Terry M 2008 Stalking the Wild Lop Hop Hora

2008 volume 80 number 4 181

Stalking the wild Lophophora PART 1 Chihuahua and Coahuila

MARTIN TERRy

Ay, Chihuahua!

Sul 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.

182 CaCtus and suCCulent Journal

east on federal Highway 45 (a well-maintained toll road equivalent to an interstate highway in the US) toward the town of Julimes, where we had GPS coordinates for the one and only Chihuahuan Lophophora locality in the entire UNAM herbari-um database. From Julimes we veered west toward the Sierra de la Amargosa, a range noted a centu-ry ago by Lumholtz as an area to which the Tarahumara had traditionally traveled from their homelands to harvest the hikuli (L. williamsii) that they used for medicinal and ceremonial purposes2. We drove fast down arroyo-cut dirt roads as the sun set, tak-ing successive forks in the 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.

The objective of our trip was to collect tis-sue samples from 15–20 representative popula-tions covering most of the Mexican portion of the geographic range of the genus Lophophora, including populations of all four (now five!) of the Lophophora species that are generally recognized as valid by Continental European systematists,

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.

This Lophophora fricii at the Laguna site has 21 ribs, which is common here and in L. diffusa in Querétaro. Most globular cacti have stable configurations of ribs at Fibonacci numbers (1, 1, 2, 3, 5, 8, 13, 21…). For instance, Astrophytum asterias and Echinocactus horizonthalonius normally have eight ribs all their lives. Lophophora williamsii starts life with five ribs, transitions to eight by the time it reaches 5–6 cm in diameter, then switches to 13 ribs as a large adult (usually > 8 cm in diameter). L. diffusa and L. fricii follow the same pattern, but as large old plants they frequently jump up to the next Fibonacci number in the series.


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.

CoahuilaAdmitting 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.

the magnificent Motel La Vaquita (the Little Cow Motel), whose sign portrayed a small happy black and white cow. After the manag-er ascertained that we wanted the room for the whole night, he gave us the key in exchange for 140 pesos (about $12.50). The room boasted a window air-condition-er (which meant you opened the window if you wanted relatively cool night air to come in from out-side), a most welcome shower, and a television, on which we watched the news until we fell asleep.

At dawn the next day we drove to Viesca, a small city at the end of the paved highway heading south-east out of Torreón. Viesca is home to known populations of Lophopho-ra fricii4, and because this species is the subject of taxonomic contro-versy5, I considered it important to obtain tissue samples for DNA anal-ysis from one or more of the popu-lations in the town’s vicinity. In fact there are populations of very un-williamsii-looking lophophoras at several spots around the perimeter of a geologic feature known as the Laguna de Viesca6, a dry depres-sion that can become an ephemer-al shallow lagoon during the rainy season. We collected tissue samples from one spot in the mountains on

TISSUE SAMPLES — What are they all about?DNA, the genetic material found in all living organisms, carries all the informa-tion that makes the organism what it is and enables it to function and repro-duce. Because the sequences of bases in DNA inevitably change over time by such processes as mutation and natural selection, different sequences often develop in populations inhabiting different geographic areas. We can therefore get an idea of the degrees of relatedness among individuals from different populations by examining and analyzing their DNA sequences.

The first step in such a study is to collect tissue samples from which to extract the DNA. In humans this is relatively simple, as DNA can be easily extracted from cells that can be painlessly scraped from inside the mouth. In many plants, leaf tissue is suitable for DNA extraction. But cacti (wouldn’t you know?) are more difficult. It’s not that there is any shortage of DNA in cactus tissues. Au contraire, there is abundant DNA in the stem, particularly in those subdermal green cells where DNA is busy choreographing photosynthesis. The problem is that cacti also produce large quantities of mucopolysaccharides (yes, that’s the same muco as in mucus), which give the inner stems of cacti their sliminess. Molecular biologists dread these long, branched chains of sugars, because they behave like a net and entangle the DNA in the test tube. DNA is therefore rather difficult to isolate from the primordial cactus goo.

