Botany of Peyote
Edward F. Anderson
Chapter 8, Peyote, The Divine Cactus, ©1980 The Arizona Board of Regents
Published by The University of Arizona Press ISBN 0-8165-0680-9
Until the 1950s there had been no careful and extensive botanical
studies of peyote. Plants which were brought into the chemist's
laboratory—or the horticulturist's greenhouse—had little or
no documentation, available only at the site of collection, about
the place of origin or other important characteristics. As a result,
plants from the same population were sometimes given different
scientific names and those of separate regions often were given
the same name. This absence of botanical understanding, primarily
due to insufficient field and laboratory studies, consequently
resulted in mistakes and confusion by historians, anthropologists,
chemists, pharmacologists, and others. For example, numerous references
have been made to peyote as belonging to the genus Anhalonium.
This name is botanically invalid, as it was applied to the group
of plants which had earlier been named Ariocarpus; hence,
the later name Anhalonium cannot be used for that group
or any other group of plants such as peyote. However, the name
Anhalonium had been employed so widely for about a century
that few people other than botanists specializing in taxonomy
were aware of the fact that the name should not have been used.
The confusion and difficulties that have resulted probably can
never be completely straightened out.
Another serious problem was created in the 1890s when the German
chemist Arthur Heffter received a shipment of poorly documented
and incorrectly identified peyote specimens for laboratory analysis.
These plants were to be the basis of some of the most important—and
confusing—pioneer chemical studies of peyote. Heffter discovered
that the plants he had received belonged to two distinct groups
based on the alkaloids present but he claimed that he was unable
to distinguish the groups on structural or morphological grounds.
Since he had no collection and field data he decided that peyote
simply consisted of two chemical forms. Jan G. Bruhn of
the University of Uppsala and Bo Holmstedt of the Swedish Medical
Research Council have thoroughly researched the literature dealing
with this period of peyote history; their conclusion is that Heffter's
batch of plants actually consisted of the two distinct species
of peyote, which do have definite alkaloid differences.[1] A better
botanical understanding of the group, as well as proper scientific
data, would have prevented the introduction of much confusing
information into the literature that has persisted for more than
seventy-five years.
This frustrating botanical chaos concerning peyote existed for
so long that in the 1950s a research biochemist interested in
the hallucinogenic properties of the plant personally financed
a graduate school program to study and determine the botanical
relationships of the group and to unscramble the nomenclature.
The botanical aspects are now much clearer.
The peyote cactus is a flowering plant of the family Cactaceae,
which is a group of fleshy, spiny plants found primarily in the
dry regions of the New World. Some of the characteristics which
one normally sees in cacti are not readily evident in peyote,
except for the obvious one of succulence. Spines, for example,
are present only in very young seedlings. However, the cactus
areole—the area on the stem that usually produces flowers and
spines—is well pronounced in peyote and is identified by a tuft
of hairs or trichomes. Flowers arise from within the center of
the plant and, like other cacti, the perianth of peyote flowers
is not sharply divided into sepals and petals; instead there is
a transition from small, scale-like, outer perianth parts to large,
colored, petal-like, inner ones. Another characteristic which
shows that peyote belongs in the cactus family is the absence
of visible leaves in either juvenile or mature plants. Leaves
are greatly reduced and only microscopic in size; even the seed
leaves or cotyledons are almost invisible in young seedlings because
they are rounded, united, and quite small. Also, the vascular
system of peyote is like that of other succulent cacti in which
the secondary xylem is very simple and has only helical wall thickening.
BOTANICAL HISTORY
 |
Fig. 8.1 — The first illustration of peyote
appeared in Curtis's Botanical Magazine
in 1847 (plate 4296). |
Peyote was first described by western man in 1560 but it was not
until the nineteenth century that any plants reached the Old World
for scientific study. Apparently the French botanist Charles Lemaire
was the first person to publish a botanical name for peyote, but
unfortunately the name that Lemaire used for the plant, Echinocactus
williamsii, appeared in the year 1845 without description
and only in a horticultural catalog. Therefore, it was necessary
for Prince Salm-Dyck, another European botanist, to provide the
necessary description to botanically validate Lemaire's binomial.
No illustration accompanied either the Lemaire name or the description
by Salm-Dyck and it was not until 1847 that the first picture
of peyote appeared in Curtis' Botanical Magazine (figure
8.1).[2]
In the second half of the nineteenth century the characteristics
and scope of the large genus Echinocactus were disputed
by several European and American botanists; gradually its limits
were narrowed and new genera were proposed to contain species
that had once been included in it. In 1886, Theodore Rumpler proposed
that peyote be removed from Echinocactus and placed in
the new segregate genus Anhalonium, thus making the binomial
A. williamsii, a name which soon became widely used throughout
Europe and the U.S.3 Much earlier (1839) Lemaire had proposed
the name Anhalonium for another group of spineless cacti,
now correctly classified as Ariocarpus.[4] Anhalonium must
be considered as a later homonym for Ariocarpus, so, according
to the International Rules of Botanical Nomenclature, it cannot
be validly used as a generic name for any plant.[5] Ariocarpus
superficially resembles peyote, but clearly is a different genus.
In 1887, Dr. Louis Lewin, a German pharmacologist, received some
dried peyote labeled "Muscale Button" from the U.S.
firm of Parke, Davis and Company in Detroit, which had obtained
the material from Dr. John R. Briggs of Dallas, Texas, earlier
that year.[6] Lewin used some of this plant material in chemical
studies and found numerous new alkaloids; he also boiled some
of the dried "buttons" in water to restore something
of their living appearance and gave them to a German botanist,
Paul E. Hennings of the Royal Botanical Museum in Berlin for study.
