Potato Remains From A Late Pleistocene Settlement in Southcentral Chile 1
Potato Remains From A Late Pleistocene Settlement in Southcentral Chile 1
Fig. 1-6. Chilean potatoes. Fig. 1. Archaeological tuber skins of Solanum maglia from Monte
Verde (epidermal and cortical views). Fig. 2. Modern specimen of S. maglia. Fig. 3. Starch grains of
1987] UGENT ET AL.: POTATO REMAINS 19
are silent in this regard, there is no actual evidence of this happening. Moreover,
as will be brought out again and discussed at greater length below, the biologic
data are in conflict with this interpretation.
The purpose of our study is to assess what is known about botanical origins of
modern-day Chilean potato species and to relate these findings to the ancient
samples of S. maglia excavated at Monte Verde.
Over 150 wild and cultivated species of Solanum, section Tuberarium, are
known from the Andes of South America. Correll (1962), whose taxonomic treat-
ment is followed here, listed nine species as being indigenous to Chile. However,
seven of these, despite their inclusion in this group, are non-tuber-bearing plants
only very distantly related to the true potato.
The only tuber-bearing species of this section that occur in Chile, according to
Correll (1962), are S. maglia and S. tuberosum, both highly polymorphic. These
were split by earlier taxonomists into a number of lesser taxa, among which were
two escaped forms of S. tuberosum (S. molinae Juz. and S. leptostigma Juz. ex
Buk.) erroneously believed by Bukasov (1933) to be ancestral to the cultivated
potato.
The wild species S. maglia is a white-flowered plant that bears small tubers,
the latter rarely more than 3-4 cm in diameter (Fig. 2). It is presently known from
low, humid, shaded areas near the edges of bogs and swamps, seepage areas,
riverbanks, ravines, and rocky slopes along the central coast of mainland Chile,
and from Chilo6 Island, where it has been recently discovered by Carlos Ramirez,
the third author of our paper. It is also known from a single quebrada locality in
west-central Argentina. All told, its Chilean distribution stretches for over 1,500
km, from Chilo6 Province (43~ in the south to Coquimbo Province (30"S) in
the north. While usually observed at elevations near sea level, this species has
been reported to ascend to 700 m in northern Chile, 2,000 m in southern Chile,
and 1,500 m in Argentina (Correll 1962).
Laufer (1938) reported that this species has long been utilized by the Araucanian
Indians of central Chile. Their ancient, original name for it, malla, found its way
first into Spanish and then into several other European languages. The Italian
spelling, maglia, was the one used by Schlechtendal (1841) when he described
the plant as new to science.
The other Chilean species, S. tuberosum (Fig. 5), is cultivated from Santiago
(330S) to Tres Montes in the Chonos Archipelago (47~ and from there south-
eastward across the Argentine border to Patagonia (5003'S). O f the five horticul-
tural groups of the common potato recognized by Dodds (in Correll 1962; see
also Table 1), the only one presently known from primitive cultivation in Chile
is Group Tuberosum. The remaining four groups, Andigena, Chaucha, Phureja,
and Stenotomum, are grown locally from northern Argentina to Venezuela. These
archaeological S. maglia, 250 • (phase contrast, polarized light). Fig. 4. Starch grains of modem S.
maglia(2n -- 24). Fig. 5. Tuber specimenof modem Chilean cultivatedpotato, S. tuberosumL., Group
Tuberosum. Fig. 6. Starch of modem Chilean S. tuberosum.
20 ECONOMICBOTANY [VOL. 41
The wild potato grows on these islands in great abundance, on the sandy, shelly soil near the
sea-beach. The tallest plant was four feet in height. The tubers were generally small, but I found
one, of an oval shape, two inches in diameter: they resembled in every respect, and had the
same smell as English potatoes; but when boiled they shrunk much, and were watery and insipid,
without any bitter taste. They are undoubtedly here indigenous: they grow as far south, according
to Mr. Low, as Lat. 50~ and are called Aquinas by the wild Indians of that part: the Chilotan
Indians have a different name for them.
Although D a r w i n did not c o m m e n t further on the possible botanical origins o f
Chilean varieties o f S. tuberosum, he did refer his readers to the still earlier travel
account by H u m b o l d t (1811-1812), wherein it was concluded that S. maglia
represents the " p r i m i t i v e t y p e " o f S. tuberosum.
