Dynamics of Andean Potato Agriculture'
STEPHEN B. BRUSH,- HEATH J. CARNEY,3 AND Z6sIMo HUAMAN4
The invention and development of agriculture created tremendous diversity
among the species selected for domestication. This diversity is still evident in
cradle areas of domestication, maintained as ancestral varieties or landraces by
traditional farmers. These centers of diversity have been recognized as imporant
since N. I. Vavilov's work 50 yr ago. Archaeologically, they are significant because
of their association with the origins of agriculture and the resulting new
way of life for human populations. Genetically, they are important to geneticists
and plant breeders as sources of germ plasm for the improvement of our modern
crop varieties and for backup crop genetic resources (Harlan, 1976; Oldfield,
1979). Moreover, they are areas where ongoing crop evolution occurs in and
around fields. They can thus provide information as to the ancestry of modern
crop cultivars and enable us to understand better the genetic architecture of our
modern domesticates.
Although extensive germ plasm collecting and archaeological and botanical
research have been undertaken in these areas of crop evolution and crop genetic
diversity, our knowledge of the dynamics and systematics of traditional agriculture
that supports this diversity remains rudimentary. Little anthropological or
ethnobotanical investigation has concerned itself directly with how farmers in
these areas identify, select, maintain and distribute the diverse genetic material
of their crops. This lack of research contrasts sharply with recent advances in
understanding the overall patterns of folk plant classification (Berlin et al., 1974;
Conklin, 1972; Witkowski and Brown, 1978) and the wealth of material on the
socioeconomic dimensions of traditional agriculture (e.g., Halperin and Dow,
1977; Wharton, 1969). The lack of detailed historical studies of genetic resources
and agriculture in areas of crop diversity hampers our ability to model the dynamics
of primitive selection, but it does not preclude attempts to extrapolate
from synchronic analysis.
There is mounting evidence from virtually every center of crop genetic diversity
of the vulnerability and loss of primitive germ plasm, a problem popularly known
as "genetic erosion," the replacement of complex assemblages of ancestral genetic
material by more uniform hybrid and high yielding varieties (e.g., Eckholm,
1978; Harlan, 1975a; Frankel and Hawkes, 1975; Myers, 1979; Oldfield, 1979).
By understanding the dynamics of the agricultural systems being affected by
genetic erosion, we might gain a better idea of how to cope with these endangered
resources (Brush, 1980). Human cultural diversity plays an essential role in the
continuing evolution of these valuable primitive crop genetic resources (Harlan,
1975a), and the crop system of agricultural people maintaining these resources
I Received 4 April 1980; accepted for publication 9 September 1980.
2 Associate Professor, Department of Anthropology, College of William and Mary, Williamsburg,
VA 23185.
3 Graduate Student, School of Natural Resources, University of Michigan, Ann Arbor, MI 48109.
4 Geneticist, International Potato Center, Apartado 5969, Lima, Peru.
Economic Botany, 35(1), 1981, pp. 70-88
? 1981, by the New York Botanical Garden, Bronx, NY 10458
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19811 BRUSH ET AL.: ANDEAN POTATO AGRICULTURE 71
provides an excellent arena for interdisciplinary ethnobotanical research. This
article reports some results of recent interdisciplinary investigation by anthropologists,
botanists and geneticists of one such crop system, that of the potato
in the Peruvian Andes. Its objective is to describe patterns of identification,
classification and distribution of primitive varieties in areas where subsistence
production predominates. The article is also concerned with the maintenance of
native varieties in areas where improved varieties have diffused during the past
3 decades.
ANDEAN POTATO
Two lines of evidence, archaeological and genetic, indicate that potatoes were
first cultivated in the central Andes. Excavations in the central highlands of Peru
indicate the tuber may have been cultivated by 5,800 B.P. (Pickersgill and Heiser,
1978, p. 821; McNeish et al., 1975, p. 30). Engel (1970) suggests an even earlier
date of 8,000-10,000 yr ago for original domestication in the central Andes. Other
archaeological evidence for Andean domestication includes the use of potatoes
as an effigy in pre-Hispanic pottery (Towle, 1961). Genetically, the richest gene
pool of potatoes, estimated by geneticists and taxonomists at 2,000-3,000 varieties,
is found in the Andes. The area thus represents a center of domestication
for the potato by Vavilov's definition (Vavilov, 1926; Harlan, 1971; Simmonds,
1976).
Europeans recognized the importance of the potato for Andean people soon
after the Spanish conquest (Hawkes, 1967). More recent students of Andean
society have noted the relationship between types of potato production and the
expansion of prehistoric states in the area (Troll, 1958). There is a growing literature
on socioeconomic aspects of potato production and use in the Andes
(Werge, n.d.). Of particular importance are recent and detailed studies of potato
production systems in the central Peruvian Andes (Franco and Horton, 1979).
The diversity of Andean cultural features associated with the potato is especially
evident in the classification, selection and distribution of species and varieties
to be discussed below. Very little ritual is associated with potatoes (Murra,
1960). A fairly elaborate variety of storage techniques and facilities are used
(Werge, 1977). During storage, potatoes are protected from worm infestations by
placing the tubers on layers of mufia (Minthostachys spp.). No native strategies
to control pest and pathogen infestation in the cultivated field were observed.
Pathogens, especially golden and cyst nematodes, are reduced by a system of
sectoral fallowing widely practiced in the Andes. Under this system (Mayer and
Fonseca, 1979), potatoes are cultivated for 1 or 2 yr, followed by short rotations
of other tubers (e.g., Oxalis tuberosa), grains (e.g., Chenopodium Quinoa) and
fallow. A period of 6 or 7 yr elapses between potato plantings. Processing is
confined to the freeze-drying of bitter species and wormy non-bitter tubers, fermentation
in running water (to produce tokosh or tongosh), sun drying, and very
limited production of starch (Werge, 1979).