Hennings noted that Lewin's plant material appeared similar to
the plant called "Anhalonium" williamsii (Echinocactus
williamsii Lemaire ex Salm-Dyck) but apparently differed somewhat
in the form of its vegetative body, namely in the characteristic
wool-filled center of the plant. Hennings decided that the dried
plant material given to him by Lewin was that of a new species,
which he formally named Anhalonium lewinii, in honor of
his colleague. His description was accompanied by two drawings,
one of the new species, A. lewinii, and the other of the
older species, A. williamsii.[7] The illustration of A.
lewinii shows a high mound of wool in the center of the plant.
Apparently, the drawing, which had been made from the dried plant
material that Lewin had boiled in water, was an incorrect reconstruction
of what had been the original appearance of the plant. When the
top of a peyote dries, the soft fleshy tissue is reduced greatly
in volume, whereas the wool does not decrease in size at all.
Therefore, the proportion of wool to what formerly was the fleshy
or vegetative part is greatly increased in the dried button. This
phenomenon presumably caused Hennings and Lewin to believe that
they had a new species of peyote when in actuality the plant material
they had studied was that of "Anhalonium" williamsii.[8]
Bruhn and Holmstedt have concluded that Lewin's plant material
known as "Anhalonium" williamsii was in fact
the southern species of peyote, Lophophora diffusa. The
specimens which Hennings described as the new species "Anhalonium"
lewinii belong to the northern species of peyote, L. williamsii.[9]
Additional confusion concerning the botanical classification of
peyote occurred in 1891 when the American botanist John Coulter
transferred peyote to Mammillaria, a genus commonly called
the pincushion or nipple cactuses Then, in 1894, a European named
S. Voss confused things even more by placing peyote in Ariocarpus,
the valid name for a distinct—and quite different—group of
plants that had been called Anhalonium also.[11] Finally,
in the same year Coulter proposed a new genus for peyote alone:
Lophophora.[12] This helped clarify the nomenclatural situation
because peyote had been included in at least five different genera
of cacti by the end of the nineteenth century. The group of plants
commonly called and used as peyote is unique within the cactus
family and deserves separation as the distinct genus Lophophora.
Beginning about 1900 numerous forms and variants of peyote were
collected in the field and sent to cactus collectors and horticulturists
in Europe and the United States. The highly variable peyote plants,
not seen as part of natural populations but as individual specimens
in pots, were often described as new and different species. None
of the taxonomic studies, however, were based on careful field
work, so little was known of the nature and range of variations
within naturally occurring peyote populations. By mid-century
greater accessibility of peyote locations in Texas and Mexico
permitted extensive field work which has shown that plants of
the genus Lophophora, especially in the north and central
regions of its distribution, are highly variable with regard to
vegetative characters (i.e. color, rib number, size, etc.). The
number and prominence of ribs, slight variations in color, and
the condition of trichomes or hairs have tended to be three of
the main characters which have delineated many of the proposed
species and varieties of peyote; however, these characters vary
so greatly even within single populations that they are an insufficient
basis for separating species—if a species is considered to be
a genetically distinct, self-reproducing natural population.
Field and laboratory studies show that there are two major and
distinct populations of peyote which represent two species:
(figure
8.2).[13] The first, Lophophora williamsii,
the commonly-known peyote cactus, comprises a large northern population
extending from southern Texas southward along the high plateau land of
northern Mexico. This variable and extensive population reaches its southern
limit in the Mexican state of San Luis Potosi where, near the
junction of federal highways 57 and 80, for example, it forms
large, variable clumps. The second species, L. diffusa,
is a more southern population that occurs in the dry central area
of the state of Queretaro, Mexico. This species differs from the
better-known L. williamsii by being yellowish-green rather
than blue-green in color, by lacking any type of ribs or furrows,
by having poorly developed podaria (elevated humps), and by being
a softer, more succulent plant.
COMMON NAMES OF LOPHOPHORA
The study of peyote has frequently been confused because the plant
has received so many different common names. Fr. Bernardino de
Sahagun first described the plant in 1560 when he referred to
the use of the root "peiotl" by the Chichimeca Indians
of Mexico.[14] The two most commonly used names, "peyote"
and "peyotl," are modifications of that ancient word.
The actual source and meaning of the word "peiotl" is
disputed and at least three theories have been proposed to explain
its etymology. Several Europeans have suggested that the term
"peyote" came from the Aztec word "pepeyoni"
or "pepeyon," which means "to excite."[15]
A derived word from this is "peyona-nic," meaning "to
stimulate or activate."
A similar proposal was made by V. A. Reko and extensively discussed
by Richard Evans Schultes; they suggested that the term "peyote"
came from the different Aztec word "pi-youtli," meaning
"a small plant with narcotic action.''[16] This somewhat
narrow interpretation of the kind of action should perhaps be
broadened to mean "medicinal" rather than "narcotic,"
as the Indians certainly would have thought of the actions of
the plant in the former context.
Probably the most widely accepted etymological explanation for
the origin of the term "peyote" was suggested by A.
de Molina, who claimed that it comes from the Náhuatl word
"peyutl," which means, in his words: "capullo
de seda, o de gusano."[17] This, translated from Spanish,
means "silk cocoon or caterpillar's cocoon." Molina's
explanation, therefore, proposed that the original word was applied
to the plant because of its appearance rather than its physiological
action. Certainly one of the most distinctive characteristics
of peyote is the numerous tufts of white wool or hair. Dried plant
material has an even greater proportion of the "silky material"
and most of it must be plucked prior to eating. The presence of
these woolly hairs seems to be of significance because some other
pubescent (hairy or woolly) plants, not even cacti, have occasionally
been called peyote by Mexicans. Examples of such non-cacti are
Cotyledon caespitosa of the family Crassulaceae and Cacalia
cordifolia and Senecio hartwegii of the family Compositae.