W h y H u m b o l d t should h a v e perceived the a b o v e close relationship between
these two species is s o m e t h i n g not difficult to u n d e r s t a n d f r o m a detailed study
o f their structures. Both species tend to resemble one a n o t h e r quite closely in
their respective habits, patterns o f leaf dissection, fruit characteristics, and corolla
shapes a n d colors. M o r e o v e r , the two plants are r e m a r k a b l y alike in having red-
dish-purple-skinned tubers. T u b e r s with similarly colored skins h a v e not been
discovered a m o n g other Solanum species o f this group.
Although plants o f S. maglia can often be distinguished f r o m S. tuberosum b y
their m o r e widely divergent s t a m e n s a n d b y their tendency to produce smaller
stems a n d tubers, these features are not at all constant. T h e only reliable attributes
o f these plants are their c h r o m o s o m e n u m b e r s a n d the relative size a n d shape o f
their starch grains (Fig. 4, 6), characters that o f course can be d e t e r m i n e d only
microscopically.
Because these species closely resemble each other, they h a v e often been confused
in b o t h the h e r b a r i u m a n d field. Darwin, for example, incorrectly d e t e r m i n e d one
1987] UGENT ET AL.: POTATO REMAINS 21
of his collections of S. tuberosum from Chile as S. maglia, an error that was also
later repeated on the same set of specimens by the taxonomists Spegazzini and
Kuntze (Correll 1962; Hawkes and Hjerting 1969). De Candolle (1886), on the
other hand, who did a careful revision of these species, classified S. maglia as a
variety of S. tuberosum. The latter treatment, however, was later overturned by
Bitter (1913), who considered the Chilean plants of S. tuberosum to be mere
varieties of S. maglia.
The above examples, although few, underscore our point that we are dealing
basically with a closely knit, polyploid complex ofintergrading forms. Comprising
the base of this complex are the diploid (2n = 24) forms ofS. maglia (=S. collinum
Dun.). These give rise to highly sterile, autotriploid (2n = 36; Rybin 1933) forms
of S. maglia through a process that involves the fusion of an unreduced gamete
with one that has been reduced. Also formed at the triploid level are hybrids of
diploid S. maglia and tetraploid (2n = 48) S. tuberosum (Okuno 1951). And,
finally, at the top of this complex are the cultivated and wild clones of S. tube-
rosum, Group Tuberosum. The latter probably arose in Chile from diploid forms
of S. maglia by spontaneous doubling of chromosomes. That initial evolutionary
step was probably followed by a long period of selection by man for higher yielding
forms.
Because the first clones of S. maglia introduced to the horticultural centers of
Europe and North America were triploid and highly sterile, most previous workers
thought this species represented an evolutionary "dead end." Bukasov (1933), for
example, who was aware that tetraploids came from diploids, failed to see a
relationship between S. maglia and S. tuberosum primarily because diploid clones
of this species were not known to him at the time. It is to his credit, however,
that he still recognized the area of southern Chile as being the original homeland
of the Group Tuberosum potatoes.
We shall now describe the area where archaeological remains of S. maglia were
found and report on methods used in their identification.
Monte Verde, situated on the mainland of Chile in the Department of Llanquihue just north of
Chilo6 Island, is 55 km WSW of Puerto Montt and 25 km from the Pacific Ocean. The buried cultural
remains at this site are at an elevation of 55 m.
Portions of the 7,000 m 2 settlement site at Monte Verde are exposed in the banks of Chinchihuapi
Creek, a tributary of the Rio de Maullin. This creek (Fig. 7) currently drains a boggy area in a humid
forest of evergreen Valdivian southern beech (Nothofagus sp.). Sixteen radiocarbon datings of wood,
bone, and charcoal samples from this site indicate that it was occupied about 13,000 yr ago (Dillehay
1985; Dillehay et al. 1982).
Interdisciplinary research by Pino (1983) on geology and by Ramirez (1983) and Diaz-Vaz (1983)
on recovered floral remains has determined that the cold, wet, humid-forest paleo-environment at the
time of human occupation was very similar to present-day climatic conditions.
According to Simpson (1979) and Vuilleumier (1971), snow line during glacial times began at about
500 m elevation in the Andes at the latitude of Monte Verde, 41~ and progressed gradually
downward to sea level until 44~ was reached. This meant that portions of Chilo6 Island and all
mainland territories to the south were capped by a sheet of glacial ice and snow. Chilo6 itself was
probably connected by a land bridge to the Chilean mainland, because ocean levels during the Pleis-
tocene were about 100 m lower than today.
