Martin Terry

Stalking the wild Lophophora

Part 1  Chihuahua and Coahuila

S
Ay, Chihuahua!
ul Ross State University gradu-
ate student Robert Hibbitts and I
set off in my old Dodge truck from
Alpine, Texas, in late May and
crossed into Mexico at Presidio-
Ojinaga, which is the only official
border crossing between El Paso, at
the westernmost corner of Texas,
and Del Rio, some 500 miles downstream. Presi-
dio, Texas is a town of about 3000 souls. Ojina-
ga, on the Chihuahua side, is considerably larg-
er, famous for its delicious asadero cheese and
its high murder rate. From here a scenic two-
lane, Mexican Highway 16, winds through the
massive mountains of northern Chihuahua to
the state capital, Ciudad Chihuahua, where we
arrived on the campus of the agricultural school
of the Autonomous University of Chihuahua on
the southwestern edge of the city. Hoping to
receive guidance with regard to the exact loca-
tions of populations of Lophophora williamsii
on the western edge of its range (and also of the
range of the genus), we were disappointed to
learn that no one had managed to locate a single
Lophophora population in the state—valuable
information nonetheless. Contrary to the report
of Robert Bye, quoting anonymous sources sug-
gesting that Lophophora occurred in “the hills
west of Chihuahua City”1, Dr Toutcha Zebgue
told us that such an occurrence was extreme-
ly unlikely. Not only had peyote not been found
there, but the geology was all wrong: igneous
rock, rather than the limestone normally pre-
ferred by Peyote.
Disappointed but undaunted, we headed south-

On a limestone mountainside south of Viesca,

Coahuila we found a montane population of L. fricii.
Most plants here were single-headed. The unusual
color of these plants is characteristic, as is their lack of
prominent raised ribs that are typical of L. williamsii.

2008 Volume 80 Number 4 181

 The exclusive Little Cow Motel, Matamoros, Coahuila. A shower-curtain-like arrangement allows customers to
hide their vehicles from public view, thus marginally decreasing the likelihood that their spouses will learn of
extramarital dalliances.

east on federal Highway 45 (a well-maintained toll The objective of our trip was to collect tis-
road equivalent to an interstate highway in the sue samples from 15–20 representative popula-
US) toward the town of Julimes, where we had tions covering most of the Mexican portion of
GPS coordinates for the one and only Chihuahuan the geographic range of the genus Lophophora,
Lophophora locality in the entire UNAM herbari- including populations of all four (now five!) of the
um database. From Julimes we veered west toward Lophophora species that are generally recognized
the Sierra de la Amargosa, a range noted a centu- as valid by Continental European systematists,
ry ago by Lumholtz as an area
to which the Tarahumara had This Lophophora fricii at the Laguna site has 21 ribs, which is common
traditionally traveled from here and in L. diffusa in Querétaro. Most globular cacti have stable
their homelands to harvest configurations of ribs at Fibonacci numbers (1, 1, 2, 3, 5, 8, 13, 21…).
the hikuli (L. williamsii) that For instance, Astrophytum asterias and Echinocactus horizonthalonius
they used for medicinal and normally have eight ribs all their lives. Lophophora williamsii starts life with
ceremonial purposes2. We five ribs, transitions to eight by the time it reaches 5–6 cm in diameter,
drove fast down arroyo-cut then switches to 13 ribs as a large adult (usually > 8 cm in diameter).
dirt roads as the sun set, tak- L. diffusa and L. fricii follow the same pattern, but as large old plants
ing successive forks in the they frequently jump up to the next Fibonacci number in the series.
road too small to appear on
the map, hoping to cover the
remaining distance to our
GPS coordinates before night-
fall. As darkness descended
we found ourselves on a road
running perpendicular to the
direction we needed to go,
so we stopped to camp about
a mile from our destination.
Ranch-style beans (eaten
cold, right out of the can) and
tortilla chips served as sup-
per, and sleeping bags on the
ground beside the truck pro-
vided a welcome buffer from
the chilly spring-night wind
of the Chihuahuan Desert.

182 Cactus and Succulent Journal

including the taxonomically contentious L.  fricii
and L.  koehresii, which are not yet accepted as
valid species by the Anglo-American oligarchy of
cactus taxonomy3. The ultimate objective was to
extract DNA from the tissue samples and use it to
generate genetic data that we believe will consti-
tute the substance of the first thorough study of
the molecular systematics of the genus Lophopho-
ra (but more on that later).
The next morning we were up at dawn, hik-
ing across the broken terrain toward our quar-
ry. But as we reached the point where our Chi-
huahuan Lophophora should have been, none
could be found. The habitat was perfect, but all
we saw were shovel-shaped holes under Larrea
nurse plants where peyote plants had been dug
up, roots and all. The absolute thoroughness of
the spoilers who dug the plants was amazing. We
could find no remaining Lophophora plants—
not even a seedling—in several hours of search-
ing in ever-widening spirals from the GPS way-
point. Poachers had removed every one.
Coahuila
Admitting defeat in Chihuahua, we exchanged
some dollars for pesos, filled the tank with diesel,
and moved on to the Laguna de Viesca in south-
west Coahuila. Taking Highway 45 southeast
past the Bolsón de Mapimí to Gómez Palacio,
then east around Torreón to the town of Mata-
moros, we arrived about midnight and stayed in

On the northern side of the Laguna de Viesca, in silty mesquite-agave flats, a much different form of
Lophophora fricii forms large clumps by repeated lateral branching of the original stem and its branches.
The branching process starts when a plant with a single crown reaches the size of a large saucer. Then
branches (consisting of small crowns) begin to erupt from the areoles at the perimeter of the parent
plant’s crown. When the new branches reach a certain size, they put down their own tap roots, making
them independent of the parent plant that produced them, and then they begin to branch in turn. The
result, which probably takes several decades to manifest itself and has no obvious endpoint, is a large
clump of dozens of more or less connected plants, ranging in size from new branches that may be no
more than a centimeter in diameter, to very large 21-ribbed plants that may exceed 10 cm across.

2008 Volume 80 Number 4 183

Habitat of the lowland form of L. fricii in the northeastern sector of the Laguna de Viesca consists of
mesquite and creosote scrub set in with small agaves. Large clumps of L. fricii are common here. The light-
colored, fine, silty soil is typical of the Laguna, which is bounded by the mountains in the background.

Tissue samples­— What are they all about?

