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CACTI<br />

<strong>OF</strong> <strong>THE</strong><br />

<strong>SOUTH</strong>-<br />

<strong>WEST</strong><br />

TEXAS, NEW MEXICO,<br />

OKLAHOMA, ARKANSAS,<br />

AND LOUISIANA<br />

by Del Weniger


CACTI <strong>OF</strong> <strong>THE</strong> <strong>SOUTH</strong><strong>WEST</strong>


NUMBER FOUR<br />

The Elma Dill Russell Spencer Foundation Series


Echinocereus lloydii.


Cacti of the Southwest<br />

Texas, New Mexico, Oklahoma, Arkansas, and Luosiana<br />

By D E L W E N I G E R<br />

U N I V E R S I T Y O F T E X A S P R E S S A U S T I N & L O N D O N


Standard Book Number 292–70000–8<br />

Library of Congress Catalog Card No. 78–104326<br />

All rights reserved<br />

Printed by Brüder Hartmann, Berlin, West Germany


To Ellen Schultz Quillin


CONTENTS<br />

Acknowledgments . . . . . . . . . . . . . x<br />

Introduction . . . . . . . . . . . . . . xi<br />

What Is a Cactus? . . . . . . . . . . . . . 3<br />

Key to the Genera of the Cacti . . . . . . . . . . 8<br />

Genus Echinocereus Engelmann . . . . . . . . . . 10<br />

Key to the Echinocerei, page 11<br />

Genus WTzlcoxia . . . . . . . . . . . . . 55<br />

Genus Peniocereus . . . . . . . . . . . . . 57<br />

Genus Acanthocereus . . . . . . . . . . . . 6o<br />

Genus Echinocactus . . . . . . . . . . . . . 63<br />

Key to the Echinocacti, page 65<br />

Genus Lophophora . . . . . . . . . . . . . 95<br />

Genus Ariocarpus . . . . . . . . . . . . . 100<br />

Genus Pediocactus . . . . . . . . . . . . . 103<br />

Genus Epithelantha . . . . . . . . . . . . . 107<br />

Genus Mammillaria . . . . . . . . . . . . . 110<br />

Key to the Mammillarias, page 112<br />

Genus Opuntia . . . . . . . . . . . . . . 159<br />

Key to the Opuntias, page 162<br />

Glossary . . . . . . . . . . . . . . . 241<br />

Index of Scientific Names . . . . . . . . . . . 243<br />

Index of Common Names . . . . . . . . . . . 248


ACKNOWLEDGMENTS<br />

Support for this study was graciously provided by a grant-inaid<br />

for research from the Society of the Sigma Xi. Appreciation<br />

is expressed to Our Lady of the Lake College which granted<br />

me time for the study and made available certain facilities for<br />

growing and photographing the plants.<br />

Gratitude is also expressed to many individuals who contributed<br />

in various ways to the completion of the present study.<br />

First among them is Mrs. Roy W. Quillin (Ellen D. Schulz) who<br />

gave constant encouragement and advice and the use of books<br />

from her fine library on the Cactaceae. Dr. E. F. Castetter of the<br />

Biology Department of the University of New Mexico contributed<br />

valuable information and some specimens from New Mexico.<br />

He also arranged for the preparation and housing of my<br />

collection of important specimens gathered in this study. Numbering<br />

over seven hundred specimens of cacti, this collection is,<br />

as a result of his interest, permanently deposited in the her-<br />

barium of the University of New Mexico.<br />

The cacti of the following herbaria were studied in their entirety,<br />

thanks to the help of the staffs and especially of the persons<br />

mentioned here: the United States National Museum of the<br />

Smithsonian Institution, Dr. Jason R. Swallen and Dr. Velva<br />

Rudd being particularly helpful; the New York Botanical Garden;<br />

the Missouri Botanical Garden in St. Louis; The University<br />

of Texas; Southern Methodist University, with appreciation<br />

to Dr. Lloyd Shinners; the University of New Mexico and Dr.<br />

Castetter, as already mentioned; and the University of Colo-<br />

rado.<br />

Many others provided valuable information and aid of one<br />

kind or another. Appreciation goes to Dr. Barton H. Warnock<br />

of Sul Ross College; Drs. Claude W. Gatewood and Bruce<br />

Blauch, then of Oklahoma State University; Mr. Charles Polaski<br />

of Oklahoma City; Mr. Homer Jones and Mr. P. M. Plimmer<br />

of Alpine, Texas; Mr. Clark Champie of Anthony, New<br />

Mexico; Mr. Fred Nadolney, Mr.-Prince Pierce, and Mr. and<br />

Mrs. Earl Newhouse of Albuquerque, New Mexico; Mr. Horst<br />

Kuenzler and Mr. Dennis Cowper of Belen, New Mexico; Mr.<br />

L. J. Holland, Mr. and Mrs. S. L. Heacock, and Mrs. Ethel B.<br />

Karr, all of Colorado; Mr. and Mrs. Walter Hems of Rogers,<br />

Arkansas; Miss Lorene Martin of the Arkansas State Plant<br />

Board; Dr. T. M. Howard, Mr. Kim Kuebel, and Mr. H. C. Lawson<br />

of San Antonio, Texas; Mr. Glenn Spraker of Houston,<br />

Texas; Sr. Orton Cerna of Rosita, Coahuila, Mexico; and Sr.<br />

Romo Ruiz of Musquiz, Coahuila, Mexico.<br />

Mr. Larry Nichols and Mr. Gibbs Milliken aided the writer in<br />

making some of the photographs. The photograph of Lopho-<br />

phora williamsii var. echinata was made by Mr. David Smith.<br />

The chromatographs mentioned in the text were run by my<br />

students, Misses Vicki Perez, Rosemary Drake, and Ethel Matthews<br />

in the Biology Department of Our Lady of the Lake College.


INTRODUCTION<br />

Portrayed among the ancient stone carvings of Mexico, woven<br />

through the legends of that land, and central in the seal of the<br />

nation itself is the strange thick stem or the spiny jointed bush<br />

of the cactus. These plants must have been ever present for the<br />

people of that land, interesting to them and of some significance<br />

to their lives as far back as we can know. How strange to realize<br />

then that this fantastic group of plants had never been<br />

seen by anyone in the so-called civilized world until after<br />

Columbus discovered America. Yet this must be true; for, except<br />

for two or three small, inconspicuous, and very uncactus-like<br />

species found in places even more unknown to early travelers<br />

in the jungles of Madagascar and Ceylon, the cacti grow naturally<br />

only in the Western Hemisphere. They are as American<br />

as corn, tomatoes, tobacco, or potatoes.<br />

So they must have been seen—and felt—by the conquistadors<br />

who drove their horses across America in search of gold.<br />

No doubt these men paid little attention to the native flora as<br />

they passed, but the cacti they would have noticed for at least<br />

two reasons. How could they have failed to see the huge thickets<br />

of these devilish plants which blocked their paths to the<br />

envisioned gold and which they had to learn to respect? And<br />

when, wandering in the arid expanses, they grew hungry<br />

and had nothing else to eat, they must quickly have learned<br />

from the Indians to spot the cactus species whose fruits were<br />

succulent and sweet to the taste. We can imagine the stories they<br />

told about these outlandish plants of the New World.<br />

These accounts must have spread very rapidly, for we already<br />

have in The Herball or Generall Historie of Plantes, by John<br />

Gerarde, published in London in 1597, the descriptions and<br />

woodcut illustrations of four cacti. Gerarde names them as follows:<br />

“The Hedgehogge Thistle,” which is clearly seen from the<br />

picture to be a Melocactus from the Caribbean area, “The Torch<br />

or Thornie Euphorbium,” which is easily recognized to be a<br />

Cereus, “The Thornie Reede of Peru,” another Cereus, and “The<br />

Indian Fig Tree,” obviously a giant Opuntia. This is already<br />

an array of cacti from widely separated parts of Central and<br />

South America.<br />

Soon there followed explorers who were more directly interested<br />

in the flora of the New World. These early explorers<br />

took living specimens of the cacti home to Europe, not only<br />

because of their uniqueness, but because they would survive the<br />

long sea voyages when other plants, except in the seed stage,<br />

would not. They could even be transported all the way without<br />

the prohibitive weight of soil on their roots.<br />

Early botanists, beginning to study these plants, faced a problem.<br />

Since the plants had been totally unknown in the Old<br />

World until this time, there were no words for them at all in<br />

classical Greek or Latin. Like John Gerarde, the botanists often<br />

thought of these plants, because of their spininess, as new sorts<br />

of thistles, and so the Greek word for thistle, Kaktos, became<br />

somehow applied to them. This has become our cactus of to-<br />

day.<br />

There were soon expeditions of botanists just to study the<br />

new plants of America. An example was the Ruiz and Paron<br />

expedition of 1777 to 1787. This expedition to Peru was commissioned<br />

by the King of Spain and encompassed ten years of<br />

exploring in most difficult terrain; Spain spent upon the venture<br />

twenty million pesetas. Thousands of plant specimens were<br />

sent back, with cacti among them.<br />

The interest in these strange plants grew and soon amounted<br />

to a “cactus craze.” The extent of the “craze” is hard for us to<br />

comprehend today. By 1800 businesses were being set up by<br />

French, Belgian, and German importers to sell quantities of the<br />

plants sent by professional collectors maintained in Central and<br />

South America. Societies and wealthy enthusiasts commissioned<br />

collectors to travel to America and bring them newer and yet<br />

more strange species. Extensive collections were soon formed,<br />

grown at great pains in greenhouses. About 1830 the Duke of


xii cacti of the southwest<br />

Bedford had such a collection of cacti at Woburn Abbey. Other<br />

famous collectors in England were the Duke of Devonshire and<br />

the Reverend Mr. H. Williams of Hendon.<br />

This cultivation of the cacti must at first have been a very<br />

expensive, very genteel hobby, open only to the rich; the plants<br />

had to be expensive after having been shipped all the way from<br />

the wilds of the Western Hemisphere, and the resources necessary<br />

to keep them alive in the climate of most of Europe before<br />

the advent of gas and electric heat must have been great. The<br />

greenhouse full of cacti may well have been one of the few<br />

warmed buildings for miles around on freezing winter nights.<br />

Borg has pointed out that the cultivation of cacti has paralleled<br />

the cultivation of orchids in many ways, and this is easy to understand,<br />

since these are two of the most exotic groups of plants<br />

which can be found.<br />

But the cultivation of the cacti became the common man’s<br />

hobby much more easily than did that of the orchids. With<br />

botanical associations and importers putting out long lists of<br />

available species, cacti became cheap and available in Europe<br />

in large numbers. Soon many a humble home had at least a few<br />

of these peculiar plants in a window somewhere. We like to<br />

think of this as wonderful and of these cherished cacti as beautiful,<br />

but we must admit that even then they were not universally<br />

appreciated. Dickens, for instance, must have had an active<br />

aversion to them. While attesting to the broadness of the interest<br />

in cacti, his description of Paul Dombey’s nurse, Mrs. Pipchin,<br />

reveals his dislike for them, for he wrote of her, “Among her<br />

failings was a fondness for cactus. In the window of her parlor<br />

were half a dozen specimens writhing round bits of lath like<br />

hairy serpents.”<br />

Perhaps Dickens was sensing something cunning and insidious<br />

in these cacti, which events proved was there. They were early<br />

grown in southern Europe, and it was found that they could be<br />

grown without protection in outdoor gardens in southern Italy,<br />

Spain, Sicily, and Greece, on the Riviera, and, of course, all<br />

along the southern shore of the Mediterranean. In these areas<br />

are still today some of the finest cactus gardens in the world,<br />

with beautiful, hundred-year-old specimens to be seen.<br />

But once introduced into the Mediterranean area, certain of<br />

the Opuntias, the hardiest and most easily spread of cacti, found<br />

the hot, arid region too much to their liking and so escaped<br />

from cultivation and established themselves as permanent residents<br />

of the area. These cacti have now spread through many<br />

Mediterranean countries. Though the time required to accomplish<br />

this was actually comparatively short, man’s memory is<br />

even shorter, for so completely are they already accepted as a<br />

normal part of the flora that in many areas one finds hardly<br />

a resident who realizes that his ancestors could not have known<br />

these immigrants. Sometimes one even sees cacti in pictures and<br />

movies supposedly reconstructing the time of Christ and the<br />

Roman Empire, or in otherwise accurate portrayals of classic<br />

Greek times.<br />

In one other place cacti escaped like this and became one of<br />

the worst plant scourges ever known. This was in Australia,<br />

where several Opuntias, in the absence of their natural enemies<br />

and in an arid situation exactly to their liking, took over millions<br />

of acres and rendered them useless for anything else. Much<br />

money and effort was spent in discovering how to control these<br />

cacti in Australia, and the effort has been largely successful.<br />

They have been eliminated from huge areas and are being kept<br />

in check in others.<br />

But these two instances of cacti escaping and invading new<br />

areas are the exceptions. Usually, when taken from their natural<br />

haunts, the cacti survive only under very precise conditions<br />

and when great care is lavished upon them by their growers.<br />

Almost none of them can survive unaided even when only<br />

transplanted from one state to another within the United States.<br />

Within the Americas, where they are at home, cacti as a<br />

group are very widespread. They range from Alberta and British<br />

Columbia in Canada on the north to Patagonia, toward the<br />

tip of South America on the south. Their greatest development<br />

in both numbers and diversity is in two areas, one along the<br />

Tropic of Cancer in Mexico and the other near the Tropic of<br />

Capricorn in South America. While flourishing the most in the<br />

American deserts, they are far from restricted to these places.<br />

Special forms are found in tropical rain forests; others abound<br />

along the seashores of the Gulf of Mexico, the Caribbean, and<br />

the Pacific Ocean; while some thrive in mountain forests and a<br />

few are at home on bleak mountain slopes to as high as fourteen<br />

thousand feet. There are also some forms which abound on the<br />

Great Plains all the way into Canada, and within the United<br />

States it is said that indigenous cacti have been found growing<br />

in every state except Maine, Hawaii, and Alaska, although in<br />

many states they are so rare and inconspicuous that their pres-<br />

ence is unsuspected by most residents.<br />

This picture of far-ranging cacti is misleading if one thinks<br />

of any single form as so far-flung. This is the range of the group,<br />

which is a huge and diverse aggregation. It is probably fruitless<br />

to try to estimate the number of cactus forms that exist, because<br />

new publications are constantly adding new-found forms to the<br />

list, as well as realigning those already known. But it is said that<br />

there are well over three thousand known species in all. Of<br />

these, almost none range across from one America to the other.<br />

A very few species, considered broadly, may range practically<br />

across a continent, and some few cover large expanses of one or<br />

the other of the Americas, but the much more usual situation is<br />

for each species to inhabit a range measured in a few hundred<br />

square miles or less. It is very common for a species to inhabit<br />

only a certain valley or mountain range, and there are numerous<br />

forms so restricted in habitat that two or three Texas-sized<br />

ranches will contain them all.<br />

As we have seen, the cacti which excited the first interest and<br />

touched off the first cactus craze were the huge and spectacular<br />

Central and South American forms. The cacti of the United<br />

States were hardly known until later. Early botanists on the<br />

east coast of the United States found only two or three very


introduction xiii<br />

inconspicuous and uninteresting little prickly pears which they<br />

dutifully recorded and no one got excited about. As they pushed<br />

west, they found little else until they got beyond the Mississippi<br />

River. But once they began to explore the West, the study of<br />

U.S. cacti began.<br />

The first great student of U. S. cacti was the famous botanist<br />

Dr. George Engelmann. He made his headquarters at the Missouri<br />

Botanical Garden, the remarkable early botanical center<br />

in St. Louis, and studied the specimens and descriptions sent in<br />

by botanists on the early governmental surveys through the<br />

West. These botanists—such as Wislizenus, Wright, Bigelow,<br />

Parry, and Poselger—were the heroes of early cactus studies in<br />

the United States, and some of their names will be met with<br />

later, since Engelmann acknowledged his debt to them by sometimes<br />

naming cacti after them. They must have been hardy souls,<br />

riding on long treks with expeditions such as the Mexican<br />

Boundary Survey, the Pacific Railroad Survey, Pike’s expedition,<br />

and others; and one must admire their stamina, gathering<br />

cacti all day on the trail, then, while the others rested, making<br />

their records and descriptions and packing up specimens of these<br />

unpleasant-to-handle plants to send back to Engelmann. Their<br />

routes took them through the heart of the cactus country of the<br />

West, and we are amazed at the number of plants found only in<br />

places almost inaccessible even a hundred years later, which<br />

they located and recorded so long ago.<br />

Beginning in about 1846, Engelmann started publishing the<br />

results of these expeditions. He faced the herculean task of listing<br />

and describing a huge population of cacti almost unknown<br />

before to the world. He coined names for the multitude of<br />

forms, worked out something of their relationships, and presented<br />

descriptions and some of the finest botanical illustrations<br />

ever made for any plants. Although his material was sometimes<br />

incomplete and so his descriptions were sometimes deficient, he<br />

gave us the first information we have of approximately two-<br />

thirds of the U. S. cacti, information so remarkably accurate<br />

that in a few cases it has taken us almost a hundred years to<br />

verify it. Modern concepts have sometimes revised his ideas of<br />

the relationships between the forms, but almost never have we<br />

found him in error when he told where a plant grew or what it<br />

looked like.<br />

The next effort at studying the cacti of the United States was<br />

made by the great botanist John M. Coulter. In 1894 he began<br />

publishing a major work on the cacti of North America. Mostly<br />

he built on the foundation laid down by Engelmann, with the<br />

benefit of much more material collected since Engelmann’s time,<br />

but he added little really new to what was already known. It is<br />

indicative of the difficulty of studying cacti that he is said to<br />

have given up the study of this group in disgust and spent the<br />

rest of his life with other plant groups after misidentifying a<br />

cactus he had earlier named himself.<br />

About the same time as Coulter’s study, a large, general work<br />

on cacti was being produced in the German language by Karl<br />

Schumann. He did firsthand work on the cacti of the then In-<br />

dian Territory and the Canadian River area, but did not travel<br />

widely in the United States, and so his work has limited value<br />

for us today in the study of U. S. cacti.<br />

A major project on cacti was then undertaken by the Carnegie<br />

Institute. Dr. N. L. Britton and Dr. J. N. Rose, with the<br />

support of that institution, undertook to list and describe all<br />

the cacti of the world. They traveled widely throughout the<br />

Americas and visited the European collections, and as a result<br />

published a four-volume work, The Cactaceae, in 1919 to 1923.<br />

Their work had probably the greatest effect of anything ever<br />

published on the study of cacti as well as on the growing popularity<br />

of these plants. They greatly revised the classification of<br />

the group, adding a multitude of new genera and species, and<br />

their beautiful volumes, with fine color illustrations, were widely<br />

circulated, giving many people, especially in the United States,<br />

their first knowledge of the beauty of the cacti. Even today,<br />

most people still think of cacti strictly in the terms of the<br />

Britton and Rose accounts.<br />

No new attempt to encompass all of the cacti in one large<br />

study was made for many years. The task had become just too<br />

big. But the great German student of cacti, Curt Backeberg, did<br />

not flinch at the challenge, and in 1958 he brought out the first<br />

volume of a new world-wide survey, Die Cactaceae. It was<br />

completed shortly before his death and comprises six large volumes<br />

of fine descriptions, with many good pictures of the cacti<br />

of the world. It is truly a monumental work.<br />

But the great diversity of the cacti and the fact, already mentioned,<br />

that the majority of them are limited to areas which are<br />

very small (often single mountain ranges or river valleys), when<br />

set against the huge expanse covered by the group as a whole,<br />

make it very difficult for any major flora or all-inclusive cactus<br />

work to be useful on a local level. Who wants to carry the four<br />

volumes of Britton and Rose or the six of Backeberg to the Big<br />

Bend of Texas or the mountains of Peru? And if these works<br />

were to give really detailed accounts of the cacti found in all<br />

of such areas, exactly the information which the local student<br />

needs, they would become encyclopedic in size. This fact explains<br />

the numerous regional publications on cacti, both articles<br />

in journals and separate books. If one is to understand the local<br />

cacti, he must refer to these publications, done by people on the<br />

scene who have studied the larger picture and then sought out<br />

and portrayed the details of the local forms he sees about him.<br />

This book is intended to be such a regional guide.<br />

In the United States there has long been a tendency to break<br />

down the treatment of cacti into state studies. States are artificial<br />

areas and their boundaries have nothing to do with plant<br />

distribution, but it has been impossible to ignore them. The present<br />

study, however, includes the cacti of five states: Arkansas,<br />

Louisiana, Oklahoma, Texas, and New Mexico. These five states<br />

make up a unit much more logically considered, cactuswise, than<br />

any one of them alone, and a unit for whose cacti there has<br />

never been a complete guide. While lists of cactus species and<br />

some good descriptions are found in the floras of the respective


xiv cacti of the southwest<br />

states (as for instance those of Wooton and Standley for New<br />

Mexico) and many articles concerning various cacti in this region<br />

are scattered all through the literature, there has been only<br />

one complete work on cacti within this area: Texas Cacti, by<br />

Ellen D. Schulz (Mrs. Roy Quillin) and Robert Runyon. This<br />

was published in 1930, covered only the cacti of Texas, and is<br />

now out of print.<br />

The need for such a guide for these five states seems, therefore,<br />

clear. While the cacti of the far Southwest have been dealt<br />

with in numerous publications—Stockwell and Brezzeale’s Arizona<br />

Cacti of 1933, Baxter’s California Cacti of 1935, Boissevain<br />

and Davidson’s Colorado Cacti of 1940, Benson’s The<br />

Cacti of Arizona of 1950, and Earle’s Cacti of the Southwest<br />

of 1963—none of these covered more than those few forms of<br />

cacti which happened to extend their range into our area from<br />

the West. And where the cacti of this five-state area have been<br />

written about in journals or magazines it has usually been done<br />

by students who lived and worked in the far West or even in<br />

Europe, and who wrote of our cacti after brief trips through<br />

this vast area or from secondhand accounts.<br />

The cacti of this area have not lacked a detailed treatment<br />

because they were fewer in number or less diverse than those<br />

of the far Southwest. If anything, they may have presented too<br />

great a challenge just because of their number. I have found<br />

some professional botanists who believed, probably because of<br />

the greater size and conspicuousness of the Arizona species plus<br />

the greater publicity they have received, that the greatest speciation<br />

in the United States occurs in the far Southwest. Actually,<br />

this is not true. Texas alone presents more species of cacti than<br />

all of the rest of the United States combined. L. Benson lists 60<br />

cactus species in Arizona. California is said to have about 20<br />

natives; and New Mexico, about 50. On the other hand the<br />

number of separate species listed here for our five-state area is<br />

119. Earle, the most recent writer on the cacti of the Southwest,<br />

lists 121 forms, counting varieties as well as separate species for<br />

all of California, Arizona, Colorado, Utah, Nevada, and western<br />

New Mexico. Within our area we find 172 different forms.<br />

Within Texas alone can be found 106 species and 142 recognizable<br />

forms.<br />

The present work, then, lists all of the forms of cacti presently<br />

known to be growing within the five states: Arkansas, Louisiana,<br />

Oklahoma, Texas, and New Mexico. They are listed by<br />

their recognized scientific names, and immediately following<br />

(in every case where such exist) are given all of the common<br />

names which could be discovered, by which the cactus is known<br />

in various localities, including the Spanish and sometimes the<br />

Indian names. Spellings of these common names show the local<br />

variations found in the literature.<br />

A description of the whole cactus plant, not meant to be tedious<br />

but made full enough to be useful for the serious student,<br />

is then given. This description is patterned on the original one<br />

of the species, but it takes into account other descriptions by<br />

more recent students plus our own observations and so may be<br />

broader than the original in the case of quantitative characters<br />

and may add information not covered in the original.<br />

Following this is outlined the known range of natural distribution<br />

for the cactus. This is given in rather general terms, partly<br />

because there is still more to be learned about the spread of some<br />

of these plants and partly because it is not meant to be a guide<br />

telling exactly which mountain slope will provide the collector<br />

with the cactus he wants. The publication of such specific information<br />

all too often has meant that the slope is bare of any<br />

cacti at all the next year. There are no laws protecting cacti in<br />

any of the states covered in this study, and the conservation of<br />

our cacti is a real problem today. While this work gives no inaccurate<br />

or misleading information about where the various<br />

cacti are to be found, the exact locations of specific populations<br />

are not usually given. Any cactus hunter worthy of the title of<br />

“cactophile,” once given the general territory where the plant<br />

grows, might rather search out the prize himself.<br />

Next follows a discussion of each cactus listed. This is a<br />

gathering together of any remarks I might have on anything<br />

unusual or especially interesting about the cactus. Included<br />

under remarks also, because these are purely my own opinions<br />

after studying the plants and the literature upon them, are suggestions<br />

concerning its relationship to its fellows, with restatement<br />

of the specific characters which distinguish it from its<br />

closest relatives.<br />

These relationships are very complex, and the concepts of<br />

them have been almost constantly changing throughout the<br />

study of the cacti, as can be quickly traced by looking at the<br />

synonymy in almost any listing. In an attempt to avoid making<br />

this part another series of synonym lists in fine print—or a dry,<br />

dead discussion of dead names-this synonymy is presented as<br />

a historical account of the vicissitudes through which individual<br />

cactus forms, individual plant names, and the ideas of individual<br />

students went from their discovery and their first<br />

statements until the present. Such a historical approach to this<br />

material has not been made before, and it can enable us to<br />

understand much about the science of botany and about men,<br />

as well as about cacti. Of course, the material is much simplified<br />

and rendered as nontechnical as possible. Nevertheless an attempt<br />

is made to evaluate fairly, from the vantage point of the<br />

present, each important author's arguments and to assign his<br />

proposal its proper place in the history of the cactus being discussed.<br />

Lastly, each cactus form is illustrated by a full-color photograph<br />

of the plant, in most cases in bloom. A few of these<br />

photographs were made in the plant's natural location, but in<br />

most cases this proved to be impractical. Most cacti bloom only<br />

a few days out of the year, and it was obviously impossible to<br />

be in a canyon of the Texas Big Bend, on the Wichita Mountains<br />

of Oklahoma, on an Indian reservation in northwest New<br />

Mexico, and on an Ozark slope in Arkansas on precisely the<br />

days when each cactus chose to bloom. Much effort, therefore,<br />

has gone into the work of locating the various forms in the


introduction xv<br />

wild, then bringing them in and growing them in such congenial<br />

environments that they have bloomed, so that they could<br />

be pictured at the right moment. Unless otherwise stated, I<br />

made all photographs myself.<br />

The soils and backgrounds visible in the pictures are, therefore,<br />

not usually the natural environments of the plants. In fact,<br />

the colors of these have often been chosen to contrast with and<br />

make as clearly visible as possible the spines and other characters<br />

of the cacti. This means that no conclusion about the environments<br />

of these plants can be drawn from these pictures.<br />

While some readers may consider this a drawback, it should be<br />

remembered that most of these cacti use protective coloration<br />

and camouflage. Pictures of them in their natural habitats, while<br />

of value to the ecologist, usually show little detail of the plants,<br />

if they are visible at all. An extreme illustration of this difficulty<br />

would be the case of Mammillaria nellieae Croiz., where<br />

the whole plant is usually totally covered by the moss which<br />

grows with it in its rock crevice; a picture of it in loco would<br />

show only the flower apparently blooming on the moss and<br />

would be useless in illustrating what the cactus is really like.<br />

In the verbal description of the flower parts, I was confronted<br />

with the choice of employing standardized color names, such<br />

as those given in A Dictionary of Color by Maerz and Paul, or<br />

of using more general terms which would be meaningful to the<br />

average reader. I decided on the latter course in the belief that<br />

what would be gained in preciseness by the use of terms from A<br />

Dictionary of Color would be counterbalanced by the incon-<br />

venience of having to consult this reference work in a library in<br />

order to determine what was meant by the shade names used.<br />

Although as full as possible a rendering of the beauty of the<br />

flowers is an aim of these pictures, this is not their only goal. It<br />

is hoped that they may also convey a real concept of the plant<br />

body itself and of the spine character; sometimes the flower is<br />

shown at less than full angle because of this other aim.<br />

In organizing this presentation, one of the biggest problems<br />

was the delineation of the genera. The widest possible range of<br />

opinions is held today by the different authorities in the field<br />

on the limits of the genera in the Cactaceae. It seems that the<br />

present extent of the knowledge of the cacti does not enable<br />

anyone to give as definite a list of cactus genera as can be made<br />

for many other plant groups. There are several very different<br />

systems of genera, each very logical in the light of a certain set<br />

of assumptions. I have attempted to re-evaluate these in the<br />

light of the latest research available. The work of Dr. Boke at<br />

Oklahoma University and results of our own chromatographic<br />

studies seem particularly important here. The resulting alignment<br />

of the genera is in no case a new one, but in some cases<br />

favors one previous proposal and in some another.<br />

Within the genera no attempt has been made to organize the<br />

species into tribes or sections, since this sort of thing—as, for<br />

instance, various proposals for the genus Opuntia—seems still<br />

to be based on conjecture, and I find little newer and more solid<br />

evidence for any of the various contradictory proposals already<br />

made.


P L AT E S<br />

Measurements given in the photograph captions are the plant body<br />

sizes of the specific plants pictured and do not, unless otherwise stated,<br />

include flowers. In most cases this size is smaller than the maximum<br />

size achieved by the species.


Echinocereus viridiflorus var. viridiflorus. Larger plant 21/4 inches tall.<br />

Echinocereus viridiflorus var. cylindricus. 41/4 inches tall. Echinocereus viridiflorus var. standleyi. 4 inches tall.<br />

1


2<br />

Echinocereus davisii. 1 inch tall.<br />

Echinocereus chloranthus var. neocapillus. 31/2 inches tall.<br />

Echinocereus chloranthus var. chloranthus. Young plant (left):<br />

mature plant, 4 inches tall (right).<br />

Echinocereus chloranthus var. neocapillus. Immature plant (center),<br />

11/4 inches tall, flanked by mature specimens.


Echinocereus russanthus. 41/2 inches tall.<br />

(above, right)<br />

Echinocereus caespitosus var. caespitosus.<br />

The white-spined “Lace Cactus,” 3 inches tall.<br />

Echinocereus caespitosus var. caespitosus. The<br />

brown-spined “Brown-Lace Cactus,” 5 inches tall.<br />

3


4<br />

Echinocereus caespitosus var. minor. Largest stem pictured, 2 inches tall.<br />

Echinocereus caespitosus var. perbellus. 2 inches tall.


Echinocereus melanocentrus. 2 inches tall.<br />

(above, left)<br />

Echinocereus caespitosus var. purpureus.<br />

31/3 inches tall.<br />

Echinocereus fitchii.<br />

21/4 inches tall.<br />

5


Plate 6<br />

(above, left) Echinocereus baileyi.<br />

White-spined. 41/4 inches tall.<br />

(above, right) Echinocereus baileyi. Brown-spined,<br />

clustering. Largest stem pictured, 21/4 inches tall.<br />

(below) Echinocereus albispinus. Tallest stem<br />

pictured, 2 inches high.<br />

Plate 7 (opposite)<br />

(above, left) Echinocereus pectinatus var. wenigeri.<br />

Stem 41/2 inches tall.<br />

(above, right) Echinocereus pectinatus var. rigidissimus.<br />

6 inches tall.<br />

(below, left) Echinocereus chisoensis.<br />

7 inches tall.<br />

(below, right) Echinocereus pectinatus var. ctenoides (right)<br />

and Echinocereus caespitosus var. caespitosus (left).<br />

Note typical ovary and fruit coverings.


8<br />

Echinocereus pectinatus var. ctenoides. 3 inches tall.


Echinocereus dasyacanthus var. dasyacanthus.<br />

Yellow-flowered. 101/2-inch stem.<br />

Echinocereus dasyacanthus var. hildmanii. Stem 5 inch tall.<br />

Echinocereus dasyacanthus var. dasyacanthus.<br />

Pink-flowered. 12-inch stem.<br />

9


10<br />

Echinocereus roetteri. 43/4 inches tall.


Echinocereus lloydii.<br />

12 inches tall.<br />

Echinocereus triglochidiatus var. triglochidiatus.<br />

8 inches tall.<br />

11


12<br />

Echinocereus triglochidiatus var. octacanthus.<br />

Stems 41/4 inches tall.<br />

(below, left)<br />

Echinocereus triglochidiatus var. gonacanthus.<br />

4 inches tall.<br />

(below, right)<br />

Echinocereus triglochidiatus var. coccineus.<br />

61/2 inches tall.


Echinocereus coccineus var. conoides. 8 inches tall. Echinocereus polyacanthus var. rosei. 8 inches tall.<br />

Echinocereus polyacanthus var. neo-mexicanus. 41/2 inches tall.<br />

13


14<br />

Echinocereus stramineus. Clump, 30 inches in diameter. Echinocereus enneacanthus var. enneacanthus.<br />

Clustered stems, 13 inches across.<br />

Echinocereus enneacanthus var. carnosus. Tallest stem 8 inches high.


Echinocereus fendleri var. rectispinus.<br />

Tallest stem pictured, 8 inches high.<br />

(above, left)<br />

Echinocereus fendleri var. fendleri.<br />

Stem 53/4 inches tall.<br />

Echinocereus dubius. Tallest stem<br />

9 inches high. Two specimens of<br />

Lophophora williamsii in right<br />

foregrounds.<br />

15


Echinocereus papillosus var. angusticeps.<br />

Tallest stem pictured, 3 inches high.<br />

16<br />

(above, right)<br />

Echinocereus papillosus var. papillosus.<br />

Sprawling stems, 2 inches high.<br />

Echinocereus pentalophus. Stems approximately<br />

3/4 inch in diameter.


Wilcoxia poselgeri. Single<br />

upright stem, 12 inches long.<br />

Echinocereus papillosus var.<br />

papillosus on ground.<br />

(above, left)<br />

Echinocereus blanckii.<br />

Stem pictured, 101/2 inches long.<br />

Echinocereus berlandieri. Tallest<br />

stems pictured, 4 inches high.<br />

17


18<br />

Peniocereus greggii. Flowers<br />

27/8 inches in diameter.<br />

(below, left)<br />

Acanthocereus pentagonus. Flower 61/2 inches<br />

long, including ovary and tube.<br />

(below right)<br />

Echinocactus horizonthalonius var. curvispina. 6<br />

inches in diameter.


Echinocactus texensis.<br />

6 inches in diameter.<br />

(above, left)<br />

Echinocactus horizonthalonius var. moelleri.<br />

7 inches in diameter.<br />

Echinocactus asterias.<br />

21/2 inches in diameter.<br />

19


20<br />

Echinocactus uncinatus var. wrightii. 8 inches tall.<br />

(above, left)<br />

Echinocactus wislizeni.<br />

14 inches in diameter.<br />

Echinocactus whipplei.<br />

3 inches in diameter.


Echinocactus mesae-verdae.<br />

13/4 inches in diameter.<br />

(below, left)<br />

Echinocactus brevihamatus.<br />

4 inches tall.<br />

(below, right)<br />

Echinocactus scheeri.<br />

21/2 inches tall.<br />

21


22<br />

Echinocactus tobuschii.<br />

25/8 inches in diameter.<br />

(below, left)<br />

Echinocactus setispinus var. hamatus.<br />

7 inches tall.<br />

(below, right)<br />

Echinocactus setispinus var. setaceus.<br />

10 inches tall.


Echinocactus sinuatus. 5 inches diameter.<br />

Echinocactus hamatacanthus.<br />

12 inches in diameter.<br />

(below, left)<br />

Echinocactus bicolor var. schottii. 4 inches in diameter.<br />

(below, right)<br />

Echinocactus flavidispinus. 2 inches in diameter.<br />

23


24<br />

Echinocactus intertextus var. intertextus.<br />

When collected, 23/4 inches in diameter.<br />

Echinocactus intertextus var. dasyacanthus.<br />

23/4 inches in diameter.<br />

Echinocactus intertextus var. intertextus. Same plant<br />

after 1 year of cultivation. 27/8 inches in diameter.<br />

Echinocactus erectocentrus var. pallidus.<br />

21/2 inches in diameter.


Echinocactus mariposensis. Green-flowered.<br />

15/8 inches in diameter.<br />

Echinocactus conoideus. 3 inches in tall.<br />

Echinocactus mariposensis. Pink-flowered.<br />

17/8 inches in diameter.<br />

25


26<br />

Plate 26<br />

(above, left) Ariocarpus fissuratus. 35/8 inches in diameter.<br />

(above, right) Lophophora williamsii var. williamsii.<br />

Largest stem 3 inches in diameter.<br />

(below) Lophophora williamsii var. echinata.<br />

Largest stem 21/2 inches in diameter.<br />

Plate 27 (opposite)<br />

(above, left) Pediocactus simpsonii var. simpsonii. 3 inches in diameter.<br />

(above, right) Pediocactus knowltonii. 1 inch in diameter.<br />

(below, left) Pediocactus papyracanthus. 2 inches tall.<br />

(below, right) Epithelantha micromeris. 11/2 inches in diameter.


28<br />

Mammillaria scheeri. 6 inches in diameter. Mammillaria scolymoides. 31/4 inches in diameter.<br />

Mammillaria echinus. Stem 21/4 inches in diameter.<br />

Mammillaria sulcata.<br />

41/8 inches in diameter.


Mammillaria ramillosa. 31/4 inches in diameter.<br />

29


Mammillaria macromeris. Young plant,<br />

not yet clustered. 3 inches tall.<br />

(below, left)<br />

Mammillaria runyonii.<br />

4 inches in diameter.<br />

(below, right)<br />

Mammillaria similis. Typical small<br />

plant; diameter 3 inches.<br />

30


Mammillaria similis. Old plant<br />

in full bloom. Diameter 12 inches.<br />

Mammillaria vivipara var. vivipara.<br />

21/4 inches in diameter.<br />

31


Mammillaria vivipara var. arizonica. 3 inches in diameter. Mammillaria fragrans 21/4 inches in diameter.<br />

Plate 32 (opposite)<br />

(above, left) Mammillaria vivipara var. radiosa.<br />

2 inches in diameter.<br />

(above, right) Mammillaria vivipara var. neo-mexicana. Large stem<br />

21/4 inches in diameter.<br />

(below, left) Mammillaria vivipara var. borealis.<br />

2 inches in diameter.<br />

(below, right) Mammillaria vivipara var. neo-mexicana.<br />

Young plants, showing juvenile spination on sides of stem and<br />

mature spines at the tips. Larger plant 2 inches in diameter.<br />

33


34<br />

Mammillaria tuberculosa.<br />

Main stem 35/8 inches tall.<br />

Mammillaria dasyacantha. Plant<br />

pictured, 3 inches in diameter.<br />

Plate 35 (opposite)<br />

(above) Mammillaria dasyacantha and<br />

tuberculosa, growing together.<br />

(below left) Mammillaria duncanii. Showing root<br />

formation. Spiny portion of the stem 1 inch tall.<br />

(below right)<br />

Mammillaria duncanii. Same plant, with fruit,<br />

after 3 months’ cultivation.


36<br />

Mammillaria albicolumnaria. Plant pictured, 27/8 inches in diameter.


Mammillaria varicolor. Largest stem<br />

12/3 inches in diameter.<br />

(below, left)<br />

Mammillaria hesteri.<br />

4 inches tall.<br />

(below, right)<br />

Mammillaria nellieae. Blooming<br />

plant, 1 inch in diameter.<br />

37


38<br />

Mammillaria roberti. Largest stem 13/8 inches in diameter.<br />

Mammillaria sneedii. Blooming stem 3/4 inch in diameter.


(above, left)<br />

Mammillaria leei. Plants in garden cultivation.<br />

Clusters 4 to 6 inches in diameter.<br />

(above, right)<br />

Mammillaria leei. Typical root formation.<br />

Large clump, 41/4 inches across.<br />

(below, left)<br />

Mammillaria pottsii. Largest stem, 11/3 inches<br />

across. M. lasiacantha var. denudata in foreground.<br />

(below, right)<br />

Mammillaria lasiacantha var. lasiacantha.<br />

Stem 1 inch in diameter.<br />

39


40<br />

Mammillaria lasiacantha var. denudata. Stem 2 inches in diameter. Mammillaria microcarpa. 2 inches tall.<br />

Mammillaria multiceps. Cluster of stems, 41/8 inches across.


Mammillaria wrightii. Stem 13/16 inches in diameter.<br />

41


42<br />

Mammillaria wilcoxii.<br />

Stem 17/8 inches in diameter.<br />

Mammillaria heyderi var. heyderi.<br />

41/3 inches across. Mammillaria heyderi var. applanata.<br />

4 inches across.<br />

Plate 43 (opposite)<br />

(above) Mammillaria heyderi var. hemisphaerica.<br />

31/2 inches across.<br />

(below) Mammillaria meiacantha.<br />

4 inches across.


44<br />

Opuntia stricta. Plant pictured 26 inches tall.<br />

Opuntia engelmannii var. engelmannii.<br />

Pad 12 inches long.<br />

Mammillaria sphaerica. Main plant, with offsets,<br />

43/4 inches across.


Opuntia engelmannii var. texana. Largest pads pictured,<br />

8 inches wide.<br />

Opuntia engelmannii var. alta. Red-flowered.<br />

Blooming pads, 51/2 inches across.<br />

45


46<br />

Opuntia engelmannii var. alta. White-flowered.<br />

Flower 31/8 inches across.<br />

Opuntia engelmannii var. cacanapa. Largest<br />

pad shown, 7 inches across.<br />

Opuntia engelmannii var. flexispina. Main pad 101/2 inches<br />

wide. White spots are cochineal insects.


Opuntia engelmannii var. aciculata. Largest pad<br />

62/3 inches across.<br />

(above) Opuntia engelmannii var. dulcis. Largest pad<br />

8 inches in diameter. This plant grew for many years in<br />

an urn by the entrance to the Old Trail Driver’s Museum<br />

in San Antonio.<br />

Opuntia engelmannii var. subarmata.<br />

Main pad 12 inches across.<br />

47


48<br />

Opuntia engelmannii var. linguiformis.<br />

Blooming pad 16 inches long. White spots<br />

are cochineal insects.<br />

(below, left)<br />

Opuntia chlorotica. Plant about 22 inches tall.<br />

(below, right)<br />

Opuntia tardospina. 9 inches tall.


Opuntia rufida. Largest pad<br />

pictured, 7 inches long.<br />

(above, left)<br />

Opuntia spinosibacca. Largest pad<br />

pictured, 53/4 inches long.<br />

Opuntia macrocentra. Largest pad<br />

pictured, 6 inches long.<br />

49


50<br />

Opuntia gosseliniana var. santa-rita.<br />

15 inches tall.<br />

(below, left)<br />

Opuntia strigil. Largest pad pictured,<br />

73/4 inches long.<br />

(below, right)<br />

Opuntia atrispina. Largest pad pictured,<br />

6 inches across.


(above) Opuntia phaeacantha var. major. Largest pad<br />

shown, 8 inches across.<br />

(below) Opuntia phaeacantha var. nigricans. Old 6-inch<br />

pad sprawling; and new, sprouting 4-inch pads.<br />

(above, left) Opuntia leptocarpa. Shown in fruit.<br />

Plant 15 inches tall.<br />

(below, left) Opuntia leptocarpa in foreground and<br />

Opuntia engelmannii var. texana in background.<br />

51


52<br />

Opuntia phaecantha var. brunnea. Wide-open flowers, 3 inches across.


Opuntia cymochila. Largest pad pictured, 4 inches across.<br />

(above, left)<br />

Opuntia phaeacantha var. camanchica. Largest pad<br />

pictured, 41/3 inches long.<br />

Opuntia phaeacantha var. tenuispina.<br />

Pads 9 inches long.<br />

53


54<br />

Opuntia compressa var. humifusa. Largest pad<br />

pictured, 3 inches across.<br />

Opuntia compressa var. microsperma. Largest pad<br />

pictured, 3 inches long.<br />

Opuntia compressa var. macrorhiza.<br />

Pad 4 inches long, blooming.<br />

Opuntia compressa var. fusco-atra. Two specimens from the same<br />

locality. Largest pad on stunted plant, 21/4 inches long;<br />

largest pad on vigorous plant, 5 inches long.


Opuntia compressa var. fusco-atra. An abnormal form known as O. macateei.<br />

Open flower 2 inches across.<br />

Opuntia compressa var. allairei. Largest pad pictured, 61/4 inches long.<br />

Opuntia compressa var. grandiflora. Largest<br />

pad pictured, 51/4 inches long.<br />

55


56<br />

Opuntia compressa var. stenochila. Largest pad pictured, 4 inches across.<br />

Opuntia ballii. Pad<br />

21/3 inches across.<br />

Opuntia pottsii. 61/2 inches tall, exclusive of fruits. Opuntia plumbea. Pad 21/4 inches long.


Opuntia drummondii. Largest pad pictured, 3 inches long.<br />

Opuntia fragilis. Clump 61/2 inches across. Opuntia sphaerocarpa. In winter condition.<br />

Largest pad pictured, 31/4 inches across.<br />

57


58<br />

Opuntia rhodantha var. rhodantha. Yellow flowered. 61/2 inches tall. Opuntia rhodantha var. rhodantha. Pink flowered. 8 inches tall.<br />

Opuntia rhodantha var. spinosior. Larger<br />

pad pictured, 41/2 inches long.<br />

Opuntia polyacantha. Typical, heavily spined.<br />

Largest pad pictured, 4 inches long.


Opuntia polyacantha. With spines few and short.<br />

Largest pad pictured, 55/8 inches long.<br />

Opuntia hystricina. Upright pad, 4 inches long.<br />

Opuntia arenaria. Largest pad pictured 23/4 inches long. Opuntia grahamii. Pictured clump, 7 inches across.<br />

59


60<br />

Opuntia Stanlyi. Largest joint<br />

pictured, 4 inches long.<br />

(below, left)<br />

Opuntia schottii. Pictured joints,<br />

each 2 inches long.<br />

(below, right)<br />

Opuntia clavata. Largest joints pictured.<br />

11/2 inches in diameter.


Opuntia imbricata var. arborescens. Branch white ripe<br />

fruits. Main pictured. 11/4 inches in diameter.<br />

Opuntia imbricata var. viridiflora. Pictured plant 15 inches tall.<br />

(above, left)<br />

Opuntia imbricata var. arborescens. Section of the<br />

main stem pictured, 11/4 inches in diameter.<br />

(below, left)<br />

Opuntia imbricata var. vexans. Branch with ripe fruits.<br />

Largest fruit pictured, 15/8, inches in diameter.<br />

61


62<br />

Opuntia spinosior. Largest stem pictured, 1 inch in diameter. Opuntia whipplei. In winter condition, 10 inches tall.


Opuntia davisii. Largest branch<br />

pictured, 5/8 inch in diameter.<br />

Opuntia tunicata. Plant pictured,<br />

13 inches across.<br />

63


64<br />

Opuntia leptocaulis. Main stem pictured, 1/4 inch in diameter. Opuntia kleinei. Plant pictured, 4 feet tall.


CACTI <strong>OF</strong> <strong>THE</strong> SOUT<strong>WEST</strong>


What Is a Cactus?<br />

Before going directly into the description of the various<br />

cacti we might pause to consider, for those who have not concerned<br />

themselves about these things before, how a cactus differs<br />

from other plants, what is so special about it, and what are<br />

some of the problems the uniqueness of its form and physiology<br />

bring to it in its natural situation and to us if we desire to raise it.<br />

It is harder to say exactly what a cactus is and how it differs<br />

from other plants than one might think. It is obvious that a<br />

cactus is a unique plant, with special problems. Imagining how<br />

it got that way, learning how to recognize it when we see it, and<br />

understanding its adaptations to its special problems—each of<br />

these presents us with special difficulties.<br />

The origin of the cactus family remains almost completely a<br />

mystery. We are balked here by the fact that there are no fossils<br />

of any cacti. So anxious have we been for such evidence that<br />

there have been several remains grasped hopefully as being cactus<br />

fossils, but all these, such as the famous one optimistically<br />

christened Eopuntia douglassii Chaney, have since proved not<br />

to be connected with the cacti at all. The most primitive and<br />

least cactus-like forms that we know are the members of the<br />

genus Pereskia, still alive, flourishing, and all-cactus, giving us<br />

no clear clues as to how they got that way. This leaves us with<br />

only theories based on comparative anatomy studies to satisfy us.<br />

Mostly because of certain flower characteristics, it is quite<br />

often assumed that cacti are related to the Rose Family. From<br />

here one can go as far as his imagination chooses to range, presuming<br />

with some that the roses of the West Indies changed in<br />

order to adapt to more arid conditions and so gave rise to the<br />

Pereskias and through them to all cacti. This is an enchanting<br />

story, and I am sure that the cactophiles are pleased with the<br />

idea that their favorites might be descendants of the rose, but I<br />

am not so certain that the rose enthusiasts are as sympathetic to<br />

the idea of appending the cacti to their queen of the flowers.<br />

Most agree that the cacti are a young group, maybe 20,000<br />

years old, and an equally big problem is how they could have<br />

developed their extreme and fantastic adaptations in such a<br />

comparatively short time.<br />

Be that as it may, the cacti are here, and one needs to know<br />

how to recognize them. This task is complicated by the fact that<br />

most of the obvious characters by which one thinks to recognize<br />

them are shared by some plant or other somewhere in the<br />

world. This is true of such things as large, fleshy stems, vicious<br />

spines, and reduction of leaves, which to an amateur mean cactus<br />

every time. There are other plants showing all of these characteristics,<br />

some of them to almost the extent the cacti do.<br />

It is a failure to recognize this fact that accounts for the popular<br />

articles, nursery ads, and many “cactus plantings,” containing<br />

mention or actual specimens of completely unrelated plants<br />

under the banner of cactus. Although they show some at least<br />

of the above characteristics, it must be stated that such things<br />

as yuccas, agaves, century plants, sotol, and so on, are not cacti<br />

at all, but extremely modified members of the Lily Family.<br />

Ocotillo and allthorn are individual residents of the desert community<br />

showing some of the same adaptations, but belonging to<br />

other plant families. Then there is the whole multitude of African<br />

plants paralleling the cacti in almost every feature of stem,<br />

rib, spine, and leaf, but all belonging to the huge, world-wide<br />

genus Euphorbia which also includes such plants as the Poinsettia.<br />

We sometimes speak of all these other plants as succulents,<br />

setting up the categories of cacti and succulents—although<br />

the cacti are also succulent, since the word merely means<br />

“fleshy.”<br />

How then do you tell a cactus? There is no easy way to do it<br />

without close observation of details and something of a bota-


4 cacti of the southwest<br />

nist’s eye. However, a cactus is always a dicot, and its two seed<br />

leaves will distinguish it at once from all those members of the<br />

Lily Family so often called cacti, since they are monocots and<br />

have only one seed leaf.<br />

Then, a feature which all cacti have and share with no other<br />

plant is the structure called the areole. All cacti have areoles<br />

quite liberally scattered over the surface, and usually arranged<br />

in rows or spirals in the most conspicuous places. These are<br />

round to elongated spots from 1/16 to sometimes well over 1/2<br />

inch in greatest measurement; their surfaces are hard, rough,<br />

uneven, and brown or blackish or else covered with white to<br />

brown or blackish wool. These areoles are now considered to be<br />

the equivalent of complex buds, and it is from these that whatever<br />

spines the cactus possesses grow. These spines, since they<br />

come from these areoles, are always arranged in clusters, which<br />

is another feature not found on other spiny plants, whose spines<br />

are produced singly from some source other than an areole.<br />

Beyond this, for the actual features separating cacti from all<br />

other plants, one has to look to the flower. Certain rather technical<br />

features of the flower are cited, such as its having sepals<br />

and petals numerous and intergrading, its having an inferior<br />

ovary with one seed chamber and having one single style with<br />

several stigma lobes.<br />

The key to understanding why the cactus is such a strange<br />

plant is the understanding of its major problem and how it<br />

solves it. This is its water problem.<br />

Most plants are great spendthrifts when it comes to water.<br />

They stand with their roots in unfailing water supplies from<br />

streams or from moisture stored in the soil between rains; and<br />

they constantly absorb quantities of water, which they use and<br />

then pass on out through their leaves into the atmosphere. This<br />

passage of water through the typical plant is so great that we<br />

call it the transpiration stream and can best think of such plants<br />

as constantly flowing fountains of water. If there is too little<br />

water available near them such plants normally solve the problem<br />

by developing extra long roots which go where the water<br />

is, and if this fails they dry out and die.<br />

But the cactus is typically a resident of the desert or else of<br />

habitats where, for one reason or another, the water supply is<br />

practically nonexistent at least part of the time. It may be<br />

because of inadequate rainfall or because the soil is too coarse<br />

or too thin to hold much water. So the cactus has a water<br />

problem which it solves in its own way.<br />

The cactus disdains to put out the extreme root systems, often<br />

drilling fifty feet deep, used by other desert plants to find the<br />

precious water which enables them to stay alive. Instead it sits<br />

and waits for the infrequent showers which ultimately do come,<br />

even in the desert, or for the rainy season. And when moisture<br />

does come the plant is ready. Its finely branching roots absorb<br />

water rapidly when it is available, and in a short time it has<br />

taken in a large amount. But there will be another drought to<br />

come, when it may be able to take in little if any water for<br />

weeks or months, so it stores this bonanza of water to the limit<br />

of its capacity, and its adaptations for great water-storing capacity<br />

form the basis for the most obvious peculiarity of the<br />

cactus.<br />

The commonest and most simple means of storing water found<br />

in these plants is by the enlarging of the stem into a thick, fleshy<br />

column or even a round ball. A cactus adapted this way becomes<br />

literally a water-filled column or ball, actually, in large<br />

specimens, a barrel of water-from which comes the common<br />

term, “barrel cactus.” The interior of such a cactus is not a reservoir<br />

of pure water into which you could dip a ladle, as cartoons<br />

sometimes show the thirsty prospector doing, but a mass<br />

of soft tissue permeated with water. Except for the supporting<br />

framework necessary in larger species, that interior is of about<br />

the consistency of a melon’s watery pulp. When rain comes it<br />

fills to the maximum with water, and in times of drought this<br />

reserve is gradually reduced. Thus, the cactus stem swells and<br />

shrinks according to the water supply, and there is always an<br />

arrangement of ribs or tubercles which make this change in bulk<br />

possible without the whole stem alternately caving in or split-<br />

ting open.<br />

In a number of the cacti where adaptations for clambering up<br />

trees, camouflage in thickets, or something else limits the thickness<br />

or size of the stems, the root may become the water-storage<br />

organ instead. In these cacti the root may become a carrot-like<br />

taproot weighing up to fifty pounds (in the extreme case of an<br />

old Peniocereus greggii specimen) or a cluster of tubers (as<br />

found on Wilcoxia poselgeri and some of the Opuntias).<br />

The cactus, then, is a plant which solves the problem of insufficient<br />

water in its habitat by storing large amounts of water<br />

within its tissues, and this explains its succulent consistency and<br />

bloated stem. Due to this habit it is internally among the softest,<br />

most delicate of plants.<br />

But standing there as almost literally a column or barrel of<br />

water in the middle of the thirsty desert brings problems requiring<br />

still further adaptations, giving us the other remarkable<br />

characteristics of a cactus. Since the cactus may have to survive<br />

for weeks or months on the moisture it has within it, it cannot<br />

afford to be a spendthrift with water like other plants. It has<br />

to give up a little at all times to stay alive, but it must sacrifice<br />

the smallest possible amount and protect its precious store<br />

against the dryness of the desert air which would otherwise<br />

evaporate it all in a matter of hours.<br />

For this reason the leaves of a cactus are reduced or eliminated<br />

altogether. After all, leaves are for the purpose of increasing<br />

the evaporative and light-absorbing surface of the plant, and<br />

the cactus needs to reduce the evaporative surface to a minimum,<br />

while certainly, in the glaring desert, it need not spread<br />

out its surface after light. Therefore, the more strictly a desert<br />

dweller it is, the more completely the leaves tend to be reduced<br />

or absent and the green stems to take over their functions. Then,<br />

its compact form is covered with a thick, waxy epidermis which<br />

is impervious to water, and even its stomata are equipped with<br />

means to reduce moisture loss. There is great variation in the


what is a cactus? 5<br />

tenacity with which individual cacti hold water, with the most<br />

extreme desert forms said to release up to six thousand times<br />

less water in a given moment than an ordinary plant of the<br />

same weight.<br />

The thick, dry, protective covering of the cactus is so deceptive<br />

to us that we seldom think of the soft, delicate, watery interior<br />

which it protects, but we may be sure the thirsty denizens<br />

of the desert, where water is life itself, are ever conscious of it.<br />

They would eat the plant immediately just for the water, if they<br />

could get at it. So the cactus has had to add protection against<br />

all the living water-seekers which surround it, and this gives yet<br />

another of its peculiarities.<br />

The almost universal solution of the cactus to this problem is<br />

to cover itself with an armament of spines. The succulent flesh<br />

is entirely covered with a system of spines so sharp and dangerous<br />

and so perfectly spreading and interlacing that neither the<br />

browsers nor the rodents can get their teeth between them to<br />

bite into the plant. The spines are never poisonous, and one can’t<br />

ascribe maliciousness to a plant for having them. They are there<br />

as necessary protection for the otherwise most delicate, most<br />

defenseless member of the desert community. It should help one<br />

to understand the spiny thing to realize that in the desert, when<br />

an injury or a malformation leaves a space wide enough for the<br />

jaws of a rabbit or even a mouse to get between the spines and<br />

start working, the cactus is soon eaten entirely. If one goes out<br />

on the desert and cuts all of the spines off a cactus, it usually<br />

disappears overnight. Ranchers have long ago learned to profit<br />

by this fact, and in some areas, by merely burning the spines off<br />

the huge prickly pears, provide their cattle with tons of free,<br />

succulent forage. The water problem, then, is directly responsible<br />

for the soft make-up of a cactus and indirectly responsible<br />

for its hard, waxy exterior and its often unpleasant but also<br />

fascinating array of spines.<br />

The cactus faces another closely related problem. Its habitats<br />

are usually extreme in their heat and the intensity of their light.<br />

The temperature on a south-facing, rocky desert slope reaches<br />

almost unbelievable heights on a summer afternoon. Even the<br />

desert reptiles are said to avoid the sun in which the extreme<br />

desert forms of cactus have to stand all day. How can they sur-<br />

vive this baking heat and searing light?<br />

Of course many cacti could not, and these grow only in the<br />

shade of thickets or trees, but the ones which stand and take it<br />

are said to depend on their own spines for shade. The spines<br />

achieve their shading effect, somewhat after the manner of a<br />

lath-house cover, by breaking the radiation up into moving<br />

strips of endurable duration. These forms also protect the exposed<br />

surface, especially the tender growing area at the top,<br />

with a covering of wool or hair, usually white and reflective.<br />

One can fairly well judge how extreme a desert situation a<br />

species comes from and how much sun it can stand by looking<br />

at how extensively this wool is developed or how complete the<br />

spine shading is.<br />

With no tender leaves, the compact body of the cactus, with-<br />

in its spiny envelope, is thus remarkably well protected against<br />

any of the natural forces or living enemies of its habitat, and it<br />

can survive in places where only the hardiest persist. Yet it has<br />

one more major problem to surmount. It must reproduce itself.<br />

And to do this it must usually produce a flower. Some of the<br />

cacti avoid this at all but the most favorable times, and depend<br />

instead upon very well-developed vegetative reproduction, but<br />

sooner or later all have to bloom.<br />

Now, a flower is an amazingly complex structure of extremely<br />

delicate parts. Flowering is a time when the plant must<br />

open and expose for the generating touch the most precious<br />

centers of its being. It is a time of vulnerability, and not even the<br />

cactus has succeeded in armoring the flower. It may swathe the<br />

bud in spines and wool, but when the moment comes it must<br />

expose the flower to the cruel desert situation as unprotected as<br />

any rose or lily. This presents another immense problem for the<br />

cactus and its way of solving it gives us both the wonderful<br />

beauty for which the flower is famous, and the extreme fleet-<br />

ingness of the flower which exasperates us.<br />

The cactus flower is almost always renowned for its size and<br />

beauty, which are said to be for the purpose of attracting insects<br />

or other flying forms across the arid distances to pollinate<br />

it. At any rate it does not seem to be beautiful for our benefit,<br />

because the flower usually lasts so short a period and blooms at<br />

such an unfavorable time that we hardly ever catch a glimpse<br />

of it and it usually is “born to blush unseen and waste its sweet-<br />

ness on the desert air.”<br />

Most cacti have flowers which open in the worst heat of the<br />

day, usually for only a few hours, and then are closed and fading<br />

before the cool of the evening begins. It is as though the<br />

plant waits until the heat of the day drives most of its enemies<br />

under the protection of some shade to unfold these tender morsels<br />

which it cannot otherwise protect. Thus, in most forms, the<br />

flower has its brief life, the reproductive act is completed by<br />

the insects which scorn the heat, and the life spark is already<br />

down within the spiny ovary before the desert cools, so that its<br />

thirsty tribes find only wilted petals for their evening meals.<br />

Many tropical and a few of our U. S. cacti reverse this schedule<br />

entirely and open their gigantic, wonderfully fragrant flowers<br />

at night to be pollinated by night-flying insects or in a few<br />

cases by bats. In most cases these species produce their flowers<br />

on tall, spiny stems where no ordinary enemy could reach them<br />

anyway, but they fade as quickly as the others, and are usually<br />

only sadly wilted remains by dawn. Only the saguaro, whose<br />

flowers are inaccessible to almost any enemy, and some other<br />

forms protected by especially long, vicious spines seem able to<br />

enjoy the luxury of longer lasting flowers.<br />

The cactus fruit, which follows the flower, is usually protected<br />

at first by spines or wool, and grows to become a berry with<br />

numerous small seeds. In some cases this dries up and the seeds<br />

are allowed to scatter, but in many species the ripe berry becomes<br />

fleshy and at the same time loses its spines or rises out of<br />

its wool covering. Here is probably the only part of a cactus


6 cacti of the southwest<br />

purposely left unprotected. It is never poisonous, and it ranges<br />

from sour to very sweet in different species. It is snapped up<br />

and carried off by animals and birds who finally get a meal<br />

from the cactus, but who pay for it by scattering the seeds far<br />

and wide. Some of the sweetest of these fruits are relished by<br />

humans. Those of the Opuntias are called “tunas”; and the<br />

“strawberry cactus” (of which there are several species) bears<br />

this common name because the flavor of the red fruits suggests<br />

that of strawberries.<br />

In all of its stages, then, the cactus is admirably adapted for<br />

survival in an arid environment, with all of these special features<br />

accentuated to the extreme in the forms inhabiting the<br />

more severe desert regions and less markedly developed in those<br />

of less extreme situations. But these same wonderful features<br />

which make the cactus so successful in the desert bring their<br />

own problems with them, limiting it in important ways even<br />

in its natural environment and making the tough desert thing<br />

one of the most vulnerable of plants when brought out of the<br />

desert into cultivation.<br />

Its very life, we have seen, depends upon the large amount<br />

of water stored within it as watery pulp. But this brings also<br />

the greatest danger to the cactus. Everyone knows how easily<br />

bruised and how quickly rotting is the watery flesh of melons<br />

and other soft fruits. A sharp blow or a gash through the protective<br />

covering of such structures causes a breakdown of the<br />

soft tissues which often spreads like wildfire throughout, leaving<br />

the whole thing a putrid, rotten mass. This is because such<br />

soft, nutrient-filled tissues form the perfect media for the<br />

growth of bacteria and all sorts of fungi.<br />

The interior of even the toughest cactus is just as vulnerable<br />

to fungi. It survives because this tender core is surrounded by<br />

the tough, fungus-resistant epidermis. The cactus is only safe<br />

when this forms an unbroken barrier covering not only the<br />

stem but the roots of the plant. But the slightest injury, any<br />

break in this epidermis, may let in a fungus, and if one gets in<br />

before the plant can repair the break with scar tissue and starts<br />

growing in the interior, the outwardly invincible old cactus<br />

will be attacked from within. It will then be quickly permeated<br />

by the fungus and will collapse into a foul, oozing thing—often<br />

literally overnight. For this reason any injury is a greater danger<br />

to a cactus than to most plants, especially if it has been<br />

removed from the desert, where fungi are not as numerous, to<br />

a more damp climate where they abound.<br />

But the fungi often gain entrance to our cacti in a more subtle<br />

way. The epidermis on the roots of these plants is necessarily<br />

thinner than that on the stems, and it is in constant contact<br />

with the fungus-populated soil. If there is very little rainfall<br />

or the soil is open and fast-draining, all will probably be well,<br />

but if there are periods of continuous rainfall, or if the soil is<br />

close-packed and remains for any time water-saturated, then<br />

the normally hard, dry, and impervious epidermis of the roots<br />

becomes wet through and softened, and loses its impermeability<br />

to the fungi. The defense barrier is dissolved, and almost any<br />

cactus whose roots lie in waterlogged soil for over twenty-four<br />

hours or so will be invaded and reduced in about that much<br />

longer to stinking carrion. This is the fate of most cacti taken<br />

in from the desert and planted in heavy yard soil in a more<br />

rainy region or else in a pot which is watered every day along<br />

with the geraniums. In general, cacti must have abundant water<br />

now and then to replenish their stored supply, but they must<br />

not stand in stagnant water at any time.<br />

We have also seen how the cactus survives not only by storing<br />

water but by being miserly with it and giving off a transpiration<br />

stream of up to several thousand times less volume than<br />

other plants. When other plants are wide-open, gushing fountains,<br />

the cactus is a dribbling faucet with extra safeguards on<br />

all the water exits of its body. This means survival in the desert,<br />

but brings problems even there and may mean death in the<br />

moisture-laden atmosphere of your garden.<br />

Plant physiologists tell us that the transpiration stream must<br />

flow unceasingly in any active plant. If the flow ceases the<br />

plant must go dormant—as some plants do in winter or severe<br />

drought—or die. They also tell us that the rate of transpiration<br />

is directly proportional to the rate of the plant’s life processes,<br />

including its growth.<br />

Relating this to cacti, we find that, having to restrict their<br />

transpiration to a minimum to conserve their stored water, the<br />

cacti are limited thereby to very slow life processes and growth<br />

as compared with other plants. When looking at a large old<br />

cactus one should appreciate the time it took, at this reduced<br />

rate, to achieve its bulk. While there is much variation, with<br />

the more extreme desert forms, naturally the most slow-growing,<br />

the variations exist even in these according to their immediate<br />

situation. It is often said that a saguaro cactus one foot<br />

tall will be about twenty-five years old, and a barrel cactus one<br />

foot in diameter between twenty and forty years old. I have in<br />

my own garden a fine specimen of Echinocactus ingens, a perfect<br />

ball just over twelve inches in diameter, which was planted<br />

at the old Shiner Cactus Garden in Laredo, Texas, as a seed,<br />

forty-five years ago. It has grown its whole life in a good<br />

situation in a plant bed, and while the species might grow<br />

somewhat faster in its native haunts in Mexico, its growth has<br />

been very nearly typical. Nor is a smaller cactus necessarily<br />

younger. I have heard it said that a peyote button two inches<br />

in diameter is ten years old. The rates of growth vary from<br />

species to species, but almost no cactus is over a fraction of an<br />

inch tall at the end of its first year, and they all must have<br />

long periods of time to achieve their potential size.<br />

The amount of water vapor in the atmosphere around a plant<br />

also affects the amount of water it will transpire. The dry air of<br />

arid regions literally drags the water out of the plants, and<br />

desert species, including cacti, have to guard their water ardently<br />

against this evaporative pull. On the other hand, the waterloaded<br />

air of humid regions is reluctant to take up more water,<br />

and the plants living in the regions must lay themselves wide<br />

open in order to promote the life-giving flow and often live


what is a cactus? 7<br />

less than the maximum span because of their inability to tran-<br />

spire enough water.<br />

Imagine the problem, then, of that cactus you brought home<br />

from the desert and planted in your nice moist yard. Adapted<br />

to hoarding its water, with its stomata small and guarded by<br />

various means against the pull of the dry desert air, what can<br />

it do in the humid air where it is now? It cannot open and lay<br />

its moisture out for the humid air. This is not its way. And<br />

since this air does not pull water out of its deep recesses, the<br />

flow will be less than it would be in the desert, so your cactus,<br />

which you thought would respond with prodigious growth to<br />

your kindness in bringing it into the moisture, may actually<br />

suffer and grow poorly because it cannot transpire and carry<br />

on its life processes here.<br />

We have also mentioned the adaptations of the cactus which<br />

enable it to live in the extreme heat and light of the desert.<br />

These may also bring severe problems to a cactus.<br />

The flesh of most cacti is more or less shaded and protected<br />

by the spines and a covering of wool or hair. In the forms<br />

which grow in extremely exposed places this covering may be<br />

developed to protect against temperatures of well over 100<br />

degrees Fahrenheit and some of the most intense light radiation<br />

found on earth. The outer tissues themselves, and even the life<br />

processes of some of these species are adapted to such extremes.<br />

It seems, for instance, that some of these cacti cannot even begin<br />

photosynthesis until the temperature reaches 75 degrees or<br />

higher. Everything in them is adjusted for high heat and light<br />

intensities.<br />

All this is fine and necessary in the desert, but what about<br />

the problems of such a cactus when you bring it home? You<br />

want to keep it in your nice cool dark house or your nice shaded<br />

garden, and you can’t understand why it does not grow, or why,<br />

if it does, it becomes all grotesque and spindly. Don’t you see that<br />

in such a situation it cannot get light and heat enough through<br />

all of its defenses to activate its high-set thermostat and stimulate<br />

its growth processes properly? Air-conditioning has marked<br />

the end of many a cactus dish-garden because the plants can<br />

hardly carry on photosynthesis, in the coolness we maintain,<br />

even in a window, and when this is coupled with what is for<br />

them little better than darkness, they may not be able to manufacture<br />

enough food even to stay alive.<br />

We have seen some of the remarkable adaptations which<br />

make cacti so fascinating and have tried to understand the<br />

problems these changes are meant to meet, as well as the special<br />

problems they can generate for cactus-growers. Successful cactus<br />

culture consists of recognizing these problems and helping<br />

the cactus meet them naturally. One does not have success with<br />

cacti by removing them from all their natural problems. More<br />

cacti in cultivation have been killed by too much kindness than<br />

by anything else. These are tough plants by nature, and all they<br />

ask is that the conditions around them remain within the ranges,<br />

rather severe by our standards, for which they are adapted.<br />

It is well to recall that there are cacti adapted for almost<br />

every sort of environment, from shady rain forest to extreme<br />

desert exposure. All of the problems of growing cacti mentioned<br />

above are less critical for those plants adapted to less severe<br />

conditions, and these can be grown much more easily than the<br />

more restricted ones; some of them can be treated quite a lot<br />

like other plants. But these are generally the less spiny, less succulent<br />

forms which are, therefore, the ones less fascinating to<br />

most of us. It is perhaps unfortunate, but unavoidable, that to<br />

grow pitahayas and barrels and other remarkable types one<br />

has to simulate to a fair degree the extreme environments of<br />

their hot, arid homes.


Key to the Genera of the Cacti<br />

The keys which are given here and before the discussion of<br />

each major genus are based as far as possible on the vegetative<br />

characters of adult individuals, but it appears to be impossible<br />

to construct workable keys for the cacti based on these alone.<br />

It was found necessary to refer in some cases to the flowers or<br />

fruits and sometimes even to the seeds. This means that the keys<br />

will be less than satisfactory in certain seasons and will not<br />

identify most juvenile forms at all. Those using the keys will<br />

need to have before them very nearly complete adult specimens<br />

of the living plants.<br />

The keys are artificial and are my own. They are binomial<br />

keys, presenting a series of choices between two alternatives. In<br />

use, one reads the first choice (1 a) and compares the specimen<br />

in question with the description. If the description at 1 a fits<br />

the specimen, he then proceeds to the number given at the end<br />

of the description and repeats the process there. If the description<br />

at 1 a does not fit the specimen, then the user abandons 1 a<br />

and moves on to 1 b, which is the alternate choice. If the specimen<br />

matches 1 b, then he moves to the number given at the<br />

end of 1 b, and so on. When this process is followed carefully<br />

with a mature plant from our area, it should lead to a description<br />

after which a plant name and page number is given. This<br />

is the name of the specimen in hand and the discussion of it will<br />

start on that page. If at any point the specimen does not fit<br />

either the a or b choice one has arrived at, there are two possible<br />

explanations. Either the user has already made a wrong<br />

choice somewhere earlier in the process, or else the cactus he has<br />

is not included in this key. Careful reconsideration of all choices<br />

should show which is the case. If the user cannot choose between<br />

alternatives by studying his specimen, he may have to secure a<br />

more mature or more complete example in order to key the<br />

form.<br />

1a. Stems of mature plants ribbed-that is, the surfaces of the stems<br />

covered with vertical or sometimes spiraling ridges which may<br />

be completely uninterrupted, undulate, or sometimes almost completely<br />

interrupted by grooves between the areoles, but which are<br />

never, on mature stems, rows of completely separate tubercles—2.<br />

2a. Plants possessing spines—3.<br />

3a. Stems of plants upright, prostrate, or clambering, with mature<br />

stems more than twice as long as they are thick; the<br />

flowers produced on the sides of the stems, with the ovary<br />

surfaces spiny; the fruit remaining fleshy and indehiscent or<br />

sometimes splitting open laterally—4.<br />

4a. Stems not more than about 6 times as long as they are<br />

thick, not over 24 inches long, upright or prostrate, but<br />

not clambering, often caespitose; the flowers produced<br />

from a rupture of the stem epidermis just above an areole<br />

—Genus Echinocereus (see key on page 11).<br />

4b. Stems when mature at least 8 to sometimes 100 times as<br />

long as they are thick, on old specimens becoming more<br />

than 24 inches long, upright or clambering, never caespi-<br />

tose; the flowers produced from within a spine areole—5.<br />

5a. Stems 1/4 to 1 inch in diameter; spines 1/32 to 1/4 of an<br />

inch long; roots tuberous—6.<br />

6a. Stems 1/4 to 5/8 of an inch thick; ribs 8; spines to 1/4<br />

or 3/8 of an inch long; roots clustering tubers; flowers<br />

purplish with short tubes and opening during the day<br />

—Genus Wilcoxia (see page 55).<br />

6b. Stems 1/2 to 1 inch thick; ribs 3 to 6; spines 1/32 to 1/8<br />

of an inch long; root a single extremely large taproot;<br />

flowers mostly white with long tubes and opening at<br />

night —Genus Peniocereus (see page 57).<br />

5b. Stems 2 to 4 inches in diameter; spines 3/8 to 2 inches<br />

long; roots fibrous—Genus Acanthocereus (see page 60).<br />

3b. Stems of plants upright, never more than twice as tall as<br />

they are thick; the flowers produced at the apex of the stem,


key to the genera of the cacti 9<br />

with the ovary surface scaly or sometimes with hair, but<br />

never spiny; the fruit opening basally or laterally<br />

—Genus Echinocactus (see key on page 65).<br />

2b. Plants spineless—7.<br />

7a. Flowers large and yellow with red centers; ovary<br />

scaly —Genus Echinocactus (see key on page 65).<br />

7b. Flowers small and pinkish; fruit never having spines<br />

or scales —Genus Lophophora (see page 95).<br />

lb. Stems of plants smooth or else tubercled—that is, covered with<br />

nipple-like projections which may be arranged in spiral rows and<br />

which may overlap due to their length, but which are never con-<br />

fluent to form raised ribs—8.<br />

8a. Plants a fraction of an inch to about 6 inches<br />

tall; stems depressed, hemispherical, or columnar,<br />

but never jointed; the spines straight or hooked<br />

but never barbed glochids; the ovaries and fruits<br />

naked or with only a few scales on them—9.<br />

9a. Plants spineless —Genus Ariocarpus<br />

(see page 100).<br />

9b. Plants spiny—10.<br />

10a. Fruit becoming dry and splitting open<br />

—Genus Pediocactus (see page 103).<br />

10b. Fruit remaining fleshy and not splitting<br />

open—11.<br />

11a. Flowers produced in the axils of the<br />

tubercles —Genus Mammillaria<br />

(see key on page 112).<br />

11b. Flowers produced from the tips of the<br />

tubercles —Genus Epithelantha<br />

(see page 107).<br />

8b. Plants several inches to sometimes 6 or more feet<br />

tall; stems jointed; at least some of the spines<br />

barbed glochids; the ovary naked or spiny<br />

—Genus Opuntia (see key on page 162).


Genus Echinocereus Engelmann<br />

The Echinocerei make up one of the largest genera of cacti,<br />

both in number of different species and in number of individuals<br />

found growing in the area of this study. Many of its members<br />

are collected and grown by cactus fanciers all over the<br />

world as great favorites because of the beauty of their flowers<br />

as well as of the plants themselves.<br />

The name of the genus is composed of two words: echinos,<br />

meaning spiny, which refers to the very spiny covering of the<br />

typical members of this genus, and cereus, which means “wax<br />

candle,” a reference to the stately appearance of the stems of<br />

the upright species.<br />

The Echinocerei are oval, conical, or cylindrical cacti, always<br />

with ribbed stems. The vertical ribs of some species are more or<br />

less divided into swellings which may be called warts or tubercles,<br />

but these are never completely separated from one another<br />

as in some other genera, so the ribs are always an outstanding<br />

character of them all.<br />

These cacti are usually very spiny, as their name implies, and<br />

these spines may be straight or curved, but are never hooked, as<br />

is common in some other groups.<br />

The stems of Echinocerei are always low as compared with<br />

many of their relatives. Most of them are well under twelve<br />

inches long when mature, and the few in the Southwest which<br />

sometimes surpass that do not usually exceed twenty-four inches<br />

long. These stems are erect in most species, but in a few they lie<br />

partly or entirely prostrate upon the ground.<br />

The plant body of some species remains a single, unbranched<br />

stem throughout life. Others cluster or branch sparingly only<br />

when very old; but many regularly form clusters of stems almost<br />

from the start. In some these clusters are made up of only<br />

a few stems, but in a few of them one plant may with age become<br />

a huge, caespitose clump of as many as a hundred or more<br />

stems, These stems are never divided into joints however.<br />

The flowers of this genus are borne on the ribs at the spinebearing<br />

areoles, developing just above the uppermost spines of<br />

the areoles, where they literally burst through the epidermis of<br />

the stem. They may be produced from almost any point on the<br />

stem, different species bearing them high or low, but most commonly<br />

they appear on the sides of the stems a little below the<br />

tips.<br />

The flowers are usually very large and beautiful, so beautiful<br />

that many fanciers pick one or another species in this genus as<br />

the most beautiful of our native cacti. However, a few of the<br />

Echinocerei have small and inconspicuous greenish flowers.<br />

The petals of some species remain only partly open, making the<br />

flowers funnel-shaped, while those of others open very widely.<br />

A perianth tube is always present. The outer surface of the ovary<br />

is always spiny and sometimes woolly as well. The stigma lobes<br />

are always green on all of our species.<br />

The fruits produced by these cacti are always fleshy, thinskinned,<br />

and often edible. Those of some species are considered<br />

delicacies. Something of their character may be imagined from<br />

the fact that a number of them are known by the common name<br />

of “strawberry cacti.” These fruits are also spiny, but the spines<br />

become loosened as the fruits mature, and may be easily brushed<br />

off.<br />

The members of the genus Echinocereus inhabit a wide belt of<br />

the North American continent from Utah and Wyoming south<br />

throughout most of northern Mexico to a little beyond the latitude<br />

of Mexico City, and from central Oklahoma and Texas on<br />

the east to the Pacific on the west. ‘Within this huge area more<br />

than eighty species have been described by various authorities,<br />

but many of these so intergrade that later students have combined<br />

various ones. The result is that almost every book or article on<br />

this genus has at least a slightly different method of listing them,<br />

depending upon the taxonomic philosophy of the writer as well


genus Echinocereus engelmann 11<br />

as upon his knowledge of these cacti. This has caused much confusion,<br />

and makes it necessary for us to deal with many authors<br />

and names in order to know exactly what plant we have before<br />

us.<br />

I will attempt to make or follow no formal classification of<br />

the species within the genus because it is too large a group and<br />

we have in the area of this study only a minority of the forms<br />

which would have to be considered in such a classification. Instead,<br />

I will group the species in a purely artificial series according<br />

to their most obvious characteristics, while concentrating<br />

upon describing the various forms properly and calling<br />

them, in the light of the most recent knowledge, by their proper<br />

names.<br />

The Echinocerei grow mostly in exposed places on dry slopes<br />

and hills in the full strength of the southwestern sun. Only a few<br />

of them prefer the shade of bushes and trees. With this sunloving<br />

characteristic and their inability to tolerate excess moisture<br />

remembered, most of them are rather easily cultivated and<br />

are, therefore, popular among collectors. They have a wide range<br />

of tolerance of cold, those of the north being very resistant,<br />

while many of those of Mexico perish by freezing when brought<br />

farther north.<br />

KEY TO <strong>THE</strong> ECHINOCEREI<br />

1a. Stems upright or sprawling, comparatively thick, being two<br />

inches or more thick when mature—2.<br />

2a. Having all three of the following characters: areoles always<br />

1/2 inch or less apart; ribs always more than 10; radial spines<br />

always more than 12—3.<br />

3a. Flowers small, 1 to 11/2 inches long and less than that in<br />

width, yellow, yellow-brown, or pinkish-red in color—4.<br />

4a. Areoles elongated; central spines 0 to 3 in number, arranged<br />

in a vertical row, and 1 inch or less in length on mature<br />

plants—5.<br />

5a. Plant globose to short-cylindric; radials 1/4 of an inch or<br />

less long—6.<br />

6a. Ribs 12 to 18; radials 16 to more than 20; plant 2<br />

inches or more tall when mature—7.<br />

7a. Spines varicolored red, brownish, or purplish-red<br />

and white —E. viridiflorus var. viridiflorus.<br />

7b. Spines yellowish —E. viridiflorus var. standleyi.<br />

6b. Ribs 6 to 9; radials 8 to 12; plant about 1 inch tall<br />

when mature —E. davisii.<br />

5b. Plant cylindrical; radials to 1/2 inch long<br />

—E. viridiflorus var. cylindricus.<br />

4b. Areoles broad oval to round; centrals 3 to 12 in number<br />

and not standing in a straight vertical row, but instead<br />

spreading outward from the center of the areole—8.<br />

8a. Radials 12 to 23; ribs hardly tuberculate; centrals<br />

3 to 5 in number on mature plants<br />

—E. chloranthus var. chloranthus.<br />

8b. Radials 30 or more; ribs markedly tuberculate;<br />

centrals 5 to 12 on mature plants—9.<br />

9a. Centrals to only 1/2 inch long; immature plants<br />

having flexible hairs instead of spines; flowers<br />

greenish-yellow to bronze and with a glossy<br />

surface —E. chloranthus var. neocapillus.<br />

9b. Centrals 3/4 to 11/4 inches long; immature<br />

plants having rigid spines from the first; flowers<br />

pale pinkish or brownish-red and with a<br />

dull surface —E. russanthus.<br />

3b. Flowers large and showy, 2 to 5 inches or more in length<br />

and width, yellow to purple in color—10.<br />

10a. Flower tube with long, cobwebby wool and hairlike,<br />

bristly spines; spines not recurved against the plant<br />

body—11.<br />

11a. Longest radial spines 3/8 of an inch or less—12.<br />

12a. Central spines 0 to 2, porrect and if more than<br />

one, then arranged in a strict vertical row—13.<br />

13a. Centrals usually missing and 1/8 inch or less long<br />

when present—14.<br />

14a. Size of mature plant stems 3 to 12 inches tall<br />

by 2 to 31/2 inches thick; flower 3 to 5 inches<br />

long and nearly as wide, petals 30 to 50,<br />

stigma lobes 8 to 22—15.<br />

l5a. Radial spines 15 to 36—16.<br />

16a. Spines white to gray or reddish, often<br />

with dark tips, but not with the outer<br />

parts of the spines conspicuously shiny<br />

purplish or black so as to give the plant<br />

a shiny blackish aspect —E. caespitosus<br />

var. caespitosus.<br />

16b. Outer parts of the spines bright, shiny,<br />

purplish or black, giving the plant a con-<br />

spicuous blackish appearance<br />

—E. caespitosus var. purpureus.<br />

15b. Radial spines 12 to 15 —E. caespitosus<br />

var. perbellus.<br />

14b. Size of mature plant stem to only 3 inches<br />

tall and 1 inch thick; flowers small, to only<br />

2 inches long and 13/4 inches wide, petals to<br />

only 20 in number and stigma lobes to only 8<br />

—E. caespitosus var. minor.<br />

13b. One black central 3/16 to 1/4 of an inch long always<br />

present —E. melanocentrus.<br />

12b. Centrals 3 to 7 and not in a strict vertical row,<br />

but spreading —E. fitchii.<br />

11b. Longest radial spines 3/8 to 1 inch—17.<br />

17a. Stems usually caespitose; areoles more than 1/8 of<br />

an inch long, having no central spines over 3/8 of<br />

an inch long—18.<br />

18a. Radials 5/8 to 1 inch long; flower rose-red<br />

—E. baileyi.<br />

18b. Radials 3/16 to 1/2 inch long; flower pale pinkish<br />

—E. albispinus.<br />

17b. Stems usually simple; areoles 1/8 of an inch or less<br />

long, having at least the main central spines 3/8 of<br />

an inch or more long on mature areoles<br />

—E. chisoensis.


12 cacti of the southwest<br />

10b. Flower tube with short wool and rigid spines—19.<br />

19a. Spines of the plant body strictly pectinate and recurved<br />

against the plant body; areoles oval to elon-<br />

gated—20.<br />

20a. Centrals 2 to 3 in a vertical row; radials slender<br />

to medium stout, white to purplish or pinkish and<br />

the plant often banded with color, but spines of<br />

individual areole not variegated—21.<br />

21a. Flower purple with white zone and green center<br />

—E. pectinatus var. wenigeri.<br />

21b. Flower orange-yellow with green center<br />

—E. pectinatus var. ctenoides.<br />

20b. Centrals none; radials stout; spines of individual<br />

areoles variegated grays or tans and red<br />

—E. pectinatus var. rigidissimus.<br />

19b. Spines of plant body spreading outward instead of<br />

being pectinate; areoles oval to round—22.<br />

22a. Radial spines 15 to 25; areoles 1/8 to 3/8 of an inch<br />

apart; flower 3 to 51/2 inches long—23.<br />

23a. Radials 16 to 25 and to 1/2 inch long; plant<br />

simple or sparingly branched —E. dasyacanthus<br />

var. dasyacanthus.<br />

23b. Radials 15 or 16 and to only 3/8 of an inch long;<br />

plant caespitose —E. dasyacanthus<br />

var. hildmanii.<br />

22b. Radial spines 10 to 15; flower 2 to 3 inches long;<br />

areoles 5/16 to 1/2 inch apart —E. roetteri (in part).<br />

2b. Never having all three of the characters listed under 2a or, in<br />

other words, having any one or more of the three following<br />

characters: areoles more than 1/2 inch apart; ribs less than 10;<br />

radials less than 12—24.<br />

24a. Flowers scarlet-red and lasting up to 4 or 5 days, with the<br />

petals firm and their edges entire—25.<br />

25a. Ribs 5 to 9; radial spines 2 to 9; centrals 0 or 1—26.<br />

26a. Spines greatly flattened and usually channeled or furrowed—27.<br />

27a. Spines extremely heavy—28.<br />

28a. Central spine absent; radials 2 to 6; areoles 7/8 to<br />

11/2 inches apart —E. triglochidiatus<br />

var. triglochidiatus.<br />

28b. One central present in at least part of the areoles;<br />

radials 6 to 8; areoles 1/4 to 3/4 of an inch apart<br />

—E. triglochidiatus var. gonacanthus.<br />

27b. Spines slender to medium thickness; radials 5 to 7;<br />

centrals 0 to 1 —E. triglochidiatus var. hexaedrus.<br />

26b. Spines round or practically so and slender to medium<br />

in thickness —E. triglochidiatus var. octacanthus.<br />

25b. Ribs 7 to 15; radial spines 7 to 16; centrals 1 to 6—29.<br />

29a. Plant clustering densely into dome-shaped masses of<br />

equal, short stems not usually over 6 inches tall—30.<br />

30a. Centrals 1 to 5; areoles 3/16 to 3/8 of an inch apart;<br />

largest central round —E. coccineus var. coccineus.<br />

30b. Centrals 3 to 5; areoles 5/16 to 5/8 of an inch apart;<br />

largest central flattened —E. coccineus<br />

var. conoideus.<br />

29b. Plant clustering sparingly to form flat clumps of un-<br />

equal stems up to 18 inches tall—31.<br />

31a. Centrals several—32.<br />

32a. Ribs 8 to 11; radials 7 to 12; centrals usually 3 or<br />

4 and variable in length from 3/4 to over 2 inches<br />

long; all spines round and medium thickness to<br />

heavy, whitish to ashy purplish-gray or reddish in<br />

color—33.<br />

33a. Ovary tube with long, flexible hair<br />

—E. polyacanthus var. polyacanthus.<br />

33b. Ovary tube with sparse and short wool<br />

—E. polyacanthus var. rosei.<br />

32b. Ribs 11 to 15; radials 8 to 16; centrals 4 to 6;<br />

spines round and white, yellow, or reddish-yellow<br />

in color —E. polyacanthus var. neo-mexicanus.<br />

31b. Central 1 —E. mojaviensis.<br />

24b. Flowers purple or yellow with red centers and delicate, last-<br />

ing only 1 or 2 days, with petals soft and more or less<br />

emarginate—34.<br />

34a. Spines opaque; flesh medium to dark or gray-green—35.<br />

35a. Mature spines opaque and varicolored or variegated<br />

with shades of brown, gray, and white streaking at<br />

least some of them; radials 5 to 12; centrals 1 to 3—36.<br />

36a. Stems more or less flaccid, wrinkled, with broad,<br />

somewhat tuberculate ribs; central spine 1, long and<br />

curving upward —E. fendleri var. fendleri.<br />

36b. Stems firm and not wrinkled; ribs narrow and not<br />

markedly tuberculate; central spines 1 to 3, the main<br />

one porrect and straight —E. fendleri<br />

var. rectispinus.<br />

35b. Mature spines opaque but not variegated or varicolored;<br />

radials 14 to 17; centrals 2 to 8—37.<br />

37a. Lowest radials as long as the lateral ones in the areole;<br />

fruits 11/4 to 2 inches long —E. lloydii.<br />

37b. Lowest radials shorter than the laterals; fruits 1/2 to<br />

7/8 of an inch long —E. roetteri (in part).<br />

34b. Spines translucent to some degree; flesh light or medium<br />

green—38.<br />

38a. Areoles 1/2 to 11/2 inches apart; ribs only slightly tuberculate<br />

on mature stems; flowers purple—39.<br />

39a. Ribs 11 to 13; areoles 1/2 to 3/4 of an inch apart;<br />

plant forming a large, regular, hemispherical clump<br />

from a single root center; flowers 4 to 5 inches long<br />

—E. stramineus.<br />

39b. Ribs 7 to 10; areoles 3/4 to 11/2 inches apart; plant<br />

forming an irregular, sprawling, or prostrate clump,<br />

often with adventitious roots; flowers 11/2 to 3 inches<br />

tall—40.<br />

40a. Stems extremely flabby; becoming prostrate and<br />

to 30 inches long; flower with 20 to 35 inner pet-<br />

als in several series and 10 to 12 stigma lobes<br />

—E. enneacanthus var. carnosus.<br />

40b. Stems sprawling, more or less flabby, and to 15<br />

inches or so in maximum length; flower with 10<br />

to 15 inner petals in one row and 8 to 10 stigma<br />

lobes—41.<br />

41a. Stem to 23/4 inches thick, comparatively firm<br />

and upright; radials 1/4 to 5/8 of an inch long<br />

and straight; areoles 1/4 to 1 inch apart<br />

—E. enneacanthus var. enneacanthus.<br />

41b. Stems 3 to 4 inches in diameter, flabby, and


genus Echinocereus engelmann 13<br />

semiprostrate; radials 3/4 to 11/2 inches long,<br />

often curving; areoles 1 to 11/2 inches apart<br />

—E. dubius.<br />

38b. Areoles 3/8 to 1/2 inch apart; ribs extremely tuberculate;<br />

flowers yellow with red centers —E. papillosus<br />

var. papillosus.<br />

lb. Stems prostrate and slender, being 1/2 to 11/2 inches thick—42.<br />

42a. Central spine present on all or most areoles and 3/8 to 2 inches<br />

long—43.<br />

43a. Stems to only 4 inches long; central spine 3/8 of an inch<br />

long; flower yellow with red center; ribs markedly tuber-<br />

culate —E. papillosus var. angusticeps.<br />

43b. Stems 6 to 14 inches long; central spine 1/2 to 2 inches long;<br />

flower purple—44.<br />

44a. Central spine 1/2 to 2 inches long, dark in color, and<br />

somewhat aimed and curved downward; flower with a<br />

dark reddish-purple throat and narrow, pointed petals<br />

—E. blanckii.<br />

44b. Central spine 1/2 to 11/2 inches long, yellowish-brown,<br />

porrect or turning upward; flower with a white throat<br />

—E. berlandieri (in part).<br />

42b. Central spine usually missing, and if present on occasional<br />

areoles only 1/4 of an inch or less in length—45.<br />

45a. Radial spines 4 to 6 and some of them 1/4 to 11/4 inches<br />

long; flower with white throat and narrow, pointed petals<br />

—E. berlandieri (immature or stunted growth form).<br />

45b. Radial spines 3 to 6 and only 1/16 to 1/4 of an inch long;<br />

flower with white throat and broad, blunt-tipped petals<br />

—E. pentalophus.<br />

Echinocereus viridiflorus Eng.<br />

“Green-Flowered Torch Cactus,” “Green-Flowered Pitaya,”<br />

“Nylon Cactus,” “New Mexico Rainbow Cactus”<br />

Description Plate 1<br />

stems: Single or forming small clusters of up to about half a<br />

dozen heads. Each stem is spherical to columnar, varying in<br />

the different forms from a sphere or cone only an inch or so<br />

across to a column as much as 8 inches tall by as much as 3<br />

inches in diameter. The surface is light green to yellowishgreen<br />

in color. There are 13 to 15 low ribs with shallow vertical<br />

grooves between them. There are definite grooves crossing<br />

the ribs between the areoles, giving them a somewhat tuberculate<br />

appearance.<br />

areoles: Small, narrow oblong to very elongated, up to<br />

about 3/8 of an inch apart on mature parts of the stem. Young<br />

areoles are covered with short, white felt, but old areoles lose<br />

this and become bare except for a small tuft of wool just<br />

above the spines of each areole where a flower has developed.<br />

spines: There are 12 to more than 20 radial spines which<br />

radiate evenly around the areole. They are straight and rigid,<br />

and lie flat upon the surface of the plant or sometimes recurve<br />

back toward the grooves between the ribs. The longer<br />

ones sometimes interlock with those of the adjacent areoles.<br />

They vary in size from very small, very weak upper ones<br />

which are bristle-like and only 1/16 of an inch long to lateral<br />

and lower ones 1/4 of an inch long in one variety and 1/2 inch<br />

long in another. There is great variation in the coloring of<br />

these radial spines. They may all be purplish-red or yellow or<br />

whitish, but the most typical pattern is for the upper and<br />

lower ones in each areole to be white while the lateral ones<br />

are reddish—although an occasional specimen displays exactly<br />

the reverse pattern. Sometimes individual spines may<br />

be white, tipped with red. All of these variations may sometimes<br />

occur on the same plant, often in zones, giving the plant<br />

a banded appearance.<br />

There is most typically one central spine, much thicker and<br />

more rigid than the radials and standing erect in the center of<br />

the areole. This spine sometimes is curved upward toward its<br />

tip and is usually about 1/2 inch long, although in one form<br />

it may be up to 1 inch long. This spine may be white with a<br />

reddish tip, half and half, or all purplish-red except for a<br />

white base. Sometimes one or even two auxiliary central<br />

spines, much shorter (only 1/16 to 3/16 of an inch long) but<br />

otherwise identical to the first, may be present, in which case<br />

the two or three centrals are arranged in a perfect vertical<br />

row. On the other hand, the areoles of many plants lack the<br />

central entirely. All the spines have bulbous bases.<br />

flowers: Small, lemon-yellow or straw-color, this often<br />

approaching chartreuse or occasionally being suffused with<br />

brown and then bronzy. These flowers are about 1 inch long<br />

by 3/4 of an inch in diameter, produced on old areoles on the<br />

sides of the stem, usually midway between the base and the<br />

top although sometimes even lower. The outer petals are<br />

linear, brownish in the midline with lemon-yellow or chartreuse<br />

edges. The inner petals are longer and become a little<br />

broader toward the tips which are more or less rounded.<br />

These inner petals are lemon-yellow to straw, usually with<br />

somewhat darker green in the midline. The edges are all entire.<br />

The stamens show the same colors. The style is somewhat<br />

longer than the stamens and crowned by 6 to 10 dark<br />

green, rather fat stigma lobes. Each areole of the ovary has<br />

short white wool and 4 to 12 white spines up to 1/4 of an<br />

inch long.<br />

fruits: These are 3/8 to 1/2 of an inch long, egg-shaped, and<br />

greenish in color, and have white wool and white spines upon<br />

them.<br />

Range. From eastern Wyoming and eastern Colorado south<br />

through eastern New Mexico to the vicinity of El Paso, Texas,<br />

and southeast through the Guadalupe Mountains into the Big<br />

Bend of Texas.<br />

Remarks. E. viridiflorus is famed as the most northerly of the<br />

Echinocerei. It grows on the bleak prairies and the foothills of<br />

eastern Wyoming and Colorado in spite of cold and extreme<br />

conditions which would kill most cacti. A few times it has been


14 cacti of the southwest<br />

reported from extreme western Kansas and the Panhandle of<br />

Oklahoma, but these reports were mostly fifty or more years<br />

ago. If it is not extinct in those areas, it is very rare today. It<br />

still may be found occasionally in the Texas Panhandle and in<br />

northeastern New Mexico, which is its type locality. The most<br />

westerly report of it seems to be just a few miles west of Santa<br />

Fe, New Mexico.<br />

Over this whole northern area of its range the cactus is a<br />

small, squat, egg-shaped, withdrawing plant, hiding as best it<br />

can in the sparse grass and practically invisible during the winter<br />

when it is greatly shrunken. The whole plant is seldom over<br />

2 or 3 inches tall in this area. Neither does it produce conspicuous<br />

flowers to give away its position in the spring. The flowers<br />

are small and greenish-yellow, and even these are borne low on<br />

the plant rather than at the top. These northern plants usually<br />

have long, up-curving central spines, but their radials tend to<br />

be shorter than those of their southern relatives. This northern<br />

form seems to he the plant Engelmann had before him which<br />

he called E. viridiflorus var. minor. He stated correctly that<br />

this variety grows from around Santa Fe, New Mexico, northeastward.<br />

South of Santa Fe the species is seen again past Socorro and<br />

Roswell, New Mexico, into the Guadalupe Mountains and at<br />

El Paso. It then becomes common east and south into the Big<br />

Bend area of Texas. But all of the southern plants studied show<br />

some differences from their northern relatives. These differences<br />

have been considered significant enough from the very<br />

beginning to warrant separate variety designations. Much of<br />

the confusion in the minds of cactophiles and, therefore, in their<br />

accounts of this species and the closely related species Echinocereus<br />

chloranthus has been due to a failure to distinguish and<br />

understand these varieties. The southern forms of E. viridiflorus<br />

are quite commonly displayed and even sold today under the<br />

name of E. chloranthus.<br />

Echinocereus viridiflorus var. viridiflorus (Eng.)<br />

Description Plate 1<br />

stems: As the species except that it is spherical to conical,<br />

growing occasionally to about 5 inches high by as much as 3<br />

inches in diameter, but usually much smaller. Usually clustering.<br />

areoles: As the species.<br />

spines: As the species, except that the centrals are often missing<br />

entirely and never observed over 1/2 inch long.<br />

flowers: AS the species, except that they are usually lemonyellow<br />

in color and more rarely brownish than in the other<br />

forms, with the petals somewhat rounded at the ends.<br />

fruits: As the species.<br />

Range. From eastern Wyoming and eastern Colorado through<br />

eastern New Mexico to the vicinity of Santa Fe, which seems<br />

to be the southern and western limit of its range. It occurs from<br />

there east into the Oklahoma and Texas panhandles and is<br />

found occasionally in the high plains of extreme northwestern<br />

Texas, but this form does not appear to come down into south-<br />

west Texas.<br />

Remarks. This is the more hardy northern form of the Species.<br />

Small and comparatively insignificant as it is, anyone who has<br />

been abroad in its range would be amazed to know how many<br />

specimens he has stepped over as it hides in the grass. It takes<br />

a real Search to locate the plant, but it can easily be grown in<br />

northern gardens where most other cacti do not survive. With<br />

its clustering habit it soon presents an attractive little clump of<br />

heads with varicolored spines.<br />

Echinocereus viridiflorus var. cylindricus (Eng.) Rümpl.<br />

Description Plate 1<br />

stems: Similar to the species except that it is cylindrical and<br />

grows to at least 8 inches tall by 3 inches in diameter. Seldom<br />

and sparingly clustering.<br />

areoles: As the species.<br />

spines: As the species, except that the radials number 14 to<br />

24, their maximum length to 1/2 inch; and the centrals number<br />

0 to 3 in a vertical row, the main central growing to a<br />

maximum length of 1 inch on some plants. On many plants,<br />

however, the main central may be entirely missing or no<br />

longer than the 1/16 to 3/16-inch auxiliary centrals.<br />

flowers: As the species, except that they are more often<br />

brownish in color and the petals are more sharply pointed.<br />

fruits: As the species.<br />

Range. Common from the area of Socorro and Roswell, New<br />

Mexico, southeast through the Guadalupe Mountains into Texas.<br />

The writer was surprised to find a stand of this variety near<br />

Mosquero, in northeastern New Mexico, which marks a great<br />

extension of its range northward. In Texas it ranges through the<br />

Davis Mountains into the Big Bend.<br />

Remarks. This is E. Viridiflorus as it appears in its southern<br />

range, a much more robust form, with radial spines about twice<br />

as long as and more interlocking than those of the northern<br />

form. Its central spines are extremely variable. Almost any collection<br />

from the Big Bend will show some plants with only five<br />

or six single centrals 3/16 to 1/8 of an inch long scattered over<br />

the whole stem, a few with no centrals at all, and others with<br />

regularly 2 or 3 centrals per areole arranged in a single vertical<br />

row. The specimens from the Hueco and Guadalupe mountains<br />

sometimes lack centrals also but more often show a single central<br />

like the species except usually to 1 inch long and curving.<br />

Thousands of these large, columnar specimens, usually dis-<br />

Rumpl -><br />

Rümpl


genus Echinocereus engelmann 15<br />

tinctly reddish of spine and bronze of flower, are shipped by<br />

dealers out of the Texas Big Bend every year. They are so obviously<br />

different from the little northern form that, since the<br />

name E. viridiflorus without variety qualification is usually applied<br />

to the northern form, these are usually distributed under<br />

the name of E. chloranthus. However, this southern variety was<br />

recognized as a distinct form long ago when it was named E.<br />

viridiflorus var. cylindricus by Engelmann. Coulter, failing to<br />

apply Engelmann’s name to it, renamed it Cereus viridiflorus<br />

var. tubulosus, basing his description on specimens from Brewster<br />

County, Texas. Both men recognized its distinctness from<br />

E. chloranthus, a distinction which has become blurred in some<br />

more recent accounts.<br />

This variety of this cactus was once one of the most common<br />

in the Big Bend region. I have been told by older collectors of<br />

beds of these cacti extending over many acres in Brewster<br />

County, Texas, in which the cacti often stood almost too close<br />

for one to walk among them. Their numbers seemed inexhaustible.<br />

There are few such stands left today, however, mostly due<br />

to the destruction wrought by the cactus dealers whose crews<br />

have for years been bringing them out by the truckloads to<br />

languish and die in dime-store bins and novelty-shop cactus<br />

assortments. I have been on some ranches in the area whose<br />

owners have allowed no cactus digging, and there they still<br />

grow in quantity; but only a few years ago I watched three<br />

big truckloads of them come out as another of these ranches<br />

was opened up and swept clean of cacti, and so they appear to<br />

be doomed to elimination as surely as the more rare species of<br />

the area.<br />

Echinocereus viridiflorus var. standleyi (B. & R.) Orcutt<br />

Description Plate 1<br />

stems: Similar to the species except becoming cylindrical and<br />

growing to about 4 inches tall, while seldom clustering.<br />

areoles: As the species.<br />

spines: As the species, except that they are a clear, rather<br />

translucent yellow, becoming sometimes whitish when old.<br />

flowers As the species.<br />

fruits: As the species.<br />

Range. South central New Mexico.<br />

Remarks. This was described as a species by Britton and Rose,<br />

but since they had not seen it flower and their description was<br />

very incomplete, it was long uncertain what the plant’s relationship<br />

really was. After having seen numerous specimens<br />

throughout their development and in bloom, I feel certain that<br />

it is a form of E. viridiflorus, as Orcutt has already suggested.<br />

I cannot distinguish its flowers from those of the species by any<br />

essential feature. It is close to the variety cylindricus in the<br />

cylindrical shape of its stems and in the character of its centrals,<br />

but its radial spines are shorter and more like those of variety<br />

viridiflorus. This leaves it distinct from these other forms only<br />

by the clear yellow color of its spines. This is a doubtful character<br />

to base even a variety upon, and it surely will not support<br />

its being listed as a separate species. It may be that it does not<br />

even warrant varietal status, but I list it separately here since<br />

anyone reading the literature will be anxious to know what the<br />

plant referred to so incompletely by almost all authors actually<br />

is.<br />

Echinocereus davisii Houghton<br />

Description Plate 2<br />

stems: Globular or nearly so and very small. This is a dwarf<br />

plant, 1/2 to 11/4 inches tall when mature. Stems occur singly<br />

as far as is known. There are 6 to 8 ribs on this tiny stem,<br />

which are rather high and broken into nearly completely<br />

separated tubercles. The color of the flesh is dark green.<br />

areoles: Narrow oval to elongated and 1/8 to 3/16 of an inch<br />

long, having very little if any wool upon them, even when<br />

young.<br />

spines: White, with the tips or sometimes the outer one-half<br />

dark brown. These spines are all radials and are 8 to 13 in<br />

number. The upper ones are shortest, slender, round, and ‘Is<br />

to 1/4 of an inch long, while the laterals are longer, stouter,<br />

usually flattened, and 3/8 to 5/8 of an inch long. The spines<br />

are straight or often somewhat curved and recurved back<br />

against the plant.<br />

flowers: Straw-yellow, about 1 inch long, not opening<br />

widely. The outer petals are narrow with midlines reddish to<br />

umber and the edges yellowish. They are pointed and entire.<br />

The inner petals are straw-yellow, and linear to slightly<br />

broadened above, with the tips sharply pointed. The filaments<br />

are green, the anthers light yellow. The style is light<br />

green and is crowned by 5 light green, heavy, curving stigma<br />

lobes. The ovary has about 12 white spines to 1/4 of an inch<br />

long on each areole, but no wool.<br />

fruits: Oval, about 3/8 of an inch long, apparently remaining<br />

green.<br />

Range. Found on only a few limestone summits a few miles<br />

south of Marathon, Brewster County, Texas.<br />

Remarks. This is one of the two dwarf cacti of Brewster County,<br />

Texas, and it is remarkable that it grows on the same few<br />

hills as the other, Mammillaria nellieae. It even grows in the<br />

same situation as that other, underneath the moss in the crevices<br />

of the limestone ledges. The tiny plant is rarely visible at all<br />

because of its mossy protection, and only sends its flowers up<br />

briefly above this layer. It is a real hands-and-knees effort to<br />

search for these diminutive plants in the expanse of the Texas<br />

hills. The best effort will be useless unless one is on the right


Rumpl -><br />

Rümpl<br />

16 cacti of the southwest<br />

hills, and these are restricted to two or three ranches near Marathon.<br />

One would think that the little cactus would be secure in its<br />

very limited range, but that is not the case. After Houghton<br />

described this plant in 1931 and others wrote about it, collectors<br />

and dealers went straight to the spot and brought out hundreds<br />

of specimens. This continued for some years, and most collections<br />

then sported some of these attractive little novelties. In<br />

this way the wild population was reduced steadily until more<br />

recently the area in which they grow has come into the hands<br />

of owners who allow no one to dig cacti at all. This is fortunate<br />

for the survival of the species, but the plant has become more<br />

and more rare in collections, and its value on the market has<br />

now become rather high.<br />

The problem here, again, is to place this cactus properly in<br />

respect to its relatives, and as always this cannot be done with<br />

absolute certainty or to please everyone. A. D. Houghton designated<br />

it a separate species when he first described it, but W. T.<br />

Marshall soon considered its similarities to E. viridiflorus too<br />

great and called it E. viridiflorus var. davisii. Anyone may take<br />

his choice of these arrangements, since it is a matter of evaluation<br />

of similarities and differences which all can see. I choose<br />

to leave it distinct from E. viridiflorus because it differs from<br />

that plant in rib and spine number and spine length, as well as<br />

in size. Of perhaps more importance to me in deciding to regard<br />

it as a separate species are the facts that the petals of its<br />

flowers are more pointed than those of E. viridiflorus, and that<br />

it has no wool on the ovary areoles while the other does. This<br />

matter of wool or no wool has been made the basis for separating<br />

others of this genus into entirely different sections, and so it<br />

certainly seems significant enough here to separate species.<br />

Although tiny in size, E. davisii is not short-lived, as one<br />

might expect. I have seen a series of specimens collected ten<br />

years ago and grown all this while by a collector who knows<br />

how to care for her plants. They have bloomed most years, but<br />

have hardly increased in size in all this time.<br />

Echinocereus chloranthus (Eng.) Rümpl.<br />

“Green-Flowered Torch Cactus,” “Green-Flowered Pitaya”<br />

Description Plate 2<br />

stems: Cylindrical, up to 10 inches high by 3 inches thick,<br />

occasionally, but not often, producing one or two branches<br />

from the main stem. There are 12 to 18 ribs which are low<br />

and definitely tuberculate at the areoles, with broad, shallow<br />

furrows between the ribs and between the tubercles. The color<br />

of the surface is pale green.<br />

areoles: Oval to circular in shape and rather large. The<br />

young areoles have much short white or yellowish wool. This<br />

gradually disappears with age so that old areoles are almost<br />

or quite bare except for the floral areole just above the spines<br />

which remains as a persistent tuft of wool. The areoles are up<br />

to 1/4 of an inch apart on the mature sides of the stems.<br />

spines: There are 12 to 38 radial spines which radiate evenly<br />

all around the areole. They are straight and rigid, and interlock<br />

with those of adjacent areoles. They may recurve back<br />

toward the plant slightly on some specimens, but when young<br />

they spread out from the plant before they assume their<br />

strictly radiating mature position. The upper 4 to 6 of them<br />

are short and weak, 1/4 of an inch or less in length. There is a<br />

gradual increase in size going around the areole, the laterals<br />

being about 1/2 inch long and the lower ones sometimes longer,<br />

even up to 3/4 of an inch. The upper radials are usually white,<br />

while the laterals and lower ones vary greatly in color, sometimes<br />

being white, yellowish, purple-red, or even variegated<br />

with white or light bases and the rest of the spine red.<br />

There is great variation in the centrals in this species. On<br />

the typical form there are 3 to 6 centrals on mature areoles.<br />

These spread from the center of the areole and are not arranged<br />

in a straight line. They are stout and rigid, somewhat<br />

translucent, and usually somewhat curved. A typical areole<br />

on a typical mature plant will have 1 or 2 upper centrals<br />

which are only 1/4 to 1/2 inch long and point upward. These<br />

are usually red or white with red tips. Then there will usually be<br />

2 centrals spreading laterally, straight or curved, about 3/4 of<br />

an inch long and all red or red with whitish or yellowish<br />

bases. Below this is one long central pointing downward and<br />

often curved, which is almost always lighter colored, white<br />

sometimes with a reddish tip, stout, rigid, and 3/4 to 11/4 inches<br />

long. Some plants have an extra central or two besides these.<br />

Young plants do not grow their centrals at all until they are<br />

4 inches or so in height, adding them gradually after that, so<br />

that it is easy to find good-sized plants with none or with<br />

only a couple of centrals. All of the spines have bulbous bases,<br />

but the bases of the radials are usually covered, until they are<br />

very old, by the wool of the areole.<br />

flowers: Funnel-shaped, not opening widely, 1 inch in diameter<br />

by about 11/4 inches long, very dark green or yellowishgreen.<br />

The outer petals have brownish midlines and green<br />

edges. The inner petals are dark green in the midlines with<br />

lighter edges, but are often suffused with brown. The petals<br />

are linear, not broadening throughout their lengths, the edges<br />

entire, and the tips sharply pointed. The filaments are light<br />

green and the anthers cream-colored. The pistil is long, green,<br />

and ends in 8 dark green stigma lobes. The areoles of the<br />

ovary have white wool and white spines about 1/4 to 1/2 inch<br />

long upon them. These flowers are produced on the sides of<br />

the stems, usually from one-half to two-thirds of the way to<br />

the top, but sometimes lower.<br />

fruits: Small, greenish, very spiny. They are usually about<br />

1/2 inch long and almost spherical.<br />

Range. A small area of southern New Mexico in Luna and<br />

Dona Ana counties, specifically from Cook’s Peak and near


genus Echinocereus engelmann 17<br />

Rincon through the Organ and the Franklin mountains into<br />

Texas and Mexico. In Texas occurring in scattered mountain<br />

areas from El Paso east about as far as Van Horn, apparently<br />

being limited to El Paso and Hudspeth counties, except for a<br />

separate variety isolated in Brewster County.<br />

Remarks. This cactus was first described by Engelmann in 1856.<br />

Typically, it is a beautiful, tall, and slender column of long,<br />

rigid, varicolored spines, these partly obscuring the surface of<br />

the stem and spreading in all directions, giving the plant an<br />

interesting unkempt appearance in contrast to the usual precision<br />

of armament of other Echinocerei. It has small green, yellow-green,<br />

or bronze-green flowers produced on the sides of the<br />

stems, very similar to those of E. viridiflorus. This is a comparatively<br />

localized form, however, growing in the mountains from<br />

about a hundred miles northwest of El Paso to about the same<br />

distance southeast of that city.<br />

E. chloranthus is one of those cacti which are very difficult<br />

to distinguish from their relatives, and I find for more confusion<br />

among cactus fanciers about it than about any other member<br />

of this genus. Somehow the names of this and the previous<br />

species were exchanged in popular usage in west Texas many<br />

years ago, and even some dealers’ lists offer them reversed in this<br />

way. In addition, another species has been entirely lost for<br />

many years by being included under this one.<br />

The striking similarity of E. chloranthus to E. viridiflorus is<br />

the main cause of the confusion. It is easy enough to distinguish<br />

between the typical E. viridiflorus var. viridiflorus, the northern<br />

cactus, and E. chloranthus. The northern plant is very small<br />

and squat, has elongated areoles with either no central spine or<br />

only 1 per areole, and has flowers with rounded petal tips. E.<br />

chloranthus, on the other hand, is tall and slender, has up to<br />

5 spreading centrals not in a row, rounded areoles, and sharply<br />

pointed petals. These differences are plain enough, and the<br />

widely different ranges of the two, separated by almost half<br />

the length of New Mexico, makes it easy to regard them as very<br />

separate forms.<br />

But the trouble lies with the form of E. viridiflorus called<br />

variety cylindricus by Engelmann, which grows in the same<br />

range as E. chloranthus and on into the Big Bend. This variety<br />

is typically cylindrical also, but somewhat stouter than E. chloranthus,<br />

with elongated areoles and with 1 to 3 centrals in a<br />

straight row. These two are close, but they can be told apart by<br />

the shape and size of the areoles and the difference in number,<br />

length, and arrangement of the central spines on mature specimens.<br />

Immature specimens are much alike, but can be recog-<br />

nized by their areoles.<br />

After one has learned to recognize the two forms the possibility<br />

of confusion is not yet over, because the names used for<br />

them have not remained standard. Marshall and T. M. Bock<br />

have proposed the combining of E. viridiflorus and E. chloranthus<br />

entirely as synonyms. Backeberg, on the other hand, chooses<br />

to combine this form with the other as E. viridiflorus var. chlo-<br />

ranthus. Future study may bring a combination of some sort,<br />

but there can be no more than conjecture about what the proper<br />

relationship of the two is, without new evidence, which is not<br />

yet at hand. In the meantime, it seems, any conscientious student<br />

of cacti will have to look very carefully at areoles and spines<br />

and come to know this interesting little cactus.<br />

Echinocereus chloranthus var. chloranthus (Eng.)<br />

Description Plate 2<br />

stems: As the species, except growing to only about 8 inches<br />

tall.<br />

areoles: As the species, except to only about 3/8 of an inch<br />

apart.<br />

spines: As the species, except always spiny and never hairy,<br />

the radials only 12 to 23 in number, and the centrals only 3<br />

to 6 in number.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. Extreme southern New Mexico in Luna and Dona Ana<br />

counties, into El Paso and Hudspeth counties, Texas, and on<br />

into Mexico.<br />

Remarks. This is the typical form of the species, but it has a<br />

restricted range, and far fewer people have seen it than suppose<br />

they have.<br />

Echinocereus chloranthus var. neocapillus Weniger<br />

Description Plate 2<br />

stems: Cylindrical, up to 10 inches high and 21/2 inches thick.<br />

It is usually single, but occasionally branches above the<br />

ground. It has 12 to 18 ribs, low, with distinct tubercles. The<br />

color is pale or yellowish-green.<br />

areoles: Oval, about 3/16 of an inch long, covered with much<br />

white or yellowish wool when young, bare when old except<br />

for a very small tuft which usually remains at the upper end<br />

of the areole where the flower is produced. The areoles are<br />

up to 1/4 of an inch apart.<br />

spines: There are 30 to 38 straight, slender radial spines.<br />

These radiate evenly around the areole, being often so crowded<br />

that they touch their neighbors most of their lengths, and<br />

they interlock with spines from other areoles. The upper ones<br />

are very slender and often only 1/8 of an inch long. From<br />

this the length increases around the areole, with laterals being<br />

up to 1/2 inch long and the lower ones only slightly shorter.<br />

They are clear translucent yellow or chalk-white, all of an<br />

areole being the same color, this usually forming bands of<br />

yellow and white on the older plants. There are 5 to 10 central<br />

spines, all heavier and straight. They spread in all direc-


18 cacti of the southwest<br />

tions from the crowded center of the areole. The uppermost<br />

ones are 1/8 to 1/4 of an inch long, and slender, while the rest<br />

of them are heavier and 1/4 to 3/4 of an inch long. They may<br />

be all translucent yellow, often with a reddish tip, or occasionally<br />

all reddish. All spines have bulbous bases. Immature<br />

plants or new branches have no spines at all, but have<br />

instead a thick covering of white, very fine, flexible hairs<br />

from 1/4 to 1/2 inch long, about 40 hairs to the areole. When<br />

the plant is 1 to 2 inches tall spines begin appearing at the<br />

tip of the stem. The juvenile hairs can often be seen on the<br />

bases of older specimens.<br />

flowers: Apparently identical with those of the species.<br />

fruits: Similar to those of the species.<br />

Range. A very small area including only two ranches 5 to 10<br />

miles south of Marathon, Texas.<br />

Remarks. This cactus was noticed by A. R. Leding in 1932, and<br />

he published an article on it in the Journal of Heredity in<br />

August, 1934, which included pictures of it. At that time he<br />

called attention to the unusual form of the juveniles, which are<br />

covered with long white hairs instead of spines, but he did not<br />

distinguish between the mature form of the plant and the typical<br />

E. chloranthus or state whether the typical form has hairs<br />

when young. He did say that comparison of plants from different<br />

localities and the discovery of whether the hairy condition<br />

is inherited might show that the form deserves specific or at<br />

least varietal rank.<br />

This article by Leding was reprinted in the October-November,<br />

1942, number of the Cactus and Succulent Journal of<br />

America, with an added note by the author, but no new conclusions<br />

were drawn.<br />

The immature plants of this variety are very striking, having<br />

no rigid spines at all, but being covered with much long white<br />

hair which is 1/4 to 1/2 inch long and produced usually 40 or so<br />

hairs to each areole. This hair is their only armament until the<br />

plants become about 11/2 inches high. At that size they suddenly<br />

begin producing the regular armament of yellow or white<br />

spines as described above. The bases of plants 11/2 to as much<br />

as 4 inches high will often show a belt of this white wool, with<br />

the typical spines above, but this hair is gradually replaced by<br />

normal Spines.<br />

The spines of adults of this form are very similar to those of<br />

E. chloranthus, but not identical. There are 30 or more radials<br />

where the typical form has only 12 to 23. These very numerous<br />

radials obscure the surface of the stem much more than do those<br />

of the typical form. Also, the centrals of this variety are more<br />

numerous and shorter. But the major difference between the<br />

two is in the juvenile stage. The young plants of typical E. chloranthus<br />

never have hairs at all, but are covered from the beginning<br />

with rigid spines. There is no possibility of confusing<br />

the two in this stage.<br />

E. chloranthus var. neocapillus is found growing in a very<br />

restricted range encompassing some hills from about 5 to about<br />

10 miles south of Marathon. Where undisturbed the plants<br />

grow in quite great numbers, but they have been removed almost<br />

entirely from much of their range. Fortunately, some of<br />

them are upon ranchland which is closed to all cactus digging,<br />

and so they still survive.<br />

Since the nearest occurrence of the typical E. chloranthus is<br />

about 150 miles northwest, there is little danger of confusing<br />

these two in the field. The many more spines, as well as the<br />

spine color and the areole shape easily set this variety off from<br />

E. viridiflorus var. cylindricus.<br />

Echinocereus russanthus Weniger<br />

Description Plate 3<br />

stems: Cylindrical, up to about 10 inches tall and 21/2 inches<br />

thick. These stems almost always branch to form clusters of<br />

up to a dozen stems. They have 13 to 18 ribs, which are low<br />

and narrow with indistinct tubercles. The color of the surface<br />

is medium green.<br />

areoles: Round at first, then broadly oval. About 1/8 of an<br />

inch long, with white wool when young, situated from 1/8 to<br />

3/8 of an inch apart.<br />

spines: All spines are very slender and somewhat flexible.<br />

There are 30 to 45 slender to very slender and bristle-like<br />

radial spines, which are very crowded and which interlock<br />

with those of adjacent areoles, giving the plant a very dense<br />

spine cover. There is a tuft of small ones at the summit (upper<br />

edge) of the areole, which are only 3/16 to 1/4 of an inch<br />

long. Moving laterally around the areole the radials become<br />

longer, the lower laterals being 1/2 to 5/8 of an inch long. All<br />

radials are white or straw-colored. There are 7 to 12 central<br />

spines spreading in all directions from bulbous bases. The upper<br />

ones are small, similar to the larger radials in size. The<br />

lower ones are from 3/4 to 11/4 inches long, though still slender<br />

and rather flexible. While there is a complete gradation<br />

of spine sizes from the smallest radial to the largest central,<br />

the centrals are distinguished from the radials not only by<br />

their position but also by their color, having at least the tip,<br />

often the upper half, and sometimes the whole spine reddish<br />

or purplish in color.<br />

flowers: About 1 inch long, funnel-shaped, not opening<br />

widely and so only about 1/2 inch in diameter. They are rustred<br />

in color sometimes with darker midlines. All segments are<br />

linear in shape, with the ends somewhat pointed, but not<br />

sharply so. The bases of all segments are lighter and the center<br />

of the flower is greenish. The stamens are pale yellow.<br />

The style is long and yellowish. There are 8 to 10 green<br />

stigma lobes.<br />

fruits: About 1/2 inch long, oval to almost spherical. They<br />

are covered with clusters of slender white spines, 10 to 12 on<br />

each areole.


genus Echinocereus engelmann 19<br />

Range. A small area of southwestern Brewster County, Texas,<br />

including the northern part of the Chisos Mountains and the<br />

country northwest of these to just past Study Butte, but not<br />

seen as far west as Terlingua or south to the Rio Grande.<br />

Remarks. I have often found it rewarding to examine the<br />

thousands of specimens which the Texas dealers in cacti have<br />

in their bins. Even a slight variation stands out when it is lying<br />

among dozens of its supposed fellows. Several times I have<br />

found in such places something new to me and have been able<br />

to trace it back to its location and discover what it was.<br />

In piles of E. viridiflorus var. cylindricus from the Texas Big<br />

Bend we repeatedly found one or two unusual specimens with<br />

far too many and too long and flexible spines to be that cactus.<br />

Usually the assortment was being sold as E. chloranthus, but the<br />

bulk of them were clearly the Big Bend forms of E. viridiflorus.<br />

These few obviously different specimens were all that could be<br />

seen to explain the confusion of names.<br />

These unusual specimens did have the round or oval areoles<br />

of E. chloranthus, and noting their long central spines I assumed<br />

at first that they were that cactus. They have been going under<br />

that name for a long time.<br />

But when I saw the actual E. chloranthus, which does not<br />

grow at all in the Big Bend, it was obvious that the two plants<br />

were really very different. All authorities have given the radial<br />

count for E. chloranthus as between 12 and 23, while this cactus<br />

has 30 to at least 45. That cactus has 3 to 6 centrals which<br />

are up to 3/4 of an inch long, while this one has 7 to 12 which<br />

are up to as much as 11/4 inches long.<br />

Such differences as these have usually been deemed sufficient<br />

to distinguish varieties in this group, and I thought at first that<br />

this would prove to be a Big Bend variety of E. chloranthus.<br />

But when the plants bloomed the new one presented a flower<br />

different in most respects from that of the other cactus. These<br />

flowers are smaller, with more narrow and linear petals, and<br />

are rust or russet-red in color. No flower of E. chloranthus ever<br />

approaches the color or the texture of these flowers, which are<br />

also the smallest I have seen on any Echinocereus. On E. chloranthus<br />

the flowers are yellow to brown or occasionally almost<br />

chocolate brown in color, but never with any reddish coloring,<br />

and their petals are firm, opaque, and glossy to the point of<br />

being almost waxy. On this Big Bend plant there is no green in<br />

the flower beyond the greenish center, the rest of the petals<br />

being essentially pale reddish, and these petals are soft, thin,<br />

somewhat translucent, and dull of surface instead of glossy.<br />

There is no mistaking the two plants when in flower, and these<br />

being so different, it seems clear that here is a species entirely<br />

separate from E. chloranthus.<br />

Once having concluded this, I cast around for what the cactus<br />

might have been named, but in all of the species and varieties<br />

listed for this group in the U. S. I have found no descrip-<br />

tion of this cactus.<br />

I did discover Engelmann’s description of a cactus from<br />

northern Mexico which seemed almost identical to this one. It<br />

was his E. longisetus. His only statement about the flowers of<br />

that cactus was “flower… said to be red.” When I saw that<br />

the type locality of his plant was in the Santa Rosa Mountains,<br />

which form the lower end of a continuous mountain chain of<br />

which the Chisos Mountains, where our cactus grows, is the<br />

northern end, I immediately thought that the two might be the<br />

same species and that it might range over this whole mountain<br />

chain.<br />

Following up this idea, several of us made trips into these<br />

difficult mountains of northern Mexico. We observed and collected<br />

E. longisetus in its type locality, where it grows in clusters<br />

of up to thirty sprawling stems, each up to 12 inches long. On<br />

the many plants observed we found the radial number always<br />

16 to 21 per areole and the centrals up to 2 inches long. These<br />

differences alone indicate that the plants are not the same. And<br />

when the true E. longisetus bloomed, there was no further<br />

question. It presented beautiful rotate flowers 2 to 21/2 inches<br />

in diameter, claret in color with white centers. They looked like<br />

a more reddish version of the E. berlandieri flowers, surely as<br />

different from the flowers of our Chisos Mountain plant as<br />

could be.<br />

We were unable to locate E. longisetus growing north of its<br />

type locality in the Sierra de Huacha or Sierra del Carmen<br />

ranges, and assume that it does not grow up into Texas. Neither<br />

did we find our cactus growing south of the northern slope of<br />

the Chisos Mountains. I assume, therefore, that these two separate<br />

species occupy only their respective escarpments of this<br />

large mountain system.<br />

E. russanthus is left, therefore, as what appears to be a separate<br />

species which, however, has always been thrown in with<br />

E. chloranthus. It occupies its own small range, where that<br />

cactus never grows, and is actually a much more beautiful plant<br />

than most of its close relatives.<br />

Echinocereus caespitosus Eng.<br />

“Lace Cactus,’ “Purple Candle,” “Classen’s Cactus”<br />

Description Plate 3<br />

stems: Spherical when very young, quickly becoming oval<br />

and then, when mature, cylindrical. These stems occasionally,<br />

in some locations, grow to 12 inches tall and 31/2 inches thick,<br />

but the typical adult size is 4 to 8 inches tall by 2 to 21/2<br />

inches thick. There is a form which never exceeds 3 inches<br />

tall and one inch thick. A plant may remain a single, erect<br />

stem all of its life, while its neighbor a few feet away may<br />

offset and branch to form a cluster of a dozen or so upright<br />

stems. This clustering habit is so common as to provide the<br />

basis for the name. There are 10 to 19 ribs which are narrow<br />

and definite and divided into distinct tubercles. The flesh is<br />

dark green.