Geological evidence from Monte Verde indicates that glacial ice sheets never reached the south side
of tbe Rio de Maullin where the site is located. Furthermore, palynological data for the 13,000-10,000
22 ECONOMIC BOTANY [VOL. 41
Fig. 7. Aerial view looking north of the Monte Verde site showing Chinchihuapi Creek, lab and
work area of the crew, and surrounding forested bogs. The buried cultural remains are located about
50 m east and west of the two long trenches cut roughly perpendicular to the creek. The southern
trench is 45 m long. The site area and adjacent terraces are being cleared of trees by lumbermen.
Maximum width of the creek is 8 m.
B.P. time interval from the nearby lake region are interpreted by Heusser (1974) and Heusser and
Streeter (1980) to represent a gradual warming from about 13,000 yr ago, culminating in temperatures
warmer than today by about 11,300 yr ago. Mercer (1976) proposed temperatures as warm during
this same interval or perhaps even warmer than those of today.
Three broad vegetational zones occurred in the Monte Verde area during late glacial times (Simpson
1979). There was a narrow belt of evergreen Magellanic Nothofagus forest along the littoral zone.
Above this, to an elevation of about 250 m, was a zone of deciduous, subantarctic Nothofagus forest,
and above the latter, there was alpine grassland to about 500 m. All three zones were open at all times
in the north to wanderings of early man.
The h u m a n settlement discovered at Monte Verde includes the remains of 12 small dwelling units,
the oldest architectural structures of this sort discovered in the Americas (Dillehay 1984). The foun-
dations are made of logs and roughly cut hardwood planks held in place by stakes driven into the
ground. Saplings placed every few feet along the foundation members defined the walls, which were
probably covered originally with animal skins.
Within the dwellings were found plant remains, stone tools, food stains, and shallow, clay-lined pits
that held burnt coals. Two large hearths, three wooden mortars, and several grinding stones were
found outside the huts. About 30 m to the west of these were found the remains of an unusual
wishbone-shaped building that may have served as a place of stone-tool manufacture, as a hide-and-
meat processing area for larger kills, and perhaps as a community shelter for the rendering of medical
care.
Aside from eating of wild potatoes, the inhabitants of Monte Verde are known to have consumed
a wide variety of other naturally occurring plant foods, including various seeds, stems, leaves, nuts,
berries, and roots. This diet was supplemented with freshwater mollusks and meat from paleocamelids,
mastodons, and smaller game (Dillehay 1984).
1987] UGENT ET AL.: POTATO REMAINS 23
Fig. 8--9. SEM micrographs of archaeological Solanum maglia starch. Fig. 8. 3,000x. Fig. 9.
4,000 x.
1987] UGENT ET AL.: POTATO REMAINS 25
chamber of the SEM and examined under magnifications ranging from 250 to
4,000 x. Polaroid film was used to capture the image directly from the screen
(Fig. 8, 9).
In S. maglia, the starch is unusual in that it is formed in four characteristic
shapes: oval, round, triangular, and rectangular (Fig. 3, 4), with the relative fre-
quency of each type indicated by the order of the above sequence. The intersection
of the arms of the interference cross falls in the geometric center of round and
rectangular grains, but towards one end of oval and triangular grains. Grain size
varies from 10 to 20 u for round grains to 35 to 80 u for the other grain types.
No other species of potato is known to have this same combination of starch
characteristics.
As a check on identification, living tubers of diploid S. maglia (2n = 24; P.I.
407408), triploid S. maglia (2n = 36; P.I. 245087), and tetraploid Chilean S.
tuberosum (2n = 48; P.I. 245317) were obtained from the USDA Potato Intro-
duction Station, Sturgeon Bay, Wisconsin. The starch of the m o d e m diploid
introduction of S. maglia (Fig. 4) is in all important respects similar to that of
our 13,000-yr-old specimen. Triploid S. maglia starch differs from both in having
the edges of the rectangular and triangular grains more irregularly formed. The
starch of S. tuberosum (Fig. 6), while obviously resembling the oval grains of S.
maglia, consists of only one grain type, this possibly derived from the latter species
by a process of human selection that included some degree of attention being
given to culinary characteristics. Significantly, no other tuber-bearing species of
Solanum are known from Chile.
COMMENTS A N D CONCLUSIONS
The earliest known potato remains in the world are those of Solanum maglia.
This species makes its appearance in the archaeological record several thousand
years before the appearance of S. tuberosum, the oldest known remains of which
come from caverns in central Peru that were inhabited about the year 8000 B.C.
(Ugent et al. 1982). Had S. tuberosum been widely grown in Chile by the time of
the earliest appearance of this wild species, its tubers would probably have been
used preferentially to S. maglia, as they are much larger.
