ají de babilla (Amazonas, Cárdenas 9423), Ají de la capitania (Amazonas, Cárdenas 9403); Peru: Aji Ayucllo (Junín, Bosland and Votava 2000).

Indigenous names

Bolivia: Pochetii (Trinitario, Cochabamba, Thomas 705), Winno, Sachimi (Yuracare, Cochabamba, Thomas 705); Colombia: Azi (Piapocos, Vichada, Rodríguez 164), Gugsobia (Amazonas, Cárdenas 9423), Kulana (And, Amazonas, Castro & Matapí 564), Kulana (Yucuna, Amazonas, Cárdenas 9403), Kuraraka (Letuama, Amazonas, Cárdenas 9401); Paraguay: Hõmpita (Ayoreo, Boquerón, Gragson 124), Nuuhá (Alto Paraguay, Schmeda 1584).

Uses

As fruits are generally extremely pungent, they are collected and stored for use as a food condiment by native populations. Some accessions of this wild pepper (“arivivi”) in Bolivia have been considered promising for their interesting agro-morphological and biochemical characteristics with potential for the development of high value products for different markets (Fundación PROINPA 2007; van Zonneveld et al. 2015). However, commercialisation is still marginal and fruits are cultivated in home gardens or gathered from nature for self-consumption or to be distributed locally (Jäger et al. 2013).

Preliminary conservation assessment

EOO (11,809,545.422 km2); AOO (1,212 km2). Capsicum baccatum var. baccatum is considered Least Concern (LC) for the time being.

Discussion

Capsicum baccatum var. baccatum is a member of the Baccatum clade and is related to C. rabenii and C. chacoense (Carrizo García et al. 2016). This entity is considered to be the wild progenitor of the cultivated C. baccatum var. pendulum and is widespread in South America. Its main centre of domestication is thought to be in the Bolivian Amazonia and inter-Andean valleys (Scaldaferro et al. 2018).

In an effort to clarify the taxonomy of C. baccatum, Eshbaugh and collaborators (Eshbaugh 1968, 1970, 1976; D’Arcy and Eshbaugh 1974; Jensen et al. 1979; Pickersgill et al. 1979; McLeod et al. 1983a, 1983b; Mitchell et al. 1989) and others (Kuriachan 1981; Egawa and Tanaka 1984; Jarret 2007; Scaldaferro et al. 2018) have studied the wild and domesticated forms of this species with a multitude of techniques (morphological, breeding, cytogenetic, biochemical, molecular, phylogeographical); the results support the wild progenitor-domesticate association proposed by Eshbaugh (1968).

Capsicum baccatum var. baccatum typically exhibits 2–3 flowers per node, rarely solitary flowers, geniculate pedicels that are erect or declining at anthesis, 5-merous flowers with white rotate or rotate-stellate corollas with greenish-yellow spots, dimorphic styles and small, globose, subglobose or ellipsoid, erect, deciduous, red fruits (Fig. 26). The domesticated genetic lines mainly differ in having larger, 5–8-merous flowers, an ovary with 2–5 locules, quite diverse fruits that vary in size, colour (green, yellow, brown, orange, red) and shape (pendent, usually elongate or of different forms) and larger seeds.

Fruiting specimens of C. baccatum var. baccatum are very similar to C. rabenii and it is sometimes impossible to distinguish the two, especially if there are no annotations about the corolla colour (in C. rabenii, corolla lobe margins are purple). However, C. baccatum var. baccatum usually has glabrescent to moderately pubescent leaves in contrast to the densely lanose pubescence found abaxially along the main veins in C. rabenii (Fig. 106B).

Capsicum baccatum var. baccatum differs from C. chacoense, with which it is sympatric in some localities of Argentina, Bolivia and Paraguay, by usually having five calyx appendages, a larger and rotate or rotate-stellate corolla with greenish-yellow pigmentation within, staminal plaques with auricles fused to the corolla and long and short styles. In contrast, C. chacoense has a calyx with 5–10 unequal appendages, entirely white and smaller corollas (4–6 mm long), staminal plaques with auricles not fused to the corolla and homomorphic styles (Fig. 46). Fruiting specimens of C. baccatum var. baccatum sometimes are difficult to distinguish from C. chacoense. The fruiting pedicels of C. baccatum var. baccatum are usually 2–3 per node and the fruiting calyces have five subequal appendages, without any evidence of a mucro below the calyx margin, which differs from the solitary fruiting pedicels and 5–10 unequal calyx appendages in C. chacoense.

Some earlier botanists submerged the epithet baccatum under C. annuum (Kuntze 1891) or C. frutescens (Irish 1898) to name specimens belonging to C. annuum var. glabriusculum or wild forms of C. baccatum, an interpretation followed by other researchers (Smith and Heiser 1957; Emboden 1961; Terpó 1966).

Although the pungency of C. baccatum is regarded as low-mild (Libreros et al. 2014; Tanaka et al. 2017), some accessions of C. baccatum var. baccatum from Bolivia have been reported as non-pungent (e.g. Chuquisaca: Manchego CBNP 04, 05 & 06; Santa Cruz: Manchego CBNP 01, 02 & 03; also see Tewksbury et al. 2006).

Cavanilles (1802) described C. microcarpum, based on material cultivated at the Real Jardín Botánico de Madrid from seeds sent from Cuba. We found two Cavanilles specimens at MA gathered in the Real Jardín Botánico de Madrid in 1802: MA-307276, separated as type material in the Cavanillesii Typi collection (Garilleti 1993; https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.ma307276?loggedin=true) and MA-307278, which is filed in the general collection. Both specimens are labelled as Capsicum microcarpum. According to Garilleti (pers. comm.), the original writing on the sheet label MA-307278 (the left label) is not from Cavanilles, but from Demetrio Rodríguez; since this plant was grown in the Real Jardín Botánico, Rodríguez could have collected it later and Cavanilles might not have seen it. Therefore, the only unequivocal original material is that with the label in Cavanilles’ handwriting (MA-307276), which is here selected as the lectotype; this sheet consists of three branches with flowers and fruit.

Willdenow (1809) based the description of C. ciliare on a specimen of unknown origin cultivated in the Botanical Garden of Berlin. At B, there is one sheet in Willdenow’s Herbarium with the script “C. ciliare, Hort. Bot. Berol. W.”; this sheet is selected as the lectotype.

Sendtner (1846) described C. microcarpum forma fruticosum, based on two collections, one from Corego de Jaragua, São Paulo (“leg. Pohl”) and the other from “prov. Sebastianopolitana, ad Lagoa de Freitas” [Rio de Janeiro], but he cited no herbaria. We have found only the specimen collected by Pohl (M-0171544), which is selected as the lectotype.

For C. microcarpum forma herbaceum, Sendtner (1846) mentioned the specimen “Mart. Mss. in Itinerario n. 132”, but cited no herbaria. We found a collection at M (M-0171543) with a handwritten description that is in agreement with Sendtner’s diagnosis and has the number 132. Therefore, this sheet is designated the lectotype.

When Kuntze (1891) proposed the combination C. annuum var. baccatum (L.) Kuntze, he was referring to specimens that actually correspond to the spontaneous forms of C. annuum (cfr. Kuntze s.n., NY barcode 01008231), but he incorrectly used the epithet baccatum L. which corresponds to a different species (C. baccatum L.).

When Terpó (1966) proposed the combination C. annuum subsp. baccatum, based on C. baccatum L., he clearly stated “Mexikanischer Wildpaprika” which actually belongs to the spontaneous forms of C. annuum.

Specimens examined

See Suppl. material 4: Appendix 4.

Capscium baccatum L. var. pendulum (Willd.) Eshbaugh, Taxon 17: 51. 1968.

Fig. 28

Capsicum pendulum Willd., Enum. Pl. [Willdenow]: 242. 1809. Type. Cultivated in the Berlin Botanic Garden, Germany “Habitat ... [Country unknown]. Cult. in Hort. Bot. Berol”., C.L. Willdenow s.n. (lectotype, designated here: B [B-W04431-01-0]).

Capsicum frutescens L. var. pendulum (Willd.) Besser, Cat. Jard. Bot. Krzemieniec: 29. 1816. Type. Based on Capsicum pendulum Willd.

Capsicum indicum Dierb. var. pendulum (Willd.) Dierb., Arch. Apotheker-Vereins Nördl. Teutschl. 30: 28. 1829. Type. Based on Capsicum pendulum Willd.

Capsicum pendulum Willd. var. majus Dunal, Prodr. [A. P. de Candolle] 13(1): 425. 1852. Type. No locality cited (no specimens cited; no original material located; Dunal may have considered this the typical variety).

Type

Based on Capsicum pendulum Willd.

Description

Erect shrubs or perennial herbs (0.60–) 1–2.5 m tall, with the main stem 1.5–2.5 cm in diameter at base, much branched from near the base and above, the branches spreading in a typical “zig-zag” appearance. Young stems 3–4-angled, fragile, dark green or green, mostly glabrous to sparsely or moderately pubescent with appressed-antrorse, short to long, simple, uniseriate, 4–9-celled, eglandular trichomes 0.5–1.3 mm long; nodes green; bark of older stems green with light brown fissures, glabrous; lenticels absent or few. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, equal or subequal in shape. Leaves membranous, discolorous, dark green above, light green beneath, glabrous to glabrescent with short eglandular trichomes in margins and long, spreading, 5–9-celled, eglandular trichomes along the primary veins and in the vein axils beneath; blades of major leaves 5–12 (–14.5) cm long, 3–5 (–7) cm wide, ovate, the major veins 5–7 on each side of mid-vein, the base asymmetric and attenuate or symmetric and rounded, the margins entire, the apex acute or acuminate; petioles 2.5–7.5 cm long, sparsely pubescent; blades of minor leaves 3–5.8 cm long, 1.3–2.5 cm wide, ovate or elliptic, the major veins 3–4 on each side of mid-vein, the base rounded, the margins entire, the apex acute; petioles 1.7–2 cm long, sparsely to moderately pubescent. Inflorescences axillary, 2–3 flowers per axil or flowers solitary; flowering pedicels 20–50 mm long, terete or angled, erect, sometimes curved, geniculate at anthesis, glabrous, glabrescent to moderately pubescent, the trichomes short, antrorse; pedicels scars inconspicuous. Buds globose, white with greenish-yellow spots, occasionally purple. Flowers 5–8-merous. Calyx 2–3 mm long, 3–4.2 mm wide, cup-shaped, thick, strongly 10-nerved, green, glabrous or glabrescent, the calyx appendages 5–6, rarely up to 8, 0.9–2.5 mm long, 0.2 mm wide, subequal, thick, erect, cylindrical, inserted close to the margin, with the same pubescence as calyx tube. Corolla 8.5–15 mm long, 12–16 (–20) mm in diameter, thick, white with greenish-yellow to tan spots and a white centre outside and within, rotate to rotate-stellate, with interpetalar membrane, lobed less than 1/3 of the way to the base, pubescent adaxially with short glandular trichomes (stalk 1–3-celled; head globose, peltate, unicellular) in the throat and base of the lobes, glabrous abaxially, the tube 4–5 mm long, the lobes 3–3.3 (–5) mm long, 3.5–6 mm wide, triangular to broadly triangular, spreading, the margins finely ciliate, the tips acute, papillate. Stamens 5–8, equal; filaments (2.5–) 3–4 mm long, white, inserted on the corolla 1.2–1.5 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 2–2.6 mm long, ellipsoid or ovoid, yellow, brownish post-dehiscent, not connivent (rarely connivent) at anthesis. Gynoecium with ovary 2.5–3 mm long, 1.6–2.5 mm in diameter, 2–5-carpelar, light green, ovoid; ovules more than two per locule; nectary ca. 1.2 mm tall, yellowish-green; styles dimorphic, short style 1.3–2 mm long, not exceeding the anthers length, long style 2.5–3.5 mm long, at the same level or slightly exserted beyond the anthers, yellowish-white, cylindrical; stigma ca. 0.2 mm long, 0.7–0.8 mm wide, discoid or bilobed, pale green. Berry 20–180 mm long, (10–) 20–40 (–50) mm in diameter, usually elongate or elongate-curved, triangular or campanulate, rarely subglobose, the base truncate or obtuse, the apex rounded, blunt or pointed, dark green or green when immature, green, bright yellow, orange, brown or red at maturity, persistent, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels (35–) 50–95 mm long, pendent, sometimes strongly curved, terete or slightly angled, widened distally, green; fruiting calyx 9–18 (–20) mm in diameter, persistent, slightly accrescent, campanulate, thick, somewhat corrugated or not, green, the appendages 0.5–2 mm long, appressed to the berry. Seeds 30–80 per fruit, (3–) 4–5.2 mm long, 3–3.8 (–4) mm wide, reniform or C-shaped, pale yellow to yellow, the seed coat smooth or slightly reticulate (SM), minutely reticulate (SEM), the cells polygonal to irregular in shape, the lateral walls straight to wavy; embryo imbricate.

Figure 28. 

Capsicum baccatum var. pendulum A plant B flower bud C flower, in pre-anthesis D flower short-styled, longitudinal section E flower long-styled, lateral view F–H flowers hexamerous showing connivent anthers and not connivent anthers I immature fruit J–M mature fruit A, B, E from Barboza 4886 C, D, I, K, L, M no specimen vouchers (cult. Pairumani, Cochabamba-Bolivia) F, G, H from Palombo 3 J from Barboza et al. 4824. Photos by G.E. Barboza.

Distribution

Capsicum baccatum var. pendulum is found from low to mid-Andean elevations, mainly in Bolivia and Peru, extending to Ecuador and Colombia in the north and reaching Argentina, Paraguay and south-western Brazil in the south (Fig. 27). Introductions to North and Central America (D’Arcy and Eshbaugh 1974; Thampi 2003), northern Brazil (Roraima, Barbosa et al. 2006), Europe (Rodríguez-Burruezo et al. 2009), Japan (D’Arcy and Eshbaugh 1974) and India (Thampi 2003; Rudrapal and Sarwa 2020) are recent.

Ecology

Capsicum baccatum var. pendulum is a cultivated plant adapted to many different ecological conditions between 150 and 3,400 m elevation.

Phenology

Flowering and fruiting all year.

Chromosome number

2n = 2x = 24 (Pickersgill 1977; Moscone et al. 2003, 2007).

Common names

Argentina: Ají (San Juan, Ariza Espinar 3214; Corrientes, Benítez 76); Varita (Salta, Krapovickas & Schinini 28134), Ají picante (Salta, Hunziker 25496); Ají vainilla (Salta, Hilgert 1363), Puta parió (Corrientes, Martínez Crovetto 11125), Varita larga (Salta, Krapovickas & Schinini 28132), Ají huevo de gallo (Salta, Hilgert 1374); Bolivia: Ají (Santa Cruz, Williams 696; Tarija, Krapovickas & Schinini 39321), Aribibe (Santa Cruz, Hurtado 296), Aribibi (Santa Cruz, Heiser C281a, La Paz, Debouck et al. 3016), Ulupica (Tarija, Krapovickas & Schinini 31056), Aji acabeche (La Paz, Hinojosa & Wásra 1133), Ají amarillo (Cochabamba, Moscone 205), Aji Picante (Santa Cruz, Krapovickas & Schinini 32133), Locato largo (Santa Cruz, Heiser C290), Ají churcu, Ají rojo, Ají redondo, Pimentón colorado, Ají colorado gigantón (Santa Cruz, Libreros et al. 2014), Asta de buey amarillo, Huacareteño Duraznal, Huacareteño naranjo, Pa púca, Plomadita amarillo, Plomadita rojo, Chicotillo grueso, Huacarateño ancho amarillo, Cola de ratón amarillo, Cola de ratón naranjo (Chuquisaca, Libreros et al. 2014), Chicotillo, Huacareteño, Huacareteño amarillo, Ají amarillo, Ají colorado, Cumbaru rojo, Asta de buey, Asta de toro, Astay toro amarillo, Astay toro rojo, Astay toro anaranjado, Puntay lanza rojo, Puntay lanza anaranjado, Astay buey rojo (Chuquisaca, Jäger et al. 2013), Lata y Toro (Cochabamba, Moscone 209); Brazil: Pimenta (Rio de Janeiro, Krapovickas et al. 23428), Pimenta ardida (São Paulo, Alves de Paiva 01), Chifre de Veado (São Paulo, Heiser BGH 952), Dedo de moça (Paraná, Freire de Carvalho s.n.; Roraima, Barbosa et al. 2006), Pimenta de Passarinho (Espírito Santo, Mori et al. 11603), Pimenta dedo-de-moça (Amapá, Pereira et al. 1819; Minas Gerais, Vianna s.n.; Paraná, Leitão s.n.), Pimenta de cheiro amarela (Amapá, Pereira et al. 1830), Pimenta de cheiro vermelha (Amapá, Pereira & Severino 1851), Pimenta-unha-de velha (Río de Janeiro, Borges 66); Chile: Chota cabra (Santiago, Krapovickas 22232); Paraguay: Ají (Paraguarí, Galander 1877), Locotito (Paraguarí, Williams et al. 127), Locotito chico (Guairá, Williams et al. 121); Peru: Ají (San Martín, Belshaw 3210), Ají escabeche (Ancash, Francia 42); Amarillo Moquegua (Lima, Velarde Nuñez 6), Amarillo Tacna (Lima, Velarde Nuñez 23), Amarillo Trujillo (Lima, Velarde Nuñez 10), Cilindro amarillo (Lima, Velarde Nuñez 13), Cilindro colorado (Lima, Velarde Nuñez 15), Colorado Tacna (Lima, Velarde Nuñez 24), Escabeche amarillo (Lima, Velarde Nuñez 11), Escabeche intermedio (Lima, Velarde Nuñez 27), Escabeche Lurín (Lima, Velarde Nuñez 2), Escabeche Moquegua (Lima, Velarde Nuñez 3), Extra Lurín (Lima, Velarde Nuñez 21), Escabeche colorado ENA (Lima, Velarde Nuñez 1), Ají Amarillo, Ají escabeche (Bosland and Votava 2000), Ayuyo, Challuaruro (San Martín, Libreros et al. 2013), Ají uña de gallina (Lambayeque, Libreros et al. 2013).

