Table of Contents
Eshbaugh, W.H. 1993. History and exploitation of a serendipitous new crop discovery. p. 132-139. In: J. Janick and J.E. Simon (eds.), New crops. Wiley, New York.
Peppers: History and Exploitation of a Serendipitous New Crop Discovery*
W. Hardy Eshbaugh
- CAPSICUM PUBESCENS
- CAPSICUM BACCATUM VAR. PENDULUM
- CAPSICUM ANNUUM VAR. ANNUUM--CAPSICUM CHINENSE
- Table 1
- Fig. 1
Few could have imagined the impact of Columbus' discovery of a spice so pungent
that it rivaled the better known black pepper from the East Indies.
Nonetheless, some 500 years later, on the quincentennial anniversary of the
discovery of the New World, chili peppers (Capsicum) have come to
dominate the world hot spice trade and are grown everywhere in the tropics as
well as in many temperate regions of the globe. Not only have hot peppers come
to command the world's spice trade but a genetic recessive non-pungent form has
become an important "green" vegetable crop on a global scale especially in
The New World genus Capsicum is a member of the Solanaceae, a large
tropical family. Various authors ascribe some 25 species to the genus but this
is only an estimate with anticipated new species to be discovered and named as
exploration of the New World tropics expands. Exploration and plant collecting
throughout the New World have given us a general but false impression of
speciation in the genus. Humans unconsciously selected several taxa and in
moving them toward domestication selected for the same morphological shapes,
size, and colors in at least three distinct species. Without the advantage of
genetic insight these early collectors and taxonomists named these many size,
shape, and color forms as distinct taxa giving us a plethora of plant names
that have only recently been sorted out reducing a long list of synonymy to
four domesticated species. The early explorations in Latin America were
designed to sample the flora of a particular region. Thus, any collection of
Capsicum was a matter of chance and usually yielded a very limited
sample of peppers from that area. Only with the advent of collecting trips
designed to investigate a particular taxon did the range of variation within a
species begin to be understood. One needs only to borrow specimens from the
international network of herbaria to appreciate what a limited sample exists
for most taxa, particularly for collections made prior to 1950. The
domesticate Capsicum pubescens, for example, that is widespread in the
mid-elevation Andes from Colombia to Bolivia, is barely represented in the
herbarium collections of the world. Most herbarium collections of
Capsicum, with the exception of Capsicum annuum holdings, are
woefully inadequate. Furthermore, besides Capsicum annuum, very little
attention has been paid to the many cultivars of each of the domesticated
species. Often material is unusable because it was collected only in fruit
neglecting the most important and critical characters associated with floral
anatomy and morphology. With the advent of germplasm collecting programs
during the past three decades, and concomitant improvement in herbarium
collections we have come to better understand the nature of variation in the
genus Capsicum. The increasing number of Capsicum herbarium
specimens permits renewed interest and debate on the proper species
One of the more perplexing questions regarding the taxonomy of Capsicum
is defining the genus (Eshbaugh 1977, 1980b; Hunziker 1979). The taxonomy of
the genus Capsicum is confounded within certain species complexes, e.g.
C. baccatum sensu lato. Major taxonomic difficulties below the
species level in other taxa, e.g. C. annuum, also exist. Armando T.
Hunziker (unpublished) is currently working on a revision of the genus. What
taxa are ultimately included in Capsicum may indeed change if the
concept of the genus is broadened to include taxa with non-pungent fruits but
with other common morphological and anatomical traits such as the nature of the
anther, the structure of nectaries, and the presence of giant cells on the
inner surface of the fruit (Pickersgill 1984).
Capsicum, as presently perceived, includes at least 25 species, four of
which have been domesticated (Table 1). An understanding of each of these
domesticates is instructive when trying to appreciate their origin and
evolution. The data from plant breeding and cytogenetics confirm that the
domesticated species belong to three distinct and separate genetic lineages.
Earlier studies suggested two distinct lineages based upon white and purple
flowered groupings (Ballard et al. 1970) but an evaluation of more recent data
argues for the recognition of three distinct genetic lineages. Although the
barriers between these gene pools may be broken down this rarely, if ever,
occurs in nature.
