the grain chenopods of the tropical american highlands

The Grain Chenopods of the Tropical American Highlands

N. W. S I M M O N D S 1

In March and April, 1963, Dr. K. S. Dodds and I visited Bolivia to study the local potatoes, a visit that was made possible by generous help from the Rockefeller Foun- dation. Several samples of the local cultivated species of Chenopodium were collected and subsequently grown and studied at this In- stitute. A collection of the Mexican grain Chenopodium, kindly provided by Dr. J. S. Niederhauser of the Rockefeller Foundation, was found to provide some points of interest m comparison with the South American plants. Since there is no general survey in English of the economic botany of these interesting plants, this paper is cast in the form of a review that incorporates many new observations.

Systematics

The three species. The three species con- cerned are: l. Chenopodium quinoa Willd. (Sp. P1. 1 (2) : 1301. 1797) 2. Ch. nuttaliae Safford (J. Wash. Aead. Sci. 8: 521-7. 1918) 3. Ch. pallidicaule Aellen (Fedde Rep. 26: 126. 1929). Vernacular names are recorded as follows. For (1), the most widely used is the Quechua name "Qfiinua" and its variants, a name which is, however, sometimes also applied to Amaranthus caudatus (7, 13). The version "Qfiinoa" (abbreviated QUIN when necessary) seems to be generally acceptable in both English and Spanish writings. In Aymar~, the plant is "Hupa" or "Jupa"; in Chilefio, "Dahue"; and in Atacamefio, "Seeksaholor" (5). For (2), the common name in modern Mexican speech is "Huauzontle" or "Guausoncle"; in Aztec, "Uauhtzontli" or "Huautzontli" (18); according to Hunziker (14), "Huautli" is also current but much confused with "Michi- huautli" which is Amaranthus hybridus. "Huauzontle" (abbreviated HUA) will be used here. For (3) the recorded vernacular

1 John Innes Institute, Hertford, England. Received for publication September 4, 1964.

names are "Cafiahua" (abbr. CAN) and its variants, especially "Cafiihua", though, in Ecuador, "Cuchi-qfiinua" has also been recorded (13).

Fig. 1. Leaf shapes. A,C,E Qfiinoa; B,D,F Huauzontle. A and B first and fourth seedling leaves, C and D eighth to tenth leaves, all from comparable glasshouse plants. E and F from field-grown plants to show the overlap of leaf shapes in the two species.

223

224 ECONOMIC BOTANY

Plan t materials. Four stocks, all bought as seed samples in markets, were used: Qfi inoa--QUIN 1, from the market of Quil- lacollo, valley of Cochabamba, Bolivia, alt. ca. 2500 m; QUIN 2 from Batallas, on the Altiplano near La Paz, alt. ca. 4000 m; Huauzontle from a market in the neighbor- hood of Mexico City, Mexico; and Cafiahua frmn La Paz, Bolivia, alt. ca. 4000 m.

P lan t characters. General descriptions of all three species are given by Hunziker (13, 14), for Huauzontle by Safford (18) and for Qfiinoa by Cfirdenas (5). Leading diagnostic characters, incorporating new observations, are summarized in Table 1, and leaf shapes are shown in Fig. 1.

All three species are plastic, and stature varies enormously in response to environment; thus, plants of Qfiinoa may range from 20 cm to over 2 m tall, depending on how they are grown. Qfiinoa and Huauzontle are of similar habit, generally reminiscent of Chenopodium album, whereas Cafiahua is a small, much branched, bushy plant.

Safford's drawings show that the leaves of Qfiinoa are somewhat more sinuate-rounded at the margins than the rather sharply

toothed leaves of ttuauzontle. This difference is certainly sometimes good (Fig. 1), but some stocks of Qfiinoa resemble Huauzontle in shape, and occasional leaves of the latter may resemble Qfiinoa (Fig. 1). The difference is, therefore, not diagnostic, even though it is sometimes characteristic.