Indigenous names

Argentina: Keuí (= picante) (Corrientes, Hunziker 7339); Bolivia: Kîî (Guaraní, Santa Cruz, Roca 0689); Paraguay: Ky y’ (Guaraní, Cordillera, Williams et al. 135), Pimenta í (Guaraní, Guairá, Williams et al. 121).

Uses

This domesticated variety, mostly known as ‘Ají’, ‘Ají amarillo’ or ‘Ají escabeche’, is an important component of the diet of the Bolivian and Peruvian native population, less so in Argentina, Brazil, Ecuador and Colombia. In Bolivia and Peru, many different pod types occur that differ both in morphological (shape, colour, size) and biochemical attributes (e.g. capsaicinoids, vitamin E, flavonoids and quercetin content and antioxidant capacity). These forms are consumed in regional cuisines as spices and vegetables, fresh or dehydrated and ground (Libreros et al. 2013, 2014).

Preliminary conservation assessment

EOO (11,296,813 km2); AOO (356 km2). Capsicum baccatum var. pendulum is a very widespread cultivated plant and can be assigned a category of Least Concern (LC).

Discussion

Capsicum baccatum var. pendulum is a member of the Baccatum clade (Carrizo García et al. 2016). Morphologically, the corollas are similar in shape and colour to those of its wild progenitor, but they are larger, reaching 20 mm in diameter at anthesis; more consistent differences are the position of the fruiting pedicels and fruits (pendent), fruits that vary in size, shape and colour, the number of seeds (up to 80 per fruit) and the larger seeds (3–5.2 mm long, 3–4 mm wide). The presence of heterostylous flowers is more frequent in var. pendulum than in var. baccatum (Barboza, pers. obs.). This is an uncommon character within Capsicum that has been observed in both domesticated (Perera and Poulos 1993; Peña-Yam et al. 2019) and wild species (C. benoistii and C. tovarii).

Many studies have been carried out that demonstrate the potential of this domesticated form in crop improvement. Capsicum baccatum var. pendulum encompasses a wide range of fruit morphology (e.g. fruit weight, size, shape and ripe colour), health-related compounds (flavonoids, polyphenols, quercetins, vitamin E, ascorbic acid, fat, amongst others) and capsaicinoids content (low to mild) (Rodríguez-Burruezo et al. 2009; Kollmannsberger et al. 2011; Libreros et al. 2013, 2014; van Zonneveld et al. 2015). Furthermore, a particularly strong vegetable-like aroma has been detected in the fruits, due to diverse volatiles in accessions from Ecuador and Peru, with additional fruity/exotic notes in some genotypes (Kollmannsberger et al. 2011).

Willdenow (1809) provided the description of C. pendulum, based on cultivated material of unknown provenance. In Willdenow’s Herbarium housed at B, we found a sheet with original material (B-W04431-01-0) consisting of a fruiting specimen with the inscription “C. pendulum, Hort. bot. Berol. W.”; this sheet is designated the lectotype.

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum baccatum L. var. umbilicatum (Vell.) Hunz. & Barboza, Kurtziana 26: 27. 1998.

Figs 29, 30

Capsicum umbilicatum Vell., Fl. Flumin.: 61. 1829. Type. Brazil. [Rio de Janeiro]: “Colitur hortis” (lectotype, designated by Knapp et al. 2015, pg. 825: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_010] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 7. 1831).

Type

Based on Capsicum umbilicatum Vell.

Description

Erect shrubs 1.50–2 m tall, with the main stem 1–1.5 cm in diameter at base, much branched from near the base, the branches spreading in a typical “zig-zag” appearance. Young stems 3–4-angled, fragile, green, mostly glabrous to sparsely pubescent with appressed-antrorse, short, simple, uniseriate, 4–5-celled, eglandular trichomes 0.3–0.6 mm long; nodes green or purple; bark of older stems green with light brown fissures, glabrous; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar in shape. Leaves membranous, concolorous or slightly discolorous, dark green above, light green beneath, glabrous to glabrescent adaxially and in margins with 5–6-celled eglandular trichomes and with long, spreading, 8–12-celled, eglandular trichomes along the primary veins and in the vein axils abaxially; blades of major leaves 5–14 cm long, 2.5–5 cm wide, ovate, the major veins 6–7 on each side of mid-vein, the base asymmetric and attenuate, the margins entire, the apex acute; petioles 2.5–7.5 (–9.5) cm long, sparsely pubescent; blades of minor leaves 4–4.5 cm long, 1.5–2.5 cm wide, ovate, the major veins 4–5 on each side of mid-vein, the base rounded, the apex acute; petioles 0.8–1.5 cm long, sparsely pubescent. Inflorescences axillary, 1–2 flowers per axil, rarely 3-flowered; flowering pedicels 25–35 mm long, angled, erect, geniculate at anthesis, rarely slightly curved and pendent, glabrescent, the trichomes short, antrorse; pedicels scars inconspicuous. Buds globose, white with greenish-yellow spots. Flowers 5–6-merous. Calyx 2–2.5 mm long, 3–3.8 mm wide, subequal, cup-shaped, thick, strongly 5–10-nerved, green, glabrescent to moderately pubescent with simple and some forked eglandular trichomes, the calyx appendages 5 or up to 8, 0.8–1.2 mm long, 0.2 mm wide, subequal, thick, erect, cylindrical, inserted close to the margin, with the same pubescence as calyx tube. Corolla 10–13 mm long, 12–14 mm in diameter, thick, white with greenish-yellow spots and a white centre outside and within, rotate-stellate with interpetalar membrane, lobed ⅓ or less than of the way to the base, pubescent adaxially with short glandular trichomes (stalk 1–3-celled; head globose, peltate, unicellular) in the throat and base of the lobes, glabrous abaxially, the tube 4 mm long, the lobes 5–6 mm long, ca. 3 mm wide, triangular or broadly triangular, spreading, sometimes with sparse eglandular trichomes abaxially, the margins finely ciliate, the tips acute, papillate. Stamens five, equal; filaments 2.5–2.7 long, white, inserted on the corolla 1–1.2 mm from the base, with auricles fused to the corolla at point of insertion; anthers 2.5–2.7 mm long, ellipsoid, yellow, not connivent at anthesis. Gynoecium with ovary 2–2.2 mm long, ca. 1.5 mm in diameter, 3 (–4)-carpelar, light green, ovoid; ovules more than two per locule; nectary ca. 1.2 mm tall, yellowish-green; styles dimorphic, short style 1.25–1.5 mm long, not exceeding the anthers length, long style 2.5–2.8 mm long, at the same level of the anthers or slightly exserted, white, cylindrical; stigma ca. 0.2 mm long, 0.6–0.7 mm wide, discoid, yellowish-green. Berry 25–40 mm long, 30–55 mm in diameter, campanulate-umbilicate, the base truncate or obtuse, the apex rounded, blunt or pointed, light green when immature, orange-red or bright red at maturity, persistent, pungent, the pericarp thick, opaque, without giant cells (endocarp smooth); stone cells absent; fruiting pedicels 25–35 mm long, pendent, sometimes strongly curved, strongly angled, widened distally, green; fruiting calyx 10–16 mm in diameter, persistent, slightly accrescent, slightly campanulate, thick, strongly nerved, green, the appendages 1–2 mm long, spreading or slightly recurved. Seeds 30–68 per fruit, 3.5–4.2 mm long, 3–4 mm wide, C-shaped, pale yellow to yellow, the seed coat slightly reticulate (SM), minutely reticulate (SEM), the cells polygonal to irregular in shape, the lateral walls straight to wavy; embryo imbricate.

Distribution

Capsicum baccatum var. umbilicatum is a cultigen and has been reported from Brazil, Colombia, Peru, Paraguay, Bolivia, Argentina and the Caribbean Islands (Fig. 27). The commercialisation of the fruits has increased enormously in different South American (Hunziker and Barboza 1998; Barboza, pers. obs.) and European markets (Hazen-Hammond 1993).

Ecology

Capsicum baccatum var. umbilicatum is a cultivated plant adapted to wet and semi-shaded places.

Phenology

Flowering from October to June; fruiting from December to July.

Chromosome number

2n = 2x = 24 (Moscone 1999; Moscone et al. 2003).

Common names

Argentina: Campanita (Salta, Barboza 164), Farolito, mitra (Distrito Federal, Melchiore s.n.); Brazil: Pimiento pitonga (Rio de Janeiro, Scaldaferro 57), Pimenta-de-cheiro-amarela (Amapá, Pereira et al. 1830), Chapéu-de-frade, Cabeça-de-frade, Chapéu-de-bispo, Pimenta-chapéu, Pimenta-de flor, Pimenta-de-cheiro (Roraima, Barbosa et al. 2006; São Paulo, Carvalho et al. 2006), Pimenta-chapeu-de-padre (São Paulo, Bernacci 2816), Pimentao fundo de garraba (Sturtevant 1888a); Jamaica and other Caribbean Islands: Jamaica hot (Miller and Harrison 1991); Peru: Rosasuchu (Andrews 1984); Western Hemisphere: Scallop pepper (Halsted et al. 1912), Rocotillo or Red squash (Andrews 1984).

Uses

The mildly hot fruits of this cultivated variety are valued for their use in dishes with tropical fruits, sauces, Caribbean fish stews, curries and chutneys (Miller and Harrison 1991) or fresh as a condiment (Andrews 1984). The fruiting plant is very showy and so it is also grown as an ornamental (Barboza, pers. obs.).

Preliminary conservation assessment

EOO (6,070,048 km2); AOO (64 km2). Capsicum baccatum var. umbilicatum is a cultigen in the Least Concern (LC) category.

Discussion

Capsicum baccatum var. umbilicatum belongs to the Baccatum clade (Carrizo García et al. 2016). Amongst the broad morphological variation of the fruits within the domesticated C. baccatum, this cultigen was described from eastern Brazil (Rio de Janeiro), based on its noteworthy and unusual morphotype, consisting in a campanulate-umbilicate, red or orange-red, pendent fruit (Hunziker and Barboza 1998), the character on which the original description was based (Vellozo 1829). The campanulate-umbilicate shape refers to the fruit having an apex with an acute protrusion arising from a central depression at the middle height of the fruit (Figs 29D, 30I, J). Due to this unusual fruit shape and bright red colour, this plant has been repeatedly discussed in the chili pepper literature. After Vellozo (1829), Sendtner (1846) mentioned C. umbilicatum as a synonym of C. annuum var. grossum. Then Irish (1898) submerged it under the ‘Red Wrinkled’ cultivar of C. annuum. Other authors illustrated and named this taxon as “Scallop pepper” (Halsted et al. 1912, Lam. 19, fig. 2) or as C. chinense cv. ‘rocotillo’ (Andrews 1984, pl. 31), respectively. It was Sturtevant (1888a) who provided a detailed description for this cultigen, based on plants obtained from seeds from Rio de Janeiro (Brazil) and stated that it was “an extremely well marked variety”. Hunziker and Barboza also grew plants of this cultivar from seeds from Paraguay and observed that the flowers were identical to the Andean domesticated C. baccatum populations. Hunziker and Barboza (1998) considered that the fruit shape was a particular variant within this taxon and proposed the name C. baccatum var. umbilicatum. In addition to the shape of the fruits, the typical giant cells of the pericarp, a derived character present in most of the Capsicum species (Carrizo García et al. 2016), are missing in this cultivar. In addition, the presence of sclereids in the epicarp and endocarp distinguish this variety from the wild and other domesticated C. baccatum (Hunziker and Barboza 1998 and see Suppl. material 1: Appendix 1). Although molecular studies (Albrecht et al. 2012b; Carrizo García et al. 2016; Scaldaferro et al. 2019) demonstrated the close affinities between domesticated lines of Capsicum baccatum, they were not sufficiently conclusive to support combining the domesticated varieties into one entity. Therefore, for those individuals that fit this distinctive and unusual fruit morphology and anatomy, we prefer to maintain the name C. baccatum var. umbilicatum.

Figure 29. 

Capsicum baccatum var. umbilicatum A reproductive branch B flower C section of the calyx showing the venation D, E eglandular trichomes of the abaxial surface of the calyx F glandular trichome of the adaxial surface of the calyx G sector of opened corolla H eglandular trichome of the corolla lobes I, J anthers, in dorsal and ventral views, respectively K gynoecium with long style L gynoecium with short style M ovary trilocular, in cross section N seed O seed, in cross section P structure of seed coat at the seed margin Q structure of seed coat at the seed body R embryo. From Rodríguez s.n. (CORD 241). Drawn by N. de Flury. Published in Hunziker (1998), reproduced with permission.

Figure 30. 

Capsicum baccatum var. umbilicatum A plant B flower bud C flower, lateral view D flower short-styled E flower long-styled F flower hexamerous G flower, seen from behind H immature fruit I, J mature fruits A–I from Barboza 5163, photos by G.E. Barboza J from Carrizo García 101, photo by C. Carrizo García.

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum benoistii Hunz. ex Barboza, PLoS ONE 14(1): 4. 2019.

Fig. 31

Type

Ecuador. Tungurahua: Baños, 3 Apr 1931, M.R. Benoist 4204 (holotype: P [P04023406]).

Description

Erect shrubs with few branches. Young stems 3-angled, light brown, glabrous or sparsely pubescent with appressed-antrorse, simple, uniseriate, 3–5-celled, eglandular trichomes 0.3–1.1 mm long; bark of older stems dark brown, angled, glabrous; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair markedly unequal in size, subequal in shape. Leaves membranous, discolorous, dark green above, light green beneath, glabrous or with sparse trichomes adaxially and abaxially, similar to those of the stems, more abundant on main veins; blades of major leaves 8.5–12 cm long, 2.8–6 cm wide, ovate or elliptic, the major veins 4–5 (–6) on each side of mid-vein, the base asymmetric and attenuate, the margins entire, the apex long-acuminate; petioles 0.5–1 cm long, glabrous or glabrescent; blades of minor leaves 2.4–6 cm long, 1.7–4 cm wide, ovate or elliptic, the major veins 3–4 on each side of mid-vein, the base asymmetric and rounded, the margins entire, the apex acute or rounded; petioles 0.1–1 cm long, glabrescent or sparsely pubescent. Inflorescences axillary, 3–6 flowers per axil; flowering pedicels 13–20 mm long, angled, filiform, pendent, non-geniculate at anthesis, moderately to densely pubescent, the eglandular trichomes long, spreading to antrorse; pedicels scars inconspicuous. Buds ovoid, colour unknown. Flowers 5-merous. Calyx 2–2.5 mm long, ca. 5 mm wide, cup-shaped, thick, colour unknown, moderately pubescent with the same trichomes as pedicels, the calyx appendages 5, 2.5–3.5 mm long, ca. 0.5 mm wide, equal or subequal, thick, erect, subulate, inserted close to the margin, with the same pubscence as calyx tube. Corolla ca. 12–13 mm long, thick, deeply stellate without interpetalar membrane, lobed nearly to the base, glabrous adaxially and abaxially, the tube ca. 3 mm long, the lobes ca. 9 mm long, ca. 2 mm wide, narrowly triangular, erect, the margins and the tips pubescent. Stamens five, equal; filaments 3–3.2 mm long, inserted on the corolla 1.5 mm from the base, with auricles fused to the corolla at the point of insertion; anthers ca. 3 mm long, ellipsoid, not connivent at anthesis. Gynoecium with ovary 1.3–1.7 mm long, 1.2–1.5 mm in diameter, subglobose; ovules more than two per locule; nectary ca. 0.3 mm tall; styles dimorphic, long style ca. 6.5 mm, short style ca. 3.6 mm, clavate; stigma 0.3 mm long, 0.5 mm wide, globose. Berry and seeds unknown.

Figure 31. 

Capsicum benoistii A flowering branch B flower C calyx D section of the calyx showing the venation E eglandular trichome of the calyx F opened corolla G gynoecium with long style H gynoecium with short style. From Benoist 4204. Drawn by N. de Flury. Published in Barboza et al. (2019), reproduced with permission.