Capsicum pubescens forms a distinct genetic lineage. This pepper, first
described by Ruiz and Pavon (1794) never received wide attention from
taxonomists until recently (Eshbaugh 1979, 1982). Morphologically, it is
unlike any other domesticated pepper having large purple or white flowers
infused with purple and fruits with brown/black seeds. Genetically, it belongs
to a tightly knit group of wild taxa including C. eximium (Bolivia and
northern Argentina), C. cardenasii (Bolivia), and C. tovarii
(Peru). Capsicum pubescens is unique among the domesticates as a
mid-elevation Andean species. Capsicum pubescens is still primarily
cultivated in South America although small amounts are grown in Guatemala and
southern Mexico, especially Chiapas. This species remains virtually unknown to
the rest of the world. A small export market seems to have reached southern
California. Two of the major difficulties in transferring this species to
other regions include (1) its growth requirements for a cool, freeze free
environment and long growing season and (2) the fleshy nature of the fruit that
leads to rapid deterioration and spoilage.
Capsicum baccatum var. pendulum represents another discrete
domesticated genetic line. Eshbaugh (1968, 1970) notes that this distinct
South American species is characterized by cream colored flowers with
gold/green corolla markings. Typically, fruits are elongated with cream
colored seeds. The wild gene pool, tightly linked to the domesticate, is
designated C. baccatum var. baccatum and is most common in
Bolivia with outlier populations in Peru (rare) and Paraguay, northern
Argentina, and southern Brazil. This lowland to mid-elevation species is
widespread throughout South America particularly adjacent to the Andes. Known
as aji, it is popular not only as a hot spice but for the subtle bouquet and
distinct flavors of its many cultivars. This pepper is little known outside
South America, although it has reached Latin America (Mexico), the Old World
(India), and the United States (Hawaii). It is a mystery as to why it has not
become much more wide spread, although the dominance of the Capsicum
annuum lineage throughout the world at an early date may be responsible.
Pickersgill (1988) has stated that "the status of Capsicum annuum, C.
chinense, and C. frutescens as distinct species could legitimately
be questioned." Several authors have previously raised this issue culminating
in the observation that "at a more primitive level one cannot distinguish
between the three species. On the one hand we treat the three domesticated
taxa as separate while the corresponding wild forms intergrade to such an
extent that it is impractical if not impossible to give them distinct taxonomic
names" (Eshbaugh et al. 1983). McLeod et al. (1979, 1983) have argued that
isoenzyme data make it impossible to distinguish between these three taxa.
From an extensive isoenzyme study of these three taxa and several other
species, Loaiza-Figueroa et al. (1989) argue that "thus far, this substitution
of alleles constitutes a good argument against the proposal that these species
form an allozymically indistinguishable association of a single polytypic
species" as advanced by Mcleod et al. (1982, 1983) and Eshbaugh et al. (1983)
in their published studies. Nonetheless, Pickersgill (1984) has pointed out
that "each domesticate intergrades with morphologically wild accessions by way
of partially improved semidomesticates. Any subdivision of the wild complex
into three taxa, each ancestral to one of the domesticates, becomes decidedly
arbitrary, although clusters corresponding to wild C. annuum, C.
chinense, and wild C. frutescens can be detected." Clearly,
Loaiza-Figueroa et al. missed the point of these earlier papers which argue for
the complexity of the problem noting that the real difficulty comes as one
approaches the more primitive forms of these taxa. Furthermore, the
Loaiza-Figueroa et al. (1989) dendrogram (p. 183) suggests that the number of
C. chinense and C. frutescens taxa included in their study is
insufficient to reach any definitive conclusion regarding the status of these
three taxa. There is a very close relationship of these three taxa based on
crossing data from several studies (Smith and Heiser 1957; Pickersgill 1980).
Stuessy (1990) has observed that "the ability to cross does not just deal with
a primitive genetic background; it deals with the degree of genetic
compatibility developed in a particular evolutionary line." As Stuessy (1990)
has inferred there can be no stronger argument for relationship than the data
obtained from plant breeding. Regardless of one's viewpoint, it is clear that
the C. annuum--C. chinense--C. frutescens complex has been
and continues to be a most difficult taxonomic morass. Some preliminary
information from the studies of Gounaris et al. (1986) and Mitchell et al.
