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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