The flowers of Cafiahua, wholly concealed within the leaves and bearing reduced numbers of stamens, appear to be virtually eleistogamous (see below); whereas those of Qfiinoa and Huauzontle, with separate females and hermaphrodites bearing exserted stigmas, are evidently adapted to some de- gree of outerossing (Fig. 2). The observation of purely female plants in Qfiinoa is new, previous authors having recorded only gynomonoecious plants. Females occur in QUIN 2 to about 40% but are absent from QUIN 1. There is no evidence as to the frequency or geographical distribution of gynodioecy in the species; comment on its significance is made below.

Perianth lobes and anthers .are basically five in all three species but with some reduction of anther numbers in Cafiahua and some split t ing of members in Huauzontle.

TABLE l CHARACTERS OF THE GRAIN CHENOPODS

Chromosome counts by G. E. Marks, that for Qfiinoa confirming an early count by Kjellmark (6), for Huauzontle confirming a count given by Singh (20), the count for Cafiahua being n e w .

Ch. quinoa Ch. nuttalliae Ch. pallidicaule Qfiinoa Huauzontle Cafiahua

Habit tall, little branched tall, little branched short, bushy, much branched

Leaves variable, sinuate-dentate more sharply toothed and small, smoothly and relatively broad, to small, relatively narrow- very broad, fleshy narrower a n d sharply er than most Qfiinoa toothed

Infloreseenees exserted leafy panicle exserted leafy panicle

Flowers gynomonoecious or gynomonoecious gynodioeeious

Perianth lobes 5 5 - 8 Stamens 5 5 ( - 7) Fruit 2 mm. diam. with fiat- 1.5 mm. diam. with flat-

tened or depressed equa- tened or depressed equa- torial band torial band smooth, tough, pale yel- smooth, tough rusty red lowish 2n : 4 x : 36 2n : 4 x : 36

Pericarp

Chromosome number

wavy,

many, minute, concealed in foliage hermaphrodite, eleistogamous

5 1 - 3

0.8 mm. diam., rounded at equator

roughened, fragile

2n : 2.r : 18

'PILE GRAIN CHENOPODS OF THE TROPICAL AMERICAN HIGHLANDS 225

Fig. 2. Inflorescences (X1.32). A gynomonoeeious Qfiinoa, B female Qfiinoa, C Qfiinoa in young fruit, D Huauzontle.

Fig. 3. The fruiting phase (x 0.4). A mature infruetescence of Qfiinoa (compact type, cf. the more open 2C), B Huauzontle, C Cafiahua (mature but minute infructeseences concealed within the subtending foliage). Rate of maturity roughly shown by this figure: A was the earliest Qfiinoa 3 months old, B the earliest Huauzontle 8 months old, C an average Cafiahua 3 months old (all glasshouse plants in small pots of the same size).

are always green, but (in the stock studied, at least) a minute ring of magenta pigment develops around the base of the perianth during fruit growth, and a magenta pig- mentatiml also develops in the immature fruit. This magenta pigmentation all fades later, the perianth becoming a brownish orange, concurrently with the developnlent of rusty pigmentation in the pericarp.

Of the distinctness of Cafiahua, there is no doubt. Qfiinoa and Huauzontle are clearly quite nearly allied but, on the present evidence, are distinguishable on leaf shape (usually), numbers of floral parts, size and shape of fruit and pigmentation; and they are also geographical]y disjunct (Fig. 5). I t is not known how constant these differential characters are, since very few collections of Huauzontle have ever been critically ex- amined. Present observati<ms agree with those of S'~fford (18), however, so the distinction may be tentatively maintained. It is known that they hybridize, nevertheless, and further genetical and systematic studies may yet indicate that they are conspeciflc.

Cafiahua is quite distinct from the other two species in its small fruits, fragile pericarp and shiny black or dark brown testa (Figs. 3 and 4) ; Qfiinoa and Huauzontle have rather similar fruits, but the latter tend to be a little smaller, they have a more prominent micropylar beak and are rusty reddish brown in colour. The fruits of Qfiinoa are basically pale yellowish but sometimes take a tinge of magenta pigment from a coloured perianth (though this fades at full fruit maturity). The upper leaves and inflorescence of Qfiinoa are variably pigmented, from none to a strong magenta (betacyanin) 2 eolour; those of Huauzontle

2 Thanks are due to J. B. Harborne for this observation. The betacyanins are anthocyanin- like pigments but are chemically quite different from the anthocyanins; they are diagnostic of a group of allied Centrospermous families in- cluding the Chenopodiaceae [review in (11)].

