The Chilean Strawberry (Fragaria chiloensis): Over 1000 Years of Domestication

Abstract

The cultivated strawberry of South America, the octoploid Fragaria chiloensis L. has a long and interesting history. While the origin of the species in Chile cannot be completely determined, it is suspected to be an introduction from North America by birds. After making landfall in Chile, the species spread from the coast into the mountains developing a number of four ecotypes over a long period. At least two native people, the Mapuche, between Rio Bio-Bio and south-central Chile, and the Picunche, between Rio Itata and Rio Bio-Bio, began the domestication process. While white and red fruited forms were developed, the white form (because of fruit size) seemed to have been preferred as the red fruited types are not mentioned frequently in the literature. At the time of the Spanish invasion of Chile, F. chiloensis was widely grown in small garden plots. Over time under the Spanish rule, larger plantings first of 1–2 ha and later of many hectares were grown. As the Spanish continued their exploration and conquest of South America, they carried F. chiloensis with them up the western coast to Peru, Ecuador and Colombia. For many years these scattered plantings were the source of fresh fruit for the burgeoning populations. The cultivated F. x ananassa Duch. ex Rozier was introduced in Chile around 1830, but the F. chiloensis was still preferentially grown. In the early 1900s, a large canning industry that served hundreds of acres of F. chiloensis planting thrived. By the 1950s, F. x ananassa began to predominate and the rise of the importance of the University of California and European developed strawberries that was impacting the world also displaced much of the traditional production. An increased awareness of this vast genetic resource arose in the 1980s and 1990s. Faculty, particularly at the Universidad de Talca, have collected and characterized germplasm that represents not only tremendous diversity but captures many of the land races that have been developed. This germplasm has been utilized in small commercial plantings (0.1–0.3 ha) and in breeding programs to further develop F. chiloensis commercial cultivars. There is still a small but vibrant community of small growers in Chile (along the North-Central coast of the Pacific Ocean) and Ecuador (mainly around Ambato) producing the land races for commercial sale in local markets. It is estimated that around 30 ha of open field plantings are cultivated in Chile with yields averaging around 3–4 tons/ha.

Full text

The Chilean Strawberry (Fragaria chiloensis): Over 1000
Years of Domestication
Chad E. Finn
1
U.S. Department of Agriculture, Agricultural Research Service, Horticultural Crops Research Unit,
3420 NW Orchard Avenue, Corvallis, OR 97330
Jorge B. Retamales and Gustavo A. Lobos
Centro de Mejoramiento y Feno
´mica Vegetal, Facultad de Ciencias Agrarias,
Universidad de Talca, Casilla 747, Talca, Chile
James F. Hancock
Department of Horticulture, College of Agriculture and Natural Resources, Michigan State University,
A288 Plant & Soil Science Building, East Lansing, MI 48824
Additional index words. Mapuche, Picunche, Fragaria 3ananassa, Ecuador, white-fruited
Abstract. The cultivated strawberry of South America, the octoploid Fragaria chiloensis,hasalongandinteresting
history. Although the origin of the species in Chile has not been completely determined, it may have been introduced from
North America by birds. After making landfall in Chile, the species spread from the coast into the mountains eventually
developing four biotypes. At least two native peoples, the Mapuche, between Rio Bı
´o-Bı
´o and south–central Chile, and the
Picunche, between Rio Itata and Rio Bı
´o-Bı
´o, began the domestication process. Although white- and red-fruited forms
were domesticated, the white form (likely because of its fruit size) may have been preferred because the red-fruited types
are not mentioned as frequently in the literature. At the time of the Spanish invasion of Chile, F. chiloensis was widely
grown in small garden plots. Under the Spanish rule, larger plantings, first of 1 to 2 ha and later of many hectares, were
grown. As the Spanish continued their exploration and conquest of South America, they carried F. chiloensis with them
up the western coast to Peru
´and Ecuador. For many years these scattered plantings were the source of fresh fruit for the
burgeoning human populations. The cultivated F.3ananassa wasintroducedinChile’
’1830 but F. chiloensis was still
preferentially grown. In the early 1900s, a large canning industry emerged serving hundreds of acres of F. chiloensis.By
the 1950s, F.3ananassa began to predominate and the rise in importance of the University of California and European-
developed cultivars displaced much of the traditional F. chiloensis production. An increased awareness of this vast native
Chilean genetic resource arose in the 1980s and 1990s. Scientists at the Universidad de Talca, associated with USDA-ARS
Plant Exploration Office-sponsored trips to Chile, and with El Instituto de Investigaciones Agropecuarias–Cauquenes in
Chile have collected and characterized germplasm that represents not only tremendous diversity, but captures many of the
land races that have been developed. This germplasm has been used in small commercial plantings (0.1 to 0.3 ha) and in
breeding programs to further develop F. chiloensis commercial cultivars. A small but vibrant community of small growers,
particularly in Chile and Ecuador, produce the land races for commercial sale in local markets. Approximately 30 to 40 ha of
open-field plantings are cultivated in Chile with yields averaging ’
’3 to 4 tons/ha. The selected F. chiloensis genotypes and
collected clones from the wild have served as a valuable source of germplasm in modern breeding programs and the
development of new cultivars with the white color and aromatic flavor typical of some of the traditional selections well
underway.