Distribution

Capsicum benoistii is endemic to a restricted area in central-southern Ecuador (Provinces of Tungurahua and Loja) (Fig. 32).

Figure 32. 

Distribution of C. benoistii, C. caatingae, C. caballeroi, C. campylopodium and C. cardenasii

Ecology

Capsicum benoistii grows in thickets in montane forests, between 1,500 and 2,600 m elevation.

Phenology

Flowering from March to May. Fruiting time unknown.

Chromosome number

Not known.

Common names

None recorded.

Uses

None recorded.

Preliminary conservation assessment

EOO (2,627.651 km2); AOO (12 km2). Considering the extent of occurrence, the area of occupancy, the few localities (3) where it was collected and the decline observed in its geographic range, we assign C. benoistii the Endangered (EN; B1+2ab(i,ii) category. The species has not been collected since 1978 despite recent intensive field explorations in the same locations (Barboza et al. 2019).

Discussion

The affinities of C. benoistii have not yet been explored and, due to the lack of data on some morphological characters, it is not assigned to any of the recognised clades (Barboza et al. 2019). This species is poorly known and information about corolla colour, fruit and seed characters and chromosome number are still lacking. However, C. benoistii is distinctive in its deeply-lobed stellate corolla (lobes three times longer than the tube, Fig. 31B, F) and in the presence of heterostylous flowers (Fig. 31G, H). These features, plus the short flowering pedicels (13–20 mm long), distinguish C. benoistii from C. geminifolium, a morphological similar species, which has campanulate corollas that are lobed less than 1/3 of the way to the base and homostylous flowers.

The presence of heterostylous flowers in C. benoistii is unusual amongst Capsicum species. It has been reported in C. chinense (Peña-Yam et al. 2019), C. baccatum (Hunziker 2001; Barboza 2013), in many cultivars (Perera and Poulos 1993) and observed in C. pubescens (see Fig. 103I–K).

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum caatingae Barboza & Agra, Syst. Bot. 36 (3): 769. 2011.

Figs 33, 34

Type

Brazil. Bahía: Cachoeira, Estação da EMPASA, Vale dos Rios Paraguaçu e Jacuipe, 12°32'39"S, 39°05'00"W, 40–120 m elev., Jun 1980, P. do Cavalo et al. 162 (holotype: HUEFS [HUESF000001216, acc. # 00920]; isotypes: ALCB [ALCB000131, acc. # 07938], RB [RB00461411, acc. # 263323]).

Description

Small trees or erect shrubs 2–4 (–6) m tall, the main stem thick, 2.5–5 cm in diameter at base and with indefinite growth up to 5 m high, few branched, the branches slender or scandent. Young stems 3–4-angled, rigid, grey, glabrescent or, more rarely, sparsely pubescent with antrorse, simple, uniseriate, 3–7-celled, eglandular trichomes 0.2–0.9 mm long, sometimes furcate trichomes 0.7–0.9 mm long or minute, simple, glandular trichomes, ca. 0.1 mm long; nodes solid, green; bark of older stems grey or brown, glabrous; lenticels light brown. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size and similar or dissimilar in shape. Leaves membranous or papery, slightly discolorous, glabrescent to moderately pubescent on both sides, with antrorse, 5–8-celled, eglandular trichomes 0.4–1.2 mm long, sometimes with branched trichomes 0.7–0.9 mm long and small glandular hairs (stalk short, 2–3-celled; head multicellular or unicellular), especially on veins abaxially; blades of major leaves 4–11.5 (–20) cm long, 1.5–2.4 (–8.5) cm wide, ovate to elliptic, the major veins 4–5 on each side of mid-vein, the base unequal and short-attenuate, the margins entire, the apex acute or somewhat acuminate; petioles (0.5–) 0.7–2.5 cm long, moderately pubescent adaxially; blades of minor leaves 1.5–2 cm long, 0.7–1.3 cm wide, ovate, the major veins 3–4 on each side of mid-vein, the base short-attenuate or truncate, the margins entire, the apex acute; petioles 0.2–0.5 cm long, moderately pubescent adaxially. Inflorescences axillary, congested, 5–13 (–20 or more) flowers per axil; flowering pedicels 7–21 (–28) mm long, terete, pendent, non-geniculate at anthesis, green with violet tones, glabrescent to moderately pubescent, the eglandular trichomes short, antrorse; pedicels scars conspicuous, somewhat corky. Buds globose to ellipsoid, cream, greenish-white or purple at the apex. Flowers 5-merous. Calyx 1.2–1.7 (–2) mm long, 2–2.5 mm wide, cup-shaped, circular in outline, thin, weakly 5-nerved, green, greenish-purple or purple, sparsely pubescent with the same antrorse eglandular and glandular trichomes of the leaves, without appendages. Corolla 4.5–6 (–8) mm long, lilac or purple, greenish-yellow at the base outside, with different tones of purple and a narrow white marginal band in the lobes and greenish-yellow to yellowish-white centre within, stellate with narrow interpetalar membrane, lobed nearly halfway to the base, pubescent adaxially with a continuous ring of small glandular trichomes (stalk short, 1–2-celled; head globose, unicellular) in the throat and base of the lobes, glabrous abaxially, the tube 2.8–3.2 mm long, the lobes 2.9–3.5 mm long, 1.7–2.4 mm wide, broadly triangular, spreading, the margins involute and finely ciliate, the tip cucullate, papillate. Stamens five, equal; filaments (0.8–)1.1–1.75 mm long, greenish-white or white, inserted on the corolla ca. 2 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.4–2.1 mm long, ellipsoid, light green or yellow, not connivent at anthesis. Gynoecium with ovary 1.1–1.4 in diameter, pale green, subglobose; ovules more than two per locule; nectary ca. 0.3–0.4 mm tall; styles homomorphic, (4.3–) 4.6–4.8 mm, exserted ca. 1 mm beyond the anthers, pale yellow or cream, clavate; stigma ca. 0.6 mm in diameter, globose or 0.15 mm long, 0.6 mm wide, discoid, light green. Berry (5–) 7–11 mm in diameter, globose, slightly flattened at the apex, green or yellowish-green when immature, red at maturity, deciduous, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels (15–) 20–25 mm long, pendent, terete, inflated, strongly widened distally and with a constriction at the junction with the calyx, green or purple; fruiting calyx 3–4 mm in diameter, persistent, not accrescent, discoid, 5 (–10)-nerved, the margin entire or sometimes easily torn, green or purple. Seeds (9–) 11–17 per fruit, 3.2–3.7 mm long, 2.2–2.8 mm wide, C-shaped, pale yellow, the seed coat reticulate and tuberculate at margins (SM), cerebelloid (SEM), the cells irregular in shape, the lateral walls wavy at margins and strongly sinuate in the central zone; embryo imbricate.

Distribution

Capsicum caatingae is endemic to the north-eastern States of Brazil (Alagoas, Bahia, Pernambuco Sergipe and northern Minas Gerais States, Fig. 32). A beautiful specimen has been cultivated on the campus of the Federal University of Viçosa (Minas Gerais) for more than 30 years (seeds from Bahia).

Ecology

Capsicum caatingae is a xerophytic species usually found in the margins of arid open Caatinga forests (Seasonal Deciduous and Seasonal Residual Forests) and in the anthropogenic Caatinga, more rarely in degraded humid forests (Floresta Estacional Decidual or Semidecidual). It is quite common in savannahs or amongst granitic and gneissic outcrops (‘inselbergs’), growing with thorny, deciduous arboreal and shrubby vegetation, between 100 and 950 m elevation.

Phenology

Flowering from December to August; fruiting from February to October.

Chromosome number

n = 12 (Pozzobon and Schifino-Wittmann 2006, as C. parvifolium); 2n = 2x = 24 (Moscone 1993; Pozzobon et al. 2006; Moscone et al. 2007; all count as C. parvifolium).

Common names

Brazil. Caraibera (Alagoas, Silva & Moura 1586), Murta (Sergipe, Silva 287), Pimenta brava (Bahía, Pinto & Bautista 104), Semente-de-macaco (Alagoas, Oliveira 7).

Uses

None recorded.

Preliminary conservation assessment

EOO (270,442.695 km2); AOO (276 km2). The large extent of occurrence and the number of localities where C. caatingae was collected indicate Least Concern (LC) category. Given its highly specialised habitat limited to the Caatinga Biome, some subpopulations may be adversely affected because deforestation has intensified rapidly in recent years due to the consumption of native firewood for domestic and industrial purposes, over-grazing and changes in the ecosystem due to pasture and agricultural expansion (MMA-Brazil 2020).

Discussion

Capsicum caatingae is a member of the Caatinga Clade (Carrizo García et al. 2016) and is closely related to C. parvifolium, with which it is sympatric in Brazil; the two species have been confused in literature (Moscone 1993; Hunziker 2001; Barboza and Bianchetti 2005; Pozzobon and Schifino-Wittmann 2006; Pozzobon et al. 2006; Moscone et al. 2007) and in herbaria (Barboza, pers. obs.). Barboza et al. (2011) clarified this confusion and re-circumscribed C. parvifolium.

Figure 33. 

Capsicum caatingae A flowering branch B branched trichome from leaf C flower D calyx E, F, G glandular trichomes of the calyx and leaf venation H eglandular trichome of the calyx I sector of opened corolla J eglandular trichome of the corolla lobes K gynoecium L ovary, in cross section M node of a fruiting branch N fruit, in cross section O seed P seed, in cross section Q embryo. From Hunziker 25233. Drawn by N. de Flury. Published in Barboza et al. (2011), reproduced with permission.

Capsicum caatingae is an unusual species with a combination of uncommon features rarely found amongst its congeners: arborescent habit, with indefinite growth of the main stem reaching up to 6 m high (15 m fide Bautista & Pinto 1023), very congested inflorescences with up to 20 or more flowers per node, corolla with varied colours (lobes white edged, then deep purple or variations of purple colour, tube with greenish-yellow to yellowish-white centre) and terete and inflated fruiting pedicels with a strong annular constriction at the junction with the calyx base (Fig. 34), similar to C. chinense and a few other species (C. minutiflorum, C. lanceolatum and C. regale).

Figure 34. 

Capsicum caatingae A flowering branch B young fruiting branch C immature fruits (note the purple pedicels) D node of a fruiting branch, some fruits already fallen down. No specimen voucher. Photos taken in Federal University of Viçosa (Minas Gerais) by C. dal Zovo (Associazione PepperFriends).

Capsicum caatingae differs from C. parvifolium in the absence of calyx appendages (five appendages in C. parvifolium), the number of flowers per node (up to 20 or more flowers vs. not more than seven flowers in C. parvifolium) and the fruit and seed colour (red berry with pale yellow seeds vs. greenish-golden yellow translucent berry with brownish-black seeds in C. parvifolium). Capsicum caatingae is sometimes sympatric with C. longidentatum from which it differs by its arborescent growth (vs. shrubby growth), the indumentum mostly of simple trichomes (vs. indumentum of branched and dendritic trichomes), the lack of calyx appendages (vs. five, rarely six, calyx appendages), the mostly purple corolla (vs. corolla white with greenish-yellow spots) and the pungent red fruit with pale yellow seeds (vs. non-pungent probably yellowish-green fruit with brown to brownish-black seeds).

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum caballeroi M.Nee, Brittonia 58 (4): 323. 2006.

Figs 35, 36

Type

Bolivia. Santa Cruz. Prov. Caballero: Parque Nacional Amboró, Cerro Bravo, 10 km al N de Comarapa, 17°49.5'S, 64°32.5'W, 2400–2500 m elev., 7–10 Apr 1994, I. Vargas C. & J.M. Camacho 3118 (holotype: USZ; isotypes: CORD [CORD00003917], MO [MO-1921597, acc. # 5959888], NY [00745836], US [00902045, acc. # 3520370]).

Description

Erect shrubs or more rarely small trees 0.70–5 (–7) m tall, with the main stem thick, 2.5–3 cm in diameter at base, few to much branched above. Young stems angled, rigid, green, glabrous to sparsely pubescent, with appressed-antrorse, simple, uniseriate, 3–4-celled, eglandular trichomes 0.07–0.2 mm long; nodes solid, green; bark of older stems brown or brownish-grey, glabrous; lenticels few, light brown. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar or dissimilar in shape. Leaves coriaceous, slightly discolorous, glabrous on both sides or sparsely pubescent along the mid-vein abaxially, with simple, eglandular trichomes 0.2–0.4 mm long; blades of major leaves 4–13.5 cm long, 1.5–4 cm wide, elliptic, the major veins 4–5 on each side of mid-vein, the base acute to short-attenuate, the margins entire slightly revolute, the apex acuminate; petioles 0.2–0.8 cm long, glabrous; blades of minor leaves 2–4.5 cm long, 1–2 cm wide, elliptic or ovate, major veins 2–3 on each side of mid-vein, the base short-attenuate, the margin entire slightly revolute, the apex acute; petioles 0.3–0.5 cm long, glabrous. Inflorescences axillary, 1–2 flowers per axil; flowering pedicels 20–40 (–50) mm long, terete, slightly pendent to pendent, non-geniculate at anthesis, green, glabrous; pedicels scars inconspicuous. Buds ellipsoid, yellow. Flowers 5-merous. Calyx 1.5–3 mm long, ca. 3 mm wide, cup-shaped, thick, pale green, sparsely pubescent with the same antrorse eglandular trichomes of the young stems, the calyx appendages 10, unequal, the five main appendages 1–3.2 mm long, the five secondary appendages 0.8–2 mm long, erect, subulate, inserted close to the margin. Corolla (10–) 11–14 (–18) mm long, 4–6 mm in diameter, thick, entirely pure yellow or lemon-yellow, campanulate without interpetalar membrane, shallowly 5-lobed; glabrous adaxially and abaxially, the tube 7–10 mm long, the lobes 3–4 mm long, ca. 2 mm wide, narrowly triangular, recurved, the margins involute and finely ciliate, the tips deeply acute, papillate. Stamens five, equal; filaments 4–6 mm long, cream, inserted on the corolla 1–2 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 2–3 mm long, ellipsoid, yellow, not connivent at anthesis. Gynoecium with ovary ca. 2 mm long, 1.5 mm in diameter, pale green, ovoid; ovules more than two per locule; nectary ca. 0.4–0.5 mm tall; styles homomorphic, 7–9 mm, scarcely exserted beyond the anthers, cream, clavate; stigma ca. 1 mm in diameter, whitish, capitate. Berry (9–) 10–16 mm in diameter, globose, slightly flattened at the apex, pale green or white when immature, bright red at maturity, persistent, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 25–50 mm long, pendent, curved, terete, strongly widened distally and with a constriction at the junction with the calyx, green; fruiting calyx 3–5 mm in diameter, persistent, not accrescent, discoid, green, the appendages (1–) 3–5 mm long, appressed to the berry. Seeds 5–21 per fruit, 3–4 (–5) mm long, 3.8–5 mm wide, C-shaped or teardrop-shaped, pale yellow or nearly white, the seed coat smooth and reticulate at margins (SM), cerebelloid-reticulate (SEM), the cells irregular and polygonal in shape, the lateral walls strongly sinuate to nearly straight at the seed margin; embryo imbricate or coiled.

Distribution

Capsicum caballeroi is an endemic species from central Bolivia (Santa Cruz, Cochabamba and Chuquisaca Departments, Fig. 32).

Ecology

Capsicum caballeroi is a rare element of montane cloud forests (Yungas and Bosque Tucumano-Boliviano Montano), found in very moist shaded wooded quebradas or at the margin of the forest between 1,000 and 2,600 m elevation.

Phenology

Flowering and fruiting probably all year long; a peak of flowering was observed in November to early January and fruiting from January to June.

Chromosome number

2n = 2x = 24 (Barboza et al. 3655, see Table 2).

Common names

Bolivia. Aribibi (Cochabamba, Fernández T. et al. 2007), Ají de monte (Santa Cruz, Vargas C. & Prado 1282), Ulupica de yunga (Santa Cruz, Vargas C. et al. 1343).

Uses

None recorded.

Preliminary conservation assessment

EOO (14,005.388 km2); AOO (84 km2). Capsicum caballeroi grows mostly in the cloud forest of the Amboró National Park and peripheral areas where the population consists of few individuals; although found in a relatively large geographical area, we observed a severe decline of both the EOO and the AOO of this species due to the continuing indiscriminate deforestation occurring in the last years; in this way, we consider C. caballeroi under threat and assign the Vulnerable category (VU; B1ab(ii,iii)).

Figure 35. 

Capsicum caballeroi A fruiting branch B flower C calyx D opened corolla E gynoecium F fruit G seed H seed, in longitudinal section. From Wood 11102. Drawn by P. Peralta.

Discussion

Capsicum caballeroi belongs to the Bolivian clade (Carrizo García et al. 2016; Barboza et al. 2019). It is easily distinguishable from any other Bolivian species in its calyx with 10 unequal appendages, its yellow tubular-campanulate pendent corollas, its long flowering and fruiting pedicels and its large, bright red fruits and pale seeds (Fig. 36D, I).

Figure 36. 