20 cacti of the southwest<br />

areoles: Small, rather oval, and quite woolly when young,<br />

becoming greatly elongated vertically and bare of wool when<br />

older. These areoles are 1/8 to 3/16 of an inch long and almost<br />

touching on shrunken, dormant plants, but up to 3/16 of an<br />

inch apart on active, well-watered ones.<br />

spines: There are 12 to 36 rigid but slender radial spines.<br />

They radiate evenly, lying almost flat over the surface of the<br />

plant. Crowded around the elongated areole in this flat position,<br />

they look, on each side of it, much like the teeth of<br />

a comb, and so are often described as being pectinate. Those<br />

at the top of the areole are very tiny, often almost bristlelike<br />

and only 1/32 to 1/8 of an inch long. The lateral ones are<br />

robust and 3/16 to 5/16 of an inch long. The lower 1 to 3 are<br />

somewhat smaller again. The spines of adjacent clusters<br />

may interlock on the longer spined forms, but do not reach<br />

each other on the shorter spined individuals. These spines may<br />

be pure white, white with brown tips, yellowish with brown<br />

tips, or all brown, or, in one form, have the outer half of<br />

each spine shining black or purplish. The plant may be somewhat<br />

banded by variations in these colors, but the spines of<br />

any single areole are never variegated.<br />

Most commonly there is no central spine, but on many specimens<br />

in some locations one can find areoles with 1 central<br />

standing straight out or 2 centrals, one above the other in the<br />

center of the areole. An occasional plant will have such centrals<br />

on the majority or even all of its areoles. These centrals<br />

are stout and firm, but only 1/32 to 1/8 of an inch long.<br />

flowers: Very large and colorful on all but the dwarf form,<br />

being 2 to 5 inches tall and 2 to 4 inches in diameter, and<br />

brilliant purple or rose pink. The flower tube is covered with<br />

white, cobwebby wool to at least 1/4 of an inch long and<br />

clusters of 10 to 14 very fine, hairlike, white, gray, or black<br />

spines, 1/4 to 3/4 of an inch long. Above these the outer segments<br />

of the flower lengthen gradually, with greenish or<br />

brownish midlines and pink edges. The petals, of which there<br />

are 30 to at least 50, arise from usually narrow reddish or<br />

reddish-brown bases (which may, however, be bright green<br />

instead) to broaden to 1/4 of an inch or more wide above.<br />

This upper part of each petal is purple or rose-pink. Its edges<br />

are more or less ragged and often notched. The tips vary from<br />

erose and rather blunt to almost entire and definitely pointed.<br />

The filaments are reddish at the bases, fading above; the<br />

anthers, cream-colored. The style is long and reddish or pinkish,<br />

crowned by 8 to 22 large, dark green stigma lobes. The<br />

dwarf form has the same flower but with a marked reduction<br />

in size and number of almost all flower aspects.<br />

fruits: Egg-shaped or almost spherical, covered with the<br />

slender spines and wool of the ovary. It remains green until<br />

it dries and splits open by one or two vertical slits.<br />

Range. As Engelmann stated, this is the most eastern of the<br />

Echinocerei. Found in hilly, mostly limestone regions from near<br />

Ponca City to near Durant, Oklahoma, on the northeast, and<br />

from there on south to the edge of the coastal plain just west of<br />

the Brazos River. Industry and Cat Spring, Texas, mark its<br />

southeastern limit. From this line—which, as Engelmann observed,<br />

approximately parallels the 96th longitude—it occurs<br />

wherever the proper conditions are found throughout central<br />

and western Oklahoma and into southeastern Colorado, as well<br />

as over all of northwestern Texas and into a little of eastern<br />

New Mexico. The southern limit of its range runs west through<br />

Texas just south of San Antonio to Eagle Pass, where it dips<br />

into Mexico; from there it curves northwest past the vicinity<br />

of Sonora and Big Spring, Texas, to just east of Carlsbad and<br />

Roswell, New Mexico. Although it occurs in only widely scattered<br />

locations in eastern New Mexico, its western limit runs<br />

approximately from Carlsbad north into eastern Colorado.<br />

Remarks. The lace cactus is probably one of the most collected,<br />

most fancied, and best known of all cacti. Almost anyone<br />

the world over who grows cacti has this plant, and it is often<br />

the favorite of the collection. The high regard in which it stands,<br />

is well earned by the beauty of the plant body with its truly<br />

lacy spines and the exquisiteness of its flowers, which are produced<br />

in profusion by a healthy plant. This cactus is well-<br />

known because it is very widespread in range and so prolific<br />

in its natural habitats as to be readily available to any dime<br />

store or nursery which wants to stock it. Furthermore, it is<br />

tolerant of the rigors of cactus culture after it is in the collection.<br />

It is the common cactus of two-thirds of Oklahoma and<br />

all of central Texas. It is not found everywhere in this wide<br />

area, but the right sort of limestone or gypsum hill often sup-<br />

ports a population of literally hundreds of individuals.<br />

The fact that this cactus has been so commonly seen has not,<br />

as one might have hoped, made for less confusion concerning it.<br />

Even yet a variety of names are bantered back and forth and<br />

each new work on cacti still is unable to make an air-tight case<br />

for what the plant really is and what its official name should<br />

be. Knowing this, I have made special effort to study thousands<br />

of specimens from all possible locations within its range, and<br />

have growing before me specimens from each general area in<br />

which it is found.<br />

Several quirks of history make for two major problems in<br />

dealing with this cactus, and each leads to its own type of confusion.<br />

I will attempt here to give as simple an account as possible<br />

of what has happened in the past in order to evaluate the<br />

conflicting views which exist today about E. caespitosus.<br />

The plant was undeniably studied by Engelmann, who wrote<br />

up several very complete and detailed descriptions of it between<br />

1848 and 1856. He originated the name Echinocereus<br />

caespitosus for his cactus. But Terscheck had in 1843 named a<br />

plant from Mexico Echinocactus reichenbachii. His description<br />

was very incomplete and could, as many have since pointed out,<br />

fit any one of several cacti in more than one genus.<br />

It is very doubtful whether the two names would ever have<br />

been associated at all except that Prince Salm-Dyck, in 1844,


genus Echinocereus engelmann 21<br />

referred to some plants in European collections as Echinopsis<br />

pectinata var. reichenbachiana. He wrote Engelmann about the<br />

possibility that these might be the same as Engelmann’s plant,<br />

and Engelmann mentioned this speculation in one account of<br />

E. caespitosus.<br />

Early writers to and including Coulter all used Engelmann’s<br />

name, E. caespitosus, which is the earliest name with an indisputably<br />

adequate description, for the plant. Things might have<br />

remained this way and all would have known what plant they<br />

were talking about, but in 1893 F. A. Haage, Jr., changed Terscheck’s<br />

useless Echinocactus reichenbachii to Echinocereus<br />

reichenbachii, and later Britton and Rose adopted this as the<br />

name they used when referring to our cactus.<br />

It is true that by the rules of nomenclature the earliest name<br />

applying to a plant must be used, but Terscheck’s description<br />

does not any more link his name to this plant than to a dozen<br />

others, while Prince Salm-Dyck’s usage of the name as a variety<br />

of E. pectinatus leads definitely away from our plant. It<br />

would therefore seem that the resurrection of Terscheck’s name<br />

and the adoption of it for our plant was probably not required<br />

by the rules. It has certainly led to continuing confusion since<br />

that time.<br />

But once done, Britton and Rose’s beautiful and popular<br />

books carried the name E. reichenbachii so well into general use<br />

that what cactus E. caespitosus actually was has been forgotten<br />

by almost everyone. Yet any serious student coming across this<br />

name so definitely set forth by Engelmann would have to do<br />

something with it. Some merely regarded it as a synonym of<br />

the other, but others tried to prove that there were two separate<br />

plants for the two names. And here the confusion had its second<br />

effect and led to the second large problem encountered in understanding<br />

this cactus.<br />

Engelmann pinpointed the eastern range of this cactus almost<br />

exactly when he said it corresponded to the 96th meridian, and<br />

he very well indicated the western extent of its range in south<br />

Texas when he said it goes no farther west than the San Pedro<br />

(Devil’s) River. But he badly underestimated the extent of its<br />

range northward and westward in the northwestern quadrant<br />

of its area when he said it occurred only to about the 100th<br />

meridian and the Canadian River. In a later note he did add<br />

that it grew from the Arkansas River to Saltillo, which exactly<br />

corrects the range northward, but he did not ever seem to know<br />

that it occurs west of the 100th meridian. Actually, the plant is<br />

found well past the western edge of the Texas and Oklahoma<br />

panhandles to the last hills east of the Pecos River in New<br />

Mexico.<br />

Now, as one might expect, over the huge extent of this<br />

range from east to west and north to south there are certain<br />

gradients of characteristics in this cactus. And since Engelmann<br />

apparently had only eastern specimens to examine—his type<br />

came from the farthest southeastern location where it grows—<br />

his descriptions are definitely slanted toward the characters<br />

most commonly found in the eastern populations and do not<br />

allow for extremes found commonly only in specimens from<br />

the part of the range he did not know existed. For example,<br />

eastern specimens almost always have 20 to 30 radial spines and<br />

only extremely rarely—I estimate from my experience maybe<br />

in one plant out of a thousand—does one find fewer than 20<br />

radials. Yet in the plants from far western Oklahoma, northwestern<br />

Texas, New Mexico, and Colorado, while radial counts<br />

of 12 to 32 are found, many localities have a vast majority<br />

showing fewer than 20 radials and only a rare plant with more<br />

than 20. Throughout central Oklahoma we find all numbers<br />

from 12 to 32 commonly on the same hillside. Also, western<br />

plants usually have less wool on the growing stem areoles and<br />

shorter spines than eastern ones. There are other less obvious<br />

gradients as well.<br />

So when students who took the name E. reichenbachii for the<br />

common eastern forms began to notice the lower spine numbers,<br />

shorter spines, and smaller amount of wool of the far northwestern<br />

forms, they tended to think they had in these northwestern<br />

plants something worthy of a name. Britton and Rose<br />

made up some new names such as E. perbellus for these forms,<br />

but others, by a strange reversal, called them E. caespitosus.<br />

Boissevain and Davidson seem to have initiated this in their<br />

work on the Colorado cacti, and others have followed them.<br />

Backeberg, in a 1941 article, made a case for a plant from the<br />

Wichita Mountains of Oklahoma as being the true E. caespitosus<br />

because of its extreme amount of clustering, but in his later<br />

works has placed that plant correctly as a form of E. baileyi.<br />

However, he still keeps E. caespitosus as separate from E.<br />

reichenbachii, saying that the spine clusters of adjacent areoles<br />

on E. caespitosus do not intertwine, while those of E. reichenbachii<br />

do. I have checked this character specifically at dozens<br />

of locations all over the range of these cacti, and have found<br />

growing together, wherever there is a large population of<br />

plants, specimens with spines interlocking and specimens with<br />

spines not interlocking. Coulter made this variation clear many<br />

years ago, saying that on E. caespitosus, “Spines may or may<br />

not interlock.”<br />

It seems obvious, therefore, that we have one definite species<br />

with a very large range over which there is gradation. The<br />

species description in respect to spine number and so forth is,<br />

therefore, broadened in order to include specimens from the<br />

part of the range Engelmann did not see. The only question<br />

really remaining is which of the two proposed names is legitimate,<br />

and since I cannot honestly tell whether Terscheck meant<br />

this cactus by his Echinocactus reichenbachii, while it is obvious<br />

that Engelmann did with his Echinocereus caespitosus, I<br />

use that name here.<br />

Engelmann noticed that the spine color of individual specimens<br />

of this cactus varied from pure white to chestnut-brown<br />

or rosy. He called the brown ones E. caespitosus var. castaneus.<br />

This would be the form called by collectors the “brown lace.”<br />

Later an attempt was made to transfer this variety to E. pectinatus,<br />

a related species, and then to equate it with E. pectinatus


22 cacti of the southwest<br />

var. rubescens. However, other characters such as the long wool<br />

and the hairlike spines of the flower tube quickly distinguish it<br />

from E. pectinatus and show this sort of combination to be in<br />

error. The fact that almost any large population of E. caespitosus<br />

studied shows a complete range of colors from white to<br />

brown with all sorts of intermediates, and each local population<br />

varying only in proportionate numbers of the various<br />

colors, indicates that the color of individual plants is only a<br />

simple genetic character and not worthy of varietal name at all.<br />

At one time members of the Oklahoma cactus society collected<br />

and studied large numbers of these cacti from the Arbuckle<br />

Mountains of that state, where this species is very common. In<br />

doing this they found several specimens which bloomed with<br />

yellowish instead of violet flowers and one plant with white<br />

flowers. For these specimens they proposed the variety names<br />

aureiflora and albiflora, respectively. There is no report of these<br />

flower colors being seen again, and they are not listed as true<br />

varieties.<br />

If we can look beyond all this confusion, we have in this<br />

species a beautiful cactus, the lace cactus, white to brown in<br />

spine color, which is deservedly a favorite of all. It is most<br />

easily confused with E. pectinatus. Sometimes individual plants<br />

of the two are very hard to tell apart by spine and stem characters,<br />

although the spines of E. caespitosus are typically not so<br />

extremely pressed against the plant nor quite so heavy as those<br />

of the other. But for absolute identification of the two one must<br />

sometimes wait for the flowers to appear. There are various<br />

differences here, the most obvious being that—on the flower<br />

tube and remaining on the fruit—E. caespitosus has much long<br />

wool and extremely thin, flexible, hairlike spines up to 3/4 of<br />

an inch long, while E. pectinatus has shorter wool and comparatively<br />

thick, rigid spines up to only 3/8 of an inch long.<br />

Echinocereus caespitosus var. caespitosus (Eng.)<br />

Description Plates 3, 7<br />

stems: As the species.<br />

areoles: As the species.<br />

spines: As the species, except that it has 15 to 36 radial spines<br />

which are usually long enough to interlock with those of adjacent<br />

areoles. In most of its range only a very rare plant has<br />

the radials fewer than 20 in number. However in the extreme<br />

northwestern part of the range individuals with 15 to<br />

20 radials become the majority in some populations.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. From approximately the 96th parallel in Texas and<br />

Oklahoma west to near Del Rio, Big Spring, and Amarillo in<br />

Texas and throughout central and western Oklahoma to the<br />

base of the Oklahoma Panhandle, but hardly if at all entering<br />

the Oklahoma Panhandle itself. The most northwesterly records<br />

of this form we have are Alabaster Caverns, Woodward, and<br />

Shattuck, Oklahoma, and Sanford and Palo Duro Canyon in<br />

Texas.<br />

Remarks. I take this to be the typical form of the species. It is the<br />

form discovered and described by Engelmann, from specimens<br />

taken in southeast Texas, as E. caespitosus. It is the form of the<br />

species which clusters the most, the name applying well for this<br />

reason. It is also the largest form the species assumes.<br />

I have broadened the spine count included in this variety<br />

from Engelmann’s 20 to 30 radials because in populations over<br />

the whole of the range given above we find occasional individuals<br />

exceeding his limits in both maximum and minimum to<br />

the extent we have indicated. Specimens with the lower numbers<br />

make up a larger proportion of the population the farther<br />

northwest one goes, but individuals falling entirely within his<br />

range of spine numbers are found all the way to the edge of the<br />

area we have indicated. I feel the plants over this wide area<br />

comprise one form varying in a continuous gradient from southeast<br />

to northwest. I consider it the typical variety. Those segments<br />

of this species which vary from this typical variety I list<br />

as separate varieties.<br />

Echinocereus caespitosus var. minor Eng.<br />

Description Plate 4<br />

stems: As the species, except that it is very much smaller,<br />

apparently reaching a maximum of only 3 inches tall and 1<br />

inch in diameter.<br />

areoles: As the species.<br />

spines: As the species, except that no centrals have been seen.<br />

flowers: Small for the group, being 2 inches tall and 11/2 to<br />

13/4 inches in diameter. They are pale lavender-pink in color,<br />

and are otherwise as those of the species, except that the<br />

petals are smaller and only 15 to 25 in number, and the<br />

stigma lobes 8 in number.<br />

fruits: As the species, except smaller.<br />

Range. Known only from the vicinity of Stockdale, Wilson<br />

County, Texas.<br />

Remarks. Engelmann thought it necessary to set this dwarf<br />

form of the lace cactus apart as a variety, and so it seems to be.<br />

The diminutive size of its body is striking and the reduction of<br />

its flower parts from the typical numbers is consistent.<br />

Warning should be given, however, that not every small lace<br />

cactus discovered is this variety. Engelmann spoke about how<br />

precocious E. caespitosus is in blooming, and it is not unusual<br />

to find a 2-inch specimen of the typical variety in bloom. In<br />

certain areas, notably around the granitic region of Llano and<br />

Ink’s Lake, Texas, are found many clusters of very small stems.<br />

These, however, all bloom with the typical huge, many-petaled


genus Echinocereus engelmann 23<br />

purple flowers, and when grown in the garden will in time take<br />

on full stem size.<br />

The cactus meant by variety minor is different in that it does<br />

not grow larger in any situation, and its flowers are small, with<br />

only around 20 petals as opposed to the 30 to 50 of the typical<br />

form of the species.<br />

Engelmann gave no range or location for his dwarf form.<br />

We have encountered it only in Wilson County, Texas, where it<br />

grows in some fields in rather dense stands.<br />

Echinocereus caespitosus var. perbellus (B. & R.)<br />

Description Plate 4<br />

stems: As the species, except that it is not over 4 inches tall,<br />

with ribs numbering 13 to 15.<br />

areoles: As the species.<br />

spines: As the species, except that the total radial number is<br />

only 12 to 15 and the central spine is always missing.<br />

flowers: As the species, except that they are not as large as<br />

the flowers of the species sometimes become. They are usually<br />

about 2 inches tall and wide.<br />

fruits: As the species.<br />

Range. The type locality is Big Spring, Texas. Individuals are<br />

found occasionally, associated with typical E. caespitosus populations<br />

north of this point in Texas, along the western edge of<br />

Oklahoma as far as Majors County, west into New Mexico as<br />

far as the Pecos River, and on into Colorado. A pure population<br />

of the form, unmixed with any other, has been found only<br />

near Muleshoe, Bailey County, Texas.<br />

Remarks. This is at best a doubtful variety, and certainly not<br />

a separate species. Its existence at all as a separate entity would<br />

probably never have been advocated except for that unfortunate<br />

failure of Engelmann to have Specimens from the northwestern<br />

part of its range when he drew up his description of<br />

E. caespitosus. Limited as he was to specimens from the east, he<br />

gave as the radial number for his species 20 to 30, which is the<br />

number on almost all examples found in central Oklahoma and<br />

Texas. But even here a rare plant will show 15 to 20 or more<br />

than 30 radials, giving us a hint that his spread is too narrow.<br />

This fact was apparently not realized for many years.<br />

So when Britton and Rose were confronted with small specimens<br />

from Big Spring, Texas, with only 12 to 15 radials, they<br />

immediately set up a new species, calling it E. perbellus, although<br />

all of its other characters which they listed equaled<br />

those of E. caespitosus.<br />

Later authorities appear to have seen no specimens of this<br />

form except the originals, and the species has stood as Britton<br />

and Rose described it until this time.<br />

In this study I have made great effort to collect this species<br />

in its type locality and to study it. Two extensive collecting<br />

trips into the Big Spring area netted numerous specimens of E.<br />

caespitosus with radials from 17 to 24 in number, but not those<br />

with the 12 to 15 of Britton and Rose. Finally, on a third extensive<br />

search of the area we collected two specimens, both<br />

small plants identical to the previous ones, but both with 12 to<br />

15 radials. We had apparently re-collected E. perbellus at last.<br />

But in the process we had established that specimens with<br />

fewer than 20 radials but more than 15 were common in the<br />

area, as they are in the surrounding countryside. We began to<br />

suspect that intermediates could be found linking these two<br />

numbers entirely and that we had no separate species here after<br />

all. Later collections throughout northwest Texas, western Oklahoma,<br />

eastern New Mexico, and Colorado established that the<br />

range of radial numbers throughout the area was 12 to 30, the<br />

most common being 17 to 26, these latter numbers appearing in<br />

the populations at almost any good location.<br />

At one location in Majors County, Oklahoma, we found<br />

specimens with radial counts as low as 12 and as high as 23.<br />

This was the same location where Caryl, in 1935, reported the<br />

first collection of E. perbellus in Oklahoma. But what we had<br />

here was no separate population of a separate form, but instead<br />

just some specimens of E. caespitosus with the lower extreme of<br />

spine number, mixed with typical specimens. At this discovery<br />

we were ready to follow Mrs. Lahman, no doubt the greatest<br />

student of Oklahoma cacti, who also in 1935 wrote, “In the<br />

meantime, I have crossed E. perbellus from my list of Oklahoma<br />

cacti until I find someone who really knows what it is.” We had<br />

about decided that Britton and Rose’s taxon had arisen out of<br />

Engelmann’s failure to realize the scope of his species and that<br />

it should be entirely relegated to the synonymy.<br />

This may still be the case, and the name may yet be dropped,<br />

but since then we have discovered a population of cacti in<br />

Bailey County, Texas, near Muleshoe, in which all of several<br />

dozen plants examined have 12 to 14 radials and no centrals.<br />

These bloomed with flowers typical, although small for the<br />

species, and have no other obvious character to distinguish<br />

them from that species. Since this is a pure population of this<br />

form, perhaps it should be recognized after all. It certainly<br />

cannot be considered a separate species, and may be only a<br />

clone. In the meantime, in the complete absence of any experimental<br />

evidence concerning any of the relationships of these<br />

plants, the most logical way to treat it so that it will not be lost<br />

and will be available for future study is to call it a variety of<br />

the species.<br />

Echinocereus caespitosus var. purpureus (Lahman)<br />

“Black Lace”<br />

Description Plate 5<br />

stems: As the species, except that it is smaller in maximum<br />

size and clusters more sparingly.


evalution -> evaluation<br />

24 cacti of the southwest<br />

areoles: As the species.<br />

spines: 14 to 22 radials and 0 to 3 centrals, as the species,<br />

except that the outer part of each spine is shiny purplish or<br />

glistening black in color.<br />

flowers: As the species, except that they are always purple<br />

in color. There were 12 stigma lobes on all specimens seen.<br />

fruits: As the species.<br />

Range. The Wichita and Glass (Gloss) mountains of western<br />

Oklahoma.<br />

Remarks: This cactus was found at Medicine Park, Oklahoma,<br />

on the eastern edge of the Wichita Mountains, and described by<br />

Mrs. Lahman, who studied the cacti of Oklahoma extensively.<br />

It is very rare and has been seldom seen again. Mr. Charles Polaski<br />

of Oklahoma City, who has the finest private collection<br />

of cacti I have seen, supplied the first specimen of it which I<br />

had the privilege to study. This specimen came from the Glass<br />

Mountains about 100 miles north of the type locality at Medicine<br />

Park. Mr. Polaski, who grew up in the Wichita Mountains<br />

near Medicine Park says he has never seen it there, and I have<br />

searched the type locality without finding either it or any form<br />

of E. caespitosus. A large area of Medicine Park is now under<br />

the waters of Lake Lawtonka, and perhaps it grew in this nowflooded<br />

area.<br />

The Glass Mountain plants show the strikingly beautiful,<br />

purple-black coloring of the spines described by Mrs. Lahman.<br />

They show all of the other characters of her plant, except that<br />

the variation of radial spines is from 14 to 22, which raises the<br />

maximum number 4 spines beyond her report. This does not<br />

seem significant in the light of the variations in spine number<br />

we have already seen in this group. The large flower is as she<br />

describes, a very beautiful deep meadow violet, except that the<br />

throat may be reddish instead of green as she described it. This<br />

also seems unimportant, since we find other specimens of E.<br />

caespitosus with flowers with either green or reddish throats.<br />

The stigma lobes on all specimens seen were 12 in number, which<br />

is strikingly regular for this group.<br />

An evaluation of all the characters of this plant seems to show<br />

only the dark coloration of the spines to distinguish it from the<br />

typical E. caespitosus. Everything else falls within the known<br />

range of the species, when it is interpreted in the broad way<br />

made necessary by including specimens of all its range. This<br />

spine color does not seem sufficient grounds for considering it<br />

a separate species, so I place it here as a variety. Whether even<br />

this is justified remains to be determined after further study.<br />

The coloring of the plant body is very beautiful, and so conspicuous<br />

as to prompt special exclamations when it is seen in a<br />

collection. We have no other cactus with this striking coloration.<br />

It is so dark that the term, “black lace,” arises spontaneously<br />

for it. Even this shows how closely it is related to the<br />

more common lace cactus. Unfortunately it is very rare. Much<br />

ranchland must be covered to discover a single specimen. It<br />

would be a good cactus to be distributed by the trade.<br />

Echinocereus melanocentrus Lowry<br />

Description Plate 5<br />

stems: Almost spherical to oval when young, becoming<br />

cylindrical. Single, or when very old, sometimes branching to<br />

include two or three side-branches. Each stem may reach a<br />

maximum of about 61/2 inches tall and about 2 inches thick.<br />

There are 10 to 13 ribs of definite, confluent tubercles. The<br />

surface color is very deep green.<br />

areoles: Practically touching to 1/8 of an inch apart. They<br />

are oval when young to elongated when old, and woolly at<br />

first, becoming bare.<br />

spines: There are 17 to 20 slender radial spines lying pectinate<br />

close to the plant surface. The upper ones are only 1/32 to 1/16<br />

of an inch long, the lateral ones gradually lengthening to a<br />

maximum in the lower laterals of 1/4 to 1/8 of an inch. The<br />

lowermost spines are somewhat shorter again. Spines of adjacent<br />

clusters may or may not interlock. There is one central<br />

spine on each areole standing out, perpendicular to the plant<br />

body, or turned slightly upward. It is straight and slender,<br />

like the radials and rises from a bulbous base, black or mahogany<br />

in color. It is 3/16 to 3/8 of an inch long.<br />

flowers: 2 to 3 inches across and tall. They are showy rosepink<br />

in color, with reddish centers. The petals are almost linear<br />

or slightly broadening over their upper parts, their ends more<br />

or less ragged. They recurve greatly as the flower ages, leaving<br />

it open extremely wide by the second day. The style is pinkish,<br />

the stigma lobes green and 12 or 13 in number. The ovary<br />

has on it cobwebby wool and very black or brown and white,<br />

very slender, flexible, hairlike spines 1/4 to 1/2 inch long.<br />

fruits: Unknown.<br />

Range. Very localized in sections of Jim Wells and Kleberg<br />

counties, Texas.<br />

Remarks: E. melanocentrus is one of those rare little cacti we<br />

find growing in extremely localized situations here and there.<br />

It is found under the extremely heavy brush of the small amount<br />

of uncleared territory there is left around Alice and Kingsville,<br />

Texas. Even here it is only found in certain spots. Much of this<br />

territory is in ranches where the owners are extremely jealous<br />

of their rights to keep outsiders out, and so few had seen this<br />

cactus until recently. Perhaps this is fortunate, because at the<br />

same time that this keeps the plant out of the public eye it<br />

preserves it in its natural setting. But the ranchers are using<br />

clearing devices and now spraying with brush-killing chemicals<br />

more and more, so the plant is not safe in any case. A few<br />

Texas dealers have found an accessible location and in the past<br />

few years I have seen possibly a hundred of the plants being<br />

distributed.<br />

This species is very similar to E. caespitosus in general appearance.<br />

It looks much like that other species with one conspicuous<br />

black central stuck on each areole. However, E. caespitosus<br />

does not grow anywhere within over 100 miles of the


genus Echinocereus engelmann 25<br />

location of E. melanocentrus, and the general aspect of the<br />

flowers of the two plants are somewhat different.<br />

The flowers of E. melanocentrus are apparently identical to<br />

those of E. fitchii, and there are other similarities to this species<br />

found farther west in south Texas, but the two will hardly be<br />

confused, since E. fitchii has 3 to 7 spreading centrals instead<br />

of only one, and its radials are not pectinate.<br />

E. melanocentrus is a little known specialty of an area which,<br />

having none of the more common representatives of this group,<br />

boasts its own<br />

Echinocereus fitchii B. & R.<br />

Description Plate 5<br />

stems: Upright and usually single, sometimes putting out<br />

only 1 or 2 side-branches when very old. These stems are to<br />

6 inches tall and 2 inches thick, medium to dark green in<br />

color, having 10 to 14 ribs of low, confluent tubercles.<br />

areoles: Round when young, becoming oval on the sides of<br />

the plant. They are small in size, almost touching to 3/16 of<br />

an inch apart, and woolly when young, becoming bare.<br />

spines: There are 20 to 25 radial spines which are white with<br />

brown tips to tan with reddish-brown or black tips. They are<br />

slender and not pectinate, but spreading outward somewhat<br />

from the plant surface. The uppers are very short, the lower<br />

laterals becoming 1/4 to 3/8 of an inch long. There are 3 to<br />

7 slender central spines which are the same color as the radials,<br />

rising from slightly bulbous bases, 1/8 to 3/16 of an inch<br />

long and spreading to the sides instead of standing out in one<br />

row.<br />

flowers: Large and showy, pink, always with dark burgundy<br />

centers. They are about 21/2 inches tall and 2 to 4<br />

inches across. The flower tube possesses much cobwebby wool<br />

and 10 to 17 hairlike, yellowish or white, dark-tipped spines<br />

to 1/2 inch long. There is much variation in petal shape, as<br />

there is in this whole group. They run from linear to spatulate,<br />

and the ends are blunt or pointed, ragged or entire. Usually<br />

the petals recurve when the flower is completely open. The<br />

long style is pink, and there are 12 or 13 green stigma lobes.<br />

The flowers are fragrant.<br />

fruits: Spherical to oval, 1/2 to 1 inch long. They remain<br />

green and covered with white wool and spines.<br />

Range. Along the Rio Grande and away from it only a few<br />

miles from near Rio Grande City northwest past Laredo to not<br />

quite as far upstream as Eagle Pass, Texas.<br />

Remarks. E. fitchii seems to be an immigrant to us from Mexico,<br />

appearing as it does just along a stretch of the Rio Grande, but<br />

it appears not to be widespread in north Mexico either. I have<br />

collected paralleling its whole range about 50 miles within<br />

Mexico without seeing one specimen of the plant that far south.<br />

It does occur in Mexico, and there are other forms farther down<br />

in that country whose relationship to this must be close, but<br />

within our own territory it has only a tenuous foothold on<br />

gravelly hillsides overlooking the Rio Grande. Much of its territory<br />

was flooded by the huge Falcon Lake, and it is now possible<br />

to collect this cactus literally from the boat, when that<br />

lake is high. It is found to the northwestern edge of Webb<br />

County, but hardly farther in that direction, its range stopping<br />

almost too abruptly to believe, about 30 miles below the lower-<br />

most collection point of E. caespitosus.<br />

E. fitchii is easily distinguished from the other cacti of this<br />

group by the fact that its spines spread outward instead of lying<br />

neatly pectinate. This is a slight difference, but apparently a<br />

sufficient one, as one never hears this cactus called a lace cactus.<br />

Its numerous spreading centrals also distinguish it.<br />

Its spines are arranged very nearly like those of E. dasyacanthus,<br />

and it does look much like a small, delicate version of<br />

that cactus, but every other character is different, and no one<br />

should confuse them.<br />

It also shows a superficial similarity to E. pectinatus, but,<br />

besides the differences in the spines, the flowers are different.<br />

The ovary with its wool and hairlike, flexible spines is entirely<br />

unlike that of E. pectinatus, which lacks the long wool and has<br />

rigid spines.<br />

There is much variation in the spine colors of E. fitchii, as<br />

indicated in the description, but all spines on any plant are the<br />

same, so there is no banding. The difference is from individual<br />

to individual. The extreme on one side is almost pure white,<br />

with only pale brown tips on the spines; the plant of the opposite<br />

extreme has spines all tan or honey-colored below, with the<br />

outer half of each spine red-brown or black. Most plants in<br />

between these extremes present a sort of salt and pepper appearance<br />

over-all.<br />

The species is easy to grow and presents beautiful flowers.<br />

These flowers vary remarkably, however, in size and petal characters;<br />

there is hardly any variation in their fine, delicate col-<br />

oring.<br />

Echinocereus baileyi B. & R.<br />

Description Plate 6<br />

stems: Globose or oblong at first, becoming cylindrical with<br />

age. They sometimes remain single, but usually form clusters<br />

which may become very dense with up to 30 stems when old.<br />

Individual stems measure up to 8 inches tall and 31/2 inches<br />

thick. They are medium green in color, with ribs narrow and<br />

somewhat tuberculate.<br />

areoles: Oval when young, and very woolly, becoming<br />

elongated when older. They usually become bare when very<br />

old, but may for a long time have a mass of dirty white or<br />

tawny wool at the upper edge of the areole, and some plants<br />

relatoinship -><br />

relationship


26 cacti of the southwest<br />

have been seen on which the areoles remained rimmed with<br />

wool. They are 1/8 to 1/4 of an inch apart.<br />

spines: There are 12 to 28 slender but rigid radial spines<br />

which are not pectinate, but spreading outward from the<br />

plant and interlocking with those of adjacent areoles. The<br />

upper ones are very small and weak, the lateral ones become<br />

progressively longer until the lower laterals are 5/8 to 1 inch<br />

long. There are 0 to 5 centrals, these often very small, sometimes<br />

only 1/8 to 3/16 of an inch long, but on those plants with<br />

well-developed centrals they may be 1/4 to 3/8 of an inch<br />

long. When there are several centrals they spread from<br />

crowded bases almost lined up vertically in the middle of the<br />

areole. The spines vary in color from pure white to yellowish,<br />

straw to rust-brown, or even rosy reddish, but all on any<br />

given plant will be the same color, giving a fine variety of<br />

colors in most stands of the species.<br />

flowers: Large and showy, 2 to 3 inches tall and 21/4 to 31/2<br />

inches in diameter. They are fuchsia in color. The petals arise<br />

from narrow red bases and their upper parts are broad and<br />

fuchsia in color, with the ends ragged or erose. The stamens<br />

are very short, and the style is short for the group. There are<br />

10 to 21, but most commonly 10 to 12, dark green stigma<br />

lobes. The ovary surface is covered with a great deal of long<br />

white wool and also has 5 to 15 hairlike, white to rusty spines<br />

1/4 to 3/8 of an inch long on each areole.<br />

fruits: Egg-shaped, 3/8 to 1/2 of an inch long. They remain<br />

green and covered with wool and bristles until they dry and<br />

split open laterally to scatter the seeds.<br />

Range. Apparently restricted to the Wichita Mountains of<br />

southwestern Oklahoma.<br />

Remarks. This is one of the most attractive of the Echinocerei,<br />

yet it is very restricted in its range and, therefore, is not so well<br />

known as some of its relatives. It is an inhabitant of the unique<br />

granitic region of southwest Oklahoma comprising the Wichita<br />

Mountains. There have been occasional reports of its having<br />

been collected outside these mountains in adjacent areas of Okla-<br />

homa and even in nearby Texas, but those reports which it<br />

has been possible to check have all proved to deal with E. caespitosus<br />

instead. As one enters the Wichita Mountains from the<br />

east, near Lawton, Oklahoma, E. baileyi is found immediately,<br />

growing in beautiful stands, often of hundreds of plants together,<br />

upon the undisturbed granite-strewn slopes and ledges.<br />

It is found in profusion almost throughout the Wichita Mountains<br />

Wildlife Refuge and north of it a few miles, then northwest<br />

to Quartz Mountain and its attendant hills. The last of this<br />

mountain chain on the northwest is Granite Mountain, arising<br />

all alone above the town of Granite, Oklahoma. All over this<br />

huge pile of red granite, in the crevices of the main mass and<br />

in between the huge boulders, are clumps of E. baileyi, but beyond<br />

this, as in every other direction from these mountains,<br />

stretch the ordinary hills and plains of Oklahoma, where E.<br />

caespitosus takes over.<br />

E. baileyi is easily distinguished from its relatives by the profusion<br />

and length of its spines. These cover the whole plant and<br />

stand out from its surface in all directions, giving it a distinctly<br />

unkempt appearance when compared with the majority of the<br />

Echinocerei. Only E. longisetus from far-off northern Mexico<br />

has such long and unruly spines, and those are extremely slen-<br />

der and flexible, while these of E. baileyi are strong and rigid.<br />

The flowers of E. baileyi are similar in many ways to those<br />

of the lace cactus, but are not quite as big and flamboyant, and<br />

are rose-red rather than purple.<br />

When a whole population of this cactus is viewed on any<br />

good mountainside in its range, it presents a remarkable selection<br />

of spine colors. There is no variation on the individual<br />

plants and, therefore, no banding at all; but individual plants<br />

vary greatly from each other in color, the range of colors running<br />

from pure white through yellowish, tan, brown, and rusty<br />

to rose-red. The effect of all these colors together in a thick<br />

stand is very attractive. Also presented is quite a variation in<br />

number and length of radial spines, from as few as 12 to as<br />

many as 28, and the longest of these from as little as 5/8 of an<br />

inch on some plants to as much as 1 inch on others. Central<br />

spine number varies also, from none on some plants to 5 on the<br />

most heavily armed ones.<br />

All of these variations, with the exception of the radial number,<br />

seem to present gradients through the range, the majority<br />

of the plants in the east being white or light color and averaging<br />

shorter spines and fewer centrals, while on the western edge of<br />

the range at Granite, the majority are rusty or reddish, have<br />

spines averaging longer, and usually have several centrals in<br />

each areole. One cannot set up ranges for these variations within<br />

the over-all range, however. These are only averages. For<br />

instance, the darkest colored, rosy-tipped spines I have seen<br />

were on a plant growing within inches of a pure white neighbor<br />

near Medicine Park, at the eastern edge of the range, while the<br />

shortest spines I measured were on a white individual on far<br />

western Granite Mountain. These variations thus appear to be<br />

the result of simple quantitative inheritance within one major<br />

unit population.<br />

Britton and Rose first named and described this cactus. Unfortunately<br />

they had the benefit of only a few specimens to<br />

examine, brought to them by Mr. Bailey; and, therefore, they<br />

had no concept of the actual limits of the species’ characters.<br />

This fact has caused much confusion, especially since most more<br />

recent students have apparently seen no more representative<br />

series of the plant than they did, and so have attempted to follow<br />

Britton and Rose’s too narrow description. Backeberg, for<br />

instance, goes to quite some trouble, apparently in order to<br />

agree with Britton and Rose who say the plant has no centrals,<br />

to try to make a case that on older areoles the spines arising<br />

in the middle of the young areoles all finally assume radial<br />

positions. This is unfortunate, as on every mountainside, even<br />

among the predominantly central-less specimens of the east, there<br />

is an occasional plant with a distinct central or two, and in the


genus Echinocereus engelmann 27<br />

west the vast majority of plants have 2 to 5 unmistakable centrals<br />

up to 3/8 of an inch long. Similarly, Britton and Rose give<br />

“about 16” as the number of radials, while the number actually<br />

varies from 12 to 28. In the description given above I have<br />

broadened the limits to include the whole range of characters<br />

found in studying the whole population over its entire known<br />

range. This seems the only way to understand it as the contin-<br />

uous entity that it is.<br />

Probably because of Britton and Rose’s too limited description,<br />

attempts have been made to segregate parts of this population<br />

as separate species. Mrs. Lahman, noticing specimens which<br />

did not conform to the original description, described and attempted<br />

to set off E. oklahomensis, distinguished by having 20<br />

to 24 radials only 5/8 of an inch long and 0 to 2 centrals. She<br />

also set up E. longispinus, with 14 to 16 radials up to 1 inch<br />

long. Specimens with both these sets of characters are easily<br />

found on almost any undisturbed slope in the Wichita Mountains,<br />

and it does not seem that they deserve even varietal designation.<br />

They just represent two extremes of the plant, un-<br />

known to Britton and Rose.<br />

A more recent attempt has been made by Backeberg to break<br />

up the species. In an article in 1941, attempting to show that<br />

there was a difference between E. caespitosus and E. reichenbachii,<br />

he turned to some white, very greatly clustering specimens<br />

of E. baileyi sent him from Oklahoma and described them<br />

as the true E. caespitosus, rediscovered after all this time. He<br />

was impressed by the dense clusters of these specimens and<br />

thought Engelmann must have meant something like this in<br />

using the name he did for his plant. It is, however, obvious that<br />

Backeberg’s specimens could not be Engelmann’s E. caespitosus,<br />

since nothing like the Oklahoma plant grows within hundreds<br />

of miles of Industry, Texas, which was the type locality of E.<br />

caespitosus. It is equally obvious to anyone knowing Oklahoma<br />

cacti that Backeberg’s photo of his plant showed only a shortstemmed,<br />

clustering specimen of E. baileyi. Backeberg, in his<br />

more recent work, Die Cactaceae, has recognized this, and now<br />

calls this plant E. baileyi var. caespitosus. It seems unnecessary<br />

to maintain even this varietal distinction, as everywhere in the<br />

Wichita Mountains the whole range of variation exists together,<br />

from single-stemmed, tall specimens to greatly clustering<br />

plants. The largest cluster I myself have counted had 30 stems<br />

5 to 6 inches tall, and grew less than two feet from a plant,<br />

identical in every respect except that it was a full 8 inches tall<br />

and showing signs of old age without offering to ever become<br />

more than a single stem.<br />

Backeberg has also set up a set of varieties based almost entirely<br />

on spine color, as follows: E. baileyi var. brunispinus<br />

having long, chestnut-brown spines; variety flavispinus with<br />

soft and pale yellow spines; variety albispinus (Lahman) Backeberg<br />

having white spines; and variety roseispinus having long<br />

but soft spines, rose at the tips. This sort of thing appears to be<br />

useless in a case of quantitatively varying characters in which<br />

there are all possible intermediates. We could as well have eight<br />

varieties on color basis as four, once this sort of thing is started.<br />

If one can look beyond the minor differences, he will see<br />

whole hillsides of granite boulders in the crevices of which<br />

grow beautiful spiny columns and clusters of columns, from as<br />

red as the rocks themselves to as white as the snow which<br />

covers them here in the winter. This is E. baileyi, one of only<br />

two or three cacti unique to Oklahoma, and surely one of which<br />

the state can be proud.<br />

It is also necessary to report here what appears to be an even<br />

more rare and remarkable variant of this variable species. Years<br />

ago Oklahoma’s outstanding cactophiles, Mr. and Mrs. Charles<br />

Polaski, discovered a strange cactus on Headquarters Mountain,<br />

near Granite, Oklahoma. Only one individual was found,<br />

and much searching of the mountain has never revealed another.<br />

But Mr. Polaski is an expert at cactus culture, and he has propagated<br />

this individual vegetatively for many years by grafting.<br />

He most kindly gave me a cutting of it, as he has to others.<br />

The plant presents real peculiarities. The grafted cuttings<br />

have become cylindrical, clustering from the base, some stems<br />

up to 2 inches thick. The surface is in very indistinct ribs composed<br />

of almost separate tubercles. The areoles have 15 to 20<br />

short, spreading, whitish spines, but are more remarkable for<br />

having, besides the spines, a mass of fine, white wool. The<br />

whole plant remains covered with this woolly development,<br />

under the spines. I puzzled over my cutting of this plant for a<br />

number of years, while I tried to persuade it to bloom, but it<br />

finally died without responding that much to my care.<br />

In 1965, Curt Backeberg described this form as a new species,<br />

calling it Echinocereus mariae, and with his article is a photograph<br />

of its bloom, which appears much like the typical E.<br />

baileyi flower.<br />

What actually is this plant? After much reflection, I feel that<br />

this individual specimen found upon Headquarters Mountain<br />

is only an aberrant, atypical individual of E. baileyi, which<br />

grows in its typical form all over the mountain. Note that the<br />

spines are much like those of E. baileyi in everything except<br />

length-they are shorter. Note also that the growing tip of<br />

E. baileyi has the same sort of woolly areoles, the wool later,<br />

with maturity, falling off. Add to this the fact that the ribs in<br />

the growing tip of E. baileyi are, as in most cacti, markedly<br />

tuberculate, only flattening out later; that Backeberg’s photo<br />

shows a flower in no visible character contrasting with those<br />

of this species; and that the grafted cuttings greatly surpass the<br />

2.3 centimeters maximum he gives for the stem diameter of the<br />

original specimen, and approach the typical stem size of E.<br />

baileyi. Notice that all of these unique characters of this cactus<br />

can be interpreted as a retention of juvenile or immature form.<br />

This makes it seem that we have here a plant retarded in some<br />

way, and the flower, when it was finally seen, not being unique,<br />

I feel it should be considered an atypical individual of E.<br />

baileyi.<br />

I therefore do not list this as a separate species here. This<br />

decision is also influenced by the following consideration. It


28 cacti of the southwest<br />

does not seem proper to erect new species upon only one individual,<br />

and I understand that this plant was never re-collected.<br />

If the modern concept of the species as a population had<br />

been followed in the past, numerous names based upon one<br />

atypical specimen only would never have been proposed, and<br />

the study of cacti would be much simpler. While the knowledge<br />

of this specimen and its existence for all these years in several<br />

collections stands as a fitting tribute to the surpassing abilities<br />

of Mr. and Mrs. Polaski in collecting and culturing cacti, it<br />

seems the one individual can hardly constitute a formal taxon.<br />

Echinocereus albispinus Lahman<br />

Description Plate 6<br />

stems: Densely clustering. Individual stems are cylindrical<br />

arid 3 to 6 inches tall, while to only 1 inch thick, with 12 to<br />

14 narrow, tuberculate ribs.<br />

areoles: Oval, very woolly when young. The wool persists<br />

for a long time, but very old areoles are bare.<br />

spines: 14 to about 20 radial spines, not pectinate against the<br />

plant surface but deflected evenly outward all around the<br />

areole. They are very slender, the uppers the shortest, being<br />

only 3/16 of an inch or so long, while the laterals are from<br />

3/8 to 1/2 inch long. There are no centrals. All spines are pure<br />

white, or sometimes white with light brown, translucent tips.<br />

flowers: Very pale pink or rose-pink, 11/2 to 21/2 inches tall<br />

and 1/4 to 3 inches in diameter. They usually open very<br />

widely so that the petals are curved backward. These petals<br />

arise from very narrow, brownish bases and broaden only<br />

slightly to a maximum width of only 1/4 of an inch. The upper<br />

parts of the petals are whitish-pink. The upper edges of the<br />

petals are very ragged, but they are pointed at the apex. The<br />

Style is white, and there are 7 to 11 light green stigma lobes.<br />

The ovary surface has much white wool and clusters of 12 or<br />

more hairlike spines up to 1/4 of an inch long, in shades of<br />

white, grays, tans, browns, to black. These flowers are usually<br />

produced below the apex of the plant and may appear well<br />

down the sides of the stems.<br />

fruits: Almost spherical, 1/2 to 3/4 of an inch across, green,<br />

said to be edible. They split open when ripe.<br />

Range. The type locality of this species is near Medicine Park,<br />

Oklahoma, in the eastern end of the Wichita Mountains. It had<br />

not been found outside of the type locality until, during this<br />

study, it was collected on a single granite ridge a few miles<br />

northwest of Tishomingo, in Johnston County, Oklahoma.<br />

Remarks. When Mrs. Lahman made her study of the cacti of<br />

the Wichita Mountains, she spoke of most of her new forms as<br />

variations of E. baileyi, even while giving several of them new<br />

species names. But two forms which she discovered she set apart<br />

most definitely from that species. After listing her five variations<br />

of the E. baileyi species, she then said, “Also there are two<br />

others which appear to be separate species and are here described<br />

as E. purpureus and E. albispinus.” We have seen that<br />

her E. purpureus was in fact separate from the E. baileyi varieties,<br />

since it appears instead to be a form of E. caespitosus.<br />

Therefore I would assume that her E. albispinus is accurately<br />

set off from E. baileyi also, instead of being just the white form<br />

of that plant.<br />

The cactus which fits her description will not be confused<br />

with E. baileyi by any observer who sees it in a living condition.<br />

Its spines are shorter, and while they are not pectinate,<br />

they stand out around the areole evenly at the angle found on<br />

some specimens of E. caespitosus. It has the manicured appearance<br />

of that species and none of the disheveled look of E.<br />

baileyi. In fact, it is much nearer in appearance to a longspined<br />

E. caespitosus than to a short-spined E. baileyi. Even<br />

from Mrs. Lahman’s photograph this is clear.<br />

However, it seems distinct from both. Its mature stems are<br />

more slender as well as shorter than those of either of the others,<br />

and its flowers are different. They are smaller and the most<br />

pale in color of any related Echinocereus. The exaggerated way<br />

in which they open back until the petal tips almost touch the<br />

outside of the ovary is also unique, as well as the low position<br />

on the stem from which the flowers usually, but not always,<br />

sprout.<br />

E. albispinus is probably the most rare of Oklahoma cacti.<br />

We have seen no evidence that this cactus has actually been<br />

observed alive by any student since Mrs. Lahman until this time.<br />

Her description and photos of it are usually reprinted without<br />

any elaboration. Backeberg, however, quickly assumed that she<br />

referred to the common white-spined E. baileyi, and so appropriated<br />

her name for his series of color-dictated varieties,<br />

making it E. baileyi var. albispinus. Not only does this sort of<br />

variation based on color alone seem ill-founded, but the use of<br />

this name in this way ignores the already-mentioned differences<br />

which set this plant off from all the E. baileyi forms.<br />

Thanks to the aid of Drs. Bruce Blauch and Claude Gatewood<br />

(until recently of the Oklahoma State University), we have<br />

collected a number of specimens of a cactus which fits Mrs.<br />

Lahman’s description very well. We feel that it is distinct from<br />

any other and stands best as a separate species between E.<br />

baileyi and E. caespitosus.<br />

It was truly surprising when this cactus turned up on a single<br />

one of the many granite ridges near Tishomingo, Oklahoma. The<br />

area in which the plant grows in that location is only a few<br />

acres and the total population is probably no more than 100<br />

plants. This is fully 100 miles from the type locality of E. albispinus<br />

or from any known collections of E. baileyi. E. caespitosus<br />

grows in profusion not many miles away, but is not found<br />

associated with this plant. This is one of those surprising turns<br />

of distribution found so often in cacti.


genus Echinocereus engelmann 29<br />

Echinocereus chisoensis Marshall<br />

Description Plate 7<br />

stems: Columnar, to 8 inches tall, but remaining rather slender,<br />

the greatest diameter seen being two inches. They are<br />

almost always simple, but may, very rarely, branch above the<br />

ground. They have 13 to 16 ribs composed of very distinct<br />

tubercles almost completely separated from each other by<br />

broad valleys. The color of the surface is deep green or bluishgreen.<br />

areoles: Very small for the group, being about 1/8 of an<br />

inch or less across. They are circular and woolly at first, becoming<br />

oval and naked when older. These areoles are about<br />

1/4 of an inch apart.<br />

spines: There are 10 to 15 very slender radial spines which<br />

radiate evenly around the areole and lie almost parallel to<br />

the plant body. They are white or gray below with the upper<br />

part red-brown or maroon. The uppermost ones are very small,<br />

only 1/16 to 1/8 of an inch long, and bristle-like. The lateral<br />

ones are progressively longer, until the lower laterals are 1/4<br />

to 3/4 of an inch long. There are also 1 to 4 central spines.<br />

One is porrect or nearly so and 1/4 to 5/8 of an inch long,<br />

while the others are more or less spreading and shorter. They<br />

are all very slender and straight, from bulbous bases. They<br />

are black or dark red-brown, usually with whitish bases.<br />

flowers: Rose in color, with reddish centers, about 21/2<br />

inches long, but never opening widely, and so only 1 to 2<br />

inches in diameter. The petals are long and these would be<br />

big flowers of 3 or more inches across if they opened back,<br />

but the petals remain almost perfectly upright. They are oblong,<br />

the bases deep red and the upper parts rose, with entire,<br />

pointed tips. The pistil is short and white. The stigma is composed<br />

of 10 small, green lobes. The surface of the ovary has<br />

some white wool and clusters of 8 to 14 white to brownish,<br />

hairlike spines.<br />

fruits: Elongated, 1 to 13/8 inches long and about 1/2 inch<br />

in diameter. They are red and fleshy when ripe, but covered<br />

with wool and bristles. When older they become dry and<br />

split open.<br />

Range. Restricted to the Chisos Mountains within the Big Bend<br />

National Park, Brewster County, Texas.<br />

Remarks. This cactus was first described by Marshall in 1940.<br />

It is an obscure species, perhaps best described as retiring in<br />

both habitat and appearance. Even among those living and<br />

working at the Big Bend National Park, it is mostly unknown.<br />

The reasons for its remaining so little known are clear. It<br />

hides shielded in the middle of clumps of brush and never seems<br />

to dare to expose itself on ledges or open spaces. Among the<br />

stems of protecting bushes its small, upright column is well<br />

camouflaged and so is seldom seen. Even its flowers are strikingly<br />

reserved for an Echinocereus. They are as large as many of<br />

the flamboyant ones, but discreetly remain almost closed even<br />

during the few hours of the afternoon when they are open at<br />

all, showing only the pale rose color of the outer petal surfaces.<br />

There is bright color within them, but it is deep in the center<br />

where only the bees see it and it attracts no one else.<br />

Although almost unknown, the cactus is not rare in its area.<br />

Marshall spoke of examining over three hundred specimens, and<br />

I found it easy to locate dozens of them, once I knew how to<br />

search for them.<br />

The relationships of the plant to other Echinocerei are difficult<br />

at best to determine. Marshall discussed its similarity to<br />

some forms of E. fendleri, but concluded that they were only<br />

superficial. On the other hand, characters of the ovary surface,<br />

rib and areole arrangement, and spine form all relate it more<br />

closely to E. pectinatus or E. caespitosus than to E. fendleri.<br />

E. chisoensis is a delicate cactus, rarely seen, but a unique<br />

resident of the Big Bend National Park, where it is, fortunately,<br />

protected and so should be with us long after many of its more<br />

flamboyant relatives are decimated through a combination of<br />

conspicuousness and lack of protection.<br />

Echinocereus pectinatus var. wenigeri Benson<br />

“Comb Hedgehog”<br />

Description Plate 7<br />

stems: Single, or sometimes in old specimens 2 or 3 to one<br />

plant. They are egg-shaped to stoutly cylindrical, growing to<br />

10 inches tall and 31/2 inches thick. Rows of distinct but confluent<br />

tubercles make up shallow ribs on the stems. The num-<br />

ber of ribs on Texas plants is 13 to 18.<br />

areoles: Broadly oval and woolly when growing at the tip<br />

of the plant, but becoming narrowly oval or elongated and<br />

bare when older, on the sides of the stem. These areoles are<br />

from almost touching to 3/16 of an inch apart.<br />

spines: There are 15 to 20 radial spines which are pectinate,<br />

spreading over the surface of the plant, the laterals actually<br />

to some extent recurved back into the grooves between the<br />

ribs. They are medium strength to rather heavy and very<br />

rigid. The upper ones are most slender and short and the<br />

laterals longer, to between 3/8 and 1/2 inch. There are 2 or 3<br />

central spines standing in a vertical row in the center of each<br />

areole. They are stout, but very short, only 1/16 to 1/8 of an<br />

inch long. All spines are white with pinkish or purplish tips.<br />

flowers: Very large and striking in appearance, 3 to 5 inches<br />

tall and broad. The outer one-half or less of each oblong,<br />

broad, blunt petal is lavender pink in color. Below that is an<br />

expanse of white extending to the narrow bases of the petals,<br />

which are green. This gives a unique, three-colored flower,<br />

pink around the edges with a distinct white zone making up


30 cacti of the southwest<br />

sometimes half of the flower, followed by a greenish center.<br />

The stigma is whitish and supports 9 to 12 large, dark green<br />

stigma lobes. The ovary surface bears some short, white wool<br />

and 6 to 18 rigid spines, white or white with dark brown<br />

tips and measuring to 3/8 of an inch long.<br />

fruits: Spherical or nearly so, about 1 inch in diameter.<br />

Fleshy at first, becoming bronze or brown in color, after<br />

which they dry and split open.<br />

Range. A local form of a Mexican species encountered only<br />

occasionally in Texas, from Del Rio on the east to just beyond<br />

Sanderson on the west, in a strip not over twenty miles wide<br />

along the Rio Grande. It is not reported from the Big Bend in<br />

Texas, but has been reported from the southern edge of New<br />

Mexico, although the report has not been confirmed.<br />

Remarks. The species E. pectinatus grows widely in Mexico, but<br />

can be regarded as established in only two areas of the U. S. It<br />

is well represented in Arizona, one form coming from that state<br />

into the extreme southwest corner of New Mexico as well; and<br />

the form just described is rare but can be found in a limited<br />

strip of Texas from Del Rio to Sanderson. Strangely, it seems<br />

to be missing in the wide expanse of the border between these<br />

locations. This leaves the Texas form of it isolated from the rest.<br />

It is never common even in that area. A full day’s tramping<br />

over the hills just north of the Rio Grande in the vicinity of<br />

the lower Pecos and Devil’s rivers will yield at best one or two<br />

specimens, if the searcher has good eyes and better luck.<br />

The Texas range of the cactus begins just west and south of<br />

the southwesternmost range of E. caespitosus and extends west<br />

just to the beginning of E. dasyacanthus. It is sandwiched be-<br />

tween these its two closest relatives.<br />

The Texas specimens of this cactus have some differences<br />

from the typical specimens from Mexico, namely, fewer ribs;<br />

more oval, more woolly, and more widely spaced areoles;<br />

whiter spine coloring; and a lower maximum number of both<br />

radial and central spines, as well as heavier spines. At first, I<br />

did not distinguish it from the Mexican E. pectinatus, but after<br />

I became familiar with the Texas plants and after I had studied<br />

literally hundreds of Mexican specimens coming from that<br />

country through our Texas cactus dealers, and had collected<br />

the plant in various parts of Mexico myself, without finding<br />

one specimen to match ours, I became convinced that we have<br />

here our own distinct form. L. Benson erroneously refers my<br />

specimens of variety ctenoides from Mexico to this variety and<br />

so credits me wrongly with having collected the variety in<br />

Mexico as well as in Texas.<br />

The cactus may well be Hooker’s E. pectinatus var. texana,<br />

but there is no way to know, and, therefore, that name has not<br />

been recognized by any recent writers. L. Benson finally described<br />

and typified the form, naming it after the author. I<br />

feel it deserves varietal rank, as he indicated.<br />

E. pectinatus var. wenigeri can easily be distinguished from<br />

E. dasyacanthus because its radials are rigidly recurved and<br />

appressed to the surface of the plant instead of spreading outward,<br />

and because its centrals are very short, and are always<br />

in one vertical line instead of being longer and spreading outward.<br />

The typical Mexican E. pectinatus is actually much closer<br />

to E. dasyacanthus than is variety wenigeri. It is equally easy<br />

to distinguish from the typical E. caespitosus, which also has<br />

more outward-standing, finer spines, and often no centrals at<br />

all. But some specimens of E. caespitosus have very nearly as<br />

recurved radial spines and 1 or 2 short centrals in the middle<br />

of the areole. It is then very hard to distinguish the two by the<br />

plant body alone. But if one has the opportunity and patience<br />

to observe the flowers of the two the distinction is obvious:<br />

E. pectinatus presents a flower tube with short wool and short,<br />

rigid spines, while E. caespitosus always has long wool and<br />

long, flexible, hairlike spines on its flower. The white zone is<br />

also always present in the flower of E. pectinatus, while the<br />

flower of the other always lacks it, since its darker purple pet-<br />

als deepen to a reddish or greenish center.<br />

Echinocereus pectinatus var. rigidissimus (Eng.) Rümpl.<br />

“Arizona Rainbow Hedgehog,” “Cabeza del Viejo”<br />

Description Plate 7<br />

stems: Thick columnar to 8 inches tail and 4 inches thick.<br />

They may be single or rarely branched, with 18 to 23 nar-<br />

row, tuberculate ribs.<br />

areoles: Elongated, and from practically touching to 1/4 of<br />

an inch apart.<br />

spines: There are 15 to 23 radial spines which are pectinate<br />

and recurved to lie fiat on the surface of the plant. They are<br />

very heavy and rigid. The upper spines of each areole are<br />

small and a translucent tan, amber, or gray. The laterals and<br />

lower spines are to 5/8 of an inch long, tan or amber, amber<br />

with red tips, or sometimes all red. The plant usually presents<br />

a banded appearance due to successions of these spine colors.<br />

There are no central spines.<br />

flowers: Identical to those of previous form except the outer<br />

parts of the segments fuchsia and often having 13 stigma<br />

lobes.<br />

fruits: As those of the previous form.<br />

Range. Extreme southwest New Mexico and south Arizona in-<br />

to Sonora, Mexico.<br />

Remarks. This is now regarded as a variety of E. pectinatus. It<br />

presents the same large, white-zoned flowers and the same<br />

general appearance, being, however, distinguished from the<br />

typical form of the species by its heavier spines and its larger<br />

stem size. The spines are always all radials, and are the longest<br />

and heaviest of this species. They interlock, and the plant body<br />

is usually hidden under this thick, rigid armament lying flat<br />

Rumpl. -><br />

Rümpl.<br />

Veijo -> Viejp


profilic -><br />

prolific<br />

genus Echinocereus engelmann 31<br />

upon it. This seems also to be the only one of this group to<br />

present a variegation of the spine color on a single areole. Typically,<br />

the upper, smaller spines of each areole are lighter, while<br />

the lower ones are more or less red. These characters make it<br />

easily distinguishable from the other varieties of the species.<br />

It will be noticed that the pattern of coloration of the spines<br />

is almost the same as those of E. viridiflorus. Because of this I<br />

have seen this cactus confused by beginners with large, robust,<br />

short-spined specimens of E. viridiflorus var. cylindricus of the<br />

Big Bend in Texas. If closer observation does not clear up this<br />

error, the big purplish flower is surely a surprise when it appears<br />

where the little greenish one was expected.<br />

The flower of variety rigidissimus is identical with that of<br />

the species. It seems, however, to be a more prolific bloomer<br />

when cared for, and so is a great favorite of growers. Numerous<br />

photographs of this plant in its glory with many of the<br />

striking flowers open at once have been printed in various<br />

books and magazines.<br />

This cactus seems to be limited in its natural range to the<br />

mountains of the southwest corner of New Mexico and southeastern<br />

Arizona, coming up out of Mexico in only that one<br />

area. Coulter once stated that it was found in west Texas, but<br />

this seems doubtful. It is a beautiful form, just managing to<br />

enter the corner of our area.<br />

Echinocereus pectinatus var. ctenoides (Eng.)<br />

Description Plates 7, 8<br />

stems: Single or sometimes clustering to half a dozen stems,<br />

each heavily cylindrical and to 6 inches tall by 3 inches in<br />

diameter, having 15 or 16 ribs greatly interrupted by tubercles.<br />

areoles: As the species.<br />

spines: As the species except that the radial number is only<br />

14 to 22 and the centrals always number 2 to 4. The spines<br />

are white with very light brown tips. The plants are never<br />

purplish or banded with color.<br />

flowers: A large and showy orange-yellow, 21/2 to 4 inches<br />

long and wide. The petals are almost linear to narrowly<br />

spatulate, the ends erose. The upper part of each petal has a<br />

bright orange midline. The lower one third of each petal is<br />

green, and the center bright green. The style is greenish white,<br />

and there are 13 dark green stigmas. The outer ovary surface<br />

has a little short white wool and 14 to 16 short, rigid spines,<br />

which are white with dark brown tips, per areole.<br />

fruits: 1/2 to 11/8 inches in diameter, spherical or egg-shaped.<br />

They are green, turning to greenish-brown when ripe, and<br />

covered with short wool and rigid spines which, however,<br />

become deciduous when the fruit ripens.<br />

Range. Given originally as from Eagle Pass to the Pecos in<br />

Texas and south to near Santa Rosa, Coahuila, Mexico. It is<br />

said to exist westward in Mexico into Chihuahua. Apparently<br />

it is now extinct in Texas.<br />

Remarks. It is a real indication of the thoroughness of the early<br />

collectors and of Engelmann’s studies that this plant was collected<br />

and described over one hundred years ago. I am persuaded<br />

that none of the more recent students of cacti since<br />

Coulter have seen it at all, but this has not kept them from<br />

writing about it, and so the confusion is extensive, as can be expected.<br />

Engelmann described the cactus in detail and pictured it well,<br />

including the flower. He had specimens from Bigelow, collected<br />

at Eagle Pass and near Santa Rosa, Coahuila, and those of<br />

Wright, collected by the lower Pecos River, which is not far<br />

from Eagle Pass. He stated that the plant “looked distinct<br />

enough from C. dasyacanthus,” and then he stated that “the<br />

flowerless plant so closely resembles C. pectinatus that it can<br />

hardly be distinguished from it except by the fewer ribs.”<br />

In spite of this, later students have, probably because of its<br />

yellow flowers, tried to link this with E. dasyacanthus, and in<br />

their enthusiasm they have published some pictures of several<br />

different forms under this name. Backeberg finally grew bold<br />

enough to come out and call the cactus E. dasyacanthus var.<br />

ctenoides.<br />

In making this study, we recognized that this plant was one<br />

of the least known of all in our area and needed clearing up<br />

more than almost any other, so we early made attempts to collect<br />

and study it. For four years we made field trips regularly<br />

to the Eagle Pass vicinity and the area of the lower Pecos River<br />

looking for it. In this time I believe that I myself covered almost<br />

every undisturbed acreage along the Rio Grande from<br />

below Eagle Pass to the Pecos looking for this cactus, and it is<br />

not now to be found. We believe the form is now extinct on<br />

the north side of the Rio Grande—if indeed Engelmann meant<br />

to imply that it was on this side of the river in the first place.<br />

Still wanting to find out what the plant really was, we then<br />

extended our field trips into the adjacent part of Coahuila,<br />

working down toward the Santa Rosa Mountains, which was<br />

the other location where Engelmann reported the plant as growing.<br />

We found nothing for many miles below Eagle Pass, but in<br />

the rugged mountains northwest of Santa Rosa (now called<br />

Ciudad Muzquiz) we suddenly found stands of small, white<br />

Echinocerei almost exactly like E. pectinatus, except for their<br />

whiter spines. We noted that they were much more inclined to<br />

cluster than any E. pectinatus we had seen, and then, noting the<br />

regularly 15 or 16 ribs and 2 to 4 centrals, we began to hope<br />

that we had found ctenoides. When the plants bloomed, they<br />

presented large, light-orange flowers, and we had our confirmation.<br />

It is obvious, as it should have been from Engelmann’s description<br />

and his illustrations, that this is a form of E. pectina


32 cacti of the southwest<br />

tus. It bears no characters half so close to E. dasyacanthus, except<br />

the flower color, and we now know that that species has<br />

flowers ranging from yellow to magenta. With this we have the<br />

parallel situation of E. pectinatus with flowers from orange-<br />

yellow to fuchsia.<br />

The question of whether this cactus should even remain listed<br />

as a separate variety will doubtless now become the order of<br />

business. We do not separate the yellow, pink, and magenta-<br />

flowered strains of E. dasyacanthus this way. But it seems that<br />

this plant has other consistently maintained characters setting<br />

it off from its species: such as its greater clustering tendency,<br />

its smaller size, its fewer ribs, and so forth.<br />

We apparently are faced here with the first cactus form rendered<br />

extinct from our area of the United States in the past<br />

hundred years. Very much of the land around Eagle Pass is<br />

farmed and the majority of the range areas which look undisturbed<br />

have actually been cleared in one way or another in the<br />

past, so this is not really surprising. But may we take warning<br />

from this and cherish our other cacti found in limited areas<br />

more actively?<br />

Echinocereus dasyacanthus Eng.<br />

“Texas Rainbow Cactus,” “Golden Rainbow Hedgehog,”<br />

“Yellow-Flowered Pitaya”<br />

Description Plate 9<br />

stems: Oval at first, soon becoming cylindrical. They grow<br />

to a maximum of 14 inches tall and 4 inches thick. These stems<br />

often remain single, but old plants quite often branch to<br />

form several heads. They have 12 to 21 narrow, tuberculate<br />

ribs.<br />

areoles: Round and with tan wool at the growing tip of an<br />

active stem, becoming oval or elliptical and bare when older.<br />

They are 1/8 to 3/8 of an inch apart.<br />

spines: There are 15 to 25 radial spines per areole, all spreading<br />

outward at various angles from the surface of the plant<br />

and interlocking with those of adjacent areoles, so that the<br />

plant surface is heavily covered by them. They are rigid and<br />

of medium thickness, the upper ones very short and the laterals<br />

to 1/2 or even 5/8 of an inch long. There are also 2 to 5<br />

robust centrals usually 1/8 to 1/2 inch long, but occasionally<br />

3/4 or even 1 inch long. These centrals stand spreading in all<br />

directions from the center of the areole, and are not lined up<br />

in a vertical row. All spines are white, yellowish, or tan,<br />

with their tips reddish or rust-brown. The variations in color<br />

shades are often found in zones up and down an individual<br />

plant, giving the typical specimen distinct bands of color,<br />

although some specimens are of one unvarying color and so<br />

are not banded at all.<br />

flowers: Very large and showy. 3 to 51/2 inches long and<br />

wide, in colors they range from yellow through pink to violet<br />

and magenta. The very long petals are spatulate, with the<br />

ends ragged and variously notched or pointed. Their bases<br />

are green. The style is long and white. There are 12 to 22<br />

large, deep green stigma lobes. The long flower tube has some<br />

short wool and 7 to 18 rigid spines to 1/2 inch long and pure<br />

white to white with reddish tips upon each areole.<br />

fruits: Spherical, 1 to 2 inches in diameter. They are covered<br />

with spines until ripe, when these become deciduous and the<br />

fruit becomes red-brown or purplish.<br />

Range. Southeastern New Mexico through southwestern Texas<br />

and the Big Bend. On the east it extends to near Sanderson and<br />

Fort Stockton, and into the Guadalupe Mountains of Texas. It<br />

follows these mountains a short distance north into New Mexico.<br />

It is common in the Big Bend and around El Paso, Texas.<br />

It has been said to extend west all the way into Arizona, but I<br />

cannot verify this.<br />

Remarks. E. dasyacanthus is the largest member of this group,<br />

the most robust in every feature. It likes to stand unprotected<br />

by the shade of any other plant on the thin soil of rocky slopes,<br />

where its thick spine cover gives it protection from the severe<br />

elements as well as from all enemies. A colony of fine old plants<br />

on a Big Bend hillside, many of them to at least a foot tall and<br />

varying from almost white to reddish or rusty brown, often<br />

banded with these colors, is a proud and handsome sight. They<br />

bloom with huge flowers, often opening well down on the sides<br />

of the plants. These flowers come in all shades from pale yellow<br />

to reddish, pink, and even deep magenta.<br />

It seems obvious that no varieties can be set up within this<br />

species on the basis of flower color. Other variations found<br />

within the limits of this species, such as banding or lack of<br />

banding with color, do not seem any more significant. Miss<br />

Clover set up the name E. steereae for the whitish, nonbanded,<br />

violet-flowered population found in the Chisos Mountains of<br />

the Big Bend, but this does not seem warranted. Rumpler tried<br />

to make this E. pectinatus var. steereae, but the radials are not<br />

pectinate, nor are the areoles elongated. If made a variety, it<br />

would have to be E. dasyacantha var. steereae, as Backeberg<br />

lists it, but even this does not seem supportable since we would<br />

have only the flower color to distinguish it.<br />

Whether or not the limits of this species have been drawn<br />

widely enough is another question. Coulter found a cactus somewhere<br />

in southeastern New Mexico which he called E. dasyacanthus<br />

var. neo-mexicanus (note that this is not the same as<br />

Wooton’s E. neo-mexicanus). Coulter described his plant as having<br />

areoles 3/8 to 9/16 of an inch apart, with stouter, “spreading<br />

radials and 4 stout centrals and larger seeds.” He apparently<br />

did not see the flowers of this plant.<br />

No one has since been sure what this cactus of Coulter’s<br />

really was, and the name should probably be eliminated, since<br />

it seems impossible to know. Most have found it irresistible to<br />

speculate, however, and I would add my own theory. Coulter<br />

Glover -><br />

Clover<br />

Rumpler: see<br />

bottom of page<br />

“ E. pectinatus v. steereae Rumpler” is an impossible<br />

combination. Probably a name confusion


genus Echinocereus engelmann 33<br />

did not know that E. dasyacanthus could have any flower color<br />

except yellow. He even used this flower color to key this plant<br />

from its relatives. So he automatically set both the species and<br />

his variety of it off from all of its red or purple-flowered<br />

relatives and apparently never thought to compare characters<br />

between the two groups he had set up, yellow-flowered and<br />

purple-flowered. Since we now know that this division will<br />

not stand, when we disregard flower color, we find that the<br />

characters of his E. dasyacanthus var. neo-mexicanus and those<br />

of the red-flowered E. roetteri are practically identical. Remembering<br />

that they are similar enough so that Engelmann had<br />

first called E. roetteri by the name of E. dasyacanthus var.<br />

minor, and that Coulter did not know the flower color of his<br />

variety, I find it likely that his variety neo-mexicanus was<br />

nothing but E. roetteri. Beyond this sort of speculation we can-<br />

not go.<br />

Benson decided to combine E. dasyacanthus with E. pectinatus<br />

as a variety of that other species, and since by the rules of<br />

nomenclature, one must use an earlier variety name for such a<br />

combination if one exists, he calls our cactus E. pectinatus var.<br />

neo-mexicanus in place of E. dasyacanthus. Even in doing this,<br />

he commented that this was unfortunate, and so it surely is, as<br />

it would leave us for our most well-known rainbow cactus only<br />

an obscure name which refers to no one knows what for sure.<br />

It seems that the species is distinct enough from E. pectinatus<br />

anyway, and the combination need not be made in the first<br />

place.<br />

Echinocereus dasyacanthus var. dasyacanthus (Eng.)<br />

Description Plate 9<br />

stems: As the species.<br />

areoles: As the species, except always round.<br />

spines: As the species, except that the centrals are not seen<br />

less than 3/8 of an inch long.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. Southeastern New Mexico and southwestern Texas east<br />

to the Guadalupe and Davis mountains and to near Sanderson,<br />

Texas. From these areas it ranges into Mexico. It is also said to<br />

range west into Arizona, but I have been unable to confirm<br />

this definitely.<br />

Remarks. This is the common, typical form of the species which<br />

is so widely appreciated by almost everyone who looks at cacti<br />

at all. It is a dominant part of almost every collection from<br />

western Texas or southern New Mexico. However, most specimens<br />

grown in gardens come from along the highways instead<br />

of from farther south in the nearly inaccessible mountains along<br />

the Rio Grande, and so most people see only yellow-flowered<br />

individuals.<br />

There is actually a gradient of flower color in the wild population.<br />

Nearly all plants growing in New Mexico and around<br />

El Paso bloom yellow or orange-yellow. In the Big Bend of<br />

Texas pink is more common; and in the lower and eastern part<br />

of the Big Bend red and purplish are almost the rule. The full<br />

range of colors is found south and west of Sanderson, with<br />

some populations there being almost entirely magenta. I saw a<br />

beautiful collection of over one hundred local specimens in a<br />

rock garden in Sanderson in which all the range of colors were<br />

blooming together, but with yellow the most rare. Each specimen<br />

I have seen has presented only one color of flower, but<br />

Marshall reported seeing a specimen bloom with yellow and<br />

purple flowers simultaneously.<br />

Echinocereus dasyacanthus var. hildmanii Arendt<br />

Description Plate 9<br />

stems: Ovate to tapering cylindrical, single at first but soon<br />

clustering to at least 6 or 8 stems, each of these to 10 inches<br />

tall and 3 inches in diameter. The flesh is dark green, and<br />

the stems have 12 to 16 very tuberculate, narrow ribs.<br />

areoles: Round to oval, woolly when young, then bare.<br />

They are 3/16 to 1/4 of an inch apart.<br />

spines: 15 or 16 radials to 3/8 of an inch long, white or gray<br />

at the bases with maroon or brownish tips. There are 3 to 5<br />

centrals 1/8 to 5/16 of an inch long, from the same color as the<br />

radials to completely maroon or dark red-brown. The spines<br />

do not band the plant with color.<br />

flowers: 4 inches tall, 31/2 inches across, deep orange-yellow,<br />

with 15 to 19 stigmas. Otherwise they are as the species.<br />

fruits: Not observed.<br />

Range. The Davis and Apache mountains of west Texas to near<br />

Pecos, Texas.<br />

Remarks. This form is very close to E. dasyacanthus, yet is sufficiently<br />

distinct from it to be set apart from it even by amateurs<br />

who don’t have any name for it. It is never a rainbow,<br />

having no banding of colors. The fewer spines do not cover<br />

its body so entirely, so the dark green flesh is always visible.<br />

Although the individual stems are smaller, the cactus is definitely<br />

a clustering one. I have actually yet to see an unbranched<br />

specimen, and the largest I have seen had 8 stems loosely clumped<br />

in a cluster well over a foot in diameter. This sort of growth<br />

is never seen in the typical E. dasyacanthus. Yet the flowers of<br />

the two are almost indistinguishable, and the other character differences<br />

between them are so minor that it seems no more than<br />

a variety.<br />

Since Arendt described it in 1892, it has, probably, not been<br />

seen by any student until now. Schumann did not mention it,<br />

and Britton and Rose seem to have confused it with E. fendleri


Rumpl -> Rümpl<br />

34 cacti of the southwest<br />

var. bonkerae, in which Backeberg followed them. It should be<br />

noted that their description when discussing the plant is of specimens<br />

collected in the Santa Catalina Mountains of Arizona,<br />

and so not of this plant. Arendt described it definitely as a<br />

Texas cactus. It is a rare form, and limited to the Davis and<br />

Apache mountains and the hills northeast of them to near the<br />

Pecos River.<br />

Echinocereus roetteri (Eng.) Rümpl.<br />

Description Plate 10<br />

stems: Single, at first rather egg-shaped, but becoming cylindrical,<br />

up to 6 inches tall and about 3 inches in diameter. The<br />

surface is bluish or grayish-green, and there are 10 to 13 ribs<br />

which are composed of definite tubercles.<br />

areoles: Oval or sometimes round, with some tan wool when<br />

young, but naked when old. These areoles are 1/4 to 1/2 inch<br />

apart.<br />

spines: There are 8 to 15 radial spines, the variation due to<br />

the plant’s frequent addition of very tiny bristle-like radials<br />

at the top of the areoles. These upper radials are only 1/8 to<br />

3/16 of an inch long. The lateral radials are of medium thickness,<br />

straight, and up to 1/2 inch long. The lowermost radial<br />

is a little shorter and weaker than the laterals. There are also<br />

2 to 5 centrals which are stout and straight, 3/8 to 1/2 inch<br />

long, and spreading in all directions. All the spines are opaque<br />

and ashy brown or almost maroon, with the tips blackish,<br />

and their bases are bulbous.<br />

flowers: Brilliant purplish in color, but only 2 to 3 inches<br />

long and not opening widely.<br />

fruits: Almost round to elongated egg-shaped, and small,<br />

being only 1/2 to 7/8 of an inch long.<br />

Range. Originally given by Engelmann as from El Paso, Texas,<br />

south in the sand hills, and said by Coulter to range from there<br />

west into Arizona and south into Chihuahua. But the only records<br />

in recent times have been from southeastern New Mexico.<br />

Remarks. This is a very obscure and difficult cactus. It has been<br />

surrounded by confusion from the beginning. Engelmann first<br />

described it as E. dasyacanthus var. minor, but later withdrew<br />

that connection and called it Cereus roetteri. Coulter probably<br />

studied it more thoroughly than anyone else, having at hand<br />

plants from Texas, Arizona, and Chihuahua. Britton and Rose<br />

seem to have been misled by Engelmann’s first linking of the<br />

plant with E. dasyacanthus. Their specimen collected at El Paso<br />

is a tiny thing only 11/2 inches tall, which seems to have no centrals<br />

at all, and appears to be a seedling of E. dasyacanthus.<br />

Their specimen from Arizona has 1 or 2 very short centrals,<br />

apparently in a row, and appears to be one of the E. pectinatus<br />

group. At any rate, since I have seen living plants of E. roetteri,<br />

their preserved specimens do not seem to me to be this plant.<br />

This may explain why their description of the cactus is erroneous<br />

in several respects. More recently Backeberg has realized<br />

the close relationship of this cactus with E. lloydii, and com-<br />

bines these two, calling that other cactus E. roetteri var. lloydii.<br />

It does seem obvious that these latter two are closely related.<br />

How closely is not clear. However, on the basis of the very few<br />

specimens of this rare cactus I have seen, I feel it is too soon to<br />

combine them. E. roetteri seems to be a much smaller, nonclustering<br />

form with other vegetative characters strikingly similar to<br />

the more robust E. lloydii. But the flowers and fruits of E.<br />

roetteri are very much smaller than those of the other, and its<br />

flowers show none of the firm, lasting characters of the claret-<br />

cup group, which those of E. lloydii display.<br />

E. roetteri is today an extremely rare cactus. The few known<br />

specimens recently collected all have come from southern New<br />

Mexico. It is surprising that with the large amount of recent<br />

study of Arizona cacti this one has not been reported there since<br />

Coulter. It is not so surprising that it is not seen around El<br />

Paso, which is its type locality, since the real estate, farming,<br />

and military developments there have already greatly reduced<br />

several much more common species of the area. For this reason<br />

it appears that it cannot be found in Texas today, surviving only<br />

farther west, in less disturbed areas.<br />

Echinocereus lloydii B. & R.<br />

Description Plate 11<br />

stems: Simple at first, but clustering and branching slowly<br />

to form clumps of up to about six stems when old, the single<br />

stems attaining a large size before clustering. Mature stems<br />

grow to 12 inches high and as much as 41/2 inches thick, and<br />

are cylindrical and bright green to gray-green in color. The<br />

ribs are 11 to 13 in number, broad, interrupted, and extremely<br />

tuberculate, being formed of broad thickenings at the areoles.<br />

There are deep furrows running down the sides of the ribs between<br />

these broad bases of the tubercles. The vertical furrows<br />

between adjacent ribs on the older parts of the stem are the<br />

deepest I have seen on any Echinocereus, being up to at least<br />

3/4 of an inch deep.<br />

areoles: Medium to large, oval to circular, with much white<br />

wool when young, but becoming practically bare with age.<br />

They are 1/2 to 5/8 of an inch apart.<br />

spines: When mature all ashy-gray to reddish-gray, but when<br />

young and growing they are a brilliant purplish-red, this color<br />

often remaining for many years on the tips of the spines. All<br />

spines are of medium stoutness, round, and straight from<br />

bulbous bases. There are 14 to 17 radial spines, the lower<br />

and lateral ones being 1/2 to occasionally 7/8 of an inch long.<br />

The upper radials are very much more slender, and much


genus Echinocereus engelmann 35<br />

shorter-down to 1/4 of an inch long. There are 4 to 8 central<br />

spines identical to the larger radials standing perpendicular<br />

to the stem and spreading slightly from the crowded<br />

center of the areole.<br />

flowers: Large and very beautiful, usually being 3 inches<br />

long and 2 to 31/2 inches in diameter. There is much variation<br />

in their color. They are most commonly scarlet, but I have<br />

specimens with petals coral-pink instead. The outer petals are<br />

greenish in the midline with entire edges and pointed tips.<br />

The inner petals are long, spatulate, with entire edges although<br />

these are sometimes somewhat notched at the tips.<br />

There are 9 to 14 green stigma lobes. The filaments are pinkish,<br />

the anthers extremely small and pinkish or rose in color.<br />

The ovary tube bears some white wool and clusters of 7 to 12<br />

firm, reddish spines which are almost equivalent in arrangement<br />

to those on the stem areoles, except that they are more<br />

slender.<br />

fruits: Egg-shaped and 11/4 to 2 inches long. They are densely<br />

covered with spines; when the fruit is ripe, these spines<br />

loosen fairly easily, but do not seem to fall unless brushed<br />

off. The ripe fruit is greenish-orange in color and fleshy.<br />

Range. Known only from the type locality, Tuna Springs,<br />

Texas, which is about 20 miles east of Fort Stockton in Pecos<br />

County.<br />

Remarks. This cactus was named by Britton and Rose in 1922<br />

from plants collected by Mr. F. E. Lloyd, in his honor. Mr.<br />

Lloyd collected many cacti in the area around Tuna Springs,<br />

and there was no other label of location on his plants but that.<br />

Tuna Springs has not existed as a town and is not on most<br />

Texas maps, so during all these years it has been impossible for<br />

interested people to go and find this cactus. The long period<br />

of time in which the plant has been lost has not been completely<br />

unfortunate, because this has meant that no one has<br />

collected it. Consequently it has grown undisturbed and rather<br />

commonly in its small area, when most of our rare species<br />

whose locales have been known are nearing extinction. But recently<br />

the fact that Tuna Springs was once a stagecoach stop<br />

a short way east of Fort Stockton has become known, and dealers<br />

have been rapidly bringing out the plants. Its continued<br />

survival under these conditions is doubtful.<br />

This is a heavy-stemmed, clustering Echinocereus possessing<br />

an interesting set of characteristics, which make it unique. In<br />

areoles, spines, and stem shape it is very close to the scarlet-<br />

flowered E. triglochidiatus-E. polyacanthus-E. coccineus group,<br />

and its flowers show certain similarities to these (such as in<br />

the pink anthers). Yet there are definite differences from these<br />

and the flowers are more purplish and less waxy or longlasting<br />

than those of the claret cups, making it appear rather<br />

intermediate between those and the purple-flowered groups.<br />

The theorists will surely play with this one, but in the meantime,<br />

it is a rare and beautiful Texas cactus recently redis-<br />

covered.<br />

Echinocereus triglochidiatus Eng.<br />

“Claret-Cup Cactus,” “Strawberry Cactus,” “King’s Cup<br />

Cactus”<br />

Description Plates 11, 12<br />

stems: Globular to cylindrical in shape with 5 to 9 broad,<br />

rounded ribs having wide, shallow grooves between them.<br />

These stems may be from 3 to 18 inches in maximum length<br />

by 21/2 to 41/2 inches thick. The surface of the stems is very<br />

soft, often markedly wrinkled, and from dark to pale green<br />

in color. These stems cluster in different numbers in different<br />

varieties, usually producing fewer than a dozen heads, but in<br />

one form an old plant occasionally becomes a large mat of<br />

up to around 50 heads. In all cases, however, the stems are<br />

loosely and irregularly clustered, a plant being made up of<br />

various-sized stems, standing or partly reclining in an irregular<br />

clump.<br />

areoles: Circular, varying greatly in size within the species<br />

from no more than 1/8 of an inch to sometimes over 1/4 of an<br />

inch in diameter. They also vary in distance from one<br />

another. The areoles always have much white wool when<br />

young, but usually become bare in age.<br />

spines: Yellowish or red when growing, becoming opaque,<br />

ashy-gray to almost black when old. They vary within<br />

the species in number, size, and shape, the range being as<br />

follows: radials from 2 to 9 in number and centrals 0 or 1,<br />

all these variously from 3/8 to 21/2 inches long, from rather<br />

slender to very thick, rounded to greatly flattened, angular,<br />

or channeled, as well as either straight or curved.<br />

flowers: 2 to 23/4 inches long and 1 to 11/2 inches in diameter,<br />

rigid, waxy, and remarkable among the cacti for their<br />

persistence, often staying open for several days and nights.<br />

The stiff, blunt petals are scarlet-red or orange-red from narrow<br />

green or whitish bases. The stamens are as long as or<br />

slightly longer than the petals, with filaments greenish below,<br />

becoming fuchsia above. The anthers are very tiny and<br />

fuchsia. They are about the same length as the style, so that<br />

they sometimes partly enclose the stigma, which has 5 to 11<br />

green lobes. The ovary surface has small areoles upon it, each<br />

with a fleshy scale-like segment, a little white wool, and 2<br />

to 6 slender, white, or white-tipped brown spines.<br />

fruits: Varying within the species. They are round or oval<br />

and from 3/4 to 1/2 inches in largest measurement, somewhat<br />

tuberculate to practically smooth, with some spines which<br />

usually become deciduous with ripening. In color they are<br />

from green to green with a pinkish cast or else bright red<br />

when ripe.<br />

Range. Taken as a species, this plant ranges over a very wide<br />

area from near Kerrville in central Texas, west through all of<br />

the Texas Big Bend to near El Paso, then north through central<br />

New Mexico into Colorado and northwestern Arizona.<br />

Remarks. There has probably been more confusion about E.


36 cacti of the southwest<br />

triglochidiatus than about any other Echinocereus. It is mostly<br />

due, I believe, to the fact that this name is the oldest among<br />

the red-flowered Echinocerei. Because of this the descriptions<br />

given under the name have been broadened from time to time<br />

to include every new form which has been put with it in combination.<br />

This process has gone on until the extreme is reached<br />

by L. Benson’s description in the Cacti of Arizona, where he<br />

describes E. triglochidiatus as having stems from 8 to 24 inches<br />

high and spines from as few as 3 to as many as 16 per areole.<br />

This sort of description is obviously drawn up to include everything<br />

in the red-flowered Echinocereus group. It really has<br />

little to do with the E. triglochidiatus as it was described by<br />

Engelmann. Since we have all degrees between these two extremes<br />

in the literature, it is not hard to see why the confusion<br />

is so great. There will never be anything but confusion here<br />

until we see how this problem came about.<br />

There exists a whole series of Echinocerei all of which have<br />

flowers similar in that their firm, long-lasting petals are scarlet-red<br />

with no blue pigments in them and their stamens light<br />

magenta. They are often called the “claret cups” because of this<br />

remarkable flower coloration. The shape of the flower and its<br />

parts is essentially the same in all of these forms, with the only<br />

difference between their flowers being minor variations in the<br />

proportions of some of the floral parts.<br />

In size, manner of growth, and details of stem and spines,<br />

however, these forms show easily as much variation as is found<br />

between many other recognized cactus species, and far more<br />

than between some. To give a general description broad enough<br />

to encompass all of them is almost to repeat the broad characteristics<br />

of the entire clustering, sparsely ribbed group of Echinocerei.<br />

This problem has caused disagreement as to how to treat these<br />

forms from the very start. On the one hand, there have been<br />

authorities who have described them all as separate species<br />

because of their diversity of stem and spine characters; on the<br />

other hand some have placed most of them together as varieties<br />

within large species groupings because of their flower similarities.<br />

A few have even proposed uniting them all into one large,<br />

variable species complex. Almost every authority has his own<br />

system at least slightly different from every other, often with<br />

different names for the same plants—and sometimes the same<br />

author has changed his system from publication to publication.<br />

All of this makes E. triglochidiatus one of the most difficult<br />

groups of cacti to understand. Synonymous names are often<br />

taken to mean different plants, and on the other hand, doubtful<br />

combinations have been made where some definite characters<br />

or other have been overlooked. This has led to the reporting of<br />

some of these forms from areas, and even from states, where<br />

they do not really occur. On the basis of this sort of thing,<br />

several of them have been almost lost sight of altogether. It has<br />

even been maintained by some that they cannot really be distinguished<br />

at all, but intergrade entirely.<br />

After much observation of the plants in their habitats and<br />

after study of available herbarium materials and the literature<br />

on them, I have concluded that, once erroneous and incomplete<br />

descriptions and doubtful records based on these are eliminated,<br />

most of the major forms long ago described in this group do<br />

exist within definite ranges and can be distinguished by con-<br />

stant characters.<br />

What are they then? Most of them are too widespread to be<br />

clones. Several of them are found in identical or very similar<br />

environments in overlapping ranges, and I have seen them preserve<br />

their distinctive characters perfectly in the uniform environments<br />

of gardens and greenhouses. Therefore, they do not<br />

seem to qualify as ecotypes or environmental modifications of<br />

a single taxon. This leaves their differences as rather clearly<br />

genetic, and the crucial questions here would be: What are<br />

their genetic relationships? Are they close enough genetically to<br />

be varieties or distinct enough, after all, to be species? But these<br />

are the questions no one can yet answer, because of a twofold<br />

lack. First, there is no standard degree of genetic relationship,<br />

at least in cacti, one side of which is a variety and the other<br />

side of which is a species. The thing has just not been worked<br />

out to this degree as yet. And, more important, we know nothing<br />

so far concerning the genetic relationships of the Echinocerei.<br />

There has been absolutely no biosystematic research reported<br />

upon them; not even chromosome numbers are known<br />

for these forms. So at the present time no one can say categorically<br />

from any research data that these are either species<br />

or varieties. Any decision is a purely arbitrary, philosophical<br />

one.<br />

So we continue to get different treatments of these forms determined<br />

entirely by each author’s concept of species. This accounts<br />

for the highly confusing spectacle we have today of two<br />

leading authorities on cacti so completely at odds that while<br />

Benson regards the claret cups all as varieties of one vastly enlarged<br />

species, Backeberg treats them just as confidently as<br />

separate small species. It is still no more than a philosophical<br />

argument.<br />

Faced with the necessity of taking a stand on this question,<br />

I am following a middle course. It seems clear that some of<br />

these forms are too close to be totally separate species. I am,<br />

therefore, following Benson in listing as varieties under the<br />

species name, E. triglochidiatus, all those forms which are close<br />

enough to the typical form that their inclusion does not cause<br />

the original species description to be altered basically. But I<br />

agree with Backeberg that the lumping of everything with the<br />

one characteristic of a firm, long-lasting flower into one species<br />

is unnecessary, and so I am leaving separate from that species<br />

all of those forms whose vegetative characters are so far removed<br />

from the original, typical form that their inclusion would<br />

cause the species to become something basically different from<br />

the original. This decision, like any made at this time about this<br />

group, is arbitrary, and we look forward to the day when someone<br />

will be able to give us experimental evidence to decide these<br />

questions.


genus Echinocereus engelmann 37<br />

From this position, then, we move on to discover and understand<br />

the varieties of this fine species.<br />

Echinocereus triglochidiatus var. triglochidiatus (Eng.)<br />

Description Plate 11<br />

stems: Clustering or branching only slowly to form small<br />

clumps often of no more than two or three stems, with an<br />

apparent maximum of a dozen or so stems per plant. The<br />

stems are of unequal lengths and loosely clustered, each one<br />

cylindrical, dark green, and usually somewhat wrinkled, the<br />

largest being up to about 8 inches high and about 3 inches in<br />

diameter. There are usually 7 ribs, but occasionally there may<br />

be 6 or 8, and they are broad, with slight swellings at the<br />

areoles and with very shallow grooves between them.<br />

areoles: Circular, but not large for the group, with white<br />

wool persisting on them, 7/8 to 11/2 inches apart, these being<br />

the more widely spaced areoles found in this group.<br />

spines: Ashy-gray to almost black, from 1/2 to 21/2 inches<br />

long. They are very stout, most of them being about 1/8 of an<br />

inch in diameter for most of their lengths. They have very<br />

much enlarged bases and are very flattened or angled, with<br />

distinct ridges or grooves running the length of the spines, the<br />

top surfaces of them even being concave in most cases. Some<br />

of the spines are straight, but some of them at least are curved,<br />

in every specimen I have seen. There are 2 to 6 spines to an<br />

areole, all of them radials, but a fair share of the areoles on<br />

any plant usually have the 3 spines from which comes the<br />

name of the plant.<br />

flowers: As the species, except to 11/4 inches long; the petals<br />

broadest at the tips and very blunt.<br />

fruits: Oval or egg-shaped, 1 to 11/4 inches long and 3/4 to<br />

1 inch in diameter. They are tuberculate. Fruits I have watched<br />

remained green until about January, when they rotted<br />

without coloring. The areoles were on the upper ends of long,<br />

broad tubercles about 1/8 of an inch wide at the top and 1/2<br />

inch wide at their bases. Each areole possessed a pinkish,<br />

fleshy scale and 2 to 5 stout, persistent spines which showed<br />

under magnification the same angled, channeled, and twisted<br />

character as the plant’s major spines. Britton and Rose, as<br />

well as Boissevain, have described the fruits as red, with<br />

deciduous spines, but it is worth remembering that these authors<br />

had already combined this with other forms known to<br />

have red fruits, so their descriptions may be of those other<br />

forms. There is no description of the fruits of this form by<br />

earlier authorities before the confusion of combinations had<br />

begun to enter the picture.<br />

Range. Northern New Mexico from near the upper Pecos River<br />

west to just beyond the Arizona boundary and north into Col-<br />

orado. Its southern limit seems to be near Albuquerque, New<br />

Mexico.<br />

Remarks. In the process of combination which has been practiced<br />

the form originally described by Engelmann as E. triglochidiatus—with<br />

its stems only up to 8 inches high and its spines<br />

very long, very heavy, greatly angled, and all radials—has almost<br />

been lost sight of, as witnessed by the fact that its range<br />

is often given today as being from Arizona throughout New<br />

Mexico and trans-Pecos Texas. When the form originally circumscribed<br />

by Engelmann is considered, however, I find it to<br />

be the form described above, and this specific cactus has a definite<br />

range much less wide than is often ascribed to it, due to<br />

the inclusion of other forms under the too-broad descriptions.<br />

When this is once understood, the confusion vanishes.<br />

The easternmost report of this plant is still Engelmann’s report<br />

of it on the Gallinas River just east of the upper Pecos, which<br />

would be near Las Vegas or Anton Chico, New Mexico, and the<br />

westernmost report of it from Fort Defiance, just within upper<br />

Arizona. It seems most common about the mountain slopes from<br />

Albuquerque to the Gallup area of New Mexico and north<br />

from there into south-central Colorado. Coulter cites a supposed<br />

collection of it in Texas by Wislizenus in 1846, and on the<br />

strength of that places its range as extending that far into Texas,<br />

but it is significant that Engelmann in 1856, having worked<br />

over Wislizenus’ material, did not mention any collections of it<br />

in Texas. This one Texas report may have led later students to<br />

mistake for it variety octacanthus, which does grow widely in<br />

Texas, and may be the reason for the survival of the idea that<br />

it grows in Texas. In much collecting throughout far west Texas<br />

and in lower New Mexico, I have never seen it growing there,<br />

nor found anyone who had, in spite of the fact that thousands<br />

of plants have been shipped out of Texas erroneously called by<br />

this name.<br />

The plant is a handsome cactus, one of New Mexico’s finest.<br />

It is very hardy, and its dark green looks its best half buried<br />

in a snowbank. It is, of course, most beautiful in the spring<br />

when it has its fiery red flowers. Their character, shared with<br />

the other red-flowered Echinocerei but unique among other<br />

cacti, of staying open for several days and nights at a time,<br />

makes the plant a favorite of all.<br />

Echinocereus triglochidiatus var. gonacanthus (Eng.) B. & R.<br />

“Claret-Cup Cactus,” “King’s Cup Cactus”<br />

Description Plate 12<br />

stems: Cylindrical, clustering very slowly and sparingly,<br />

usually only 2 or 3 stems to a plant and the most I have seen<br />

in one cluster being 6. Each stem has 7 to 9 ribs which are<br />

somewhat rounded, with shallow grooves between them and<br />

with distinct, rounded swellings of the ribs at the areoles.<br />

There are two separate populations of this form identical ex-


38 cacti of the southwest<br />

cept for size. In the widespread northwestern population the<br />

stems are only 3 to 6 inches high and to about 21/2 inches<br />

thick, but there is a localized population restricted to the<br />

vicinity of the White Sands National Monument, where the<br />

stems grow to a maximum of at least 18 inches tall by 41/2<br />

inches thick.<br />

areoles: These are very large for the group, 1/4 to 3/8 of an<br />

inch in diameter, with a great deal of wool when young, but<br />

losing much of it with age. They are about 1/4 to 3/4 of an<br />

inch apart.<br />

spines: The spines of this cactus are most striking. They are as<br />

heavy as or heavier than those of variety triglochidiatus, and<br />

there are more of them. At least some of them are always<br />

curved, bent, and twisted, and they are conspicuously angled,<br />

ridged, and furrowed—the largest ones often having 6 or 7<br />

flattened surfaces and deep grooves. They are yellowish,<br />

mottled or tipped with black, when young, and then gray to<br />

almost black when old. There is commonly one very heavy<br />

central spine which usually has 6 or 7 angles, and is curved<br />

and twisted, 1 to 21/2 inches long, and around 1/8 of an inch<br />

thick. There are 6 to 8 radial spines, the lower 7 radiating<br />

fairly evenly and being the shortest of the spines. The upper<br />

radial is usually about equal to the central spine and may be<br />

even longer. It is often mistaken for a second central because<br />

of its size.<br />

flowers: As the species, except that they are larger, being<br />

about 21/2 inches long, with the petals widening more gradually<br />

from their narrow bases and their tips not as broad as<br />

those of the plant’s relatives. The ovary surface has slender<br />

white bristles at least 1/2 inch long upon it.<br />

fruits: Unreported by anyone except Earle, who describes<br />

them as spiny, globose, and green with a pink blush.<br />

Range. This cactus is always rare, and is found in a comparatively<br />

small area around Zuni and Gallup, New Mexico, extending<br />

from there to just within Arizona and into the southwestern<br />

corner of Colorado around Cortez and Dolores. There<br />

is also a separate population of the variety in southern New<br />

Mexico, where it grows in the White Sands region between the<br />

San Andres and Sacramento mountains.<br />

Remarks. This is a very beautiful cactus, yet one rarely seen.<br />

Its spines are very remarkable and set it off from any of its<br />

relatives. They are heavy enough to be worthy of some massive<br />

barrel cactus. It is closest to variety triglochidiatus, but the<br />

spines are always much more numerous and much more ridged,<br />

grooved, and flattened.<br />

Since its discovery, the study of this cactus has gone through<br />

a history of confusion. It was named by Engelmann; and, at<br />

first, Coulter followed in keeping it distinct. But later Coulter<br />

and Nelson broadened their description of it to include characters<br />

of variety triglochidiatus, and they were followed in this<br />

by others. Plants with fewer ribs and spines were then called by<br />

this name, and of course when plants which were really variety<br />

triglochidiatus were called E. gonacanthus, the supposed range<br />

of variety gonacanthus was enlarged. Britton and Rose then<br />

took the step which logically followed from the blurring of<br />

distinctions between the two plants and placed E. gonacanthus<br />

for the first time as a variety of the other. Different combinations<br />

were also made. Boissevain and Davidson, in their Colorado<br />

Cacti, picture and describe a cactus collected near Corte,<br />

Colorado, which appears to be this form, but which they call<br />

E. coccineus var. octacanthus. This confuses the cactus with that<br />

very different cactus from a far-off part of Texas. But even<br />

today the idea persists that E. gonacanthus occurs in Texas.<br />

This seems to be a perfect example of what happens when<br />

details of stem and spine structure are regarded as insignificant,<br />

for this is a constant form occurring in a definite, small range,<br />

which any person interested in cacti would want to be able to<br />

identify and refer to specifically. Yet, because of the confusion,<br />

I have found typical specimens of it, collected near its type<br />

locality, in herbaria under four different names. It is probably<br />

best considered as only a variety of the larger species, but un-<br />

less it is kept distinct no order will be possible in this group.<br />

Throughout its range in northwest New Mexico and Colorado<br />

variety gonacanthus grows as a small plant with stems only to<br />

about six inches tall, precocious only in its spination. But there<br />

is a strange quirk in its range and its response to a different<br />

environment.<br />

A cactus was collected long ago by Wooton at the White<br />

Sands area of southern New Mexico, which had 7 radial spines<br />

and one much-flattened central, and which he, therefore, called<br />

E. gonacanthus. I have seen this specimen, and in its preserved<br />

state it certainly could pass for our plant. However, its occurrence<br />

so far from the ordinary range of the variety seems<br />

strange and might make us doubt that it is our plant. Our<br />

doubts are increased when we find that the White Sands cactus<br />

grows to a majestic 18 inches tall by 41/2 inches in thickness,<br />

making it probably the largest Echinocereus in our Southwest.<br />

I went into the study of this White Sands cactus with these<br />

doubts. I studied the cacti growing in their natural habitat.<br />

They have been fairly common and are still to be found here<br />

and there within the White Sands National Monument, but<br />

they have for the most part been eliminated in the rest of their<br />

territory, which lies mostly within the military’s missile range,<br />

from wide areas of which the vegetation seems to have been<br />

systematically eliminated.<br />

After observing numerous specimens of the cactus I was forced<br />

to conclude that they are identical to the much smaller variety<br />

gonacanthus in every respect but stem size and I wondered<br />

how one could explain the great difference in size alone.<br />

Fortunately this question was answered when cactophile friends<br />

in both Albuquerque and Colorado showed me what had<br />

happened to specimens of the White Sands cacti which they had<br />

brought out and planted in their gardens. In each case, during<br />

one year in their gardens, stems 12 inches or more tall when


genus Echinocereus engelmann 39<br />

collected had shrunk to half that size or sometimes even smaller,<br />

and then had proceeded with the slow, limited growth and<br />

branching seen in the northern wild population.<br />

Obviously, we have at the White Sands a local population of<br />

variety gonacanthus isolated some way in a very unusual habitat<br />

which enables it to grow to sizes it cannot match or even<br />

maintain anywhere else. This makes it probably the most remarkable<br />

example of environmental effect upon growth of any<br />

cactus within our area. Except for this, the variety is a smaller,<br />

heavy-spined cactus, a rare and beautiful resident of the north-<br />

west corner of our area, where it grows in small clumps under<br />

the cedar trees on the higher, sandy hills.<br />

Echinocereus triglochidiatus var. octacanthus (Muehlenpf.)<br />

Marshall<br />

“Strawberry Cactus,” “Claret-Cup Cactus”<br />

Description Plate 12<br />

stems: Branching or clustering to form clumps, of from 2 or<br />

3 to sometimes as many as 50 stems in some old plants. The<br />

stems are globular to cylindrical, bright green or pale, yellow-<br />

green, and plump, with shiny surfaces. They are of varying<br />

heights and somewhat loosely clustered within the clump.<br />

There is a distinct but gradual increase in the maximum size<br />

of the plants from one end of its range to the other, the sterns<br />

hardly ever exceeding 5 inches in height in the northern part<br />

of the range, but often standing 9 to 12 inches high in the<br />

southern part, particularly in the eastern Big Bend and along<br />

the Devil’s River. They are always comparatively thick, the<br />

largest ones being 41/2 inches in diameter. They have 5 to 9<br />

wide, shallow ribs with slight enlargements at the areoles, but<br />

with no—or only slight—cross-furrows between areoles.<br />

areoles: Round, about 3/16 of an inch across, and 3/8 to 11/4<br />

inches apart. At first they have much short white wool on<br />

them, but later they become bare and are entirely filled by<br />

the swollen bases of the spines.<br />

spines: All spines on this plant are slender or medium in<br />

thickness, actually measuring from only 1/32 to 1/16 of an<br />

inch in diameter, always round and straight or nearly so,<br />

and having very bulbous bases. When young they are yellowish,<br />

often with distinct red streaks and shadings, especially<br />

toward the bases. When old they may remain yellowish<br />

or darken to gray or almost black. The amount of darkening<br />

seems to depend upon how much sun they get: I have collected<br />

plants from sunny ledges which had almost black<br />

spines, only to have the new spines remain yellowish or gray<br />

when grown with some shade. There are 3 to 9 radials, very<br />

nearly equal in size on any areole, or with the upper ones<br />

only slightly smaller. They are from 1/2 to 11/4 inches long.<br />

There may be one central spine standing out at a right angle<br />

to the stem, 5/8 to 11/2 inches long, round and scarcely any<br />

heavier than the radials, but many plants lack the central<br />

entirely.<br />

flowers: As the species, except for the following limitations:<br />

they are smaller, being only 11/2 to 2 inches tall, with the<br />

petals short and blunt; the ovary tube has almost no wool, but<br />

has clusters of 2 to 6 slender spines up to 1/2 inch long; the<br />

stigma lobes are 5 to 8 in number and light green.<br />

fruits: Round, smooth, red, edible berries about 3/4 to 11/2<br />

inches in diameter. They have a few spines which soon fall<br />

off, leaving them naked by the time they are ripe.<br />

Range. A band of territory about 100 miles wide from central<br />

Texas west through the Big Bend, perhaps to near El Paso. Specifically,<br />

the northeastern limit of this strip is just east of the<br />

Colorado River near Lampasas, Texas, the southeastern limit<br />

about Kerrville, Texas. From there it is found quite commonly<br />

westward past junction and Del Rio. The northern edge of the<br />

range proceeds on south of Fort Stockton to the Davis Mountains,<br />

while the southern limit, after dropping into Mexico at<br />

Del Rio, takes in all of the Big Bend. The western limit of the<br />

range is not so definitely known, due to confusion of the names<br />

and incompleteness of the older records. The most western<br />

strictly verifiable reports seem to be from Presidio County on<br />

the south and near El Paso on the north.<br />

Remarks. This plant was first named Echinopsis octacanthus<br />

in 1848 by Muehlenpfordt, from a plant collected by Dr. F.<br />

Roemer in Texas. In 1849 Engelmann described the same Texas<br />

plant and named it Cereus roemeri. Muehlenpfordt had in the<br />

meantime named an entirely different New Mexico plant Cereus<br />

roemeri, and from here on one can anticipate confusion.<br />

In 1896 Coulter wrote of a plant fitting this description,<br />

calling it Cereus octacanthus, but stating that it occurred from<br />

extreme southwestern Texas around El Paso northwestward<br />

through New Mexico into Utah. It seems impossible now to<br />

guess what plant he was referring to. But from this time on it<br />

was assumed that Echinocereus octacanthus, as Britton and Rose<br />

called it, occurred in New Mexico. Perhaps the reason lies in the<br />

coincidence of both a Texas plant and a different New Mexico<br />

plant having been given the same name, Cereus roemeri, and<br />

in the superficial similarity between the Texas plant and a Colorado<br />

and New Mexico cactus called E. coccineus. To add to the<br />

confusion, Boissevain and Davidson, in their Colorado Cacti,<br />

mistook for variety octacanthus both variety gonacanthus, a<br />

primarily Colorado cactus which can have the same number of<br />

ribs and spines but which is vastly different in most other details,<br />

and E. mojaviensis, another similar species which reaches<br />

from the West into the corner of Colorado, and called their<br />

amalgamation of these two E. coccineus var. octacanthus.<br />

As a start in putting the confusion straight, I will say that I<br />

have not seen a plant definitely known to be from either Colorado<br />

or New Mexico having the combination of characters set


40 cacti of the southwest<br />

out in the early, carefully circumscribed descriptions and outlined<br />

above for this plant. I believe it to be entirely a Texas<br />

cactus, except for an extension into northern Mexico.<br />

A further confusion was building, however. In 1856 Engelmann<br />

created a name, E. paucispinus, for a Texas form with 5<br />

to 7 ribs and 3 to 6 round, slender radials. This form he said<br />

usually lacked a central, but might have an occasional round,<br />

porrect one.<br />

It was soon noticed that the rib and spine numbers of the<br />

new E. paucispinus matched those of the New Mexico E. triglochidiatus,<br />

which also lacks centrals. Soon the differences between<br />

the two were overlooked, and the Texas plant was being<br />

spoken of as E. triglochidiatus. However, when both plants are<br />

seen together the difference between them is obvious. The color<br />

and surface of the stems are very different. The spines of the so-<br />

called E. paucispinus are always round and not more than 1/16<br />

of an inch thick-truly insignificant on the broad ribs; while<br />

those of E. triglochidiatus are always greatly flattened, ridged<br />

or even grooved, and twice to four times as thick-easily the<br />

most striking thing about that plant. On these characteristics<br />

alone it is easy to separate them.<br />

As I have found no real intermediate between these two<br />

plants to confuse them, I have also found no specimens of either<br />

of them growing in that wide expanse separating the New<br />

Mexico E. triglochidiatus from the Texas E. paucispinus. There<br />

is about 200 miles between their known ranges. However, this<br />

difference in ranges means that few people have seen them together<br />

and realized the difference, with the result that in Texas<br />

this plant is almost entirely known under the name of the<br />

other. Britton and Rose are largely responsible for this confusion,<br />

since they united the two, but even in doing so they<br />

stated their opinion that E. paucispinus should perhaps be restored<br />

for the Texas plants. And they did us the service of<br />

picturing side by side a true E. triglochidiatus var. triglochidiatus<br />

and the Texas plant, even though their Texas specimen<br />

appears to be a badly damaged variety octacanthus.<br />

And here we may have a clue to unravel some more of the<br />

confusion. In erecting the species E. paucispinus, the Texas form<br />

with fewer ribs, fewer but round spines, and no central spine,<br />

Engelmann appears to have merely set apart and given a name<br />

to the lower end of variety octacanthus’ range of variations.<br />

For a long time I have tried to keep the two separate, but it<br />

just cannot be done. Too many times, on too many hills, the<br />

whole range of characters from 5 to 9 ribs, 3 to 9 radials, and<br />

1 or no central have been found growing happily together. The<br />

flowers and fruits appear to be identical, and it seems that the<br />

two must be combined entirely.<br />

So we have, when we see the whole picture, variety octacanthus,<br />

an entirely or at least primarily Texas cactus which<br />

was once called E. roemeri and so was confused with the true<br />

E. roemeri of New Mexico and Colorado. Then those of its individuals<br />

with fewer ribs and spines were called E. paucispinus<br />

and immediately confused with the very different New Mexico<br />

form, variety triglochidiatus.<br />

In the extreme northeastern part of its range, variety octacanthus<br />

is a small cactus with stems only 3 to 5 inches high, one<br />

of the weakest growths of any in this group. However, in this<br />

area it is adjusting to the wettest habitat any of the red-<br />

flowered Echinocerei succeed in colonizing, and this may be the<br />

price. But around Sabinal and Uvalde, Texas, one begins to<br />

come across clumps of this cactus much more robust in size of<br />

the stems. These specimens grow larger as one goes west through<br />

Del Rio, Langtry, and Sanderson, until from Del Rio west to<br />

south of Marathon and Alpine one occasionally finds majestic<br />

clumps with massive stems to at least 12 inches tall. This appears<br />

to be the plant’s response to the more arid conditions of<br />

this part of its range. If we are correct in our interpretation of<br />

the immense effect of the White Sands area’s conditions on<br />

variety gonacanthus, then we should not be surprised at a similar<br />

though less extreme and more gradual effect of the environ-<br />

ment on this related cactus.<br />

The spines in this western area are more uniformly dark gray,<br />

which one would expect, due to the more general exposure to<br />

the sun here. I was gratified to find that a specimen taken from<br />

a canyon south of Alpine, where it was shaded most of the<br />

day, had the same light, reddish-yellow spines as those growing<br />

under the shade of the eastern junipers, although it stood almost<br />

10 inches high, completely overshadowing its eastern counter-<br />

part in size.<br />

Variety octacanthus is definitely more resistant to rot than<br />

most others in this genus. I have seen plants of it growing and<br />

healthy beside rotted plants of E. enneacanthus and E. caespitosus,<br />

where their ranges all meet around Sabinal, Texas. For<br />

this reason it would probably be a better cactus for use in<br />

eastern gardens than many others of the genus.<br />

Echinocereus triglochidiatus var. hexaedrus (Eng.)<br />

Boissevain & Davidson<br />

Description<br />

stems: Few in the clump, each 4 to 6 inches high and 2 to<br />

21/2 inches in diameter, with 6 obtuse ribs having wide, shal-<br />

low grooves between them.<br />

areoles: Only 1/16 to 1/8 of an inch in diameter, which<br />

makes them among the smallest in this group. They are about<br />

1/2 to 5/8 of an inch apart, and woolly when young.<br />

spines: Slender, with bulbous bases, and distinctly angular<br />

or flattened. Each areole usually has 6 radial spines, but Engelmann<br />

says one specimen had 7 and another 5. The lower<br />

radials are the shortest spines, being 3/8 to 7/8 of an inch long,<br />

and yellowish-red in color. The upper radials are longer,<br />

about 5/8 to 11/8 inches long, stouter, and darker in color.


genus Echinocereus engelmann 41<br />

The central spine is missing, although there may be one, and<br />

Engelmann mentions finding one plant with two centrals.<br />

The centrals are 7/8 to 13/8 inches long and definitely flat-<br />

tened, but not very thick.<br />

flowers: Unknown.<br />

fruits: Unknown.<br />

Range. Known only from about 15 miles west of Zuni, New<br />

Mexico.<br />

Remarks: This cactus was described by Engelmann from plants<br />

collected by Bigelow in 1853. No one has reported seeing it<br />

since that time. Coulter described it many years later from<br />

Bigelow’s preserved specimens. The writer has not seen these, so<br />

the description given above is a combination of Engelmann’s<br />

and Coulter’s.<br />

It is hardly possible to do more than guess about the real<br />

relationships of this cactus to the other forms it resembles, since<br />

we know so little about it; so I merely list it as did the only<br />

two authorities who had it before them. Since then various<br />

authors have given it as a form of one or the other related<br />

species, but little can be proved.<br />

I do find in the U. S. National Herbarium one specimen collected<br />

by Standley in the Tunitcha Mountains of New Mexico,<br />

not far north of Zuni, which fairly well fits Engelmann’s description<br />

of E. hexaedrus. It was labeled by Standley as E. paucispinus,<br />

but that is doubtful, since it has the very flattened<br />

spines which E. paucispinus (var. octacanthus) does not have,<br />

and it was collected very far from that plant’s area in Texas.<br />

It could be the only collection of variety hexaedrus since the<br />

type.<br />

It would be a real service for someone to rediscover and<br />

study this cactus, if it still exists.<br />

Echinocereus coccineus Eng.<br />

“Aggregate Cactus,” “Bunch-Ball Cactus,” “Turk’s Head<br />

Cactus,” “Heart Twister,” “Red-Flowered Hedgehog Cac-<br />

tus”<br />

Description Plates 12, 13<br />

stems: This cactus always consists of a cluster of short, equal-<br />

sized stems tightly packed to form a dense, hemispherical<br />

mass 1 to 6 feet in diameter and containing, in old specimens,<br />

up to several hundred heads. Each head or stem is from only<br />

2 to 6 inches high and up to about 21/2 inches in diameter,<br />

with 8 to 11 ribs which are either practically straight or<br />

often composed of pronounced tubercles or projections on the<br />

tips of which the areoles are found.<br />

areoles: These are large, circular to more or less oval in<br />

shape, and only 3/16 to 1/2 inch apart—the closest areoles<br />

among the clustering, few-ribbed Echinocerei. They are woolly<br />

when young, becoming bare.<br />

spines: All spines on one variety of this cactus are slender to<br />

medium in thickness, often almost bristle-like, straight, and<br />

round. In color they are white, gray-white, or straw; sometimes<br />

the centrals are brownish when new. On the other form<br />

of the species the central is stouter, darker, and definitely<br />

flattened. Due to the closeness of the areoles and the length<br />

of the spines, the general appearance is of a mass of whitish<br />

bristles, from which fact comes the common name, “hedgehog<br />

cactus.” There are 8 to 12 radial spines, 1/4 to 11/8 inches<br />

long, which all more or less stand out from the stem of the<br />

plant, the upper ones being the shorter ones. There are also 1<br />

to 4 centrals: in the one variety, these are always round, very<br />

slender to medium in thickness, 3/8 to 13/4 inches long, and<br />

standing out nearly perpendicular to the stem; in the other<br />

variety the main central is flattened, a little stouter, directed<br />

downward, and 1 to 3 inches long.<br />

flowers: Deep crimson to orange-red in color, around 11/2<br />

to 2 inches long and opening from 1 to 2 inches in diameter.<br />

The stamens are fuchsia and shorter than the petals. The<br />

stigma lobes are green and 6 to 12 in number. The tube surface<br />

has white wool and 8 to 11 slender white spines, 1/4 to<br />

1/2 inch long, in each areole.<br />

fruits: Red, juicy, with deciduous bristles.<br />

Range. Growing in a wide territory from Colorado south past<br />

Raton, New Mexico, down along the upper Pecos River to<br />

about even with Santa Fe and Albuquerque, and from there<br />

west into Arizona and Utah.<br />

Remarks. The “bunch-ball cactus” was among the first collected<br />

in our area and is distinctive enough, with its slender,<br />

round spines and its manner of growing as a tight, perfectly<br />

regular ball of stems that it has not been so much confused with<br />

others of the group as some. However, it also has had too many<br />

names applied to it. Engelmann first called it E. coccineus, and<br />

then, later, because of a change of genus name, he called it<br />

Cereus phoeniceus. He also had once named a plant Mammillaria<br />

aggregata, which he did not describe in any detail. Coulter,<br />

assuming correctly that this was our cactus, called our plant<br />

Cereus aggregatus. Later authorities got back to the first definite<br />

name, E. coccineus, until, in an attempt to combine this cactus<br />

with others by reducing it to a variety, L. Benson found it necessary<br />

to rename it yet again as E. triglochidiatus var. melanacanthus,<br />

resurrecting for it an obscure varietal name once coined<br />

by Engelmann but ignored since as referring to no form really<br />

distinct from the species.<br />

There has been much confusion of E. triglochidiatus var. octacanthus<br />

with E. coccineus because they both have round spines.<br />

Since the two do not grow at all in the same areas, the differences<br />

between them have been easily overlooked, but they are


42 cacti of the southwest<br />

very different plants, with hundreds of miles and other forms<br />

separating their ranges, and when they are closely compared it<br />

is hardly possible to confuse them.<br />

Echinocereus coccineus var. coccineus (Eng.)<br />

Description Plate 12<br />

stems: As the species, except that the ribs undulate by having<br />

definite and pronounced tubercles at the areoles.<br />

areoles: As the species, except that they are to only 3/8 of<br />

an inch apart.<br />

spines: As the species, except that all spines are round, the<br />

centrals all slender, spreading, and to only 7/8 of an inch long.<br />

flowers: As the species, except that the stigma lobes number<br />

only 6 to 8.<br />

fruits: As the species.<br />

Range. As the species.<br />

Remarks. This is the typical form of the species. It was apparently<br />

very common in northern New Mexico at one time, growing<br />

equally well under the trees on the lower mountains and in<br />

grassy valleys, forming huge mounds of dozens of stems. It has<br />

been a great favorite in that area because of the beauty of the<br />

large, symmetrical clumps it forms, its brilliant flowers, and its<br />

hardiness. For this reason many plants were dug up, and most<br />

of them were allowed to languish and die in boxes and planters.<br />

As a result it is now much easier to find it growing in gardens<br />

of the cities of that area than on the range, and one has to<br />

search long in remote spots to find an old specimen rounded up<br />

into a massive hemisphere. All one usually finds in any easily<br />

accessible place are young specimens of only a few heads, which<br />

give little clue to the magnificent things they could become if<br />

left unmolested.<br />

Echinocereus coccineus var. conoideus Eng.<br />

“Beehive Cactus”<br />

Description Plate 13<br />

stems: 3 to 6 inches high, 2 to 21/2 inches in diameter, the tips<br />

being markedly smaller in diameter, this giving them a conical<br />

shape. When old, they form large, rounded mounds of 30<br />

to 40 nearly equal stems, somewhat similar to but larger in<br />

individual stem size than those of the typical species form.<br />

There are 9 to 11 ribs on each stem, which are almost straight<br />

with at most very small swellings at the areoles and with<br />

rather deep furrows between them.<br />

areoles: Large, woolly when young, later almost bare, 1/4 to<br />

1/2 inch apart.<br />

spines: There are 9 to 12 slender radial spines which are very<br />

uneven, the upper 2 or 3 being only 1/4 to 1/2 inch long, while<br />

the lateral and lower ones are larger, up to 11/8 inches long.<br />

They are all round, with enlarged bases, and are white to<br />

straw-colored. There are 3 or 4 central spines. The upper ones<br />

are as the radials in shape and color and not much if any<br />

longer than the longest radials. The lower central, however,<br />

is directed downward, instead of standing perpendicular to<br />

the stem, and is usually somewhat curved. This lower central<br />

is a little stouter than the other spines, although still slender<br />

for the group, and different from the other spines on the<br />

areole in being definitely flattened, often to the point of being<br />

quadrangular. It is darker in color, often being yellowish<br />

with a brown base, or else ashy-gray. It is 1 to 3 inches long.<br />

flowers: These are a little larger than some in this group,<br />

being about the same length and color, but opening all of 2<br />

inches in diameter, and there are 9 to 12 green stigma lobes.<br />

fruits: As the species.<br />

Range. From along the upper Pecos River in north central New<br />

Mexico north into southern Colorado and west into Arizona.<br />

Remarks. Variety conoideus is very close to variety coccineus,<br />

and there seems more justification for uniting these two, as Britton<br />

and Rose suggested but did not do, than any of the other<br />

forms listed here. Boissevain and Davidson stated that transitional<br />

forms existed between these two but kept them as separate<br />

species, a position which seems hard to maintain. However,<br />

the typical forms of the two are so different that it seems there<br />

must be some distinction made between them, so we are going<br />

back to Engelmann’s first opinion and placing this plant once<br />

again as a variety of the other. Variety conoideus has the longer<br />

spines and less tuberculate ribs, with more widely separated<br />

areoles, and it is a larger, more robust plant in general. In particular,<br />

the long, flattened, and darker central spine serves to<br />

identify it quickly.<br />

The plant was first named Cereus roemeri by Muehlenpfordt,<br />

but Engelmann did not seem certain about Muehlenpfordt’s description;<br />

and besides he had in the meantime applied the name<br />

roemeri to the Texas cactus we know as E. triglochidiatus var.<br />

octacanthus, so he was obliged to rename this New Mexico<br />

cactus. He used the name Cereus phoeniceus var. conoideus,<br />

and later, when he decided to elevate it to species rank, Cereus<br />

conoideus. Muehlenpfordt’s description seems very clear, however,<br />

and his name, being the oldest, is the one which should<br />

be used if the plant is treated as a species. The Texas plant can-<br />

not go under this name.<br />

But since the plant we are considering seems much more accurately<br />

placed as a variety, by the rules of nomenclature it<br />

must carry the first varietal name applied to it. This is quite<br />

clearly Engelmann’s variety conoideus.<br />

Most of what was said about the growth habits of E. coccineus<br />

would apply to this cactus as well, except that it seems<br />

to grow in higher mountains and on more rocky locations. This


genus Echinocereus engelmann 43<br />

one has also been gathered from the field too widely and is now<br />

hard to find growing wild.<br />

Echinocereus polyacanthus var. rosei (Wooton & Standley)<br />

“Red-Goblet Cactus,” “Pitahaya”<br />

Description Plate 13<br />

stems: This is another cactus forming flat, loose clumps of<br />

unequal stems. The clumps may become large, with up to 50<br />

stems, but this is rare and they are usually smaller, with only<br />

a dozen or fewer heads. Each stem is cylindrical, but somewhat<br />

tapering over almost the whole length toward the smaller,<br />

rather pointed tip. They grow to at least 10 inches long<br />

and 4 inches thick at the base, when in ideal situations, but<br />

remain much smaller in poor environments. The color is a<br />

lighter or paler green than some of its relatives. The number<br />

of ribs is variable, from 9 to 11, these broad with very shallow<br />

grooves between them on the older parts of the stems,<br />

but sharp with fairly prominent tubercles or swellings at the<br />

areoles and deep, narrow grooves between them at the tips.<br />

areoles: This plant has probably the most conspicuous areoles<br />

of the group. When young they are about 1/4 of an inch<br />

across, circular, bulging outward, with much white or yellowish<br />

wool covering the bases of the new spines. As they get<br />

older they become more typical, nearly flat, and lose much<br />

of their wool. They are from 3/8 to 1 inch apart.<br />

spines: The spines of this cactus are undoubtedly the most variable<br />

in size of any in the group. I have seen plants with large<br />

centrals over 2 inches long and other plants with no spines<br />

over 1/2 inch long. In spite of this variation, the spines are<br />

constant in being straight, of medium stoutness, and always<br />

round. There are 7 to 10 radials 3/8 to 1 inch long, radiating,<br />

the lower ones usually almost twice as long as the upper ones.<br />

There are 3 to 5 centrals on mature plants. Young seedlings<br />

always have only 1 central per areole, and in some mature<br />

specimens occasional areoles will have only the 1 central, but<br />

this is never typical of the whole mature plant. These centrals<br />

may be 1/2 to 2 inches long, depending upon the specimen,<br />

but they are always round, from somewhat enlarged<br />

bases, and spreading at various angles from the areole. All<br />

spines are reddish, ashy-gray, or occasionally dark, purplish-<br />

gray, with the centrals usually somewhat darker than the<br />

radials.<br />

flowers: Variable in color, including tints from pale red to<br />

orange, these often in the same flower. These shades are unique<br />

among the firm, long-lasting flowers of this group. Otherwise<br />

the flowers are typical of the group, 11/2 to 21/2 inches<br />

long, with short, rigid petals broadening and blunt at the<br />

ends. They have fuchsia stamens and the stigma lobes number<br />

7 to 10. On the ovary are brownish or yellowish spines, 1/8<br />

to 5/8 of an inch long, with reddish tips, plus some short<br />

wool.<br />

fruits: 3/4 to 1 inch long, greenish-purple when ripe, with<br />

deciduous spines.<br />

Range. From Mexico northward over west Texas and much of<br />

New Mexico, into southern Colorado and southeastern Arizona.<br />

The northeastern limit of the known range is near San Antonio,<br />

Texas, and so far as is known the northern edge of the range<br />

runs past Rocksprings, Texas, to the Davis Mountains and then<br />

turns sharply north through the Guadalupe Mountains into<br />

New Mexico, where it follows the Pecos River, goes past Las<br />

Vegas, New Mexico, and along the eastern edge of the mountains<br />

into Colorado. The range seems to come back south out of<br />

Colorado east of the Continental Divide, and does not cross<br />

this demarcation until near Lordsburg, New Mexico, where it<br />

moves into southeast Arizona.<br />

Remarks: E. polyacanthus var. rosei is the most common red-<br />

flowered Echinocereus throughout southern New Mexico and<br />

far southwest Texas. It is easily found on the lower slopes of<br />

the Franklin and Organ mountains and in similar places west<br />

into Arizona. It is less common but may be found occasionally<br />

in the mountains of central New Mexico and north into Colorado.<br />

It is even less common, but has also been collected in the<br />

Texas Big Bend and the Davis and Guadalupe mountains. Its<br />

eastern limit is hard to establish, as occasional specimens which<br />

must be referred to this species have turned up in widely separated<br />

places over a very large area. I have a living specimen<br />

with very short spines but otherwise typical from near Rock-<br />

springs, Texas, and there is in the U.S. National Herbarium a<br />

specimen labeled as collected by Tourney at San Antonio in<br />

1897, but I cannot find it growing in the area now.<br />

Because of the variability of its spines and its very wide range<br />

this cactus has been widely confused with others of its relatives.<br />

In northern New Mexico and Colorado it is sometimes mistaken<br />

for E. coccineus, from which it can be distinguished by the facts<br />

that it never grows in the compact, dome-shaped clusters nor<br />

has such slender spines as that plant, or for E. coccineus var.<br />

conoideus, from which it can be told by the fact that its spines<br />

are never flattened as is the lower central of that plant. In<br />

Texas its immature growth and certain stunted, atypical specimens<br />

have been called E. triglochidiatus var. octacanthus, in<br />

spite of the fact that Engelmann in his early description was so<br />

thorough that he mentioned that the immature plants have only<br />

1 central per areole. The atypical forms can be distinguished by<br />

the fact that they have more ribs than variety octacanthus.<br />

Engelmann did leave room for confusion to enter by giving<br />

two slightly varying descriptions for the plant he named E.<br />

polyacanthus. His original description was of plants collected<br />

in Chihuahua, Mexico, by Wislizenus. This was a narrow description<br />

of a very localized cactus found only there. It is<br />

unique for the whole group of scarlet-flowered Echinocerei in<br />

having long wool on the flower tube. In his later writings En-


44 cacti of the southwest<br />

gelmann used the same name, but broadened the description<br />

somewhat, as well as the range. In the later descriptions he said<br />

the cactus was a common plant at El Paso, and did not mention<br />

the long wool on the flower. Since no specimens have ever been<br />

collected at El Paso or anywhere in the U.S. with this long<br />

wool, its seems clear that, after further study, Engelmann meant<br />

to include all of this form, whether having long or short wool,<br />

under this name. Coulter took it this way, and so included in<br />

the range of the species New Mexico and Arizona.<br />

Wooton and Standley returned to restricting E. polyacanthus<br />

to the local Chihuahuan form, and set up the New Mexico representatives<br />

having only short wool as two new species, E. rosei<br />

and E. neo-mexicanus.<br />

Britton and Rose faced the problem of relating these forms.<br />

Dr. Rose made the famous trip to the type locality of E. polyacanthus<br />

at Cosihuiriachi, Chihuahua, and collected specimens<br />

there, establishing to their satisfaction that the original type<br />

specimens were actually different from our forms, at least in<br />

having the long wool on the flower tube. They, therefore, followed<br />

Wooton and Standley in their names, E. rosei and E.<br />

neo-mexicanus, for the two U.S. forms which lack this unique<br />

character. Since then no one has used the name E. polyacanthus<br />

for any U.S. cactus except Benson, who ignores the distinction<br />

and calls the plants of New Mexico and Arizona E. triglochi-<br />

diatus var. polyacanthus.<br />

Helia Bravo, in her book, Las Cactaceas de Mexico, has a<br />

good illustration of the true E. polyacanthus.<br />

It seems necessary to follow most recent students in separating<br />

the U.S. forms in some way from the unique Chihuahuan cactus,<br />

which must carry the name E. polyacanthus. But it seems<br />

equally true that the U. S. forms are very close to it, close<br />

enough so that Engelmann had justification in lumping them all<br />

together. The best way to show the whole picture appears to be<br />

by regarding the U. S. forms as varieties of the species. Our<br />

common cactus of such wide range in the U. S. then becomes<br />

E. polyacanthus var. rosei.<br />

Echinocereus polyacanthus var. neo-mexicanus (Standley)<br />

Description Plate 13<br />

stems: As those of variety rosei, except to only about 3<br />

inches thick and having 11 to 15 ribs.<br />

areoles: As those of variety rosei, but closer, being 3/8 to<br />

5/8 of an inch apart.<br />

spines: Radial spines variable in number from 8 to 16, but<br />

usually 10 to 13. These are slender; straight; round; white,<br />

straw, or yellow; and to 5/8 of an inch long. The spreading<br />

centrals number 4 to 6 on mature plants, and are straight,<br />

round, slender, and usually 3/4 to 11/4 inches long. In color<br />

they are yellowish below with the outer parts or the tips<br />

reddish or blackish.<br />

flowers: As those of variety rosei, except usually smaller<br />

and of burnt-orange to yellow coloring. The petals are not as<br />

wide as those of the other variety and are, as Wooton put it,<br />

“almost acute,” if one emphasizes the almost.<br />

fruits: As those of variety rosei.<br />

Range. Fairly common around Las Cruces, New Mexico, and<br />

found occasionally at least to Socorro, in central New Mexico.<br />

Remarks. This plant is very close to the previous variety, but is<br />

easily recognized by the more numerous ribs and the more<br />

numerous, lighter colored, and more slender spines. Where va-<br />

riety rosei presents a gray or purplish-gray appearance, the<br />

spine covering of variety neo-mexicanus is always yellowish,<br />

and sometimes strikingly so. It does not appear to be a separate<br />

species, but it does definitely appear to stand in some way distinct<br />

from the other more widespread form. Collections made<br />

around Las Cruces are easily divisible into the two types.<br />

This is a beautiful cactus, usually large and impressive in<br />

growth, with the long-lasting flowers which make it a favorite.<br />

Near El Paso and Las Cruces, in the few areas which have been<br />

too remote for the casual collector or which have been in some<br />

way protected, there are still to be seen whole slopes dotted<br />

with the clumps of tall heads covered in April and May with<br />

beautiful goblet-shaped flowers, and here and there within the<br />

cities transplanted clumps are to be seen gracing yards and gardens<br />

most beautifully. Variety neo-mexicanus is cold-resistant<br />

and would be a good garden cactus in other colder areas, except<br />

that it is definitely a desert form and so must be protected<br />

from moisture more carefully than some of the other species of<br />

this group.<br />

It is worth mentioning, for the sake of any hobbyist who<br />

might be growing the plant inside to protect it from the winter<br />

dampness of their areas, that this cactus, like many other winter-<br />

hardy forms, must have some cold during the winter in order<br />

to trigger the blooms. In south Texas I have seen plants which<br />

had grown for several years without a suggestion of a flower<br />

bloom profusely for the first time after the stimulus of one of<br />

those freezes which are so destructive to southern cacti.<br />

Echinocereus enneacanthus Eng.<br />

“Strawberry Cactus,” “Pitaya”<br />

Description Plate 14<br />

stems: 3 to 30 inches long and 11/2 to 4 inches thick, and<br />

cylindrical, tapering somewhat over the last one-third of the<br />

length to a rather pointed tip. These stems grow in loose<br />

clusters of a few to as many as 100 in a large plant. New<br />

stems multiply as side branches at or just above the ground<br />

level, so their first growth tends to be lateral, after which<br />

they turn upward. This results in all of the stems around the<br />

edges of a large clump being long and curving, the lower


genus Echinocereus engelmann 45<br />

part often lying flat on the ground and the upper part standing<br />

erect. In color they are bright green, and the flesh of the<br />

plant is soft, even flabby, giving them a more or less wrinkled<br />

appearance and causing them to appear actually withered<br />

in very dry periods or in the winter. There are 7 to 10<br />

ribs on each stem, which are low and broad with shallow<br />

grooves between them and with slight to rather pronounced<br />

tubercles or swellings at the areoles.<br />

areoles: Circular, about 1/8 of an inch in diameter, with<br />

much gray wool on them when young, some of which remains<br />

on the older areoles. These are placed 1/4 to 11/2 inches<br />

apart on mature stems.<br />

spines: All spines on this plant are rigid and slender to fairly<br />

stout, rise from enlarged bases, and are light-colored, being<br />

white to straw-colored or very light brown. All spines are<br />

distinctly translucent, having a horny appearance with age.<br />

When very young and while still growing, the spines are a<br />

delicate pink color which fades quickly as they mature. There<br />

are 7 to 12 white radial spines radiating evenly, straight or<br />

slightly curved back toward the stem, but varying greatly in<br />

length and thickness. The upper ones tend to be shortest, only<br />

1/4 to 5/8 of an inch long, while the lateral and lower ones are<br />

usually longer—1/2 to as much as 1 inch long. There is one<br />

stout central spine standing perpendicular to the surface of<br />

the stem or slightly deflexed. It has a very bulbous base and<br />

is round and white when young, but becomes darker, especially<br />

toward the base, and more or less flattened when old.<br />

On immature tips this central is very little longer than the<br />

radial spines, but with age it usually continues to grow, becoming<br />

heavier and longer, up to 11/4 or even 2 inches long.<br />

Many plants are found with 2 extra central spines above this<br />

main one, these spreading upward and remaining shorter.<br />

flowers: Large and beautiful, opening widely, 2 to 3 inches<br />

in height and about the same in diameter, purple-red in<br />

color. There are 10 to 20 short outer petals with brownish-<br />

green centers and pinkish, crinkled edges. The inner petals<br />

are in 1 to 3 rows, 12 to 35 in number and oblong, linear,<br />

or spatulate in shape. The edges of these petals are entire or<br />

toothed, and the tips pointed or blunt. The stamens are much<br />

shorter than the petals, the filaments greenish, and the anthers<br />

yellow. The style is white, the stigma lobes green, long and<br />

slender, 8 to 12 in number. The tube of the flower has white<br />

wool and white bristle-like spines up to 1/2 inch long upon it.<br />

fruits: About 1 inch long, almost spherical, greenish to<br />

brownish or purplish, with bristle-like spines which fall<br />

off easily. The flesh of the fruit is edible and very delicious.<br />

Range. Along the Rio Grande from near that river’s mouth to<br />

the Big Bend. It has seldom been found west of the Pecos River<br />

or more than 50 miles north of the Rio Grande in any area.<br />

However, there have been isolated reports of it from near San<br />

Angelo, Kerrville, San Antonio, and Raymondville, Texas,<br />

which would mark the northern and eastern extremes of its<br />

range. Coulter says it occurs west of Texas into Arizona, but I<br />

have found no record of it ever having been collected in New<br />

Mexico.<br />

Remarks. This is the common Echinocereus of a wide area of<br />

the lower Rio Grande Valley. It is found occasionally farther<br />

west, but only becomes common near Del Rio. It is very common<br />

from the Devil’s River to Eagle Pass, large clumps of it<br />

dotting the gravelly ground on the low hills just north of the<br />

Rio Grande wherever these hills have not been cleared. It is so<br />

common as to be a pest at Laredo and for about 50 miles north<br />

and east of there, with clumps under almost every bush on<br />

whatever of the range is well-drained and has not yet been<br />

cleared. The practice of “rooting” or “chaining” the range with<br />

large machinery in order to clear out the brush seems to do<br />

away with this, as well as with most other small cacti, although<br />

it merely does a favor to the large Opuntias of the region,<br />

whose pads are scattered by the machines and immediately take<br />

root as new plants. The result is that you now have to hunt for<br />

unspoiled range where the pitayas still are allowed to grow<br />

and the giant Opuntias have not taken over. Below Laredo this<br />

cactus is found commonly to near Rio Grande City, but below<br />

there it is found only rarely.<br />

E. enneacanthus was noticed and collected on the very early<br />

survey of the Rio Grande area by Wislizenus, and named by<br />

Engelmann in 1848. It is a distinct form which has seldom been<br />

confused with any other. However, there seem to be more or<br />

less distinct varieties within its population, which are listed<br />

below.<br />

When its clusters are covered with dozens of its large, lively<br />

colored flowers (usually during April), the species is a beautiful<br />

cactus, appreciated by almost everyone. It is most appreciated<br />

by those who take its common name of strawberry cactus literally,<br />

for the greenish-brown fruits have a flavor very similar to<br />

strawberries. Where it grows profusely the fruits are actually<br />

gathered, the spines brushed off, and the flesh eaten with cream<br />

and sugar.<br />

This cactus is fairly hardy, being able to stand a temperature<br />

considerably below freezing, but it cannot stand excess moisture.<br />

If it is in some way protected from winter rains and allowed to<br />

wither in dryness during the winter, it can live quite far north<br />

of its natural range. Then, when moisture is given to it again in<br />

the spring, the reward will be quick appearance of many large<br />

flowers.<br />

Echinocereus enneacanthus var. enneacanthus (Eng.)<br />

Description Plate 14<br />

STEMS: As the species, except that they grow to only about 12<br />

inches long, 23/4 inches thick, and are much less flabby, with<br />

only slight tubercles at the areoles.


Rumpl -> Rümpl<br />

46 cacti of the southwest<br />

areoles: As the species, except that they are to only 1 inch<br />

apart.<br />

spines: As the species.<br />

flowers: As the species, except with only 10 to 15 outer<br />

petals and only 12 to 15 inner petals in one row. These petals<br />

are oblong or linear in shape, with edges entire and tips<br />

pointed. There are only 8 to 10 stigma lobes.<br />

fruits: As the species.<br />

Range. As the species.<br />

Remarks. This is the typical variety of the species which is com-<br />

mon over all of its range. Its smaller stems to only 12 inches<br />

long are prostrate for only the first part of their length, and the<br />

plant tends to form rather tidy, firm clumps.<br />

Echinocereus enneacanthus var. carnosus (Rümpl.)<br />

K. Schumann<br />

Description Plate 14<br />

stems: Becoming at least 16 and sometimes as much as 30<br />

inches long by 3 to 4 inches in diameter. These very flabby<br />

stems grow out laterally from the cluster and when large lie<br />

fully prostrate on the ground, only the very tips turning upward.<br />

There are 8 or 9 broad ribs on each stem, with rather<br />

pronounced tubercles.<br />

areoles: 3/4 to 11/2 inches apart.<br />

spines: There are 8 or 9 radials, the uppers only 3/8 of an<br />

inch long (often missing entirely), and the laterals 1/2 to 3/4<br />

of an inch long. There is usually only 1 porrect central 1/2 to<br />

2 inches long, but rarely there may be 2 short upper centrals<br />

besides. The centrals are round when immature and flattened<br />

when old.<br />

flowers: Very large and full. There are 13 to 20 short, greenish<br />

outer petals and 20 to 35 inner petals in 3 rows. These<br />

inner petals are spatulate, the edges crinkled and toothed,<br />

with the tips not pointed. The narrow bases are greenish, the<br />

broader upper parts fuchsia or reddish-purple. The style is<br />

white and short. There are 10 to 12 green, linear stigma<br />

lobes.<br />

fruits: As the species.<br />

Range. Occurring only about Laredo and Eagle Pass, Texas.<br />

Remarks. There is no experimental evidence to show the relationship<br />

of this form to the typical variety enneacanthus. Its<br />

range is entirely within that of the species, and there is no way<br />

at present to know how much of its distinctness is due to environment.<br />

It does exist as a recognizable form, however, and<br />

in its fullest development a very remarkable one. I have seen<br />

large plants fully five feet in diameter, their outer stems spreading<br />

over the ground like giant starfish arms. The sight of such<br />

green snake-like forms nearly a yard long running out from<br />

under a mesquite tree is unforgettable. The large, extremely full<br />

flowers of this form are as remarkable as the stems. With their<br />

several rows of petals they are truly the roses of the cacti.<br />

Small specimens of variety carnosus may be distinguished<br />

from the typical form by the excessive flabbiness of the stems,<br />

the more distant areoles and, of course, the surpassing flowers.<br />

This form has apparently been known by the name E. enneacanthus<br />

var. major Hort.<br />

Echinocereus stramineus (Eng.) Rümpl.<br />

“Strawberry Cactus,” “Organo,” “Pitaya”<br />

Description Plate 14<br />

stems: Up to at least 10 inches tall and 31/2 inches thick,<br />

tapering gradually over most of their lengths to a rather<br />

pointed apex. They have 11 to 13 ribs, which are rather<br />

sharp, with fairly deep furrows between them and with slight<br />

tubercles or enlargements at the areoles. These stems cluster<br />

very freely by multiplying from the base, thus forming large,<br />

compact clumps of up to 100 or more equal stems. Such a<br />

large plant has the form of a hemisphere, often 2 or 3 feet<br />

across and nearly as high.<br />

areoles: On this cactus they are small, round, white, with<br />

much wool when young, and 3/8 to 3/4 of an inch apart.<br />

spines: All white to straw-colored and translucent, slender to<br />

medium in thickness from thickened or bulbous bases, covering<br />

the plant profusely. When very young at the tip of the<br />

stem, the spines are a very delicate pink in color, this becoming<br />

straw-colored for a while and then fading quickly to<br />

whitish. The radials vary in number from 7 to 14, and in size<br />

from 3/8 to 11/2 inches long, these extremes in size often being<br />

found on the same areole with the lower being the longer<br />

ones. They are all round and either straight or curved. The<br />

centrals vary from 2 to 5 in number and are to 31/2 inches<br />

long, slender for their length, and round or slightly flattened.<br />

They are usually a little darker than the radials, and may<br />

be straight or curved. Usually the lower central is perpendicular<br />

to the stem surface, while the others spread upward<br />

at various angles and interlock with those of other areoles.<br />

flowers: The flowers of this cactus are very large and beautiful,<br />

and are produced in large numbers. They are 4 to 5<br />

inches tall by 3 to 4 inches in diameter, and purple-red in<br />

color. There are 10 to 15 pointed outer petals with green<br />

centers and pink edges. There are 15 to 20 inner petals which<br />

are longer than the outer ones, with narrow bases and broadening<br />

to 1/2 inch wide or more toward the tips. These bases<br />

are a bright red which blends gradually to a bright rose toward<br />

the tips. The edges of these inner petals are ragged and<br />

toothed, and the blunt ends are sometimes notched. The fila-<br />

Rumpl -> Rümpl


Forster -> Förster<br />

Rumpl -> Rümpl<br />

genus Echinocereus engelmann 47<br />

ments are short and red. The anthers are yellow. The style is<br />

long and red, and on top of it there are 10 to 13 long, green<br />

stigma lobes. The tube of the flower is very long, with many<br />

white, bristle-like spines upon it.<br />

fruits: These are spherical, 11/2 to 2 inches long, purplish to<br />

red when ripe, with deciduous bristle-like spines. They are<br />

edible.<br />

Range. From El Paso almost to the lower Pecos River in Texas,<br />

and extending deep into Mexico. It is said to have been collected<br />

in southern New Mexico, but I have not been able to<br />

verify this.<br />

Remarks. E. stramineus is one of the most beautiful Echinocerei.<br />

Its domelike, compact manner of growth is shared by no<br />

other cactus within its range, and only by the more western<br />

and northern E. coccineus. It grows commonly only on the upper<br />

slopes of the sandy hills east of El Paso and on the limestone<br />

ridges in and around the Big Bend National Park. In these<br />

places its large clumps show up from a distance as glistening,<br />

whitish, or golden balls on the almost bare crowns of these hills.<br />

When it blooms in the spring its flowers form brilliant diadems<br />

of color for these otherwise drab rises. Its flowers are among<br />

the largest and most numerous of any Echinocereus. I have<br />

counted 40 of these brilliant purple blossoms on one clump at<br />

one time. Its fruits are also delicious, with a taste similar to and<br />

said by some even to surpass that of strawberries.<br />

I have not seen this cactus from anywhere west of the Franklin<br />

Mountains near El Paso, and there are no recent reports of<br />

its collection west of there, although there are old reports of it<br />

from southern New Mexico and even Arizona. It occurs only<br />

occasionally east of the Hueco Mountains of Texas, in an area<br />

south of Marfa, Alpine, and Marathon, Texas, extending into<br />

the Big Bend National Park, where it is once again almost as<br />

common as farther west. Near Langtry, Texas, seems to be its<br />

eastern limit. It grows widely in Mexico, and is considered by<br />

some to be the same as E. conglomeratus (Förster) Maths. of<br />

Mexico.<br />

This species must have the very dry, rocky conditions of its<br />

hillsides. For this reason it seldom survives when transplanted<br />

to gardens unless extra care is taken to give it sandy soil and to<br />

shield it from moisture.<br />

Echinocereus dubius (Eng.) Rümpl.<br />

“Strawberry Cactus,” “Pitaya”<br />

Description Plate 15<br />

stems: Cylindrical, tapering at the upper end to a somewhat<br />

pointed tip. They are up to at least 15 inches long and 3<br />

inches thick, light green in color and very soft and flabby.<br />

They have 7 to 10 broad, rounded ribs with shallow furrows<br />

between them and very Slight enlargements at the areoles.<br />

These stems branch and cluster very slowly to form loose,<br />

irregular clumps which I have never seen with more than 8<br />

heads. The stems branch at all angles from about the ground<br />

level, and the heavy, flabby stems seem too soft to stand upright,<br />

so the clump is usually partly sprawling or semiprostrate—never<br />

rounded, compact, and regular as is that of E.<br />

stramineus.<br />

areoles: These are circular, about 1/4 of an inch in diameter<br />

and 1 to 11/2 inches apart, having much white wool when<br />

young, but losing all of it with age.<br />

spines: All of the spines are white to light brown and somewhat<br />

translucent when young, becoming opaque when old.<br />

They grow from enlarged bases. The radial spines are 5 to 9<br />

in number, often very irregular in size, being almost bristlelike<br />

to medium in thickness, 1/2 to 11/2 inches long, the upper<br />

ones shorter than the lower ones and these upper ones sometimes<br />

pushed aside or eliminated entirely by the large bases<br />

of the centrals. There are 1 to 5 very conspicuous central<br />

spines curving or spreading in all directions from mature<br />

areoles. They are very large, 11/2 to 3 inches long and from<br />

1/16 to 1/8 of an inch in thickness. They are also markedly<br />

flattened and sometimes ridged. It should be mentioned that<br />

the development of these centrals is very slow, so that young<br />

areoles often have only 1 or 2 shorter, round centrals, which<br />

makes them look almost exactly like typical E. enneacanthus<br />

spines. And when the plant is growing in conditions not entirely<br />

favorable to it these spines do not achieve their full<br />

development; thus large stems will sometimes have entirely<br />

juvenile spines. I have collected such plants, especially near<br />

the Devil’s River, which I would have taken for the other<br />

species except for the more robust, lighter green, and more<br />

flabby stems, and only after two full years in better growing<br />

conditions have these plants gone ahead to produce the typi-<br />

cal large growth of central spines.<br />

flowers: Magenta in color. Otherwise similar to those of<br />

E. stramineus, but not nearly so beautiful because they are<br />

much smaller in size, and because they fail to open widely<br />

and are produced in comparatively sparse numbers, each<br />

clump having only a few blossoms each year. They are only<br />

2 to 3 inches long and about 2 inches across. There are about<br />

10 outer petals, green in the centers with pinkish edges. There<br />

are about 10 inner petals with entire edges, narrow bases<br />

which broaden considerably and then end in a broad tip with<br />

a prolonged point at the apex. The bases are greenish with an<br />

orange area above that shading into magenta on the broader<br />

ends of the petals. The filaments are brownish and the anthers<br />

yellow. The style is long and white, while the stigma lobes<br />

are 8 to 10 in number, and green. The ovary tube has white<br />

spines 1/4 to 3/4 of an inch long, and has almost no wool upon<br />

it.<br />

fruits: Globular, 1 to 1/2 inches long, with many deciduous<br />

spines. It is said to be as edible as that of E. stramineus.