DNA, the genetic material found in all living organisms, carries all the informa- the magnificent Motel La Vaquita
tion that makes the organism what it is and enables it to function and repro- (the Little Cow Motel), whose sign
duce. Because the sequences of bases in DNA inevitably change over time by portrayed a small happy black
such processes as mutation and natural selection, different sequences often and white cow. After the manag-
develop in populations inhabiting different geographic areas. We can therefore
er ascertained that we wanted the
get an idea of the degrees of relatedness among individuals from different
populations by examining and analyzing their DNA sequences. room for the whole night, he gave
The first step in such a study is to collect tissue samples from which to us the key in exchange for 140
extract the DNA. In humans this is relatively simple, as DNA can be easily pesos (about $12.50). The room
extracted from cells that can be painlessly scraped from inside the mouth. In boasted a window air-condition-
many plants, leaf tissue is suitable for DNA extraction. But cacti (wouldn’t you er (which meant you opened the
know?) are more difficult. It’s not that there is any shortage of DNA in cactus window if you wanted relatively
tissues. Au contraire, there is abundant DNA in the stem, particularly in those
cool night air to come in from out-
subdermal green cells where DNA is busy choreographing photosynthesis. The
problem is that cacti also produce large quantities of mucopolysaccharides side), a most welcome shower, and
(yes, that’s the same muco as in mucus), which give the inner stems of cacti their a television, on which we watched
sliminess. Molecular biologists dread these long, branched chains of sugars, the news until we fell asleep.
because they behave like a net and entangle the DNA in the test tube. DNA is At dawn the next day we drove
therefore rather difficult to isolate from the primordial cactus goo. to Viesca, a small city at the end of
One way of minimizing the problems associated with the cactus-stem the paved highway heading south-
polysaccharides is to use liquid nitrogen to disrupt the tissue in the lab. But that
east out of Torreón. Viesca is home
assumes that one has access to liquid nitrogen. Another option is to use tepal
tissue instead of stem tissue. But that assumes that one will find flowers in the to known populations of Lophopho-
field. A third recourse is to collect the epidermis of the stem. The epidermis of ra fricii4, and because this species
Lophophora is exquisitely thin, so that in a sample the size of a thumbnail one is the subject of taxonomic contro-
may get only a milligram or two of dried epidermis. But we have found that to versy5, I considered it important to
be enough to yield sufficient DNA for analysis. obtain tissue samples for DNA anal-
Our strategy was to collect tissue samples from ten plants per population, ysis from one or more of the popu-
with the constraint that we would not sample any plants located less than 10
lations in the town’s vicinity. In fact
meters from plants already sampled. The intent of this 10-meter minimum dis-
tance rule was to exclude plants most likely to be related as parents and prog- there are populations of very un-
eny. Having selected a plant for sampling, the person doing the sampling would williamsii-looking lophophoras at
cut (with a clean knife blade) a piece of tissue about the size of a thumbnail and several spots around the perimeter
weighing 100–200 milligrams from the crown (the above-ground part of the of a geologic feature known as the
stem) of the plant. The tissue sample was then placed in a snack-size zip-lock Laguna de Viesca6, a dry depres-
plastic bag labeled with the code for the location and the specimen number. sion that can become an ephemer-
The samples were carried in my backpack until we returned to the truck, where-
al shallow lagoon during the rainy
upon they were transferred to an ice chest, where they remained cooled until
we reached the lab where the DNA was extracted. A photo was taken of each season. We collected tissue samples
cactus from which a sample was taken, and notes were made of the plant’s from one spot in the mountains on
physical measurements and habitat description, including companion plants.

184 Cactus and Succulent Journal

the southwest side of the Laguna and from anoth- sampled are also strikingly different from each
er in the silty mesquite-creosote-agave flats on its other. One is a rock-loving, montane population;
northeastern perimeter. the other occurs in loose alluvial soils of lowland
These plants are notably different from typ- flats. In the montane population the plants are
ical specimens of L.  williamsii. In stem mor- predominantly solitary (single-stemmed); in the
phology, in particular the flattened tubercles, bed of the laguna, the mature plants are largely
the “diffuse” bound-
aries between adja-
cent ribs, and the hor-
izontal “double chin”
basal folds around
the stems of the larg-
er plants, Laguna
de Viesca plants are
more similar to L. dif-
fusa than to L.  wil-
liamsii, which nor-
mally shows sharp-
ly demarcated, raised
ribs, with no horizon-
tal folds of tissue run-
ning perpendicular to
the ribs around the
base of the stem. And
where L. williamsii is
normally found grow-
ing in frank limestone
or calcareous soils,
L.  fricii plants from
the southwest side
of the Laguna were Lophophora williamsii flowering on the south slope of a limestone
growing from crev- mountain near Cuatro Ciénegas, Coahuila. The color and morphol-
ices in a hard type ogy of these plants, as well as the limestone mountainside they inhabit,
of limestone unfa- are virtually identical to plants and habitats in Trans-Pecos Texas.
miliar to me, and L  On the northern slope of a limestone mountain north of Saltillo, Lophophora
those from the north- williamsii is found under nurse shrubs, cryptic and partly covered with fallen
east side of the Lagu- leaves from the nurse shrubs. There were mature peyote plants here, but they
na were growing in were neither large nor abundant, suggesting that this population had been
loose, silty, alluvial harvested in the not-too-distant past. That is hardly surprising, given the easy
soil with no apparent access provided by the major highway connecting Saltillo and Monterrey.
limestone influence.
Chemical differenc-
es are known from
phytochemical work
done on these spe-
cies as well. The pre-
dominant alkaloid in
L. fricii is (non-hallu-
cinogenic) pellotine,
the same as in L. dif-
fusa and L.  koeh-
resii—not mesca-
line, the most abun-
dant peyote alkaloid
in L. williamsii7.
The two popula-
tions of L.  fricii we

2008 Volume 80 Number 4 185

 caespitose, forming clumps approaching a meter of those pressurized products that squirts white
in length. Although we were unable to find flow- goo in a tire to plug a leak, we were able to reach a
ering plants under the parched conditions we ranch house where they had a bicycle pump that
encountered in late May, further differences may filled the wounded tire with enough air to get us
yet be revealed: it will be interesting to see if back to the nearest vulcanizador.
there are substantial genetic differenc-

O
es between these two pop- Mescaline ur next stop was Cua-
ulations, separated by only tro Ciénegas, a lovely
about 30  km but marked- oasis in central Coa-
ly distinct in their chemis- huila, where it was relative-
try, morphology, and habit, as ly easy to locate a healthy
well as their habitat. population of L.  william-
sii from the list of localities

N
othing beats the exhil- in the UNAM database. The
aration that comes plants were growing in crevices
with new heights of and on natural terraces along the
accomplishment, such as Pellotine slopes of limestone hills. Mor-
when we managed three phologically and ecologically
flat tires in one day. The they were indistinguishable
first one was occasioned by from plants that occur in
yours truly looking at the west Texas. This is not sur-
scenery of a small hill arising prising when you consider
unexpectedly out of the flats that Cuatro Ciénegas is only
of the Laguna de Viesca, instead 250  km from the southern tip of
of looking at the road. A little stump of a the Big Bend.
long-dead mesquite tree on the side of the Our next stop was on the northern
dirt road was sufficient to stab a gaping sidewall outskirts of Saltillo, on Highway 40 in the direc-
puncture in the right-front tire. That left us with tion of Monterrey. L. williamsii was not abundant
a shoddy spare tire on the road and a forever-use- at this site, but in the course of an hour and a half
less flat in the bed of the truck. Not thrilled with we were able to collect stem tissue samples from
the prospect of a second flat and no functional ten individuals separated from each other by a
spare in this remote corner of Coahuila, we decid- distance of at least 10 meters. Most of the plants
ed to head for the big city of San Pedro to buy a were found growing under nurse shrubs and
new tire. We ended up buying two, but attempts were partially obscured by leaf litter. In the more
to locate a used wheel for my old truck, so as to northern regions of its range L. williamsii shows
have two mounted spare tires rather than one, an absolute preference for the south slopes of hills
met with failure. and ridges8. At the lower latitude around Saltillo,
As we started back to the Laguna de Viesca an in contrast, the plants occurred on both northern
impressive storm arose—no trace of rain, but the and southern slopes, interspersed with some fab-
afternoon sky was dark with dust. We stopped at a ulous specimens of Ariocarpus retusus.
roadside vegetable stand and fought the wind and That night in Saltillo we met the bus to col-
grit until we found refuge in the shack of the ven- lect one of my former students, Lia Carrasco, who
dor, whose cantaloupes were marvelously sweet joined us for the rest of the trip, to be continued
and juicy. A particularly strong gust brought dust in Part 2: “Zacatecas, San Luís Potosí, Nuevo
into the shack as we were devouring our sample León, and Tamaulipas.” 
slices. The vendor smiled and observed, “These
violent dust storms are the thundershowers of
San Pedro.” References
We got our second flat on the sharp gravel road 1 Bye RA. 1979. Hallucinogenic plants of the Tarahumara.
back to the Laguna. That puncture was a small, J Ethnopharmacol 1: 23–48. 2 Lumholtz C. 1902. Unknown Mexico.
repairable hole in the tread, and we decided to Scribner’s Sons, New York. 3 Anderson EF. 2001. The Cactus Family.
Press, Portland, Oregon. 4 Habermann V. 1975. Two red
wait until the next day to fix it. An hour later, as Timber
flowering species of Lophophora. Cact Succ J (US) 47: 123–127.
sundown approached, we stopped to camp for the 5 Anderson EF. 1996. Peyote, the Divine Cactus. 2nd ed. University of
night and heard the hiss of our third flat tire. For- Arizona Press, Tucson. 6 Bohata J, Myšák V, Šnicer J. 2005. Genus
tunately, Robert had brought a small air pump Lophophora Coulter. Kaktusy (Special 2): 1–45. 7 Štarha R. 1997.
Appendix IV, Chemický rozbor rodu Lophophora, pp 85–90 in Grym
that ran off the DC of the truck’s electrical sys- R. Rod/Die Gattung Lophophora. Vydavateľstvo Roman Staník:
tem, and the next morning, with the help of one Bratislava. 8 Terry M. Personal observation.