(1989) suggest that molecular data may be useful in resolving this and other
taxonomic questions. For the present, I have chosen to recognize the
Capsicum annuum complex and the Capsicum chinense complex as two
distinct domesticated species. Where C. frutescens fits into this
scenario remains to be resolved. William G. D' Arcy, A.T. Hunziker, and
others may solve the problem by merging the three taxa under a single taxonomic
entity. Taxonomists and formal taxonomy are having a very difficult time
coping with what is a complex and dynamic evolutionary process. The problem is
heightened by the economic importance of Capsicum and the requirement
that not only the domesticated species be named properly but that the several
cultivars receive taxonomic recognition.
Capsicum annuum is the best known domesticated species in the world.
Since the time of Columbus, it has spread to every part of the globe. The
non-pungent form, bell pepper, is widely used as a green vegetable. Another
non-pungent form, "pimento," is also present throughout much of the globe. The
hot spicy forms of this species have come to dominate the spicy foods within
Latin America and the rest of the world. Capsicum annuum probably
became the dominant pepper globally in part because it was the first pepper
discovered by Columbus and other New World explorers (Andrews in press). This
taxon was the first Capsicum species taken to Europe and quickly spread
to other regions.
Capsicum chinense was also discovered at an early date and spread
globally but to a lesser extent than C. annuum. The more limited global
expansion of this species is most probably related to its later discovery in
South America and the competitive edge enjoyed by C. annuum which was
firmly established in the Old World before C. chinense was introduced
A discussion of the geography of Capsicum touches on two questions. The
first relates to the origin of the genus Capsicum and the second to the
origin of the domesticated taxa. The area of origin of Capsicum cannot
be resolved until we understand the nature of the genus. If we accept the
genus as currently circumscribed and limited to pungent taxa, then a clear
center of diversity is to be found ranging from southern Brazil to Bolivia
(McLeod et al. 1982; Eshbaugh et al. 1983; Pickersgill 1984). However, if the
genus is reconstituted to include other non-pungent taxa, another center of
diversity may be recognized in Central America and southern Mexico.
Ultimately, our definition of the genus Capsicum and what species it
includes will determine our view of its center of origins and whether the genus
is monophyletic or polyphyletic. The emerging molecular studies of J.D. Palmer
and R.G. Olmstead should give us a better sense of where Capsicum
belongs within the framework of the Solanaceae.
Determining the place of origin of the genus and each of the domesticated
species is at best a problematic exercise. In 1983, I stated that "it appears
that the domesticated peppers had their center of origin in south-central
Bolivia with subsequent migration and differentiation into the Andes and
Amazonia." This is a condensation of a highly speculative hypothesis (McLeod
et al. 1982). From that hypothesis Pickersgill (1989) later suggested that I
(Eshbaugh 1983) argued that all the domesticated taxa arose in Bolivia.
Without question, I could have stated this idea more clearly. We (McLeod et
al. 1982) have speculatively hypothesized that Bolivia is a nuclear center of
the genus Capsicum and that the origin of the domesticated taxa can
ultimately be traced back to this area. That does not imply that each of the
domesticated species arose in Bolivia. Clearly, evidence supports a Mexican
origin of domesticated C. annuum while the other domesticated species
arose in South America. Nonetheless, the ancestry of the domesticates can be
traced to South America. While McLeod et al. (1982) have hypothesized a
Bolivian center of origin for Capsicum there is no evidence for a
polyphyletic origin of the genus as now understood.
Evidence suggests that C. annuum originally occurred in northern Latin
America and C. chinense in tropical northern Amazonia (Pickersgill
1971). Capsicum pubescens and C. baccatum appear to be more
prevalent in lower South America (Fig. 1). Thus, at the time of discovery, the
former two species were exploited while the later two species awaited a later
discovery and remain largely unexploited outside South America today.