Cooking is almost always done by simple boiling or steaming, and skins are
not eaten. A favorite variation in cooking is the construction of sod and fieldstone
ovens (watia or pachamanca) for roasting potatoes at harvest time. Chili peppers
(Capsicum spp.) are the usual condiment with steamed potatoes, and special
herbal concoctions, often based on wakatay (Tagetes minuta), are served with
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72 ECONOMIC BOTANY [VOL. 35
the tubers. Although nutritional data on subsistence-level farmers in the Andean
highlands is inconclusive, little clinical evidence exists to indicate problems of
malnutrition or protein deficiency in Andean populations with a high dependency
on potatoes (Pic6n-Reategui, 1976). The basic culinary standard for potatoes is
dry matter content. High dry matter is esteemed, and such potatoes are described
as machka in Quechua or harinosa (floury) in Spanish. A puzzling aspect of this
preference is the inverse relationship between dry matter and protein content
(Gray and Hughes, 1978). Why should people who depend on potatoes, a low
protein food, prefer those varieties that are lowest in protein? A possible explanation
for this is that amongst primitive potato cultivars there is great variation
in dry matter and protein content. A number of them have high dry matter and
high protein content (Huaman, 1978).
CLASSIFICATION OF CULTIVATED POTATOES
Potatoes are included in the section Petota of the genus Solanum (Hawkes,
1978). That section includes 154 wild tuber-bearing species, 90 of which are found
in the Andes of Peru and Bolivia (Hawkes, 1978). Today, 3 different taxonomic
systems are used by scientists working with cultivated potatoes. The Dodds system
divides the cultivated potatoes into 3 species (Dodds, 1962), and a Russian
system recognizes 18 cultivated potato species (Lechnovich, 1971). The third,
and most widely used system, was devised by Hawkes and classifies 8 cultivated
species (Hawkes, 1978). In all 3 systems cultivated potatoes are divided into 4
ploidy levels, ranging from diploid (2n = 24) to pentaploid (2n = 60). The Hawkes
taxonomic system was utilized in this research.
By far the most ubiquitous species in the Andes is the tetraploid Solanum
tuberosum subsp. andigena (Juz. et Buk.) Hawkes. Also commonly found are
the diploids S. stenotomum Juz. et Buk., S. goniocalyx Juz. et Buk., and S.
phureja Juz. et Buk., as well as the triploid S. x chaucha Juz. et Buk. Similarly
widespread are the 2 frost-resistant species generally grown at higher altitudes
and processed by freeze-drying to remove glycoalkaloids. These 2 species are the
triploid S. x juzepczukii Buk. and the pentaploid S. x curtilobum Juz. et Buk.
Another diploid, S. ajanhuiri Juz. et Buk., is endemic to southern Peru and
northern Bolivia.
The origin of the potato grown in Europe and North America, S. tuberosum
subsp. tuberosum (Juz. et Buk.) Hawkes, has been the object of much debate
(Hawkes, 1967; Salaman, 1949; Simmonds, 1976). The prevailing opinion holds
that this subspecies, adapted to long photoperiod, resulted from European selection
of andigena varieties brought to Europe following the Spanish conquest of
the Andes. Recent cytogenetic research, however, suggests confirmation of an
earlier theory that the progenitors of modern tuberosum varieties were from
southern Chile-from the Chiloe Archipelago (Grun et al., 1977). Genetic material
from European and North American tuberosum varieties has been introduced
to the central Andes by the diffusion of new varieties from national and
international breeding programs.
The diploid S. stenotomum is widely regarded as the most primitive of the
domesticated species (Hawkes, 1967). The other 7 cultivated species were derived
through various processes including mutation, mitotic and meiotic polyploidizaThis
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19811 BRUSH ET AL.: ANDEAN POTATO AGRICULTURE 73
tion, intra- and interploidy hybridization, and selection. It appears that the process
of potato evolution, through polyploid development from primitive diploids,
has been one of increasing genetic isolation of cultivated from wild potatoes.
There is evidence that variation in the cultivated potatoes has been increased by
hybridization between S. stenotomum and some wild potatoes (Hawkes, 1962;
Huaman, 1975). At the diploid level, hybridization is favored by a mechanism of
self-incompatibility. At the tetraploid level, on the other hand, hybridization in
nature is more difficult because of self-compatibility. Undoubtedly, man's role as
a selective agent is felt on all levels. Characteristics such as lack of tuber dormancy
in S. phureja, frost resistance in S. ajanhuiri, S. x juzepczukii and
S. x curtilobum, good palatability in S. goniocalyx and adaptation to long daylight
in subsp. tuberosum are probably examples of traits selected for by humans.
Botanically, only 3 of the 8 cultivated species are identified without much
difficulty by means of morphological characteristics. In the rest of the cases,
chromosome counts to determine ploidy level are necessary to arrive at certain
species identification. Below these levels, there are vast numbers of distinct
clones, numbering in the thousands. The International Potato Center has developed
a set of 53 plant and tuber descriptors to determine the identity of any single
clone (Huaman et al., 1977). As in species identification, however, morphological
characteristics are often uncertain, and some chemotaxonomic techniques such
as electrophoresis are used to establish other identifying criteria for clones (Desborough
and Peloquin, 1968; Stegeman and Loeschcke, 1976). Electrophoresis
relies on the fact that each individual potato clone has a unique combination of
proteins which are separated according to electrical charge and molecular weight.
As with other crops in their cradle areas of domestication, potato variation in
the Andes is expressed by the existence of numerous landraces of native varieties.
These are adapted to the conditions of traditional agriculture, such as low soil
fertility, low plant populations, and low yield (Harlan, 1975b). Although variable,
they are locally identified and named. In the case of the vegetatively propagated
potato, landraces found within a single locality are comprised of a number of
discrete genotypes, or clones, which are individually named.