HISTORY
The cultivated strawberry (Fragaria
3ananassa Duch. ex Rozier) originated
from an accidental cross of the white-fruited
Chilean strawberry [F. chiloensis (L.) Mill.
subsp. chiloensis f. chiloensis]andthemeadow
strawberry (F. virginiana Mill. subsp. virginiana)
that occurred in a Royal Botanical Garden in
France (Darrow, 1966; Staudt, 1999). Fragaria
virginiana is an octoploid (2n=8
x= 56)
species native throughout much of North
America where there is sufficient moisture.
Fragaria virginiana’s movement to Europe
cannot be pinpointed to a single event or
person; rather, it seems to have occurred many
times from the settlers from present-day
Virginia up to those along the St. Lawrence
River and may have occurred as early as
when Jacques Cartier explored this region in
1523 (Darrow, 1966; Wilhelm and Sagen,
1974). On the other hand, the colorful history
of the journey of F. chiloensis,alsoanocto-
ploid, to Europe is well described (Fig. 1;
Darrow, 1966; Wilhelm and Sagen, 1974).
Fragaria chiloensis is native to the fog belt
along the Pacific Ocean from Alaska through
British Columbia to the central California
coast and as a disjunct distribution in Chile
along the Pacific Ocean and into the Andes
Mountains from lat. 35°30#S to long. 47°33#S
and between sea level and 1850 m elevation
(del Pozo and Lavı´n, 2005; Lavı´n et al., 2000).
Migrating birds are presumed to have carried
seeds from the North American to South
American coast. Once established on the
Chilean coast, the species moved inland and
differentiated into different forms, often tak-
ing on the phenotype of F. virginiana grow-
ing inland in North America.
In Chile, the mouth of the Biobio River is
near current-day Concepcio´ n. The native peo-
ple in this region, the Picunche, to the north
of the Biobio River, and the Mapuche, to the
south of the Biobio, cultivated strawberries
more than 1000 years ago (Hancock et al.,
1999). The Picunche, who were largely an
agrarian society, were believed to have taught
the Mapuche, who were primarily hunter–
gatherers, about agriculture, including straw-
berry cultivation. Further to the north, the
invading Incas either received or took supe-
rior plants from the Picunche to grow in their
gardens. The fruit in each of these cultures
were consumed fresh, dried, or prepared as
medicines (De Moesbach, 1992). Like other
cultures with access to native sweet fruits, the
people fermented juice of the strawberry.
Labarca (1994) reported that the Mapuche’s
favorite fermented drink (lahuen˜e mushca)
was made from the small, red-fruited native
strawberry called ‘‘llahuen’’ or ‘‘lahuen
˜e.’’
Although there were many uses for the small
red form, over time two distinct cultivated
types began to stand out: a large white and an
improved, but not necessarily large, red form.
Received for publication 25 Jan. 2013. Accepted
for publication 5 Mar. 2013.
This paper was part of the workshop ‘‘A Survey on
the Contribution of Indigenous People of North
America to Horticulture’’ held 1 Aug. 2012 at the
ASHS Conference, Miami, FL, and sponsored by
the History of Horticultural Science Working Group
(HIST).
1
To whom reprint requests should be addressed;
e-mail Chad.Finn@ARS.USDA.gov.