Capsicum caballeroi A plant B leaf pairs C major leaf D flower bud E flower F flower, in front view G immature fruit H mature fruit I mature fruit, in cross section, showing the seeds A from Barboza et al. 4907 B, C, E, F, H Barboza et al. 3655 D, G, I from Barboza et al. 4908. Photos by G.E. Barboza and S. Leiva González.

Capsicum caballeroi is morphologically most similar to C. piuranum, an endemic species from northern Peru and can be distinguished from that species in its calyx (green calyx with 10 unequal linear appendages vs. purple or greenish-purple calyx with five equal subulate appendages), its corolla lobes (recurved vs. spreading) and its fruits and seeds (red pungent fruits with pale yellow or nearly white seeds vs. orange to red non-pungent fruits with dark brown to black seeds). The fruits and the seeds of C. caballeroi are larger than those of C. piuranum. Capsicum caballeroi is sympatric with C. minutiflorum; they share yellow corollas and red fruits, but are distinguished by the coriaceous leaves, calyx with 10 unequal appendages, campanulate and larger corollas (10–14 mm long) and fruits (9–16 mm in diameter) of C. caballeroi vs. the membranous leaves, calyx with five equal or subequal appendages, stellate, smaller corollas (6.5–8.5 mm long) and fruits (7–10 mm in diameter) of C. minutiflorum (Fig. 87).

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum campylopodium Sendtn., Fl. Bras. (Martius) 10: 144. 1846.

Figs 37, 38

Capsicum gracilipes Dunal, Prodr. [A. P. de Candolle] 13(1): 418. 1852. Type. [Brazil]. Rio de Janeiro, 1834, C. Gaudichaud 513 (lectotype, designated here: G-DC [G00131901]; isolectotypes: F [F neg. F0BN002869], MPU [MPU013436 fragment], P [P00410015, P00410016]).

Capsicum salicifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 418. 1852. Type. Brazil. Rio de Janeiro: “In provinciã Rio de Janeiro, Serra dos Órgãos”, Oct. 1833, A.-C. Vauthier 528 (lectotype, designated here: G-DC [G00131881]; isolectotypes: CORD [CORD00006953], F [F neg. 6846 ex G-DC + F0093724F fragment, acc. # 644821], GH [GH00077007], MPU [MPU023040 fragment], P [P00410013, P00410014]).

Type

Brazil. “Brasilia”, [no date], F. Sellow 6 (lectotype, designated by Barboza 2011, pg. 29: P [P00410022]; isolectotypes: B [destroyed, F neg. 2865], CORD [CORD00006952, fragment of lectotype], K [K000585891 right plant], F [v0076865F, acc. # 648993, fragment of holotype], LE [LE01072483]).

Description

Erect subshrubs or shrubs, 0.5–2 m tall, with the main stem thick, up to 2.5 cm in diameter at base, much branched above, the branches dichotomously spreading in a typical “zig-zag” appearance. Young stems striate, fragile, green, glabrous to glabrescent, with antrorse, curved, simple, uniseriate, 3–4-celled eglandular trichomes 0.3–0.5 mm long; nodes solid, green; bark of older stems dark brown, glabrous; lenticels few, light brown. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar in shape. Leaves membranous, slightly discolorous, green or dark green above, light green beneath, glabrous or sparsely pubescent with appressed-antrorse, 3–5-celled, eglandular trichomes 0.2–0.4 mm long on both surfaces; blades of major leaves 4–11.5 (–20) cm long, 1.5–2.4 (–8.5) cm wide, elliptic to ovate, the major veins 5–7 on each side of mid-vein, the base attenuate and unequal, the margins entire, the apex acuminate to long-acuminate; petioles 0.5–1.7 cm, glabrescent or glabrous; blades of minor leaves 1.7–3.5 (–4.5) cm long, 0.5–2 cm wide, elliptic to ovate, the major veins 3–4 on each side of mid-vein, the base attenuate, the margins entire, the apex acute; petioles 0–0.5 cm, glabrous. Inflorescences axillary, 2–5 (–7) flowers per axil, rarely flowers solitary; flowering pedicels 9–14 mm, very thin, delicate, terete to slightly striate, erect to slightly spreading, geniculate at anthesis, entirely green or reddish basally, glabrescent, the eglandular trichomes short, antrorse; pedicels scars inconspicuous. Buds ovoid, cream or with greenish-yellow spots. Flowers 5-merous. Calyx (1–) 1.2–1.6 mm long, 1.4–1.5 mm wide, hemispherical, circular in outline, very thin, green, glabrous or rarely glabrescent, without appendages. Corolla 4.5–6.5 (–8) mm long, (6–) 6.4–7.5 (–11) mm in diameter, mostly cream outside, white or cream with two large golden yellow or ochraceous spots on each lobe and part of the limb and cream or white centre within, stellate with narrow interpetalar membrane, lobed more than 1/3 to nearly halfway to the base, pubescent adaxially with a continuous ring of glandular trichomes (stalk long, 2–3-celled; head globose, peltate, unicellular) in the throat and base of the lobes, glabrous abaxially, the tube 2–4 mm long, the lobes 1.8–3.5 (–4.3) mm long, 2–4 mm wide, triangular or broadly triangular, spreading, the margins finely ciliate, the tips acute, cucullate, papillate. Stamens five, unequal in length; three filaments short 1.5–2.3 mm long, the two longer 1.9–3 mm long, white or cream, inserted on the corolla 0.6–1 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 0.8–1.7 mm, ellipsoid, yellow, not connivent at anthesis. Gynoecium with ovary 0.7–1 mm long, 0.9–1.3 mm in diameter, light green, ovoid; ovules two per locule; nectary ca. 2.2 mm tall; styles homomorphic, 2.3–4 mm, somewhat exserted beyond the anthers, cream, clavate, slightly curved; stigma 0.1–0.2 mm long, 0.4 mm wide, discoid, pale green. Berry 3–5 mm long, 5–7 mm in diameter, globose-depressed, green when immature, greenish-golden yellow at maturity, deciduous, pungent, the pericarp thin, translucent, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 14–25 mm, pendent, angled, slightly widened distally, green; fruiting calyx 3–4.5 mm in diameter, persistent, not accrescent, discoid, green. Seeds 4 (–6) per fruit, 3.7–3.9 (–4) mm long, 3–3.3 mm wide, C-shaped or reniform, brownish-black, the seed coat smooth or faintly reticulate and tuberculate at margins (SM), reticulate-cerebelloid with pillar-like outgrowths at margins (SEM), the cells rectangular or polygonal at margin and irregular in seed body, the lateral walls straight to wavy at margins and sinuate in the central zone; embryo imbricate.

Distribution

Capsicum campylopodium is an endemic species from south-eastern Brazil (Rio de Janeiro, Minas Gerais and Espírito Santo States, Fig. 32).

Ecology

Capsicum campylopodium is a typical component of the coastal Atlantic Forest (Mata Atlântica) and of some remnants of interior forests of the same biome. It is found in small colonies of a few individuals in shady or semi-shady places, sometimes also in sun, along roadsides or trails of the Ombrophilous Forest (Floresta Ombrófila Densa Submontana and Montana), between 100 and 1,200 m elevation.

Phenology

Flowering from late September to March; fruiting from December to April.

Chromosome number

n = 13 (Pozzobon and Schifino-Wittmann 2006), 2n = 2x = 26 (Moscone et al. 2003, 2007; Pozzobon et al. 2006).

Common names

Brazil. Pimenta da Serra (Rio de Janeiro, Kuhlmann 6288).

Uses

None recorded.

Preliminary conservation assessment

EOO (58,586.312 km2); AOO (212 km2). Capsicum campylopodium is a relatively widespread species that occurs in many formally protected areas, such as Parque Nacional da Tijuca, Parque Estadual da Pedra Branca, Estação Ecologica Estadual de Paraiso, Reserva Ecologica de Rio das Pedras, Parque Municipal Ecológico da Prainha, Estação Biológica Caratinga, amongst others (see Suppl. material 4: Appendix 4). Based on the extent of occurrence and the number of localities (ca. 50), we assign a category of Least Concern (LC). However, most collections are from a much-visited place in Rio de Janeiro (Parque Nacional da Tijuca) and from Serra dos Órgãos, with a serious problem of forest fragmentation in some areas (Freitas et al. 2006; Nehren et al. 2009) which may represent a threat to this species.

Discussion

Capsicum campylopodium belongs to the Atlantic Forest clade (Carrizo García et al. 2016). It is characterised by its small flowers, hemispherical calyx with a circular outline, white corolla with golden yellow or ochraceous spots, locules with only two ovules and depressed fruits with not more than four seeds (Fig. 38). In herbarium material, the morphology of the fruits is not always preserved. However, in living material, the number of seeds per locule (2), the size of the seeds (large in proportion to the size of the fruit) and the arrangement of the seeds (parallel, but not touching) results in a slightly depressed fruit, rather than spherical (Figs 37I, J, 38C), which is a more common fruit shape in other species.

Figure 37. 

Capsicum campylopodium A flowering branch B eglandular trichome of the leaf C flower D calyx E section of the calyx showing the venation F flower, upper view G sector of opened corolla H gynoecium I fruit J fruit, upper view K fruit, in cross section L anatomical detail of the pericarp (note the giant cell in the mesocarp) M seed N seed, in cross section O structure of seed coat at the seed margin P structure of seed coat at the seed body Q embryo. From Hunziker 25116. Drawn by L. Sánchez. Published in Hunziker (2001), reproduced with permission.

Figure 38. 

Capsicum campylopodium A plant B flower C immature fruit D–F diagrams of different stages of fruit development D ovary, in cross section, showing the locules and the number of ovules E young fruit, in cross section (the lateral arrows indicate the fruit is flattened around the centre) F mature depressed fruit (one locule), in longitudinal section, showing the two seeds occupying the whole locule A from Barboza et al. 2057, photo by G.E. Barboza B, C from Bianchetti et al. 511, photos by L. Bianchetti.

Capsicum flexuosum, C. schottianum and C. campylopodium all lack calyx appendages and are sometimes extremely difficult to distinguish from one another due to some characters being poorly preserved in herbarium specimens. Capsicum campylopodium and C. schottianum both have clearly geniculate pedicels, a calyx with five evident nerves and greenish-golden yellow fruits with brownish-black to black seeds. The distinction of C. campylopodium from C. schottianum is based on calyx shape and size (hemispherical and ≤ 1.5 mm in diameter vs. cup-shaped and > 2 mm in diameter in C. schottianum), corolla colour (mostly white with large golden yellow spots within vs. white usually with purple and greenish-yellow spots within), fruit shape (globose-depressed vs. globose or subglobose) and number of seeds (four, very rarely six vs. ≥ six). The separation of C. flexuosum, the most distinctive species of the three, is based primarily on its lack of geniculate pedicels and its having a calyx with ten evident nerves, white corolla with yellowish-green spots within and red fruits (Fig. 63).

Dunal (1852) described C. gracilipes, based on three Gaudichaud specimens citing “v.s. in h. DC et h. Mus. Paris”; these are now housed at G-DC (G00131901) and P (P00410015, P00410016). All three collections are fruiting specimens, the one at G-DC (G00131901) is the best-preserved and here is designated lectotype.

For C. salicifolium, Dunal (1852) mentioned in the protologue the Vauthier 528 collection he saw in G-DC and P. We found these specimens and other duplicates. We select the best-preserved specimen (G00131881) as the lectotype.

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum carassense Barboza & Bianch., PhytoKeys 140: 127. 2020.

Figs 39, 40

Type

Brazil. Minas Gerais: Catas Altas, RPPN Serra do Caraça, trilha da gruta de Lourdes, após a capelinha, 20°05'41"S, 43°28'52"W, 1386 m elev., 26 Oct 2014, J.R. Stehmann, L.L. Giacomin, G.E. Barboza & S. Knapp 6347 (holotype [two sheets]: BHCB acc.#174038 [BHCB0019940_1, BHCB0019940_2]; isotypes: CORD [CORD00006968], RB [RB01220059, acc. # 674586], MBM).

Description

Erect shrubs (0.8–) 1–2 (–3) m tall, with the main stem somewhat thick and sparsely branched, the branches dichotomous and spreading horizontally. Young stems 3–4-angled, fragile, green, moderately to densely pubescent with uncinate and antrorse, simple, uniseriate, 3–5 (–6)-celled, eglandular trichomes 0.2–0.7 mm long, yellowish-brown when dried; nodes green or purple; bark of older stems brown, striate, pubescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar in shape. Leaves membranous to chartaceous, discolorous, dark green above, paler beneath, moderately pubescent especially on the veins, with simple trichomes like those of the stem and sparse or frequent glandular trichomes (stalk unicellular; head multicellular) adaxially and abaxially; blades of major leaves 6–16 cm long, 0.9–2.5 cm wide, narrowly elliptic to lanceolate, the major veins 6–8 on each side of mid-vein, the mid-vein prominent and the secondary veins obscure, the base attenuate, the margins entire, the apex acute to obtuse; petioles 0.2–0.6 cm long, moderately pubescent; blades of minor leaves 2.9–3.9 cm long, 0.5–0.8 cm wide, narrowly elliptic, the major veins 2–3 (–4) on each side of mid-vein, the base attenuate, the margins entire, the apex obtuse; petioles 0.2–0.4 cm long, moderately pubescent. Inflorescences axillary, 2–4 flowers per axil; flowering pedicels (12–) 15–20 (–22) mm long, slightly angled, erect to spreading, geniculate at anthesis, green, moderately pubescent, the eglandular trichomes short or long, antrorse to spreading; pedicels scars inconspicuous. Buds ellipsoid, cream with greenish-yellow spots. Flowers 5-merous. Calyx 1.2–1.6 mm long, 2.5–3 mm wide, cup-shaped, thin, light green to cream, moderately pubescent with antrorse, curved, 3–5-celled, eglandular trichomes and sparse short glandular trichomes (stalk short, unicellular; head dark, elongate, multicellular), the calyx appendages five, (2.8–) 3–4 (–5) mm long, subequal, thick, erect, cylindrical, inserted very close to the margin. Corolla (8–) 10–12 mm long, 13–20 mm in diameter, thick, white with greenish-yellow spots outside, mostly with large purple spots on the lobes and the throat and cream centre within, stellate with abundant interpetalar membrane, lobed halfway or less of the way to the base, pubescent adaxially with a continuous ring of long glandular trichomes (stalk 2–3-celled; head globose, peltate, unicellular) in the throat and base of the lobes, glabrous abaxially, the tube 4.5–5 mm long, the lobes 4.5–6.5 mm long, 5–8 mm wide, broadly triangular to triangular, the margins densely pubescent, the tips cucullate. Stamens five, subequal; filaments 2.7–3.1 (–4.1) mm long, white, inserted on the corolla ca. 1 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.5–1.9 mm long, ellipsoid, blue, not connivent at anthesis. Gynoecium with ovary 1.3–1.5 mm long, ca. 1.2 mm in diameter, light green, subglobose to ovoid; ovules more than two per locule; nectary ca. 0.3 mm tall; styles homomorphic, 4.3–5 (–7) mm long, barely exserted beyond the anthers, white, clavate; stigma ca. 0.2 mm long, ca. 0.7 mm wide, discoid, cream. Berry 6–7 mm in diameter, globose-depressed, green when immature, greenish at maturity, deciduous, pungent, the pericarp with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 18–25 mm long, pendent and slightly curved, slightly angled, widened at the apex, green; fruiting calyx ca. 4 mm in diameter, persistent, not accrescent, discoid, yellowish-green, the appendages spreading, green. Seeds 7–13 per fruit, 3.5–4 mm long, 2.5–3 mm wide, ellipsoid to reniform, brownish-black to black, the seed coat deeply reticulate and slightly tuberculate at margins (SM), reticulate with small pillar-like outgrowths at margins (SEM), the cells polygonal in shape, the lateral walls straight to wavy; embryo imbricate.

Figure 39. 

Capsicum carassense A flowering branch B–D leaf morphology E eglandular trichome of the stem F glandular trichome of the calyx G, H flower buds in different stages of development I flower J opened corolla K fruit L fruiting calyx. From Bianchetti et al. 1364. Drawn by L. Bianchetti. Published in Barboza et al. (2020a), reproduced with permission.

Figure 40. 

Capsicum carassense A plant, showing the typical lanceolate leaves B inflorescence with geniculate pedicels C flower, in front view D fruit A–C from Stehmann 6344 D from Agra 7268. Photos by J.R. Stehmann. Published in Barboza et al. (2020a), reproduced with permission.

Distribution

Capsicum carassense is endemic to south-eastern Minas Gerais State (Brazil), growing mainly in the Serra do Caraça and other nearby mountainous areas (Fig. 41).

Figure 41. 

Distribution of C. carassense, C. ceratocalyx and C. chacoense

Ecology

Capsicum carassense inhabits the understorey of the semi-deciduous montane Atlantic Forest, in a shaded and moist environment, between 1,000 and 1,390 m elevation.

Phenology

In flower from October to January, also in May; fruiting in December, February and April.

Chromosome number

Not known.

Common names

None recorded.

Uses

None recorded.