In considering the question of origin of each particular domesticated species
two issues must be considered. First, what wild progenitor is the most likely
ancestor of each domesticated species and second, where is the most probable
site of domestication?
Capsicum pubescens ranges throughout mid-Andean South America. An
analysis of fruit size of this domesticate indicates that fruits of a
statistically smaller size occur in Bolivia, while fruits from accessions
outside Bolivia on the average are somewhat larger suggesting that Bolivian
material approaches a more primitive size (Eshbaugh 1979).
Eshbaugh (1979, 1982) has argued that the origin of this domesticate can be
found in the "ulupicas," C. eximium and C. cardenasii. Clearly,
these two taxa are genetically closely related to each other and C.
pubescens. Natural hybrids between these taxa have been reported and
evaluated (Eshbaugh 1979, 1982). Furthermore, the two species that show the
highest isoenzyme correlation with C. pubescens, C. eximium and
C. cardenasii, occur primarily in Bolivia (Eshbaugh 1982; McLeod et al.
1983; Jensen et al. 1979). All three of these taxa form a closely knit
breeding unit with the two wild taxa hybridizing to give fertile progeny with
viable pollen above the ninety percent level. Crosses between the wild taxa
C. eximium and C. cardenasii and the domesticate C.
pubescens most often show hybrid pollen viability greater than 55%. These
factors lend to the conclusion that domesticated C. pubescens originated
in Bolivia and that C. eximium--C. cardenasii is the probable
ancestral gene pool. This does not prove that these two taxa are the ancestors
of C. pubescens but of the extant pepper taxa they represent the most
logical choice. One perplexing question remains to be investigated and that is
the origin of the brown/black seed coat in domesticated C. pubescens, a
color unknown in any of the other pepper species.
Capsicum baccatum var. pendulum is widespread throughout lowland
tropical regions in South America. It ranges from coastal Peru to Coastal,
Brazil. The wild form, recognized as C. baccatum var. baccatum,
has a much more localized distribution but still ranges from Peru to Brazil.
These two taxa have identical flavonoid (Ballard et al. 1970; Eshbaugh 1975)
and isoenzyme profiles (McLeod et al. 1979, 1983; Jensen et al. 1979) and are
morphologically indistinguishable except for the overall associated size
differences found in the various organ systems of the domesticated taxon
(Eshbaugh 1970). The wild form of Capsicum baccatum exhibits a high
crossability index with domesticated C. baccatum var. pendulum
with the progeny typically exhibiting pollen viability in excess of 55 percent
(Eshbaugh 1970). The greatest center of diversity of wild C. baccatum
var. baccatum is in Bolivia leading to the conclusion that this is the center
of origin for this domesticate.
Can we ever unscramble questions about the origin and evolution of the C.
annuum--C. chinense--C. frutescens species complex?
Pickersgill (1989) states that there is an "overwhelming likelihood of at least
two independent domestications of the chile peppers of this complex." She also
notes that one "may ... argue about whether wild forms of this complex should
really be assigned to different species, and indeed whether domesticated C.
annuum and domesticated C. chinense are really conspecific." I
would agree that the evolutionary lineage of C. annuum--C.
chinense--C. frutescens complex is intimately linked but I would
further emphasize that when, where, and how they diverged is obscured in
antiquity and that the extant wild forms of these three taxa are so similar as
to make them very difficult to separate. One might well ask whether, at a
minimum, C. chinense and C. frutescens are conspecific or grades
within the same species.
In contrast, a reasonably clear picture emerges on origin and progenitor of
C. annuum. Capsicum annuum has its center of diversity in Mexico
and northern Central America with a local, and more recent distribution in
parts of South America. The wild bird pepper, Capsicum annuum var.
aviculare, ranges from northern South America (Colombia) into the
southern United States and Caribbean. Crossing studies indicate that the wild
bird pepper is genetically the most closely related taxon to domesticated C.
annuum (Emboden 1961; Smith and Heiser 1957; Pickersgill 1971).
Pickersgill (1971), using karyotype analysis, suggests that the origin of
domesticated C. annuum is to be found in southern Mexico. Pickersgill
et al. (1979) also provided a detailed phenetic analysis of the C.
annuum--C. chinense--C. frutescens complex and the difficulty
of separating these taxa at the most primitive level is apparent.