186 Cactus and Succulent Journal


Stalking the wild Lophophora
Part 2 
Zacatecas, San Luis Potosí, Nuevo León, and
Tamaulipas
H
eading south from Saltillo, Fed-
eral Highway 54 leaves Coa-
huila and cuts across the east-
ern edge of the state of Zacate-
cas. Our first attempt to find a
lophophora after crossing the
border brought us to GPS coor-
dinates now in the middle of a
newly plowed field. The area was generally dis-
turbed, and we found no Lophophora william-
sii in the surrounding brush, although the hab-
itat was superficially similar to the Tamaulipan
thornscrub that makes up what members of the
Native American Church call the “Peyote Gar-
dens” of South Texas.
In the vicinity of San Tiburcio, Zacatecas,
we found a typical population of L.  william-
sii, replete with all the usual companion plants
of the Chihuahuan Desert, including candelilla
(Euphorbia antisyphilitica), lechuguilla (Agave
lechuguilla), leatherstem (Jatropha dioica),
and tasajillo (Opuntia leptocaulis). We parked
in a pulloff on the side of the highway and fol-
lowed an old ranch road leading back into the
desert scrub. Within 50 meters of walking we
found our first small, mature cluster of L.  wil-
liamsii right beside the road beneath a mes-
quite tree. It turned out that the road followed a
contour of particularly good limestone soil, and
the only plants we found for the first hour or so
222 Cactus and Succulent Journal
Martin Terry
were in that stratum. They were not abundant,
and only when we were about to leave the site
did Robert find a denser stand of plants on the
northern slope of the next low limestone hill to
the south.
San Luis Potosí
It seems as if everyone in the world who has
any interest in Lophophora gravitates to the
flats west and southwest of Real de Catorce in
San Luis Potosí. We likewise succumbed to this
attraction, partly to collect DNA samples from
a well-known population, and partly to assess
the impact of many years of “narcotourism”
and other commercial enterprises that depend
on the harvesting of peyote. On one side of the
road where we stopped to investigate there was a
newly plowed field. On the other side of the road
was what appeared to be an old agricultural field
with vestiges of plowed rows, now regrown with
creosote bush and little else. There was some
native brush in a strip running parallel to the
road, and there we found a few small specimens
of L. williamsii, but we were desperately trying
to find just ten plants to complete our DNA sam-
pling when a goatherd came walking along with
about forty goats. We chatted about the margin-
al state of the goat business and local attitudes
about peyote. He said that in spite of the suppos-
 L. williamsii is sparse in the alluvial flats below Real de Catorce, San Luis Potosí, reportedly due to chronic
over-harvesting concomitant to “narcotourism” and other commercial peyote harvesting enterprises,
despite large signs warning: “The extraction and illegal trafficking of peyote is a federal offense.”

edly strict enforcement of laws to punish out-
siders who might extract peyote from this area
(which is protected as Wirikuta, the sacred land
where the Huicholes come annually to gath-
er peyote) there were people who had hauled
out great quantities of peyote from local popu-
lations for sale in some unspecified distant mar-
ket. I told him that we were searching for pey-
ote for a scientific study, but that we were hav-
ing difficulty finding enough plants. He looked
down at the ground where we were talking and
pointed with his herding stick: “There’s one.”
And then there was another, and another, until
we had our ten tissue samples. The sound of the
goat bell disappeared into the deepening dusk as
we walked back to the truck. We found a small
restaurant in the nearby town and enjoyed some
local cuisine and a cold beer before heading for
Matehuala.
My pickup began running hot as we approached
the town of El Cedral (The Cedar Grove), so we
bought some antifreeze and stayed at a conve-
nient hotel near the Pemex station. I spent most
of the next day getting a new water pump locat-
ed, purchased, and installed in Matehuala while
Robert and Lia processed a fraction of each of
the tissue samples we had collected, grinding
the tissue up into a solution designed to pre-
serve DNA in the field until it can be extracted
K  continued on page 227