Preliminary conservation assessment

EOO (483.4 km2); AOO (32 km2). Capsicum carassense is considered Endangered (EN, B1ab(iii,iv)). We suggest this because of its very restricted geographic distribution, as well as the increasingly degraded habitat quality, especially associated with the extensive iron mining activities in the region (Barboza et al. 2020a).

Discussion

Capsicum carassense belongs to the Atlantic Forest clade (Barboza et al. 2020a). This species is morphologically very similar to C. mirabile with which it has been confused in herbaria. Both species share a similar habit, the geniculate pedicels at anthesis, the number of calyx appendages, the shape and colour of the corolla, the colour and pungency of the fruits and the blackish seeds. They can be easily distinguished by the moderate to dense indumentum, the narrowly elliptic to lanceolate major leaves with apex acute to obtuse, the shorter petioles (up to 0.6 cm long), the cream buds with greenish-yellow pigmentation, the longer calyx appendages (up to 5 mm) and the larger corolla (13–20 mm in diameter) in C. carassense, compared to C. mirabile which has sparse pubescence (young stems sometimes glabrous), major leaves elliptic (rarely narrowly elliptic) to ovate with apex acuminate to long-acuminate, longer petioles (up to 2.5 cm), purple or greenish-purple buds, shorter calyx appendages (up to 3 mm) and smaller corolla (9–13 mm in diameter) (see also Table 4).

Table 4.

Differences between C. mirum, C. cornutum, carassense and C. mirabile.

Character C. mirum C. cornutum C. carassense C. mirabile
Indument/trichomes (stems and leaves) Densely pubescent/trichomes antrorse Densely pubescent/trichome spreading Moderately to densely pubescent/trichomes antrorse Glabrate to sparsely pubescent/trichomes antrorse
Major leaf shape Mostly elliptic, apex acute to acuminate Ovate to widely elliptic, apex acuminate Narrowly elliptic to lanceolate, apex acute to obtuse Elliptic to ovate, rarely narrowly elliptic, apex acuminate to long- acuminate
Major leaf length/width ratio 2.5–3 2.4–5 (4–) 5–10 (–16) (2–) 2.5–4 (–4.9)
Petioles length 0.8–2.5 cm 0.3–0.8 cm 0.2–0.6 cm 0.7–2.5 cm
Pedicels length 12–17 mm (22–) 25–35 mm (12–) 15–20 (–22) mm (13–) 16–25 mm
Buds colour Purple White with green and purple spots Cream with greenish-yellow spots Purple or greenish-purple
Calyx appendages 10, subequal, spreading, long, (1.7–) 2–3.2 mm (5–) 7–10, unequal, erect or spreading, short to long, 0.5–6 mm 5, subequal, long, erect, (2.8–) 3–4 (–5) mm 5, subequal, erect, short to long, (0.4–) 0.5–1.5 (–3) mm
Corolla size 6–8 mm long, 11–14 mm in diameter (8–) 9–14 mm long, 18–22 mm in diameter (8–) 10–12 mm long, 13–20 mm in diameter (6–) 7.5–12 mm long, (9–) 10–13 mm in diameter
Corolla colour Almost entirely purple and a thin white border within White with small purple or reddish-brown spots within Mostly with large purple spots and a thin white border within Mostly with large purple spots and a thin white border within

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum cardenasii Heiser & P.G.Sm., Brittonia 10(4): 195. 1958.

Figs 42, 43

Type

Cultivated at Indiana University greenhouse from seeds sent by M. Cárdenas from market in La Paz, Bolivia, 15 Aug 1956, C.B. Heiser Jr. 4196 (Paul Smith Ac.-1793) (lectotype, designated here: IND [IND1000063, acc. # 139347]; isolectotype: IND [IND1000064, acc. # 139348]).

Description

Erect shrubs or subshrubs, 0.8–2 (–2.5) m tall, with the main stem 1–1.5 cm in diameter at base, much branched from near the base, the fragile branches in a typical “zig-zag” appearance above. Young stems strongly angled, green, glabrescent with sparse appressed-antrorse, simple, uniseriate, 4–6 (–7)-celled eglandular trichomes 0.08–0.6 mm long and minute, simple, glandular trichomes (stalk short; head dark); nodes green; bark of older stems greyish-white or with light brown-green fissures, glabrescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair subequal in size and shape. Leaves membranous, slightly discolorous, glabrescent, with sparse eglandular trichomes similar to those on stems and many small glandular trichomes (stalk unicellular; head dark, multicellular) on both surfaces, the glandular trichomes more abundant along mid-vein abaxially; blades of major leaves 3–5 (–6.5) cm long, 1.4–2.5 cm wide, narrowly ovate or ovate-lanceolate, the major veins 3–4 on each side of mid-vein, the base attenuate, the margins entire, the apex acute; petioles 1.2–2 cm long, glabrous or glabrescent; the blades of minor leaves 2–3.5 cm long, 0.9–1.2 cm wide, narrowly ovate or ovate-lanceolate, the major veins 2–3 on each side of mid-vein, the base attenuate, the margins entire, the apex acute or obtuse; petioles 0.5–0.8 cm long. Inflorescences axillary, 2–3 flowers per axil or flowers solitary; flowering pedicels 8–18 (–22) mm long, angled, erect to slightly spreading, geniculate at anthesis, entirely green, or purple distally, with moderate small glandular trichomes (stalk transparent, uni-bicellular; head dark, multicellular) and sparse short, antrorse eglandular trichomes; pedicels scars inconspicuous. Buds ellipsoid or ovoid, lilac or violet. Flowers 5-merous. Calyx 1–3 mm long, 2–3 mm wide, cup-shaped, thick, green or green with violet spots, moderately pubescent with the same glandular and eglandular trichomes as the pedicels, the calyx appendages five, 1–2 mm long, 0.3 mm wide, subequal, thick, erect or spreading, cylindrical, inserted close to the margin, sparsely pubescent with the same trichomes as the calyx tube. Corolla (6–) 6.5–12 mm long, 8–11 (–13) mm in diameter, thick, almost completely violet or lilac, but white at the base and along the main veins outside and within, sometimes greenish-yellow spots near the base within, campanulate with interpetalar membrane, lobed 1/3 or less of the way to the base, the tube 7–9 mm long, pubescent adaxially with short glandular trichomes (stalk 1–2-celled; head globose, unicellular) up to near its base, glabrous abaxially, the lobes (1.5–) 3–3.2 mm long, 2–2.4 mm wide, triangular, erect or spreading, alternating with five minute interlobes, glabrous adaxially and abaxially, the margins papillate, the tips acute, papillate. Stamens five, equal; filaments (4–) 6–7 mm long, whitish or lilac, inserted on the corolla 1.5–2 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.5–1.8 mm long, ellipsoid, lilac or bluish, not connivent at anthesis. Gynoecium with ovary 1.5–1.85 mm long, 1.2–1.6 mm in diameter, green, ovoid or pear-shaped; ovules more than two per locule; nectary 0.4–0.6 mm tall, light green; styles homomorphic, 4.5–5.7 mm long, exserted ca. 1 mm beyond the anthers, lilac or purple, clavate; stigma ca. 0.2 mm long, 0.8 mm wide, discoid or globose, pale green. Berry 6–10 mm in diameter, globose or subglobose, green when immature, orange-red to bright red at maturity, deciduous, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 10–24 mm long, pendent, strongly angled, slightly widened distally, usually green; fruiting calyx 2–4 mm in diameter, persistent, not accrescent, discoid, green, the appendages 1–2.5 mm long, ca. 0.3 mm wide, appressed to the berry, spreading or reflexed. Seeds (4–) 5–13 per fruit, (2.5-) 3–4.2 mm long, (2.2–) 2.5–2.8 mm wide, C-shaped or subglobose, pale yellow to brownish-yellow, the seed coat reticulate to obscurely reticulate (SM), mostly cerebelloid (SEM), the cells irregular in shape, the lateral walls strongly sinuate in the central zone, rectangular to subpolygonal at margins; embryo imbricate.

Distribution

Capsicum cardenasii is a narrow endemic species restricted mainly to the highlands of La Paz Department (Bolivia, Fig. 32). Only one collection from Tarija, probably introduced.

Ecology

Capsicum cardenasii is a typical component of the warm and dry hillsides and remnants of forests in the inter-Andean valleys, growing preferentially in open places between cactus and Cassia, at 2,400–3,000 m elevation. It is cultivated by local people on small farms for local or family use of the fruits (Barboza, pers. obs.).

Phenology

Flowering from December to March; fruiting from February to April.

Chromosome number

n = 12 (Heiser and Smith 1958); 2n = 2x = 24 (Pickersgill 1977; Moscone et al. 2007; Scaldaferro et al. 2013, 2016).

Common name

Bolivia. Ulupica (La Paz, Heiser & Smith 4196).

Indigenous name

Bolivia. Uaika (Aymará, La Paz, Barboza 4881).

Uses

The fruits are harvested directly from wild plants and marketed locally on a small scale (Jäger et al. 2013), mainly in La Paz, Bolivia. People consume dehydrated or fresh fruits in the preparation of a hot sauce called ‘Jallpa huayka’ (in aymara), ‘uchu llajfua’ (in quechua) (Heiser and Smith 1958; Cárdenas 1969), ‘llajwa’ (in quechua) or ‘llaswa’ (most popular), a mix of ‘tomato’, ‘onion’, ‘ulupica’ or other ‘chiles’ and aromatic herbs. Fruits are also preserved in vinegar or in oil and used as pickle (‘escabeche’) or they are cooked in boiling water before being sold (National Research Council 1989; Barboza, pers. obs.).

Preliminary conservation assessment

EOO (1,032.864 km2); AOO (32 km2). Capsicum cardenasii is a geographically isolated species from the dry valleys of Luribay (Prov. Loayza), not far from La Paz; based on its extent of occurrence and the number of localities (6), it is assigned a status of Endangered (EN; B1ab(iii,iv)). It is harvested by local people; its area of distribution is poorly known.

Figure 42. 

Capsicum cardenasii A flowering branch B eglandular trichome of the calyx C glandular trichome of the corolla D glandular trichome of the adaxial surface of the calyx E flower F section of the calyx showing the venation G sector of opened corolla H gynoecium I fruit J anatomical detail of the pericarp (note the giant cell in the mesocarp) K seed L seed, in cross section M structure of seed coat at the seed margin N structure of seed coat at the seed body O embryo A–H from Eshbaugh 1527 I–O from Eshbaugh 2046 J. Drawn by N. de Flury.

Discussion

Capsicum cardenasii is resolved within the Purple corolla clade (Carrizo García et al. 2016). More recent preliminary phylogenetic evidence showed that C. pubescens is sister to this clade (Carrizo García et al. 2019; CCG, pers. obs.), thus circumscription of these taxa is under revision (CCG, pers. obs.; see under C. pubescens description). The fruits of C. cardenasii are very similar to those of C. eximium and C. eshbaughii, both also known as “ulupica”. The three species can be differentiated by their general pubescence and corollas. Capsicum eshbaughii (Fig. 58) has a dense pubescence of long furcate glandular trichomes and stellate white corollas with greenish-yellow spots around the throat (rarely purple lines in the lobes). In contrast, C. cardenasii and C. eximium have sparse to moderate pubescence of eglandular simple trichomes and minute simple glandular trichomes. In addition, C. cardenasii has lilac to purple, broadly campanulate corollas (Fig. 43C–F), while C. eximium has mostly purple, lilac or magenta, stellate corollas with greenish-yellow pigmentation within (Fig. 60H–J).

Figure 43. 

Capsicum cardenasii A plant B flower bud C, D flower, in lateral view E, F flower, in front view (note the different colouration inside the corolla) G immature fruits H mature fruit A, B, D, E, G from Barboza 4882, C, F, H from Barboza 4881. Photos by G.E. Barboza.

Phytogeographically, Eshbaugh (1979) suspected that C. cardenasii and C. eximium could be sympatric on the eastern margin of the range of C. cardenasii. He had observed some intermediate plants between both taxa in the Luribay Valley (Eshbaugh 1976, 1979, but specimens not cited) and, at that time, no collections of the typical C. eximium were known from that area (Eshbaugh 1979). We recently collected C. eximium (e.g. Barboza 4885) in Luribay, very near to the sites where C. cardenasii grows abundantly. Luribay Valley deserves to be explored intensively to document the presence of natural hybrids in the area and to find out if their level of fertility is as high as in experimental crosses (Heiser and Smith 1958; Eshbaugh 1976). Evidence of hybridisation was found while attempting to identify and cytogenetically characterise these Capsicum species, using a molecular cytogenetic approach (seeds from Gene Bank, Nijmegen University, The Netherlands); however, the fertility of the hybrid has not been possible to ascertain (Scaldaferro 2019).

Eshbaugh and Smith (1971) also obtained successful crosses between C. cardenasii and C. eshbaughii (e.g. IND 139349), though F1 and F2 hybrids were less fertile than with C. eximium. The narrow distributions of Capsicum cardenasii and C. eshbaughii are allopatric, with the first species in north-western Bolivia (La Paz: Luribay) and the second concentrated in central-eastern Bolivia (mainly Santa Cruz: Samaipata). Their distributions are separated by nearly 700 km of distance, making hybridisation in the wild highly unlikely.

Although the number of collections of C. cardenasii obtained in the field are scarce (8), the ease with which the seeds of this species germinate and produce fertile plants explains the large numbers (> 20) of specimens (and duplicates) gathered from plants in cultivation that are housed in many herbaria (see Specimens Examined).

The type collection of C. cardenasii consists of flowering specimens (two sheets dated 15 Aug 1956 at IND) obtained from seeds bought at the La Paz (Bolivia) marketplace; it is supposed that fruits came from warm, dry places along the Río Abajo, near La Paz, at 2400 m altitude (Heiser and Smith 1958). Of the two specimens in IND, that with barcode 1000063 is the most complete, is labelled type and is here designated as the lectotype. There are specimens distributed in other herbaria (e.g. CORD, IND, LIL, US) with the same collection number as the lectotype (Heiser 4196, Paul Smith Acc. 1793), but these have different dates of collection and should not be considered as duplicates of the lectotype.

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum ceratocalyx M.Nee, Brittonia 58 (4): 326. 2006.

Fig. 44

Type

Bolivia. La Paz: Prov. Sud Yungas: 7.5 km (by road) from Huancané on road to San Isidro, 16°21'S, 067°30'W, 2225 m elev., 10 May 2001, M. Nee, L. Bohs, S. Knapp & J.M. Mendoza F. 51778 (holotype: LPB [LPB0003514]; isotypes: CORD [CORD00004289], MO [MO-2078805, acc. # 5959885], NY [01085523], USZ).

Description

Erect shrubs 0.80–3 m tall, much branched above. Young stems angled, green, glabrous to sparsely pubescent, with antrorse, curved, simple, uniseriate, 2–4 (–5)-celled, eglandular trichomes 0.09–0.5 mm long; nodes solid, green; bark of older stems brown, glabrous; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair markedly unequal in size, similar in shape. Leaves coriaceous, slightly discolorous, glabrescent on both sides with sparse eglandular trichomes similar to the ones of the stems, mainly along the mid-vein abaxially; blades of major leaves 10–22 cm long, 4–7 cm wide, elliptic, the major veins 6–7 on each side of mid-vein, the base attenuate and unequal, the margin slightly revolute, the apex long-acuminate; petioles 1–2 (–3) cm long, glabrous; blades of minor leaves (3–) 5.5–7.5 cm long, 2–2.7 cm wide, elliptic, the major veins 3–4 on each side of mid-vein, the base short-attenuate, the margin slightly revolute, the apex acute; petioles 0.5–0.6 cm long, glabrous. Inflorescences axillary, congested, (4–) 8–10 (–12) flowers on a short rachis; flowering pedicels 10–23 mm long, strongly angled and nearly winged, erect, geniculate at anthesis, green, glabrous; pedicels scars prominent and corky. Buds ovoid, yellow. Flowers 5-merous. Calyx 1.8–2 mm long, ca. 3 mm wide, cup-shaped, slightly 5-nerved, sparsely pubescent, with the same antrorse eglandular trichomes of the young stems and small glandular trichomes (stalk unicellular; head multicellular), the calyx appendages (3–) 5, 0.25–2.5 mm long, subequal, thick, notoriously incurved, spreading, flattened laterally, glabrescent, inserted close to the margin. Corolla 6–8.5 mm long, ca. 5 mm in diameter, yellow with green spots within, stellate to broadly campanulate with interpetalar membrane, lobed nearly or more than the halfway to the base, the tube 3–4 mm long, pubescent adaxially with sparse glandular trichomes (stalk uni-bicellular; head unicellular), glabrous abaxially, the lobes 3.2–3.7 mm long, 2–3 mm wide, triangular, erect, glabrous adaxially and abaxially, the margins involute, the tips acute, papillate. Stamens five, equal; filaments 1–2 mm long, inserted on the corolla 1–1.5 mm from the base, with auricles fused to the corolla at the point of insertion; anthers 1.4–1.8 mm long, ovoid, not connivent at anthesis. Gynoecium with ovary 1.8–2 mm long, 1.3–1.5 mm wide, ovoid; ovules more than two per locule; nectary ca. 0.3 mm tall; styles homomorphic, 5.8–6.5 mm, clavate; stigma 0.1–0.2 mm long, ca. 0.5 mm wide, discoid. Berry 8–11 mm in diameter, globose, slightly flattened at the apex, green when immature, bright red at maturity, persistent, pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels (15–) 25–30 mm long, erect, conspicuously angled, winged and widened distally; fruiting calyx 3–5 (–7) mm in diameter, persistent, not accrescent, discoid, the appendages 2–5 mm long, incurved. Seeds 13–26 per fruit, 4–5 mm long, (2.6–) 3–4 mm wide, C-shaped or teardrop-shaped, brownish-yellow to brown, the seed coat reticulate (SM), reticulate-cerebelloid (SEM), the cells irregular in shape, the lateral walls strongly sinuate in the seed body, wavy at margins to nearly straight near hilum; embryo coiled.