418 HORTSCIENCE VOL. 48(4) APRIL 2013

Although relatively uncommon, native white
forms have been found in multiple sites in
southern Chile (Hancock et al., 1999). Larger
and larger selections of the white form ended
up dominating the cultivated types and when
comparisons of the modern F.3ananassa
cultivated types and the land race types are
made, the modern types genotypically align
with the white forms (Hancock et al., 1999;
Hokanson et al., 2006; Lavin, 1997) more
closelythanwithNorthAmericanF. chiloensis.
Apparently, whereas strawberries were well
known as a native crop and they were commonly
grown in garden plots by the native peoples,
there was not a concerted effort to cultivate
them in large quantities.
When the Spaniards invaded western South
America, they came into conflict with the
Mapuche in 1536, who kept them largely north
of the Bı´o-Bı´o River until the 1880s. The large
elite strawberries were considered a bounty of
conquest and moved with the Spaniards north
to Cuzco, Peru
´in 1557, and to Ecuador before
1789 (Popenoe, 1921). During the colonial
period, strawberry production on a somewhat
larger scale (1 to 2 ha) began to take place
near Renca, just north of Santiago (Hancock
et al., 1999). Surprisingly, although the Chilean
strawberries were introduced to Peru
´in 1557,
and although they were noted by explorers, it
took until 1712 for these superior clones to be
introduced to Europe from Chile (Darrow,
1966; Popenoe, 1921). Ame´de´e Francois
Fre´zier, an engineer in the French Army In-
telligence Corps, was commissioned by King
Louis XIV of France to conduct a reconnais-
sance mission to Chile and Peru
´. The King was
apparently determined to keep his grandson
on the Spanish throne and therefore wanted to
know as much as possible about all parts of
the Spanish empire (Darrow, 1966). Although
not a botanist, Fre´ zier, ‘‘had the botanist’s
impulse for collecting’’ and collected unusu-
ally large-fruited strawberry plants that he
transported back to France (Fig. 2; Darrow,
1966). As the story goes, he nursed the plants
carefully with limited water on the six-month
voyage and returned with five plants to Mar-
seille (Popenoe, 1921). Frezier gave two plants
to the ship’s owner, one plant to the Royal
Garden, one to the minister of fortifications,
and he kept one for himself (Popenoe, 1921).
Most evidence points to strawberries hav-
ing been primarily a garden crop for the native
Chileans, but during the Spanish colonial
period, there were larger 1- to 2-ha plots of
cultivated types raised in coastal valleys from
west of Chilla´ n (lat. 36°34#0$S) to the Isle of
Chiloe´ (lat. 41°40#Sto43°40#S) (Hancock
et al., 1999). Most production was geared
toward local markets until the early 1900s and
fruit was usually transported to the markets by
mule. Popenoe (1921) famously commented
on the quality of the 1920 cultivars grown in
the United States to the native types that had
been taken from Chile and planted in Huachi,
Ecuador (lat. 0°18#S): ‘‘What sorts have we,
may I ask, which could be thrown into boxes
holding 30 to 35 quarts, carried seven or eight
miles on mule back, worked over by hand and
packed in two to six-quart baskets, and then
shipped down to a tropical seaport, there to be
kept in the market for two to three days at
a temperature of 70 to 85 degrees? Even with
such treatments as this, the Guachi strawberry
(ed. Huachi/Guachi is a small community near
Ambato, Ecuador) holds up well. Retaining
its shape and texture to an extent altogether
unknown among northern strawberries.’’
As canning technology developed around
the world, a large canning industry using
F. chiloensis developed near Nueva Imperial,
Chile (lat. 38°43#59$S) and a somewhat smaller
one near Corral, Chile (lat. 39°52#02$S) be-
tween 1900 and 1950. In the mid-1990s, culti-
vars from Europe and from the United States
began to displace the native strawberry (Hancock
Fig. 1. Origin and dispersal of Fragaria chiloensis (Wilhelm and Sagen, 1974).