L. williamsii lays low at a site north of Doctor Arroyo, Nuevo León.

2008 Volume 80 Number 5 223

 the
Hitchhiker’s
Guide
T
to molecular systematics
he accompanying article
can be read as a picaresque
account of the quixotic
adventures of botanists zig-
zagging through northeast-
ern Mexico from one set of GPS coordinates to the
next, collecting DNA samples of various populations
of cacti in the genus Lophophora and simultaneously
taking stock of the conservation status of those
populations. But what is the point of collecting those
DNA samples? In Part 1 we described the process of
extracting DNA from cactus tissue, but then what
does one do with the extracted DNA?
The adjective “genetic” (from the noun “genesis”)
refers to origins. DNA can be considered the point
of origin of the processes that create and maintain
the structure and function of living organisms.
Traditional taxonomy (which concerns itself with
naming and classifying organisms according to their
similarities and differences) and systematics (which
concerns itself with the relationships among organ-
isms based on their evolutionary history) traditional-
ly made use of morphological characters (the visible
form of the plant) as the basis for evaluating relat-
edness among different organisms. In other words,
up until the 20th century, if we wanted to compare
different species and assess how closely related they
were, we would look at their anatomical structures as
the basis for comparison. (This was particularly con-
venient, because it also worked for fossils of long-
extinct species.)
Then we became more adept in organic chemis-
try and found that closely related species of plants
would produce identical or closely related chemicals
(such as alkaloids), and in the 1960s there was a
blossoming of plant systematics based on these phy-
tochemical characters.
Meanwhile, the picture became more complicated
as ethologists made systematists aware of behavioral
differences that could be used to compare related
species, while biochemistry was producing a genera-
tion of protein chemists who taught the systematists
to do electrophoresis (a technique which separates
molecules migrating through a gel by differences in
224 Cactus and Succulent Journal
their electrical charge, shape, and/or size) to separate
proteins such as enzymes, which could be used to
detect distinctions between related genera or species,
and sometimes even between subspecies/varieties.
All of those types of characters and techniques
focused upon various aspects of the phenotype of the
organism, which is the tangible or detectable expres-
sion of its underlying genotype. The genotype consists
of the DNA sequence (sequence of nucleotides in the
DNA) of a defined and specific part of the total genet-
ic makeup (the genome) of the organism. While the
structure of DNA had been elucidated by the middle
of the 20th century, it wasn’t until the last quarter of
the century that DNA sequencing techniques became
widely available. Once it was possible to analyze the
exact sequence of nucleotides in DNA—and thus
to know the genotype itself rather than its pheno-
typic manifestations—most of the older techniques
became obsolete, at least in the minds of the emerg-
ing army of molecular biologists. Why, they reasoned,
should one work with old, blunt instruments to obtain
data that would at best yield an indirect, partial, and
often ambiguous expression of the genotype, when
one could now analyze the genotype itself, directly,
totally and unambiguously, thereby cutting through
to the ultimate genetic truth?
This view of the primacy of DNA research in US
biological and medical science gained quick accep-
tance in circles that controlled the purse strings
of government funding of academic research. (As
Shakespeare might have put it, the DNA’s the thing
Wherein I’ll catch the funding from the King.) And
this in turn led to schisms in academic institutions,
where the “Haves” (the well funded molecular biolo-
gists) were enviously derided as “gene jockeys” by the
I
“Have-nots” (the organismal biologists) who contin-
ued to do biological research on a shoestring in the
traditional ways. (Incidentally, as an organismal biolo-
gist who revels in the mysteries of the whole organ-
ism and simultaneously appreciates the analytical
power that DNA research allows, I refuse to participate
in this still-smoldering war.)
n practical terms, the first ques-
tion to be answered is, what
sort of DNA locus is most likely
to be useful for this specific
problem? The answer is largely
determined by taxonomic hierarchy. For instance,
if one is sorting out relationships among families
within an order, it might be appropriate to use slowly
evolving segments of DNA such as genes that code
for functional proteins. The sites for viable mutations
(the stuff of which evolution is made) are restricted in
such a gene to nucleotides whose replacement does
not cause severe or lethal dysfunction of the protein
that is the product of the gene. That means that it
takes a relatively long time (evolution is measured in
geologic time) for enough viable mutations to occur
and become established in the gene pool, so that
significant sequence differences can be seen between
organisms representing different higher taxa.
If one is working at the species level, however,
DNA segments of protein-coding genes are not likely
to show sufficient change over the relatively short
time it takes for species to evolve from a common
ancestor. What is needed is rapidly mutating DNA.
Such DNA is selectively neutral, meaning that a muta-
tion in such a locus (segment of DNA) will have no
effect on the survival or reproductive success of the
organism. A neutral mutation has no effect on essen-
tial proteins, so the mutation has a reasonable chance
of being preserved in the gene pool.
What kinds of DNA are presumed to be selectively
neutral? (Here’s where things get complicated.)
One type is known as spacer DNA, found between
tandomly arranged genes. The function of spacer
DNA is to separate “busy” genes that are subject to
frequent transcription (reading). Spacer DNA is not
itself transcribed to RNA as genes are.
Another type of selectively neutral DNA is tran-
scribed. Coding segments of a gene (exons) are sepa-
rated from each other by segments of DNA called
introns, which are transcribed to RNA along with
the exons of a gene, after which the RNA segments
corresponding to the introns are edited out and
the transcribed gene fragments spliced together to
make a continuous, finished messenger RNA, which
is then translated into a protein. Introns may not
consist entirely of selectively neutral DNA, but major
portions of many introns are thought to be muta-
tionally neutral.
A third useful type of DNA locus is called a micro-
satellite. Such loci are likewise thought to be largely
selectively neutral. Microsatellites consist of simple
DNA-sequence repeats arranged in tandem; for
instance, GAGAGAGA. These constitute the fastest-
evolving type of DNA loci, because not only are they
subject to “normal” mutations that occur at the rate
of one mutation per 10,000 to 100,000 DNA base
pairs per generation, but microsatellites also have
their own accelerated type of mutation. When DNA
is being replicated, the enzyme responsible for mak-
ing a new DNA strand from the template of an old
DNA strand is called DNA polymerase. What is pos-
tulated to happen with microsatellites is that when
DNA polymerase is confronted with so many tandem
repeats (like GAGAGAGA…), the enzyme undergoes
“polymerase slippage,” meaning that it either “drops
a stitch” (that is, deletes one of the GA repeats in the
example) or inserts one too many “stitches” (that
is, inserts an extra GA repeat in the example). This
process often results in a microsatellite locus with
10 or 15 different alleles (different DNA sequences
2008 Volume 80 Number 5 225
within the locus), all of which would differ from each
other by the number of simple sequence repeats (the
number of tandem GA repeats in the example given).
Such a locus would be described as highly polymor-
phic and would likely be a sensitive genetic marker
for analyzing either population structure or phyloge-
netic structure at the species or subspecies/variety
level. (Are you still with me?)
So how do we generate usable data from our DNA
samples using microsatellites? First, we go through a
lengthy lab procedure to “capture” microsatellite loci
from the DNA of our target species and grow bacterial
clones containing the various microsatellites captured
(the more loci used, the more sensitive and accurate
the results will be). Then we sequence the microsatel-
lite loci and use the sequences to design primers. If
the primers work—that is, if they succeed in produc-
ing millions of copies of a given microsatellite in PCR
(polymerase chain reaction)—then we send them to
someone who puts fluorescent labels on the primers.
At that point, after testing the fluorescent-labeled
primers to confirm that they work, another PCR is run,
using the fluorescent primers and DNA from our field-
collected tissue samples. The products of that reaction
are then put into an expensive electrophoresis instru-
ment which detects the alleles of a given microsatel-
lite by the fluorescence provided by the fluorescent
primers. That process gives the exact length of each
allele to the nearest single DNA base pair, so that you
know how many different alleles you have detected
in the population sampled, as well as the frequency of
each allele in the population. It also tells you whether
each individual plant sampled is a heterozygote (hav-
ing different alleles on its two chromosomes that
contain the locus) or a homozygote (having the same
allele on both chromosomes). The latter information
in turn tells you about the breeding system of the
plant, with a high percentage of heterozygotes in the
population indicating a high degree of outcrossing
(individual plants fertilizing other individuals) and a
high percentage of homozygotes indicating a high
degree of inbreeding (self-fertilization being the
extreme of inbreeding).
OK, then, skipping the rest of the population
genetic analysis, how do we use the microsatellite
dataset to determine how the various geographically
defined groups of individuals sampled are related
to each other phylogenetically? The short answer
consists of one word: software. There are a number
of programs that analyze microsatellite datasets
and produce trees showing phylogenetic affinities
among the various operational taxonomic units that
we input. Would you like to know how many species
of Lophophora there are? So would we, but we don’t
have the dataset completed yet, much less the analy-
sis. Please stay tuned for the results of the analysis,
coming soon at a Haseltonia near you.
226 Cactus and Succulent Journal
I  L. koehresii near Tula, Tamaulipas. Plants here grow in the mud beneath large nurse shrubs, are almost all
solitary, and are relatively small compared to other species of Lophophora. Can you spot the African-native
succulent, Kalanchoe?
J Near Miquihuana, Tamaulipas, the peyote plants are nearly all caespitose (clump forming).

K  Wild Lophophora continued from page 223

in the lab. Matehuala was already hot at the end
of May, and by four o’clock in the afternoon we
were all quite ready to get on the road and stir
up a breeze, when finally the truck was declared
“listo.”
Nuevo León
We headed for the fair city of Doctor Arroyo, and
then north for a few miles, where we pulled into
a small village nestled against some low moun-
tains. A rain shower was just ebbing, and we had
about an hour of daylight left. It didn’t take long
to find the first L. williamsii, but we were pleas-
antly amazed by the abundance and diversity of
other cactus species. We gawked over all these
species that none of us had ever seen before and
took many pictures. Eventually one of the villag-
ers showed up to see what we were doing. When
he found out we were interested in cacti he gave
us a brief ethnobotanical tour of the common
local species of edible and medicinal plants. This
was delightful and informative, but used up pre-
cious daylight.
The GPS coordinates we had were mislead-
ing; following them, I ended up high on a moun-
tain overlooking the village, where I got a fabu-
lous view of the sun setting, but encountered no
Lophophora. In the end, most of the L. william-
sii plants we found were right along the road
that skirted the edge of the village. The last two
samples were collected in the dark. We had an
unmemorable supper out of cans from the back
of the truck, and slept soundly on the ground
until dawn arrived in a mist.
After attaining a caffeinated semblance of