Figure 44. 

Capsicum ceratocalyx A fruiting branch B node with pedicels scars and base of two pedicels C flower D calyx E opened corolla F fruit G seed H seed, in longitudinal section A–E from Beck 28089, F–I from Seidel & Hinojosa 1267. Drawn by P. Peralta.

Distribution

Capsicum ceratocalyx is endemic to the Bolivian Departments of La Paz and Cochabamba (Fig. 41).

Ecology

Capsicum ceratocalyx is known from few collections, all from moist montane forest (Yungas) with little disturbance, between 700 and 2,500 m elevation.

Phenology

Flowering and fruiting from November to July.

Chromosome number

Not known.

Common names

None recorded.

Uses

None recorded.

Preliminary conservation assessment

EOO (3,612.392 km2); AOO (32 km2). Capsicum ceratocalyx is an endemic with relatively small EOO and AOO from the montane forests of the Bolivian Yungas, an ecoregion that deserves urgent special conservation efforts (Mueller et al. 2002). The few collections (9) come from unprotected areas and the populations are threatened by expanding agriculture. For these reasons, it is assigned a status of Endangered (EN; B1ab(iii,iv)).

Discussion

Capsicum ceratocalyx has been assigned to the Bolivian clade (Carrizo García et al. 2016; Barboza et al. 2019), with C. minutiflorum as its sister species. Capsicum ceratocalyx is a poorly known component of the mid-altitudes of the Bolivian Yungas. It has large coriaceous leaves, a calyx with 3–5 incurved and spreading appendages, yellow corollas and winged fruiting pedicels. The affinities of this species need to be further studied, since new preliminary data, based on genome-wide DNA sequences, suggest this species may form an isolated lineage (CCG, pers. obs.).

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum chacoense Hunz., Darwiniana 9(2): 228. 1950.

Figs 45, 46

Capsicum chacoense Hunz. var. tomentosum Hunz., Darwiniana 9(2): 235. 1950. Type. Argentina. Chaco: [Dept. Independencia], Colonia J.J. Mármol, 31 Dec 1946, F. Buratovich 117 (holotype: LIL [acc. # 173409); isotype: B [B10-1067916]).

Type

Argentina. Chaco: [Dept. Primero de Mayo] entre Colonia Benítez y Resistencia, 12 Mar 1945, A.T. Hunziker 7340 (lectotype, designated here: CORD [CORD00003920]; isolectotypes: CORD [CORD00003919, CORD00087962]).

Description

Low compact shrubs or subshrubs 0.40–1 (–2.5) m tall, much branched from a thick basal rootstock, with the main stem 2–3 cm in diameter at base, the branches expanded and divaricated, in a typical “zig-zag” appearance. Young stems strongly 3–4-angled, fragile, green or purple, sparsely to densely pubescent with antrorse and more or less rigid or spreading and flexuous, simple, uniseriate, 3–7-celled, eglandular trichomes 0.03–0.8 (–1.4) mm long, rarely branched trichomes 1–7 mm long; nodes green; bark of older stems brown, glabrescent to glabrous; lenticels sparse. Sympodial units unifoliate or difoliate, the leaves geminate; leaf pair more or less similar in shape and size. Leaves membranous, slightly discolorous or concolorous, glabrescent to densely pubescent with eglandular trichomes similar to the stems on both surfaces and margins; blades of all leaves 2–6 (–8) cm long, 1–3.5 (–5) cm wide, ovate, narrowly ovate or rarely elliptic, the major veins 3–4 on each side of midvein, the base attenuate and asymmetric, the margins entire, the apex long-acuminate; petioles 0.5–2 (–3.5) cm long, glabrescent to densely pubescent. Inflorescences axillary, flower solitary; flowering pedicels (5–) 10–25 (–40) mm long, strongly angled, erect, geniculate at anthesis, green, scarcely to moderately pubescent; pedicels scars inconspicuous. Buds globose or ovoid, white. Flowers 5-merous. Calyx 1.2–2 mm long, 2–2.5 mm wide, cup-shaped, thick, green, moderately pubescent with the same eglandular trichomes as stems, the calyx appendages 7–10 (rarely 5), unequal, rarely subequal, the five main appendages longer, 0.5–1.5 mm long, 0.3 mm wide, alternating with 2–5 shorter secondary appendages up to 0.7 mm long, thick, erect or slightly spreading, cylindrical or slightly compressed, inserted close to the margin, sparsely pubescent with the same trichomes as calyx tube. Corolla 4–6 (–9) mm long, 9–11 mm in diameter, thick, entirely white, stellate with interpetalar membrane, lobed 1/2 or less of the way to the base, glabrous adaxially and abaxially, the tube 2–4 mm long, the lobes 2–2.6 (–3.1) mm long, 2–2.4 mm wide, triangular, spreading, the margins papillate, the tips acute, papillate. Stamens five, equal; filaments 0.8–1.5 mm long, white, inserted on the corolla 1–1.3 mm from the base, with conspicuous auricles free, not fused to the corolla at the point of insertion; anthers (0.9–) 1.2–1.5 mm long, ellipsoid, yellow or cream, not connivent at anthesis. Gynoecium with ovary 1.5–2.3 mm long, ca. 1.7 mm in diameter, light green, ovoid; ovules more than two per locule; nectary ca. 0.4 mm tall, light green; styles homomorphic, 2.4–2.8 (–4) mm long, exserted ca. 1–1.5 mm beyond the anthers, white, cylindrical; stigma ca. 0.2 mm long, 0.2 mm wide, cream or light green, globose. Berry 7–10 mm in diameter, globose or (6–) 8–14 mm long, 5–8 mm in diameter, ellipsoid, green or green with blackish spots when immature, orange to bright red at maturity, deciduous, pungent, in some populations, non-pungent, the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 15–20 mm long, erect, strongly angled, widened distally, green; fruiting calyx 4–5 mm in diameter, persistent, not accrescent, discoid, green, the appendages 0.5–1.6 mm long, spreading or slightly recurved. Seeds 14–25 per fruit, 3.4–4 mm long, 2.8–3 mm wide, C-shaped, rarely subglobose or reniform, pale yellow, the seed coat smooth to reticulate (SM), reticulate-cerebelloid (SEM), the cells irregular in shape, the lateral walls strongly sinuate in seed body, nearly straight at margin; embryo imbricate or coiled.

Distribution

Capsicum chacoense is a widespread species usually confined to Chaco vegetation, which extends from Bolivia and Paraguay to central Argentina (Fig. 41).

Ecology

Capsicum chacoense is a typical component of the dry and subhumid Gran Chaco forests ranging from the inter-Andean valleys at higher elevations to the lower Chaco forests in northern Argentina and Paraguay. Capsicum chacoense is found in the undergrowth, beneath the shade of nurse shrubs or trees, from 50–2,700 m elevation.

Phenology

Flowering from late October to March and April, fruiting from January to May or June.

Chromosome number

n = 12 (Moscone 1992); 2n = 24 (Pickersgill 1977; Moscone 1990; Moscone et al. 1993, 1995, 2007).

Common names

Argentina: Ají (Catamarca, Brizuela 1127; Chaco, Aguilar 568; Córdoba, Fernández 17; Corrientes, Nicora s.n.; Santiago del Estero, Soriano & Barret 3592), Bolita (Santiago del Estero, Perrone s.n.), Cumbaré (Córdoba, Castellanos 217), Cumbarí (Catamarca, de Ance 81; Corrientes, Ibarrola 2988; La Rioja, Hunziker 4807), Lají (Córdoba, Lorentz s.n.), Pipí (Formosa, Morel 910), Putaparió (Chaco, Gaillard s.n.; Córdoba, Cocucci 4966; Tucumán, Schreiter 1927), Putaqueteparió (Córdoba, Hawkes et al. 3308), Uchuca (Catamarca, Vervoorst 3535), Ají cumbarí (Catamarca, Schickendantz 74; Chaco, Schulz 2; Córdoba, Castagnino s.n.; La Rioja, Giacomelli 130), Ají chuca (Chaco, de Ance 81), Ají fuerte (Córdoba, Villafañe 400), Ají picante (Córdoba, Botta 115), Ají quitucho (Catamarca, Capparelli 158), Ají uchiquita (La Rioja, Hunziker 4807), Ají uchuco (Catamarca, Falcone & Castellanos 262), Ají del campo (Chaco, Schulz 92; Córdoba, Castellanos 217; La Rioja, Hunziker 5088; San Luis, Anderson 1447; Santa Fe, Krapovickas 762), Ají del monte (Catamarca, Cabrera 1132; Chaco, Fortunato 1445; Córdoba, Spegazzini s.n.; Salta, Ayarde & Sidán 299; Santiago del Estero, Bartlett 20435; Tucumán, Schreiter 1927), Ají mala palabra (Santiago del Estero, Crespo s.n.), Ají puta parió (San Juan, Cortez 156), Ají quitucho dulce (Salta, Hunziker 1578), Picante del monte (Jujuy, Joergensen s.n.), Pimiento del monte (Chaco, Meyer 8561); Bolivia: Aribibi (Santa Cruz, Navarro & Vargas C. 261), Ají del zorro (Chuquisaca, Saravia Toledo 10343); Paraguay: Ají (Boquerón, August 18), Cambarí (Central, Rojas 10821), Cumbary (Capital, Rojas 14325).

Indigenous names

Argentina: Atéshuk (Chorote, Scarpa 2007), Awarañink¨i’¨i’ (Tapieté, Montani and Scarpa 2016), Chemmak’-raík’ (= picante) (Mocoví, Scarpa and Rosso 2014), Ke-ig (Guaraní, Corrientes, Ibarrola 2988), Kodae (Pilagá, Formosa, Martínez Crovetto 31, Arenas 3043), Ko’rae (Pilagá, Filipov 1994), Ko’rai’ (Pilagá, Formosa, Arenas 1956), Pájanak (Chorote, Scarpa 2007), Pa:nãn (Salta, Arenas 2121), Quihiy (Chaco, Escobar 19); Bolivia: Aguara keu (Santa Cruz, Bourdy 2000), Kî mi’ (Santa Cruz, de Michel 2568); Paraguay: Atés (Boquerón, Arenas 1688), Atic (Presidente Hayes, Arenas 1546), Ciaq taqatic (Presidente Hayes, Arenas 2377), Hũpita (Boquerón, Arenas 1853), Naatikgit (Boquerón, Arenas 436), Yemade (Toba, Presidente Hayes, Williams et al. 140).

Uses

The pungent fruits are locally used as spicy food additives for both local and indigenous people (National Research Council 1989; Arenas and Scarpa 2007; Biurrun et al. 2007; Scarpa 2007; Martínez Crovetto 2014; Montani and Scarpa 2016; Saur Palmieri et al. 2018). In regional markets, fruits are sold fresh or as pickles in oil or vinegar (Barboza, pers. obs.). Fruits are also used in traditional medicine (see Table 3).

Preliminary conservation assessment

EOO (1,724,002 km2); AOO (952 km2). Capsicum chacoense is widespread across subtropical Chaco forests from Bolivia to Paraguay and is assigned the Least Concern (LC) category.

Discussion

Capsicum chacoense belongs to the Baccatum clade (Carrizo García et al. 2016). It has a peculiar dwarf habit, profusely branched from the thick rootstock with small leaves, solitary flowers, entirely white corollas, filaments with conspicuous free auricles at point of insertion to the corolla and abundant red mature fruits per plant. Its most variable character is pubescence. Typically, plants are sparsely pubescent, but densely tomentose populations grow in north-eastern Argentina (Chaco) and Paraguay (Alto Paraguay), which, in the past, were recognised as a separate variety, C. chacoense var. tomentosum (Hunziker 1950). Some of these tomentose populations were included in Hunziker’s circumscription of C. microcarpum var. tomentosum (= C. rabenii) (Hunziker 1950), leading to confusion between the two taxa.

A second variable feature is the number and degree of development of the calyx appendages. The most common condition is the presence of calyces with 10 unequal appendages, the five main ones longer and alternating with five shorter, all of them usually well-developed (that is, appendages exceed the truncate calyx edge). In some cases, the shorter appendages vary from 2–5, with some exceeding the calyx edge and others scarcely noticeable, reduced to a mucro. Calyces with this second pattern create confusion in the identification of fruiting specimens, especially when all short appendages are not well-developed. Often, these specimens are annotated in herbaria as C. baccatum from which C. chacoense can be distinguished by its smaller and entirely white corollas.

The fruits of C. chacoense are locally very appreciated for their flavour and high pungency (see references in Uses). However, C. chacoense is naturally polymorphic for the production of capsaicinoids, such that completely pungent and completely non-pungent individuals co-occur in some Bolivian populations (Tewskbury et al. 2006); non-pungent fruits have also been recorded in Argentinean populations (e.g. Salta, Hunziker 1578). Tewksbury et al. (2008b) demonstrated that the pungency variation found in C. chacoense may be an adaptive response to selection by a microbial pathogen. Thus, capsaicinoids protect the fruits and seeds from fungal pathogens (Fusarium) that severely reduce seed viability.

There is little information about which birds, the most effective dispersers of Capsicum seeds, eat C. chacoense fruits. Information from a herbarium label (Bolivia, Debouck 3016) suggests that small parrots eat the fruits of this species. Tewskbury et al. (2006) mentioned Elaenia parvirostris ‘fiofío pico corto’ (Fam. Tyrannidae) and Turdus amaurochalinus ‘zorzal chalchalero’ (Fam. Turdidae) as the major dispersers of Capsicum seeds in Bolivia.

In the protologue of C. chacoense, Hunziker (1950) indicated “Typus speciei. (ATH)” [Armando Teodoro Hunziker] referring to his own Herbarium now housed at CORD. In CORD, there are three mounted sheets of Hunziker 7340, two of them (sheets A and B) are labelled as “holotype” (barcodes CORD00003919, CORD00003920) and the third one (CORD00087962) has a label in Hunziker’s hand, indicating it is a duplicate specimen (probably with the intention to be donated elsewhere, but now mounted and accessioned at CORD). Sheet B contains a complete fruiting young individual with a label where the word Typus was handwritten by Hunziker as an indication that it should be the holotype; we are designating this sheet B (CORD00003920) as the lectotype.

Figure 45. 

Capsicum chacoense A root B fruiting branch C eglandular trichome of the leaf D flower E sector of opened corolla F gynoecium G fruit H seed I seed, in cross section J embryo A, C–F from Hunziker 18572 B, G–J from Hunziker et al. 25388. Drawn by N. de Flury. Published in Barboza (2013), courtesy of the Board of the Instituto Darwinion (San Isidro, Buenos Aires, Argentina), reproduced with permission.

Figure 46. 

Capsicum chacoense A plant B flower bud C flower, in pre-anthesis D flower, in front view E immature fruit F mature fruit A–F from Barboza et al. 4910. Photos by G.E. Barboza.

Specimens examined

See Suppl. material 4: Appendix 4.

Capsicum chinense Jacq., Hort. Bot. Vindob. 3: 38. t. 67. 1776.

Fig. 47

Capsicum cerasiforme Mill., Gard. Dict. ed. 8, no. 5. 1768. Type. Cultivated at the Chelsea Physic Garden, London, seeds from Spanish West Indies (no specimens cited).

Capsicum cerasiforme Lam., Tabl. Encycl. 2: 26. 1794, nom. illeg., not Capsicum cerasiforme Mill. (1768). Type. Brazil. “E Brafilia” (lectotype, designated here: P [P00357733]).

Capsicum luteum Lam., Tabl. Encycl. 2: 26. 1794. Type. India. “Ex India. Sonnerat” P. Sonnerat s.n. (lectotype, designated here: P [P00357731]).

Capsicum cerasiforme Willd., Enum. Pl. [Willdenow] 1: 242. 1809, nom. illeg., not Capsicum cerasiforme Mill. (1768). Type. Cultivated in the Berlin Botanic Garden, Germany, “Habitat…..” Capsicum cerasiforme [sheet] 1, herb. Willdenow (lectotype, designated here: B [B-W04427-01-0]).

Capsicum milleri Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 563. 1819, nom. illeg. superfl. Type. Based on Capsicum cerasiforme Mill. (cited in synonymy).

Capsicum indicum Dierb. var. luteum (Lam.) Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 24. 1829. Type. Based on Capsicum luteum Lam.