One way of minimizing the problems associated with the cactus-stem polysaccharides is to use liquid nitrogen to disrupt the tissue in the lab. But that 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 field. A third recourse is to collect the epidermis of the stem. The epidermis of Lophophora is exquisitely thin, so that in a sample the size of a thumbnail one may get only a milligram or two of dried epidermis. But we have found that to be enough to yield sufficient DNA for analysis.

Our strategy was to collect tissue samples from ten plants per population, with the constraint that we would not sample any plants located less than 10 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-eny. Having selected a plant for sampling, the person doing the sampling would cut (with a clean knife blade) a piece of tissue about the size of a thumbnail and weighing 100–200 milligrams from the crown (the above-ground part of the stem) of the plant. The tissue sample was then placed in a snack-size zip-lock plastic bag labeled with the code for the location and the specimen number. The samples were carried in my backpack until we returned to the truck, where-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 cactus from which a sample was taken, and notes were made of the plant’s physical measurements and habitat description, including companion plants.

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.



the southwest side of the Laguna and from anoth-er in the silty mesquite-creosote-agave flats on its northeastern perimeter.

These plants are notably different from typ-ical specimens of L. williamsii. In stem mor-phology, in particular the flattened tubercles, 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 growing from crev-ices in a hard type of limestone unfa-miliar to me, and those from the north-east side of the Lagu-na were growing in loose, silty, alluvial soil with no apparent 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

sampled are also strikingly different from each other. One is a rock-loving, montane population; the other occurs in loose alluvial soils of lowland flats. In the montane population the plants are predominantly solitary (single-stemmed); in the bed of the laguna, the mature plants are largely

Lophophora williamsii flowering on the south slope of a limestone mountain near Cuatro Ciénegas, Coahuila. The color and morphol-ogy of these plants, as well as the limestone mountainside they inhabit, are virtually identical to plants and habitats in Trans-Pecos Texas.L On the northern slope of a limestone mountain north of Saltillo, Lophophora williamsii is found under nurse shrubs, cryptic and partly covered with fallen leaves from the nurse shrubs. There were mature peyote plants here, but they were neither large nor abundant, suggesting that this population had been harvested in the not-too-distant past. That is hardly surprising, given the easy access provided by the major highway connecting Saltillo and Monterrey.


caespitose, forming clumps approaching a meter in length. Although we were unable to find flow-ering plants under the parched conditions we encountered in late May, further differences may yet be revealed: it will be interesting to see if there are substantial genetic differenc-es between these two pop-ulations, separated by only about 30 km but marked-ly distinct in their chemis-try, morphology, and habit, as well as their habitat.

Nothing beats the exhil-aration that comes with new heights of

accomplishment, such as when we managed three flat tires in one day. The first one was occasioned by yours truly looking at the scenery of a small hill arising unexpectedly out of the flats of the Laguna de Viesca, instead of looking at the road. A little stump of a long-dead mesquite tree on the side of the dirt road was sufficient to stab a gaping sidewall puncture in the right-front tire. That left us with a shoddy spare tire on the road and a forever-use-less flat in the bed of the truck. Not thrilled with the prospect of a second flat and no functional spare in this remote corner of Coahuila, we decid-ed to head for the big city of San Pedro to buy a new tire. We ended up buying two, but attempts to locate a used wheel for my old truck, so as to have two mounted spare tires rather than one, met with failure.

As we started back to the Laguna de Viesca an impressive storm arose—no trace of rain, but the afternoon sky was dark with dust. We stopped at a roadside vegetable stand and fought the wind and grit until we found refuge in the shack of the ven-dor, whose cantaloupes were marvelously sweet and juicy. A particularly strong gust brought dust into the shack as we were devouring our sample slices. The vendor smiled and observed, “These violent dust storms are the thundershowers of San Pedro.”

We got our second flat on the sharp gravel road back to the Laguna. That puncture was a small, repairable hole in the tread, and we decided to wait until the next day to fix it. An hour later, as sundown approached, we stopped to camp for the night and heard the hiss of our third flat tire. For-tunately, Robert had brought a small air pump that ran off the DC of the truck’s electrical sys-tem, and the next morning, with the help of one

of those pressurized products that squirts white goo in a tire to plug a leak, we were able to reach a ranch house where they had a bicycle pump that filled the wounded tire with enough air to get us

back to the nearest vulcanizador.