2008 Volume 80 Number 5 227

 consciousness, we walked around the village
to say our goodbyes to our lecturer/host of the
previous evening, then fired up the truck and
struck off to the east.
Tamaulipas
Miquihuana is a town on the western edge of
Tamaulipas, at about the same latitude as Ciu-
dad Victoria on the other side of the Sierra Madre
Oriental. The population of L.  williamsii near
Miquihuana was one that Ted Anderson exam-
ined in his PhD thesis1. It is a unique population
that continues to puzzle and fascinate students of
Lophophora to this day. It appears that virtually
100% of the plants are caespitose, and the plants
begin to sprout lateral branches as seedlings2,
whereas the plants in other Lophophora popu-
lations generally start such branching only after
reaching maturity, if at all. In terms of stem mor-
phology and color, the Miquihuana plants bear a
striking resemblance to the plants of the “Peyo-
te Gardens” of South Texas, some 300 km to the
north. In terms of their breeding system, howev-
er, they seem to have a closer affinity to the pop-
ulations around El Huizache, about 120 km to the
south. The northern plants are known to be autog-
amous3 (self-fertile), whereas greenhouse obser-
vations4 suggest that the Miquihuana plants, like
those of El Huizache to the south, are obligate out-
crossers (self-sterile). So how could we possibly
neglect to include the Miquihuana population in
our DNA-based phylogenetic study of the genus?
Locating the plants in Miquihuana was another
matter. The only GPS coordinates we had led us to
a roadside location where we were unable to find a
single plant in an hour of careful searching, though
the habitat looked reasonable for Lophophora.
We put away the GPS data, and I fell back on my
imprecise memory of the plants’ location from a
visit some years before. After a chain of succes-
sive conversations with local people, we finally
found someone who knew where the peyote grew.
But then we had to get permission to collect our
samples, which involved talking with a local offi-
cial who, when I produced a blanket permiso that
had been skillfully crafted by Héctor Hernández
in anticipation of such situations, stared at it for
a moment, then handed it back to me and asked
me to read it to him. It only slowly dawned on me
that this man could not read. The only condition
he imposed on us was that we not divulge the loca-
tion of the population to anyone, as such publicity
could only result in problems, including increased
risk of decimation of the rather small population,
which the local people value greatly and harvest
sustainably for therapeutic use (particularly as a
228 Cactus and Succulent Journal
topical analgesic for sore muscles). We collected
our tissue samples from the relatively dense pop-
ulation of multi-stemmed plants growing among
small agaves and Larrea, said our thanks and
goodbyes, and departed.
From Miquihuana, the gravel road took us east-
ward, up and over an arm of the Sierra Madre Ori-
ental, through forests of arborescent yuccas in the
high canyons, and down in a southeasterly direc-
tion until we hit Highway 101. We turned south,
heading to the northernmost known population of
L. koehresii in the town of Tula. We got to the spot
after a hard thundershower, just as it was getting
dark, and slept in the truck parked by the side of
the road.
The next morning we awoke to a thorough-
ly soaked desert. The L.  koehresii were there,
all right—some of them within a few steps of the
road, covered with wet mud. Others nearby had
been washed to a pristine green by the rain. It was
the first time I had ever seen Lophophora grow-
ing in mud flats, but it would not be the last. One
of the hallmarks of L. koehresii is its ecological
disposition to spurn the (often hilly) limestone
habitat of L. williamsii in favor of low-lying mud
flats. In spite of the fact that there was consider-
able human foot traffic through the area, probably
associated with nearby agriculture, we saw no sign
of harvesting. This may be a direct result of the
fact that, like L. fricii and L. diffusa, L. koehresii
is lacking in pharmacologically active concentra-
tions of mescaline. These non-williamsii species
of Lophophora have gained the reputation among
Huicholes as “the peyote that makes you sleepy”5,
which is perfectly compatible with data indicating
that pellotine is the principal alkaloid in these spe-
cies6. Pellotine was marketed as a sleeping aid in
the early 20th century, before it was rendered eco-
nomically obsolete by the advent of barbiturates,
which proved much less expensive to manufac-
ture7. By the time we finished exploring the Tula
population and collecting the required tissue sam-
ples, we were all—like many of the Lophophora
koehresii—well covered with mud.
Continued in Part 3: “San Luis Potosí (again),
Querétaro, and Mexico City.” 
References
1 Anderson EF. 1961. A Taxonomic Revision of Ariocarpus, Lophophora,
Pelecyphora, and Obregonia (Family Cactaceae). PhD Thesis,
Claremont Graduate School, Claremont, California. 2 Koehres
G. Personal communication. 3 Rowley G. 1980. Pollination syndromes
and cactus taxonomy. The Cactus and Succulent Journal of Great
Britain 42: 95–98. 4 Koehres G. Personal communication. 5 Williams
B. Personal communication. 6 Starha R, Kuchyna J. 1996. Analysis
of Mexican Populations of Lophophora (Cactaceae). Acta Facultatis
Rerum Naturalium Universitas Ostraviensis 156: 67–70. 7 Perrine D.
2001. Visions of the Night: Western Medicine Meets Peyote 1887-
1899. The Heffter Review of Psychedelic Research 2: 6–52.

Stalking the wild Lophophora
PART 3 San Luis Potosí (central),
Querétaro, and Mexico City
W
e continued south on High-
way 101, leaving Tamau-
lipas and entering San
Luis Potosí just before we
hit Highway 80, on which
we turned east toward El
Huizache. The latter is a
village at the intersection
of Highways 57 and 80. It is also the landmark
for the population that Ted Anderson selected as
the source of his neotype specimen to represent
the species Lophophora williamsii. I had visited
this population in 2001, and it was immediate-
ly apparent, now six years later, that the popula-
tion had undergone some changes for the worse.
There was evidence that plants were being dug
up entire (including the roots, as opposed to the
sustainable practice of removing the “button,”
or aboveground portion of the stem, and leav-
ing the large subterranean portion of the stem
to resprout (see sidebar Where’s the goods) and
the average size and density of the plants had
visibly decreased compared to what I had seen
six years before, with the clumps of caespito-
se plants being similarly reduced in size. There
was also new agricultural activity in the middle
of the Lophophora habitat, where fields plowed
for marginal agriculture had replaced Chihua-
huan Desert. An undernourished burro brayed at
us—or perhaps it just brayed, at no one in par-
310
MARTIN TERRY
ticular—before retreating into the brush. It was
not an appealing environment to spend time in,
and as soon as we had collected our samples and
taken our photos, we left, heading further east-
ward on Highway 80.
We stopped after a short distance to check
a friend’s GPS record of what was reported
to be “L. williamsii.” And we did indeed find
Lophophora there, on both sides of the highway,
but it was L. koehresii, not L. williamsii. This
was another mud-flat population, and while the
plants were not exactly abundant, we were able
to find enough to meet our quota of tissue sam-
ples without difficulty. Here again, there was no
evidence that the L. koehresii had been harvest-
ed, despite the fact that it was a heavily traf-
ficked area with much human activity.
Robert noticed that just a short hike up from
the highway was a limestone ridge that looked
like typical habitat for L. williamsii, so we decid-
ed to check it out. Bingo! On the lower slopes, in
alluvial limestone soil, we found just a couple of
specimens of L. williamsii, but a few meters far-
ther up in a limestone outcrop we found classic
L. williamsii habitat and what had been a fair-
ly dense population of the species. Unfortunate-
ly we arrived a few weeks too late to see the pop-
ulation in its full glory. The landscape had been
devastated. Massive quantites of whole plants
had been dug up and removed. Seedlings and
CACTUS AND SUCCULENT JOURNAL
+ * Near El Huizache, San Luis Potosí, L. williamsii may be solitary or caespitose. This is the neotype locality
for L. williamsii. Its fame and easy accessibility, at the junction of two major federal highways, render it an
obvious target for commercial cactus harvesters, and we found a number of their familiar excavations where
entire plants had been dug up and removed. The plants here are expected to be genetically different from
the L. williamsii that occur in the US, because the Texas plants are apparently 100% self-fertile, whereas the
plants from El Huizache are reported to be obligate outcrossers (that is, a given plant can set viable seed only
if it is fertilized by another plant). That means the Texas plants are highly inbred with minimal genetic diver-
sity, while the El Huizache plants are expected to show far more genetic diversity within a given population.