In contrast to what De Candolle (1885) reported, the tubers of S. maglia are
not bitter tasting. A single tuber of this species (P.I. 407408), obtained from the
USDA Potato Introduction Station, Sturgeon Bay, Wisconsin, was taste-tested by
the first author after it was boiled in water for 30 min. This 2-cm-long tuber
proved similar in flavor to those produced by m o d e m varieties of S. tuberosum.
The second author reports that he has eaten various types of wild potatoes on
his Chilean archaeological expeditions, including those of S. maglia, and that
none was bitter. Moreover, the local inhabitants of south central Chile claim they
eat all varieties of wild, edible potatoes. It should be noted that although bitter
tubers are produced by some wild species of this genus, these kinds occur much
farther north in the central Andes.
The edible tubers of S. maglia might very well have been gathered by the early
inhabitants of southern Chile along with other wild tubers in terrestrial and coastal
aquatic zones for several thousand years prior to the initiation of agriculture.
Domestication of this plant in subsequent millenniums (perhaps in response to
26 ECONOMIC BOTANY [VOL. 41
the need for producing more food for an expanding population) could have brought
a b o u t t h e s p e c i a l set o f d i f f e r e n c e s a s s o c i a t e d w i t h S. tuberosum. T h e s e i n c l u d e
a m o r e r o b u s t h a b i t , a h i g h e r t u b e r y i e l d , a n d a t e n d e n c y t o f o r m a single, l a r g e r -
grained starch type. All of the above could have been brought about as a result
of spontaneous doubling of the chromosome number, followed by many years of
human selection.
L a s t l y , it m u s t b e p o i n t e d o u t t h a t d o m e s t i c a t i o n o f t h e p o t a t o m i g h t h a v e
t a k e n p l a c e in a t l e a s t t w o w i d e l y s e p a r a t e d a r e a s o f S o u t h A m e r i c a : t h e c o a s t a l
forests of southern Chile and the central Andes. In each, distinct cultigens would
h a v e a r i s e n as a r e s u l t o f d o m e s t i c a t i o n o f d i f f e r e n t w i l d species. A l t h o u g h clas-
sified f o r m e r l y as d i f f e r e n t species, t h e s e c u l t i g e n s t o d a y h a v e b e c o m e so i n t e r -
mixed at the tetraploid level in Europe and North America that they are best
r e g a r d e d a s c o m p r i s i n g b u t t w o h o r t i c u l t u r a l g r o u p s o f a h i g h l y v a r i a b l e species,
S. tuberosurn L.
ACKNOWLEDGMENTS
We thank the National Science Foundation, National Geographic Society, Goettingen Foundation,
West Germany, Universidad Austral de Chile, Valdivia, and the University of Kentucky, Lexington,
for supporting research at the Monte Verde site9 Special thanks are extended to Professor Andres
Contreras, Instituto de Agronomla, Universidad Austral de Chile, for sharing his knowledge of wild
potatoes9 In addition, we acknowledge the help of Dr. John Bozzola, Electron Microscopy Center,
Southern Illinois University, Carbondale, and the support of the SIU Department of Botany.
LITERATURE CITED
Anonymous. 1966. R.H.S. colour chart. Published by the Royal Horticultural Society, London.
Bitter, G. 1913. Solana nova vel minus cognita. Repert. Spee. Nov. Regni Veg. 12:452-453.
Bukasov, S.M. 1930. The cultivated plants of Mexico, Guatemala and Colombia9 Bull. Appl. Bot.
Genet. P1.-Breed. (Trudy Prikl. Bot.) Suppl9 47:513-523.
9 1933. The potatoes ofSouth America and their breeding possibilities. Bull. Appl. Bot. Genet.
P1.-Breed. (Trudy Prikl. Bot.) Suppl9 58:1-192.
Correll, D.S. 1962. The potato and its wild relatives. Texas Research Found., Renner, TX.
Darwin, C. 1845. A naturalist's voyage9 Journal of researches into the natural history and geology
of the countries visited during the voyage of H.M.S. Beagle round the world9 John Murray,
London.
De Candolle, A. 1885. Origin of cultivated plants9 Appleton, New York.
9 1886. Nouvelles recherches sur le type sauvage de la pomme de terre. Arch. Sei. Phys. Nat.,
Ser. 3, 15:425-438.
Diaz-Vaz, J. 1983. Identificaci6n de madera Pleistocenica de Monte Verde, Chile9 Unpublished
research report of the Monte Verde Project, on file in the Dept. of Anthropology, Universidad
Austral de Chile, Valdivia.
Dillehay, T. D. 1984. A late Ice-Age settlement in southern Chile. Sci. Amer. 251:106-117.