HORTSCIENCE VOL. 48(4) APRIL 2013 419

et al., 1999). Although the circumstances are
unknown, the breeding records for the USDA-
ARS, Horticultural Crops Unit indicate that
in 1947 and 1948, selections from the USDA-
ARS program, OSC 2074, OSC 2127, and
OSC 2138, were named ‘Quillan’, ‘Maullin’,
and ‘Ancud’, respectively, in Chile (unpub-
lished USDA-ARS breeding records). Like
with so many places around the world, the
meteoric rise of the California strawberry in-
dustry led to California cultivars being adopted
around much of the world and Chile was no
exception. Most of Chile as well as Peru
´and
Ecuador, where the native F. chiloensis was
once such an important part of production,
are now planted in genotypes that trace their
pedigree to California germplasm and are
grown using the same annual, plasticulture
system developed in California. Nonetheless,
small pockets of production of pink- and
white-fruited genotypes of F. chiloensis pro-
duction have held on in places like Curepto
(lat. 35°08#S), Putu
´(lat. 35°13#S), Curanipe
(lat. 35°50#S), Chovelle´n (lat. 35°53#S),
Contulmo (lat. 38°00#S), Pure´n (lat. 38°01#S),
Puerto Saavedra (lat. 38°46#S), and the Island
of Chiloe´ (lat. 41°40#Sto43°40#S), Chile;
Cuzco (lat. 13°45#S), Peru; and Huachi Grande
(lat. 1°18#S), Ecuador (Finn et al., 1998; Lavı´n
et al., 2000).
CURRENT PRODUCTION
There has been in increasing interest in
expanding production of F. chiloensis in Chile
to find a niche in the market as well as to take
advantage of a resurgent interest in new crops
(Table 1; Retamales et al., 2005). This expan-
sion is hampered by selections/land races that
are low-yielding and, when compared with our
modern cultivars, very soft, small, and tender-
skinned with a very short shelf life. Traditional
production systems have ranged from fields
where plants are relatively evenly spaced in
all directions, as might be found in Ecuador
today, to systems very comparable to tradi-
tional matted row systems (Finn et al., 1998).
Researchers are exploring whether improve-
ments in irrigation, fertilization, pest control,
or perhaps environmental modification using
high tunnels or greenhouse, maybe with hy-
droponics, might expand the production sea-
son and provide significant improvement in
yield and fruit quality over the traditionally
or currently used systems. Currently, the
F. chiloensis genotypes grown in Chile have
a range of skin color from almost white to
slightly red. When these are grown in tunnels
with ultraviolet-filtering plastic, the fruit are
whiter than they would be if grown in open-
field production. The crop is sold mostly
fresh to markets no more than 70 km away.
Some fruit is shipped as far as Santiago
(Chile) from the more southern production
areas, but most of the fruit that leaves the
production region is processed as jams or
spreads for pastries and the lowest quality
material fruit is mixed with white wine to
create a very aromatic and refreshing drink.
Commercial Chilean F. chiloensis fields
are concentrated near Putu
´(lat. 35°13#S,
Regio´ n del Maule), Contulmo (lat. 38°00#S;
Regio´ n del Bı´o-Bı´o), and Pure´n (lat. 37°35#
S; Regio´ n de la Araucanı´a). Fields are estab-
lished using runners harvested from a grower’s
own plantings, leading to potential problems
with moving viruses and other diseases into
new fields. Plantings usually last five to eight
years before they are replanted. Most plant-
ings are less than 0.1 ha in size with a total
estimated of 30 to 40 ha in Chile. Plantings
established from lat. 35 to 36°S, where
rainfall is typically 500 to 600 mm, are
irrigated, whereas those from lat. 37 to 42°S,
where 900 to 1200 mm of rainfall is more
common, are usually not irrigated. The nu-
trient program primarily consists of manure
applications, although calcium carbonate is
applied to soils where the pH is too low for
good production. The leading disease prob-
lems are strawberry leaf spot (Ramularia
tulasnei Sacc.), powdery mildew (Oidium
sp.), and several viruses. Other than diseases,
aphids (primarily Chaetosiphon fragaefolii
Cockerell) and snails (primarily Helix aspersa
O.F. Muller) are the predominant pests. Expan-
sion of production of F. chiloensis is desirable
Fig. 2. Fraise du Chili or Frutilla. From A. Fre´zier (1717), A voyage to the South Seas and along the coasts
of Chile and Peru in the years 1712, 1713, and 1714 as reproduced in Wilhelm and Sagen (1974).
Table 1. Comparison of Chilean native strawberry and commercial strawberry (FAOSTAT, 2013;
Hancock, 1999).