These plants have little in common with the peyote cactus except
for their pubescence and the fact that sometimes they have been
used medicinally.
The Mexican word "piule," which is generally translated
to mean "hallucinogenic plant," may have come indirectly
from the word "peyote." R. Gordon Wasson, who has studied
many hallucinogenic plants and fungi, suggested that "peyotl"
or "peyutl" became "peyule," which was further
corrupted into ''piule.''[18] "Piule" is also applied
to Rivea corymbosa (Convolvulaceae).
Other names which are apparently variations in spelling (and pronunciation)
of the basic word "peyote" or "peyotl" include:
"pejote," "pellote," "peote," "Peyori,"
"peyot," "pezote," and "piotl."
The many tribes of Indians who use peyote also have words for
the plant in their own languages. However, many also know and
use the word "peyote" as well. Some of the tribes and
their common names are:
Comanche—wokowi or wohoki
Cora—huatari
Delaware—biisung
Huichol—hícouri, híkuli, hícori, jícori, and xícori
Kickapoo—pee-yot (a naturalization of the term "peyote"
into their language )
Kiowa—seni
Mescalero-Apache—ho
Navajo—azee
Omaha—makan
Opata—pe jori
Otomi—beyo
Taos—walena
Tarahumara—primarily híkuli, but also híkori, híkoli, jíkuri, jícoli, houanamé, híkuli wanamé, híkuli walúla saelíami,
and joutouri
Tepehuane—kamba or kamaba
Wichita—nezats
Winnebago—hunka
Numerous other common names have been applied to Lophophora.
These include:
Biznaga (= carrot-like or worthless thing)—commonly applied
to many globose cacti
Cactus pudding
Challote—used principally in Starr County, Texas, one of the
major collecting sites for peyote in the United States
Devil's root
Diabolic root
Dry whiskey
Dumpling cactus
Indian dope
Moon, the "bad seed," "p"—these names have
been applied to peyote by drug users in the United States in the
late 1970s
Raíz diabólica (= devil's root)
Tuna de tierra (= earth cactus)
Turnip cactus
White mule
Part of the confusion with regard to the numerous common names
for Lophophora is because they are frequently applied to
and/or taken from cacti of other genera or plants from another
family entirely.
PLANTS CONFUSED WITH OR CALLED PEYOTE
Two factors have led to the confusion of various plants and the
name peyote: (1) a similarity of appearance because of pubescence,
a globose shape, or growth habit, and (2) a similar physiological
effect or use for medicinal or religious purposes. In fact, most
of the plants that are sometimes called "peyote" possess
both of these characters.
Many alkaloid-containing cacti are commonly called "peyote"
but they are not in the genus Lophophora, and, even though
some of the alkaloids are the same, probably they have few or
no physiological actions similar to the true peyote. Cacti that
have at one time or another been called "peyote" or
the Spanish diminutive "peyotillo" are:
Ariocarpus fissuratus—more frequently called "living
rock" or "chautle" but also "peyote cimarr6n."
A. kotschoubeyanus—usually called "Pezuna de venado"
or "pata de venado."
A. retusus—usually called "chautle" or "chaute."
Astrophytum asterias—surprisingly similar in appearance
to Lophophora.
A. capricorne—also called "biznaga de estropajo."
A. myriostigma—called "peyote cimarr6n," "mitra,"
and "birrete de obispo" (bishop's cap or miter).
Aztekium ritterii—another small, globose cactus with
superficial resemblance to Lophophora.
Mammillaria (Dolichothele) longimamma—sometimes called
"peyotillo."
M. (Solisia) pectinifera
Obregonia denegrii
Pelecyphora aselliformis—commonly called "peyotillo"
and sold as such in the native markets. Contains some of the alkaloids
possessed by Lophophora, including small amounts of mescaline.
Strombocactus disciformis—similar in appearance to Lophophora
and occurring in the same general area as L. diffusa.
Turbinicarpus pseudo pectinata
Other plant families, including the Compositae, Crassulaceae,
Leguminosae, and Solanaceae, also have representatives that occasionally
are called "peyote." A member of the Compositae was
first described as a type of peyote by the Spanish physician,
Francisco Hernandez, in his early study of the plants of New Spain.[19]
In his book he described two peyotes: the first, Peyotl Zacatecensi,
clearly was Lophophora, whereas the other, Peyotl Xochimilcensi,
apparently was Cacalia cordifolia, a Compositae which had
"velvety tubers" and was used medicinally. Other sunflowers
of the closely-related genus Senecio have also been called
such things as "peyote del Valle de Mexico" and "peyote
de Tepic."
"Mescal" is the correct name for the alcoholic beverage
obtained from the century plant, Agave americana, but was
also used by missionaries and officials of the Bureau of Indian
Affairs for peyote. Possibly this was an attempt to confuse Congressmen
and the public into thinking that peyote was an "intoxicant"
similar to alcohol, but it just may have been a case of incorrect
information perpetuated unwittingly.
The name "mescal beans" has also been applied incorrectly
to peyote but actually is the common name of Sophora secundiflora
of the Leguminosae. The beans of this plant contain cytisine,
a toxic pyridine that causes nausea, convulsions, hallucinations,
and even death if taken in too large quantities.[20] The colorful
red beans have been used for centuries both in Mexico and the
United States by the Indians for medicinal and ceremonial purposes,
and sometimes the seeds of this desert shrub are worn as necklaces
by the leaders of peyote ceremonies. The stimulatory and hallucinatory
nature of these beans probably led to the confusion with peyote,
especially when the latter occasionally was called "mescal."