Rumpl -> Rümpl<br />

48 cacti of the southwest<br />

Range. Near the Rio Grande from near El Paso to the mouth<br />

of the Devil’s River, and extending into Mexico.<br />

Remarks. E. dubius is very closely related to E. stramineus,<br />

which grows in similar situations, but usually at higher elevations.<br />

E. dubius can hardly be mistaken for the other cactus,<br />

however, since it has fewer branches and ribs and poorer floral<br />

development, and it never grows in the huge, dome-shaped<br />

masses characteristic of E. stramineus. No authority seems to<br />

have suggested any combination of the two. E. stramineus is<br />

much more popular with collectors because only in the size of<br />

its spines does E. dubius excel; and yet, because it has more ribs<br />

and closer areoles, E. stramineus appears enclosed in its spines,<br />

while the stems of E. dubius are not obscured by its heavier,<br />

larger ones. Only on the hills just north of the Big Bend National<br />

Park have I seen these two species growing together; here<br />

the young plants are sometimes hard to distinguish, but the<br />

large, mature ones are easily told apart.<br />

E. dubius also is close to E. enneacanthus var. carnosus, in<br />

its stem structure, but it does not ever become so huge as that<br />

other form, and the flower developments of the two are the<br />

opposite extremes of the group.<br />

This cactus usually grows wherever the rocky hills spread out<br />

their lower slopes toward the Rio Grande. Its favorite place<br />

is the edge of the sandy river valley at the base of the hills.<br />

Just west of Sierra Blanca, Texas, where the highway starts to<br />

climb out of the sandy valley into the Sierra Blanca range it is<br />

very abundant. Along the Rio Grande below Presidio it is also<br />

common to beyond the Big Bend National Park. The specimens<br />

east of Presidio have longer, straighter, more stout and ridged<br />

central spines than those farther west, until one gets beyond the<br />

Park. East of the Big Bend Park it is very rarely seen, and the<br />

atypical specimens I found on the lower Devil’s River must<br />

represent its eastern limit, as well as its weakest spine development.<br />

Borg, in his book, Cacti, apparently is in error when he says<br />

that it is found in the sandy wastes of southeastern Texas, since<br />

there is no record of it anywhere except in the far southwestern<br />

part of the state.<br />

Echinocereus fendleri (Eng.) Rümpl.<br />

“Fendler’s Pitaya,” “Fendler’s Hedgehog Cactus,” “Purple<br />

Hedgehog,” “Strawberry Cactus,” “Torch Cactus,” “Sitting<br />

Cactus,” “Pink-Flowered Echinocereus”<br />

Description Plate 15<br />

stems: This cactus grows as a small, loose clump of upright<br />

stems which may take various shapes, from short and almost<br />

oval to longer and cylindrical, usually tapering and therefore<br />

often somewhat conical. These stems are not usually<br />

over 12 inches tall, but may reach a maximum of 18 inches<br />

tall and about 4 inches thick in one variety. The surface is<br />

dark green and soft, often being wrinkled in appearance.<br />

There are 8 to 16 ribs, which are broad, somewhat wrinkled,<br />

and have rather conspicuous swellings around the areoles.<br />

areoles: Circular, not large, with some white wool when<br />

young, but later becoming bare. Up to only 1/2 inch apart on<br />

mature growth.<br />

spines: Rather stout from bulbous bases. This is the only<br />

purple-flowered Echinocereus I know within our area which<br />

has spines that can be called truly variegated in color. At<br />

least some of the spines on each plant have brown and white<br />

or black and white coloring in streaks along them. There are<br />

5 to 12 radial spines which are round or sometimes slightly<br />

angled and variegated brown and ashy-gray with white<br />

streaks. The lower ones are the stoutest and are 1/2 to 1 inch<br />

long; the upper lateral ones are bristle-like, usually white and<br />

only 1/4 to 1/2 inch long. The radial is missing entirely from<br />

the top of the areole. There is usually one central spine which<br />

is longer, more stout, round or only very slightly flattened.<br />

This spine is from 3/4 to 3 inches long, dark brown or black<br />

fading to gray when old, except in one variety where it<br />

reaches only 5/16 of an inch long. Some varieties of the species<br />

sometimes present 2 additional centrals.<br />

flowers: Large, to at least 3 inches in height and diameter,<br />

and a beautiful violet-purple. There are broad, greenish outer<br />

petals with violet edges. The inner petals are long, somewhat<br />

variable in shape, and violet-purple. The bases of these are<br />

narrow and dark purple-red, the petal broadening from this<br />

to a blunt or pointed tip. The filaments are green and the<br />

anthers light yellow. The style is whitish and only a little<br />

longer than the stamens. The stigma has 9 to 16 dark green<br />

lobes. The tube of the ovary has some white wool and many<br />

spines which are white or white with brown tips and mea-<br />

sure to at least 3/8 of an inch long.<br />

fruits: Almost spherical, 1 to 11/2 inches long, purplish,<br />

covered with spines which fall easily from it. It is fleshy and<br />

edible.<br />

Range. Extreme southwestern Texas, all of western New Mexico<br />

and into Arizona, Colorado, and Mexico. Its eastern range apparently<br />

stops at a line from near the upper Pecos River in<br />

New Mexico to just east of El Paso, to near Presidio, Texas.<br />

Within Texas it has been collected only in the Franklin Mountains<br />

just within the border of that state and in extreme western<br />

Presidio County.<br />

Remarks. E. fendleri is one of the most beautiful Echinocerei.<br />

Its flowers are large and of very delicate rose shades which are<br />

not matched by any of its relatives within our area. The colors<br />

of its spines are also unique for us, being truly variegated. The<br />

brown and whitish shadings in them I have not seen elsewhere<br />

except in some specimens of the more western E. engelmannii.<br />

The clumps formed by E. fendleri within our territory are not


genus Echinocereus engelmann 49<br />

large, and they do not flower as profusely as some, but it is a<br />

very worth-while cactus.<br />

Engelmann, in first describing and naming the cactus a hundred<br />

years ago, described a form of it with only 5 to 7 radial<br />

spines and no centrals, calling it variety pauperculus, but said<br />

also that on every specimen he had seen of this form some<br />

areoles had normal spines, so he suspected it was only an atypical<br />

growth. I have seen specimens from both New Mexico and<br />

Texas with this atypical spination, but each was a stunted or<br />

injured plant which could well have suffered some impairment<br />

to the spine growth, and each which I was able to grow under<br />

good conditions later put on typical spines. So I feel that this<br />

cannot stand as a distinct variety, and I therefore include this<br />

in the description of the species.<br />

E. fendleri must be considered as one of the more western<br />

Echinocerei, only dwelling a comparatively short distance into<br />

our territory. It reaches a great development in Arizona where<br />

there are at least four varieties besides the typical one. Our<br />

New Mexico plants, however, are mostly the typical form, as<br />

would be expected, since its type locality is near Santa Fe, New<br />

Mexico. Only in extreme southwestern New Mexico do we find<br />

one of these other varieties.<br />

Echinocereus fendleri var. fendleri (Eng.)<br />

Description Plate 15<br />

stems: As the species, except that the stems are usually around<br />

6 inches tall and reach a maximum of 12 inches, with only 9<br />

to 12 ribs.<br />

areoles: As the species.<br />

spines: As the species, except that the radials number only 5<br />

to 10, and the single central is 1 to 2 inches long and usually<br />

curving upward.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. As the Species.<br />

Echinocereus fendleri var. rectispinus (Peebles) L. Benson<br />

Description Plate 15<br />

stems: Forming small clumps of up to 6 or 8 comparatively<br />

firm, columnar stems, up to about 8 inches tall. There are 8 to<br />

11 ribs on these stems, which are narrower than on the typical<br />

form and not tuberculate.<br />

areoles. Similar to the species.<br />

spines: There are 10 to 12 radials very similar to the typical<br />

except that they are usually lighter in color. There is 1 main<br />

central spine which is often accompanied by 1 or 2 upper<br />

accessory centrals. The main one is stout and porrect and<br />

straight instead of curving upward. It is 1/2 to 11/4 inches<br />

long. The accessory centrals are only 1/4 to 3/4 of an inch<br />

long.<br />

flowers : Similar to those of the Species, with stigmas numbering<br />

10 to 13.<br />

fruits: As the species.<br />

Range. Extreme southwestern New Mexico and extreme southeastern<br />

Arizona. In New Mexico reported, so far, only from<br />

western Hidalgo County.<br />

Echinocereus papillosus A. Linke<br />

“Yellow-Flowered Echinocereus,” “Yellow-Flowered Alico-<br />

che”<br />

Description Plate 16<br />

stems: This plant consists of 2 or 3 to sometimes, in one<br />

variety, dozens of clustering and branching stems forming a<br />

very loose clump. The stems are slender, soft, and weak, apparently<br />

unable to entirely support their own weight, and so<br />

they lean and sprawl at awkward angles, although seldom<br />

being actually prostrate. Each stem is deep green in color, up<br />

to 10 inches long, and 1 to 23/4 inches thick. There are 7 to 9<br />

ribs which are extremely tuberculate, formed of series of<br />

conical enlargements about 3/8 of an inch high, with the<br />

areoles on the tips of them, these enlargements being separated<br />

by deep valleys which almost completely interrupt the<br />

ribs.<br />

areoles: Small, bare, 3/8 to 1/2 inch apart, crowning the<br />

tubercles.<br />

spines: Slender but rigid, straight, and round, from bulbous<br />

bases, white to brownish or yellowish in color. There are 7 to<br />

11 radial spines radiating evenly. They are whitish to yellow-<br />

brown, usually with brownish bases. The lower and lateral<br />

ones are longest and heaviest, being up to 1/2 inch long. The<br />

upper 2 or 3 are very much shorter and very slender, almost<br />

bristle-like. There is 1 central spine about 3/4 of an inch long,<br />

not much more stout than the radials, but having a very<br />

bulbous base and standing perpendicular to the stem surface.<br />

It is brownish to sometimes bright yellow, often with a dark<br />

brown base, a yellow zone in the middle, and a brown tip.<br />

flowers: Large, beautiful, and delicately fragrant, 21/2 to 4<br />

inches in diameter and height. The outer petals are oblong,<br />

reddish in their centers to yellowish at the edges-which are<br />

ragged. The inner petals lie in 2 to 4 rows, and are long, rising<br />

from narrow, bright orange-red bases, giving the center of<br />

the flower a striking red color. The upper part of the inner<br />

petal is much wider, with ragged edges and a tip either


50 cacti of the southwest<br />

somewhat pointed or blunt. This upper part of the petal is<br />

yellow, shading to almost white at the edges. The midline of<br />

each petal on the dorsal side has some unique feathery ridges<br />

and furrows extending almost to the tip. The filaments are<br />

reddish, the anthers light yellow. The style is white, and the<br />

stigma lobes are green, 10 to 13 in number, long, and broad,<br />

with a furrow running the length of the underside of each.<br />

The ovary surface has reddish scales and white spines up to<br />

about 1/4 of an inch long.<br />

fruits: Greenish, covered with short bristles.<br />

Range. A comparatively small part of south Texas bounded by<br />

a line from about 20 miles east of Laredo northeast into McMullen<br />

County, then southeast to near Alice, and from there south<br />

to within a few miles of Edinburg and back to Laredo.<br />

Remarks. In south Texas is found a cactus unique among the<br />

Echinocerei of the U.S. in having large, fragrant, yellow flowers<br />

with red centers. The plant itself is insignificant and not particularly<br />

attractive in its growth, but it is highly prized for its<br />

beautiful flowers. It would undoubtedly be a favorite with cac-<br />

tus collectors except for two facts.<br />

First, it grows in a comparatively small area of south Texas,<br />

and this not right along the Rio Grande where most collecting<br />

has been done. The center of its range is Duval, Jim Hogg, and<br />

upper Starr counties. From there it extends not quite to the Rio<br />

Grande Valley at any point on the west and south, and not<br />

quite to the coastal plain on the east. This interior country<br />

where it grows is dense brush country through which few<br />

people travel and in which fewer care to collect. This area presents<br />

some of the most difficult brush to move around in that<br />

I have encountered anywhere. And even here the cactus is an<br />

inconspicuous plant usually well-hidden under the chaparral or<br />

the tall Opuntias of the area. So it is seldom found except by<br />

the hardy.<br />

The second reason for its being so little known is that the<br />

plant is one of the most difficult to grow out of its natural<br />

habitat. It is found only on the light, sandy, limestone loam of<br />

the area. It cannot tolerate the darker soils of the Rio Grande<br />

Valley, or even of the bluffs overlooking the valley. Its range<br />

starts where this light sandy loam starts, a few miles to 50<br />

miles from the river, and at the same time it is unable to grow<br />

in the soils of the coastal plain on the east. I have the word of<br />

an experienced cactus dealer and grower in Laredo that it will<br />

not grow unprotected in that city only 20 miles from its natural<br />

range, and also that of a long-time grower of cacti in San Antonio<br />

that it will not live out-of-doors in that city less than a<br />

hundred miles above its range. I know of a grower much farther<br />

west who grows it with some success, but it is in desert soil and<br />

in real desert conditions. If conditions such as this are not painstakingly<br />

provided, the very soft flesh of the plant will rot almost<br />

overnight. I find it necessary, myself, to give it the most<br />

careful treatment to keep it alive and flowering, but the reward<br />

for this sort of care is great when one succeeds in getting a<br />

healthy plant to produce its huge yellow and red flowers.<br />

Long ago Hildmann described a variety of E. papillosus<br />

which he said has spines pink to red, passing to brownish. He<br />

called it variety rubescens. I have not found this variety, or<br />

seen anyone who knows it. Cactophiles might keep it in mind<br />

and try to discover whether it is truly a distinct variety.<br />

Echinocereus papillosus var. papillosus (A. Linke)<br />

Description Plates 16, 17<br />

stems: As the species, except each plant a clump of only up<br />

to 10 or 12 stems.<br />

areoles: As the species.<br />

spines: As the species.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. As the species.<br />

Remarks. This is the typical form of the plant, and it is discussed<br />

above. Only in one small area will the dwarf form, described<br />

next, be encountered.<br />

Echinocereus papillosus var. angusticeps (Glover) Marshall<br />

“The Small Papillosus”<br />

Description Plate 16<br />

stems: As the species except that it is markedly smaller, attaining<br />

the size of only 3 to 4 inches tall and 1 to 11/4 inches<br />

in diameter, but occurring in dense clusters of many stems.<br />

While 1 to 3 or 4 dozen stems to the plant is common, Miss<br />

Glover tells of one specimen containing 95 of these small<br />

stems. The stems are sprawling to upright.<br />

areoles: As the species.<br />

spines: As the species, except that they are shorter, the cen-<br />

tral spine reaching only 3/8 of an inch long.<br />

flowers: As the species except that they are usually slightly<br />

smaller and have a larger number of petals. The petals are<br />

usually more blunt at the tips. The yellow coloring is more<br />

greenish and the red paler.<br />

fruits: As the species.<br />

Range. Northern Hidalgo County, Texas.<br />

Remarks. While this cactus does not seem to be a separate spe-<br />

cies, it is definitely recognizable as a separate variety. It is a<br />

very interesting, dwarf-like form, found in only the portion of<br />

Hidalgo County above Edinburg, Texas, where it grows under<br />

mesquite and brush thickets.


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Rümpl<br />

genus Echinocereus engelmann 51<br />

Echinocereus pentalophus (DC) Rümpl.<br />

“Alicoche,” “Lady-Finger Cactus”<br />

Description Plate 16<br />

stems: Light green and rather flabby, up to 12 inches long,<br />

but usually only 4 to 6 inches, slender, from 1/2 to 11/4 inches<br />

in diameter. The ribs number 4 or 5 on old stems, and are<br />

low, with very shallow furrows between them so that the<br />

stem in cross-section is almost a perfect square or pentagon.<br />

There are very low tubercles at the areoles. Young growing<br />

tips have much more pronounced tubercles and much more<br />

distinct furrows between the ribs, and in some specimens the<br />

stem does not flatten out much until very old. These slender<br />

stems branch and bud off new stems at any point and at any<br />

angle, forming an unorganized mass, most of which is prostrate<br />

on the ground. Young branches often are upright for a<br />

while, but soon bend over unless supported by something.<br />

These prostrate branches root at the areoles touching the<br />

ground and the whole mass thus grows rapidly, a very old<br />

plant becoming 10 or even 15 feet in diameter.<br />

areoles: Very small, less than 1/16 of an inch in diameter,<br />

with yellowish wool when young, then bare. They are spaced<br />

3/8 to 3/4 of an inch apart.<br />

spines: Very short and slender, but rigid, from bulbous bases.<br />

There are 3 to 6 radials which are 1/16 to 1/2 inch long, rosecolored<br />

when growing, then lightening to brownish with dark<br />

tips and finally becoming gray with dark tips when old.<br />

There is often no central, but one may be present, which is<br />

somewhat stouter than the radiais, darker in color, and from<br />

3/8 to 1/2 inch long. This central points upward if present, and<br />

sometimes the upper radial assumes this direction without<br />

moving from its position.<br />

flowers: 3 to at least 4 inches in diameter as they open<br />

widely. The outer petals have greenish midlines and pink<br />

edges. The inner petals are about 18 in number and long, their<br />

bases rather narrow and the upper part broadening to a<br />

maximum of 3/4 of an inch wide toward their tips. The edges<br />

are entire, the end of the petal blunt and often somewhat<br />

notched, but still having a small point at the apex of the<br />

midline. The basal third to one-half of each petal is white<br />

shading to yellowish, while the upper part is a light cerise.<br />

This results in a beautiful flower, cerise-pink with a large<br />

whitish center. The filaments are greenish, the anthers yellow.<br />

The style is composed of 10 to 16 linear, olive-green lobes<br />

with a yellow furrow on the ventral side of each. The ovary<br />

wall has much long white wool and many brown, bristle-like<br />

spines, giving it a woolly covering.<br />

fruits: Egg-shaped, green, 1/2 to 3/4 of an inch long, covered<br />

with white wool and brown spines.<br />

Range. Northeastern Mexico, extending across the Rio Grande<br />

into its lower valley up as far as Rio Grande City. It nowhere<br />

occurs over 15 or 20 miles north of the river.<br />

Remarks. The most common Echinocerei of deep south Texas<br />

and the lower Rio Grande Valley on both sides of the river are<br />

several forms of prostrate cacti. These all have very slender<br />

stems, usually 1 inch or less in diameter. They branch very<br />

profusely, forming masses of these stems, most of which lie entirely<br />

or partly on the ground, rooting where they touch the<br />

ground, and spreading the plant outward in this way. In some<br />

of them there are upright stems or tips of stems when young,<br />

but in all at least the old stems are prostrate. They are all known<br />

locally as the alicoches.<br />

In spite of the lowly forms of their growth, these are interesting<br />

cacti, and are justly famous for the beauty of their large<br />

pinkish flowers. These flowers, often 4 inches or more in diameter,<br />

are particularly startling when numbers of them appear<br />

upon a prostrate mass of these slender stems.<br />

The number of forms in this group is not large, but they were<br />

among the very early cacti described, and the differences between<br />

them not being strikingly obvious, there has been a his-<br />

tory of confusion over them.<br />

E. pentalophus was described very early, in 1826, by De<br />

Candolle. At that time he also described three varieties of it,<br />

variety simplex, variety subarticulatus, and variety radicans. It<br />

is very difficult to know today just what these varieties were,<br />

but we can see a hint here that the species is somewhat variable<br />

and that identification of these forms will be confusing. In<br />

1837, Pfeiffer ascribed the new names of variety propinquus<br />

and variety leptacanthus to the first two of De Candolle’s varieties<br />

respectively, and he was followed in this by Salm-Dyck<br />

in 1850. Then, in 1898, Schumann dropped the species name and<br />

spoke of Cereus leptacanthus.<br />

In the meantime, Engelmann had coined a new name for the<br />

plants he studied, calling them Cereus procumbens. We may<br />

perhaps understand why Engelmann did this if we notice a<br />

statement he made in his Synopsis of Cactaceae in 1856. He said,<br />

C. pentalophus is “…similar, but an erect plant.” Thinking<br />

this, he naturally gave the prostrate plants he had before him a<br />

new name. But it seems clear that in the variety simplex of E.<br />

pentalophus as described by De Candolle, there is a form with<br />

more numerous upright branches and with central spines, and<br />

that the variety subarticulatus was a more uniformly prostrate<br />

plant with no central spines on most areoles, which seems to fit<br />

Engelmann’s specimens. Accordingly, E. procumbens is considered<br />

widely to be a synonym of E. pentalophus, and perhaps<br />

more specifically, of E. pentalophus var. subarticulatus.<br />

More recently, there has been mentioned a Cereus runyonii<br />

from near the mouth of the Rio Grande. It has never been described<br />

completely, but it seems to have the same characters as<br />

E. pentalophus except for the fact that it has underground stems<br />

with areoles and short spines on them. I do not believe that this


Rumpl -><br />

Rümpl<br />

52 cacti of the southwest<br />

species has been validly published, its mention being only in a<br />

list published in 1926 by Orcutt, but the area where it was<br />

found is one of very sandy coastal dunes, and since these cacti<br />

lie prostrate, it seems easy to account for the unusual occurrence<br />

of underground stems by the drifting action of wind-blown<br />

sand covering up the usual prostrate stems of the ordinary E.<br />

pentalophus.<br />

This is the cactus called the lady-finger cactus because of the<br />

small size of its stems, which are perhaps the slenderest of the<br />

genus. But anyone happening upon a clump of these stems, pros-<br />

trate and spreading over and under each other to cover the<br />

ground like a mass of green, intertwining snakes, might think<br />

of a less pleasant name for it. However, when this mass covers<br />

itself with dozens of large, cerise flowers it is a different matter.<br />

It is a favorite because of the beauty of its flowers and because<br />

it is among the easiest of cacti to grow. Wherever even a piece<br />

of it touches sandy, well-drained soil it roots and grows, and<br />

with its prostrate, spreading habit it soon fills a window box<br />

or overflows a pot.<br />

Echinocereus berlandieri (Eng.) Rümpl.<br />

“Berlandier’s Alicoche”<br />

Description Plate 17<br />

stems: A sprawling, clustering, and branching plant with the<br />

older parts of the stems prostrate, but with the growing tips<br />

and sometimes complete stems erect. The stems are deep green<br />

to bright green and slender, not flabby, up to about 6 inches<br />

long and 1/2 to 1 inch thick, although not usually over 3/4 of<br />

an inch thick. There may be 4 to 6 ribs, composed of rows of<br />

distinct, conical tubercles at the areoles with rounded notches<br />

between them often interrupting the ribs entirely. These rows<br />

of tubercles usually spiral about the stem, and often stop and<br />

start abruptly, so that the upper part of the stem may have<br />

either more or fewer ribs than the lower part of it.<br />

areoles: Round, 1/8 of an inch in diameter, having much<br />

white wool when young, but becoming entirely bare with age.<br />

They are from 3/8 to 5/8 of an inch apart.<br />

spines: Slender to rather stout, round, from bulbous bases,<br />

very light yellow to white, and rather translucent, with<br />

slightly brownish bases and very minutely brown-tipped<br />

when young, becoming all gray and opaque when old. There<br />

are 6 to 8 radial spines, slender and yellowish white, some-<br />

times with tips and bases brownish. They are 1/4 to 11/4 inches<br />

long, the lower 3 being longest, and the upper 3 being much<br />

shorter and more bristle-like, except on areoles without a<br />

central, where the upper radial often is longer and heavier<br />

and moved into the areole almost to the position of an upward-pushing<br />

central. On typical plants there is 1 central<br />

on most areoles. It is much stouter and longer than the radials,<br />

being 1/2 to 11/2 inches long, and stands almost perpendicular<br />

to the stem or else turned upward. The areoles low on the<br />

stems tend to have short, weak centrals or none at all, while<br />

those toward the tips have robust, long centrals, which often<br />

gives the plant a curious top-heavy look. Some specimens are<br />

weaker in armament and on them centrals are found only on<br />

occasional areoles, but I have never seen a plant of this species<br />

wholly without centrals.<br />

flowers: Said to be 2 to 4 inches in diameter in some descriptions,<br />

but all I have seen were 3 to 5 inches across, while<br />

only 2 inches tall, opening very widely. The petals are in<br />

only one row, always very long and narrow, not usually over<br />

3/8 of an inch wide at any point. They have very gradually<br />

tapering, pointed tips with entire edges. The bases, and at<br />

least the lower one-third of the petals’ length are whitish,<br />

while the tips are cerise-pink. The stigma lobes are 7 to 11 in<br />

number. The surface of the ovary has very short white wool<br />

and many longer, weak, dark spines.<br />

fruits: Green, egg-shaped, about 3/4 of an inch long, covered<br />

with long, dark bristles, but having very little, and very<br />

short wool.<br />

Range. Throughout most of south Texas below a line drawn<br />

from the Rio Grande just northwest of Laredo to near Uvalde,<br />

Texas, then southeast to just below San Antonio and down to<br />

Corpus Christi. Most common along the Nueces River and the<br />

lower Rio Grande.<br />

Remarks. Berlandier’s alicoche is very close to the true alicoche,<br />

E. pentalophus, and has often been confused with it. However,<br />

it is a more northern form occurring over a wide area of south<br />

Texas, while the other is restricted very closely to a small section<br />

of the lower Rio Grande Valley. E. berlandieri was first<br />

discovered and described from along the Nueces River, where<br />

E. pentalophus never grows. It has flowers of the same color<br />

as the other cactus, but they are larger, with fewer, more narrow<br />

and pointed petals, fewer stigma lobes, and only short<br />

wool on the ovary surface. It also is a more upright-growing<br />

plant, never with all stems prostrate and never branching or<br />

rooting except near the base of the stems. Its stems are shorter,<br />

more slender, more tuberculate, and less flabby. It has longer<br />

and more numerous spines, usually with long and robust central<br />

spines on most areoles, while E. pentalophus usually has no<br />

centrals at all and at most sometimes produces a very much<br />

shorter and weaker one on an occasional areole.<br />

It must be admitted that there have been found plants which<br />

seem to be puzzling intermediates, if only such things as spine<br />

character are observed. The secret to this seems to be that it is<br />

not uncommon to find immature or stunted specimens of E.<br />

berlandieri which present only the shorter spines typical for<br />

E. pentalophus, but when I have grown such plants in good<br />

conditions they have, within a year, put on the greater arma-


genus Echinocereus engelmann 53<br />

ment typical of this species, as well as shown the narrower,<br />

more pointed petals and shorter wool on the flower tube which<br />

are a sure sign of this species.<br />

The two forms are so close that some have thought that perhaps<br />

they should be combined, but since we really know nothing<br />

experimentally about their actual relationship, any combination<br />

would be only conjectural at this stage. A combination<br />

here would make the nomenclature extremely confusing. It may<br />

well be that E. berlandieri is the plant referred to as C. pentalophus<br />

variety simplex by De Candolle and variety propinquus<br />

by Pfeiffer, since that form was supposed to have central<br />

spines. This could then be the plant referred to by Engelmann<br />

as the “erect” C. pentalophus. But if this is so, it is hard to see<br />

why he went on to give it a new species name at all.<br />

E. berlandieri has been further confused with the next species<br />

we will take up, E. blanckii. Britton and Rose went so far as to<br />

claim that these two very distinct and different species were<br />

synonymous. They even feature a clear photograph of E. berlandieri<br />

as E. blanckii, while at the same time having a correct<br />

color plate of E. blanckii. We do not know how much Coulter’s<br />

and even Engelmann’s descriptions of E. berlandieri were influenced<br />

by a failure to distinguish between these two species,<br />

since neither of those authors mentions E. blanckii at all.<br />

E. berlandieri, when once identified certainly and separated<br />

from its relatives, is a handsome plant forming small thickets<br />

of slender, sprawling stems standing at least 6 inches high. When<br />

blooming, a big clump of it will have dozens of the very large<br />

flowers with beautiful, two-colored, sharp-pointed petals.<br />

It may formerly have been common over its wide range, but<br />

it now is seldom seen anywhere (and then in only small clusters)<br />

except on the low banks overlooking the sandy plains of<br />

the Rio Grande delta below Brownsville, where it is still found<br />

under almost every bush. Still, an occasional specimen is found<br />

much farther north. I found a fine specimen growing 20 miles<br />

south of D’Hanis, Texas, which puts it only about 50 miles<br />

southwest of San Antonio. I have also found several nice specimens<br />

a few miles north and east of Laredo, Texas. These points<br />

apparently are near the extreme northwestward limit of its<br />

range.<br />

Echinocereus blanckii (Poselgr.) Palmer<br />

“Alicoche”<br />

Description Plate 17<br />

stems: At first erect, but later sprawling. They are very soft<br />

and usually wrinkled, twisted, and contorted when old,<br />

bright green in color when in perfect health, but rapidly<br />

fading and turning yellowish-red when overexposed to the<br />

sun. The stems are 3/4 to 11/2 inches in diameter and to at<br />

least 14 inches long. There are 5 to 8 ribs. On young, growing<br />

tips these are composed of rows of practically unconnected<br />

tubercles, but on older parts of the stems these ribs are continuous,<br />

with the tubercles less prominent and with almost no<br />

depressions between tubercles or adjacent ribs, making the old<br />

stems almost perfect hexagons to octagons in cross sections.<br />

These ribs do not usually spiral, but individual ribs may stop<br />

or start at any point on the stem, increasing or decreasing<br />

the number of them. These stems sometimes branch above the<br />

ground, but not profusely. The plant produces most of its<br />

stems in clusters from its large underground root. Since they<br />

come forth from very small bases often only 1/4 of an inch in<br />

diameter at the ground level, if for any reason the soil is<br />

moved away from them, these young stems appear standing<br />

on very spindly bases, from which they abruptly widen to<br />

the typical size.<br />

areoles: Small, less than 1/8 of an inch in diameter, 1/4 to 1/2<br />

inch apart. They have a very little short wool when young,<br />

and then become bare.<br />

spines: Radial spines 6 to 9 in number, very slender and<br />

translucent, 3/16 to 7/8 of an inch long, radiating evenly. The<br />

3 radials on each side of the areole are the longest, and are<br />

white, sometimes with minute black tips when young. The<br />

lower radial is the shortest and is very weak, almost bristle-<br />

like. The uppermost radial is very dark brown when young,<br />

in striking contrast to the other white radials, and fades<br />

gradually to almost gray when very old, usually keeping at<br />

least a brown tip. There is 1 stouter central spine with a<br />

bulbous base. It stands perpendicular to the stem at first, but<br />

then turns downward and often is slightly curved downward.<br />

This central spine is from 1/2 to 2 inches long. It is<br />

translucent and variegated when young, with zones of very<br />

dark brown to light brown and almost white alternating<br />

throughout its length. When old it fades to almost the same<br />

gray as the other spines.<br />

flowers: Funnel-shaped, not usually opening widely. About<br />

2 inches tall by 21/4 to 31/2 inches across in their natural,<br />

partly spread state. The ovary surface has pointed, reddish,<br />

scalelike segments and clusters of white to brown spines on<br />

it, with a little short wool. The outer petals have purple<br />

midlines with lavender-pink edges, and are narrow and<br />

pointed, with smooth, unbroken edges. The inner petals are<br />

about 26 in number, widening a little to 3/8 or 1/2 inch across<br />

near the tips, their margins smooth and their tips sharply<br />

pointed. The upper four-fifths of each of these is light rose<br />

in color, while the base darkens to carmine, giving the flower<br />

a dark, reddish throat. The filaments match the center of the<br />

flower. The anthers are orange-yellow. The pink style is little<br />

longer than the stamens. There are 8 to 11 long, slender, and<br />

very light green stigma lobes.<br />

fruits: Practically unknown. There seem to be no descriptions<br />

of them except for the very general statement by Miss


54 cacti of the southwest<br />

Schulz in Texas Cacti that, “The fruit is greenish, globose,<br />

and covered with small spines.”<br />

Range. Northeastern Mexico and southern Texas, particularly<br />

in Starr and Hidalgo counties. Reported otherwise only once,<br />

and this from near Carrizo Springs, Texas, in Dimmit County.<br />

Remarks. E. blanckii is a very distinct species of prostrate<br />

Echinocereus. It was first named and described by Poselger in<br />

a German publication in 1853. Strangely, however, neither Engelmann<br />

nor Coulter mentioned it in their studies, so it was<br />

overlooked for many years. Then, in 1922, Britton and Rose<br />

confused it with E. berlandieri, stating that the two were<br />

synonymous. They published a photograph obviously of E.<br />

berlandieri as E. blanckii, but at the same time also published<br />

the best color plate of E. blanckii in bloom that I have seen. As<br />

a result of this, E. blanckii is still confused with the others of<br />

this group. I have seen numerous pictures of both E. pentalophus<br />

and E. berlandieri called E. blanckii in popular magazines and<br />

in catalogs but, due to its rarity, have not seen any good picture<br />

of the true species except the one of Britton and Rose, one in<br />

Borg’s Cacti, and one in Texas Cacti by Ellen Schulz.<br />

This species is easily distinguished from both E. berlandieri<br />

and E. pentalophus by the thicker stems with more ribs, the<br />

more numerous radial spines with the lower one smallest and<br />

the upper one dark colored. The downward-turning, variegated<br />

central spines are also very distinct from anything found in the<br />

others. When the flower is present there is no possibility of<br />

mistaking this cactus, as it has an over-all darker color with a<br />

dark center where the others have strikingly whitish centers, it<br />

opens only partly while they open extremely wide, and its<br />

petals are narrower, more pointed, and more numerous than in<br />

the other prostrate Echinocerei. Also, it normally blooms in<br />

February or March, about a month earlier than the others, but<br />

buds on disturbed plants may remain half-formed for more than<br />

two months and then open normally much after their usual<br />

season.<br />

E. blanckii is primarily a Mexican species, and it occurs in<br />

Texas mainly in Starr and Hidalgo counties. The only record I<br />

find of its occurrence outside these two counties is a single specimen<br />

which is incomplete but which appears to be of this<br />

species in the herbarium of The University of Texas. This is<br />

labeled as collected 18 miles southwest of the Dimmit-Frio<br />

county line, which would be near Carrizo Springs, Texas, well<br />

above Laredo. On the basis of this specimen its range would be<br />

rather large. Without it the range would be restricted to Starr<br />

and Hidalgo counties. But nowhere is it common, and, in recent<br />

years I know of only a dozen or so plants being brought in by<br />

professional dealers who comb the countryside continually. It is<br />

an exceedingly inconspicuous plant in the field, and it cannot<br />

stand much sun. The stems quickly turn yellow and red and<br />

wilt if exposed to the sun, so it is limited to the shady thickets<br />

where it is very hard to find. With the increase in the practice<br />

of clearing the brush cover, it will probably be one of the first<br />

species to become extinct on our side of the Rio Grande. This<br />

is unfortunate, as it is a beautiful cactus producing many fine<br />

flowers when given the conditions it requires.<br />

A possible reason for its scarceness may be this plant’s apparent<br />

inability to produce fruits. These have never been described<br />

except for the incomplete description given by Schulz,<br />

and the form may be practically sterile. A fine specimen forming<br />

a clump all of 10 feet across is fully protected within the<br />

Santa Ana National Wildlife Refuge, and over a number of<br />

years of observation this has been seen to bloom profusely, but<br />

has never set a fruit. The reason for this is unknown.


Genus Wilcoxia B. & R.<br />

The genus Wilcoxia was erected by Britton and Rose in 1909.<br />

Some earlier writers had included its members in Cereus and<br />

some in Echinocereus. Since 1909 the species in the genus have<br />

been dealt with in various ways. Berger placed them in the<br />

genus Peniocereus, while Benson returned both of these genera<br />

to the genus Cereus, and others have considered each of these<br />

a subgenus of that large genus.<br />

Whatever they have concluded about the genus, all recent<br />

writers have agreed that while these species seem to form a<br />

rather good grouping, the separation of the genus as such from<br />

other genera is not as clear-cut as they would like. Most say<br />

that any final word on the genus will have to await further<br />

investigation. With this word of caution about its status, we<br />

use the genus in the way most authorities are using it at the<br />

present time.<br />

Five species in this genus are usually recognized. They are<br />

almost entirely Mexican, only one species coming into the<br />

United States at all, and that one only into southern Texas.<br />

They are also inconspicuous cacti. In form they are very<br />

slender-stemmed, sparingly branched bushes. The stems, about<br />

5/8 of an inch or less in diameter, grow very long, but are weak,<br />

so the plants seldom attain much size unless they have trees or<br />

shrubs to recline upon. If growing in a thicket, they clamber<br />

over the other plants, some species then becoming up to three<br />

yards or more long. But the slender stems are usually well hid-<br />

den among the branches of the supporting plants.<br />

Each individual grows from a cluster of tuberous roots which<br />

provide the water storage for these cacti, whose stems are too<br />

slender to handle that function adequately.<br />

The spines are very short, 1/4 of an inch or less long, and are<br />

appressed to the stems. There is also more or less hair on the<br />

areoles.<br />

The flower is large and beautiful, bell-shaped or funnel-<br />

shaped, but with a short tube. It is reddish to purplish in color,<br />

and diurnal. The ovary surface is scaly, woolly, and covered<br />

with bristly or hairlike spines which remain on the fruits. The<br />

seeds are black.<br />

By way of summarizing the differences between the genus<br />

Wilcoxia and those genera closest to it encountered in our area,<br />

we can list the following points: genus Wilcoxia differs from the<br />

genus Echinocereus by having much more slender stems, by its<br />

clambering habit, by never being caespitose, by having fascicled,<br />

tuberous roots, by producing its flower from within its spine<br />

areole instead of from a rupture of the stem epidermis just<br />

above the areole, and by the difference of seed form. Wilcoxia<br />

differs from the genus Peniocereus by having even more slender<br />

stems with 8 ribs where that genus has only 3 to 6 ribs; by having<br />

fascicled tubers while that has a single extremely large taproot;<br />

by producing red to purple flowers with short perianth<br />

tubes, which open during the day, instead of white or very occasionally<br />

rose flowers with long tubes, which open nocturnally;<br />

by having fruit with wool and bristles instead of with rigid<br />

spines; and by the difference in seed form.<br />

Wilcoxia poselgeri (Lem.) B. & R.<br />

“Pencil Cactus,” “Dahlia Cactus,” “Sacasil”<br />

Description Plate 17<br />

roots: A cluster of dahlia-like tuberous roots usually half a<br />

dozen or more in number. Each tuber is rather spindle-shaped,<br />

up to about 4 inches long and about 1 inch in diameter.<br />

stems: Long, very slender, and sparingly branched to form a<br />

very weak bush standing by its own strength sometimes to<br />

2 feet tall, but when supported by the branches of a thicket


56 cacti of the southwest<br />

sometimes attaining a height of 3 or even 4 feet. Each stem<br />

is round with 8 very low and inconspicuous ribs and is from<br />

1/4 to 5/8 of an inch thick. The upper parts of the stems are<br />

green, but the lower parts become brown and woody.<br />

areoles: Very small and very close together, with some white<br />

wool.<br />

spines: There are 9 to 12 very slender radial spines only about<br />

1/16 to 3/16 of an inch long. They lie perfectly flat against the<br />

surface of the plant and are white or gray in color. There is<br />

1 central spine about 1/4 of an inch long, which is turned<br />

upward against the upper radials. It is white, whitish tipped<br />

with brown, or sometimes all dark.<br />

flowers: Borne on the sides of the stems, very near to their<br />

tips. The flowers are funnel-shaped and 11/2 to 2 inches wide<br />

by 2 to 21/2 inches long, deep pink in color. The outer petals<br />

are narrow with greenish midribs and pink edges. Inner petals<br />

are broader but still linear and sharply pointed, deep pink or<br />

rose shading to lighter edges. The stamens are pale yellow. The<br />

stigma has 8 long green lobes. The ovary surface has reddish<br />

scales upon it and has a dense covering of long wool and<br />

black-and-white hairlike bristles. Each flower opens about<br />

noon and closes before night, usually for 2 or 3 days.<br />

fruits: Practically oval in shape, becoming nearly dry and<br />

remaining covered with the wool and bristles. The seeds are<br />

black.<br />

Range. From western Hidalgo County, Texas, along the Rio<br />

Grande to slightly beyond Laredo, Texas, and in adjacent Mexico.<br />

It has not been reported more than about 30 miles away<br />

from the Rio Grande on the U. S. side.<br />

Remarks: The pencil cactus is a fascinating member of this<br />

family, but so inconspicuous and actually camouflaged in its<br />

native habitat that few people have seen it. It is rather common<br />

in its range, but always grows in thickets where its slender stems<br />

lie upon those of the bushes and trees. The support of these<br />

thickets seems practically essential to it, since individuals un-<br />

fortunate enough to be growing in the open rarely attain a<br />

height of over a foot or so. Within thickets in the same area<br />

one finds many individuals 2 or 3 feet tall and sometimes even<br />

more. Yet these are extremely hard to see. I have found that<br />

almost any good thicket in the area will have one, but about<br />

the only way to discover it is just to sit down and study the<br />

tangle of branches until it becomes apparent.<br />

Once discovered, the cactus usually prompts exclamations of<br />

wonder. It presents the most slender stems of any U. S. cactus<br />

except Opuntia leptocaulis. These may extend for several feet,<br />

and branch, but they do not increase in size with age, remaining<br />

almost exactly the thickness of a lead pencil. Its common name<br />

is well chosen. But in spite of this slenderness, the plant may<br />

reach quite a good size, with a dozen or more branches. And once<br />

having gotten so large, it can really amaze us with the production<br />

of up to two or three dozen of its large, beautiful flowers<br />

at a time.<br />

Each plant grows from a cluster of fleshy roots. It does best<br />

in sandy, loose soil, and can be grown quite easily in cultivation<br />

if treated much like other cacti of the region. However the individual<br />

plants seem to lose vigor after a few years. While they<br />

may remain alive for up to ten years or so, they seem to largely<br />

stop growing or blooming after the first three to five years. It<br />

seems that there must be a maximum size and number of the<br />

fleshy tubers and that these are not replaced with age. Mrs. Roy<br />

Quillin (Ellen Schulz), long a student of these cacti, always<br />

keeps several beautiful specimens of this cactus growing in pots,<br />

but she keeps them vigorous by cutting off the upper branches<br />

every few years and, after rooting these for her new starts,<br />

throwing away the whole of each plant which has passed its<br />

prime. Others grow them very successfully to very large size by<br />

grafting them upon other cacti.<br />

Wilcoxia poselgeri has been known in the past as Echinocereus<br />

poselgeri Lem., and as Cereus poselgeri Coult. Even earlier it<br />

was known under the name Cereus tuberosus Poselgr. and Echinocereus<br />

tuberosus Rümpl. It is the only U. S. species of Britton Rumpl. -><br />

Rümpl.<br />

and Rose’s genus Wilcoxia.


Genus Peniocereus (Berger) B. & R.<br />

This is another small genus erected by Britton and Rose. The<br />

name means something like thread cereus, referring to the slen-<br />

der stems of all the members.<br />

Before Britton and Rose made this separation, the group had<br />

been part of the large genus Cereus. It is very hard to show<br />

significant characters to distinguish this genus Peniocereus from<br />

several other closely related genera. Perhaps its standing as a<br />

separate genus cannot be well justified, but the old genus Cereus<br />

has been so subdivided and reduced that it would be impossible<br />

to weigh the question of putting it together again without a<br />

restudy of many Central and South American genera, and that<br />

places the question beyond the scope of this book. It should be<br />

noted that Benson, in his 1950 study, placed these small genera<br />

back into Cereus, but that at the same time Backeberg and<br />

others have gone on very rapidly carving new genera out of<br />

the territory formerly covered by the original genus Cereus. It<br />

appears to be a matter which will be decided by the turn of<br />

taxonomic philosophy.<br />

One cannot even state the number of species in the genus<br />

Peniocereus definitely or give an unequivocal set of characteristics<br />

for the genus, because these will be different depending<br />

upon whose limits to the genus one elects to follow. The number<br />

of species will be from two in the case of the most restrictive<br />

authors to seven in the case of Backeberg, who places back into<br />

this genus such things as Marshall’s genus Neoevansia and some<br />

species from the genus Acanthocereus.<br />

Therefore, we can only give characteristics of this genus in a<br />

very general way here. We can say that the members of the<br />

genus all have a single, extremely large, fleshy taproot, from<br />

which grow one to several slender stems which are ribbed at first<br />

but then become round. All have fragrant, nocturnal flowers<br />

with long perianth tubes, the flowers produced from within the<br />

spine areole, and all have very short spines on the stems, and<br />

rigid spines on the fruits.<br />

A comparison of this genus with the genus Wilcoxia was<br />

made in the discussion of that genus. The other genus in our<br />

area to which it is most closely related is Acanthocereus. The<br />

differences between these two genera are as follows: Peniocereus<br />

has a huge, fleshy taproot, stems ribbed but becoming<br />

round when old, very short spines, and more elongated fruits,<br />

while members of Acanthocereus have fibrous roots, stems always<br />

markedly ribbed, much longer spines, and fruits more<br />

nearly spherical.<br />

Peniocereus greggii (Eng.) B. & R.<br />

“Arizona Queen of the Night,” “Texas Night-Blooming Cereus,”<br />

“Deer-Horn Cactus,” “Chaparral Cactus,” “Sweet-<br />

Potato Cactus”<br />

Description Plate 18<br />

roots: Each plant has a single, huge, fleshy taproot. This is<br />

roughly carrot-shaped or turnip-shaped, but often takes on<br />

the shapes of crevices between rocks and other obstacles<br />

through or around which it grows, and so may be greatly<br />

distorted. A typical plant of medium age will possess a root<br />

8 to 12 inches long and 3 to 5 inches in diameter at its<br />

thickest point, but very old plants have been reported with<br />

the root up to 2 feet in diameter and weighing up to 125<br />

pounds.<br />

stems: Slender, erect, and sparingly branched to form a weak


58 cacti of the southwest<br />

bush, 1 to reportedly at least 9 feet tall. Young branches are<br />

dark, dull, grayish-green, quickly becoming reddish when in<br />

too much sun. They are from 1/2 to 1 inch thick, with 3 to 6<br />

very strong ribs. Old stems do not increase in size, but rather<br />

shrink to a smaller diameter, and in the process the surface<br />

loses its areoles and ribs and becomes brownish, woody, and<br />

circular in shape.<br />

areoles: Very tiny and closely situated on tiny tubercle-like<br />

prominences crowning the ribs. There is much white wool at<br />

the growing tips, some of which remains when the areoles<br />

are older. They are round to elliptical in shape.<br />

spines: Each areole has 6 to 9 spreading radials and 1 or sometimes<br />

2 centrals. All of these are blackish fading to gray;<br />

they are very short, being only 1/32 to 1/8 of an inch long, but<br />

stout and rigid, from bulbous bases.<br />

flowers: Strictly nocturnal. Produced from well down on the<br />

sides of the branches, these flowers are large and beautiful, as<br />

well as extremely fragrant. In size, they are 5 to 8 inches<br />

long by 2 to 3 inches in diameter; in color, whitish with the<br />

outer perianth segments somewhat tinged with brown; in<br />

shape, having a very long, very slender perianth tube, with<br />

the inner perianth segments spreading very widely and the<br />

outer segments usually recurved. The ovary surface has tubercles,<br />

and the areoles on these bear short, rigid spines. The<br />

outer surface of the elongated tube has scales and longer,<br />

more bristle-like spines. The perianth segments are lanceolate<br />

and sharply pointed. The stamens are erect, exserted, with the<br />

anthers cream in color. The style is slender; the stigma, white.<br />

fruits: These are 2 to 3 inches long, ovoid in form, with the<br />

upper end attenuated and ending in the persistent, dried remains<br />

of the perianth, which is sometimes referred to as the<br />

“beak.” Some authors give the size of the fruit as 5 to 6 inches<br />

long, but this includes the dried perianth, which is not actually<br />

part of the fruit and which usually hangs downward<br />

anyway. The surface of the fruit is strongly tuberculate, each<br />

tubercle bearing a round, black areole about 1/8 of an inch in<br />

diameter. These areoles bear short, black, rigid spines. The<br />

color of the surface remains bright green until after they are<br />

very ripe and the pulp is a beautifully contrasting magenta.<br />

Later as the fruits soften and start to disintegrate the surface<br />

becomes a brilliant orange-red. The seeds are black, very<br />

broadly obovate, and about 1/8 of an inch or a little more in<br />

greatest diameter.<br />

Range. Trans-Pecos Texas, west through southern New Mexico,<br />

across southern Arizona, and in adjacent Mexico.<br />

Remarks. Peniocereus greggii is one of the most fascinating of<br />

the cacti. It is one of only two “night-blooming cereus” species<br />

found in our area. It is justly famous for its large, extremely<br />

fragrant, nocturnal flowers—so famous that one finds a rivalry,<br />

as seen in its names: both Arizona queen of the night and Texas<br />

night-blooming cereus.<br />

These flowers are not the only amazing things about the cactus.<br />

The blooms come out on the sides of very slender stems. The<br />

branching of these stems creates a spindly bush which always<br />

grows up within a thicket of other plants. The other plants<br />

about it provide not only the support which is necessary if it is<br />

to stand over a foot or two tall, but also the light shade which<br />

it likes, and a shield of stems and branches among which it is<br />

extremely well camouflaged.<br />

The cactus is always rare in all of its range, and this fact,<br />

together with its camouflage, makes it almost completely foolish<br />

to start out anywhere looking specifically for it. The chances are<br />

overwhelmingly against one’s finding a specimen. But the plant<br />

has to face the crucial few days when it must expose itself in<br />

flower, and this has been the downfall of many a fine specimen.<br />

The standard way of searching for it has often been to<br />

search the desert at night with good lights, under which the<br />

large white flowers stand out like signs. Or, even more simply,<br />

some merely follow the wonderful fragrance of these flowers<br />

back to the plant. This scent is said to carry sometimes up to<br />

one-fourth of a mile in the desert.<br />

Once the plant is located and the searcher sees its slender<br />

stems, he has still not seen perhaps the most amazing part, for<br />

like the iceberg this plant has most of its bulk underground.<br />

The bush above ground may be one foot or several feet tall. This<br />

depends almost entirely on the conditions in which it is growing<br />

and not on its age, since the stems die back very quickly during<br />

severe seasons and are usually destroyed whenever they grow<br />

past the limit of whatever chaparral they may be surrounded by.<br />

So any specimen of this cactus which is old enough to stand a<br />

foot or more high will normally have a taproot 6 inches long<br />

and 3 inches thick at the minimum, and without being any taller<br />

it may as easily be an ancient specimen with the root weighing<br />

dozens of pounds. There is hardly any way to predict the size of<br />

this enormous root, where the water storage of this plant is<br />

taken care of so that the stems may remain so inconspicuously<br />

slender among those of the thicket.<br />

Bearing this in mind and noting the fact that it grows invariably<br />

in very rocky or very hard, clayey soil, it is a coura-<br />

geous person who starts to dig up this cactus. The root conforms<br />

to the shape of rocks and other roots it has to grow between,<br />

and getting it out is usually a good half-day’s work for a strong<br />

man.<br />

Once acquired, keeping the plant alive is also difficult. The<br />

huge root is very vulnerable to fungus if it is kept in soil even<br />

a little too damp. If rotting is avoided, the plant seems to<br />

decline gradually over a few years’ time in greenhouse cultivation<br />

or in pots. I have only been able to keep my own specimens<br />

healthy, growing, and blooming, even in the climate of San Antonio,<br />

when they are planted in outdoor beds shielded from the<br />

rain. It is no wonder they are so seldom seen in collections.


genus Peniocereus 59<br />

Engelmann, at one time, distinguished two varieties of this<br />

species, which he called variety cismontanus and variety transmontanus.<br />

Few have since tried to separate them, and we have<br />

seen no clear division in any of the material we have studied.<br />

Kuntze, in 1919, described plants of this species collected at<br />

Organ, New Mexico, which had light purple flowers instead<br />

of white. He established the new variety roseiflorus for these,<br />

but I will not list this variety separately, for two reasons. We<br />

have already seen how unwise it is to erect varieties on flower<br />

color alone, and there seems to be no other difference in these<br />

plants from the typical. Also, I know of no record since then of<br />

the collection of specimens with purplish flowers.


Genus Acanthocereus (Berger) B. & R.<br />

Here is a small genus of somewhere around a dozen species<br />

carved out of the huge old genus Cereus. No one seems to dispute<br />

this one. Whether it is so little criticized because it is more<br />

of a natural group or because so little is certainly known about<br />

it is a question which might occur to anyone reading the<br />

numerous but remarkably incomplete and often contradictory<br />

statements about its members. From Linnaeus on we have had<br />

not a lack of, but actually too many, references to these plants.<br />

What we do lack is enough good data on which solid decisions<br />

can be based.<br />

The members are more or less shrubby plants. The stems grow<br />

upright at first, but usually cannot support their own weight<br />

for long, and thus recline upon some support—usually other<br />

plants—or else become more or less prostrate and thicket-form-<br />

ing. Supported stems may grow to at least twenty feet tall<br />

and branch sparingly, but prostrate stems throw up many upright<br />

branches. Stems may be from an inch or so to 4 inches in<br />

diameter. Mature stems have 3 to 7 conspicuous ribs. The areoles<br />

are not close, and bear strong and rigid spines. The flowers are<br />

nocturnal, large, and white, with the perianth tube long and<br />

the ovary usually spiny. The fruit is round or nearly so, spiny<br />

or with the spines deciduous. The seeds are black or brown.<br />

This is a group of tropical, lowland cacti. They are never<br />

found far from a coast, and seem to thrive best on semi-arid<br />

coastal plains. However they can tolerate much more moisture<br />

than most cacti, and when given it their rate of growth is often<br />

amazing. I have had one species, when planted in an outdoor<br />

bed where it got sufficient water, produce a 6-foot stem in one<br />

summer’s growing season. But they are most severely limited<br />

by cold, being among the most tender of the cacti. A frost will<br />

kill the tips of the stems, and at 32 degrees Fahrenheit the<br />

whole of the plant above the ground is killed, although the<br />

roots may sprout again.<br />

The combination of conditions these plants need is found in<br />

various coastal areas in Central and South America. There are<br />

species of Acanthocerei native in eastern Mexico, Guatemala,<br />

and Panama, and in northern, coastal regions of Colombia,<br />

Venezuela, and Bahia, Brazil. In the U.S. they have a precarious<br />

hold along the coast in south Texas and in Florida. Besides<br />

these locations, they have been introduced in Cuba, the islands<br />

of St. Thomas and St. Croix, and probably in some other areas.<br />

Acanthocereus pentagonus (L.) B. & R.<br />

“Triangle Cactus,” “Night-Blooming Cereus,” “Organo,”<br />

“Pitahaya”<br />

Description Plate 18<br />

roots: Fibrous.<br />

stems: At first upright, later reclining, becoming practically<br />

prostrate and rooting to form low thickets unless supported.<br />

If supported, it grows to at least 6 feet tall. Such an upright<br />

stem may branch once or twice and the branches, if supported,<br />

rebranch similarly to attain a total height of at least<br />

20 feet. The thickness of the stems is extremely variable: from<br />

11/4 to 4 inches in diameter. Mature stems have 3, 4, or 5<br />

ribs so high and narrow that the stem has only an extremely<br />

small central axis and is essentially a triangle, quadrangle,<br />

or pentagon with deeply concave sides. These winglike ribs<br />

are from about 1/2 to 2 inches high. Their summits are somewhat<br />

tuberculate at first, but become almost smooth. ‘The<br />

surface is light to medium green.<br />

areoles: Round, 1/8 to 3/8 of an inch across, bulging on slight<br />

prominences on the summits of the ribs. They are spaced from


genus Acanthocereus 61<br />

about 3/4 to at least 2 inches apart, and have very short,<br />

whitish wool.<br />

spines: There are 5 to 7 radial spines which radiate rather<br />

evenly around the areole; 1 to 3 upper ones are from 3/16 to 1<br />

inch long. There are sometimes 2 upper laterals and always 2<br />

lower laterals, each 3/4 to 1 inch long. There is always one<br />

lower radial which is more slender than the laterals and 1/4<br />

to 3/4 of an inch long. There are also 1 to 4 central spines; 1<br />

is always a lower central, porrect or slightly deflexed and<br />

3/4 to 11/2 inches long. There may be 0 to 2 laterally directed<br />

centrals 3/4 to 2 inches long, and 0 or 1 upper central 1 to<br />

21/4 inches long. All spines are medium to heavy in thickness,<br />

straight and round, from bulbous bases. They are light brown<br />

when growing, then rough and gray when old.<br />

flowers: Nocturnal, extremely large and showy, but only<br />

slightly if at all fragrant. They are white in color, from 51/2<br />

to 8 inches long, and about 51/2 to 6 inches wide when fully<br />

expanded. The ovary is hardly expanded more than the long<br />

green tube, which is only 1/3 to 3/4 of an inch in diameter.<br />

There are conspicuous areoles on slight prominences rather<br />

closely placed on the ovary and becoming very widely spaced<br />

as they proceed up the tube. These have white wool in them<br />

and one to several rigid spines each; these spines are very<br />

short on the ovary but become progressively longer as they<br />

proceed up the tube until they are to about 1/4 of an inch long<br />

on the upper tube areoles. The outer perianth segments are<br />

greenish, the inner ones white. All are lanceolate and pointed.<br />

The stamens are shorter than the perianth segments, with the<br />

filaments white and the anthers straw in color. The stigma is<br />

white, with 10 to 12 close-standing lobes.<br />

fruits: Oval to rather egg-shaped, about 3 inches long by 2<br />

inches in diameter. They are slightly tuberculate, with 1 to 4<br />

spines per areole, and they are bright red and edible when<br />

ripe. The seeds are obovate, about 1/8 of an inch (3 millime-<br />

ters) in size, and bright, shining black in color.<br />

Range. In Texas in a few locations only a few miles from the<br />

coast in Kenedy, Willacy, and Cameron counties. It is reported<br />

to be also a native in southern Florida, along the east coast of<br />

Mexico, in coastal parts of and on islands near Guatemala,<br />

Panama, Colombia, Venezuela, and Bahia, Brazil, and to have<br />

been widely introduced in other places, among them Cuba,<br />

parts of the Virgin Islands, and reportedly southern Louisiana.<br />

Remarks. The triangle cactus is a very beautiful cactus, but a<br />

very difficult one to deal with in a study with only the scope<br />

of this one. It is clearly a tropical cactus which has at best a<br />

precarious foothold in the United States and which never attains<br />

its full growth here. There are many questions about it which<br />

could only be answered by a wide survey of this genus in several<br />

countries of tropical America.<br />

The very name of the cactus is the first of these questions<br />

which we cannot evaluate satisfactorily here. Cacti similar to<br />

this were seen very early, and we have numerous very old<br />

names accompanied only by notes too incomplete to be conclusive<br />

and by no type specimens. We are in the embarrassing<br />

situation of having too many names which might refer either<br />

to this cactus or to some entirely different one.<br />

Linnaeus started this problem. In 1737 he named a Cactus<br />

pentagonus and gave a very brief description of it. He gave<br />

no location for this cactus except, “Habitat in America.” The<br />

description could apply to this plant, but there are difficulties.<br />

A main obstacle is that he describes its stem as being 5-angled,<br />

which our plant never is in the U. S. But in a new note in<br />

1753, Linnaeus modified this point a little by saying that the<br />

stem was sub-5-angled (subquinquangularis).<br />

Haworth, in 1813, applied this name to a Central American<br />

plant, but, interestingly enough, no later authorities used the<br />

name for many years. Engelmann, in referring to a Mexican<br />

cactus which must have been the same as ours, ignored Linnaeus’<br />

name entirely and called the plant Cereus variabilis, while<br />

Coulter refers apparently to the same Mexican plant under the<br />

name originated at about the same time by Pfeiffer, Cereus<br />

princeps.<br />

princips -> princeps<br />

In the meantime there had been a host of other names which<br />

some have thought referred to our plant and some have<br />

thought were meant for entirely different forms. Among them<br />

were Cactus pitajaya Jacquim, published in 1761, Cereus prismaticus<br />

Wildenow, 1813, Cereus undulosus DC, 1828, Cereus<br />

acutangulus Otto, 1837, Cereus baxaniensis Karwinsky, 1837,<br />

Cereus nitidus SD, 1850, Cereus dussii Schumann, 1899, Cereus<br />

sirul Weber, 1904, and so on. Only very extensive study of much<br />

material and very astute evaluation of tiny clues could ever<br />

determine whether any one of these names is a synonym for the<br />

cactus we have in our area or a legitimate name of some closely<br />

related form.<br />

It is obvious that there was no agreement on a name for<br />

this cactus up to this century. Perhaps the most remarkable fact<br />

to be seen from the above list is that all of the authorities<br />

seemed to avoid the use of Linnaeus’ name, Cactus pentagonus.<br />

Did they consider the name unusable because the description<br />

was so brief and lacked even a type locality for the plant it<br />

was supposed to name?<br />

It remained for Britton and Rose, in 1909, to revive the old<br />

name and apply it for the first time to our Texas cactus, adapting<br />

it to their new genus as Acanthocereus pentagonus. They<br />

were able to do this by stating that the plant has 3 to 5 ribs,<br />

thus enabling it to fit the number mentioned by Linnaeus fairly<br />

well. Due to their wide influence, our plant has been known<br />

from their time until recently as A. pentagonus, and most have<br />

happily forgotten, if they ever knew, about the whole 150 years<br />

of controversy over names.<br />

We do not want to be blamed for ending a period of pleasant<br />

stability by introducing some doubt once again. It was Backeberg<br />

who, in his recent great work on cacti, had the courage<br />

to restudy the literature and who came up with a disquieting


62 cacti of the southwest<br />

decision. He it is who now tells us that Britton and Rose went<br />

back to the wrong name by Linnaeus for this cactus.<br />

Backeberg does not feel that the assumption is warranted that<br />

Linnaeus had our plant before him in describing his Cactus<br />

pentagonus. It must be admitted that this can be no more than<br />

an assumption and that for 150 years none of the authorities<br />

assumed it. But isn’t this better than the maze of names we are<br />

faced with without it? What can be offered positively?<br />

Backeberg is ready with an answer. He says that Cactus<br />

pentagonus L. is “unidentifiable.” Then he refers us to Hummelink,<br />

who has already stated that the first name, Acanthocereus<br />

pentagonus, of Britton and Rose could only have been Cactus<br />

tetragonus L.<br />

The point is that in the same publication in 1753 Linnaeus<br />

relisted his Cactus pentagonus and also described a new form,<br />

Cactus tetragonus, with stems 4-sided. This plant is even more<br />

inadequately described than the first, but it does have a more<br />

proper number of angles to the stem for our cactus. And even<br />

more important, Linnaeus pinpoints the location of it as<br />

“Curaçao, America.” This does give something to work on.<br />

Backeberg makes the flat statement that our plant is the same<br />

as the one which grows on Curaçao, and so he considers the<br />

question of names closed. He lists our cactus as Acanthocereus<br />

tetragonus (L.) Hummelink.<br />

I have not so far read any discussion of Backeberg and<br />

Hummelink’s idea, and no doubt there will be rebuttals. In the<br />

meantime I am unable to make a choice between the two proposals.<br />

I have been able to examine no specimens from Curaçao,<br />

and it seems that only a very detailed comparison of specimens<br />

from this type locality and our plants can give any basis on<br />

which to determine their relationship. In the present state of<br />

our knowledge, I feel it best to repeat the use of Britton and<br />

Rose’s name, at the same time pointing out the question about<br />

it and warning any serious cactus student that he may or may<br />

not in the future find the tide of opinion turning toward<br />

Backeberg’s theory.<br />

The triangle cactus is a beautiful one. It seems to be the only<br />

U. S. cactus which can outdo the large Opuntias in rate of<br />

growth. I have seen well-established plants produce stems 5 and<br />

6 feet tall in one growing season, and since some of these fall<br />

over and root where they touch the ground, in a few years such<br />

plants become large thickets.<br />

This is its mode of growth in the wild. It grows in the very<br />

dense brush just back from the coast in south Texas, sprouting<br />

under the trees and bushes which present an almost unbroken<br />

cover in this area. Here the large stems start upward, and if<br />

they find a tree trunk to lean against or attain a tree limb<br />

before they bend of their own weight, they go clambering up<br />

the tree and branch sparingly in it. Those branches which do<br />

not succeed in remaining upright touch the ground again and<br />

by rerooting where they touch, help the cactus to spread. By<br />

combinations of this method and seeding, in several Texas<br />

locations the cactus forms an understory plant so thick among<br />

the trees and bushes that it is difficult to walk through the<br />

stands. On summer nights the brush in such a place is so covered<br />

with hundreds of the huge white flowers that the sight is glo-<br />

rious.<br />

But often when one goes to such a place the sight is not so<br />

wonderful. This cactus is one of the most tender to frost of any<br />

I have grown. It is far more tender than most of the larger<br />

Mexican Cerei. So if one goes to such a natural stand of it in<br />

south Texas in the spring after a winter in which there has been<br />

a freeze in the area, one finds the trees draped with the dry,<br />

brown skeletons of the stems and only new shoots starting again<br />

from the roots. It seems that throughout its range in Texas the<br />

plant is killed back nearly to the ground by freezes every few<br />

years. The cold limits its range, and nowhere north of Corpus<br />

Christi can the plant be grown successfully without protection.<br />

It is definitely a tropical species barely hanging on in our area.<br />

All specimens which I have seen growing wild in Texas have<br />

been uniform in having the mature stems 3-angled—hence the<br />

name, triangle cactus. Only in cultivated specimens have I<br />

seen the stems 4-angled, and these I suspect came from Mexico.<br />

I have never seen a 5-angled mature stem in a U. S. specimen.<br />

However, this is apparently not typical of the species in the<br />

rest of its range. Many stems on the plant as it grows in Mexico<br />

have 4 ribs and some are seen there with 5, while the plant<br />

growing on Curaçao is described as having only 4 or more ribs.<br />

This and other differences may be due to the fact that in<br />

Texas we actually have only young sprouts of the plant. Here,<br />

in a lucky time when it does not get frosted for several years it<br />

may attain 8 feet or so tall, but then comes a freeze and it has<br />

to start over again. Only a hundred miles or so south along<br />

the Tamaulipas coast, where it never gets frozen, the cactus<br />

grows 20 feet or more up into the large trees and is a truly<br />

impressive giant.<br />

The largest stand of this cactus which I have seen is in<br />

Cameron County, Texas, in the brush about a mile and a half<br />

inland from the waterway called Callo Atascosa. It is almost<br />

within sight of the Atascosa Wildlife Refuge, and it is unfortunate<br />

that it is not within this protected area. Such a stand is a<br />

unique extension of the tropical flora into our area.


Genus Echinocactus Link & Otto<br />

Most of the cacti in the genus Echinocactus live up to the<br />

meaning of the name. Some of them present among the strongest,<br />

most rigid spines found on any cacti, and most of them are<br />

covered with as complete a spine cover as is found anywhere.<br />

Their main spines are often made especially troublesome by<br />

being hooked at the end, but, because they are never barbed<br />

along the shaft they are actually not as vicious as the much<br />

more slender spines of the Opuntias. Although these heavy<br />

spines are a feature of most of the Echinocacti, a few of them<br />

present more slender and flexible spines, and there are even a<br />

few spineless members of the group.<br />

These cacti are often known as the “barrel cacti,” and this<br />

term is a good one if it reminds us of their heavy, fleshy bodies<br />

and we do not let it limit our concept of them to something only<br />

barrel-shaped or barrel-sized. In size they may actually range<br />

from the huge, truly barrel-like species usually thought of under<br />

this name and sometimes weighing hundreds of pounds, to<br />

miniature forms essentially the same but, in some cases, only<br />

a few inches high when mature. In shape they are typically<br />

globular, although they may be very flattened, hemispherical,<br />

or sometimes heavily cylindrical.<br />

The exteriors of these cacti are typically firm and solid. They<br />

are shaped into from 8 to more than 20 vertical or spiraling<br />

ribs, which may be broad or narrow, high or low, smooth and<br />

even throughout their lengths or undulating, notched, or cross-<br />

furrowed, but never completely interrupted. These ribs may be<br />

thought of as partly or completely fused tubercles, but with the<br />

fusing process always clearly visible. They are never rows of<br />

completely separate tubercles. The areoles are on the summits<br />

of these ribs.<br />

The presence of these ribs distinguishes the Echinocacti handily<br />

from the tubercled cacti, but not from the Cerei. For the<br />

characters which set these apart and the characters which a<br />

botanist uses to be sure of his genera, we have to look to the<br />

reproductive structures. All of the Cerei produce the flowers<br />

on the sides of the stems, have a flower tube prolonged above<br />

the ovary, and also have a spiny ovary surface. Our Echinocacti,<br />

however, produce their flowers at or near the apex of the<br />

plant, have no distinct floral tube, and the ovary bears scales<br />

and sometimes wool, but not spines.<br />

It is harder to state differences between the Echinocacti and<br />

their other relatives. Ribs versus tubercles will usually do it,<br />

giving us a handy way to tell them from any of the Mammillarias<br />

and the members of the genus Pediocactus or genus Ariocarpus,<br />

but the genus Lophophora has what are best thought of<br />

as low ribs. These are small, spineless forms, different from the<br />

Echinocacti in various ways, but for clearly observable differences<br />

one has to look to their ovaries and fruits, which are<br />

naked, with no appendages of any kind. Most of the Mammillarias<br />

can also be separated from the Echinocacti by having<br />

such naked fruits, but several of them may have some scales on<br />

their fruits.<br />

The genus Echinocactus is used here in practically the old and<br />

original sense given it by Link and Otto. It was originally described<br />

as a large and complex grouping of ribbed cacti which<br />

produced their flowers at or near the apex of the plant, where<br />

the blossoms grew out of the upper edges of the young spine-<br />

bearing areoles, and whose ovary and fruit surfaces were to<br />

some degree scaly. Schumann organized a series of subgenera<br />

within it, and later Britton and Rose divided the old genus up<br />

into a whole array of separate small genera, leaving the original<br />

name, Echinocactus, to cover in their system only a few species<br />

of large barrel cacti.<br />

The dividing process has continued until we have had at least


64 cacti of the southwest<br />

twelve genera carved out of the old genus, among them Echinocactus<br />

(in the sense of Britton and Rose), Ferocactus, Homalocephala,<br />

Hamatocactus, Glandulicactus, Astrophytum, Sclerocactus,<br />

Thelocactus, Ancistrocactus, Neolloydia, Echinomastus,<br />

and Coloradoa.<br />

Due to the large influence of Britton and Rose’s publications<br />

these are no doubt the names that most cactus fanciers are familiar<br />

with today. Most people probably think of them as distinct<br />

and definite groupings, even though almost all would be<br />

hard-pressed if they had to try to tell the differences among<br />

them. They are annoyed when someone comes along who thinks<br />

that what they have known as Ferocactus johnsonii is actually<br />

Echinomastus johnsonii, or that Ferocactus uncinatus should<br />

have an entirely new genus made for it; and they sometimes<br />

make uncomplimentary remarks about taxonomists who bring<br />

up such subjects. This is because they are blissfully unaware of<br />

the uncertainty of these numerous small genera.<br />

Actually, these genera were erected upon very small differences<br />

and these differences are used to divide them so arbitrarily<br />

that, for instance, the presence or absence of wool on the ovary<br />

is taken to be important enough to divide the woolly Echinocactus<br />

(sensu B. & R.) from the nonwoolly Ferocactus, while the<br />

equally woolly ovary of Homalocephala is considered of no<br />

significance and the single species placed in this latter genus is<br />

linked in this system no more closely to Echinocactus than to<br />

Ferocactus. Or Glandulicactus is a genus recently erected almost<br />

entirely, as the name implies, because of the presence of glands<br />

on the plants, even though other plants left in three other<br />

genera of the group may have equally obvious glands.<br />

These small genera erected upon this sort of extreme use of<br />

the dividing rather than the synthesizing method are so<br />

poorly defined that their actual history is one of continual shifting<br />

of species from one to another and continual disagreement<br />

among the authorities who have tried honestly to define them.<br />

Echinocactus uncinatus, for instance, has been placed during<br />

the fifty years since the move to break up the original genus<br />

into five different genera, Britton and Rose placing it in Ferocactus,<br />

Knuth putting it in Echinomastus, Marshall in Thelocactus,<br />

Buxbaum in Hamatocactus, and finally Backeberg<br />

making the new genus Glandulicactus for it. Various other species<br />

have been shifted from genus to genus and the genera themselves<br />

have been both combined and redivided from time to<br />

time. The result is that the index of any recent book on cacti<br />

is used only with difficulty because it either lists many of these<br />

forms under different genera from the book last consulted or<br />

else is rendered overly long by listing each species repeatedly<br />

under the various genera each has been placed into by one<br />

authority or another.<br />

This situation looks, on the face of it, like the dividing process<br />

gone to excess, and this attitude toward the problem has<br />

been taken by one contemporary authority. Dr. L. Benson, in<br />

his book, Arizona Cacti, has ignored these newer genera and<br />

placed them all back into the old genus Echinocactus. This<br />

greatly simplifies things, and would seem a welcome move, if it<br />

can be justified.<br />

Some twenty years have passed since Benson’s recombination<br />

of these genera and there should be something more to be said<br />

on the problem. However, we actually find no more prospect<br />

of agreement on how to classify this group today than ever.<br />

Backeberg has recently published his large work with the genetic<br />

divisions at their all-time extreme. Dr. Benson has published<br />

studies on another genus, Pediocactus, in which he has applied<br />

a similar combining to some other small genera, but he has not<br />

dealt again with the Echinocacti themselves. We are left with<br />

the two fundamentally opposing views, each with a top expert<br />

in the field backing it; thus, we must make a choice for our-<br />

selves between them.<br />

To help in doing this we must look about for some newer research<br />

which might shed light on the problem. Remarkably<br />

little actual research has been reported recently on these cacti,<br />

but there has been a little. I do not want to place too much<br />

weight upon this small bit of evidence, but we must use it in<br />

choosing between the two very different approaches to this<br />

group.<br />

The most significant recent research is that of Dr. Norman<br />

H. Boke at the University of Oklahoma. He has undertaken<br />

very fine anatomical studies, particularly of the shoot and<br />

areole formation in various species of cacti. Of particular importance,<br />

he has shown that the famous groove which in some<br />

cacti extends above the spine areole-in some running a long<br />

way upward and in some a shorter distance-is really only an<br />

extension of the spine areole itself and so all flowers which<br />

come out of the upper edge of an unelongated spine areole or at<br />

the end of this groove are essentially coming from the same<br />

position in relation to the areole. He has shown that the areole<br />

in all such cases is the same in essential development and structure,<br />

described by the term “monomorphic,” as opposed to the<br />

term “dimorphic” for situations where the flower areole is separated<br />

from the spine areole entirely. This is important, since<br />

it makes all but academic the huge discussions of grooves versus<br />

merely elongated areoles and short grooves versus long grooves<br />

which have figured in trying to divide the genera within this<br />

group. The areole in all of the various members of this large<br />

group is uniformly monomorphic, a fact which is of some significance.<br />

On most of the other characters used to divide the Echinocacti<br />

into the many genera proposed, much has been written in<br />

the past, but there is no new evidence. Some authorities could<br />

be quoted on each distinguishing character proposed; and other,<br />

equally eminent students, on reasons why the character is of<br />

doubtful value.<br />

However, what little recent research has been done has made<br />

it clear that if the theory of dividing into separate genera on the<br />

basis of each and every little difference prevails, the familiar<br />

genera of Britton and Rose will, for the most part, not stand<br />

as generally used today. Instead there will have to be newer and


genus Echinocactus 65<br />

more logical realignments and even some more splittings. Backeberg<br />

has already embarked upon that sort of future with his<br />

genus Glandulicactus. An indication of the future, if this line<br />

is followed, can be seen in a statement by Dr. Boke summarizing<br />

his research on only three species in the group. He says,<br />

The anatomical data obtained in this study emphasize the<br />

similarities between Homalocephala texensis and Echinocactus<br />

horizonthalonius as well as the difference between these<br />

two species and Echinocactus grusonii. The combined list of<br />

characters suggests to the author that if E. horizonthalonius is<br />

to be retained in the genus Echinocactus, Homalocephala<br />

should be returned to the same genus. It further suggests that<br />

E. horizonthalonius could, with equal logic, be referred to the<br />

genus Homalocephala. The outcome will depend upon taxonomic<br />

opinion and also upon the results of investigations on<br />

the developmental anatomy of other species now included in<br />

the genus Echinocactus.<br />

These further investigations have not yet been reported, but<br />

since it seems clear that after they are there will probably have<br />

to come a very far-reaching and so far unpredictable realignment<br />

of the genera already proposed, it seems undesirable to<br />

perpetuate genus names in this book which have so little meaning<br />

now and which have such unpredictable futures. For this<br />

reason I am inclined to follow Benson in a conservative approach,<br />

going back to the usage of the older, inclusive genus<br />

Echinocactus, at least until more work is done. There is a trend<br />

toward synthesis as opposed to division in plant taxonomy and<br />

the distinct possibility exists that a more complete understanding<br />

will show that these recent genera are better thought of as subgenera<br />

or tribes, as Schumann first meant them to be. For these<br />

reasons I am referring them all back to the genus Echinocactus.<br />

The Echinocacti, understood in this way, are especially interesting<br />

to cactus fanciers because of their heavy bodies and conspicuous,<br />

often beautiful spine covers. They include all of the<br />

huge barrel cacti growing within the United States, which inspire<br />

such feelings of awe in us—except the huge saguaro. Because<br />

they are such favorites, collectors like especially to grow<br />

them.<br />

But fanciers must always remember that these are the real old<br />

desert rats. If one were to rate the cacti on their adaptation for<br />

survival in extreme desert situations, the finest and most beautiful<br />

of these Echinocacti would be among the most specialized<br />

for the extremes of heat and drought. And one should always<br />

remember that this makes the big, tough-looking specimens one<br />

most admires among the least capable of surviving in our cool,<br />

moist gardens and patios. It is a sad sight to see a venerable old<br />

desert barrel turning to mush in an over-watered situation. We<br />

feel sorry for a person who has to watch a desert planting of<br />

these cacti going to pieces in his yard, but there are unscrupulous<br />

nurserymen who will extract large prices for selling and installing<br />

fine old plants in places where they cannot possibly live.<br />

Many growers have shown that all of these can be grown almost<br />

anywhere if they are protected and cared for properly, in which<br />

case their beauty is well worth the trouble they take, but anyone<br />

wishing to grow them must go to the trouble of learning their<br />

requirements and providing for them properly.<br />

Knowing that some will disagree with the treatment of this<br />

group which I am using and will wish to use instead the microgenera<br />

of Britton and Rose, and realizing that the decision is<br />

at this time largely an arbitrary one, I add with the listing of<br />

each species in this group the alternate name which would be<br />

the valid one at the present time under that system.<br />

KEY TO <strong>THE</strong> ECHINOCACTI<br />

la. Plants spiny—2.<br />

2a. Central spines never more than one—3.<br />

3a. Central spine hooked if present, or else absent—4.<br />

4a. Radial spines 7 or 8 in number, 3/4 to 2 inches long, the<br />

upper 4 or 5 of them straight and flattened, the lower 3<br />

round and hooked; central spine 2 inches or more long;<br />

flower red-brown —E. uncinatus var. wrightii.<br />

4b. Radial spines 8 or more, 3/8 to 11/4 inches long, all round<br />

and straight; central spines absent or less than 13/4 inches<br />

long; flower cream or yellow in color—5.<br />

5a. Radials 10 to 19; central spine always present and 1/4 to<br />

13/4 inches long; flower large and yellow with red throat<br />

—6.<br />

6a. Stem becoming to 5 inches in diameter; radial spines<br />

10 to 13 in number; flowers 2 to 3 inches tall<br />

—E. setispinus var. hamatus.<br />

6b. Stems becoming to only 3 inches in diameter; radial<br />

spines 12 to 19; flowers 13/4 to 2 inches tall<br />

—E. setispinus var. setaceus.<br />

5b. Radials 8 to 11; central usually absent and when present<br />

(in rare cases) only 1/2 inch or less long; flower cream to<br />

pale yellowish without red coloring —E. mesae-verdae<br />

(in part).<br />

3b. Central spine present and straight—7.<br />

7a. Mature plants large, 6 to 12 inches in diameter; spines very<br />

heavy and cross-ridged; radials 5 to 8 in number; central<br />

deflexed—8.<br />

8a. Ribs 5 to 13; areoles 1/2 to 7/8 of an inch apart; stigma<br />

lobes 6 to 10; fruits imbedded in much long wool, soon-<br />

drying, and not bright colored—9.<br />

9a. Spines extremely heavy, very flattened and very severely<br />

recurving against the plant; stems remaining<br />

at most pyramid-shaped instead of cylindrical, with<br />

the ribs noticeably tuberculate —E. horizonthalonius<br />

var. curvispina.<br />

9b. Spines not so heavy, only somewhat flattened and not<br />

recurving against the plant; stems becoming columnar,<br />

with the ribs hardly tuberculate —E. horizonthalonius<br />

var. moelleri.


66 cacti of the southwest<br />

8b. Ribs 13 to 27; areoles 1 to 11/4 inches apart; stigma lobes<br />

10 to 17; fruits standing exposed beyond the wool, very<br />

slow drying, and bright red in color —E. texensis.<br />

7b. Mature plants not so large, 11/2 to 4 inches in diameter;<br />

spines rigid but not extremely heavy or cross-ridged; radials<br />

8 to 16 in number; central porrect or turned upward<br />

—10.<br />

10a. Central usually present on only some areoles and 1/2<br />

inch or less long; radials 8 to 11 in number and 1/2 inch<br />

or less long —E. mesae-verdae (in part).<br />

10b. Central always present, 5/8 to 7/8 of an inch long; ra-<br />

dials 10 to 16 in number and to 7/8 of an inch long<br />

—E. erectocentrus var. pallidus.<br />

2b. Central spines more than 1 on mature plants—11.<br />

11a. Lowermost central spine hooked; upper ones straight—12.<br />

12a. Mature plants massive in size; radial spines 12 to 20, all<br />

but the lower 3 of them slender, flexible, bristle-like, and<br />

white; central spines very heavy and cross-ringed<br />

—E. wislizeni.<br />

12b. Mature plants small to large, but not massive, radial<br />

spines all rigid; centrals various but not cross-ringed—13.<br />

13a. Mature plants large, 7 to 12 inches thick; largest cen-<br />

tral spines 2 to 6 inches long—14.<br />

14a. Ribs very high and broad and composed of massive,<br />

rounded tubercles 11/2 to 2 inches tall and in diameter<br />

at their bases; central spines 4 to 8 in number,<br />

all round or nearly so and smooth<br />

—E. hamatacanthus.<br />

14b. Ribs high but acute, composed of indistinct tubercles<br />

only about 3/8 of an inch across; central spines 4,<br />

all flat and pubescent —E. sinuatus.<br />

13b. Mature plants small, not over 6 inches and usually 4<br />

inches or less thick; largest central spines 2 inches or<br />

less long—15.<br />

15a. Radials brown, yellowish, or tan, fading to gray;<br />

upper centrals conspicuous for forming an erect “V”<br />

of straight, diverging spines; flowers greenish or<br />

yellowish—16.<br />

16a. Radials 12 or more, ribs 13; flowers greenish and<br />

not opening widely—17.<br />

17a. Roots fibrous; radials 12 to 14; flowers green<br />

suffused with rose —E. brevihamatus.<br />

17b. Roots composed of a long, fleshy, white tap-<br />

root; radials 13 to 28; flowers plain green<br />

—E. scheeri.<br />

16h. Radials 7 to 12; ribs 8; flowers yellow and open-<br />

ing widely —E. tobuschii.<br />

15b. Radials all but the lower one on each side of the<br />

areole white; upper centrals not forming a conspicuous,<br />

erect “V”; flowers rose, purplish, pink, white,<br />

or rarely pale yellowish —E. whipplei (in part).<br />

11b. All central spines straight—18.<br />

18a. Radial spines predominantly white—19.<br />

19a. Radials 25 to 36; plant small, 31/2 inches or less tall<br />

—E. mariposensis.<br />

19b. Radials 7 to 16; plants becoming larger than 31/2 inches<br />

tall—20.<br />

20a. Radials 1/2 to 1 inch long; centrals somewhat flattened<br />

and 1 to 2 inches long —E. whipplei (in part).<br />

20b. Radials 1/4 to 1/2 inch long; centrals round and 3/8<br />

to 11/8 inches long —E. conoideus.<br />

18h. Radial spines with various strong colors besides white<br />

—21.<br />

21a. Ribs 8 in mature plants —E. bicolor var. schottii.<br />

21b. Ribs 12 to 14—22.<br />

22a. Spines with bright red zones and some of them flattened<br />

and 1 inch or more long on mature plants;<br />

flowers large and very bright rose-pink<br />

—E. flavidispinus.<br />

22h. Spines gray or yellowish to dull purplish or reddish,<br />

but not bright red; all spines round and none over<br />

3/8 of an inch long; flowers small and pale pinkish<br />

—23.<br />

23a. Lowermost porrect central only 1/8 to 3/16 of an<br />

inch long; ribs 3/4 to 1 inch wide —E. intertextus<br />

var. intertextus.<br />

23b. Lowermost porrect central 1/4 to 5/8 of an inch<br />

long; ribs narrower, only 5/8 to 3/4 of an inch apart<br />

—E. intertextus var. dasyacanthus.<br />

1b. Plants spineless —E. asterias.<br />

Echinocactus horizonthalonius Lem.<br />

“Turk’s Head,” “Devil’s Head,” “Eagle Claws,” “Bisnagre,”<br />

“Bisnaga de Dulce,” “Bisnaga Meloncillo”<br />

Description Plates 18, 19<br />

stems: Each plant is at first a depressed hemisphere which<br />

later elongates to become a pyramid or a short cylinder in<br />

shape. It grows to a maximum size of about 8 inches tall and<br />

8 inches in diameter, and is almost always single, but rarely<br />

forming 2 or very rarely 3 stems. There are almost always 8<br />

ribs, but reports have been made of 5 to 13 ribs. These are<br />

very broad and rounded, with shallow grooves between them,<br />

and when old they are more or less interrupted by shallow<br />

cross-furrows. They may be vertical or spiraling. The color of<br />

the surface is a dull gray-green. Young plants have only a<br />

very little short wool at the apex, but old plants have a tuft<br />

of long wool filling the apex.<br />

areoles: Spherical or nearly so and 1/2 to 7/8 of an inch apart.<br />

Having much long wool when young, but nearly bare when<br />

old.<br />

spines: There are 5 to 8 radial spines. They radiate rather<br />

evenly around the areole, except that there is no lower radial<br />

present. In one form the radials are strongly recurved against<br />

the plant. Those of adjacent areoles interlock extensively.<br />

There is also one central spine on mature plants, which is<br />

more or less strongly deflexed and curved downward upon


genus Echinocactus 67<br />

the others. All spines are from 3/4 to 11/2 inches long, heavy<br />

to very heavy from enlarged bases, almost round to distinctly<br />

flattened. Their surfaces are rough and more or less cross-<br />

ridged. Their color is brownish or reddish fading to grayish<br />

or sometimes almost black.<br />

flowers: Very brilliant rose-red in color. They are 2 to 3<br />

inches broad and long. The ovary is covered with small scales<br />

and dense white or pink wool. The outer segments are short,<br />

narrow, and sharply pointed, sometimes ending in a blackish<br />

spine. The inner petals are longer, somewhat lance-shaped,<br />

with notched, toothed, or ragged edges, but usually ending in<br />

a spinelike point. There are very many short yellow stamens.<br />

The style is pink. There are 6 to 10 long stigma lobes which<br />

are reddish or pink on the lower side with the upper or inner<br />

sides salmon to somewhat orange or even olive.<br />

fruits: Oblong, to 11/4 inches long by about 1/2 inch thick,<br />

with some scales on the surface and enclosed in the long wool<br />

at the apex of the plant. This fruit dries from the tip downward<br />

so that when it is ripe the lower part is usually reddish<br />

and soft while the upper part is brown and dry. This upper,<br />

dry part usually breaks off, leaving the base and many seeds<br />

imbedded in the long wool. The seeds are about 1/8 of an inch<br />

long, somewhat irregular and angular in shape, with large,<br />

depressed hila. The surface is rough and dark brown.<br />

Range. All of trans-Pecos Texas, most of southern New Mexico<br />

and on into Arizona and Mexico.<br />

Remarks. With this species we venture into the fascinating<br />

group of the barrel cacti. While this is not one of the largest of<br />

the group, it is as tough as they come, its surface almost as hard<br />

as leather and its spines so strong, rigid, and spreading that one<br />

of its local common names, “eagle claws,” is apt.<br />

This little barrel grows on arid, rocky hilltops and slopes<br />

where there is no brush and very little grass to shade it. One<br />

may encounter it in such exposed places once he has passed<br />

westward over the Pecos River anywhere below Roswell, New<br />

Mexico. It is especially common in parts of the Davis, Guadalupe,<br />

Franklin, and other mountains of southern New Mexico<br />

and southwest Texas, where places may still be found with<br />

dozens of fine old specimens dotting the hillsides. However, it<br />

is much more usual now, over most of its range, to find an isolated<br />

plant and to search the surrounding area without finding<br />

a companion to this one somehow surviving individual. The most<br />

likely places to see this species in quantity today are in the bins<br />

of cactus dealers and the beds of nurserymen.<br />

It is hard for us now to imagine the wealth of such plants<br />

that our unspoiled deserts once boasted and the mayhem man<br />

has visited upon them in a relatively short time. Mr. Ernest<br />

Braunton, in an article published in the Cactus and Succulent<br />

Journal in 1933, tells of counting 500 cacti of this species uprooted<br />

and dumped into a ravine during a single clearing and<br />

building project that year in El Paso, Texas. He called then for<br />

some sort of conservation of such plants, but was not heard, and<br />

now one has to go far into the out-of-the-way places of the<br />

mountains to find any of these cacti at all. Such is the situation<br />

almost everywhere, and the individuals we do find seem to be<br />

only the lucky ones which have so far been missed by the diggers.<br />

Since this cactus does not hide under any brush or survive<br />

well in thick grass, it is especially vulnerable and the motives<br />

for taking it have been several.<br />

As one of its Spanish common names implies, in the past its<br />

flesh has been used for the making of cactus candy. It is probably<br />

used little if at all for this in the U. S. today, but now it is<br />

taken for use in the so-called desert plantings fashionable for<br />

yards and gardens. But in this it usually suffers almost as certain<br />

a death as in the candy manufacture. It is so strictly a<br />

desert plant that it does not thrive well even in the yards of<br />

New Mexico or west Texas; only those willing to give it the<br />

most specialized situation should try to grow it. In the meantime,<br />

it becomes more and more scarce as we are fascinated by<br />

the tough old fellows and bring them from their deserts.<br />

This species apparently received two names in the same year.<br />

In 1839 it was named E. horizonthalonius by Lemaire and E.<br />

equitans by Scheidweiler; however, the first of these two names<br />

has long been the one used for it. Engelmann attempted to set<br />

those specimens with central spines apart as variety centrispinus,<br />

but this division was long ago dropped as unnecessary since it<br />

would comprise almost all mature specimens seen and leave out<br />

all juvenile ones. However, there do seem to be two recogniz-<br />

able varieties which are consistent.<br />

Echinocactus horizonthalonius var. curvispina SD<br />

Description Plate 18<br />

stems: As the species, except that old specimens remain short<br />

and pyramid-shaped instead of cylindrical and have ribs<br />

which are very flat but quite noticeably interrupted by cross-<br />

furrows and, therefore, tuberculate.<br />

areoles: As the species.<br />

spines: As the species, except that they are always very heavy,<br />

very flat, and very severely recurved against the plant, with<br />

no spines projecting outward.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. Practically limited to west Texas. From the lower Pecos<br />

River to the Davis and Guadalupe mountains.<br />

Remarks. The Texas members of this species are mostly of this<br />

form. I have examined hundreds of Davis Mountain specimens<br />

without seeing a single plant diverging from this description. In<br />

southeastern New Mexico there are less definite examples which


68 cacti of the southwest<br />

appear sometimes as intermediates. Engelmann stated, in “Corrections<br />

to the Cactaceae of the Boundary,” that these two<br />

forms had “entire identity” and could scarcely be called varieties,<br />

but that the intermediates seem to require them. Yet when<br />

one moves so short a distance from the Franklin Mountains as<br />

the Selden and Portrillo mountains he finds all of the specimens<br />

there clearly of the next variety.<br />

Variety curvispina is, therefore, the eastern form of the species.<br />

The extremely heavy, recurved spines on the squat, pyramidshaped<br />

barrel gives us one of the most beautiful forms found in<br />

the cacti. It is a plant remarkable for the unvarying tidiness and<br />

symmetry of its form and it gives a fascinating expression of<br />

unyielding toughness. Still, its spines are so appressed against its<br />

surface that one may hold an old giant of this variety in the<br />

flat of the hand without getting a prick from it. I have seen<br />

workmen unloading a truckload of these plants by casually<br />

tossing them down from one to another like balls-all bare-<br />

handed. This is something one had better not try with the more<br />

western variety of the species.<br />

Echinocactus horizonthalonius var. moelleri Haage Jr.<br />

Description Plate 19<br />

stems: As the species, except that with age it becomes more<br />

columnar than pyramid-shaped and taller but not so large in<br />

diameter. The ribs are high but with very shallow cross-<br />

grooves; the tubercles are more fused and often are hardly<br />

visible at all.<br />

areoles: As the species.<br />

spines: As the species, except that the spines are not so heavy,<br />

being only somewhat flattened, and straighter; not recurving<br />

against the plant but rather standing out at angles from its<br />

surface.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. From the Franklin and Guadalupe mountains west into<br />

Arizona.<br />

Remarks. This is the western form of the species. By comparison<br />

to the other variety it is smaller in diameter but taller and more<br />

columnar, with sharper, higher, less interrupted ribs. But the<br />

most noticeable difference is in the spines, which in this variety<br />

show none of the tidiness of the others, but stand out at all<br />

angles in chaotic, interlacing masses. Where the one may be<br />

handled easily, woe betide anyone who thinks to pick up this<br />

variety by other than the roots, as there are spines aimed in<br />

every direction and if he seeks to withdraw from a point in one<br />

direction he will usually back into an opposing point nearby or<br />

even be caught between the spines, which often have a pincers<br />

action. These spines are also much less heavy or flattened than<br />

those of the other variety.<br />

Echinocactus texensis Hopff.<br />

[Homalocephala texensis (Hopff.) B. & R.]<br />

“Devil’s Head,” “Horse Crippler,” “Candy Cactus,” “Manco<br />

Caballo,” “Viznaga”<br />

Description Plate 19<br />

stems: Very broad. Greatly flattened to sometimes dome-<br />

shaped. This cactus grows to a maximum of about 12 inches<br />

across, and such large plants rise from only 2 to occasionally<br />

8 inches high. Stems are usually single, but the plants occasionally<br />

produce 2 or 3 equal stems. When injured at the summit,<br />

they often produce a cluster of small heads on top of the<br />

old one. The surface is dark green. The ribs are prominent and<br />

acute, normally 13 or 14, 20 or 21, or 27 in number. The apex<br />

of the stem is filled with some long, white wool.<br />

areoles: Triangular to inverted heart-shaped, 1/4 to 3/8 of an<br />

inch in greatest diameter, and covered with white or gray<br />

wool. They are located about 1 to 1/4 inches apart.<br />

spines: Reddish or brownish-gray, becoming whitish when<br />

old. Only a little flattened, they are ringed by regularly occurring<br />

ridges, and are very heavy and rigid. There are 6 or 7<br />

radial spines from 3/8 to 2 inches long. There are typically 2<br />

diverging upper radials which are comparatively small and<br />

short, 2 lateral radials which are very heavy and long—often<br />

the longest spines—and a pair of lower diverging radials<br />

which are again smaller. Occasionally there may be one additional<br />

radial which is also comparatively small and directed<br />

straight upward. All the radials may be straight and spreading<br />

to strongly recurved. There is also one central spine which<br />

is the stoutest spine, often 1/8 of an inch thick at the base and<br />

sometimes much more than that. It is from 3/4 to 23/4 inches<br />

long, deflexed, and from straight to recurved or sometimes<br />

slightly hooked.<br />

flowers: These are bell-shaped, 1 to 21/4 inches in diameter<br />

and similar in length, slightly fragrant and very beautiful,<br />

each flower displaying an interesting range of shades. The<br />

ovary is densely covered with long white wool and many<br />

short, sharp-pointed but soft, blackish scales. These scales<br />

lengthen as they progress upward, while the wool thins. The<br />

outer perianth segments are short, narrow, and sharp-pointed.<br />

Their midribs are fleshy and greenish or brownish, ending in<br />

a brownish point, while their edges are greenish to whitish,<br />

fringed, and more or less covered with a web of wool. The<br />

inner perianth segments are narrowly lanceolate from narrow<br />

bases. These bases are red. A pale rose midline extends up the<br />

petal, darkening noticeably as it nears the apex, where it ends<br />

in a pronounced mucro which is purplish or brownish. All of<br />

the expanded part of the petal is pale lavender, salmon, pink<br />

or sometimes almost white, depending upon the individual<br />

plant. The edges of the petals are fringed to the tips, feathery<br />

in appearance. The filaments are reddish to pinkish, the anthers<br />

pale yellow. The style is yellowish or pinkish. The stigma


genus Echinocactus 69<br />

has 10 to 17 rather long yellowish or pinkish lobes, each often<br />

having a red stripe on the lower side.<br />

fruits: Spherical to oval, 3/4 to about 11/2 inches long. They<br />

remain fleshy and become brilliant red. After a very long<br />

time on the plant they usually split open vertically on one<br />

side. Scattered over the surface are the dried and hardened,<br />

bristle-like ovary scales, each with a tuft of white wool in its<br />

axil. On the top of the fruit clings the dried remains of the<br />

perianth. The seeds are black, kidney-shaped, slightly less<br />

than 1/8 of an inch in length.<br />

Range. All of Texas west of a line from approximately the<br />

mouth of the Colorado River to near Fort Worth and ‘Wichita<br />

Falls, except the Texas Panhandle and the extreme western tip<br />

of the state beyond the Guadalupe Mountains. Also in extreme<br />

southwest Oklahoma and southeastern New Mexico as far north-<br />

west as Roswell. Extending deep into adjacent Mexico.<br />

Remarks. It is fitting for this plant to carry the name texensis,<br />

for it has one of the widest ranges of any cactus in that state,<br />

and it does not venture far into any other state, barely entering<br />

Oklahoma and only penetrating a corner of New Mexico. It is<br />

a low, flat, retiring cactus content to stay under the grass, so<br />

although it is fairly common over a wide range, many people<br />

do not know it. Yet when one has once discovered it, he is not<br />

likely to forget the plant.<br />

The impression this cactus gives is one of elemental, even<br />

brutal strength. It squats low to the ground, usually only 2 to 5<br />

inches or so high, its surface is hard and unyielding, and it is<br />

covered with a loose system of not too many but some of the<br />

most robust and rigid spines found on any of our cacti. There<br />

it sits, and it seems to dare anyone to come its way. No wonder<br />

the ranchers of two nations call it in two languages by the name<br />

“Horse Crippler,” for it is said that it sits there unseen in the<br />

grass, and if a running horse steps on it, the rigid spines will<br />

penetrate the tender underside of the hoof and cripple the horse.<br />

Its very strength brings its downfall, since most ranchers regularly<br />

uproot any of these pests they see on their ranges, and<br />

this has greatly reduced the numbers of these cacti found in<br />

many areas. However, the species is still very common in certain<br />

fields from within sight of the Gulf around Corpus Christi<br />

northwest through central Texas and in extreme southeast New<br />

Mexico. To either side of this broad southeast to northwest band<br />

one finds it less frequently, but it ranges over a wide area.<br />

This species is remarkable because, even though it is one of<br />

the toughest of the barrels in structure, it can stand lots more<br />

moisture than most others of this group. For this reason it is<br />

much better adapted to cultivation than most of its relatives<br />

and can also stand more cold than most of them. It is a favorite<br />

among collectors and growers.<br />

The species was first described and named Echinocactus<br />

texensis in 1842. Three years later Engelmann described it<br />

again and designated it Echinocactus lindheimeri. There were<br />

few other names coined for it, but E. platycephalus by Muehlen-<br />

pfordt and Melocactus laciniatus by Berlandier are two syn-<br />

onyms.<br />

There have been two varieties of this cactus proposed, variety<br />

gourgensii Cels and variety longispinus Schelle. Since it grows<br />

over so wide a range it is surprising there are not more localized<br />

forms deserving varietal rank, but the species is remarkably<br />

stable. There is a general gradient of spine size on these plants<br />

as one moves from southeast to northwest over its range. Those<br />

specimens from southeast Texas have more slender and usually<br />

somewhat shorter spines than those from farther northwest.<br />

This is not surprising, and the first of the above proposed<br />

varieties seems to be no more than a segment of this gradient<br />

which cannot really be set apart.<br />

However, there is one local population which seems to be<br />

rather distinct from the rest, and which may deserve varietal<br />

rank. In the Big Bend of Texas, in Brewster County, one finds a<br />

population of this species which is identical to the rest except<br />

for the following points: its surface is more gray-green than<br />

typical, and its central spines run from 2 to 23/4 inches long on<br />

mature individuals. This plant’s physiology is also so different<br />

from that of the typical specimens from the rest of Texas that<br />

while the typical specimens have been growing, blooming, and<br />

fruiting each year in my beds, my examples of this form have<br />

sat right beside them for five years with a minimum of growth<br />

and never a bloom. This may be variety longispinus Schelle,<br />

but since I have never succeeded in seeing its flowers and fruits<br />

and cannot give more details about it, it is not so listed here.<br />

After what looked like a history free of any arguments over<br />

its taxonomic place, this cactus caught the eye of Britton and<br />

Rose and they took it out of the genus Echinocactus and erected<br />

a new genus, Homalocephala, for it. This has remained a monotypic<br />

genus.<br />

There seems to have been little if any discussion of the validity<br />

of this new genus. It was accepted as a logical result when the<br />

original genus Echinocactus was broken up, and so it may be if<br />

one is to follow in that step to no one knows yet what fragmentation.<br />

However, I have noted in the discussion of the genus<br />

Echinocactus that some of the most significant recent research<br />

on cacti has involved this species. Dr. Boke’s summary of his<br />

anatomical research results are so important here that we should<br />

refer to them again. He says, “The combined list of characters<br />

suggests to the author that if E. horizonthalonius is to be retained<br />

in the genus Echinocactus, Homalocephala should be returned<br />

to the same genus. It further suggests that E. horizontha-<br />

lonius could, with equal logic, be referred to the genus Homalocephala.”<br />

Faced with this alternative, I choose to return Homalocephala<br />

texensis to the genus where it was originally thought to belong,<br />

rather than to further fragment the original genus and eliminate<br />

it from our area altogether by making it a Homalocephala<br />

horizonthalonius.<br />

I find a great similarity between Echinocactus texensis and<br />

a plant brought out of Mexico by a dealer. I saw several dozen


70 cacti of the southwest<br />

specimens of this Mexican plant, some of them up to about 16<br />

inches in diameter but still only about 8 inches tall. The ribs,<br />

areoles, and spines of this plant are almost identical to those of<br />

our plant, except that the spines are longer and more yellow in<br />

color, and the centrals stand more upward. The plant was being<br />

sold as Echinocactus victoriensis, but I have been able to learn<br />

little more about it. It does not seem to be Ferocactus rafaelensis<br />

(Purpus) B. & R. This plant appears to be the most closely re-<br />

lated to our Texas cactus of anything I have yet seen.<br />

Echinocactus asterias Zucc.<br />

[Astrophytum asterias (Zucc.) Lem.]<br />

“Sea-Urchin Cactus”<br />

Description Plate 19<br />

stems: Extremely flat, depressed disc-shaped to sometimes<br />

low dome-shaped. Mature plants are from 2 to occasionally<br />

6 inches in diameter by less than I to at most 21/2 inches tall.<br />

This plant body, which is always simple, is divided by very<br />

narrow but distinct vertical grooves, into 8 broad, almost flat<br />

ribs. These ribs actually form triangular sections of the stem.<br />

Up the center of each section runs a line of areoles which are<br />

on no projections and separated by no cross-grooves of any<br />

kind. The surface of the plant is a dull green, and scattered<br />

over it are tiny, less than pin-head-sized clusters of very short,<br />

whitish wool.<br />

areoles: Circular, a little less than 1/8 to 1/4 of an inch across,<br />

and filled with dense wool, at first straw-colored, then gray.<br />

They are located about 1/4 to 3/8 of an inch apart.<br />

spines: None.<br />

flowers: Yellow with orange throats. 2 to 31/2 inches across<br />

and about 2 inches tall. Opening widely. The ovary is densely<br />

covered with scales which have blackish, bristle-like points at<br />

their summits and much cobwebby wool in their axils. The<br />

outer perianth segments are short, narrow and pointed,<br />

greenish in color, and covered on their outer surfaces with<br />

short fuzz. The inner segments are long, slightly spatulate,<br />

from narrow orange bases. The upper parts are clear yellow,<br />

the edges entire and the tips from entire and slightly pointed<br />

to somewhat erose and irregular. The filaments are orange at<br />

their bases and yellow above. The anthers are yellow. The<br />

style is yellowish and the stigma has 10 to 12 yellow lobes.<br />

fruits: Oval, about 1/2 inch long. Densely covered with spines<br />

and wool. Becoming dry while on the plant and finally<br />

breaking off at or near the base. The seeds are black or<br />

nearly so, shiny, about 1/16 of an inch long.<br />

Range. Entering Texas from Mexico only in the lower Rio<br />

Grande Valley, where it is found in a few locations in Starr and<br />

Hidalgo counties.<br />

Remarks. Strange as it may seem, this is a diminutive barrel<br />

cactus. It is unmistakably the dwarfed relative of a group of<br />

large, columnar barrels found in Mexico and it is still very<br />

closely related to the biggest barrels of all. But this one has<br />

apparently survived by being small, inconspicuous, and retiring.<br />

It even dispenses with the typical heavy covering of spines and<br />

shows that survival is possible without these. This is said to be<br />

accomplished by having the whole body suffused with some<br />

chemical compounds distasteful to all enemies so that it can<br />

remain unarmored among them. It also has scattered over its<br />

surface to a greater or lesser degree on different specimens a<br />

series of clusters of short white wool, said to substitute to some<br />

extent for the lack of the shade which other cacti get from their<br />

spines.<br />

The form of this cactus is unique and remarkably beautiful.<br />

It projects a very short distance above the ground even when<br />

plump and water-filled. When water is deficient it shrinks to a<br />

mere flat disc which hardly projects above the ground at all<br />

and may be almost covered by the sand. Its common name<br />

comes from the fact that its broad, nearly flat ribs and its shape<br />

make it look almost exactly like the skeleton of a sea urchin<br />

denuded of its spines.<br />

The cactus has had at best a precarious foothold on the north<br />

side of the Rio Grande. It has been found on the dry hills overlooking<br />

the river and never more than about 15 or 20 miles<br />

north of that river. And we have in recent times not been kind<br />

to this immigrant. The cactus has been a favorite among collectors,<br />

partly because it is a beautiful curiosity without spines.<br />

Dealers have for many years scoured the counties where it<br />

grows for the plant, and have uprooted it by the thousands for<br />

the trade. At the present time it takes a very good guide to show<br />

one the few locations in Texas where it still grows in numbers,<br />

and the chances are rather good that when one gets there a<br />

cactus digger will already have eliminated the population. With<br />

the widespread clearing of the area which is now going on, this<br />

cactus may well be eliminated in Texas, although it is abundant<br />

in Mexico.<br />

The species was long ago called Echinocactus asterias by<br />

Zuccarini. Over a hundred years ago Lemaire separated out<br />

some cacti formerly in that genus, including this one, into a new<br />

genus, Astrophytum. Coulter later thought this unwarranted<br />

and returned them to Echinocactus. K. Schumann agreed with<br />

him, but listed Astrophytum as a subgenus of the genus Echinocactus.<br />

Most authors since then have treated Astrophytum as a separate<br />

genus, although very few of the books on U.S. cacti have<br />

included it at all, since only this one species is found in the<br />

U. S. and that only in a very small region of south Texas.<br />

Buxbaum, the great theorizer about cacti, decided that the<br />

Astrophytums do not have any close relationship to the Echinocacti<br />

at all, but instead he considers them the most northerly of<br />

the South American cactus group. He considers the closest relatives<br />

to the sea-urchin cactus to be the members of the South


genus Echinocactus 71<br />

American genus Frailea. Others, including Backeberg, have<br />

opposed this idea most strenuously, and their arguments show<br />

good reasons why it is after all an Echinocactus. It does seem<br />

that if this, with its scaly, woolly ovary, is not a member of the<br />

Echinocacti then the distinguishing characters of that group are<br />

of little validity and the group can hardly exist on any level at<br />

all.<br />

Echinocactus wislizeni Eng.<br />

[Ferocactus wislizeni (Eng.) B. & R.]<br />

“Barrel Cactus,” “Fishhook Barrel,” “Candy Barrel,”<br />

“Visnaga,” “Biznaga,” “Biznaga de Agua”<br />

Description Plate 20<br />

stems: Spherical at first, then ovate or conical, and finally<br />

cylindrical. This plant becomes very massive, up to at least<br />

4 feet tall, and is said to have reached 6 feet tall. It grows<br />

about 2 feet in diameter. There are 13 to 25 ribs which are<br />

sharp, up to about 11/4 inches high, and a little undulate<br />

because of the slight bulging at the areoles. The color of the<br />

surface is dark green.<br />

areoles: 5/8 to 1 inch long and 3/4 to 1/4 inches apart,<br />

elliptic to linear. Those areoles which have not produced<br />

flowers have the upper end prolonged as a narrow groove;<br />

but after blooming this upper part of the areole is permanently<br />

broadened and connected to the lower, spinous part<br />

by a more narrow neck. It has a short brownish wool which<br />

fades to gray and then is mostly lost on old areoles. Some<br />

glands are present.<br />

spines: There are 12 to more than 20 slender radial spines.<br />

The upper and lateral ones are flexible, bristle-like, whitish,<br />

and about 1 to 2 inches long. The lower 3 are a little shorter<br />

and rigid, approaching the character of the centrals. There<br />

are 4 very strong centrals 11/2 to 3 inches long, yellowish to<br />

red or purplish-red, and all ringed by conspicuous annular<br />

ridges. The upper three are straight, spreading upward, and<br />

are round to somewhat flattened. The lower one stands porrect<br />

or slightly deflexed, is usually hooked downward at the end<br />

(but on rare specimens is nearly straight), is much heavier<br />

than the other centrals, and is somewhat to greatly flattened.<br />

flowers: Variable shades of yellow, gold, orange, or red,<br />

about 2 inches long by 2 to 3 inches across. The ovary is<br />

covered with scales which are green, edged in white. The<br />

outer segments of the perianth are short, triangular-shaped to<br />

ovoid, with pointed apexes, the midlines greenish to reddish<br />

or yellowish, and the edges entire and lighter colored. The<br />

inner segments are linear and sharply pointed, the edges<br />

slightly irregular or erose. They are most commonly orange-<br />

red on the midlines shading to conch-shell pink on the edges,<br />

but they may be all yellow or all red. The extremely numer-<br />

ous filaments are yellow or red, the very small anthers<br />

yellow. There are 18 to at least 26 long, pointed, erect stigma<br />

lobes which are in yellow flowers yellow, in red flowers<br />

reddish below and yellowish above.<br />

fruits: Oblong, 1/4 to 21/4 inches long, practically covered<br />

with white-edged scales. They are yellowish and fleshy at<br />

first, but remain long on the plants, becoming finally dried<br />

and hard. The seeds are nearly 1/8 of an inch long, black,<br />

with the surface rough but not tuberculate.<br />

Range. From Arizona east through the mountains of southern<br />

Hidalgo and Luna counties, New Mexico, to the Organ and<br />

Franklin mountains of Dona Ana County, and extending into<br />

Texas only in the Franklin Mountains near El Paso. Also in<br />

adjacent Mexico.<br />

Remarks. This is the largest cactus in our area, and the one<br />

which is truly barrel-like in its dimensions. An old specimen of<br />

this cactus is awe-inspiring for its sheer bulk and the impression<br />

of age and strength it presents. It is the tough old patriarch of<br />

the exposed mountain slopes in our semidesert regions. Such a<br />

mountainside still dotted with its population of old barrels is<br />

an unforgettable sight. The effect is much like that of viewing<br />

a herd of buffalo or a flock of whooping cranes, and the chance<br />

of experiencing it today is almost as unlikely. Each of these is<br />

a giant among its kind, formidable to confront—yet today each<br />

is, as a species, only a pitiful remnant of what it once was. One<br />

is lucky now to find an individual candy barrel, and there are<br />

very few places left in our area where one can see a population<br />

of old specimens still undisturbed in their natural glory.<br />

There are several reasons why such great, formidable species<br />

have fared so badly in the last hundred years, and understanding<br />

them can teach us much about this cactus and about ourselves.<br />

Some of these reasons arise out of the fact that, taken<br />

individually, these organisms are not as invulnerable as they<br />

appear. They are all approaching extremes in development and<br />

adaptation to very special environments, and while this gives<br />

them great survival powers in these special environments, it<br />

makes them conspicuous targets for new enemies which enter<br />

the community and it renders them ungainly misfits which<br />

cannot maintain themselves when the environment is changed<br />

or they are pulled out and put somewhere else.<br />

Modern man is the new enemy which has recently entered<br />

the environment of each of these, and they have fallen in terrible<br />

numbers before his attack. But why did he come before them as<br />

such a ruthless destroyer? Precisely because of their size and<br />

majesty.<br />

Each of them was so big that an individual presented a mass<br />

of flesh usable for food and other purposes. As they slaughtered<br />

the buffalo for meat and hides, they slaughtered the barrel<br />

cactus to use its watery pulp. The few that were sacrificed to<br />

the thirst of prospectors caused no problem. The real inroads<br />

upon this plant came with its use for making candy. This became<br />

so widespread that it became known by the name, candy barrel.