Capsicum indicum Dierb. var. humifusum Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 26. 1829. Type. Based on Capsicum cerasiforme Mill. (cited in synonymy).

Capsicum indicum Dierb. var. ochranthum Dierb., Arch. Apotheker-Vereins Nordl. Teutschl. 30: 27. 1829. Type. Based on Capsicum cerasiforme Willd. (cited in synonymy).

Capsicum dichotomum Vell., Fl. Flumin.: 61. 1829 (“1825”); Fl. Flumin. Icon. 2: t. 9. 1831 (“1827”), as “Capsicum conicum”. Type. Brazil. [Rio de Janeiro]: “Colitur hortis” (lectotype, designated by Knapp et al. 2015, pg. 824: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_012] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 9. 1831).

Capsicum odoriferum Vell., Fl. Flumin.: 61. 1829 (“1825”); Fl. Flumin. Icon. 2: t. 8. 1831 (“1827”). Type. Brazil. [Rio de Janeiro]: “Colitur hortis” (lectotype, designated by Knapp et al. 2015, pg. 824: [illustration] Original parchment plate of Flora Fluminensis in the Manuscript Section of the Biblioteca Nacional, Rio de Janeiro [cat. no.: mss1198651_011] and later published in Vellozo, Fl. Flumin. Icon. 2: t. 8. 1831).

Capsicum conicum Vell., Fl. Flumin. Icon. 2: t. 9. 1831. (“1827”), nom. illeg., not Capsicum conicum G.Mey. (1818). Type. Based on Capsicum dichotomum Vell.

Capsicum toxicarium Poepp., Not. Natur- Heilk. 32: 228. 1832. Type. No locality cited (no specimens cited, no original material found). Peru. “Peruvian hispan., Maynas alto”, Jan 1830, E.F. Poeppig 2220 (neotype, designated here: W [acc. # 0102196].

Capsicum cereolum Bertol., Hort. Bonon. Pl. Nov. 1: 6. t. 2. 1838. Type. Cultivated in Bologna, Italy “Nascitur in Brasilia unde semina ad nos attulit, et comiter dedit Eq. NUNNEZIUS” (lectotype, designated here [illustration]: Bertoloni, Hort. Bonon. Pl. Nov. 1: 6. t. 2. 1838).

Capsicum cordiforme Mill. var. subangulosum Fingerh., Monogr. Capsic.: 29. 1832. Type. No locality cited (no specimens cited; lectotype, designated here: [Fingerhuth, Monogr. Capsic. Tab. IX d. 1832]).

Capsicum cordiforme Mill. var. majus Fingerh., Monogr. Capsic.: 30. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. IX e. 1832).

Capsicum cordiforme Mill. var. minus Fingerh., Monogr. Capsic.: 30. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. X a. 1832).

Capsicum cordiforme Mill. var. olivaeforme Fingerh., Monogr. Capsic.: 30. 1832. Type. No locality cited (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. X b. 1832).

Capsicum cordiforme Mill. var. globosum Fingerh., Monogr. Capsic.: 30. 1832. Type. “Crescit in Indiis et America meridionali” (no specimens cited; lectotype, designated here [illustration]: Fingerhuth, Monogr. Capsic. Tab. X c. 1832).

Capsicum ustulatum Paxton, Paxton’s Mag. Bot. 5: 197. 1838. Type. Cultivated at Chatwsworth House, England (no specimens cited; lectotype, designated here: [illustration], Paxton, Paxton’s Mag. Bot. 5: [un-numbered plate opposite] 197. 1838).

Capsicum cerasiforme Willd. var. maurocarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 422. 1852. Type. No locality cited (no specimens cited; lectotype designated here: [illustration] “SolanumCapsicum dictum perenne minuserectum” Weinemann, Phytanthoza Iconogr. 4: 349, t. 930 c. 1745).

Capsicum oxycarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 426. 1852. Type. Brazil. Rio de Janeiro, 1831, M. Blanchet 90 (holotype: G-DC [G00200070]; isotype: MPU [MPU023041]).

Capsicum cordiforme Mill. var. subsulcatum Dunal, Prodr. [A. P. de Candolle] 13(1): 427. 1852. Type. No locality cited (no specimens cited; lectotype designated here: [illustration] Fingerhuth, Monogr. Capsic. Tab. IX c. 1832).

Capsicum annuum L. var. chinense (Jacq.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum chinense Jacq.

Capsicum annuum L. var. luteum (Lam.) Alef., Landw. Fl.: 132. 1866. Type. Based on Capsicum luteum Lam.

Capsicum annuum L. var. milleri (Roem. & Schult.) Alef., Landw. Fl.: 133. 1866. Type. Based on Capsicum milleri Roem. & Schult.

Capsicum grossum Willd. var. cerasiforme (Lam.) C.B.Clarke, Fl. Brit. India 4: 239. 1883. Type. Based on Capsicum cerasiforme Lam.

Capsicum annuum L. var. cerasiforme (Mill.) Irish, Rep. (Annual) Missouri Bot. Gard. 9: 92. 1898. Type. Based on Capsicum cerasiforme Mill.

Capsicum frutescens L. var. cerasiforme (Mill.) L.H.Bailey, Gentes Herbarum 1: 129 1923. Type. Based on Capsicum cerasiforme Mill.

Capsicum assamicum J.Purkay. & Lok.Singh, Ozean J. Appl. Sci. 5(1): 56. 2012. Type. India. Assam: Tezpur, 157 m, 25 May 2008, Purkayastha & Singh, DRLT 12 (holotype: CAL [by error, BSI in protologue, n.v.]; isotype: CAL [by error, DRLT and BSI in protologue, n.v.]).

Type

Cultivated in Vienna, Austria [“Hort. Bot. Vindob.”], N.J. von Jacquin s.n. (lectotype, designated here: W [acc. # 0080115]).

Description

Low, erect, short-lived subshrubs or rarely shrubs, 0.5–1.5 (–2.5) m tall, with the main stem (0.5–) 0.8–1.5 cm in diameter at base, few to much branched from near the base. Young stems 4-angled, fragile, green or greenish-brown, glabrous to glabrescent, with spreading, simple, uniseriate, (2–) 4–5-celled, eglandular trichomes 0.07–0.7 mm long; nodes solid, green or purple; bark of older stems light brown or green with light brown stripes, glabrous to sparsely pubescent; lenticels absent. Sympodial units difoliate, the leaves geminate; leaf pair unequal in size, similar in shape. Leaves membranous, concolorous or slightly discolorous, dark green above, pale green below, glabrescent to sparsely pubescent, with simple, 4–9-celled, eglandular trichomes 0.6–1.2 mm long, especially along the mid-vein or with a tuft of trichomes in the basal vein axils abaxially; blades of major leaves (4–) 5.25–10 (–15.5) cm long, 2.4–4 (–7) cm wide, ovate to elliptic, the major veins 4–5 (–6) on each side of mid-vein, the base attenuate or truncate and rather unequal, the margins entire, the apex short-acuminate, acuminate or acute; petioles 0.5–3 (–3.5) cm, glabrous to sparsely pubescent; blades of minor leaves similar in shape, 4–5.5 cm long, 2–2.5 cm wide; petioles 0.7–1.5 cm long, with the same pubescence as the major leaves. Inflorescences axillary, 2–4 (–5) flowers per axil, occasionally solitary flower; flowering pedicels 12–20 (–30) mm long, angled, erect or slightly spreading, geniculate at anthesis (wild forms) or pendent and non-geniculate (domesticated forms), green or green with purple lines, glabrous to moderately pubescent, the eglandular trichomes short, antrorse; pedicels scars conspicuous, slightly corky. Buds ellipsoid, cream or greenish-white. Flowers 5–6-merous. Calyx 1–2.5 (–3) mm long, 3–4 mm wide, cup-shaped, green, pentagonal or hexagonal in outline, the main veins strongly marked, the calyx appendages absent or with 5–6 mucro-like appendages, glabrous to moderately pubescent with similar short or long eglandular trichomes as the stems. Corolla 5 (–6)-merous, (5–) 6.5–8 mm long, 10–15 (–20) mm in diameter, dull white or greenish-white, occasionally with purple spots outside and within, stellate with interpetalar membrane, lobed nearly 2/3 of the way to the base, glabrous adaxially and abaxially, the tube (2–) 2.5–3 mm long, the lobes 3.5–5 mm long, 2–3.5 mm wide, triangular, spreading, the margins papillate, the tips acute, cucullate, papillate. Stamens 5 (–6), equal; filaments 1–1.3 mm long, white, cream or purple, inserted on the corolla 1–1.5 mm from the base, with auricles fused to the corolla at the point of insertion; anthers (1–) 1.38–2.05 mm long, blue or bluish-grey, very rarely yellow or greenish-white, broadly ellipsoid or ellipsoid, connivent or not connivent at anthesis, the connective sometimes wide and clearer. Gynoecium with the ovary 2–2.5 mm long, 2.5–3.5 mm in diameter, 2–3 (–4)-carpelar, light green, subglobose; ovules more than two per locule; nectary ca. 0.5 mm tall; styles heteromorphic, 3–4.1 mm, included, at the same level to the stamen length or exserted ca. 2 mm beyond the anthers, lilac or white, cylindrical; stigma ca. 0.15 mm long, ca. 0.5 mm wide, minute, discoid, pale green or pale yellow. Berry < 10 mm in diameter, subglobose and orange to red (in wild populations), highly variable in size, shape and colour (in semi-domesticated or domesticated cultivars): subglobose or triangular, 10–20 mm in diameter, to long-triangular or campanulate, 30–60 (–100) mm long, 20–30 mm in diameter, some blocky, with the apex pointed, blunt or long-acuminate and upcurved and the base obtuse or truncate, green, yellow, brown or purple when immature, pale yellow, yellow, dark brown, orange, red or vermilion-scarlet at maturity, deciduous or persistent, very pungent (sometimes non-pungent), the pericarp thick, opaque, with giant cells (endocarp alveolate); stone cells absent; fruiting pedicels 15–45 (–55) mm long, thick, angled, strongly widened distally, erect and rigid (wild) or pendent and curved (domesticated); fruiting calyx 5–10 mm in diameter, persistent, not accrescent, discoid or shallowly cup-shaped, sometimes reflexed, with a strong annular constriction at junction with the pedicel (wild and domesticated), sometimes the margin ripped, green. Seeds 14–35 per fruit, (2.7–) 3–4 mm long, (2.5–) 3–3.5 mm wide, C-shaped or subglobose, pale yellow or nearly white, the seed coat smooth (SM), cerebelloid (SEM), the cells irregular in seed body, polygonal at margins, the lateral walls sinuate in the seed body, straight at margins; embryo imbricate.

Figure 47. 

Capsicum chinense (A–H from wild plants I–M from domesticated plants) A plant B flower buds C flower on geniculate pedicel D flower, in front view E flower, in full anthesis F immature fruit G fruiting pedicel H mature fruit I flowering branch J flowers on pendent pedicels K, L immature fruits M mature fruit A–H no specimen vouchers (cult. in Banco de Germoplasma de Hortaliças, Embrapa/Hortaliças, Brasília-DF, Brazil), photos taken in situ by L. Bianchetti I–M no specimen vouchers (cult. in Pairumani, Cochabamba-Bolivia), photos taken in situ by G.E. Barboza.

Distribution

Capsicum chinense wild forms were thought to originally occur in the north-central Amazon Basin lowlands (Brazil), where domestication is thought to have occurred (Pickersgill 1971; Eshbaugh 1993). Pickersgill (1984) stated that wild C. chinense is confined to the lowlands of the Amazon, Orinoco and eastern Brazil, while Moses et al. (2014) suggested that populations in Central America and the Caribbean may have been primarily derived from progenitors from the Upper Amazon Region and later diverged through geographical isolation. Wild populations can still be found in the nature, but they are difficult to locate or are restricted to remote areas. Wild C. chinense (indigenous name: Pimi’ró) have been found in Roraima State (Brazil) (Barbosa et al. 2002, 2008; Busso et al. 2003; Nascimento-Filho et al. 2007; Romero da Cruz and Forni Martins 2015), which confirms the occurrence of probable ancestors in the eastern lowlands of Amazonia (Fig. 48).

Figure 48. 

Distribution of C. chinense.

Capsicum chinense has been introduced into United States of America (Bosland and Votava 2000), Mexico through Cuba (González Estrada et al. 2010; Ruiz-Lau et al. 2011; López Castilla et al. 2019) and Central and South America where is found cultivated or escaped from cultivation; it has also been introduced outside the Americas (Eastern Europe, Africa, Asia: China, Japan, Taiwan, India and others) mainly by Portuguese explorers (Eshbaugh 1983; Andrews 1993; Meghvansi et al. 2010).

Ecology

Capsicum chinense grows in wet tropical and subtropical forests where native communities cultivate it around their chakras or in-home gardens, between 100 and 800 (–1,800) m elevation. Wild populations in northern Brazil grow in lowlands and disperse spontaneously in fallow agriculture areas, before the beginning of the rainy period (Barbosa et al. 2002).

Phenology

Flowering and fruiting all year in most parts of its range.

Chromosome number

n = 12 (Pickersgill 1966; Pozzobon and Schifino-Wittman 2006); 2n = 2x = 24 (Pickersgill 1977; Moscone et al. 2003, 2007; Pozzobon et al. 2006; Romero da Cruz and Forni Martins 2015).

Common names

Argentina: Ají bolita (Salta, Hilgert 2060). Belize: Jabanero pepper (Corozal, Balick 2211); Bolivia: Ají (Beni, Williams 937; La Paz, Williams 774), Aji mote (Santa Cruz, Krapovickas & Schinini 32488), Ají soliman (Cochabamba, Thomas 1093), Ají trompillo (Cochabamba, Thomas 704), Ají grande rojo (Beni, Rivero 428); Brazil: Baianinha (Rondônia, Teles Walter 579), Canaimé, Chumbinho, Malaguetão, Morangão, Moranguinho, Murici, Murupi, Pimentãozinho, Peão-amaerlo, Peão-vermelho, Peixe-boi, Pimenta amarela, Pimenta-moranga, Pimenta murici, Pimenta ornamental, Pimenta-roxa, Chifre-de-carneiro, Chifre-de veado, Cumari-do-Pará, Esporão-de-galo, Olho-de-cará, Olho-de-carangueijo, Olho-de-mutum, Olho-de-peixe (falsa), Olho-de-peixe (verdadeira), Pimenta-vermelha longa, Unha-de-gato, Pimenta-de-cherio amarela (ardosa), Pimenta-de-cheiro longa, Pimenta-de mesa-ardosa (Roraima, Barbosa et al. 2006), Pimenta (Goiás, Amaral 139; São Paulo, Hoehne s.n.), Habanero vermelha (São Paulo, Lélis et al. 12), Pimenta açaí (Amapá, Pereira et al. 1918), Pimenta acerola (Amapá, Pereira et al. 1816), Pimenta-biquinho (Mina Gerais, Goiás, Carvalho et al. 2006), Pimenta boliviana (Rondonia, Teles Walter et al. 572), Pimenta camapú (Amapá, Pereira et al. 2011), Pimenta cambuí (Bahía, Medeiros et al. 196), Pimenta-doce (Rondônia, Teles Walter 567), Pimenta habanero (Carvalho et al. 2006), Pimenta-murupi (Região Norte, Carvalho et al. 2006), Pimenta queimosa (Amapá, Pereira et al. 1905), Pimenta do Curupira (Roraima, Barbosa et al. 2002; Nascimento Filho et al. 2007), Pimenta amarela ardosa (Amapá, Pereira et al. 1908), Pimenta de bode (Região Centro-Oeste, Carvalho et al. 2006), Pimenta de cheira (Pickersgill RU72-175), Pimenta de cheiro (Bahía, Medeiros et al. 182; Rio de Janeiro, Gonçalves & Kaketru 2; Paraíba, Gadelha Neto 1748), Pimenta queimosa amarela (Amapá, Pereira et al. 1904), Pimenta roxa alongada (São Paulo, Hoehne s.n.), Pimenta de cheiro amarela (Amapá, Pereira et al. 1901), Pimenta de cheiro legítimo (Río de Janerio, Freire de Carvalho s.n.), Pimenta de cheiro redondinha (Pereira et al. 2011), Pimenta de cheiro tacacá (Amapá, Pereira et al. 2011), Pimenta-de-cheiro vermelha (Amapá, Pereira et al. 1831; Roraima, Barbosa et al. 2006), Pimenta-dedo-de-papagaio (Bahía, Medeiros et al. 200), Pimenta de cheiro de panela (Amapá, Pereira et al. 2011); Colombia: Ají (Meta, Quevedo et al. 1816, Santander, Rosado 01), Trompito (Vichada, García Guzmán s.n.), Aji amarillo (Amazonas, Cárdenas et al. 9439), Ají pecas (Caquetá, Cárdenas et al. 9311), Aji trompito (Caquetá, Cárdenas et al. 9356), Ojito de Lucía (Caquetá, Cárdenas et al. 9331), Ají de babilla (Amazonas, Cárdenas et al. 9431); Costa Rica: Chile, Chile chiricano, Chile panameño (Bohs 2015); Ecuador: Ají (Napo, Ríos 429; Morona-Santiago, Villegas & Meneses 16), Ají botoncillo (Guayas, Holmgren & Heilborn s.n.); French Guiana: Péppé (Cayenne, Fleury 124); Jamaica: Scotch bonnet (Jamaica, Bosland and Votava 2000); Mexico: Habanero (Quintana Roo, Palma Gutiérrez 85-69), Rosita (Yucatán, López Castilla et al. 2019), Chile cimarrón (México, Hinton 4336), Chile costeño (Campeche, Ramírez A. 56), Chile habanero (Yucatán, Ucan 4652), Chile xalapeño (Quintana Roo, Palma Gutiérrez 85-25), Chile de árbol (Michoacán, Soto Núñez & Silva R. 3842); Panamá: Ají Chombo (Bosland and Votava 2000); Peru: Ají (Lambayeque, Llatas Quiroz 3454; San Martín. Plowman & Schunke V. 11695), Arequipeño (Lima, Velarde Núñez 8), Charapita (Iquitos, Bosland and Votava 2000), Chinchi-uchu (Peru, Andrews 1984), Esticana (Cuzco, Ferreyra 16384), Limos (Lima, Velarde Núñez 16), Pucunuchu (San Martín, Plowman & Schunke 11497), Ají amarillo (Pasco, Huamán et al. 0317), Ají bravo (Loreto, Plowman 2457), Aji Charapa (Iquitos, Bosland and Votava 2000), Ají Dulce (Loreto, Plowman 2458), Ají mutecillo (Amazonas, Campos & Vargas 6945), Ají rojo o ají chanchamayo (Pasco, Oxapampa, Huamán et al. 0320), Ají Panca (Bosland and Votava 2000), Ají Pucomucho (Bosland and Votava 2000), Ajuju aji (Loreto, Hormia 2194), Colorado Cuzco (Lima, Velarde Núñez 4), Colorado Trujillo (Lima, Velarde Núñez 9), Munición uchu (Loreto, Williams 5225), Rojo Arequipa (Lima, Velarde Núñez 12), Ají corazón de pollo (Madre de Dios, Pesha Baca 95); Puerto Rico: Rocotillo (Bosland and Votava 2000); Trinidad and Tobago: Congo (Bosland and Votava 2000); United Stated of America: Datil (Florida, Bosland and Votava 2000), Rocotillo (Texas, Andrews 1984).