The probable relationship of the old mescal bean ceremony and
the modern peyote cult also may have led to confusion by white
men; this relationship is discussed in chapter 2.
Peyote has also been referred to as the "sacred mushroom";
this confusion probably is the result of the similar appearance
of dried peyote tops and dried mushrooms. Also, there are some
mushrooms that can produce color hallucinations similar to those
of peyote. The Spaniards first misidentified peyote as a mushroom
late in the sixteenth century when they stated that the Aztec
substance "teonanacatl" and peyote were the same; this
mistake was perpetuated by the American botanist, William E. Safford.[21]
He and other reputable scientists insisted that there was no such
thing as the sacred mushroom "teonanacatl"; they believed
that it was simply the dried form of peyote. The problem was resolved
when hallucinogenic mushrooms were rediscovered in 1936 and definitely
linked to early Mexican ceremonies. In recent years at least fourteen
species of hallucinogenic mushrooms have been identified in the
genera Psilocybe, Stropharia, Panaeolus, and Conocybe
of the family Agaricaceae. It is evident that they are well known
to Mexican Indians.[22]
Another plant that has occasionally been confused with peyote
is "ololiuhqui," which is now classified as Rivea
corymbosa of the Convolvulaceae. Ololiuhqui has been
widely used by Indians in the Sierra Madre Occidental of Mexico
for many purposes, such as an aphrodisiac, a cure for syphilis,
an analgesic, a cure for colds, a stimulating tonic, a carminative
(to relieve colic), a help for sprains and fractures, and for
relief of pelvic cramps in women.[23] Recent studies have shown
that the plant contains several potent chemicals which are ergot
alkaloids closely related to LSD.[24] Thus, the effects are somewhat
similar to those of peyote: stimulatory at first and later producing
color hallucinations. Indians could easily see many "divine"
actions resulting from ingestion of the seeds of Rivea
and it is not difficult to understand why they and others may
have confused it with peyote, another "divine" plant.
Several Mexican plants and fungi are hallucinogenic like Lophophora.
The following summary gives the ancient Mexican name, the botanical
name or names, the plant or fungus family, and one or more of
the main psychoactive substances: [25]
Picietl = Nicotiana rustica L. (Solanaceae)
A species of tobacco which contains nicotine.
Teonanacatl = Psilocybe spp.
Panaeolus campanulatus L. var. sphinctrinus (Fr.)
Bresad.
Stropharia cubensis Earle
Conocybe spp.
(All of the above are in the family Agaricaceae)
The psychoactive substances are psilocybin and psilocin.
Pipiltzintzinli = Salvia divinorum Epling&Javito (Labiatae)
The psychoactive principle of this plant is as yet undetermined.
Ololiuhqui = Rivea corymbosa (L.) Hall. fil. (Convolvulaceae)
Tlitlitzen = Ipomoea violacea L. (Convolvulaceae)
Both of the above members of the Convolvulaceae contain the ergot
or Iysergic acid alkaloids;
LSD is a synthetic derivative and
is not believed to occur naturally.
Marijuana is one of the best-known and most widely used substances
currently classified as a hallucinogen. However, there is serious
question whether it actually is a hallucination-producing plant
(at least in the way that it is used by most people)—and it
is of Old World origin. Marijuana is obtained from the genus Cannabis
of the angiosperm family Cannabaceae.[26] It is psychoactive but
has quite different effects than does peyote.
MORPHOLOGY
Morphological studies, including microscopic examinations, have
provided much information about the evolution and relationships
of the cacti. Investigations of both vegetative and reproductive
parts support the proposal that Lophophora is a distinct genus
consisting of two species.
Vegetative parts—The growing point or apical meristem,
located in the depressed center of the plant, is relatively large
and similar to those found in other small cacti. The young leaf,
which arises from the meristem, is difficult to distinguish from
the expanding leaf base and subtending axillary bud. The leaf
base, usually separated from the actual leaf by a slight constriction,
grows rapidly to become the podarium, rib, or tubercle. Thus,
the leaf base functions as the photosynthetic or food-producing
part of peyote. With sufficient magnification the vestigial leaves
of seedlings are often large enough to be identified, but they
are never more than a microscopic hump in the vegetative shoot
of mature peyote plants.
Spines occur only on young seedlings; adult plants produce spine
primordia but they rarely develop into spines.
The caespitose or several-headed condition of the peyote cactus
apparently occurs through the activation of adventive buds that
appear on the tuberous part of the root-stem axis below the crown.
Such growth often is the result of injury and almost always occurs
if the top of the plant is cut off. However, some populations
of peyote seem to have a greater tendency to develop the caespitose
condition than do others.
Epidermal cells, usually five-to six-sided and papillose (nipple-like),
have cell walls only slightly thicker than those of the underlying
parenchyma cells. Sometimes a hypodermal layer can be recognized
early in development, but as the stem matures it does not become
specialized and never differentiates from the underlying palisade
tissue. Normally the epidermis is covered by both cuticle and
wax; the latter substance is primarily responsible for the blue-green
or glaucous coloration of L. williamsii. Stomata are abundant,
especially on the younger, photosynthetically active parts of
the vegetative body. They are paracytic and usually subtended
by large intercellular spaces. The subsidiary cells of a stoma
usually are about twice the size of neighboring epidermal cells.