Native agriculture associated with this diverse genetic material has 3 consequences:
(a) the maintenance of numerous genotypes over space and time, (b)
the wide distribution of particular genotypes, and (c) the generation or amplification
of new genotypes. In order to understand these consequences, major cultural
patterns of traditional potato agriculture in the Andes must be examined in
relation to the genetic diversity of that crop. These patterns include the identification
and naming of varieties, the selection and planting patterns of different
varieties, and the exchange of varieties within and between localities. Another
significant feature of native agriculture relevant to genetic diversity of potatoes
is the presence of wild and weedy potatoes in and around fields. This promotes
the introgression of germ plasm from those sources into cultivated stocks (Ugent,
1968).
Two principal types of studies have attempted to document the diversity and
distribution of native potato cultivars in Peru. First, germ plasm has been collected
for plant breeding since 1925 by numerous national and international expeditions
(Hawkes, 1941; Ochoa, 1975a). Second, in order to detail more accurately
and comprehensively the varietal frequency and distribution of native
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74 ECONOMIC BOTANY [VOL. 35
cultivars, ethnobotanical studies on a much smaller scale have been carried out
in Peru (Jackson et al., 1980) and Mexico (Ugent, 1968). There remains, however,
a dearth of information concerning the selection and planting patterns of traditional
farmers responsible for the frequency and distribution of cultivars. The
naming system of Andean farmers for native potato varieties has been examined
by several scholars. Hawkes published over 1,000 potato names from the Andean
region (Hawkes, 1947), and large individual collections from more specific regions
have been published (Vargas, 1949, 1956). La Barre (1947) analyzed some 200
names from the Aymara around Lake Titicaca, noting the taxonomic structure
of this nomenclature.
Our research attempts to relate botanical and cultural aspects of traditional
Andean agriculture in a systematic fashion and to address several issues not
included in earlier work. These issues are: a) the consistency of the native system
of classification, b) the spatial distribution of names and types, c) the relationship
between farmer preference and the frequency of genotypes, and (d) the agricultural
regimes of small farmers applied to traditional varieties and to new hybrids
and high yielding varieties.
STUDY AREA AND SAMPLING
Ancestral types of potatoes are grown throughout the Andes from Venezuela
to northern Chile, with the greatest concentration of diversity in the central Andes
of southern Peru and northern Bolivia. In the Peruvian Andes, where this research
took place, they are grown at elevations between 3,000-4,200 m. This effective
range of potato cultivation can, in most areas, be subdivided into 3 zones: 1) a
commercial zone, 2) a zone of primitive, ancestral classes of non-bitter potatoes,
and 3) a zone of bitter potatoes. Between 3,000-3,600 m, growing conditions are
optimal, especially because of lack of frost. In most major valley systems between
these elevations, potato production is often commercial, oriented to producing
marketable, edible potatoes and "seed" potatoes with modern agricultural technology:
the heavy use of commercial fertilizers and insecticides, machinery, and
improved, high-yielding varieties for "seed." Even small scale, subsistence-oriented
potato production in this zone relies primarily on these improved varieties
(Franco and Horton, 1979). Above 3,600 m, peasant, subsistence-oriented farming,
utilizing more traditional agricultural technology and primitive cultivars of
potato is found. This technology usually involves communal control of agricultural
lands, cooperative labor, a short cultivation/long fallow cycle, the use of
traditional Andean farm implements such as the chakitaklla (footplow), and a
lack of dependency on nonlocal inputs of chemicals and energy (Brush, 1977;
Gade, 1975). This area may be divided into 2 zones. Between 3,600-3,900 m nonbitter
potatoes are produced. Above 3,900 m bitter, frost-resistant potatoes are
grown and converted into chuno by freeze-drying. Most households in this area
produce both types of potatoes, as well as other Andean and European crops.
Two general study areas were chosen for research: the Mantaro Valley region
around the city of Huancayo in the central Peruvian highlands and the Vilcanota
Valley region around the city of Cuzco in southern Peru. In each area, natural
vegetation zones of the traditional potato-producing areas were similar, varying
between moist and wet tropical montane forest zones of the Holdridge system
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1981] BRUSH ET AL.: ANDEAN POTATO AGRICULTURE 75
(O.N.E.R.N., 1976). Generally, as one moves westward from the central Mantaro
and Vilcanota Valleys, rainfall decreases to 600 mm, and frost occurs in May or
June. As one moves eastward, rainfall averages 800-1,000 mm, and frost usually
occurs in July or August. Topsoils along the eastern slopes are considered more
fertile. In both the Mantaro and Vilcanota areas, the greatest diversity of potatoes
occurs along the eastern slopes, and potatoes lose predominance to cereals as
one moves westward (Mayer, 1977).
Within both regions, certain areas have been heavily influenced by changes in
the types of "seed," technology and household economy occurring with the rise
of commercial potato agriculture. These changes are especially evident close to
the major urban centers, Huancayo and Cuzco, and along major roads.
In both the Mantaro and Vilcanota areas, villages were visited for the purposes
of collecting potatoes and their associated nomenclature, observing and discussing
farming methods applied to potato cultivation, and mapping fields for the
distribution of genotypes. Information was collected to allow analysis of name
and varietal distribution across 5 different social and spatial levels: field, household,
village, microregion and region. Certain villages were sampled because they
occupied different points on the same "seed" distribution networks. In the Mantaro
region (Fig. 1) collecting was done in 25 villages over an area of roughly
4,000 km.2 Villages were sampled in microregions (defined by small river valleys)
and over broad, ecologically and linguistically diverse areas. Two villages were
selected for more intensive research. Here collecting was done from a larger
number (10-15) households, visits were made throughout the research period
(September 1977-June 1978), and formal and informal interviews were conducted.