72 cacti of the southwest<br />

Man’s slaughter of these organisms could be more easily excused<br />

if it had been limited by even such uses of them, but it<br />

has not been. The very appearance of them apparently invokes<br />

a reaction in modern man, and he has turned his weapons upon<br />

them in a frenzy of destruction. It is as though before their<br />

formidable majesty he developed an inferiority complex which<br />

issued in an urge to kill. The extent of this factor in the slaughter<br />

of the buffalo is easily seen, but few realize that it has<br />

played a big part in the disappearance of the barrel cactus.<br />

One of the few places where I have seen a stand of old barrels<br />

was in a small canyon so deep in the Franklin Mountains that<br />

a mountain lion shadowed us as we proceeded up it. Here I<br />

found myself on a steep slope with several dozen fine old cacti<br />

up to four feet tall in view at the same time. But I had arrived<br />

there a week or so too late. As I moved from one to another<br />

of these old monarchs, each perhaps half a century old, I found<br />

that each had been neatly decapitated or else the apex split<br />

open by one deft stroke of a machete or some such instrument.<br />

The exposed flesh of each one was melting away in the heat.<br />

Since then that canyon is a sad place to anyone knowing what<br />

was needlessly destroyed.<br />

I did not understand the reason for such a destruction until<br />

some time later when I stood and observed people filing through<br />

the cactus house of the New York Botanical Garden. Here I<br />

saw a man push against the fence which protected the plants<br />

and, gesturing excitedly toward a beautiful old barrel cactus,<br />

roar for the benefit of his young son, “Look at that big, ugly,<br />

——— ——— thing!! God, if I could only get at it, you’d see<br />

how I’d fix it! I’d tear it apart!!” Only then did I realize how<br />

such great organisms have been sacrificed to the ego of modern<br />

man.<br />

More indirectly, but quite as effectively they have been sacrificed<br />

to man’s meddling with their environment. When he<br />

moves in and tampers with it, they, being so specialized, are<br />

usually the first to go. And, finally, the unwise treatment of<br />

those who do appreciate the beauty of these cacti takes its toll.<br />

Fine old giants are pulled up and planted in lush yards where<br />

they cannot live. The extent of this activity is so great that I<br />

have seen them arrive by the truckload at nurseries in climates<br />

where they cannot possibly be kept alive out-of-doors. Against<br />

all of this destruction the buffalo and the whooping crane are<br />

now protected, but not the candy barrel within our area.<br />

There have been a few problems about this species’ taxonomy.<br />

Engelmann first described and named it. He also described and<br />

named an Echinocactus emoryi, which has been the subject of<br />

much confusion since some have thought it was a synonym of<br />

this and some have thought not, and he also had an E. wislizeni<br />

var. decipiens, but neither of these forms appears in the<br />

area covered by this study and so we do not have to evaluate<br />

them here. Apparent later synonyms for E. wisli zeni were<br />

E. falconeri Orcutt and E. arizonicus Kunze.<br />

Everything was rather stable, taxonomically, concerning this<br />

plant until Britton and Rose. These authors, however, in<br />

breaking up the old genus Echinocactus erected a new genus<br />

Ferocactus and placed in it this plant, along with various<br />

others. It has generally been known since their time as Fero-<br />

cactus wislizeni.<br />

But Britton and Rose’s genus Ferocactus has not fared very<br />

well. A number of species which they included in it have since<br />

been removed by other students to other genera, among them<br />

their Ferocactus johnsonii to Echinomastus, F. hamatacanthus<br />

to Hamatocactus and F. crassihamatus and F. uncinatus to<br />

Glandulicactus. It is obvious that Britton and Rose’s concept of<br />

the genus Ferocactus was not satisfactory and that the genus<br />

has come to be, at best, another microgenus. The only real character<br />

which can be cited to set wislizeni off from the Echinocacti<br />

is the absence of wool on the ovary and fruit. If this is considered<br />

a character less essential than it takes to support a genus<br />

all by itself, as Link and Otto obviously regarded it, then we<br />

would have little if anything else definite to uphold the genus<br />

Ferocactus. Because of these considerations I am using the older<br />

genus name for this plant.<br />

Echinocactus uncinatus var. wrightii Eng.<br />

[Glandulicactus uncinatus var. wrightii (Eng.) Backbg.]<br />

“Turk’s Head,” “Cat-Claw Cactus,” “Brown-Flowered<br />

Hedgehog,” “Texas Hedgehog”<br />

Description Plate 20<br />

stems: Oval, up to 8 inches tall and 41/2 inches thick, but<br />

usually much smaller. These stems are almost always single,<br />

but occasionally they may produce 1 or 2 branches at or near<br />

the base. The surface is bluish-green with a gray glaucescence<br />

and is formed into 9 to 13 ribs. These ribs are fairly high<br />

and conspicuous, separated by broad grooves, and consist of<br />

rather distinct but partly fused tubercles. The tubercles themselves<br />

are rather remarkable in shape. The areole is on the<br />

upper slope of the tubercle, which is prolonged below or<br />

ventral to the areole into a chinlike swelling which more or<br />

less overhangs the upper end of the areole on the next tubercle.<br />

The abrupt drop-off from this prominent chin before<br />

the next tubercle produces a sharp cross-indentation of the<br />

rib when it is old. On the new tubercles the chin and this<br />

cross-indentation are not yet so definite.<br />

areoles: Elongated oval, to about 5/8 of an inch long, the<br />

upper end prolonged into a narrow extension which has been<br />

called a groove. The spines all grow from the broader lower<br />

portion of the areole, while the flower is produced from the<br />

narrower upper end of it. In the still more narrow neck of<br />

the areole joining these two regions there are produced several<br />

yellowish glands. The whole areole contains gray or slightly<br />

yellowish wool.<br />

spines: All spines are very heavy and are at first red, later<br />

crassahamatus<br />

-> crassihamatus


genus Echinocactus 73<br />

reddish-brown, straw-colored, or grayish, with the points<br />

remaining darker. There are 7 or 8 radial spines which radiate<br />

rather evenly. The upper and lateral 4 or 5 are straight,<br />

flattened, and from 3/4 to 2 inches long. The lower 3 radials<br />

are 3/4 to 11/4 inches long, round, or nearly so, more or less<br />

curved, and hooked at their tips. There is one central spine<br />

from 2 to at least 4 inches long, which stands porrect or<br />

turned upward, and is very heavy, angled, usually twisted,<br />

and hooked at the tip.<br />

flowers: Maroon to garnet in color. They are funnel-shaped,<br />

not opening widely, 3/4 to 11/4 inches long, and about 1 inch<br />

across. The narrow ovary is covered with short, broad scales<br />

with brown centers and broad white, membranous edges. The<br />

outer perianth segments are triangular to long triangular with<br />

bluntly pointed tips and white, entire, but often crinkled,<br />

edges. The maroon to garnet inner segments are linear, their<br />

edges sometimes lighter colored and often irregular or toothed<br />

above, with the apex slightly pointed, irregular, or sometimes<br />

even squared off. The filaments are brown or maroon, the<br />

anthers cream-colored. The style is brown, the stigma having<br />

10 to 14 broad, fat lobes which are cream-colored above and<br />

maroon below.<br />

fruits: Oblong to egg-shaped, 5/8 to 1 inch long. They are<br />

for a brief time pale reddish between the whitish scales which<br />

very nearly cover them. They soon become dry and colorless,<br />

usually remaining on the plant for a long time in this condition.<br />

The seeds are about 1/16 of an inch long, curved, and<br />

compressed, with the hila basal. The surface of the seeds is<br />

finely tuberculate.<br />

Range. Southern New Mexico, south Texas, and adjacent<br />

Mexico.<br />

Remarks. Although commonly designated as a hedgehog cactus<br />

rather than a barrel, this cactus is a smaller one of the latter<br />

group with spines to rival any.<br />

The range given above for this form is less specific than for<br />

most. This is because, while the cactus has been found over a<br />

huge area, it is apparently common today hardly anywhere,<br />

and we suspect that it has been eliminated from some of its<br />

former range. It is still to be found in limited numbers on dry<br />

hills from El Paso to Van Horn and on down the Rio Grande<br />

past Presidio, Texas. This is a far cry from Coulter’s statement<br />

of 1896 that it was then “abundant from El Paso, Texas, to the<br />

Pecos,” but it does seem to be the center of this cactus’ range<br />

and the only area where one can count on finding it today.<br />

From here there are isolated records of the plant for great<br />

distances in all directions, even though none of them seem to<br />

represent established populations at these faraway points.<br />

Coulter stated its range as, “Extending almost to the mouth of<br />

the Rio Grande.” After several years of extensive observation<br />

almost all along that river, the idea of this cactus existing any<br />

more below the Big Bend had about become unbelievable to<br />

me, when suddenly I found one specimen of the cactus growing<br />

happily near Falcon Dam in Starr County, far down toward<br />

the mouth of the river from any place I had seen it before. No<br />

more could be found in the area, and we have still found no<br />

record of the plant in the more than 300 miles between this<br />

collection and the Trans-Pecos records. However, this does<br />

cor roborate to a great extent Nealley’s old record of the plant<br />

having been taken near Rio Grande City.<br />

In all other directions we find similar very widely scattered<br />

records, some of them not duplicated in many years. Northeast<br />

of the central range there is Lloyd’s record of the plant just east<br />

of Fort Stockton, Texas, which I have not been able to duplicate.<br />

Going north and west of El Paso there are few and very<br />

widely scattered records of the cactus in New Mexico. Very far<br />

to the northwest. Wooton and Standley list it as found at Pena<br />

Blanca, New Mexico, which is only some 25 miles or so south-<br />

west of Santa Fe. While it has been found west of El Paso in<br />

southern New Mexico, it apparently does not reach Arizona.<br />

We are faced with the fact that there must once have been a<br />

very huge area over which this cactus ranged, but that today,<br />

everywhere but in the central range as outlined above, it is so<br />

rare that if one locates a single specimen he should consider<br />

himself very lucky.<br />

Our U. S. form is not the typical form of the species, Echinocactus<br />

uncinatus Galeotti. That form has 3 or 4 central spines<br />

and a difference in the seed form, and seems restricted to<br />

Mexico. Engelmann realized the difference in our form and<br />

called our variety wrightii. There has been no disagreement<br />

about this since, except that some authors have not bothered<br />

to tack on the varietal name, merely using the species name for<br />

our plant. No one has produced any evidence that I have seen<br />

of the typical form being found in the U. S. One man, the late<br />

Mr. Fred Leasure, an El Paso teacher who for most of a lifetime<br />

collected, studied, and dealt in cacti from that area, once told<br />

me that he had found a population of the typical form with<br />

several centrals, somewhere in the mountains near El Paso. I<br />

was never able to learn any details about this.<br />

Most of the disagreement over this plant has been about what<br />

genus to put it in, and the history of these arguments is a rather<br />

disillusioning one.<br />

Galeotti, in the beginning, placed the species in the old genus<br />

Echinocactus. There was no question at all through all of the<br />

earlier treatments of the cactus until Britton and Rose broke<br />

up the large genus. When they did, they put this cactus in their<br />

new genus Ferocactus. It does have some close similarities to the<br />

other species placed in that genus, among them being a very<br />

scaly ovary which is bare of wool. But differences can be cited<br />

too. Once these small similarities and differences began to be<br />

considered significant on the generic level, the gate was down,<br />

and the confusion which has resulted has not enhanced the reputation<br />

of cactus scholarship.<br />

Knuth, in 1935, was just as certain as Britton and Rose had<br />

been about Ferocactus that the cactus belonged in the equally<br />

new genus Echinomastus. He had his reasons also. But by 1941


74 cacti of the southwest<br />

Marshall published the plant as Thelocactus uncinatus, with his<br />

set of arguments backing this name. By this time Backeberg had<br />

also published his solution of the problem, which was to erect<br />

an entirely new genus for the species. He justified this to a large<br />

extent by the presence of glands in the areoles of the species,<br />

and so his new name was Glandulicactus uncinatus.<br />

In 1951 Buxbaum published yet another disposition of the<br />

species. He was very sure that it should be in the genus Hamatocactus.<br />

His reasons were good enough to persuade Marshall, and<br />

in 1957 he published his new opinion that the species should<br />

really be called Hamatocactus uncinatus. However, Backeberg<br />

remained unconvinced and in his still more recent large work<br />

maintains the genus Glandulicactus and adds some other species<br />

to it.<br />

One seems to be able to take his own choice of all these genera<br />

for this species, with perhaps Hamatocactus and Glandulicactus<br />

having a slight edge in taxonomic opinion at the present time.<br />

It would take a long digression to outline the points for and<br />

against each genus. They are all very detailed, slight differences<br />

of morphology which tend to balance each other off too well,<br />

leaving the decision between them at best somewhat arbitrary.<br />

I myself feel that these are all no more than microgenera and<br />

that the argument may well be left with the theorists while we,<br />

by harking back to the old genus Echinocactus of Link and<br />

Otto, can find a name so usable and stable that it will not be<br />

different in every book picked up.<br />

This is a rare species, a tough, desert-loving one, and therefore<br />

one which is not very easy to cultivate. It rots very quickly<br />

from too much moisture, but if this problem is taken care of, it<br />

can grow slowly and produce a number of small but interesting<br />

flowers which are very unusual for their brownish hues.<br />

Echinocactus whipplei Eng. & Big.<br />

[Sclerocactus whipplei (Eng. & Big.) B. & R.]<br />

“Devil’s Claw Barrel”<br />

Description Plate 20<br />

stems: From practically spherical to an ovate or even short-<br />

cylindrical shape. These stems are usually single and up to 6<br />

inches tall by 4 inches in diameter, but occasionally forming<br />

small clusters of 2 or 3 stems, and said to have reached 12<br />

inches tall and 6 inches in diameter. They have 13 to 15 ribs<br />

composed of conspicuous tubercles 1/4 to 3/8 of an inch high.<br />

These tubercles are almost completely separated near the stem<br />

apex, but are quite fused lower down on the stem. The sur-<br />

face is dark green.<br />

areoles: Circular at first, but the growth of the flower out<br />

of the upper edge of each produces a narrower extension of<br />

it upward which persists, leaving the areole from then on<br />

elliptical ovate, or even narrowly ovate. Areoles range in size<br />

from about 3/16 to slightly over 1/4 of an inch long, and have<br />

much white wool.<br />

spines: There are 7 to 11 straight, evenly radiating radial<br />

spines 1/2 to 1 inch long. They are rather slender, compressed,<br />

and all but 1 lower lateral on each side white in color. These<br />

2 lower laterals are gray or brownish in color. There are 4<br />

central spines on matured specimens, although immature ones<br />

have no centrals. The lowermost central is porrect to a little<br />

deflexed, strong, round to somewhat flattened, hooked at the<br />

end, and 1 to 2 inches long. It is gray or tan to reddish-<br />

purple, usually streaked with white. The two lateral centrals<br />

are similar but straight. The upper central is lighter in color,<br />

sometimes whitish, more flattened (sometimes very much so),<br />

and straight or nearly so.<br />

flowers: Very beautiful, more or less funnel-shaped blooms<br />

of varying colors: mostly fuchsia, but sometimes purplish,<br />

pink, white, or even yellowish. They are about 1 to 2 inches<br />

in diameter and nearly the same length. The short ovary has<br />

a few short, greenish, triangular scales with membranous,<br />

crinkled edges and short hairs in their axils. They intergrade<br />

into the outer perianth segments, which are almost lanceolate,<br />

greenish in the midline with the edges whitish, membranous,<br />

and crinkled. The inner perianth segments are fuchsia, pink,<br />

white, or yellowish; lanceolate; with their ends pointed and<br />

the edges entire or nearly so. The filaments are pink, rose, or<br />

yellow, the anthers orange. The style and stigma are greenish<br />

or rose, the stigma with 5 to 10 lobes.<br />

fruits: Oblong, about 1/4 to 1/2 inch long. Green to pinkish<br />

and fleshy at first. The surface has several scales upon it, with<br />

small tufts of wool in their axils. When ripe it becomes dry<br />

and then opens by splitting all the way around at or near the<br />

base.<br />

Range. The northwestern corner of New Mexico into north-<br />

eastern Arizona, the very southwestern corner of Colorado, and<br />

on into Utah. In New Mexico mostly limited to San Juan, Mc-<br />

Kinley and Sandoval counties, but recorded once from upper<br />

Socorro County.<br />

Remarks. This is a beautiful though diminutive barrel cactus<br />

which is so retiring in both its appearance and its range that<br />

many have never seen it. It looks like just another clump of<br />

dried grass and sticks in the field, and although it occurs over a<br />

wide area of several states, it is adapted to such extremely arid<br />

conditions that one has to go far into the desolate hills and Indian<br />

reservations to find it. For these reasons it has not been<br />

brought out and sold as commonly as many other cacti. This is<br />

probably just as well, since with its adaptation to such extreme<br />

conditions it is extra sensitive to moisture and most people are<br />

disappointed who try to keep it alive outside of the arid south-<br />

west.<br />

The species was first dealt with by Engelmann and Bigelow,<br />

who named it Echinocactus whipplei. The only change in its


genus Echinocactus 75<br />

name since was that made by Britton and Rose who decided to<br />

put it in one of their separate microgenera. They called it Sclerocactus<br />

whipplei, and this name has been widely used.<br />

The basis for this separation of Sclerocactus from the other<br />

Echinocacti has always been at best vague, and in 1950, when<br />

L. Benson was acting upon his own taxonomic principle of synthesis<br />

above splitting, he dropped this new genus and used the<br />

original name. But in 1966 Benson published a series of articles<br />

in which he resurrected the genus Sclerocactus again and broadened<br />

it to include species never before in it. This was done,<br />

however, without the advancing of any new facts to make the<br />

move appear more necessary now than before. The decision<br />

either way seems purely arbitrary, and since we have not seen<br />

a single new character mentioned since Britton and Rose’s<br />

doubtful ones to justify putting their little grouping on a par<br />

with the genus Echinocactus I see no real reason to regard it<br />

as more than a subgenus.<br />

In his series of articles Benson distinguishes three varieties of<br />

this species. The form he designates variety roseus does not grow<br />

in our area and so I do not deal with it here. He designates as<br />

Sclerocactus whipplei var. whipplei a form which he says is<br />

found only in Arizona, growing to only 3 inches tall, and<br />

having the upper central white, flat, and 1/16 to 1/8 of an inch<br />

wide at the base, and, so far as is known, bearing only yellow<br />

flowers. Then he has a third variety, Sclerocactus whipplei var.<br />

intermedius. This is the one found in all four states which grows<br />

to 6 inches tall, has the upper central less flattened and only<br />

1/24 to 1/16 of an inch thick at the base, and has purple, rose,<br />

pink, or white flowers. If these varieties are distinct all New<br />

Mexico specimens known so far would be the latter, as no one<br />

has reported yellow flowers in the state.<br />

I have been unable to follow these varieties in every case.<br />

Some New Mexico plants seem to duplicate all of the characters<br />

of variety whipplei so closely that they would, except for<br />

blooming with fuchsia flowers, be that variety. Since I have<br />

avoided basing any variety on flower color and the other distinguishing<br />

characters of these two varieties seem to intergrade<br />

in New Mexico specimens, I am not listing these varieties definitely<br />

here. More study of this species will be needed before a<br />

final word on them can be given.<br />

Echinocactus mesae-verdae (Boissevain) L. Benson<br />

[Sclerocactus mesae-verdae (Boissevain) L. Benson]<br />

“Mesa Verde Cactus”<br />

Description Plate 21<br />

stems: Depressed- globose to short-cylindrical in shape, with<br />

mature plants usually from 11/2 to 3 inches tall and wide,<br />

but said to have achieved 7 inches tall. There are 13 to 17<br />

ribs. On young plants they are indistinct, composed of tubercles<br />

almost completely separated from one another, but on<br />

older plants these tubercles become confluent, with deep<br />

grooves between the ribs and lesser cross-furrows between the<br />

tubercles. The surface is pale grayish-green.<br />

areoles: Ovate and 1/8 to 1/4 of an inch long, containing<br />

much wool at first yellowish and then fading to gray.<br />

spines: There are 8 to 11 radial spines spreading irregularly<br />

around the areole. They are straight or slightly curved, round<br />

or nearly so, tan or straw-colored, about 3/8 to 1/2 inch long,<br />

and rigid. Central spines are usually missing, but on very rare<br />

individuals are present—then described as being one per<br />

areole, 1/2 inch or less in length, gray with a dark tip, porrect<br />

to ascending, and straight or sometimes hooked.<br />

flowers: Cream to whitish in color, funnel-shaped, 3/4 to 11/4<br />

inches wide and tall. The ovary and tube have a few broadly<br />

triangular scales placed high on them, but have no wool. The<br />

outer perianth segments are from triangular to oblanceolate<br />

and brownish with yellowish, entire margins. Inner perianth<br />

segments are cream or whitish from greenish bases, oblanceolate,<br />

entire or somewhat erose at the tips. The filaments<br />

are green or yellowish-green, the anthers yellow. The stigma<br />

has 6 to 8 light green lobes. The flowers are fragrant.<br />

fruits: Very small: about 3/16 of an inch long, and cylindrical<br />

in shape. They are greenish at first, becoming brownish<br />

and dry and finally splitting open with an irregular, transverse<br />

opening near the middle of the fruit. The seeds are<br />

black, 1/8 of an inch or a little more in the longest measure-<br />

ment.<br />

Range. A very small area in the extreme southwest corner of<br />

Colorado and the extreme northwest corner of New Mexico,<br />

actually comprising a strip of territory only about fifty miles<br />

long from near Cortez, Colorado, past Mesa Verde to slightly<br />

southwest of Shiprock, New Mexico.<br />

Remarks. This is one of the rarest of our cacti. It was discovered<br />

by Boissevain near Cortez, Colorado. He remarked about its<br />

rarity and the fact that it is adapted to drought beyond any<br />

other Colorado cactus, so much so that he could not keep it<br />

healthy even in his Colorado garden. Actually it is adapted to<br />

an extremely alkaline soil, as well as to extreme dryness. The<br />

combination of its rarity and the difficulty encountered in trying<br />

to grow the plant artificially means that few have ever seen<br />

it. Yet it does grow down into New Mexico, and is a most interesting,<br />

if rather prosaic cactus inhabitant of our area.<br />

In his first description of the cactus, Boissevain thought of it<br />

as representing a new genus. Going all the way with pride of<br />

state, he gave it the name Coloradoa mesae-verdae, certainly<br />

one of the most geographical plant names ever. It seems difficult<br />

to justify Coloradoa as a genus, and in 1951 L. Benson moved<br />

the plant to the genus Echinocactus. This would seem to have<br />

given the species a place to rest comfortably, but in a recent set<br />

of articles Benson has reinstated the genus Sclerocactus again,<br />

after having himself dropped it, and now he assigns this cactus


76 cacti of the southwest<br />

to that genus. Thus this cactus has begun its journey, following<br />

the others in looking for a generic home. We hope it is not as<br />

long a journey as it has been for some others of the Echinocacti.<br />

In view of the extreme rarity of this cactus and the great<br />

difficulty which is encountered in keeping it alive after taken,<br />

it would seem useless and wanton to endanger the species by<br />

collecting in the wild specimens which are probably doomed<br />

by being taken anyway. Perhaps someone can grow these plants<br />

from seeds, and thus condition them to cultivation, after which<br />

this rarity can take its place in collections.<br />

Echinocactus brevihamatus Eng.<br />

[Ancistrocactus brevihamatus (Eng.) B. &R.]<br />

“Fishhook Cactus”<br />

Description Plate 21<br />

roots: Fibrous.<br />

stems: Globose, egg-shaped, or, when older, columnar. The<br />

maximum size seems to be about 5 inches high by 31/2 inches<br />

in diameter, and no clustering has been reported. The surface<br />

is a very dark, dull green. There are 13 straight or slightly<br />

spiraling ribs which are made up of rows of tubercles almost<br />

completely separated from each other by deep notches. These<br />

tubercles are up to about 1/2 inch tall from bases nearly as<br />

broad, but are compressed from side to side to leave an uninterrupted<br />

groove between the ribs.<br />

areoles: Almost linear. The lower end of the areole is slightly<br />

expanded and contains the spine cluster. Above this is a<br />

narrow groovelike portion of the areole containing 1 to several<br />

glands. At the upper end of this groove the flower is<br />

produced, and after flowering this portion remains a little<br />

expanded. The whole areole thus measures 1/4 to 1/2 inch long<br />

and runs from three-fourths to all of the way to the base of<br />

the tubercle. These areoles are at first filled with white wool,<br />

which is mostly lost on old areoles.<br />

spines: There are 12 to 14 rather heavy radial spines which<br />

spread out slightly from the plant. They are from 3/8 to 1<br />

inch long, the upper ones the longer. They are opaque, tan<br />

with dark brown tips at first, then turning gray. There are<br />

typically 4 centrals, but occasionally 1 or 2 more. The uppermost<br />

is erect in front of the upper radials, slender, straight,<br />

its upper surface flat while its lower side is rounded. This<br />

spine is 3/4 to 11/4 inches long. On either side of it are 2 or<br />

more centrals diverging upward and similar except growing<br />

to 13/4 inches long. These upper centrals are all colored like<br />

the radials except that their flattened upper sides may be<br />

rust-colored in some specimens. The lowermost central stands<br />

out perpendicular to the plant surface, is heavier, distinctly<br />

flattened, and hooked, and measures 3/4 to 1 inch long. It is<br />

yellowish-brown, rust, or dark brown on its flat upper side,<br />

lighter below, with the hook dark brown.<br />

flowers: Green in color, suffused with rose. They are 1 to<br />

15/8 inches long, but only 1/2 to 3/4 of an inch wide, since<br />

they can hardly open because of the spines around them. The<br />

ovary of the flower has upon it fewer than a dozen small<br />

scalelike segments with fringed edges. Next there are about<br />

8 outer perianth segments which are oblong with bluntly<br />

pointed ends. There are slightly over a dozen inner segments<br />

which are almost linear, about 5/8 of an inch long, and<br />

pointed. These are dark green with a faint rose midline on<br />

the outer side and dull rose fading to green edges on the inner<br />

surfaces. The filaments are bright rose. The anthers are yellow.<br />

The style is short, and there are 10 or 11 rose-pink stigma<br />

lobes.<br />

fruits: Egg-shaped, green, becoming faintly pinkish when<br />

very ripe. They are 5/8 to about 1 inch long. There are several<br />

scales upon the surface of each fruit. They do not become<br />

completely dry, but remain somewhat fleshy until they disintegrate.<br />

The seeds are dark brown to blackish.<br />

Range. Along the Rio Grande from near the mouth of the Pecos<br />

River to near Eagle Pass, occurring northeast almost to Uvalde<br />

and Brackettville, Texas, in the Anacacho Mountains, but not<br />

found otherwise more than a few miles north of Del Rio or up<br />

the Devil’s River.<br />

Remarks. This species is a member of a very closely related<br />

group. However it has remained distinct in almost the whole of<br />

the literature since its first description by Engelmann, being<br />

regarded as a separate species by all except Weber who reduced<br />

it to a variety of the more widely known species, Echinocactus<br />

scheeri.<br />

This plant is immediately told from E. scheeri by its fibrous<br />

roots, its darker green body, its fewer and longer opaque radial<br />

spines, and its flowers which are suffused with rose. Many take<br />

the name brevihamatus to mean that it has the lower, hooked<br />

central spine shorter than those of its relatives, but this leads<br />

them into difficulty, since the centrals of E. scheeri are often<br />

much shorter. Engelmann coined the name because of “the<br />

shortness of the hook,” not the shortness of the spine, by which<br />

he meant that the hook itself is not curved as far around as in<br />

the typical E. scheeri. This is still an observation of doubtful<br />

value, since I have seen specimens of E. scheeri on which the<br />

spines failed to hook at all.<br />

The species is limited to a comparatively small area in four<br />

counties of Texas along the middle course of the Rio Grande. It<br />

is found growing in the scattered clumps of low vegetation on<br />

rocky hillsides overlooking the river and on alluvial soil between<br />

the hills, never far from the Rio Grande except where it<br />

spreads into the Anacacho Mountains.<br />

Engelmann included this species and its relatives in the genus<br />

Echinocactus. They were left there, a subgenus Ancistrocactus<br />

being erected by Schumann for them and some other species<br />

later put in a number of other genera, until Britton and Rose<br />

broke up the genus. When they did this Britton and Rose ele-


genus Echinocactus 77<br />

vated Ancistrocactus to an entirely separate genus containing<br />

only this and a couple of other species. Their arrangement is<br />

naturally best known today. But if, in order to methodize some<br />

of the chaos which has resulted from Britton and Rose’s break-<br />

up of that genus we go back to the original genus Echinocactus<br />

these are surely part of it.<br />

Echinocactus scheeri SD<br />

[Ancistrocactus scheeri (SD) B. & R.]<br />

“Fishhook Cactus,” “Root Cactus”<br />

Description Plate 21<br />

roots: E. scheeri grows from a long, fleshy, white taproot<br />

which is 1/4 to 1/2 inch in diameter and may be from a few<br />

inches to as much as 3 feet long. This root may occasionally<br />

branch to form several the same size.<br />

stems: Globular at first, becoming quickly columnar or even<br />

club-shaped with the upper end often twice the diameter of<br />

the lower half. These stems grow to at least 7 inches tall and<br />

3 inches in diameter, and are single until very old, when they<br />

sometimes branch from the base to form clumps of 6 or 8<br />

stems. The flesh is medium to dark green in color. There are<br />

13 straight or spiral ribs which are composed of only slightly<br />

connected, almost perfectly conical or only slightly compressed<br />

tubercles to 1/2 inch tall and the same width at their<br />

bases on large specimens.<br />

areoles: Broadly ovate on unflowering tubercles. On flower-<br />

ing tubercles the ovate areoles are prolonged by a short<br />

groovelike extension above, at the upper end of which the<br />

flower appears. There are several glands in this groove. The<br />

length of this groove and, therefore, of the areole depends to<br />

a great degree on the age of the plant. Typically it extends<br />

from the spinous part of the areole at the summit of the tubercle<br />

about one-half of the way down the upper side of the<br />

tubercle; but in immature plants the groove does not form at<br />

all; in young flowering plants it is short; and on old plants it<br />

often extends three-fourths or more of the way to the axil.<br />

The areoles, therefore, vary from about 1/8 to as much as 3/8<br />

of an inch long. They have much white wool in them at first.<br />

spines: There are 13 to 28 very slender radial spines, radiating<br />

strictly and tending to recurve toward the plant. They are 1/4<br />

to 1/2 inch long on young plants, but grow to as much as 11/8<br />

inches long on some very old specimens. They are a very light,<br />

translucent yellowish shade, with the tips red-brown. There<br />

are also 3 or 4 centrals as follows: 2 straight upper centrals<br />

diverge as they stand erect to form a distinct V. They are 3/4<br />

to 2 inches long, distinctly flattened, the upper side being<br />

brown to dark mahogany-brown, while the lower or outer<br />

side is tan to whitish. There may or may not be 1 more upper<br />

central bisecting the V formed by the other 2 and similar to<br />

them except shorter and more slender. The lowermost central<br />

stands out perpendicular to the stem; it is stout, flattened<br />

above or sometimes almost round, and almost always hooked.<br />

It varies on different plants from 1/2 to 11/2 inches long. In<br />

color it is variegated, being mostly dark brown or black<br />

above and light brown or whitish below.<br />

flowers: Plain green or yellow-green. About 1 inch long,<br />

but only 1/2 to 3/4 of an inch wide due to their inability to<br />

open farther because of the surrounding spines. There are<br />

about 12 fringed scales on the ovary. There are about 8 outer<br />

segments of the perianth, which are linear with blunt ends,<br />

deep green with yellowish edges. There are about 13 inner<br />

segments, to about an inch long and almost 1/4 of an inch<br />

wide, with pointed ends. These are bright green in some specimens<br />

or yellowish in others. The color of the filaments varies.<br />

In some plants they are green, in some yellowish, while in a<br />

few they are pink or even reddish. The anthers are yellow or<br />

pale orange. The style is somewhat longer than the stamens.<br />

The stigma lobes vary in number from 5 to 10, in length from<br />

3/16 to 1/4 of an inch, and in color from green to yellowish,<br />

cream or even brown, sometimes the brown ones having a<br />

faint pink flush at the tips.<br />

fruits: 3/4 to 13/8 inches long, not including the old perianth<br />

which persists upon them. They are club-shaped. There are 12<br />

to 24 scales widely spaced upon each fruit. They remain green<br />

for the whole of the summer, but when very ripe they finally<br />

turn yellowish tinged with pink. They ultimately disintegrate,<br />

sometimes splitting open vertically on one side as they rot.<br />

The seeds are about 1/16 to of an inch long, very dark mahoganybrown<br />

with the surface dull instead of shining due to extremely<br />

fine pitting. They are globular, compressed from side<br />

to side, with large, deeply concave hila.<br />

Range. In Texas south of a line from approximately Eagle Pass<br />

to Pleasanton to near Kingsville, also ranging deep into Mexico.<br />

Remarks. This species was described before any of its relatives<br />

by Prince Salm-Dyck. As Britton and Rose mention, there is<br />

doubt as to what specimens the description was originally based<br />

upon. Engelmann made his description, the first detailed one,<br />

from obviously very immature specimens, since he gave as the<br />

maximum size a mere 2 inches tall. His plants also were from<br />

Eagle Pass, the northern extreme of their range. Coulter, Britton<br />

and Rose, and even Backeberg have descriptions which are essentially<br />

repetitions of Engelmann’s, although the latter two did<br />

increase the maximum size to 4 inches tall.<br />

I have studied literally hundreds of these plants in their habitats<br />

and also as they have come to San Antonio dealers before<br />

being sent all over the world. I have seen plants from Eagle Pass<br />

on the north to Brownsville on the south and numbers from<br />

Mexico. As a result I find that I must enlarge the description in<br />

several respects.<br />

I have in my collection a plant which consists of a single stem


78 cacti of the southwest<br />

7 inches tall and club-shaped, being 3 inches in diameter near its<br />

top and only 2 inches thick near the ground. I have seen several<br />

clusters formed by the branching of such large stems near the<br />

ground, the largest with 8 heads forming a cluster about a foot<br />

across. This species can be a much larger and more majestic plant<br />

than we have been told.<br />

Very early in my study of it, I noticed wide variations in the<br />

number of radial spines on different plants—considerably wider<br />

than the limits heretofore given. Engelmann’s 11 to 18 was narrowed<br />

by Britton and Rose to 15 to 18, which number Backeberg<br />

repeats. On the other hand, in one group of 50 plants at a<br />

single location near Zapata, Texas, I found specimens with all<br />

numbers of radials from 13 and 14 on one plant to 28 on all<br />

areoles of another. Many plants varied widely in the number<br />

on the various areoles of the same plant. For instance, one had<br />

18, 19, 21, 23, and 24 on its different areoles. I must state, however,<br />

that every plant I studied in the vicinity of Eagle Pass had<br />

between 14 and 17 radials, showing why Engelmann did not<br />

give the higher numbers in his description.<br />

I was fortunate to be able to examine much larger specimens<br />

than the earlier authorities, and I found the spines on these to<br />

be somewhat longer also. For instance, some of these old plants<br />

have radials to 11/4 inches long where Engelmann described<br />

them as to only 1/2 inch on his little plants and everyone has<br />

dutifully followed him. I found great variation in the lower<br />

central spine also, from only 1/2 inch long on some plants to 11/2<br />

inches on others. I have one good big plant 5 inches tall which<br />

keeps its short but stout 1/2-inch centrals and also one 4-inch<br />

plant whose centrals are all 11/2 inches long, so central length is<br />

not entirely related to the size of the plant.<br />

The flowers of the species vary also. The petals may be dark<br />

green or green suffused with yellow so as to be almost golden.<br />

The filaments may be various colors, and the stigma lobes vary<br />

greatly in number, size, and color. All of the several flowers of<br />

any single specimen are consistent in these characters, however.<br />

The fruits of the species are all similar, as described, and they<br />

are all alike in having the long, white taproot of almost unvarying<br />

size over a distance of up to several feet. This root is<br />

said to be unique among cacti. Many people, not knowing of it,<br />

pull the cactus up instead of digging it out. The root is so slender<br />

that it breaks off easily and they may not realize what is missing<br />

when they take the plant home or why it languishes so long<br />

before it re-establishes its roots.<br />

The species grows over a wide area, including much of Mexico,<br />

and it seems clear that this is an aggregate containing several<br />

forms which could perhaps be separated out as varieties, if<br />

anyone wished to make a complete study of them. Britton and<br />

Rose have a form which appears at first glance to be one of<br />

these. It is their Echinocactus megarhizus Rose, from near Victoria,<br />

Mexico. It has the fleshy root, 20 or more radials, and 4<br />

centrals quite like those of E. scheeri. However, they describe<br />

its seeds as black and shining, which would set it apart from<br />

our cactus at once.<br />

I have seen two specimens of E. scheeri in which the lower<br />

centrals were not hooked at all.<br />

It should be noticed that E. scheeri may not be separated<br />

from E. brevihamatus on the basis of how far down the tubercle<br />

the groove extends, as Engelmann seemed to think, since on the<br />

large, old plants of E. scheeri, which Engelmann’s measurements<br />

indicate he did not have before him, the groove goes fully as far<br />

as on the other species.<br />

The plant has been one of the most common on gravelly hill-<br />

sides from Eagle Pass south, becoming very common around the<br />

area of Falcon Lake. However, dealers have kept after it relentlessly.<br />

It is found in almost every box of cacti in dime and curio<br />

stores, and more recently large tracts of its territory are being<br />

cleared, which spells its doom in those areas. We hope the conspicuous<br />

V formed by its upper centrals means victory for it<br />

over its enemies and that this attractive little fishhook cactus<br />

will be with us a long time to come.<br />

This, along with the last species, will be in the genus Echinocactus<br />

or the genus Ancistrocactus according to the philosophy<br />

of the person defining these genera.<br />

Echinocactus tobuschii (Marsh)<br />

[Ancistrocactus tobuschii Marsh]<br />

Description Plate 22<br />

roots: Short, not too well developed taproots which are turnip-shaped,<br />

tapering, and brown rather than white.<br />

stems: Low, flattened hemispheres. The largest I have seen<br />

was about 31/2 inches in diameter and nearly as tall. I have<br />

seen only one double plant, but have been told of clusters of<br />

8 and 10 heads having been found. The flesh is dark green.<br />

There are 8 broad, flat ribs made up of pyramidal tubercles<br />

to almost 1/2 inch tall from greatly flattened, quadrangular<br />

bases to as much as 5/8 of an inch wide.<br />

areoles: Linear or very nearly so. From the spine cluster at<br />

the lower end of the areole there extends a narrow groove inward<br />

and upward usually half to three-fourths of the length<br />

of the tubercle to the floral part of the areole. There is some<br />

white wool when the areole is young, but this is later lost.<br />

There are also 1 or 2 glands in the groove of each areole.<br />

spines: There are 7 to 12 slender radial spines which are 5/8<br />

to 3/4 of an inch long, equal in length on any given areole,<br />

yellowish in color, becoming gray with age, the tips being a<br />

little darker. There are 3 to 5 centrals. The 2 upper centrals<br />

are always diverging to form an erect V in front of the upper<br />

radials. These upper centrals are to 11/2 inches long, flattened,<br />

and ridged. There may or may not be 1 other upper central<br />

bisecting the V formed by the first two. If it is present it is<br />

similar to the others, except more slender, growing to only<br />

7/8 of an inch long, and recurving somewhat back toward the


genus Echinocactus 79<br />

plant. Very rarely on some areoles there may be 2 of these<br />

erect centrals in the V. There is always 1 lower central standing<br />

out perpendicular to the plant or directed upward a little.<br />

It is stout, hooked, angled and ridged, arid to 1 inch long. All<br />

centrals are translucent yellowish to gray in color.<br />

flowers: 1 to 11/2 inches long and to almost as wide, opening<br />

almost completely. The ovary and tube have many scalelike<br />

segments upon them which are greenish, triangular in shape,<br />

with yellowish, entire edges. These intergrade to almost linear<br />

outer perianth segments, with blunt tips, midlines greenish<br />

tinged with brown, and with yellow, entire edges. The inner<br />

segments are about 20 to 25 in number, clear citron or golden<br />

yellow, with no variation at midline or edges. They are<br />

shorter than the outer segments, almost spatulate, coming to<br />

pointed tips often having a very small, soft spine at the apex.<br />

The filaments are yellowish, the anthers pale orange. The<br />

style is green and up to 1/4 of an inch longer than the stamens.<br />

The stigma has 5 to 9 yellow or whitish lobes which are very<br />

small at first, but when expanded become over 1/8 of an inch<br />

long.<br />

fruits: Elongated egg-shaped, about 1 inch long, greenish in<br />

color, flushing pinkish when very ripe. There are numerous<br />

small scales upon it. The seed is almost spherical, very dark<br />

brown and shiny, with a large hilum.<br />

Range. Known only from a very small area in the Texas hill<br />

country from just above Vanderpool to near Ingram and Mountain<br />

Home, Texas.<br />

Remarks. This little cactus was discovered by H. Tobusch in<br />

1951 and described by Marsh in 1952. It is one of the rarest<br />

forms in the Southwest, known only from an area not more than<br />

30 miles long. I have succeeded in finding scattered plants in the<br />

type area, but there appear to be stands of them in only a couple<br />

of places and I would estimate that the population does not<br />

comprise over a few hundred plants in all. This should be remembered<br />

by any who are tempted to take any number of them.<br />

They could become extinct very easily.<br />

These plants might at first be mistaken for small specimens of<br />

E. brevihamatus with fewer ribs made up of broader tubercles<br />

and fewer radial spines. The range of that other species comes<br />

to about 50 miles southwest of this plant at the nearest point<br />

that I could find. Any idea that they might be the same species<br />

is dispelled, however, when they bloom. The bright yellow,<br />

broadly opening flowers with entire edges on the outer perianth<br />

segments and with greenish-yellow filaments, long styles, and 5<br />

to 9 stigma lobes found on E. tobuschii are greatly different<br />

from the green suffused with rose, hardly open flowers of the<br />

other form, with their rose-pink filaments, short styles, and 10<br />

or 11 rose-pink stigma lobes. E. tobuschii seems to be a distinct<br />

form limited to the environs of the canyons cutting into the<br />

edge of the Edwards Plateau.<br />

The author of the original description of this cactus was<br />

already a victim of the confusion which exists about the cactus<br />

genera. In his description he used the dodge so often resorted<br />

to when one is confused and called the cactus Mammillaria<br />

(Ancistrocactus) tobuschii. Of course a review of the characteristics<br />

will make it clear that it is not a Mammillaria. It is<br />

definitely one of the group known widely as the genus Ancistrocactus,<br />

which group is obviously part of the Echinocacti. As this<br />

group is returned to the genus Echinocactus this species must<br />

necessarily follow, and once its relationship to the other Echinocacti<br />

is seen, its difference from the Mammillarias should also be<br />

clear.<br />

Echinocactus setispinus Eng.<br />

[Hamatocactus setispinus (Eng.) B. & R.]<br />

“Fishhook Cactus,” “Hedgehog Cactus,”<br />

“Twisted-Rib Cactus”<br />

Description Plate 22<br />

stems: Hemispheric at first, later egg-shaped, and finally<br />

columnar. They may be single or may branch around the base<br />

to form clusters of up to 7 or 8 heads. There is great variation<br />

in size, some individuals reaching a maximum of 3 inches<br />

tall, while others may reach 12 inches. One form never exceeds<br />

3 inches in diameter, while another attains 5 inches.<br />

The color of the surface is light green in one form and dark<br />

green in another. There are 13 sharp ribs which are about 3/8<br />

of an inch high, undulating, with shallow cross-furrows<br />

between the areoles. On one form these ribs usually spiral, on<br />

another they are straight and vertical.<br />

areoles: At first elliptic or egg-shaped and about 3/16 of an<br />

inch long. They very soon elongate to nearly twice that<br />

length and becoming obovate as the floral part of the areole<br />

develops above the original spine-producing part. After the<br />

flower and fruit are gone, the floral part of the areole remains,<br />

with several glands within it. Then, as it gets older this<br />

floral part contracts to become a narrow groove running<br />

about 1/8 of an inch inward and upward from the spine-<br />

bearing part of the areole. There is quite a lot of yellowish<br />

wool in the young areole, most of which is lost with age.<br />

spines: The radial spines are 10 to 19 in number, round,<br />

slender, bristle-like to somewhat rigid, and straight or recurved<br />

toward the plant. They are from 3/16 to 11/4 inches<br />

long, the upper ones the longest. They are dark mahogany,<br />

yellow, or whitish in color. There is one central spine which<br />

stands perpendicular to the plant surface, is round, weak,<br />

hooked, and 1/4 to l3/8 inches long. This central is translucent<br />

yellow fading to gray on one form, while on another it is<br />

opaque brown.<br />

flowers: Ivory or cream with red centers and extremely<br />

fragrant; 13/4 to 3 inches tall and 21/4 to 3 inches wide. The<br />

ovary is cylindrical with a few scales on it and the outer


80 cacti of the southwest<br />

perianth tube has many triangular scales which are green,<br />

brownish, or reddish-green, with white or yellowish, fringed<br />

edges. The outer perianth segments are triangular to oblong<br />

with blunt ends, or sometimes with their upper parts much<br />

broader than the bases and flaring into earlike projections on<br />

each side just below the blunt tip. They have greenish<br />

midlines with yellowish, fringed edges. The inner segments<br />

are spatulate and ivory or cream with red bases. Their edges<br />

are somewhat ragged and often toothed, the tips pointed. The<br />

filaments are reddish, weak, and swirled. The anthers are<br />

cream-colored or pale yellow. The style is long, thick, and<br />

greenish-yellow. There are 5 to 11, but usually 9 or 10 stigma<br />

lobes, which are yellow or pale orange, to 1/4 of an inch long,<br />

rough, and blunt. They usually curve in all directions.<br />

fruits: Spherical or nearly so, 3/8 to 3/4 of an inch across.<br />

Bright scarlet in color with a smooth, shiny skin and almost<br />

no scales. They remain upon the plants for a long time, often<br />

a year, if not disturbed, before they finally dry up and split<br />

open down one side, releasing the seeds. The seeds are about<br />

1/16 of an inch long, black, with a finely pitted surface,<br />

somewhat irregular in shape, having been described variously<br />

as globose and club-shaped. The large hilum at or near the<br />

end of the seed is surrounded by a broad lip with an unpitted,<br />

shining surface.<br />

Range. Growing over a wide area of central and south Texas<br />

and on into Mexico. Known to grow north as far as San Saba,<br />

Lampasas, and near Georgetown, Texas. From there the northeastern<br />

boundary of its range seems to be approximately the<br />

Colorado River. Rare near the mouth of that river, it becomes<br />

more and more common going south along the lower Texas<br />

Coast all the way to Brownsville. On the west it is not found so<br />

far north, but is seen around the mouth of the Devil’s River,<br />

which seems to be about the limit of its northwestern range.<br />

Remarks. Echinocactus setispinus is one of the most common<br />

cacti in south Texas, and also one of the most colorful. It will<br />

bloom practically the whole summer if happily situated. It is at<br />

the same time one of the most easily grown, being very resistant<br />

to rotting, and so is a great favorite for indoor growing as a<br />

potted plant. I have seen large specimens which had lived for<br />

many years in the windows of business establishments, planted<br />

all these years in flat dishes with soil not over two inches deep.<br />

I have also seen numerous dish gardens and planters originally<br />

planted with an assortment of various cacti, in which, after<br />

some months, only this species remained alive. Here at last is<br />

the cactus which I can recommend for the amateur grower who<br />

cannot provide the specialized conditions other cacti require<br />

and who wants to grow a cactus on his windowsill. It does not<br />

like the full sun, and it tolerates more moisture than most.<br />

In the extreme western part of its range this cactus is found<br />

growing near another species of this genus, E. sinuatus, and is<br />

often confused with it. The two are readily distinguished,<br />

however, by the fact that E. sinuatus has broader, not so sharply<br />

edged ribs, fewer radial spines, and 4 central spines at least<br />

some of which are always flattened.<br />

E. setispinus was originally considered by all authorities as<br />

part of the genus Echinocactus. When Schumann set out sub-<br />

genera for that large genus he placed it in his subgenus<br />

Ancistrocactus. But Britton and Rose segregated this species<br />

from the rest in a new genus, Hamatocactus. Other species were<br />

added to this new genus by other authors, and for a time its<br />

favor in taxonomic opinion was at full tide. Then the tide began<br />

to turn as they began to subtract from the proposed genus<br />

Hamatocactus. Buxbaum put one of its members back into<br />

Ferocactus, while adding E. uncinatus to it. Backeberg, of<br />

course, took E. uncinatus out again as he erected a new genus<br />

for it. Meanwhile Hester had decided on the basis of seed<br />

characters that all of those species put in Hamatocactus really<br />

belonged in Thelocactus. This sort of confusion has not yet been<br />

resolved, and seems to be the result of trying to define the genera<br />

too closely. Since no new character has come to light which<br />

will separate these microgenera cleanly and no one has managed<br />

a combination of characters which will organize them<br />

logically, the return to the inclusive old genus Echinocactus<br />

seems indicated here too, at least until these smaller groups can<br />

be redefined in meaningful terms.<br />

Echinocactus setispinus var. hamatus Eng.<br />

[Hamatocactus setispinus var. hamatus (Eng.) B. & R.]<br />

“Twisted-Rib Cactus,” “Fishhook Cactus,”<br />

“Hedgehog Cactus”<br />

Description Plate 22<br />

stems: Hemispheric when young, becoming columnar when<br />

old, usually occurring singly, but occasionally branching at<br />

the base to form a clump of several stems. The stems grow up<br />

to 12 inches tall and 5 inches in diameter. They are dark, dull<br />

green in color, and have 13 very sharp ribs which undulate<br />

somewhat, but are not interrupted by cross-grooves. These ribs<br />

are about 3/8 of an inch high, and usually spiral by twisting<br />

sideways between the areoles. There are hardly distinct<br />

enough swellings at the areoles to be called tubercles, but the<br />

ribs are slightly higher at each areole than between them.<br />

areoles: As the species.<br />

spines: There are 10 to 13 radial spines which are very slender,<br />

bristle-like, and rather flexible. They are straight and<br />

radiate evenly. They are all translucent yellow at first, but<br />

when mature they develop as follows: the lower 3 become<br />

3/8 to 3/4 of an inch long and dark mahogany-brown on some<br />

plants, honey-yellow on others; the lateral 2 or 3 on each<br />

side become 5/8 to 1 inch long and white in color, only the<br />

very tips of these sometimes remaining honey-yellow; the 3<br />

to 5 uppermost ones become 3/8 to 11/4 inches long, dark


genus Echinocactus 81<br />

mahogany-brown or variegated shades of brown in color.<br />

There is 1 central spine which stands out perpendicular to the<br />

stem and is hooked. It is 1 to 13/4 inches long, round, and<br />

very slender, being so weak as to be easily flexible. It is<br />

brown, often very dark, but with the tip usually lighter.<br />

flowers: As the species, except that they are 2 to 3 inches<br />

tall and 21/2 to 3 inches wide and have the inner perianth segments<br />

almost linear, to about 1/2 of an inch wide. The edges<br />

are crinkled but not toothed or fringed, and the tips are<br />

pointed.<br />

fruits: As the species.<br />

Range. The northern and eastern part of the species range.<br />

There is no record of this variety south of Eagle Pass along the<br />

Rio Grande until near Brownsville. The southwestern limit of<br />

its range seems to be a large arc from Eagle Pass on the north<br />

swinging east of Cotulla, Texas, then south to near Alice, and<br />

on down to some point in the lower Rio Grande Valley not far<br />

west of Brownsville.<br />

Remarks. The species contains such well marked and constant<br />

variations that from its very first descriptions it has been found<br />

necessary to speak of varieties. Engelmann set up two of these,<br />

but remarked that “many forms” had been collected. It does<br />

seem that his two varieties are basic, however. Even though<br />

there is variation within each of them, they seem to be the only<br />

two consistently distinct and separable varieties.<br />

E. setispinus var. hamatus is the larger, more grand form of<br />

the species, and it also occurs over much the greater range of<br />

the two. At its prime it is a beautiful, bulky plant up to a foot<br />

high, dark green, with very slender, flexible spines, the central<br />

one of each cluster a perfect little fishhook as round and slender<br />

as a real fishhook of its size. When it blooms, which if it gets<br />

moisture enough is all summer from April to October, it produces<br />

its large, extremely fragrant, yellow and red flowers several<br />

at a time, and soon the upper part of the plant is also adorned<br />

by the scarlet fruits. Just recently, near San Antonio, I happened<br />

upon 13 plants of this variety growing under one mesquite tree,<br />

each with 3 to 6 flowers open. As other specimens were under<br />

almost every tree and shrub in every direction, both the sight<br />

and the fragrance of this unspoiled field was delightful.<br />

The ribs of this cactus are very sharp and not interrupted<br />

between the areoles. They almost always spiral by twisting sideways<br />

between each pair of areoles. It is from this that the plant<br />

gets the name twisted-rib cactus. This twisting becomes more<br />

pronounced as the cactus loses water during drought or settles<br />

with old age.<br />

This variety grows over much of south-central Texas. It is<br />

common along the upper Colorado River and in the hill country<br />

of Texas from just north of Austin to some 30 miles or so south<br />

of San Antonio and from there on west. It is again very common<br />

along the lower Texas coast from Corpus Christi to<br />

Brownsville. It does not grow along the Rio Grande any farther<br />

than 50 miles or so above its mouth until it is found again near<br />

Eagle Pass, from which point it apparently grows at least as<br />

far west as the mouth of the Devil’s River. I have also seen<br />

beautiful specimens of this cactus which were said to have come<br />

from Coahuila, Mexico approximately south of Sanderson,<br />

Texas. This is perhaps not surprising since Coulter said it grows<br />

in Coahuila and even in Chihuahua. However I have found no<br />

verifiable records of it in Texas west of the Devil’s River.<br />

In the gap along the Rio Grande where this cactus, variety<br />

hamatus, does not grow, and limited to that gap, is found the<br />

other distinct variety of the species, variety setaceus. This is a<br />

much smaller form with much more slender stems, straight ribs<br />

and heavier, more rigid, as well as more numerous spines. The<br />

best means of distinguishing the two varieties will be pointed<br />

out after that other variety is described. A good photo of a young<br />

specimen of variety hamatus is presented by Britton and Rose<br />

on page 104 of Volume 3 of The Cactaceae, while the other<br />

variety is well illustrated on the following page.<br />

Echinocactus setispinus var. setaceus Eng.<br />

[Hamatocactus setispinus var. setaceus (Eng.) B. & R.]<br />

“Fishhook Cactus,” “Hedgehog Cactus”<br />

Description Plate 22<br />

stems: Hemispheric when very young, becoming egg-shaped,<br />

and when very old often becoming columnar. Some plants<br />

branch early to form clusters of up to 7 or 8 heads, while<br />

others remain single throughout life. Clustering plants do not<br />

usually grow over 3 inches tall, while single plants may become<br />

12 inches in height. The maximum diameter in either<br />

case seems to be about 3 inches. The surface is a light green<br />

color. There are 13 sharp ribs which are straight, vertical,<br />

and shallow, being less than 1/4 of an inch deep, but which<br />

are interrupted by cross-furrows between the areoles. There<br />

are almost no thickenings at the areoles, which could be called<br />

tubercles.<br />

areoles: As the species.<br />

spines: There are 12 to 19 radial spines which are round and<br />

rigid, straight or often recurved back toward the plant. The<br />

lower 3 or 4 are 3/16 to about 1/2 inch long and translucent<br />

honey-yellow, sometimes with their bases reddish-brown. The<br />

lateral 3 to 5 on each side are 3/8 to 3/4 of an inch long and<br />

whitish with their tips translucent yellow. The upper 3 to 5<br />

radial spines are 5/8 to 11/8 inches long, translucent honeyyellow,<br />

often with dark red-brown bases. There is 1 central<br />

spine which stands approximately perpendicular to the stem,<br />

is round, hooked, weak but rigid, and 1/4 to 11/2 inches long.<br />

On some plants this central is translucent honey-yellow becoming<br />

gray with age, while on others it is partly or all dark<br />

red-brown.<br />

flowers: As the species, except that they are only 13/4 to 2


82 cacti of the southwest<br />

inches tall by 21/4 to 3 inches wide and the inner segments<br />

are spatulate.<br />

fruits: As the species.<br />

Range. Starr and Hidalgo counties, Texas, and south into<br />

Mexico.<br />

Remarks. Although the descriptions of various authors are confusing<br />

due to the failure of some to distinguish between this and<br />

the previous variety, this plant seems clearly to be E. setispinus<br />

var. setaceus of Engelmann. It can readily be recognized by its<br />

straight, shallow ribs, its shiny, pale green color, its more numerous<br />

and more rigid radial spines, and its slender stems never<br />

observed to be over 3 inches in diameter.<br />

This variety is found in great numbers in a very restricted<br />

area of south Texas. It suddenly appears on gravelly hills near<br />

Falcon Dam, and is common on such undisturbed slopes nearly<br />

to Mission, Texas. On a specimen in the U. S. National Museum<br />

collected by Robert Runyon, who no doubt knew south Texas<br />

cacti more completely than any other man, I find a note that<br />

this cactus occurs in only Starr and Hidalgo counties. This is an<br />

area where variety hamatus is not found.<br />

It seems almost impossible to discuss variety setaceus in any<br />

more detail without noting that within it there are two obvious<br />

growth forms. They grow together in the same fields, and I have<br />

found the two forms actually within inches of each other, in<br />

which case the difference between them is striking.<br />

One of them always remains a single stem and becomes when<br />

old a very slender column up to 12 inches tall but only 2 to 3<br />

inches thick. This form has the upper radials and the hooked<br />

central variegated with brown or sometimes wholly dark brown.<br />

The centrals are 1/2 to 11/2 inches long.<br />

The other form never grows as a single tall column, but begins<br />

branching around the base when about 21/2 inches tall. An old<br />

plant will be composed of up to 7 or 8 egg-shaped stems, the<br />

largest of which is only 3 inches tall. This form has all its spines<br />

shorter and honey-yellow, becoming gray when old. There is no<br />

brown in them. The hooked centrals are only 1/4 to 3/8 of an<br />

inch long, but they are heavier than on the other form.<br />

These two forms seem to correspond in many ways to two<br />

varieties in the literature, the tall one with darker spines to<br />

E. setispinus var. cachetianus (Monv.) Knuth, and the shorter<br />

form with yellow spines to E. setispinus var. mierensis K. Schumann,<br />

but there are discrepancies in each case in the descriptions,<br />

and it seems impossible to apply the names with certainty<br />

to these or any other plants. Even if they were applied to these<br />

forms, they could not be other than growth forms or forma.<br />

Two good photographs of the single, columnar type of variety<br />

setaceus supplied by Robert Runyon are reproduced as figures<br />

113 and 114 on page 105 of Britton and Rose’s The Cactaceae,<br />

Volume 3.<br />

This variety is not nearly so massive and spectacular as the<br />

previous variety, and is not as well adapted for the amateur’s<br />

growing. It is much less tolerant of moisture and shade. Its flesh<br />

is much more firm and lighter in color and does not present the<br />

soft, deep green, and the luxuriantly massive appearance of the<br />

variety hamatus. Its flowers are somewhat shorter than those<br />

of the other variety, but deeper in the shade of their yellow.<br />

Echinocactus sinuatus Dietrich<br />

[Hamatocactus sinuatus (Dietrich) Orcutt]<br />

“Lower Rio Grande Valley Barrel”<br />

Description Plate 23<br />

stems: Spherical, becoming conical and finally elongated<br />

ovate with a definitely pointed tip, and up to at least 12<br />

inches tall and 8 inches thick. It is usually single, but some<br />

old plants produce one or two branches at the base. The surface<br />

is very dark, dull green or even blue-green. There are 13<br />

ribs which are deep but compressed. They are 1 to 11/4 inches<br />

deep, undulating, acute and sharp between the areoles, and<br />

raised into indistinct tubercles which are somewhat rounded<br />

but only about 3/8 of an inch across at the areoles.<br />

areoles: Round or slightly oval and about 1/4 to 3/8 of an<br />

inch in longest measurement at first. The flower comes out of<br />

the upper end of the areole, leaving afterward a short groovelike<br />

extension of the areole forward which is usually about<br />

1/8 of an inch long and so broad as to be almost oval. This<br />

has some wool and several large, elongated glands in it. The<br />

areoles are about 1 inch apart.<br />

spines: There are 8 to 12 spreading radial spines which vary<br />

consistently in all specimens as follows: the lowermost 1 is<br />

3/8 to 11/8 inches long, round or slightly flattened, sometimes<br />

slightly hooked, red or purplish with a translucent yellow tip;<br />

the spines on each side of this one are similar in color but 5/8<br />

to 13/4 inches long, slightly flattened, and also sometimes<br />

slightly hooked; the 2 lateral radials on each side are flat,<br />

straight, 1 to 21/4 inches long, yellowish, and often slightly<br />

banded, becoming gray and rough with age; the upper 1 to<br />

3 radial spines are round, 11/4 to 21/8 inches long, straight, the<br />

most slender of the radials, reddish with yellow zones or all<br />

yellow, becoming gray with age. There are 4 centrals which<br />

are all flat and pubescent. The three upper ones spread upward<br />

in front of the upper radials and are 11/4 to 21/2 inches<br />

long, straight, yellowish with reddish zones, becoming gray<br />

with age. The lowermost central is approximately perpendicular<br />

to the stem, hooked, very flat and wide, 2 to 31/2 inches<br />

long and to 1/8 of an inch wide, reddish, becoming purplish-<br />

gray with age.<br />

flowers: 2 to 3 inches long and wide, clear lemon or greenish-<br />

yellow without red centers, and hardly fragrant. The ovary<br />

has a few rounded and fringed scales. The outer petals are<br />

short, greenish with reddish-brown midlines and greenish-<br />

yellow, fringed edges. Their tips are pointed, but they are


genus Echinocactus 83<br />

wide and irregular in shape otherwise. The inner petals are<br />

long, with entire edges, sharply pointed, all clear lemon yellow<br />

with narrow yellow bases. The filaments and anthers are<br />

all yellow. The style is yellow and the stigma has 8 to 10<br />

lemon-yellow lobes which are rough and about 3/8 of an inch<br />

long.<br />

fruits: Green, oval or egg-shaped, about 1 inch long and 3/8<br />

of an inch wide at the widest part near the center. There are<br />

about 6 to 12 small scales upon each fruit, and the old flower<br />

parts persist. When undisturbed these fruits remain on the<br />

plant for months, finally rotting, at which time they usually<br />

split open and the fermenting pulp containing the seeds spews<br />

out. The seeds are about 1/25 of an inch long, almost globular<br />

except compressed and prolonged beaklike at one end surrounding<br />

the small, sunken hilum. The surface is slightly<br />

shiny, with small but comparatively widely spaced pits all<br />

over it.<br />

Range. From near Brownsville in the lower Rio Grande Valley<br />

in a narrow strip north along the river to Eagle Pass, Texas.<br />

Spreading northeast of Eagle Pass beyond Brackettville to the<br />

Montell and Camp Wood, Texas, area and west along the Rio<br />

Grande to the mouth of the Devil’s River,<br />

Remarks. This cactus has been confused with other forms to a<br />

remarkable extent, considering its definite differences from its<br />

relatives. Its existence has even been denied by some more recent<br />

writers-one of the most strange omissions in all of cactus<br />

study.<br />

It was first described as Echinocactus sinuatus by Dietrich<br />

in 1851. Poselger, in 1853, referred to it as E. setispinus var.<br />

sinuatus, and perhaps also as E. setispinus var. robustus. Engelmann,<br />

a few years later, had no doubt about its standing as a<br />

separate species, and wrote an especially full description of it<br />

with the stated purpose to show this. Coulter understood the<br />

plant well, and followed Engelmann and Dietrich. Weber, however,<br />

in 1902, listed an E. longihamatus sinuatus, by which he<br />

must have meant to place this cactus as a variety of E. hamatacanthus.<br />

Then came Britton and Rose, who took a cue from<br />

Weber but then took one of their long steps and said that the<br />

plant was the same as their Ferocactus hamatacanthus. They<br />

even reproduced a photograph of E. sinuatus as their illustration<br />

of Ferocactus hamatacanthus, the picture in The Cactaceae, Volume<br />

3, page 144, figure 152. Marshall, following them, speaks<br />

of sinuatus as an extreme form of Hamatocactus hamatacanthus.<br />

Backeberg apparently did not have specimens to study first<br />

hand, but recognized from the previous literature that it was<br />

probably a separate species, and so listed it with a call for<br />

further study to be made of it.<br />

I had found mention by Schulz and others of the big fishhook<br />

barrel, E. hamatacanthus having been collected near<br />

Brownsville and “on the clay dunes near the Texas Coast.” I<br />

had never understood how that cactus, a native of the Big Bend,<br />

could turn up in such a different environment down in the lower<br />

Rio Grande Valley. When I found my first foot-high specimen<br />

of E. sinuatus near Zapata, Texas, I didn’t know what it was,<br />

but I knew at a glance that it wasn’t any form of E. setispinus<br />

or E. hamatacanthus. I doubt that anyone would confuse them<br />

who had actually seen all three together. I have since seen<br />

dozens of specimens all the way from the edge of the Gulf to<br />

the mouth of the Devil’s River, and they are consistently dis-<br />

tinct.<br />

As briefly as possible, E. sinuatus may be distinguished from<br />

E. setispinus by the following comparisons: E. sinuatus grows<br />

to 8 inches thick; has ribs 7/8 to 11/8 inches deep; 8 to 12 radial<br />

spines, some of them flattened; 4 centrals, all flattened and more<br />

or less pubescent; flowers which are never red-centered and<br />

hardly fragrant; fruits 7/8 to 1 inch long, oval or egg-shaped,<br />

and green until they rot; and seeds 1/25 of an inch long, shining,<br />

with very fine pits and small hila which have no collars. E.<br />

setispinus, on the other hand, grows to only 5 inches thick; has<br />

ribs 1/4 of an inch or less in depth; 10 to 19 radial spines, all<br />

round and bristle-like; only 1 central spine, which is round and<br />

smooth; flowers which are red-centered and very fragrant;<br />

fruits spherical, 3/8 to 3/4 of an inch across, and scarlet; seeds<br />

about 1/16 of an inch long, dull with closely set pits and large<br />

hila having wide collars.<br />

The following distinguish E. sinuatus from E. hamatacanthus:<br />

E. sinuatus grows to a maximum of about 12 by 8 inches; has<br />

13 compressed ribs the summits of which are 3/8 of an inch wide<br />

at the most; 8 to 12 radials, some of which are greatly flattened;<br />

4 centrals, all flattened and more or less pubescent; flowers<br />

which are lemon yellow with 8 to 10 stigma lobes; fruits 7/8 to<br />

1 inch long, oval to egg-shaped, and green; seeds 1/25 of an inch<br />

across. E. hamatacanthus, on the other hand, grows to at least<br />

24 by 12 inches; has 13 to 17 massively rounded ribs the summits<br />

of which are 3/4 of an inch or more across; 8 to 14 radial<br />

spines which are all round or very nearly so, and smooth; 4 to<br />

8 central spines which are smooth instead of pubescent, round<br />

or with only the lower one flattened on top; flowers which are<br />

entirely straw or pale yellow with or without red centers, with<br />

11 to 14 stigma lobes; fruits 1 to 2 inches long, egg-shaped or<br />

oblong, and brownish-red when ripe; and seeds about 1/16 of an<br />

inch long with dull surfaces.<br />

E. sinuatus is a beautiful species first encountered on the first<br />

solid ground back of the beach in deep south Texas. It may<br />

once have been common around and above Brownsville, but<br />

few undisturbed areas in which it can be seen remain there now.<br />

It may be found on a strip of territory a few miles wide all<br />

along the river up to Eagle Pass, being very common in some<br />

areas around Roma and Zapata. It is rare north of Zapata to<br />

north of Eagle Pass, but is occasionally seen. In some parts of<br />

the Anacacho Mountains and in the hills at the edge of the<br />

Edwards Plateau north and east of Brackettville, Texas, it is<br />

again rather common, although these northern specimens do not<br />

grow nearly as tall as they do farther south, preferring to remain<br />

spherical in shape. It is fairly easily found in various hilly


84 cacti of the southwest<br />

areas west of Eagle Pass to the mouth of the Devil’s River,<br />

which seems to be its westward limit, but it does not grow any<br />

distance north along that river. I have also seen numerous specimens<br />

collected in northern Mexico, some to almost as far south<br />

as Monterrey.<br />

Echinocactus hamatacanthus Muehlenpf.<br />

[Hamatocactus hamatacanthus (Muehlenpf.) Knuth]<br />

“Turk’s Head,” “Visnaga,” “Biznaga Costillona,” “Biznaga es<br />

Pinosa,” “Biznaga Ganchuda,” “Biznaga Limilla,” “Biznaga<br />

de Tuna”<br />

Description Plate 23<br />

stems: Large and heavy, hemispherical or almost spherical,<br />

becoming columnar and to a maximum of at least 2 feet tall<br />

and 1 foot in diameter. These stems are usually single, but<br />

occasionally become double or triple by branching, and when<br />

injured sometimes form larger clusters. The surface is dull<br />

green or gray-green. There are usually 13, but may occasionally<br />

be as many as 17 massive, broad ribs 11/2 to 2 inches high<br />

and wide. These ribs are divided between the areoles into very<br />

distinct, rounded tubercles 11/2 to 2 inches tall and the same<br />

width at their bases. The rounded tops of these tubercles are<br />

about 3/4 of an inch wide on mature plants. The areoles are<br />

on the tops of these massive tubercles.<br />

areoles: About 1 to 11/2 inches apart, oval to oblong, and<br />

about 3/8 to 1/2 inch long. There are very wide, felted grooves<br />

running inward and upward about 3/8 of an inch from the<br />

spinous parts of the areoles on mature plants, but these are<br />

not present on young plants which have not yet bloomed.<br />

There are very large, elongated glands in these grooves. The<br />

flower comes from the end of this groovelike extension of the<br />

areole.<br />

spines: There may be 8 to 14 radial spines, but usually are 10<br />

or more of them, as follows in appearance: the lowermost<br />

spine is 3/4 to 2 inches long and variegated reddish at first,<br />

turning gray-brown with age; the 3 laterals on each side are<br />

1 to 3 inches long and the same color or whitish. All of these<br />

are round or only very slightly flattened, straight, and radiating<br />

evenly. The 3 to 5 upper radials do not radiate, but<br />

stand spreading upward. They vary on different plants from<br />

as short as 3/4 of an inch to as much as 31/4 inches long, are<br />

round, or practically so, slender, straight, and reddish or<br />

gray, often variegated. There may be 4 to 8 round or somewhat<br />

flattened, smooth central spines in each areole. There<br />

is one extremely large lower central which stands out approximately<br />

perpendicular to the stem, but which usually<br />

curves and twists in any direction. It is hooked at the end,<br />

entirely round or else round below and flattened on its upper<br />

surface, 2 to at least 6 inches long, a heavy spine but some-<br />

what flexible due to its great length. It is yellow mottled with<br />

red or else all dull red at first, often indistinctly annulate,<br />

becoming gray-brown with age. There are always 3 more<br />

upper centrals which stand spreading upward. They are<br />

straight, round, relatively slender, reddish often mottled with<br />

yellow, later turning gray-brown, and 1 to 31/2 inches long.<br />

These 4 are all the centrals on young plants, but with age<br />

2 or 3 upper centrals are added which are shorter and more<br />

slender, standing erect just back of the previous upper cen-<br />

trals.<br />

flowers: Entirely straw to yellow or the outer parts thus<br />

with red centers, 21/4 to 4 inches tall, 23/4 to 3 inches wide,<br />

and very fragrant. The ovary and tube have very many<br />

small triangular scales on them; Engelmann, having counted<br />

them, said there are 30 to 60 of these. Their centers are reddish<br />

or brownish, while their edges are greenish-yellow,<br />

crinkled, and may or may not have a few twisted cilia on<br />

them. The very many outer perianth segments—Engelmann<br />

says there are 55 to 80 of them—range all the way from<br />

short, scalelike ones to full-length, oblong ones. They all have<br />

reddish midlines, the outer parts being greenish and the edges<br />

being yellowish. There are about 30 inner petals which are<br />

long, wide, and pointed, the edges entire or often toothed<br />

irregularly. These inner petals may be all yellow or yellow<br />

with red bases. The filaments will be yellow in all yellow<br />

flowers, but reddish in those with red centers. The anthers<br />

are yellow. The style is yellow and longer than the stamens.<br />

There may be 11 to 14 lobes in the stigma, which are about<br />

1/4 of an inch long, yellow, rough, and usually much curved<br />

and twisted.<br />

fruits: Egg-shaped to oblong, 1 to 2 inches long, not including<br />

the persistent perianth. There are 30 or 40 scales on<br />

the fruit, each greenish edged in white. The fruit remains<br />

green all summer and fall and then during the winter it ripens,<br />

becoming at that time a brownish-red color. The seeds are<br />

practically round, about 1/16 of an inch long, black, with the<br />

surface pitted.<br />

Range. Along and never many miles north of the Rio Grande<br />

from the mouth of the Devil’s River all the way to El Paso,<br />

Texas. Occurring very rarely west of El Paso for perhaps 50<br />

miles along the southern border of New Mexico. Growing very<br />

abundantly in Mexico.<br />

Remarks. Echinocactus hamatacanthus is the second largest,<br />

second most splendid member of this genus in the United States.<br />

It has often been called E. longihamatus. Although this latter<br />

name may have been coined by Galeotti first, it was used without<br />

description and so most have agreed that Muehlenpfordt’s<br />

hamatacanthus has precedence.<br />

This species may be distinguished from its already described<br />

relatives by its great rounded ribs composed of massive tubercles<br />

swelling around each areole. Both E. setispinus and E. sinuatus<br />

have sharp ribs without these large rounded tubercles. The


genus Echinocactus 85<br />

same characteristic distinguishes it from the only other cactus<br />

in our area presenting such a massive size and having hooked<br />

spines—Echinocactus wislizeni—since that large barrel cactus<br />

has very large but uninterrupted, sharp ribs.<br />

Engelmann described three varieties of this cactus. However,<br />

since they overlap and there are many intermediates which one<br />

cannot assign to any one of them with any certainty, they do<br />

not stand distinct as do those of E. setispinus. They are no doubt<br />

just the extreme variations in spine characters of the species.<br />

Variety crassispinus Eng. has its central spines when typical relatively<br />

heavy and the most flattened of this species. It is the<br />

form which Engelmann first called Echinocactus flexispinus. He<br />

and other authorities consider it to be found only in Chihuahua,<br />

but Coulter assigned some Texas specimens to this variety. Variety<br />

gracilispinus Eng. is the most common form in Texas and<br />

Mexico, having its centrals comparatively slender and the hooked<br />

one only slightly flattened. Variety brevispinus Eng. is the form<br />

having the central spines shorter than the others, hardly if any<br />

longer than the radials, and all spines round. Almost all young<br />

specimens could pass for this variety, and a few mature ones also.<br />

Coulter states that this is the form found west of El Paso in New<br />

Mexico.<br />

One often finds in the literature the name Brittonia davisii<br />

applied to this species and credited to Dr. A. D. Houghton. Marshall,<br />

in his Cactaceae made this Hamatocactus hamatacanthus<br />

var. davisii. However, in an article in the Cactus and Succulent<br />

Journal of America in 1944, Marshall relates that shortly before<br />

his death, Houghton wrote him that he had never published the<br />

name at all, and so in this article Marshall agrees with Borg in<br />

dropping the name entirely.<br />

E. hamatacanthus is a beautiful cactus, becoming one of the<br />

largest in the Southwest. Everything about it grows in grand<br />

proportions. This is not a cactus for dish gardens or window<br />

ledges, and it should not be dwarfed in a cramped pot. It needs<br />

space, in return for which it will grow slowly into a massive<br />

barrel with perhaps the longest spines of any in our area. It is<br />

not so easy to grow as its relatives from farther east in Texas,<br />

being more liable to rot if given too much water. It is more<br />

completely a desert species, and one must remember this. Neither<br />

is it as resistant to freezing as our other cacti, and it can only at<br />

great risk be left unprotected during the winter anywhere north<br />

of its range.<br />

This species has been shunted about between genera more than<br />

its other close relatives. Of course, it was first described as an<br />

Echinocactus. But when Britton and Rosa broke up that genus,<br />

they did not assign this species to Hamatocactus with the others,<br />

but instead to the genus Ferocactus. This was because they concluded<br />

that this species had enough scales on its ovary to belong<br />

there. It was Knuth who finally transferred it to Hamatocactus,<br />

where it seems most obvious that it would have to remain unless<br />

those two microgenera are actually so close as to be merely<br />

subdivisions of the one actual genus, Echinocactus. But more<br />

recently Buxbaum has maintained that it is really a Ferocactus,<br />

while Hester shows how impossible it is to keep any of these<br />

microgenera separate by asserting that all of the members of<br />

Hamatocactus, including this one, should be included in Thelo-<br />

cactus.<br />

Echinocactus bicolor var. schottii Eng.<br />

[Thelocactus bicolor var. schottii (Eng.) B. & R.]<br />

“Glory of Texas”<br />

Description Plate 23<br />

stems: Egg-shaped or conical to almost columnar, sometimes<br />

to 10 inches tall and 5 inches in diameter, but usually smaller.<br />

These stems are usually single, but very old plants sometimes<br />

form small clusters of 3 or 4 heads by branching from the<br />

base. There are 8 broad, flat ribs which are composed of wartlike<br />

tubercles to about 1/2 inch high from almost perfectly<br />

square bases up to 3/4 of an inch wide.<br />

areoles: Oval or nearly round with yellow wool at first,<br />

later egg-shaped on immature plants. On adult tubercles the<br />

floral part of the areole forms a short groove about 1/8 to 3/16<br />

of an inch long and often so wide as to make the areole as a<br />

whole obovate in shape. In old plants glands are often visible<br />

in the areoles.<br />

spines: There are 12 to 18 radial spines on each areole. The<br />

upper 1 to 4 of these radials are erect, straight, flattened, and<br />

3/4 to 21/4 inches long. They are yellow when young, becoming<br />

gray with age. The lateral and lower radial spines are all<br />

round, 1/2 to 11/4 inches long, and varicolored, the bases of<br />

them being gray, the middle zones dark red, and the ends<br />

yellowish. The lower radials often recurve a little back toward<br />

the plant. There are 3 or 4 straight central spines. The<br />

uppermost stands erect just in front of the upper radials. It<br />

is 1 to 31/2 inches long, very flat and broad—often 1/8 of an<br />

inch wide—and flexible. Standing erect beside this one are 1<br />

or 2 other centrals, flat on some plants but round on others,<br />

and not quite so long. These erect centrals are yellow at first,<br />

becoming gray. The lower central stands perpendicular to the<br />

plant or is turned downward. It is perfectly round or oval,<br />

stout and rigid, and 3/4 to 23/4 inches long. It is gray, red, and<br />

yellow like the lower radials at first, becoming all gray when<br />

very old.<br />

flowers: 2 to 3 inches long, 3 to 4 inches across, opening<br />

widely with petals usually recurving backward. They are<br />

brilliant fuchsia with scarlet throats and a shining, satiny surface.<br />

The sepals vary from short, rounded scales on the ovary<br />

wall to more elongated, oblong sepals above. All have greenish<br />

midlines and whitish, fringed edges. The inner petals are<br />

oblong from narrow bases, about 3/8 of an inch wide at the<br />

widest point, the margins entire but crinkled, the tips pointed<br />

and recurving. Their bases are bright scarlet, the upper three-


86 cacti of the southwest<br />

fourths of each one a bright, satiny fuchsia. The filaments are<br />

bright scarlet, matching the petal bases. The anthers are yellow.<br />

The style is pink and a little longer than the stamens.<br />

The stigma has 8 to 11 rough, blunt lobes, light rose to brownish-pink<br />

in color.<br />

fruits: About 3/8 to 1/2 inch long, becoming dry and splitting<br />

open by means of an irregular basal pore. The seeds are about<br />

1/16 of an inch long, almost globular, with very large hila.<br />

Range. Occurring in Texas in two widely separated areas, one<br />

near and up to about 20 miles north of the Rio Grande in Starr<br />

County, and the other around Lajitas and above Candelaria,<br />

Texas, in Brewster County in the Big Bend. No record of collections<br />

between these widely separated points comes to light.<br />

Remarks: This is primarily a Mexican species. We are very fortunate<br />

that it steps across our border in two widely separated<br />

places, where we can claim it as the glory of Texas. The name is<br />

well deserved. The bright-colored, variegated spines are attractive,<br />

and when the cactus blooms, it presents undoubtedly the<br />

brightest and most exotic flower of any in our area. The fuchsia<br />

petals with their scarlet bases are satiny in texture and colorful<br />

beyond description. It is unfortunate that they are so sensitive<br />

that they will only open in the full heat of the southwestern<br />

summer afternoon and close again permanently as the sun begins<br />

to fall. They are so sensitive that they start to close visibly when<br />

a cloud temporarily covers the sun. I have found that it requires<br />

fast action to photograph them, since they close in a short time<br />

when taken out of the greenhouse and placed outside where the<br />

temperature is 10 degrees cooler, even when the sun is still full<br />

upon them. Because of this sensitivity few people have seen these<br />

exquisite flowers each one of which is open only three or four<br />

hours of only one afternoon.<br />

It is quite common to consider the plants from the two different<br />

areas in Texas as different forms, those from Starr County<br />

usually being called E. bicolor and those from the Big Bend E.<br />

bicolor var. schottii or E. bicolor var. tricolor. I have examined<br />

many specimens from both areas, and although the southern<br />

ones are smaller with correspondingly shorter and less colorful<br />

spines, there is no essential difference between them. I am convinced<br />

that they must all be considered the same form, the west-<br />

ern one only being more robust in every respect.<br />

The species was first described from Mexican specimens by<br />

Galeotti in 1848, under the name Echinocactus bicolor. When<br />

Engelmann first described the Texas plants, he found them<br />

enough different from the original description that he erected a<br />

new variety, calling the Texas plants E. bicolor var. schottii. He<br />

stated that the Texas specimens differed from the species by<br />

having more radials and having the upper centrals greatly flat-<br />

tened and the longest spines on the plants.<br />

I have examined many plants from both Texas and the area<br />

of the type in Mexico, and find Engelmann’s distinction between<br />

the Texas specimens and the type of the species partly right and<br />

partly in error. There is no consistent difference in number of<br />

radials between the two. On Texas specimens I have found 12<br />

to 18 radials, while on Mexican specimens I have found 10 to<br />

18. Engelmann must have seen only examples of the lower<br />

number in his Mexican plants. There is, however, a great difference<br />

in the length of the radials between the two types. The<br />

Mexican form shows lower radials 1/4 to 1/2 inch long, laterals<br />

5/8 to 3/4 of an inch, and the upper radials 1/2 to 3/4 of an inch<br />

long, while the same spines on Texas plants run lower and lateral<br />

radials 1/2 to 11/4 inches and the uppers 3/4 to 21/4 inches<br />

long.<br />

In the matter of central spines, Engelmann was quite right,<br />

and this is the primary distinction overlooked by many. The<br />

Texas plants have a round or oval lower central which is rather<br />

stout and 3/4 to 23/4 inches long. Then they have 2 or 3 upper<br />

centrals which are erect, the uppermost one being very flat, 1/16<br />

to 1/8 of an inch wide, flexible, and 1 to 31/2 inches long—always<br />

longer than the lower central on a mature plant. The<br />

Mexican plants of the type area, on the other hand, have this<br />

uppermost central round or oval like the lower, very stout, and<br />

awl-shaped, not flattened or flexible, and only 3/4 to 11/8 inches<br />

long—never longer than the lower central. In general, the Mexican<br />

plant has short, very stout, and rigid spines, while the Texas<br />

form is more or less covered by its longer, more slender, more<br />

flattened, flexible spines.<br />

The difference between the two forms is certainly not great,<br />

but it seems enough that they can be recognized once they are<br />

understood. No doubt they are not separate species, but it does<br />

seem that Engelmann’s variety is valid, even though Britton and<br />

Rose, as well as some others, have ignored it. I find no record<br />

of the typical E. bicolor which occurs in Mexico or any of the<br />

several other varieties of it having been found north of the Rio<br />

Grande. Variety schottii, however, does grow quite some dis-<br />

tance down into Mexico.<br />

E. bicolor var. tricolor is a name proposed by Schumann for<br />

the specimens with the most brilliant red spines. Such extremely<br />

red-spined plants show no other differences from the other Texas<br />

specimens, however, and are found growing right among the less<br />

brightly colored specimens in the Big Bend, so this does not seem<br />

a valid basis for a variety, and the name probably should be<br />

dropped. Backeberg described a variety texensis. After observation<br />

of many specimens, it seems to me that plants fitting his<br />

description cannot be set off from the variety schottii, but intergrade<br />

with the others and must be included within that variety.<br />

This species was regarded as an Echinocactus as long as that<br />

genus was recognized in its original sense. Later Schumann<br />

erected a subgenus Thelocactus for it which Britton and Rose<br />

elevated to a separate genus. This separation was made because<br />

of the few scales on the ovary and its lack of wool, and because<br />

of a misinterpretation of the areole. Britton and Rose thought<br />

the flower originated in a separate floral areole separated by a<br />

groove from the spine areole. It has since been shown that this<br />

whole structure is one monomorphic areole and that there is no<br />

distinction between the areoles of this plant and those of the


genus Echinocactus 87<br />

most typical Echinocacti. Unless only those species are left in<br />

the genus Echinocactus which have wool on the ovary, this<br />

species must be included along with the rest.<br />

Very large and beautiful specimens of this cactus 8 and even<br />

10 inches tall are now being brought out of the more inaccessible<br />

mountains of the Big Bend by resourceful commercial gatherers.<br />

They are much larger and have much longer spines than the<br />

earlier writers ever saw, but are otherwise the same. Unfortunately<br />

this is a very fastidious desert cactus, and these large old<br />

specimens grown to such splendid size because of the most perfect<br />

desert environment in these very arid mountains will not<br />

live and grow further in the less favorable environment of a<br />

pot or garden. It is a questionable practice of the dealers to offer<br />

for sale these huge old plants which cannot live long. They thrill<br />

the collector, but must disappoint him when they soon die. Even<br />

the small specimens, which are sold widely, are impracticable<br />

for all but the collector who can duplicate the desert conditions<br />

for them. There is a saying that they will not live over 3 years<br />

in a pot, and in most cases I believe this is longer than can be<br />

expected. I have disproved the saying by keeping one in a pot<br />

4 years now, but this must be credited to extremely alkaline,<br />

limestone soil and to its position in the full Texas sun in a greenhouse<br />

where the temperature climbs to between 110 and 120<br />

degrees every day of the summer. Treated in this seemingly inhumane<br />

way, the cactus thrives and shows its appreciation for<br />

the heat and sunshine by blooming all summer. Its earliest<br />

flower has opened on April 12, and its latest on September 16.<br />

It seems cruel to subject such a delicately flowering plant to<br />

this kind of treatment, but if it is provided, this plant will show<br />

all of its glory.<br />

Echinocactus flavidispinus (Backbg.)<br />

[Thelocactus flavidispinus Backbg.]<br />

Description Plate 23<br />

stems: Hemispherical at first, becoming columnar and sometimes<br />

branching at the base or, if injured, at the point of<br />

injury. Becoming at least 4 inches tall and 3 inches in diameter.<br />

There are 13 ribs composed of rows of conical tubercles<br />

to about 3/8 of an inch high. These are distinct in young specimens,<br />

but become somewhat confluent on older plants. The<br />

color of the surface is light green or even yellowish-green.<br />

areoles: Oval at first, but after flowering, ovate, the upper<br />

part prolonged into a short groove containing glands.<br />

spines: There are 14 to 20 radial spines which are recurved<br />

against the plant. They are all yellow at first, or yellow<br />

streaked with red. Later the lower and lateral ones remain<br />

round, only 1/4 to 3/4 of an inch long, and become bright red<br />

in their middle zones with yellow tips, while the uppermost<br />

becomes flattened, cream to gray in color, and to 1 inch long.<br />

Juvenile plants possess only the radials, and these are all round<br />

at first. When the plant is a little older there appears 1 strong,<br />

round central which stands perpendicular to the stem or<br />

turned downward; it is pubescent, all yellow or yellow at the<br />

base and tip with bright red in the middle, and 3/8 to 7/8 of an<br />

inch long. Mature plants add 3 more centrals turned upward:<br />

2 of them are like the radials in all respects, while the uppermost<br />

becomes 1 to 11/2 inches long, flattened, more or less<br />

curved toward the plant, and all yellow, fading to gray with<br />

age.<br />

flowers: About 11/2 inches tall and 3 to 4 inches in diameter<br />

when fully open. The scales on the ovary and the outer perianth<br />

segments have brownish-green midlines shading to whitish,<br />

entire edges and scarlet bases. The inner segments have<br />

very narrow scarlet bases widening to bright rose-pink or<br />

fuchsia upper parts, which have entire edges, are very sharp-<br />

pointed, and do not recurve. The filaments and anthers are<br />

yellow. The style is yellow or pink, and there are 11 stigma<br />

lobes which are scarlet at their bases fading to yellowish at<br />

their tips.<br />

fruits: Not seen.<br />

Range. Known only from near Marathon, Texas, particularly<br />

in the Pena Blanca Mountains in the upper part of the Big<br />

Bend region.<br />

Remarks. This cactus was first described as Thelocactus bicolor<br />

var. flavidispinus by Backeberg in 1941. Later Backeberg elevated<br />

it to a separate species. He apparently had before him<br />

only the younger plants, since he described the species as having<br />

only 1 central. Examination of many specimens in their native<br />

habitat shows that the juveniles have no central, the younger<br />

adults for a number of years after flowering have only one, and<br />

old plants produce 4 centrals on all areoles, as described above.<br />

This cactus is easily distinguished from Echinocactus bicolor<br />

var. schottii or the typical Echinocactus bicolor, neither of<br />

which grows anywhere near the very limited range of this form.<br />

The 8 ribs of E. bicolor and 13 of E. flavidispinus are an unvarying<br />

difference, and the shorter, more yellow, and all round<br />

spines—except for the one upper central on old plants—characterize<br />

E. flavidispinus.<br />

The flowers of E. flavidispinus are basically similar to those<br />

of E. bicolor var. schottii, but differ in details which are obvious<br />

if the two are seen blooming together. This flower is a much<br />

lighter rose color, its petals are much more pointed and do not<br />

open so widely. The filaments are not red, but yellow, and the<br />

stigma lobes are more slender and less colorful.<br />

Backeberg did not describe the fruits of this species, and in<br />

examining hundreds of specimens in their native locations and<br />

in growing examples for several years I have seen many flowers<br />

but no fruits. I do not know what if any significance there is<br />

to this. The plants in their natural situations suffer very often<br />

from some sort of injury to their growing tips, and it may be<br />

that some insect rapidly devours the young fruits. At any rate,<br />

they do not seem to have been observed up to this time.


88 cacti of the southwest<br />

In much of its range, I found that fully one-half of the plants<br />

had suffered the injury mentioned above. Their growing tips<br />

were partly destroyed, perhaps by an insect or by a severe<br />

freeze, and they were reproducing at this point 3 or 4 small,<br />

spherical branches with short, yellow, juvenile spines upon<br />

them. Several of these, when grown in my greenhouse for two<br />

years, have now put out the one reddish central heralding maturity,<br />

but none of them has yet produced the other 3 centrals<br />

obvious on the older, original stem.<br />

The description given for Thelocactus wagnerianus Berger, of<br />

eastern Mexico, seems very close to this plant. Backeberg discussed<br />

the relation of the two in his 1941 article, but did not<br />

conclude them to be synonymous. However, he did not realize<br />

that the Texas plant has 3 or 4 centrals when fully mature, as<br />

does the Mexican one. The relationship between the two needs<br />

study, but I have not been privileged to see the Mexican cactus,<br />

and so cannot offer anything on it.<br />

Echinocactus intertextus Eng.<br />

[Echinomastus intertextus (Eng.) B. & R.]<br />

“The Early Bloomer,” “White-Flowered Visnagita”<br />

Description Plate 24<br />

stems: Always occurring individually and without branching.<br />

They are spherical at first, becoming egg-shaped or conical, and<br />

when very old, short, thick columns. The maximum size seems<br />

to be 4 to 5 inches tall and 31/2 to 4 inches in diameter. There<br />

are almost always 13 ribs, although 12 and 14 are said to<br />

have occurred. These are distinct and broad but low, 5/8 to 1<br />

inch wide and only 1/4 to 3/8 of an inch deep on a large plant.<br />

The tubercles making up these ribs are distinct, being almost<br />

entirely separated by deep cross-grooves. They are conical at<br />

first, but on the older sides of the stem the deep cross-grooves<br />

separating them make the bases square. The summits of the<br />

tubercles are prolonged below the areole into a sharply peaked,<br />

chinlike ridge running back so far as to almost overhang the<br />

next lower tubercle, and at the cross-groove between them this<br />

ridge terminates suddenly with a straight drop into the pitlike<br />

depression thus formed for the axil.<br />

areoles: The young areoles are rather large, slightly oval,<br />

very woolly, becoming bare when old. The mature areole becomes<br />

elongated by a woolly groovelike extension which runs<br />

inward and upward from the spinous portion all the way to<br />

the base of the tubercle, where it meets the cross-groove between<br />

the tubercles. The flower is produced from the end of<br />

this extended areole in the axil of the tubercle. There is usually<br />

left after flowering and fruiting a tuft of longer, yellow-<br />

ish-white wool in this depressed axil.<br />

spines: The spines are all round with slightly enlarged bases.<br />

They are all dull gray or yellowish at the bases, with the up-<br />

per one-half or so darkening into purplish or reddish-brown<br />

tips. There are 16 to 27 radial spines which radiate evenly,<br />

recurve slightly and lie tightly against the plant surface on<br />

one form and spread outward on another. The upper ones are<br />

much the weakest spines, being almost bristle-like and 3/16<br />

to 5/8 of an inch long. The lateral radials are heavier and<br />

longer, being up to 7/8 of an inch long. The 3 to 5 centrals are<br />

similar to the radials but a little heavier and slightly darker<br />

in color. 2 to 4 of these stand erect in front of and against the<br />

upper radials in one form or spread upward in the other form.<br />

They are 1/2 to 7/8 of an inch long. The lowermost central<br />

stands straight out from the center of the areole. On one form<br />

it is heavy, but very short, being only 1/8 to 3/16 of an inch<br />

long. On the other form it is not so heavy, but is 1/4 to 5/8<br />

of an inch long. Juvenile plants have 16 to 18 strictly radiating<br />

radial spines and no centrals. Then 1 central appears and<br />

the others follow.<br />

flowers: Salmon to white in color, 3/4 to 1 inch long and 1/2<br />

to 1 inch in diameter, the state of the plant determining how<br />

widely they open. On plants growing with a minimum of<br />

water the spines will prevent the petals from opening out,<br />

but on plants well expanded with water the spines will allow<br />

them to open widely. There are about half a dozen small<br />

scales on the ovary. The outer perianth segments are from<br />

very short to about 3/4 of an inch long and 3/16 of an inch<br />

wide. They have pink midlines with very pale pink edges,<br />

and are pointed, with the edges entire or sometimes somewhat<br />

toothed and ragged. The inner petals are about 3/4 of an<br />

inch long and about 1/8 to 3/16 of an inch wide, whitish at<br />

their bases and with a very pale pink midline shading to<br />

white edges. They are pointed, often with a tiny, soft spine at<br />

the apex, and their edges are usually irregular. The filaments<br />

are greenish and the anthers yellowish. The style is greenish,<br />

but the stigma has 6 to 12 bright pink to brilliant purple-red<br />

lobes.<br />

fruits: Small, 3/8 to 1/2 inch in diameter and globular to<br />

somewhat oblong, with the old perianth persistent upon it.<br />

It becomes dry and brown without coloring, and then it<br />

splits open all around its base and the upper part falls off,<br />

releasing the seeds. It has a few scales on its surface. The<br />

seeds are about 1/16 of an inch or slightly larger, black and<br />

shining, with a rough surface. They are nearly kidney-shaped<br />

and have large hila.<br />

Range. From the Texas Big Bend and lower Davis Mountain<br />

region west to El Paso and the Franklin Mountains. They occur<br />

from there west in a narrow strip along the lower border of<br />

New Mexico into the southeastern corner of Arizona, and also<br />

in adjacent Chihuahua and Sonora, Mexico.<br />

Remarks. The white-flowered visnagita is an interesting little<br />

cactus venturing into our area from Mexico. Nowhere does it<br />

manage to survive very far within our territory. Its deepest<br />

penetration is about 100 miles above the Rio Grande around


genus Echinocactus 89<br />

Alpine and Fort Davis, Texas, where one form has been fairly<br />

common in the past, but is now much more rarely seen, due no<br />

doubt to the activities of collectors and dealers who have long<br />

made Alpine their headquarters. Lower in the Big Bend. occasional<br />

plants may still be found, but it is nowhere abundant.<br />

It may once have existed much farther east in Texas than this,<br />

as early reports mention its collection near the mouth of the<br />

Pecos River, but no specimens have been reported east of Alpine<br />

and the Big Bend National Park for many years. This form is<br />

occasionally found along the very southern edge of New Mexico<br />

and into southeastern Arizona. Another form of the same species<br />

ventures up through the Franklin and Organ mountains as far<br />

into New Mexico as Rincon and Lake Valley.<br />

The cactus is well named the early bloomer. So far as I have<br />

observed it is the first cactus in its locale to bloom each year. I<br />

have had plants bloom in my garden in San Antonio as early as<br />

February 13.<br />

Concerning the growing of E. intertextus in cultivation I<br />

must be discouraging. It is an attractive small cactus, especially<br />

for its very early flowers which are unusual in that, while the<br />

petals are very pale in color, the brilliance is in the bright colored<br />

stigma. But this is one of our most particular cacti in regard<br />

to its growing requirements. It cannot tolerate water. It<br />

must be kept almost dry at all times or it will rot immediately.<br />

Only when given soil which will not hold moisture, and much<br />

hot sun will it grow and bloom. It should be remembered that<br />

this plant does not grow in partial shade like most small cacti,<br />

but is found unprotected in the full sun and heat.<br />

Echinocactus intertextus was first described by Engelmann,<br />

and there has been little disagreement about it until recently.<br />

The only instance of confusion seems to have been when Coulter<br />

apparently mistook an Arizona specimen of it for a Cereus and<br />

named it Cereus pectinatus centralis. Schumann then called it<br />

Echinocereus pectinatus centralis. This is of course absurd, as<br />

the plant has no real similarity to a Cereus.<br />

Britton and Rose naturally took this species out of the genus<br />

Echinocactus along with most others, and based their new<br />

genus Echinomastus on it. While this name has been widely<br />

used, there has never been much certainty about this proposed<br />

genus among taxonomists. It has been combined with various<br />

other genera. Because of the long, groovelike extension of the<br />

areole and the tubercles more separated from one another than<br />

most of those in the Echinocacti, some have played with the<br />

idea of calling this cactus a Coryphantha, but no one has actually<br />

done so officially. In his book, Arizona Cacti, Benson returned<br />

the genus Echinomastus to Echinocactus, but he is now proposing<br />

to merge it with the genus Neolloydia. Thus the species<br />

seems off on the rounds of the microgenera like so many other<br />

species, pausing next as Neolloydia intertextus, unless Echinocactus<br />

in its original sense, the only meaningful genus designa-<br />

tion we have had for all these plants is maintained.<br />

Engelmann’s description of this species included only the typical<br />

form of it and did not include the other variety which he<br />

himself originated. The best taxonomical practice would require<br />

us to designate this typical form a variety also and to broaden<br />

the species description to include both varieties of the species,<br />

which has been done here.<br />

Echinocactus intertextus var. intertextus (Eng.)<br />

Description Plate 24<br />

stems: As the species, except that it grows to only 5 inches<br />

tall and that the ribs are broader, being 3/4 to 1 inch wide.<br />

areoles: As the species.<br />

spines: As the species, except that the radial spines always<br />

radiate evenly, recurving and lying tightly against the plant<br />

surface, while the upper radials are to only 1/2 inch long, the<br />

lateral radials to only 3/4 of an inch long, and the lower<br />

radials to only 3/8 of an inch. Also, the upper centrals stand<br />

erect in front of and against the upper radials, while the<br />

lower, porrect central is very heavy and only 1/8 to 3/16 of an<br />

inch long.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. From the Texas Big Bend and lower Davis Mountains<br />

west past El Paso in a narrow strip along the lower border of<br />

New Mexico into Arizona.<br />

Remarks. This is the typical form of the species, and it occurs<br />

over a wide range. Engelmann’s original description of the<br />

species was restricted to this form, and most books and articles<br />

refer to this only when they use the species name.<br />

This variety appears as a single small hemisphere or short<br />

column made of distinct ribs composed of curiously shaped<br />

tubercles as described under the species. Its areoles are close<br />

together, and the plant is encased in its purplish-red spines, all<br />

of which lie flat against the surface, except for the one very<br />

short and stout lower central which stands straight out, but<br />

which is so short as to be almost unnoticed. The spines meet and<br />

interlock over the top of the plant and completely enclose the<br />

growing tip, which is very woolly when active. The flowers<br />

come out of this woolly summit and often have a terrible time<br />

opening because of the interlocking spines.<br />

The effect of the environment upon the form of the flower<br />

in this variety was dramatically impressed upon me by one<br />

plant which I collected just at blooming time. It had apparently<br />

had a very dry winter, and when it bloomed for me the plant<br />

was still quite shrunken due to low water content. It bloomed<br />

profusely in spite of this, and had as many as 6 flowers at a<br />

time on the rather flat summit of the stem. But the spines were<br />

so many and so tightly interlocking around them that none of<br />

these flowers could open properly. Their petals managed to<br />

stand straight up, but that was all, and one had to look directly<br />

down into the tube about 3/8 of an inch across which they


90 cacti of the southwest<br />

formed to see the bright purplish stigma. This same plant flour-<br />

ished in cultivation, however, and exactly a year later it<br />

bloomed again in my greenhouse. But now it was nicely filled<br />

out and plump, and the swelling of the stem had pushed the<br />

spines over the summit much apart, so the flowers could now<br />

open widely. I was amazed to see the petals open all the way<br />

back above the spines until the flowers were rotate and at least<br />

1 inch across.<br />

Echinocactus intertextus var. dasyacanthus Eng.<br />

[Echinomastus intertextus var. dasyacanthus (Eng.) Backbg.]<br />

Description Plate 24<br />

stems: As the species, except that they grow to 6 inches tall<br />

and that the ribs are somewhat narrower, being only 5/8 to 3/4<br />

of an inch wide.<br />

areoles: As the species.<br />

spines: As the species, except that all spines are longer and<br />

more spreading than on the typical variety. The upper radials<br />

are 1/2 to 5/8 of an inch long and very slender, the lateral<br />

radials 5/8 to 7/8 of an inch long and heavier, the lower ones<br />

1/2 to 5/8 of an inch long and rather heavy. The upper centrals<br />

spread upward instead of lying appressed against the radials<br />

as in the other form and are 3/4 to 7/8 of an inch long. The<br />

lower central stands practically perpendicular to the stem, as<br />

in the other form, but is no heavier than the upper centrals<br />

and is 1/4 to 5/8 of an inch long.<br />

flowers: As the species.<br />

fruits: Apparently identical with the species.<br />

Range. A narrow belt of mountainous territory from near Lake<br />

Valley and Rincon, New Mexico, south at least to El Paso and<br />

probably into Mexico. Most common in the Franklin and Organ<br />

mountains near Las Cruces and El Paso. Apparently not coming<br />

into Texas beyond the extent of the foothills of the Franklin<br />

Mountains.<br />

Remarks. This is merely a variety of the species. Engelmann<br />

considered it that when he first named it, and it was so considered<br />

until Britton and Rose elevated it to species rank, calling<br />

it Echinomastus dasyacanthus. From that time on collectors<br />

have been constantly struggling to distinguish what they were<br />

led to believe were two nicely distinct species. It has been a great<br />

contribution of Backeberg to return this form to varietal status.<br />

Collectors should not be surprised if they have trouble telling<br />

variety dasyacanthus from variety intertextus. The only real differences<br />

are matters of maximum stem size and spine character.<br />

The most obvious difference is the length and character of the<br />

lowermost, porrect central spine. Near El Paso and in New<br />

Mexico just north of El Paso the plants collected in the Franklin<br />

and Organ mountain foothills all have the longer, more spreading<br />

spines, with the lower central 1/4 to 5/8 of an inch long. This<br />

contrasts nicely with the typical variety intertextus of the Texas<br />

Big Bend and the very lower edge of New Mexico with its generally<br />

shorter, sharply appressed spines and its lower central very<br />

heavy but only 1/8 to 3/16 of an inch long. But there are definitely<br />

intermediates. They come particularly from the area of Presidio<br />

and Candelaria, Texas. As adults some of these are hard to assign<br />

to either variety with certainty.<br />

It had seemed to me that even a separate varietal rank for a<br />

form merging so closely into the typical was hard to justify,<br />

and I might have been tempted to merge them more closely if I<br />

had not been shown juveniles of the two forms. These were grown<br />

in quantity from seed by Mr. Clark Champie of Anthony, New<br />

Mexico-Texas. When 1 inch in diameter both forms were almost<br />

hemispherical to slightly conical in shape. But the character of<br />

the spines at this stage is completely different one from the<br />

other. Variety intertextus has all of its 16 to 18 radials heavy,<br />

3/16 of an inch long, opaque purplish-gray and appressed flat<br />

against its surface. One does not feel the spines in handling this<br />

cactus, they lie so flat upon the plant. At the same age and size<br />

the spines of variety dasyacanthus are only about half as heavy,<br />

nearly twice as long, translucent yellowish to reddish-brown, and<br />

stand spreading well out at all angles from the plant so that it<br />

is, indeed, a very dangerous little ball to handle. The difference<br />

between them at this age, before they get any centrals at all, is<br />

much more striking than it is when they are adults, and no one<br />

could look at the flats of seedlings at Mr. Champie’s establishment—hundreds<br />

of variety intertextus consistently the same on<br />

one hand and on the other hand hundreds of variety dasyacanthus<br />

so different—without realizing that here is a difference<br />

which must be recognized, even if it is within the one species.<br />

Echinocactus erectocentrus var. pallidus (Backbg.)<br />

[Echinomastus pallidus Backbg. nom. prov.]<br />

Description Plate 24<br />

stems: Single until very old, then occasionally producing<br />

several short branches just above the ground. The stem is globose<br />

at first, becoming oblong or short columnar and growing<br />

to a maximum of 6 inches tall and 4 inches in diameter when<br />

old. It has 13 spiraling ribs when young, this number increasing<br />

by branching of the ribs to a maximum of at least 21 on<br />

large stems. These ribs are up to 5/8 of an inch deep and composed<br />

of definite tubercles which, however, vary greatly in<br />

shape on the same plant. Some are compressed from side to<br />

side and are only 1/4 of an inch wide, while others are up to<br />

1/2 inch broad at their bases. The tubercle is prolonged as a<br />

short, sloping ridge running downward from the areole. Often<br />

rown -> brown


genus Echinocactus 91<br />

this ridge rises a little to form a second shorter, chinlike projection<br />

behind the main tubercle and then ends abruptly by<br />

falling to a definite though narrow cross-furrow between the<br />

tubercles.<br />

areoles: Elongated and very woolly at first, then becoming<br />

nearly round and almost bare, except for a woolly groove<br />

which extends inward and upward from the spinous portion<br />

to the floral portion of the areole in the axil, which is often<br />

almost overhung by the chin of the next higher tubercle.<br />

spines: Very light straw-colored with pale brown tips when<br />

young, becoming darker with the tips sometimes dark purplishbrown<br />

on old plants. There are 10 to 16 round, rigid radial<br />

spines which all spread out at an angle from the plant surface.<br />

The 5 or 6 upper radials spreading erect are the longest spines<br />

of the plant, 3/4 to 7/8 of an inch long. In front of these at the<br />

very tip of the areole or occasionally scattered to as much as<br />

halfway down the groove to the axil there will often be 1 to<br />

3 additional very tiny spines to as little as 1/16 of an inch long.<br />

The lateral radials spread outward and are about 3/4 of an<br />

inch long. The lower 2 to 4 radials spread almost perpendicular<br />

to the plant surface and are 3/8 to 3/4 of an inch long.<br />

There is one central spine which is always turned upward to<br />

stand in front of the upper radials. It is round and a little<br />

heavier than the radials, has a bulbous base, and is 5/8 to 7/8 of<br />

an inch long.<br />

flowers: 1 to 11/2 inches wide and tall, white in color. There<br />

are a few whitish scales on the ovary. The perianth segments<br />

on the lower tube are small and scalelike with arrowheadshaped<br />

edges. These gradually lengthen up the tube until they<br />

become oblong, blunt-tipped segments about 3/16 of an inch<br />

wide. They have greenish-brown midlines and whitish, entire<br />

edges. The inner petals are cream-colored or pure white, only<br />

1/8 of an inch wide, and pointed, with entire edges. The filaments<br />

are green or whitish, the anthers yellow. The stigma has<br />

6 to 10 slender, light green lobes.<br />

fruits: Spherical or nearly so, about 1/4 of an inch in diameter.<br />

They are light green, sometimes with pinkish areas when<br />

ripening, becoming dry and papery when ripe. The perianth<br />

persists and there are a few whitish scales on the fruits. They<br />

split open along one side when mature. The seeds are black,<br />

finely tuberculate, about 1/10 of an inch long, with a very large,<br />

concave hilum.<br />

Range. Known only from lower parts of the Texas Big Bend.<br />

Scattered populations occur from near Terlingua, Texas, just west<br />

of the Big Bend National Park, to northwest of Ruidosa, Texas.<br />

Remarks. This cactus was apparently first discovered by J. P.<br />

Hester during his wonderfully thorough field study of cacti of<br />

the Texas Big Bend region. He published a description of it in<br />

the Cactus and Succulent journal of America in 1939, not naming<br />

it however, but assigning it two numbers in his own numbering<br />

system—this because he was inclined to think there were two<br />

forms here instead of one. No one again took notice of the plant<br />

for many years. This is not remarkable, since it only grows in<br />

the more inaccessible part of the Big Bend.<br />

The next mention of the cactus appears to be by Backeberg in<br />

his Die Cactaceae, Vol. 5. He apparently had only a few specimens<br />

sent to him in Europe, and his data on them was so meager<br />

that he gave the location of them as “U.S.A. (Arizona?).” He<br />

had not seen the flowers or fruits. It is obvious, however, from<br />

his rather complete description that he was describing this cactus.<br />

He showed uncertainty about exactly what to do with the plant,<br />

and took care of it by calling it Echinomastus sp. (Echinomastus<br />

pallidus nom. prov.). Its students it seems, have experienced an<br />

unusual reluctance to name this cactus, the first one to describe<br />

it assigning it only a number and the second one only a provi-<br />

sional name.<br />

It will be clear to anyone who has seen both of them together<br />

that this cactus is very close to the Arizona cactus, Echinocactus<br />

(Echinomastus) erectocentrus Coult. At a glance the two look<br />

alike, and the characteristics of stem and spines are almost alike.<br />

The differences are only quantitative. E. erectocentrus grows to<br />

8 inches tall, while our cactus has been seen to only 6 inches.<br />

The radials of the former are appressed rather tightly against the<br />

plant surface, while our Texas plant has its radials more spreading.<br />

The centrals are the same, except that on the Arizona plant<br />

they sometimes reach 1 inch, while on the Texas plant they have<br />

not been observed over 7/8 of an inch long.<br />

But if the differences of stem and spines are too small to he<br />

significant, the flowers and fruits of the two present more definite<br />

differences. E. erectocentrus has pink flowers and all writers<br />

describe it with 8 or 9 pink to deep purple stigma lobes. No<br />

one has previously described the flowers of our Texas cactus<br />

except for Hester’s statement that they were pure white. They<br />

are indeed pale cream-colored or pure white, with 6 to 10 light<br />

green stigma lobes. The fruits of the two differ in shape, those of<br />

E. erectocentrus being cylindrical and 1/2 to 5/8 of an inch long<br />

by 1/4 of an inch wide, while those of our Texas form are perfectly<br />

or very nearly spherical and only 1/4 of an inch across, but<br />

they both split alike along one side to release their seeds.<br />

The differences outlined above do not seem to indicate two<br />

distinct species, but neither do I think they can be ignored. I<br />

think the plant is best regarded as a separate variety of the Arizona<br />

species. Since it is essentially a slightly smaller form of the<br />

other and more pale in all of its coloring of spines and flowers,<br />

the use of Backeberg’s proposed species name for it as the name<br />

of the variety seems most appropriate.<br />

Roads have recently been built into the lower Big Bend areas<br />

where variety pallidus grows. It is never common there, but is<br />

found on widely scattered limestone ridges from near Terlingua<br />

up along the Rio Grande to beyond Ruidosa, Texas. Where it had<br />

been seen until recently by only a very few people, now that the


92 cacti of the southwest<br />

area is opened this has become one of the standard offerings of<br />

the Texas cactus dealers, going out under almost every sort of<br />

name. Anyone who has searched for this cactus and who knows<br />

how scarce it is, when he looks down into a bin of literally<br />

hundreds of these specimens in a dealer’s stock must realize with<br />

sadness how efficient their digging is and in what danger all the<br />

cacti lie. This will probably be another in that procession of cacti<br />

suddenly sold in every dime store and nursery until its newly<br />

opened territory is stripped and the form again becomes unknown<br />

except to specialists.<br />

This cactus has all of the characteristics of its relatives, and is<br />

like most of them in being very exacting in its requirements of<br />

full sun and heat and very little water.<br />

Echinocactus mariposensis (Hester)<br />

[Echinomastus mariposensis Hester]<br />

Description Plate 25<br />

stems: Single, practically globose, egg-shaped, or short oblong.<br />

These stems grow to a maximum of 31/2 inches tall by<br />

about 2 inches in diameter, but are usually smaller. Small<br />

plants have 13 ribs, but as they mature the number of ribs<br />

increases to 21. These are usually twisted and wrinkled into<br />

more or less distinct but small tubercles. The surface is light,<br />

often yellowish-green.<br />

areoles: At first practically spherical and about 1/8 of an<br />

inch across, with much short brownish wool. At maturation<br />

the areole extends forward as a narrow groove on the upper<br />

side of the tubercle, the flower being produced at the end of<br />

this groove in the axil of the tubercle, where it is accompanied<br />

by long wool and a tuft of persistent white bristles in<br />

the axil.<br />

spines: There are 25 to at least 36 radial spines. These radiate<br />

evenly, are rigid, and are from 3/16 to 3/8 of an inch long. They<br />

are pure, shining white to gray, sometimes tipped with light<br />

brown. There are 4 to 7 centrals. The upper 3 to 6 of these<br />

spread upwards or are often somewhat appressed against the<br />

upper radials, are comparatively heavy, and are 1/2 to 3/4 of<br />

an inch long. The lower central is porrect or curving downward,<br />

heavy, but only 3/16 to 1/2 inch long. The centrals are<br />

whitish, gray, or pale yellow below, with their distal sections<br />

usually light brown or a striking bluish-gray.<br />

flowers: About 3/4 to 11/4 inches in diameter and length,<br />

opening funnel-shaped or wider. The ovary and tube of each<br />

flower have a dozen or so whitish scales. The outer perianth<br />

segments have somewhat erose edges. The inner segments are<br />

somewhat spatulate, their tips bluntly pointed and sometimes<br />

notched or toothed, There are two distinct flower colors found<br />

in this species. One has the outer perianth segments with green<br />

midribs and white edges and the inner segments with light<br />

green midlines and white edges. The other has the outer segments<br />

with brown midlines and pink edges and the inner<br />

petals pink fading to whitish at the edges. The stamens are<br />

cream-colored, sometimes with the filaments pinkish. The style<br />

is long and greenish or brownish, the stigma with 5 to 8 green<br />

lobes.<br />

fruits: Globose or oblong, up to 3/8 of an inch long. They are<br />

yellowish-green at first, becoming dry, and then splitting open<br />

on one side. They have a few scales upon them. The seeds are<br />

slightly over 1/16 of an inch long, ovate, and black.<br />

Range. Known only from hills a short distance north and north-<br />

west of Terlingua, Texas, in the southwest corner of Brewster<br />

County.<br />

Remarks. This dainty little cactus is the smallest of the barrel<br />

cacti in our area—usually little larger than a golf-ball—but it<br />

is definitely one of the Echinocacti. It is far from spectacular in<br />

any way and, with a very limited range in very rough country,<br />

it is not surprising that it was not discovered until 1945 and is<br />

still very little known.<br />

J. Pinckney Hester, the tireless explorer of the Texas Big Bend,<br />

discovered the cactus and described it in great detail. He found<br />

it first on hills overlooking the site of the once famous quicksilver<br />

mine called the Mariposa Mine, and named it after that<br />

mine. It has still not been reported very many miles from that<br />

site. He described it as an Echinomastus, and it is clearly a close<br />

relative of those species usually put in that microgenus. It will<br />

probably share in whatever decision is finally reached when the<br />

question about that group of plants is settled. It also shares most<br />

of the growth characteristics of those plants, which means that<br />

although it is small and delicate in appearance, it is just as tough<br />

a desert species as its bigger relatives. It is not found naturally<br />

hiding in any shade, but grows in the open in the thin layer of<br />

soil overlying hot, exposed limestone ridges. In cultivation it<br />

must be kept drier and in brighter sun than most of the other<br />

small species of our area, if it is to survive.<br />

The range of this species is very small and the population<br />

must not be great, so specimens should not be taken out in large<br />

numbers. In fact, it is a wonder that the population has not already<br />

been depleted. It was one of the most terrible experiences<br />

of this study to come upon at least a thousand specimens of<br />

small cacti, mostly this rare species, gathered by someone and<br />

left to die in a pile on a hill only a few miles from the old<br />

Mariposa Mine. I was told that this was probably a cache left<br />

by a professional cactus-digging crew for the dealer to pick up<br />

with a truck—but a cache which he missed. At any rate, the<br />

cacti were mostly burned to a crisp by the time I saw them and<br />

nothing could be done for them, but perhaps they did not die in<br />

vain if my telling of them here makes us a little more conscious<br />

of the plight of these little species.


genus Echinocactus 93<br />

Echinocactus conoideus (DC) Poselgr.<br />

[Neolloydia conoidea (DC) B. & R.]<br />

Description Plate 25<br />

stems: Globular to egg-shaped at first, becoming conical or<br />

cylindrical, to at least 4 inches tall and 23/4 inches thick. The<br />

stem of a plant may remain simple, but it often sprouts near<br />

the base, or even from higher on the sides, to produce 2 or 3<br />

branches. The surface is dull gray-green and shaped into 8 or<br />

13 indistinct ribs composed of spiral rows of almost completely<br />

separate tubercles. These tubercles are to about 1/2 inch<br />

long, conical in shape, from bases broad but somewhat compressed<br />

horizontally by their crowded, almost overlapping<br />

position.<br />

areoles: Circular, about 1/16 of an inch in diameter and with<br />

white wool when new. It soon becomes enlarged by the formation<br />

of a narrow groovelike extension forward from the<br />

spinous part of the areole. On a mature areole this groove<br />

extends to the axil of the tubercle, where it broadens into a<br />

larger felted area from which the flower comes. After blooming<br />

the original circular part of the areole remains as the<br />

spine-bearing portion, usually losing its wool, and the felted<br />

groove runs upward and inward from this.<br />

spines: There are 10 to 16 radial spines per areole, all radiating<br />

rather evenly, straight, rigid, white fading to gray, and<br />

1/4 to about 1/2 inch long. There are also 1 to 4 spreading central<br />

spines 3/8 to slightly over 1 inch long, straight, rigid,<br />

blackish when young, fading to gray. The lowest of these is<br />

the longest and heaviest one.<br />

flowers: Beautiful violet or violet pink in color, 1 to 2 inches<br />

in diameter and about 1 inch tall, opening rather widely. The<br />

ovary surface and tube seem most commonly naked of scales,<br />

but occasionally the ovary has one or two small, rounded,<br />

white-edged scales upon it. The outer perianth segments have<br />

pink centers with whitish, entire edges. The inner segments<br />

are violet or pinkish-violet all over and are lanceolate, with<br />

pointed tips and entire edges. The stamens are bright orange.<br />

The stigma has 5 to 7 long, white or yellowish lobes.<br />

fruits: Spherical, yellowish or reddish at first, drying and<br />

becoming brown, after which they seem to remain until broken<br />

open by some outside force. Most of them seem to be naked,<br />

but on a few the dried remains of 1 or 2 tiny scales may he<br />

recognized. The seeds are about 1/16 of an inch in diameter,<br />

black, tuberculate, with large basal hila.<br />

Range. Widely found in central Mexico, extending north into<br />

Texas to a distance of about 30 miles or so along the arc of the<br />

Rio Grande from near Del Rio west to somewhere near Boquillas.<br />

Remarks. Echinocactus conoideus had long been known from<br />

Mexico, where it grows over a huge area, but it was not known<br />

at first to be in the United States. Engelmann apparently saw<br />

specimens from Texas, but did not seem to connect them with<br />

the Mexican species. Instead, he called the Texas plant Mammillaria<br />

strobiliformis.<br />

When Britton and Rose dealt with it they first erected a new<br />

genus, Neolloydia, for it, and then described the Texas specimens<br />

they saw as a new species separate from the others, calling<br />

it Neolloydia texensis. This is therefore the name under which it<br />

has been most widely known in the U. S.<br />

It was Boedeker who first questioned the correctness of setting<br />

off the Texas plants as a separate species, saying that they<br />

were actually only the northern form of the original Mexican<br />

species, Echinocactus conoideus. But it is hard to change a usage<br />

so widely followed as this of Britton and Rose. While most<br />

authorities since Britton and Rose have not listed the species<br />

texensis separately, most collectors still use the name.<br />

Being very concerned with what might be the correct name<br />

for this plant, we have studied many specimens from both Texas<br />

and Mexico. We have found that while the range of variations<br />

in the Mexican plants is much greater, it wholly encompasses<br />

anything we have found in Texas. Since we have seen duplicated<br />

every character of the Texas plants in those from the general<br />

type area in Mexico, we do not feel that the Texas plants can<br />

even be considered to form a distinct variety.<br />

This species and its close relatives stand at the opposite end of<br />

the Echinocacti from the huge barrels. These are the only ones<br />

to which it really seems incongruous to apply the term “barrel<br />

cactus.” Their ribs are indistinct and made up of very nearly<br />

separate tubercles and their ovaries have at best very few scales<br />

upon them and often none at all. They are undeniably the nearest<br />

to the next large group of cacti, the Mammillarias.<br />

Because of this position, these cacti have been classified in almost<br />

all possible ways. In its original description it was named<br />

by De Candolle Mammillaria conoidea. In a few years it had<br />

also been saddled with a remarkable number of other names,<br />

among them M. diaphanacantha by Lemaire, M. inconspicua by<br />

Scheidweiler, M. echinocactoides by Pfeiffer, who must already<br />

have noticed its similarity to the Echinocacti, M. scheeri by<br />

Muehlenpfordt (1845 non 1847), and M. strobiliformis by Engelmann.<br />

When the large genus Mammillaria was divided up, it<br />

then naturally had to be given the name Coryphantha conoidea,<br />

since it has grooved tubercles, and Orcutt took care of that.<br />

But there had been another line of reasoning about its proper<br />

relationship, and Poselger had already renamed it Echinocactus<br />

conoideus, soon followed by Kuntze with Cactus conoideus.<br />

As if these weren’t already genera enough to choose from,<br />

Backeberg more recently took some of the relatives of this cactus<br />

and some formerly called Thelocactus species and created a new<br />

genus, Gymnocactus, for them. But now the most recent move is<br />

to revitalize the genus Neolloydia by returning these to it and<br />

by adding to them other species formerly in the genera Thelocactus<br />

and Echinomastus. Once again we seem to see the splitting<br />

process gone to its extreme, giving us more and more Un-


94 cacti of the southwest<br />

stable microgenera until lately a move has begun to recombine<br />

these again.<br />

The main point seems to be the question of which major group<br />

this cactus and its close relatives belong in, Echinocactus or<br />

Mammillaria. On this the decision seems to have been made<br />

some time ago. It is many years since anyone has considered this<br />

form a Mammillaria. While it is admittedly the more doubtful<br />

of them, all recent students seem to have placed it among the<br />

Echinocacti. This is because the ovary has at least one or two<br />

scales in at least some specimens, the fruit becomes dry, and it is<br />

possible to speak of ribs on at least most of the species involved.<br />

Beyond this, little can be said except that the recent move to<br />

combine the much more clearly Echinocactus species in Thelocactus<br />

and Echinomastus with these in their own microgenus,<br />

Neolloydia, draws these more solidly than ever into the Echinocactus<br />

group.<br />

Considering this species as an Echinocactus, then, I choose to<br />

use the name of that large genus for it-at least until the microgenera<br />

are a little more stabilized.<br />

Echinocactus conoideus is not ever very common in Texas.<br />

Where it occurs it is usually in stands of several dozen speci-<br />

mens, but these stands are in widely scattered locations. It grows<br />

on rocky hillsides, more or less in the open.<br />

Its range does not go at any point very far north of the Rio<br />

Grande. It is most easily found around Sanderson, Texas, and<br />

in the very eastern edge of the Big Bend. Our knowledge of the<br />

eastern limits of its range were changed by finding a nice stand<br />

of the cactus on a hillside overlooking Goodenough Springs,<br />

which is just north of the Rio Grande south of Comstock, Texas.<br />

This is the most eastern record of this species of which I know<br />

personally, although there are reports of it from near Del Rio.<br />

The cactus has been eliminated at beautiful Goodenough Springs<br />

and its whole surrounding area has been submerged with the<br />

filling of the lake behind Amistad Dam.<br />

I have no knowledge of this cactus west of the eastern Big<br />

Bend. Although I have seen a herbarium specimen labeled “near<br />

El Paso,” I have been unable to confirm that the species grows<br />

that far west.<br />

This is a fairly easy cactus to grow, not quite so badly affected<br />

by moisture as are most of the other members of this group.<br />

It is not a vigorous grower or a prolific bloomer, but the flowers<br />

it does produce are of a very beautiful shade and rather large.


Genus Lophophora Coult.<br />

We come here to one of several cactus genera which seem<br />

to lie between the Echinocacti and the Mammillarias. Although<br />

there have been attempts in the past to submerge them in first<br />

one and then the other of these larger groups they seem to defy<br />

either combination. The reasons for this may seem rather technical<br />

to the nonspecialist, but they are the stuff out of which cac-<br />

tus taxonomy is constructed.<br />

Although some of its members were first described by Lemaire<br />

as Echinocacti, the significant points which seem to rule the<br />

Lophophoras out of the genus Echinocactus are the facts that<br />

the ovary and fruit on them are entirely naked and that the<br />

fruit remains always fleshy. These characters would put them<br />

in agreement with the Mammillarias, but they are even more<br />

clearly set apart from that genus by the facts that their stems<br />

are ribbed and that their monomorphic areoles produce the<br />

flowers from the apexes of young tubercles rather than from<br />

the axils. So this small genus is left by all recent students to<br />

stand alone.<br />

There are only a very few species in this genus, and as yet<br />

little agreement exists as to exactly how many they number.<br />

Most authors list two and some three or four, but there is no<br />

standardization of species and varietal arrangement, so no defi-<br />

nite figure can be given.<br />

The members of Lophophora are small, globose, or depressed<br />

globose cacti growing from comparatively large, carrot-shaped<br />

taproots. Usually the stem of the plant is to about 3 inches in<br />

diameter, and although one form sometimes reaches to about 5<br />

inches, they stand no more than 2 inches above the ground. The<br />

stems of an individual may be single or may sometimes branch<br />

from the base to form large clusters.<br />

The surfaces of these cacti are blue-green, usually with much<br />

gray glaucescence. There are no spines at all after the early seedling<br />

stage. The very broad and flat ribs are composed of some<br />

of the broadest, flattest, most confluent tubercles seen anywhere.<br />

The areoles are small and round, with long white to yellowish<br />

wool which tends to persist. The flowers are small, bell-shaped,<br />

and pink, pale rose, white, or rarely pale yellowish. The fruits<br />

are club-shaped and rose-pink or reddish.<br />

The insignificant little members of this genus have been famous<br />

out of all proportion to their size and appearance as far<br />

back as we can trace them. They are the sacred plants of the<br />

Indians best known by the ancient Indian name, peyotl, which<br />

has become the peyote of common usage. This is all because<br />

these plants contain in their flesh a group of alkaloids which,<br />

when taken into the human body, have remarkable effects upon<br />

the nervous system.<br />

The history of man’s use of and society’s reaction to these<br />

alkaloids is a fascinating study in itself. From ancient times to<br />

the present, Indians of Mexico and the U.S. Southwest have<br />

eaten these cacti specifically for the effects they have on their<br />

senses. An idea of how ancient and how widespread the practice<br />

has been can be grasped by noting that these plants have been<br />

called in different cultural periods and different Indian societies<br />

by all of the following names: peyotl, teonanacatl, tlalcoyote,<br />

uocoui, xicorl, seni, and hicore or jiculi. All during the history<br />

of the area, the Indians have given the cactus a very special<br />

veneration, eating it in very special religious ceremonies. This<br />

was because it gave them remarkable sensations which they were<br />

convinced came from their Great Spirit and marvelous visions<br />

which they were certain were glimpses—granted them by the<br />

Great Spirit—of ultimate reality itself.<br />

In the meantime others who did not venerate its visions as<br />

quite so god-given discovered peyote and came to enjoy its relaxing<br />

effect. Many Mexicans came to relish a little of it now<br />

and then, just for the relaxation and sense of well-being it<br />

brings, and in central Mexico most good markets will include


96 cacti of the southwest<br />

among the herbs a stock of the mescal buttons, as the plants<br />

have also come to be called after mescaline, the most famous of<br />

its alkaloids. In the U. S. and in Europe many years ago word<br />

of its effects got out, and for years now some people have experimented<br />

with it, there being an extensive literature built up<br />

through the years on its effects.<br />

Cactus dealers, particularly in southern Texas, where the<br />

plants have been easily available, have sold the mescal buttons<br />

in quantities for at least 40 years.<br />

But very early, in the U.S., this ran up against the typical<br />

American suspicion of anything so mystical and strange as the<br />

experiences produced by eating the peyote button. Moves were<br />

made long ago to outlaw the sale and even the possession of<br />

these cacti, and for a while they were declared narcotics and<br />

prohibited by edict. But these moves failed for two reasons. In<br />

the first place it was found impossible to maintain that they<br />

were narcotics under any meaningful definition of that term.<br />

And in the second place it was found a direct breach of the<br />

principle of religious freedom to deny the Indians this key part<br />

of their religious ceremony. Peyote could no more be denied to<br />

these Indians than sacramental wine could be denied the Christians.<br />

This move therefore soon collapsed, and peyote again became<br />

legal in all but California, where a state prohibition of it<br />

has hung on to the present, although it has been only sporadi-<br />

cally enforced.<br />

This was the fascinating peyote cactus, then, eaten for millenniums<br />

by the Indians as their means to induce and intensify the<br />

mystical experience, relished casually as a harmless relaxing<br />

agent by many a humble Mexican, and beyond that tried occasionally<br />

by the curious all over the world in order to ex-<br />

perience its strange effects—until the past few years.<br />

Then suddenly, only a few years ago, our little cactus was<br />

catapulted into the limelight, where it is now discussed in everything<br />

from the most technical medical and psychological journals<br />

to the best art and literary magazines to the most lurid<br />

sensation-promoting newspaper.<br />

This spurt of interest in the cactus seems to have been touched<br />

off by the success of chemists in synthesizing some of the alkaloids<br />

found in the peyote. The most famous of these synthetic<br />

products similar to the compounds in peyote is the now wellknown<br />

LSD. This synthetic compound is extremely potent and<br />

does very strange things to the nervous system of the user.<br />

Knowledge of LSD has been general among psychologists and<br />

a few others for some years, but it was treated as a dangerous<br />

drug and used sparingly for research until recently. However,<br />

the accounts which did appear concerning LSD stimulated much<br />

interest, and many turned to the comparatively very mild, natural,<br />

unconcentrated, layman’s version of this sort of agent—<br />

peyote. Their motives in eating peyote were various, including<br />

a desire to share in the mystical experience many have always<br />

invoked by all sorts of means, from fasting to Zen, a desire<br />

among artists to profit from the heightening to the visual and<br />

auditory senses which it is well-authenticated that peyote brings,<br />

a desire to gain the feeling of cleansing and well-being which it<br />

is universally testified that peyote leaves in the user, and, of<br />

course, a good share of just plain curiosity.<br />

All this boded only ill for our little cacti. These little species<br />

which had foregone the use of spines to protect themselves from<br />

their enemies and instead saturated their flesh with a set of<br />

alkaloids unpalatable to most animals in order to survive, suddenly<br />

found themselves taken by the hundreds and thousands<br />

just for these unique protective agents they had developed. The<br />

demand became so great that the countryside was systematically<br />

sacked of all its peyote. Five or six years ago I knew<br />

thousands of acres in the lower Rio Grande Valley where peyote<br />

grew in profusion under almost every shrub, but visits to one<br />

after another of these locations now show them barren of even<br />

a surviving specimen after the crews of gatherers have been<br />

through. The plant now only survives north of the Rio Grande<br />

in a few small areas.<br />

For all of these years dealers have sold peyote openly to anyone<br />

who wanted it, with the full knowledge of the authorities.<br />

In the past few years I have been at establishments where hundreds<br />

of the plants were being shipped when governmental<br />

agents visited and observed the business and heard them assure<br />

the cactus dealers that they were doing nothing wrong. This was<br />

because various court cases had established that the active substances<br />

in these plants were not narcotic or intoxicating substances<br />

and because there was no evidence of ill effects from the<br />

eating of the plant.<br />

But very recently the picture has changed dramatically. LSD<br />

has escaped the laboratory and is being indulged in indiscriminately<br />

by all sorts of people. This synthetic substance is very<br />

powerful, has sometimes very violent effects, and a case can<br />

easily be made that it is a danger when misused. Moves for its<br />

control appear justified, and are under way. But how does that<br />

involve peyote, which does not even contain this dangerous<br />

synthetic substance?<br />

Fuzzy thinking seems to be indicated when peyote is involved<br />

at all. LSD, the alkaloids in peyote, and those found naturally<br />

in various other plants are all lumped together under the suddenly<br />

very emotion-charged term “hallucinogens,” because all<br />

of them are capable of producing hallucinations in the user.<br />

There is little, if any evaluation of these various substances.<br />

Every magazine and paper making any attempt to follow the<br />

trends has had articles on one of the prime subjects of discussion<br />

today, the hallucinogens. In almost all of these articles little or<br />

no effort is made to differentiate among them and deal with the<br />

subject of their possible good and bad qualities, their potential<br />

or lack of potential for harm, and the question of whether each<br />

one in its own right should or should not be prohibited to the<br />

public. They are all lumped together as hallucinogens and they<br />

seem destined to stand or fall with LSD.<br />

The Food and Drug Administration recently invoked the 1938<br />

Federal Food, Drug, and Cosmetic Act and ruled that the sale<br />

or use of any of them is the illegal sale or use of drugs. Under


genus Lophophora 97<br />

this new ruling you may not dig our little cactus, sell it to be<br />

eaten, or eat it yourself—unless you arc a member of the Native<br />

American Church, the legal name of the Indian religious group<br />

in the U. S. using it in their ceremonials. Note that the naturally<br />

occurring peyote or the substances in it in their naturally occurring<br />

concentration have never been ruled narcotic, incapacitating,<br />

or even intoxicating, and that it has never been stated to<br />

have harmful effects either mental or physical. It has apparently<br />

suffered from illogical association in the minds of the authorities<br />

with the man-made substances LSD and mescaline, which<br />

undeniably can be dangerous, and the 1938 Food and Drug Act,<br />

which could as easily be applied to aspirin or wine as this, has<br />

been used to suppress it. Because the distinction between peyote<br />

and LSD has been overlooked, we are once again in the hard-<br />

to-maintain position of prohibiting the use of a substance as too<br />

bad for the general public which we allow a chosen group to<br />

use as a supreme good in their religious services. Can this posi-<br />

tion be maintained now any more than it was before?<br />

Of course, this prohibition clearly works for the good of the<br />

cacti themselves. Without it the species of this genus which<br />

grows in south Texas would soon have become extinct in the<br />

U.S., but now the wholesale digging has stopped and the cactus<br />

has been granted a reprieve. From the standpoint of the cacti,<br />

this is good.<br />

But the cactophile is faced with a peculiar situation. While it<br />

is specifically made clear that in all of the United States but<br />

California—and most recently some other states also—he may<br />

acquire and grow a few peyotes as garden or pot plants, if he<br />

should eat one of his plants, or if he should allow someone else<br />

to eat one of them, he would be guilty of violating a federal drug<br />

control act. This special dispensation for him (no doubt granted<br />

to avoid raising his ire) is appreciated, but he must weigh carefully<br />

the question of whether, under this sort of ruling a bed of<br />

peyotes is not too dangerous to have around. Perhaps, if he wants<br />

to keep his collection complete, including the peyote species, he<br />

should join the Native American Church—just to be safe. The<br />

cactus dealer is, of course, in double jeopardy. He is assured that<br />

he can sell a few peyotes singly or in small numbers to those who<br />

wish to grow them, but that he is a lawbreaker if anyone eats<br />

any of the peyotes he sells. Faced with being prosecuted as a<br />

“pusher’ at any moment if he sells peyotes, most dealers I know<br />

have stopped handling these plants entirely.<br />

We are, of course, happy at the prospect that these interesting<br />

cacti may be saved from almost certain extinction by the new<br />

ruling, and are not primarily concerned here with whether or<br />

not anyone should be free to use them to “expand his consciousness.”<br />

But we do note that if the present rulings had gone into<br />

effect a few years earlier and put these species beyond the pale<br />

of respectability at that time, the research which has gone into<br />

these plants would have been well-nigh impossible. This is undoubtedly<br />

the most peculiar situation that has ever arisen concerning<br />

any cactus.<br />

It has been said that somewhere among the cacti you will<br />

find almost every kind of strangeness, and that this is the secret<br />

of the interest cacti generate in people. There is truth in this,<br />

and here, in the genus Lophophora we have another kind of<br />

strangeness. Here we have our “notorious” cacti, and with them<br />

the study of cacti acquires all the exciting elements of mysticism<br />

and sinister intrigue and danger. These are the cacti for those<br />

who thrive on such things.<br />

Lophophora williamsii (Lem. in SD) Coult.<br />

“Peyote,” “Mescal Button,” “Whisky Cactus,” “Dry Whisky”<br />

Description Plates 15, 26<br />

roots: Each plant grows from a large carrot-shaped taproot<br />

the same diameter at its top as the stem and tapering slowly<br />

below, being usually 3 to 5 inches long.<br />

stems: Each plant begins as a single stem and sometimes remains<br />

so, but often clusters greatly to form in one variety<br />

sometimes up to 50 stems to one specimen. ‘These stems, hemispherical<br />

or usually depressed-globular, grow to about 5 inches<br />

in diameter, but not over about 2 inches tall. The flesh of the<br />

stem is soft and flabby, the surface blue-green, usually with a<br />

gray glaucescence. There are 5 to 13 very broad, very low<br />

ribs separated by narrow grooves. These ribs may be straight<br />

or sinuous. Each rib is more or less divided into tubercles. At<br />

the apex these tubercles are fairly distinct, but lower on the<br />

stem they are only very slight projections or almost entirely<br />

obliterated, with or without small wrinkles to indicate their<br />

limits.<br />

areoles: Round, or nearly so, and small, only about 1/8 of an<br />

inch in diameter and located 1/4 to 11/4 inches apart on the<br />

summits of the tubercles. At first each areole is filled with<br />

much long white or yellowish wool, so that the usually depressed<br />

summit of the stem is more or less filled with the wool<br />

of the close-standing young tubercles. With age the wool usually<br />

turns gray and may or may not be worn off to leave the<br />

areole with merely a tuft of short wool. The flowers are produced<br />

from within these areoles at the summits of the young<br />

tubercles.<br />

spines: The plant is spineless after the very young seedling<br />

stage.<br />

flowers: Small, usually pale pink or whitish, but said to be<br />

rarely rose or pale yellowish. They are bell-shaped and 1/2 to<br />

1 inch in diameter. The small ovary is naked. The outermost<br />

perianth segments are greenish with entire edges. The inner<br />

segments are almost linear and are pale pink to rose, white,<br />

or yellowish. The filaments are whitish, the anthers yellow.<br />

There are 3 to 7 reddish or yellowish stigma lobes.<br />

fruits: Club-shaped, 3/8 to 3/4 of an inch long, pale pink or<br />

very pale rose in color when ripe, and remaining fleshy and


98 cacti of the southwest<br />

indehiscent. The seeds are 1/16 of an inch or slightly less in<br />

length, with basal hila.<br />

Range. Extending from a very wide range in Mexico across the<br />

Rio Grande a short distance into south and southwest Texas. In<br />

south Texas it occurs in Hidalgo, Starr, and Zapata counties; in<br />

west Texas it occurs in a few locations in Brewster County in<br />

the Big Bend. There are records of many years ago of the species<br />

having been taken near Laredo and near the mouth of the Pecos<br />

River, but the plant does not seem to be in these areas now.<br />

Remarks. This is the famous peyote. Individual stems are often<br />

called mescal buttons. Some plants remain single until very<br />

large, while others sprout new heads or buttons all around them<br />

almost from the beginning. I once collected an old plant of the<br />

latter type which was 18 inches across with almost 50 heads.<br />

Since it is said that a button 2 or 3 inches in diameter takes<br />

around 10 years to grow, it is easy to see that such old plants<br />

are very venerable. With the great interest in peyote of recent<br />

years it is almost impossible to find such old plants in the wild<br />

any more.<br />

There is much variation in the rib shape, number, and size,<br />

and some in flower features. This plant has been greatly studied<br />

for a long time, and a confusing series of taxa have been set up<br />

for it. After observing many of the plants in the field and thousands<br />

of them in dealers’ bins and markets from our area all the<br />

way to Chihuahua and central Mexico, and after growing and<br />

flowering them ourselves, we have come to the conclusion that<br />

many of the proposed varieties are unnecessary and that the<br />

species found in the U.S. needs to be divided into only the<br />

following two taxa.<br />

Lophophora williamsii var. williamsii (Lem. in SD)<br />

Description Plate 26<br />

roots: As the species.<br />

stems: As the species, except that they grow to only 3 inches<br />

or less in diameter and the tubercles are less distinct than in<br />

the other form, being smaller, only about 5/8 to 3/4 of an inch<br />

across their bases. They cluster, usually quite early and extensively<br />

when old.<br />

areoles: As the species, about 1/4 to 5/8 of an inch apart.<br />

spines: As the species.<br />

flowers: As the species, except that the sepals and petals<br />

each grow in fewer than 3 series and the stigma lobes are 3 to<br />

5 in number.<br />

fruits: As the species.<br />

Range. Central Mexico to south Texas. In Texas it is restricted<br />

at the present time to Hidalgo, Starr, and Zapata counties.<br />

Remarks. This is the typical form of the species. Since it was<br />

described too narrowly in the early accounts, a whole series of<br />

variety names grew up around it. These include the following:<br />

Anhelonium lewinii Hennings, which became Lophophora williamsii<br />

var. lewinii (Hennings) Coult.; Lophophora williamsii<br />

var. typica Croiz.; var. pluricostata Croiz.; and var. caespitosa<br />

Y. Ito. I have found specimens falling within the limits of all<br />

of these proposed varieties within single populations of both<br />

Mexican and Texas plants, and they all intergrade, so it seems<br />

most logical to broaden the description of the typical form to<br />

include all limits found together and stop trying to interpret<br />

simple genetic traits as varieties.<br />

This is the form of the species growing in south Texas, the<br />

form which has smaller stems but which forms large clusters of<br />

these smaller stems when old. It is the more tender form, preferring<br />

to grow under the partial shade of the brush, shrubs, and<br />

trees, and rather easily damaged by frost. Two or three degrees<br />

below freezing will usually kill it. It does not now appear north<br />

or west of southern Zapata County, but old records suggest that<br />

it may have once grown in Webb County.<br />

There was also described by Fric as variety texana a form<br />

with 14 ribs, but the locality of it is not known and I can find<br />

no record of another collection with 14 ribs. A variety lutea<br />

(Rouhiers) Croiz. has been proposed for specimens with yellowish<br />

flowers, but we have long ago seen that it is very risky to<br />

base even a variety on flower color alone.<br />

Lophophora williamsii var. echinata (Croiz.)<br />

Description Plate 26<br />

roots: As the species.<br />

stems: As the species, except remaining single or at most<br />

dividing to form 2 or 3 heads in very old specimens. These<br />

stems are larger than in the typical form, growing to at least<br />

5 inches in diameter. The tubercles are more conical and<br />

larger, the bases being 3/4 to 11/4 inches across.<br />

areoles: A little larger than the typical form, with more<br />

wool, but otherwise the same.<br />

spines: Spineless after the seedling stage, as the species.<br />

flowers: As the species, except that the sepals and petals are<br />

more numerous and usually in 3 series each.<br />

Range. Northern Mexico, extending from Chihuahua and Coahuila<br />

into the Texas Big Bend in lower Brewster County.<br />

Remarks. This is a larger, tougher form of the species. There is<br />

little difference in the structure of the two except that in all<br />

respects the stems of this form are heavier and larger, although<br />

not clustering to any marked degree. This form is found growing<br />

on dry, exposed hillsides of the Big Bend where the lower


genus Lophophora 99<br />

Rio Grande Valley form would be burned to a crisp. It can also<br />

survive the much more severe cold of the Big Bend. I have several<br />

times had the smaller form from south Texas freeze in San<br />

Antonio, while this form growing in the same bed showed no ill<br />

effects.<br />

This form does seem distinct enough to be recognized, and it<br />

has been called a separate species, but this hardly is warranted.<br />

It seems at best a variety of the species, but a stable one. It is<br />

L. williamsii in the sense of Schultes (Cactus and Succulent Jour-<br />

nal, 1940), Britton and Rose’s description of the species seems<br />

to include this form along with the typical form.<br />

This is clearly the form which grew nearest to the Indians of<br />

Arizona, New Mexico, and northwestern Mexico, and is the one<br />

they originally hunted and used for their religious experiences.<br />

It has been noted that this form seems to have more of the alkaloids<br />

in it than are in the typical variety, since a small plant<br />

of this form when consumed will give more effect than the<br />

same-sized button of the typical variety.


Genus Ariocarpus Scheidweiler<br />

This is a small genus containing about half a dozen very<br />

strange cacti, one of which is found in Texas, the rest in Mexico.<br />

The body of an Ariocarpus consists of one or occasionally a<br />

cluster of low, flattened stems from only about 2 inches in diam-<br />

eter and not projecting above the soil level at all in some<br />

forms to as much as 10 inches across and 5 inches tall in one<br />

form. This stem sits on top of a large, carrot-like taproot.<br />

The surface of the stem does not have ribs, but is divided into<br />

very distinct, usually imbricated but noncoalescent tubercles.<br />

These are very firm, often have a horny, rough epidermis, and<br />

are of peculiar shapes, usually more or less triangular and flat-<br />

tened above. There are no spines after the first seedling growth.<br />

The members of this genus are unusual among cacti of our<br />

area because they flower in the fall of the year, usually from<br />

September to December. The flowers come from the woolly<br />

axils of the young tubercles at the center of the plant. They<br />

open widely, are diurnal, and are white, yellowish, or purplish<br />

in color. The ovary and fruit are both naked; the fruit is fleshy<br />

at first, becoming dry at maturity and disintegrating, leaving<br />

the seeds in the wool at the center of the plant.<br />

This genus was described and named Ariocarpus by Scheidweiler<br />

in 1838. The next year Lemaire redescribed it, calling it<br />

Anhalonium, and many students, including Engelmann and<br />

Coulter, thought that Lemaire’s name had precedence, so for<br />

many years there was confusion over these names.<br />

This is another genus which falls into the gap between the<br />

Echinocacti and the Mammillarias, or rather, which has some<br />

characters typical of each of these major groups but will not<br />

rest easily in either.<br />

The members have fruits which become dry, as do those of<br />

the Echinocacti, and some of them have monomorphic areoles<br />

also, but they have never been considered Echinocacti. This is<br />

partly because they have no ribs and because they have naked<br />

ovaries.<br />

They actually seem to be closer to the Mammillarias. In fact,<br />

for most of his life Engelmann persisted in including them in<br />

the genus Mammillaria. This is because they have a tuberculate<br />

surface and naked ovaries, and some species of them have dimorphic<br />

areoles with the nonproducing spinous portion of the<br />

areole at the tip of the tubercle and the floral portion separated<br />

from it at the base. However, the tubercle characters are very<br />

different; the flowers come from the apex of the stem instead<br />

of from older tubercles away from the apex; the fruits become<br />

dry and open; and there are differences of seed structure; all of<br />

which seems to separate these plants from the Mammillarias as<br />

well. Coulter remarked with obvious relish in 1896 that Engelmann<br />

had “finally come” to the opinion that these must be kept<br />

distinct from Mammillaria.<br />

The confusion over this sort of thing had no more than subsided<br />

when in 1925 Berger noted an obvious difference between<br />

certain members of the genus. Most have no groove on the upper<br />

surface of the tubercle, but two have a woolly groove on it.<br />

Berger seized upon this difference and proposed that those with<br />

grooves should be removed from the genus Ariocarpus and put<br />

into a new genus, Roseocactus.<br />

Since that time there has been a history of disagreement over<br />

Berger’s proposal. In brief, Marshall did not think that the difference<br />

was fundamental enough to warrant completely separating<br />

the plants into different genera, and proposed that Roseocactus<br />

be put back into Ariocarpus as a subgenus. Buxbaum<br />

seemed to agree with Marshall. The main recent champion of<br />

Berger’s view was Backeberg, who backed it vociferously in his<br />

large work on cacti.<br />

Only very recently was detailed study of the mode of development<br />

of the tubercles and areoles carried out and the information<br />

acquired, together with other factors, applied to the<br />

problems. However, it seems already to have brought some welcome<br />

clarification, as well as results of significance to the classi-


genus Ariocarpus 101<br />

fication of some other cactus groups. Edward F. Anderson made<br />

these studies and reported on them in a series of articles begin-<br />

ning in 1961.<br />

He found that in all of this group there is a single original<br />

growing point for the areole development which is located near<br />

the base of the tubercle instead of at its tip. From this single<br />

point develop all areolar structures, including the spinous portion<br />

(if represented), groove (if present), and floral portion. But<br />

there are differences in the different species in the way these<br />

develop from the original growing point. In Ariocarpus (Roseocactus)<br />

fissuratus, the floral development is at the base, with<br />

elongation of the tubercle occurring beneath the vestigial spinous<br />

part of the areole and thus drawing that part of the areole<br />

out into the already mentioned groove. In Ariocarpus retusus,<br />

the type species of the genus Ariocarpus, the rudimentary spinous<br />

part of the areole soon separates from the floral part and<br />

elongation between them then leaves the floral part in its basal<br />

position, while it pushes the spinous part to near the tip of the<br />

tubercle, where it persists as a woolly spot. In Ariocarpus<br />

trigonus the elongation occurs ahead of the spinous portion<br />

and never allows it to separate from the floral part at the base<br />

at all.<br />

The usual interpretation of these events requires us to call the<br />

elongated, groovelike areoles of A. fissuratus and also the short,<br />

basal areoles of A. trigonus, whose meristems do not divide,<br />

monomorphic. At the same time the areoles of A. retusus, where<br />

the floral and spinous parts separate, are entirely dimorphic.<br />

Essentially this same difference has, since Britton and Rose,<br />

been made the reason for separating the Coryphanthas out of<br />

the genus Mammillaria, and if it is so fundamental a difference<br />

as some have thought, it should also make mandatory the division<br />

of the genus Ariocarpus as well. But Anderson carried on<br />

many other investigations of seedling development, seed structure,<br />

other aspects of stem anatomy, fruit composition, and hybrid<br />

reactions, and concluded from these that they should all<br />

make up one genus, Roseocactus being at most a subgenus. The<br />

lack of importance of those details of areole structure which<br />

have been used so much in separating cacti is further indicated<br />

by Anderson’s report of A. fissuratus individuals without the<br />

groove and A. retusus individuals which have no spinous portion<br />

of the areole at all. This means that in at least A. retusus<br />

both the monomorphic and the dimorphic areoles occur in the<br />

same species, a situation which Dr. Boke has found also in certain<br />

Coryphanthas. This study has far-reaching implications for<br />

the taxonomy of other groups, where, it seems, too much em-<br />

phasis has been put on grooves or their absence.<br />

The members of Ariocarpus are retiring species, often not rising<br />

above the ground level at all, with usually horny and discolored<br />

surfaces which make them almost invisible. They are<br />

very difficult to find in their native haunts, but this is the point<br />

of their method of growth. They have no spines, and they depend<br />

instead upon being so insignificant as to be overlooked,<br />

upon camouflage, and upon some unpalatable alkaloids in their<br />

flesh for their survival. They are so unusual in their appearance<br />

that most people find it hard to believe they are cacti at all.<br />

Ariocarpus fissuratus (Eng.) K. Schumann<br />

“Living Rock,” “Star Cactus,” “Star Rock,” “Sunami,”<br />

“Chautle,” “Peyote Cimarron”<br />

Description Plate 26<br />

roots: A carrot-like taproot.<br />

stems: Entirely flat to somewhat rounded and depressedglobose,<br />

usually level with the ground or rising to only 1 inch<br />

or so above it. Covered with very crowded and overlapping<br />

tubercles which have bases broad and flattened and upper<br />

surfaces flattened and triangular in shape and 1/2 to about 3/4<br />

of an inch long. These upper surfaces are crossed by numerous<br />

small fissures which give them a warty appearance. The epidermis<br />

of the plants is always very firm and is gray-green<br />

when young or well-watered, but in the older plants or in the<br />

usual desert situation it is yellowish or brownish and, over<br />

most of the surface, hard, horny, and dead-appearing. The<br />

apex of the stem has long wool often almost entirely covering<br />

the younger tubercles. Plants usually have a single stem, but<br />

occasionally they branch to form a cluster of several to as<br />

many as a dozen stems.<br />

areoles: Each areole is at first circular and at or near the<br />

base of the young tubercle, filled with a dense mass of woolly<br />

hairs, but by the time it is easily visible on the elongated<br />

tubercle it is stretched into a conspicuous woolly groove running<br />

from the axial floral part to the tip of the tubercle on the<br />

upper surface. Mature areoles are thus linear and 3/8 to about<br />

5/8 of an inch long, except on very rare individuals on which<br />

the areoles do not elongate, remaining instead in the axil of<br />

the tubercle.<br />

flowers: From the axils of young tubercles at or near the<br />

center of the stem, where they arise out of the long wool.<br />

They are 1 to 2 inches broad, opening rather widely, but only<br />

about 1 inch tall. In color they are from almost white to pink<br />

or magenta. The ovary is naked and short; the outer perianth<br />

segments are almost linear with pointed tips, brownish or<br />

greenish with whitish, entire edges. The inner segments are<br />

pinkish or purple with whitish edges and are rather oblong<br />

from narrow bases, the tips with small, hairlike points at the<br />

apex. The filaments are white, the anthers bright orange. The<br />

style and stigma are white, with 5 to 10 lobes.<br />

fruits: Oval, about 1/4 to 5/8 of an inch long, pale green or<br />

whitish at first, becoming dry and disintegrating while still<br />

mostly buried in the long hair at the apex of the plant. The<br />

seeds are about 1/16 of an inch long, black, with rough sur-<br />

faces.<br />

Range. Northern Mexico into Texas. In Texas it is found along


102 cacti of the southwest<br />

the Rio Grande from the mouth of the Pecos River to near Presidio,<br />

but never penetrating more than a few miles into the state<br />

except in the Big Bend, where it occurs almost as far north as<br />

Alpine.<br />

Remarks. This is the plant to which the name “living rock”<br />

truly applies. It likes to grow on barren, rocky slopes where it<br />

survives burning up by hardly if at all projecting above the<br />

ground level and by having its surface covered with a thick,<br />

horny, brownish epidermis. It appears that its water storage is<br />

in the thick taproot on which the stem merely forms a flattened<br />

cap, and that the result of desiccation during the dry seasons is<br />

the shrinking of this root. This in turn seems to pull the stem<br />

down into the ground as it shrinks, so that it has even less surface<br />

to suffer from the cruel elements in these most exposed of<br />

all habitats in our region. The result is that the plants do not<br />

project any higher than the rocks all around them, and the<br />

brown, horny epidermis, broken into irregular warts by the<br />

many fissures from which the plant gets its name, looks just as<br />

dead and mineral as anything on the slope. I have repeatedly<br />

had the experience of walking around on what I thought was<br />

an unoccupied slope until I saw the first one of these cacti and<br />

then realized that I had been treading all over them, never<br />

knowing they were anything but rocks. Fortunately they are so<br />

hard that stepping on them does not damage them.<br />

The species is most interesting for its unusual appearance—<br />

quite uncactus like—and it blooms with a fine flower at a time<br />

of the year when most other cacti are through blooming. It<br />

interests people greatly, but I do not often advise it for growing<br />

in the usual cactus garden. Most people cannot keep it alive<br />

because it is one of the most extreme of the desert-adapted. It<br />

grows on the most exposed brows of the most arid ridges in<br />

west Texas, and cannot tolerate much moisture at all, or any<br />

amount of shade. Few people have the courage to “mistreat”<br />

this cactus with enough sunlight, heat, and dryness to keep it<br />

alive, healthy, and blooming. They want to pamper it until it is<br />

nice and soft and green, but A. fissuratus is a hard, rocklike<br />

thing, and it will not change, except to melt into rot if it is not<br />

kept in a situation approaching its desert home.<br />

As mentioned in discussing the genus, Engelmann long thought<br />

that this cactus was a Mammillaria: his first name for it was<br />

Mammillaria fissurata. Later Lemaire used the name Anhalonium<br />

engelmannii for it. K. Schumann first put it in Ariocarpus.<br />

The only other name applied to this species was Berger’s Roseocactus<br />

fissuratus.<br />

There is a form known as Ariocarpus fissuratus var. lloydii<br />

(Rose) Marshall. It is characterized by having the stems higher,<br />

more rounded, and larger in maximum size, the tubercles more<br />

rounded and with less distinct fissures. This variety is apparent-<br />

ly found only in Mexico.<br />

The living rock may have been more common in the past in<br />

the eastern part of its Texas range, but it is now very difficult<br />

to find east of the Big Bend. There it is still fairly common.


Genus Pediocactus B. & R.<br />

Originated for one species of cactus, this genus has grown<br />

through the years until it now contains at least seven species.<br />

The increase has come about in two ways. Four of its species<br />

have been discovered and described only within the past twenty<br />

years, and three of these less than ten years ago. This is, therefore,<br />

the only group of cacti in the United States which has had<br />

major additions to it in the last few years. New understanding<br />

has come with these new species. They have tended to fill gaps,<br />

and this has resulted in the combining under the over-all genus<br />

Pediocactus of four small genera: Pediocactus sensu B. & R.,<br />

Utahia B. & R., Navajoa Croiz., and Toumeya B. & R. This action<br />

has been taken by Dr. Benson only in the last few years,<br />

and since there has not yet been time for any opposing interpretations<br />

to appear it may be regarded as still somewhat tentative.<br />

However it seems to be the best system to take into account<br />

all the newer forms.<br />

This enlarged version of the genus Pediocactus would be<br />

characterized as follows: the stems are either single or branching<br />

sparingly; flattened, spherical, or cylindrical; usually very<br />

small but in one form up to 6 inches in diameter and height.<br />

The surface of the stem is covered with small but prominent,<br />

noncoalescent tubercles, spirally arranged. Areoles are small and<br />

entirely on the tips of the tubercles, sometimes with glands pres-<br />

ent. The spines are variable. The flowers are bell- or funnelshaped.<br />

The ovary is naked or with 2 or 3 small scales, sometimes<br />

these having a few hairs or bristles in their axils. The outer<br />

perianth segments are fringed to entire. The fruits are green at<br />

first, often changing to tan or yellowish, and then becoming<br />

dry. They are naked or have several small scales. In shape they<br />

are from nearly spherical to almost club-shaped. The fruits are<br />

dehiscent, opening by a ring around the apex, by a lateral slit<br />

on the upper side, or sometimes rather irregularly by both of<br />

these. Seeds are black or gray, the surfaces rough or shiny, but<br />

always textured when seen under the microscope.<br />

The members of this genus are once again cacti which fall between<br />

the major groups, the Echinocacti and the Mammiliarias,<br />

overlapping each to some extent. Most technical discussions of<br />

these cacti have become involved with trying to balance the<br />

characters in which they coincide with the one group against<br />

the characters in which they agree with the other. This began<br />

even with Engelmann, who had the type species of this genus<br />

as an Echinocactus, but who said that this species, with some<br />

others, “forms a small section of Echinocacti with the appearance<br />

of Mammillarias named by Prince Salm Theloidei.” Although<br />

he insisted that they were still “true” Echinocacti, he<br />

repeated that they “… constitute the closest and most imperceptible<br />

transition to Mammillaria subgenus Coryphantha.”<br />

To show the reasons for the divergent opinions over these<br />

cacti in the past, and the way they overlap both adjacent major<br />

cactus groups while actually falling outside of either one, we<br />

will mention here the most significant of the characters involved.<br />

They share with the Echinocacti the following points: the<br />

areoles are monomorphic with the flower coming at or near the<br />

tips of the tubercles; the flowers are similar to those of the<br />

Echinocacti, the ovary often with 2 or 3 tiny scales and occasionally<br />

these with a few bristles in their axils; the fruit becomes<br />

dry and splits open—but they differ from the Echinocacti<br />

by having no ribs and by having mucilage cells, which are not<br />

found in any recognized Echinocactus. On the other hand they<br />

share with the Mammillarias the following characters: the<br />

stems are tubercled instead of ribbed; the ovary is sometimes<br />

naked or has only 2 or 3 scales—while differing from them by<br />

producing the flowers from a monomorphic areole at the tip of<br />

the tubercle and by having dry, dehiscent fruits. In possessing<br />

mucilage cells, as Dr. Boke has pointed out, they look toward<br />

the Echinocerei. The result of all this is that this genus is left<br />

standing with those few others which are somewhat alone, outside<br />

of any of the major groups. Buxbaum has considered it


104 cacti of the southwest<br />

significant, because of its peculiar combination of characters, as<br />

an ancestor of other groups, but other scholars have disputed<br />

his theories on this.<br />

The members of this genus can be as exasperating to the ordinary<br />

cactophile as to the taxonomist. If it is difficult to view<br />

them in the proper systematic niche, it is even harder to view<br />

them in their native habitat. They are all extremely retiring<br />

cacti. They are usually so well camouflaged in their natural environment<br />

that there are places where it is more rewarding to<br />

hunt for them by feel than by sight. And it is not easy to find<br />

their locations. With the exception of one species, they all occupy<br />

very small ranges, several only a few miles in extent, and<br />

some are noted more for their rarity than for anything else.<br />

Each is restricted to a particular soil type or geologic formation,<br />

and some are associated with one other specific plant. The one<br />

species which is more widespread is usually a high mountain<br />

inhabitant where only the hardy collectors will come across it.<br />

So these are especially challenging little cacti not seen by<br />

many people and perhaps fully appreciated by only the specialist.<br />

However, they are part of the huge group known as cacti<br />

and they contribute to its amazing diversity.<br />

Pediocactus simpsonii var. simpsonii (Eng.) L. Benson<br />

“Mountain Cactus”<br />

Description Plate 27<br />

stems: Globose or sometimes even a little elongated in the<br />

growing season, but usually depressed and often almost flat<br />

in the winter. These stems are up to 5 inches in diameter and<br />

1 to 6 inches tall. They are almost always single, but on rare<br />

occasions clustering. The surface is covered with spirally arranged<br />

tubercles which are conical or sometimes somewhat<br />

pyramidal, 1/4 to 5/8 of an inch long. The color of the surface<br />

is light green.<br />

areoles: The areoles, situated on the tips of the tubercles, are<br />

circular or nearly so. When young, at the apex of the plant,<br />

they are large, to 3/16 of an inch in diameter, with much long<br />

white wool, but when older they shrink to 1/8 of an inch or<br />

so and lose most or all of their wool.<br />

spines: There are 15 to 30 radial spines which are white or<br />

whitish, rigid and straight, but very slender, radiating, and<br />

1/4 to 1/2 inch long, the shortest and most slender of them<br />

being at the top of the areole. On mature plants there are 5<br />

to 11 widely spreading central spines. These are heavier than<br />

the radials, rigid, straight or nearly so, and 3/8 to 3/4 of an<br />

inch long. They are whitish, cream, or pale yellow below,<br />

with the outer half of each darkening to brown or red-brown.<br />

flowers: Bell-shaped, opening rather widely. They are 5/8 to<br />

1 inch in diameter and length, and are pale pink, pale purplish,<br />

whitish, or yellowish in color. The ovary has several<br />

small scales near its top. The outer perianth segments are<br />

broadly rounded, greenish with pink to whitish, somewhat<br />

fringed, ragged, or notched edges. The inner segments are<br />

pink, pale purple, whitish, or yellowish, almost linear, with<br />

pointed tips and entire edges. The stamens are yellow, the<br />

stigma with 5 to 7 yellowish lobes.<br />

fruits: 1/4 to 3/8 of an inch long. Almost spherical to shortcylindrical.<br />

The fruits are green, sometimes suffused with reddish,<br />

later becoming dry. When ripe they split open somewhat<br />

irregularly along the upper side. The seeds are gray or<br />

black, the surface rough, 1/16 to 1/8 of an inch long in largest<br />

measurement.<br />

Range. Occurring far north of our area in Idaho, Montana,<br />

Nevada, Utah, and Colorado. From these states it enters Ari-<br />

zona and the northern mountains of New Mexico.<br />

Remarks. This cactus is a mountaineer. It is seldom if ever<br />

found below 6,000 feet in altitude, and ranges happily up to<br />

10,000 feet in the high Rockies. There are reports of its having<br />

been collected even above that altitude, but I have not been<br />

able to verify them. However, one thing is certain. This cactus<br />

scorns the extreme cold and the snow of the high mountains.<br />

Give it a sunny south slope for light and warmth in the summer<br />

and it will gladly sit all winter in the snows. It can stand an<br />

amazing amount of cold and moisture such as would kill most<br />

other cacti almost over night.<br />

The adaptation which this cactus has made to the high mountain<br />

climate is all gain in those mountains. Yet with every such<br />

gain something is usually lost, and this cactus has almost completely<br />

lost the ability to live where most of its fellows reside,<br />

in the hot, dry desert below the mountains. It can be kept alive<br />

and healthy in gardens in central New Mexico, but this is about<br />

its limit. It languishes and dies in two years or so in the low<br />

altitude and greater heat of San Antonio, even when kept more<br />

moist than its relatives. Those trying to grow this cactus should<br />

remember and try to simulate its mountain habitat, or they will<br />

have trouble.<br />

This species was first discovered and named by Engelmann.<br />

He observed it as a result of several expeditions over many<br />

years and described it very fully. However, he was from first<br />

to last convinced that it was an Echinocactus, and called it al-<br />

ways Echinocactus simpsonii.<br />

Coulter followed Engelmann entirely, but in 1893 the plant<br />

was called by M. E. Jones Mammillaria simpsonii, this indicating<br />

that already its position between the two major groups was<br />

becoming noticed. K. Schumann found a plant in Colorado<br />

which he called Mammillaria purpusii, and this is thought to<br />

have been the same plant.<br />

In 1913 Britton and Rose first set this species off from both<br />

the Echinocacti and the Mammillarias. They originated a new<br />

genus for this species alone, calling it Pediocactus simpsonii. The<br />

genus name is not at all apropos to this mountain cactus and<br />

seems to have been prompted by a doubtful report that the


genus Pediocactus 105<br />

cactus was once found on the plains of Kansas. However, it has<br />

remained as the valid name for this cactus, and a whole group<br />

of others have joined this species in the genus.<br />

Engelmann’s very complete description of the species is that<br />

of what, to be taxonomically correct, we must call variety simpsonii.<br />

Besides this Engelmann described a smaller variety which<br />

he called variety minor, and Coulter added a variety robustior<br />

from Nevada, Oregon, and Colorado.<br />

Although everyone has mentioned it, variety minor has not<br />

ever been described very completely. It appears to be a smaller<br />

form of the species found in Colorado, but there are few definite<br />

characters to establish it. It has been said that some specimens<br />

found in northwestern New Mexico might be this form,<br />

but as there are doubts about them, the variety is not listed as<br />

definitely one within our area. It should be noticed in this connection<br />

that a young, immature specimen of Mammillaria borealis<br />

which has not flowered and on which the grooves of the<br />

mature areoles have not yet formed fits the description of variety<br />

minor very well. This may be the explanation of some of the<br />

specimens of variety minor.<br />

Be that as it may, the typical variety of the species is found<br />

in the mountains of northern New Mexico. It is a beautiful<br />

cactus and one of our most hardy forms for cold climates.<br />

Pediocactus knowltonii L. Benson<br />

“Knowlton Cactus”<br />

Description Plate 27<br />

stems: Very small, 1/2 to 1 inch in diameter. These stems are<br />

depressed-globular or globular, a fraction of an inch to a<br />

maximum of 11/2 inches tall. Individuals usually have single<br />

Stems, but sometimes they form small clusters. Each stem is<br />

covered by small tubercles only 1/16 to 1/10 of an inch long.<br />

areoles: Almost circular at first, becoming elongated oval.<br />

These areoles are very small, being only about 1/24 of an inch<br />

in length. They have much white wool at first, which be-<br />

comes shorter with age, but is quite persistent.<br />

spines: There are 18 to 24 radial spines, which lie pectinate<br />

or even recurve somewhat and are 1/24 to 1/16 of an inch long.<br />

They are somewhat flattened and magnification reveals fine<br />

hairs upon them. In color they are white, pinkish, or reddish-<br />

tan.<br />

flowers: Opening widely and when fully open about 3/4 of<br />

an inch across by about 3/8 of an inch long. They are pinkish<br />

in color, with the ovary naked. The outer perianth segments<br />

are entire and blunt, the inner segments somewhat pointed.<br />

The stamens are yellow. The stigmas are 4 in number and rose-<br />

purple.<br />

fruits: Egg-shaped or somewhat club-shaped, about 3/8 of an<br />

inch long, becoming tan, dry, and dehiscent. The seeds are<br />

black and about 1/16 of an inch long.<br />

Range. Known only from one area in northwestern New Mexico,<br />

near the Los Pinos River just south of the New Mexico-<br />

Colorado border.<br />

Remarks. This is one of the very small, very inconspicuous new<br />

species in this genus which have only recently been discovered.<br />

It was first described in 1960 by Dr. Benson from plants dis-<br />

covered by the late Mr. Fred G. Knowlton.<br />

The cactus is quite clearly a Pediocactus. Very small, with<br />

nothing outstanding in stem, spine, or flower development, it<br />

has no special interest because of any feature except its rarity.<br />

However, for those who are intrigued by this, it is rare enough<br />

to make up for all it lacks otherwise, because it is definitely in<br />

the running for the most rare cactus of our area. It has been<br />

found so far only on gravelly hills near the Los Pinos River in<br />

New Mexico. Its population is apparently small, even there,<br />

and collectors should avoid decimating the species.<br />

Pediocactus papyracanthus (Eng.) L. Benson<br />

“Paper-Spined Cactus,” “Grama-Grass Cactus,” “Toumeya”<br />

Description Plate 27<br />

stems: Ovate or nearly so when young, becoming cylindrical.<br />

Most often the stem remains single, but old plants sometimes<br />

give off several branches by proliferation of several areoles<br />

on the sides of the stem. The surface of the stem is covered by<br />

dark green tubercles 1/8 to 3/16 of an inch long when mature.<br />

areoles: Round or nearly so, very small on immature stems,<br />

but up to 3/16 of an inch long on robust stems. With yellowish<br />

wool at first, this becoming gray and short, but persisting.<br />

Sometimes active areoles have 1 to several pinkish glands on<br />

the upper edge of each.<br />

spines: There are 6 to 9 radial spines which are straight, rigid,<br />

and flattened, and which radiate evenly. They are from less<br />

than 1/8 of an inch to as much as 1/4 of an inch long; the<br />

lowermost is heavier, wider, and longer than the others. They<br />

are white or gray in color, often snowy white. There are also<br />

1 to 4 central spines. The lowermost central, which seems always<br />

to be present after the very early stage, is greatly flattened,<br />

up to 1/10 of an inch wide at the base, flexible, papery<br />

in texture, and always to some extent twisted and curved. On<br />

young areoles it tends to stand at least somewhat upward,<br />

but lower down on the sides of the stem it may be aimed in<br />

any direction. It is from 3/4 to 11/4 inches long, usually mottled<br />

brown at first, fading to pale gray or whitish. Mature<br />

plants usually have 1 to 3 upper centrals, curving upward.<br />

They are similar to the main central except shorter and very<br />

much more slender, as well as usually less flattened.