Fig. 4. Fruits. A Qfiinoa, B Huauzontle, C Cafiahua and D Chenopodium album for comparison: on left, fruits enclosed in perianths; on right perianth removed, except for C in which the fragile pericarp has also been stripped.

THE GRAIN CHENOPOI)S Ol~ T H E TROPICAL AMER1CAN HIGHLANDS 227

1 0 - -

0

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/_-- /.=-

u

m

m

m p---

2 0 ~ 1 - "~ ' -

3o

4 0 ~ 0 Fig. 5. Distribution of the cultivated Chenopods. Area of Huauzontlc stippled; of Qfiinoa shaded where rare or extinct, solid black in present area of caltivation; tlle last 'lrea corresponds approximately with the distribution of ('~lfi~lhua. Outline based on a map in the Oxford Atbls and reproduced by kind permission of the Ox- ford University Press.

228 ECONOMIC BOTANY

A note on Chenopodium pueblense. This plant was described by Reed (17) as a pot- herb cultivated in Mexico under the name Cuahzontli. The description agrees, so far as it goes, with Ch. nuttalliae, and the plant had the same chromosome number (2n ---- 36). the method of preparat ion for eating is the same as that described by Safford (18) for Huauzontle. There seems to be no doubt, therefore, that Ch. p~eblcn.+e is merely a later synonyn for Ch. n~tttalli~e.

Geographical distr ibution. The distribu- tions of the three species in cultivation are as follows (Fig. 5) :

Q~iinoa: is common in the high country of Peru and Bolivia to over 4000 m alti tude; it is r~are and declining in Argentina (Deps. Ju juy and Salta) , Chile and Ecuador; and has been lost in Colombia in comparatively recent years (1, 7, 13, 15, 16). The latitu- dinal range was 5~ to 30~ but the plant is common now only between about 5 ~ and 20~ according to Sauer (19), it once ex- tended to Chiloe (43~

Huauzontle: occurs in the States of Mi- choaSan, Oaxaca, Vera Cruz and Tamauli- pas in Mexico (17-24~ but cultivation is generally declining (13, 18).

Ca~ahua: occurs with Qfiinoa in the cen- tre of its range in Peru and Bolivia, but is much less common and is elsewhere recorded only as very rare in Ecuador (13, 23).

The only one of the three seriously tried as a crop outside the native area of distribution is Qfiinoa. I t showed some promise as a grain plant for land between 7,000 and 8,000 feet in Kenya (8) but apparent ly came to nothing there (3). I t is said to have been used as a foodstuff in Germany during the first World War (2) and could undoubt- edly be adapted to cultivation in temperate countries if there were a need for it.

Bio logy

Seed biology. The few published figures on Qfiinoa frui t weights range between 250 and 520 fruits per gm (190 to 400 mg per 100) (1, 7, 8). Four recent valu(,s were 240, 310, 410, 540 per gm (190 to 420 mg per 100), in good agreement. The fruits of Huauzontle are somewhat lighter (690 per gin, 146 mg per 100) and those of Cafiahua lighter still (2080 per gin, 48 mg per 100).

Freshly harvested Cafiahua fruits showed a transient dormancy which was readily brok- en by treatment with gibberellic acid and disappeared naturally in less than three months; Qfiinoa and ttuauzontle were virtually completely non-dormant. Germination and seedling growth rate are extraordinarily rapid in all three species, a rapidi ty which is often not reflected in subsequent flowering and fruiting. Reed (17) found also very rapid germination in his Ch. pl~eble~lse.

I t is sometimes said that Qfiinoa seed washed and scrubbed in preparat ion for milling loses viability, but this is certainly not always so, for several washed samples acquired in South American nmrkets were found to give good germination.