In the Vilcanota area, collections were made from 4 villages. Here, fieldwork
occurred in 2 periods, at planting and during the florescence of the potato fields
(November and March).
Tuber samples of native cultivars were treated in 3 fashions. First, a collection
of 486 tubers, from 4 villages and 29 farmers was planted; species and data on 18
stem, flower and tuber characteristics were recorded to determine genetic similarity
or dissimilarity. Information from field mapping was used to augment the
number of plants and names to be compared by over 500 cases. Second, tubers
from the test plot and a large number of tubers collected from farmers during
harvest were described according to 9 tuber characteristics and scored against a
standard. Third, tubers were grouped according to local name, and tubers within
groups were compared in 2 ways: by visual assessment and by electrophoresis.
About 700 samples, divided among 86 groups, were compared by one or both of
these methods. Forty groups and 200 samples were subjected to vertical slab
electrophoresis (Stegeman and Loeschcke, 1976). The total sample contains 262
named varieties from the Mantaro region and 45 from the Vilcanota region.
From January through March, when potato fields were blooming, 19 fields were
mapped, 11 in the Mantaro region and 8 in the Vilcanota region. Fields were
chosen to reflect degrees of selection, from highly heterogeneous to generally
homogeneous plantings. Three different sampling strategies were employed: 1)
all plants, particularly in smaller fields containing great variety and mixture of
clones, 2) transects-alternate rows, or every few rows, in the more homogeneous
fields, and 3) quadrats, often combined with transects, especially in larger fields
or groups of fields where varieties appeared to be clustered. In all fields, native
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76 ECONOMIC BOTANY [VOL. 35
:, < ~~~~~~~~~~~~ o5 lokm
:. lcoyaii.
V ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~( P_ bk::
% 3nsmeit Agro-lifa Zon Toia otn os oct
a Intensiva .ultivotin of indigenous teto Andasm(3 rc
(3400)
I Reseorch nates
(38000 (300
%
HUANCAY0-O
(306), '
.. (3840)
(Cu:zo Legend
Montaro ecu re Pititayom Chongowa noted a
Volley 40 m / (3764 Alto
I intermediate Agro life Zone (Tropical Montane Moist
* Intensive cultivation of indigenous potato varieties
0Research sites
Fig. 1. Location of areas of potato cultivation and research sites in Mantaro Valley region.
informants provided local names and other information about the field. For all
potato varieties encountered, the native name was noted, and plant and tuber
characteristics were recorded using the International Potato Center's descriptors.
Species were determined according to morphological indicators and by cytogenetic
investigation to determine ploidy level (Tarn, 1967).
FOLK TAXONOMIES OF CULTIVATED POTATOES
The native classification of potatoes in the areas investigated involves 4 levels
arranged taxonomically, beginning at the folk generic level and descending to the
subvariety level. The term papa is used universally for all tuber-bearing Solanum
species, although regional dialects may use another term, as in the Mantaro region
where akshu replaces papa. As a folk genus, potatoes are distinguished from
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1981] BRUSH ET AL.: ANDEAN POTATO AGRICULTURE 77
other Andean tubers (e.g., Oxalis tuberosa and Ullucus tuberosa). Below this
level, potatoes are grouped into 4 distinct folk species, according to 4 criteria: a)
cultivation, b) edibility, c) processing, and d) frost resistance. In the Mantaro
region these 4 folk species are:
1) akshu: cultivated non-bitter potatoes with little or no frost resistance.
2) shiri-akshu: cultivated bitter potatoes which must be processed by freezedrying
to remove glycoalkaloids before eating; frost resistant.
3) kurao-akshu: edible, semidomesticated (uncultivated) potatoes found in midaltitude
maize fields; no processing required and not frost resistant.
4) atoqpa-akshu: unedible, wild potatoes; some frost resistant. Literally,
"fox's potato," perhaps to distinguish them from cultivated potatoes that
are sometimes fed to dogs.
At the folk species level cultural divisions are clearly related to botanical
species divisions. All wild, tuber-bearing Solanums are lumped into the atoqpaakshu
class; shiri-akshu is comprised of S. x juzepczukii and S. x curtilobum;
and the other domesticated potato species grown in the area are grouped together
as akshu. The botanical components of the semidomesticate kurao-akshu have
not been established, but they may be escaped S. tuberosum subsp. andigena.
Below the folk species level, the akshu and shiri-akshu groups divide into
locally named varieties, based on tuber characteristics. The former has up to 100
of these named varieties in a single locality, while the latter generally has only
4. These are distinguished according to tuber skin color, meal color and consistency,
tuber shape and configuration of the eyes. Some plant features, such as
flower and stem color are included in the naming system, but this is rare. During
field mapping, however, it became apparent that a number of clones were widely
recognized for their plant characteristics as well as for their tubers.
The primacy of tuber shapes and colors in the native classification is apparent
in many names. Examples include calhuay ("weaver's shuttle") for a long flat
variety, mishipasinghan ("cat's nose") for a round, compressed cultivar with
shallow eyes concentrated in the center, lumchipamundana ("potato which
makes the young bride weep") for a knobby, convoluted variety, turumangia
("rainbow") for a multicolored type, and chunchupafiahuin ("eyes of a jungle
person") for a dark-skinned potato with light colored eyes.
The average farmer growing these varieties can name about 35 types, a number
which varies according to the experience and interest of the individual. In a single
locality, 50-70 potato names may be found, but perhaps 10-20% are synonyms.
In gathering potato nomenclature, some informants were obviously casual and
careless in naming tubers, while others were meticulous and conscientious. The
latter were generally able to discuss the characteristics of a particular tuber and
why it should or should not be classed as a particular variety. This indicates the
presence of implicit rules in the naming and classification of tubers. Rules were
also evidenced when a group of farmers was presented with a set of tubers to
name. Detailed and sometimes animated discussion took place concerning the
tuber in question. A name would be suggested and debated: "no this cannot be
a huayru because it is too flat and not mealy enough, perhaps it is an ichipsa.'"