Trichomes are persistent for many years in the form of tufts of
hairs or "wool" arising from each areole. They tend
to be uniseriate on the younger areoles but are often multiseriate
on older ones.[27]
Ergastic substances are evident in the cortex of peyote. Usually
they are druses of calcium oxalate which often exceed 250 microns
in diameter, but which rarely are found within one millimeter
of the epidermal layer. These anisotropic crystals can be easily
seen if fresh or paraffin-embedded sections are examined in polarized
light. Mucilage cells do not occur in the vegetative parts of
peyote but are found in flowers and young fruits.[28]
The chromosome number of peyote, like most other cacti, is 2n
= 22. The root tip chromosomes are quite small, and apparently
there is no variation from the basic chromosome number of the
Cactaceae which is n= 11.
Reproductive parts—Peyote flowers, in contrast to those
of other cactus genera such as Echinocactus and most of
the Thelocacti, have naked ovaries or the absence of scales on
the ovary wall, a character shared with the flowers of Mammillaria,
Ariocarpus, Obregonia, and Pelecyphora. Thus, in Lophophora
all floral parts are borne on the perianth tube above the ovule-containing
cavity. The flower color of Lophophora varies from deep
reddish-pink to nearly pure white; those of L. diffusa
rarely exhibit any red pigmentation, making them usually appear
white or sometimes a light yellow because of the reflection of
yellow pollen from the center of the flower. Development of peyote
flowers is much like that of Mammillaria.
Pollen of Lophophora is highly variable. Pollen of the
Dicotyledonae tend to have three apertures or pores, while those
of the Monocotyledonae usually have only one aperture. Peyote
pollen varies greatly in aperture number, the northern population
having 0-18 and the southern population 0-6. Though the grains
are basically spheroidal and average about 40 microns in diameter,
the varying numbers of colpae or apertures produce about twelve
different geometric shapes. Such a variety from a single species
or even population is rare in flowering plants. The pollen of
L. diffusa has less variation than that of L. williamsii;
it also has a much higher percentage of grains that are of the
basic tricolpate (three-aperturate) type. Thus, the basic dicotyledon
pattern is best observed in the southern population, whereas more
complex grains occur in the northern localities. Small, tricolpate
grains probably are more typical of the ancestors of the cacti
and the more elaborate geometric designs of L. williamsii
seem to represent greater evolutionary divergence and specialization.[29]
Fruits of peyote are similar to those of Obregonia and
Ariocarpus in that they develop for about a year and then
elongate rapidly at maturity. The fruits of Lophophora
and Obregonia usually have only the upper half containing
seeds whereas they are completely filled with seeds in Ariocarpus.
The seeds of Lophophora are black, verrucose (warty), and
with a large, flattened, whitish hilum. They are virtually identical
to those of Ariocarpus and Obregonia although there
are some minor structural differences of the testa.
Lophophora seems to stand by itself in possessing a particular
combination of morphological characters unlike any other group
of cacti. Its nearest relatives appear to be the genera Echinocactus,
Obregonia, Pelecyphora, Ariocarpus, and Thelocactus.
The character of seeds, seedlings, areoles, and fruits certainly
support the contention that peyote belongs in the subtribe Echinocactanae
(sensu Britton and Rose) rather than in the more recently
proposed "Strombocactus" line of Buxbaum. Perhaps the
poorly understood genus Thelocactus may be the single most
closely related group.[30]
BIOGEOGRAPHY
The genus Lophophora is one of the most wide-ranging of
all the plants occurring in the Chihuahuan Desert; it has a latitudinal
distribution of about 1,300 kilometers (800 miles), from 20 degrees,
54 minutes to 29 degrees, 47 minutes, North Latitude
(figure 8.2).
Within the United States L. williamsii is found in the
Rio Grande region of Texas. There is a small population occurring
in western Texas near Shafter; it occurs in the Big Bend region,
and then it is found in the Rio Grande valley eastward from Laredo.
Peyote extends from the international boundary southward into
Mexico in the basin regions between the Sierra Madre Occidental
and the Sierra Madre Oriental to Saltillo, Coahuila; this vast
expanse of Chihuahuan Desert in northern Mexico covers about 150,000
square kilometers (60,000 square miles). Just south of Saltillo
the range of peyote narrows, is interrupted by mountains, and
then expands again eastward into the foothills of the Sierra Madre
Oriental and westward into the state of Zacatecas. It extends
southward nearly to the city of San Luis Potosi where its distribution
terminates (figures 8.3 and 8.4). The southern population of peyote,
that of L. diffusa, is restricted to a high desert region
in the state of Queretaro. This area of about 775 square kilometers
(300 square miles) is isolated from the large northern populations
by high, rugged mountains (figures 8.5 and 8.6).
Three factors apparently are responsible for the discontinuous
distribution of Lophophora between the large northern and
the smaller southern population: (1) extensive saline flats in
the Rio Verde region east of the city of San Luis Potosi, (2)
formidable mountains: the Sierra Gorda extension of the Sierra
Madre Oriental, and (3) high elevations even in the broad valleys.
The relatively high desert area in Queretaro apparently is an
isolated pocket of the Chihuahuan Desert.
There are great elevation differences from the north to the south
within the total range of peyote; the Rio Grande peyote occurs
at an elevation of about 50 meters (150 feet), but in the southern
portion of its range in the state of San Luis Potosi it is found
at nearly 1,850 meters (6,000 feet) elevation. The elevation of
the southern population in Queretaro is about 1,500 meters (5,000
feet).[31]
It is unclear to what extent human beings have affected the distribution
of peyote. There are areas where man has collected large quantities
of the plant, such as near Laredo, Texas; near Matehuala, San
Luis Potosi; and in the dry desert valley area of Queretaro. In
1961 I collected L. diffusa in a region near the road going
north from Vizarron, Queretaro; in 1967 I returned to the same
area but could find no peyote. Farmers living nearby told us that
about a year earlier a man from a nearby village whom they called
a "Padre" hired workers to collect all of the peyote
that they could find in the region. The farmers didn't know why
the man had wanted so many plants or what he planned to do with
them, but I doubt that they were used for religious or medicinal
purposes. Probably they were sold to cactus collectors—or perhaps
even destroyed. Fortunately, peyote is a common and widespread
plant and it occurs in many areas that are almost inaccessible.