106 cacti of the southwest<br />

flowers: Bell-shaped, not opening very widely, whitish in<br />

color. They are 3/4 to 1 inch long and wide. The ovary usually<br />

has a few small, toothed scales upon it, but may be bare. The<br />

outer perianth segments are triangular in shape with their<br />

edges entire or ragged, but not fringed. Their midlines are<br />

dark brownish, the edges whitish. The inner perianth segments<br />

are practically white. The stamens are cream-colored.<br />

The style and stigma are also cream-colored, with 4 or 5<br />

stigma lobes.<br />

fruits: Almost spherical, 3/16 to 3/4 of an inch long, becoming<br />

tan and dry at maturity and splitting open by a dorsal slit<br />

and a ring at the top. They are with or without a few scales.<br />

The seeds are black, shiny, but with a fine texture under<br />

magnification. These seeds are up to about 1/8 of an inch long.<br />

Range. A limited area in north-central New Mexico and an<br />

even more limited area in northeastern Arizona. In New Mexico<br />

it is found in a few scattered locations from near Santa Fe to<br />

the Sandia Mountains near Albuquerque. The range in New<br />

Mexico and the range in Arizona are not continuous.<br />

Remarks. This has long been regarded as perhaps the rarest cac-<br />

tus in our area. It is not profitable to try to decide which of<br />

several species actually is that, but P. pipyracanthus certainly<br />

is one of the least seen of them all.<br />

The rarity of its collection is partly due to its scarcity, but<br />

also to its excellent camouflage. This species grows in open<br />

grasslands where it appears to be almost always, if not always,<br />

associated with the grama grasses, from which comes the common<br />

name, “grama-grass cactus.” It usually grows in or near<br />

clumps of these grasses, and in this situation the broad, papery<br />

central spines look just like dried grass leaves and the cacti<br />

themselves like clumps of grass.<br />

Engelmann first described this cactus as a Mammillaria. Later,<br />

after closer examination, he observed that the flowers arose<br />

from the unelongated areoles at the tips of the young tubercles,<br />

and so observed that the plant was really an Echinocactus, as<br />

the genus was then understood. Coulter, therefore, classified it<br />

in the genus Echinocactus.<br />

We have already discussed in the introduction to the genus<br />

Pediocactus why this cactus and its relatives could not very<br />

logically remain in the genus Echinocactus. Britton and Rose<br />

appear to have been correct in separating it from that genus.<br />

They erected a special new genus, Toumeya, for this species<br />

alone. It has stood there, all alone, until recently.<br />

The very recent study of these plants by L. Benson has shown<br />

what appear to be good reasons for expanding the genus Pediocactus<br />

to include this cactus. New species only recently discovered<br />

are in several respects intermediate between the two<br />

original species, Pediocactus simpsonii and Toumeya papyracantha,<br />

and it seems that at best Toumeya should be reduced to<br />

a subgenus or section of Pediocactus.


Genus Epithelantha (Weber) B. & R.<br />

There seems to be but one species of this genus in the United<br />

States. It is a very small, but distinctive cactus.<br />

The whole stem of this cactus is covered with very many,<br />

very tiny tubercles—apparently the smallest tubercles of any<br />

United States cactus. Hiding these almost entirely from view<br />

are very many tiny spines. The growing tip of the stem is in the<br />

form of a rather distinct depression which is filled with a great<br />

deal of hairlike wool and covered over by the converging, later<br />

deciduous tips of the longer spines. This makes it very difficult<br />

to observe the formation of the tubercles, areoles, and flowers,<br />

but the way these are formed has assumed much importance and<br />

has been studied very closely. This is because taxonomically al-<br />

most everything hinges upon them.<br />

Originally Engelmann described this cactus as Mammillaria<br />

micromeris. In most of its characters it is a perfectly good Mammillaria.<br />

Later, however, something unusual was noticed about<br />

the cactus. It produces its flower not in the axil of the tubercle,<br />

but at the top of it. Mammillarias otherwise produce their flowers<br />

from halfway down the dorsal side of the tubercles to deep<br />

in the axils.<br />

When this was noticed it was assumed that the flower was<br />

produced from within a single, unlengthening, monomorphic<br />

areole on the tip of the tubercle. This is the situation in the<br />

Echinocacti. Because of this difference, Weber seemed unable to<br />

come to a real conclusion about this cactus, listing it once as a<br />

Mammillaria, once as Echinocactus micromeris, but also coining<br />

a new name, Epithelantha, for it. He apparently did not officially<br />

describe this latter as the name of a new genus, however.<br />

Britton and Rose then took the name Epithelantha and applied<br />

it to a new and separate genus. This genus, because of the supposed<br />

production of the flower from within the spine areole,<br />

has usually been placed in the subtribe Echinocactanae, although<br />

its other features, such as the naked fruits and lack of ribs, seem<br />

to point more toward the Mammillarias.<br />

Recently Dr. Norman H. Boke has done most thorough studies<br />

of cactus anatomy and development, and studied this species<br />

very carefully. In the course of his studies he has discovered<br />

that this cactus does not produce its flower from within a monomorphic<br />

spine areole after all. The blossom is, in fact, produced<br />

after a division of the meristem into a determinate spinous portion<br />

and a separate, indeterminate floral or vegetative meristem.<br />

This gives essentially a dimorphic areole, very different from<br />

those of the Echinocacti. It is actually more removed from the<br />

Echinocactus arrangement than is that of the many Mammillarias<br />

often set apart as Coryphanthas because they usually have<br />

monomorphic areoles elongating toward the axils instead of dimorphic<br />

areoles. The situation in this cactus can be interpreted<br />

as good dimorphic Mammillarian areoles in which the floral<br />

meristems merely remain at the tops of the tubercles. Dr. Boke<br />

notes Moran’s remark that for many years no one has linked<br />

Epithelantha to Mammillaria, but Boke’s conclusion is that a<br />

strong case for doing just this can be built.<br />

This possibility is very attractive, since the cactus is in so<br />

many ways a better Mammillaria than many of the Mammillarias<br />

themselves. It does seem that the work of Boke has made it<br />

impossible to classify it any longer with the Echinocacti, and<br />

that it points it toward the Mammillarias. Yet the fact remains<br />

that its flower is produced at the top of the tubercle, which is a<br />

trait not found in other members of that genus, and this difference<br />

in itself may be justification for keeping the cactus sepa-<br />

rate from the genus Mammillaria.<br />

As a separate genus based upon this cactus, Epithelantha<br />

seems, like Lophophora and Ariocarpus, to fall somewhere between<br />

the two major genera, Echinocactus and Mammillaria. It


108 cacti of the southwest<br />

is worth noting in this connection that the Epithelanthas Possess<br />

alkaloids similar to those of Lophophora and Ariocarpus,<br />

which seems to link them in some way.<br />

I, therefore, leave this genus in this difficult middle area.<br />

Buxbaum has made elaborate schemes in attempting to relate<br />

these plants phylogenetically, but others have pointed out that<br />

entirely different schemes from his could be devised which<br />

would appear just as logical as his, if different assumptions were<br />

made to start with. I am not primarily interested here in such<br />

phylogenetic schemes, so I merely list this as a small genus be-<br />

cause it seems in some way a separate entity among the cacti.<br />

Epithelantha micromeris (Eng.) Weber<br />

“Button Cactus,” “Mulato”<br />

Description Plate 27<br />

stems: Spherical or spheroid, usually with a depressed top.<br />

Usually only 1/2 to 1 inch in diameter, but sometimes to 13/4<br />

inches. Plants usually consist of a single stem, but are occasionally<br />

seen with double, and rarely with triple stems. These<br />

stems are covered with tiny, wartlike or somewhat conical<br />

tubercles about 1/20 of an inch long.<br />

areoles: Dimorphic. The spinous areole is at the tip of the<br />

tubercle, is small and at first has much long hair, which is<br />

later lost. The flower comes from a separate floral areole<br />

which appears adjacent to the spinous areole, also at the top<br />

of the tubercle.<br />

spines: There are about 20 to 40 slender but rigid spines per<br />

areole. They are in one series on immature plants, but in<br />

several series on mature areoles. The outermost series on mature<br />

plants are the stronger, and the upper ones of these are<br />

often 1/4 to 5/16 of an inch long when first produced, with the<br />

outer half of each swollen in diameter until they are somewhat<br />

club-shaped, but with the tips acute. These form an incurving<br />

tuft of long spines over the shorter spines and hairs<br />

of the new growth in the more or less depressed top of the<br />

mature plant. The enlarged outer sections of these spines break<br />

off with time, however, and the hairs are shed, leaving the<br />

sides of the plants with only short spines about 1/16 to 1/8 of<br />

an inch long. All spines are white, sometimes with faintly<br />

gray tips.<br />

flowers: Very small, only about 3/16 to 1/4 of an inch long<br />

and 1/8 to 3/16 of an inch wide, only partly rising above the<br />

long wool and spines in the top of the plant and only partly<br />

opening. They are pale, whitish-pink. The ovary is rather<br />

clavate, greenish-yellow, and naked. There are 3 to 5 outer<br />

perianth segments which have greenish-brown or pinkish midlines<br />

and pinkish-white, somewhat notched or eroded edges,<br />

in some specimens bearing a few short cilia. There are 5 inner<br />

petals which are whitish-pink with entire edges. The stamens<br />

are greenish-white. The style and stigma are greenish-white,<br />

with 3 or 4 stigma lobes.<br />

fruits: Club-shaped and 3/8 to 3/4 of an inch long by 1/8 to<br />

3/16 of an inch in diameter, red, fleshy, and naked, with the<br />

perianth remains persistent upon them. The seeds are black,<br />

about 1/16 of an inch (11/2 millimeters) long.<br />

Range. In the U.S. forming a large arc with the east end in<br />

Medina County, Texas, near San Antonio, running west from<br />

there, barely remaining within Texas at the mouths of the Devil’s<br />

and Pecos rivers and perhaps leaving Texas briefly before turning<br />

northward through Brewster County in the Big Bend. From<br />

there following the Davis and Guadalupe mountains northwest<br />

into New Mexico, passing through the Sacramento Mountains<br />

to have its northwest end in the Capitan Mountains of New<br />

Mexico. The western limit in Texas seems to be the Hueco Moun-<br />

tains just east of El Paso.<br />

Remarks. The tiny, tidy little button cactus reminds me of a<br />

very spherical button indeed, and even more of a white, fuzzy<br />

marble or ping-pong ball. It sits on exposed ridges and hillsides,<br />

a little white globe among the whitish limestone rocks. It is also<br />

the only cactus I have ever seen growing in a river bed. I have<br />

on several occasions found populations of dozens of specimens<br />

growing among the water-piled rocks in the almost always<br />

bone-dry beds of west Texas streams. Once, however, I found<br />

them within ten feet of perpetually running water in the bed<br />

of a major stream. Specimens in this sort of situation have extensive<br />

root systems running for several feet in all directions<br />

through the loose rocks and pebbles among which they stand.<br />

These roots anchor them against the rushing water which must<br />

entirely cover them at times, so long as the whole rock jam is<br />

not dislodged. The almost perfect roundness of the plants is<br />

what usually betrays their presence on hillsides and ledges where<br />

the surrounding rocks are almost never rounded, but in a deposit<br />

of water-rounded rocks they are doubly hard to see. This<br />

may be why they have not been reported from such a habitat<br />

before.<br />

There seems to be only a single form of this genus in the U.S.<br />

A separate form of this species, variety greggii, was also described<br />

by Engelmann but it is a larger form found only in<br />

Mexico. Many have regarded this as merely a synonym, but I<br />

have concluded that this larger and distinct form does exist.<br />

Beginning about 100 miles south of the Rio Grande on the road<br />

to Saltillo, I have seen this other form, which is unlike anything<br />

I have seen in Texas. I have often found stems of this form to<br />

3 inches in diameter. Dr. Boke, in the report of his study of Epithelantha,<br />

gives some interesting details of anatomical differ-<br />

ences he found between this Mexican form and our U.S. cactus.<br />

H. Bravo, Marshall, Backeberg and others have described a<br />

whole list of other Epithelantha forms found in Mexico. I have


genus Epithelantha 109<br />

seen some of these, and I am sure that some of them are distinct<br />

from our U. S. cactus, but as they do not enter the United States,<br />

I will not deal with them here. Perhaps it should be stressed<br />

that they do not grow in the United States, since some of them<br />

do from time to time enter the country in the trade. Because of<br />

this fact, one should be careful about assuming that any plant<br />

he buys as E. micromeris is actually the U. S. form. I have often<br />

seen the densely clustering Mexican form, with clumps of 10 to<br />

20 heads and beautiful rose-colored flowers 2 to 3 times as large<br />

as those on our cactus, sold simply as E. micromeris, when in<br />

reality it is quite different. No specimen like this has ever been<br />

recorded from within the U. S.


Turnefort -> Tournefort<br />

Genus Mammillaria Haw.<br />

The members of this genus are for the most part comparatively<br />

small or sometimes extremely tiny cacti. The plant stems vary<br />

in different species from depressed and almost flat to globular<br />

or sometimes even columnar in shape, and are often referred to<br />

as “heads.” In some species these remain single, while in many<br />

others they multiply from the base to become caespitose, one<br />

individual thus sometimes forming a large clump of these<br />

“heads.” In a few species the stems may branch sparingly from<br />

higher up on the stem.<br />

Each stem is entirely covered by a system of nipple-like projections<br />

called tubercles. These are usually arranged in spiral<br />

rows, but in a few cases are more loosely organized. These tubercles<br />

are usually cylindrical or conical, but sometimes may<br />

have more or less quadrangular bases and sometimes are mildly<br />

keeled below.<br />

Very early the knowledge of cacti progressed to the point<br />

where it became obvious that the huge assortment of forms they<br />

present could not be left in the one catchall genus Cactus L. By<br />

the middle of the eighteenth century Miller felt it necessary to<br />

divide the lot. By using the four old names of Tournefort, he<br />

separated out many cacti into Pereskia, Opuntia, and Cereus,<br />

leaving the rest in the genus Cactus. By 1812 even this narrowed<br />

genus Cactus was too broad, and Haworth abandoned it<br />

entirely, erecting five new genera out of it, one of which was<br />

Mammillaria, including all of the unjointed, tubercled cacti.<br />

Discoveries of new species continued, and as even this genus<br />

came to include a myriad of forms, the process of subdivision<br />

began all over again. Engelmann proposed two sections of the<br />

genus Mammillaria. He had section Coryphantha, which he<br />

characterized as having grooved tubercles, green fruits, and yellow<br />

or brown seeds, and section Eumammillaria with groove-<br />

less tubercles, scarlet fruits, and black or blackish seeds. Lemaire<br />

very soon elevated the section Coryphantha to a separate genus.<br />

Many concurred—although not all, as for instance Berger, who<br />

left this group as a subdivision which he rechristened Eu-coryphantha.<br />

This was the situation, rather uneasy and not wholly satisfactory<br />

to anybody, when Britton and Rose presented their<br />

major study, and they swept it all away by dividing the old<br />

genus Mammillaria into a whole spectrum of new and much<br />

smaller genera. Their names are in constant use and are most<br />

familiar to us today. The old section Coryphantha became the<br />

genera Coryphantha, Escobaria, Neobesseya, and others, and<br />

the old genus Mammillaria was eliminated as the rest of its<br />

forms were separated out into new genera such as Dolichothele<br />

and Neomammillaria. It seemed that the process of subdivision<br />

had been carried to its logical conclusion by this courageous<br />

leap of Britton and Rose, and almost the whole cactophile<br />

world adopted their array of new genera with surprising speed<br />

and many sighs ‘of relief.<br />

But the genera of Britton and Rose were not to go unchallenged<br />

for long. They were assaulted from two directions. As<br />

early as 1931 Fosberg questioned the basis for separating Escobaria<br />

from Coryphantha and concluded that the two should be<br />

recombined. This was an early expression of a desire for simplification<br />

by recombination. Many people had already found the<br />

genera of Britton and Rose so hard to tell from one another<br />

that it was often more difficult to determine the genus of a<br />

specimen directly than it was to determine its species first; and<br />

some had noted that the distinguishing characteristics of these<br />

genera were not always consistently present.<br />

But at the same time the trend to still more subdivision was


genus Mammillaria 111<br />

continued by various students. J. Pinkney Hester conducted<br />

very detailed studies of the seeds of cacti, and concluded that<br />

their variations did not well uphold the alignment of Britton<br />

and Rose’s genera, but actually, if regarded as diagnostic characters,<br />

would require a new realignment. As a result, in 1941,<br />

he shifted some species from one to another of these genera and<br />

erected such new genera as Escobesseya.<br />

Backeberg had already started subdividing further with his<br />

subgenera Subgymnocarpae and Neocoryphantha. Buxbaum<br />

conducted large studies of the cacti and proposed his own new<br />

subgenera, such as Pseudocoryphantha. He also proposed major<br />

theoretical schemes of cactus evolution which would appear to<br />

indicate radical new alignments of the species in this group.<br />

Thousands of words have been written concerning Buxbaum’s<br />

phylogenetic theories, but we do not need to study them here,<br />

because no one has yet actually followed his lead and there has<br />

been no essentially new scheme for classifying this group since<br />

Britton and Rose.<br />

What we do have at the present time are two opposing philosophies<br />

of classification giving two different concepts of this<br />

group, the same as they do of the Echinocacti. One considers<br />

very small differences in plants to be adequate bases for establishing<br />

genera and this results in lists of very slightly varying<br />

microgenera. This attitude is well expressed in Backeberg’s major<br />

work, where all of Britton and Rose’s genera are perpetuated<br />

and even some new ones added. ‘The other attitude is the more<br />

conservative one that a genus should be a major group based<br />

upon some rather obvious and very fundamental differences.<br />

This attitude regards the newer genera based on very small differences<br />

as no more than sections or subgenera, or at most micro-<br />

genera of an entirely different level from such larger plant<br />

genera as for instance Euphorbia. This approach had recent expression<br />

in L. Benson’s Arizona Cacti where all of these proposed<br />

genera were recombined once again into the original genus<br />

Mammillaria.<br />

Every serious cactus student is faced today with the battle<br />

between these opposing views, and even the amateur is affected<br />

by it, since, in order to be conversant, he often has to remember<br />

two or even more names for each of his cacti.<br />

It cannot be said that either view is established at this time.<br />

The present study does not presume to answer a major taxonomic<br />

question such as this. It is not even addressed to such a<br />

purpose. I would have preferred to avoid the issue entirely, but<br />

under the circumstances even to list a series of species is to take<br />

sides.<br />

Since a decision was thrust upon me, I wished to make it as<br />

intelligently as possible, so before making my decision I have<br />

studied the arguments for each view and then applied to the<br />

problem the most recent evidence to come to my attention. After<br />

the most exhaustive study of which I am capable, I feel constrained<br />

to follow here the recombination of these cacti under<br />

the genus Mammillaria and to consider this genus in the older<br />

and larger sense. The results of research reported since the publication<br />

of the last major work on these cacti has figured largely<br />

in my decision, so it may be of value to mention that newer<br />

evidence here.<br />

Most significantly, the old distinction between those plants<br />

with grooved tubercles and those with grooveless tubercles<br />

which prompted Engelmann to make the first division of the<br />

group into two sections and which is still so much emphasized<br />

that all artificial keys use it, seems to have failed us. Dr. Norman<br />

H. Boke, in a series of very detailed studies of cactus shoot<br />

form and development, has recently shown that both Coryphantha<br />

erecta and C. clava, two common Mexican species, may<br />

have grooved and grooveless tubercles on the same mature heads<br />

at the same time, or may change the form of their growth back<br />

and forth from the one to the other. This would appear to make<br />

it impossible to classify these particular species in either the<br />

proposed genus Coryphantha or Neomaimmillaria and to make<br />

it possible for a given specimen to fulfill the characteristics of<br />

both of these genera at once, which would seem to cast real<br />

doubt upon the divisions themselves.<br />

In terms more technical but more meaningful to the botanist,<br />

the grooved group have areoles monomorphic, which means<br />

producing from a single meristem not only both vegetative<br />

structures (leaf primordia and spines) but also the later reproductive<br />

structures (flowers and branches), while the grooveless<br />

forms have areoles dimorphic, with two separate meristems, one<br />

producing only vegetative structures at the summit of the tubercle<br />

and the other producing only reproductive structures,<br />

usually at the axil of the tubercle. This distinction appeared at<br />

first to be an essential one, dividing the whole group handily,<br />

but here again Dr. Boke was able to show that in the two species<br />

mentioned above, the areoles may be either monomorphic or<br />

dimorphic on the same adult head of the same specimen.<br />

Since these distinctions have broken down, there apparently<br />

remains no character by which the large group formerly known<br />

as genus Mammillaria can be divided into two major subdivisions.<br />

Such things as sepals fringed versus sepals entire, fruit<br />

green at maturity versus fruit red at maturity, fruit with a few<br />

scales versus fruit naked, and details of seed form all show exceptions<br />

on one side or the other in all major subdivisions which<br />

have been proposed.<br />

But what about the status of the array of small genera erected<br />

within this large group of tubercled cacti? As already mentioned<br />

the division into Coryphantha and Escobaria was challenged<br />

almost immediately by Fosberg. The distinguishing characters<br />

usually given of green fruit on the one hand and red fruit on the<br />

other obviously does not work, because the fruits of some Coryphanthas<br />

become brownish or reddish when very ripe and those<br />

of several Escobarias remain green barely flushed with apricot<br />

on the sunny side. Nor is seed color always reliable to separate<br />

these two proposed groups. One searches in vain for a valid<br />

reason why Fosberg has not been followed and why these two


abilitiy -> ability<br />

112 cacti of the southwest<br />

groups have been allowed to stand so long in most of the litera-<br />

ture.<br />

To make a long story short, all other distinguishing characters<br />

proposed for these microgenera have proved as uncertain. We<br />

have, therefore, been left with only such quantitative characters<br />

as long tubercles versus short tubercles, tubercles grooved all the<br />

way versus tubercles grooved more or less of the way, flowers<br />

predominantly yellow versus flowers brownish through pink to<br />

purple, and so on, which hardly seem adequate to distinguish<br />

genera—and there are exceptions to all of them anyway. Attempts<br />

to separate on this sort of basis have resulted in a constant<br />

shifting of species from one to the other genus and finally<br />

in the proposal of Escobaria subgenus Pseudocoryphantha Buxbaum<br />

and subgenus Neocoryphantha Backbg., as well as of<br />

genus Lepidocoryphantha Backbg., for those which burst out of<br />

the closely drawn genera.<br />

The other genera which Britton and Rose proposed for this<br />

group fare little better. We have seen that monomorphic versus<br />

dimorphic areoles and grooved versus ungrooved tubercles will<br />

not divide them. Neither will flower color, since we have the<br />

whole range of colors in the proposed genus Coryphantha, in<br />

Neobesseya, and in Britton and Rose’s strictly drawn Neomammillaria.<br />

Fringed versus nonfringed sepals and even various de-<br />

grees of fringing in the same species are found in both Coryphanta<br />

and Neomammillaria. Seed form fails also, with both<br />

Escobaria and Neomammillaria showing the whole range of<br />

seed coats, shapes, and hilum positions so completely that Buxbaum<br />

has to theorize parallel evolution within each of these<br />

groups because of it.<br />

It seems that there are no characters left strong enough upon<br />

which to erect genera and that the whole group is best considered<br />

one genus, as originally conceived. Dr. Boke’s judgment<br />

after his research would seem justified:<br />

In any event, it is my opinion that the discovery of a combination<br />

of areole monomorphism and areole dimorphism in<br />

Coryphantha clava and C. erecta weakens one of the principal<br />

distinctions between the Mammillarias (sensu lato) and<br />

other tubercled cacti. I think that it also indicates a cautious,<br />

conservative approach in delimiting genera in these cacti.<br />

He who can take the larger view will find in the genus Mammillaria<br />

a rich and diverse group of cacti presenting almost<br />

every sort of interesting variation on the theme of the small,<br />

tubercled cactus body. As a group they present all of the challenges<br />

to his ability at collecting, classifying, and culturing<br />

cacti which the most ardent cactophile can desire.<br />

For those who are fascinated by the Britton and Rose type of<br />

genus divisions and who want to concern themselves with this<br />

sort of thing, as well as for those who may be familiar with<br />

only those plant names, I have added for each form described<br />

here the name which seems to be the most valid one under the<br />

microgenus system.<br />

KEY TO <strong>THE</strong> MAMMILLARIAS<br />

1a. Diameter of stems on mature plants 2 inches or more and the<br />

length of the tubercles 1/4 of an inch or more—2.<br />

2a. Stems on mature plants hemispherical to flattened, always as<br />

broad as they are tall on normal specimens and usually much<br />

greater in diameter than in height—3.<br />

3a. Areoles always dimorphic, with the spinous portion at the<br />

tip of the tubercle and the floral portion in the axil of the<br />

tubercle and having no groove connecting them—4.<br />

4a. Central spines 1 or 2 per areole, short and always straight;<br />

outer perianth segments entire—5.<br />

5a. Color of the plant surface deep green or blue-green;<br />

tubercles firm and their bases quadrangular and more<br />

or less keeled; flowers whitish, rose, or pinkish—6.<br />

6a. Radial spines 5 to 9; tubercles strongly keeled and to<br />

Is of an inch long; plant to 12 inches in diameter<br />

—M. meiacantha.<br />

6b. Radial spines 9 to 26; tubercles with bases quadrangular<br />

but not so strongly keeled; plants to about 5<br />

inches in diameter—7.<br />

7a. Radial spines 20 to 26 —M. heyderi var. heyderi.<br />

7b. Radial spines 9 to 20—8.<br />

8a. Radial spines 14 to 20 —M. heyderi<br />

var. applanata.<br />

8b. Radial spines 9 to 13 —M. heyderi<br />

var. hemisphaerica.<br />

5b. Color of plant surface light yellowish-green; tubercles<br />

flabby and egg-shaped to cylindrical; flowers brightly<br />

yellow —M. sphaerica.<br />

4b. Central spines 1 to 4 and at least some of them hooked;<br />

outer perianth segments fringed—9.<br />

9a. Radial spines 8 to 15; flowers bright purple with about<br />

20 inner perianth segments and 11 yellow stigma lobes<br />

—M. wrightii.<br />

9b. Radial spines 14 to 22; flowers paler pinkish-purple<br />

with about 40 inner perianth segments and 5 to 9 green<br />

stigma lobes —M. wilcoxii.<br />

3b. Areoles normally and predominantly monomorphic and prolonged<br />

into a groove extending halfway or more toward the<br />

axil of the tubercle on mature stems, with the flower pro duced in<br />

the end of this groove—10.<br />

10a. Tubercles equal in size or nearly so on a given stem and<br />

arranged regularly; central spines 0 or 1 per areole and<br />

1/4 to 5/8 of an inch long; flowers greenish-yellow, brown-<br />

ish, or pink; fruits scarlet when ripe—11.<br />

11a. Flowers greenish, greenish-yellow, or brownish, sometimes<br />

streaked with pink; the outer perianth segments<br />

fringed —M. similis.<br />

11b. Flowers pure pink without stripes or zones of various<br />

colors; the outer perianth segments not fringed, but en-<br />

tire —M. rosiflora.<br />

10b. Tubercles unequal in size and shape on a given stem and<br />

arranged irregularly; centrals 1 to 4 per areole and 3/4 to


genus Mammillaria 113<br />

2 inches long; flowers purplish or rose-pink; fruits re-<br />

maining greenish when ripe —M. runyonii.<br />

2b. Stems on mature plants spherical to columnar, usually taller<br />

than they are broad and often markedly so—12.<br />

12a. Having at least one and often several hooked central spines;<br />

areoles dimorphic with the spinous portion at the tip of the<br />

tubercle and the floral portion in the axil of the tubercle,<br />

the two never connected by a groove —M. microcarpa.<br />

12b. Without hooked centrals; areoles mostly or entirely monomorphic,<br />

the flower always produced at the end of the undivided<br />

areole which is prolonged into a groove running at<br />

least part of the way down the tubercle—13.<br />

13a. Having fleshy taproots and 2 to 8 central spines to 21/4<br />

inches long; flowers purplish —M. macromeris.<br />

13b. Having no fleshy taproots and having central spines not<br />

over 15/8 inches long—14.<br />

14a. Having one brownish gland in the groove formed by<br />

the elongated areole —M. bella.<br />

14b. Without glands—15.<br />

15a. Centrals 0 to 4 on mature plants—16.<br />

16a. Centrals present—17.<br />

17a. Centrals not hooked—18.<br />

18a. Flowers yellow, orange-yellow, or yellow with<br />

red centers—19.<br />

19a. Stems single or very sparingly branched;<br />

flowers yellow or orange-yellow, sometimes<br />

reddish when fading, but not yellow with<br />

red centers; radials and centrals to at least<br />

3/4 of an inch long—20.<br />

20a. Plants large and robust, to at least 6<br />

inches tall and 4 inches or more in diameter<br />

when old; tubercles 1/2 to 1 inch long<br />

—21.<br />

21a. Radials 6 to 16; outer perianth segments<br />

lacerated and more or less<br />

fringed<br />

—M. scheeri (in part).<br />

21b. Radials 14 to 28; outer perianth segments<br />

entire and smooth<br />

—M. scolymoides.<br />

20b. Plants small, to a maximum of 3 inches<br />

tall or wide; tubercles 3/8 to 1/2 inch long<br />

—M. echinus (in part).<br />

19b. Stems greatly branching by new heads arising<br />

from the grooves in old tubercles all<br />

around their bases to form large masses of<br />

often dozens of stems; flowers yellow with<br />

red centers —M. sulcata (in part).<br />

18b. Flowers pale pink to deep rose-purple<br />

—M. ramillosa.<br />

17b. One central hooked —M. scheeri (in part).<br />

16b. Centrals absent—22.<br />

22a. Tubercles over 1/2 inch long; radials 3/8 to 3/8 of<br />

an inch long; flowers yellow with red centers<br />

—M. sulcata (stunted or atypical plant).<br />

22b. Tubercles 1/2 inch or less long; flowers not yel-<br />

low with red centers—23.<br />

23a. Radials 3/8 to 1 inch long; flowers yellow<br />

—M. echinus (stunted or atypical plant).<br />

23b. Radials only 1/8 to 1/2 inch long; flowers light<br />

purple —M. hesteri.<br />

15b. Centrals 4 to 17 on mature plants—24.<br />

24a. Fruits green or greenish when ripe, sometimes becoming<br />

brownish or apricot on part of the surface<br />

when very ripe, but never bright red—25.<br />

25a. Radial spines 12 to 20—26.<br />

26a. Flowers rose-purple to deep purple and 1 inch<br />

or more long and wide; stigma lobes 7 to 12;<br />

at least some of the spines over 1/2 inch long<br />

—27.<br />

27a. Tubercles 3/8 to 1 inch long; centrals pale<br />

mottled, and brown-tipped, but never<br />

blackish over half or more of their lengths;<br />

flowers 11/2 to 21/2 inches across when fully<br />

opened; seeds 11/2 to 13/4 millimeters long<br />

with ventral hila—28.<br />

28a. Radials to only 5/8 of an inch or so long;<br />

stigma lobes 7 or 8 in number and rose-<br />

purple in color; seeds about 11/2 millimeters<br />

long, dark brown, with the hila<br />

small —M. vivipara var. vivipara.<br />

28b. Radials to around 1 inch long; stigma<br />

lobes 8 to 10 in number and white in<br />

color; seeds about 13/4 millimeters long,<br />

light brown in color, with the hila large<br />

—M. vivipara var. arizonica.<br />

27b. Tubercles 1/2 inch or less long; centrals very<br />

dark brown or purplish black over one-half<br />

to all of their lengths; flowers 1 to 11/2<br />

inches across when fully opened; seeds 2<br />

millimeters long with subbasal hila<br />

—M. vivipara var. borealis.<br />

26b. Flowers white, pink, or very pale rose in color<br />

and about 3/4 of an inch long; stigma lobes 5<br />

or 6; all spines 1/2 inch or less long<br />

—M. varicolor.<br />

25b. Radial spines 20 to at least 60—29.<br />

29a. Flowers 1 inch or more in length and width;<br />

spines strong and somewhat flexible; seeds 1/2<br />

to 21/2 millimeters long—30.<br />

30a. Flowers deeply and brilliantly purple or<br />

else reddish-purple; radial spines white or<br />

white tipped light brown—31.<br />

31a. Flowers 2 inches or more long and broad;<br />

radials 20 to 30; seeds dark brown—32.<br />

32a. Tubercles about 3/4 of an inch long;<br />

centrals 4 to 7 and straw-colored or<br />

whitish; flowers deep purple with 7 to<br />

9 rose-colored, blunt stigma lobes; seeds<br />

oval, brown, 2 to 21/2 millimeters long,<br />

with the surfaces very finely pitted<br />

—M. vivipara var. radiosa.<br />

32b. Tubercles about 1/2 inch long; centrals<br />

6 to 12 and purple-black or dark


114 cacti of the southwest<br />

brown; flowers reddish-purple with 6<br />

to 10 pure white or slightly pinkish,<br />

somewhat pointed stigma lobes; seeds<br />

21/2 to 3 millimeters long, markedly<br />

reniform and much flattened, with<br />

their surfaces shiny smooth and un-<br />

pitted —M. fragrans.<br />

31b. Flowers 1 inch or only slightly more in<br />

width and length; radials 20 to at least<br />

60; seeds light brown —M. vivipara<br />

var. neo-mexicana.<br />

30b. Flowers yellowish suffused with purple;<br />

radial spines straw-colored with reddish<br />

tips —M. vivipara var. deserti.<br />

29b. Flowers less than 1 inch in length and width;<br />

spines heavy but very brittle and glassy,<br />

breaking at slight pressure; seeds I millimeter<br />

long —M. albicolumnaria.<br />

24b. Fruits bright red or scarlet when ripe—33.<br />

33a. Stems hardly over 2 inches in diameter; length<br />

of radial spines 1/8 to 3/8 of an inch; 4 to 7 centrals<br />

with one conspicuous, heavy central standing<br />

porrect or turned downward a little; flowers<br />

over 1 inch in diameter, opening widely, lavender-white<br />

or very pale purplish in color, with 5<br />

or 6 white stigma lobes —M. tuberculosa<br />

(in part).<br />

33b. Stems to 23/4 inches in diameter; radials 1/2 to<br />

1 inch long; 7 to 17 centrals without a conspicuous,<br />

heavy, porrect one; flowers 3/4 of an inch<br />

or less in diameter and not opening widely,<br />

pinkish in color streaked with brown and with<br />

4 or 5 very green stigma lobes —M. dasyacantha.<br />

1b. Diameter of stems on mature plants less than 2 inches and<br />

length of tubercles 1/4 of an inch or less—34.<br />

34a. Radial spines 13 to 18; all spines club-shaped, remaining<br />

thick almost their whole length and coming to a point very<br />

suddenly —M. nellieae.<br />

34b. Radial spines 20 or more; spines not club-shaped, but awl-<br />

shaped, bristle-like or hairlike—35.<br />

35a. Centrals present—36.<br />

36a. All spines rigid—37.<br />

37a. Having a fleshy taproot—38.<br />

38a. Single or sparingly clustering; radials 24 to 36 in<br />

number; tubercles grooved all of the way from the<br />

tips to the axils by the greatly elongated, mono-<br />

morphic areoles; stigma lobes bright yellow<br />

—M. duncanïi.<br />

38b. Densely clustering with dozens of heads in a typical<br />

plant; radials 40 to 85 in number; tubercles<br />

either ungrooved or grooved to around 1/2 of their<br />

lengths; stigma lobes pure white —M. leei (in part).<br />

37b. Roots all fibrous—39.<br />

39a. Stems single to sparingly branching; 1 to 2 inches<br />

thick and standing well over 3 inches tall when<br />

old—40.<br />

40a. Centrals from very bulbous bases; flowers coming<br />

from the sides of the stems, 1/2 inch or less<br />

in diameter and dark red in color; tubercles not<br />

grooved —M. pottsii.<br />

40b. Centrals not conspicuously bulbous; flowers<br />

produced at the summit of the stem, an inch or<br />

more in diameter and clear lavender or very<br />

pale purplish in color without darker midlines<br />

—M. tuberculosa (in part).<br />

39b. Stems branching greatly into large masses, but each<br />

stem no more than 11/4 inches thick or 3 inches<br />

tall—41.<br />

41a. Radial spines 20 to 30, gray or straw-colored<br />

tipped with brown; centrals 5 to 10, 3/8 to 5/8 of<br />

an inch long and dark brown, red-brown, or<br />

black for most of their lengths; stigma lobes<br />

green —M. roberti.<br />

41b. Radial spines 24 to 85 and all white in color;<br />

centrals 6 to 22, 3/8 of an inch or less in length;<br />

stigma lobes white—42.<br />

42a. Radials 24 to 45 —M. sneedii.<br />

42b. Radials 40 to 85 —M. leei (in part).<br />

36b. Radial spines soft, flexible, and hairlike<br />

—M. multiceps.<br />

35b. Centrals absent; radials 40 to 80 in number, slender and<br />

bristle-like but rigid—43.<br />

43a. Spines pubescent under magnification —M. lasiacantha<br />

var. lasiacantha.<br />

43b. Spines smooth and naked under magnification<br />

—M. lasiacantha var. denudata.<br />

Mammillaria scheeri Muehlenpf.<br />

[Coryphantha scheeri Lem.; Coryphantha muehlenpfordtii<br />

(Poselgr.) B. & R.]<br />

“Long-Tubercled Coryphantha,” “Needle ‘Mulee’”<br />

Description Plate 28<br />

stems: Spherical at first, becoming egg-shaped or somewhat<br />

conical when old. These stems grow to 9 inches tall by 51/2<br />

inches in diameter. They are usually single, but sometimes<br />

have one or two branches at the base. The surface is bright<br />

green or even yellowish-green. It is formed into large, rather<br />

soft, spreading tubercles. These arise from broad bases often<br />

3/4 of an inch or more across, becoming cylindrical above and<br />

reaching an over-all length of 3/4 to 11/2 inches.<br />

areoles: Monomorphic. When young the areoles are very<br />

woolly. Upon maturing they consist of a roundish spinous<br />

portion at the tip of the tubercle and a long, narrow, deep,<br />

groovelike portion running part or all of the way down the<br />

upper side of the tubercle. In the groove are often one or<br />

more brownish glands.<br />

spines: 8 to 16 stout to very stout radial spines radiating or<br />

slightly spreading outward. These are from 1/2 to about 11/4<br />

inches long, the lower ones the heaviest and longest. They<br />

are round and proceed from slightly bulbous bases. There are


genus Mammillaria 115<br />

also 1 to 5 central spines 3/4 to 11/2 inches long, stouter than<br />

the radials. One is very stout and stands porrect. This spine<br />

may be straight, curved downward slightly over its whole<br />

length, or in some individuals straight with a hooked tip. The<br />

other centrals are more nearly like the radials and spread upward<br />

in front of them. When growing, all spines are reddish<br />

or brownish with blackish tips. On some specimens they remain<br />

for a long time yellowish—especially the main central—<br />

but on others they very soon become ashy-gray.<br />

flowers: About 2 inches long and wide. Orange, bronze, or<br />

bronzy-yellow at first, darkening to reddish as they wilt. The<br />

outer perianth segments are lacerated and sometimes this effect<br />

on their edges approaches a true ciliated fringe. The 20<br />

or so inner segments are narrow below, widening to their<br />

broadest near the tips, which are often somewhat pointed and<br />

also toothed. The stamens are orange, the style short and<br />

with 6 to 10 yellowish or flesh-colored stigma lobes.<br />

fruits: Egg-shaped to almost club-shaped, 1 to 11/2 inches<br />

long. The surfaces are smooth. They remain greenish. The seeds<br />

are dark red-brown with shiny, smooth surfaces. They are<br />

about 1/8 of an inch long and are flattened ovate.<br />

Range. Extreme west Texas and the southern edge of New<br />

Mexico into Arizona and Chihuahua. Specifically, from the<br />

Pecos River near where it enters Texas to the Davis Mountains,<br />

Marfa, and Presidio, Texas, and west. In New Mexico apparent-<br />

ly not north of the Organ Mountains, but found from there west<br />

into Arizona.<br />

Remarks. This cactus was well described by Engelmann as a<br />

“stately plant.” It is the largest in bulk of the Mammillarias in<br />

our area. A fine old specimen actually gives more of an impression<br />

of a barrel cactus than of this group. But unless we are<br />

aware of the size which is attained by some of the Mammillarias<br />

in Mexico we may have an erroneous concept of the Mammillarias<br />

as all small, delicate cacti. It is fortunate that this large<br />

species enters our area to set things straight.<br />

However it is unfortunate that so few people see this fine<br />

cactus. Although it has quite a large range in the United States<br />

it is nowhere common. I have collected it numbers of times in<br />

widely distant places, but I have never seen a stand of this<br />

species, and, in fact, never collected more than one plant at any<br />

given location. The small number of specimens of this species in<br />

any dealer’s stock makes it seem likely that the plant nowhere<br />

grows in quantity. So it remains a little-known species highly<br />

prized by the cactophile who happens to possess one.<br />

Perhaps the scarcity of the plant has led students over the<br />

years to write about it from insufficient observation. Certainly,<br />

the naming of this species has been royally confused—with the<br />

confusion persisting right down to the present.<br />

Muehlenpfordt apparently started things off in 1847 by describing<br />

a Mexican plant and naming it Mammillaria scheeri.<br />

Unfortunately the type of his plant is unknown. When Engelmann<br />

studied the cacti of Texas and wrote up this cactus he was<br />

not at first sure that he had the same plant, but knew that he<br />

had something very similar, so he called the Texas plants M.<br />

scheeri var. valida. However, after he had seen Mexican specimens<br />

he became convinced that there was only one form involved<br />

and stated in “Corrections to the Cactaceae of the<br />

Boundary” that the variety agrees exactly with Mexican specimens<br />

of M. scheeri, and so seems to have withdrawn the idea<br />

of a separate variety.<br />

Even before this, however, in 1853, Poselger had redescribed<br />

the cactus as an Echinocactus. By an interesting coincidence he<br />

used the name of the original describer for it. Poselger’s name,<br />

therefore, came out Echinocactus muehlenpfordtii. This name<br />

Engelmann, Lemaire, Coulter—everyone in that century, and<br />

even Wooton and Standley in this—ignored. They all regarded<br />

the original name of M. scheeri as valid and used it. Lemaire<br />

very early split this cactus off from the Mammillarias as a Coryphantha,<br />

but even in doing this he called it Coryphantha scheeri.<br />

But when Britton and Rose made their big break with tradition<br />

they broke cleanly. They resurrected Poselger’s name (when<br />

he mistook the species for an Echinocactus) and rechristened<br />

the cactus Coryphantha muehlenpfordtii. This is therefore the<br />

name for the cactus which these great popularizers of the cacti<br />

have caused to be the most familiar in recent years.<br />

But the return of this species to the genus Mammillaria cannot<br />

be merely a shifting to Mammillaria muehlenpfordtii, since<br />

Forster, in 1847, used that name in a way that makes it a synonym<br />

for Mammillaria celsiana Lem., a Mexican species. We see<br />

no problem in returning to the first name of all, which was<br />

Mammillaria scheeri, but L. Benson did. He returned this species<br />

to its original genus, but in his 1950 work on Arizona cacti<br />

he chose to use still another obscure name for the species, a<br />

name which had been coined by Cory and which Marshall had<br />

mentioned. It was Coryphantha engelmannii, and Benson’s<br />

name for the plant, therefore, became Mammillaria engelmannii<br />

(Cory) Benson. The reason for this latest substitution may be the<br />

fact that the type of Muehlenpfordt’s M. scheeri is lost, but if<br />

we drop all original names for all species of which the type is<br />

lost, the roster of botanical names will be greatly altered.<br />

One may take his pick among all of these names and have the<br />

cactus as a memorial to any one of those early cactus students<br />

he wishes, but whichever name finally prevails, the plant is a<br />

large, beautiful, and proud cactus which must have a dry, sunny<br />

situation, but which if treated properly contributes a spate of<br />

remarkably beautiful flowers of a color unusual among the<br />

cacti.<br />

A closely related form is Mammillaria robustispina Schott.<br />

This is very well named, since it has the stoutest spines of any<br />

Mammillaria in the United States. Its spines actually equal those<br />

of many Echinocactus horizonthalonius specimens in thickness.<br />

Benson’s 1950 description of M. robustispina is so broad that<br />

some Texas specimens of M. scheeri could fit it, and Earle has<br />

recently fallen into this trap, combining it with our species as<br />

Coryphantha muehlenpfordtii var. robustispina. Benson’s dis-


Bodeker -> Boedeker<br />

116 cacti of the southwest<br />

tribution map of M. robustispina implies that its range enters<br />

New Mexico, and Earle states its distribution to include southwestern<br />

New Mexico. I do not know whether there has been a<br />

referral of some ordinary New Mexico M. scheeri specimens to<br />

this other form or not, but I do know that the real Mexican<br />

M. robustispina specimens which I have seen are very different<br />

from our cactus, and I can find no record of that form’s having<br />

been collected in New Mexico.<br />

Mammillaria scolymoides Scheidweiler<br />

[Coryphantha scolymoides (Scheidweiler) Boedeker]<br />

Description Plate 28<br />

stems: Single, spherical to conical, and to at least 6 inches<br />

tall and 4 inches in diameter. The surface is deep bluish-green,<br />

with some grayish glaucescence. The tubercles are to 1 inch<br />

long, tapering from bases as wide as the length, firm, and<br />

overlapping upward.<br />

areoles: When mature the areoles are prolonged beyond the<br />

spinous part into deep grooves extending all the way to the<br />

bases of the tubercles. These grooves are bare except for a tuft<br />

of wool at the base of each, in the axil of the tubercle.<br />

spines: Radial spines 14 to 28 in number, 3/4 to 11/8 inches<br />

long, evenly radiating except at the top of the areole where<br />

they are so crowded together as to be bunched into a bundle.<br />

There are 1 to 4 heavy central spines, usually 3 of them turned<br />

upward and spreading in front of the upper bundle of radials<br />

and the lower one standing out conspicuously and<br />

curving downward from the middle of the areole. These centrals<br />

are all about 1 inch long. All spines are yellowish and<br />

hornlike when young, then gray with dark tips.<br />

flowers: About 2 inches long, clear yellow at first and later<br />

darkening to a reddish color, but not having a red center. The<br />

outer petals are short, pointed, with entire edges, and total<br />

about 20 in number. The inner petals are similar except longer,<br />

wider, and about 35 in number. The filaments are pinkish,<br />

the anthers pale orange-yellow. The stigma has 9 rough, yellowish<br />

lobes about 1/4 of an inch long.<br />

fruits: Not seen. They failed to develop during the four years<br />

that a number of the plants bloomed in my garden.<br />

Range. Some of the higher elevations of the Big Bend of Texas:<br />

most common in the Glass Mountains near Alpine. There was<br />

an early collection made near the lower Pecos River which was<br />

referred to this species.<br />

Remarks. This form has been a source of much confusion, and<br />

its position in the classification is still not settled, but it does<br />

appear as a definitely recognizable form separate from anything<br />

else found in Texas, and so must be listed here. Scheidweiler<br />

first described it in 1841 from specimens collected in<br />

Mexico, and Engelmann followed him. Then later a small plant<br />

was collected on the Pecos River in Texas which Engelmann<br />

classified as M. scolymoides. Coulter saw plants from various<br />

locations in Texas, New Mexico, and Mexico which he put<br />

under this name. Both Engelmann and Coulter, however, felt<br />

enough uncertainty about their classifications to suggest the<br />

possibility that their plant might be only a form of Mammillaria<br />

cornifera DC, a species which occurs in Mexico. Schumann<br />

relegated it to Mammillaria radians DC, another Mexican species.<br />

Britton and Rose took Engelmann’s and Coulter’s suggestions<br />

as certainty and merely listed it as a synonym of Coryphantha<br />

cornifera without describing it, so it is almost unknown<br />

since their time. This may be proper, or perhaps it may end up<br />

as Mammillaria cornifera var. scolymoides, but regardless of the<br />

name, this is a cactus which west Texas collectors are bound to<br />

find and wonder about.<br />

The cactus is fully as near to Mammillaria echinus as to any<br />

other species. Small specimens are almost indistinguishable from<br />

that cactus except for their longer and heavier spines and tubercles.<br />

Many have been distributed under that name. But anyone<br />

who has ever seen a large, mature specimen cannot but<br />

recognize the difference between them. M. scolymoides grows<br />

at least twice as large as M. echinus and in its prime is fully as<br />

tall and as impressive as M. scheeri, only remaining a little<br />

smaller in diameter than that species. In regard to maximum<br />

size, my description exceeds that of Engelmann, which has been<br />

the one copied by most other students, because Engelmann apparently<br />

saw only a small specimen and had no idea of the size<br />

it can attain.<br />

All specimens I have collected have come out of the Glass<br />

Mountains, a small range of mountains northeast of Alpine,<br />

Texas. Although I have seen many specimens in dealers’ stocks<br />

which apparently came from other mountains deeper in the Big<br />

Bend, I have been unable to trace them any more exactly, since<br />

they were brought out of these mountains by crews of laborers<br />

who roam these areas digging cacti for the trade.<br />

I have collected two specimens of this cactus which, although<br />

showing all the other characters of the species, lacked the central<br />

spines. One of them had been badly damaged and was recovering<br />

from a serious wound on one side. Later, under cultivation,<br />

both of them put on normal centrals on new growth. I<br />

believe, therefore, that this is an abnormal growth form reverting<br />

to the immature condition under bad situations. As such it<br />

would be parallel to the same sort of reversion which occurs in<br />

the species M. echinus and which was the basis for the supposed<br />

species M. pectinata.<br />

Mammillaria echinus Eng.<br />

[Coryphantha echinus (Eng.) B. & R.]<br />

Description Plate 28<br />

stems: Usually spherical, but sometimes becoming egg-shaped


genus Mammillaria 117<br />

or conical when old. The plants almost always consist of<br />

single stems, but sometimes branch at the base when old to<br />

form small clumps of up to about 6 stems. These stems are 2<br />

to 3 inches tall when mature. They are covered with very<br />

firm, conical tubercles about 3/8 to 1/2 inch long, which turn<br />

upward and somewhat overlap those above them.<br />

areoles: At first nearly round, remaining so on some specimens<br />

until the plants are very large. But when matured normally<br />

the areoles of the plants are elongated by a groovelike<br />

extension which may extend only part of the way or all of<br />

the way to the base of the tubercle. The flower is produced<br />

from the end of this groove. There is white or brownish wool<br />

in the grooves at the tip of the plant where they are growing,<br />

but on the sides of the plant the grooves are bare except for<br />

a tuft of wool remaining at the base of each groove.<br />

spines: There are 16 to 30 radial spines growing around the<br />

edge of each areole. They are 3/8 to 1 inch long, and radiate<br />

evenly all around the areole except at the upper edge, where<br />

they are more numerous and bunched. They lie flat against<br />

the plant or even curve back toward the plant a little, interlocking<br />

with spines of neighboring areoles. They are slender,<br />

but very rigid, round or sometimes slightly flattened from<br />

side to side. When young they appear yellowish and partly<br />

translucent like the material of horn, and when old they become<br />

gray. They usually have black or dark brown tips. There<br />

are also 3 or 4 much thicker central spines growing from the<br />

center of the typical adult plant areole. These have enlarged<br />

bases, are round, and extend about 1/2 to 3/4 of an inch in<br />

length, usually being gray with black tips; but a plant is occasionally<br />

found on which these centrals are black almost all<br />

the way to their bases. Two or three of these spines are turned<br />

upward so that they lie directly upon the upper radial spines.<br />

The lowest central spine always stands straight out from the<br />

center of the areole. It is thick and conspicuous, usually perfectly<br />

straight, but sometimes curves downward. Immature<br />

plants lack the central spines entirely, having only the radial<br />

spines, and sometimes poor growing conditions will keep a<br />

plant from growing centrals when it is otherwise mature. An<br />

injury or a reversal of good conditions will sometimes cause<br />

new growth on a typical plant to be without these centrals.<br />

flowers: At least 3 inches across and to 2 inches tall, the<br />

color a very clear sulphur yellow. The outer petals are narrow<br />

and brownish-green, with yellowish, smooth edges. The 20 to<br />

30 inner petals are about 3/8 to 5/8 of an inch wide, pointed,<br />

with the edges slightly ragged toward the tips. They are clear<br />

yellow in color. The filaments are rose-pink. The anthers are<br />

bright orange. The style is about the length of the stamens.<br />

The stigma is made up of 10 to 12 rough, cream-colored lobes<br />

about 3/16 to 1/4 of an inch long.<br />

fruits: Oval or egg-shaped, to nearly 1 inch long, light green<br />

in color. The remains of the flowers stay upon them. The seeds<br />

are dark brown, smooth surfaced, kidney-shaped, and about<br />

11/2 millimeters long.<br />

Range. An area of southwest Texas from near the mouth of the<br />

Pecos River up to about the level of Fort Stockton, and from<br />

there southwest into the mountains of the lower Big Bend area.<br />

I find no record of its occurrence west of Marfa, Texas.<br />

Remarks. This is a very common little cactus in a rather limited<br />

area of west Texas. It seems to be confined to the eastern part<br />

of the Big Bend. Early students collected it as far east as the<br />

lower Pecos River, but it seems not to have been seen that far<br />

east for many years.<br />

The cactus, when typical, sits as a small sphere or cone only<br />

2 or 3 inches high. Its spines cover it and enclose it completely.<br />

On each areole there is one heavy central spine up to 1 inch<br />

long standing straight out, perpendicular to the surface of the<br />

plant, or nearly so. This gives the cactus a striking resemblance<br />

to the Echinoderm called the sea urchin, and for this reason<br />

Engelmann gave it the name it bears.<br />

Early descriptions of the species always listed it as not clustering,<br />

but dealers have been bringing many specimens out of<br />

the more inaccessible mountains of the lower Big Bend where<br />

they must have better conditions, and these are sometimes<br />

branched around their bases to form small clusters of up to half<br />

a dozen heads.<br />

A hundred years ago, when Engelmann first studied these<br />

cacti, he described Mammillaria echinus very plainly. At the<br />

same time he also described another form identical in every way<br />

to this one, except having no central spines. He called this form<br />

M. pectinata, from the comblike appearance of the radial spines.<br />

From that time on there has been discussion by every student of<br />

these two forms, some stoutly maintaining them to be separate<br />

species and others trying to combine them as varieties.<br />

As a result of growing these cacti and observing them over a<br />

period of years, I have satisfied myself that the form called<br />

M. pectinata is only an immature or abnormal growth form of<br />

M. echinus, and that the names are, therefore, synonymous. Immature<br />

specimens of M. echinus under about 1 inch in diameter<br />

always lack central spines and are identical to M. pectinata in<br />

armament. Central spines normally appear gradually on new<br />

areoles of these plants as they grow past 11/2 inches or so in<br />

size.<br />

But what about larger specimens and even blooming specimens<br />

described by Engelmann as having no centrals, on which<br />

he based his M. pectinata? This sort of thing has been seen by<br />

any student who has looked over any number of these plants,<br />

since it is fairly common among them. But I have found that a<br />

normal M. echinus with large centrals can often be caused to<br />

put out new growth lacking centrals merely by inflicting an<br />

injury on it, or giving it poor growing conditions. For example,<br />

I have a plant which I bought from a dealer as M. pectinata. It<br />

had 3 spherical heads, each about 11/2 inches across, and on no<br />

areole was there a central spine. But upon looking closely I


118 cacti of the southwest<br />

could see that these three heads grew from a single short base.<br />

This was the remains of what had once been a large M. echinus<br />

which had suffered so severe an injury to its upper part that<br />

only this short section had been left. But it had unmistakably<br />

been M. echinus, since the old spines of it, when dug out from<br />

under the edges of the new growth, had a full set of centrals in<br />

each cluster. The new growth it had put out after its injury<br />

consisted of 3 perfect miniature heads with no centrals. Now,<br />

two years later, these new heads have reached a size where they<br />

are maturing, and the first centrals are showing up upon them,<br />

proving that when grown they will form a triple-headed M.<br />

echinus which has grown from the one form to the other twice<br />

in its lifetime.<br />

In confirmation of this, I have a specimen of typical M. echinus<br />

collected near Marathon, Texas. It has been growing in a<br />

pot for several years. Something in the transplanting or the lack<br />

of some necessary materials since has caused this one plant to<br />

put out growth entirely without centrals for the past two years.<br />

In other words, the base of this plant is now M. echinus and its<br />

summit is now M. pectinata. It has been blooming every year,<br />

proving that this growth without centrals, although typical of<br />

the immature, can also occur on mature, flowering plants. It<br />

seems obvious, then, that the two forms are one and the same<br />

cactus.<br />

Mammillaria sulcata Eng.<br />

[Coryphantha sulcata (Eng.) B. & R.]<br />

“Nipple Cactus,” “Finger Cactus,” “Pineapple Cactus”<br />

Description Plate 28<br />

stems: Rapidly and densely clustering, new stems growing<br />

from the grooves on old tubercles around the base of the<br />

plant. Old plants are often masses several feet across, and<br />

made up of dozens of heads. Individual stems are spherical,<br />

usually with somewhat flattened tops, and often wider than<br />

tall. Mature stems will be 3 inches across and about 11/2 to<br />

3 inches tall. The surface is dark green and soft, divided into<br />

tubercles which are nearly cylindrical, the bases on old ones<br />

becoming somewhat broadened. They curve upward to overlap<br />

slightly on old specimens, but it is more common for them<br />

to stand out from each other. Because they are very soft,<br />

when the plant is shrunken from lack of water they often sag<br />

downward and give the plant an untidy, irregular appear-<br />

ance. Mature tubercles are to about 3/4 of an inch long.<br />

areoles: Practically round and very woolly at first and remaining<br />

so on immature growth. On mature growth they are<br />

elongated by the formation of groovelike extensions which<br />

reach to the axils of the tubercles. These grooves are filled<br />

with much white or yellowish wool when young, making the<br />

top of the plant woolly, but they are almost bare when old.<br />

spines: There are 8 to 15 radial spines 3/8 to 5/8 of an inch<br />

long, which radiate evenly from the edge of the areole. They<br />

are round and rather heavy, yellowish and partly translucent<br />

at first, then gray with black or dark red-brown tips. There<br />

may be no central spine, or to as many as 3 per areole, this<br />

much variation often being found on the same plant. If the<br />

central is single, it usually stands perpendicular to the stem.<br />

If two are present, one is usually perpendicular or turned a<br />

little downward, while the other is turned upward. When<br />

three are present the one is still in the perpendicular or downturning<br />

position, while the other two spread upward, sometimes<br />

almost against the upper radials. The centrals are the<br />

same length as the radials, but heavier. They are usually the<br />

same color, but on occasional plants they will be streaked<br />

with black almost to their bases on only the upper sides.<br />

flowers: 2 or 3 inches across and 1 to 2 inches tall. They<br />

vary from a greenish-yellow to a dark golden-yellow. The<br />

centers are usually very bright red, but on rare specimens this<br />

red is a paler brownish-red or even almost nonexistent. The<br />

outer petals are short and greenish with yellowish, smooth<br />

edges. There are 20 to 30 of them. The inner petals are long<br />

and rather narrow from narrow, red bases, the upper part<br />

being golden and the same width as the outer petals, tapering<br />

gradually to a definite point. The upper margins may be entire<br />

or sparingly ragged and toothed. There are about 25 inner<br />

petals. The filaments are bright red, except in the rare flowers<br />

almost without red, where they are greenish. The anthers are<br />

yellow. The style is equal in length to the stamens on some<br />

plants and somewhat longer on others, greenish, and ending<br />

in 7 to 10 yellow, greenish-yellow, or cream-colored lobes.<br />

fruits: Oblong, green, about 3/4 of an inch long. The seeds<br />

are light brown, about 1/16 of an inch (11/2 millimeters) long,<br />

with the surface pitted.<br />

Range. South-central Texas within a triangle the points of<br />

which lie just north of Houston on the east, near Fort Worth<br />

on the north, and near the Pecos River on the west.<br />

Remarks. This cactus does not present any striking form of<br />

growth. Its stems are not large, and even the old clumps which<br />

may be up to two feet or more across are low, uneven clusters<br />

presenting no particular beauty of appearance. It likes to grow<br />

under the junipers, oaks, and brush plants of the limestone hills<br />

at the edge of the Edwards Plateau. It is rarely found in the<br />

more open country farther north, but there are a few records<br />

of its having been collected to near Fort Worth. It apparently<br />

never grows on the coastal plain or in deep south Texas.<br />

If undistinguished ordinarily, this cactus shows its beauty<br />

when it blooms. Its flowers are large and usually of a satiny,<br />

golden color unmatched by any other species I have ever seen.<br />

With the red centers and sheaves of red filaments these make<br />

some of the finest blooms on any of our cacti.<br />

Very rarely one comes across a plant of M. sulcata which has<br />

a more greenish-yellow flower with the red center almost com-


genus Mammillaria 119<br />

pletely lacking and only represented by a faint brownish-red<br />

shading at the very base of the petals. I believe that this is the<br />

cactus which was at hand when early students described it first<br />

as Mammillaria similis robustior Eng., later as M. wissmannii<br />

Hild., and still later as Neobesseya wissmannii (Hild.) B. & R.<br />

I therefore present the probability that these names are all synonyms<br />

of M. sulcata. See the discussion under Mammillaria si-<br />

milis for my reasons for this suggestion.<br />

The beauty of its flowers, together with the fact that it is<br />

not strictly a desert species, should make this cactus the logical<br />

choice for dish gardens and windowsill growing. It does not rot<br />

so easily as most, and so can stand much more water in the soil<br />

and humidity in the air. If given a limestone soil it should do<br />

well and bloom in almost any situation.<br />

After giving this cactus the name M. sulcata in his original<br />

description, Engelmann later renamed it M. calcarata, but it<br />

must be called by its earliest name.<br />

Mammillaria ramillosa (Cutak)<br />

[Coryphantha ramillosa Cutak]<br />

Description Plate 29<br />

roots: Entirely fibrous.<br />

stems: Single or very sparingly clustering, to 31/2 inches in<br />

diameter, spherical or nearly so. The tubercles are about 3/4 of<br />

an inch long, tapering from very much flattened bases which<br />

are wider than the tubercles are tall. These tubercles are firm,<br />

are arranged regularly, and overlap upward very much. The<br />

color is dark green.<br />

areoles: Monomorphic, becoming linear and extending as<br />

grooves to the axils of the tubercles. Woolly at first, but bare<br />

when mature except for a small tuft of wool at the end of<br />

each groove in the axil of the tubercle.<br />

spines: There are 14 to 20 slender radial spines 3/8 to about<br />

1 inch long. They radiate evenly around the areole, are flattened,<br />

usually curving and twisted, and gray in color, often<br />

with darker tips. There are 4 central spines which are also<br />

slender, but which are round, 1 to 15/8 inches long, gray mottled<br />

with brown, and spreading and curving in all directions.<br />

The radials are pinkish when growing and the centrals winecolored.<br />

flowers: Varying from pale pink to deep rose-purple in color.<br />

They are about 21/2 inches long and 2 inches wide, often unable<br />

to open so fully because of the long spines around them.<br />

The outer petals vary from small, green scales on the flower<br />

tube to greenish-purple, linear, unfringed segments. The inner<br />

petals are about 1 inch long and to 3/16 of an inch wide, the<br />

lower half white, while the upper half is pink to rose-purple.<br />

The filaments are white, the anthers pale orange. The style is<br />

about the length of the stamens. There are 6 or 7 stigma lobes<br />

1/8 to 1/4 of an inch long.<br />

fruits: 3/4 to 1 inch long, oval or egg-shaped, green, covered<br />

with minute white or silvery, hairlike scales which give them<br />

a silvery sheen. The old flower parts remain upon them. The<br />

seeds are brown, less than 1/16 of an inch (1 millimeter) long,<br />

very broad above with flattened sides below and a long, nar-<br />

row point of attachment toward the end of the ventral side.<br />

Range. The southeastern corner of Brewster County, Texas,<br />

from near the Stillwell Ranch along the Rio Grande to below<br />

Sanderson, Texas. Also found in a very small area close to the<br />

Rio Grande within the Big Bend National Park, just down-<br />

stream from Mariscal Canyon.<br />

Remarks: This cactus presents a dilemma to those who would<br />

subdivide the genus. By all of its vegetative characters it is a<br />

relative of the large, yellow-flowered Mammillarias just listed,<br />

which are often called Coryphanthas, yet its large flower is<br />

definitely purplish and reminds one of Mammillaria macromeris,<br />

which is placed by Backeberg in a separate genus, Lepidocoryphantha.<br />

Most have been inclined to regard it as a closer<br />

relative of the latter cactus, but if two separate genera are made<br />

here, this cactus will hardly fit into either of them: the purple<br />

flower looks odd in the one and the plant lacks the fringed<br />

sepals and the taproot of the other. When one recalls that the<br />

yellow-flowered Mammillaria scheeri of the first group has<br />

more or less fringed sepals anyway, the only conclusion seems<br />

to be that any division on the basis of fringed or entire sepals or<br />

of flower color is certainly less than generic in significance.<br />

M. ramillosa is distinguished from the purple-flowered M.<br />

macromeris, which grows close by it, by the facts that M. ramillosa<br />

forms firm, compact, spherical, squatty stems, while<br />

M. macromeris is more cylindrical in shape, with longer, looser,<br />

flabbier tubercles. The spines of the former are more slender,<br />

especially the round centrals. It has only small fibrous roots,<br />

instead of the large taproot of the other, and never forms the<br />

large carpets of plants which that other species does, even in<br />

its most favorable locations. Most specimens I have seen of<br />

M. ramillosa have been single stems, but sometimes it clusters<br />

sparingly.<br />

The significance of the minute hairs on the fruit surface has<br />

not been determined. The fruit was not seen by the original describer,<br />

and so far as I know these hairs have not been noticed<br />

before. They are too tiny to be easily visible to the naked eye<br />

except by the silvery sheen they impart to the whole surface of<br />

the fruit. They are, however, so numerous that the surface of the<br />

fruit approaches the condition described as being hirsute. I have<br />

seen such a covering of hairs on no other cactus fruit. Because<br />

they are so tiny I find it hard to think of them as scales or hairs<br />

in the same sense as those on the fruits of the Echinocacti. If<br />

one did equate them with those this would open up a whole<br />

new realm of speculation for the theorists.<br />

M. ramillosa was discovered in 1936 by Mr. A. R. Davis, a


120 cacti of the southwest<br />

great collector of the Big Bend cacti. It was described by Ladislaus<br />

Cutak in 1942. It grows on the limestone hills along the<br />

Maravillas and Reagan canyons of southeastern Brewster County<br />

and along a few other smaller canyons farther east. This is<br />

very inaccessible territory, and very few of the cacti have been<br />

brought out, so that the plant is almost unknown even today. It<br />

can be most easily seen in a small location in the Big Bend<br />

National Park where it is fortunately protected. This seems to<br />

represent a second location for the plant and is perhaps the<br />

second and discontinuous range mentioned by Davis.<br />

The cactus is a rather attractive one, presenting a tidy and<br />

symmetrical growth. Its slender spines, however, are extremely<br />

untidy, curving and spreading at all angles, and making it look<br />

much like a bunch of dead, gray grass stems, which must be its<br />

particular form of camouflage.<br />

Mammillaria macromeris Eng.<br />

[Coryphantha macromeris (Eng.) Lem.]<br />

“Long Mamma”<br />

Description Plate 30<br />

roots: Adult plants have long, fleshy taproots.<br />

stems: Usually short columns to about 4 inches tall, but<br />

sometimes to 8 inches; to about 3 inches thick. These stems<br />

may remain single, but usually cluster by the formation of<br />

new stems from the grooves on the lower tubercles. The clusters<br />

often form flat mats of stems, but occasionally round<br />

up to form hemispherical mounds. Seeds often germinate and<br />

grow close around the bases of mature plants. This, with the<br />

branching, sometimes produces masses of the plants up to sev-<br />

eral feet across. The stems are made up of very soft, flabby,<br />

long tubercles, arranged regularly and standing upright at the<br />

top of the stem, but usually flattened and either curving upward<br />

or curving and standing out at various angles on the<br />

sides of the stems. These tubercles are 3/4 to 11/2 inches long,<br />

cylindrical, and dark green in color.<br />

areoles: Each tubercle on an adult plant is grooved with a<br />

deep groove which usually runs about halfway down the tubercle<br />

length, but sometimes goes all the way to the axil. This<br />

groove is almost entirely bare when old, but has a little short<br />

white wool in it when young. It is the linear extension of the<br />

areole forward from the spinous portion, and the flowers, or<br />

lower on the stem the new shoots, arise from within this<br />

groove.<br />

spines: There are 10 to 18 slender radial spines. They are 5/8<br />

to 2 inches long, the upper ones tending to be the longer ones.<br />

They are gray in color, sometimes mottled with pale brown.<br />

They are flattened and ridged and usually twisted and curved.<br />

There are also 2 to 8 heavier central spines on mature<br />

plants, these 1 to 21/4 inches long, spreading in all directions<br />

from the center of the areole. They are mottled brown, dark<br />

brown, purplish, or black. They grow angled and often<br />

ridged, either straight or twisted and curved, from bulbous<br />

bases.<br />

flowers: Light purple or pinkish in color, 2 inches tall and<br />

wide. These flowers would usually open wider if they were<br />

not prevented from opening fully by the long spines around<br />

them. The green ovary has several white-edged, fringed scales<br />

upon it. The 20 to 30 outer petals are short, narrow, and<br />

greenish-brown with pink, fringed edges. The 20 to 25 inner<br />

petals are about 11/4 inches long and 3/16 of an inch wide, the<br />

midlines deep pink and the edges almost white. These edges<br />

are rough and often fringed, and the ends, while pointed, are<br />

also toothed and notched. The filaments are rose-colored, the<br />

anthers pale orange. The style is a little longer than the stamens.<br />

The stigma has 7 to 10 rough, white lobes about 3/16 of<br />

an inch long. These lobes are slender, and each has a smooth,<br />

pinkish point like a soft spine on the end.<br />

fruits: Almost round or broadly egg-shaped, 5/8 to 1 inch<br />

long, greenish in color. The seeds are practically spherical,<br />

light brown in color, and smooth.<br />

Range. Southern New Mexico into Mexico and southeast along<br />

the Rio Grande in Texas to Rio Grande City.<br />

Remarks. This cactus is very common around Las Cruces, New<br />

Mexico, and is easily found around Presidio, Texas. Otherwise<br />

it occurs sporadically over a large area, including the area<br />

along the course of the Rio Grande where it becomes rather<br />

common again as far as Rio Grande City, Texas. It grows on<br />

the stony crests and sides of low hills, where it is almost always<br />

found under the light shade of shrubs and trees. In such places<br />

it sometimes forms mats of closely standing plants extending<br />

over a number of square feet.<br />

M. macromeris is remarkable for its long, soft, loosely standing<br />

tubercles. These, however, give it a rather untidy appearance,<br />

just the opposite of the neat, symmetrical body we admire<br />

in most cacti, and cause it to be less of a favorite with cactus<br />

fanciers than some others. It does have a beautiful flower, how-<br />

ever, and can be a very fine pot plant.<br />

This cactus shares with many others the habit of budding new<br />

stems from the grooves of old tubercles. There is nothing unusual<br />

about this, since the vegetative meristem is in these monomorphic<br />

areoles located at or near the end of this elongated<br />

areole. But the grooves usually extend only halfway or so down<br />

the long tubercles in this plant, and the new little stems appear<br />

quite out of place bursting out of the grooves only a short distance<br />

below the ends of the tubercles instead of out of the axils.<br />

There have been attempts to make much of this peculiarity, even<br />

to using it to set this species off in its own new genus, this<br />

called by Backeberg genus Lepidocoryphantha. But sober reflection<br />

makes it seem obvious that this is no really essential<br />

difference from any of the other Mammillarias having monomorphic<br />

areoles. In fact, Mammillaria sulcata sometimes has


genus Mammillaria 121<br />

similarly short areoles and in this case branches in the same<br />

way.<br />

There have been described some varieties of M. macromeris,<br />

but the cactus is very variable, and it is hard to separate forms<br />

exactly. Specimens with light central spines, longer and straighter<br />

than the average, seem to be the basis for the variety longispina<br />

of Schelle, and specimens with black or very dark, shorter,<br />

and more twisted and curved centrals may be his variety nigrispina,<br />

but they both grow together, and the differences seem of<br />

doubtful significance. The plants do seem to become smaller in<br />

their maximum size in the eastern part of their range, and also<br />

to average fewer centrals than those farther west, but the flowers<br />

are identical, and no varieties have been set up for this<br />

difference.<br />

Engelmann stated that Mammillaria dactylothele Labouret<br />

was a variety of this species, but Britton and Rose consider it a<br />

synonym.<br />

M. macromeris will grow very well if given perfect drainage<br />

and enough heat and sunlight. It blooms well when healthy, but<br />

one should remember that it needs room to develop its long taproot,<br />

so will not do well in a shallow pot. When specimens are<br />

collected the long roots are often broken off, as they grow deep<br />

into very hard and rocky soil. The plants may live after this,<br />

but will usually sit without growing or flowering for years until<br />

they can replace their taproots. This is why large plants of this<br />

species are often so disappointing in performance after being<br />

brought in.<br />

Mammillaria runyonii (B. & R.)<br />

[Coryphantha runyonii B. & R.]<br />

“Runyon’s Coryphantha,” “Dumpling Cactus”<br />

Description Plate 30<br />

roots: Having a large, succulent, carrot-shaped taproot.<br />

stems: Highly caespitose by producing many irregularly sized<br />

and shaped stems from the top of the taproot, the whole mass<br />

of these stems sometimes up to 18 inches or more across. The<br />

tubercles are so irregularly arranged on most specimens that<br />

one can hardly speak of separate stems at all, but on other<br />

plants the mass of tubercles is divided roughly into irregular<br />

heads up to about 2 inches tall and 11/2 inches thick. The<br />

tubercles, which stand at almost any angle, vary in size on<br />

the same plant, but most of them are around 3/8 of an inch<br />

long with the maximum being about 3/4 of an inch long. They<br />

are very soft, tapering from very broad bases which are<br />

often wider than the tubercle is long.<br />

areoles: The spinous portion of the areole is small and practically<br />

circular, but the mature areole is prolonged from this<br />

in the form of a groove running about halfway down the<br />

tubercle, but never to the axil. The flower is produced from<br />

the end of this short groove which is naked after completion.<br />

spines: Usually there are 6 to 11 radial spines, but there is a<br />

tendency for some individuals to produce very many very<br />

small tubercles, each having as few as 3 radials. The radials<br />

are usually 1/4 to 1/2 inch long, but may occasionally grow to<br />

1 inch long. They are evenly spaced around the areole, are<br />

straight and round, and yellowish in color at first, then turning<br />

gray. There may be 1 to 4 straight, round central spines<br />

3/4 to 2 inches long, which spread in all directions from the<br />

center of the areole. These are gray mottled with brown.<br />

flowers: Purplish or rose-pink, 13/4 inches tall and 2 inches<br />

in diameter, opening widely, the petals sometimes even recurving.<br />

The outer petals are greenish or purplish, covered<br />

and fringed with white hairs. The inner petals are about 3/16<br />

of an inch wide, long and pointed, the edges smooth except<br />

for being somewhat toothed toward the tips. The filaments<br />

are pinkish, the anthers pale orange. The style is a little longer<br />

than the stamens, and pale pink. The stigma has 6 to 10 lobes<br />

which are white, 1/8 to 3/16 of an inch long, fairly thick, and<br />

not pointed.<br />

fruits: Green, 1/2 to 3/4 of an inch long, oval. The seeds are<br />

brown.<br />

Range. The lower Rio Grande Valley from near Roma, Texas,<br />

to near Brownsville, and into adjacent Mexico.<br />

Remarks. This is probably the most untidy and ill-organized<br />

of all cacti with normal growth. It sometimes takes on the appearance<br />

of a cristate or monstrous growth. Assuming that it<br />

starts from a single stem, branching must occur very quickly,<br />

since I have never seen a plant so small that it was not already<br />

a cluster of tiny heads. These heads branch so fast and so irregularly<br />

that it is typical for the larger plants to present a completely<br />

disorganized mass of small tubercles at all angles. The<br />

whole thing is very low to the ground, not over 2 or 3 inches<br />

high, but may be quite broad. Mr. Runyon, its discoverer, said<br />

that a large plant, with its fleshy root, might weigh 50 pounds.<br />

One should hardly look for such huge specimens now, however,<br />

as all of these old clumps seem to have been uprooted long ago,<br />

and one should be thrilled to find any specimen over 6 inches<br />

in diameter today.<br />

M. runyonii grows on gravelly hillsides overlooking the Rio<br />

Grande, and to a few miles north of the river. It may have<br />

grown in the lower flats of the valley also, but very little of the<br />

valley proper is still uncultivated, so its locations are now few<br />

and far between. It always prefers partial shade, and will not<br />

do well in full sun.<br />

The plant was first described by Britton and Rose after their<br />

break-up of the genus Mammillaria, so it has been almost universally<br />

known as Coryphantha runyonii.<br />

In 1934 Werdermann described a Coryphantha pirtlei found<br />

in Starr County, Texas. The description of that plant does not<br />

present any character which lies outside the descriptions already<br />

given for M. runyonii or which would enable one to distinguish<br />

it from that species, except that his plant was said to occur


122 cacti of the southwest<br />

sometimes as a single stem. Starr County is the area where M.<br />

runyonii is most common, and I have not been able to find there<br />

any specimens which I could separate from the rest of the population<br />

and designate as a separate form. At the same time I<br />

must admit that I have not found there any individual with a<br />

single stem. Since M. runyonii is rather variable, I am inclined<br />

to consider this name as a synonym, or at best a possible variety<br />

of M. runyonii, but there can be no certainty about the plant<br />

until it is re-collected.<br />

Mammillaria similis Eng.<br />

[Neobesseya similis (Eng.) B. & R.]<br />

“Nipple Cactus”<br />

Description Plates 30, 31<br />

roots: Fibrous.<br />

stems: Occasionally growing individually, but usually clustering,<br />

often to form large, irregular masses up to a foot or<br />

more across when old. Each stem is spherical, usually wider<br />

than it is tall, growing up to 4 inches in diameter, but the<br />

average being 2 or 3 inches. The stem is covered by tubercles<br />

up to 7/8 of an inch long on large plants. These are cylindrical<br />

on the upper part of the plant, but the older ones on the<br />

sides of the stem become somewhat flattened and overlap up-<br />

ward a little.<br />

areoles: On immature plants round or oval. On mature<br />

growth each tubercle is deeply grooved its whole length on<br />

the upper side with a groove which is an extension of the<br />

areole into or nearly to the axil. The whole areole is at first<br />

filled with white wool, but it later becomes bare except for a<br />

tuft which persists at the base of the groove in the axil, where<br />

the vegetative meristem is.<br />

spines: There are 10 to 17 slender radial spines. They vary in<br />

length with the size and age of the plant, from 1/8 to 1/2 inch<br />

long. They are fairly equal on any one areole, except that the<br />

1 or 2 uppermost ones are usually much more slender than<br />

the others. There is often no central spine, but about as often<br />

there is one central which is very slightly longer and heavier<br />

than the radials. It either stands straight out, perpendicular<br />

to the plant surface, or else is turned upward sharply in front<br />

of the upper radials. All of the spines are gray to white, sometimes<br />

with brown tips, and when young are covered with tiny<br />

white hairs or scales only visible under a magnifying glass,<br />

but these hairs are soon worn off and the spines then often<br />

appear smooth, yellowish, and hornlike.<br />

flowers: 1 to 2 inches tall and the same in diameter, with<br />

very narrow, linear, sharply pointed petals. The color of these<br />

flowers varies from plant to plant from greenish, bronze,<br />

gold, gold streaked with pink, or pale, almost clear yellow.<br />

Most commonly all of these shades are mixed in the same<br />

flower. There are 15 to 20 outer petals which are greenish<br />

with fringed, yellowish or whitish edges. There are 20 to 25<br />

inner petals which are even narrower than the outer, about<br />

1/8 of an inch wide near their bases and tapering very gradually<br />

to a sharp point. These are clear yellow, gold, chartreuse,<br />

or bronze, often with pinkish midlines, and with unfringed,<br />

smooth edges. The filaments may be white, light green, or<br />

yellow. The anthers may be yellow or pale orange. The style<br />

is very much longer than the stamens and green or yellowish<br />

in color. The stigma has 4 to 6 green or yellowish lobes, which<br />

are 1/8 to 1/4 of an inch long and which in some plants are<br />

slender and smooth, while in others they are thicker and<br />

rough.<br />

fruits: Spherical to oval in shape, 3/8 to 3/4 of an inch long.<br />

They remain green and below the spines of the plant between<br />

the tubercles for nearly a year. The second spring they turn<br />

bright scarlet. They then remain where they are and shrivel<br />

and harden unless plucked out from their protected places by<br />

birds or animals. The seeds are approximately 1/16 of an inch<br />

(11/2 to over 2 millimeters) across, are almost spherical in<br />

shape, and black in color, with pitted surfaces.<br />

Range. Generally, Kansas and eastern Colorado south through<br />

Oklahoma and north Texas to near San Antonio. More specifically,<br />

I have found records of this form’s occurrence in a long<br />

but narrow belt from south-central Kansas through central<br />

Oklahoma into Texas, including the Dallas and Fort Worth area<br />

and on down to near San Antonio and Columbus. I find no<br />

record of it west of a line from San Antonio to approximately<br />

Oklahoma City, and must regard the Colorado specimens as<br />

either an isolated population or else an extension of the Mon-<br />

tana and Idaho population, which is a form separate from ours.<br />

Remarks. As most people have met with this cactus, it is a small,<br />

irregular clump of spherical stems producing flowers with narrow,<br />

sharp-pointed petals of a curious greenish-yellow color<br />

often striped with browns and pinks. It has appealed mostly as<br />

a curiosity, growing in areas where few if any other cacti are<br />

found. Only a few people have seen it in its glory, an old clump<br />

a foot or more across, made up of dozens of little heads, and<br />

covered with 20 or 30 flowers in its blooming period. When<br />

seen this way it is recognized as a beautiful cactus to be cherished.<br />

The cactus grows equally well in loamy places on the central<br />

plains or on calcareous hilltops. It does not like the full sun in<br />

the summer, shrinking up and even dying when totally unshaded.<br />

In its native habitat it is shaded just enough by the prairie<br />

grass growing over it on the plains and by shrubs and brush on<br />

the hills. It is much more tolerant of moisture than most cacti,<br />

but will still rot quickly if placed in poorly drained, heavy soil.<br />

It is rather common in the hills on all sides of Austin, Texas,<br />

and is met with occasionally in a band of territory about 100<br />

miles or so wide which includes Waco, Fort Worth, and Dallas,<br />

Texas. It may be found, but is not at all common, north of this<br />

in a band of about the same width through central Oklahoma,


genus Mammillaria 123<br />

until it becomes rather common again between Ponca City and<br />

Tulsa, Oklahoma. From there north rare populations have been<br />

found in Kansas, mostly near the Arkansas River and below<br />

Hutchinson, Kansas. I find no record of it north of this.<br />

There has been much confusion about Mammillaria similis<br />

from the beginning, and it is not all cleared up even today. The<br />

first cactus of this sort described was a plant from the upper<br />

Missouri River area. It was described very briefly by Nuttall in<br />

1818, called Cactus mammillaris. In 1827 Sweet renamed this<br />

plant Mammillaria missouriensis.<br />

In 1849 Engelmann secured this plant from Fort Pierre on<br />

the upper Missouri, which is near Pierre, South Dakota. He also<br />

felt it necessary to rename the cactus, and called it Mammillaria<br />

nuttallii, after its discoverer. But Engelmann already had a very<br />

similar cactus discovered in central Texas. In 1845 he had named<br />

this Texas cactus Mammillaria similis. So we have had from this<br />

early time two forms, one commonly known as M. missouriensis<br />

(Engelmann’s M. nuttallii), discovered in the far north, and the<br />

other, M. similis, found deep in central Texas. No doubt every<br />

serious cactus student has struggled with these two, with the<br />

questions of whether they are different or not, how they are<br />

related, and what to call them.<br />

We must credit Engelmann with wrestling with these problems<br />

himself. By 1856 he had studied many more specimens<br />

from wide areas, and by then he must have concluded that these<br />

two forms were very close to each other, for he came up with a<br />

new naming system for them. The northern M. missouriensis he<br />

now called M. nuttallii var. borealis, and said that it grew from<br />

Montana and South Dakota to Nebraska, western Kansas, and<br />

eastern Colorado. The southern M. similis he now called M. nuttallii<br />

var. caespitosa, ranging, he said, from the Kansas River to<br />

New Braunfels, Texas. The point is that he made them varieties<br />

of the same species.<br />

From this time on some chose to call them two separate species<br />

and some to call them varieties of the same species, while at<br />

the same time placing them in almost every conceivable genus.<br />

The result was that Poselger in 1853 used for our Texas form<br />

the name Echinocactus similis; Small in 1903 used Cactus similis;<br />

Britton and Rose in 1913 and again in 1921, Schulz in 1930,<br />

and Backeberg in 1960 used Neobesseya similis; while Watson<br />

in 1878 used Mammillaria missouriensis var. caespitosa; Coulter<br />

in 1896 used Cactus missouriensis var. similis; and Schumann in<br />

1898 used Mammillaria missouriensis var. similis. Whether it is<br />

correct to place this form as a separate species or as a variety of<br />

the other form seems impossible to state categorically even today.<br />

I list it here as a separate species, mostly because the recent<br />

studies have been rather consistent in doing this, but I summarize<br />

the actual known differences between the two so that the<br />

reader may come to his own decision.<br />

There is no actual difference of stems or tubercles between the<br />

two except that the southern M. similis usually grows to a larger<br />

maximum size, but this may well be due to the better growing<br />

seasons in the south. The spines are essentially the same in num-<br />

ber and character in both, except that, once again, the maximum<br />

length is slightly greater in the southern form. The flower diameter<br />

and length of the northern M. missouriensis is about 1 inch,<br />

while M. similis flowers vary from 1 inch across and long in its<br />

extreme upper range in Oklahoma and Kansas to 2 inches in<br />

central Texas, with climate seeming definitely to make the difference,<br />

since plants from Kansas brought to San Antonio and<br />

grown in my collection doubled the size of their flowers the<br />

second year.<br />

One flower character is given as definitely different in the<br />

two forms. M. missouriensis is stated to have 2 to 5 stigma lobes<br />

which are very short, about 1/12 to 1/20 of an inch long, while<br />

M. similis has 4 to 6 lobes 1/8 to 1/4 of an inch long. This does<br />

not seem to be influenced by environment. The size of seeds,<br />

which are only about one-half as big in the former as the latter,<br />

also seems significant. But these things are about all that can be<br />

definitely given to divide them—a fact which leaves them so<br />

close that a good case can be made for calling them mere vari-<br />

eties of the same species.<br />

Anyone collecting in the Southwest need not worry about<br />

distinguishing between the two, however, as it seems certain<br />

that the northern form does not come into Oklahoma or Texas.<br />

From southern Kansas down all of them appear to be the same<br />

southern M. similis.<br />

Most books repeat the listing, as either a variety of this species<br />

or as a closely related separate species, of a cactus which I<br />

do not believe exists as a separate form at all. Engelmann first<br />

described this most doubtful form as a variety, Mammillaria<br />

similis var. robustior, and then later as M. nuttallii var. robustior.<br />

Others, such as Watson and Coulter, followed him in this,<br />

calling it M. missouriensis var. robustior, but did it on the<br />

strength of nothing more than Engelmann’s preserved specimens,<br />

which were collected in 1845 and 1847 by Lindheimer and by<br />

Bigelow. In 1898 Hildmann elevated it to a species for the first<br />

time, calling it Mammillaria wissmannü. Everyone has followed<br />

him since then, and the species is listed in all the books on the<br />

Texas area, although most of the authors have failed to collect the<br />

plant and have merely followed Britton and Rose, who copied<br />

the earlier description by Engelmann, reproduced a drawing<br />

said to be of the form from Blühende Kakteen, and called the<br />

cactus a Neobesseya. The only new material on this cactus since<br />

Engelmann’s time is a photograph in Backeberg’s 1960 work<br />

purported to be of the plant and credited to Thiele.<br />

Over a period of years I have searched for this cactus throughout<br />

central Texas, where it is supposed to be found. I have not<br />

been alone, as a perennial project of most Texas cactophiles is<br />

the unending search for the “yellow-flowered Neobesseya” having<br />

the large flowers with the unfringed outer petals. I have<br />

followed many leads, only to find everything M. similis with<br />

the smaller flower having the fringed outer petals and fewer<br />

stigma lobes than M. wissmannii is supposed to have. I have<br />

never found, nor do I know of any collector who has found the<br />

Neobesseya with the gold-yellow flowers having 7 or 8 stigma


124 cacti of the southwest<br />

lobes and unfringed outer perianth segments, although I have<br />

seen many specimens of typical M. similis called this.<br />

I had puzzled over this situation for a long time, when I was<br />

shown some plants of M. sulcata from near Kerrville, Texas,<br />

which were not quite typical. The flowers on these specimens<br />

were more greenish-yellow than the pinard-yellow I had seen<br />

before in that species, and the red of the center was almost entirely<br />

lacking, only represented by a faint brownish stain in the<br />

very center of the flower. Their collector gave me an idea when<br />

he speculated that he had in them a cross between M. sulcata<br />

and M. similis. This I doubt. They were quite clearly M. sulcata,<br />

but his speculation presented a new possibility, which I entertain<br />

reluctantly, but which I cannot ignore. It seems highly<br />

likely that all of the M. similis var. robustior or M. wissmannii<br />

found by Lindheimer and Bigelow in this same area and described<br />

by Engelmann were really only these somewhat atypical<br />

M. sulcata, lacking or nearly lacking the red centers in the<br />

flowers.<br />

In support of this theory it should be noted that the size and<br />

shape of the two are the same, the number and size of the radial<br />

spines fall within the same range, and the 0 to 3 centrals are the<br />

same in all respects in both. Add to this the gold-yellow color<br />

of the flower which is stressed for both, the unfringed petals<br />

and 7 or 8 stigmas described for both, and the case could easily<br />

be considered closed. Another remarkable coincidence is that<br />

M. wissmannii is expressly stated to sprout new stems from the<br />

grooves of old tubercles, a character which is very striking in<br />

M. sulcata but not noticeable in M. similis.<br />

I can cite no clear-cut differences noted so far which could<br />

separate M. wissmannii from M. sulcata. The fruit character<br />

should do this, since, if M. wissmannii is so closely related to<br />

M. similis, its fruit would be red, while the other’s fruit stays<br />

green. But there has never been a description of the color of the<br />

fruit on M. wissmannii, so we can settle nothing there. The seeds<br />

of the two are both described, however, but are practically the<br />

same size. Since those of M. wissmannii are merely described as<br />

“dark,” we cannot know whether they were actually black like<br />

those of M. similis or dark brown like those of M. sulcata.<br />

In short, I can find no proof that anyone has ever seen anything<br />

other than some M. sulcata in which the red was lacking<br />

in the flower, and which were, therefore, thought to be another<br />

cactus. When I put the specimens preserved in the United States<br />

National Museum under the name Neobesseya wissmannii side<br />

by side with specimens of M. sulcata I could not separate them<br />

by any character showing in the preserved state. Since the range<br />

in which the two plants are supposed to grow is practically the<br />

same, the evidence seems to point to the assumption that they<br />

are one and the same. This would make Mammillaria (Neolloydia)<br />

wissmannii merely a synonym of M. sulcata. It should at<br />

least clear up the situation for collectors, in whose gardens I<br />

have so many times seen the more brightly yellow-flowered<br />

specimens of M. similis arbitrarily labeled M. wissmannii merely<br />

because no one can find anything else to bear this label.<br />

M. similis is an eastern cactus, not strictly a desert habitant.<br />

It is, therefore, much better suited to the ordinary cactus collection<br />

or window garden than most. It also has the fine feature<br />

of being able to take most ordinary winter temperatures over a<br />

wide area of the U. S. without protection, so long as it is growing<br />

in a well-drained situation. It is probably the best cactus to<br />

be grown by the amateur who does not want to bother with the<br />

intricacies of cactus culture.<br />

Mammillaria rosiflora (Lahman)<br />

[Neobesseya rosiflora Lahman]<br />

Description<br />

stems: Identical to those of M. similis except reaching a maxi-<br />

mum of only about 23/4 inches across and clustering more<br />

sparingly to form only small clumps.<br />

spines: There are 13 to 16 radial spines to 5/16 of an inch long,<br />

equal around the areole except for the uppermost 2 of them,<br />

which are very short and so slender as to be almost bristlelike.<br />

They are gray with brown tips, and have minute white<br />

hairs on the spines when they are young. There is apparently<br />

always one central spine which is about 1/4 of an inch long<br />

and turned upward.<br />

flowers: About 11/2 inches across and tall. They are a pure<br />

pink, without the stripes and zones of various colors found in<br />

the flowers of the plant’s relatives. The outer petals are narrow,<br />

but thick and fleshy, and they are not fringed. The inner<br />

petals are so narrow as to be almost threadlike. They are very<br />

weak and limp, spreading and bending in all directions. The<br />

filaments are deep pink, the anthers yellow. The stigma has<br />

4 to 6 white lobes.<br />

fruits: Round to egg-shaped, crimson, 3/8 of an inch long.<br />

The seeds are almost circular, black, pitted on the surface,<br />

and less than 1/16 of an inch (11/2 millimeters) long.<br />

Range. Known only from an area about 80 miles long from<br />

near Tulsa to near Ponca City, Oklahoma.<br />

Remarks. This form was discovered by Mrs. Marion Sherwood<br />

Lahman, a great student of Oklahoma cacti, and described by<br />

her in 1939. She found it in only one location, a prairie just<br />

west of Tulsa, Oklahoma. Professors Blauch and Gatewood of<br />

Oklahoma State University collected it again near Ponca City,<br />

Oklahoma, in 1962, and graciously supplied me with information<br />

about it, as well as with a cutting of the plant. I am very<br />

sorry to admit that what ability I may possess at cacti culture<br />

failed me in this case. I was unable to grow my cutting to blooming<br />

stage, or even long enough to secure an adequate photograph<br />

of it. I find no recorded collection of it between Tulsa and Ponca<br />

City, but it must be assumed that it grows as a rare form in the<br />

area.<br />

There is hardly enough information concerning this cactus to


genus Mammillaria 125<br />

decide definitely its relationships to other species. In both instances<br />

mentioned above there were found numerous plants of<br />

M. similis growing within a matter of feet of the M. rosiflora<br />

specimens. No character of plant body or growth has been<br />

found to distinguish the two. Without the flowers one cannot<br />

segregate them at all, but M. rosiflora has flowers of clear pink,<br />

unadulterated by any other colors, with extremely narrow,<br />

weak, entirely unfringed petals, which set it apart when in<br />

bloom from even the pinkish-flowered M. similis, with its petals<br />

striped and streaked with browns, grays, greens, or yellows, and<br />

its outer segments fringed. The flowers of M. rosiflora are strikingly<br />

beautiful, about the same shade of pink as a typical flower<br />

on Echinocactus texensis. They remind one somewhat of those<br />

of Mammillaria vivipara, but there seems no close relationship<br />

here. The same peculiarities of flower which set this form apart<br />

from M. similis also set it apart very clearly from the pinkish-<br />

flowered populations of M. missouriensis found in Montana and<br />

called by Britton and Rose Neobesseya notesteinii.<br />

It will be a worth-while achievement when someone can again<br />

locate and collect this very rare form so that more may be learned<br />

about it. Unfortunately a very large proportion of the flat<br />

prairies of Oklahoma where it is found have been turned to<br />

cultivation, reducing greatly the chances that this cactus will get<br />

adequate study.<br />

Mammillaria vivipara (Nutt.) Haw.<br />

“Spiny Star,” “Ball Cactus,” “Pincushion”<br />

Description Plates 31, 32, 33<br />

stems: Spherical to columnar and single to greatly clustering<br />

by offset heads from around the base. Individual stems measure<br />

from about 2 to at least 5 inches tall and wide. The surface<br />

is deep green in color and is divided into many cylindrical<br />

or tapering tubercles which are regularly arranged and<br />

spreading or slightly overlapping upward.<br />

areoles: Round or nearly so on immature plants, but the<br />

upper part of the areole elongated on mature plants into a<br />

narrow groove running from the spinous portion of the areole<br />

at the tip of the tubercle to the reproductive part in the<br />

axil of the tubercle. When young these areoles are filled with<br />

much white or yellowish wool, but when old, on the sides of<br />

the stems, the wool is usually worn off and they are bare.<br />

spines: Various forms have from 12 to at least 60 radial<br />

spines. These are slender, straight, radiating rather evenly<br />

around the areole. They are firm, 3/8 to about 1 inch long,<br />

white, white-tipped dark, or grayish with brown tips. There<br />

are also 3 to 12 central spines which are straight, spreading,<br />

rather slender to medium in thickness and 1/4 to 11/4 inches<br />

long. These centrals are from whitish or yellowish with dark<br />

tips through mottled brown to partly or completely dark<br />

brown or purplish-black.<br />

flowers: Purple, violet, or rose, usually very bright in color,<br />

but one form has pale purplish coloring. From about I to 21/4<br />

inches in length and diameter. The perianth segments are<br />

lance-shaped to linear, the outer ones greenish and fringed<br />

and the inner ones purple and entire. The stigma has 5 to 10<br />

white to rose-purple lobes which are obtuse in some forms<br />

and pointed or even mucronate in others.<br />

fruits: Oval in shape, 1/2 to 1 inch long, not including the<br />

dried perianth, which persists, remaining green or in some<br />

cases becoming brownish when very ripe. The seeds are<br />

approximately 1/16 of an inch (1 to 21/2 millimeters) long, almost<br />

spherical to oval or curved ovate. They are light or dark<br />

brown and with the hila ventral or subbasal, concave or con-<br />

vex, and oval to linear in the different forms.<br />

Range. Including for the species as a whole a huge area of the<br />

North American continent from the Dakotas, Nebraska, Kansas,<br />

Oklahoma, and Texas on the east through all the states<br />

southwest to California and deep into Mexico.<br />

Remarks. It seems almost impossible for any but taxonomists<br />

to think of this species as a single unity. Collectors of cacti rarely<br />

get to collect in more than a fraction of its vast range and<br />

rarely see all of the forms together at one time, and so they<br />

usually pick up one or two of this species’ varieties and then go<br />

to the verge of distraction trying to determine which forms they<br />

have. This is often hard to determine even with all of the stages<br />

of all of the varieties before one, and is almost impossible when<br />

one has only the vegetative stages of one or two varieties to<br />

study.<br />

Perhaps because of this difficulty of getting an overview of<br />

the whole wide-ranging species, its various forms have more<br />

often than not been thought of as separate species themselves,<br />

even though Engelmann, the original describer of most of them,<br />

recognized that they were merely varieties of one large unity<br />

and his first descriptions list them that way. Subsequent writers<br />

could not conceive of them that way, and soon they were all<br />

listed as separate species. The listing of these supposed species<br />

by most authors over a long period as though they were completely<br />

and obviously distinct has been a great disservice to<br />

cactophiles, many of whom have been turned into feuding rivals<br />

over these forms, thinking of them as separate species. It was<br />

clearly of prime importance to be able to identify them correctly,<br />

and yet they are so closely related that with only limited<br />

material to study, mistakes were bound to be made. I have even<br />

heard of cactus clubs being split over the interpretation of some<br />

of these forms. If only the fact could be remembered that this<br />

is one very large and variable species and that the different<br />

forms, which we can all notice but which are so close that we<br />

can only understand them after a great deal of study, are no<br />

more than very close and perhaps to some degree even intergrading<br />

varieties, then perhaps cactophiles would be less con-<br />

cerned about placing a given specimen exactly.<br />

It would no doubt be better if those who can collect only a


126 cacti of the southwest<br />

form or two of this species would content themselves with<br />

knowing that they have Mammillaria vivipara. However, knowing<br />

the marvelous inability of the cactophile to be satisfied with<br />

this sort of halfway identification, I am following this species<br />

description with an account, as clear as I can make it, of each<br />

of the varieties of this species occurring in our area. But I must<br />

warn those who would follow to the varietal level that here<br />

they are embarking upon some of the most stormy seas of controversy<br />

in all of cactus study, that all of the winds of argument<br />

about these forms are still not spent, and that there are those<br />

who will maintain that even in these varieties which seem the<br />

most certain we have no secure harbors. But no doubt this, which<br />

is one of the most common little cacti over a huge part of this<br />

continent, deserves the effort it takes to understand its confusing<br />

varieties.<br />

It is important before going any farther to understand that<br />

Mammillaria vivipara (Nutt.) Haw. is not to be equated with<br />

Cactus viviparus Nutt., which became the Coryphantha vivipara<br />

(Nutt.) B. & R. The former is from the beginning an inclusive<br />

taxon, taking in, as described by Haworth, all of the then-<br />

known range of spine numbers, plant sizes, amount of clustering,<br />

and geographical area of the plant. On the other hand the<br />

latter, in the sense used by Nuttall and Britton and Rose, is only<br />

a very strictly delimited northern form of the species, one segment<br />

of the total species, and in modern taxonomic usage<br />

should be called Coryphantha vivipara var. vivipara (Nutt.)<br />

B. & R., if one follows their generic scheme, or Mammillaria<br />

vivipara var. vivipara (Nutt.) if the species is returned to this<br />

genus.<br />

Mammillaria vivipara var. vivipara (Nutt.)<br />

[Coryphantha vivipara (Nutt.) B. & R.]<br />

Description Plate 31<br />

stems: As the species, except that the stems are always practically<br />

globular or even depressed-globular, never columnar,<br />

and they grow to only about 21/2 inches in diameter.<br />

areoles: As the species.<br />

spines: There are 12 to 21 radial spines on each areole. These<br />

are slender, 3/8 to 5/8 of an inch long, and very white, sometimes<br />

with brown tips. There are 1 to 8 central spines which<br />

are usually 1/2 to 3/4 of an inch long, and only slightly heavier<br />

than the radials. There will always be 1 to 3 centrals directed<br />

outward and downward. These are the heaviest spines. The<br />

rest of the centrals will be standing upright at the top of the<br />

areole, spreading fanwise in front of the upper radials, and<br />

varying in thickness, some of them being almost as fine as the<br />

radials themselves. There is variation in the color of these<br />

centrals from plant to plant. Most commonly they are light<br />

brown or honey-colored, often mottled with whitish, and<br />

often with darker tips or other darker zones, but never with<br />

blackish coloring.<br />

flowers: Deep magenta or purple in color, not opening widely,<br />

11/8 to 13/4 inches tail and 11/2 to 2 inches across, having<br />

very narrow petals. The outer petals are greenish or brownish<br />

with pink edges fringed part or all the way to the tips by<br />

long, pink cilia. The petals are about 1/8 of an inch wide near<br />

the base, tapering gradually to a sharp point. The inner petals<br />

are 3/4 of an inch long and 1/16 to 1/8 of an inch wide, with<br />

edges smooth or sometimes very finely toothed, tapering over<br />

their length to a sharp point. There are at least 30 outer petals,<br />

about 17 to 20 of them elongated and the rest so short as<br />

to be almost like fringed scales. There are about 30 to as many<br />

as 47 inner petals. The filaments are green at their bases, the<br />

upper parts becoming pink. Anthers are bright orange. The<br />

style is the same length as the stamens, or sometimes slightly<br />

longer. The Stigma has 7 to 10 dark rose-purple lobes, which<br />

are slender and about 3/16 of an inch long. They are always<br />

somewhat pointed, and may have spinelike mucrones on the<br />

ends, but do not always have these.<br />

fruits: Oval in shape, 1/2 to 3/4 of an inch long, light green<br />

in color, becoming brownish when old. The seeds are light<br />

brown in color, their surfaces finely pitted. They are approximately<br />

1/16 of an inch (1 to 11/2 millimeters) in length and<br />

very thick so as to be almost round, except for the concave<br />

hila.<br />

Range. From Canada south through Montana, the Dakotas,<br />

Nebraska, and Kansas across the northwestern corner of Oklahoma<br />

into extreme northwestern Texas, and across eastern Colorado<br />

into northeastern New Mexico.<br />

Remarks. M. vivipara var. vivipara was discovered very early,<br />

its first description having been made by Nuttall in 1813 under<br />

the name Cactus viviparus. It is the small, northern cactus found<br />

in parts of Canada and the northern plains where few other<br />

cacti grow. Apparently it is immune to freeze damage, so it<br />

grows anywhere in the north where there is not too much moisture<br />

and where the soil is right for it. It is quite easily rotted by<br />

continued dampness in the wrong soil, its requirement of alkaline<br />

soil no doubt keeping it from growing any farther east than<br />

the western Great Plains.<br />

Within our area of study it is limited to the northwest, as<br />

though it cannot survive the severe heat in which its relatives<br />

revel. It is found in scattered locations over all of Oklahoma<br />

northwest of a diagonal line running from near Enid to the<br />

southwest corner of the state. It is the form of the species found<br />

in the upper Texas Panhandle, but the Red River and its upper<br />

tributaries seem to mark the southern edge of its penetration in<br />

Texas. It grows in the northeastern corner of New Mexico, particularly<br />

around Raton, and down nearly as far southwest as<br />

Santa Fe. This large area of Oklahoma, Texas, and New Mexico<br />

is its home, where it grows on sandy hills and plains; it is not a


genus Mammillaria 127<br />

mountain species, and its range stops short of the high moun-<br />

tains in both Colorado and New Mexico.<br />

The close relatives of M. vivipara var. vivipara with which it<br />

is often confused will be described in detail following this discussion.<br />

It may not be distinguished from them by any one characteristic,<br />

and from some of them it can hardly be told at all<br />

when it is not in flower or fruit. It can be said that it has fewer<br />

radial spines than most other forms of the species, but this fact<br />

alone will not always separate it from all of them. Part of the<br />

confusion over these varieties has arisen from attempts to distinguish<br />

them on radial number alone.<br />

When it blooms, variety vivipara can be recognized by the 7<br />

to 10 stigma lobes which are long, slender, and pointed, sometimes<br />

to the extent of having spinelike tips, as well as being very<br />

beautiful dark carmine to pink in color, but never white.<br />

For still more certainty about the identification of his plant,<br />

the collector must nurture it further until it forms its fruits. He<br />

can then check the seeds it produces, and here find what appears<br />

to be the most certain character to distinguish it. Variety<br />

vivipara has medium brown seeds which are egg-shaped to<br />

almost round except for concave, oval hila, and which have<br />

roughly pitted surfaces. They are approximately 1/16 of an inch<br />

(1 to 11/2 millimeters) long. This distinguishes the variety from<br />

its close relatives, all of whose seeds are different in size and<br />

detail.<br />

Within itself the variety varies. The description mentions that<br />

the color of the centrals varies from plant to plant. However,<br />

they never have blackish zones, and they are always some shade<br />

of brown, usually predominantly honey-colored. Neither does the<br />

amount of clustering the plants do have any significance, as this<br />

varies greatly. The first descriptions cautioned us that it grows<br />

either singly or profusely clustering; nevertheless, the two types<br />

of growth appear so different that all through the years there<br />

have been attempts to separate them as two forms, the simple<br />

and the clustering. Yet I have plants taken from only a few feet<br />

of each other, identical in every respect except that one specimen<br />

has grown a single stem while its neighbor has grown as<br />

many as 7 heads.<br />

Perhaps the most ambitious attempt to divide up the species<br />

on this sort of character was made about 25 years ago by Mrs.<br />

Marion Sherwood Lahman, one of the most ardent collectors<br />

and students of Oklahoma cacti. She described a total of 6 species<br />

of this group in Oklahoma, including 2 new ones, but all of<br />

them fall within the limits of the species M. vivipara in the sense<br />

used by Haworth, and as she separated them on only such general<br />

and variable characters as amount of clustering, it appears<br />

that her taxa have only doubtful significance.<br />

When dealing with this cactus it must always be remembered<br />

that the name, Mammillaria (Coryphantha) vivipara has been<br />

and still is used with two very different meanings. Some use this<br />

name to refer specifically to the cactus which grows in Nuttall’s<br />

type locality on the upper Missouri River. This is the exact form<br />

which grows, as Engelmann well stated in his earliest writings,<br />

from the Yellowstone and Missouri rivers down to Santa Fe and<br />

no farther southwest, and which has only 12 to about 20 radial<br />

spines. But as new, very similar forms were discovered, they<br />

were all designated by Engelmann as merely varieties of one big<br />

species, so they were listed as M. vivipara variety or subspecies<br />

borealis, neo-mexicana, radiosa, and so on. So as the name M.<br />

vivipara came to mean the species encompassing all of these<br />

varieties, writers—Haworth started it, followed by Engelmann<br />

himself in his later writings—began to give a description under<br />

this name so broad as to take in everything (as for instance saying<br />

M. vivipara had 12 to 36 or more radials). It is obvious that<br />

here the name is being used to refer to a different entity than<br />

the limited northern form. The practice of keeping the original,<br />

typical form separate by calling it M. vivipara var. vivipara<br />

would have minimized the confusion which resulted, but this<br />

practice was not followed at that time, so the form we are discussing<br />

here got lost among its own relatives as its name was<br />

applied to them all.<br />

When understood in this sense, the variety vivipara is a small,<br />

retiring cactus which is content to huddle unseen in the prairie<br />

grass of the northern plains, but which is amazingly hardy and<br />

which produces very beautiful flowers in their season. It is seldom<br />

common, and as the grass of northwestern Texas and<br />

western Oklahoma has been trimmed ever shorter by overgrazing,<br />

these little plants more and more lose their protection and<br />

are carried home as curiosities, thus becoming more and more<br />

scarce. I observed this order of events myself on the outskirts of<br />

Amarillo, Texas. Some years ago I searched for the plant in a<br />

particular area there without success. Then, a few years ago, as<br />

that city was increasing in size, homes were built in the area<br />

where I had unsuccessfully searched, and the ranchland became<br />

small lots for ponies. After their intensive grazing, when I visited<br />

in one of the new homes, I saw from the window several specimens<br />

of the cactus in the almost table-bare lot. Even more recently<br />

I noticed that all of these specimens had been taken. With<br />

this process and with the cultivation of so much prairie, one has<br />

now to look for this cactus in rough places.<br />

Mammillaria vivipara var. radiosa Eng.<br />

[Coryphantha radiosa Rydberg]<br />

Description Plate 32<br />

stems: As the species, except always ovate to columnar when<br />

mature and very rarely and sparingly clustering. These stems<br />

grow to 5 inches tall and about 3 inches in diameter. The<br />

tubercles are cylindrical and to about 3/4 of an inch long.<br />

areoles: As the species.<br />

spines: As the species, except that the radial spines are always<br />

white, sometimes tipped with brown, very slender, to 1/2 inch<br />

long, and 17 to 30 in number. The central spines are 4 to 7 in


128 cacti of the southwest<br />

number, spreading from the center of the areole in no set pattern.<br />

Most commonly there will be 1 or 2 rather heavy downturning<br />

centrals and the rest will stand erect almost against<br />

the upper radials, but I have seen some specimens with only<br />

4 centrals and all of them erect—the lower ones lacking entirely.<br />

These centrals are light in color, honey- or straw-col-<br />

ored, often with large zones of white.<br />

flowers: About 2 inches across and 21/4 inches tall, very deep<br />

violet in color. The petals are 1 to 11/4 inches long and very<br />

narrow, being about 1/8 of an inch across. The outer ones are<br />

40 to 50 in number, with about 17 of these elongated, and are<br />

greenish in color and fringed; while the inner ones are 30 to<br />

40 in number and violet with entire edges. The filaments are<br />

deep violet, while the anthers are deep orange. The style is at<br />

least 3/16 of an inch longer than the stamens, and pink. The<br />

stigma has 7 to 9 rose-colored lobes which are 3/16 to 1/4 of an<br />

inch long and fairly slender, but not pointed.<br />

fruits: About 3/4 of an inch long, oval, and green in color.<br />

The seeds are rather dark brown with a finely striated surface,<br />

more than 1/16 of an inch (2 to 21/2 millimeters) long, and<br />

oval, with the ventral surfaces convex and the hila linear in<br />

shape.<br />

Range. From the southern slopes of the Arbuckle Mountains in<br />

Oklahoma south to a line from near Austin, Texas, west past<br />

Fredericksburg to Sonora, Texas. The area comprises a triangle<br />

with its three points near Springer, Oklahoma, and Austin and<br />

Sonora, Texas.<br />

Remarks. This is a form which usually passes for Mammillaria<br />

vivipara var. vivipara. Admittedly it differs from that variety<br />

only in minor characters of the plant body, and very careful<br />

observation alone will separate them. Variety vivipara always<br />

has globose stems and 12 to 21 radials, while the stems of variety<br />

radiosa become elongated and columnar and it has 17 to 30<br />

radials. The difference of flowers is also minor, the flowers of<br />

variety radiosa being larger—in fact definitely the largest and<br />

most deeply colored of the various forms in this species found<br />

in our area—but beyond that about all that can be said is that<br />

the stigma lobes of variety radiosa are never pointed as they are<br />

in variety vivipara.<br />

I would surely not blame anyone for thinking we were splitting<br />

spines here if we listed these as two separate species: they<br />

are difficult even as varieties. It would be much easier to consider<br />

these two cacti the same and be done with it, but a hundred<br />

years ago Engelmann saw a consistent difference in the<br />

seeds of the two cacti, and we must recognize it. The seeds of<br />

variety radiosa are unmistakably different, almost twice as large<br />

in each direction as those of variety vivipara; they are full and<br />

convex on all surfaces, as well as darker brown, and their surfaces<br />

are very finely pitted, where the seeds of variety vivipara<br />

are small, flattened, and concave, and lighter brown with more<br />

coarse pitting. There is more difference between these two than<br />

among the seeds of all the various and otherwise vastly differ-<br />

ent Echinocerei, and this species provides the only instance I<br />

know where cacti almost identical in other respects have such<br />

basically different seeds.<br />

Amazing confusion has arisen out of Engelmann’s changing<br />

schemes to relate all the forms in this species. In his Plantae<br />

Lindheimerianae in 1850, before he was combining, he had M.<br />

vivipara described as it is still found in Nuttall’s type locality,<br />

and at the same time he described as a separate species a Mammillaria<br />

radiosa with the type locality, “Sterile, sandy soil on<br />

the Pierdenales” (the Pedernales River in central Texas). This<br />

plant he described as having 20 to 30 radials, being columnar<br />

and having different seeds from M. vivipara. It is a very distinct<br />

form, treated very fully later by Coulter, and is exactly<br />

the form we are discussing here.<br />

But in trying to combine forms, Engelmann later confused an<br />

originally clear picture. First, in the Synopsis of the Cactaceae,<br />

he reduced M. radiosa to a variety of M. vivipara and put some<br />

newly noticed forms, borealis, neo-mexicana, and texana as<br />

subvarieties under it, in so doing expanding his description of<br />

radiosa and no longer reserving this name for the Texas form.<br />

But at the same time he did not lose the identity of the Texas<br />

form, since it remained as the subvariety texana. Then, still later,<br />

swinging back a little the other way, in the Cactaceae of the<br />

Mexican Boundary he elevated radiosa to a subspecies and the<br />

three taxa under it to varieties.<br />

Coulter even later returned M. radiosa to species status again,<br />

and added to it numerous other varieties found all the way to<br />

California, giving it the broadest definition of all.<br />

This name, therefore, has many meanings in the literature, but<br />

in the present study I use it in the sense of the original description<br />

in the Plantae Lindheimerianae. It is in no way a synonym<br />

of M. vivipara, as many have thought, but is a definite Texas<br />

and Oklahoma form not found anywhere west of Texas, and its<br />

only synonym among Engelmann’s names is the subvariety name,<br />

texana.<br />

It is one of the largest forms of this species, has large, loosestanding<br />

tubercles, and extremely fine flowers. It is a localized<br />

form of the species and, since no other variety of this cactus<br />

grows in its area, identification of field-collected plants should<br />

not be a problem.<br />

This may possibly be the Oklahoma form called Coryphantha<br />

radiosa by Lahman, but her description is so incomplete that it<br />

is hard to be certain.<br />

Mammillaria vivipara var. neo-mexicana Eng.<br />

[Coryphanta neo-mexicana (Eng.) B. & R.)<br />

“Spiny Star,” “Pincushion,” “New Mexico Coryphantha,” “Estria<br />

del tarde”<br />

Description Plate 32<br />

stems: As the species, sometimes growing as single stems, but


genus Mammillaria 129<br />

usually forming small clusters. Individual stems are spherical<br />

at first, becoming short columns when old. The maximum size<br />

seems to be 31/2 to 4 inches tall by 21/2 inches thick. The tubercles<br />

measure to 1/2 inch long and are cylindrical or sometimes<br />

somewhat flattened from top to bottom and crowded close to-<br />

gether.<br />

areoles: As the species.<br />

spines: Each areole has 20 to 40 slender radial spines to 3/8 of<br />

an inch long, white, sometimes with brown or purplish tips.<br />

These form a fringe of spines around each areole and interlock<br />

with those of neighboring clusters to hide almost completely<br />

the flesh of the plant. There are also 6 to 15 central spines on<br />

each areole of the adult plant. These are white or yellowish<br />

with purplish to light brown tips, grow to 3/4 of an inch long,<br />

and spread out in all directions and at all angles from the<br />

center of the areole.<br />

flowers: About 1 inch tall and broad, and violet in color,<br />

sometimes shading at the center to deep rose. The outer petals<br />

are narrow, pointed, and fringed. The inner petals are also very<br />

slender and sharp pointed, having smooth edges, and are very<br />

deep violet in color. The filaments are green at the bases, becoming<br />

pink above. The anthers are orange. The style is as short<br />

as the stamens and reddish in color. The stigma lobes are 7 to<br />

10 in number, pure white, short, fat, and not pointed.<br />

fruits: Green, oval in shape, and 5/8 to 15/8 inches long. The<br />

medium brown seeds are approximately 1/16 of an inch (about<br />

2 millimeters) long, ovate or almost reniform, with the bodies<br />

curved around the ventral, concave, linear hila. The surfaces of<br />

the seeds are pitted.<br />

Range. The mountain areas of central Colorado and New Mexico,<br />

south into Mexico, west into Arizona and east into Texas<br />

from El Paso and the Guadalupe Mountains to near Fort Stockton.<br />

The plant is very rarely, if ever, found in Texas south or east<br />

of Marfa.<br />

Remarks, This is a comparatively distinct member of this closely<br />

knit group. It is fairly easily distinguished from its relatives by<br />

the more numerous spines, the more closely crowded and smaller<br />

tubercles and the smaller flowers with the short, heavy, white<br />

stigma lobes only about 1/8 of an inch long. Its seeds are also<br />

distinctive.<br />

Mammillaria vivipara var. neo-mexicana is the mountain member<br />

of the species. Engelmann once remarked that it was found<br />

in the Sandia Mountains at a 13,000-foot altitude. I do not know<br />

that it has been seen since at this extreme altitude, but it grows on<br />

the hills, and its range takes in all of the high mountains of New<br />

Mexico and the Guadalupe and Davis mountains of Texas.<br />

At first, Engelmann called this form subvariety neo-mexicana of<br />

his Mammillaria vivipara subspecies radiosa, and later he spoke<br />

of it as a variety of the species. He apparently never did consider<br />

it a separate species. Small was the first one to elevate it to a<br />

separate species, calling it Cactus neo-mexicanus, and Nelson<br />

then placed it back into the Mammillarias as a full species. Britton<br />

and Rose followed, but called it Coryphantha neo-mexicana.<br />

It is clearly one of the spiny stars most removed from the typical<br />

M. vivipara var. vivipara, but since the earliest studies concluded<br />

it to be part of that species and recent usage seems to have come<br />

back to that opinion, we list it here as a variety.<br />

It also became common to list Engelmann’s other two subvarieties,<br />

borealis and texana as synonyms of neo-mexicana. To do this<br />

is to be so superficial as to overlook the differences which Engelmann<br />

noticed and outlined carefully when he first separated<br />

them, differences of spine numbers and colors, stigmas, flowers,<br />

and seeds. To insist on their existence as separate forms is not to<br />

deny that there may be intermediate specimens—as Engelmann<br />

cautions—nor to state flatly that they are all entirely separate<br />

species. The latter position obviously cannot be maintained successfully,<br />

and no doubt it was this problem, arising when all of<br />

these forms were assigned species rank, that caused the attempt<br />

on the part of some students to dismiss them as synonyms. But<br />

each of these forms has a separate, continuous range within<br />

which it presents consistent characteristics, and we feel constrained<br />

to list them as distinct varieties rather than enlarging<br />

the description of this form and making it almost meaningless<br />

in an attempt to contain them.<br />

Mammillaria vivipara var. borealis Eng.<br />

“Sour Cactus”<br />

Description Plate 32<br />

stems: As the species, but usually single and only rarely found<br />

in small clusters. Individual stems are usually spherical, but<br />

when old become more or less egg-shaped. The maximum size<br />

seems to be about 3 inches tall by 2 inches in diameter. The<br />

tubercles grow to about 1/2 inch long, are closely crowded,<br />

and overlap upward.<br />

areoles: As the species.<br />

spines: Having 12 to 22 white, slender, short radial spines<br />

and 3 to 6 central spines which are arranged much like those<br />

of the typical variety vivipara, one usually turned downward<br />

and the other 3 to 5 spreading upward just in front of the<br />

upper radials. These centrals, or at least the upper halves of<br />

them, are wholly purplish-black or maroon.<br />

flowers: Small, only about 1 to 11/2 inches tall and the same<br />

in diameter. The petals are 3/4 of an inch or less in length and<br />

1/8 or more in width. The outer petals, about 20 in number,<br />

are brownish and fringed; the inner ones are lavender, deep<br />

pink, or pale violet, and entire. All petals are sharply pointed.<br />

The filaments are pink and the anthers pale orange. The style<br />

is about the same length as the stamens, and the stigma has 8<br />

to 12 rather long, slender, pale pink lobes, which are blunt.


130 cacti of the southwest<br />

fruits: Egg-shaped, to about 3/4 of an inch long, greenish,<br />

often becoming brownish or even faintly flushed with brownish<br />

red when very ripe. The seeds are very light brown or<br />

straw-colored, approximately 1/16 of an inch (2 millimeters or<br />

so) long, oval in shape, with the surface finely pitted. The<br />

hilum is near the end of the seed and is round, or nearly so.<br />

Range. New Mexico from northwest of Santa Fe past Zuni<br />

northwest into Colorado, Arizona, and Utah.<br />

Remarks. This small cactus of the northwestern New Mexico<br />

mountains is apparently limited to the western side of the Continental<br />

Divide. It has been generally overlooked by students,<br />

and we usually find it included in their files of several other<br />

forms, while their descriptions of those other forms have therefore<br />

usually been broadened to include it, leading to much con-<br />

fusion.<br />

Engelmann first called this cactus Mammillaria vivipara var.<br />

radiosa subvar. borealis. A number of later writers ignored it<br />

entirely, but Britton and Rose listed it as a synonym of their<br />

Coryphantha neo-mexicana. Backeberg seems to leave it as a<br />

synonym of that form, which, however, he reduced to a variety<br />

of the species. How it ever got so closely connected with variety<br />

neo-mexicana is hard to understand. It seems to be the nearest<br />

relative west of the Continental Divide of the typical northeastern<br />

form, variety vivipara, even as variety radiosa can be<br />

regarded as the nearest southerly relative of that typical variety.<br />

It is much further removed from variety neo-mexicana by several<br />

characters. It is closer to variety radiosa than to any of the<br />

southern or western varieties; but it remains a much smaller<br />

plant in over-all size and in size of tubercles; it has fewer radial<br />

spines, darker-colored centrals, and probably the smallest flowers<br />

of the group, while variety radiosa has the largest. The size<br />

of seeds is about the same in these two cacti, but otherwise they<br />

differ.<br />

Its relationship to the other western forms of this complex<br />

species is more difficult, and since the variety borealis has almost<br />

universally been ignored, we find almost no attempts to<br />

deal with it in any recent works on these cacti. The reason for<br />

this fact is quite interesting.<br />

First of all, in trying to see the reason for resurrecting this all<br />

but forgotten name and to understand in general the western<br />

forms of this species, which venture into the area covered by<br />

our study far more than most people realize, we must face and<br />

deal with one of the most remarkably persistent errors in all of<br />

cactus study. Almost throughout the literature on cacti we run<br />

across the name Mammillaria aggregata. Britton and Rose<br />

thought there was a plant of this group in New Mexico which<br />

they listed under the name Coryphantha aggregata, and Mrs.<br />

Lahman even had it growing in Oklahoma. There are whole<br />

files under this name in most herbaria. It persists to the present,<br />

Benson listing M. vivipara var. aggregata (which apparently<br />

included the form variety borealis) in his scheme, while Backeberg<br />

used Coryphantha vivipara var. aggregata for a clustering<br />

form of New Mexico and on west. We will never understand<br />

the western forms until we understand this persistent name aggregata.<br />

Engelmann coined the name in 1848 in Emory’s Report, for<br />

a plant observed on the Gila River. His description of this plant<br />

was very incomplete, as he did not see its flowers or fruits. He<br />

said it made hemispherical clusters of several feet across with up<br />

to 100 to 200 heads. He started the whole misunderstanding by<br />

a simple speculation that Mammillaria aggregata, “appears to be<br />

allied to M. vivipara.” How the plant he saw on the Gila ever<br />

reminded him of the small, flat clusters of the species M. vivipara<br />

in the first place is hard to see, but it did, and the name<br />

has been applied to that species very often, even down to the<br />

present.<br />

Engelmann had made a snap judgment on this plant without<br />

even seeing flowers or fruits. He was wrong, and as soon as he<br />

knew he had made an error he corrected it manfully. Concerning<br />

this report, he wrote in the Report of the Ives Exploration<br />

in 1861, as follows: “Cereus phoeniceus, Pacif. Rail. Rep. Synop.<br />

Cact., Echinocereus coccineus Wisliz. Rep. N. Mex., note 9. This<br />

is Mammillaria aggregata, Emory’s Report, 1848, and the ‘Aggregated<br />

Cactus’ of the explorers of the western parts of New<br />

Mexico and the Gila regions.”<br />

How anything could be more clear than the above statement<br />

is hard to see. Engelmann is stating that the name is an error,<br />

actually a synonym of Echinocereus coccineus, and that there is<br />

no Mammillaria which should bear the name aggregata at all.<br />

Early students such as Coulter, Rydberg, and Wooton were<br />

aware of the true identity of this plant and did not list the name<br />

among the Mammillarias. But Britton and Rose apparently<br />

overlooked the retraction of the name and set up a species,<br />

Coryphantha aggregatus. What their plant of this name was apparently<br />

cannot be known for sure.<br />

Engelmann would no doubt be astounded could he know that<br />

one hundred years after his retraction of the name it is still listed<br />

in the most respected works on the cacti of the area. This<br />

illustrates the harm that can be done by the easily tossed-off<br />

speculation, and should certainly prompt authors to restrain<br />

themselves from casual speculations.<br />

Since all modern accounts of the M. vivipara complex in New<br />

Mexico and Arizona which I have examined still list this defunct<br />

name, usually as a vague catchall, it becomes our task to<br />

sort out the specimens filed under the name and place them in<br />

the valid taxa in which they really belong. When we do that we<br />

find that they all settle into the valid taxa comfortably, but that<br />

the process resurrects some almost forgotten varieties and that it<br />

extends the known range of some others much farther east than<br />

is usually recognized.<br />

How variety borealis, which hardly clusters at all, was included<br />

in a supposed variety aggregata of hemispherical masses<br />

to 200 heads is hard to see, but once we look for it we find it<br />

there. Or specimens may be classified under several other names.<br />

For instance, Boissevain’s Coryphantha radiosa is mainly va-


genus Mammillaria 131<br />

riety borealis, but his description of it is skewed a little to include<br />

M. vivipara var. arizonica. He never suspected that vari-<br />

ety arizonica grew in Colorado, but some of the specimens upon<br />

which he based his C. radiosa were clearly this form.<br />

If we can clear away the accretion of the years of confusion<br />

over such errors, we find growing, never profuse but surprisingly<br />

widespread, here and there upon the hills and lower mountains<br />

of northwest New Mexico and southwest Colorado and<br />

into northeast Arizona as well as Utah, a small, usually single-<br />

stemmed spiny star with few and very weak radials and few<br />

but very dark-colored centrals. It has small flowers, pale by<br />

comparison with most of its relatives, and very light brown<br />

seeds. It is retiring in habit and appearance, and probably does<br />

not mind greatly being overlooked so generally or else being<br />

merged with its more robust neighbors. Its name is rather fitting,<br />

as it is the most northern form of this species other than<br />

the typical variety vivipara.<br />

Mammillaria vivipara var. arizonica (Eng.)<br />

[Coryphantha arizonica B. & R.]<br />

“Sour Cactus,” “Arizona Coryphantha”<br />

Description Plate 33<br />

stems: Simple or occasionally clustering sparingly to form to<br />

3 or 4 stems. The individual stems are spherical to ovate or<br />

conical and robust, becoming sometimes to 4 inches in diameter<br />

and as much as 5 inches tall. The tubercles are cylindri-<br />

cal and rather loose-standing, and to 1 inch long.<br />

areoles: As the species.<br />

spines: With 13 to 20 radial spines which are slender but<br />

rigid and long, varying from about 5/8 to 3/4 of an inch as a<br />

rule, but said to have reached 11/4 inches in length. They are<br />

whitish, but not pure white, usually with brown tips. The<br />

centrals are 3 to 6 in number, spreading, rather stout, about<br />

5/8 to 7/8 of an inch long, and yellow or gray below with deep<br />

brown or purplish-brown above.<br />

flowers: Large and showy, about 2 inches in diameter and<br />

length, rose-pink to rose-purple in color. The 30 to 40 outer<br />

segments are slender and fringed and greenish-brown. The 40<br />

or so inner segments are linear and entire and sharply pointed.<br />

The stigma has 7 to 10 white, blunt lobes.<br />

fruits: Oval and green. The seeds are approximately 1/16 of<br />

an inch (about 13/4 millimeters) long, broadly egg-shaped,<br />

curved, and beaked around the short, oval, greatly concave,<br />

ventral hila. They are light brown in color, and pitted.<br />

Range. Widely in the far West, and east from northern Arizona<br />

into the extreme northwestern corner of New Mexico. Apparently<br />

restricted in New Mexico to the area bounded by an arc<br />

drawn from near Gallup past Blanco Trading Post and into<br />

Colorado near Farmington.<br />

Remarks. One of the reasons the New Mexico members of this<br />

species complex have been so hard to understand is that it has<br />

been assumed that this variety does not grow in that state, but<br />

that it is strictly an Arizona specialty. Such may be the effect<br />

of a name.<br />

But Engelmann in his original description gave no limit to<br />

this plant’s eastern penetration, saying only, “from the Colorado<br />

eastward,” and the detailed maps of its range, such as that<br />

in Benson’s Arizona Cacti show it covering the whole of northeastern<br />

Arizona right up to the border. Since we know that cacti<br />

do not respect state boundaries, it is only logical to expect it to<br />

spill over into neighboring New Mexico. This it does, from about<br />

Gallup northward, and when we recognize this fact and take<br />

the specimens which really fit this taxon from the collection of<br />

various forms put under the erroneous name, Mammillaria aggregata,<br />

as well as from where they have been deposited under<br />

other names, we have a nice array of records for the cactus over<br />

the extreme northwestern corner of New Mexico.<br />

M. vivipara var. arizonica is a robust form of the species in<br />

stem size, spination, and flowers. Although it clusters only sparingly,<br />

its stems are large and strong, with long tubercles. Its<br />

spines are not numerous as in some forms, but are stouter than<br />

those of any form yet discussed under this species. The spines,<br />

together with the loose-standing tubercles, give it the appearance<br />

of a bold, open-growing, well-protected plant which contrasts<br />

in appearance with the smaller, more delicate, more retir-<br />

ing variety borealis and variety vivipara. It more resembles variety<br />

radiosa of Texas, but has much more rigid and darkercolored<br />

spines. Its flowers are also the only other ones approaching<br />

in size and brilliance those of variety radiosa. However,<br />

they are more pinkish than the deep violet blossoms of the Texas<br />

form. On the other hand, the seeds of variety arizonica are the<br />

closest in size and shape to those of variety vivipara of any<br />

other form, but they are much lighter in color.<br />

Mammillaria vivipara var. deserti (Eng.) non L. Benson<br />

[Coryphantha deserti (Eng.) B. & R.]<br />

Description<br />

stems: As the species, except apparently always simple and<br />

unbranched. These stems are at first globose, later becoming<br />

oval or egg-shaped and growing to a maximum of about 4<br />

inches tall by 31/2 inches thick. They have short, cylindrical<br />

tubercles to about 1/2 inch long which are close-standing and<br />

compact, but not overlapping.<br />

areoles: As the species.<br />

spines: Having 22 to 35 radial Spines which are 3/8 to 3/4 of<br />

an inch long, spreading almost pectinate and interlocking<br />

with those of adjacent tubercles. The upper and lower radials<br />

are rather slender, but the lateral ones are stout and rigid.