We do not know the nature of the market the destroyers

were supplying with these plants. We do know, however,
that it was a mescaline market.
small juveniles had been dug up, discarded, and
left to die. The poachers had been careless and
dropped a few of the uprooted adult plants along
the trail on the way out. We replanted as many
of the uprooted plants as we could find and col-
lected the tissue samples we needed, but this
scene of destruction was sickening, and we left
without delay. We do not know the nature of
the market the destroyers were supplying with
these plants. We do know, however, that it was
a mescaline market. We know this because the
poachers walked right through the population of
L. koehresii—without touching them—on their
way to the ridge where they found and thorough-
ly pillaged the population of L. williamsii. They
knew exactly what they were looking for, and it
was not pellotine (see sidebar A cactus at the
pharmacy).
The positive aspect of the situation was the
clear finding of two populations, one of L. koeh-
resii and one of L. williamsii, separated by no
more than 500 meters, but completely distin-
guishable morphologically, ecologically, and phy-
tochemically (the difference in alkaloid profiles
is reported in the analytical work of Štarha6 and
confirmed commercially by the poachers’ accu-
WHERE’S THE GOODS?
The concentration of mescaline (the hallucino-
genic chemical) in peyote varies among popula-
tions. It also varies with time of year, plant age,
and tissues sampled. Some investigators have
found considerably less mescaline in the roots
of plants from some populations, for example,
but others have found mescaline concentrations
in the root and stem to be similar. I suspect that
this disparity is attributable not so much to
variation in the methods of analytical chemistry,
but rather to widespread misunderstanding of
how much of the subterranean portion of the
plant is actually stem, how little is actually root,
and where the thin transition region between
these tissues is. Specific tissue concentrations
of alkaloids constitute another one of those
questions that needs careful and comprehen-
sive evaluation, taking into account geography
and time of year, as well as tissue type. And the
answer to this question is not merely academic,
as harvesting the tuberous subterranean portion
of the plants hampers new vegetative growth
and therefore jeopardizes the population.

2008 VOLUME 80 NUMBER 6 311

 rate, selective behavior in the field). I
predict that our DNA data will show an
equally clear distinction between these
two seemingly sympatric, but in fact eco-
logically allopatric, species.
We spent the night in Ciudad del Maíz,
where cooked food, a shower, and bed
were welcome amenities. Next morn-
ing, refreshed, we hit the highway head-
ing south on a good gravel road, stop-
ping to sample more L. koehresii popula-
tions near Las Tablas and San Francisco.
The Las Tablas population is one that has
been known for many years—but not as
a population of L. koehresii. Ted Ander-
son included Las Tablas among the pop-
ulations he sampled for his PhD thesis,
but he did not recognize that the speci-
mens there were anything other than an
unusual form of L. williamsii7. And I con-
fess that when I first saw these plants in
2001, my reaction was similar: They’re
just L. williamsii plants with a differ-
ent flower color, living in lowland alluvial
soil instead of upland calcareous soil. But
these are not taxonomically significant
differences in such a highly polymorphic
species as L. williamsii...or so I thought.
Once one’s eye has learned to recognize

* L. koehresii east of El Huizache, growing mostly in mud near the highway. The large specimen was found
growing beneath a large Opuntia leptocaulis. - Scene of destruction east of El Huizache. L. williamsii was
uprooted en masse by commercial cactus harvesters, who dropped these plants on the trail between the
highway and the peyote population. Interestingly, they walked right through a population of L. koehresii,
confirming that species as a non-drug plant. Please note: peyote can be sustainably harvested. It is
not necessary to remove the root of the plant to harvest the “button.” If left behind, the subterranean
portion of the stem will often sprout new stems that eventually grow to become harvestable crowns.2

312 CACTUS AND SUCCULENT JOURNAL


* Lophophora koehresii near Las Tablas, San Luis
Potosí. Ted Anderson and I both mistook these
plants for L. williamsii on first sight, but we would
not now make the same mistake. Morphological
differences include the relatively small size of the
mature adult stems of L. koehresii and the relatively
large size of its flowers, which may have tepals
twice the length of those on L. williamsii. The fruits
of L. koehresii are essentially spherical, in contrast
to the cylindrical fruits of L. williamsii. The seeds of
L. koehresii are larger than those of L. williamsii, and
the two species have markedly different topologies
of their seed surfaces as viewed under scanning
electron microscopy. Other differences include
stem color: generally a dark green in L. koehresii;
blue green to gray green to butternut-brown green
in L. williamsii. Large specimens of L. koehresii
may develop a conspicuous “double-chin-like”
horizontal fold of tissue at the base of the crown,
similar to those seen in large specimens of
L. diffusa that cannot support the weight of their
own upper stem (especially during drought), but
such a fold of tissue is never seen in L. williamsii,
which has much more rigidly constructed ribs.
2008 VOLUME 80 NUMBER 6
A CACTUS AT THE PHARMACY
Pellotine, which was marketed as a sedative/
hypnotic about a century ago by Boehringer &
Sohn in Germany, was obtained by isolating the
alkaloid from an extract of Lophophora diffusa
(at that time confusingly known as Anhalonium
williamsii. What we now know as L. williamsii was
then known as Anhalonium lewinii). When the
Bayer company discovered how to synthesize
barbiturates (starting in 1911), the drugs proved
so cheap to manufacture—and so effective—that
the extraction of pellotine from a field-collected
Mexican cactus of unpredictable availability was
no longer commercially competitive, so pellotine
disappeared from the pharmaceutical market.
Eventually Späth4 synthesized pellotine, and
Brossi and others5 later discovered an improved
procedure for synthesizing it, but the drug was
never brought back into commercial use. It is
interesting that pellotine is the second most
abundant alkaloid (after mescaline) in L. william-
sii, but it is by far the most abundant alkaloid in
the other species of Lophophora (accounting for
70–90% of total alkaloid content), and in those
species mescaline is present in only trace concen-
trations—not high enough to have pharmaco-
logical effects from ingestion of the cactus.
the differences between L. koehresii and L. wil-
liamsii, the two taxa remain indelibly demarcat-
ed and impossible to put back into the same con-
ceptual taxonomic container (see sidebar It’s in
the ribs). Particularly conspicuous is the diffu-
sa-like rib morphology of L. koehresii, charac-
terized by shallow, sinuous sulci separating adja-
cent ribs. Successive tubercles within each rib
are connected with each other in such a way that
repeated tandem hourglass shapes radiate from
the center of the crown, and the tubercles of any
given rib are in conspicuous alternation with the
tubercles of the two adjacent ribs. Such differ-
ences cannot, however, be discerned from des-
iccated herbarium specimens, even by the most
experienced cactus experts. One has to see the
plants alive in the well-defined ecological niches
of their natural habitats to appreciate how differ-
ent they really are.
Querétaro
From the southernmost population of L. koeh-
resii near Río Verde in San Luis Potosí, we
faced a long afternoon of driving—south down
Highway 89 to Jalpan, and then south toward
Vizarrón on Highway 120—to reach the land of
Lophophora diffusa in the state of Querétaro.
We went to the first roadside population indi-
313
 * L. diffusa in the northern portion of its range. Note the shallow, “diffuse,” sinuous grooves between the ribs,
especially toward the base of the crown.
cated by our GPS coordinates and immediately
found the plants just fifty meters from the high-
way. L. diffusa has an affinity for gravelly alluvial
soils of coarse sand, in or near major creek beds
(which were all dry when we were there in May).
It seemed surprisingly easy to collect our tissue
samples from what I had anticipated to be a dif-
ficult species to find. Of course it helps if you
know where to start looking.
314
We spent the night at a hotel in Vizarrón with-
in walking distance of an obsolete GPS location
for L. diffusa on the eastern edge of the town.
The location was now an urban vacant lot filled
with trash and a few opuntias, but no lophopho-
ras. We talked to a few of the townspeople about
peyote in the area, and none of them seemed to
know of anywhere in the immediate vicinity of
the town where the cactus could still be found.
CACTUS AND SUCCULENT JOURNAL
 GEOGRAPHIC DISTRIBUTION OF LOPHOPHORA

