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.
Mescaline
also called -3,4,5-trimethoxyphenethylamine,
naturally occurring
alkaloid, the active principle contained
in the flowering heads of the
peyote cactus (species Lophophora williamsii)
of Mexico and the
southwestern United States, that has been
used as a drug to induce
hallucination. The mescaline molecule is
related structurally to two
hormones secreted by the adrenal glands,
adrenaline and
noradrenaline; both are catecholamine compounds
that take part in
the transmission of nerve impulses. Mescaline
was isolated as the
active principle of peyote in 1896, and its
structural resemblance to
adrenaline was recognized by 1919.
In experiments mescaline requires 2 to 3 hours
for onset of action,
and its effects sometimes last for more than
12 hours. The
hallucinatory effects vary greatly among
individuals and even for a
particular individual from one drug session
to the next. The
variations seem to reflect such factors as
the mood and personality
of the subject and the setting in which the
drug is administered.
Hallucinations are usually visual, less often
auditory. Side effects
include nausea and vomiting. Mescaline is
prepared from the
peyote cactus by extraction and purification,
but it can be
synthesized.
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