In some cases, a dozen names were suggested and rejected by reference to numerous
criteria. In others, a tuber was immediately recognized and named without
debate.
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78 ECONOMIC BOTANY [VOL. 35
TABLE 1. PERCENTAGES OF VISUALLY SIMILAR AND ELECTROPHORETICALLY IDENTICAL
TUBERS IN LIKE-NAMED GROUPS.
Visual Electrophoresis
No. of % No. of %
Spatial level comparisons similarity comparisons identity
Locality 372 81.2 231 75.8
Microregion 512 71.7 185 40.5
Region 3,694 74.3 967 48.7
The fourth, and lowest, taxonomic level in the native classification is that of
subvariety. This is based on secondary tuber characteristics, principally variations
in tuber color. Examples are puca suito, yurac suito and yana suito which
have morphologically similar tubers but with red, white and purple skin color,
respectively. Twenty percent of all varieties have subvarieties, varying from 2-
8. Subvarieties are not as widely recognized or agreed upon as the varietal denomination
and may be considered of minor importance in the overall scheme.
It appears to be equivalent to native recognition of polymorphism among closely
related types.
It may be assumed that the naming system of farmers is closely related to their
knowledge of potatoes on such matters as selection (Rosch and Lloyd, 1978).
Consistency of the naming system, therefore, is a key issue. If the system were
found to be random or highly capricious, the relevance of native selection procedures
for understanding the botanical structure of Andean potato agriculture
would be in question. Our research indicates, however, that the naming system
is not random.
A clear indication of the consistency of the naming system is based on visual
assessment of the similarity of tubers in like-named groups. The results of this
comparison are reported in Table 1 along with results of electrophoretic comparison
of tubers in the same groups. Tubers are compared at 3 spatial levels:
those from the same locality, those from different localities in a single microregion
(clustered according to geographic features such as a river valley), and those from
the entire Mantaro survey region (4,000 km2). Like-named tubers from the same
locality were judged to be morphologically similar and electrophoretically identical
at a high rate. It seems probable, therefore, that potatoes given the same
name by farmers in one village are clones. This probability, however, falls off
sharply as one leaves a locality. Even at the microregional level, most electrophoretic
comparisons between like-named tubers showed them to be dissimilar.
The native classification, however, is consistent at all 3 spatial levels when judged
by the criteria of tuber similarity. Potato names are, thus, meaningful both within
and beyond village boundaries. Andean farmers have a particular type of tuber
in mind when a name is given, and this association holds over a large region.
Biologically, these names are generally meaningful within a village and less so
beyond.
Beyond different farmer aptitudes in naming, variation in the native classification
may be linked to 3 factors. First is the very nature of folk taxonomy. This
is based on implicit and usually unexpressed rules rather than on explicit ones.
Second is the polymorphism of potato clones and the effects of different enviThis
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1981] BRUSH ET AL.: ANDEAN POTATO AGRICULTURE 79
ronmental conditions on the development of tubers. Third is the possibility of
inter- and intra-ploidy hybridization producing new clones. This occurs within
cultivated fields, and the presence of wild species surrounding fields makes the
introgression of wild germ plasm into cultivated stocks likely, especially at the
2x level. Andean farmers are familiar with "ground keeper," or uncultivated,
potatoes in their fields. These are referred to as kipa throughout Peru. Farmers
are generally unaware of the possibility of sexually reproduced potatoes developed
from true botanical seed and associate kipa with unharvested tubers from
previous plantings. Although no formal mechanism for the incorporation of new
clones into the native system was evident, an informal mechanism may exist.
While most cultivars seem to be named according to relatively narrow, albeit
implicit, rules, a few names are applied to tubers of dubious similarity. In some
cases, these names appeared almost to constitute residual categories. New and
unusual clones might easily be classified under these names. Many of these names
were applied to diploid and triploid tubers. The former are likely to hybridize
with wild germ plasm, and the latter are products of hybridization between diploids
and tetraploids.
The relevance of the native practice of naming potato varieties to the botanical
structure of Andean potatoes is indicated in 2 ways. First, there are indications
of a close relationship between tuber morphology and genotypic identity in this
and other research (Jackson, 1977). Therefore, a naming and selection system
based on tuber morphology, such as the native Andean classification of potatoes,
should have definite biological implications. Second, ethnographic information
indicates that potatoes are usually exchanged and distributed according to native
name. Exchange of potatoes between families was commonly referred to in terms
of specific locally named varieties. Farmers in one village generally know which
households have certain named varieties and their particular agronomic, culinary
and perhaps commercial characteristics. When asked about a particular variety,
farmers often reported that they did not have it but that it could be found in a
particular household and where it was cultivated. Furthermore, it was reported
that traders and farmers from other villages, sometimes distant, often request
particular named varieties. Indeed, place names were commonly attached to certain
varieties.
SELECTION AND MAINTENANCE OF NATIVE VARIETIES
As Harlan (1975b) points out, landraces of traditional crops are products of
human selection for such characteristics as color, flavor, texture, and storage
quality. The importance of these characteristics to Andean potato farmers is
indicated in their naming system for potatoes. Selection may be observed in the
planting patterns of potato fields and in the spatial distribution of particular varieties
within a region. Although studies of native potato fields in Peru (Jackson
et al., 1980) and Mexico (Ugent, 1968) have demonstrated field heterogeneity,
our research indicates that the selection of ancestral varieties is more complex
than previously thought. We found fields of all types of potatoes to be generally
orderly. Rows were well kept, with relatively few weeds and little evidence of
wild species within fields. At the margins, however, wild species were sometimes
common. In the Mantaro region, "seed" tubers were usually planted individually.