132 cacti of the southwest<br />

They are all gray, the larger ones usually tipped with brown.<br />

There are 5 to 10 central spines, the lower 3 to 5 of these<br />

being porrect or spreading outward, heavy, awl-shaped, and<br />

1/4 to 3/8 of an inch long. The upper 2 to 5 of them spread upward,<br />

are more slender, and are 1/2 to 5/8 of an inch long. All<br />

centrals are straw- or honey-colored below with red-brown<br />

tips.<br />

flowers: Small, about 1 inch long and wide. In color these<br />

flowers are pale pinkish suffused with cream or tan. The outer<br />

segments are brownish and fringed; the inner ones are entire<br />

and often described as straw-colored with pinkish tips. There<br />

are 5 or 6 whitish stigma lobes.<br />

fruits: Short, oval, about 1/2 inch or so long. They remain<br />

green in color for a long time, but are sometimes suffused<br />

with brown or even faint maroon when very ripe. The seeds<br />

are 1/16 of an inch (11/2 to 2 millimeters) long, dark brown,<br />

pitted, obovate, and somewhat curved around the round or<br />

oval, convex or flat, ventral hila.<br />

Range. Entering New Mexico from southern Arizona. Found<br />

in the strip of New Mexico west and south of Silver City.<br />

Remarks. Originally described as a California cactus, but soon<br />

known from Nevada, this form was later traced east across<br />

Arizona. Now we find a population along the Arizona border<br />

and a short distance into New Mexico which seems clearly this<br />

plant, marking another penetration into our area by a western<br />

form and giving this cactus a very wide range.<br />

The cactus as it grows in New Mexico and as described above<br />

is not to be confused with another plant described under this<br />

name in many accounts of the western cacti. Borg and Boissevain<br />

equate this cactus with Mammillaria alversonii Coult., a<br />

larger, clustering form restricted to southern California. Benson<br />

gives for M. vivipara var. deserti (Eng.) L. Benson a description<br />

almost totally unlike that of either Engelmann or Coulter and<br />

with limits which would appear to exclude the type itself, listing<br />

15 to 20 slender white radials and only 3 to 5 centrals—<br />

white, tipped with brown. Only the small, straw-colored flower<br />

of his description is the same as in the earlier descriptions,<br />

and it seems obvious that whatever his plant may be it is not<br />

the same as what Engelmann and Coulter meant by M. deserti—<br />

nor is it the cactus we see in southwestern New Mexico.<br />

Be that as it may, we have a cactus with a definite range in<br />

New Mexico which duplicates the more western form originally<br />

described as M. deserti. It is a robust but single-stemmed cactus<br />

whose short, compact tubercles are almost completely covered<br />

by its very numerous, interlocking spines. Only variety neomexicana<br />

has such a profuse covering of spines, and the spines<br />

of that cactus are white and much more slender. This cactus,<br />

with its ashy-gray radials and its brown centrals which are all<br />

relatively heavy, gives quite a different impression. It most resembles,<br />

of any form in this species, variety arizonica, but is<br />

much more squat and compact, with much shorter tubercles, and<br />

many more and much more appressed, more stout spines than<br />

on that relative. The form, spines, and general appearance of<br />

variety deserti are similar enough to those of Echinocactus intertextus<br />

var. dasyacanthus Eng. that I have several times felt very<br />

foolish for at first mistaking it for that cactus of an entirely<br />

different genus.<br />

Originally treated as a separate species, this cactus has been<br />

incorporated, along with others, into the broad Species complex<br />

of M. vivipara. If this placement is correct, it is the most distant<br />

from the typical form of that species, both in its characters and<br />

its range. The population of this cactus found in New Mexico<br />

forms a distinct entity there, and must be taken into account in<br />

gaining any adequate concept of this complex as it occurs in<br />

that state. Whether or not one sees it as the same cactus as the<br />

California variety deserti, which I believe it to be, no collection<br />

of New Mexico cacti is complete without it.<br />

Mammillaria fragrans (Hester)<br />

[Coryphantha fragrans Hester]<br />

Description Plate 33<br />

stems: Almost always occurring singly, very rarely with one<br />

or two branches from the base. Becoming conic to cylindrical<br />

when older, and growing to a maximum of 8 inches tall by<br />

about 3 inches in diameter. The tubercles are 3/8 to 5/8 of an<br />

inch long, usually being around 1/2 inch on typical plants.<br />

They are soft and oval in shape, being flattened dorsally and<br />

somewhat broadened above the base to about 3/8 of an inch<br />

wide before they taper toward the tips.<br />

areoles: Monomorphic, mature areoles lengthening to form<br />

grooves extending nearly to the bases of the tubercles, with<br />

white wool in the grooves when they are young, but later<br />

bare.<br />

spines: There are 20 to 30 slender radials to 1/2 inch long, and<br />

white in color. There are 6 to 12 centrals to 5/8 of an inch<br />

long, which spread at all angles from the center of the areole.<br />

They are mostly purple or very dark brown, only the bases of<br />

them being gray; with age they usually become entirely black.<br />

flowers: About 11/2 to 2 inches across and tall, cup-shaped<br />

or even bell-shaped, magenta or reddish-purple in color. The<br />

outer petals are greenish with pink edges fringed with long<br />

white hairs. The inner petals are narrow, pointed, reddish-<br />

purple shading to light pink at the edges from greenish bases,<br />

with edges entire or very slightly ragged near the tips. The<br />

filaments are greenish below to pink above. The anthers are<br />

orange. The style is about the length of the stamens or a little<br />

longer. The stigma has 6 to 12 pure white to pinkish lobes<br />

which are long and more or less pointed, but rather thick.<br />

fruits: Greenish, becoming yellow-green when ripe, oval in<br />

shape, and to 1 inch long. The seeds are kidney or bean shaped,<br />

approximately 1/8 of an inch (21/2 to 3 millimeters) long, and


genus Mammillaria 133<br />

much flattened. Their translucent surface is rich red-brown<br />

and shiny smooth, with no pitting, but with a pattern of minute<br />

checks showing through from inner layers when viewed<br />

with a microscope. The hilum is ventral or nearly so, oval or<br />

oblong.<br />

Range. The Big Bend of Texas, from near Sanderson on the east<br />

to the Rio Grande west of Alpine, but apparently not north of<br />

the southern slopes of the Davis and Van Horn mountains.<br />

Remarks. Here is yet another form which only the closest observation<br />

will reveal to be separate from the complex group just<br />

listed. It apparently was not distinguished from M. vivipara<br />

var. neo-mexicana by any of the earlier students, which is not<br />

surprising since its armament of spines comes within the range<br />

of that variety, and the accounts of some other students can<br />

only be understood by assuming that they failed to distinguish<br />

it from M. vivipara var. radiosa. For these reasons it was overlooked<br />

until described as a separate species by Hester in 1941.<br />

Mr. Hester carried on extensive studies of the seeds of cacti, and<br />

no doubt his discovery of this plant with seeds so much larger<br />

than those of M. vivipara var. neo-mexicana and different in<br />

almost every detail of shape and surface first caused him to<br />

notice its other less marked differences from that plant. It was<br />

in the same way, by noticing the remarkable seeds, that I first<br />

learned that my supposed variety neo-mexicana specimens from<br />

near the Rio Grande in the Big Bend were different from those<br />

collected farther northwest. Much later I learned that I had<br />

stumbled upon Hester’s cactus.<br />

There are few definite or measurable characters of the plant<br />

body by which to recognize the cactus before it fruits. It is a<br />

much larger cactus than variety neo-mexicana, growing double<br />

the height of that species when at its maximum, and it also differs<br />

in that it almost never branches. Its tubercles are also unusual<br />

in that they broaden out above their bases and are flattened<br />

on their upper sides, this giving them a curious ballooned<br />

appearance which seems unique in this form, but which is not<br />

always obvious in specimens suffering from drought or winter<br />

shrinkage. Its central spines are very dark, the dark coloring<br />

extending farther down on them than on any other relatives<br />

except perhaps some of the northwestern specimens of M. vivipara<br />

var. borealis.<br />

The flower of M. fragrans differs from its relatives in several<br />

minor ways. Its petals are shorter. Its stigmas are intermediate<br />

in length between those of the eastern M. vivipara var. radiosa<br />

and the western variety neo-mexicana, but are usually pure<br />

white, and only sometimes a light pink. Its flowers are very<br />

fragrant with a sweet scent from which comes its name, while<br />

those of variety neo-mexicana have almost no scent and those<br />

of the eastern M. vivipara forms have a very strong and pungent,<br />

but very green scent which is not sweet.<br />

Hester seems to have failed to distinguish sometimes between<br />

his own new form and the old form, M. vivipara var. radiosa,<br />

and as a result he gives the range of M. fragrans as extending<br />

almost to Fort Worth, Texas, and into southern Oklahoma. In<br />

actuality M. fragrans differs almost as completely from variety<br />

radiosa as it does from the other M. vivipara varieties, and the<br />

range of variety radiosa stops just northeast of where the range<br />

of this plant begins.<br />

This cactus may also be part of the M. vivipara complex. It<br />

is similar enough to it so that the assertion would be easy. Yet<br />

with no real evidence for the combination, and with the seeds<br />

so different, such a combination seems premature.<br />

This is a rare cactus restricted to the lower Big Bend of Texas,<br />

but on some hillsides or in some alluvial valleys it may grow in<br />

fairly extensive populations. Perhaps it extends into Texas from<br />

Mexico, but this is not known. During some limited trips into<br />

the very rough mountains across from the Texas Big Bend I did<br />

not observe the cactus.<br />

Mammillaria tuberculosa Eng.<br />

[Escobaria tuberculosa (Eng.) B. & R.]<br />

Description Plates 34, 35<br />

stems: Globular to egg-shaped at first, becoming upright cylinders<br />

1 to 2 inches thick, said to grow to 7 inches tall, but<br />

not usually exceeding 5 inches and often much shorter. The<br />

stems are single until large, then sprout slowly around their<br />

bases to form small clusters. The color of the surface is a dull<br />

gray-green. The stem is composed of many separate tubercles<br />

usually about 3/8 of an inch long, but sometimes not over 1/4<br />

of an inch. These tubercles are practically cylindrical when<br />

young, but their bases broaden horizontally so that they become<br />

somewhat rhomboid and about as wide as they are tall<br />

when old. They are somewhat crowded, and mature tubercles<br />

turn upward so that they overlap each other.<br />

areoles: Small and round on immature stems. On mature<br />

stems a groove runs all the way to the base of each tubercle<br />

on its upper side, as the linear extension of the areole from<br />

the spinous portion at the tip of the tubercle to the floral part<br />

of it in the axil. When young, this groove is filled with wool,<br />

but when old only a tuft of this white wool remains in the<br />

axillary end of the groove.<br />

spines: On each areole there are 20 to 30 radial spines, which<br />

are slender but stiff, radiating in all directions around the<br />

areole. They are white and vary in size, those at the top of the<br />

areole being very small bristles, while those around the sides<br />

and bottom of the areole are to 3/8 of an inch long and rather<br />

firm. There are 4 to 9 central spines which are gray-white<br />

with purplish ends; 4 of these are always much heavier than<br />

the radials, with 3 of them spreading upward and to 5/8 of an<br />

inch long, while the lowest of these 4 always stands abruptly<br />

outward or turns a little downward. This lowest central is the<br />

heaviest spine of all, and is conspicuous. On young plants<br />

there are only these 4 centrals, but with age there may be 3


134 cacti of the southwest<br />

to 5 more centrals added above the others, standing upright<br />

with the earlier upper ones; they are never quite as heavy or<br />

as long as the first and are intermediate between those and<br />

the radials. There is a definite tendency for the spines to be<br />

shed from the older part of the stem, leaving the base bare of<br />

spines.<br />

flowers: 3/4 to 13/8 inches across and 3/4 to 1 inch tall; opening<br />

widely. The color of the flower is a very delicate lavender-<br />

white or extremely pale purple. The outer petals are greenish-<br />

brown with almost white, fringed edges. They are to 3/4 of an<br />

inch long and 1/8 of an inch wide, and there are 16 to 18 of<br />

them. The inner petals are 10 to 15 in number, 3/4 to 1 inch<br />

long, and only 1/8 of an inch wide, very pale lavender fading<br />

to almost white at the edges. The filaments are cream, while<br />

the anthers are cream or pale yellowish. The style is white, the<br />

same length or just longer than the stamens. The stigma is<br />

made up of 5 or 6 white lobes which may be short and thick<br />

or slender and to as much as 3/16 of an inch long.<br />

fruits: Egg-shaped to oblong, about 3/4 of an inch long, and<br />

bright red. The seeds are very small, usually around 1/2 milli-<br />

meter long, and brown in color, with the surface pitted.<br />

Range. The mountains of the Big Bend region of Texas, and<br />

from there southward into Mexico, westward across southern<br />

New Mexico, and into southern Arizona.<br />

Remarks. Mammillaria tuberculosa is usually spoken of as one<br />

of the most common cacti of far west Texas and southern New<br />

Mexico. In the trade, plants are shipped by the hundreds from<br />

this area under the name Escobaria tuberculosa. However, it<br />

appears that the species is actually much less common than is<br />

usually thought, at least today, and that most of those specimens<br />

now in collections under this name are actually another species.<br />

The reasons for this confusion are several, and we must understand<br />

them if we are to understand this species. There are several<br />

plants known commonly as Escobarias which are very close in<br />

most of their characteristics. They are small plants with many<br />

spines and small flowers, and it requires painstaking work, usually<br />

with a magnifier, to recognize the characters which divide<br />

them. We are often reluctant to go to this trouble, and the one<br />

thing seized upon to distinguish M. tuberculosa from its relatives<br />

with a minimum of effort—indeed, the character for which it is<br />

named—is an unfortunate choice, since at least two other forms<br />

show this fully as much as it does. This is the tendency of these<br />

cacti to shed their spines on their older tubercles at the bases of<br />

the stems. These old tubercles then become corky and dead-<br />

appearing, rather unsightly bumps. Perhaps my own experience<br />

with these plants should show the problem, as well as illustrate<br />

the differences between the confusing forms.<br />

I remember well when I collected plant after plant, mature<br />

specimens of which showed this naked, rather ugly base—so,<br />

knowing no better, I had to call them all M. tuberculosa. Many<br />

of these specimens bloomed, and I thought I had the species.<br />

They matched those my collector friends called by that name<br />

and the bins of dealers were full of the same plants under the<br />

same name.<br />

But what about its relatives—M. dasyacantha, for instance?<br />

Try as I might, I couldn’t find this closely related form. Everyone<br />

was vague about it, and finally I was persuaded that it was<br />

merely a minor variation of M. tuberculosa, for all practical<br />

purposes the same.<br />

During all of this time I had one specimen which appeared<br />

slightly different from the others. I was dissatisfied with it because,<br />

year after year, it refused to bloom, even when growing<br />

beside all of the others which put out their small brownish-pink<br />

flowers in numbers. Finally, one June, this plant bloomed—not<br />

with flowers like the others, but with flowers nearly twice as<br />

large and opening out widely where the others remained at best<br />

bell-shaped, and with a clear whitish color where the petals of<br />

the others were pinkish with definite brown midlines. Its 5 or 6<br />

white stigma lobes contrasting with the 5 very green stigmas of<br />

the common plants emphasized the difference.<br />

Only after discovering these differences did I get my magnifier<br />

and compare the spines of these plants, and to my delight I<br />

found that my newly flowered specimen had 5 centrals with the<br />

upper ones standing upright against the upper radials while the<br />

lower one stood out and slightly downward and was especially<br />

heavy. Examination showed that my common, brownish-flowered<br />

plants all had 7 or more comparatively slender centrals<br />

spreading irregularly. Much later I was able to observe that my<br />

reluctant bloomer had brown seeds, while my other plants had<br />

black seeds. I had M. tuberculosa and M. dasyacantha side by<br />

side, both with naked, corky bases.<br />

M. tuberculosa, when known from its relatives, is apparently<br />

a plant much less often seen than it is thought to be, while that<br />

similar cactus coming in such numbers out of the Big Bend and<br />

called so widely by this name is really its relative, M. dasyacantha.<br />

M. tuberculosa is easily distinguished when in bloom by<br />

the details of its larger and much more delicately colored flowers,<br />

but it blooms reluctantly in cultivation. It is much more<br />

difficult to distinguish without the flowers, but the arrangement<br />

of the central spines—particularly the presence of the stout<br />

lower central standing out by itself—is the best clue. In general,<br />

it is a smaller plant than M. dasyacantha, its spines having purplish<br />

ends instead of brown, and not as completely obscuring<br />

the stem as do those of its relatives.<br />

This cactus has also been referred to by the name Mammillaria<br />

strobiliformis, and this name still persists in some authors,<br />

making it necessary for us to review the origins of these names<br />

so that we may evaluate them.<br />

In 1850 Scheer described what most agree was this cactus,<br />

giving it the name M. strobiliformis. His description, however,<br />

was vague, and in 1856, when Engelmann first studied the plant,<br />

he thought he had a different cactus and originated a new name<br />

for it, M. tuberculosa. Later, realizing his mistake, Engelmann<br />

tried to retract his own name, stating, in the “Corrections to the<br />

Cactaceae of the Boundary”: “M. tuberculosa is clearly identical


genus Mammillaria 135<br />

with M. strobiliformis Scheer in Salm. Hort. Dyck (1850), as I<br />

have ascertained by a careful examination of the original specimens<br />

(now dead) in the collection of Prince Salm. Mr. Scheer’s<br />

name, having the priority, must be substituted for mine.”<br />

It would seem that nothing could be more clear, and the periodic<br />

return to the use of the name M. strobiliformis even to today<br />

is based upon this statement. But the ways of taxonomic<br />

priority often lead through mazes of cross-references, and it was<br />

soon noted that the name M. strobiliformis had already been<br />

twice used before Scheer’s use of it. In 1848 Muehlenpfordt had<br />

applied the name to what is generally agreed to be the already<br />

described M. sulcata Eng. Then, also in 1848, in the Wislizenus<br />

Report, Engelmann himself had described a plant under that<br />

same name. There has been rare agreement through the years<br />

since that Engelmann’s M. strobiliformis of 1848 is actually<br />

Echinocactus (Neolloydia) conoidea, first described by De Candolle<br />

in 1829. This leaves it of little interest to us here, but all<br />

of this does render the M. strobiliformis of Scheer twice a homonym<br />

and, therefore, an invalid name for the plant we are discussing,<br />

and it leaves Engelmann’s M. tuberculosa as, after all—<br />

unless there is another turn to the maze which we have missed—<br />

the first name for this plant valid under taxonomic rules.<br />

Borg repeats Quehl’s list of five varieties under this species,<br />

but the descriptions of these are very general and seem almost<br />

impossible to distinguish in actuality. Several of them appear to<br />

indicate the two forms listed next in our study, seeming to show<br />

characters clearly outside the limits of this species. However, I<br />

have noticed that the New Mexico specimens of this species are<br />

often slightly different from the Texas ones, being smaller, with<br />

more rounded, often almost spherical, fruits. Further study of<br />

these differences might lead toward some of Quehl’s varieties or<br />

might show these differences as only environmental.<br />

The name Escobaria orcuttii appeared in a catalog, without<br />

description, in 1925. In 1926 Rose mentioned a Neolloydia orcuttii.<br />

It was said by him to have 15 centrals, but a rose-colored<br />

flower and 6 long, white stigmas. These characters would seem<br />

to give a curious intermediate between M. tuberculosa and M.<br />

dasyacantha. But no location for the plant was ever given except<br />

“U.S.A. (Texas?).” The plant has not been reported by any<br />

recent students, and it seems best to dismiss it entirely, at least<br />

until it can be found and described exactly.<br />

Mammillaria dasyacantha Eng.<br />

[Escobaria dasyacantha (Eng.) B. & R.]<br />

Description Plates 34, 35<br />

stems: Practically spherical when young, becoming elongated<br />

and cylindrical when older. The young plants Consist of single<br />

stems, and they often remain this way all their lives, but occasionally<br />

old plants sprout at their bases to form small clusters<br />

of several stems. They sometimes grow to a maximum of<br />

8 inches tall and 23/4 inches thick. Their tubercles are cylindrical<br />

and straight in young plants, but become definitely flattened,<br />

somewhat overlapping, and 3/8 to 1/2 inch long on old<br />

plants. The oldest tubercles on the bases of the plants usually<br />

become bare of spines and remain as discolored, dead-appear-<br />

ing bumps.<br />

areoles: The spine-bearing portions of the areoles are at the<br />

tips of the tubercles, the floral part in the axils, and the<br />

tubercles are marked all the way to their bases by grooves,<br />

woolly when young but nearly naked when old, which are<br />

the narrowed portions of the areoles connecting the two extremities.<br />

Immature areoles consist of merely the small, round-<br />

ed spinous portion.<br />

spines: Radial spines 25 to at least 35 in number, radiating<br />

all around the areole. These are white in color, usually about<br />

1/2 inch long, but sometimes to almost a full inch in length.<br />

They are very slender, the upper ones being almost bristlelike.<br />

There are also from 7 to as many as 17 centrals which<br />

fill the center of each areole and spread irregularly in every<br />

direction. These are slender, some of them being almost as<br />

slender as the radials. They vary in length, most being about<br />

5/8 of an inch long, but occasionally some are as short as 3/8<br />

of an inch, and some—particularly the upper ones—sometimes<br />

grow to a full inch. The lower parts of them are white,<br />

while the tips are reddish-brown.<br />

flowers: 1 inch tall and 1/2 to 3/4 of an inch across, not opening<br />

widely, whitish to pale pink in color. The outer petals are<br />

1/2 to 5/8 of an inch long, 3/16 of an inch wide, and pointed,<br />

with the edges fringed. The midlines of these are greenish-<br />

brown, fading to whitish at the edges. There are 8 to 10 of<br />

them. There are 13 to 16 inner petals, which are the same<br />

width and length, and taper gradually to pointed tips. They<br />

have brownish midlines; the edges are tan, whitish, or pink.<br />

The filaments are pale pink or cream, the anthers bright yellow.<br />

The style is bright green and ends in 4 or 5 very short,<br />

very green stigma lobes which are deeply grooved on their<br />

ventral sides.<br />

fruits: The fruits of this plant are egg-shaped to somewhat<br />

elongated, 1/2 to 3/4 of an inch long, and dark red or scarlet.<br />

The seeds are small, practically round, about 1/2 to 1 millimeter<br />

long, pitted deeply on their surfaces, and a shiny black<br />

color.<br />

Range. The mountains of the lower Big Bend region of Texas,<br />

extreme southern New Mexico, and south into Mexico.<br />

Remarks. In extreme southwestern Texas this is the common<br />

species of the group often called Escobarias. It is also the largest<br />

species of the group. It is found on slopes and ledges in the<br />

mountains, and even on the summits of the higher mountains,<br />

often being found growing in the moss-filled crevices of high,<br />

exposed rock surfaces. In fact, some of those locations show<br />

many young plants without any mature ones, apparently be-


136 cacti of the southwest<br />

cause the seeds have germinated where there is too little soil to<br />

support them to maturity. I had to bring home some of these<br />

miniature specimens from rocky crevices near the summits of<br />

the Chisos Mountains and grow them several years before they<br />

put on mature spination, bloomed and fruited, and I knew they<br />

were this species. It is interesting to speculate about the means<br />

by which seeds are distributed to these high locations. I would<br />

suspect that birds deposit them there after eating the fruits of<br />

lower-growing mature plants.<br />

The distinguishing characters of M. dasyacantha were discussed<br />

in connection with the preceding species, the other species<br />

with which this one is so widely confused, and under whose<br />

name I have seen scores of these cacti being distributed. Without<br />

flowers M. dasyacantha is recognized by its comparatively<br />

thicker stems, its numerous, slender central spines spreading irregularly<br />

and without a conspicuous, heavier central standing<br />

outward or downward in the areole. With its flowers it is easy<br />

to distinguish, because its flowers do not open widely, and they<br />

have conspicuously green stigmas, fewer outer petals, and more<br />

as well as broader inner petals than the flowers of M. tuberculosa.<br />

When fruited it may be told by its shiny black seeds which<br />

are so different from those of M. tuberculosa that Hester felt it<br />

necessary to propose a separate genus, Escobesseya, with this<br />

species as its type species.<br />

It seems important to note that the juvenile plants of M. dasyacantha<br />

are so markedly different in almost all characters from<br />

the adults as to be taken for other forms. These young plants are<br />

spherical, sometimes even with depressed tops. When very young<br />

their tubercles do not possess any grooves, they have 18 to 22<br />

white, translucent radial spines 1/8 of an inch long, and they<br />

have uniformly only 2 central spines per areole, one pointing<br />

directly upward and one downward, each 3/16 of an inch long.<br />

By the time the plants get to be about 1 inch in diameter, the<br />

new tubercles begin to have short grooves, the radials begin to<br />

be up to 1/4 of an inch long, and there begin to be 4 centrals to<br />

the areole arranged as points of a cross and to 1/2 inch long.<br />

When the plant is about 13/4 inches in diameter and height more<br />

centrals start to appear, and soon the plants begin to lengthen<br />

out and take on their adult appearance.<br />

Although this cactus is fairly common, it is a plain one in all<br />

of its stages, and the extent to which both this and the previous<br />

species have been ignored by almost everyone may be seen by<br />

the fact that neither of them seems to have been given a com-<br />

mon name, even in the local area.<br />

Mammillaria duncanii (Hester)<br />

[Escobesseya duncanii Hester]<br />

Description Plate 35<br />

roots: Having one, or often several fleshy, carrot-like tap-<br />

roots 1/4 to 1 inch in diameter and sometimes to as much as a<br />

foot long before tapering.<br />

stems: Practically spherical to broadly ovate or somewhat<br />

conical, to about 2 inches tall by 11/4 inches in diameter. The<br />

stems are usually single, but a few double- or triple-stemmed<br />

specimens have been seen. The stem is covered by small tubercles<br />

about 1/8 of an inch long.<br />

areoles: Monomorphic, being elongated into a groove from<br />

the spinous portion on the tips of the tubercle to the floral<br />

portion in the axil, with some white wool in the axillary portion<br />

at first.<br />

spines: There are 24 to about 40 radial spines which are slender,<br />

straight, 3/16 to 3/8 of an inch long, and white or white<br />

with very slightly brownish tips. There are 3 to 16 central<br />

spines, although the usual number is about 8. These spread<br />

very widely and are around 3/8 of an inch long, white, tipped<br />

with light brown.<br />

flowers: 5/8 to 3/4 of an inch long and about 1/2 inch wide,<br />

not opening widely. They are pale pink, or whitish with pinkish<br />

zones. The ovary is smooth and rounded. The outer perianth<br />

segments are pointed, the outermost with a few cilia on<br />

their margins. They are pinkish with whitish edges. The inner<br />

segments are entire, pointed, their midlines pink and their<br />

edges whitish. The filaments are whitish, the anthers pale<br />

orange. The stigma has 4 to 6 yellow or sometimes almost<br />

chartreuse lobes.<br />

fruits: Club-shaped, 1/2 to 3/4 of an inch long and about 3/16<br />

of an inch wide, not including the persistent perianth, becoming<br />

bright red when ripe. The seeds are practically spherical,<br />

black in color, with the surface pitted and the hila basal.<br />

Range. A small area of Brewster County, Texas, a few miles<br />

west and north of Terlingua, Texas. Said to have been collected<br />

also in New Mexico, but this fact has not been verified.<br />

Remarks. This obscure little cactus was described in 1945 by the<br />

very meticulous observer of Big Bend cacti, J. Pinckney Hester.<br />

He described it under the name of Escobesseya duncanii. It has<br />

hardly been seen since.<br />

I was fortunate to receive a number of specimens of this species<br />

from Mr. Homer Jones, a dealer friend in Alpine. He had<br />

gotten them from the type locality a few miles north of Terlingua,<br />

Texas, and was so good an observer himself that he recognized<br />

he had something unusual. Although both Mr. Jones<br />

and I have searched in the area since that time, we have not<br />

been so fortunate as to collect the cactus again. Neither has it<br />

appeared in the large number of cacti from the general region<br />

passing through his business. I believe that it is extremely rare.<br />

Along with the giving of the type locality in Brewster County,<br />

Texas, in his original description of the plant, Mr. Hester<br />

added that it was also found “in a low range of mountains a<br />

few miles west of Hot Springs, New Mexico.” This has prompted<br />

much searching for the species in New Mexico. I have searched


genus Mammillaria 137<br />

for it there myself without success. Some have reported finding<br />

it in New Mexico, but the two or three of these plants from<br />

New Mexico labeled thus which I have been privileged to examine<br />

in herbaria have definitely not been this form. They appear<br />

to be the smaller form which M. tuberculosa often takes in<br />

New Mexico; one of them with brown seeds which prove this<br />

identification. Therefore, I think that the question of whether<br />

the plant grows in New Mexico is still in doubt. Hester may<br />

have overstated the range of the cactus because of misidentifying<br />

some New Mexico specimens, as he did that of M. fragrans<br />

by including with it some specimens of M. vivipara var. radiosa.<br />

Another interesting possibility is that the New Mexico in his<br />

article is a misprint and he meant to refer to Hot Springs, Texas,<br />

which is in Brewster County. However this may be, it seems<br />

necessary to doubt the existence of this cactus in such a discontinuous<br />

range in New Mexico until obvious specimens of it come<br />

forth from that state, and there seem to be none so far.<br />

In the general characteristics of its spines, its flowers, its fruits<br />

and seeds, the cactus is very close to M. dasyacantha. Except for<br />

certain peculiarities, it might pass for a dwarf form of that cactus.<br />

Its seeds are so close to those of M. dasyacantha that Hester<br />

placed it with that species in his proposed new genus, Escobesseya,<br />

apart from all others. It is, however, set off from M. dasyacantha<br />

by the unusual fleshy taproot, by its smaller size and its<br />

yellow stigma lobes. It is a most interesting, but very unobtru-<br />

sive little rarity.<br />

Mammillaria albicolumnaria (Hester)<br />

[Escobaria albicolumnaria Hester]<br />

“The White Column,” ‘The Silverlace Cactus”<br />

Description Plate 36<br />

stems: Oblong or cylindrical almost from the beginning,<br />

nearly always single, only a few plants having been seen<br />

with 2 or 3 stems. These stems are said to grow to 10 inches<br />

tall, but are rarely seen over 5 inches. The maximum diameter<br />

seems to be about 21/2 inches. The tubercles of the stem are<br />

about 3/8 of an inch long, tapering to points from rhomboid<br />

bases about as wide as they are tall. The bases of old plants<br />

become naked of spines in most cases.<br />

areoles: Monomorphic, but elongated on mature stems to<br />

run the whole length of the tubercle, the spinous part at the<br />

tip, the floral part in the axil, and the rest of the areole a<br />

narrow groove connecting the two. On younger specimens the<br />

areoles usually extend only part way down the tubercles.<br />

There is white wool in the grooves when they are young, but<br />

most of this disappears with age.<br />

spines: Each areole has 25 to sometimes at least 35 radial<br />

spines, which are fine and some of them almost bristle-like<br />

but which are very rigid and brittle. They are to 3/8 of an inch<br />

long, are very white in color, and are somewhat translucent.<br />

There are also 11 to as many as 17 central spines to 5/8 of an<br />

inch long. They are heavier than the radials and fill up the<br />

center of the areole, spreading in all directions. They are pink<br />

or very light red when growing, then translucent white with<br />

red-brown tips. The spines are all very brittle.<br />

flowers: Small, not opening widely, but remaining funnelshaped.<br />

They are pink in color and are usually only 3/8 to 5/8<br />

of an inch wide and 3/4 to 1 inch tall. The outer petals are<br />

greenish-brown in the midlines with whitish edges fringed all<br />

the way to the pointed tips. There may be 16 to 26 of them.<br />

The inner petals are 25 to 26 in number, pale pink to whitish,<br />

narrow, and pointed in shape, and their edges are not fringed.<br />

They are about 1/8 of an inch wide. The filaments are white<br />

and the anthers yellow. The style is shorter than the stamens<br />

and pink in color. The stigma is made up of 3 to 7 white or<br />

pale pink lobes which are short and thick.<br />

fruits: On this cactus the fruits are oblong or club-shaped,<br />

to 5/8 of an inch long and 1/4 of an inch thick. The lower half<br />

of the fruit remains greenish in color when ripe, but the upper<br />

half becomes a pale yellowish or apricot color. The seeds are<br />

brown, pitted on the surface, and about 1 millimeter long.<br />

Range. Found only in the extreme southwestern corner of Brewster<br />

County, Texas, near the localities of Terlingua and Lajitas.<br />

Remarks. This little cactus was not found by the early students<br />

who hunted cacti in Texas, probably because of the limited size<br />

and the ruggedness of the area in which it grows, one of the<br />

most inaccessible locations in the Big Bend. It was first described<br />

in 1941, and very few people saw it for some years after that.<br />

In recent years, however, dealers have had crews of men combing<br />

the mountains of the Big Bend for cacti, and hundreds of<br />

this species have gone into the trade, mostly under the name of<br />

Mammillaria (Escobaria) dasyacantha.<br />

Some students would like to combine this cactus with the<br />

three just listed into one species, but it is impossible to do this<br />

without ignoring the differences in their seeds which are great<br />

enough so that Hester, who no doubt observed more of them<br />

alive than any other student, felt it necessary even to place them<br />

in different genera. And this cactus is the one of the four which<br />

has spines and flowers most like M. dasyacantha but seeds like<br />

M. tuberculosa, so that it cannot well be placed as a variety of<br />

either.<br />

In general plant and spine characters this form seems very<br />

much like M. dasyacantha. It never grows as large, however,<br />

and usually does not cluster. It is set off from that species by<br />

the translucent whiteness and the brittleness of its spines, which<br />

break easily upon handling, rather than by their numbers or arrangement.<br />

I have often demonstrated this character by breaking<br />

spines off a specimen with the pressure of a finger. This may<br />

seem a small point upon which to base a species, but there are<br />

other confirming characters. Its many more numerous petals,


138 cacti of the southwest<br />

white stigmas, greenish-apricot fruits, and brown, different<br />

shaped seeds set it apart effectively from M. dasyacantha, with<br />

its fewer petals, green stigmas, scarlet or deep red fruits, and<br />

black seeds.<br />

Backeberg, on the other hand, considers it as synonymous with<br />

Quehl’s variety durispina of M. tuberculosa. However, the spines<br />

themselves very easily show it is not M. tuberculosa, as it has<br />

too many of them and lacks the heavy lower central which is<br />

characteristic of that cactus. Add to that the great difference in<br />

flower form, size, number of petals; the pale color of our plant’s<br />

fruits as compared with the rich red of those of M. tuberculosa;<br />

and the size of its seeds, which are up to twice those of that<br />

species, and it seems it cannot be put under that species either.<br />

So M. albicolumnaria seems to stand, a charming little “white<br />

column” from the very wildest mountains of the Texas Big Bend,<br />

which has become such a favorite with collectors as to be the<br />

only one of these four to possess a generally used common name.<br />

Mammillaria varicolor (Tieg.)<br />

[Escobaria varicolor (Tieg.) Backbg.]<br />

Description Plate 37<br />

stems: Apparently single except when injured, in which case<br />

they branch to form several heads. Normal stems are eggshaped,<br />

to 5 inches tall and nearly as broad with tubercles to<br />

about 1/2 inch long, conical from broadly flattened bases, turning<br />

upward to overlap greatly when old.<br />

areoles: On mature stems the areoles are elongated to run as<br />

narrow grooves from the spinous portions at the tips of the<br />

tubercles to the floral portions in the axils. They are naked<br />

except for large tufts of white wool at the floral region. This<br />

wool makes the growing tips of the plants very fuzzy, and it<br />

persists on old tubercles, making the axils woolly.<br />

spines: There are 15 to 20 very slender, white, semitranslucent<br />

radial spines. Some of them are to 1/4 of an inch long, but<br />

many of them, especially the upper ones, are weak bristles as<br />

short as 1/8 of an inch long. There are usually 4, but occasionally<br />

5 centrals, which are to about 1/2 inch long, heavier than<br />

the radials, and yellowish at the bases, with the upper part of<br />

each central brownish or purplish. They are semitranslucent<br />

and hornlike. The lower central stands out, perpendicular to<br />

the plant body or downward, while the 3 or 4 upper centrals<br />

spread fanwise in front of the upper radials.<br />

flowers: Almost pure white, pink, or very pale rose in color.<br />

They are 3/4 of an inch long and 11/4 inches in diameter. The<br />

outer petals are 11 or 12 in number, 3/4 of an inch long,<br />

fringed all the way on their edges, and brownish with pinkish<br />

edges. The inner petals are 26 or 27 in number, 1 inch long<br />

and 1/8 of an inch wide, pointed, their edges smooth, and<br />

various in color, as above. The filaments and anthers are<br />

bright yellow. The style is a little longer than the stamens and<br />

white. The stigma has 5 or 6 white lobes which are about 1/8<br />

of an inch long and slender.<br />

fruits: Ellipsoidal or sometimes slightly curved club-shaped,<br />

with the wilted perianth persistent. Approximately 1/2 to 5/8<br />

of an inch long by only 3/16 to 1/4 of an inch in diameter. In<br />

color, when ripe, they are a medium bright rose-red. They<br />

ripen in August.<br />

Range. Originally given only as southwestern Texas. I have<br />

collected specimens from only a small range running from near<br />

Marathon, Texas, to about 12 miles north of Alpine, Texas.<br />

Remarks. This is a very obscure little cactus which has no doubt<br />

been mistaken many times for other forms. It takes a keen eye<br />

to recognize it from the stem characters alone. When it blooms,<br />

however, the differences of the flowers from all other forms are<br />

obvious. Tiegel says that the flowers vary from almost white to<br />

carmine rose, for which variation he named it. Most which I<br />

have seen were very nearly pure white to a very pale rose in<br />

color. This is vastly different from the magenta or purple of the<br />

other forms most similar to it. Beyond this the yellow filaments<br />

and less numerous stigma lobes distinguish it from them.<br />

In his original description of it Tiegel called it Coryphantha<br />

(Escobania) varicolor, apparently not wishing to commit himself<br />

as to which of these no doubt too strictly drawn genera it<br />

really belonged with. There was good reason for doubt as to<br />

which of these microgenera it really belonged to, since the fruits<br />

are practically the only means of distinguishing the two microgenera,<br />

and the fruits of this species were never described. Backeberg,<br />

however, considered it an Escobaria. Since I have observed<br />

the fruits, which proved to be red in color, if I thought it necessary<br />

to place it in either of these microgenera, I would have to<br />

agree with Backeberg and place it in Escobaria.<br />

Tiegel described the cactus as always consisting of a single<br />

stem, which seems to be correct in its normal growth. However,<br />

the finest specimen I have is a very large old plant which was<br />

apparently injured after reaching a diameter of about 3 inches,<br />

and which then put out a cluster of five new heads. I have observed<br />

numerous specimens with such form in the undisturbed<br />

wild population, but each of them shows clearly an injury to<br />

the original growing tip.<br />

Tiegel only lists the range of his plant as southwestern Texas.<br />

This takes in an immense territory, and is of little value in trying<br />

to locate the cactus. It has been so seldom reported since his<br />

description that little by way of more specific range can be<br />

given. Even Backeberg remarks that he had one specimen sent<br />

to him from Texas, but does not locate its place of origin.<br />

The only specimens I have seen have been those I collected in<br />

a small area from the Pena Blanca Mountains southeast of Marathon,<br />

Texas, to the hills just north of Alpine. Whether the range


genus Mammillaria 139<br />

is any larger than this remains to be seen. It must be rare in all<br />

of its range.<br />

Mammillaria hesteri (Wright)<br />

[Coryphantha hesteri Wright]<br />

Description Plate 37<br />

stems: Spherical to egg-shaped, to 3 inches tall and nearly as<br />

wide. They may occur singly, but usually cluster very rapidly<br />

to form irregular clumps up to more than a foot across. Each<br />

stem is covered by tubercles which are cylindrical when young,<br />

then tapering and turning upward from broad, flattened bases<br />

and up to 1/2 inch long when old.<br />

areoles: Roundish on immature stems, then on mature stems<br />

elongating to form grooves running almost all of the way to<br />

the axils, with the flowers produced at the ends of these<br />

grooves. Maturing stems show all degrees of the lengthening<br />

of the areoles. There is only a very small tuft of wool in the<br />

base of the groove at first, which usually soon disappears.<br />

spines: On mature heads there are 14 to 20 radial spines. They<br />

radiate evenly around the areole, but the upper ones are much<br />

longer and heavier than the others, the lower ones being only<br />

1/8 to 1/4 of an inch long, while the upper ones are often 1/4 to<br />

1/2 inch long. In each areole there is one spine markedly heavier<br />

than the rest and standing upright against the more slender<br />

upper radials behind it. This has been considered just a heavier<br />

radial by all who have described the cactus, but I have<br />

known people to think it a central spine and so misidentify<br />

the plant. The distinction here is very fine, as the spine does<br />

seem to arise from the interior of the areole within the circle<br />

of the radials. On young stems there are only 12 or 13 short,<br />

equal, radiating spines. On plants from the type locality all<br />

spines are very white and translucent at first, later developing<br />

gray opaqueness with tips of red or purplish-brown. However,<br />

specimens from near Sanderson, Texas, all have gray or<br />

tan bases, with the upper two-thirds of each spine red or purple-brown.<br />

All spines are more or less flattened.<br />

flowers: Mauve, 1 to 11/2 inches across and tall. The outer<br />

petals, about 8 in number, grow to about 3/16 of an inch wide,<br />

are greenish to pinkish, and are fringed with long, white<br />

hairs. The inner petals, about 22 to 27 in number, grow to<br />

about 1/2 inch long and 3/16 of an inch or slightly less in width,<br />

are bluntly pointed, and have edges smooth or slightly ragged.<br />

They are mauve, darkest at the tips. The filaments are<br />

white to rose, and the anthers are orange-yellow. The style is<br />

slightly longer than the stamens and greenish or yellowish<br />

pink. There are 4 to 6 short, stout, rough, cream-colored or<br />

white stigma lobes.<br />

fruits: Unknown.<br />

Range. A small area a few miles southeast of Alpine, Texas, and<br />

another small area a short distance west of Sanderson, Texas.<br />

Remarks. This and the next species are two of several diminutive<br />

cacti occurring in very limited areas just south and southeast<br />

of Marathon, Texas. Each of these areas is so small as to be<br />

only a few miles across, and is all included in only two or three<br />

ranches.<br />

M. hesteri was first discovered by J. P. Hester, and he wrote<br />

briefly of it in 1930. It was officially described by Wright in<br />

1932 as Coryphantha hesteri. Immediately, even though it is a<br />

small and rather insignificant cactus, it became a sought-after<br />

novelty. The large clusters which used to be available all seem<br />

to have been removed, and now one has to search well to find<br />

even a small specimen. Since it occurs in such limited areas, it<br />

would be very easy for it to be exterminated by too much gathering.<br />

I am glad to learn that one of the ranches on which it<br />

grows is now closed by the owner to all cactus digging. This<br />

should give it an opportunity to increase its sadly depleted<br />

population.<br />

Only within the past three years a new stand of this cactus<br />

was discovered a short distance west of Sanderson. The plants<br />

from this location are identical to those of the type locality except<br />

that their spines are much darker in color and the stems<br />

are more egg-shaped and thus taller.<br />

In spite of the fact that I have grown this cactus for six years<br />

and it has bloomed most of these years, my specimens have<br />

never set fruit. Neither have I been fortunate enough to find<br />

fruits on wild plants. Therefore I cannot contribute the description<br />

of the fruits of this species, which apparently has never<br />

been given.<br />

Mammillaria nellieae (Croiz.) Croiz.<br />

[Coryphantha minima Baird, Coryphantha nellieae Croiz.]<br />

Description Plate 37<br />

stems: Egg-shaped or cylindrical and very small, usually<br />

under 1 inch tall and to 3/4 of an inch in diameter, but occasionally<br />

to as much as 13/4 inches tall. The stems are usually<br />

single, but sometimes branch above the base. The tubercles<br />

covering the stem are conic and up to about 1/16 of an inch<br />

long.<br />

areoles: Each tubercle has a broad, deep, naked groove run-<br />

ning the full length of its upper side, which is the linear,<br />

monomorphic areole running from the spinous portion at the<br />

tip of the tubercle to the floral portion in the axil.<br />

spines: There are 13 to 15 definite radial spines which are 1/16<br />

to 1/8 of an inch long and which lie flat against the plant all<br />

around the areole. They vary from rather slender to fairly<br />

heavy. Besides this there are 3 other spines, much heavier and


140 cacti of the southwest<br />

to about 1/4 of an inch long which stand upright in front of<br />

and curve slightly backward to lie directly upon the upper<br />

radials. Two of these are heavy and spread upward, forming<br />

a rather distinct V. The third is always somewhat smaller,<br />

and bisects the V formed by the first two. Some authorities<br />

have called these 3 spines centrals and others have considered<br />

them only extra-heavy radials, this latter reckoning giving a<br />

maximum of 18 radials. All spines are pinkish when growing,<br />

then becoming yellowish until fading to gray. All spines are<br />

also unique in shape, being round and maintaining their full<br />

thickness from the bases to the very tips, these tips being then<br />

suddenly sharp-pointed. This gives the spines somewhat of a<br />

club-shaped appearance.<br />

flowers: Rose-purple in color, about 3/4 of an inch tall and<br />

5/8 to 1 inch in diameter. The outer petals are short and greenish<br />

with pink, fringed edges. The inner petals are about 3/8 of<br />

an inch long and about 1/8 of an inch wide, rose-purple, and<br />

bluntly pointed, with smooth edges. The filaments are greenish<br />

and the anthers pale orange. The style is short, and the<br />

stigma has 8 green lobes.<br />

fruits: Egg-shaped and very small, being usually 1/8 to 1/4 of<br />

an inch long, green in color, sometimes with a faint yellowish<br />

blush, with the dried perianth remaining upon them. The seeds<br />

are about 1/2 millimeter long, blackish, with smooth surfaces.<br />

Range. Known only from the type locality a few miles south of<br />

Marathon, in Brewster County, Texas.<br />

Remarks. This is another of the dwarf cacti found in Brewster<br />

County, Texas. It grows in crevices of limestone ledges on a few<br />

hills just south of Marathon. It is often so imbedded in the moss<br />

of these crevices that the plant itself is almost wholly covered.<br />

In such situations it is surprising to see the beautiful purplish<br />

flowers rising up above the moss from the wholly unseen cactus<br />

below. There are only a few slopes upon which the plants grow<br />

openly and exposed.<br />

These little cacti are interesting to see when grown in proper<br />

surroundings so that they are healthy, but this is not easy to<br />

arrange. They need very good drainage so that they do not rot,<br />

but also must have frequent waterings, since these tiny plants<br />

cannot store much water within themselves. They need heat and<br />

sun, but appreciate being shaded with something which breaks<br />

the direct rays of the sun just a little. I know of no collector who<br />

has succeeded in keeping specimens of this species alive more<br />

than a couple of years, yet the life-span of the plant must be<br />

much longer than that.<br />

This cactus is tiny, but the problems with its names are large.<br />

It has become almost universally known by the name, Coryphantha<br />

nellieae, but the name Coryphantha minima was given<br />

to it in its first description and has precedence. The first of these<br />

names was given to it in a description written by Croizat in the<br />

January and February, 1934, issue of Torreya. Strangely, Croizat<br />

seemed totally unaware that Ralph O. Baird had already<br />

described the cactus under the name Coryphantha minima in<br />

the American Botanist, 37 (4), 150–151, 1931. Unaccountably,<br />

all authorities since that time have omitted mention of this earlier<br />

description by Baird and used only the later name. Baird’s<br />

description is in every way more complete and official than<br />

Croizat’s, except that it lacks the Latin description, but this cannot<br />

disqualify it, as 1931 antedated the rule requiring Latin de-<br />

scriptions.<br />

Most recently, Backeberg transfers the species to Escobaria,<br />

calling it Escobaria nellieae (Croiz.) Backbg. He says the fruit is<br />

unknown. This error would have been avoided if he had referred<br />

to Baird’s description where the fruit was accurately described<br />

as being green, which is a Coryphantha characteristic.<br />

Croizat himself had already made the standard argument over<br />

whether the cactus is a Coryphantha or an Escobaria unnecessary<br />

by writing, “In the course of a recent study of the Cactaceae<br />

I have come to the conclusion that the following transfer<br />

may be desirable: Mammillaria nellieae (Croiz.) comb. nov.—<br />

Cory. nellieae Croiz. in Torreya 34: 15. 1934.” His conclusion<br />

here is in line with our approach to this group, but it would<br />

seem that under this combination the name of this cactus should<br />

by the rules be rather Mammillaria minima. This would be true<br />

except for one fact, and because of this Baird must lose even in<br />

this combination. There had already been a previous Mammillaria<br />

minima put forth by Reichenbach in Terscheck, Supp. Cact.<br />

Verz. 1, for a Mexican cactus. Reichenbach’s cactus is now listed<br />

as a variety of M. elongata DC., but since the name has been<br />

used once, it cannot be used again for another plant, so we are<br />

left after all with Croizat’s M. nellieae as the name for our diminutive<br />

Texas plant.<br />

The species is easily recognized by the remarkable appearance<br />

of the spines, which lie tight against the surface of the plant and<br />

preserve their thickness right up to the points, so that they appear<br />

very thick for their length and are more like tiny clubs<br />

than ordinary spines.<br />

Mammillaria roberti (Berger)<br />

[Escobaria runyonii B. & R., Coryphantha roberti Berger]<br />

“Runyon’s Escobaria,” “Junior Tom Thumb Cactus”<br />

Description Plate 38<br />

stems: Spherical at first, becoming oblong and to about 3<br />

inches tall by 11/4 inches thick. Branching very fast and irregularly<br />

to form large, low clumps of dozens of stems. The tubercles<br />

covering the stems taper from cylindrical bases and<br />

are from 3/16 to 1/4 of an inch long.<br />

areoles: Round and at the tips of the tubercles on immature<br />

stems. On mature stems they develop from oval to almost<br />

linear by being prolonged from the spinous portion at the tip<br />

of the tubercle into a very narrow, very woolly groove running<br />

from halfway to all the way down the upper side of the


genus Mammillaria 141<br />

tubercle. Flowers and new branches are produced from the<br />

end of this groove, at or near the axil of the tubercle.<br />

spines: There are 20 to 30 slender radial spines which are 3/16<br />

to sometimes almost 1/4 of an inch long, white in color, usually<br />

barely tipped with brown. There are 5 to at least 10 central<br />

spines filling the center of each areole and spreading in<br />

all directions. These are 3/8 to 5/8 of an inch long, the upper<br />

ones being much the longer, and are slender. These centrals<br />

have white bases, but the upper three-fourths of each one is<br />

dark brown, red-brown, or black.<br />

flowers: Very inconspicuous in color, 3/4 of an inch in diameter<br />

and 5/8 of an inch tall, but opening widely. All petals<br />

are slender and pointed, buff or tan, with midlines of reddishbrown.<br />

The outer ones are fringed and the inner ones may be<br />

either with or without fringes. The filaments are pink and the<br />

anthers yellow. The style is about 1/8 of an inch longer than<br />

the stamens and reddish-brown. The stigma has 5 or 6 short,<br />

thick lobes which are green, sometimes yellow-green or even<br />

brownish-green.<br />

fruits: Spherical to egg-shaped, 1/4 to 3/8 of an inch long,<br />

scarlet in color.<br />

Range. The Rio Grande Valley and its overlooking hills from<br />

near McAllen, Texas, to the mouth of the Pecos River. Also<br />

found along the west side of the lower Devil’s River.<br />

Remarks. This is undoubtedly one of the most perfectly camouflaged<br />

of the cacti. Even large clumps a foot or more in diameter<br />

are only a couple of inches high, and the dark-tipped<br />

spines blend perfectly with the rocky soil around them and the<br />

grasses usually growing up through them and partly covering<br />

them. Even the flower is a blending color and gives no aid in<br />

locating the plant. Only the scarlet fruit is allowed to advertise,<br />

no doubt so that birds and rodents will come and eat them in<br />

order to scatter the seeds.<br />

The cactus was discovered by Robert Runyon, a fine student<br />

of the cacti of the lower Rio Grande Valley, and it was first<br />

described by Britton and Rose as Escobaria runyoni. It grows<br />

upon the rocky hills overlooking the Rio Grande, being most<br />

common near Rio Grande City and around the lower part of<br />

what is now the Falcon reservoir. It is still fairly common there,<br />

partly because it takes a real search to find it, and partly because<br />

it is too insignificant in size and appearance to appeal to<br />

those who take home the more beautiful cacti.<br />

While searching along the lower parts of the Devil’s River in<br />

West Texas for another cactus, we were very surprised to find,<br />

growing on the west side of that river, numerous specimens of<br />

a cactus identical to this one in all respects. It is clearly the same<br />

cactus, and represents a large increase of the known range of<br />

this cactus. But this left a large gap between this Devil’s River<br />

location and the old locations known on the lower Rio Grande,<br />

where there was no record of its having been seen. This prompted<br />

careful searching, and we have now located the cactus grow-<br />

ing all the way past Laredo to Eagle Pass and near Del Rio, so<br />

there is in reality one continuous range, much longer than at<br />

first thought.<br />

There have been problems over the name for this cactus, all<br />

because of the confusion which has existed over genera. In an<br />

article in 1929, Berger sought to transfer the whole genus Escobaria<br />

to Coryphantha. In so doing he met a problem with Escobaria<br />

runyoni, as there already existed a Coryphantha runyonii.<br />

He attempted to solve the difficulty by re-naming this cactus<br />

Coryphantha roberti, using the first name of Mr. Runyon for it<br />

instead of his surname.<br />

In 1931 Fosberg attempted to do the same thing again and<br />

met with the same problem. He coined another new name, Coryphantha<br />

piercei for this plant.<br />

This system of combination has not caught on, however, perhaps<br />

because it did not go far enough. We feel that a combination<br />

cannot be avoided, and Britton and Rose’s name cannot be<br />

preserved, no matter what form this combination takes. Under<br />

our proposal to return both of these microgenera to the original<br />

genus, Mammillaria, this species becomes Mammillaria roberti,<br />

since this was the first substitute name suggested. This seems<br />

especially fitting since it preserves the reference to Dr. Runyon,<br />

as was intended.<br />

Mammillaria sneedii (B. & R.)<br />

[Escobaria sneedii B. & R.]<br />

Description Plate 38<br />

stems: To only about 2 inches long and 3/4 of an inch thick.<br />

These small cylindrical stems branch and cluster very greatly<br />

to form masses of as many as 100 heads on old specimens. The<br />

individual stems are composed of many tiny, cylindrical,<br />

green tubercles 1/16 to 3/16 of an inch long.<br />

areoles: Round or nearly so on immature stems, lengthening<br />

somewhat on mature stems to form short grooves running<br />

from the spinous portion of the areoles at the tips of the<br />

tubercles to at most one-third of the way down toward the<br />

axils. Flowering or branching occurs at the ends of these short<br />

grooves. There is a little white wool in the groove.<br />

spines: The tubercles are almost entirely obscured by the many<br />

white spines. There are 24 to 45 radiating outer spines, which<br />

are about 1/8 of an inch long and very slender, but rigid. There<br />

are 13 to 17 central spines, which are slightly heavier than the<br />

radials and 1/8 to 5/16 of an inch long. These centrals are all<br />

spreading and lie against the radials, except for the middle<br />

one out of the center of the areole, which often stands outward<br />

and downward at an angle. All spines are white when<br />

mature, but pinkish when growing, giving the tip of a growing<br />

stem a pinkish color.<br />

flowers: Small—about 1/2 inch tall and broad—not opening


142 cacti of the southwest<br />

widely, and pink to pale rose in color. The outer petals are<br />

narrow, with fringed edges. The midlines are rose and they are<br />

edged in very pale pink. The inner petals are the same color,<br />

but paler. Their edges are fringed at least half of the way to<br />

the pointed tips. There are also sometimes notches in the otherwise<br />

entire margins of these petal ends. The filaments are pink,<br />

the anthers bright orange. The style is longer than the stamens.<br />

The stigma has 3 or 4 slender, white lobes.<br />

fruits: About 1/4 of an inch thick, almost spherical, but a<br />

little longer than they are thick. They become a deep pink<br />

when ripe. The seeds are brown, pitted on the surface, and<br />

less than 1/16 of an inch (about 1 millimeter) long.<br />

Range. Known only from the Franklin Mountains between El<br />

Paso, Texas, and Las Cruces, New Mexico.<br />

Remarks. The tiny, white stems of this little cactus are insignificant<br />

individually, but in an old plant made up of dozens of<br />

them, the whole may be a clump up to a foot or more in diameter,<br />

which is very impressive. Unfortunately, very few persons<br />

have ever seen such mature plants, and it is unlikely that<br />

anyone will see such a sight in the future unless he grows the<br />

plant himself.<br />

The cactus was brought down out of the higher elevations of<br />

the Franklin Mountains and shown to Britton and Rose in 1921.<br />

They first described it in their large work, saying it was known<br />

from only a single station in those mountains, and that only five<br />

plants had been found. Immediately many went up looking for<br />

it, and no doubt dozens of specimens were brought down, dispersed<br />

to gardens, and never heard of again. Some photographs<br />

of large clusters were published in various journals, but it is<br />

doubted that any of these large plants still survive, and in recent<br />

years even small specimens growing wild have been almost impossible<br />

to find. The species is no doubt another of those well on<br />

the way to extinction, and the future for it would look very dark,<br />

except for the work of Mr. Clark Champie of Anthony, New<br />

Mexico-Texas. He is the first dealer propagating these Texas and<br />

New Mexico species from seed instead of relying upon stripping<br />

nature’s gardens for his stock. Located as he is at the foot of the<br />

Franklins, he has secured mature plants and has many seedlings<br />

of M. sneedii, so collectors can continue to have the plant from<br />

his nursery.<br />

Mammillaria leei (Böd.)<br />

[Escobaria leei Böd.]<br />

Description Plate 39<br />

roots: Fibrous on young individuals and mostly fibrous when<br />

older, but usually with a single definite taproot developed<br />

from the original center of an old clump. This taproot is about<br />

3/16 to 1/4 of an inch thick and usually runs for some inches<br />

before reducing.<br />

stems: Almost spherical at first, later becoming mostly club-<br />

shaped, but sometimes cylindrical, to a maximum of about 3<br />

inches long and about 11/8 inches thick, but ordinarily only<br />

about 1 to 11/2 inches tall and 3/4 of an inch thick. These small<br />

stems branch and proliferate very rapidly, forming irregular<br />

clumps of up to several hundred very tightly packed stems,<br />

such clumps a number of inches across, but only about 2 inches<br />

high. Each stem is covered with cylindrical tubercles up to<br />

about 3/16 of an inch long, although often smaller.<br />

areoles: Consisting on many stems of only circular or elliptic<br />

areoles on the ends of the tubercles. Whether this represents<br />

nonflowering, immature stems was not observed, but this condition<br />

sometimes persists on some of the larger stems. On other<br />

stems, more or less regardless of stem size, the areoles are<br />

elongated into grooves extending from the spinous portions at<br />

the ends of the tubercles to around half of the distance to the<br />

tubercle bases. The flowers and branches observed came from<br />

the ends of these grooves. The whole areole is at first very<br />

woolly, and the wool remains in the grooves. The center of<br />

the spinous part of the areole is often, but not always, very<br />

convex and bulging outward. In such cases the spines are<br />

mostly pushed back in position, where they radiate around<br />

and form a sort of frame for this convex areole center, this<br />

giving the stems in such cases a unique, knobby appearance.<br />

spines: There are very many, very tiny, white radial spines.<br />

Counting them is difficult, but there seem to be from 40 to at<br />

least 90 per areole. These are appressed against the surface of<br />

the plant and often recurve between the tubercles. There are<br />

also 6 or 7 stouter central spines, white or Sometimes white<br />

with pale brownish tips. These spines are irregular in length,<br />

usually about 1/8 of an inch long, but sometimes a few may be<br />

longer. One of these centrals stands perpendicular to the plant<br />

body in the center of the areole, but it may be a very short<br />

one. The rest radiate in front of the radials.<br />

flowers: Not opening widely. They are about 3/4 of an inch<br />

long and 1/2 inch wide. In color they are deep pink suffused<br />

with brownish. The outer petals are brown or greenish-brown<br />

with light edges fringed by white cilia. The inner petals are<br />

deep pink in the midlines, the edges lighter and entire. The<br />

stigma lobes are 4 to 6 in number, short, and white.<br />

fruits: Oblong or somewhat club-shaped, about 1/2 inch long<br />

by 1/8 to 1/4 of an inch thick. They remain greenish tinged<br />

with brownish or faintly pinkish when ripe. The seeds are<br />

more or less pear-shaped, dark brown, and less than 1/16 of an<br />

inch (slightly over 1 millimeter) long.<br />

Range. Known only from several canyons about 30 miles southwest<br />

of Carlsbad, New Mexico.<br />

Remarks. This is a very peculiar little cactus. It was apparently<br />

first collected in 1924 by W. T. Lee, whose name it bears. Since<br />

that time specimens of it have lain in the U. S. National Herbarium,<br />

two peculiar, unnamed little specimens to tantalize any-


genus Mammillaria 143<br />

one working through that collection. One of those specimens is<br />

a whole cluster with about a hundred stems, most of them hard-<br />

ly thicker than a lead pencil in their dried condition.<br />

Several have dealt with these herbarium specimens to some<br />

extent, but only recently does it appear that the cactus has been<br />

re-collected. Dr. Rose, in 1925, labeled one of them Neomammillaria<br />

leei, which would have been its earliest name, but he<br />

apparently never wrote up the description of this new species.<br />

It therefore fell to Bödecker to make the first description of the<br />

cactus in the literature. He did this in 1933, calling it Escobaria<br />

leei. His description was very incomplete, based apparently<br />

upon only these two dead specimens.<br />

Within the past few years several of us have tracked down<br />

the plant and collected it in the type locality. In 1966 Castetter<br />

and Pierce published an article on the species, giving the first at<br />

all complete description, and showing that the plant grows quite<br />

a bit larger in stem than Bödecker thought. Castetter and Pierce,<br />

however, in spite of the fact that the fruits remain essentially<br />

green, leave the cactus as an Escobaria.<br />

We find this form especially interesting and feel that, in the<br />

light of the genus problem, it is worthy of much more study. On<br />

many stems it remains with unelongated areoles. What is an immature<br />

condition in all cacti seems to persist sometimes throughout<br />

the life of the stem in this cactus. We have never been able<br />

to observe whether such ungrooved stems flower, only having<br />

seen flowers on those with elongated, obviously monomorphic<br />

areoles. It seems difficult to believe that so many stems would<br />

remain immature and never bloom. If they do bloom, it might<br />

be significant to know where the flowers are produced, since the<br />

groove is absent.<br />

The cactus is close to M. sneedii, and the ranges of the two are<br />

not far apart. I was at first tempted to think of them as two<br />

manifestations of the same cactus, but knowledge of their details<br />

seems to put this out of the question.<br />

M. leei comes from a very limited area, so far as is known<br />

consisting of only a few canyons. The population cannot be<br />

large. It has been fortunate, no doubt, in being practically overlooked<br />

until very recently. Let us hope that collectors do not<br />

now descend upon it and eliminate it, as they seem to have done<br />

to M. sneedii.<br />

Mammillaria bella (B. & R.)<br />

[Escobaria bella B. & R.]<br />

Description (adapted from Britton and Rose)<br />

stems: Clustering, the heads about 23/8 to 31/4 inches long and<br />

cylindrical. The tubercles are 5/8 to almost 7/8 of an inch long,<br />

marked by a hairy groove, near the center of which there is<br />

a brownish gland.<br />

areoles: Not known.<br />

spines: Radial spines said to be several, whitish, and less than<br />

1/2 inch long. The central spines are 3 to 5 in number, brown,<br />

and the longest to at least 7/8 of an inch long.<br />

flowers: To 3/4 of an inch across, petals pinkish with pale<br />

margins, narrow and pointed, the outer ones fringed. The filaments<br />

are reddish, the stigma green.<br />

fruits: Not known.<br />

Range. Collected “on hills of Devil’s River, Texas.”<br />

Remarks. This cactus was collected by Rose and Fitch before<br />

1914, and described by Britton and Rose in 1922. There is no<br />

record of its having been collected again. Numerous searches<br />

along the Devil’s River have turned up no plant with the unique<br />

characteristic which Britton and Rose emphasize for this one—<br />

the possession of the gland in the groove of the tubercles—in<br />

conjunction with so small a stem size, such large tubercles, and<br />

a smallish, pinkish flower. The nearest I have come to it were<br />

some specimens found in the area, which roughly approach<br />

these characters but which turned out to be seedlings of Mammillaria<br />

macromeris.<br />

This being the situation, the plant can only be listed upon the<br />

authority of Britton and Rose as a cactus once collected in Texas.<br />

There is not even any way to restudy the form. But it seems that<br />

their authority requires us to list the cactus, and I repeat their<br />

description here in case—and in hope—someone may yet suc-<br />

ceed in finding it where all of us have failed.<br />

Mammillaria pottsii Scheer<br />

Description Plate 39<br />

stems: Cylindrical, to about 11/4 inches in diameter and to at<br />

least 8 inches tall, although usually only 4 to 5 inches high. It<br />

was reported by Coulter to grow to 12 or 14 inches, but this<br />

is doubtful. These stems may remain single, but sometimes<br />

branch above the ground to form small, irregular clusters of<br />

3 or 4 heads. The tubercles covering the stems are conical to<br />

egg-shaped, 1/8 to 1/4 of an inch long, rather closely crowded,<br />

and somewhat overlapping.<br />

areoles: Dimorphic. The spinous portion on the tip of the<br />

tubercle is small, round, and especially filled with spines because<br />

of the enlarged bases of the centrals. It at first contains<br />

much rather long, white wool, but is later bare. The flower<br />

develops in or near the axil of the tubercle, together with long<br />

wool and sometimes a bristle or two, which remain as an axil-<br />

lary tuft.<br />

spines: There are 30 to 40 radial spines which are white, very<br />

slender, but straight and rigid. These are about /lo of an inch<br />

long and remarkable for being entirely equal in size as they<br />

radiate around the areole. The number of central spines may<br />

vary from 6 to 12 from plant to plant, but the number seems<br />

to be identical in all areoles of any one plant. These centrals<br />

spread evenly in all directions from greatly enlarged, bulbous


144 cacti of the southwest<br />

bases. The lower and lateral ones are all 1/4 to 3/8 of an inch<br />

long and straight, but the upper ones are 1/2 to 5/8 of an inch<br />

long and curve upward. The lower ones are gray with their<br />

very tips brownish. The upper ones are gray with as much as<br />

the outer half of each dark brown, purple, or a peculiar bluish<br />

color; this color, as they curve upward, gives the tip of the<br />

plant over which they converge a purplish color.<br />

flowers: Small, 3/8 to 1/2 inch long and wide, bell-shaped, the<br />

petals not opening widely, but then recurving toward the tips.<br />

They are deep red, maroon, or rust red in color. They develop<br />

around the upper sides of the stem instead of at the summit of<br />

it. The outer petals are very broad, but with pointed tips, the<br />

margins usually ragged or irregularly toothed, but not fringed.<br />

These outer segments are red or maroon with pale, cream-<br />

colored edges. The inner petals have red to rust-red midstripes<br />

with rose to pinkish, smooth margins, and pointed tips. The<br />

stamens are cream-colored. The style is reddish, and there are<br />

4 or 5 cream-colored or yellow stigma lobes.<br />

fruits: Club-shaped, up to 5/8 of an inch long, pale or light<br />

red. The seeds are blackish, pitted, almost oval, and less than<br />

1/16 of an inch (not quite 1 millimeter) in length.<br />

Range. The U. S. range is limited to the mountains deep in the<br />

Big Bend region around Terlingua, Texas, but the plant is widely<br />

distributed in Mexico.<br />

Remarks. This is a distinctive cactus unlike anything else found<br />

in the United States. Its appearance readily distinguishes it from<br />

all the other Mammillarias we have in this country, and when it<br />

blooms we have no other cactus with a flower even remotely<br />

like this one’s small, bell-shaped, red blossom. It seems to be our<br />

only representative of the large group of Mammillarias found<br />

in Mexico, with small, bell-shaped flowers. In the United States,<br />

therefore, it should be one of the most easily recognized of all;<br />

yet it has a history of serious confusion with other forms.<br />

It seems that this cactus was first described by Scheer in the<br />

Salm-Dyck publication of 1850. Perhaps the uncertainty started<br />

there, as that description mentions wrinkles and a very slight<br />

groove on the tubercle. This, once the distinction between grooved<br />

and grooveless had become paramount, would of course make<br />

one think it something else instead of a Mammillaria in the narrow<br />

usage of that term. Poselger, in 1853, collected a plant with<br />

a groove, which Britton and Rose think must have been Mammillaria<br />

(Escobaria) tuberculosa, but which Poselger himself<br />

thought was our cactus, although he called it Echinocactus pottsianus.<br />

But when, in the same year, he apparently collected our<br />

plant which does not actually have a groove, he thought he had<br />

a new species and renamed it Mammillaria leona.<br />

From that time the confusion spread. Engelmann seemed to<br />

have had a different plant in mind when describing M. pottsii,<br />

since he said that it has large flowers and grows on the Rio<br />

Grande below Laredo. Coulter and others more or less followed<br />

him in this, their descriptions, therefore, being confusing. All<br />

the way to the present the confusion has remained. Berger, followed<br />

by Borg, finally called some Mexican plant with a “narrow<br />

groove” Coryphantha pottsii, and so the plant has been<br />

put into three different genera. Backeberg has only extended the<br />

confusion when he says that Mammillaria pottsii was an Escobaria,<br />

and so uses for our plant the name Mammillaria leona<br />

Poselgr. once again.<br />

Britton and Rose seemed to have been among the first to realize<br />

the source of the confusion. They got back to our cactus<br />

again in their description, but they were very limited, having<br />

only one Texas record of this cactus to work with.<br />

Craig states most clearly the cause of the difficulty in saying<br />

that the tubercles of this cactus become wrinkled and shrunken<br />

when dehydrated—as do most—and that since the original description<br />

mentions both a slight groove and many wrinkles,<br />

Scheer must have had before him a shrunken, atypical plant,<br />

and thus the groove he mentioned to start the whole confusion<br />

was not the natural one of an elongated areole. Through this<br />

overemphasis on a groove, then, the original name became applied<br />

to other grooved cacti of the Coryphantha and Escobaria<br />

microgenera, while the Texas plant, although seldom seen at all,<br />

was known under Poselger’s name of M. leona. This seems to<br />

have been the situation until Britton and Rose reapplied the<br />

original name to the plant again. Since both Craig and Marsden,<br />

two of our greatest authorities on the Mammillarias, see no difficulty<br />

in using Scheer’s name, I follow them here.<br />

Be that as it may, the plant which grows in Texas and which<br />

we will call by the name of M. pottsii is a beautiful cactus of one<br />

to several slender, whitish columns over which is thrown a peculiar<br />

bluish cast by the unusual coloring of the upper centrals.<br />

It most nearly resembles M. tuberculosa, but is much more beautiful<br />

because of its more regular and tidy spines. The greatly<br />

enlarged bases of its central spines are a quick clue for recognizing<br />

this plant. When it blooms its flowers are fine little bellshaped,<br />

deep-red blossoms produced in a ring around the upper<br />

part of the stems. They are small flowers, but they are charming,<br />

showing that cactus flowers can be small and demure as well as<br />

large and flamboyant.<br />

Because of the confusion about M. pottsii which has reigned<br />

for so long, many of the older accounts of its range cannot be<br />

trusted. It does not, for instance, grow in Texas below Laredo,<br />

as Engelmann and Coulter thought. Britton and Rose also make<br />

the peculiar statement that it comes from the “highlands of the<br />

Rio Grande.” No doubt the mountains of the Big Bend, where it<br />

does grow and through which that river flows, are highlands,<br />

but it does not grow in the mountains of northern New Mexico,<br />

the “highlands” in which the Rio Grande actually arises. To be<br />

specific, all specimens of which I know came from the moun-<br />

tains deep in the Big Bend around Terlingua, Texas.<br />

The cactus grows well in limestone soil which is well-drained,<br />

and is often used in dish or windowsill gardens. It is, however,<br />

a desert species, and needs much sun and cannot tolerate much<br />

dampness.


genus Mammillaria 145<br />

Mammillaria lasiacantha Eng.<br />

Description Plates 39, 40<br />

stems: Usually single, but sometimes forming Small clusters of<br />

2 or 3 heads. Each stem is spherical, conical, or egg-shaped<br />

and small, the maximum appearing to be about 2 inches in<br />

diameter and only slightly taller. The plant is covered by many<br />

small, cylindrical tubercles to about 3/16 of an inch long and<br />

1/8 of an inch in diameter. The axils between these are bare.<br />

areoles: The spinous portions of the dimorphic areoles are on<br />

the tips of the tubercles and are small and somewhat eggshaped.<br />

The floral portions of the areoles remain in the axils<br />

of the tubercles, the flowers thus being produced in the axils.<br />

spines: Each areole produces 40 to 60 or more white radial<br />

spines which lie flat, interlocking with those of the neighboring<br />

areoles and covering the surface of the plant. These spines<br />

are so numerous that they cannot all lie in one plane around<br />

the areole, and so they are produced in several series forming<br />

3 or 4 concentric rings in the areole, one radiating layer of<br />

them lying upon the next. They are from about 1/16 to 3/16 of<br />

an inch long. They may have rough, even pubescent surfaces<br />

caused by varying numbers of almost microscopic white hairs<br />

upon them, or they may be smooth of surface.<br />

flowers: 1/2 to 3/4 of an inch long and the same diameter,<br />

opening widely. The color is whitish with each petal marked<br />

by a conspicuous tan, red-brown, or purplish-red midstripe.<br />

The outer petals are slender, with the ends either rounded or<br />

bluntly pointed, their margins usually somewhat irregular.<br />

The inner petals are oblong, rather wide, being 1/16 to 1/8 of an<br />

inch Wide over most of their lengths, and the tips are smooth<br />

and rounded or else ragged. The filaments are long and yellowish,<br />

the anthers yellow. The style is greenish and slightly<br />

longer than the stamens. The stigma has 4 or 5 small, greenish<br />

or yellowish lobes.<br />

fruits: Bright scarlet, club-shaped, and 3/8 to 5/8 of an inch<br />

long. The remains of the flower persist upon them, and the<br />

surfaces of the fruits are naked. The seeds are oval in shape,<br />

black with pitted surfaces, and less than 1/16 of an inch (about<br />

1 millimeter) in length.<br />

Range. Entering the U. S. from Mexico along a stretch of the<br />

Rio Grande from south of Sanderson, Texas, west to El Paso,<br />

and growing from Sanderson northwest past Fort Stockton<br />

through the Guadalupe Mountains west of Carlsbad, New Mexico,<br />

and from the Rio Grande at El Paso into the Sacramento<br />

Mountains near Almagordo, New Mexico, which seems to be the<br />

northern limit of the range.<br />

Remarks. This is another of our tiniest cacti. It usually grows as<br />

a little ball up to an inch or two in diameter, and is perfectly<br />

white due to the huge number of white spines entirely covering<br />

its surface. Very rarely it clusters, but to only 2 or 3 heads. It is<br />

found growing on the tops of hills where the soil is thin, often<br />

wedged in between limestone rocks, and it will not prosper un-<br />

less there is much limestone in its soil. It is usually more or less<br />

covered by surrounding grasses, but does not seem to require<br />

shade, as I have collected plants which were totally unprotected<br />

from the sun and have grown them this way for several years.<br />

No doubt the huge number of its spines provide its flesh its own<br />

partial shade.<br />

It is a real indication of the thoroughness of the early explorers<br />

that they even noticed this tiny cactus, and remarkable observation<br />

on Engelmann’s part that he separated this species into two<br />

other very similar varieties. I have met few cactophiles or even<br />

dealers who have taken the trouble to do this, since it involves<br />

using a magnifying glass to distinguish them.<br />

As a species, M. lasiacantha has had a comparatively uneventful<br />

taxonomic history. Only one dissenting voice seems to have<br />

been raised concerning its position as a Mammillaria. In 1951<br />

Buxbaum, because of seed characters and the fact that the axillary,<br />

floral part of the areole develops first instead of the spinous<br />

part—this the reverse of many others in the genus—separated<br />

this cactus and some of its relatives into a new genus which<br />

he called Ebnerella, but he has not so far been followed in this.<br />

There has perhaps been enough confusion over the varieties of<br />

this cactus to make up for the unanimity concerning the species.<br />

Much has been written about M. lasiacantha which is confusing<br />

because different authors using this name may be referring to<br />

different forms. Britton and Rose, for instance, mean by this<br />

name the form which should more accurately be called variety<br />

lasiacantha of the species. On the other hand, some authors have<br />

clearly seen only the variety denudata and write of the species<br />

from knowledge of only this variety. Ranges given are particu-<br />

larly confused because of this sort of thing.<br />

The species is another one of that large variety of tiny cacti<br />

which we find growing in west Texas and southern New Mexico,<br />

in the most exposed and unlikely places for such small, delicate-<br />

appearing plants. It is these inconspicuous forms for which one<br />

has to search carefully which give this area such a surprising<br />

array of cactus species.<br />

Mammillaria lasiacantha var. lasiacantha (Eng.)<br />

Description Plate 39<br />

stems: As the species, except to only about 1 inch in maximum<br />

diameter and height.<br />

areoles: As the species.<br />

spines: As the species, except that the average number is in<br />

the upper number range for the species, and that the surfaces<br />

of the spines are always roughened by at least some to very<br />

many almost microscopic cilia or trichomes. Many specimens<br />

have the spines wholly pubescent.<br />

flowers: As the species, except about 3/4 of an inch in length


146 cacti of the southwest<br />

and width, the main petals about 1/8 of an inch wide, and the<br />

stigmas more yellowish than greenish.<br />

fruits: As the species.<br />

Range. From near Fort Stockton, Texas, and the Davis Mountains<br />

north in the Guadalupe Mountains west of Carlsbad, New<br />

Mexico, to between Carlsbad and Alamogordo, New Mexico.<br />

Remarks. This form appears to have been the type for the species.<br />

Then later, when he described the two varieties within the<br />

species, Engelmann called this form M. lasiacantha var. minor.<br />

His name for the variety would be most appropriate, but the<br />

rules of taxonomy, if this is the type form, require it to be called<br />

instead variety lasiacantha.<br />

The spines of variety lasiacantha have on them a growth of<br />

almost microscopic “hairs” which give them a fuzzy appearance<br />

under a lens. Individual specimens vary in the amount of this<br />

hair, but it is always there to some extent, and in some plants it<br />

is so thick as to be a velvety covering on the spines. Specimens<br />

of the other variety, M. lasiacantha var. denudata, lack this hair<br />

entirely or have only a few scraggly cilia, and their spines are,<br />

therefore, smooth and shiny.<br />

If this matter of pubescent versus smooth spines were the only<br />

difference between these two forms, it would seem that the various<br />

students who have combined them would be justified, but<br />

there are also other differences. As indicated by Engelmann’s<br />

name for it, variety minor, the rough-spined form is smaller in<br />

maximum size. Its maximum observed size is about 1 inch, while<br />

specimens of the smooth-spined form 2 inches in diameter are<br />

rather easy to find. The smallness of the over-all size probably<br />

keeps us from evaluating this difference properly. It is a difference<br />

in maximum size of double—which would be very noticeable<br />

in two giant barrel cacti. At the same time, the flowers of<br />

this smaller form are larger, with wider petals, than those of the<br />

larger variety.<br />

Admittedly these differences may intergrade. There are very<br />

clearly a few cilia on some specimens of variety denudata. This<br />

would certainly preclude calling them separate species, as did<br />

Britton and Rose, but this is not an obstacle in varietal relationships.<br />

When I note that I have found no specimen of the small,<br />

truly pubescent, large-flowered variety lasiacantha from south<br />

of the Davis Mountains, and that in hundreds of specimens from<br />

the bins of Big Bend dealers collected in the lower Big Bend<br />

which I have examined with a magnifying glass, and even among<br />

those doubtful specimens from that area which I have grown<br />

and flowered not a one has been other than variety denudata,<br />

I feel that we have here two legitimate varieties which should<br />

be recognized.<br />

At present, however, dealers and collectors alike, knowing<br />

only that there are supposed to be two forms, seem to be separating<br />

their plants by some sort of magic into two supposed<br />

categories, and the lots I see under one name or the other usually<br />

have little meaning. Actually, I know of no way to separate the<br />

two varieties without using a magnifier, and even beyond that,<br />

in some borderline cases, growing the specimens to flowering to<br />

compare the flowers.<br />

The failure to do this makes for much confusion. In general,<br />

most people have seen the next variety and very few have actually<br />

had variety lasiacantha. Huge numbers of variety denudata<br />

have been and still are being sold out of the lower Big Bend,<br />

where they are abundant, while few dealers have worked out<br />

of the more northerly range of variety lasiacantha, where that<br />

variety is relatively rare anyway. The study of these two forms<br />

by Britton and Rose suffered from this same situation. Although<br />

they listed the plants as two separate species, they admitted that<br />

they had not themselves seen the more rare M. lasiacantha.<br />

Dr. Boke, in his very valuable anatomical and developmental<br />

study of M. lasiacantha, did not distinguish between the two<br />

varieties at all. His specimens were collected in several locations,<br />

all but one of which were in the lower Big Bend, where all he<br />

would presumably have collected were variety denudata, so the<br />

bulk of his work must have been done with this form. But some<br />

of his plants came from a ridge 9 miles east of Alpine, Texas,<br />

where he could conceivably have picked up a few specimens of<br />

variety lasiacantha. This possibility becomes interesting because<br />

he mentions that he found mucilaginous areas in all but 2 or 3<br />

of the many specimens which he examined. He could suggest no<br />

reason why 2 or 3 of his specimens lacked this mucilaginous<br />

system. Could it be that these 2 or 3 without it were his only<br />

specimens of variety lasiacantha, and that this is another difference<br />

between these varieties? Restudy would be necessary to<br />

determine whether this is true.<br />

Mammillaria lasiacantha var. denudata Eng.<br />

Description Plates 39, 40<br />

stems: As the species, except with the maximum size at least<br />

2 inches tall and wide.<br />

areoles: As the species.<br />

SPINES: As the species, except smooth and shiny, with usually<br />

no cilia or trichomes upon them, and at most on some specimens<br />

a few scattered “hairs” upon the spines.<br />

flowers: As the species, except to about 1/2 inch in diameter,<br />

petals almost linear and only about 1/16 of an inch wide, and<br />

stigma with more greenish than yellowish lobes.<br />

fruits: As the species.<br />

Range. From Mexico into the mountains of the Big Bend in<br />

Texas, along the Rio Grande from near Sanderson to Canutillo,<br />

Texas, just beyond El Paso. It also occurs north past Alpine and<br />

Marfa, Texas, into the southern part of the Davis Mountains.<br />

Remarks. This is the more common southern form of the species.<br />

It is very common in parts of the Big Bend region, and thousands<br />

of the plants come out of that area into the trade every


genus Mammillaria 147<br />

year. On the west it has been collected at Sierra Blanca and at<br />

Canutillo, Texas, but I find no record of it in New Mexico. On<br />

the east, Orcutt once reported it from Langtry, Texas, but it<br />

does not seem to have been found there since.<br />

Although these two varieties are very close, this is a larger<br />

plant when grown, and has smaller flowers. The two are cer-<br />

tainly not separate species, as some have maintained.<br />

Mammillaria multiceps SD<br />

“Hair-Covered Cactus,” “Grape Cactus”<br />

Description Plate 40<br />

stems: Spherical to egg-shaped or sometimes short-columnar.<br />

Matured, blooming heads measure from 1/2 to almost 2 inches<br />

in diameter and 2 inches tall, producing off-sets extremely<br />

rapidly so that a typical plant consists of a dozen to more<br />

than a score of different-sized heads forming a large, low,<br />

matlike clump. The tubercles are conical or cylindrical, spreading,<br />

and up to 3/8 of an inch long.<br />

areoles: Dimorphic. The spinous portion on the end of the<br />

tubercle is round, with white wool at first, later naked. The<br />

floral or vegetative portion in the axil of the tubercle produces<br />

with the flower some wool and usually several long,<br />

twisted, white, hairlike bristles which persist.<br />

spines: Radial spines number 30 to more than 60, and are<br />

said to have sometimes reached 80. These are produced in<br />

several series. The outer ones are white, hairlike, very fine,<br />

entirely flexible, and usually curved and twisted. When<br />

straightened they are 1/4 to 5/8 of an inch long. The inner<br />

series become progressively shorter and heavier until the<br />

innermost are straight, rigid bristles spreading in all directions,<br />

white or light yellow in color, and usually somewhat<br />

pubescent. In the center of the areole are 4 to 9 comparatively<br />

heavy central spines with bulbous bases. They are straight,<br />

rigid, 1/4 to 3/8 of an inch long, and spreading in all directions.<br />

In color they are white to yellow or brown at their bases, the<br />

upper parts and tips of them varying on different plants from<br />

whitish or light honey-yellow to dark red-brown or almost<br />

black. These are more or less pubescent under a lens. There is<br />

also an outer series of centrals which are lighter in color and<br />

more slender than the inner ones, and sometimes approach so<br />

closely to the heaviest inner radials that on some plants the<br />

point of transition from one to the other is not clear. Count-<br />

ing these, there may be up to 12 centrals.<br />

flowers: About 3/4 of an inch long and the same diameter,<br />

brownish-yellow or almost tan in color, with pinkish to<br />

mauve-rose streaks. The petals have cream or tan edges shading<br />

into pinkish or dull rose midlines. They are usually unfringed,<br />

but the outer ones may occasionally have a few cilia<br />

on their edges. The filaments are yellowish or white, the an-<br />

thers yellow. The style is cream-colored and short. There are<br />

3 to 8 cream-colored to yellow stigma lobes.<br />

fruits: About 1/2 to 3/4 of an inch long, scarlet, egg-shaped<br />

to club-shaped, the old flower parts persisting upon them.<br />

The seeds are black, pitted, approximately 1/16 of an inch<br />

(about 11/2 millimeters) long.<br />

Range. Entering Texas from Mexico all along the Rio Grande<br />

from its mouth to the mouth of the Pecos River, extending north<br />

along the Gulf Coast of Texas to near Rockport, but growing<br />

only in a narrow band of coastal plain extending not over a few<br />

miles from the beach. Up the river from Brownsville, Texas, it<br />

never grows north of the sandy hills overlooking the Rio Grande<br />

Valley anywhere below Eagle Pass, but near that city it leaves<br />

the vicinity of the river to grow on the summits of the limestone<br />

hills of the Edwards Escarpment past Brackettville and Uvalde,<br />

almost to Bandera and Rocksprings, Texas.<br />

Remarks. This little cactus can be recognized easily by the flex-<br />

ible, hairlike radial spines which cover its surface and from<br />

which it gets its name, the “hair-covered cactus.” No other<br />

small, unribbed cactus in the Southwest has such spines.<br />

The individual stems of this cactus are small, usually around<br />

an inch or only a little more in diameter and height, but they<br />

multiply very rapidly by branching. Often a head an inch in<br />

diameter will surround itself with as many as a dozen little offsets.<br />

The result is soon a low clump sometimes more than a foot<br />

across, made up of many heads.<br />

In Texas this cactus grows in two widely differing habitats.<br />

It grows in or near the Rio Grande Valley, usually in rich, deep,<br />

lowland soil where it seems to prefer the shelter of thickets, or<br />

else between the clumps of coastal grasses on the low, flat coastal<br />

plain almost within sight of the Gulf. It is a far cry from such<br />

places to its other habitat, where it grows on the crests of rocky<br />

hills, usually wedged into crevices of limestone ledges, often<br />

where there hardly seems soil enough to support it, and often<br />

even on the undersides of the ledges, where it is shaded by the<br />

rocks. It loves the highest brows of the Edwards Escarpment<br />

north of Uvalde and Brackettville, Texas, but it is handicapped<br />

here by its susceptibility to freezing. During the years when the<br />

winters in this area are mild its clumps proliferate and become<br />

large, but after one of the colder winters which occasionally visit<br />

this area, the plants are a sad sight, severely frozen back and<br />

often half dead. The plant seems to be able to survive almost as<br />

far north as Rocksprings, Texas, but no farther.<br />

Even though it sometimes produces clumps up to more than<br />

a foot across, M. multiceps is always an inconspicuous cactus.<br />

Even though produced in profusion, its flowers are small and<br />

not highly colored, Engelmann calling them “dirty yellow.” Its<br />

red fruits provide the only real color it has. However it is interesting<br />

for the large clusters it forms, and a clump of the little,<br />

hairy heads is attractive. Because of its habit of growing from<br />

shallow crevices of rock ledges, it makes fine, healthy growth in<br />

little pockets of soil in rock gardens and on the sides of irregular


148 cacti of the southwest<br />

rock patio walls, wherever the temperature does not get too cold<br />

for it.<br />

This particular little cactus of Texas and northern Mexico was<br />

first described by Prince Salm-Dyck in 1850, and he called it<br />

Mammillaria multiceps. It would have seemed that it could have<br />

had a history unconfused by name troubles, since no one has<br />

doubted that it is a Mammillaria of the best form, but such was<br />

not to be the case.<br />

Very much earlier, in the beginnings of cactus study, a similar<br />

cactus had been found and described in Cuba and some other<br />

West Indies islands. Miller first named this other cactus Cactus<br />

proliferus, way back in 1768. Lamarck renamed it Cactus glomeratus<br />

in 1783, and De Candolle in 1803 called it Cactus pusillus.<br />

In 1812 Haworth placed it more properly as Mammillaria prolifera.<br />

This already discovered West Indian cactus is very close to<br />

our form, and this fact was soon noticed. When Engelmann saw<br />

our Texas plant and wrote of it in 1856, he said that it “seems<br />

scarcely distinct” from the West Indian one, and so he called<br />

our Texas cactus Mammillaria pusilla var. texana. Since then<br />

the Texas form has been called in the literature almost every<br />

possible combination of variety texana or variety multiceps<br />

with each one of the various names already given for the West<br />

Indian cactus. At the same time there have always been others<br />

who have maintained that the two are separate, and so used<br />

simply Mammillaria multiceps for our cactus.<br />

It seems that the first thing necessary in trying to arrive at<br />

the correct name for this cactus is to establish once and for all<br />

how close to the West Indian cactus it is. Although this would<br />

seem to be simple, it is in reality not easy to do. It is almost<br />

impossible today to get specimens from Cuba and the islands in<br />

that area. I have not had the good fortune to observe living<br />

plants from the islands, and herbarium material from there is<br />

very scarce. This means I can only rely upon the statements of<br />

other students concerning the similarity or differences between<br />

these cacti.<br />

Looking back into the statements of these students, we find<br />

that the early students, who seemed to have observed the West<br />

Indian plants enough that Engelmann called it “well-known”<br />

all agree that it has only 12 to 20 radial spines. Coulter states<br />

that it differs from our mainland form “only in the very much<br />

fewer (12 to 20) radial spines, although numerous specimens,<br />

both dried and living, were examined for additional characters.<br />

This difference, however, is so constant and striking that, taken<br />

together with the wide geographical separation, it should stand<br />

as varietal.” He therefore agreed with Engelmann in making<br />

our form a variety of the other.<br />

Since that time it seems that students have not been able to<br />

decide whether this difference between 12 to 20 radials on one<br />

hand and 30 to 80 on the other hand is justification for two<br />

different varieties or two different species. There has, altogether,<br />

been a hundred years of discussion of this problem, and there<br />

still is no agreement. I am choosing to list our form as a separate<br />

species. However this decision, I wish to make clear, is not on<br />

the basis of any first-hand observation or any new data, but<br />

only on the strength of its being listed this way by Britton and<br />

Rose, Craig, and Backeberg. These are the most thorough stu-<br />

dents to have dealt with it recently.<br />

There remains still another problem concerning this cactus.<br />

There seem to be two rather distinct growth forms of it in Texas<br />

and northern Mexico. The two forms seem identical except that<br />

one of them has the outer parts of the centrals dark brown, red-<br />

brown, or black, while the other has the centrals all whitish or<br />

else translucent, honey-yellow. Plants of these two different<br />

colorings seen side by side are conspicuously different, and the<br />

difference has been noticed by several authors.<br />

Engelmann, Coulter, and Britton and Rose seem none of them<br />

to have seen both forms. The descriptions by the first two do not<br />

include characters of the yellow-spined form at all. Of course,<br />

Britton and Rose’s very broad description of the West Indian<br />

M. prolifera would include it, but they apparently knew nothing<br />

of that species being in either the United States or Mexico.<br />

Schulz and Runyon only listed M. multiceps, but stated, “There<br />

are two distinct varieties. The most common has brown spines.<br />

The other has gray or nearly white spines. The flowers and fruits<br />

of the two varieties are very much alike.” This was probably the<br />

first recognition of the existence of the two forms.<br />

Borg was probably the next to refer to the two forms. He<br />

described the yellow-spined one briefly but clearly, but he sadly<br />

confused things by calling this so far unnamed, yellow-spined<br />

form M. prolifera var. texana Eng., while he reserved for the<br />

dark-spined form the name M. prolifera var. multiceps SD.<br />

It seems that Borg makes an error here. It is unfortunate that<br />

he equates Engelmann’s variety texana with the yellow-spined<br />

form of this species, because Engelmann expressly stated the<br />

plants he meant by that name to have “Interior spines… in<br />

young or weakly specimens whitish with dark tips, in robust<br />

ones yellow at base, brown upwards, and almost black at tip.’<br />

It seems clear that variety texana is the same plant as Salm-<br />

Dyck’s M. multiceps.<br />

Is there then a reason for describing the yellow-spined form as<br />

a separate variety at all? When I first discovered it I thought<br />

that there was. All of the specimens I have seen from the summits<br />

of the high hills in the Texas hill country are of this coloring,<br />

and as I had seen only the dark-spined form from the very<br />

different coastal and Valley habitat, I thought that this could<br />

be a separate form with its own northern range. But this idea<br />

proved to be wrong. I was very surprised to find the two forms<br />

growing together in Potrero Canyon in the Santa Rosa Mountains<br />

of Coahuila, Mexico. This showed that the dark-spined<br />

form could also grow in high, rocky situations, and I soon confirmed<br />

this in other Mexican locations. When I later saw the<br />

yellow-spined form, formerly seen only on high hills, growing<br />

in a habitat about as completely the opposite as could be found,<br />

on the coastal plain only a short distance from Copano Bay and<br />

the Gulf of Mexico, I realized that the ranges and habitats of


genus Mammillaria 149<br />

these two cannot be separated. They actually grow together on<br />

the same hillsides, in a few cases. They appear to be merely<br />

growth forms, perhaps simple phenotypes of the same variable<br />

population. They are much like the two color phases found in<br />

the same populations of Echinocereus caespitosus.<br />

Given the kind of care used on any but the most extreme desert<br />

forms of cacti, even in a small pot, this little cactus will grow<br />

and proliferate its small heads into an interesting cluster. It<br />

blooms freely for many weeks at a time, and for months after<br />

that its brilliant, scarlet fruits will be popping out to decorate it.<br />

Mammillaria microcarpa Eng.<br />

“Fishhook Cactus,” “Pincushion Cactus,” “Sunset Cactus”<br />

Description Plate 40<br />

stems. Single until old, then clustering at the base or sometimes<br />

branching above the base to form small clumps of several<br />

stems. Each stems grows to at least 6 inches, but usually<br />

to only about 3 inches tall; each 1 to 31/2 inches in diameter<br />

and practically spherical, conical, or somewhat cylindrical in<br />

shape. The tubercles are cylindrical when young, then conical<br />

from more or less quadrangular bases which are sometimes<br />

flattened from side to side when old; the younger tubercles<br />

up to about 1/4 of an inch long and the same width at their<br />

bases, the old tubercles up to nearly 1/2 inch long.<br />

areoles: Dimorphic. The spinous portion, at the tip of the<br />

tubercle, is small, round or oval, with some white wool at<br />

first, but later bare. The floral or vegetative part, in the axil,<br />

is without wool, and so the axils are bare.<br />

spines: There are conspicuous, slender, white radial spines and<br />

dark-colored, hooked central spines. There are 20 to 30 radials<br />

radiating evenly from the areole, lying flat upon the surface<br />

of the plant, and interlocking with those of neighboring areoles.<br />

They are very slender, but stiff, often slightly hairy under<br />

a magnifying glass, and 1/8 to 1/2 inch long. The upper and<br />

lower spines of each areole are the shortest, usually about 1/4<br />

of an inch long, with an occasional one only 1/8 of an inch,<br />

while the lateral spines are the longer. Each areole has 1 to 3<br />

centrals. The lowermost one of these centrals stands out perpendicular<br />

to the plant surface, is 1/4 to 3/4 of an inch long,<br />

red-brown to almost black, with the base the lightest color and<br />

the dark point strongly hooked. The upper centrals stand upright<br />

just in front of the upper radials, forming a V if there<br />

are two of them, but there may be only one. These are never<br />

hooked, are usually whitish over the lower half, with the upper<br />

half red or yellow-brown.<br />

flowers: Rose-purple in color, 3/4 to 11/8 inches long and<br />

wide. Usually there are a number of these flowers forming a<br />

ring around the new growth of the plant. They open widely.<br />

The outer petals are short, oblong or conical, having brown-<br />

ish midlines and pink, fringed edges. The inner petals are<br />

longer, about 3/4 of an inch long, slender (being about 3/16 of<br />

an inch wide at their widest), with smooth edges and either<br />

bluntly pointed or sharply pointed tips. These petals may be<br />

solid pink in some specimens, or in others have deeper midlines<br />

fading to almost white edges. The filaments are pink, the<br />

anthers pale orange. The style is very long, easily 1/4 of an<br />

inch above the stamens, making the pistil about equal to the<br />

length of the longest petals. There are 6 to 10 slender, light<br />

green stigma lobes about 3/16 of an inch long.<br />

fruits: Scarlet when ripe, oval to club-shaped, 3/4 to 1 inch<br />

long, with the dried flower parts remaining upon them. These<br />

fruits develop rather slowly, there is a long blooming season,<br />

and those started late in the summer remain as small, green,<br />

oval structures under the spines of the plant until during the<br />

winter they dry up without ripening properly at all. For this<br />

reason the fruits have sometimes been called dimorphic. The<br />

seeds are black, pitted, almost round, and less than 1/16 of an<br />

inch (1 millimeter or less) in diameter.<br />

Range. Extreme western Texas from the very northwestern corner<br />

of Presidio County to the Franklin Mountains near El Paso,<br />

and across the extreme southern part of New Mexico into Ari-<br />

zona and Mexico.<br />

Remarks. This is the dainty little fishhook cactus we so often<br />

see in dimestore bins and dish gardens. It is rather common in<br />

the places where it grows, and its white, spine-covered surface<br />

with the slender, dark brown centrals standing out for all the<br />

world like fishhooks makes it a sure seller for the novelty it<br />

presents. Those who succeed in providing it the perfect drainage<br />

and hot sun it needs will have the added reward, quite without<br />

warning some day in the summer, of a whole ring of fine, rose-<br />

purple flowers circling the crown of the plant.<br />

Mammillaria microcarpa was first mentioned in Engelmann’s<br />

Emory Report of 1848. Engelmann at that time only had a<br />

drawing of a plant seen on the expedition in Arizona. Because<br />

this drawing showed extremely tiny fruits only about 1/8 of an<br />

inch long, Engelmann remarked that the species should be called<br />

M. microcarpa, as no other cactus has so small a fruit. This drawing,<br />

incidentally, shows no hooked central spines.<br />

In his later studies, Engelmann examined many specimens of<br />

our cactus, and in his writings described it completely. However,<br />

he apparently did not ever connect it with the cactus of<br />

Emory’s report, since he named it Mammillaria grahami and<br />

only referred to it under this name.<br />

The cactus was known by this name, M. grahami, by all writers<br />

up to Britton and Rose. In their large work on cacti, they<br />

went back to the earlier name, M. microcarpa, for this species.<br />

This was because, in 1922, at Dr. Rose’s request, a Mrs. Ross<br />

visited the locality in Arizona which was thought to be the same<br />

spot where the cactus figured in the drawing made on Emory’s<br />

expedition was said to have been abundant, and there she found<br />

our cactus growing. This was taken by Britton and Rose as con-


pellosperma -><br />

phellosperma<br />

150 cacti of the southwest<br />

clusive evidence that the drawing was of this plant, and they<br />

resurrected the old name, M. microcarpa, to replace the one<br />

which had been used for the past 70 years.<br />

There were naturally many who objected. They pointed out<br />

that the 1848 drawing and description does not show or mention<br />

the most conspicuous feature of the plant, the hooked central<br />

spines—a remarkable omission—and that the 1/8-inch fruits<br />

of that description are surely not the 3/4- to 1-inch fruits of our<br />

cactus. These were undoubtedly the reasons why Engelmann felt<br />

it necessary to give this cactus the different name of M. grahami<br />

when he came across it. In reply to Britton and Rose’s argument<br />

that since our cactus grows at the location where Emory’s plant<br />

was collected, the two must be the same, it was argued that since<br />

Mrs. Ross’s determination of the exact spot 70 years later was at<br />

best a guess, and since in many cases moving only a few hundred<br />

yards from a location gives a different cactus, we have too little<br />

reason indeed to maintain from the supposed identity of location<br />

that our plant with the long, dark, hooked centrals and<br />

fruits nearly an inch long is M. microcarpa of the early report,<br />

which lacked dark, hooked spines and which had tiny fruits.<br />

But such was the veneration in which the work of Britton and<br />

Rose was held, that soon everyone was using the name M. microcarpa<br />

for this species; I have not found an author in the past 30<br />

years going back to the other name. It is, therefore, in spite of<br />

my own suspicion that the original M. microcarpa was some entirely<br />

different plant, but in deference to the unanimous use of<br />

this name by all other writers of recent years, that I list this<br />

cactus under that name.<br />

This cactus grows in abundance in the Franklin Mountains<br />

near El Paso, Texas. Occasionally it is found in the mountains<br />

northwest of Presidio, Texas, particularly near Candelaria, Texas.<br />

In New Mexico it is encountered in the mountains across the<br />

very southern part of the state, particularly in the Tortugas and<br />

Selden mountains near Las Cruces, and in the Peloncillo Mountains<br />

in the southwestern corner of the state.<br />

This cactus has very close relatives in M. phellosperma Eng., of<br />

Arizona, California, and Utah, and M. sheldonii B. & R., of Sonora,<br />

Mexico. M. milleri B. & R. is considered to be a synonym.<br />

Mammillaria wrightii Eng.<br />

Description Plate 41<br />

stems: Solitary, the part above the ground 11/2 to 3 inches in<br />

diameter and varying in shape according to environmental<br />

conditions. When moist and growing it is hemispherical or<br />

even spherical, when desiccated and dormant it is very much<br />

flattened and even depressed on top. The stem is prolonged<br />

below the ground into a tapering, top-shaped base which is<br />

similar to a short taproot, but which has never been interpreted<br />

that way. The surface above the ground is covered by coni-<br />

cal or almost cylindrical tubercles 1/4 to 7/8 of an inch long,<br />

whose bases are round and whose axils are naked.<br />

areoles: Dimorphic. The spinous portion on the tip of the<br />

tubercle is round, with short white wool. The floral part is in<br />

the axil.<br />

spines: There are 8 to 15 radial spines which lie flat upon the<br />

surface of the plant, or very nearly so. They are 3/16 to 5/8 of<br />

an inch long, and vary in color. They may be all white, or<br />

they may be white with brown tips. There is a tendency for<br />

the uppermost of them to be darker, and sometimes these are<br />

all brown. The bases are somewhat bulbous, and the spines<br />

have more or less of a fine, hairy pubescence upon them visible<br />

only with a lens. There are 1 to 4, but usually 2 or 3 central<br />

spines. These are red-brown or blackish, slender, 3/8 to 1/2<br />

inch long, usually all hooked, although it is said that sometimes<br />

the upper one may be straight. These spines also have<br />

some fine pubescence upon them.<br />

flowers: Bright purple in color. They are about 1 inch long<br />

and the same in diameter when open fully. There are about 13<br />

outer segments which are somewhat triangular in shape, about<br />

3/8 of an inch long by 3/16 of an inch wide, pointed, purplish-<br />

green, with light-colored, fringed edges. The inner petals are<br />

about 20 in number, almost linear, about 3/4 of an inch long<br />

by 1/8 to 3/16 of an inch wide, pointed, and purple. The filaments<br />

are greenish at their bases, pinkish above. The anthers<br />

are pale orange. The style is greenish, becoming pinkish above.<br />

There are 11 lemon-yellow stigma lobes.<br />

fruits: About 3/4 to 1 inch long, oval to broadly obovate,<br />

about 5/8 of an inch in widest diameter, dull purple in color.<br />

These fruits often occur well to the sides of the plant. The<br />

seeds are large, more than 1/16 of an inch (about 2 millimeters)<br />

long, almost round, and black, with pitted surfaces.<br />

Range. Reported from several points in a narrow strip of southcentral<br />

New Mexico, the northernmost of these points being<br />

near Anton Chico, New Mexico, another in the mountains east<br />

of Carrizozo, and others in the Organ Mountains near Las<br />

Cruces, New Mexico.<br />

Remarks. This cactus and the next are always surrounded by<br />

confusion. The reason for this is easy to understand when it is<br />

realized that they are very similar in all of their characteristics—<br />

so much so that it is extremely hard to know which of them one<br />

has if he sees only one of them by itself. Since they are so rare<br />

that few have seen them at all, and very few students have actually<br />

seen both species together for real comparison, the descriptions<br />

of them have sometimes been inconsistent and they<br />

have sometimes been mistaken for each other so that the limits<br />

of their ranges are blurred. We will discuss their relationship<br />

after they have both been described.<br />

It can be said that M. wrightii is a very rare little cactus of<br />

the lower mountain slopes in south-central New Mexico. It is so<br />

rare that mass, organized searches for it have, on several occa


genus Mammillaria 151<br />

sions, failed to turn up a single specimen, but at the same time,<br />

individuals looking casually over the hills in its territory have<br />

occasionally stumbled upon a specimen or two. Besides the fact<br />

of its rarity, another reason for this sort of thing happening is,<br />

no doubt, the fact that the cactus has a large base underground,<br />

and in the late summer, when it goes dormant, it protects itself<br />

by shrinking down almost to ground level, becoming very flat.<br />

It is very hard to see in this shrunken condition in which it stays<br />

throughout the winter.<br />

Hallenbeck, after a most concerted hunt in 1934, involving<br />

many people and covering a huge territory, finally found a<br />

stand of nearly fifty specimens of what may have been this cactus<br />

near Anton Chico, New Mexico. He promptly took them<br />

all, explaining his action by saying that he believed those were<br />

all that existed and he felt bound to take every one in order to<br />

preserve the species. I think no better criticism of this sort of<br />

wholesale gathering of rare plants need be made than to note<br />

that no one of his collected plants or any of their offspring can<br />

be traced today, but that there have been a few specimens collected<br />

in the wild in the years since then, proving that nature is<br />

the best preserver of her children after all.<br />

As a result of this sort of thing, M. wrightii is more than ever<br />

a very rare little cactus of the mountains and hills of south-<br />

central New Mexico. As far as I know, the species is not on the<br />

market, and fortunate is the cactophile who ever gets to see one.<br />

I want to express my appreciation to Mr. Ed Nadolney of Albuquerque,<br />

New Mexico, for letting me have several fine specimens<br />

of this cactus from his living collection, as I have not been<br />

fortunate enough to collect the species myself.<br />

Mammillaria wilcoxii Toumey<br />

Description Plate 42<br />

stems: Solitary, rounded to nearly spherical without a large<br />

underground base, to 6 inches in diameter, but often much<br />

smaller. The tubercles are 3/8 to 3/4 of an inch long, conical<br />

from somewhat flattened bases about 1/4 of an inch wide. The<br />

axils are naked.<br />

areoles: Dimorphic. The spinous portion on the tip of the<br />

tubercle is oval and very small, with some short wool. The<br />

floral portion is in the axil.<br />

spines: There are 14 to 22 radial spines which are 3/8 to 5/8 of<br />

an inch long, slender and straight, white with light brown tips<br />

or sometimes the whole spine light brown, these radiating<br />

from the areole or spreading outward slightly, interlocking<br />

with those of neighboring areoles to almost obscure the view<br />

of the tubercles. There may be 1 to 5 spreading centrals, at<br />

least one of which is hooked, but in most cases there are 2 or<br />

3 of them and they are all hooked. These are brown, slender,<br />

and 1/2 to 11/4 inches long. Under a lens all spines show a<br />

variable amount of downlike pubescence.<br />

flowers: Pink, pale rose, or very pale purple in color. They<br />

are 1 to 11/4 inches long, about 15/8 inches wide when open<br />

fully. There are about 20 outer perianth segments which are<br />

brownish-green, almost linear, 3/4 of an inch long by about 1/8<br />

of an inch wide, pointed, and fringed with white hairs. There<br />

are up to about 40 inner petals in two rows, which are long<br />

and narrow, usually to 11/4 inches long by 1/8 of an inch wide,<br />

and pointed, and have pink midlines and cream-colored edges.<br />

The filaments are white, the anthers light orange. The style is<br />

greenish and the stigma has 5 to 9 green lobes.<br />

fruits: Pink to greenish-purple, obovate, 5/8 to 1 inch long<br />

and to about 1/2 inch thick. The seeds are black, pitted, almost<br />

round, and to 1/16 of an inch (about 13/4 millimeters) long.<br />

Range. Southeastern Arizona and Sonora, into southwestern<br />

New Mexico. It is certain that this plant is found in the mountains<br />

along the extreme southwestern edge of New Mexico, but<br />

how far east it extends is uncertain due to the failure of authorities<br />

to agree on the identity of several specimens. It seems certain<br />

it has been collected as far east in New Mexico as Silver<br />

City and near Deming, and it has been said to have been seen as<br />

far as the Franklin Mountains, but this location is doubtful.<br />

Remarks. Mammillaria wilcoxii is a more western form than<br />

M. wrightii, Arizona being its main locale. It does grow in New<br />

Mexico, but it is so rare there as to be an oddity anywhere it is<br />

found.<br />

The rarity of these two forms has made it difficult to study<br />

them, and they have been often confused. It is hard to establish<br />

ranges upon the records of only a few collections, yet this is all<br />

that have been made of either of these two cacti in New Mexico.<br />

They are so rare that few people have been fortunate enough to<br />

see the two forms side by side. This makes for such confusion as<br />

must have occurred in Wooton’s study, since he lists only M.<br />

wrightii, and then says his specimens came “from the mountains<br />

of the western side of the state, but it is reported from the upper<br />

Pecos region east of Santa Fe.” We must assume that he did not<br />

distinguish the two forms, thought he was seeing only M. wrightii,<br />

and so stated that this species occurred along the western edge of<br />

New Mexico, where no one else has ever collected it.<br />

W. Taylor Marshall became convinced that the cacti collected<br />

by Hallenbeck near Anton Chico were M. wilcoxii, and so would<br />

extend the range of this species that far northeast, although all<br />

he had were Hallenbeck’s photographs, and Hallenbeck’s own<br />

descriptions of his plants seem to contradict the theory. Marshall<br />

also maintained that some specimens collected in the Organ<br />

Mountains were M. wilcoxii, although their collector, Dr. H. V.<br />

Halladay, as well as Ladislaus Cutak, who studied them, were<br />

equally certain they were M. wrightii. In this way the ranges of<br />

the two have both been overextended and confused. Since there<br />

is no real evidence to the contrary, it seems most logical that M.<br />

wilcoxii is restricted to the southwestern corner of New Mexico,


152 cacti of the southwest<br />

while M. wrightii is found only in the south-central mountains<br />

of the state.<br />

The difference between the two cacti can be summarized as<br />

follows: M. wrightii has an enlarged underground base, 8 to 15<br />

radials, and short centrals, while its flowers are about 1 inch<br />

across, have 13 triangular sepals, about 20 inner petals, and 11<br />

yellow stigma lobes. M. wilcoxii has no such enlarged base, has<br />

14 or more radials, and longer centrals, while its flowers are at<br />

least 1 inch and usually more in diameter, with about 20 linear<br />

sepals, up to 40 inner petals, and 5 to 9 green stigma lobes. The<br />

general appearance of the two has not been better shown and<br />

contrasted than by a fine photograph of the two forms sitting<br />

side by side which is on page 183 of Marshall and Bock’s book,<br />

Cactaceae.<br />

It may be, as some have suggested, that these two forms are<br />

mere varieties of the same species. If that is so, there may be<br />

intermediate forms, but his has not been demonstrated so far.<br />

Mammillaria heyderi Muehlenpf.<br />

“Nipple Cactus,” “Biznaga de Chilitos,” “Little Chilis”<br />

Description Plates 42, 43<br />

stems: Each plant consists of a single stem which is a flattened<br />

hemisphere up to 5 inches in diameter and up to about 2<br />

inches tall. It grows from a heavy base under the ground<br />

which is not prolonged into a taproot, but which abruptly<br />

issues in fibrous roots. The stem is covered by firm tubercles<br />

neatly arranged in spiral rows, the axils between them being<br />

very woolly when young. This wool disappears on most plants<br />

on the older sides of the stem, but on an occasional specimen<br />

it persists and Sometimes seems to become even thicker between<br />

the old tubercles. The mature tubercles are up to about<br />

1/2 inch long, the tips round and conical from pyramidal,<br />

quadrangular bases about 1/4 of an inch wide. The ventral side<br />

of this base is definitely keeled. The sap of this plant is milky.<br />

areoles: Dimorphic. The spinous portion on the tip of the<br />

tubercle is round, small, with a little white wool at first, then<br />

naked. The floral portion is in the axil, with much white wool<br />

at first.<br />

spines: There are 9 to at least 26 radial spines radiating evenly<br />

like the spokes of a wheel from each areole. They are 3/16 to<br />

1/2 inch long, the lower being longer and stouter than the<br />

upper ones, although all of them are slender and weak. The<br />

small upper ones are whitish with red-brown tips, while the<br />

larger lower ones are all red-brown on most specimens. There<br />

is usually only one central spine which stands straight out<br />

from the center of the areole and is short—only 1/8 to 3/8 of<br />

an inch long. It is all dark reddish-brown, or reddish-brown<br />

at base and tip with a zone of lighter brown in the center. On<br />

rare Specimens there are found two identical central spines,<br />

one turned upward and the other deflexed.<br />

flowers: Often occurring a number at a time arranged in a<br />

circle around the newer growth at the center of the plant.<br />

Each is 3/4 to 11/2 inches tall and wide, brownish, pinkish, or<br />

very pale purple shading to white. The outer petals have a<br />

greenish or brownish midline which shades into whitish,<br />

smooth, unfringed edges and a pointed tip. The inner petals<br />

have brownish-pink to pale rose midlines with whitish or<br />

pink edges which are entire to slightly ragged toward the<br />

pointed tips. The filaments are whitish to deep pink. The anthers<br />

are yellow. The stigma has 5 to 10 light green, cream-<br />

colored, or tan lobes.<br />

fruits: Bright carmine red in color, club-shaped, and 1/2 to<br />

11/2 inches long. They usually ripen about a year after their<br />

inception, so there is often a ring of them on the plant at the<br />

same time as the flowers. The seeds are less than 1/16 of an<br />

inch (about 1 millimeter) long, reddish-brown, and pitted.<br />

Range. As a species ranging north out of Mexico over a huge<br />

area. The northern definitely known limit runs from near Corpus<br />

Christi on the Texas Gulf Coast to near San Antonio, then<br />

almost straight north to Jackson and Greer counties in the extreme<br />

southwestern corner of Oklahoma. From there it seems to<br />

turn back southwest, running somewhere near Carlsbad, New<br />

Mexico, and from there west through southern New Mexico into<br />

Arizona. There are old reports of the cactus from the Oklahoma<br />

Panhandle and from southeastern Colorado, but I could not<br />

verify these reports.<br />

Remarks. The cactus described as Mammillaria heyderi by Mueh-<br />

lenpfordt was the nipple cactus. He originally described it as<br />

having 20 to 22 radial spines, but it soon became apparent that<br />

the number of these radials is much more variable than that.<br />

Because of this there has been confusion. Engelmann handled<br />

the specimens having radial numbers outside these limits at two<br />

different times in two different ways, once as separate species<br />

and once as varieties of one larger species. Authorities have differed<br />

on this point until this day.<br />

It seems to be rather clear now, however, that we have here<br />

one large, wide-ranging species within which are several very<br />

close forms which can hardly be considered more than varieties,<br />

and that is the way I am handling them here.<br />

This cactus grows as a more or less flattened hemisphere of<br />

very regular tubercles always arranged tidily in spirals. It never<br />

grows as large in diameter as M. meiacantha, its nearest relative<br />

in our area, but it is often less flattened and, therefore, taller.<br />

It is usually found under the partial shade of trees, shrubs, or<br />

tail grass, and cannot do well in the full, unbroken sunshine. It<br />

can tolerate more water than many cacti, and so is more easily<br />

grown out of its normal environment than many species. When<br />

it blooms, with its pinkish or brownish-white flowers—on healthy,


genus Mammillaria 153<br />

big plants, often a dozen or more of these form a flowery diadem<br />

around the top of the plant—it is a beautiful sight. If another<br />

ring of the carmine red fruits from last year’s flowering is<br />

also present outside the ring of flowers, as is often the case, then<br />

the sight is truly spectacular.<br />

The Spanish common name for the plant, biznaga de chilitos,<br />

and its English equivalent, little chilis, refer to the fruits, which<br />

are edible and flavorful and greatly relished by the knowing<br />

natives of its locality, as well as by the birds, which will even<br />

enter my greenhouse to get at them.<br />

There is quite a bit of variation in the color of the flowers in<br />

this species. Often there seem to be two main flower colors, one<br />

a pinkish midline of the petals fading to white edges, and the<br />

other a brownish or even greenish-brown midline with white<br />

edges to the petals. There have been attempts to separate the<br />

varieties on this basis, but this cannot be done, since one often<br />

finds identical plants in a single population blooming side by<br />

side with the two flower colors.<br />

I must mention here a few specimens collected in central and<br />

northern Texas which present a peculiarity of armament for this<br />

species. These were fairly large plants and typical M. heyderi in<br />

every characteristic of stem and tubercles. They were remarkable<br />

for having all radials pure white instead of more or less colored<br />

and tipped with brown. Also, the number of the radials varied<br />

from 20 to 26 per areole, with many areoles having 23 or 24.<br />

Even more remarkable on these plants was the fact that about<br />

one out of five of the areoles had two centrals instead of one.<br />

These two centrals were the same as the ordinary single central<br />

spine found standing perpendicular to the stem on the other<br />

areoles, but where there were two, one was turned sharply upward<br />

and the other as sharply downward. Neither of these was<br />

a misplaced radial, since the ring of radials was unbroken around<br />

them. Both the number of radials and the possession of two centrals<br />

in some areoles put these specimens beyond the limits so far<br />

given for M. heyderi.<br />

It is impossible to be sure from these few specimens whether<br />

this represents an extreme form of M. heyderi and the species<br />

should be expanded to include these characters, or whether they<br />

represent a new form. But since I have other specimens of M.<br />

heyderi var. heyderi on which occasional areoles have 23 or 24<br />

radials, indicating that the maximum in the usual descriptions<br />

is too low, and I also have a single specimen of M. heyderi var.<br />

applanata on which a few areoles possess the two centrals, indicating<br />

that this may be a seldom expressed character carried in<br />

all of these forms, I am inclined to assume the former. So I have<br />

expanded the description of the species to include these characters.<br />

These specimens cannot be equated with the Mexican M.<br />

heyderi var. waltheri (Bödecker) Craig, although they do in<br />

some ways link that form more closely with the other varieties<br />

of the species.<br />

Expansion of the limits of this species there has had to be, and<br />

it is hard to know where this should stop. It must not go on un-<br />

til the concept of the species is meaningless. L. Benson lists M.<br />

heyderi as occurring in very small areas of extreme southeast<br />

Arizona. His description, however, does not agree with any previous<br />

description of the cactus. He describes it as having conical<br />

tubercles, 2 to 15 radial spines, and 2 or 3 central spines. This<br />

combination of characters is not given for any previously de-<br />

scribed U. S. species in the literature.<br />

Mammillaria heyderi var. heyderi (Muehlenpf.)<br />

Description Plate 42<br />

stems: As the species.<br />

areoles: As the species.<br />

spines: As the species, except radials 20 to 26 in number.<br />

flowers: As the species, except only 3/4 to about 1 inch long<br />

and broad, and the stigma lobes 6 to 8 in number.<br />

fruits: As the species, except 3/4 to 11/2 inches long.<br />

Range. From near Brownsville, Texas, south and west into<br />

Mexico and north to the west of a line running past San Antonio<br />

to near Fredericksburg, Texas, occurring from this northeastern<br />

limit west through Texas to near Carlsbad, New Mexico,<br />

on the north, and past El Paso over all of southern New Mexico<br />

and into Chihuahua on the south.<br />

Remarks. This was the first one of this exceedingly close group<br />

to be described. It was the form actually described as M. heyderi<br />

by Muehlenpfordt, and so is the typical variety of the species.<br />

It also seems to have the widest range of the three U. S. varieties<br />

in the species.<br />

There should be little difficulty in recognizing this variety. It<br />

is the only cactus occurring in the southwest U. S. with milky<br />

sap and more than 20 radial spines.<br />

Mammillaria heyderi var. applanata Eng.<br />

Description Plate 42<br />

stems: As the species.<br />

areoles: As the species.<br />

spines: As the species, except the radial spines 14 to 20 in<br />

number.<br />

flowers: As the species, except to only about 1 inch in size,<br />

and having stigma lobes 5 to 10 in number.<br />

fruits: As the species.<br />

Range. From near Austin and Fredericksburg, Texas, south over<br />

most of south Texas and west into the Big Bend. Also collected


154 cacti of the southwest<br />

in numbers in extreme southwestern Oklahoma in Harmon,<br />

Jackson, and Greer counties. It is the form which has been reported<br />

from Cimarron County in the Oklahoma Panhandle and<br />

from southeastern Colorado, but I cannot verify these reports.<br />

Remarks. This cactus was considered a separate species and then<br />

again called a variety of M. heyderi by Engelmann, who first<br />

described it. Then Coulter, Wooton, and others considered the<br />

two synonymous. Since that time most students have listed it as<br />

a separate species, following Britton and Rose, although most<br />

have followed them also in appending a note that the two may<br />

represent only races or varieties. This seems much more likely.<br />

The cactus has also been known in the past under the name M.<br />

texensis Lab.<br />

Everything mentioned concerning the manner of growth of<br />

M. heyderi applies as well to this cactus. It cannot be separated<br />

from the typical variety except by counting spines. In part of<br />

their range they may grow together. However, the extents of<br />

their ranges are different. Variety applanata grows farther east<br />

in south Texas, since variety heyderi has not been found on the<br />

Texas coast and variety applanata has. One of the peculiarities<br />

of variety applanata is the apparently isolated population of it<br />

found in at least three counties of southwestern Oklahoma. There<br />

is no record of the cactus having been found anywhere in the<br />

approximately 300 miles separating this Oklahoma population<br />

from the regular range of the cactus in central Texas.<br />

Near Roma, Texas, a large number of large variety applanata<br />

and variety applanata were found growing together. Among<br />

these plants was one specimen of variety applanata with 18 and<br />

19 radial spines in all areoles and usually with one typical central,<br />

but on 6 areoles of the plant there were 2 equal centrals 3/8<br />

of an inch long, one pointed upward and one exactly opposing<br />

it in a downward direction. It is impossible to collect more<br />

specimens in this interesting area, as irrigation wells were already<br />

completed at the time, and a square mile of territory upon<br />

which these and almost the whole list of other cacti native to<br />

that area were growing was being cleared when the collection<br />

was made.<br />

Drs. Claude W. Gatewood and Bruce Blauch of Oklahoma<br />

State University have written me of a specimen collected by<br />

them at Mangum, Oklahoma, which has 14 to 16 radials and on<br />

a few areoles the same arrangement of 2 centrals. The possession<br />

of 2 centrals on at least a few areoles is obviously a character<br />

which can crop up at any place in this species, but which has<br />

not been mentioned in connection with it.<br />

Mammillaria heyderi var. hemisphaerica Eng.<br />

Description Plate 43<br />

stems: As the species.<br />

areoles: As the species.<br />

spines: As the species, except radials only 9 to 13 in number.<br />

flowers: As the species, except almost always pinkish white<br />

instead of purplish or greenish white and somewhat larger<br />

than in the other forms, being 1 to 11/2 inches tall and wide.<br />

Stigma lobes 5 to 8.<br />

fruits: As the species, except averaging smaller than those of<br />

the other forms.<br />

Range. From eastern Mexico along the Texas Gulf Coast from<br />

Brownsville to the vicinity of Corpus Christi. Extending inland<br />

in south Texas not over 30 to 50 miles, but extending northwest<br />

from the upper end of its coastal limit as far as Atascosa County,<br />

Texas. Also collected in numbers in Jackson and Greer counties<br />

of southwestern Oklahoma, and reported from the Organ<br />

Mountains of New Mexico, although this latter record is not<br />

certain.<br />

Remarks. M. heyderi var. hemisphaerica is in certain respects<br />

different from its relatives within the species, but the similarities<br />

are much more obvious. It has, however, never been considered<br />

synonymous with them. Even when Coulter was calling variety<br />

applanata the same as M. heyderi, he found it necessary to list<br />

hemisphaerica as a separate cactus.<br />

Variety hemisphaerica has the smallest number of radials of<br />

the M. heyderi group. This provides the main clue for recognizing<br />

it, as any cactus with milky sap, the single short central, and<br />

only 9 to 13 radials found in our 5-state area is likely to be this<br />

variety, although there are two other cacti which at least come<br />

close to this area and show the same features.<br />

Among the things which separate this plant from variety applanata<br />

and variety heyderi are the facts that the flower is noticeably<br />

larger and the fruits usually smaller than those of its<br />

relatives. This conflicts with Backeberg’s statement that the<br />

flowers of hemisphaerica are smaller than those of its relatives,<br />

but among many specimens from United States and also from<br />

the type locality in Mexico which I have examined I have always<br />

found the flowers larger. Engelmann also states that the<br />

seeds are smaller, but Craig gives their dimensions as the same,<br />

and I have not been able to notice any difference in the seeds of<br />

the three varieties. Neither is it true that the stem of this variety<br />

is more rounded and hemispherical than those of variety heyderi<br />

and variety applanata, as its name would imply. The shape of<br />

the stem in this variety, as in the others, depends entirely upon<br />

whether the environment is favorable or unfavorable. The tallest,<br />

most rounded specimen of this species I have seen is a luxuriant,<br />

large specimen of variety applanata, while the flattest specimen<br />

I have seen was a desiccated plant of variety hemisphaerica<br />

found on the dry bluff just back of the beach near Corpus<br />

Christi.<br />

The main locality for variety hemisphaerica is the Gulf Coast<br />

from its type locality within Mexico north to Corpus Christi.


genus Mammillaria 155<br />

Here it grows to the very edges of the sand dunes along the<br />

beach, as close to the water as the bushes grow which it must<br />

have over it to protect it from the full sun it can hardly tolerate.<br />

As you go west from the Gulf into the south Texas brush<br />

country you soon lose this species, except in a narrow strip extending<br />

along the Nueces River and into Atascosa County.<br />

It is a real surprise, therefore, to collect this same cactus growing<br />

rather commonly, along with variety applanata, in Jackson<br />

and Greer counties of extreme southwestern Oklahoma, but there<br />

it is, having skipped over all of Texas between these widely discontinuous<br />

ranges.<br />

There have been supposed collections of variety hemisphaerica<br />

in the Organ Mountains of New Mexico. This is indeed a greatly<br />

different environment from the south Texas coast, and if this is<br />

our cactus, makes for one of the most curious distributions of<br />

all cacti. I cannot either confirm or deny this occurrence of the<br />

variety in New Mexico, since all I have seen are the herbarium<br />

specimens from New Mexico, and these lack the flowers. It<br />

should be remembered, however, that there grow in Arizona<br />

and nearby Mexico two species of Mammillarias very similar in<br />

most respects, with the same number of radial spines as variety<br />

hemisphaerica. Merely counting spines would not separate this<br />

variety and those two: Mammillaria macdougalii and M. gummifera.<br />

The best means of distinguishing these from our cactus is<br />

by the fact that M. macdougalii and M. gummifera have the<br />

outer perianth segments of their flowers fringed with cilia, while<br />

variety hemisphaerica has smooth edges on these segments. The<br />

fruits of M. macdougalii also fail to become bright red, remaining<br />

greenish with only the upper parts turning rose. Without<br />

having seen the flowers and fruits of these Organ Mountain<br />

specimens, I feel that it is at least as likely that they are one of<br />

these more southwestern forms as that they are variety hemisphaerica,<br />

since finding one of those in the Organ Mountains<br />

would prove far less an extension of their known ranges and<br />

environments than to find the Gulf Coast cactus there. Perhaps<br />

the reader may be fortunate enough to secure this New Mexico<br />

form and, seeing its flowers, be able to settle the question, which<br />

I feel must stand unanswered at present.<br />

Mammillaria meiacantha Eng.<br />

“Biznaga de Chilitos,” “Little Chilis”<br />

Description Plate 43<br />

stems: Each plant consists of a single circular stem up to as<br />

much as 12 inches across, but greatly depressed so that it rises<br />

to a maximum of 1 or 2 inches above the ground. This is the<br />

top of a large underground base which might be interpreted<br />

as an extremely short taproot. The almost flat surface of the<br />

stem above the ground is covered by firm, dark green or blue-<br />

green tubercles arranged in spiral rows. These tubercles are<br />

pyramidal from quadrangular bases having their ventral<br />

angle exaggerated into a keel. They vary from about 1/2 inch<br />

long and 3/8 of an inch wide at the base on small specimens to<br />

a maximum of about 7/8 of an inch long and 5/8 of an inch<br />

wide on large plants. A milky sap is exuded at any wound.<br />

areoles: Dimorphic. The spinous portion on the tip of the<br />

tubercle is nearly round and about 1/8 of an inch across, with<br />

white wool at first, but later bare. The floral portion is in the<br />

axil of the tubercle, with white wool at first, which is later<br />

lost.<br />

spines: There are 5 to 9 stout radial spines which instead of<br />

lying flat upon the surface of the plant stand more or less<br />

spreading. They are 1/4 to 1/2 inch long. The lower ones on<br />

each areole are the longest and heaviest. They are pinkish at<br />

first, fading to gray or yellowish, and always having black<br />

or dark brown tips. There is one central spine which is similar<br />

to the radials except often a little darker and only about 1/4<br />

to 3/8 of an inch long. It stands out almost perpendicular to<br />

the surface of the plant, or quite often is turned upward almost<br />

with the upper radials.<br />

flowers: Opening widely, 1 to 13/4 inches long and wide,<br />

often a number of them forming a ring of blooms just outside<br />

the newer center of the plant. The 12 to 14 outer segments of<br />

the perianth have reddish-brown midlines with pinkish, entire<br />

edges. The 14 to 16 inner petals have purplish midlines and<br />

pink or white edges. These are to 1/4 of an inch wide, with<br />

pointed tips and edges often very lightly notched toward the<br />

tip. The filaments are white or pink, while the anthers are<br />

cream or yellowish. The style is longer than the stamens. There<br />

are 6 to 9 light green stigma lobes.<br />

fruits: Elongated and club-shaped, deep rose or scarlet in<br />

color. They are about 7/8 to 11/4 inches long. The seeds are reddish-brown,<br />

less than 1/16 of an inch (less than 1 millimeter)<br />

long, and pitted.<br />

Range. From northern Mexico throughout southwestern Texas<br />

west of a line running from near Sanderson, Texas, along the<br />

eastern edges of the Davis and Guadalupe mountains to near<br />

Carlsbad, New Mexico, and throughout about the lower one-<br />

fourth of New Mexico into Arizona.<br />

Remarks. This is the largest Mammillaria found in the U. S., but<br />

certainly not the most conspicuous. Although most specimens<br />

are about 5 to 8 inches in diameter, it is still possible to find old<br />

plants up to 12 inches across—that is, if one has eyes sharp<br />

enough to find this cactus at all. Although growing easily to<br />

pie-plate size, M. meiacantha is usually so flat as to be near the<br />

shape of an inverted pie-plate, and in very dry seasons its large<br />

fleshy base shrinks and literally pulls the plant into the ground<br />

so that it is common for its tubercles to be level with the soil or


156 cacti of the southwest<br />

only slightly above it. The surrounding grass and weeds then<br />

make its discovery a real accomplishment.<br />

This species has numerous close relatives from which it is not<br />

easily distinguished. Various characters have been tried as distinguishing<br />

ones, but most of them can be matched by the extreme<br />

forms of its relatives. M. meiacantha has sharply quadrangular<br />

and keeled tubercles, but the bases of the tubercles of<br />

M. heyderi and M. melanocentra are also somewhat quadrangular.<br />

It has milky sap, but so do those other species, although<br />

on them one must often pierce the base of the stem itself to get<br />

a flow of this white sap, while in M. meiacantha it flows from<br />

any pin-prick on any tubercle. The flowers and fruits are quite<br />

similar to those of the other species, except a little larger than<br />

those of the western varieties of M. heyderi.<br />

For distinguishing M. meiacantha, then, it seems we must come<br />

back to its spines. It has only 5 to 9 stout radials, which number<br />

distinguishes it from all the M. heyderi group, as well as from<br />

M. macdougalii and M. gummifera, all of these having from 9<br />

to more than 20 slender, almost bristle-like radials. Only M.<br />

melanocentra, a Mexican species, has spines similar in number<br />

and thickness, but here the length of spines distinguishes, as<br />

M. meiacantha has radials only 1/4 to 1/4 inch long and centrals<br />

only 1/4 to 3/8 of an inch long, while M. melanocentra has radials<br />

3/4 to 11/2 inches long and the central 3/4 to 11/4 inches long,<br />

making separation easy.<br />

With only such details of spines to separate these closely<br />

related forms, it is natural that there has been a history of uncertainty<br />

about how they should be classified and how closely<br />

they should be linked in the taxonomy. Engelmann, who first described<br />

a number of them, remarked that M. meiacantha might be<br />

only a variety of M. heyderi, but this idea did not appear likely<br />

enough to any of those who studied these plants after him to be<br />

followed by any of them. All left it as a separate species until<br />

1945, when Craig, in his Mammillaria Handbook, combined M.<br />

meiacantha with the long-spined Mexican species, M. melanocentra,<br />

as M. melanocentra var. meiacantha. This seems a more<br />

likely relationship, and he was followed in this by Marsden.<br />

Backeberg, however, more recently leaves the cactus as a separate<br />

species. What future taxonomists will do, and whether the<br />

cactus will be called a variety, or remain a species does not yet<br />

seem possible to predict.<br />

At any rate, M. meiacantha is a definite resident of west Texas<br />

and southern New Mexico which anyone will want to recognize.<br />

It is more specifically a desert plant than is M. heyderi, with<br />

which most are more familiar. It can stand the full sun better,<br />

and often grows totally unshaded except for the sparse grasses<br />

of the Big Bend hillsides, while M. heyderi grows only under<br />

bushes and shrubs and suffers in full sun. But being more of a<br />

desert cactus, it also rots more quickly than the others if watered<br />

too much. This must be kept in mind when it is cultivated.<br />

There seems some confusion about the range of M. meiacantha,<br />

or else it is not now growing over large parts of its former ter-<br />

ritory. Engelmann said it was found throughout New Mexico,<br />

and Coulter, another student of the past century, stated that it<br />

grew then from the Guadalupe River of Texas on west. I can<br />

find no evidence of its having been seen east of the Big Bend<br />

and the Davis Mountains in Texas, or north of approximately<br />

the southern one-fourth of New Mexico since their times.<br />

Mammillaria sphaerica Dietrich<br />

[Dolichothele sphaerica (Dietrich) B. & R.]<br />

Description Plate 44<br />

roots: A thick, soft, fleshy taproot sometimes up to 1 inch<br />

thick.<br />

stems: Light green in color, spherical, often depressed at the<br />

top. These heads usually produce new ones on all sides rapidly,<br />

so that a mature plant usually consists of a low mass up<br />

to nearly a foot in diameter composed of irregularly clustering<br />

stems. The tubercles making up the stem of a vigorous<br />

plant are very soft, spreading loosely, and ranging from 1/2<br />

to 11/4 inches long. They are cylindrical, usually tapering toward<br />

the end, and are not grooved and not broadened at the<br />

base. When a plant suffers from insufficient water or too<br />

much sun, however, these same tubercles will shrink to as<br />

little as 1/4 of an inch long and become firm, regular in size,<br />

and closely packed, giving the plant a very different ap-<br />

pearance.<br />

areoles: Dimorphic. The spinous portion on the tip of the<br />

tubercle is small and circular, with some short wool at first,<br />

becoming bare. The floral portion in the axil of the tubercle<br />

has some hairs which may or may not persist after the flower.<br />

spines: There are 12 to 15 radial spines which are slender and<br />

weak. They radiate evenly around the areole, but on the<br />

same areole they will usually vary in length from the three<br />

lower ones which are as short as 3/16 of an inch to the thicker<br />

lateral and upper ones which are 3/8 to 5/8 of an inch long.<br />

They have brownish, enlarged bases, and the upper parts are<br />

smooth, hornlike, and yellow in color, becoming gray with<br />

age. There is one central standing out perpendicularly to the<br />

plant, it being similar to the radials except slightly thicker<br />

and 3/8 to 1/2 inch long.<br />

flowers: Clear lemon-yellow in color, about 2 to 21/2 inches<br />

in diameter and height. The ovary is cylindrical, light green,<br />

and naked. There are a total of about 25 petals. The outer<br />

ones are short, narrow, linear, and pointed. The next ones are<br />

long—the longest on the flower, broadening toward the point<br />

to about 3/8 of an inch wide, with brownish midlines and yellow,<br />

smooth edges. The innermost petals broaden from very<br />

narrow bases to about the same width, but are shorter. They


genus Mammillaria 157<br />

are lemon-yellow. The filaments are pale orange and swirled<br />

in position, the anthers of the same color being tucked in<br />

around the style. There are 8 long, rough stigma lobes which<br />

are yellowish. The flower is fragrant.<br />

fruits: Seldom seen, as the plants do not seem to set many<br />

fruits even in the native habitats. When produced they are<br />

egg-shaped, about 1/2 inch long, persisting on the plant as<br />

green structures between the tubercles until at least the following<br />

winter, when they turn maroon, although they have<br />

usually been eaten by animals and disappeared before this. The<br />

seeds are black, with pitted surfaces.<br />

Range. South Texas below a line from near Laredo to approximately<br />

Corpus Christi, and on into Mexico.<br />

Remarks. This cactus grows under the bushes in the thickets of<br />

the south Texas brush country. Its low clusters are rather common,<br />

wherever the brush has never been cleared, below the longitude<br />

of Laredo on the west and Corpus Christi on the east,<br />

but it is much more often found, at least today, in the western<br />

part of its range than nearer the coast. Where the brush has at<br />

any time been cleared and then has grown back again, as in vast<br />

areas of this territory, the vegetation may look the same, but<br />

there is little use searching for our cactus, as it cannot survive<br />

the sun without its protecting shrubs, and it does not seem to be<br />

able to reestablish itself under the new growth.<br />

Mammillaria sphaerica can most quickly be distinguished<br />

from the other Mammillarias of Texas, which it superficially<br />

resembles, by its color. Its surface is a very light, almost sickly<br />

yellow-green, even when most healthy, while all of the other<br />

Mammillarias we have are dark green or even bluish-green.<br />

When growing in its habitat or with perfect conditions, M.<br />

sphaerica has large, soft, loose, spreading tubercles. The looseness<br />

of these and the variation in their sixes on the several heads of<br />

a robust clump make it hard to distinguish the individual heads<br />

of such a clump. It often looks like one disorganized stem. But<br />

if water or shade is withdrawn from such a plant, or if it is<br />

transplanted, the tubercles will shrink to less than one-half their<br />

former size and be pulled together until the plant appears entirely<br />

different, being now a cluster of small, spherical, compact<br />

heads of short, firm tubercles. This latter appearance seems<br />

to be the condition from which all of the early descriptions were<br />

made.<br />

The flowers of this species are large, a beautiful, clear yellow<br />

in color, and produced in large numbers, on a healthy specimen.<br />

It is not uncommon for a good specimen to have a dozen blooms<br />

at a time, covering it entirely. I had trouble photographing this<br />

species, trying to get a good specimen in bloom with few enough<br />

flowers so that the plant body would still be visible.<br />

Because of its beautiful flowers plus the liking of this species<br />

for shade and its rather good tolerance of coolness and moisture,<br />

this should be a much better cactus for growing in northern and<br />

eastern climates than the strictly desert species. One must remember,<br />

however, that it cannot stand freezing, and so would<br />

have to be well-protected when grown in the north.<br />

M. sphaerica has a very close relative in northern Mexico,<br />

called M. longimamma. The two are very similar, and might<br />

even be considered varieties of the same species. The latter, however,<br />

is larger in most respects than our Texas form, its tubercles<br />

to 2 inches long, its spines to 1 inch long, and its flowers<br />

about the same size. The two may easily be confused, and two<br />

characters only are usually given by which to distinguish them.<br />

M. sphaerica has smooth spines and the filaments of its flowers<br />

are swirled, while M. longimamma has rough or hairy spines<br />

and straight filaments.<br />

These cacti have, since Britton and Rose’s fragmentation of<br />

the genera, often been considered to make up a separate genus<br />

called Dolichothele. The validity of this separation is still the<br />

subject of much discussion.<br />

Through the years of discussion of this problem, the reasons<br />

given for the separation of these plants from Mammillaria have<br />

boiled down to only one real one. It is undeniable that the flowers<br />

of these plants have a longer and more constricted flower<br />

tube than the typical Mammillarias, and this is the whole reason<br />

upon which Buxbaum, for instance, insists the genus Dolichothele<br />

should be kept separate. He insists that in these plants the<br />

tube is a solid, closed column, and that this is essentially dif-<br />

ferent from the more hollow tubes of other cacti.<br />

The evaluation of this character is a very technical job for<br />

a botanist. Suffice it to say here that while Buxbaum considers<br />

this “column-building” of the tube an essentially distinct character<br />

isolating these plants from all other cacti, others have<br />

noted that there are degrees of lengthening and closing of the<br />

tube among cacti, and that these cacti are not the only ones<br />

showing this feature to at least some degree.<br />

Backeberg states flatly that Buxbaum erred in saying that this<br />

character fully isolated the Dolichotheles from all others. He<br />

devotes several pages of very technical discussion in Die Cactaceae<br />

to this problem, and there is no reason to repeat his<br />

points here, but he concludes with statements that, strictly speaking,<br />

all arguments for the separation of these cacti are removed,<br />

and that after the inclusion of the genus Phellosperma and<br />

Krainzia into Mammillaria again by Buxbaum, there exists no<br />

ground for separating Dolichothele from Mammillaria either.<br />

His argument is so effective, in my opinion, that I fail to see<br />

why, after making it, he has gone ahead and listed the genus<br />

Dolichothele anyway in his own account. I only differ from<br />

him by acting upon his arguments and placing these cacti back<br />

into Mammillaria. The whole point of his discussion seems to be<br />

that opinions can at this stage differ, and that the final solution<br />

of “the Doiichothele problem” cannot even yet be presented.<br />

With this I heartily agree. I only differ in feeling that much<br />

confusion would have been avoided if separations had not been<br />

made until the evidence for them had been definite in the first


158 cacti of the southwest<br />

place, and that, since the evidence for this particular separation<br />

is not clear even yet, the best thing for the service of clarity<br />

would be to drop the separation which has always stood, at<br />

best, on doubtful grounds. If cactus genera are to be meaningful<br />

it seems they should be based on something more tangible than<br />

this, or taxonomy will always remain the bugaboo it is now<br />

for the cactophile, and we will continue to be the target of jokes<br />

about “cactusization,” defined by C. V. Morton as “the process<br />

of fragmenting or even pulverizing large genera of plants<br />

a popular sport among numerous fanciers of the Cactaceae.”