1985. The cultural relationships at Monte Verde: a late Pleistocene site in south-central
Chile9 In A. L. Bryan, ed., New evidence for the Pleistocene peopling of the Americas. Center
for the Study of Early Man, Univ. Maine, Orono, ME.
, M. Pino, E. M. Davis, S. Valastro, Jr., A. G. Varela, and R. Casamiquela. 1982. Monte
Verde: radiocarbon dates from an early-man site in south-central Chile. J. Field Archaeol. 9:
547-550.
Hawkes, J. G., and J. P. Hjerting. 1969. The potatoes of Argentina, Brazil, Paraguay, and Uruguay.
Oxford Univ. Press, London.
Heusser, C.J. 1974. Vegetation and climate of the southern Chilean Lake District during and since
the last Interglaciation. Quat. Res. 4:290-315.
1987] UGENT ET AL.: POTATOREMAINS 27
, and S. S. Streeter. 1980. A temperature and precipitation record of the past 16,000 years in
southern Chile. Science 210:1345-1347.
Humboldt, A. 1811-1812. Essai politique sur le royaumo de la Nouvelle-Espagne. F. Schoell, Paris.
Laufer, B. 1938. The American plant migration. Part I: The potato. Field Mus. Nat. Hist., Anthropol.
Ser. 28:1-132.
Mercer, J.H. 1976. Glacial history of southern-most South America. Quat. Res. 6:125-166.
Okuno, S. 1951. Cytological studies on potatoes, with some remarks on genetical experiments. Part
I. Jap. J. Genet. 26:70-103.
Pino, M. 1983. La geologia del sitio de Monte Verde. Unpublished research report of the Monte
Verde Project, on file in the Dept. of Anthropology, Universidad Austral de Chile, Valdivia.
Ramirez, C. 1983. Informe de las paleo-plantas de Monte Verde, Chile. Unpublished research report
of the Monte Verde Project, on file in the Dept. of Anthropology, Universidad Austral de Chile,
Valdivia.
Rybin, V. A. 1933. Cytological investigation of the South American cultivated and wild potatoes,
and its significance for plant breeding. Bull. Appl. Bot. Genet. P1.-Breed. (Trudy PriM. Bot.)
II. 2:3-100.
Schlechtendal, D. F. L. 1841. Hortus halensis. C. A. Schwetschke, Halle.
Simpson, B.B. 1979. Quaternary biogeography of the high montane regions of South America. In
W. E. Duellman, ed., The South American herpetofauna: its origin, evolution, and dispersal.
Monogr. Mus. Nat. Hist., Univ. Kansas, No. 7, Lawrence, KS.
Ugent, D. 1970. The potato. Science 70:1161-1166.
, S. Pozorski, and T. Pozorski. 1982. Archaeological potato tuber remains from the Casma
Valley of Peru. Econ. Bot. 36:182-192.
, and M. P. Verdun. 1983. Starch grains of the wild and cultivated Mexican species of Solanum,
subsection Potatoe. Phytologia 53:351-363.
Vavilov, N.I. 1951. The origin, variation, immunity and breeding of cultivated plants. Chron. Bot.
13:1-366. [Originally published in 1935; transl, from Russian by K. S. Chester.]
Verdun, M.P. 1982. Starch grains: a taxonomic character in Solanum (Tourn.) L., section Tuber-
arium. Master's Thesis. Southern Illinois Univ., Carbondale, IL.
Vuilleumier, B.S. 1971. Pleistocene changes in the fauna and flora of South America. Science 173:
771-780.
Whistler, R. L., and E. F. Paschall. 1967. Starch: chemistry and technology. Academic Press, New
York.
Book Review
American Ginseng: Green Gold. W. Scott Persons. Exposition Press of Florida, 1701 Blount
Rd., Pompano Beach, FL 33069. 1986. 172 pp. $19.15 (paper).
Growing Panax quinquefolium L. is not easy. Here is a specific, hands-on account of its
cultivation covering three basic methods: in open fields under artificial shade; in a natural
forest setting simulating the wild growing conditions by scattering seeds with a m i n i m u m
land disturbance; and in well-cultivated and tended beds prepared in the woods under
hardwood timber. Satisfactory woods-grown ginseng can, according to the author, produce
a crop worth $30,000 in five or six years with little capital investment on only half an acre!
Besides the growing manual there are interesting other parts, including ginseng life history,
trade, medicinal properties, and hunting of wild populations, this is a worthwhile and
entertaining treatment of America's premier herb.
WALTER H. LEWIS, WASHINGTONUNIVERSITY, ST. LOUIS, i O 63130