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80 ECONOMIC BOTANY [VOL. 35
In Cuzco, the more heterogeneous fields were planted in clumps containing 3-5
"seed" tubers. Selection involves planting some fields with relatively homogeneous
stands of preferred varieties, as well as the maintenance of heterogeneity
in others.
Three types of selection of potatoes by Andean farmers may be observed.
First, on the level of folk species, non-bitter potatoes (papa or akshu in the
native nomenclature) are grown separately from bitter potatoes (luki in southern
Peru, shiri in the central region). In some fields around Cuzco, these varieties
were interplanted, but this was never observed in the Mantaro region. Second,
within the non-bitter group, certain preferred varieties are selected and grown in
uniform plantings, apart from the heterogeneous collections in other fields. Third,
high yielding varieties originating from breeding stations are often grown separately
from native varieties. Selection may be observed in fields, in the terminology
for fields, in the technology applied to different fields, and in the farmer's
objective in planting different fields.
Within the native variety class of potato, selection of non-bitter potatoes establishes
2 kinds of fields. The first are fields where varieties are mixed and
randomly planted. In the central highlands, such fields are referred to as chacro.
Throughout Peru, this is the most common kind of potato field. "Seed" for this
kind of field is selected for "seed" quality and size rather than for variety. The
traditional planting procedure is to sow "'seed" tubers into holes opened by the
footplow. As many as 5 small tubers may be dropped into a single hole. When
planting new land, potatoes are often planted into holes opened in untilled soil,
a practice known as tikpa. After roughly 6 wk, the sod around each clump is
turned over and mounded against the plants. The fertilizer generally applied at
planting is animal dung. The objective for planting these heterogeneous fields is
almost invariably to produce potatoes for home consumption. The mixture of
colors, shapes, textures, and flavors enhances a diet in which many meals consist
entirely of potatoes garnished with a hot pepper sauce. The native varieties are
also believed to retain palatability and viability as "seed" over many months of
storage.
Varietal heterogeneity of native potato fields is one of their most important
attributes. The crop evolution of the cultivated potato is closely linked to the
mixture of species and genotypes which promotes hybridization and crossing
between ploidy levels and among clones. Table 2 summarizes the frequencies of
ploidy levels and cultivars in ten fields sampled in the Mantaro Valley. The predominance
of tetraploid tubers is typical for fields throughout the Andes. Pentaploids
are not found in this sample since they are bitter, frost resistant potatoes
grown at higher altitudes. Fig. 2 depicts a quadrat from a mixed field outside the
village of Umasbamba, north of Cuzco. This field was planted by an officer in
the local religious hierarchy, its harvest intended for a fiesta. Forty-six named
varieties were randomly planted in the field, with 2 clones frequently growing
from a single planting.
Besides these heterogeneous collections of native varieties, it is common for
farmers throughout the Andes to select and maintain more uniform plantings of
native potatoes. Selection of this type is done by farmers who market native
cultivars or who are involved in "seed" distribution networks of native varieties.
Selection is by "'seed" quality and by variety. It usually occurs at harvest when
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1981] BRUSH ET AL.: ANDEAN POTATO AGRICULTURE 81
ROW -
0 ED iZ 0
O5 . 0 IU
(4x)0W ?9 4x 4) (4x)
C 8( mur bol yano ( ) 'usi 3) mur wijoO reoineto
CE) O* () ( (x)T (4C)CTO zo a 90 CQ0Oe44 )O
KEY- (p tika boli p muru kousi puk
(4x) (4x) (4x) (4x)
7muru boli syona ksu'usi muru wiroajo renociminto
(4x) (4x) (4x) (4x)
olpuka boli 9 puka suytou ykna wiroajo machonors
(4x) (4x) (4x) (4x)
0 kompis simon suytu 0alka wayrtito o inkolo
(4x) (4x) (4x) (2x)
oalka kompis qyelna wakoto 0puka wayotit ero maetlo
(4x) (4x) (4x) (4x)
(3puka kompis Q
(4x) (4x) (4x) (3x)
0 yurac kompis ql
(4 x) (4 x) (4 x) (4 x)
0 wayruru olo0nes )achancas papa
(4 x) (4 x) (4 x)
Fig. 2. Quadrat of native varieties of potato in field near Chinchero, Cuzco Dept., alt. 3,820 m.
"seed" tubers of different varieties are sorted and stored in separate mounds.
Selection may also occur at planting time when "seed" tubers are desprouted.
Selection of these preferred varieties relates to their demand in markets and trade
networks for "seed" tubers. Important characteristics affecting this demand are
culinary quality and special agronomic qualities such as frost or late blight (PhvThis
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82 ECONOMIC BOTANY [VOL. 35
TABLE 2. CUMULATIVE PLOIDY AND CULTIVAR FREQUENCIES FOR MANTARO VALLEY
SAMPLE.
Number of plants per ploidy level Total
Cultivars observations
Field no. per field 2x 3x 4x per field
1 23 27 40 511 578
2 19 30 15 115 160
3 11 30 7 115 152
4 7 235 12 247
5 6 35 45 80
6 13 24 11 136 171
7 11 5 391 396
8 5 217 217
9 30 82 18 486 586
10 18 14 16 273 303
Total 212 377 2,301 2,890
Percent 7.34 13.04 79.62 100
tophthora infestans) resistance. The development of roads, markets and urban
centers in the Andean highlands encourages small farmers to market at least some
of their crops. The demand for native potato varieties is evidenced by the fact
that they bring a substantially higher price than the improved, high yielding ones.
Selection of these preferred native varieties depends also on their demand in
"seed" distribution networks which antedate more recent market development.
These networks exist because some potato producing areas regularly lose "seed"
to frost, drought and late blight. "Seed"-producing areas on the wetter and relatively
frost free eastern slopes of the Andes export "'seed" tubers westward to
communities bordering the high, cold and dry plateaus of the western ranges.