Capsicum chinense remains the least understood of the four domesticated
taxa with respect to center of origin and probable progenitor. If one maps the
range of forms in C. chinense, it is clear that amazonian South America
is the center of diversity of this species. Furthermore, C. chinense
does occur sporadically throughout the Caribbean. It is likely that C.
chinense spread into the Caribbean at a later date since the diversity of
taxa is more limited in that region than in amazonian South America. In
considering the progenitor of C. chinense, one is bewildered by the
evidence. It has been suggested that C. frutescens, in its primitive
form, may be the ancestor of C. chinense (Eshbaugh et al. 1983).
However, one needs to ask whether C. frutescens is merely a weedy
offshoot of C. chinense or C. annuum. It is clear that the three
species, C. annuum, C. frutescens, and C. chinense, hybridize
with each other. They form a morphological continuum especially at a primitive
level (McLeod et al. 1979). Genetic evidence from isoenzymes also confirms the
close relationship of these three taxa (McLeod et al. 1983; Jensen et al.
The spread of domesticated peppers throughout the world during the 500 years
since discovery is truly a phenomenon. Two of the domesticated species, C.
annuum var. annuum and C. chinense have been widely utilized
on a global scale. Both C. baccatum var. pendulum and C.
pubescens have been extensively exploited in South America but remain
largely confined to that market. Given both the unique qualities and flavors
of these later two species they each represent a potential source for future
Of special interest to those working with peppers is the use and exploitation
of the wild species. Wherever wild taxa of Capsicum occur, humans use
them for their hot properties. In a few cases, exploitation of wild species
has reached a commercial level. Capsicum praetermissum is collected and
sold commercially in parts of Brazil (reported by correspondents). Capsicum
chacoense and C. eximium are collected and bottled and marketed
throughout southern Bolivia (pers. observ.). Fresh C. cardenasii is
harvested and transported to the La Paz, Bolivia market for sale (pers.
observ.). In Mexico and the southwestern United States wild C. annuum
var. aviculare, the chiltepin, has been locally used for many years
(Nabhan et al. 1989). More recently, a commercial market has developed for
chiltepin. A large amount of this wild species is now harvested and sold to
the gourmet food market. Nabhan et al. (1989) indicate that "currently
chiltepin is almost completely wild harvested." They note that "as much as 12
tons of chiltepines may be harvested from a single Sonoran municipio in a good
year, but total harvest may vary from perhaps 8 to as high as 50 tons."
While the quantity of C. eximium, ulupica, being harvested in southern
Bolivia is unknown, there is an extensive commercial trade in bottled whole
peppers. Bolivians have not attempted to commercially plant wild plants of
C. eximium, but Nabhan et al. (1989) indicate that incipient cultivation
of C. annuum var. aviculare was initiated with extensive planting
of the chiltepin by Sonoran farmers in the 1980s. The manipulation of these
two wild species in each setting has led to some significant changes for the
wild species. In both the case of C. annuum var. aviculare and
C. eximium, larger fruit size has been selected for in the incipient
area of cultivation and manipulation. Sonoran farmers are selecting for larger
fruit size in the wild chiltepin. In C. eximium, there is a
statistically significant larger fruit form of this ulupica in the zone of
exploitation when compared to regions where the fruit is not widely collected
(Eshbaugh 1979, 1982). In both cases we are witnessing incipient or
semi-domestication of the wild species.
Apparently, a market exists for the exploitation of peppers for the medicinal
properties of capsaicin and several companies are pursuing such investigations.
Two of the more interesting products to come to market in the last five years
are the prescription drug Zostrix (Genderm registered trade mark), an analgesic
cream, containing 0.025% capsaicin that is used topically to treat shingles and
to provide enhanced pain relief for arthritis patients and Axsain (GalenPharma
registered trade mark) that contains 0.075% capsaicin and is used for relief of
neuralgias, diabetic neuropathy, and postsurgical pain. Both products are
believed to work by action on a pain transmitting compound called substance
Several pepper species, because of their unique fruit shapes and bright fruit
colors, have been widely used as ornamentals. The presence of capsaicin,
however is a potential hazard.