Indigenous names

Bolivia: Bidó (Tacana, La Paz, Williams 1177), Cheti (Trinitario, Cochabamba, Thomas 704), Ta’ (Beni, Davis 1066), Hchetgi (Trinitario, Cochabamba, Thomas 1092), Tata (Yuqui, Cochabamba, Martínez et al. 246), Uchu (Quechua, La Paz, Williams 772), Winno (Yuracare, Cochabamba, Thomas 1093), Winno manera (Yuracare, Cochabamba, Thomas 704); Brazil: Pimi’ró (Macuxi, Roraima, Barbosa et al. 2002; Nascimento Filho et al. 2007); Colombia: Aati (Curripaco, Guainía, Espina et al. 182); Chamouiju (Colono Huitoto, Caquetá, Cárdenas et al. 9331), Doehoe (Amazonas, Cárdenas et al. 9439), Duaó (Caquetá, Cárdenas et al. 9385), Gubsovia (Makuna, Amazonas, Cárdenas et al. 9431), Kukunarí (And, Amazonas, Castro & Matapí 565), Jade deé (Mui, Amazonas, Castro & Ramírez 132), Kukunari (Amazonas, Castro & Matapí 565), Fakiki deé (Mui, Amazonas, Castro & Rodríguez 309), Lole-ra (Norte de Santander, Solange 8), Tada tarre (Sáliba, Casanare, Camargo 020), Wijichi-taku (And, Amazonas, Castro & Matapí 561); Ecuador: Atalba ucho (Kichwa, Sucumbíos, Moya & Reyes 207), Ampi (Shuar, Zamora-Chinchipe, Santín et al. 102), Ccoma (Cofán, Sucumbios, Cerón 20848), Giimo (Oncaye, Napo, Davis & Yost 993), Giimo (Huaorani, Napo, Miller & Yépez 651), Giimoñabu (Huaorani, Napo, Miller & Yépez 651), Guimo (Huaorani, Orellana, Naranjo & Freire 329), Giimohue (plant) (Huaoroni, Napo, Miller & Yépez 651), Guimuwe (Huaorani, Orellana, Freire & Naranjo 612), Hueapia (Secoya, Cerón 62013), Imiá (Shuar Jívaro, Pastaza, Lewis et al. 13960), Jinia (Shuar, Morona Santiago, Pujupet RBAE 1005), Juminialinae ([Guahibo?] Casanare, Camargo 020), Khoma (Kofán, Napo, Pinkley 230), Munisión (Kichwa, Orellana, Carrillo & Reyes 434), Ochabia (Siona, Sucumbíos, Miranda & Moya 437), Piujimia (Orellana, Herrera & Guerrero 186), Ucho (Quechua, Napo, Miller et al. 2404), Úchu (Achuar Jívaro, Pastaza, Lewis et al. 13960), Aji ucho (Quechua, Orellana, Reyes & Carrillo 773), Anya bia (= snake pepper) (Napo, Vickers 178), Arara uchu (Quechua, Orellana, Kohn 1613), Araya ucho (Quechua, Sucumbíos, Reyes & Moya 234), Araray-uchu (Quechua, Napo, Balslev & Irvine 4598), Biruti uchu (Quechua, Orellana, Kohn 1611), Butun uchu (Quechua, Napo, Ríos & Oña 430), Chipiri khuma (Kofán, Napo, Pinkley 256), Cua Khoma (Kofán, Napo, Pinkley 544), Palanda uchu (Quechua, Orellana, Kohn 1612), Puca uchu (Quechua, Orellana, Kohn 1896), Sasi uchu (Quechua, Orellana, Kohn 1611), Tiupi tio (Cayapa, Esmeraldas, Kvist 40566), Tɛsi khuma (Kofán, Napo, Pinkley 257), thothao khuma (Kofán, Napo, Pinkley 259), Tota cu’ma (Kofán, Napo, Cerón et al. 5849), Uchu muyu (Quechua, Napo, Ríos 429), Wea bia (Siona, Napo, Vickers 115), Bula muyu uchu (Quechua, Orellana, Kohn 1910), Dio tape fin chuno (Cayapa, Esmeralda, Kvist 40586); Guyana: Há-ka-pu-tá (Cuyuni-Mazaruni, Tillett & Tillett 45424); Tor-tor-oi-ma (Cuyuni-Mazaruni, Tillett & Tillett 45627); Mexico: Habanero ik (Maya, Quintana Roo, Gutiérrez 24), Maax-ik (Maya, Quintana Roo, Gutiérrez 388), X-Kat ik (Quintana Roo, Palma Gutiérrez 85-24); Peru: Imia (Achual Jívaro, Loreto, Lewis et al. 11207), Tots (Yanesha, Pasco, Huamán et al. 0317), Yaájima (Amazonas, Kayap 652); Surinam: Ah-se-se (Kaxuyana, Kwamala, Plotkin 139), Pepra (Sranam, Kwamala, Plotkin 139), Pom-we (Tirio, Kwamala, Plotkin 139).

Uses

Capsicum chinense has medicinal, ornamental and food uses. Indigeneous communities and rural people widely cultivated this species around their homes to consume the fruits raw, cooked or used as a spice (Barbosa et al. 2002). The consumption of the fruits of this species has spread globally and are highly appreciated (sweet and pungent) due to its fleshiness and distinct aroma (Kurian and Starks 2002; Basu and De 2003; Gahungu et al. 2011) in sauces and powders.

In addition to medicinal properties indicated in Table 3, C. chinense has spiritual uses which have sometimes been recorded by researchers on herbarium specimen labels. For example, in Ecuador (Comunidad Oncaye, Davis & Yost 993), fruits were used by a shaman’s wife in the termination of an ayahuasa session, (Comunidad Bataburo, Freire & Naranjo 612), the entire plant is used to cure “espanto” (fearfulness) and (Comunidad Indillama, Reyes 502) and the plant is used to cure illness caused by the “mal viento”, an evil spirit of the air. In Peru (Comunidad Palcazú, Huamán et al. 317), the smoke of a single dried fruit is used to help children when they are tearful. In Brazil (Roraima; Comunidad Macuxi, Barbosa et al. 2002), the common name of the species is an allusion to ‘curupira’, a supernatural being, guardian of the Amazon forests in Tupi mythology (Gomez Platero and Palma Ehrichs 2011). In addition, outside the Americas, C. chinense has a broad spectrum of ethnopharmacological applications and medicinal properties (e.g. see Meghvansi et al. 2010).

Preliminary conservation assessment

EOO (18,657,305 km2); AOO (712 km2). Capsicum chinense is a very widespread cultivated species worldwide; we assign the status of Least Concern (LC).

Discussion

Capsicum chinense belongs to the Annum clade (Carrizo García et al. 2016). It is most popularly known as Habanero (Mexico and United States of America) or Scotch Bonnet (Caribbean Islands), but it has numerous common and indigenous names in South America (see above and Baba et al. 2016), as well as landrace and cultivar names (e.g. Carolina Reaper, Bhut Jolokia, Red Savina, Congo pepper, NuMex Suave, NuMex Trick-or-Treat). Many of the names refer to differences in the fruit’s pungency, shape or colour (Bosland and Votava 2000; Votava and Bosland 2004; Bosland and Baral 2007; Bosland et al. 2012; Bosland and Coon 2015).

Ever since Smith and Heiser (1957) first recognised C. chinense as a cultivated species in the genus, its recognition as an independent entity and as a member of the C. annuum complex (C. annuum-C. frutescens-C. chinense) has been debated in literature (see Baral and Bosland 2004; Eshbaugh 2012 and references therein), with arguments based on morphological, cytogenetic, biochemical (isoenzymes), phylogenetic and reproductive evidence. Recently, Raveendar et al. (2017) established the phylogenetic relationships of 13 complete chloroplast genome sequences belonging to the Solanaceae (five Capsicum species included). This study showed that the C. chinense cp genome is much closer to C. annuum var. glabriusculum, wild progenitor of C. annuum, than any other sampled Capsicum species (C. annuum var. annuum, C. frutescens, C. baccatum and C. lycianthoides). This indicates that members of the C. annuum complex share a common gene pool as was suggested by Eshbaugh (1983), Ibiza et al. (2012) and others.

An unequivocal morphological delimitation for C. chinense is difficult for two reasons: first, intermediate well-established types occur between wild and domesticated forms, probably due to intraspecific hybridisation (Pickersgill et al. 1979) and second, some specimens are hardly indistinguishable from their closest relatives (C. frutescens and C. annuum) at flowering or fruiting stage (Smith and Heiser 1957; Eshbaugh 1976; Baral and Bosland 2004; Pickersgill 2016). Capsicum chinense is a short-lived subshrub or rarely shrub with 2–4 (–5) flowers per node, flowering pedicels that can be erect or spreading and geniculate at anthesis (mostly in wild forms) or pendent and non-geniculate (mostly in domesticated cultivars), a calyx with strongly-marked main veins that lacks appendages or is pentagonal or hexagonal in outline with 5–6 mucro-like appendages, a corolla that is dull white or greenish-white, fruits that are variable in shape, size and colour, deciduous or persistent, a fruiting calyx with a strong annular constriction at the junction with the pedicel and a clearly discoid fruiting calyx. The discoid calyx with a distinct annular constriction is the most conspicuous and consistent feature of C. chinense (Fig. 47).

Morphologically, Baral and Bosland (2004) found C. chinense produces primarily pendent flowers and calyces with a circular constriction, while C. frutescens produces erect flowers and calyces without a constriction. Another character found in literature to distinguish these species is the degree to which the style is exserted beyond the anthers (D’Arcy 1973; Eshbaugh 2012). It is most difficult to distinguish these two species at the flowering stage. The great number of specimens of C. chinense, analysed in this treatment, showed that the position of the flowering pedicels and whether or not the style is exserted more than 1 mm beyond the anthers can be quite variable and prevents confident species assignment (Peña-Yam et al. 2009). Calyx characters can be helpful in the identification of flowering specimens. The flowering calyx in both C. chinense and C. frutescens is deeply cup-shaped, but in C. chinense, the main nerves protrude remarkably from the calyx surface (Fig. 47C, J, L), while in C. frutescens, they are often completely immersed in the calyx surface (Fig. 68C, G) or may protrude distally. In both species, these nerves may slightly exceed (less than 1 mm) the calyx edge. In fruit, these species differ clearly. Capsicum chinense has erect and uniformly widened pedicels (wild/domesticated forms, Fig. 47F–H) or pendent pedicels with one or two swellings distally (domesticated/cultivated forms, Fig. 47L–K), but in both cases, a noticeable circular constriction at the junction with the fruiting calyx is present. In addition, the calyx flattens completely (discoid calyx) and remains appressed or reflexed to the fruit base (Fig. 47H, M). In contrast, in C. frutescens, the pedicels are usually erect and widen in a similar way to the wild form of C. chinense, but the circular constriction is totally absent and the fruiting calyx remains deeply cup-shaped, housing the narrowed base of the elongate fruit (Fig. 68G, H).

The differentiation of C. chinense from C. annuum (wild or domesticated specimens) is sometimes also difficult, especially at the fruiting stage. Wild forms of C. chinense with small red fruits and erect pedicels could be confused with some domesticated (or semi-domesticated) specimens of C. annuum var. glabriusculum, which share the same pedicel position and fruit characteristics (size, shape and colour), although wild forms of this variety have smaller fruits than the typical C. chinense. The presence/absence of the constriction in the pedicel/calyx junction (lacking in C. annuum var. glabriusculum) allows the assignment to one or other taxon. Furthermore, C. chinense is a more robust plant with larger leaves and flowers than C. annuum var. glabriusculum.

During its long history of cultivation, C. chinense fruits have developed a broad range of variation due to active selection by growers, differing cultivation methods and adaptation to the environment (Barbosa et al. 2010); in addition, there may have been historical and geographical isolation of subpopulations (Bharath et al. 2013). This variation has been observed along its distribution (Bharath et al. 2013; Baba et al. 2016; Moreira et al. 2018), where elongate, campanulate and blocky fruit shapes are more predominant than triangular or rounded shapes. Similarly, red fruits, followed by orange or yellow ones, are the most common, with rare occurrences of white, orange-yellow, black or brown fruits.

The pungency and aroma of this species are also remarkable. The “Carolina Reaper” cultivar is reputed to be the world’s hottest chile, which rates at an average of 1,641,183 Scoville Heat Units (SHU) (https://www.guinnessworldrecords.com/world-records/hottest-chili, accessed on 29 October 2019) and is a favourite amongst hot-pepper lovers. At the other extreme, the fruits of the low pungency varieties (pimenta-doce, pimenta-de-cheiro longa, pimenta-de-cheiro amarela) found in Brazil (Roraima) are used by indigenous and non-indigenous communities and are preferentially used in stews or salads, for preparing jellies or liqueurs or for food dish decoration (Barbosa et al. 2006). The fruits are also distinguished by their distinct volatile profiles (depending on the cultivars), which give them a powerful exotic-fruity aroma (Rodríguez-Burruezo et al. 2010; Garruti et al. 2013 and references therein; Patel et al. 2016).

Baral and Bosland (2004) provided morphological, phylogenetic (RADP) and reproductive (sexual compatibility) evidence to differentiate C. chinense from C. frutescens, while a study of genetic diversity (SSR and AFLP) (Ibiza et al. 2012) has shown that, although C. chinense and C. frutescens are closely related, the molecular characterisation obtained is sufficient to differentiate the individual members of these species. Intermediate phenotypes and intermediate genetic accessions were explained by hybridisation, since gene flow could be occurring in sympatric areas of these two species (Baral and Bosland 2004; Ibiza et al. 2012).

Jacquin (1776) described C. chinense, based on cultivated material. In the protologue, he stated that he saw this species cultivated on the Island of Martinique (Lesser Antilles), a small island in the Caribbean Sea, but he called it C. chinense, alluding to the region to which he thought the plant was native. We found probable original material only at W (W-acc. # 0080115). This sheet has two labels, one smaller at the lower left corner with the script “Capsicum sinense”, probably in Jacquin’s filius writing (see D’Arcy 1970) and another at the lower right corner where “Capsicum sinense” and “Hort. bot. Vind.” are handwritten by an unknown hand; this sheet is also annotated “Hb. Jacq.” in a different hand and clearer ink. We designate this sheet (W-acc. # 0080115) as the lectotype for this species.

Capsicum cerasiforme was published by Lamarck (1794) with only the type locality “E Brafilia” cited. At P, we found a collection in Lamarck’s Herbarium with a label indicating that it belongs to “Capsicum cerasiforme” (P00357733) which we designate as the lectotype.

For C. luteum, Lamarck (1794) cited in the protologue “Ex India. Sonnerat” and the vernacular names ‘piment jaune’ and ‘Le piment de Mozambique’. At P, we found a sheet in Lamarck’s Herbarium with origi