Flowering and fruit ing. All three species are highly plastic in their response to ex- ternal environment. Generally, the more re- stricted the rooting spaee the more quickly the plants come into flower. But there is also in Qfiinoa, at least, a large genetic ele- ment involved. Thus, QUIN 1 flowered and fruited well when crowded in small boxes in the glasshouse during the summer of 1963 but, in the field, formed enormous plants over 2 m tall which had only just started to flower when killed by autumn frosts; QUIN 2 in parallel flowered and fruited only a little more quickly in the glasshouse than QUIN 1 but, in the field, attained only about 150 cm and flowered and fruited well. The difference between the two stocks was apparent very early in the greater seedling growth rate of QUIN 1.

The single stock of Cafiahua that was available flowered and fruited in the glasshouse more quickly than the quickest Qdi- noa, that of Huauzontle later than the latest Qfiinoa (cf. Fig. 3). Responses to root- restriction were similar to those of Qfiinoa.

A glasshouse experiment in which half- boxes of young plants of Qfiinoa were subjected to normal daylight (about 14 horn's) as ag~dnst a short day regime (9 hours) for two weeks showed that the short day treatment hastened flowering by a few days, but that this was accompanied by a distinct stunting due to a reduction of the light in- tegral, exactly as found by Fuller (9). There is no support for Sivori's finding (21) that Qdinoa is a short day plant, but it might

T t t E GRAIN CHENOPOI)S OF T t I E TROPICAL AMERICAN H I G H L A N D S 229

:Fig. 6. The effect of light on Qfiinoa. Three weeks after sowing the right-hand half of the box was subjected to a 9-hour day, the left-lmnd remaining in about 18 hrs. Short-day lmlf pale, stunted and quick to flower (see text).

be noted that he used material from Ecua- dor, so the possibility that the species is heterogeneous in response cannot be ex- cluded. The conclusion is, therefore, that flowering is mediated by a nutritional-geno- typic interaction rather than by day length.

At flowering, the hermaphrodite flowers of Qfiinoa and Huauzontle opeu widely, and the extruded anthers burst at a touch; female flowers hardly open, hut the stigmas are extruded and give a characteristic whiskery appearance to the inflorescence (Fig. 2). After pollination, the hermaphrodite perianth closes again and only at full inaturity of the frui t does the perianth (of all flowers) shrivel and deflex, exposing the ripe frui t which readily falls off. tIowever,

infrueteseenees harvested before this late stage bear mature (i.e., viable) fruits which, if threshed off, fall enclosed in the perianth. Thus, whole plants can be harvested in bun- dies and the complement of' ripe grain pre- served without too much shatterino'. No doubt this non- (or delayed-) shattering habit has been developed by human selection in the ehenopods, as in other cereals.

Cafiahua has a rather different behaviour. The inflorescenees are concealed in the foli- a~e, the flowers hardly open at all and, at maturity, the fruits, still mostly enclosed in the perianths, are freely shed when the plant is shaken, in this more closely resembling what one expects of a wiht Chenopodium (Fig. 3).

230 ECONOMIC BOTANY

Variability. With such plastic plants as these chenopods, the only satisfactory way of studying variability is by the experimental culture of mass collections. C/rrde- has (5) grew a number of Qfiinoa collections from the altiplano region of Peru and Bo- livia, but no other experimental studies have been reported. Information is, therefore, very scanty.

Qfiinoa is, even to casual inspection, very variable, individuals in any one plot com- monly showing a range of stature, inflorescence type and pigmentation. Such variation was very evident in the altiplano collection, QUIN 2; in pigmentation, for example, some plants were unpigmented, others had magenta axil spots, others had various levels of pigmentation in the young leaves and in- floreseences, with or without axil spots. By contrast, QUIN 1 was nearly uniform in habit, was but little variable in pigmentation and of much later maturity than QUIN 2. The difference in maturity is of obvious possible ecological significance but cannot yet be interpreted. The first impression, therefore, is of locally great variability within a field but of possible ecological differ- entiation--which would be expected a priori in response to the diversity of habitats in which the crop is grown.