Our next stop was Cua-tro Ciénegas, a lovely oasis in central Coa-

huila, where it was relative-ly easy to locate a healthy population of L. william-

sii from the list of localities in the UNAM database. The

plants were growing in crevices and on natural terraces along the slopes of limestone hills. Mor-phologically and ecologically they were indistinguishable from plants that occur in west Texas. This is not sur-prising when you consider

that Cuatro Ciénegas is only 250 km from the southern tip of

the Big Bend.Our next stop was on the northern

outskirts of Saltillo, on Highway 40 in the direc-tion of Monterrey. L. williamsii was not abundant at this site, but in the course of an hour and a half we were able to collect stem tissue samples from ten individuals separated from each other by a distance of at least 10 meters. Most of the plants were found growing under nurse shrubs and were partially obscured by leaf litter. In the more northern regions of its range L. williamsii shows an absolute preference for the south slopes of hills and ridges8. At the lower latitude around Saltillo, in contrast, the plants occurred on both northern and southern slopes, interspersed with some fab-ulous specimens of Ariocarpus retusus.

That night in Saltillo we met the bus to col-lect one of my former students, Lia Carrasco, who joined us for the rest of the trip, to be continued in Part 2: “Zacatecas, San Luís Potosí, Nuevo León, and Tamaulipas.”

RefeRences

1 Bye RA. 1979. Hallucinogenic plants of the Tarahumara. J Ethnopharmacol 1: 23–48. 2 Lumholtz C. 1902. Unknown Mexico. Scribner’s Sons, New York. 3 Anderson EF. 2001. The Cactus Family. Timber Press, Portland, Oregon. 4 Habermann V. 1975. Two red flowering species of Lophophora. Cact Succ J (US) 47: 123–127. 5 Anderson EF. 1996. Peyote, the Divine Cactus. 2nd ed. University of Arizona Press, Tucson. 6 Bohata J, Myšák V, Šnicer J. 2005. Genus Lophophora Coulter. Kaktusy (Special 2): 1–45. 7 Štarha R. 1997. Appendix IV, Chemický rozbor rodu Lophophora, pp 85–90 in Grym R. Rod/Die Gattung Lophophora. Vydavateľstvo Roman Staník: Bratislava. 8 Terry M. Personal observation.

pellotine

Mescaline


Stalking the wild Lophophora PART 2 Zacatecas, San Luis Potosí, Nuevo León, and Tamaulipas

MARTIn TeRRy

Heading 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

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 PotosíIt 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-


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

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.”

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

K continued on page 227


T 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

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 “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.)I 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

the hitchhiker’s guideto molecular systematics


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

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.



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 LeónWe 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

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


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.

TamaulipasMiquihuana 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

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.

310 CACTUS AND SUCCULENT JOURNAL

MARTIN TERRY

We 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-

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

Stalking the wild LophophoraPART 3 San Luis Potosí (central),

Querétaro, and Mexico City

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

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-

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.


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


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.

QuerétaroFrom 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-

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.

A CACTUS AT THE PHARMACYPellotine, 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.


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.

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.

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.


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

PEYOTE AND THE LAWDrugs 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.

GEOGRAPHIC DISTRIBUTION OF LOPHOPHORA

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

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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

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.

Mexico CityHaving 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

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.


cactus-DNA nerds—the sort of stuff you’ll read in

Haseltonia when the DNA study is completed.

Border EpilogueThe 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

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.

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.

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


BOOK REVIEW BRIAN KEMBLE

Eric 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

which each species is assigned. Each taxon’s chromosome number (where known) is pro-vided. Species distribu-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.

The genus Echeveria by John Pilbeam. 2008. The British Cactus & Succulent Society. Hard cover, 333 pp. ISBN 0 902099 80 9. $94.95.

* 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.

Documents

Terry M 2008 Stalking the Wild Lop Hop Hora