Traditional "seed" distribution, involving yearly treks of up to 50 km to "seed"producing
areas, is being supplanted by trucks and centralized markets, but
"'seed" still moves over the same range. The distribution of preferred "seed" by
regular networks has produced a number (5-8 per region) of cosmopolitan varieties,
clones grown by virtually every household and uniformly named over several
thousand square kilometers. Recent work by Ochoa (1975b) and Jackson
(1977) indicates that certain varieties are distributed throughout Peru and are likenamed
in widely dispersed geographic and linguistic areas. These varieties are
often selected and grown separately, as demonstrated in Fig. 3, depicting the
fields of a farmer in the hamlet of Aymara, Huancavelica Department. This farmer
sells potatoes to "seed" suppliers dealing with farmers from the Chongos Alto
area, 30 km to the west. As the map shows, he practices a high degree of selection
in the 4 fields below his house. Field 1 contains sections of 90% murunki (subsp.
andigena) separated by a thin corridor of 44% mariva (tuberosum x andigena)
and 22% duraznillo (subsp. andigena). Murunki is a highly cosmopolitan clone
in the Mantaro region; duraznillo is less so; and mariva is a hybrid-improved
variety found frequently in urban markets. Field 2 is planted with 80% pujuya
(S. phureja) which the farmer reported to be particularly tolerant of frosts. Pujuya
is common in the inventories of farmers in the Chongos Alto area who report
receiving "seed" from the Aymara area. Field 3 is planted in bitter shiri potatoes
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1981] BRUSH ET AL.: ANDEAN POTATO AGRICULTURE 83
(S. x curtilobum and S. x juzepczukii) for making chuno. One-half of Field 4
consists of 90% yana huancuy (subsp. andigena), while the other half consists
of 90% yana shuitu (subsp. andigena), both highly prized native varieties in the
Huancayo market. This farmer maintains 2 other similar fields, removed from his
house. He is, however, atypical in not maintaining a heterogeneous chacro field.
Native varieties, dealt with above, are distinguished as papa de regalo ("gift
potatoes") or papas de color ("colored potatoes") from the improved varieties,
known as papa blanca ("white potatoes") or papa mejorada ("improved potatoes").
The latter have diffused widely in the Andes after their initial appearance
in the early 1950s, and now appear in fields of subsistence farmers in remote
villages. In areas with easy access to urban markets, improved varieties have
made deep inroads into the total stock of native tubers. In many areas, farmers
grow both native and improved varieties in separate fields, using different technologies
and for different objectives.
Improved varieties, when selected and grown separately, are usually grown as
a cash crop. Their chief advantage is their yield, often 2 or 3 times that of native
varieties. This is due to the resistance bred into them, especially to Phytophthora
infestans, and to their relative responsiveness to fertilizers. Selection of specific
improved varieties depends primarily on the cost and availability of "seed" and
on their marketability. Small farmers, who also grow a subsistence crop of native
varieties, often plant improved potatoes in previously cultivated soil using oxen
and a method referred to as chacma, rather than the tikpa method. Yield is the
single most important criterion for this cash crop, and farmers utilize more fertilizer,
especially manufactured types, on improved varieties than on fields of
native varieties destined for home consumption. Likewise, there is a greater
investment in insecticides and fungicides for these fields.
While improved varieties are selected for their commercial value, native ones
are selected according to the demands of subsistence agriculture. Three major
reasons are cited by farmers for retaining native potatoes: taste, storage quality,
and "seed" viability. For people who eat potatoes as the primary food 2 or 3
times a day, culinary appeal is essential. Improved potatoes are regarded as
insipid, watery, palatable only in soups, fried, or to undiscriminating potato eaters.
The native varieties are preferred because of their floury consistency and
high dry matter content. Native potatoes also retain their palatability after months
of storage, while improved varieties often become bitter. This may be due to the
fact that most bred varieties have a white or cream skin which turns green with
bitter alkaloids when exposed to light. The heavily pigmented skin of native
varieties protects them, to some extent, from becoming green.
Finally, native varieties are maintained by small farmers because of their
"seed" viability. These varieties produce viable "seed" year after year, while
improved varieties depend on having "seed" replenished after 1-3 yr. Farmers
observe that improved potato varieties "degenerate" rapidly, losing yield and
producing deformed tubers. Native varieties, on the other hand, maintain acceptable
yields and tuber forms indefinitely. This difference may relate to the fact
that "seed" from improved varieties often originates from lower altitudes where
they are infected with virus by aphids, while native varieties are grown at high
altitudes where aphid populations are greatly reduced. The use of improved varieties,
however, is associated with greater expense in acquiring "seed" and
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84 ECONOMIC BOTANY [VOL. 35
44%Mariva (4x)
220% Duraznillo(4 x
17 % Cante6a (4x)
17 % Tarma (4x)
98% Tarma
(4x)
98% Tarma
( 4 x ) _ _ _ _ _ _ _ _ _ _ _ _ _
80% Pujuya (2 x)
IO % Puka Pulush (2 x)
10%Others
90% Yana Shuitu 90%Yana Huancuy
(4x) (4x)
(3 x)
Shiri (5 x)
0 10 20m
S~treamn
Fig. 3. Diagram of a farmer's fields planted in selected native potato varieties. Aymara, Huancavelica
Dept., alt. 4,050 m.
dependency on commercial "seed" distribution systems. Purchased "seed" potatoes
are sought from areas distant or ecologically distinct from the area of
cultivation. Thus farmers from lower elevations seek "seed" potatoes from higher
zones, and vice versa.