However, we may see considerable disturbance and loss of peyote
populations in areas easily reached by man.
ECOLOGY
The Chihuahuan Desert where peyote occurs is a type of warm-temperate
desert biome. This region has considerable variation in both topography
and vegetation, which has prompted ecologists to describe numerous
subdivisions. Unfortunately, these subdivisions are not alike
nor have they received the same names. Following the classification
of the Mexican botanist, Jerzy Rzedowski, peyote occurs primarily
in two subdivisions of the Chihuahuan Desert: ( 1 ) the microphyllous
desert scrub, which has shrubs that are leafless or have small
leaves and are represented by such plants as Larrea tridentata,
Prosopis laevigata, and Flourensia cernua; and (2)
the "rosettophyllous" desert scrub, with many plants
bearing rosettes of leaves, such as Agave lecheguilla and
Yucca spp.[32] Probably neither of these vegetation subdivisions
can be considered climax communities, nor even formations, because
there is continuous mixing of the two life forms. Since there
is such confusion between these two subdivisions, perhaps Cornelius
H. Muller's general term "Chihuahuan Desert Shrub" should
be used to describe the general area in which peyote occurs.[33]
The well-isolated southern population apparently is outside the
region normally included within the Chihuahuan Desert. However,
the presence of Larrea tridentata and other plants typical
of this type of desert is an indication that it should, indeed,
be included within the Chihuahuan Desert.
The soils of the Chihuahuan Desert Shrub are limestone in origin
and have a basic pH, from 7.9 to 8.3. These soils can also be
characterized as having more than 150 ppm (parts per million)
calcium, at least 6 ppm magnesium, strong carbonates, and no more
than trace amounts of ammonia. The soils test negatively for iron,
chlorine, sulfates, manganese, and aluminum. Phosphorus and potassium
vary somewhat throughout the range, but in most localities occur
in trace amounts or are not present at all. Soils from the southern
locality in Queretaro are not different from those to the north.[34]
As stated earlier, peyote occurs in diverse habitats of the Chihuahuan
Desert, and no particular plants are associated with it in all
localities. Only Larrea tridentata (creosote bush) is found
in more than 75 percent of the peyote sites studied; other plants
commonly found with peyote and their percentage of occurrence
in the sites analyzed are: [35]
Jatropha dioica (leatherplant)—70 percent
Echinocereus spp. (hedgehog cactus)—70 percent
Opuntia leptocaulis (pipestem cactus)—70 percent
Prosopis laevigata (mesquite)—70 percent
Agave lecheguilla (lechuguilla)—50 percent
Echinocactus horizonthalonius (eagle claws cactus)—50
percent
Mammillaria spp. (fishhook or nipple cactus)—50 percent
Flourensia cernua (tarbush)—50 percent
Acacia spp. ( acacia )—40 percent
Condalia spp. (lotebush)—40 percent
Coryphantha spp.—40 percent
Neolloydia spp.—40 percent
Yucca filifera (yucca)—40 percent
Hamatocactus spp.—40 percent
The following plants, supposedly typical of the Chihuahuan Desert,
occurred in less than 40 percent of the peyote sites studied:
Coldenia canescens
Euphorbia antisy phylitica ( wax plant )
Koeberlinia spinosa (crucifixion thorn)
Of course not all perennial plants growing with peyote have been
cited, but this information indicates that peyote occurs over
a broad range of vegetation types within the Chihuahuan Desert.
The climatic data from the regions in which peyote grows have
been analyzed to obtain an "index of aridity." Using
the index of aridity devised by Consuelo Sota Mora and Ernesto
Jauregui O. of the University of Mexico, [36] peyote is found
to tolerate a very wide range of climatic conditions: precipitation
ranges from 175.5 mm up to 556.9 mm per year, maximum temperatures
vary from 29.1 degrees centigrade to 40.2 degrees, and minimum
temperatures range from 1.9 to 10.2 degrees centigrade. There
is also a variation in the time of year that precipitation occurs.
Rains typically fall in the late spring and summer in the Chihuahuan
Desert, but in certain areas some winter rains do fall. There
are peyote populations in both types of areas, so probably they
should be classified as being in intermediate rather than strictly
summer rainfall regions. The modified index of aridity, which
is based on the relationship of temperatures and precipitation,
shows that Lophophora exhibits a wide range of aridity,
between 64.0 and 394.0. It also appears that the index of aridity
is related to elevation, although there are some definite exceptions,
such as in Queretaro, where there is a relatively high elevation
(about 1,500 meters or 5,000 feet) but an index of aridity that
is over 115. This southern habitat, though of high elevation,
may be especially arid because of the proximity of surrounding
high mountains which cause a more intensified rain shadow.
CHARACTERISTICS OF PEYOTE POPULATIONS
Peyote consists of populations that are not only wide-ranging
geographically, but which are also variable in topographical locations,
appearance, and methods of reproduction. Commonly peyote is found
growing under shrubs such as Prosopis laevigata (mesquite),
Larrea tridentata (creosote bush), and the rosette-leafed
plants such as Agave lecheguilla; at other times, however, it
grows in the open with no protection or shade of any kind. In
some areas, such as in the state of San Luis Potosi, peyote sometimes
grows in silty mud flats that become temporary shallow fresh-water
lakes during the rainy season. In west Texas peyote has even been
found growing in crevices on steep limestone cliffs.