Item Fragaria 3ananassa F. chiloensis
World area planted (ha) 280,000 75–80
Yield (tons/ha) 50–70 3–5
Length of harvest season (month) 3–6 1–2
Planting duration (years) 1–2 6–8
420 HORTSCIENCE VOL. 48(4) APRIL 2013

but faces problems with low plant availabil-
ity, poor nursery plant quality (including an
accumulation of viruses), low yields, and the
lack of a structured market to distribute this
unique crop. The marketing model being used
for the white-fruited strawberry called pine-
berry in Europe offers a marketing model that
may be applicable in Chile (<http://www.
vitalberry.eu/pineberries/>).
CURRENT USES IN BREEDING
The Universidad de Talca (Chile) has an
ongoing breeding effort toward developing
white- and pink-fruited, pure F. chiloensis
types for the export market. This work is
based primarily on germplasm collected by
scientists sponsored by the USDA-ARS Plant
Exploration Office to collect in Chile (Cameron
et al., 1991, 1993; Lavin, 1997; Lavı´n et a l.,
1993) and who were accompanied by research-
ers from El Instituto de Investigaciones
Agropecuarias–Cauquenes (Lavı´n et al., 1993).
At the beginning of this century, this work was
expanded by collections done by the Uni-
versidad de Talca (Retamales et al., 2005),
who in collaboration with C.E. Finn (USDA-
ARS, Corvallis, OR) gathered germplasm
from southern Chile (lat. 39 to 43°S). From
those germplasm collections, several pure
F. chiloensis crosses using white- and red-
fruited parents were made to develop impro-
ved selections from this species. The Universidad
de Talca plans to release three F. chiloensis
cultivars in 2013 with white-, pink-, or salmon-
colored fruit. The tragic earthquake in 2010
destroyed much of the university’s laboratories
and germplasm collections (in vitro and potted
material) and progress on different research
areas has been slowed. The USDA-ARS
(Corvallis, OR) continues to work on devel-
oping white-fruited, pure F. chiloensis types
as well as working with Michigan State Uni-
versity in recreating F.3ananassa with a
much broader germplasm base than the orig-
inal source.
In 1996, the ‘Huachi Grande’ clone of
F. chiloensis was collected on a commercial
farm near Huachi Grande (Finn et al., 1998).
This genotype has been used extensively in
the USDA-ARS Corvallis and Michigan State
University breeding programs and was pre-
sumed to be similar to, or the same, genotype as
Darrow’s ‘Ambato’ (Darrow, 1952). Darrow
hadused‘Ambato’incrossesintheUSDA-
ARS Beltsville program but very little seems to
have come of that effort (Darrow, 1966).
This genotype was growing on volcanic,
extremely well-drained soils. It cyclically
ripened fruit that were large by ‘‘wild’’ stan-
dards but were in the 7- to 10-g size range in
the off-season. The fruit of ‘Huachi Grande’
was pinkish white, soft, and had a tender skin
but the fruit are extremely aromatic with a very
nice flavor. ‘Huachi’ does not survive out-
doors in Michigan as a result of cold tem-
peratures and struggles but lives through
Oregon’s very wet winters. When used in
crosses with adapted germplasm, selections
that are very large (20 to 30 g) with adapta-
tion, fair yield, and fruit that are flavorful
but soft can readily be selected in the first
generation hybrids between F. chiloensis and
F.3ananassa.
Hancock et al. (2001a, 2001b) identified
other outstanding F. chiloensis clones for the
strawberry ‘‘supercore’’ group of 38 elite ge-
notypes that represents the diversity present
in F. chiloensis and F. virginiana.This
‘‘supercore’’ has since been extensively eval-
uated and used for germplasm enhancement,
especially as part of the cultivated strawberry
reconstruction project (Hancock et al., 2001a,
2001b, 2005, 2008, 2010; Stegmeir et al.,
2010). PI 236579, PI 551746, and PI 602567,
in addition to ‘Huachi Grande’ (PI 612318),
that trace to Chilean heritage have been par-
ticularly valuable parents and their genes are
in advanced selections for commercial trial
(USDA-ARS, National Clonal Germplasm
Repository, 2013).
One of the amazing genetic resources
contributed to the world from South America
has been F. chiloensis. A critical component
of the cultivated strawberry, it is still im-
portant as a cultivated species andas a source
or terrific germplasm for breeding modern
strawberries.
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