In August, 1980, an expert consultative group, under the auspices of the IBPGR
(International Board for Plant Genetic Resources), met at CATIE (Centro
Agronomico Tropical de Investigacion y Ensenanza) in Turrialba, Costa Rica, to
discuss the status of Capsicum germplasm collections and to map a
strategy for future collecting and management of these resources (Genetic
Resources of Capsicum 1983). The discussions led to a plan to systematically
collect Capsicum throughout New World paying particular attention to the
wild species most closely related to the domesticated taxa. The efforts of the
past decade have resulted in a significant accumulation of pepper germplasm
(seeds) that is now stored in various collections. Eshbaugh (1980a, 1981,
1988) has detailed the history of Capsicum germplasm collecting prior to
1980 and discussed the collecting efforts of peppers in Bolivia. Capsicum
germplasm collections are now maintained in a number of facilities in the
United States, as well as Mexico, Costa Rica, Bolivia, and Brazil.
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*The investigations cited and drawn upon in this paper were supported most
recently by a grant from the National Science Foundation (BSR 8411136). I am
indebted to many colleagues for their thoughts and comments regarding the
evolution of the genus Capsicum, although I accept sole responsibility
for the comments in this paper. I especially acknowledge Charles B. Heiser,
Jr. who started me on a course of study into the evolution of the genus
Capsicum that has lasted more than 30 years. My wife, Barbara, has been
with me on every step of this journey and deserves a special thanks for all her
support, patience, and understanding.
Table 1. Synopsis of the genus Capsicum
zThe following Capsicum species have been omitted: C.
anomalum, C. breviflorum, and C. ciliatum following the earlier
suggestion of Eshbaugh (1983). Also, C. flexuosum Sendt. has been
treated as a variety of C. schottianum by Hunziker. The treatment of
C. frutescens L. remains to be resolved and some may choose to retain it
as a distinct species of Capsicum while others submerge it into C.
chinense, as suggested in this paper. Finally, C. eximium var.
tomentosum Eshbaugh & Smith is so distinctive that it may deserve
Capsicum ||New world distribution|
|annuum L. ||Colombia north to southern United States|
|baccatum L. ||
Argentina, Bolivia, Brazil, Paraguay, Peru|
|buforum Hunz. ||Brazil|
|campylopodium Sendt. ||South Brazil|
& Smith ||Bolivia|
|chacoense Hunz. ||Argentina, Bolivia,
|chinense Jacq. ||Latin and South America|
(Rusby) Hunz. ||Bolivia, Peru|
|cornutum (Hiern) Hunz. ||South
|dimorphum (Miers) O.K. ||Colombia|
|dusenii Bitter ||
|eximium Hunz. ||Argentina,
|glapagoensis Hunz. ||Ecuador|
Hunz. ||Colombia, Ecuador|
|hookerianum (Miers) O.K. ||
|lanceolatum (Greenm.) Morton & Standley ||Mexico,
|leptopodum (Dunal) O.K. ||Brazil|
(Rusby) Hunz. ||Argentina, Bolivia, Paraguay|
|mirabile Mart ex. Sendt ||
|parvifolium Sendt. ||Colombia, Northeast Brazil,
|praetermissum Heiser & Smith ||South
|pubescens Ruiz & Pav. ||Latin and South
|scolnikianum Hunz. ||Peru|
|schottianum Sendt. ||
Argentina, South Brazil, Southeast Paraguay|
|tovarii Eshbaugh, Smith
& Nickrent ||Peru|
|villosum Sendt. ||South Brazil|
yThis table has been developed, adapted, and modified from Hunziker
1956; Eshbaugh 1977, 1980b; and Pickersgill 1984.
||Fig. 1. Distribution of domesticated species of
Capsicum (solid circles) and wild progenitor taxa (open circles).
Capsicum chinense and C. frutescens are combined together
because they may represent the same taxon. Copyright Bul. Torrey Bot. Club,
Last update September 9, 1997