The appearance of peyote also varies widely, especially in the
species L. williamsii. In some cases the plants occur as
single-headed individuals and in others they become caespitose,
forming dense clumps up to two meters across with scores of heads.
Plants in Texas do not seem to form clumps as often as those in
the state of San Luis Potosi, but plants with several tops can
arise as the result of injury by grazing animals or other factors.
Many-headed individuals are also produced by harvesting the tops.
In Texas, for example, collectors normally cut off the top of
the plant, leaving the long, carrot-shaped root in the ground;
the subterranean portion soon calluses and in a few months produces
several new tops rather than just a single one like that which
was cut off.
The number of ribs present in a single head varies widely, rib
number and arrangement apparently being in part a factor of age,
as well as a response to the environment. Rib number within a
single, genetically identical clone may vary from four or five
in very young tops up to fourteen in large, mature heads (figure
8.4). At other times there are bulging podaria instead of distinct
ribs. Field studies have shown that rib number and variation apparently
are due to localized interactions between genotype and environment.
Because of the high degree of variation occurring in a single
population, rib characteristics alone are of little value in the
delimitation of formal botanical taxa.
Reproduction occurs mainly by sexual means. The plants flower
in the early summer, and the ovules, which are fertilized during
that season, mature into seeds a year later. The fruit which arises
from the center of the plant late in the spring or early in the
summer rapidly elongates into a pink or reddish cylindrical structure
up to about one-half inch in length. Within a few weeks these
fruits mature; their walls dry, become paper thin, and turn brownish.
Later in the summer, usually as a result of wind, rain, or some
other climatic factor, the fruit wall ruptures and the many small
black seeds are released. The heavy summer rains then wash the
seeds out of the sunken center of the plant and disperse them.
Another method of reproduction in peyote is by vegetative or asexual
means. Many plants produce "pups" or lateral shoots
which arise from lateral areoles. After these new shoots have
attained sufficient size they can often root and survive if broken
off. If these new portions successfully grow into new plants,
they are genetically identical to their parents. Surprisingly,
peyote plants rarely rot if injured or cut, so excised pieces
will readily form adventitious roots and can become independent
plants.
EVOLUTION OF PEYOTE
The evolutionary history of the cacti is not documented by fossils
because their succulent vegetative parts did not lead to preservation
as fossils in the dry climate. The highly specialized cactus has
few distinctive characteristics that probably were present in
distant ancestors, but it does appear that the tropical leafy
cactus, Pereskia, may represent a form that has changed
little from the non-cactus ancestral types. It and many of the
more specialized cacti have many characteristics similar to the
other ten families of the order Caryophyllales (Chenopodiales)
in which the cacti are often placed. Most of these families, for
example, have a curved embryo, the presence of perisperm rather
than endosperm, either basal or free-central placentation, betalain
pigments rather than the usual anthocyanins, anomalous secondary
thickening of the xylem walls, and succulence.[37]
The evolutionary picture from Pereskia is only hazy at best, although
Pereskiopsis seems to represent an intermediate form in
the Opuntia line. The "barrel" or "columnar"
cacti, on the other hand, show virtually no links to one another
or to any of the more "primitive" cacti such as Pereskia
or Pereskiopsis. Apparently the living representatives
of the cacti are terminal points of a highly branched evolutionary
history, and ancestors no longer exist. Therefore, we must work
with characters of living representatives to draw any conclusions
regarding the past evolutionary history of the cacti, a procedure
of speculation at best.
Certain evolutionary trends appear evident in the two species
of peyote. Pollen of L. diffusa, because of its higher
percentage of the basic tricolpate type of grain, could be considered
more primitive than that of L. williamsii. Likewise, James
S. Todd and other chemists have shown that certain of the more
elaborate alkaloids are either absent or in lesser amounts in
L. diausa.38 This, they feel, indicates that L. diffusa
may not have evolved and diversified to as great an extent chemically
as has L. williamsii. Also, the greater variation of the
vegetative body of L. williamsii, in addition to more varied
habitats and a wider distribution, perhaps show a more diverse
and highly evolved gene pool.
Lophophora probably arose from a now-extinct ancestor that
occurred in semi-desert conditions in central or southern Mexico.
Morphological and chemical diversity may have then appeared in
various populations as they slowly migrated northward into drier
regions which were being created by the slow uplift of mountains.
Perhaps L. diffusa represents one of the earlier forms
that became isolated in Queretaro, whereas L. williamsii
spread more extensively to the northward, producing new combinations
of genes that eventually led to a distinct but highly variable
species having somewhat different pollen, vegetative characters,
and alkaloids from the peyote populations to the south.
CULTIVATION
Peyote is easily cultivated and is free-flowering. On the other
hand, one must be very patient if he wishes to grow peyote from
seed, as it may take up to five years to obtain a plant that is
15 millimeters in diameter. At any stage, however, peyote can
be readily grafted onto faster-growing rootstocks, and this usually
triples or quadruples the plant's rate of growth. Japanese nurserymen,
for example, have obtained peyote plants large enough to flower
within a period of 12-18 months by grafting the young seedlings
onto more robust root stocks.
To insure the obtaining of fertile seed, it is advisable to out-cross
peyote plants by transferring with forceps some stamens containing
pollen from the flower of one plant to the stigma of the flower
of another.
Propagation can also be accomplished by removing small lateral
tops from caespitose individuals. The cut button or top should
be allowed to callus for a week or two and then planted in moist
sand or a mixture of sand and vermiculite. It is wise to dip the
freshly cut portion in sulfur to facilitate healing. Rooting is
best done in late spring or early summer. Eventually a new root
system will develop from the top; the old root will produce several
new heads to make a caespitose individual.