El Paso Lophophora williamsii

U S A Lophophora fricii
Lophophora alberto-vojtechii
? ? Lophophora koehresii
? Lophophora diffusa
San Antonio
?
? ?
c o
x i
e
M

f
o
?
M

l f
G u
E

?
P

0 100 200 km
a

I 0 50 100 mi
c
i

C
f

20oN
i
c

O O
c Mexico City
e
a
n

110oW 105o

This map is the most accurate (based on a rigorous requirement for documentation of localities) and phylo-
genetically complete distribution map for the genus Lophophora now available. It is based on documented
voucher specimens (brown dots) from the UNAM database supplied by Héctor Hernández, Billie Turner’s
Atlas of the Vascular Plants of Texas, personal communications from Gerhard Koehres, Jaroslav Bohata, and
Jürgen Menzel, my own field observations, and herbarium specimens I have personally examined. Older
maps tend to portray the range of L. williamsii as being more extensive, particularly with regard to the
placement of its northwestern boundaries. Such marginal regions are indicated by question marks in the
present treatment due to the lack of voucher specimens. All these areas merit further exploration but are

So the next day we went up the highway a few
kilometers to a “fresh” location indicated by a
friend’s GPS coordinates for a population that
had been visited recently and confirmed to exist.
This population was also on both banks of a large
dry creek, but most of the plants were obvious-
ly on private land. The owner’s dog barked at us
from a distance until the owner came out to see
what was going on. I walked down a broad, open
arroyo to talk with him. He did not tell us to
remove ourselves from his ranch, nor did he say
that he preferred that we not take tissue samples
from his peyote plants. But he did want to see
the written permiso from the Mexican authorities
authorizing our research activities. So I told him
I’d go back to the truck and get the papers, but
when I was about 100 meters away, he shouted
2008 VOLUME 80 NUMBER 6
PEYOTE AND THE LAW
Drugs are placed in the DEA’s Schedule I because
they are deemed to have “high potential for
abuse… no currently accepted medical use… [and]
lack of accepted safety data for use… under medi-
cal supervision.”8 Although such scheduling may
seem unjustified with regard to what is actually
known of its effects, unlicensed possession of any
part of L. williamsii—including its seeds—carries
non-trivial criminal penalties in the US. Our British
friends, and many other cactus growers around
the world, are welcome to grow peyote, and it is
frequently found on seed lists of our sister societies
and foreign nurseries. For reasons which are not
entirely obvious, peyote has now been banned in
France, Italy, the Czech Republic, Russia, Poland,
and most recently Australia, much to the conster-
nation of cactus enthusiasts in those countries.
315
 * L. diffusa—a population in the central portion of its range with large, all-white flowers similar to those
seen on L. koehresii. - We stayed at the Motel Santa Rosalía in Camargo, Chihuahua, en route from Mexico
City to the Texas border, only because my student’s full first name happens to be Rozalía (Lía for short,
though it is unclear whether she would identify with the title ‘Santa’). The limestone mountains visible in the
background are suspected (not confirmed) to constitute habitat for the elusive L. willliamsii of Chihuahua.

of them were in full bloom (even in the apparent

absence of recent rain), their large white flowers
contrasting with the distinctive yellowish green
hue of the crowns of the plants.

and waved “never mind”—or at least I interpret-
ed his wave and shout in that sense. In any case,
we finished gathering our samples without delay,
to avoid any further complications. The plants
at this location tended to be caespitose, form-
ing clumps up to 30 cm in diameter, and many
Mexico City
Having sampled all the populations we were
going to include in our DNA study, the next
phase of the trip was to proceed to the Nation-
al Autonomous University of Mexico (UNAM)
in Mexico City and extract DNA from the tis-
sue samples in the laboratory facilities provided
by my friend and colleague, Héctor Hernández.
That little maneuver was designed to eliminate
the non-trivial problem of transporting peyote
tissue (a Schedule I controlled substance—see
sidebar Peyote and The Law) from Mexico into
the US, which has not legally been done in the
last 35 years, and which would require permisos
involving new and untested regulatory pathways
on the Mexican side. The solution to the prob-
lem was to separate the DNA from the psycho-
active alkaloids (particularly mescaline), and to
wash the latter down the drain in the lab in Mex-
ico City, so that we would be transporting only
pure DNA back to the US. The laboratory phase
of the trip in Mexico City was a mixture of hard
work and great company, as we found a warm
reception from Héctor Hernández and family and
from his colleagues at the molecular biology lab
of the Instituto de Biología at UNAM, where in
about ten days we were able to develop and apply
a viable way of making commercial DNA extrac-
tion kits work on our samples without the use of
liquid nitrogen. But that is a technical story for