Genus Opuntia Miller<br />

Placed last in this account is the large genus, Opuntia. Those<br />

who deal in matters of primitive versus advanced and theories<br />

of development tell us it should really be the first United States<br />

genus considered. The Opuntias are generally regarded as more<br />

primitive than the cacti we have already enumerated, and they<br />

also certainly deserve first place for their success. In over half<br />

of our states Opuntias are the only cacti found, and it is these<br />

cacti which enable us to say that cacti grow over almost the<br />

whole of the U. S. It is also the Opuntias which have escaped<br />

and flourished when introduced in such faraway parts of the<br />

world as the Mediterranean and north African countries, where<br />

they have become in many places a common part of the scenery,<br />

and Australia, where they have become the classic examples of<br />

plant invaders.<br />

But in spite of all this I am putting the genus Opuntia at the<br />

back of the book. This is in deference to the fact that most<br />

people find them the least interesting of the cacti, and many<br />

people associate them too strongly with unpleasantness to be<br />

able to appreciate them at all. Even many cactophiles lack interest<br />

in them. It is not at all uncommon to find a fancier with<br />

a large collection of almost all other types of cacti, which includes<br />

not a single Opuntia, or only a few of the more exotic<br />

forms of the genus. Some collectors will travel hundreds of miles<br />

to get a new barrel cactus and on the way pass by twenty species<br />

of Opuntias all in sight from the highway without even<br />

stopping to study them.<br />

There are very real reasons for this lack of interest in or actual<br />

avoidance of the Opuntias. I do not deny that. These cacti<br />

bring it upon themselves. The Opuntias make no attempt to<br />

please, and actually seem to be the experts in every means of<br />

antagonizing other organisms. They have several built-in fea-<br />

tures not shared by other cacti, which lose them friends.<br />

No doubt their most effective feature in provoking our dislike<br />

is one of their most obvious distinguishing characteristics—<br />

their possession of glochids. Glochids are special spines produced<br />

by this genus, usually in great numbers. They are distinct from<br />

the ordinary spines, which these cacti usually have in profusion<br />

as well, although there may be intermediates.<br />

The ordinary, larger spines of a cactus are dangerous enough,<br />

may inflict real injuries, and in cases where they are hooked or<br />

positioned at opposing angles on the plant, may hold one all<br />

too firmly, but they are not ordinarily barbed, and so they can<br />

usually be extracted from clothing or flesh rather easily if one<br />

is careful to remove them at the same angle they went in. But<br />

not so the glochids. These are comparatively very slender, often<br />

almost invisible they are so thin, and during development the<br />

cells covering the surface of each glochid loosen on their posterior<br />

edges so that on the completed glochid they stand as hundreds<br />

of firm, scalelike structures, each aimed obliquely to the<br />

rear—literally forming hundreds of tiny barbs to hold this tiny<br />

spear in whatever soft tissue it may pierce with its sharp point,<br />

and to rend and tear if there is an attempt to remove it. Add to<br />

this the fact that glochids, instead of being solidly attached to<br />

the plant like the other spines, become when mature so loosely<br />

held that they come off the areole at the slightest touch, and<br />

you have here instruments of torture parallel to and equally as<br />

diabolical as the spines of the porcupine. Man or beast usually<br />

comes away from a brush with an Opuntia bearing in his flesh<br />

whole clusters of these tiny, almost invisible glochids, which<br />

may not have been felt at all when they pierced the flesh, but<br />

which are firmly imbedded there—not deep, not even through<br />

the skin—but each one well set and with its shaft projecting so<br />

that any slightest touch to it or pressure on it makes the tiny<br />

barbs tear, and produces a pain like a needle prick. They are<br />

especially maddening because they are often so tiny that you can<br />

hardly see what is causing all the pain, and so delicate that if<br />

you do find them and take hold of them to remove them, their<br />

ends usually break off, leaving the tips imbedded, with only


is -> his<br />

160 cacti of the southwest<br />

the stumps to be bumped and cause continual pain. If one cannot<br />

get the maddening things out at once before they work in<br />

and set their barbs, the only really effective thing to do for a<br />

mass of tiny glochids in the flesh is to shave them off with a<br />

close-shaving razor. Once the projecting ends are removed this<br />

way, the tips in the skin will not be felt again and will soon<br />

work out.<br />

Some Opuntias make use of barbs on their larger spines as<br />

well, and on some of them a whole branch comes off and remains<br />

stuck to animal or human which has touched it, hitching<br />

a ride to a new location this way. It is an extremely unpleasant<br />

task to pull one of these branches off when its spines are well<br />

imbedded.<br />

So who can blame the person who wouldn’t touch an Opuntia<br />

with a ten-foot pole, let alone have it in his greenhouse,<br />

especially when many of them have a habit of spreading out<br />

into good-sized bushes and so taking up much room? Add to<br />

this the fact that they are the most difficult of all cacti to classify,<br />

the most variable and, therefore, difficult to recognize,<br />

and you have reasons enough for avoiding them.<br />

Yet there is much to be said for the Opuntias as well. They<br />

have many quite interesting features, may be very beautiful, and<br />

are also the most widely used for food of any cacti. And then<br />

they are undoubtedly the most challenging of the cacti. So<br />

there are those most dedicated of cactophiles who wade right<br />

into the thickets of the Opuntias, so to speak, and study them<br />

out. When to these people who are dedicated to understanding<br />

all cacti are added all of the people who are more likely to see<br />

an Opuntia in the back pasture or by the roadside than any<br />

other cactus, and who would like to know something about it,<br />

there must be a large number who will find interest in the admittedly<br />

difficult account of this complex, creeping, sprawling<br />

group, whose relationships are sometimes almost as hard to understand<br />

as its thickets are to penetrate. We offer them here, at<br />

the end of our account, as the final challenge in cactus study.<br />

Usually mentioned characteristics of the Opuntias are the<br />

possession of jointed stems; cylindrical or conic leaves on young<br />

stems; the presence of glochids; the production of spreading,<br />

rotate flowers with more or less sensitive stamens and with<br />

areoles, which often produce glochids and spines, on the ovaries;<br />

fruits with thick rinds; and seeds comparatively large, rounded<br />

in one plane and flattened in the other, while covered by hard,<br />

bony, light-colored arils.<br />

There is usually little difficulty in recognizing an Opuntia<br />

and telling it from the cacti of the other genera. One or another<br />

of the features just listed is almost always so obvious that one<br />

could hardly miss it. So we do not need to dwell here on the<br />

characteristics of the genus.<br />

But once a person knows he has an Opuntia, his problems have<br />

only begun, if he wishes to classify it further. This is because<br />

the process of sorting out the Opuntias within the group is one<br />

of the most difficult in all taxonomy. Several things contribute<br />

to this difficulty.<br />

One of these is the fact that the Opuntias seem to react to<br />

differences in their environments more quickly and with more<br />

drastic growth-form changes than do other cacti. In the Echinocacti<br />

or Echinocerei the plant usually grows much the same as<br />

any other of its own species as long as the environment is tolerable<br />

at all, and then ordinarily, if the environment becomes so<br />

changed that it cannot put out typical growth, it just fails to<br />

grow at all. On the other hand, the Opuntia in a bad situation<br />

will grow on, but grow in a form often so radically different<br />

from the typical that one would hardly suspect the environmentally<br />

modified specimen to be of the same species as a typi-<br />

cal one.<br />

As a result of this fact, where, for instance, in the former genera<br />

one can list maximum and minimum spine numbers, spine<br />

lengths, and spine characters within definite, rather narrow, and<br />

unvarying limits, and often use these to recognize species, one<br />

has to list the spines of Opuntias within widely varying limits<br />

and one must be very careful in any attempt to delineate species<br />

of Opuntias by their spine characters. A simple experiment, if<br />

only once carried out, would keep a person from ever again<br />

depending on Opuntia spines remaining the same. It is easy to<br />

take half a dozen different, typically spiny Opuntias and by<br />

growing them a few years in a very shady, moist situation, end<br />

up with half a dozen very nearly indistinguishable, spineless or<br />

only weakly spined specimens. All too often such an atypical<br />

plant has been given a separate name or has been mistaken for a<br />

normally weak-spined species. The literature is, therefore, a mo-<br />

rass of conflicting reports.<br />

Other characters of the Opuntias, such as stem size or even<br />

stem shape, flower and fruit size, and so on, are also capable of<br />

being influenced by the environment. Where an Echinocactus or<br />

a Mammillaria either puts out a standard-sized flower and fruit<br />

or else just waits to make the effort until the environment is<br />

more favorable, an Opuntia in a poor situation usually does not<br />

mind at all putting out a flower half as big as typical, and may<br />

go ahead to ripen a fruit also half as big as it should have been.<br />

Great care must be taken to allow for this variability.<br />

Perhaps the only effort to approach this problem properly<br />

was made by Dr. David Griffiths, who, with U. S. experiment<br />

stations at his disposal, once started an ambitious program of<br />

growing all the Opuntias of the Southwest in identical situations<br />

and of raising seedlings of them all in uniform and differing environments<br />

in order to discover what really are the constant<br />

characters of these cacti. But this sort of a program would take<br />

many years to yield really definite answers, and although Dr.<br />

Griffiths was able to gain some hints in the years he carried it<br />

on, no one continued his fields of Opuntias, and no such attempt<br />

has been made again. We are therefore left to work out indirectly<br />

what the solid characters of these species are, and it is no easy<br />

task.<br />

To the foregoing must be added the fact that some of the<br />

Opuntia species cover huge ranges of territory and within these<br />

huge territories show various different forms not directly due to


Cornyopuntia -><br />

Corynopuntia<br />

genus Opuntia 161<br />

environment. It seems clear, for instance, that the large prickly<br />

pears of the U.S. can only be understood if a few very broad<br />

species are recognized. One of these, Opuntia engelmannii, with<br />

its range running from at least Louisiana—and some would say<br />

even from Florida—to California, would be one of the widest<br />

ranging of any U.S. cactus. And within this huge entity are a<br />

number of varieties which have caused great confusion, as some<br />

have described them as separate species, and others have ignored<br />

them entirely, assuming them to be the mere results of local environments.<br />

It is hard to find agreement as to how to interpret<br />

such a situation, and experimental evidence is so far almost to-<br />

tally lacking.<br />

In this study, when faced with this sort of situation, I have<br />

taken the attitude that no described form can be ignored and<br />

relegated to the synonymy without investigation. So we have<br />

made great efforts to locate and study each one described for<br />

this area, no matter how obscure. And wherever we have found<br />

a form which exists as a population with a definable range,<br />

which does not seem to be due to a local environmental factor,<br />

but which does not seem distinct enough to be clearly a separate<br />

species, I have listed it as a variety. Only by keeping these forms<br />

thus in sight can we keep them available for the biosystematic<br />

studies which are long overdue and which will someday give us<br />

a clearer understanding of the Opuntias.<br />

The genus Opuntia is an old one, and it has been subdivided.<br />

The system for the U. S. members has usually taken the form of<br />

a division into 2 or 3 groups based mostly on the forms of the<br />

stems and partly on spine characters. These groupings were some<br />

of them made very early and the divisions called subgenera or<br />

sections of the genus. The earliest divisions were into subgenus<br />

Cylindropuntia Eng., with stems tending to be cylindrical and<br />

more or less tuberculate, and subgenus Platyopuntia Weber,<br />

having the stems to some degree flattened. Cylindropuntia was<br />

then divided and those members of it which had cylindrical<br />

stems were left under that name, while those which had<br />

more or less club-shaped stems were placed in a subgenus called<br />

variously Clavatopuntia by Fric, Clavatae by Berger, and most<br />

recently Corynopuntia by Knuth. This arrangement would give 3<br />

groups of Opuntias in the United States, and there are several<br />

others in Central and South America.<br />

Little fault can be found with these groupings. They seem<br />

natural groups within the genus. But granting that, we are soon<br />

faced with the old genus problem, since some authors have recently<br />

elevated these groupings, originally sections or subgenera<br />

of the genus Opuntia to the level of genera themselves, thus<br />

doing away with the old genus entirely, or like Earle, limiting<br />

the genus Opuntia to the forms which made up the former subgenus<br />

Platyopuntia.<br />

What are we to do about the newer system which divides this<br />

genus into several genera? Here once more it seems I must make<br />

a decision based almost totally upon my own philosophy and<br />

my concept of a genus. It seems rather clear that there is a natural<br />

grouping into cylindrical and flat-jointed Opuntias, although<br />

a few species are close to intermediates even here. The division<br />

between cylindrical and club-shaped is not as clear. There are<br />

some forms which show an ability to adopt somewhat of either<br />

stem form, and are difficult to place definitely in either. Other<br />

differences which have been searched out between these two<br />

groups—such as spines sheathed or not sheathed—have not<br />

proved too reliable either.<br />

After much study of this problem it seems to me that while<br />

these groupings represent something natural, and have some<br />

significance, they hardly represent essential enough differences<br />

to be the bases for different genera. To use the worn, but it seems<br />

important, analogy from the genus Euphorbia once again, the<br />

stems of Euphorbia obesa, Euphorbia grandicornis, and Euphorbia<br />

prostrata seem every bit as essentially different as do those<br />

of our three Opuntia groups. We have also, in our laboratory,<br />

run a series of chromatographic studies on Opuntias and found<br />

no evidence of chemical groupings to parallel these divisions, as<br />

one would expect if the differences were generic. So we have<br />

concluded that while these groupings may be useful, and may<br />

form the basis for subdividing the old genus Opuntia, they<br />

should probably not supplant it.<br />

I am not concerning myself here with the long lists of series<br />

or sections into which this genus has been divided by various<br />

authors. Each series published so far contradicts the others, and<br />

I have found enough faults in those of Britton and Rose and<br />

also in those of Backeberg, for instance, that I feel such detailed<br />

classification of the group is still premature. We just don’t understand<br />

the genetics of these plants, the way they vary, and the<br />

significance of their various characters well enough yet. Perhaps,<br />

if their many forms are not lost sight of, after experimentation<br />

is carried on it will then be possible to set up meaningful<br />

series of the Opuntias.<br />

Those hardy enough, and possessing enough of the true botanist’s<br />

curiosity which will let him pass up no plant, however<br />

thorny, will find a huge opportunity in the study of the Opuntias.<br />

In sheer numbers of their populations and extents of their<br />

ranges they excel. Nor does one have the disappointing experience<br />

with these, that he all too often has with the other cacti, of<br />

traveling to a spot where a species is supposed to grow profusely<br />

only to find that some collector or dealer has stripped the area<br />

bare of every specimen which once was there. So few bother to<br />

bring home Opuntias that one can travel through country most<br />

ruthlessly sacked of cacti and still find the Opuntias untouched.<br />

Even the inroads of agriculture have not adversely affected the<br />

majority of them. Root-plowing, chaining, and most of the<br />

other practices used to clear the range of the chaparral have usually,<br />

while exterminating the other cacti, merely torn the Opuntias<br />

apart and distributed their stems widely, each one of which<br />

then rooted, and the Opuntia population has, therefore, multiplied.<br />

So the opportunity for studying them is perhaps greater<br />

today in our area than ever before. Only a few obscure species<br />

limited to very small ranges have been reduced by clearing of<br />

fields and agricultural practices, although in the future the pic-


162 cacti of the southwest<br />

ture will probably be very different. All the big guns of the<br />

chemical industry are being aimed at the chaparral, and the<br />

mass spraying of the range with some of the newer herbicides,<br />

barely started now, clearly could reduce these plants to a vanishing<br />

point. But the student of the Opuntias can take heart in<br />

knowing that, tough as these plants are, they will be among the<br />

last wild plants to go, and he will be one of the last students of<br />

wild plants to survive.<br />

The flat-stemmed Opuntias are known almost universally as<br />

prickly pears. Tough and thorny as they are, these are the main<br />

food-producing cacti. Tons of their stems are fed to cattle, particularly<br />

in Texas and Mexico, where sophisticated methods are<br />

used to burn the spines off so that the flesh is available to the<br />

animals. And these plants are used extensively for human consumption<br />

as well. The young pads, before their tissues have<br />

hardened or their spines been produced are relished. Called nopalitos,<br />

these are eaten widely in Mexico, where they are usually<br />

breaded with cornmeal and fried, and in Texas, where they<br />

are more often boiled and added to a sort of omelet. In the<br />

spring they are often on sale at markets in San Antonio and the<br />

rest of south Texas, and in San Antonio they are served in some<br />

Mexican restaurants, as well as being sold canned in some supermarkets.<br />

But the most widely used part of the Opuntia is probably<br />

the fruit, called tuna. Certain species producing large,<br />

sweet fruits are cultivated as crops in Mexico, and this was one<br />

of the reasons for these plants being introduced into some other<br />

parts of the world where they have become such pests.<br />

The large, upright, cylindrical-stemmed Opuntias are known<br />

collectively as chollas. I do not know of any use of these chollas<br />

for food. They are far too woody and tough, and their fruits<br />

are not edible. But certain species of them make very large and<br />

beautiful bushes, and are widely grown as ornamentals. Almost<br />

any inhabited area of the Southwest will have some specimens<br />

of these species planted just for their beauty. They may be prized<br />

highly in places where almost no other shrubs or trees will grow.<br />

And anyone who has looked over the typical curio store knows<br />

well the lamp bases and other trinkets made of the curious, reticulated<br />

wood of these chollas.<br />

So, whether regarded as friends or enemies, the Opuntias are<br />

easily some of the most strange, fascinating, and challenging of<br />

plants.<br />

KEY TO <strong>THE</strong> OPUNTIAS<br />

1a. Joints flattened; spines not covered with papery sheaths (Platyopuntias)—2.<br />

2a. Plants upright and bushy, 2 to 7 feet tall, sometimes spreading<br />

and forming thickets, but with no branches prostrate, and with<br />

upright branches attaining a length of three pads or more—3.<br />

3a. Areoles 3/4 to 21/2 inches apart, with the majority on any pad<br />

one inch or more apart—4.<br />

4. Possessing spines—5.<br />

5a. Spines all round —O. Stricta (in part).<br />

5b. At least one spine of each areole flattened—6.<br />

6a. Pads conspicuously tuberculate by raised areoles; fruits<br />

spiny, with I to 5 rigid spines up to 1 inch long on at<br />

least each upper areole, and these fruits becoming dry<br />

when ripe —O. spinosibacca.<br />

6b, Pads not markedly tuberculate; fruits naked or nearly<br />

so—7.<br />

7a. Spines when mature whitish, yellow, red-brown, or<br />

mottled, but with no part of any spine black—8.<br />

8a. Pads circular or variously shaped, but not over<br />

twice as long as broad—9.<br />

9a. Seeds 1/16 to 3/16 of an inch (11/2–41/2 millimeters)<br />

in diameter—10.<br />

10a. Spines on old areoles on old pads not increasing<br />

in number beyond the normal 10 or so of<br />

mature pads—11.<br />

11a. Areoles and glochids normal in appearance<br />

and not exaggerated in development when<br />

old—12.<br />

12a. Fruits spherical, oval, or broadly pear-<br />

shaped, with little or no constriction below,<br />

with the umbilicus not constricted<br />

but flat or nearly so and as broad or<br />

nearly as broad as the widest part of the<br />

fruit; flowers yellow to red, but never<br />

yellow with red centers—13.<br />

13a. Fruits 2 to 31/2 inches long, deep burgundy<br />

in color when ripe; seeds 1/8 of<br />

an inch or a little more (3–4 millimeters)<br />

in diameter; stigmas green—<br />

14.<br />

l4a. Spines deep brown at their bases,<br />

becoming white or whitish toward<br />

the tips; leaves 1/4 to 3/8 of an inch<br />

long; fruits edible<br />

—O. engelmannii var. engelmannii.<br />

14b. Spines and glochids all bright yellow<br />

or yellow with the bases to<br />

sometimes all but the tips red-<br />

brown; leaves 1/2 to 1/2 inch long;<br />

fruits not pleasant to the taste<br />

—O. engelmannii var. texana.<br />

13b. Fruits 1 to 2 inches long, bright red to<br />

purplish-red when ripe; seeds 1/16 to<br />

3/16 of an inch (11/2–41/2 millimeters)<br />

in diameter; stigmas white to greenish<br />

—15.<br />

15a. Spines heavy and rigid, the longest<br />

spine usually about 11/2 inches long<br />

and only rarely reaching 21/2<br />

inches; fruits purplish-red—16.<br />

16a. Fruits spherical or nearly so<br />

with no constriction below and<br />

always spineless; seeds about 3/16


genus Opuntia 163<br />

of an inch (4–41/2 millimeters)<br />

in diameter; spines red-brown<br />

or with at least the bases red-<br />

brown; flowers small with dark<br />

green stigmas —O. engelmannii<br />

var. cyclodes.<br />

16b. Fruits not spherical, but broadly<br />

egg-shaped with some slight<br />

constriction below and having<br />

numerous glochids and often a<br />

few spines 1/2 to 5/8 of an inch<br />

long on them; seeds extremely<br />

small, being 1/16 of an inch or<br />

slightly more (11/2–21/2 millimeters)<br />

in diameter; spines entirely<br />

yellow; flowers large with<br />

stigmas light greenish-white<br />

—O. engelmannii var. alta.<br />

15b. Spines slender and long, as well as<br />

somewhat flexible, the main spines<br />

being 2 to 3 inches long; fruits not<br />

so purplish, but more bright red;<br />

stigmas either dark green or white<br />

—17.<br />

17a. Pads thick and light or yellowish-green;<br />

main central spines<br />

all sharply deflexed and very<br />

flexible and 1/2 to 3 inches long;<br />

fruit broad egg-shaped to club-<br />

shaped, with some constriction<br />

below; seeds about 1/8 of an inch<br />

(3–31/2 millimeters) in diameter;<br />

stigmas dark green<br />

—O. engelmannii<br />

var. flexispina.<br />

17b. Pads thin and blue-green in<br />

color; main spines porrect, 2 to<br />

3 inches long and conspicuously<br />

longer than the other spines;<br />

fruits spherical to oval with no<br />

constriction below; seeds between<br />

1/16 and 1/8 of an inch<br />

(2–3 millimeters) in diameter;<br />

stigmas white —O. engelmannii<br />

var. cacanapa.<br />

12b. Fruits elongated and club-shaped, with<br />

the base definitely constricted and the<br />

umbilicus also constricted, much narrower<br />

than the widest part of the fruit<br />

and deeply to very deeply pitted; stig-<br />

mas yellowish—18.<br />

18a. Spines bright yellow or yellow with<br />

light brown bases; flowers all yellow;<br />

fruits purplish or plum in color;<br />

seeds 3/16 of an inch or less (31/2–41/2<br />

millimeters) in diameter<br />

—O. tardospina.<br />

18b. Spines brown below with whitish<br />

above, never bright yellow; flowers<br />

orange-yellow with red centers;<br />

fruits bright scarlet-red in color;<br />

seeds n/is of an inch or more (4–5<br />

millimeters) in diameter<br />

—O. phaeacantha var. major<br />

(in part).<br />

11b. Areoles and glochids exaggerated in development<br />

when old—19.<br />

19a. Areoles enlarging to 1/2 inch in diameter<br />

on old stems and bulging outward<br />

to form a sort of cylindrical projection<br />

to 1/2 inch high, with a compact tuft of<br />

short glochids on the summit of it; pads<br />

large, mostly round, and blue-green in<br />

color —O. engelmannii var. dulcis.<br />

19b. Areoles large and bulging outward<br />

noticeably, with glochids very long<br />

and scattered loosely throughout the<br />

areole, forming starlike clusters almost<br />

covering the surfaces of the mature<br />

pads; pads smaller and more obovate<br />

and with the color more dull dark<br />

green than is typical for the species<br />

—O. engelmannii var. aciculata<br />

(in part).<br />

10b. Spines on old areoles on old trunks increasing<br />

greatly to 20 or 30 per areole, covering<br />

the older parts of the plants with a com-<br />

plete covering of spines —O. chlorotica.<br />

9b. Seeds more than 3/16 of an inch (41/2 milli-<br />

meters or more) in greatest diameter<br />

—O. phaeacantha var. camanchica<br />

(when unusually large and ascending).<br />

8b. Pads greatly elongated so that at least some of<br />

them on a normal plant are at least twice as long<br />

as they are broad —O. engelmannii<br />

var. linguiformis.<br />

7b. Spines when growing black or bright orange, when<br />

mature black or dark blackish-brown, at least at<br />

the base—20.<br />

20a. Fruits 3/4 to 11/2 inches long, oval or ovate, with<br />

no noticeable constriction below and the umbilicus<br />

rather deeply pitted, yellowish to scarlet<br />

when ripe; seeds 1/8 to 3/16 of an inch (3–41/2<br />

millimeters) in diameter; spines heavy, rigid,<br />

very angular, and 1 to 6 per areole, found on<br />

most areoles, but to only 2 inches or less in<br />

length; glochids becoming long and average in<br />

number on edge areoles; pads thin and blue-<br />

green to yellow-green in color, not becoming<br />

reddish —O. phaeacantha var. nigricans.<br />

20b. Fruits 11/4 to 2 inches long, elongated oval or<br />

egg-shaped, with some slight constriction below<br />

and a shallowly pitted umbilicus; seeds more<br />

than 3/16 of an inch (about 5 millimeters) in<br />

diameter; spines heavy, rigid, almost round to


164 cacti of the southwest<br />

somewhat flattened, 2 to 4 per areole in only<br />

the upper areoles, but to 3 inches long; glochids<br />

few at first, becoming very many and very long<br />

in the edge areoles; pads thick and pale green,<br />

often reddish in color or spotted with reddish<br />

around the areoles —O. phaeacantha<br />

var. brunnea.<br />

4b. Spineless—21.<br />

21a. Pads elongated obovate to elliptic or even spindle-<br />

shaped; surface of pads smooth and shining, not glau-<br />

cous; leaves only 1/8 to 1/4 of an inch long; ovaries<br />

narrow and elongated; fruits elongated, pear-shaped,<br />

constricted at the base and with deeply pitted um-<br />

bilicus —O. Stricta (in part).<br />

21b. Pads round or oval or even broader than long, surface<br />

dull and glaucous; leaves 3/8 to 5/8 of an inch long;<br />

ovaries short and stout; fruits spherical to broadly<br />

pear-shaped, with little or no basal Constriction and<br />

umbilicus flat or only slightly concave—22.<br />

22a. Surface blue-green; areoles small; glochids very few<br />

and short —O. engelmannii var. subarmata.<br />

22b. Surface bright green or dark green, but not blue-<br />

green; areoles enlarged and bulging; glochids very<br />

long in loose, spreading clusters —O. engelmannii<br />

var. aciculata (when spineless).<br />

3b. Areoles 1/4 to 11/4 inches apart, with the majority on any pad<br />

less than 1 inch apart—23.<br />

23a. Spineless—24.<br />

24a. Pads pubescent; glochids very many but very short and<br />

minute, hardly exceeding the wool in the bulging areoles;<br />

fruits remaining greenish and rather dry when<br />

ripe —O. rufida.<br />

24b. Pads not pubescent; glochids few but larger; fruits<br />

bright red or orange-red and fleshy when ripe<br />

—O. macrocentra (in part).<br />

23b. Spines present—25.<br />

25a. Spines black or brown, sometimes with gray but never<br />

with yellow coloring, to 4 per areole in a few of the<br />

upper areoles only, 2 to 5 inches long, somewhat flexible,<br />

and at least one spine flattened; pads usually purplish<br />

in color; seeds 3/16 to more than 1/4 of an inch<br />

(41/2–7 millimeters) in diameter —O. macrocentra<br />

(in part).<br />

25b. Spines yellow in their outer zones, with black or brown<br />

bases, not flexible, from 3/4 to 4 inches, but usually less<br />

than 2 inches long; pads sometimes reddish, but not<br />

distinctly purplish in color; seeds around 1/8 of an inch<br />

(21/2–4 millimeters) in diameter—26.<br />

26a. Spines 1 to 3 on a few upper areoles only, 1/2 to 2<br />

inches, but said to have attained 4 inches long; bases<br />

of spines brown; spines round; pads usually with<br />

reddish coloring; glochids many but very short<br />

—O. gosseliniana var. santa-rita.<br />

26b. Spines 3 to 16 on many or all areoles; never over<br />

13/4 inches long; bases black or dark red-brown, at<br />

least when young; pads yellow-green without purple<br />

or reddish coloring; glochids many and long—27.<br />

27a. Spines 6 to as many as 15, consisting of 1 to 5 rigid<br />

main spines which are either round or flattened<br />

and 3/16 to 3/4 of an inch long, plus up to 10 lower,<br />

bristle-like spines 3/16 to 3/4 of an inch long<br />

—O. strigil.<br />

27b. Spines 3 to 6, consisting of about 2 rigid, round<br />

main ones 1/2 to 11/4 inches long and 2 or 3 lower,<br />

bristle-like spines 3/16 to 5/8 of an inch long<br />

—O. atrispina.<br />

2b. Plants growing prostrate or sprawling, or if ascending, never<br />

standing upright over 2 or 3 pads or 12 to 20 inches tall—28.<br />

28a. Pads 6 to 10 inches long in maximum size—29.<br />

29a. Areoles 3/8 to 7/8 of an inch apart; fruits more or less<br />

spiny and becoming dry when ripe—30.<br />

30a. Spines 1 to 10 per areole; fruits 11/2 to 2 inches long<br />

and broadly club-shaped, spiny above or spineless—31.<br />

31a. Spines 1 to 4 in only the upper areoles of the pads<br />

—O. rhodantha var. rhodantha (in part).<br />

31b. Spines 4 to 10 in all or nearly all areoles of the pads<br />

—O. rhodantha var. spinosior.<br />

30b. Spines 10 to 15 in all or nearly all areoles; fruits 7/8 to<br />

11/4 inches long and obovate to broadly club-shaped,<br />

spiny —O. hystricina (in part).<br />

29b. Areoles 3/4 to 2 inches apart; fruits remaining fleshy and<br />

spineless—32.<br />

32a. At least some areoles with more than one spine; pads<br />

almost circular to broad egg-shaped; seeds thin or<br />

average in thickness—33.<br />

33a. Spines 1 to 5, slender to medium thickness, round or<br />

nearly so with no lower bristle-like spines present<br />

—34.<br />

34a. Fruits 11/2 to 31/2 inches long, club-shaped, with<br />

pronounced constriction at the base and with the<br />

top narrow and deeply pitted; seeds about 1/8 of<br />

an inch (21/8–4 millimeters) in diameter, with nar-<br />

row rims and the body of the seed rather thick<br />

—O. leptocarpa.<br />

34b. Fruits 11/4 to 23/8 inches long, broadly club-shaped,<br />

with the base constricted and the umbilicus somewhat<br />

pitted; seeds about 3/16 of an inch (4–5 millimeters)<br />

in diameter —O. phaeacantha var. major<br />

(in part, when stunted).<br />

33b. Spines 1 to 8, main spines heavy to very heavy and<br />

flattened, often with lower bristle-like spines present;<br />

fruits 1 to 2 inches long, oval to very broadly egg-<br />

shaped, with little or no constriction at the base and<br />

with broad, flat, or shallowly pitted umbilicus; seeds<br />

3/16 to 1/4 of an inch (41/2–6 millimeters) in diameter,<br />

with wide rims —O. phaeacantha var. camanchica<br />

(in part).<br />

32b. Spineless or with only one spine in a few areoles; pads<br />

elongated egg-shaped to spindle-shaped; seeds very<br />

thick —O. compressa var. allairei.<br />

28b. Pads 2 to 6 inches long—35.<br />

35a. At least some areoles of the pad more than 11/4 inches<br />

apart—this is a stunted, abnormally small specimen of<br />

some larger species, and it will probably be impossible to<br />

identify it with this key.<br />

35b. Areoles 3/16 to 11/4 inches apart—36.


erinaceae -><br />

erinacea<br />

genus Opuntia 165<br />

36a. Pads solidly attached to each other and not detaching<br />

to come away from the plant at a touch—37.<br />

37a. Spines to 5 or more per areole and found on 1/2 or<br />

more areoles of mature pads; stigmas bright green<br />

—38.<br />

38a. Fruits remaining fleshy when ripe and oval or oblong<br />

in shape; seeds not thick for their diameters<br />

—39.<br />

39a. Pads thin with definitely constricted or even<br />

attenuated bases; spines round or nearly so,<br />

slender and flexible and mostly deflexed; glochids<br />

red-brown and short; seeds slightly over<br />

1/8 to 1/4 inch (31/2–6 millimeters) in diameter,<br />

with narrow to medium rims —O. phaeacantha<br />

var. tenuispina.<br />

39b. Pads thickish, round or nearly so with no definite<br />

constrictions at their bases; spines at least<br />

somewhat flattened, medium to thick and always<br />

rigid, glochids long and yellow or light<br />

brown; seeds 3/16 to just over 1/4 inch (41/2–61/2<br />

millimeters) in diameter, with wide rims<br />

—O. cymochila.<br />

38b. Fruits dry when ripe, various in shape—40.<br />

40a. Pads elongated oval to clavate, less than half as<br />

broad as they are long, and very thick; roots<br />

stolon-like; seeds about 1/4 of an inch (6–7 millimeters)<br />

in diameter —O. arenaria.<br />

40b. Pads round or oval to broadly egg-shaped or<br />

oblong, always more than half as broad as they<br />

are long and not unusually thick; roots normally<br />

fibrous—41.<br />

41a. Pads more or less prostrate, sometimes with<br />

young pads ascending, but with older ones<br />

reclining; at least main spines rigid and stout<br />

—42.<br />

42a. Surface of pads noticeably tuberculate and<br />

often wrinkled; glochids few to average<br />

and short; main spines 0 to 5 per areole and<br />

porrect to deflexed, 2 inches or less long,<br />

all rigid or often becoming greatly elongated,<br />

soft, flexible, and hairlike, particu-<br />

larly toward the bases of the pads<br />

—O. polyacantha (in part).<br />

42b. Surface of pads not noticeably tuberculate<br />

or wrinkled; glochids many and often long;<br />

main spines 1 to 8 per areole, spreading,<br />

heavy, but moderately flexible, and to 4<br />

inches long —O. hystricina (in part).<br />

41b. Pads all erect or ascending, with not even old<br />

pads reclining; spines slender, to 4 inches long,<br />

and moderately flexible —O. erinacea.<br />

37b. Spines 0 to 4 or rarely 5 per areole on only the upper<br />

edge areoles or at most on those of the upper one-<br />

half of the pad—43.<br />

43a. Fruit remaining fleshy; stigmas yellow or whitish<br />

—44.<br />

44a. Fruits egg-shaped to club-shaped with the bases<br />

noticeably constricted—45.<br />

45a. Pads ascending from a central trunklet, blue-<br />

green or gray-green in color, usually with<br />

some purplish blotching around the areoles;<br />

root a large central taproot 1/2 to 11/2 inches in<br />

diameter and 12 inches or longer, not tuber-<br />

like enlargements on otherwise fibrous roots;<br />

flowers deep red to purple—46.<br />

46a. Plants standing 6 to 12 inches tall; pads<br />

medium thickness and to 4 or 5 inches long,<br />

glabrous, blue-green in color, the surface<br />

flat without raised areoles; glochids numerous<br />

and yellow in color; fruits 11/2 inches<br />

or more long; seeds just over 3/16 to 1/4 of<br />

an inch (5–6 millimeters) in diameter<br />

—O. pottsii.<br />

46b. Plants standing less than 6 inches tall; pads<br />

thin and to only 21/2 inches long, glaucous<br />

gray-green in color, the surface tuberculate<br />

by raised areoles; glochids numerous and<br />

bright red-brown in color; fruits 5/8 to 1<br />

inch long. Seeds 1/8 of an inch or a little<br />

more (3–4 millimeters) in diameter<br />

—O. plumbea.<br />

45b. Pads prostrate or sprawling, not ascending<br />

from a central trunklet, thick, yellow-gray of<br />

deep green, but not blue-green in color; roots<br />

fibrous or fibrous with tubers on them, but<br />

with no central taproot—47.<br />

47a. Pads gray-green or yellow-green and glaucous—48.<br />

48a. Pads to 51/2 inches long; spines to 21/4<br />

inches long and slender to medium in<br />

thickness; glochids greenish-yellow to<br />

straw or light brown, average in number,<br />

and to 1/4 of an inch or less long; flowers<br />

large and yellow in color; fruits 11/2 to<br />

21/2 inches long; seeds 3/16 to 1/4 of an<br />

inch (5 millimeters plus) in diameter<br />

—O. compressa var. stenochila.<br />

48b. Pads to 4 inches long; spines to 23/4<br />

inches long and thick to very thick; glochids<br />

bright yellow, very numerous and<br />

conspicuous, and to 1/2 inch long on old<br />

pads; flowers small and reddish in color;<br />

fruits 3/4 to 1 inch long and very slender;<br />

seeds 1/8 of an inch or a little more (about<br />

31/2 millimeters) in diameter —O. ballii.<br />

47b. Pads deep green, surface shining—49.<br />

49a. Plant entirely prostrate, with all pads reclining<br />

by the end of their first season;<br />

pads very thick, very wrinkled when dehydrated,<br />

but not tuberculate by elevations<br />

at the areoles; spines 0 to 3 per<br />

areole on the upper areoles and 1 inch or<br />

less long, heavy, straight, and round;<br />

inner petals 10 to 14 in number<br />

—O. compressa var. humifusa.<br />

49b. Plants sprawling, with very old pads


sparcely -> sparsely<br />

166 cacti of the southwest<br />

sometimes reclining, but most pads ascending,<br />

pads not so thick as the last,<br />

somewhat to extremely tuberculate by<br />

raised areoles; spines 0 to 5, often some<br />

of them over 1 inch long and often the<br />

main spine somewhat flattened; inner<br />

petals 5 to 9 in number—50.<br />

50a. Spineless; flowers very large (4 to 5<br />

inches in diameter); fruits 2 inches or<br />

more long; pads very tuberculate<br />

—O. compressa var. grandiflora.<br />

50b. Spines 1 to 5; flowers average size (2<br />

to 3 inches in diameter); fruits 1 to 2<br />

inches long—51.<br />

51a. Seeds about 3/16 of an inch (4–5<br />

millimeters) in diameter—52.<br />

52a. Glochids brown or yellowish; surface<br />

only somewhat tuberculate;<br />

spines 1 to 5; inner petals 7 to 9<br />

in number —O. compressa<br />

var. macrorhiza.<br />

52b. Glochids bright red-brown; surface<br />

very tuberculate; spines 1 to<br />

3; inner petals 5 to 7<br />

—O. compressa var. fusco-atra.<br />

51b. Seeds only around 1/16 of an inch<br />

(11/2–2 millimeters) in diameter<br />

—O. compressa var. microsperma.<br />

44b. Fruits oval with no constriction below<br />

—O. phaeacantha var. camanchica<br />

(when stunted).<br />

43b. Fruits becoming dry when ripe; flowers yellow,<br />

pink, or rose; stigmas bright green—53.<br />

53a. Stigma lobes 4 to 6; fruits 3/4 to 13/8 inches long,<br />

spineless, and without constriction below—54.<br />

54a. Fruits spherical to oval; seeds slightly over<br />

3/16 of an inch (5 millimeters) in diameter,<br />

thickish, with narrow, acute rims<br />

—O. sphaerocarpa.<br />

54b. Fruits oblong; seeds from 1/4 to just over 1/16<br />

of an inch (6–8 millimeters) in diameter, flat<br />

and not thick for the size, with wide rims<br />

—O. polyacantha<br />

(in part, when sparcely-spined).<br />

53b. Stigma lobes 8 to 12; fruits 11/2 to 1/4 inches<br />

long, broad club-shaped with some tapering<br />

constriction below; upper areoles spiny or else<br />

entirely spineless —O. rhodantha<br />

var. rhodantha (in part).<br />

36b. Pads loosely attached so that they separate at a touch;<br />

pads small, thick, and almost cylindrical when young<br />

—55.<br />

55a. Fruits fleshy, elongated, and club-shaped, 11/4 inches<br />

or more long; seeds just under 3/16 of an inch (4<br />

millimeters) in diameter; pads to 41/2 inches long,<br />

stigmas yellowish —O. drummondii.<br />

55b. Fruits dry, oval or broadly egg-shaped, 1/2 to 1 inch<br />

long; seeds just over 5/16 of an inch (5 millimeters)<br />

in diameter; stigma lobes bright green —O. fragilis.<br />

1b. Joints not flattened but cylindrical or club-shaped; spines with<br />

either conspicuous or rudimentary papery sheaths—56.<br />

56a. Joints club-shaped or egg-shaped and very tuberculate; plants<br />

prostrate or ascending but 12 inches or less in height; spine<br />

sheaths rudimentary on tips of immature spines only and<br />

usually not present on adult spines (Corynopuntias)—57.<br />

Cornyopuntias -><br />

Corynopuntias<br />

57a. Joints elongated club-shaped, prostrate to ascending, usually<br />

curved; spines to more than 11/4 inches long—58.<br />

58a. Main central spine very heavy, flat, wide, and roughly<br />

cross-striated—59.<br />

59a. Joints 31/2 to 6 inches long; tubercles 1 to 13/4 inches<br />

long —O. stanlyi.<br />

59b. Joints 1 to 3 inches long; tubercles 1/2 to 3/4 of an inch<br />

long —O. schottii.<br />

58b. Main spines round or nearly so, slender to medium thickness,<br />

not striated —O. grahamii.<br />

57b. Joints short club-shaped or egg-shaped; main spines to only<br />

11/4 inch long —O. clavata.<br />

56b. Joints cylindrical; plants upright or ascending, never prostrate,<br />

bushy, and 1 to 12 feet tall; spine sheaths conspicuous,<br />

at least on newly matured main spines (Cylindropuntias)—60.<br />

60a. Width of current year’s joints 3/4 to 2 inches—61.<br />

61a. Tubercles 3/4 to 11/2 inches long; color deep green at all<br />

times—62.<br />

62a. Fruits globose and about 1 inch in diameter, always<br />

tuberculate—63.<br />

63a. Main spines 3/4 to 11/4 inches long; spine sheaths<br />

whitish or straw-colored; fruits spineless when mature—64.<br />

64a. Flowers purple or reddish (reported as rarely yellowish<br />

or white); growth treelike and 3 to 8 feet<br />

tall —O. imbricata var. arborescens.<br />

64b. Flowers pale greenish with lavender shading; a<br />

small bush 1 to 3 feet tall —O. imbricata<br />

var. viridiflora (in part).<br />

63b. Main spines 2 to 21/2 inches long, with glistening<br />

silvery sheaths; flowers greenish yellow; growth low,<br />

1 to 2 feet tall; fruits with sheathed spines to 1 inch<br />

long —O. tunicata.<br />

62b. Fruits obovate to globose, 1 to 2 inches in diameter,<br />

becoming nearly or entirely smooth when ripe; main<br />

spines to no more than 5/8 of an inch long<br />

—O. imbricata var. vexans.<br />

61a. Tubercles 3/8 to 3/4 inch long—65.<br />

65a. A large bush 3 to 12 feet tall; spines 6 to 25 in number,<br />

mostly 3/8 of an inch or less long and only rarely to 5/8<br />

of an inch, spine sheaths inconspicuous and soon falling<br />

off; color of plant surface often purplish in severe conditions<br />

—O. spinosior.<br />

65b. Low spreading or mat-forming bushes 1 to 2 feet tall;<br />

spines 3 to 12 in number, maximum spine length 3/4 to<br />

11/4 inches, spine sheaths loose, persistent, and conspicuous;<br />

color of plant surface light green —O. whipplei<br />

(in part).<br />

60b. Width of current year’s joints 3/4 of an inch or less—66.<br />

66a. Plants low-growing, 1 to 3 feet tall; width of current


genus Opuntia 167<br />

year’s joints 3/8 to 3/4 of an inch, with distinct tubercles<br />

—67.<br />

67a. Largest spines 11/4 inches or less long (usually less than<br />

1 inch)—68.<br />

68a. Erect, more or less trunked plants; joints strictly<br />

cylindrical; spine sheaths not conspicuous and shining;<br />

flower greenish-purple or lavender with reddish<br />

stigma lobes —O. imbricata var. viridiflora (in part).<br />

68b. Ascending or spreading, but without real trunks,<br />

often mat-forming plants; joints slightly clavate by<br />

being constricted at the bases. Spine sheaths conspicuous<br />

and shining; flowers pale yellow; stigmas green-<br />

ish —O. whipplei (in part).<br />

67b. Largest spines 11/2 to 2 inches long; tubercles 5/8 to 1<br />

inch long —O. davisii.<br />

66b. Plants bushy and growing 3 to 6 feet or more tall; width<br />

of current year’s joints 1/8 to 1/2 inch; tubercles indistinct<br />

or sometimes actually absent—69.<br />

69a. Joints 1/4 to 1/2 inch thick, with tubercles indistinct to<br />

distinct, but always present—70.<br />

70a. Joints 3/16 to 1/2 inch in diameter; flowers greenish-<br />

purple and 1 to 11/4 inches in diameter; fruit red and<br />

with indistinct tubercles or else smooth when ripe,<br />

3/4 to 1 inch long; spines to 1 inch long —O. kleiniae.<br />

70b. Joints 3/16 to 3/8 of an inch in diameter; flowers unknown;<br />

fruits yellowish and distinctly tuberculate<br />

when ripe, 9/16 to 3/4 of an inch long; spines 1 to 21/2<br />

inches long —O. vaginata.<br />

69b. Joints 1/8 to 1/4 of an inch thick, with tubercles at best<br />

indistinct and often absent; flowers yellow or greenish-<br />

yellow and 1/2 to 7/8 of an inch in diameter; fruits red<br />

or orange-red and smooth when ripe —O. leptocaulis.<br />

Opuntia stricta Haw.<br />

“Pest Pear”<br />

Description Plate 44<br />

stems: An erect plant, but much branched and diffuse instead<br />

of treelike. Usually 2 to 3 feet tall, having elongated, ovate to<br />

elliptical or even spindle-shaped pads usually 6 to 9 inches<br />

long, but sometimes to 14 inches in length. These pads are of<br />

medium thickness, bluish-green and glabrous when young, but<br />

becoming light green and somewhat glaucous when old. Its<br />

leaves are 1/8 to 3/16 of an inch long.<br />

areoles: Very elongated in young pads, becoming oval when<br />

older. They are 1/4 of an inch or a little more in length, and<br />

situated 11/4 to 21/2 inches apart.<br />

spines: Variable. These plants are often entirely spineless, but<br />

some specimens have 1 to 3 spines per areole. When present<br />

the spines are porrect or spreading, 3/8 to 13/4 inches long,<br />

straight, stiff, and of medium thickness. They are round in<br />

cross-section, yellow in color, sometimes becoming slightly<br />

mottled with brown, but not having brown bases.<br />

glochids: Few and very short on young pads, becoming<br />

more with age, but never becoming conspicuous. Remaining<br />

comparatively short, growing to a maximum of about 3/8 of<br />

an inch long. They are yellow or straw-colored.<br />

flowers: About 3 to 4 inches in diameter, clear lemon-yellow<br />

in color. The ovary is slender club-shaped and 11/2 to 3 inches<br />

long, with a few glochids in its areoles. The perianth segments<br />

are pointed spatulate, greenish at their bases, and clear<br />

yellow above. The filaments are greenish below, becoming<br />

yellow above, and the anthers are yellow. The short, greenishyellow<br />

style has 6 to 8 fat, white or very pale greenish-white<br />

stigma lobes tipping it.<br />

fruits: Light pinkish-red to rather bright carmine-red when<br />

ripe. In shape elongated club-shaped and 11/2 to about 23/4<br />

inches long by 3/4 to 1 inch thick at the broadest upper part.<br />

The umbilicus on the summit of the fruit is deeply pitted. The<br />

flesh is sweet and edible. The seeds are about 1/8 of an inch<br />

(21/2–3 millimeters) in diameter, thin, regular, and with very<br />

narrow but fairly thick rims.<br />

Range. This is primarily a Caribbean species, found on Cuba,<br />

Haiti, and the West Indies, but widely spread to South America<br />

and Australia. It is found in the United States in south Florida,<br />

at Houma, Louisiana, and at the entrance to Galveston Bay.<br />

Remarks. This was one of the earliest of our Opuntias to be<br />

studied. It was first described under the name, Cactus strictus<br />

in 1803, and then again as Opuntia stricta in 1812, both by<br />

Haworth. It has often been called Opuntia inermis. This is because<br />

of a picture, apparently of this plant, which was published<br />

by De Candolle in 1799 labeled Cactus opuntia inermis, but<br />

this is not generally taken to have constituted an official description.<br />

The cactus grows widely in the Caribbean Islands and the<br />

West Indies, which seem to be its native location, but it has been<br />

cultivated on the west coast of South America, and was taken to<br />

Australia. It is the cactus which ran wild over millions of acres<br />

of New South Wales and Queensland and became perhaps the<br />

most famous of plant invaders in modern times. There it took<br />

over huge tracts of both agricultural and grazing lands, making<br />

them unfit for any use, and earned its designation as the pest<br />

pear. Huge programs were instituted to control it, and it has<br />

more recently been brought under a measure of control in Australia.<br />

It is most fortunate that this cactus has not been so successful<br />

in the U.S. It has been found at scattered locations in Florida,<br />

at one location at Houma, Louisiana, and at one location in<br />

Texas, this being at the lower end of Galveston Bay. It is not<br />

known whether it was introduced to these places in modern<br />

times or not, but the places where it has been found, at least in<br />

Louisiana and Texas, are both precisely the places where early<br />

shipping came ashore, and thus it looks like the cactus was<br />

brought, either knowingly or as a stowaway, on early boats


168 cacti of the southwest<br />

from its Caribbean home. Anyway, the plant did not spread<br />

and invade to any extent anywhere in the U. S., and in fact it<br />

seems to have practically failed here. It has not been re-collected<br />

in many years in Louisiana, and can now be found only on Galveston<br />

Island and the Bolivar Peninsula in Texas.<br />

The Galveston Island specimens, which comprise the only pop-<br />

ulation of other than isolated individuals we know of in our<br />

area, is found as a solid row of thickets along the top of the<br />

high sea-wall embankment between the city of Galveston and<br />

Fort Point at the north end of the island. Here it grows near<br />

the massive concrete bunkers and gun emplacements thrown up<br />

to guard the entrance to Galveston Bay, and looking at it<br />

ranged near the ruins of these fortifications it is easy to imagine<br />

that it might have been planted there by the defenders as one<br />

more barrier to deter invaders. It would at least make the route<br />

of infiltrators more difficult. It does seem strange that anyone<br />

planting such a barrier would use this cactus instead of the<br />

native, larger, and much more spiny Opuntia engelmannii var.<br />

texana which must have been growing nearby, as it is today, but<br />

the defenders probably knew little about the country beyond<br />

their island and might well have brought the pest pear they had<br />

seen in the Caribbean here for this use.<br />

Opuntia stricta must be compared carefully with two varieties<br />

of O. engelmannii which grow on the eastern Texas Coast,<br />

and with which it is often confused, variety texana and variety<br />

alta. When they are all in flower or fruit these three are easily told<br />

apart. O. stricta is distinguished from both of the others by its<br />

elongated, club-shaped ovary, since that of the others is obovate<br />

or inverted cone-shaped and is usually as broad or nearly as<br />

broad as it is long, and also by the resulting fruits which in O.<br />

stricta are elongated club-shaped with deeply pitted summits<br />

and light red or carmine-red in color, while those of the O. engelmannii<br />

varieties are broad obovate to almost spherical with<br />

the umbilici flat or practically so and the color deep purplish or<br />

burgundy-red. The fruits of O. stricta are sweet and edible,<br />

which also distinguishes it from O. engelmannii var. texana,<br />

whose fruits are insipid and unpleasant, but this will not distinguish<br />

it from O. engelmannii var. alta, whose more rounded<br />

fruits are also sweet.<br />

The most clear difference between O. stricta and these other<br />

two cacti which can be seen when only vegetative characters<br />

are present, is the elongated ovate, elliptical, or spindle-shaped<br />

pads, which contrast rather clearly with the circular to very<br />

broadly ovate pads of all O. engelmannii forms in normal<br />

growth. O. stricta also has many fewer spines, and these always<br />

completely round, while at least the main spines of any normal<br />

O. engelmannii form are flattened at least at the bases.<br />

O. stricta is admittedly close to O. engelmannii var. alta,<br />

which grows profusely almost all along the Texas coast, but<br />

careful observation can separate them. We have been interested<br />

to find in our laboratory that O. stricta presents a basically different<br />

chromatograph map from any of these other forms.<br />

This is one cactus which we are not especially happy to have<br />

to include in this account. We believe it to have been an introduction<br />

to our area, and it seems fortunate that it has not spread.<br />

Opuntia engelmannii SD<br />

“Nopal,” “Tuna,” “Engelmann’s Prickly Pear,” “Flaming<br />

Prickly Pear”<br />

Description Plates 44, 45, 46, 47, 48<br />

stems: Always growing upright, forming a bush which may<br />

be compact with a definite central trunk or open and diffuse<br />

without a main trunk. These bushes commonly attain a height<br />

of 3 to 6 feet, and some varieties sometimes stand over 10 feet<br />

tall when in optimum environments. Individual pads are<br />

thick and circular to broadly egg-shaped. They are 8 to 12 or<br />

even 14 inches long when mature. The surface is medium<br />

green or sometimes slightly blue-green, often becoming pale<br />

yellowish-green with age. It is covered more or less thickly<br />

with a whitish bloom which easily rubs off. The leaves are<br />

about 3/16 to 1/2 inch long.<br />

areoles: Small and oval or oblong at first, usually becoming<br />

round and enlarging to 1/4 of an inch or more with maturity.<br />

Sometimes they produce much wool and other tissue to form<br />

a hemispherically bulging or even columnar structure when<br />

very old.<br />

spines: Very variable. Typically there are 1 to 5 per areole<br />

on newly matured pads, but sometimes the number increases<br />

to as many as 10 or 12 during the second year. The main<br />

spines are heavy and rigid on typical forms, but on some<br />

forms they are more slender and more or less flexible. The<br />

larger spines are always flattened to some degree, often great-<br />

ly so, and often also somewhat twisted and curved, but these<br />

are usually surrounded by smaller spines which may be round<br />

and straight. The several main spines are heavy and spreading<br />

and curving downward and outward, 3/4 to 21/2 inches long,<br />

except for one form on which they are porrect and to 3 inches<br />

long. The rest of the spines spread in all directions around<br />

these and are shorter, as well as more slender. All the spines<br />

are variable in color, ranging on different specimens from<br />

entirely yellow, yellow with only slightly brownish bases<br />

through yellow mottled with brown, whitish with dark brown<br />

bases, to deep reddish-brown only slightly yellowish above.<br />

They always show more or less of an annular pattern.<br />

glochids: Tending to be few in number on the sides of the<br />

pads, but often becoming rather numerous on the edges of<br />

old pads. They are coarse and rigid, almost like the smaller<br />

fixed spines, 1/8 to sometimes 5/8 of an inch long, spreading<br />

loosely, and in one form having an exaggerated development<br />

of them, becoming a large raylike cluster. They may be all<br />

yellow, yellow mottled with brown, or all red-brown.


genus Opuntia 169<br />

flowers: Large and showy and produced in profusion. They<br />

are 21/2 to 4 inches tall and wide, clear yellow, orange, or<br />

bright red in color, the whole perianth one color on any one<br />

flower and never variegated. The ovary is variable in length<br />

from 1 to 3 inches, but is always thick for its length, obovate<br />

or inverted cone-shaped, often as wide at the top as it is long.<br />

The outer perianth segments are short and varying in shape.<br />

The inner segments are 8 to 10 in number, long, to about<br />

11/2 inches wide with the tip the broadest part, entire, blunt,<br />

but usually with a slight point at the apex. The filaments are<br />

long and cream-colored. The anthers are the same color. The<br />

stamens are very sensitive, moving at the slightest touch. The<br />

style is whitish, the stigma made up of 5 to 10 heavy lobes<br />

ranging from dark green to pure white in the different varieties.<br />

fruits: Varying within the species. Dark red, usually becoming<br />

dull purplish or burgundy when very ripe. They are<br />

completely spherical to broadly pear-shaped, with no constriction<br />

or else with a slight one at the base, but with the<br />

umbilicus perfectly flat or nearly so. Ranging in size from 1<br />

to 31/2 inches long in the different forms of the species. Also<br />

differing in the character of the fruit pulp, this in some varieties<br />

being edible and in some not, as noted in their descriptions.<br />

The seeds small to tiny, 1/16 to 3/16 inch (11/2–41/2 millimeters)<br />

in greatest diameter and comparatively thin, with<br />

narrow rims.<br />

Range. When all of its varieties are included, O. engelmannii is<br />

apparently one of the widest ranging cacti of our continent. In<br />

this broad sense, it grows from the Gulf Coast all the way to<br />

the Pacific and deep into Mexico; more specifically, in our area,<br />

from Brownsville all along the Gulf Coast into southwestern<br />

Louisiana, then back northwest past Dallas, Texas, to extreme<br />

south-central Oklahoma, which is its northernmost penetration.<br />

From there its range dips down to the vicinity of Abilene, Texas,<br />

after which it turns northwestward again into New Mexico,<br />

passing near Santa Fe, and from there going slightly southwestward<br />

into Arizona. It is found almost universally south of this<br />

line into Mexico.<br />

Remarks. Almost every student writing about this cactus has<br />

prefaced his remarks with something about this being a most<br />

misunderstood species or some such warning of difficulty to be<br />

expected. The warning is worth heeding, as only the more persistent<br />

cactophile will ever achieve any satisfactory understanding<br />

of this species. Its relationships are so subtle that no brief<br />

survey will ever reveal them and no one can ever identify all of<br />

its forms at a glance.<br />

Most descriptions of O. engelmannii have started, “Originally<br />

described as…” and then gone on to show necessary broadening<br />

of the original descriptions. This is because the plant was<br />

originally described by Prince Salm-Dyck and later by Engelmann<br />

from a few specimens out of Chihuahua. Immediately<br />

the broadening began. Engelmann himself first extended the<br />

species to include some California material, and later stated that<br />

it grew east all the way to the mouth of the Rio Grande and<br />

included those forms found there. Our description above is the<br />

broadened one very similar to the later ones of Engelmann and<br />

that of Britton and Rose.<br />

In this huge range, spanning over half the lower width of the<br />

continent, it is not surprising that there are many local populations<br />

which are different enough one from another to have been<br />

taken at one time or another for separate species. There have<br />

been various grand theories as to why so much variation occurs,<br />

most of them hinging upon assumed hybridization of an original,<br />

lately invading stock with various already existing local<br />

cacti. The first difficulty with these theories is in answering the<br />

question of what the original, invading stock was, where it<br />

came from and how it managed to invade so wide a territory.<br />

One of the pet ideas these days is that it was one of the hardy,<br />

edible prickly pears from Mexico, and that it was carried north<br />

throughout the area by early Spanish missionaries and settlers,<br />

after which it escaped and commenced the hybridizing procedure.<br />

This is an interesting and ingenious theory, but there is<br />

no actual evidence for it at all. Nor has there been shown any<br />

actual hybridization experiment to prove that this process could<br />

give the results this theory would require. It is not within the<br />

scope of this work to pass on such theories, the attempt here<br />

being, rather, to enumerate the forms we do find growing so<br />

that others may explain them if they will, but it does seem after<br />

studying them that such elaborate theories are unnecessary. Although<br />

the genetics of the cacti is still largely unknown, modern<br />

genetic principles would lead us to expect just such variation of<br />

any such widely ranging species as we find this one to be, due<br />

to the simple segregation of local populations in different environments.<br />

One must be prepared for finding these variations<br />

whenever he goes into the widely different locations where this<br />

plant grows, and must do something more than close his eyes to<br />

them or explain them away if he is to make any sense out of<br />

the cactus world.<br />

In general, O. engelmannii is the largest and strongest of our<br />

U.S. prickly pears. While in some severe locations it remains<br />

a straggly bush of hardly more than 3 feet in height, in much<br />

of its territory it is not unusual for it to become a giant of 6<br />

feet tall, and a south Texas variety occasionally stands more<br />

than 10 feet tall. Each plant is a whole thicket of giant pads<br />

comparable in size and shape to skillets, and seeming almost as<br />

indestructable. The strength of the plant is expressed also in its<br />

numbers. It is not unusual for it to take over a stretch of rangeland,<br />

and in much of south Texas there are thousands of square<br />

miles where it grows to the exclusion of almost everything else.<br />

Most of the ranchers hate it with a fury and work hard to destroy<br />

it, but for all their efforts they hardly affect it. This cac-<br />

tus can be imagined reacting with glee to the widespread rooting<br />

and chaining of the range, which eliminates much of the competing<br />

brush and most other cacti, since all of its branches and


170 cacti of the southwest<br />

pads torn apart and scattered by the machines only root where<br />

they are distributed and thus the cactus is multiplied while its<br />

competitors for space are being eliminated.<br />

At those drought times when the ranchers have nothing else<br />

left to feed their cattle, they suddenly look at O. engelmannii<br />

with a new appreciation, and move upon it with everything<br />

from common blow-torches to huge gas flame-throwers on<br />

wheels, burning off the spines which are the plant’s only protection.<br />

The hungry cattle soon catch on, and it is not an uncommon<br />

sight in south Texas and northern Mexico to see herds<br />

of cattle eagerly following the burner and jostling each other<br />

to get at the juicy pads while they are still hot from the torch<br />

which seared away their armor. But even this does not long affect<br />

our indestructable giant. Almost immediately new pads<br />

start rising from the old stump, and soon it is back to its old<br />

size.<br />

The newer technique of herbicide broadcasting could change<br />

this picture completely, as not even this giant cactus can survive<br />

the chemicals which destroy trees and all the rest of the range<br />

brush. At the time when this means is used widely enough to<br />

eliminate our giant prickly pear, however, there will be no flora<br />

of the Southwest left except grasses, and most ecologists doubt<br />

that even the animal life or the climate of the region could sur-<br />

vive such an impoverishment.<br />

This cactus is so commonly seen over much of its range that<br />

people hardly think of it, or else so dislike it—almost everyone<br />

in the area has his own story of most painful injury from it—<br />

that many cannot appreciate its beauty even when that does<br />

appear. But the tough giant does have its tender moments, and<br />

then, for usually a month of the spring, it produces every day<br />

a new crop of large, roselike flowers, to the delight of bees and<br />

all others who can evade the spines and appreciate the blooms.<br />

These flowers vary greatly in color, those of some individuals<br />

being clear yellow, those of others deep red, and there are found<br />

all intermediate shades of orange between these two extremes.<br />

While all shades are sometimes found together in the same field,<br />

there seems to be a gradient in this color distribution, the red<br />

being very common in south Texas, while it is more rare and<br />

most specimens bloom yellow or orange in New Mexico.<br />

The burgundy or deep red fruits resulting from these flowers<br />

are large and juicy, but vary in quality of the pulp, those of<br />

some of the varieties being very flavorful, while those of others<br />

are unpleasant. This probably is one reason why in some areas<br />

the fruits have all disappeared soon after they ripen while in<br />

other areas they remain on the plants so long that ripe fruits are<br />

often found nearly all winter.<br />

O. engelmannii as it occurs in southern New Mexico is very<br />

much like the original descriptions, since those were based on<br />

plants from neighboring Chihuahua. These plants usually have<br />

very heavy and whitish spines with dark brown bases. As one<br />

moves away from that locality either west or east he finds some<br />

differences in the specimens he sees. Both in Texas and California<br />

the predominant color of the spines becomes more yellow<br />

than white, and there are other differences in the various local<br />

populations.<br />

In Texas, particularly, we find many different variations to<br />

confuse us, as we might expect in the widely differing environments<br />

of that state’s huge area, and added to that we find one<br />

major mistake made very early in the study of this cactus which<br />

still affects us and distorts our understanding of the plant today.<br />

After describing O. engelmannii from Chihuahuan specimens<br />

and before understanding its wide range, Engelmann was sent<br />

specimens of cacti from New Braunfels, Texas. They were sent by<br />

Mr. Lindheimer, an early botanist living there. Some of these<br />

pads were from plants Lindheimer described to him as 6 feet tall<br />

with trunks, others from plants he said grew low and spreading.<br />

Unfortunately he sent Engelmann only fruits of the low-growing<br />

plants, and these were slender club-shaped with a deeply<br />

pitted umbilicus, very different from the fruits of the Chihuahuan<br />

O. engelmannii already described. Knowing no more than<br />

this about the New Braunfels plants, Engelmann described the<br />

whole thing as one variable new species growing either tall or<br />

low and spreading, and called it O. lindheimeri.<br />

Engelmann was too hasty in this, and, with his usual honesty,<br />

he admitted it later. In his Synopsis of the Cactaceae of the U. S.<br />

he stated without reservation: “O. lindheimeri Eng. Pl. Lindh.,<br />

is partly this same plant [O. engelmannii, which he had just<br />

been discussing], partly a hybrid form between it and perhaps<br />

O. rafinesquii, with narrow clavate fruit.” So he who erected<br />

the species, O. lindheimeri, himself abolished it, stating clearly<br />

that the large plants of New Braunfels are O. engelmannii and<br />

the low form with the clavate, deeply pitted fruits which threw<br />

him off in the first place, is a different plant which he regarded<br />

as some sort of hybrid form. This low-growing form was later<br />

called a separate species and named by Mackensen O. lepto-<br />

carpa.<br />

The point of all this is that there is nothing left to be referred<br />

to by the name O. lindheimeri. That has resolved naturally into<br />

O. engelmannii and the then yet unnamed smaller species, incidentally<br />

extending O. engelmannii’s range far across Texas<br />

where Engelmann in his earlier works did not state it to be.<br />

All this seems perfectly clear, yet the name, O. lindheimeri<br />

still persists in usage among Texans so much that all large prickly<br />

pears in the state are usually called by that name. Harder yet<br />

to understand, most botanical works still use that name for the<br />

Texas specimens of this species, just because they are from Texas.<br />

Britton and Rose show no consciousness of O. lindheimeri’s<br />

synonymy with O. engelmannii. While they discuss its variableness<br />

at length, they seem never to have read Engelmann’s renunciation<br />

of the name, made within a few years of his having<br />

coined it.<br />

And as the error persists it compounds. Backeberg, most recently,<br />

knows well the reference to the upright, trunked form<br />

and the low one, but chooses not only to ignore Engelmann’s<br />

statement that the New Braunfels plant is O. engelmannii and<br />

to keep the name O. lindheimeri for it, but then insists that the


genus Opuntia 171<br />

two forms found at New Braunfels are all one dimorphic species.<br />

Then, on this already discredited, supposedly dimorphic<br />

species, he erects a whole new section of the genus, section Lindheimeriana<br />

Backbg., into which he places together this already<br />

evaporated species and the reportedly polymorphic forms of the<br />

Galapagos Islands, merely because they are supposedly very<br />

variable. Before he is through, then, his O. lindheimeri has also<br />

become polymorphic and has taken in all the forms found at<br />

the mouth of the Rio Grande which Engelmann had already<br />

stated were O. engelmannii.<br />

Because O. engelmannii was for so long thought of as a practically<br />

unvarying species inhabiting only southern New Mexico<br />

and perhaps some of adjacent Arizona, the slightly differing<br />

forms found in Texas and the far West were constantly given<br />

names as new species. Engelmann started this himself, and others<br />

have continued it until we have a whole series of supposed species<br />

difficult to distinguish and place logically. Griffiths con-<br />

tributed a long list of these.<br />

Dr. David Griffiths, sent here by the Australian government,<br />

was associated with the U.S. Department of Agriculture for some<br />

years, studying the prickly pears all the way from Texas to California<br />

in order to try to find some natural enemies of them<br />

which could be introduced into Australia to aid in the fight<br />

against those which had established themselves and become such<br />

a scourge there. He collected scores of specimens and grew them<br />

in the U.S. Experiment Stations in Texas and in California,<br />

which were at his disposal. He took this wonderful opportunity<br />

to carry on detailed taxonomic studies of the Opuntias.<br />

His was undoubtedly the most extensive study of this group<br />

made to this day. As a result he published many papers upon<br />

them and described numerous new species, apparently producing<br />

a new species name for nearly every variation of pad, fruit, or<br />

even flower color he found.<br />

Britton and Rose published their large work on cacti soon<br />

after Griffiths’ work, but they apparently did not have the patience<br />

to study through all of his forms, and it seems they chose<br />

to list a few of them, selected almost at random, and relegated<br />

all the rest, with hardly a mention, to synonymy. Since then<br />

most others have followed in ignoring them, which is all that<br />

can be done unless a great deal of time is spent in the field recovering<br />

them, much of this time in quite inaccessible areas.<br />

I was happily following this easy path, until the ranchers<br />

brought me up short. I was calling all of the big prickly pears<br />

the same name, when some of the ranchers with whom I was<br />

looking over the pastures objected and even took offense, pointing<br />

our that this one was good and tender cattle food, while<br />

that one was so tough and fibrous the cattle could not eat it<br />

even after the spines were burned off, that this one grew only<br />

on one ranch while that one was only on a neighbor’s, and<br />

so on. I felt acutely the derision aimed my way when I had no<br />

separate names for the various forms the ordinary cowboys<br />

could distinguish at a glance. This sort of prodding made me<br />

dig up the original descriptions of all these ignored forms from<br />

Engelmann through Griffiths. I studied the original Griffiths<br />

collection in the U. S. National Museum, and then headed into<br />

the hills to try to locate them. It has taken much effort over<br />

several years, but usually they were where they were supposed<br />

to be, and many of them were distinct, once the effort was<br />

made to understand them. I can now name my rancher friends’<br />

different prickly pears to everyone’s satisfaction.<br />

Of course, the taxonomists will not be satisfied until the exact<br />

relationships of all these forms are understood, and I do not<br />

pretend to have any final word on that. Griffiths made a great<br />

start which could have led to that, but unfortunately all he was<br />

able to do was grow these forms together in the experiment<br />

stations a few years and report on the stability of their characters<br />

in these uniform situations, proving these characters more<br />

than just environmental adaptations. There is no record of anyone<br />

having done any breeding experiments or of any other such<br />

studies on them at all since then. In fact, many of these forms<br />

have probably not been collected again until this time. It is<br />

exactly for this reason that they are all dealt with here and the<br />

most specific data possible concerning them is given: otherwise<br />

they were all but lost. It is hoped that this will interest some of<br />

the newer experimental taxonomists in them and make it possible<br />

for them to find the exact forms for their detailed modern<br />

studies. Then and only then can we know exactly how to deal<br />

with them taxonomically. In the meantime it is my conviction<br />

after observing them widely that some of them more accurately<br />

deserve varietal status than species rank, and so they are listed<br />

that way here. Some of them, such as those based upon flower<br />

or spine color alone seem clearly to be only synonyms or at<br />

best forma, and these will be mentioned here, but not listed as<br />

separate taxa.<br />

Those readers desiring only a general knowledge of the Opuntias<br />

and who are satisfied with knowing what “that big prickly<br />

pear is that we see all over the desert,” and who are not worried<br />

about the detailed differences encountered within that<br />

wide ranging plant, need go no further into these varieties. The<br />

large, upright prickly pear they see all across the southern expanses<br />

of the U.S. is for the most part Engelmann’s prickly<br />

pear.<br />

Opuntia engelmannii var. engelmannii (SD)<br />

Description Plate 44<br />

stems: As the species, except usually only 3 to 5 feet tall.<br />

areoles: 1 to at least 2 inches apart. They are larger than in<br />

some forms of the species, being 3/16 to 3/8 of an inch across<br />

during the first year and sometimes enlarging even further<br />

when older. They are oval to round and more or less bulging,<br />

with brown wool.<br />

spines: As the species, except that the main centrals are less<br />

than 21/2 inches long, heavy, and much flattened, and all


172 cacti of the southwest<br />

spines are at first brown or deep red-brown at their bases,<br />

shading to lighter above, until the outer parts are an opaque,<br />

chalky white or pale straw-colored, the whole spine usually<br />

fading entirely to a plain gray with age.<br />

glochids: As the species, except yellow, brownish, or mottled<br />

yellow and brown in color and often increasing greatly<br />

to form conspicuous, open, spreading clusters on the edges of<br />

older pads.<br />

flowers: As the species, except mostly yellow and only occasionally<br />

orange or reddish, having the ovary only 1 to 11/2<br />

inches long and the stigma of 8 to 10 dark green lobes.<br />

fruits: 2 to 3 inches long, globose to sometimes broadly pear-<br />

shaped, with little or no constriction below and a flat, wide<br />

umbilicus on top. In color they are deep, dull burgundy when<br />

ripe. The pulp is juicy and edible. The seeds measure 1/8 to not<br />

quite 3/16 of an inch (3–4 millimeters) in diameter, with nar-<br />

row rims.<br />

Range. From Chihuahua and western Coahuila, Mexico, into<br />

southwestern Texas, southern New Mexico, and southeastern<br />

Arizona. Specifically, in our area, west of a line running from<br />

near Eagle Pass, Texas, to near Austin, and then on into extreme<br />

south-central Oklahoma. From there south of a line to Abilene,<br />

Texas, and on into New Mexico between Hobbs and Clovis, to<br />

the vicinity of Santa Rosa and Anton Chico, New Mexico. Beyond<br />

this the northern boundary retreats rapidly southwestward,<br />

passing near Santa Fe, but then apparently crossing into<br />

Arizona in the mountains west of Silver City, New Mexico.<br />

Remarks. This is the form of the species which was first described,<br />

and which is meant when the name is used in the narrow<br />

sense, as by all those who separate from it the primarily<br />

Texas forms—nowadays lumping these together under the name<br />

O. lindheimeri—and the far western forms which are given various<br />

other names. This description attempts to make it possible<br />

to distinguish this typical form of the species from all of its related<br />

varieties, although it must be remembered that the distinction<br />

between these forms is not a distinction between separate<br />

species, and that there are possible intermediates which will be<br />

hard to place. This is the reason why it has seemed more logical<br />

to place this, even though it is the originally described form, as<br />

a varietal subdivision apart from and under the broader, all inclusive<br />

species description. By this method the confusion over<br />

these plants is reduced, at least for me.<br />

If any one character can be mentioned which will serve to<br />

set this variety apart from all the rest to follow, it is the whiteness<br />

of the spines, usually set off against their distinctly brown<br />

bases. This whiteness is usually conspicuous and only rarely and<br />

in young spines shades into a straw-color toward the tip, while<br />

in all the other varieties which stand separate in not only this<br />

but various other ways, the predominant color is a distinct yel-<br />

low which does not even bleach to a real white.<br />

Earle, in his recent book, Cacti of the Southwest, refers to this<br />

variety and recognizes one of the interesting distinctions between<br />

it and some of its related varieties when he states, “A<br />

tasty jelly can be made of the fruits from southeastern Arizona,<br />

those in other parts of its range have a ‘flat’ taste.” Those from<br />

other areas with the “flat” taste do not belong to this variety.<br />

It is interesting that tenderness of pads does not accompany this<br />

tastiness of fruit in this variety. Its supporting tissues are too<br />

tough for cattle to master, so in the areas where this variety is<br />

the only one growing, it is not the custom to “burn pears” as it<br />

is in those areas inhabited by some of its relatives.<br />

There have been proposed at various times, as varieties or<br />

even species, some segregates which seem to actually fall within<br />

the limits of this variety. These seem to represent one or another<br />

of the extremes it takes under various conditions, but they do<br />

not have consistent, qualitative differences from it. It seems best<br />

to regard them merely as synonyms of this variety, but still to<br />

mention them so that further study of them may be made by<br />

interested workers. I add a short discussion of those found in the<br />

area of this study.<br />

O. valida Gr. seems to represent the most spiny extreme of the<br />

variety. There seems to be no way to distinguish it from the<br />

typical specimens except that the spines average a little longer<br />

and a few more numerous per areole. Some specimens in most<br />

any large population of the variety will be apt to show this<br />

armament.<br />

On the other hand, Griffiths proposed two species which seem<br />

to be only specimens of this variety with minimal size of pads<br />

and numbers of spines. They are O. gregoriana and O. gomei.<br />

The first he found near El Paso, and the second in the Platt<br />

National Park in south-central Oklahoma. This latter is, I believe,<br />

the most northeasterly record of the plant, and its smallness<br />

as well as its paucity of spines seems a natural stunting. I<br />

have more typical specimens of the variety collected not far<br />

from the Platt Park.<br />

A particular problem is presented by a form growing in the<br />

San Andreas Canyon of the Sacramento Mountains just south<br />

of Alamogordo, New Mexico. Griffiths, with his usual thoroughness,<br />

called our attention to it and, as was his habit, erected a<br />

new species on it. He called it O. dillei.<br />

As seen growing in the type locality, a very small area where<br />

it comprises only a small population (Griffiths said he saw a<br />

dozen plants only, and there are few more in evidence today)<br />

this certainly seems a distinct cactus—more so than some others.<br />

The yellow-green pads are massive, often 12 or 14 inches long,<br />

as wide or wider, and up to 1 inch thick. These huge, cart-wheel<br />

pads are often completely spineless, only occasionally having 1<br />

or 2 spines up to 1 inch long scattered here and there. It only<br />

remotely suggests a huge, bald O. engelmannii, and for this reason<br />

there has been more of a tendency for writers to treat it<br />

separately from the typical form than in the case of some others<br />

of Griffiths’ many species. However, a special factor enters here<br />

which seems to change the picture.


genus Opuntia 173<br />

Griffiths mentions that all of the specimens he saw of this description<br />

were growing in places inaccessible to livestock, and<br />

his published photo of the plant shows it growing out of the<br />

sides of steep canyon walls. Observations in the area seem to<br />

confirm that all of the population are restricted to such rocky<br />

ledges, which constitute a rather special environmental situation.<br />

As was his habit, Griffiths took some cuttings of these plants<br />

and grew them in his experimental plots. He found that, “Under<br />

cultivation the species becomes much more spiny than indicated<br />

above.” This sort of change did not take place with many of his<br />

other forms when they were cultivated, and it should make us<br />

suspect that we have here only an environmentally caused<br />

growth form instead of a genetically different plant. We should<br />

perhaps already have suspected this from the knowledge that<br />

growing out of vertical or nearly vertical rocky cliffs has the<br />

effect of reducing spines on other Opuntias as well. For instance,<br />

spineless O. macrocentra specimens are sometimes found on cliffs,<br />

with typical, spiny specimens above or at the base of the same<br />

cliffs. And there is O. laevis Coult., said to be restricted to cliffs<br />

in southern Arizona and southwestern New Mexico. While its<br />

presence only upon cliffs is generally interpreted as the result of<br />

livestock having eaten the individuals growing in the accessible<br />

places because they were unprotected by spines, I take this to be<br />

the same effect of cliff-hanging working on the normally spiny<br />

O. phacacantha.<br />

Be that as it may, Griffiths himself says, “O. dillei is most<br />

closely related to O. engelmannii, from which it differs in rarity<br />

of its spines, which are very conspicuous on that species.” It<br />

seems obvious that when, in cultivation, this difference is wiped<br />

out, we lose the real basis for making this a separate form. He<br />

does further state that the fruits and seeds are also different, but<br />

in typical O. engelmannii var. engelmannii I have found fruits<br />

and seeds which seemed in every essential detail identical with<br />

both his description and his photograph of those organs given<br />

for O. dillei.<br />

It seems best, therefore, since we want to avoid listing environmentally<br />

induced aberrations as taxa, to consider O. dillei<br />

here as merely a synonym of the typical variety. I take the<br />

space to discuss it in the hope that this will prompt someone to<br />

do ecological and genetic studies on it which will give us definite<br />

answers as to why the small population in that particular<br />

canyon develops differently. This might clear up other such problems<br />

for us at the same time. Engelmann, for instance, mentioned<br />

another growth form collected by Mr. Wright on Limpia<br />

Creek in the Davis Mountains of Texas, which he noticed but<br />

apparently did not think distinct enough for naming. I have<br />

seen and collected that form still there in Limpia Canyon, 100<br />

years later. Far better, no doubt, to leave it nameless in this<br />

group which has already had too many names, but it will be<br />

wonderful when someone discovers why in the Opuntias we<br />

have so many and such confusing forms. Until they do we must<br />

keep track of even such apparent synonyms as O. dillei, and<br />

require everyone to make plain whether he is referring broadly<br />

to the species O. engelmannii or precisely to the more closely<br />

limited O. engelmannii var. engelmannii when he refers to Engelmann’s<br />

prickly pear.<br />

Opuntia engelmannii var. cyclodes Eng.<br />

DESCRIPTION<br />

stems: As the species, except that it is a much spreading bush<br />

only 3 to 5 feet high and usually wider than it is tall, without<br />

a well-defined central stem. The individual pads are as those<br />

of the species, except only 6 to 8 inches long, usually circular,<br />

but sometimes very broad oval or egg-shaped. The surface is<br />

bright green or yellow green with some glaucescence.<br />

areoles: As the species, except never quite as large in maximum<br />

size as in some forms, and only 1 to 11/2 inches apart.<br />

spines: As the species, except for the following details. All<br />

spines are deflexed, relatively slender, and pale yellow or<br />

straw-colored with or without red or reddish-brown bases.<br />

There are usually 1 or 2 spines per areole on the upper half<br />

or so of the pad, but sometimes up to 6 spines. A typical<br />

areole’s armor consists of 1 large, flattened spine 3/4 to 13/4<br />

inches long, plus occasionally an additional shorter spine below<br />

which is only 3/8 to 1 inch long; but some specimens show<br />

up to 4 of the larger spines spreading downward and 2 of the<br />

lower, shorter ones.<br />

glochids: As the species, except on current growth usually<br />

missing entirely. On older pads side areoles show few and<br />

very short glochids, if any, but edge areoles produce some<br />

yellow glochids up to 1/2 inch long.<br />

flowers: As the species, except smaller, being usually only<br />

about 2 inches in diameter and length, with bright orange<br />

anthers and 6 or 7 dark green stigma lobes.<br />

fruits: AS the species, except always spherical or practically<br />

so, only 1 to 13/4 inches in diameter, purplish when fully ripe,<br />

but not very juicy and not pleasant to the taste. The seeds are<br />

about 3/16 of an inch (4–41/2 millimeters) in diameter, the<br />

largest found in this Species, and with broader rims than is<br />

typical for the species.<br />

Range. From the Chisos Mountains in the Texas Big Bend to<br />

near Anton Chico, New Mexico, and back down to Stein’s Pass,<br />

where the northern limit passes into Arizona.<br />

Remarks. ‘This variety of the Species was the first recognized of<br />

all which were to follow, having been set up by Engelmann as<br />

early as he listed the species, but it has suffered varying fortunes<br />

at the hands of later authorities. Engelmann encountered it at


174 cacti of the southwest<br />

Anton Chico, New Mexico, at the northern edge of the species’<br />

range. Coulter, in the next serious study, said that the variety<br />

ranges over all of southern New Mexico from El Paso to Stein’s<br />

Pass. This would have left us with a clear idea of the plant in<br />

New Mexico, at least, but then Wooton unaccountably stated<br />

“the fruit of the New Mexico plant is never globose but ellipsoid<br />

to slightly obovoid, about twice as long as broad,” and also<br />

stated that he had never seen the variety. Britton and Rose, on<br />

the other hand, do not doubt the existence of spherical fruits,<br />

but state that the characters of the variety are not constant and<br />

render it a synonym of the species. All since have ignored it entirely,<br />

except perhaps Benson, who mentions that there is a possible<br />

segregate of O. engelmannii with circular joints, smaller<br />

flowers, and spines of this color; but he says the status of the<br />

plant is undetermined and he never actually links it with this<br />

name.<br />

It is not surprising, then, that when Miss Anthony found, not<br />

in New Mexico, but in the Big Bend of Texas a plant of almost<br />

exactly this description, she did not connect it with this old variety,<br />

but erected a new taxon for it. There is a common failure<br />

to relate Texas and New Mexico cacti. No doubt because it was<br />

in Texas and because it had yellowish spines, she related it to the<br />

old, defunct name for the Texas forms of the species, and called<br />

it O. lindheimeri var. chisoensis. I have studied the preserved<br />

specimens of Miss Anthony’s type, as well as the plant growing<br />

in all stages of flower and fruit in the Chisos Mountains, her<br />

type locality. I have also studied both preserved specimens and<br />

the living plants from Engelmann’s type locality of his variety<br />

near Anton Chico, New Mexico, and I cannot but think them<br />

the same. Various other specimens have been seen from widely<br />

scattered points in southern New Mexico, bearing out the existence<br />

of the plant in exactly the range Coulter claimed for it.<br />

The Big Bend material, classed with it, only extends its previous<br />

range 250 miles or so down along the Rio Grande from its previously<br />

known existence at El Paso.<br />

As for the question, raised by Britton and Rose and most<br />

others since, of whether this form is distinct enough from the<br />

type of the species to deserve varietal rank, a reading of Backeberg’s<br />

unequivocal statement of how far Miss Anthony’s variety<br />

is from the type of O. lindheimeri, which we have seen is more<br />

properly O. engelmannii, should answer that. With him the<br />

question would seem to be instead whether this should not be set<br />

up as a separate species entirely. Charting a middle course between<br />

these opposing views, I choose to leave it here as Engelmann<br />

first represented it, a recognizable variety occurring, it is<br />

true, wholly within the range of typical O. engelmannii var.<br />

engelmannii, but separate from it by several definite characters.<br />

Variety cyclodes grows, so far as has been seen, always on the<br />

lower slopes of high mountain ranges, where it often forms large<br />

populations. Although in several situations I have seen it growing<br />

within a few miles of the typical variety engelmannii, I have<br />

never seen the two mingled in one stand.<br />

Opuntia engelmannii var. texana (Gr.)<br />

Description Plates 45, 51<br />

stems: As the species.<br />

areoles: As the species, except with less wool than in some<br />

of the other varieties.<br />

spines: As the species, except usually somewhat more slender<br />

than those of the typical variety and always yellow instead<br />

of white in their lighter color. That color varies from entirely<br />

bright, translucent yellow to yellow with brown bases or even<br />

sometimes almost entirely red-brown with only yellowish<br />

mottling or yellowish tips. The number of spines varies from<br />

1 to 8 per areole on the upper areoles of the pad.<br />

glochids: As the species, except perhaps fewer in number<br />

than in the typical variety.<br />

flowers: As the species, except usually around 3 to 4 inches<br />

in diameter. The stigma lobes number 6 to 9 and are dark<br />

green.<br />

fruits: Within the range of the species, but always 2 to 31/2<br />

inches long, globose to sometimes broadly pear-shaped, with<br />

little or no constriction below and a flat, wide umbilicus.<br />

They are a deep, dull burgundy in color when fully ripe. The<br />

pulp is juicy but flat and rather unpleasant to the taste. The<br />

seeds measure 1/8 to not quite 3/16 of an inch (3–4 millimeters)<br />

in diameter, with narrow rims.<br />

Range. All of south and central Texas east into western Louisiana.<br />

Extending north to about Tyler, Texas, and the Dallas<br />

area, then southwest to the Texas Big Bend and southeastern<br />

New Mexico, as well as into Mexico.<br />

Remarks. Certainly the difference between this form and the<br />

typical Opuntia engelmannii var. engelmannii is slight. It is hard<br />

to see how the two could ever be considered as separate species,<br />

yet this is the plant which is today almost universally referred to<br />

as O. lindheimeri, in opposition to O. engelmannii. Remembering<br />

Engelmann’s own denial of this, I am glad to be free to recombine<br />

the two and be done with the tedious arguments for leaving<br />

them two species. This plant is much more understandable<br />

in its proper place within that large, variable species complex,<br />

and only thus can we understand Engelmann’s statement that<br />

O. engelmannii grows all the way to the Gulf Coast.<br />

But can we merge it entirely with the typical form and forget<br />

it as a synonym? Although it is close to the typical variety and<br />

there may be some intermediates hard to assign between the two,<br />

when we make the separation between them varietal instead of<br />

specific, this need not hinder us, and there do seem reasons to<br />

assign this plant a separate place beside the other variety. Summarized,<br />

the distinctions between the two are as follows: variety<br />

texana has translucent yellow instead of opaque white as the<br />

dominant spine color; it has more slender spines, larger leaves,<br />

somewhat larger flowers and fruits, fruit pulp flat and unpleasant<br />

to the taste instead of tasty, pads with soft enough suppor-


genus Opuntia 175<br />

tive tissues to be edible by grazing animals instead of so tough<br />

as to be unusable in this fashion. Admittedly these differences<br />

are all minor, and it is largely in deference to my rancher friends<br />

that I first took the distinction seriously, but in the area where<br />

both forms grow together, a rancher will often know the two<br />

varieties at a glance. This one is the widely used cactus cattle<br />

food of south Texas and northern Mexico.<br />

It is hard to establish the exact range of this variety because<br />

many reports and herbarium specimens are too incomplete to<br />

make determination exact. I have not been able to collect the<br />

variety myself east of the lower Brazos River in south Texas,<br />

but it apparently has extended much farther east. O. anahuacensis<br />

Gr., from Griffiths’ description, would seem to have been<br />

this variety growing near the mouth of the Trinity River, but<br />

repeated searching of that area has not revealed it growing<br />

there today. A related variety, O. engelmannii var. alta, is in<br />

profusion in the area, but it does not fit his description. Assuming<br />

that the name Griffiths gave to his plant meant that he<br />

found it near the town of Anahuac, I have searched that immediate<br />

vicinity repeatedly, but all around that town the land<br />

has long been farmed, and no cacti survive except variety alta<br />

growing on waste islands nearby. Griffiths’ plant may have<br />

been growing in what is now the cultivated area.<br />

Engelmann and others mention such a plant as this from<br />

western Louisiana, and I have seen one good specimen, definitely<br />

variety texana, from Cameron, Louisiana, although I do<br />

not know whether it was native there or introduced. I also have<br />

had a specimen which appeared to be this variety from the<br />

piney woods just north of Alexandria, Louisiana, but it was extremely<br />

stunted and unhealthy, and did not survive for me long<br />

enough that its true form could be determined beyond question.<br />

On the western side of its range, it is equally hard to establish<br />

an exact limit to this plant’s range. It has been so long the rule<br />

to call all western forms O. engelmannii and eastern ones O.<br />

lindheimeri, that few have thought to look for more subtle distinctions.<br />

For this reason the descriptions of O. engelmannii have<br />

been broadened enough to include any of this variety which<br />

might have appeared in the West—this being the reason for the<br />

statement found in more recent books that the spines of that<br />

species are either white or yellowish. Herbarium specimens in<br />

O. engelmannii folders from New Mexico show specimens which<br />

I would suspect are actually variety texana. The variety is fairly<br />

common in the Big Bend of Texas, particularly in the mountains<br />

all the way from the Chisos to the Guadalupes, occasionally<br />

growing together with the typical variety engelmannii. I have<br />

also seen it in the Sacramento and Organ mountains of New<br />

Mexico. From his descriptions and keys, I believe what Wooton<br />

called O. lindheimeri, collected by him in the Guadalupe Mountains<br />

of New Mexico was this variety, and it also appears to be<br />

the plant of the Organ and Tortuga mountains in southern New<br />

Mexico generally called O. wootonii Gr. This latter form is supposed<br />

to have a special shape of the pad, but it seems to come<br />

within this variety, as I have duplicated its shape in Texas specimens<br />

growing in typical populations of the variety. It must<br />

be noted here that Miss Anthony’s O. engelmannii var. wootonii<br />

cannot be the same cactus.<br />

In the territory from San Antonio (which is the type locality)<br />

to Laredo, Texas, variety texana is very common, and here it<br />

often has spines almost wholly red-brown, with only the tips<br />

slightly yellowish. This coloring formed the basis for Griffiths’<br />

O. ferruginispina, but it seems only a minor color variation of<br />

the same plant. Upon the more typically colored specimens in<br />

this same area Griffiths once erected another supposed species,<br />

O. sinclairii, but it also is only a synonym of this variety.<br />

Mackensen also tried to separate this variety further, taking the<br />

extremes of the population as it is found at San Antonio and<br />

calling them O. convexa, O. griffithsiana, and O. reflexa, but the<br />

differences between these are minor, and after living at San Antonio<br />

for 6 years, I have seen so many intermediates between<br />

them that it does not seem they represent more than individual<br />

variations.<br />

Although it is another species which is generally known by<br />

the name, pest pear, this variety is the worst pest pear of south<br />

Texas. Only in a few local situations and along the Texas coast<br />

do other varieties take its place as dominant in the brush which<br />

has developed from the overgrazing of the range. Its increase in<br />

recent times has been dramatic.<br />

I have seen isolated specimens typical of this variety in all<br />

ways except with fruits very large and strongly clavate. I suspect<br />

that they may be diseased fruits, as are the extremely large<br />

and clavate ones sometimes found on O. compressa var. fuscoatra<br />

which were the basis for the supposed species, O. macateei.<br />

There is a strong temptation to say that this form of variety<br />

texana with the elongated fruits is Griffiths’ O. pyrocarpa, but<br />

his description does not otherwise fit this plant and does not seem<br />

to indicate a member of this species at all. I have not seen any<br />

plant which does conform to his description under that name,<br />

even in the type locality.<br />

Opuntia engelmannii var. alta (Gr.)<br />

Description Plates 45, 46<br />

stems: As the species, except usually forming a more definite<br />

trunk, and averaging taller than the typical variety, being<br />

commonly 3 to 6 feet tall and sometimes growing to more<br />

than 10 feet. The pad surface is more yellow-green than on<br />

most of the other varieties, and the surface is somewhat irregular,<br />

often giving the pad edge an irregular outline. This<br />

latter effect is not marked enough, however, to be considered<br />

a tuberculate surface.<br />

areoles: As the species.<br />

spines: As the Species, except for the following differences.


176 cacti of the southwest<br />

On the average the spines are more numerous, being 1 to 6 on<br />

most areoles of new pads, and ordinarily 8 to 10 or occasionally<br />

even 12 on very old pads. All spines are always pure<br />

translucent yellow in color, with no brown at the bases. The<br />

spines stand more porrect in the areole, not spreading so<br />

widely. The main spines are flattened, but not so greatly as in<br />

the typical variety, often only the bases showing this character;<br />

the smaller ones are usually round. On new growth the<br />

spines are usually about 1 inch long, but on older pads the<br />

main ones may sometimes elongate up to a maximum of 21/2<br />

inches long.<br />

glochids: As the species, except always bright yellow in<br />

color.<br />

flowers: As the species, except even more variable in color<br />

than those of most of the other varieties. They may be found<br />

from very pale cream through yellow, orange, and red to<br />

dark purplish-red. The stigma lobes are pale green, sometimes<br />

almost white. The ovary is noticeably tuberculate.<br />

fruits: Purplish-red in color, as the species. 1 to 2 inches long.<br />

In shape they are broadly oval or egg-shaped, usually with<br />

some slight constriction below and having the umbilici flat to<br />

shallowly pitted, but narrower than the widest part of the<br />

fruit. There are usually numerous glochids and often a few<br />

slender spines 1/4 to 3/4 of an inch long on these fruits until<br />

they are fully ripe, when they fall off. The pulp is deep red<br />

and edible. The seeds are quite small, being from less than 1/16<br />

of an inch to only slightly more (11/2–21/2 millimeters) in<br />

diameter, with almost no rims.<br />

Range. All along the Texas Gulf Coast, entering the state from<br />

Mexico, and found in almost continuous stands on the islands,<br />

just above the salt flats or back of the dunes all the way to the<br />

Sabine River. Said to be native in coastal Louisiana also, but<br />

not so numerous there, and also said to have been introduced<br />

all the way to Florida. Found inland in the Rio Grande Valley<br />

about as far as Mission, Texas, but nowhere else more than about<br />

20 miles or so from the Gulf.<br />

Remarks. We have here a variety easily confused, if only spines<br />

are considered, with the more spiny individuals of O. engelmannii<br />

var. texana, but clearly different, if we will take the<br />

trouble to observe such things as fruits and seeds.<br />

O. engelmannii var. alta grows in huge thickets around the<br />

mouth of each river entering the Gulf on the Texas coast. Such<br />

places are often most easily approached by boat—an unusual<br />

experience in cactus hunting—for the numerous islands in the<br />

bays and the first clayey rises above the salt flats are often so<br />

covered with huge thickets of this cactus, many individuals<br />

standing 6 to 10 feet tall, that one can hardly set foot on them.<br />

The cactus often grows so close to the water that one can reach<br />

it from the boat itself. Or some may prefer to observe it where<br />

it grows as smaller and more scattered individuals just back of<br />

the beach sand dunes or spotted here and there on the coastal<br />

plain a few miles from the baysides. Large stands of it may be<br />

easily and comfortably observed in the Santa Ana National<br />

Wildlife Refuge by the lower Rio Grande and in the Aransas<br />

National ‘Wildlife Refuge on the Gulf.<br />

When students of cacti have been faced with these huge populations<br />

of this cactus they have reacted variously to what they<br />

saw. Engelmann, in Pl. Lindh., lists “O. engelmannii… at the<br />

mouth of the Rio Grande, with almost globose fruits, innumerable<br />

small seeds and very luscious deep red pulp.” He obviously<br />

had this variety before him, and we notice that he had no hesitancy<br />

in assigning it to the species O. engelmannii, not feeling<br />

it necessary to set it apart from the western type in any way.<br />

The next person to study the coastal cacti was Dr. Griffiths.<br />

He evidently spent much time with them, and he got thoroughly<br />

entangled in trying to work out the variations he found on the<br />

Gulf. He had no way to indicate each form he came across except<br />

by describing it as a new species, and where in some other<br />

areas he did us great service this way, here his method seemed<br />

to carry him too far. He soon had a whole list of supposed<br />

species growing in this same strip along the Texas coast.<br />

When Britton and Rose were faced with this profusion of<br />

species, they made short work of it, writing as follows:<br />

Dr. Rose has examined all this region and is of the opinion<br />

that only one species of this series exists there, and this we believe<br />

is to be referred to O. lindheimeri. It is very common<br />

about Brownsville and Corpus Christi, where it forms thickets<br />

covering thousands of acres of land. It is very variable in habit,<br />

being either low and widely spreading or becoming tall<br />

and tree-like, sometimes 3 meters high, with a definite cylindric<br />

trunk. Plants from these two extremes, if studied apart<br />

from the field, might be considered as different species, but<br />

in the field one sees innumerable intergrading forms… Decided<br />

differences in the flower colors have been pointed out<br />

in the original descriptions, and we have observed them in<br />

greenhouse specimens, but they do not correlate with other<br />

characters.<br />

It seems to me that Britton and Rose were right in reducing<br />

the bulk of this coastal form to only one taxon, but I follow<br />

Engelmann in referring it to O. engelmannii, since O. lindheimeri<br />

is an invalid name for type material from two different<br />

species. Yet I am not ready to regard this Gulf Coast form as<br />

identical with the typical variety engelmannii, nor even with<br />

variety texana which sometimes grows near it. When considered<br />

together as one entity, this form presents definite and constant<br />

characters which set it off enough from all others of the O. engelmannii<br />

complex to make it one distinct variety within that<br />

species.<br />

In order to help serious students who may want to follow the<br />

many segregates of this variety which Griffiths recognized and<br />

check them themselves, I add a list of the more important of his<br />

species here included as synonyms of this variety, together with<br />

the most outstanding character he gave for each one:<br />

O. cyanella Gr. Flower brick-red, fading to purplish.<br />

Growing at the mouth of the Rio Grande.


genus Opuntia 177<br />

O. gilvoalba Gr. Flower very pale, being yellowish-white<br />

or cream-colored. This rare form was collected about 30 years<br />

ago by Mr. Fred Lawson, the well-known cactus dealer of<br />

San Antonio, Texas. His original plant still is growing in San<br />

Antonio, and is now a huge, treelike specimen about 12 feet<br />

high and broad and with several trunks oval in shape and<br />

about 2 feet in circumference. Countless cuttings from this<br />

specimen have been distributed worldwide. I have heard it<br />

referred to as O. lawsonii, and have been told that the name<br />

was once published, but have failed to find this publication.<br />

Found near the mouth of the Rio Grande.<br />

O. deltica Gr. Flower yellow, fading to reddish. Spines<br />

not quite so numerous as in some specimens of the variety.<br />

Fruit broader and more spherical than average. Found here<br />

and there in almost the whole range, but the type locality was<br />

near Brownsville.<br />

O. laxiflora Gr. Flower orange-red to purplish-red. From<br />

near Brownsville.<br />

O. bentonii Gr. Spines usually fewer and shorter than<br />

typical for the variety, but ordinarily 1 to 5 on each areole.<br />

Pads usually darker green. Flowers yellow. Fruit oval to<br />

rather elongated obovate. The more elongated of these fruits<br />

are the most nearly like those of variety texana found in the<br />

area, but the same plant at the same time will display oval<br />

fruits more typical of variety alta—as shown in Griffiths’<br />

own photograph. Said by Griffiths to grow from the Brazos<br />

River into southwestern Louisiana and to be in cultivation in<br />

Florida. This form has been listed as a synonym of O. Stricta<br />

by Britton and Rose and also by Backeberg, but it is much<br />

larger and more spiny than that plant and does not have its<br />

narrow, club-shaped, constricted, and deeply pitted fruits.<br />

The whole question of the relationship of our variety to those<br />

large Opuntias of the eastern Gulf Coast has hardly been explored.<br />

It is my suspicion that once our Texas coastal variety<br />

is looked at carefully as an entity separate from the more<br />

western varieties, and this form is compared with, for example,<br />

O. dillenii (Ker-Gawier) Haw., we may find the rela-<br />

tionship very close.<br />

O. engelmannii var. alta, taken to include all of those listed<br />

above, is probably the largest and most robust of the Texas<br />

cacti. A strictly coastal species, it surprises many people by<br />

growing so near the water. It is not useful in any stage. Its<br />

fruits are edible, but smaller than those of the more western<br />

edible varieties and full of a huge number of very small seeds.<br />

Its large pads are too woody and tough to be fed to cattle.<br />

Opuntia engelmannii var. cacanapa (Gr.)<br />

Description Plate 46<br />

stems: As the species, except usually more profusely branching,<br />

forming a broader, more bushy plant which is usually a<br />

whole thicket in itself. The pads also remain more bluish and<br />

glaucous than those of the other forms, and are usually smaller<br />

and more regular in shape than in some of the others,<br />

being normally round or only slightly oval and usually only<br />

5 to 8 inches in diameter, rarely larger than 10 inches. These<br />

pads are also thinner than is typical for the species.<br />

areoles: As the species, except a little smaller than is typical.<br />

spines: Fewer in each areole than the typical variety usually<br />

shows, having commonly only 1 on each upper side areole of<br />

the pad with 1 to 3 and rarely to 6 on the edge areoles. These<br />

spines are usually all bright yellow, only occasionally showing<br />

brown coloring in the bases. The main spine is sharply and<br />

rigidly porrect, flattened, and 1 to 3 inches long. The other<br />

spines are 3/4 to 2 inches long, slightly flattened or round, and<br />

also nearly porrect.<br />

glochids: As the species, except very few, the sides of the<br />

pads often showing none, and the edge areoles only boasting<br />

a few scattered ones.<br />

flowers: As the species, except apparently always yellow,<br />

and in all examples observed with the stigma lobes white or<br />

yellowish instead of green. The blooms come several weeks<br />

later than the other varieties of the same species in the same<br />

area.<br />

fruits: Spherical to very broadly obovate with no constriction<br />

at either end. They are small for the species, being only<br />

1 to 2 inches long, averaging slightly over 1 inch. In color<br />

they are dark red, with less purple in the hue than is typical<br />

for the species. The umbilicus is large and perfectly flat. The<br />

fruit pulp is juicy, but tasteless and apparently is not eaten<br />

by animals, since these fruits remain on the plants long after<br />

those of other varieties are gone.<br />

Range. Along the Rio Grande in northern Zapata County and<br />

over the whole western half of Webb County, Texas. It also<br />

occurs in the Chisos Mountains of Brewster County, Texas. Both<br />

of these populations represent only small invasions into the<br />

U.S. from northern Mexico.<br />

Remarks. If we were to apply the appealing but too simple suggestion<br />

that we should separate only Opuntia forms different<br />

enough to be recognizable from the moving automobile, this<br />

would still be a separate form. Once it is noticed, the more<br />

bushy growth of smaller, more bluish pads with their long, porrect,<br />

bright yellow spines can be picked out of a mixed stand<br />

at a glance, even when one is riding by at good speed. The difference<br />

from the other O. engelmannii varieties with which it<br />

often grows in the same thickets is so constant that it must be<br />

listed separately, yet the fact seems clear that it also falls within<br />

the larger species. The plant is very common around Laredo,<br />

Texas, where it makes up in some places, particularly very near<br />

the Rio Grande, half of the population of large Opuntias cov-<br />

ering the uncleared range.<br />

The collection of this form in the Chisos Mountains of the


178 cacti of the southwest<br />

Texas Big Bend was a real surprise and very confusing until we<br />

observed this plant growing rather widely in northern Coahuila,<br />

Mexico. We found it as far south as Sabinas, Coahuila. It was<br />

then easy to see that our two Texas locations represent extensions<br />

of its territory northward at the two ends of its range<br />

where the Rio Grande dips south, while it apparently does not<br />

grow to the river along its northward arc between these two<br />

points.<br />

O. engelmannii var. cacanapa is closest to variety cyclodes,<br />

from which it can readily be distinguished by its longer and<br />

more slender, more porrect spines, larger flowers with white<br />

instead of green stigmas, brighter red fruits with much smaller<br />

seeds, and thinner, blue-green instead of thick, bright green or<br />

yellow-green pads.<br />

Dr. Griffiths first encountered this form early in his studies<br />

near Encinal, Texas, which is at the extreme northern edge of<br />

its range. He gave a rather sketchy but clear enough description<br />

of it then under the name, O. cacanapa. Several years later he<br />

redescribed it as he found it in the center of its U.S. range<br />

around Laredo, Texas, with a new name, O. tricolor. The two<br />

are clearly the same, and although the plant is more commonly<br />

known under the second name, the original one should be ap-<br />

plied to it.<br />

This is one of the most beautiful of the O. engelmannii varieties,<br />

striking along the roadside with its round, frosty bluish<br />

pads and long, perfectly porrect, bright yellow spines.<br />

Opuntia engelmannii var. flexispina (Gr.)<br />

Description Plate 46<br />

stems: As the species, except smaller than some forms, growing<br />

only to about 4 feet tall and spreading, without a truly<br />

central trunk. The pads are mostly yellow-green, with a minimum<br />

of glaucescence, often becoming quite shiny.<br />

areoles: As the species.<br />

spines: As the species, except averaging longer, the main<br />

spines being 2 inches or more long, as well as more slender<br />

to the point of being somewhat flexible. All spines are yellow<br />

with or without brown bases, and all slope sharply downward<br />

or recurve tightly against the surface of the pad.<br />

glochids: As the species.<br />

flowers: As the species, except apparently always yellow<br />

and with the stigmas dark green.<br />

fruits: 11/2 to 2 inches long, broad to rather elongated eggshaped,<br />

with some constriction below. The seeds are about 1/8<br />

of an inch (3 millimeters) in diameter.<br />

Range. Known only from Webb and Zapata counties, Texas,<br />

where it is never common, but may be found occasionally from<br />

near Zapata to about 40 miles northwest of Laredo, always near<br />

the Rio Grande.<br />

Remarks. This is one of Griffiths’ forms which, while certainly<br />

not a separate species in the current concept, does appear in a<br />

definite area among the regular population. It seems possible to<br />

distinguish it from the typical O. engelmannii only on spine<br />

characters, which are never stable enough in Opuntias to be the<br />

sole basis for separating species.<br />

I list this form here as a variety only because its unique spination<br />

is obviously not environmentally caused, and this seems the<br />

best way to retrieve it from the synonymy where it has been<br />

all but lost, so that the newer techniques of taxonomy may<br />

someday be used upon it. This might reveal its true relationship<br />

to the typical form of the species and to other varieties in the<br />

midst of which it grows.<br />

The spines on this plant are much more slender than is typical<br />

for the species, and this is all the more striking since they are<br />

longer. The main spines become 2 to 3 inches long. These spines<br />

are slender enough to be somewhat flexible to the touch, hence<br />

the name of the plant. If anyone wonders about listing this<br />

form separately from the other varieties, let him try to imagine<br />

an O. engelmannii with flexible spines, and I think he will realize<br />

that this plant requires some sort of separate treatment.<br />

These spines become markedly deflexed as they mature, and on<br />

old pads all lie pressed downward almost against the surface of<br />

the pad.<br />

The cactus is encountered occasionally on the dry hills overlooking<br />

the Rio Grande both above and below Laredo, but is<br />

never abundant.<br />

Opuntia engelmannii var. aciculata (Gr.)<br />

Description Plate 47<br />

stems: As the species in growth except not observed becoming<br />

over about 4 feet high. The pads are typical, except smaller,<br />

being 5 to 8 inches long. They are also a deep, vivid green<br />

seldom seen in the other varieties.<br />

areoles: As the species, except more closely set than typical,<br />

being from only 1/2 to 11/4 inches apart, and with somewhat<br />

more brown wool. The young areoles on the sides of the pads<br />

are not as small as is typical and seem more round, but the<br />

edge areoles are typical in size, all seeming, however, much<br />

more conspicuous than ordinary because of the exaggerated<br />

glochid formation.<br />

spines: Often completely lacking. However, many plants<br />

produce 1 to 3 spines on the upper and edge areoles. Only<br />

occasionally a plant will produce a full armament of 3 to 8<br />

spines on old stem pads. These spines are typical for the species,<br />

except more slender and all more strictly deflexed or<br />

spreading downward. They are brown with yellow tips or<br />

mottled.<br />

glochids: Very conspicuous and formidable. They are bright


genus Opuntia 179<br />

red-brown or mottled or bright brown with yellow tips, and<br />

vary in length from 1/8 to 1/2 inch in the same areole and<br />

spread outward from all parts of the areole so as to form a<br />

large, loose, starlike cluster up to 1/2 inch across. These pronounced<br />

clusters are found on all areoles, not just on the<br />

edges of the pads.<br />

flowers: As the species, varying from clear yellow through<br />

orange to brick-red, and sometimes almost magenta. The stigmas<br />

have dark green lobes.<br />

fruits: Oval to broadly pear-shaped with a little constriction<br />

below and with the umbilici flat or nearly so. They are deep<br />

purplish-red in color. The seeds are 1/8 of an inch or slightly<br />

more (3–31/2 millimeters) in diameter.<br />

range. Apparently limited to Webb County, Texas, northwestward<br />

along the Rio Grande to about 50 miles above Laredo. It<br />

has been observed also in Nuevo Leon, Mexico, southwest of<br />

Laredo.<br />

remarks. This has been a much misunderstood cactus. Originally<br />

discovered by Griffiths, it would probably have remained<br />

unnoticed like most of his other finds except that Britton and<br />

Rose chose to list it as a separate species instead of lumping it<br />

with his other forms in the synonymy of O. engelmannii. Kept<br />

in this way before the public while the other forms were overlooked,<br />

the striking beauty of the spineless specimens with their<br />

areoles like stars of which the widely spreading glochids are the<br />

bright rays made it a favorite of growers. It has been cultivated<br />

almost around the world by collectors, and has recently been the<br />

only strictly Texas Opuntia cultivated and listed for sale in the<br />

catalogs of the large California dealers. All this testifies to its<br />

uniqueness, and when a spineless specimen of the darker flower<br />

color is crowned with its deep red blooms it is certainly one of<br />

the most beautiful of the Opuntias, worthy of a place in any<br />

cactus collection.<br />

This spineless form, which is the one cultivated, is seldom<br />

linked in the minds of cactus growers with the common, spiny<br />

O. engelmannii. Griffith seemed to have had only this or specimens<br />

with an occasional short spine before him, so his original<br />

description does not reflect the whole range of spination in the<br />

form. Britton and Rose mentioned that it may have several<br />

spines, but that they were deciduous. Observation in the type<br />

locality, however, shows some specimens with up to 8 long, deflexed<br />

spines on older pads. These are quite like the spines of the<br />

O. engelmannii complex in character and nearest to those of O.<br />

engelmannii var. flexispina. When it is observed that the flowers<br />

and fruits are practically identical to those of some other<br />

members of this group, it seems clear to us that we have here<br />

another variety of that complex with only minor differences<br />

from the typical form except for the exaggerated glochid formation.<br />

Backeberg’s O. aciculata is this form, but the plant he describes<br />

and pictures as O. aciculata var. orbiculata is another cactus entirely,<br />

clearly of the O. phaeacantha species.<br />

O. engelmannii var. aciculata is encountered a few miles east<br />

of Laredo, Texas, and north of that city along the Rio Grande,<br />

where it grows only on the tops of the gravelly hills. It is most<br />

common in the vicinity of the Dolores Ranch. Formerly one<br />

could drive north of Laredo on the river road and find it growing<br />

rather profusely on the shoulders of that unimproved road<br />

where it passes the Dolores Ranch, but recently that road was<br />

widened and paved, and the shoulders scraped bare of all vegetation<br />

in the process. Now one can only catch glimpses of the<br />

cactus through the ranch’s very high deer-proof fence and must<br />

drive on 40 to 50 miles above Laredo, where it is not nearly so<br />

common, to observe or collect it. I know of no other place where<br />

it can be found without a rather strenuous expedition into rough<br />

rangelands. It has been reported to me by some Texas collectors<br />

that it has been found southeast of this area below Hebbron-<br />

ville, Texas, but I have been unable to verify this.<br />

Opuntia engelmannii var. dulcis (Eng.) Schumann<br />

Description Plate 47<br />

stems: As the species, except growing more low and spreading<br />

than typical. The tallest plant observed was about 31/2<br />

feet tall.<br />

areoles: As the species at first, but becoming very large, often<br />

to 1/2 inch in diameter when old and containing a large amount<br />

of gray wool in the form of a raised doughnut with spines<br />

and long glochids on the outer edge of it and a tuft of short<br />

glochids in the center. This wool proliferates with age until on<br />

older pads each areole contains a raised doughnut of wool<br />

which becomes a columnar growth extending outward sometimes<br />

up to 1/2 inch, with spines and long glochids on the edge<br />

of it and the tuft of short glochids in its center. At a touch<br />

the whole structure of a mature areole dislodges from the surface<br />

of the plant as a unit, instead of glochids being dislodged<br />

individually, the whole cluster of spines and glochids with<br />

the woolly center sticking to clothing or flesh like a burr and<br />

leaving the surface of the plant with a small pit where the<br />

areole had been.<br />

spines: As the species, except usually 1 to 3 and never over<br />

6 per areole in number. In color they are brown at the base,<br />

the upper parts whitish to pale straw-yellow, all fading to<br />

dirty tan.<br />

glochids: Mottled brown and yellow or all yellow in color.<br />

Arranged in a unique way as follows: around the lower edge<br />

of the young areole is a semicircle of scattered, spreading glochids<br />

which are coarse and formidable, becoming with age al-<br />

most bristle-like and up to 1/4 of an inch long. With age the<br />

upper part of the areole is nearly ringed with these also. The<br />

center of the new areole contains the bulging gray wool, at<br />

first with a mass of short, very fine glochids growing out of


180 cacti of the southwest<br />

it. The wool seems to overtake these smaller glochids and<br />

grows over them except in the center of the areole, where it<br />

remains depressed, forming the doughnut of gray wool. Out<br />

of this depressed center there continues to grow the tight<br />

cluster of very fine, short glochids not at all like the heavy<br />

ones on the edge of the areole. These center glochids seem to<br />

stay just ahead of the wool as it grows outward to form the<br />

curious projection in the middle of the areole.<br />

flowers: As the species. Yellow and orange colors have been<br />

seen. The stigmas are dark green.<br />

fruits: Egg-shaped with a slight constriction below and the<br />

umbilici broad but slightly pitted. They are 11/4 to 13/4 inches<br />

long, and purplish in color, and are said by Engelmann to be<br />

sweet and more edible than most others of this group, a statement<br />

which is borne out by the fact that they are so soon removed<br />

from the plants by animals that fully ripened fruits<br />

are seldom seen in the field. The seeds are 1/8 of an inch or a<br />

little more (3–4 millimeters) in diameter.<br />

Range. Originally said by Engelmann to have been collected at<br />

Presidio del Norde and Eagle Pass, Texas. Collected in this study<br />

only about 50 miles southeast of Eagle Pass, Texas, growing on<br />

the hills overlooking the Rio Grande and again about 20 miles<br />

southeast of Anahuac, Nuevo Leon, Mexico, which is about<br />

60 miles southwest of Laredo, Texas. Although we were not<br />

able to find it there, a specimen said to have been collected some-<br />

where near Presidio, Texas, is growing in the garden of Mr.<br />

Homer Jones, the cactus dealer at Alpine, Texas.<br />

Remarks. It is another testimony to the thoroughness of the<br />

early students of cacti that a hundred years ago they found and<br />

described carefully this unusual form. Engelmann made the first<br />

description from Bigelow’s specimens. Coulter also worked with<br />

the same preserved specimens and with the living plants which,<br />

remarkably, were still growing at the Missouri Botanical Garden<br />

30 years later. He was the first to recognize that it was actually<br />

only another form of our large species; so, using the other name<br />

for the species, he called it O. lindheimeri var. dulcis. Schumann<br />

studied it also, and first made the combination of names I am<br />

using.<br />

Wooton still later claimed our plant as growing in southeast<br />

New Mexico, but his description and picture shows an entirely<br />

different cactus, and to our knowledge it has not yet actually<br />

been found in New Mexico.<br />

Britton and Rose, perhaps seeing that Wooton’s plant seems<br />

nothing but a rather typical O. engelmannii, ignored the form<br />

completely, calling it a synonym of their O. lindheimeri. Everyone<br />

has followed in this since, as the plant is rare and was ap-<br />

parently not seen again. Even Griffiths makes no mention of it.<br />

We happened upon the cactus growing rather sparingly in the<br />

very small areas described above. The areole structure with its<br />

remarkable wool and glochid formation is most striking on old<br />

plants and surely warrants listing the plant so that it is not<br />

overlooked entirely as it has been in all the studies since Britton<br />

and Rose. Other than in the unique areole structure, however,<br />

the plants characters are all similar to those of the species to<br />

which it belongs. Something like this remarkable areole structure<br />

is found again in quite another plant, however. On old pads of<br />

O. atrispina there usually develops something like the doughnut-<br />

shaped elevation of wool, but in that plant the edge areoles then<br />

proliferate the growth centers until they finally have several of<br />

these elevated circles in each areole.<br />

Opuntia engelmannii var. subarmata (Gr.)<br />

“Flap-Jack Cactus”<br />

Description Plate 47<br />

stems: Growing as a large, upright, compactly branching bush<br />

to at least 6 feet high and often as broad. Individual pads are<br />

round to obovate or often broader than they are long. They<br />

are to at least 12 inches long and wide, blue-green with much<br />

white glaucescence on them at first, changing when very old<br />

to yellow-green. They are firm and heavy, sometimes up to<br />

1 inch thick.<br />

areoles: As the species, except smaller than in some forms.<br />

spines: None on young, active pads. On very old pads which<br />

have enlarged to form a trunk or main branch there may occasionally<br />

be one spine on some areoles. It is 1/2 inch or less<br />

long, mottled brown or gray, slender, and flattened, as well<br />

as deflexed in position.<br />

glochids: As the species, except very few in number.<br />

flowers: As the species.<br />

fruits: Very broad egg-shaped with a slight constriction below<br />

and the umbilicus flat or nearly so. They are 2 to 21/2<br />

inches long, and dark red to almost purple in color. The seeds<br />

are a little more than 1/16 to just over 1/8 of an inch (2–31/2<br />

millimeters) in diameter, with wider rims than are found in<br />

most other varieties of the species.<br />

Range. Never common, but encountered occasionally along the<br />

Rio Grande from near Laredo, Texas, to the Devil’s River.<br />

Remarks. Here is a large, robust O. engelmannii, standing sometimes<br />

higher than a man’s head, and displaying huge, smooth<br />

pads, yet totally without spines except some small ones on old<br />

trunks. These pads are so broad and round that they naturally<br />

evoke the name, flap-jack cactus, long used for them. The question<br />

naturally occurs to one, how can a plant which appears so<br />

luscious stand perfectly spineless but untouched in a pasture<br />

where the animals eat any ordinary O. engelmannii to the very<br />

ground as soon as its spines are burned off ? The ranchers answer<br />

that the flap-jack cactus survives because it is so tough that<br />

the animals can’t chew it. And a person’s first attempt to lop<br />

off a branch of the plant confirms this assertion very quickly.<br />

The skin of the pads itself is very tough and the supporting vas-


genus Opuntia 181<br />

cular tissue is the most rigid I have seen in any U.S. Platyopuntia.<br />

One has to go to the large chollas to equal its strength. It<br />

takes a hatchet or some similar tool to sever even a young pad<br />

from this giant. Spines are rendered unnecessary by the tough-<br />

ness of this plant’s tissues.<br />

The cactus is never common, and there is no indication that<br />

it grows in stands anywhere. It is encountered here and there<br />

within its range as a proud individual usually towering over the<br />

lower O. engelmannii forms around it. These occasional individuals<br />

have been found along the Rio Grande from the Devil’s<br />

River on the northwest to within about 25 miles of Laredo,<br />

Texas, on the southeast.<br />

The listing of the plant here as a variety of O. engelmannii<br />

does not imply that all about its relationship is known, any<br />

more than about the other forms of this species listed as varieties.<br />

If anything, it really implies the opposite. The cactus does<br />

present general plant body, flower, and fruit characters so similar<br />

to those of O. engelmannii as to make it difficult to let it<br />

stand as a separate species. Yet it does not seem proper to lose<br />

it in that species by synonymy. Its unique features are more<br />

than just the one of spinelessness—they include its manner of<br />

growth, its robustness, the firmness of its tissue mentioned above,<br />

and so on, so that it seems it has to be regarded as more than a<br />

simple genetic form emerging here and there. The varietal status<br />

seems best until it is further studied.<br />

We find no real intermediates between O. engelmannii var.<br />

subarmata and the other varieties of the species. Griffiths, as we<br />

have seen, described what would sound like an intermediate,<br />

calling it O. dillei. But that was from canyons of the Sacramento<br />

Mountains of New Mexico, where we do not find our form, and<br />

its character of having few spines appears definitely to be an<br />

environmental one so that it seems necessary to regard it as a<br />

synonym of the typical variety. The spinelessness of our present<br />

form is not due to environment, as it often is found in the field<br />

surrounded by extremely spiny examples of the species, and it<br />

puts on no additional spines in cultivation. The point is also<br />

made that O. dillei survives only where it is inaccessible to livestock<br />

which eat it otherwise, while variety subarmata scorns<br />

such enemies.<br />

Opuntia engelmannii var. linguiformis (Gr.)<br />

“Cow’s Tongue Cactus,” “Lengua de Vaca”<br />

Description Plate 48<br />

stems: Growing upright or sprawling, often 3 to 5 feet high.<br />

Individual pads narrowly ovate to linear, varying often on<br />

the same plant from a minimum of about 8 inches long by 6<br />

wide to a greatly exaggerated length of sometimes as much as<br />

36 inches long by only 4 inches wide at the widest point at or<br />

near the base of the pad. Otherwise these stems are as the<br />

species.<br />

areoles: As the species.<br />

spines: As the species, except often shorter than typical.<br />

glochids: As the species.<br />

flowers: A weak bloomer. Some plants fail to bloom at all<br />

and are reproduced only vegetatively, but other plants bloom<br />

fairly well in good situations. The flowers are identical to the<br />

species. All observed, however, were orange or red in hue with<br />

dark green stigmas.<br />

fruits: As the Species in both fruits and seeds.<br />

Range. Originally found and definitely known to grow wild<br />

only in Bexar County, Texas. The original location was near the<br />

southeast quadrant of San Antonio’s Loop 13 and Loop 410,<br />

just south of the small towns of China Grove and Sayers.<br />

Remarks. The cow’s tongue cactus has long been a favorite with<br />

gardeners. The strange shape of the exaggerated pads is very<br />

striking, and countless yards in South Texas have somewhere in<br />

a corner a bush of this cactus. It grows well in the whole area<br />

and reproduces itself vegetatively by the rooting of dislodged<br />

pads, but it is not grown for its flowers, as many people have<br />

never seen it bloom, and some specimens, it seems, cannot even<br />

be pampered into blooming.<br />

The striking shape of the pads is obviously due to the continued<br />

activity of each pad’s growing tip. It appears that this<br />

growing tip somehow escapes the biological restriction which<br />

would normally stop growth early in the pad development on<br />

most cacti, and thus the pads of this form continue to lengthen,<br />

perhaps as long as they live. There is reason to believe that the<br />

more healthy the plant is the more rapidly the growing tip proliferates<br />

in relation to the lateral growth, and so the more exaggerated<br />

the shape becomes. Nothing is known of the internal<br />

physiology or the genetic basis for this unusual growth pattern,<br />

but the plant seems to put all its energy into the lengthening of<br />

the pads at the expense of flowering. Fruits with seeds are occasionally<br />

produced, but the seedlings have never been observed.<br />

This may represent some mutant form of O. engelmannii which<br />

reproduces itself entirely vegetatively. There is an accompanying<br />

loss of strength, as the form is more damaged by insect pests<br />

than the Species, as well as more easily killed by frost.<br />

The original plants came from southeastern Bexar County,<br />

Texas. In the type locality most of the land is now farmed, and<br />

few of the wild population have survived. Only an occasional<br />

specimen may still be found persisting in a fence-row. But innumerable<br />

examples are in the yards of San Antonio and most<br />

of South Texas to as far north as the vicinity of Austin, beyond<br />

which the cold will hardly allow it to be grown.<br />

It is hard to know now whether the form ever grew wild in<br />

other than the Bexar County location. San Antonians like to<br />

believe it is their own unique contribution to the cactus world,<br />

and it may well be. I have been told of the plant’s growing wild<br />

in various other places, but all of these I have been able to check<br />

have been near old farm sites or in some such situations where


182 cacti of the southwest<br />

they could well have escaped from some old cultivation. Britton<br />

and Rose mention having seen similar plants from near Brownsville,<br />

Texas, but do not indicate whether they were native there.<br />

Opuntia chlorotica Eng. & Big.<br />

“Clock-Face Prickly Pear”<br />

Description Plate 48<br />

stems: Erect, bushy, becoming 6 or 7 feet tall in good situations,<br />

but often smaller. They form a definite, rounded trunk<br />

quite early, often when only 2 or 3 feet tall. The individual<br />

pads are circular to very broadly obovate, with little or no<br />

constriction below, 6 to 12 inches long, pale yellowish-green<br />

with some whitish glaucescence when young.<br />

areoles: Small to medium-sized and spaced about 1 inch<br />

apart on young and growing pads, but on the very old pads<br />

increasing in size, often to 1/2 inch and until the trunk is almost<br />

entirely covered by these enlarged areoles. In the old<br />

areoles much wool appears, producing on the trunk a thick<br />

layer which has sometimes been called a tomentum.<br />

spines: Pure yellow in color, 1/2 to 2 inches long, more or less<br />

flattened, medium in thickness, and mostly deflexed. There<br />

are 1 to 6 spines per areole on young, growing pads, but they<br />

continue to be produced on older pads, until on the main<br />

stems and trunk the total may run 20 to 30 per areole, these<br />

large clusters of deflexed spines often almost wholly covering<br />

the trunk with a formidable mass of spines unique in the<br />

Platyopuntias.<br />

glochids: Few and short on new pads, but also increasing<br />

greatly with age until becoming a spreading mass of rigid<br />

bristles, many up to 1/2 or 3/4 of an inch long, in each en-<br />

larged areole. They are always yellow or straw in color.<br />

flowers: Clear yellow in color, 11/2 to 3 inches in diameter.<br />

The ovary is 11/8 to 11/2 inches long and very wide, usually 1<br />

inch or more in diameter, tuberculate, with each areole having<br />

a cluster of short, brown glochids and occasionally 1 or 2<br />

bristles up to 1/2 inch long. The stamens are yellow, and the<br />

stigmas have 10 or 12 fat greenish-yellow lobes.<br />

fruits: Spherical to ovate, 11/2 to 2 inches in length, with a<br />

shallowly pitted umbilicus. In color they are reddish-purple.<br />

The seeds are around 1/8 of an inch (21/2–31/2 millimeters) in<br />

diameter.<br />

Range. The mountains of southwestern New Mexico west of the<br />

Rio Grande on into Arizona and said to extend into California<br />

and Sonora, Mexico.<br />

Remarks. This cactus is very close to some forms of O. engelmannii,<br />

so close that there is a temptation to reduce it to the<br />

status of a variety of that species. As far as I know, however,<br />

no authority has yet actually done this, although several have<br />

discussed the possibility. I have found no one expert enough to<br />

tell the 1-year-old pads of this cactus, when removed from the<br />

bush, from those of some specimens of O. engelmannii var.<br />

texana, for instance, in every case. There seems to be little difference<br />

except that the areoles of O. chlorotica are closer together<br />

on the average. And the 2-or 3-year-old pads of O. chlorotica<br />

are almost identical with those of some O. engelmannii<br />

var. flexispina specimens. But after that, the older, trunk-forming<br />

parts of this cactus go on in the growth of their areoles and<br />

the production of spines beyond anything seen in the O. engelmannii<br />

forms. The flowers are practically identical in all visible<br />

aspects to those of O. engelmannii, and the seeds are essentially<br />

the same. The fruits of O. chlorotica seem more consistently<br />

round and they have more areoles with more glochids on them,<br />

as well as a little more deeply pitted umbilicus.<br />

This cactus is a beautiful inhabitant of the high mountains in<br />

southwestern New Mexico, being especially common and robust<br />

on the slopes of western Sierra, Grant, and Hidalgo counties,<br />

where the massive covering of bright yellow spines on the lower<br />

parts of old plants rivals in shining brilliance those of some of<br />

the chollas.<br />

Opuntia tardospina Gr.<br />

Description Plate 48<br />

stems: An erect cactus, diffuse and spreading instead of bush-<br />

like. Lacking any central stem or trunk, it produces many<br />

ascending blanches, forming a small thicket 2 to 3 feet tall<br />

when mature. The pads are round to broadly pear-shaped, 6<br />

to 12 inches long, medium in thickness. The surface is glaucous<br />

and bright to yellowish-green.<br />

areoles: Conspicuous from the first, being almost round and<br />

3/16 of an inch or more long when young, often increasing to<br />

round and 3/8 of an inch in diameter when old, bulging with<br />

much wool. They are situated 3/4 to 15/8 inches apart.<br />

spines: As observed in the field, 1 or 2 per areole in number,<br />

in only a few upper areoles of the pad. The main spines are<br />

3/4 to 11/2 inches long, rather heavy and flattened, deflexed or<br />

even recurved tightly against the pad. The second spine, when<br />

present, is a small lower spine also deflexed and only 3/16 to<br />

7/8 of an inch long. The spines are all translucent yellow with<br />

or without brown bases. O. tardospina is said by the discov-<br />

erer to become much more spiny under cultivation.<br />

glochids: Few at first, soon becoming many to very many<br />

in all areoles, 3/16 to 1/2 inch long, yellow or sometimes yellow<br />

mottled with brown.<br />

flowers: 21/2 to 3 inches in diameter and pure lemon-yellow<br />

in color. The ovary is 13/4 to 2 inches long, club-shaped, 3/4 of<br />

an inch wide, with a few very short glochids on it. The style


genus Opuntia 183<br />

is shorter than the stamens. The stigma has 7 white or yellow-<br />

white lobes.<br />

fruits: Broadly egg-shaped to club-shaped, with rather noticeable<br />

constriction of the base and with the umbilici pitted<br />

fairly deeply. They are 11/2 to 21/4 inches long and purplish-<br />

red when ripe. The pulp is not very juicy, is slow ripening,<br />

and is unpalatable. The seeds are 1/8 to 3/16 of an inch (3–41/2<br />

millimeters) in diameter.<br />

Range. In the hills along the Colorado River in central Texas<br />

from near Austin to Llano and Lampasas, Texas, and southwest<br />

to upper Uvalde County.<br />

Remarks. This is one of Griffiths’ numerous discoveries which<br />

has been very difficult for students to confirm, but which has<br />

seemed from his description unique enough that, although we<br />

can tell from their handling of it that most scholars never saw<br />

the plant, they have consistently retained the species with little<br />

comment.<br />

The amount of living material which Griffiths had before him<br />

when he made his description must have been small. He, in fact,<br />

states that the description is mainly from the type plant. This<br />

plant was collected near Lampasas, Texas. Griffiths’ reliance on<br />

so little material, the failure of more recent students to show<br />

evidence that they had their own material, and the difficulty we<br />

encountered in this study in finding it, all show that the cactus<br />

is rare in its type locality. Perhaps this is because the type lo-<br />

cality is the most northerly location at which it has been found.<br />

The fact that Griffiths based his description upon a plant from<br />

the extreme edge of its range, where the soil and water conditions<br />

are not severe, and even there upon a plant from the “valley<br />

lands,” may explain why it has been so hard to recognize the<br />

plant again. Even in that area it has been impossible to rediscover<br />

specimens with pads and fruits quite as big as his maximum,<br />

and farther southwest, where the conditions are much<br />

more rocky and arid, the specimens found are much smaller.<br />

But the species has truly outstanding characteristics, and once<br />

a person realizes that the size descriptions given are for a seldom-attained<br />

maximum and starts looking for these characters<br />

in a smaller plant, he will find O. tardospina here and there in<br />

the hills from the type locality southwest along the edge of the<br />

Edwards Plateau.<br />

The most remarkable things about the cactus are the large,<br />

bulging, dark-colored areoles, the conspicuous, loose clusters of<br />

uneven glochids, and the yellow and deflexed, but late-appearing<br />

and soon discolored spines. Griffiths did not describe the<br />

flowers, but they seem always to be clear yellow, fading to<br />

pinkish. I have always found them appearing early in the spring,<br />

being over and gone before those of the other species growing in<br />

the same area arrive.<br />

Neither did Griffiths describe the seeds of the plant. Britton<br />

and Rose erroneously state them to be 5 millimeters (over 3/16 of<br />

an inch) broad, and Backeberg follows them in the description.<br />

But then, Britton and Rose give a description almost completely<br />

different from the original and ignore all of the most conspicuous<br />

details of areoles, glochids, and spines. Their whole description,<br />

including seeds, would fit an O. phaeacantha var. camanchica<br />

with minimum armament, and they show to represent their plant<br />

a drawing of a plant from north of Dallas, Texas, where that<br />

other cactus is the main large Opuntia to be found. Seeds<br />

produced by the specimens collected in this study were rather<br />

smaller in diameter, very different from those of O. phaeacantha<br />

var. camanchica, and in the range of those of O. engelmannii,<br />

which is its nearest relative. Our plant also shows some similarities<br />

to O. atrispina, another of Griffiths’ species, whose range<br />

begins just beyond where that of O. tardospina stops on the<br />

southwest.<br />

Opuntia spinosibacca Anthony<br />

Description Plate 49<br />

stems: A strictly upright bush, its base becoming more or less<br />

trunklike by much thickening of the old pads. Many branches<br />

arise from this short base, each composed of rows of pads<br />

small for so large a cactus. These branches standing close together<br />

make this a compact, tidy shrub up to at least 4 feet<br />

tall. The pads are usually pear-shaped with rather pronounced<br />

constrictions at their bases, but occasionally they are<br />

almost round. They are from 4 to 7 inches long and 3 to 6<br />

inches wide, of average thickness for their size, but conspicuously<br />

tubercled so that each areole is on the summit of a<br />

small elevation. This gives the edge of the pad an unusual,<br />

slightly notched outline. The color of the pads is a light green<br />

or yellowish-green.<br />

areoles: Situated 3/4 to 11/2 inches apart, the areoles are<br />

oval to round, small on the faces of young pads, but to 1/4 of<br />

an inch in diameter on the edges of older pads.<br />

spines: There are 1 to 8 spines per areole, with all or almost<br />

all areoles spiny. As is common in Opuntias, lower areoles<br />

have fewer spines and the number rises in areoles toward the<br />

upper edge of the pad. A fully spined areole has 1 to 5 main<br />

spines 3/4 to 23/4 inches long spreading in all directions. These<br />

spines are heavy to very heavy, slightly or greatly flattened,<br />

often twisted and curved. They are red-brown or dark brown<br />

over the lower two-thirds of their lengths, then gray above,<br />

with the tips yellow and translucent. Below these main spines<br />

are 1 to 4 lower spines spreading downward, straw-colored<br />

mottled with brown. These range from 3/8 to 5/8 of an inch<br />

long, and are very slender bristles.<br />

glochids: Very few on this cactus. There appear to be none<br />

on the sides of most pads and only 6 to 12 in each areole of<br />

the upper edge. These few, however, are up to 1/2 inch long<br />

and light brown in color, often with yellow tips.<br />

flowers: Standing about 2 inches tall, but not opening widely,<br />

so normally 2 inches or less across. In color, they are


184 cacti of the southwest<br />

bright orange-yellow with red centers. The ovary is 1 inch<br />

long, broadly vase-shaped, slightly tuberculate, and 3/4 of an<br />

inch across at its top, which is its widest point. The petals are<br />

very wide, the edges usually somewhat ragged, but with a tiny<br />

point at the apex. They are orange-yellow, with their bases<br />

bright red, and this red may flush up the midline most of the<br />

way to the top. The stamens are very pale yellowish. The style<br />

is long and white. The stigma is composed of 7 to 9 light yellowish<br />

lobes.<br />

fruits: 1 to 13/4 inches long, 1/2 to 1 inch thick, oval or egg-<br />

shaped, and slightly tuberculate. There is no constriction at<br />

the base, which is broadly rounded. There is a rather pronounced<br />

constriction at the top, above which the upper<br />

edge flares widely to enclose the broad, very deeply concave<br />

flower scar. All of the areoles on the edge and a few down<br />

farther on the sides of the fruit have 1 to 4 spines on them.<br />

These are 1/4 to 1 inch long, rigid, round to slightly flattened,<br />

colored much as are the spines on the pads. Spreading straight<br />

out from the upper edge of the fruit or turning slightly downward,<br />

they surround the whole upper part of it with a formidable<br />

armament. When mature the fruits take on a very light<br />

greenish-yellow color. They never show any red or purple<br />

shading. Almost at once after yellowing, the fruits begin to<br />

dry out. This starts below and proceeds upward, the skin becoming<br />

tan and papery in appearance and wrinkling slightly,<br />

but not enough to change the shape or reduce the over-all<br />

size. When opened, the dried fruit presents a mass of very<br />

tightly packed seeds with the outer skin of the fruit resembling<br />

a peanut hull. The seeds are remarkably irregular. They<br />

may be almost any shape from perfectly round to elongated<br />

and from a little less than 3/16 to just over 1/4 of an inch (4–<br />

61/2 millimeters) in diameter. They are usually thin or very<br />

thin, but may be flat or sometimes greatly twisted. The rim is<br />

from medium width to very broad, 1/2 to 11/2 millimeters<br />

wide, varying on its way around the seed.<br />

Range. Known only in the small area extending a few miles<br />

from near the ranger station at Boquillas toward the Hot Springs,<br />

all in the Big Bend National Park, Brewster County, Texas.<br />

Remarks. It is remarkable that this unique cactus was not described<br />

until 1956. It presents a set of characteristics truly<br />

amazing when all combined in the one cactus. It is no wonder<br />

that theorists have not yet decided where its relationships lie.<br />

It has a tall, upright, bushy manner of growth found in the O.<br />

engelmannii group, but the compactness of its form and the<br />

smallness of its pads contrast sharply with those upright but<br />

spreading Opuntias with their large pads. It has a very yellow-<br />

green color of the surface, which I have seen otherwise only in<br />

O. chlorotica and a few specimens of O. engelmannii. Its spines<br />

appear more similar to those of some O. phaeacantha forms. It<br />

has a tubercled surface unique to itself, or equaled only on some<br />

of the small, crawling Opuntias. And most unusual of all, it has<br />

fruits which do not ever color any shade of red or purple like<br />

our other large forms and which are similar to those of the<br />

small, dry-fruited Opuntias and the Corynopuntia by being<br />

truly spiny, by becoming faintly yellowish when mature, and<br />

then by becoming dry and hard and papery. This is undoubtedly<br />

the platypus of the Opuntias. Miss Anthony discovered O. spinosibacca<br />

during her study of the Opuntias of the Big Bend,<br />

and it is clearly the greatest achievement of her study.<br />

The reason for the late date of its discovery no doubt is the<br />

fact that this prickly pear seems to grow only on a few remote<br />

hills bordering the Rio Grande in the very deepest tip of the<br />

Big Bend. Until comparatively recently there would have been<br />

no way to enter its area except by the most strenuous expedition.<br />

But fortunately for the cactus fancier, the Parks Department’s<br />

new paved road to Boquillas Canyon runs right through<br />

this area, and one may see ranks of these tidy, compact bushes<br />

drawn up stiffly on the limestone ledges of the hills he skirts<br />

without even leaving the asphalt. Fortunately also, the range of<br />

this cactus is within the Big Bend Park, where it is protected<br />

and will no doubt be flourishing when many of the desert spe-<br />

cies are gone the way of the bulldozer and the herbicide.<br />

Opuntia rufida Eng.<br />

“Blind Pear”<br />

Description Plate 49<br />

stems: Growing erect and bushy with a definite trunk, and 2<br />

to 6 feet tall. The pads are 4 to 8 inches long, round to broadly<br />

egg-shaped, or sometimes wider than they are long. They are<br />

thick, with flat surfaces, pale gray-green to somewhat yellowish-green<br />

in color. The surface appears dull due to being<br />

covered with a very short pubescence.<br />

areoles: Conspicuous, being round and large, 3/16 to 1/4 of an<br />

inch in diameter, and crowded, being 1/2 to 1 inch apart.<br />

spines: None.<br />

glochids: Very many, but very short and extremely slender.<br />

They are only about 1/16 of an inch long, and so crowded in<br />

the areole as to form a dense hemispherical tuft. This is red-<br />

brown to light brown, often fading to grayish with age.<br />

flowers: 2 to 21/2 inches in diameter, orange-yellow in color.<br />

The outer petals are almost linear or at least lanceolate, the<br />

inner ones obovate, with the ends erose or notched. The stamens<br />

are whitish. The style is somewhat longer than the stamens,<br />

and there are 5 to 11 dark green stigma lobes. The<br />

ovary is 1 to 11/8 inches long and about 5/8 of an inch across<br />

at the top, which is its widest part. It is tuberculate.<br />

fruits: Almost spherical. They are about 1 to 11/4 inches in<br />

length, and greenish-red in color. The pulp is greenish. The<br />

seeds are between 1/16 and 1/8 inch (2–3 millimeters) in diam-<br />

eter, irregular in shape, with narrow rims.<br />

range. Crossing from Mexico into the United States only in<br />

Cornyopuntia -><br />

Corynopuntia


genus Opuntia 185<br />

Texas, and there growing only on rocky mountainsides along<br />

the Rio Grande from near Presidio, through the Big Bend National<br />

Park.<br />

Remarks. This is a large, beautiful, and distinctive Mexican spe-<br />

cies which enhances our flora by crossing into our territory.<br />

However, it does not venture far into the United States. Only<br />

along a stretch of the Rio Grande something over a hundred<br />

miles long beginning near Presidio and ending at about the eastern<br />

edge of the Big Bend National Park is it to be found on our<br />

side of the stream, and nowhere is it found over about 20 miles<br />

north of the river.<br />

This narrow strip of territory comprises the most rugged and<br />

inaccessible mountains of the Big Bend, and in them O. rufida<br />

is to be found growing on rocky mountainsides and high ridges.<br />

Most typically it is seen as proud bushes growing out of almost<br />

vertical cliffs far above one, where it would take hours at least<br />

to climb up for a close look. In the past a full-scale expedition<br />

was needed to get to its area, but more recently, in the Big Bend<br />

National Park, some roads and trails lead past good stands of<br />

the cactus. And only very recently the very beautiful new road<br />

constructed from Presidio down along the river to the Park has<br />

opened up most of its U. S. range to all. Along this road there<br />

are several places where the cactus can be found close enough to<br />

the road to be accessible to hardy cactophiles. The best stands<br />

I have observed which are readily accessible are near Boquillas<br />

and just off the highway west of Terlingua, Texas.<br />

Because of its inaccessibility, early students had few specimens<br />

to refer to, and their descriptions of O. rufida were far<br />

from complete. This led to confusion. It was particularly confused<br />

with another smaller Mexican cactus, O. microdasys (Lehmann)<br />

Pfeiffer, because both of these have pubescent surfaces<br />

and lack spines. It does relate to that species, but O. microdasys<br />

is a low, spreading, or at best only a partly erect species without<br />

a trunk and with everything in miniature as compared to<br />

O. rufida. O. microdasys was and still is the most commonly<br />

cultivated Opuntia. Almost every dime store has starts of it for<br />

sale. So often was it taken for our larger cactus that Griffiths<br />

stated that in his time specimens of O. microdasys were still being<br />

distributed for O. rufida in American collections. Working<br />

from these specimens and probably never having seen the real<br />

O. rufida alive, Schumann called the cactus O. microdasys var.<br />

rufida, and this name has hung on for the form of O. microdasys<br />

with red glochids. This is, however, a completely different thing<br />

from the real O. rufida which grows in Texas—and yet, at the<br />

beginning of this study it was only this O. microdasys with red<br />

glochids which I found in the few existing Texas collections and<br />

which I naturally took for O. rufida until I saw the real thing<br />

growing in the wild. I have still never seen the real O. rufida in<br />

cultivation. All cultivated or herbarium material going under<br />

that name should be checked carefully before being accepted as<br />

the real thing.<br />

It is also very easy to confuse the true O. rufida with a cactus<br />

found growing in Mexico, which Griffiths named O. macrocalyx.<br />

This cactus grows around Saltillo, Coahuila, and I have<br />

seen it as far north as Monclova, Coahuila. It might be taken<br />

for a low, spreading, trunkless variant of O. rufida. Its pads are<br />

fully as big and very similar, except that they are darker green<br />

in color. Its growth form is mostly ascending, but is spreading<br />

and diffuse in its branching, without any central trunk, forming<br />

thickets usually about 2 to 3 feet in height. Its areoles are even<br />

closer together than those of our cactus, more like those of O.<br />

microdasys in spacing. Its flower is similar to that of O. rufida,<br />

but its fruit is elongated and 2 inches or more in length. It is<br />

not necessary here for us to evaluate its relationship to O. rufida,<br />

but the two should not be confused.<br />

Griffiths considered that O. rufida resembled O. glosseliniana<br />

var. santa-rita (Gr. & Hare) Benson more closely than any of<br />

these, except for the lack of spines. No one has studied this relationship.<br />

Anthony, in her study of the Big Bend Opuntias,<br />

describes a new variety, O. rufida var. tortiflora, which Backeberg<br />

does not regard as valid. It is interesting that on seeing her<br />

actual preserved specimen of this plant, I would have called it<br />

a spineless example of O. glosseliniana var. santa-rita. Whether<br />

future studies can link the two more closely will be interesting<br />

to watch.<br />

Opuntia macrocentra Eng.<br />

“Purple Prickly Pear”<br />

Description Plate 49<br />

stems: Upright, forming a small bush up to 2 or 3 feet high,<br />

but without any trunklike main stem. The pads are 4 to 8<br />

inches in length, round, broader than they are long, or sometimes<br />

broadly egg-shaped. These pads are comparatively thin,<br />

being often only 1/2 to 5/8 of an inch thick. They may be<br />

medium green to yellowish-green when shaded and well<br />

watered, but more often they are purplish over-all or at least<br />

spotted and streaked with purple. In the winter and in the<br />

heat of the summer this purple coloration is usually very pro-<br />

nounced.<br />

areoles: Small and inconspicuous on the sides of the pads,<br />

enlarging to about 1/4 of an inch in the edges. They vary in<br />

the distance of their spacing, on some smaller plants being only<br />

3/8 to 3/4 of an inch apart, while on larger, more robust plants<br />

they may be 3/4 to 11/4 inches apart.<br />

spines: Few in number, being limited to the very upper and<br />

edge areoles, but these long and conspicuous. There are only 1<br />

to 4 spines as a maximum, with 1 or 2 per armored areole<br />

being most common. Of these, 1 to 3 are very exaggerated<br />

spines 2 to 5 inches long, the upper round and the lower one<br />

flattened or often even grooved. They are so slender for their<br />

length that they are flexible and usually are twisted and bent.<br />

rufia -> rufida


186 cacti of the southwest<br />

They are all black or else blackish below with usually gray<br />

or brown zones at the center and tips. There is sometimes 1<br />

lower deflexed spine 3/4 to 11/2 inches long.<br />

glochids: Very few to average in number, short, 1/16 to 3/16<br />

of an inch long on the sides of the pads and not increasing in<br />

number but sometimes increasing in length to 1/2 inch on the<br />

edges. They are brown or red-brown in color.<br />

flowers: Approximately 3 inches in diameter, light yellowish<br />

with red centers. The ovary is short and egg-shaped, with<br />

some glochids on it. The stamens are yellow, and the stigma<br />

has 5 to 11 yellow or light greenish-yellow lobes.<br />

fruits: 3/4 to 11/2 inches long, spherical to broadly vase-<br />

shaped, with no constriction at the base and with the apex<br />

deeply pitted. They become bright red or orange-red in color<br />

when ripe. The seeds are about 3/16 of an inch (4–5 millimeters)<br />

in diameter, flat but twisted and irregular, with a<br />

wide, undulating rim.<br />

Range. West of a line from Brewster County in the lower Big<br />

Bend of Texas northwest through the Davis Mountains to near<br />

Van Horn, Texas, and from there north to near Roswell, New<br />

Mexico. In New Mexico all south of a line from near Roswell<br />

northwest at least as far as Socorro, then back southwest where<br />

it enters southeast Arizona.<br />

Remarks. O. macrocentra is one of the most striking Opuntias.<br />

The sight of this bushy prickly pear with the beautiful purple<br />

pads and extremely long, blackish spines is one of those desert<br />

visions which remain so visibly in a visitor’s memory. And it is<br />

remarkable that the more severe the desert climate the more<br />

vivid the coloring of the purple prickly pear becomes. Actually,<br />

the beautiful hue is merely the means this cactus uses to survive<br />

severe conditions, whether they be heat or cold or desiccation.<br />

It survives by building up large amounts of these bright pigments<br />

at its surface to shield its tender flesh. This means of protection<br />

is widely used by cacti, as many of them turn reddish in<br />

severe conditions. This cactus only excels in the amount and<br />

richness of its coloration, but that is enough to render it memorable.<br />

The spines of this cactus are few in number, but among the<br />

longest of this genus, being normally 2 to 4 inches but sometimes<br />

to 5 inches long. They are, however, slender for their length and<br />

flexible. At least one of them is always flattened, which seems<br />

to set this form off from O. gosseliniana var. santa-rita (Gr. &<br />

Hare) Benson, with which it is often confused. Benson’s insistence,<br />

strange in the light of all early descriptions, that the spines<br />

are round contributes to this error.<br />

O. macrocentra is very common, even forming the dominant<br />

Opuntia in much of the lower Big Bend of Texas. In the area of<br />

the Big Bend National Park it grows profusely, the small bushes<br />

often standing on rocky slopes where almost nothing else can<br />

survive, burnt purple by the sun in summer and frozen as purple<br />

in the winter, but still erect and unvanquished. Here in the Big<br />

Bend, in the most severe exposures, the pads are often smaller<br />

with closer areoles than typical, but nearby, where there is a<br />

little protection and moisture will be the larger, greener pads<br />

with areoles farther apart. Miss Anthony, in her study of Opuntias<br />

of the Big Bend designated the specimens she found with<br />

everything in miniature as O. macrocentra var. minor, but it<br />

does not seem this is other than the typical form stunted by environment.<br />

In the Hueco Mountains on the Texas-New Mexico border<br />

there has been noticed another population smaller than typical<br />

and having lighter colored spines than typical. This form, which<br />

has not been proved distinct either, has been referred to as O.<br />

macrocentra var. castetteri, but I have not found this name’s<br />

official publication.<br />

In southern New Mexico and Arizona O. macrocentra grows<br />

typically, as in the Big Bend, but as one goes farther north anywhere<br />

in its range, where the heat and aridity is not so severe,<br />

the species often has larger pads than farther south and presents<br />

much less of the purple coloring in the summer. Spination is also<br />

less heavy here, occasionally reduced to none. In the Davis<br />

Mountains of Texas, on a protected ridge where moisture oozed<br />

out of the rock ledges most of the time, I collected specimens<br />

with broad, fan-shaped pads, bright green in the summer and<br />

perfectly spineless. This would no doubt be the extreme effect<br />

of easy conditions on the species. But these northern plants get<br />

their turn to deck out in color when winter hits them, often<br />

standing beautifully purple against the white snow.<br />

There is so much variation in the shape and color of the fruits<br />

and the size of the seeds in various individuals of this species<br />

that I suspect the variations might fall into definite segregates<br />

within it, but so far I have not been able to correlate these differences<br />

with any other constantly varying characters or any<br />

definite populations. This would bear much more study.<br />

The species is very close to and often confused with the cactus<br />

O. phaeacantha var. brunnea Eng., which grows with it in the<br />

Big Bend and around El Paso. This other form often shows reddish<br />

coloring, sometimes almost as purple as O. macrocentra,<br />

and also has long, blackish-based spines. Almost every herbarium<br />

collection I have seen contains a scattering of this form in<br />

its O. macrocentra file, and some descriptions of O. macrocentra,<br />

beginning with those of Britton and Rose who did not recognize<br />

variety brunnea, have been broadened to include characters of<br />

that other plant. Before being satisfied that he knows the purple<br />

prickly pear one should study the description of that other form<br />

and be sure he can distinguish the two.<br />

Opuntia gosseliniana vat. santa-rita (Gr. & Hare) Benson<br />

Description Plate 50<br />

stems: An erect, bushy shrub 2 to 5 feet high with a distinct,<br />

though short trunk. The pads are thin and mostly circular


genus Opuntia 187<br />

but varying occasionally to be very broadly egg-shaped or<br />

sometimes broader than they are long. Their surfaces are glaucous<br />

blue-green, with reddish coloring around the areoles and<br />

along the edge; this coloring is more pronounced during<br />

winter.<br />

areoles: Oval to round and only 1/8 of an inch or slightly<br />

more in diameter when young, but increasing with age to<br />

about 1/4 of an inch. They are bulging when young, with much<br />

wool and many short glochids. The growth of the wool and<br />

glochids on old stem pads sometimes produces an elongation<br />

of the areole structure outward somewhat like that seen to an<br />

extreme degree in O. engelmannii var. dulcis. The areoles are<br />

spaced about 1/2 to 1 inch apart.<br />

spines: Not numerous, new pads often being spineless or with<br />

1 spine present on only a few edge areoles. Older pads may<br />

have up to 3 spines in the upper areoles only. These spines are<br />

brown or red-brown at the bases with the upper parts yellowish<br />

or mottled with yellow. In shape they are cylindrical, in<br />

position spreading or deflexed, in size slender and 3/4 to 4<br />

inches long, although usually about 11/2 to 2 inches.<br />

glochids: Many on all areoles, but very short, being usually<br />

only about 1/8 of an inch long, forming with the wool a compact,<br />

bulging mass in the areole. They are yellow to brown in<br />

color.<br />

flowers: Yellow, with white or yellowish-white stigma lobes.<br />

fruits: Oval to oblong, said to often be curved, 1 to 13/8<br />

inches long, 3/4 of an inch or a little more in diameter, with<br />

little or no constriction below and with a pitted apex. They<br />

are purple or purplish-red in color. The seeds are 1/8 of an<br />

inch or less (2–3 millimeters) in diameter, flat, with narrow,<br />

acute rims.<br />

Range. Long known to extend out of Sonora, Mexico, into extreme<br />

southeastern Arizona. It has been found more recently in<br />

extreme southwestern New Mexico, where it is probably limited<br />

to Hidalgo County. It is found again in Texas near the Rio<br />

Grande between Presidio and the Big Bend National Park.<br />

Remarks. Everyone has been cautious about declaring this a<br />

New Mexico plant, although it has seemed it should enter out<br />

of adjacent Arizona into that state. Now, however, it is definitely<br />

known to be there, having been collected a number of<br />

times. I am even more reluctant to claim it as a Texas plant,<br />

but I have found a few specimens growing on the mountains<br />

near the Rio Grande below Presidio, which I have followed all<br />

the way through fruiting and cannot distinguish from plants I<br />

have which were gathered in the Santa Rita Mountain type<br />

locality itself.<br />

O. gosseliniana var. santa-rita shows similarities not just to<br />

one, but to several other cacti. The reddish coloring of its pads<br />

makes it seem at first to be close to O. macrocentra, but in almost<br />

everything else it is very distinct from that cactus, growing<br />

more arborescent, having a similar number of spines but these<br />

very different in shape and color, having different color of both<br />

flowers and fruits, and having entirely different, much smaller<br />

seeds. In these seeds, as well as in its size and manner of growth<br />

it is somewhat like O. engelmannii, this and O. rufida being the<br />

only other Opuntias in our area equaling O. engelmannii var.<br />

alta and var. tricolor in the smallness of their seeds, but it is<br />

different from that species in various other ways. However, the<br />

seeds, almost identical to those of O. rufida, give us another<br />

link, and it is a strong one. In all main appearances this cactus<br />

is very much like a spiny O. rufida, and the discovery of variety<br />

santa-rita growing in that plant’s range in Texas requires us to<br />

take a second thought about this. Of course, there are differences<br />

besides the spines between them, variety santa-rita not being<br />

pubescent and having a different hue of pad, not having its<br />

glochids quite so fine, having white instead of green stigmas and<br />

different colored fruits, but, nevertheless, it may be the most<br />

closely related to O. rufida of any cactus in our area.<br />

Originally, in the first description of this cactus, Griffiths and<br />

Hare emphasized another similarity which they thought so<br />

strong that they named the plant O. chlorotica var. santa-rita.<br />

About the same number of similarities and differences separate<br />

O. chlorotica and variety santa-rita as the others. Britton and<br />

Rose were heroic enough, if it may be called that, to ignore all<br />

these and let the plant stand as Opuntia santa-rita, without<br />

comment. Marshall followed them. But more recently Benson<br />

has chosen to link the form up with yet another species, Opuntia<br />

gosseliniana. This is a much more spiny cactus found in Sonora<br />

near the Pacific. I do not know that plant, and since all of the<br />

students from Arizona and California who should know both<br />

forms best now choose to make this combination, I will trust<br />

that the reasons for it are even stronger than the ones which<br />

could be given for combining it with any of the above.<br />

This cactus is beautiful, but seems to be more affected by environmental<br />

factors than most large Opuntias, suffering quickly<br />

from both extreme drought or too much moisture, unusual cold,<br />

and so on. Perhaps for this reason it is never common and only<br />

isolated individuals are encountered anywhere in the U. S.<br />

Opuntia strigil Eng.<br />

Description Plate 50<br />

stems: Erect, growing as a diffuse shrub without trunk, usually<br />

about 2 feet tall. The pads are 3 to 8 inches long, round<br />

to broadly egg-shaped, medium in thickness with no constriction<br />

at the base. The surface is pale green or yellow-green<br />

in color.<br />

areoles: 1/4 to 3/4 of an inch apart and therefore appearing<br />

crowded on the sides of the pads, but often running together<br />

on the edges. They are round or oval, 3/16 to 3/8 of an inch<br />

across, and often become elevated by the addition of much<br />

gray wool.