Until the necessary experimental studies have been made, it does not seem profitable to embark upon systematic subdivision founded upon characters which may well turn out to be themselves variable or trivial and assessment of the published systematic work is, therefore, impossible. Hunziker (13) recognized three colour forms (ft. viri- descen.% lutescens, rubesccns) and a fourth (f. melanospermum) having black fruits rounded at the edges; the last sounds very distinct and would probably well repay study of living plants. C~rdenas (5) recognized three varieties based on inflorescence habit: var. amarantiforme (with diffuse panicles, 9 forms), var. glomer~data (con- gested panicles, 9 forms) and var. inter- media (intermediate panicles, 2 forms). Dennler (7) thought that leaf pigmentation and leaf content of caleimn oxalate were correlated with altitude, the greater the height, the more pigment and oxalate; but he gave no detailed observations.

There is no information available on the variability of Huauzontle.

Cafiahua seems to be rather little variable, though it has hardly been investigated. The La Paz stock studied varied in eolour (stems pink or green) and in testa colour (brown or black--exposed by loss of the fragile pericarp) ; plant and seed colors varied in- dependently in this material, though Hun- ziker (13) regarded them as correlated, rec- ognizing a colourless f. typicum, a lightly pigmented f. purpureum and a very dark f. melanosl)erm~m.

The key to the variability of any crop plant lies in an understanding of the breeding system. Flower behaviour (reviewed above) and breeding experiments (yet far from complete) suggest that all three spe- c ies -and especially Cafiahua--are predonfi- nautly inbreeders and that populations, therefore, tend to be made up of mixtures of inbred lines. Occasional crossing would be expected--and certainly occurs--in gynomonoecious Qfiinoa, while the gynodioecious QUIN 2 is evidently adapted to a still higher level of outbreeding. Genetic studies will be reported elsewhere.

Origins. All three are native American crops developed by Indian agriculturists in pre-Columbian times. Early Spanish writings make it clear that Huauzontle and Qfiinoa at least, in their respective areas, were very important food plants (14). For Qfiinoa, there are a number of archaeological re('ords from Peru, Chile and Argentina (22). The ancestors of all three must, therefore, have been native American wild species of Chenopodiun,--though none has been identified. There are several references in the literature to wild forms, but none has been properly investigated; and it is un- known whether these records refer to es- caped eultivars, ancestral wild species or totally distinct plants. C~rdenas (5) uIen- tions a wild form of Qfiinoa under the Ay- mar~ name "Isualla" and Hunziker (13) states that Cafiahua is wild in the north Ar- gentine at ca. 3700 m altitude; the latter author believed Ch. hlrcinum Schrad. to be the aneester of Qfiinoa.

Cultivation and Use Cultivation. Qfinoa is a summer crop in

Fig. T. (~flino:~ (.ountr.v. Tlw edge of lh(' |~olivian :lltil)l:~m~ near A('h:a.:J~.hi, on tlw ~s:Lv ~)v~.r t . ,~or:lta. Al~itud~, :lh(mt 14(100 f~'('t.

the Andes, hcin~ sown in ,~cl)t('mh*'r-Now~m- her, cx(,(,ptionally as early as ,tun(, at lower ~dtitudes (4, 7). It is harvested about M~u'(,h at'ter a / rowth l)eriod of four t() six months. Seed is hroad('ast or |)lantcd in rows and estimates of sced rates are 20 kg per hectare for the former ~uid 2-10 kg per he('tare for the latter method. At m~lturity, but before the grain sh(,ds, the plants ar(' cut, bundled, dried, fl~filed and the / ra in is winnowed. Yields vary greatly and the impression on the altiplano is that they arc mostly very h)w. According to Alberts (4), ml average yield is about 840, rising execptionally to 3000 kg per het'tarc (7). Albcrts (4) states that Cafiahua is slightly quicker to grow and mature than Qfiinoa and is, therefore, favoured for cultivation at higher altitudes; Gareia (10) gives the maturity period of Cafiahua as four to five months in the Puno region of Peru.

~fiin()a and ('~fflahua ;ll)l)~,ar t() h,, nlth(,v hardy plants tolerant of the poor ~t~my soils on whi(,h th(,y ,lr~,, i)erl'[)r(.( *, ()t'tcn culivatcd (Fig. 7), and tolerant also of droug'ht and mo(h,ratc frosts. They ar(, am)- l);~rcntly littl(, attacked by pests or dis- cases, though birds can be a nuisance at the Qfiino~l harv(~.~t (4) (this is true also in K(,nya (S) and tho English sparrow rc~ldily ,~(l~pis itself to (dfiinoa).