Soil conditions for the cultivation of peyote are not too critical.
As the natural soil for peyote is of limestone having a basic
pH, one should provide adequate calcium, insure that the soil
is slightly basic, and provide good drainage. Peyote should be
watered frequently (every four to seven days) in the summer but
very little or none at all in the winter. Fertilizer should be
applied while the plants are being watered during the growing
season, especially May through July.
Peyote hosts few insect pests and does not need to be treated
differently from other cultivated cacti and succulents with regard
to pesticides.
Greenhouse-grown peyote plants sometimes develop a corky condition;
this brownish layer often covers most of the plant and is not
natural. Its cause is not known.
The propagation of seeds is a rewarding experience but requires
great patience. Seeds should be sowed on fine washed sand and
then covered with one to two millimeters (about one-eighth inch)
of very fine sand. Cover the flat or pot with a plastic bag or
plate of glass and place an incandescent light (60 watt) or Grolux
lamp about twelve inches above the sand. These provide both heat
and light. The sand should be kept moist to insure that the humidity
is high and that the young plants will not dry out as they first
sprout. Germination usually occurs within two or three weeks but
growth of the seedlings is exceedingly slow. The plants should
be transplanted and thinned after they are about one centimeter
(one-fourth of an inch) in diameter.
Most states, as well as the federal government, now prohibit the
possession of peyote (see chapter 9), and apparently one is in
violation of the law even if peyote is grown as part of a horticultural
collection.
NOTES TO CHAPTER 8
1. Jan G. Bruhn and Bo Holmstedt, "Early Peyote Research:
An Interdisciplinary Study," pp.384-85.
2. William Jackson Hooker, "Tab. 4296. Echinocactus Williamsii."
3. Theodor Rumpler, Carl Friedrich Forster's Handbuch der Cacteen
kunde, p. 233.
4. Charles Lemaire, Cactearum Genera Nova et Species Nova en
Horto Monville, pp. 1-3.
5. J. Lanjouw and others (eds.), International Code of Botanical
Nomenclature, p. 50.
6. Bruhn and Holmstedt, "Early Peyote Research," pp.
358-60.
7. Paul Hennings, "Eine giftige Kaktee, Anhalonium lewinii
n. sp.," p.411.
8. Edward F. Anderson, "The Biogeography, Ecology, and Taxonomy
of Lophophora (Cactaceae)," pp. 305-06.
9. Bruhn and Holmstedt, "Early Peyote Research," pp.
384-85.
10. John M. Coulter, "Manual of the Phanerogams and Pteridophytes
of Western Texas," p. 129.
11. A. Voss, "Genus 427. Ariocarpus Scheidw. Aloecactus,"
p. 368.
12. John M. Coulter, "Preliminary Revision of the North American
Species of Cactus, Anhalonium, and Lophophora,"
pp. 131-32.
13. Anderson, "Biogeography, Ecology, and Taxonomy,"
pp. 299-303.
14. Fr. Bernardino de Sahagun, Historia General de las Cosas
de Nueva Espana, X, p. 118.
15. Richard Evans Schultes, "Peyote (Lophophora williamsii)
and Plants Confused with It," pp.61-88.
16. Ibid.
17. A. de Molina, Vocabulario de la Lengua Mexicana, p.
80.
18. R. Gordon Wasson, "Notes on the Present Status of Ololiuhqui
and the Other Hallucinogens of Mexico," pp. 166-67.
19. Francisco Hernandez, De Historia Plantarum Novae Hispaniae,
pp. 70-71.
20. Richard Evans Schultes and Albert Hofmann, The Botany and
Chemistry of Hallucinogens, p. 99.
21. William E. Safford, "An Aztec Narcotic," p. 311.
22. Schultes and Hofmann, Botany and Chemistry of Hallucinogens,
pp. 36-39.
23. Hernandez as reported in Schultes, "Peyote and Plants
Confused with It," p.74.
24. Schultes and Hofmann, Botany and Chemistry of Hallucinogens,
pp. 144-51.
25. Wasson, "Notes on Ololiuhqui," pp. 164-75.
26. Richard Evans Schultes, William M. Klein, Timothy Plowman,
and Tom E. Lockwood, "Cannabis: An Example of Taxonomic
Neglect," pp.360-62.
27. Norman H. Boke and Edward F. Anderson, "Structure, Development,
and Taxonomy in the Genus Lophophora," p. 573.
28. Ibid., pp.573-74.
29. Edward F. Anderson and Margaret S. Stone, "A Pollen Analysis
of Lophophora (Cactaceae)," pp. 77-82.
30. Boke and Anderson, "Structure, Development, and Taxonomy,"
p. 577.
31.Anderson, "Biogeography, Ecology, and Taxonomy,"
pp. 301-02.
32. Jerzy Rzedowski, "Vegetacion del Estado de San Luis Potosi,"
pp. 219-20.
33. Cornelius H. Muller, "Vegetation and Climate of Coahuila
Mexico," p. 38.
34. Anderson, "Biogeography, Ecology, and Taxonomy,"
p. 302.
35. Ibid., pp. 302-03.
36. Consuelo Soto Mora and Ernesto Jaurequi O., Isotermas Extremas
e Indice de Aridez en la Republica Mexicana, pp. 26-28.
37. Arthur Cronquist, The Evolution and Classification of Flowering
Plants, pp. 177—80.
38. James S. Todd, "Thin-layer Chromatography Analysis of
Mexican Populations of Lophophora (Cactaceae)," pp.
395-98.
Figure 8.2
Natural distribution of the two species
of peyote, Lophophora williamsii and Lophophora diffusa.
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