316 CACTUS AND SUCCULENT JOURNAL


cactus-DNA nerds—the sort of stuff you’ll read in
Haseltonia when the DNA study is completed.
Border Epilogue
The last vignette of our experience on this cactus
trip involves an incident that occurred upon our
reentry into the US at the Presidio Port of Entry
in Far West Texas. Because I had among my DNA
samples a few grams of non-mescaline-contain-
ing tissue from the three Lophophora species not
included in the legal definition of peyote, I had
consulted on the phone with my DEA contact in
the El Paso office to get his guidance on how to
avoid problems at the border. He kindly put me
in touch with the Director of the Port of Entry,
who in turn alerted the USDA person on his staff,
who would be expecting me to pass through the
Port of Entry on a specified afternoon to help
with the inspection of my cactus tissue and to
avoid unpleasant encounters “of the third kind.”
Unfortunately, the USDA person so-informed
was ill that day, and the only other person who
could carry out the “Aggie” inspection function
was not coming in until four o’clock in the after-
noon. So what actually happened was that I duly
declared the cactus tissue and the cactus DNA
as required, and the random inspector I talked
to thought this was the most exciting thing that
had happened in Presidio in weeks. Soon I had
a half dozen young men in dark blue uniforms
standing in a semicircle at the back of my pick-
up, staring at the samples and firing questions at
me. When their routine questions failed to elicit
anything more than a boring response from me,
one of them puffed up his chest and asked, “Do
you have FDA authorization to be doing research
with this DNA?” To which I responded, “Well, as
a matter of fact, I do, but FDA authorization is
not required to bring plant DNA into this coun-
try.” And he responded, most wonderfully, “It
is if you want to bring it in through this Port
of Entry.” About that time a supervisor showed
up and broke up their sport, and the dialogue
returned to a more rational plane. The senior
officials were cordial and as helpful as they could
be, given the uncomfortable uncertainties that
evidently, in their minds, surrounded plant DNA.
In the end, we were able to clear customs in a
mere two and a half hours, including the time it
took to radiograph my entire old truck. “Just as
a formality.”
REFERENCES
1 Trout K. 1999. Sacred Cacti: Botany, Chemistry, Cultivation &
Utilization. Second Edition. Better Days Publishing, Austin. 2 Terry
2008 VOLUME 80 NUMBER 6
IT’S IN THE RIBS
There are conspicuous differences in crown mor-
phology between L. williamsii (left) and L. koehresii
(right). Ribs of L. williamsii are raised and clearly
divided by straight, deep sulci (grooves). Each rib
resembles a pie slice. Tubercles, denoted by areoles
bearing tufts of hairs, may be individually raised,
forming a radial row of elevations along the already
raised rib. Ribs of L. koehresii are flat and subtly
divided by sinuous, shallow sulci. They show an
alternation of widening at each tubercle and nar-
rowing between tubercles, forming a radial series
of connected figure-eights. Tubercles, denoted by
areoles bearing tufts of comparatively few, shorter
hairs, are relatively flat. The tubercles of adjacent
ribs tend to be offset, so that a wide tubercle in
one rib is situated between the narrow intertuber-
cular portions of the two adjacent ribs.
M and Mauseth JD. 2006. Root-shoot anatomy and post-harvest
vegetative clonal development in Lophophora williamsii (Cactaceae:
Cacteae): Implications for conservation. Sida, Contributions to Botany
22: 565–592. 3 Perrine D. 2001. Visions of the night: Western medicine
meets peyote, 1887–1899. The Heffter Review of Psychedelic Research 2:
6–52. 4 Späth E. 1922. Über die Anhaloniumalkaloide. V. Die Synthese
des Anhalonidins und des Pellotins. Monatshefte fuer Chemie 43:
477–484. 5 Brossi A, Schenker F, Leimgruber W. 1964. Synthesen in
der Isochinolinreihe. Neue Synthesen der Cactusalkaloide Anhalamin,
Anhalidin, rac. Anhalonidin und rac. Pellotin. Helv Chim Acta 47: 2089–
2098. 6 Štarha R, Kuchiňa J. 1996. Analysis of Mexican Populations
of Lophophora (Cactaceae). Universitas Ostraviensis Acta Facultatis
Rerum Naturalium, Physica-Chemia 156: 67–70. 7 Anderson EF.
1961. A Taxonomic Revision of Ariocarpus, Lophophora, Pelecyphora,
and Obregonia. PhD Thesis, Claremont College. 8 Code of Federal
Regulations, Title 21, Section 812.
ACKNOWLEDGMENTS
This research project, including the field work described in this
article, is the product of many contributors, most notably Héctor
Hernández, who both encouraged and made administratively pos-
sible our fieldwork and lab work in Mexico, my students Robert
Hibbitts and Lia Carrasco, and my colleagues Keeper Trout and
Bennie Williams at the Cactus Conservation Institute. Additional
photos of the Mexican Lophophora species and their habitats can be
viewed at www.cactusconservation.org/CCI/cslm00.html.
Heartfelt thanks are also due to Geoff Bailey, Jaroslav Bohata, Kendall
Craig, Gerhard Koehres, Guadalupe Martínez, Jürgen Menzel, John
Miller, Woody Minnich, Tim Mullet, and Vojtěch Myšák for help in
mapping, locating, and/or collecting tissue samples from populations
of Lophophora. Unpayable IOUs of gratitude are due to the late Ted
Anderson and Dick Schultes for inspiration and laying the ground-
work for those of us who follow. Funding for the project was provid-
ed in the form of a grant from the Research Committee of the Cactus
and Succulent Society of America and a Research Enhancement
grant from Sul Ross State University.
317
 BOOK REVIEW BRIAN KEMBLE
The genus Echeveria by John Pilbeam. 2008. The which each species is
British Cactus & Succulent Society. Hard cover, 333 assigned. Each taxon’s
pp. ISBN 0 902099 80 9. $94.95. chromosome number
(where known) is pro-
vided. Species distribu-

E
ric Walther’s monograph Echeveria was pub-
lished in 1972, well over a decade after his
death. In the years since it has remained the
most thorough work on the genus. However, there has
long been a need for an update of Walther’s efforts,
for several reasons. First, there have been many new
species described since his death in 1959, and a
number of the species he recognized have since been
lumped with others. In addition, the majority of the
pictures in his book are black-and-white, and this is
less than ideal for a genus with such an astonishing
array of foliar color. To compensate for the paucity
of color illustrations, Walther carefully described the
subtle hues of his subject matter. Unfortunately, he
used a color chart that is not included in the book,
and whose tantalizing names—vetiver-green, rosa-
lane-purple, ochraceous-salmon—leave us to wonder
exactly what colors are meant.
In light of these shortcomings, John Pilbeam’s
new book is a welcome addition to the literature on
the genus. For the first time we have color photos
of all the species (save a few orphans not in cultiva-
tion for which photos could not be located), which
Pilbeam and his contributors went to great lengths
to assemble. While the result of all this effort consti-
tutes a somewhat uneven set of photographs—some
old and some new, some in habitat and some in cul-
tivation, and with some flat-bed scans thrown in for
good measure, the results serve to ably illustrate
more species than many thought possible when the
project began*.
The genus Echeveria is up-to-date taxonomical-
ly, and for each species there is a thoughtful com-
mentary and description, notations on distinguish-
ing features, a recounting of its area of occurrence,
and some words on cultivation. There are lists cover-
ing publication dates for each species, taxa published
since Walther’s book, and a run-down of the series to
tions are listed by state
(for those in Mexico) or
by country (for those
elsewhere), and a series
of maps for each series
in the genus depicts the
states or countries with-
in which each species
can be found—rather
less specific than might
be desired, but proba-
bly the best you can do
with a genus of plants typically found in small, wide-
ly scattered, and remote populations.
Myron Kimnach notes that many of the photos in
the book that were taken by John Trager were not
from the Huntington, but rather Myron’s personal
collection. And although he provided locality data for
most of these, this information, or at least the near-
est town, does not appear in the book’s figure cap-
tions, which is regrettable, as the data would have
lent the illustrations more value.
Interestingly, Pilbeam assigns cultivar names to
as-yet-unnamed wild populations (undescribed spe-
cies or varieties) in a section at the end of the book.
While probably acceptable, this is also rather uncon-
ventional.
All this adds up to the most complete work avail-
able on Echeveria, essential for students of the genus
and highly informative for the botanist and hobby-
ist alike. In addition to being well-researched, it is
a beautiful book which will doubtless serve as a key
reference on this group for many years to come. Per-
haps its least appealing aspect is the price, which at
$95 may strike some as steep; no doubt the dollar’s
current weakness has not been helpful here. Howev-
er, this is a book worth stretching the budget for. I
recommend it highly.

* One small correction: two photographs of Echeveria schaffneri are incorrectly credited to me; they were taken by Chad Davis,
horticulturist at the Desert Botanical Garden in Phoenix.

LETTER

“ Due to worldwide downward financial trends and tumbling markets, even avant-garde artist Cristo has felt the
crunch—scaling down his latest project to daub red paint on small cacti in a remote area of Mexico.”
It’s been about 15 years since the home of Ariocarpus bravoanus (CSJ 80–5, pp 220–221) has been invaded and sacked.
Cactus collectors may justify their desire to possess these plants because of their great love. But no one would believe it if
they heard that the plants stripped from their homes in the wild were going to be returned. That’s not going to happen.
Not one. So what can be done now?
In cultivation ariocarpus aren’t really that slow. We should be seeing flowering sized seedlings on sale tables by now. If
you have any of these treasures in your collection, search around between the tubercles and extract the seeds likely to be
hiding there. Scatter the seeds around the base of the plant, or give them to someone who grows cacti from seed. If you
have more than one A. bravoanus, exchange pollen between them when they flower in the fall, and scatter the seeds in
with the adults. Or give them away the following spring. If you have only one, find someone you can exchange pollen with
(locally or by mail) to produce some seeds. This is an important project for the true lover of cacti.
And always use good sense when purchasing wild plants. Mexican cacti are probably not going to be accompanied by
a salvage permit. —Fred Gaumer

318 CACTUS AND SUCCULENT JOURNAL