Utilization. The l)rinmry use of Qfiinoa is as ~l flour-yicl(ting grain. Analytical data are given by Dcnnler (7) and White ct a]. (24). Al)l)roximate average fi/ures are: protein 1-1-15%, fat 4%, carbohydrate 51- 58%. The protein content is higher than in most cereal g'rains, and it has been shown (24) to be of excellent quality (equiwdent to that of whole dried milk) in rat-feeding trials. For Cafiahua, there are analytical

Fig. 8. Qfiiaoa cultivated on the altipla~lo.

data very similar to those for Qfiinoa (10, 24).

The grain of' Qfiinoa is generally held to contain a toxic bitter principle, though the literature does not mention this for Cafiahua or Huauzontte. According to Dennler (7), the bitter principle is a saponin, but it does not seem to have been studied chemically. The saponin is removed by repeated wash- ing in ~lkaline water coupled with pound- ing and rubbing to abrade the pericarp; but sometiriJes the grain is milled and the flour itself' is washed (14).

The uses to which these plants are put are intimately bound up with the lore and cus- toms of the people that grow them, and no atteml)t will be made to enumerate them

here. Basically, all are grain plants which yielA a nutritious flour made into bread, biscuits, porridge, etc. or fermented, alone or in various combinations, to yield chicha. Accessory products are spinach from the foliage and an alkaline ash from the stems, lliptu, which is favoured for chewing with coea leaves.

0onelusions In the central Andes, maize is grown

wherever the environment allows, frmn low levels up to ca. 10,000 feet altitude. Above this height, the people depend on hardier I)lants for food, notably potatoes, the minor tubers (Oxali,~, Tropaeolum and Ullucus), beans (Vicing), harley (grown as green fod- der for stock) and the chenopods, Qfiinoa

Fig. 9. Q6inoa in England (John Innes Institute, July 1964). A--QUIN 1, late flowering, leafy growth, little pigmented; B--QUIN 2, early flowering, smaller and less leafy, very variable in pigmentation.

2 3 4 ECONOMIC BOTANY

and Cafiahua. I t seems to be a reasonable speculation that the ehenopods were selected locally from wild ancestors by people who needed a nutritious grain to supplement the starchy tubers but who had no adaptable wild grasses fronl which to create cereal grains. The relatively great size of Qfiinoa and Huauzontle fruits in comparison with those of other ehenopods, together with the non-shattering inflorescence habit, nmst be taken to be "cultivated characters" developed by human selection, exactly as in the cereal grasses. On this basis, Cafiahua must be assumed to be rather little advanced as a crop.

Old World agriculturists also used ehenopods as grains, as shown by the Iron Age bog burials of Jutland and many other archaeological remains in Europe. Helbaek (12) reviewed the prehistoric use of Cheno- podium album and concluded that it was reaped fronl stubbles and fallow land rather than cultivated. In India, forms of this species are still grown, often in mixtures with grain amaranths, in hilly areas of the north- west (alt. 5000-7000 ft.) (20). But nowhere in the Old World did the species become an important er(~p probably because, at the altitudes at which agriculture was practiced, there was always an equally good or better cereal grass available. I t may be that, if there had been an Old World agriculture at the immense altitudes characteristic of the Andes, then Chenopodium album would have become a crop, as the ancestor of Qfiinoa did; this, however, is sheer speculation and it could equally well be argued that it was the Old World agriculturist's good luck and/ or skill in breeding cereal grasses that saved him from cultivating chenopods. 3

Summary A general review, incorporating many new

observations, of the American grain chenopods is given. The species are Ch. quinoa (Qfiinoa), Ch. nuttalliae (Huauzontle) and Ch. pallidicaule (Cafiahua); the first two are related tetraploids (2n = 36), and the third is a quite distinct diploid. Huauzontle

3 He does, however, sometimes cultivate buck- wheats which were developed in the harsh climates of central Asia and it might be interesting to explore the analogy (if any) between Chenopodium and Fagopyr~m as grain crops.

is Mexican, and the other two are Andean plants. Brief surveys of systematics, biology, cultivation and utilization of the three species are given. Qfiinoa is the principal native grain of the high central Andes and is still of considerable local importance.