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19811 BRUSH ET AL.: ANDEAN POTATO AGRICULTURE 85
NATURAL SELECTION SOCIO-ECONONIIC FACTORS
Environmental Stresses: Regional and Local Economy
-pathogens -land, labor, and capital available
-pests -market and road development
- frost -production costs
-drought -transportation
-commercial demand and price
-extension programs
DIVERSITY IN FIELDS FARMER SELECTION
Biological Maintenance: Subsistence Criteria
-mutation -culinary preference
-hybridi zation -storage quality
-amphidiploidy -seed viability
-agronomic characteristics
Market and Trade Criteria
- demand and price
______________________ _ -storage quality
CULTIVATION PRACTICES
Proximity of Wild and
Weedy Species: K_\
-weeding
-hedgerows SEED NETWORKS
-crop rotation Traditional Exchange
Tillage -kinship system
- ritual obl igat ions
-diverse altitudinal fields
Commercial Exchange
-native varieties
- improved varietiesl
Fig. 4. Summary of the selection and maintenance of potatoes by Peruvian highland peasants.
CONCLUSION
The primary components of Andean potato agriculture are summarized in Fig.
4, a flow chart of the factors affecting the selection, maintenance and distribution
of potato germ plasm in central Peru. This summary outlines the interaction of
human and biological factors contributing to genetic diversity of potatoes in the
area.
Botanists have suggested that native Andean potato fields are dynamic evolutionary
systems (Jackson, 1977; Iltis, cited in Ugent, 1970). Inter- and intra-ploidy
hybridization is made possible by the planting practices of Andean farmers reported
here and elsewhere. Our research indicates, however, that selection and
distribution of native potato varieties is far more complex than the random planting
of numerous varieties. The maintenance of these numerous varieties is neither
casual nor random. A regular system of nomenclature, organized in a taxonomic
manner, accompanies this cultivation. Specific cultivars are identified according
to an implicit set of criteria involving tuber and sometimes other characteristics.
Selection of varieties relates primarily to culinary characteristics, but specific
biological fitness is recognized in certain clones. The adaptation of native varieties
to local conditions is revealed in the difference between native and hybrid or
improved varieties. The latter depend on frequent replenishment of "seed" to
remain viable. The maintenance of clonal heterogeneity is a regular part of native
agricultural practices, as is the distribution of a few cosmopolitan clones through
"'seed" networks. The cultural and biotic features of Andean potato agriculture
are thus interwoven and express a complex symbiotic relationship between man
and plant.
There are indications, however, that this relationship is deteriorating. In many
areas, subsistence-oriented potato agriculture is being supplanted by commercial
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86 ECONOMIC BOTANY [VOL. 35
agriculture relying on hybrid varieties. The genetic erosion of native varieties is
increasing at an alarming rate (Ochoa, 1975a). At the least, multidisciplinary
information on native agriculture and actual cultivars grown on the local and
regional level should be collected before the irreplaceable agricultural heritage of
Andean farmers and the invaluable clonal richness of native potato varieties are
lost forever. Any efforts to preserve the diversity of native Andean potatoes must
take into account the need to retain the cultural practices which maintain these
cultivars.
ACKNOWLEDGMENTS
This research was supported by grants from the National Science Foundation
(Grant BNS76-83067), the College of William and Mary, and the International
Potato Center. We are thankful for the kind assistance of the staff of the International
Potato Center during fieldwork, especially Douglas Horton, Carlos
Ochoa, Peter Schmiediche, Wilman Galindez, and Robert Werge. We are also
grateful to those who made suggestions for the fieldwork phase and commented
on earlier drafts of this paper: Brent Berlin, Daniel Gade, Joseph Casagrande,
Enrique Mayer, Gustavo Javier, H. David Thurston, Hugh Iltis, David Reed,
Margery Oldfield, Janis Alcorn, Billie Jean Isbell, Valentine Cusihuaman, and
Benjamin Orlove. Thanks, also, to Christine Kurtz for drafting the map and
figures. Finally, we are indebted to the farmers of the Mantaro and Cuzco regions
of Peru who graciously shared their knowledge with us and allowed us to tramp
through their potato fields.
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BOOK REVIEW
Cashew. J. G. Ohler. 260 pp. illus. Koninklijk Instituut voor de Tropen, Ams
Dfl 40.00.
J. G. Ohler has put together an impressive and very useful monograph on the cashew
(Anacardium occidentale), based on 20 years of personal experience with this crop. The
volume, illustrated with many charts and photographs (including some in color), begins
with the early history of the cashew and brings the reader up to the present with a discussion
of its use as an edible tree-nut crop and for the production of cashew nut-shell
liquid (CNSL). The latter product, 20-25% of total nut weight, is primarily utilized in the
brake linings of automobiles. In the second chapter, world production data for the past
and present, along with future projections, are offered.
Twenty seven pages are dedicated to the botany and morphology of cashew, which is
andromonoecious, having both perfect and staminate flowers in the same panicle. A thorough
summary of agronomic and cultural practices is offered, including climatic requirements
(in general a temperature no less than 7?C and rainfall 1,000-2,000 mm annually).
The volume also serves as an invaluable reference to potential growers of the crop, elucidating
basic diseases and pests that might be expected in a plantation. An interesting
section is devoted to the processing of the nuts, where precautions must be taken to expel
all of the toxic CNSL. The "fruits" (i.e., fleshy, swollen pedicels) of this species, known
as "apples," produce ajuice that is very agreeable in taste and contains two to three times
as much vitamin C as citrus juice. It is estimated that many of the 2.5-5 million tons of
"'apples" produced each year go unused, left to rot under the trees. This appears a great
waste, as syrup, wine, brandy, canned fruit, pickles, and jam can also be manufactured
from the fresh "apples." Four hundred nineteen references are listed at the end of the
text, an asset to the reader who might wish to peruse the available literature. The author
is to be congratulated for a well-organized and enjoyable book that will now serve as the
major work on this important but presently under-utilized food plant.
MICHAEL J. BALICK, NEW YORK BOTANICAL GARDEN, BRONX, NEW YORK 10458
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