Literature Cited 1. Anonymous. 1909. Kew Bull. misc. Info.

1909: 425-7. 2. - - . 1930. Kew Bull. 1930:332 3. 3. --1946. Dep. Agric. Kenya, Ann. Rep.

1946: 133. 4. Alberts, H. W. 1946. Quinoa--ancicnt

food crop in South America. Agric. Am. 7: 150-2.

5. CSrdenas, iV[. 1944. DescripeiSn preliminar des las variedades de Chenopodi~lm q~dnoa de Bolivia. Rev. Agric. Bolivia 2: 13-26.

6. Darlington, C. D., and A. P. Wylie. 1955. Chromosome Atlas of Flowering Plants. George Allen and Unwin, Ltd., London.

7. Dennler, G. 1936. Reismelde und Inkn- weizen. Tropenpflz. 38: 97-108.

8. Elmer, L. A. 1942. Quinua (Chenopodi~m q~dnoa). E. Afr. agric. Jour. 8: 21-3.

9. Fuller, H. 1949. Photoperiodie responses of Chenopodi~lm quinoa and Amaranthus cat, daters. Amer. J. Bot. 36: 175-80.

10. Gareia, M. J. 1953. Estudio qfiimico de la Cafiihua (Chenopodi~m pallidicmde Aellen). Informe 85 Esta. exp. Agric. Peru 1953, 20 pp.

11. Harborne, J. B., and N. W. Simmonds. 1964. The natural distribution of the phenolic aglycones. In: Biochemistry of Phenolic Compounds: pp 77-127. Aca- demic Press, London and New York.

12. Helbaek, It. 1960. Comment on Chenopo- dium album as a food plant in pre- history. Ber. Geobot. Inst. Eidg. Teehn. Hochsch., Stilting Rubel 31: 16-19.

13. Hunziker, A. T. 1943. Las cspecies ali- menticias de Amaranthus y Chenopadium cultivadas pot los Indios de Am('rica. Rev. Argentina Agron. 10: 297-354.

14. - - . 1952. Los pseudocereales de la Agricultura indigena de America. Univ.

Nac. de CSrdoba, Buenos Aires. 104 pp. 15. Looser, G. 1943. Chenopodium quinoa, un

cultivo que despareee de Chile. Rev. Argentina Agron. 10: 111-3.

16. - - . 1946. Cultivos de Qfiinoa en Chile. Rev. Argentina Agron. 13: 285.

17. Reed, H. S. 1950. A new species of Cheno- podium from Mexico. Madrofio 10: 139-42.

THE GRAIN CHENOPODS OF THE TROPICAL AMERICAN tIIGHLANDS 235

18. Safford, W. E. 1918. Chenopodium nuttalliae, a food plant of the Aztecs. Jour. Wash. Acad. Sci. 8: 521-7.

19. Sauer, C. O. 1950. Cultivated plants of South and Central America. Smith- sonian Inst. Bur. AnL EthnoI. Bull. 143: vol. 6, 487-543.

20. Singh, H. 1961. Grain amaranths, buck- wheat and chenopods. Ind. Coun. agrie. Res. Cereal Crop Ser. 1 :46 pp.

21 Sivori, E. M. 1947. Fotoperiodismo de

Chenopodium quinoa Willd. Darwiniana 7: 541-51.

2'2. Towle, M. A. 1961. The Enthnobotany of pre-Columbian Peru. Chicago. p. 36.

23. u C. C. 1938. Nora etnobotanica sobre la Cafiihua. Rev. Argentina Agron. 5: 225-30.

24. White, P. L. et al. 1955. Nutrient content and protein quality of Quinua and Cani- hua, edible seed products of the Andes Mountains. Jour. Agric. Food Chem. 3: 531-4.

the grain chenopods of the tropical american highlands

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