188 cacti of the southwest<br />

spines: Possessing an array of 5 to 10 very slender, almost<br />

bristle-like spines 1/4 to 3/4 of an inch long around the lower<br />

half of each areole. These are dark brown at their bases, fading<br />

to red-brown and finally, toward the tips, becoming yellow<br />

or whitish, or else they are all whitish. Some plants have<br />

1 to 3 central spines on each upper and edge areole, but many<br />

individuals lack these. When present they are 3/4 to 13/4 inches<br />

long, slender to medium strength, round or somewhat flattened,<br />

straight and erect on the edges while more or less deflexed<br />

and bent in various directions on the sides of the pads.<br />

They are dark brown below, medium or red-brown and rather<br />

translucent in the middle zone and translucent yellowish beyond<br />

that, the three colors contrasting conspicuously.<br />

glochids: Each mature areole has a loose, spreading cluster<br />

of slender yellow glochids in the upper part of it. They are<br />

from 1/8 to 1/2 inch long, the largest ones standing upward<br />

from the upper part of the areole opposite the lower radiating<br />

spines and almost equaling them in length, contrasting sharp-<br />

ly with them in color.<br />

flowers: Small to medium in size, being 11/2 to 21/2 inches<br />

across and 11/4 to 11/2 inches tall. The ovary is short, with<br />

leaves and a few long bristles on its areoles. The petals are<br />

lemon-yellow to cream-yellow, with a slight orange cast to<br />

the midline, their ends 1 inch broad, blunt, and entirely erose<br />

or with a very slight point at the apex. The stamens are<br />

cream-colored; the stigmas with 6 cream-colored or very pale<br />

greenish-white lobes. The flowers fade to pinkish or salmon as<br />

they wilt.<br />

fruits: Spherical or practically so, and very small, usually<br />

only 1/2 to 7/8 of an inch long. They have a few very slender,<br />

brown bristles on them, and broad, flat or slightly depressed<br />

umbilici. They are bright red in color. The seeds are 1/8 of an<br />

inch or slightly larger (3–4 millimeters) in diameter, regular,<br />

and flat, with narrow, acute rims.<br />

Range. A comparatively small, triangular area of Texas from<br />

just west of Fort Stockton on the northwest to near Ozona on<br />

the northeast, and narrowing to the vicinity of Longfellow,<br />

Sanderson, and Dryden on the south.<br />

Remarks. There has long been a lack of specific information<br />

about this cactus. It is so unusual in appearance that all students<br />

have given adequate descriptions of it, but none seem to have<br />

known where it actually grows. This was probably because<br />

Engelmann, in first describing it, said simply, “In crevices of<br />

limestone rocks, between the Pecos and El Paso, Texas.” This<br />

was repeated by Coulter and by Britton and Rose, all of whom<br />

apparently saw only the type specimens.<br />

Much time spent in the huge area covered by that statement<br />

has enabled me to pinpoint its range rather closely. It does not<br />

occur in far west Texas at all. It first appears as one leaves the<br />

mountains of west Texas traveling east. At the edge of its western<br />

range it will be found as compact little upright bushes standing<br />

on the flat tops of limestone ridges and buttes just west of<br />

Fort Stockton on the north and near Longfellow, Texas, on the<br />

south. In these locations it is fairly common, but only on the<br />

tops of the hills. East of Fort Stockton it is found here and there<br />

on the hills or high ground, but less and less commonly all the<br />

way beyond the Pecos River to its easternmost collection just a<br />

few miles west of Ozona, Texas, this giving an east-west range<br />

at this latitude of about 100 miles. The range is much narrower<br />

south of this, running only from near Longfellow to about Dryden,<br />

a distance of only about 30 miles. It has never been seen on<br />

the Pecos River beyond a few miles from the site of the old Fort<br />

Lancaster, which must have been the type locality where Wright<br />

first found it in 1851.<br />

Over all of its range O. strigil is always found associated with<br />

O. phaeacantha var. nigricans, the two forms contrasting beautifully<br />

in almost all characters as they stand together on the tops<br />

of the hills.<br />

O. strigil is one of the most unusual of the Opuntias. Its close-<br />

set areoles with their numerous, very slender bristles give it an<br />

appearance of gracefulness and an entirely misleading aspect of<br />

delicacy which make it among the most attractive in any collection<br />

of Opuntias. Its fruits are among the smallest of any Opuntia<br />

in the Southwest. Its small, compact bushes are only at home<br />

on the very thin soil overlying limestone outcrops. It seems<br />

never to form thickets, probably because no arm of the bush<br />

ever touches the ground so there can be no rooting of still attached<br />

parts of the plant. It doesn’t spread from the original<br />

base.<br />

The relationship of O. strigil to other Opuntias is not clear. It<br />

has small seeds similar to those of the O. engelmannii complex,<br />

but is unlike those forms in almost all other characters. On the<br />

other hand, O. phaeacantha var. nigricans, which grows with it<br />

over all of its range, has seeds hardly larger, and some specimens<br />

of variety nigricans approach rather closely to O. strigil in the<br />

character of their smaller spines and glochids. However, this<br />

cactus has very different flowers and fruits. Near Dryden, Texas,<br />

the extreme southeastern edge of O. strigil’s range is close to the<br />

extreme southwestern tip of O. atrispina’s range. In this area are<br />

found some specimens which are almost intermediate between<br />

these two closely related forms. The relationship between the<br />

two is worthy of more study.<br />

Opuntia atrispina Gr.<br />

Description Plate 50<br />

stems: An ascending, spreading plant 2 to 3 feet tall, with<br />

joints round to broadly egg-shaped, 4 to 7 inches long by 31/2<br />

to 6 inches wide, with pads thin to medium thickness, and<br />

with the areoles usually somewhat raised. The surface is rather<br />

shiny and bright or yellow-green, occasionally with some<br />

reddish color around the areoles on specimens growing in<br />

severe exposure.


genus Opuntia 189<br />

areoles: Slightly elongated oval and 1/8 to 3/16 of an inch<br />

long when young, to egg-shaped or almost round and 1/4 of<br />

an inch or more in diameter when older. Situated 1/4 to 11/4<br />

inches apart on the pad. The areoles on this plant have the<br />

peculiar habit of proliferating when old until each areole<br />

contains 2 to as many as 6 or 8 circular growing centers, the<br />

glochids arranged in these bunches and the areoles then becoming<br />

very big and bulging. After this sort of growth the<br />

edge areoles become to 1/2 inch in diameter and almost touch<br />

each other.<br />

spines: 3 to 6 in number on the upper 1/2 to 2/3 of the pad,<br />

the main spines with black or very dark red-brown bases<br />

shading into a zone of brown followed by yellow toward the<br />

tips. The smaller spines are similar except lighter in color.<br />

There are usually 1 or 2 main spines 1/2 to 11/4 inches long,<br />

heavy and rigid, round, porrect or deflexed, and bent downward,<br />

but in some specimens these main spines may be missing<br />

entirely. There is below them another central 3/8 to 11/8<br />

inches long, identical to them except strongly recurved downward.<br />

There will then be 1 to 3 lower, deflexed and spreading,<br />

thin and bristle-like spines 3/8 to 5/8 of an inch long. All<br />

of these spines seem to fade to a dull gray-brown or blackish<br />

within a year or so and often the old spines seem to become<br />

brittle so that many of them are broken or missing entirely on<br />

old pads.<br />

glochids: Yellowish or brownish or mottled at first, becoming<br />

dull blackish-brown when old. There are average to many<br />

on young pads, 1/8 to 1/2 inch long, in irregular, untidy clusters,<br />

becoming very many and filling the areoles on old edge<br />

areoles.<br />

flowers: Deep chrome-yellow with a somewhat greenish<br />

center when first open. Soon fading to flesh-color or apricot.<br />

They usually stand only 2 to 21/2 inches in diameter, but are<br />

sometimes smaller, down to 1 inch in diameter. The ovary<br />

tube is small, 3/4 to 1 inch long, slightly tuberculate, with<br />

some glochids on it. The stamens are cream-colored; the style<br />

white. The stigma has 7 or 8 yellow or slightly greenish-<br />

yellow lobes.<br />

fruits: Bright red or yellowish-red in color, with pulp rather<br />

dry and greenish. In shape they are almost round to broadly<br />

pear-shaped, with little or no constriction below and with the<br />

umbilici from almost flat to sometimes quite deeply pitted.<br />

Extremely variable in size, ripe fruits having been measured<br />

which were from 3/8 to 15/8 inches long. The seeds are 1/8 of<br />

an inch or a little more (3–4 millimeters) in diameter, comparatively<br />

thick, with narrow rims.<br />

Range. A narrow strip of southwest Texas about 20 miles wide<br />

from the Anacacho Mountains in the southwest corner of Uvalde<br />

County west past Del Rio, the mountains of the Devil’s and<br />

Pecos rivers, to near Dryden, Texas. We have collected it again<br />

in some hills just east of Sabinas, Coahuila, Mexico, which in-<br />

dicates that the range extends on into Mexico for an unknown<br />

distance.<br />

Remarks. This is another of Griffiths’ Texas discoveries, first<br />

found near the Devil’s River, where we find it today only near<br />

the mouth of that river. This is the center of that narrow strip<br />

along our border in which it grows.<br />

I believe the knowledge of this cactus has been clouded for<br />

must by an unfortunate lack in the original description and a<br />

subsequent misunderstanding. Dr. Griffiths described the spines<br />

of his plant very completely as to color and size, but unaccountably<br />

failed to make any statement at all about their shape. Britton<br />

and Rose apparently put great reliance on the color of<br />

spines, stated by Griffiths to be for this species, “Jet black to<br />

reddish brown at base with yellow rips.” One would think that<br />

this would be striking enough to distinguish any plant, so when<br />

Dr. Ruse, looking over the type locality, found a plant with jet<br />

black spine bases, he thought he had O. atrispina. This plant had<br />

heavy, very flat spines, and Britton and Rose, therefore, incorporated<br />

into their description the statement, “Principle ones<br />

[spines]… flattened.” Once added in this way to the description,<br />

the supposed trait of flat spines has remained in all subsequent<br />

write-ups, even being featured in Backeberg’s description<br />

of the plant and his key.<br />

It is easy to see how this idea of O. atrispina having flat<br />

spines got started and persists. There is a cactus at the Devil’s<br />

River with very heavy and very conspicuously flattened, often<br />

even triangular black-based spines, their upper parts whitish.<br />

It is the cactus one would probably notice first when one visits<br />

the type locality. I thought it was O. atrispina and grew it in<br />

my own collection for some time for that species. This plant<br />

grows quite commonly from Del Rio west to at least the Davis<br />

Mountains and thence northwest into New Mexico. I even tried<br />

at one time to persuade some New Mexico cactophiles that they<br />

had O. atrispina in their state, fortunately with little success.<br />

However, I would have been right if this were Griffiths’ cactus.<br />

At this time I already had a cactus for which I could not find<br />

a name, which I kept finding on the higher elevations along<br />

about a hundred miles of the Rio Grande. It had spines blackish<br />

below with yellow rips, but these spines were always round.<br />

When it bloomed and fruited, it fitted Griffiths’ description<br />

exactly. Only when I began to suspect that this was the true<br />

O. atrispina and that Britton and Rose had seen the wrong<br />

plant, did I check everything on their plant with flat spines<br />

against Griffiths’ description, and when I did I found that its<br />

areoles were too far apart to fit O. atrispina, it had larger<br />

orange-yellow flowers with bright red centers and much too<br />

large fruits. It couldn’t be the plant Griffiths described. I then<br />

understood why their figure illustrating O. atrispina looks so<br />

little like it as to be no better than misleading.<br />

The true O. atrispina has been virtually unknown all these<br />

years while the other plant has been taken for it. This other cac-<br />

tus, also unknown by its right name all this time due to the


witholding -><br />

withholding<br />

190 cacti of the southwest<br />

confusion, is actually O. phaeacantha var. nigricans Eng. O.<br />

atrispina, as rightly known, has no trait of flower, fruit, or<br />

pad relating it to the O. phaeacantha group. It is actually closest<br />

to O. strigil, whose range adjoins that of this cactus on the west.<br />

As I have mentioned in discussing that species, there are some<br />

specimens which appear almost intermediate between these two<br />

around Dryden, Texas. It also has some characters in common<br />

with O. tardospina, whose range almost but not quite meets its<br />

own on the northeast.<br />

A remarkable peculiarity of this cactus, noted by Griffiths,<br />

is the way its fruits differ so widely in size. Healthy, well-<br />

watered plants produce fruits 1 to 15/8 inches long, yet one often<br />

sees in the field specimens having fruits 3/4 of an inch or so in<br />

length. Quite by accident I learned that this variation is due to<br />

the environment. A pot containing one of these plants once got<br />

moved back in my garden by mistake, just after flowering, to<br />

where it got almost no water for the rest of the summer. I first<br />

noticed it when its fruits ripened to a beautiful red when only<br />

1/2 inch in length. Experimentation since then has shown that<br />

by withholding water at flowering and fruiting time I get on this<br />

plant as many flowers as usual, but these reduced to as little as<br />

1 inch in diameter, and I have had these small flowers produce<br />

tiny fruits only 3/8 of an inch long yet ripening normally and<br />

containing 1 or 2 normal seeds. This is very different from most<br />

Opuntias, which in my experience either fail to bloom at all<br />

under such conditions or have regular-sized flowers which either<br />

manage to produce normal-sized fruits or abort them entirely.<br />

Opuntia leptocarpa Mackensen<br />

Description Plate 51<br />

roots: Usually tuberous, but in Some whole populations they<br />

are fibrous.<br />

stems: Large, robust pads which stand upright at first, but<br />

are too weak to support each other, so that when a branch<br />

attains the length of 2 or 3 pads it falls over and lies on the<br />

ground, thus keeping the plant a low, sprawling one at most<br />

1 to 11/2 feet high. The pads are broadly oval or egg-shaped<br />

to sometimes almost spindle-shaped, with a gradual but not<br />

pronounced constriction at the base. They are 4 to at least 10<br />

inches long and to 7 inches wide, but thin for the size, which<br />

probably accounts for their being so often bent over by the<br />

weight of new pads sprouting from them. They are bright,<br />

deep green with a shiny surface when healthy, but somewhat<br />

glaucous and yellow-green when suffering from drought or<br />

sunburn.<br />

areoles: Small and oval or round on the sides of the pads,<br />

enlarging somewhat, but not conspicuously on the edges. They<br />

are located 3/4 to 2 inches apart.<br />

spines: 1 to 5 per areole, found on upper and edge areoles<br />

only. Occasional plants are almost spineless. The main spines<br />

are yellow or straw mottled with brown, the lower and smaller<br />

spines gray. All spines are slender to medium thickness,<br />

round or only slightly flattened. A fully armed edge areole<br />

will present 1 main central 1/2 to 11/4 inches long, porrect or<br />

slightly deflexed; 1 to 2 upper spines standing at any angle,<br />

1/2 to 1 inch long; and 1 or 2 slender, deflexed lower spines<br />

1/4 to 1 inch long.<br />

glochids: Brown to red-brown in color, few and short, 1/16<br />

to 1/8 of an inch long on the sides of the pads, to medium in<br />

number and sometimes to 1/4 of an inch long on the edges of<br />

the pads. They are never conspicuous.<br />

flowers: 21/2 to 3 inches across, yellow or orange with maroon<br />

to bright red centers. The ovary is 11/2 to 2 inches long,<br />

narrowly club-shaped, and tuberculate, with short red glochids<br />

on it. The petals have a more pronounced point at the<br />

apex than on most Opuntias. The anthers are cream in color.<br />

The stigmas consist of 6 to 8 white or greenish-white lobes.<br />

fruits: Very large, being 2 to 31/2 inches long and 1 to 15/8<br />

inches thick. They are club-shaped with a pronounced or even<br />

exaggerated constriction of the base and narrow and deeply<br />

pitted umbilici. They become old red when fully ripe, but the<br />

flesh often remains greenish and sour until late fall, when it<br />

finally becomes red and sweet. The seeds are less than 1/8 to<br />

not quite 3/16 of an inch (21/2–4 millimeters) in diameter. They<br />

are regular in shape, flat but rather thick bodied, with nar-<br />

row, acute rims.<br />

Range. An area of south Texas bounded, so far as is presently<br />

known by the following points: Langtry, Texas, and the vicinity<br />

of the mouth of the Devil’s River on the west, near Austin on<br />

the north, Flatonia on the east, Mustang Island in front of Corpus<br />

Christi Bay to near Brownsville on the southeast, and La-<br />

redo on the southwest.<br />

Remarks. Mackensen, in originally describing O. leptocarpa,<br />

stated that it was intermediate between O. macrorhiza and O.<br />

lindheimeri [O. engelmannii]. His remark was one of those<br />

purely subjective pronouncements which contribute nothing but<br />

confusion, and much confusion has entered here.<br />

Having the combination of such large pads and fruits with<br />

such small seeds and growing, as it does around San Antonio,<br />

in the same fields with O. engelmannii, the careless observer, encouraged<br />

in this direction by Mackensen’s statement, sees it as<br />

just a low, prostrate form of that upright, bushy giant. On the<br />

other hand, to some such observers casually alerted by Mackensen’s<br />

statement, its flowers and fruits may seem essentially only<br />

oversized versions of those of O. macrorhiza, while its often<br />

tuberous roots will surely strengthen this idea. I find people<br />

about equally divided as to which of these other species to call<br />

it. This is unfortunate. Closer observation will show that its flowers,<br />

fruits, and roots are essentially different from those of O.<br />

engelmannii, while its pad character and size, glochids, spines,<br />

and seeds put it clearly outside the limits of O. macrorhiza. Nat-


genus Opuntia 191<br />

ural hybrid or what have you though it may turn out to be,<br />

although it is somewhere between the two, yet it is neither of<br />

them, and in the preoccupation with relating it to those two,<br />

the fact has not been mentioned that its closest similarity is actually<br />

to the O. phaeacantha group. But it seems clear that with<br />

our present knowledge such speculation gets us nowhere.<br />

This plant probably figured in Engelmann’s confusion which<br />

resulted in his contradictory description of his now long-defunct<br />

O. lindheimeri as “erect, robust or low and prostrate,” and accounts<br />

for his attempt to set that off from E. engelmannii at all.<br />

He said in Plantae Lindheimerianae, “the fruit which Lindheimer<br />

has sent as belonging to this species [O. lindheimeri] resembles<br />

very much that of O. vulgaris, 2–21/2 inch long, slender,<br />

with a deep umbilicus, very different from that of the following<br />

species [O. engelmannii].” It should be noted that the plants<br />

sent him by Lindheimer were from New Braunfels, Texas, within<br />

the range of O. leptocarpa. Later, after gaining more knowledge<br />

of the New Braunfels plants, Engelmann said, “O. lind-<br />

heimeri Eng. Pl. Lindh. is partly this same plant [O. engelmannii],<br />

partly a hybrid form between it and perhaps O. rafinesquii<br />

[O. compressa], with narrow clavate fruit.” While the low,<br />

prostrate form referred to above, that first caused Engelmann<br />

to set up a species O. lindheimeri, could conceivably have been<br />

O. phaeacantha var. camanchica, which is found about New<br />

Braunfels also, that plant does not have the long, narrow,<br />

clavate fruits he mentioned. These are found on O. leptocarpa,<br />

and we still find in the area in question the large, upright O.<br />

engelmannii with large but broad and flat-topped fruits and the<br />

low, sprawling O. leptocarpa with large but narrow and deeply<br />

pitted fruits. Both have similar large pads and small seeds, but<br />

they have different flowers and roots, and so, as Engelmann<br />

discovered and admitted long ago and as any careful observer<br />

will verify, they cannot be combined into the one nonexistent<br />

species O. lindheimeri.<br />

Speculation about this plant’s relationship to O. phaeacantha<br />

var. major might prove to be more fruitful. A careful checking<br />

of the characters of the two show few really clear differences<br />

between them. About all that can be demonstrated to distinguish<br />

them is O. leptocarpa’s lower maximum height and much more<br />

sprawling, almost prostrate aspect, its fruits being larger and<br />

its seeds smaller. While O. leptocarpa has tuberous roots in most<br />

specimens seen and O. phaeacantha usually fibrous, we know<br />

from other species that this seldom if ever can be a diagnostic<br />

character. There have been found O. phaeacantha specimens in<br />

New Mexico with tuberous roots. At this writing I am leaving<br />

the two separate, but further study is needed to see if they are<br />

really more closely related.<br />

In few places in its whole range does O. leptocarpa exist in<br />

large stands. It is most widespread and common in Comal<br />

County, Texas, which is the type locality, and in Bexar County.<br />

Outside of these counties we find scattered small populations<br />

with many miles between them, the main ones being just west<br />

of Falls City, Texas; near Devine and Flatonia; in the sand<br />

dunes at Port Aransas on Mustang Island; in the sandy river<br />

bottoms at Laredo, Texas; and in the flats from Del Rio to<br />

Langtry.<br />

Opuntia phaeacantha Eng.<br />

“New Mexico Prickly Pear,” “Brown-Spined Prickly Pear,”<br />

“Tulip Prickly Pear”<br />

Description Plates 51, 52, 53<br />

roots: Almost always fibrous, but some specimens have been<br />

found with tuberous roots.<br />

stems: Growing either as a large, robust, ascending cactus 2<br />

to 3 feet or more high, trunkless, and forming dense thickets<br />

sometimes up to 6 or 8 feet across, or else as a low, prostrate<br />

plant with most of its pads resting their edges on the ground<br />

and so only 1 to 11/2 feet in height. The pads are flat, almost<br />

spherical to broadly egg-shaped or sometimes even club-shaped<br />

from 4 to 9 inches long and 3 to 7 inches wide. The surface<br />

is glaucous and bluish-green or deep green when young, usually<br />

becoming bright shiny green or yellow-green when older,<br />

and sometimes taking on a purplish cast around the areoles<br />

and pad edges during the winter.<br />

areoles: Small to medium and oval to round when young,<br />

becoming round and enlarged to 1/4 of an inch or so when<br />

older. They vary from 3/4 to 2 inches apart.<br />

spines: 1 to 8 spines per areole on only the very upper areoles<br />

of the pad or sometimes on the upper 3/4 or more of each pad.<br />

In length the spines vary from 3/4 to 3 inches long. They<br />

spread in almost all directions from the areole, with the upper<br />

spines large and heavy, the uppermost of these usually<br />

round and at least the main porrect or deflexed central flattened.<br />

There are usually some small, weak spines below these<br />

main ones. All spines are yellow, gray, or mottled toward the<br />

tips, with the bases brown, red-brown, or black.<br />

glochids: Few to average in number and short on young<br />

pads, becoming usually quite numerous to very many on the<br />

edges of old pads and 1/2 to 3/8 of an inch long. In color they<br />

are pale brown to red-brown.<br />

flowers: 2 to 3 inches in diameter, bright yellow or orange<br />

with red to maroon centers. The ovary varies from only 3/4<br />

of an inch long in some forms to as much as 13/4 inches in<br />

others, always with some short red-brown glochids on it. The<br />

stamens are yellowish or cream in color. The style is long and<br />

pinkish or whitish. The stigmas have 6 to 10 short, fat, whitish,<br />

yellowish, or very pale greenish lobes.<br />

fruits: Variable within this large complex: from oval or<br />

goblet-shaped with little or no constriction at the bases and<br />

the umbilici somewhat pitted to flat, to pear-shaped or even<br />

club-shaped with pronounced constriction of both the bases


192 cacti of the southwest<br />

and umbilici. In size they range from 1 to 21/2 inches long;<br />

in color, from yellowish-red to bright scarlet or cherry-red<br />

to deep purplish-red. The seeds measure from 1/8 to 1/4 of an<br />

inch (3–6 millimeters) in diameter.<br />

Range. A very large area covering many states. In our area it<br />

grows over approximately the western halves of Oklahoma and<br />

Texas and all of New Mexico.<br />

Remarks. This is the second major complex of U. S. Opuntias.<br />

It includes a number of forms so different as to require separate<br />

treatment. Their existence renders any general description like<br />

the above almost useless, and yet they intergrade so often that<br />

they have caused much confusion. Most of them have been described<br />

at one time or another as separate species, yet they<br />

hardly seem that distinct, and there have been many schemes<br />

for grouping them. At the present time there is no agreement<br />

as to how they should be treated, the writings of the authorities<br />

ranging all the way from those who would give most of them<br />

species rank to those who speak of O. phaeacantha in general<br />

terms such as the above and would forget almost all of the segregates.<br />

The method followed here is to give the above general<br />

description, beyond which anyone satisfied with merely knowing<br />

O. phaeacantha in this general way need not go, but then to<br />

describe below the segregates which we find definite and distinguishable<br />

with known ranges, listing them as varieties. In this<br />

way we hope to satisfy the serious student and also to keep the<br />

knowledge of these forms alive until more about their real relationships<br />

can be learned.<br />

O. phaeacantha, all of these forms included, may equal or<br />

even exceed O. engelmannii in the amount of territory it covers<br />

in the United States. While it does not venture as far into the<br />

southeast as does that other species, it does range practically to<br />

the Pacific, and extends its territory far north through Nebraska,<br />

Colorado, and Montana. It is an extremely hardy, often<br />

very conspicuous cactus over most of the western United States.<br />

Opuntia phaeacantha var. major Eng.<br />

Description Plate 51<br />

stems: Ascending and spreading. Growing without a central<br />

trunk to at least 3 feet tall. The pads are round or nearly so,<br />

only rarely being very broadly pear-shaped, and 4 to 8 inches<br />

long. Their surfaces are flat, deep green to yellow-green in<br />

color, often with some purplish around the areoles, and with<br />

only very slight glaucescence, usually becoming shiny when<br />

older.<br />

areoles: As the species, except smaller and more elongated<br />

oval when young than some phaeacantha forms, and spaced<br />

1 to 2 inches apart.<br />

spines: Fewer, shorter, and lighter in color than most forms<br />

of the species possess. There are typically 1 to 3 spines on only<br />

the upper edge areoles, with only a few individuals showing<br />

to 4 spines on the areoles of about the upper half of the pad.<br />

The 1 or 2 main spines are porrect or spreading, 3/4 to 17/8<br />

inches long, averaging about 11/4 inches, flat or round, with<br />

usually the uppermost one round and the lower one flattened.<br />

These are gray, whitish, or straw with brown or red-brown<br />

bases. There may or may not be 1 or 2 lower, deflexed, bristle-<br />

like spines, 1/4 to 3/4 of an inch long and grayish in color.<br />

glochids: Very few and short, or even none on the sides of<br />

the pads. The edges of old pads develop an average number<br />

of them to 1/4 of an inch or longer and yellow or brownish<br />

in color.<br />

flowers: As the species, except that the ovary is usually 11/4<br />

to 13/4 inches long.<br />

fruits: 11/2 to 21/2 inches long, club-shaped, with noticeable<br />

constriction of the bases, and constricted, pitted umbilici. In<br />

color, they are light red to purplish-red. The pulp is juicy, but<br />

sour to the taste. The seeds are about 3/16 of an inch (4–5 millimeters)<br />

in diameter.<br />

Range. From at least as far north as the vicinity of Santa Fe,<br />

New Mexico, south and east over the southeastern corner of<br />

New Mexico into far west Texas from El Paso to at least Fort<br />

Stockton.<br />

Remarks. Engelmann felt it necessary after describing O. phaeacantha<br />

in very general terms to break the species up into varieties,<br />

and one of these was his variety major. He described<br />

this variety rather poorly, merely stating that it had suborbiculate<br />

pads to 8 inches long and spines fewer, shorter, and paler<br />

than typical for the species. Coulter, working from the same<br />

specimens as Engelmann, while abandoning all of Engelmann’s<br />

other varieties of the species, felt this one alone was so distinct<br />

as to require listing. Wooton apparently means this variety only<br />

and is eliminating all of the lower, very spiny, and very common<br />

New Mexico forms of the species when he writes of O.<br />

phaeacantha, since he says it grows to 1 meter tall and “is the<br />

common sub-erect species of the north and central part of the<br />

state of New Mexico, but nowhere very abundant.” Britton and<br />

Rose, on the other hand, were the first to eliminate all of these<br />

varieties and spoke of O. phaeacantha only in the most general<br />

way.<br />

Early in this study, after we had seen many small, very spiny<br />

plants of this species with fat, hardly constricted fruits, we began<br />

looking for the reason why Engelmann and Coulter in their<br />

descriptions of the species both called the fruits “slender,” “Pyriform,”<br />

with “base much constricted,” and so on, as well as why<br />

Wooton described it as so large and so uncommon. We sensed<br />

something more than meets the eye in the usual present-day accounts<br />

of the species here, and began looking for some special<br />

form.<br />

In this search we finally found the form described above<br />

growing in a garden collection in New Mexico. It was a large,<br />

spreading plant 3 feet high and had grown in the few years it


genus Opuntia 193<br />

had been in the garden to an impenetrable thicket over 6 feet<br />

across. The gardener could not remember where he had collected<br />

it, beyond the fact that it was in the mountains near<br />

Santa Fe, which is exactly the type locality given by Engelmann<br />

for his variety major. It was the first O. phaeacantha we<br />

had seen with actually club-shaped fruits, deeply pitted at the<br />

apex. The fruits were up to 21/2 inches long. In studying this<br />

plant, we soon realized that we must have the form which was<br />

the basis for Engelmann’s variety major.<br />

Wooton was certainly correct in stating that this form is not<br />

abundant anywhere, as much searching has never enabled us to<br />

find it growing wild anywhere in northern or central New<br />

Mexico. The nearest to that area where we have ourselves found<br />

it growing is west of Roswell, New Mexico, where we have<br />

found numerous individuals growing on the eastern foothills of<br />

the Capitan Mountains. We have also found isolated individuals<br />

of this form growing near Lovington and southeast of Carlsbad,<br />

both in the southeastern corner of the state. Griffiths’ herbarium<br />

specimens in the United States National Museum appear to increase<br />

the southeastern range of the variety still more, as he has<br />

what appear to be this same plant from near Fort Stockton,<br />

Texas, and from El Paso. We have not been able to collect it<br />

in Texas ourselves, however.<br />

The true extent of the range of this variety is very hard to<br />

discover, since, because of the general tendency to submerge any<br />

brown-spined Opuntia under the catch-all name O. phaeacantha<br />

and dismiss it, there are too many incomplete descriptions in the<br />

literature, and herbarium collections are hopelessly mixed. There<br />

are specimens from Arizona which may be this variety, but one<br />

can hardly be sure from a single pad, and we hesitate on this,<br />

since we have not found it anywhere in western New Mexico.<br />

In Texas the relationship between this variety and O. leptocarpa,<br />

which has almost identical flowers and fruits, but which<br />

does not grow so robustly and which usually has tuberous roots<br />

as well as smaller seeds, should be considered. While we have<br />

not seen variety major in far northern New Mexico, we do find<br />

a form of O. phaeacantha rather intermediate between this and<br />

O. phaeacantha var. camanchica in Colorado, growing at higher<br />

elevations and farther north than variety camanchica does in<br />

that state. It has spines, fruits, and seeds much like variety major,<br />

but is much smaller and more low-growing than this variety<br />

appears farther south. We do not know whether this difference<br />

is due to the environment and variety major should actually be<br />

considered to range into Colorado or whether this is another<br />

segregate, but we assume this is the cactus from Colorado described<br />

by Coulter as O. mesacantha var. oplocarpa, which<br />

might, therefore, be a synonym of variety major or might be a<br />

very closely related Colorado form. It is our opinion that O.<br />

expansa Gr. is a synonym of variety major.<br />

This is a rather rare variety of O. phaeacantha, but an important<br />

one to the understanding of the species. The ignoring of<br />

it in most recent accounts of the species has made much of the<br />

early literature on this species hardly understandable. We do not<br />

pretend to know the details of its relationship to the other forms<br />

of this complex, but the knowledge of it must be kept alive<br />

until all this is worked out. The tendency to overlook it entirely<br />

or to casually dismiss it as a hybrid between O. phaeacantha and<br />

O. engelmannii without presenting any evidence that this hybridization<br />

actually occurs will surely not satisfy any thorough<br />

student.<br />

Opuntia phaeacantha var. nigricans Eng.<br />

Description Plate 51<br />

stems: A large bush 2 to 3 feet high when mature. The branches<br />

are upright but without any central stem and so it spreads<br />

outward to form small thickets. The pads are normally 4 to 8<br />

inches but sometimes to 10 inches long, round to broadly egg-<br />

shaped, with little or no constriction below. These pads are<br />

thin to medium thickness. In color they are at first a very<br />

glaucous blue-green, but when older become yellow-green.<br />

The two colors often contrast sharply between the upper and<br />

lower pads of the same plant.<br />

areoles: 3/4 to 17/8 inches apart. They are oval to round and<br />

small to medium sized on the sides of the pads, increasing to<br />

3/8 of an inch and becoming round on the edges of the pads.<br />

spines: There are 1 to 6 spines per areole on the upper 1/2 to<br />

almost all areoles of the pad. There is 1 main central spine<br />

which is only rarely missing, 1/2 to 2 inches long, porrect to<br />

deflexed in position, medium thickness to very heavy, very<br />

flattened or even triangular in shape. This spine is mostly<br />

black or blackish-brown, with usually a zone of gray or purplish-gray<br />

above and then a dark tip. There will usually be 2<br />

lateral spines very similar to this one, spreading and often<br />

curving outward. There will usually be 1 upper spine (which,<br />

however, may be missing) the same length as the others, erect<br />

or porrect in position, and similar to them except apparently<br />

always round. Below these main spines will be 0 to 4 lower,<br />

deflexed, slender, often bristle-like spines 1/4 to 3/4 of an inch<br />

long, usually gray with bases and tips somewhat blackish.<br />

glochids: Red-brown, brown, straw, or mottled in color.<br />

They are very variable in number, from almost none to very<br />

many on the sides of the pads, but almost always becoming<br />

many to very many and 1/4 to 3/4 of an inch long on the<br />

edges of the pads.<br />

flowers: As the species, except that the ovary is only 3/4 to<br />

1 inch long.<br />

fruits: Oval to goblet-shaped, with some slight constriction<br />

at the base and a rather deeply pitted apex. In size they are<br />

3/4 to 11/2 inches long. They become deep purplish-red. The<br />

seeds are 1/8 to almost 3/16 of an inch (3–4 millimeters) in diameter,<br />

flat, but usually very irregular in shape.<br />

Range. Apparently comprising a narrow but very long strip of


194 cacti of the southwest<br />

territory entering Texas from the south, where we have collected<br />

the plant in Coahuila, Mexico, and proceeding far north<br />

into New Mexico. The eastern and western boundaries of this<br />

strip are approximately as follows: entering Texas between Del<br />

Rio on the east and the Chisos Mountains on the west, proceeding<br />

northward past Fort Stockton on the east and Alpine on the<br />

west, then to the eastern foothills and the western slopes of the<br />

Guadalupe and Capitan mountains in southern New Mexico,<br />

and finally to the sides of the abrupt escarpment just east of<br />

Mosquero, New Mexico, in Harding County, which is its most<br />

northeasterly known range, and to the hills between Albuquerque<br />

and Cuba, New Mexico, which is its most northwesterly<br />

known penetration.<br />

Remarks: This cactus often forms dense thickets on the sides and<br />

tops of rocky ridges or steep slopes. It is a beautiful plant in the<br />

spring of the year when its pads are new and a wonderful bluish-<br />

green color, their smooth, glaucous surfaces contrasting with the<br />

heavy, blackish spines. But later in the summer, when the extreme<br />

heat and dryness have had their effect, one would hardly<br />

recognize it as the same plant, for its pads have become by then<br />

a yellowish-green from which they will not change for the rest<br />

of their lives.<br />

Because it grows in the type locality of that plant and because<br />

both have blackish coloring in the spines, this form has<br />

been widely mistaken for O. atrispina Gr., particularly since<br />

Britton and Rose seem to describe this plant under that name.<br />

However, any careful reading of Griffiths’ description of O.<br />

atrispina will show that this cannot be the cactus he had in<br />

mind, and even a casual comparison will separate the two.<br />

When we come to establish what this cactus really is, we find<br />

only one account in the literature which will fit it, and that is<br />

Engelmann’s description of O. phaeacantha var. nigricans. Admittedly<br />

his description of this form is so incomplete that we<br />

can only guess at what he had before him, yet we cannot ignore<br />

the statements of a botanists of his stature, so when we find a<br />

cactus fitting his description perfectly as far as it goes and still<br />

existing in the area of his type, we must presume that we have<br />

in this cactus the plant he meant. When we notice that the flowers,<br />

which he did not describe, are on our plant the large, orange-<br />

yellow blooms with red centers of the O. phaeacantha complex,<br />

and that the fruits fall within the limits of that species, then I<br />

feel that we can be certain we have before us the O. phaeacantha<br />

var. nigricans of Engelmann, and that it is a wide-ranging vari-<br />

ety of this very large and complex species.<br />

It has been the fashion for many years to regard this variety<br />

as merely a synonym of the species broadly considered, yet any<br />

reasonable amount of observation will show how different this<br />

form is from the other varieties here listed. The flower has a<br />

remarkably short ovary (about 3/4 of an inch long), which is<br />

distinctly different from the l1/4- to 11/2-inch ovary of some others<br />

in this complex. Its spines are as others in the group in<br />

having the main central flattened and the uppers round, and so<br />

on, but only variety nigricans has such dark colored, such heavy<br />

and such greatly flattened lower centrals, and no other form of<br />

the species has the heavy, flat, spreading laterals of this variety.<br />

It is interesting to note that almost every recent work which<br />

has dropped O. phaeacantha var. nigricans has then described<br />

this plant separately under the name of O. atrispina.<br />

After studying both her description and her preserved specimens,<br />

I believe that Miss Anthony’s O. engelmannii X phaeacantha<br />

of her Big Bend study is this cactus. I have heard it<br />

called such a hybrid by others, and if one likes to speculate on<br />

hybrids, this would be a likely candidate. The bluish-green of<br />

its young pads, the possession of heavy, curving lateral spines,<br />

the grayness instead of yellowness of the upper parts of its<br />

spines, much about its fruits and seeds all remind one of O.<br />

engelmannii, but no form of O. engelmannii has red-centered,<br />

yellow flowers. Here again experimental work with these plants<br />

to establish something of their genetics is all that would clarify<br />

things further, and renaming various forms as hybrids before<br />

we even know whether the species involved can interbreed<br />

would not seem really to advance our understanding.<br />

On the tops of many limestone ridges from just west of Fort<br />

Stockton, Texas, to around Sanderson, this plant grows together<br />

with O. strigil, the two such contrasting Opuntias making a<br />

strikingly beautiful sight together.<br />

Opuntia phaeacantha var. brunnea Eng.<br />

Description Plate 52<br />

stems: To about 3 feet tall, consisting of upright branches<br />

without a central trunk. Pads 4 to 8 inches long, round or<br />

even wider than they are long, usually with no constriction<br />

below, but sometimes broadly egg-shaped with some lower<br />

constriction. Their color is slightly blue-green when very<br />

young, when older glaucous yellow-green, usually with some<br />

purplish-red mottling on the edges of the pads and these often<br />

suffused with this color in winter. The pads are rather thick.<br />

areoles: Small to medium and elongated to almost round on<br />

the sides of the pads, enlarge to about 3/16 of an inch or so<br />

and become almost round on the edges of old pads. They are<br />

spaced 3/4 to 15/8 inches apart.<br />

spines: 1 to 5 per areole on only the upper edge or to at most<br />

the upper third of the pad. The main central spine is 1 to 3<br />

inches long, porrect or deflexed, straight or often twisted and<br />

bent, medium to heavy, and round to slightly flattened. There<br />

will usually, but not always be 1 to 3 upper spines, 1 to 21/2<br />

inches long, erect or spreading upward, medium to very<br />

heavy, round to very flat and twisted. These main spines are<br />

all blackish, chocolate-brown or red-brown at the bases, their<br />

upper parts tan or grayish. There may be 1 or 2 lower, deflexed,<br />

slender, and often bristle-like spines, 3/16 to 1 inch<br />

long. These are either round or flattened, and gray in color.


genus Opuntia 195<br />

glochids: Brown to straw in color, almost none to an average<br />

number but very short on the sides of the pads. On the edges<br />

there are few at first, later often increasing to very many and<br />

growing to 3/4 of an inch long.<br />

flowers: As the species.<br />

fruits: 11/4 to 21/2 inches long, oval to club-shaped with some<br />

constriction below and a shallowly concave umbilicus above.<br />

They are dull red in color. The seeds measure 3/16 of an inch<br />

or a little more (41/2–51/2 millimeters) in diameter.<br />

Range. Southwest Texas from the Anacacho Mountains east of<br />

Del Rio along the Rio Grande through the Big Bend at least to<br />

El Paso, Texas. Also northward from the Big Bend through the<br />

Davis Mountains to the southern parts of the Guadalupe and<br />

Organ mountains, where it enters New Mexico a short distance.<br />

Remarks. Engelmann had an O. phaeacantha var. brunnea from<br />

El Paso, which was never redescribed after his time and which<br />

was apparently never taken seriously by anyone except Griffiths.<br />

In Griffiths’ preserved collection I find very good specimens<br />

of this form collected at El Paso and at Fort Hancock and<br />

labeled by him with this name, but relabeled otherwise since<br />

then by others. I find the variety quite abundant in the Anacacho<br />

Mountains, growing here and there around Langtry,<br />

Texas, and on into the eastern Big Bend. It becomes quite numerous<br />

again west of Terlingua, Texas, and continues all along<br />

the Rio Grande to El Paso. There are some fairly good stands of<br />

it in the Davis Mountains and scattered individuals along the<br />

Texas-New Mexico line at the Guadalupe, Hueco, and Franklin<br />

mountains, where it enters New Mexico for a short distance.<br />

Engelmann described the form very incompletely, but did<br />

mention most of the important features which distinguish it<br />

from the other varieties of the species. The greater glaucescence,<br />

thicker joints, and purplish-red winter coloring he mentions are<br />

all important points in recognizing it. He remarks that the spines<br />

are longer than typical for the species, and with the spines<br />

usually to 3 inches long this is important. He calls the spines<br />

greatly angled, which is true of most of the main ones, but in<br />

this cactus one notices that it is the upper centrals which are the<br />

most greatly flattened instead of the lower central-a reversal<br />

of the usual order in the species. Beyond this he does not go,<br />

but the flowers and fruits are typical for the species, and the<br />

seeds larger than those of most of the other varieties.<br />

The coloring this plant assumes in severe weather, when linked<br />

with the 3-inch spines, misleads many observers into thinking it<br />

is O. macrocentra. When I first realized the existence of this<br />

form, I checked back through those of my own specimens which<br />

I had called O. macrocentra and found that I had included several<br />

of this cactus under that name. I am convinced that herbarium<br />

collections filed under O. macrocentra often contain<br />

specimens of this cactus, and that some descriptions of that plant<br />

have been overly broadened and made almost useless because<br />

of this confusion.<br />

To distinguish these two cacti from only vegetative parts is<br />

not always easy, but one clue is that the coloring of O. phaeacantha<br />

var. brunnea is not ever actually purple, as is that of the<br />

other, but really a dull, dark, purplish-red. In gardens of west<br />

Texas collectors I have several times had my attention called to<br />

the fact that one of a collector’s O. macrocentra specimens will<br />

be a beautiful purple while the other one beside it will be a very<br />

different purplish-red. This is because they have collected for<br />

that plant a specimen of this variety which is all but unknown<br />

today after reposing in the synonym of O. phaeacantha for 100<br />

years.<br />

There are other differences besides differences of hue, but they<br />

take close observation. The pads of variety brunnea are thick,<br />

while those of the other are thin. While the areoles of this variety<br />

are 3/4 of an inch or more apart, with an average over 1<br />

inch apart, those of the other species are always less than 1 inch<br />

apart. The spines of this variety are long, but never as long as<br />

the maximum on O. macrocentra. Although they easily equal<br />

in length those on some less robust specimens of that other species,<br />

they are more brown than black, and are rigid rather than<br />

flexible like those others. The flowers of the two are very similar,<br />

but the fruits are different. O. macrocentra’s fruits are to<br />

only 11/2 inches long, with no constriction below, deeply pitted<br />

above, and bright red to orange-red. Those of variety brunnea<br />

are to 21/2 inches long, with some constriction below, shallow<br />

umbilicus, and dull, dark red color.<br />

It is my opinion that when Britton and Rose gave for O.<br />

macrocentra fruit to 23/8 inches long and purple, they did it<br />

because they had included in their specimens some which were<br />

actually of this variety. This idea seems to be borne out by the<br />

figure they featured to illustrate O. macrocentra. I could never<br />

visualize O. macrocentra from their figure, while it is a passable<br />

representation of an O. phaeacantha var. brunnea pad. More recently<br />

both Backeberg and Earle have followed in giving such<br />

overly broadened descriptions of O. macrocentra.<br />

Anthony’s O. tenuispina from the Big Bend also appears to<br />

be this cactus. There are reasons to think that Rose’s O. chihuahuensis<br />

may also have been this variety, but it is so incompletely<br />

described that one cannot be sure what it was. Wooton and<br />

Standley considered the plants of the O. phaeacantha complex<br />

found in the Organ Mountains just within New Mexico to be<br />

O. chihuahuensis, saying that they exactly match the type specimens<br />

of Rose’s plant. They also say that a specimen from that<br />

location classified by Coulter himself as his O. mesacantha var.<br />

oplocarpa is the same. Specimens that I have collected in this<br />

area exactly match the description given above for variety<br />

brunnea, and the evidence seems good that both of the above<br />

are this variety. These plants are certainly different from the<br />

Colorado plants called variety oplocarpa by Coulter. If that is<br />

a valid variety at all it apparently does not enter New Mexico<br />

from Colorado.<br />

The question of how far variety brunnea ranges into New<br />

Mexico is a hard one. Assuming that the above-mentioned plants<br />

are synonyms, Wooton mentions them from only the Organ


196 cacti of the southwest<br />

Mountains, but Britton and Rose talk as though they were more<br />

widespread over southern New Mexico, extending north out of<br />

Chihuahua. I have seen this variety in New Mexico only in the<br />

Guadalupe and Organ mountains, and this not over 40 miles<br />

or so from the border. I have never seen this plant west of El<br />

Paso, but recently I was sent a specimen from the Santa Rita<br />

Mountains of Arizona which appears to be it. Whether further<br />

study will enlarge its range westward remains to be seen.<br />

Opuntia phaeacantha var. camanchica (Eng.)<br />

Description Plate 53<br />

roots: Although in most cases the roots are fibrous, occasional<br />

specimens have tubers upon their roots.<br />

stems: Growing as a low, semiprostrate, spreading cactus. It<br />

produces large pads, but these usually lie on their edges by the<br />

second season to form chains of reclining pads. By the rooting<br />

of these pads there are formed clumps sometimes 3 to 5 feet<br />

across. The pads are often 8 or 9 inches in length, so a leaning<br />

branch may rise to a foot or even a little higher, but this is<br />

about the maximum height. The pads vary from perfectly<br />

round through egg-shaped, to occasionally broad club-shaped<br />

with constricted bases. They vary from 3 to 9 inches long,<br />

with slightly less width. The average size would be about 6<br />

inches long by 5 inches wide. The surfaces are flat and the<br />

thickness medium, which is slightly less than 1/2 inch thick.<br />

They are rather deep green in color with some glaucescence,<br />

sometimes taking on a reddish cast around the areoles and on<br />

the edges during the winter.<br />

areoles: These are oval or oblong to round. They are about<br />

3/16 of an inch across on the sides of the pads, enlarging to<br />

about 1/4 of an inch on the edges. They are from 3/4 to 11/2<br />

inches apart.<br />

spines: There will be from 1 to 8 spines on the upper 3/4 or<br />

more of the areoles, only the lowest areoles not bearing spines.<br />

The lowermost spine-bearing areoles usually have one spine,<br />

with the number increasing upward until the very upper and<br />

edge areoles have 3 to 8 spines. A fully armed areole will<br />

contain 1 to 5 straight upper spines 3/4 to 3 inches long. These<br />

spread in all directions upward and outward, are of medium<br />

thickness, entirely round on some specimens, flattened on<br />

others, their bases and usually the lower half brown, redbrown<br />

or blackish-brown, followed by a zone of tan, gray,<br />

purplish-gray, or whitish, and ending in yellowish, semitranslucent<br />

tips. It will also contain 1 to 5 lower spines turning or<br />

spreading downward from the lower part of the areole. The<br />

largest one of these may be up to 11/4 inches long, flattened,<br />

and rather heavy, but the rest of them will range smaller in<br />

size and thickness until they are hardly more than deflexed<br />

bristles. All of these lower spines are whitish or gray with<br />

brown tips.<br />

Plants not vigorous for some reason, such as those growing<br />

in shade, may put out pads as big as those on normal, robust<br />

plants, but will be deficient in spines. It is common then for<br />

only the upper third of the areoles to bear spines at all, and<br />

for these to have only 1 to 3 spines. In this case there will be<br />

2 lower, deflexed spines 3/8 to 3/4 of an inch long, gray with<br />

lighter tips on most of the armed areoles, and only an occasional<br />

areole on the very edge may have 1 of the heavier,<br />

brown upper spines to about 11/4 inches long.<br />

glochids: Somewhat variable. Some specimens will have almost<br />

none, some an average number, and sometimes on old<br />

pads of the same plant they may become very conspicuous.<br />

From 1/8 of an inch or nearly that short on the sides of the<br />

pads to at least 3/8 of an inch long on the edges of old pads.<br />

They are straw, tan, or brownish in color.<br />

flowers: As the species, except longer, with the ovary 11/4 to<br />

13/4 inches long.<br />

fruits: Bright cherry-red to dull plum-red when ripe, with<br />

juicy, pleasant-tasting flesh. Oval in shape, not constricted<br />

at the base, 1 to 21/2 inches long by 3/4 to 11/2 inches wide. The<br />

umbilicus is a rather narrow surface which is flat or only very<br />

shallowly concave. The seeds are 3/16 to 1/4 of an inch (41/2–6<br />

millimeters) in largest measurement. They are very irregular in<br />

shape, perfectly flat and thin, or much twisted and thicker,<br />

round to square or even sometimes almost triangular. They<br />

have conspicuous rims 1 to 11/2 millimeters wide, but sharp.<br />

These rims are usually wavy and irregular.<br />

Range. Out of Kansas and Colorado to the north over western<br />

Oklahoma, Texas, and all of eastern New Mexico. More specifically,<br />

it occurs in Oklahoma west of a line running approximately<br />

from Blackwell to Ardmore, and in Texas west of a line<br />

from near Fort Worth south to the Balcones Fault in the vicinity<br />

of San Antonio. It is not known to grow south of San Antonio,<br />

and occurs rather commonly east of the central mountains in<br />

New Mexico, but is seldom if ever seen west of central New<br />

Mexico.<br />

Remarks. This is the smallest, most prostrate, most spiny version<br />

of the species. It occurs primarily east of the more upright,<br />

bushy varieties’ mountain ranges, preferring to grow on the<br />

high plains and hills east of the mountains. In their healthy,<br />

robust, typical forms these varieties are easy to tell apart. In<br />

immature, stunted, atypical conditions it is often impossible to<br />

distinguish them one from the other. This is, however, true of<br />

most Opuntias, even separate species. When he undertook the<br />

study of our cacti, Engelmann found it necessary to distinguish<br />

this form. He listed it as a separate species, but he remarked that<br />

it differed from O. phaeacantha “only in habit.” This is essentially<br />

true, but exactly what that means and what significance<br />

there is to it cannot really be told even yet, 100 years later.


camanchia -><br />

camanchica<br />

genus Opuntia 197<br />

It is true that the upright growth of the other varieties and<br />

the prostrate growth of this variety are hardly definite enough<br />

characters upon which to separate species. Shade or poor growing<br />

conditions will render them all almost identical in a couple<br />

of seasons. This smaller cactus called variety camanchica is also<br />

normally more spiny than its larger relatives, but the same poor<br />

conditions will cause it to develop fewer spines. Fruit shape<br />

seems a minor point also, yet minor as these things may be individually,<br />

when taken together they add up to two clearly different<br />

cacti. No matter how closely they may coincide when<br />

they can’t grow normally, when they are grown side by side in<br />

the same good environment, they are separate entities. I have<br />

seen this in a well-arranged old garden, where on one hand was<br />

upright O. phaeacantha var. major 3 to 4 feet high and lightly<br />

spined, while 6 feet away a plant of prostrate O. phaeacantha<br />

var. camanchica made an impenetrable mat, about 1 foot high<br />

covered with its much more numerous spines. Fruits were characteristically<br />

different on these two plants also.<br />

So there is a difference, even though, beginning with Britton<br />

and Rose, numerous authorities have made these and the other<br />

forms listed here as varieties synonymous. Borg has stood almost<br />

alone in setting this cactus off from the others in the species<br />

as a separate subspecies. No one quite knows, even yet, the<br />

basis for these differences or what their significances are. The<br />

kind of differences we see do not seem great enough to fit our<br />

modern ideas of separate species, and so we agree in the downgrading<br />

of such as this form from that of species rank. But then<br />

what is it, if not a species? Is camanchica an intermediate of<br />

some kind? Is there a hybridization, and if so between what<br />

forms? Or are these merely individual variations within a species?<br />

There is even yet not one particle of evidence, no breeding<br />

records, no hybridization experiments, no genetic studies on<br />

these plants, so we cannot answer such questions as these. We<br />

only know that we are faced with the separate forms and need<br />

a way to refer to them. Hesitating to call them species, suspecting<br />

that they are in some way more closely related than that,<br />

we use the only other rank currently in vogue for such close<br />

relationships, and our cactus becomes O. phaeacantha var. Camanchica.<br />

This then, is the prostrate but large-jointed, heavy-spined<br />

prickly pear which Engelmann found growing on the Llano<br />

Estacado, which we know is common in western Oklahoma,<br />

northwestern Texas, the panhandles of both these states, and<br />

into New Mexico, and which Coulter stated grew “from southern<br />

Colorado through western Texas, New Mexico and Ari-<br />

zona.”<br />

As might be expected in any form with such a wide range,<br />

there are many minor variations in local races. There have been<br />

attempts to give various of these species or varietal rank, but it<br />

does not seem they can be maintained. Much confusion has resulted.<br />

Some of these local forms which can best be regarded as<br />

synonyms but which are often referred to separately should be<br />

mentioned here to help a beginner past the confusion.<br />

In central Texas a form with somewhat fewer spines than<br />

typical and more often having tuberous roots, but otherwise<br />

indistinguishable from this variety was described and named by<br />

Rose O. mackensenii. Because of its occasional tuberous roots he<br />

seemed to associate it with O. compressa rather than the O.<br />

phaeacantha group, but otherwise it is clearly of this complex.<br />

It is the form seen commonly around Austin, Texas, and west<br />

through the Texas hill country on the southern edge of the variety’s<br />

range. Perhaps its fewer spines may be due to marginal<br />

growing conditions on the edge of the range.<br />

O. phaeacantha var. camanchica has often been called. O.<br />

tortispina, a name coined by Engelmann for a form with spines<br />

lighter-colored and more twisted than is typical. The two cannot<br />

be separated, and I have a number of times, in northwestern<br />

Oklahoma, seen the two supposedly different forms growing<br />

together along with all sorts of intermediates between them. The<br />

lighter form is more common in northern Oklahoma and Kansas.<br />

Miss Anthony unaccountably uses this name for he variety<br />

with the regular dark spines as it is found in the Big Bend.<br />

There grows in Colorado the nearest thing we have seen to an<br />

intermediate between variety camanchica and the upright varieties<br />

of O. phaeacantha. These Colorado plants have the low-<br />

growing form of this variety, but far fewer spines as well as<br />

elongated, clavate fruits and small seeds such as it never has.<br />

Most of the characters of this Colorado plant are like those of<br />

variety major, except that it lacks the size and erectness of that<br />

variety. It seems to be the form described by Coulter as O.<br />

mesacantha var. oplocarpa, but may represent only the effect<br />

of the severe northern climate and altitude on variety major. At<br />

the same time, variety camanchica grows very spiny and very<br />

typical up along the east side of the Colorado mountains. However,<br />

we have seen a few specimens from Oklahoma and north-<br />

west Texas of variety camanchica which approached this Colorado<br />

form by having longer than typical fruits. Although it is<br />

so incompletely described that no one can be sure, O. pyrocarpa<br />

Gr. may have been such a variety camanchica with minimum<br />

spines and long fruits.<br />

Indistinct and problematical as variety camanchica may be<br />

to a taxonomist, it is very real and very much a plant to be<br />

taken into consideration over huge areas of the high plains. Here<br />

it is the largest Platyopuntia to be found, and often quite a<br />

pest. The grass it protects from grazing animals often grows up<br />

to practically hide it, but everything moving must be alert for<br />

it or suffer the consequences of blundering into it.<br />

Opuntia phaeacantha var. tenuispina (Eng.)<br />

Description Plate 53<br />

roots: Previous accounts did not describe the roots, except<br />

for Wooton, who said they were sometimes tuberous. Roots of<br />

those living plants examined in this study were fibrous.


198 cacti of the southwest<br />

stems: A low, semiprostrate, spreading plant. Occasionally<br />

the most ascending branches of the plant may extend to about<br />

1 foot high. The pads are 3 to 8 inches long and to 4 inches<br />

wide, pear-shaped, with the bases noticeably constricted. The<br />

pads are rather thin, smooth, and dark blue-green or bright,<br />

shiny green in color.<br />

areoles: The areoles are oval to practically round, and from<br />

slightly over 1/8 to about 3/16 of an inch across. They are closely<br />

set on the pads, varying from 3/8 of an inch apart low on<br />

the pad to one inch apart higher up. Some full-sized pads<br />

have no areoles over 3/4 of an inch apart.<br />

spines: There are found 1 to 6 spines on all but the very lowest<br />

areoles of the pad. Areoles up to about the middle of the pad<br />

will carry 1 to 3 slender, whitish spines from a mere fraction<br />

of an inch to as much as 1 inch long. These all grow downward<br />

and lie spread almost against the pad surface. The most<br />

fully armed areoles around the upper edge of the pad will<br />

carry 1 or 2 main spines 1 to 21/2 inches long, usually white<br />

or gray, always with brown tips and often with pale brownish<br />

bases. They are usually turned severely downward, but occasionally<br />

one may stand fairly erect on the edge of the pad.<br />

These main spines are straight, slender, round or practically<br />

so, firm instead of bristle-like, but slender enough to be easily<br />

flexible. Below them in the areole will be 1 to 4 shorter, de-<br />

flexed spines like those found lower on the pad.<br />

glochids: Bright reddish-brown fading to dirty brown, rather<br />

numerous, forming a dense tuft in the center of the areole,<br />

but very slender and short, being only 1/8 of an inch or slightly<br />

longer on any areole. They hardly extend beyond the<br />

brown wool of the areole. The effect of this is to give the<br />

areoles the appearance of bulging outward.<br />

flowers: As the species, except only 21/2 to 3 inches across,<br />

and when old fading gradually to a pale pinkish. There are 6<br />

to 8 green stigma lobes.<br />

fruits: Bright red in color, oblong or elliptic, with little if<br />

any constriction at the base, but with a very deeply pitted<br />

umbilicus. From 1 to 15/8 inches long and about 3/4 of an inch<br />

thick. There are usually brown glochids on the areoles of the<br />

fruits, and sometimes the upper edge areoles have 1 to 3 slender,<br />

whitish spines, tipped brown and up to 3/8 of an inch long,<br />

on them. The seeds are very irregular in outline, sometimes<br />

round and almost even, but more often elongated in one axis,<br />

twisted, or otherwise unevenly shaped. This gives quite a variation<br />

in measurement, individual seeds from the same plant<br />

ranging from hardly more than 1/8 to 1/4 of an inch (3–6 millimeters)<br />

in greatest diameter. They are of average thickness.<br />

Range. Originally stated to be limited to the sandhills around<br />

El Paso, from the village of Dona Ana, above Las Cruces, New<br />

Mexico, to the village of San Elizario below El Paso, in Texas.<br />

In spite of numerous claims to a broader range, there seem to be<br />

no indisputable records of the plant outside of this limited area.<br />

Remarks. In his study of cacti 100 years ago, Engelmann described<br />

and named a small prickly pear growing on the sand-<br />

hills around El Paso. It had several distinctive characters, namely<br />

its constricted pads, its rather numerous, light-colored, and<br />

slender, flexible spines which were mostly pointed downward<br />

on the pads, its bright brown or reddish-brown but short<br />

glochids, its close-standing areoles, and its oblong, deeply-pitted<br />

fruits. Engelmann considered it a separate species and called it<br />

O. tenuispina.<br />

Coulter repeated Engelmann’s description and range for the<br />

plant, apparently no other specimens of it having come to light<br />

in the 40 years intervening before his own study but those first<br />

seen by Engelmann. It appears from this that it was a rare plant<br />

limited to the habitat of the Rio Grande Valley sands around<br />

El Paso and Las Cruces.<br />

In his own accounts a little later, however, Wooton made a<br />

strange statement which opened the gates to much confusion,<br />

saying, “This is the most common species in the lower Rio<br />

Grande Valley on the heavier soils.” How he transferred the<br />

cactus from the sand hills to the heavier soils is unknown, and<br />

the photograph he reproduced for the plant is so poor that it<br />

does not tell us whether he was confusing this plant with another<br />

or not.<br />

At any rate, Britton and Rose apparently did become so confused.<br />

Although they saw the real thing and give us probably<br />

the best picture of the cactus so far existing, their description<br />

shows that they also included with it something quite different,<br />

something with fewer spines. Having brought in some other<br />

plant, they included this other plant’s range and said the species<br />

even grows west into Arizona.<br />

Following them, the range has been enlarged by other writers<br />

until the plant has sometimes been considered a common cactus<br />

of all southern New Mexico and Arizona. Finally Anthony<br />

called a rather large, bushy cactus of almost entirely different<br />

description growing in the Big Bend of Texas O. tenuispina. We<br />

have concluded that her plant was instead O. phaeacantha var.<br />

brunnea.<br />

Nothing can be definite until the question is decided: is O.<br />

tenuispina the rare plant of very limited habitat it at first appeared<br />

to be, or is it really a common plant of wide range? Is<br />

the broadening of range and description a natural result of<br />

wider study or is there another cactus close enough in general<br />

characters to have become confused with it and so explain the<br />

increase of range?<br />

A partial answer to the question can be found by looking at<br />

Rose’s own specimens as they are still preserved. Upon searching<br />

them out, we find that all of the specimens he called by this<br />

name from outside the original small range have been redetermined<br />

by other students since as ordinary O. tortispina. This is<br />

the white-spined form O. phaeacantha var. camanchica sometimes<br />

takes, and I believe some of the confusion has arisen be-<br />

cause of mistaking this cactus for O. tenuispina.


genus Opuntia 199<br />

There is an even better explanation for the trouble with this<br />

cactus. One of the problems of Opuntia study is the fact that<br />

most Opuntias, when immature or not growing in conditions<br />

very favorable to them, fail to put on their complete spination.<br />

The very characters of spines upon which so much depends in<br />

recognizing them do not show at all on such stunted specimens.<br />

Anyone who has tried to collect and grow Opuntias knows that<br />

all too often his various beautiful types, painstakingly gathered,<br />

end up in a few years of cultivation practically indistinguishable.<br />

And he will find that all of the O. phaeacantha group show in<br />

this imperfect growth form a few slender, whitish, rather deflexed<br />

spines and lack their main and heavy upper spines. If too<br />

little light is part of the trouble there is also a tendency for the<br />

pads of all of them to lengthen and produce narrowed bases.<br />

Now this imperfect form of all these varieties happens to fit<br />

somewhat nearly the description of O. tennispina, especially the<br />

very general and simplified descriptions of Wooton and Britton<br />

and Rose, which have been the ones most widely copied by recent<br />

writers. The details of areoles and so on, having been left<br />

out of their descriptions and so not being referred to, it has been<br />

easy to imagine that many weakly spined specimens of the other<br />

O. phaeacantha varieties were this cactus. In this way, we believe,<br />

the supposed range of O. tenuispina has grown and grown,<br />

while the conception of the plant has become diluted so that<br />

more and more plants have been taken in.<br />

The actual O. tenuispina, however, when properly recognized,<br />

is one of our rarest prickly pears, and it is now probably very<br />

near extinction. Most of the sandy strip along the Rio Grande<br />

where it had grown is now included in the remarkable megalopolis<br />

which is fast fusing Las Cruces, El Paso, and Juarez, and<br />

most cacti there are already gone or appear doomed as the<br />

build-up proceeds across the sands toward the slopes of the<br />

Franklin Mountains. It may be that in the future there will be<br />

no further opportunity to study the problem of this form.<br />

I have not been so fortunate as to collect this cactus myself,<br />

in spite of quite a lot of wandering around the sands of<br />

its area. However, I am indebted to Mr. Charles Polaski, the<br />

remarkable collector and grower of cacti who has been the<br />

source of most of the Oklahoma specimens in European collections,<br />

for making it possible for me to describe and photograph<br />

the plant. He has a whole bed of the variety grown from a collection<br />

he made, “west of El Paso.” We feel most certain it is the<br />

plant in question, and it may represent the only living specimens<br />

in cultivation today. It represents what we feel to be at least a<br />

very distinct variety with definite form not due to stunting or<br />

other environmental influence. It has kept its characteristics unchanged<br />

both in Mr. Polaski’s yard in the rather severe Oklahoma<br />

climate, and in my garden in San Antonio. I feel most fortunate<br />

to be able to include it in this study at all, even though<br />

I cannot give any further data on presently existing wild populations.<br />

Perhaps this will prompt someone to locate and study<br />

them further, if they still exist at all.<br />

It is my opinion that this form is the smallest, most low-<br />

growing of the O. phaeacantha complex, and so I list it here as<br />

a variety of that species.<br />

Opuntia cymochila Eng.<br />

Description Plate 53<br />

roots: Described as fibrous, and usually so, but occasional<br />

examples with tuberous roots have been found.<br />

stems: Prostrate. Growing as a spreading mat of small pads<br />

rooting where they lie either flat on the ground or resting on<br />

their edges. The height of the clump will be the height of the<br />

most upright pad, which may be 6 or 8 inches, while the width<br />

of the clump is usually 1 to 4 feet. Pads are usually round or<br />

sometimes even wider than they are long, but occasionally become<br />

slightly egg-shaped. These pads are rather thick for their<br />

size, averaging about 1/2 inch thick. They are from 21/2 to<br />

occasionally 6 inches long and to 4 inches wide. In the dry<br />

season or in the winter the pads are much shrunken and take<br />

on a very wrinkled appearance, but in a moist growing season<br />

they fill out and become flat. They are deep glossy green<br />

when filled with moisture, but when dry or suffering from<br />

freezes, they become lighter green often suffused with a reddish<br />

color.<br />

areoles: These are oval or round, small or medium in size,<br />

and usually 3/8 to 3/4 of an inch apart, but have been found<br />

1 inch apart.<br />

spines: Robust plants will have a total of 1 to 7 spines on<br />

almost all areoles or sometimes on those of only the upper<br />

half of the pad. Of these spines, 1 to 3 are main central spines<br />

which are 1 to 23/4 inches long, fairly heavy, round to flattened<br />

and often twisted, but straight. Most of these spread in<br />

a downward direction, but sometimes one or two of them<br />

may stand straight out from the areole or even ascend. From<br />

the lower part of the areole there will spread downward 1 to<br />

5 small spines 3/16 to 1 inch long. These are slender, almost<br />

bristle-like to fairly stout and very slightly flattened. All<br />

Spines are white or gray with the very tips light brown and<br />

translucent, and the bases brownish. Occasionally the brownish<br />

coloring extends up most of the length of the spine. Less<br />

healthy plants growing in excessively dry or exposed places<br />

and all new pads before maturity will lack the main upper<br />

spines entirely and will present only the lower, deflexed<br />

spines, these 1/4 to 1 inch long, slender and bristle-like, and<br />

pure white with brownish tips.<br />

glochids: Yellow, straw, or dirty brownish-yellow in color.<br />

They are not too numerous in young areoles, but usually increase<br />

to very many in old areoles. They stand out in a com-


200 cacti of the southwest<br />

pact cluster in the middle of the areole and are comparatively<br />

long, 3/16 of an inch at first, becoming often 1/2 inch long on<br />

old pads.<br />

flowers: About 3 inches in diameter and 21/2 inches tall. Usually<br />

wholly chrome-yellow in color, but sometimes with the<br />

center golden-brown. The outer petals are short and pointed,<br />

the inner ones broad at the ends with a tiny point at the apex.<br />

The stamens are very pale yellow, the style long and pale<br />

green. There are 9 green stigma lobes.<br />

fruits: Purplish-red in color, oval or broadly egg-shaped, 1<br />

to 11/2 inches long and 3/4 to 1 inch in diameter at the thickest<br />

point. The base is not constricted at all, or only very slightly<br />

so on some plants. The umbilicus is narrow and forms a deep<br />

pit. The seeds are large, a little over 3/16 to just over 1/4 of an<br />

inch (5–61/2 millimeters) in diameter, of medium thickness,<br />

irregular in shape. In the same plant there will be seeds practically<br />

round, elongated, and almost triangular, and sometimes<br />

they are greatly twisted. The body of the seed is depressed<br />

in its center with a ridge around its edge. The rim of<br />

the seed is wide to very wide (1–2 millimeters), but very<br />

sharp. It often undulates, but there is no notch at the hilum.<br />

Range. The whole of the Oklahoma and Texas panhandles and<br />

adjacent New Mexico west to the mountains and south into west<br />

Texas. Extending east from the panhandles about as far as<br />

Cheyenne, Woodward, Alva, and Manchester, Oklahoma, and<br />

north into west and central Kansas and eastern Colorado. It<br />

also grows south in Texas to a line running from near Abilene<br />

approximately to Big Spring and Pecos, then dipping sharply<br />

south past Fort Davis to just south of Marfa, the southernmost<br />

point at which the cactus has been recorded. From there the<br />

range retreats northward abruptly, entering New Mexico east<br />

of the Sacramento Mountains.<br />

Remarks. O. cymochila was first described and named by Engelmann<br />

as a small cactus of the Comanche Plains. When he<br />

later erected his large species O. rafinesquii he included this<br />

plant in it as O. rafinesquii var. cymochila. When Coulter substituted<br />

mesacantha for rafinesquii, he left our cactus as O. mesacantha<br />

var. cymochila. These men would, therefore, connect<br />

this cactus with O. compressa, which is the presently accepted<br />

name for that widespread Opuntia complex.<br />

Wooton, in his study of New Mexico cacti, says instead that<br />

the cactus, except for its having succulent instead of dry fruits,<br />

resembles O. polyacantha.<br />

Britton and Rose, in their study, made O. cymochila a synonym<br />

of O. tortispina Eng. and were followed in this by Borg.<br />

Backeberg connected the two, but still distinguished between<br />

them and so has O. tortispina var. cymochila. Since both Boissevain<br />

and Benson consider O. tortispina closely related to or<br />

even a variety of O. phaeacantha, a view with which I agree,<br />

this gives us yet another way of relating this plant.<br />

There are plausible reasons for each of the above combina-<br />

tions, yet obviously they can’t all be correct. The cactus remains<br />

recognizably distinct from them all, and since there seem to be<br />

about as many reasons for one combination as for another, it is<br />

probably better not linked any more closely to any one of them<br />

than to the others. We therefore leave it standing separate among<br />

them until we have better evidence for one relationship or<br />

another.<br />

The mature and robust O. cymochila is easily distinguished<br />

from the actual O. compressa and all the varieties of that cactus,<br />

some of which may grow in its range, by the possession of<br />

spines on all or nearly all areoles. O. compressa and its varieties<br />

have spines only on the upper parts of the pads. This seems a<br />

simple distinction, but a chance for confusion enters with the<br />

fact that young or stunted specimens of O. cyrnochila often lack<br />

the major spines, and the appearance of this weakly bespined<br />

growth is almost identical to that of some forms of O. compressa.<br />

I have collected plants, particularly just north of Big<br />

Spring, Texas, which seemed perfectly good examples of O.<br />

compressa until grown for a couple of years in good conditions,<br />

when they put forth more numerous spines than that cactus ever<br />

has. These plants, by then, had also displayed the fruits of O.<br />

cymochila, rather oval fruits with little or no constriction of the<br />

bases and with very large, relatively thin seeds. These features<br />

are greatly different from the narrow, clavate fruits of the other<br />

plant, with its much smaller, very thick seeds.<br />

It is obvious that O. cymochila is distinct from the dry, spiny-<br />

fruited O. polyacantha, and I do not believe that Wooton ever<br />

meant they should be combined, but it is surprising how closely<br />

a stunted plant of O. cymochila with pads all shrunken and<br />

with only depressed, white spines can approximate this other<br />

species when seen on some dry Canadian River bluff. Here<br />

again, a chance to grow well and put out its real spination as<br />

well as its purplish, naked, fleshy fruits will show it as so entirely<br />

different from the other with its dry, spiny fruits that<br />

in one’s garden it is hard to believe they could ever have looked<br />

alike.<br />

The problem of relating this cactus properly to the O. phaeacantha<br />

group is more complex. It might have fallen in line<br />

neatly as a smaller relative of O. phaeacantha var. camanchica,<br />

much of whose range it shares, except for the interjection of<br />

another name and description which has brought nothing but<br />

confusion for this area.<br />

At the same time and in the same publication in which he<br />

first put forth O. cymochila, Engelmann also described an O.<br />

tortispina. He gave the same range for the two, the Comanche<br />

Plains. Coulter repeated the description of the latter, but said<br />

that O. tortispina grows all the way to Nebraska. Significantly,<br />

he did not combine it as a variety of his O. mesacantha (O.<br />

compressa), as he did O. cymochila. Wooton ignored O. tortispina<br />

entirely, but Britton and Rose took this to be the proper<br />

name for the common cactus growing from Wisconsin and Dakota<br />

all the way to New Mexico, and decided O. cymochila was<br />

the same plant. They were followed in this by others. We must


genus Opuntia 201<br />

take time to see what O. tortispina was before we can determine<br />

if it is our plant or how they are related.<br />

Engelmann’s description of O. tortispina had some characters<br />

in common with O. cymochila. He said it was a practically<br />

prostrate cactus with round or nearly round pads. It had straw-<br />

colored or brownish glochids. It had upper spines 11/2 to 21/2<br />

inches long and white or white with yellowish tips and bases.<br />

Its fruits were oval and not constricted. All this is similar in the<br />

two plants. But in size of pad O. tortispina was said to be larger<br />

than O. cymochila. Engelmann had before him pads 6 to 8<br />

inches long with areoles 1 to 11/2 inches apart. This larger plant<br />

also had a total of 6 to 8 spines, 1 or 2 more than a robust O.<br />

cymochila has. Much has been made of the fact that Engelmann<br />

said his O. tortispina had spines compressed, angled, sometimes<br />

even channeled and twisted, and named the plant for this feature.<br />

Its fruits were also longer, being 13/4 to 2 inches long, with<br />

at most a shallow depression at the top.<br />

Everyone remembers the twisted-spine character, but most of<br />

the other characters Engelmann gave for that plant have been<br />

ignored. I believe a careful reading of the description shows that<br />

Engelmann’s O. tortispina has characters found quite often in<br />

O. phaeacantha var. camanchica, but except for the whitish<br />

spine color, its characters really give us a much more robust<br />

plant with larger pads, more widely spaced areoles, and fruits<br />

and seeds more similar to that cactus than to O. cymochila.<br />

This leaves us with O. cymochila as a distinct form occurring<br />

over a vast area of the high western plains. Where it grows in<br />

more hilly country it still is usually found on the flats of the<br />

valleys rather than on steep slopes. It often dots the flat plains<br />

of the panhandles where the sod has not been broken. It has<br />

kept its hold through the dust storms of the area, often now<br />

perched on top of low mounds formed by the drifting dust<br />

which was held by its older pads and through which its newer<br />

ones have grown. In more grassy places and especially on the<br />

rough breaks along the Canadian and other rivers of the area,<br />

it is often very hard to find, as the grass grows up in the clump<br />

and entirely shields the low pads from view. One can learn to<br />

spot it by looking for the mounds of longer grass here and there<br />

on the prairie caused by the protection from grazing animals<br />

the grass gets from the unseen pads between which it grows.<br />

Opuntia compressa (Salisbury) Macbride<br />

“Low Prickly Pear,” “Smooth Prickly Pear”<br />

Description Plates 54, 55, 56<br />

roots: Fibrous, often entirely so, but often with spindle-<br />

shaped to spherical tubers upon these fibrous roots. The plant<br />

never has a central taproot.<br />

stems: A low-growing species with pads entirely prostrate or<br />

spreading upward at first and then reclining on their edges<br />

with age. The height of the plant thus is usually 1 foot or less,<br />

but occasionally in some forms it grows to 18 inches tall. The<br />

pads are round or even broader than long to egg-shaped or<br />

sometimes spindle-shaped, some with and some without constriction<br />

below. They are typically 1 to 6 inches long, by 1 to<br />

5 inches wide, but occasionally in some forms they may be to<br />

7 or 8 inches long on exceptionally robust specimens. They<br />

are of average thickness to very thick for their size, tuberculate<br />

by raised areoles or smooth and flat, dark green to pale<br />

or yellowish-green, some forms with purplish spots at the<br />

areoles in the winter.<br />

areoles: Small and inconspicuous, 1/16 to 1/8 of an inch in<br />

diameter and elongated oval to circular at first, becoming<br />

with age more circular and sometimes to 1/4 of an inch across.<br />

They are spaced 3/16 to 11/4 inches apart.<br />

spines: Entirely spineless or with spines on less than the upper<br />

half of each pad. Areoles of the lower half to two-thirds of<br />

each pad are always spineless. The armed areoles with 1 to 5<br />

spines are arranged as follows. There may be 1 or 2 straight<br />

main spines which stand porrect, somewhat deflexed or erect,<br />

1/4 to 21/4 inches in length, rather slender to heavy, round or<br />

flattened, their color from almost pure grayish-white to yellowish<br />

or mottled above with the bases brown or red-brown,<br />

or occasionally the whole spine brown. There may be present,<br />

but more often will be lacking, one lower, deflexed spine 3/16<br />

to 1 inch long, slender, similar in color. Some forms have occasionally,<br />

besides these, 1 or 2 bristle-like spines spreading<br />

downward, only 1/8 to 3/8 of an inch long, and white in color.<br />

glochids: Few, short, and fine at first, comprising a compact<br />

tuft in the middle of the areole, but usually increasing greatly<br />

in number on old pads and becoming to 1/4 or even 3/8 of an<br />

inch long. As numerous on the sides of the pads as on the<br />

edges, or even more numerous. They are greenish-yellow,<br />

straw, brown, or bright red-brown in color.<br />

flowers: 2 to 5 inches in diameter, pale yellow to orangeyellow,<br />

or orange-yellow with red centers. The ovary is elongated<br />

obovate to clavate and is 1 to 21/4 inches long. The<br />

anthers are cream-colored. The stigma lobes are 4 to 9 in<br />

number, white or yellowish in color.<br />

fruits: Clavate with constricted or greatly prolonged bases<br />

and narrow, noticeably depressed to deeply pitted umbilici,<br />

red-brown in color. The seeds are round, flat, but thickbodied,<br />

with narrow, acute rims. They are 1/8 to 1/4 of an inch<br />

(3–6 millimeters) in diameter and 2 to sometimes 4 millimeters<br />

thick.<br />

Range. Including all of the forms at least tentatively assigned<br />

to the species, ranging over the entire U. S. except approximately<br />

the northwest third and the state of Maine.<br />

Remarks. When all of its varieties are considered together, this<br />

small, prostrate, inconspicuous prickly pear is no doubt the leading<br />

candidate for the most wide-ranging cactus in the U. S. It is


202 cacti of the southwest<br />

known to occur from New England southwest through all states<br />

to the Rocky Mountains, and from there on west through New<br />

Mexico and Arizona probably into California. It is said to be<br />

absent only from the state of Maine and those states west of the<br />

Continental Divide and north of Arizona and California.<br />

This wide-ranging, adaptable cactus was naturally among the<br />

earliest noticed, and should be the easiest to recognize and name,<br />

but it actually presents the greatest confusion to serious students<br />

of cacti. Even finding the proper name for the species is enough<br />

to discourage the hardiest cactophile. It was first christened<br />

Cactus opuntia by Linnaeus, and shortly thereafter Cactus compressus<br />

by Salisbury. Soon it was called Cactus humifusus by<br />

Rafinesque. By the rules of nomenclature, after the genus Opuntia<br />

was erected, its name could not remain Opuntia opuntia, although<br />

some books have called it that, so Salisbury’s name, as<br />

revised by Macbride into Opuntia compressa is now accepted as<br />

valid, although Prince Salm-Dyck had in the meantime coined<br />

yet another name, Opuntia Vulgaris, for it.<br />

But this is only the beginning of the naming problem. Any<br />

plant growing over so huge an area is likely to present various<br />

localized forms, and whatever level of relationship they represent,<br />

few will be satisfied with a view so general that it ignores<br />

them. I know from experience that few collectors who have side<br />

by side two of the widely differing forms this species can show<br />

are satisfied when I tell them they are both O. compressa. They<br />

want to know more exactly what they have. So from the first<br />

these differing forms had to be studied and named, and here the<br />

real confusion enters. We will deal with these variations which<br />

come within our area of study one by one.<br />

Rafinesque seemed to be the first to notice these different<br />

variations. He treated them all as separate species, erecting first<br />

of all a species, O. humifusa, for the western, Mississippi Valley<br />

form. Later he broke down even this, his original western species,<br />

into several others, O. humifusa (in a more restricted sense), O.<br />

caespitosa, and O. mesacantha, plus others, but his descriptions<br />

of these latter variations were so vague that no one can really<br />

identify plants from them, so that Engelmann, in exasperation<br />

with him and failing to go back to the earlier names, proposed<br />

instead to start over, calling the whole complex O. rafinesquii.<br />

This seems an illogical solution, and most students have returned<br />

to the name O. compressa as the valid one for the species, with<br />

mostly newer names having been proposed for the varieties.<br />

The description given above is a broadened one drawn to include<br />

all of the varieties presently assigned to the species complex.<br />

It is not the description of the form restricted to the eastern<br />

seaboard, which was the type variety and which is still<br />

commonly referred to as O. compressa, but which, under the<br />

modern system, should be given a varietal rank. This typical<br />

variety does not enter the area of this study, and so is not described<br />

separately here. The question of what forms to include<br />

in the species complex is at present a very arbitrary one, with<br />

very little solid evidence for or against the inclusion of some<br />

forms. I am drawing the species boundaries as they seem most<br />

understandable to me. Future study may very well revise the<br />

limits of this species further.<br />

Opuntia compressa var. humifusa (Raf.)<br />

Description Plate 54<br />

roots: Usually fibrous, but occasionally with peanut-shaped<br />

or spindle-shaped tubers on these fibrous roots. The plant<br />

never has a fleshy taproot.<br />

stems: Prostrate, never ascending over 1 or 2 pads high. The<br />

pads are round to broadly oblong, usually 2 to 4 inches long,<br />

but sometimes reaching 5 inches. The pads are thick and flat<br />

when healthy, becoming wrinkled when desiccated, but never<br />

being tuberculate by elevations at the areoles. The surface is<br />

dark or bright green and shining, sometimes having some purplish<br />

coloring at the areoles in winter or drought.<br />

areoles: Small, being only 1/16 to hardly 1/8 of an inch long<br />

and enlarging little if any with age. They are elongated oval<br />

to round in shape, and the opposite of being bulging outward<br />

with wool, they are usually rather concave, pitlike depressions<br />

on the surface of the flat pad. They are spaced 1/2 to 1<br />

inch apart.<br />

spines: Most often the plant is spineless. If spines are present<br />

they will be found in only a few upper edge areoles and will<br />

number only 1 to 3 per areole. They ordinarily consist of 1 or<br />

2 spreading spines up to 1 inch long. Rarely there will be 1<br />

lower, deflexed, and very small one. The main spines are<br />

round, straight, and heavy, whitish to brownish in color.<br />

glochids: Few and short at first, comprising a small, compact<br />

tuft of minute bristles less than 1/8 of an inch long in the<br />

small areoles. Often they do not increase at all on old pads,<br />

but in some specimens they lengthen to 3/16 of an inch or so<br />

with age. They are red-brown to yellowish in color.<br />

flowers: Yellow, usually, but not always, with red centers or<br />

else streaked with red. They are 2 to 3 inches in diameter and<br />

13/4 to 3 inches tall, including the relatively long, slender<br />

ovary. On the ovary there are some glochids. There are 10 to<br />

14 broad inner petals. The filaments are orange, often conspicuously<br />

so, and the anthers are cream-yellow. The style is<br />

short, and there are 4 to 8 white or very pale greenish-yellow,<br />

fat, grooved stigma lobes.<br />

fruits: Elongated club-shaped or oval above, always with a<br />

constricted, slender, and markedly elongated base and deeply<br />

pitted apex. They are 11/8 to 2 inches long, 1/2 to 3/4 of an inch<br />

thick at the thickest part, which is near the top, the whole<br />

lower one-third at least being only 1/4 to 3/8 of an inch thick.<br />

These fruits are greenish for a long time, but tardily become<br />

apricot or plum-red or brownish red, the pulp remaining<br />

greenish and sour in poor growing conditions, but when growing<br />

in the best conditions becoming reddish and sweet. The


genus Opuntia 203<br />

seeds are 3/16 of an inch or just under (31/2–41/2 millimeters)<br />

in diameter, thick to very thick, with narrow rims.<br />

Range. Entering our area from the north and east, coming into<br />

Louisiana from Mississippi, into Arkansas from Mississippi, Tennessee,<br />

and Missouri, and extending throughout Arkansas and<br />

northern Louisiana into Texas and probably into Oklahoma. Its<br />

southwestern limit seems to be a line running approximately<br />

through Alexandria, Louisiana, to the vicinity of Silsbee, Texas.<br />

From there its known range turns sharply north, remaining east<br />

of the Trinity River until it nears Dallas. From there it seems to<br />

retreat back toward Arkansas, except for an apparent occurrence<br />

of it near Madill, Oklahoma, which opens up the possibil-<br />

ity that it ranges widely over eastern Oklahoma.<br />

Remarks. There has been continuous discussion for 150 years<br />

about how different the O. compressa of the eastern seaboard is<br />

from that found in the Mississippi Valley. One thing is apparently<br />

agreed upon from Rafinesque through the last article<br />

treating the problem. All seem to agree that there is a difference<br />

between the two forms, although it is frustrating to find so little<br />

anywhere by way of a concrete statement of how they actually<br />

differ.<br />

The early students regarded the two as entirely separate species.<br />

The eastern one, apparently found only east of the Appalachian<br />

Range, is usually called O. compressa. It is the type form<br />

of the species, and considered as part of the bigger complex,<br />

would be the typical variety of it. But how do we treat the<br />

form of the Mississippi Valley, which is the one concerning us<br />

directly? Rafinesque divided it into his several separate species<br />

before he was through. It was his inability to identify these by<br />

their descriptions that caused Engelmann to abandon them all<br />

and rename the Mississippi Valley form O. rafinesquii. Engelmann<br />

then added new names for what he regarded as new spe-<br />

cies he found farther west.<br />

Coulter, regarding this as an unsatisfactory dodge, and working<br />

from the fact that Rafinesque said his O. mesacantha grew<br />

from Kentucky to Louisiana, took this name, O. mesacantha, for<br />

what he considered a major western species in opposition to the<br />

eastern O. compressa. He ended up with nine named subspecies<br />

in his western complex under that name.<br />

More recently it has been regarded as improbable that all of<br />

these forms are separate species and even as doubtful that many<br />

of them are separate forms at all. It has come to be considered<br />

more correct to regard those which are separate forms as varieties<br />

of the one huge species, O. compressa. This has caused the<br />

most recent works to eliminate all the names of Rafinesque and<br />

most of those of Engelmann and of Coulter. The change has<br />

made for greater clarity, except when one turns to various floras<br />

and handbooks which still often use the different names of the<br />

above authors, apparently at random. There has been no standardization<br />

of the varietal names. The result is that most beginners,<br />

not knowing which are synonyms, go into the field with<br />

twice as many names as there are cacti, and are soon frustrated<br />

when they can’t find them all. With the aid of the above admittedly<br />

over-simplified summary and the following discussion<br />

of our form, it is hoped some of this can be avoided.<br />

In the geographical territory we are considering we are not<br />

concerned with the typical, eastern variety. However, we do<br />

have throughout Arkansas and much of Louisiana and coming<br />

into Oklahoma and Texas what appears to be the midwestern<br />

or Mississippi Valley variety of the cactus. We need the correct<br />

varietal name for it. The first person to deal with this particular<br />

form was Rafinesque. However incompletely he described it, it<br />

seems clear that he meant this cactus by his O. humifusa. Engelmann<br />

described our form, but, as already mentioned, his solution<br />

of renaming it entirely to O. rafinesquii is unsatisfactory.<br />

Coulter, who next described it, returned to a Rafinesque name,<br />

but chose a more obscure one, O. mesacantha, which we cannot<br />

pin down to anything we know at all. We are left with Rafinesque’s<br />

first name for the Mississippi Valley form. Regarding it<br />

now as a variety instead of a species, we get as a name for the<br />

low prickly pear of this area O. compressa var. humifusa.<br />

By whatever name, this is an inconspicuous little prickly pear<br />

found mostly in sandy places or on the drier hillsides jutting<br />

out of the piney woods. In such places it sometimes forms mats<br />

covering the ground. Occasionally plants are found on wooded<br />

hillsides under the trees, but here they are more straggling and<br />

pads tend to be somewhat etiolated, in which case they become<br />

larger and more elongated than typical, but still remain fat and<br />

thick.<br />

I have plants from various locations in Arkansas. In Hempstead<br />

County, near Tokio, they grow in sandy bottoms. There<br />

are fine examples to be seen in Benton County in the northwest<br />

corner of the state, growing east of Rogers, Arkansas. They are<br />

not as common in the eastern part of the state, but have been<br />

found near Batesville in the north and Warren in the south.<br />

In Louisiana the plant is less common and only a few records<br />

of it there have been found. It appears to grow best in the north-<br />

west part of the state, having been found just north of Alexan-<br />

dria, near Winnfield, and near Minden.<br />

The variety enters Texas from Louisiana, but only grows along<br />

the eastern edge of this state. The southwesternmost record of it<br />

is from near Silsbee, Texas, in Hardin County. It does not progress<br />

farther to enter the coastal plain or the central Texas limestone<br />

hills, being restricted almost entirely to the eastern piney<br />

woods. It grows quite profusely under the trees at Woodville,<br />

Texas, and has been found again at various places north of there.<br />

It is rather common at Longview, Texas, this being the apparent<br />

basis for Griffiths’ O. nemoralis.<br />

The question of this variety’s occurring in Oklahoma is still<br />

an open one. Engelmann and Coulter both state that, when they<br />

wrote, it had not been found west of the western boundary of<br />

Missouri and Arkansas. We have seen that it progresses farther<br />

west than that in Texas, but there have not been any good records<br />

of it from Oklahoma cited in more recent works, although<br />

Waterfall, in his “Catalog of the Flora of Oklahoma,” lists O.


204 cacti of the southwest<br />

rafinesquii as well as O. macrorhiza, and must mean this variety.<br />

Not having found it in many trips into Oklahoma, I would<br />

be inclined to doubt its growing there, except for one population<br />

of plants which I found growing in the Tishomingo Wildlife<br />

Reservation near Madill, Oklahoma. These seem to be referable<br />

only to this variety, even though they are growing so far<br />

west. Perhaps future study will show that the variety is found<br />

in very scattered spots over the eastern hills of Oklahoma.<br />

O. compressa var. humifusa has small, shiny, dark green pads<br />

which are very thick. They become very wrinkled in winter or<br />

drought, in which case they also often get purple blotches at the<br />

areoles, but the pads are smooth-surfaced and never appear<br />

tuberculate by elevation of the areoles. In fact, more often the<br />

small areoles actually seem to be in slight depressions of the sur-<br />

face. In this and in the fact that neither the areole nor the<br />

glochids increase noticeably in size with age or on the edge of<br />

the pad, as do those of most of its relatives, it is like the more<br />

eastern, typical variety. It is also a very prostrate form. When<br />

it is growing in the same garden bed with them, the difference<br />

is striking between this form with its pads all reclining once<br />

they have reached maturity and variety macrorhiza, variety<br />

fusco-atra, and variety allairei, all of which have arms more or<br />

less turning upward from old pads that much prefer to touch<br />

only their edges to the ground.<br />

It has often been stated that this variety always has fibrous<br />

roots. In fact, these roots have been used as the main character<br />

to distinguish it from the more western forms. Britton and Rose<br />

separate this form from O. macrorhiza in their key by roots<br />

fibrous versus roots tuberous alone. But handy as this would be,<br />

it just isn’t true that variety humifusa always lacks tubers. While<br />

fibrous roots are the most common, I have found small, peanut-<br />

shaped tubers up to about 3/4 of an inch in diameter and 11/2<br />

inches long on specimens in Arkansas and in Louisiana, as well<br />

as in Missouri. The character of the roots cannot be used to separate<br />

forms in this species.<br />

Opuntia compressa var. macrorhiza (Eng.) Benson<br />

Description Plate 54<br />

roots: Having a basically fibrous root system without a<br />

central taproot, but these slender fibrous roots often having<br />

spherical to spindle-shaped tubers on them. These tubers may<br />

be from a fraction of an inch up to Sometimes 3 inches in<br />

diameter. They may occur in clusters on the various branching<br />

roots, or may sometimes be found in a series of several one<br />

after another along the same root. Plants lacking tubers completely<br />

are not uncommon.<br />

stems: Prostrate or nearly so, with old pads mostly leaning<br />

on their edges on the ground, and with newer pads only<br />

temporarily upright, the whole plant being 6 inches to 1 foot<br />

high. The pads are almost round to elongated obovate, often<br />

with some constriction at the bases, but this not pronounced.<br />

These pads are medium in thickness to rather thick; the surface<br />

is quite tuberculate by raised areoles when growing, and<br />

these elevations remain more or less noticeable on old pads,<br />

which may also become wrinkled. In color they are medium,<br />

shining green when young, becoming dull, dark green when<br />

old. The pads are normally 2 to 5 inches long, but when growing<br />

under shading vegetation they sometimes become etiolated<br />

to 6 or even 8 inches long.<br />

areoles: Oval or almost round and 1/8 of an inch or so across<br />

at first, enlarging to 1/4 of an inch or more on old pads. They<br />

are 1/2 to 11/4 inches apart.<br />

spines: Spines are present on the upper one-fourth of the pad<br />

or less, but the pads are occasionally but not often totally<br />

spineless. The most common spination is 2 to 3 spines per<br />

areole, but very robust plants present up to a maximum of 5<br />

spines on some edge areoles. On young or weak plants all<br />

spines are round or with only the bases somewhat flattened,<br />

but robust specimens have the main spine conspicuously flattened.<br />

Completely armed areoles present 1 main central spine<br />

which is porrect or a little deflexed, slender to medium thickness,<br />

straight, round if weak but flattened if robust, and 1/4 to<br />

13/4 inches long. There may be, but will not always be, 1 or 2<br />

upper spines porrect to erect, 3/8 to 1 inch long, slender, weak,<br />

and always round. These main spines are gray or whitish,<br />

with often the bases and sometimes the tips brown or red-<br />

brown, fading to all gray and rough when old. On robust<br />

specimens there will sometimes be 1 or 2 lower spines spreading<br />

below which are very slender, thin, whitish bristles 1/8 to<br />

3/8 of an inch long.<br />

glochids: Bright red-brown to dirty straw in color, medium<br />

in number, and 3/16 to 1/4 of an inch long, in a compact clump<br />

at each areole at first. On old pads they often grow to 5/16 of<br />

an inch long and form large clusters.<br />

flowers: Orange yellow with red centers. They are 21/2 to 3<br />

inches in diameter and 3 to 31/2 inches tall, the ovary being<br />

11/4 to 21/4 inches long, with some reddish-brown glochids on<br />

it. There are 7 to 9 inner petals which are very broad from<br />

narrow, red bases. The stigma has 4 to 9 yellowish, fat stigma<br />

lobes.<br />

fruits: Elongated club-shaped, 11/4 to 21/2 inches long and 3/4<br />

to 1 inch in diameter at the thickest part. The base is greatly<br />

constricted and prolonged. The top is shallowly to deeply<br />

pitted. These fruits become light red to purplish-red, often<br />

however remaining greenish very late in the season. The pulp<br />

is greenish and tasteless until fall, finally becoming colorless<br />

or slightly reddish and sweet to the taste. The seeds are from<br />

just under 3/16 to just under 1/4 of an inch (4–51/2 millimeters)<br />

in diameter. They are thick or very thick, their rims of me-<br />

dium width but acute.<br />

Range. Western Arkansas on the east through all of Oklahoma


genus Opuntia 205<br />

and all of Texas west of a line from near Tyler to near Houston.<br />

The plant grows almost throughout New Mexico and on into<br />

Colorado and Arizona.<br />

remarks. This is a very wide-ranging plant, but, being small<br />

and marked by no unique feature, it is usually ignored or actually<br />

avoided even by cactus fanciers. Over a huge area it is the<br />

common prostrate prickly pear that lies in the grass or under<br />

thickets and, except for a few days each year when it has beautiful<br />

flowers, has little to offer to anyone except thorns for those<br />

who don’t look before they step.<br />

Information about the exact range of this plant and its relationships<br />

is hard to be sure about even today, because it so nearly<br />

approaches other forms in appearance and because the name has<br />

been so often misapplied that printed records of it and even<br />

labeled herbarium specimens are often actually something else.<br />

One has to wade through a mass of extraneous material to assemble<br />

a picture of the true form. Also, the early descriptions<br />

were largely taken from too few and often stunted specimens,<br />

and so there has had to be a broadening of the original concept<br />

almost from the beginning, a broadening which has threatened<br />

to go on until it seemed almost all prostrate prickly pears would<br />

be herded under this name. Only by a tracing of the history of<br />

the form can its present status be understood.<br />

When Engelmann made the first concerted study of the western<br />

forms of this species, he found in the west, beyond the O.<br />

compressa forms he had seen in the east, several small, prostrate<br />

cacti. Almost without exception he set each one of these up as a<br />

separate new species. One of them was a plant sent him by Lindheimer<br />

from the upper Guadalupe River at New Braunfels,<br />

Texas. Engelmann described it from Lindheimer’s specimens and<br />

called it O. macrorhiza. These specimens were, by any character<br />

he gives for them—size of pad, number of spines, character of<br />

spines, length of spines, size of fruits—less than robust. I have<br />

seen some of these Lindheimer specimens, and they are small,<br />

weak examples such as one can find in the type locality, but<br />

which one has to search out on the most exposed, dry, rocky<br />

ledges. The plant usually attains a much greater size and development<br />

even in the type locality. I cannot but conclude that<br />

these type specimens are atypically underdeveloped individuals<br />

with characters nearer to those of the more eastern O. compressa<br />

var. humifusa than are found in more typical individuals. But<br />

all of Engelmann’s specimens from New Braunfels happened to<br />

have tuberous roots, and he seized upon this as a distinguishing<br />

feature, actually stating that the plant is really different from<br />

the eastern form mainly by the roots.<br />

Taking Engelmann’s limited description and following his<br />

statement, some students have had real trouble with these two<br />

forms. For instance, Britton and Rose happily keyed them: roots<br />

fibrous = the eastern form, roots tuberous = O. macrorhiza. It<br />

would be wonderful if it were that simple, but following this in<br />

the field gives difficulty. In numerous fields in Texas, Oklahoma,<br />

and Arkansas, I would be forced by this to say that I have the<br />

two forms growing as close as 6 feet apart and all intermixed<br />

over the whole population. Even in the type locality of O. macrorhiza<br />

both kinds of roots are found together. So, if this were<br />

true, not only would we have variety humifusa growing all the<br />

way into New Mexico when all have stated that it hardly goes<br />

west of Arkansas and Louisiana, but we would in honesty be<br />

required to combine the two. The only conclusion is that either<br />

plant can have either wholly fibrous roots or roots with tubers<br />

present, the western form, it is true, having tubers more often<br />

than not, while the eastern one usually lacks them. But the character<br />

of the roots obviously cannot be used to separate the two<br />

cacti.<br />

If this is true, then why not combine the two forms entirely?<br />

It seems remarkable that this has never been attempted. But per-<br />

haps the fact that the western form so often surpasses Engelmann’s<br />

original description in other characters has enabled all<br />

to see that it is actually different in other ways.<br />

Coulter was quick to see that the description of this form had<br />

to be broadened, and he set about it, raising the spination maximums<br />

in number, length, and stoutness, but still not realizing<br />

the maximums of fruit size sometimes attained, or the fact, unnoticed<br />

until Griffiths observed it, that the main spine could be<br />

flattened when fully developed.<br />

Because the plant, as originally described, was a less than<br />

robust thing, more robust specimens were sometimes described<br />

as new species when they were noticed. Engelmann himself<br />

started this, for instance, with his O. fusiliformis of Kansas and<br />

Nebraska. All students since have considered this merely a synonym.<br />

Only the very long fruits, said to be sometimes 2 to 21/2<br />

inches long distinguish it from the other, and I have collected<br />

variety macrorhiza from a dozen places, including the type<br />

locality, with fruits this size. More recently Mackensen called<br />

some long-spined examples he found at Kerrville, Texas, O.<br />

roseana, but I find nothing in this locality outside the limits of<br />

variety macrorhiza as it grows in its own type locality not very<br />

far away.<br />

But the broadening of the characters of the plant cannot go<br />

on indefinitely, to include everything. I have given in this description<br />

the full maximum of each character that we have found<br />

in and near the type locality close to which we have lived and<br />

in which we have observed constantly for 7 years. I have excluded<br />

from consideration any described or preserved materials<br />

from any other place which do not, in all their features, fall<br />

within the limits attained in that type area.<br />

This gives us in O. compressa var. macrorhiza a very widely<br />

distributed plant. Its easternmost record would be that mentioned<br />

by Coulter of near Fayetteville, Arkansas, and Demaree’s<br />

collections from near Bismarck and from Magnet Cove, Hot<br />

Springs County, Arkansas. We also have the plant collected<br />

near Jasper, in Newton County, and near Little Rock, Pulaski<br />

County. These records give us a range of the whole northwestern<br />

part of that state. It has apparently not been collected in<br />

Louisiana, and the known collections of it in Texas give as an


206 cacti of the southwest<br />

eastern limit excluding the whole eastern edge of the state, it<br />

first appearing along a line roughly from Tyler—where Griffiths<br />

collected it and dubbed it O. sanguinocula—to Houston. It apparently<br />

ranges practically all of the rest of Texas, being very<br />

common in the central and northern parts, but occurring in scattered,<br />

usually very sandy, localities elsewhere. I have collected<br />

it on the Bolivar Peninsula—where, however, it was growing<br />

only on and around the grounds of old Fort Bolivar, and one is<br />

tempted to assume it may have been introduced—in the Corpus<br />

Christi area, as well as near Harlingen in the southern extremity<br />

of the state. At the other end of the state, in the Chisos Mountains<br />

and near Muleshoe, Texas, well up toward the northwestern<br />

edge of the state, I have come across isolated populations<br />

which seem clearly to be this cactus. It ranges practically all, if<br />

not all of Oklahoma, being very common as far west as Alva<br />

and Woodward. It occurs in scattered localities in New Mexico,<br />

becoming quite common in the northern mountains of that state,<br />

and I have seen good specimens of it from both Arizona and<br />

Colorado.<br />

It will be noticed that this range does give us both variety<br />

macrorhiza and variety humifusa in an overlapping area in<br />

western Arkansas and probably some of Oklahoma. When the<br />

two are so close, how, without the roots to decide for us automatically,<br />

can we tell them apart? Admittedly in weak speci-<br />

mens without blooms it can sometimes be almost impossible, and<br />

everyone should be honest enough to admit that he can’t identify<br />

an incomplete or atypical specimen with certainty. This would<br />

reduce the confusion. Also, if we think of these two as varieties<br />

instead of separate species, we need not be surprised if there are<br />

specimens apparently intermediate. But in identifying typical<br />

specimens there is little difficulty. The salient differences between<br />

them are as follows: variety macrorhiza has tuberculate<br />

surfaces to the pads, up to 5 spines with the main spine flattened<br />

and to 13/4 inches long when spination is completely developed,<br />

flowers with only 7 to 9 inner petals. On the other hand, variety<br />

humifusa has a flat surface to the pad, a maximum of 3 spines<br />

which are heavier but always round and to only 1 inch long,<br />

and flowers with 10 to 14 inner petals. The two are very close<br />

together, and there may be intergrading. They surely do not<br />

warrant being considered separate species. This form seems most<br />

logically considered, therefore, as the more western member of<br />

the O. compressa complex, and so is known today by most taxonomists<br />

as O. compressa var. macrorhiza.<br />

Opuntia compressa var. microsperma (Eng.) non Benson<br />

Description Plate 54<br />

roots: Mostly fibrous with small tubers on them, but sometimes<br />

entirely fibrous.<br />

stems: Prostrate, 6 to 8 inches high, with most pads resting<br />

one edge on the ground. The pads are broad to elongated egg-<br />

shaped, 1 to 4 inches long by 3/4 to 21/2 inches wide, rather<br />

thick, with the surface very tuberculate by elevations of the<br />

areoles. The color is medium green with purplish shading<br />

around the areoles.<br />

areoles: Small, round or oval, 1/16 to 1/8 of an inch long, on<br />

top of distinct elevations. They are situated 1/4 to 3/4 of an<br />

inch apart.<br />

spines: Sometimes spineless, but more often with 1 to 3 spines<br />

on the upper areoles of the pad. The one or two main spines<br />

are very slender, round, and all white, or with the bases<br />

brown and the upper parts white. Occasionally there is 1<br />

lower, deflexed, bristle-like spine, 1/8 to 3/4 of an inch long<br />

besides the main spines.<br />

glochids: Greenish-yellow or straw color, many in all areoles,<br />

at first to 3/16 of an inch long, but when older becoming<br />

to 1/4 of an inch long.<br />

flowers: Identical with those of the previous variety.<br />

fruits: Very elongated club-shaped. They are 1 to 11/4 inches<br />

long by 3/8 of an inch thick at the widest part, which is near<br />

the top, the base being very constricted. The apex is deeply<br />

pitted. The fruits on my specimens remain green all summer<br />

and through September, after which they dry without turning<br />

color. The seeds are from just under 1/16 to just under 1/8 of an<br />

inch (1/2–21/2 millimeters) in diameter, smooth, and thick,<br />

with almost no rims.<br />

Range. Collected 10 miles north of Campbellton, Atascosa<br />

County, Texas, growing in deep sand near the Atascosa River.<br />

Remarks. Engelmann originated this variety over 100 years ago,<br />

but he gave almost no description of it except to emphasize its<br />

extremely small seeds. He also neglected to give any location<br />

for it. Coulter, working with Engelmann’s materials before him,<br />

both the preserved specimens and living plants still being cultivated<br />

at that time at the Missouri Botanical Garden, was impressed<br />

enough by the tiny seeds to continue listing it as a sepa-<br />

rate form. But after that all authorities dropped it into the syn-<br />

onymy.<br />

Most recently the name has been revived and used in an entirely<br />

new sense. When Benson decided to treat the common,<br />

prostrate, Mississippi Valley Opuntia—until then always regarded<br />

as a separate species and called O. humifusa, O. rafinesquii,<br />

or O. mesacantha—as a variety instead of a species, he, in<br />

an attempt to be taxonomically proper, called that form O. compressa<br />

var. microsperma. His stated reason was that this was the<br />

earliest varietal name in existence under that species. This is true,<br />

but in using it thus he ignores the fact that Engelmann erected it<br />

for the very special form with the tiny seeds and meant it to<br />

refer to that form alone, as the meaning of the name clearly<br />

shows. To use this name for the ordinary, larger seeded, common<br />

form is to say that this seed difference is of no significance, and<br />

to apply this name to the large-seeded form is an error which<br />

every person who can tell the meaning of this simple name can


genus Opuntia 207<br />

recognize. It blurs a distinction which was considered significant<br />

by those earlier students of cacti.<br />

In the course of this study many collections of the species O.<br />

compressa were made over several states. Time and again the<br />

ordinary forms with seeds 3 to 5 millimeters in diameter were<br />

found, and we had only wondered vaguely about Engelmann’s<br />

variety microsperma with the tiny seeds. Since we knew the<br />

seeds of the species well, it was a real surprise when Mr. Kim<br />

Kuebel, an interested friend and cactus collector, brought in a<br />

plant of the O. compressa complex which had tiny seeds only<br />

11/2 to 21/2 millimeters in diameter. They equal in size the smallest<br />

Opuntia seeds we have seen, but are very thick. It was obvious<br />

to us that we must have the plant referred to by Engel-<br />

mann, but probably not collected again until this time.<br />

We studied the plant carefully, trying to discover the details<br />

of its relationship. It is certainly only a variety of the large O.<br />

compressa complex. Within that, however, its closest affinities<br />

are to the more western O. compressa var. macrorhiza rather<br />

than to the eastern variety humifusa. This is shown by the tuberculate<br />

surface of the pads, the more numerous glochids, more<br />

slender spines, and so on. The fruit is remarkable for never turning<br />

red, but then many others of this species only turn light red<br />

or apricot, and sometimes this is only a blush on the sunny side<br />

of the fruit.<br />

It is impossible to give a range for this form, as Engelmann’s<br />

specimens carried no location at all. We have so far found this<br />

plant in only one location, as given above. Typical variety macrorhiza,<br />

with seeds 41/2 to 5 millimeters in diameter was grow-<br />

ing nearby.<br />

Listing the variety microsperma as a separate form on the<br />

basis of only one population would not ordinarily have been<br />

justified, but this is a form which we know existed somewhere a<br />

hundred years ago. Whether the unique, small seeds are an environmentally<br />

induced or a genetically stable character, which<br />

has occurred only in this location or, as seems more likely, crops<br />

up here and there, no one knows. By listing the form instead of<br />

ignoring it and appropriating the name for one of its relatives,<br />

we hope that we may prevent its again being lost, with no one<br />

ever knowing more about it, and that we may alert collectors to<br />

be on the watch for it. Thus we may someday learn what its<br />

true significance is.<br />

Opuntia compressa var. fusco-atra (Eng.)<br />

Description Plates 54, 55<br />

roots: Either fibrous or about half of the time with small<br />

tubers upon the fibrous roots.<br />

stems: Prostrate, usually resting at least one edge of each pad<br />

on the ground, only to 6 or 8 inches high. The pads are almost<br />

round to broadly club-shaped with some constriction at the<br />

base, usually 2 to 4 inches long and 2 to 3 inches wide, but<br />

rarely to at least 6 inches long and 4 inches wide. They are<br />

thin or of medium thickness, noticeably tuberculate by elevations<br />

of the areoles. The surfaces of the pads are shiny and<br />

bright green to deep green or sometimes rather blue-green.<br />

areoles: Small and elongated when young, becoming round<br />

and larger to about 1/4 of an inch in diameter on old pads.<br />

They are spaced 3/16 to 11/4 inches apart, varying with the<br />

size of the pads; both extremes have been observed on the<br />

same plant.<br />

spines: Usually having 1 to 3 spines on only upper areoles of<br />

the pad, but rarely having a total of 4 or 5 by the addition<br />

of one or two very small, bristle-like spines at the lower edge<br />

of the areole. There is 1 main spine porrect or nearly so, slender<br />

to medium thickness, round or practically so, 1/4 to 2<br />

inches long. There may be 1 or 2 upper spines above this and<br />

similar to it except only 3/8 to 1 inch long, and sometimes 1<br />

lower spine deflexed below and 3/8 to 5/8 of an inch long.<br />

Spreading below these are occasionally 1 or 2 very slender<br />

bristles 1/8 to 1/2 inch long. All spines are yellowish when<br />

growing, but when hardened become completely dark brown<br />

or gray with brownish bases.<br />

glochids: Many, in erect, compact clusters in all areoles, be-<br />

coming very many and to 1/4 of an inch long when old. In<br />

color they are very bright, shining red-brown when young,<br />

fading to dark, dirty brown when old.<br />

flowers: Completely sulphur yellow or yellow with red centers.<br />

They are very variable in size, depending upon environmental<br />

conditions, from 13/4 to 4 inches in diameter and 2 to<br />

3 inches tall, with very slender ovaries 11/8 to 13/8 inches long,<br />

having some brown glochids on them. There are very few but<br />

broad inner petals, usually only 4 or 5, the total number of<br />

perianth segments usually being only 9. The stigma has 3 to 5<br />

white lobes.<br />

fruits: Elongated club-shaped with constricted bases and<br />

deeply pitted tops. They are typically 1 to 11/2 inches long,<br />

but sometimes to 2 inches, and 5/8 to 7/8 of an inch in diameter<br />

at the thickest point. The seeds are about 3/16 of an inch<br />

(4–5 millimeters) in diameter, fairly thick, with narrow, acute<br />

rims.<br />

Range. The coastal plain, sandy areas near the beaches and on<br />

islands of the Texas coast from the western edge of present-day<br />

Houston to beyond Brownsville. The plant is never found more<br />

than a few miles from the Gulf, except in the valleys of the<br />

Colorado and Brazos rivers, where it occurs inland at least 75<br />

miles, its known range in this area running from Houston west<br />

to Sealy in Austin County, and to Altair in Colorado County.<br />

Remarks. This is almost certainly only an extreme form of the<br />

O. compressa complex, but, although it was noticed and described<br />

over 100 years ago by that amazingly thorough student,<br />

Engelmann, it has been so rarely collected since then that little


208 cacti of the southwest<br />

has been known about it. It is very close to O. compressa var.<br />

macrorhiza, which is found occasionally in most of its range<br />

and sometimes, as just west of Houston, grows side by side with<br />

it. Still, the two are rather distinct, and there is no reason to<br />

confuse them. Variety fusco-atra can be recognized at once by<br />

its combination of numerous brilliant red-brown glochids, very<br />

marked elevation of the areoles, and extreme reduction in num-<br />

ber of flower segments.<br />

In the type locality, stated to be the “sterile prairies west of<br />

Houston,” which begin approximately where the pine trees leave<br />

off, at about the present western city limits of Houston in the<br />

vicinity of U. S. Route 90, and extend past Addicks and Katy,<br />

the plant is very common. However, this very flat territory<br />

gives all the appearance of being very arid, in spite of the high<br />

rainfall of the area, and here the plants appear stunted, the pads<br />

usually only 2 to 3 inches long. This no doubt accounts for the<br />

tiny size given in the original description. Where growing near<br />

the beaches the cactus becomes much larger.<br />

Although the cactus is to be encountered here and there along<br />

most of the Texas coast south of the type locality, it is never<br />

common anywhere else except around Copano Bay in the Rockport<br />

area. At Fulton Beach, particularly, it grows in profusion<br />

right down to the water’s edge. And here it appears to develop<br />

a remarkable environmental form, which has of course been<br />

called a separate species.<br />

In their monumental work, The Cactaceae, Britton and Rose<br />

described a new cactus, Opuntia macateei. It was collected at<br />

Rockport, Texas, in 1910. It was described as a small, prostrate<br />

plant with orbicular to obovate, glabrous, dull green joints which<br />

were small and somewhat tuberculate, and as having 1 to 3<br />

brownish spines up to 1 inch long. Nothing was said about the<br />

root form. The distinctive and really remarkable characteristic<br />

of this cactus was its production of flowers 3 to 4 inches long,<br />

including slender, subcylindric ovaries 2 to 23/8 inches long.<br />

These ovaries were said to bear leaves up to 1/2 inch long. We<br />

spent much time attempting to find and study this form.<br />

It soon became evident that very little was known about the<br />

cactus. We studied the type specimen. It consists of one small<br />

pad with one flower, the flower being typical for the O. compressa<br />

complex, except that it possessed an extremely elongated,<br />

practically cylindric ovary slightly over 2 inches long but only<br />

about 1/2 inch in diameter, bearing several long leaves upon it.<br />

No evidence of any more recent collections of the plant was<br />

found. Later writers, even including Backeberg, have merely<br />

repeated the original description for the species, apparently<br />

without seeing further specimens. No one has even known<br />

enough about the cactus to propose relating it to any other<br />

form.<br />

We then attempted to collect the living plant, making repeated<br />

trips to the type locality at Rockport, Texas, for this purpose.<br />

We found numerous specimens of variety fusco-atra, which<br />

is very common throughout that area, and of variety macrorhiza,<br />

which is less common there but occurs occasionally. We studied<br />

variety fusco-atra especially, as it was early noted that the descriptions<br />

of this plant and of O. macateei were identical as regards<br />

all features mentioned except the structure of the ovary,<br />

that of variety fusco-atra being described as leafless and only<br />

about 1 inch long, less than half as long as the minimum for the<br />

ovary of O. macateei. Britton and Rose used this difference as<br />

the sole means to distinguish these two forms.<br />

For several years we found only the usual forms with typical<br />

short ovaries in the area. Then, in May, 1963, we discovered<br />

some plants producing elongated fruits. These made up a population<br />

of only a few dozen plants growing in almost pure sand<br />

inside a motte of live oak trees. The location was 41/2 miles north<br />

of Rockport, Texas, and about 1/5 mile back from the waterfront<br />

known as Fulton Beach.<br />

The fruits on these specimens were 11/2 to 3 inches long, markedly<br />

clavate from very narrow bases, often widening near the<br />

ends to 1 inch in diameter. They were often curved, and the<br />

younger examples possessed several leaves up to 1/2 inch long. It<br />

was too late in the season to observe the flowers of these plants.<br />

These specimens were identical to the numerous stands of variety<br />

fusco-atra common in the area, but the fruits certainly<br />

were different. No description has ever been given of the fruits<br />

of O. macateei, but we could easily visualize these greatly elongated<br />

fruits developing from the markedly elongated ovaries of<br />

that form, so we felt rather certain that we had found that<br />

cactus.<br />

But almost immediately we noticed that the elongated fruits<br />

of our specimens remained green instead of coloring, and actually<br />

turned whitish with the passing of time. Soon one of them<br />

split open with one longitudinal furrow running the length of<br />

the upper, broadened part of the fruit. This furrow exposed a<br />

mass of black fungus growth resembling smut. Other fruits ruptured<br />

the same way. When we opened some of the fruits before<br />

they ruptured, we found them all to be sterile, the aborted<br />

ovules being much enlarged.<br />

It was impossible to make further studies that season, but the<br />

area was watched closely at blooming time the next year. All<br />

cacti observed in the vicinity of Rockport bloomed and fruited<br />

normally with short, fertile ovaries, except those in the same<br />

motte of live oaks observed to have been atypical the previous<br />

year. Within this particular motte, which comprises probably<br />

two acres of dense tree growth, there are several dozen specimens<br />

of prostrate cacti. When they bloomed, many of their flowers<br />

produced the long ovaries described for O. macateei; these<br />

flowers were then followed by the large, clavate, sterile fruits<br />

observed the previous year. However, this year we noted that<br />

in some cases the same plant produced both elongated flowers<br />

and fruits and the normal flowers and fruits of variety fuscoatra,<br />

sometimes both on the same pad. There were also some<br />

specimens within the population which produced only entirely<br />

typical flowers and fruits.<br />

It seems obvious that we have here an abnormal growth of<br />

ovary and fruit resulting in sterility, rather than a separate


genus Opuntia 209<br />

taxonomic form. Since some plants are found with both normal<br />

and abnormal ovaries, these sometimes side by side on the same<br />

joint, it does not seem that there could be a genetic basis for the<br />

difference. In vegetative characters and also in unaffected flowers<br />

and fruits the plants are clearly variety fusco-atra, and we<br />

feel that O. macateei must be regarded as merely a diseased form<br />

of that species.<br />

The cause of the diseased condition has not proved easy to<br />

isolate. While the determination of the causative agent does not<br />

have to be made in order to understand the plants, we have been<br />

working with specialists in order to try to locate it. A fungus<br />

was at once suspected, but so far it has been possible to culture<br />

no organism from the abnormal fruit tissues except some common<br />

Aspergillis and Penicillium forms. It seems hard to imagine<br />

that such common fungi could have such drastic effects upon<br />

the most common cactus of the area in only one such localized<br />

habitat. Yet the fungi are in the abnormal fruits. But might they<br />

not be merely taking advantage of a breakdown of the tissues<br />

due to something else?<br />

A new factor appeared in the affected plants the next year<br />

which we must have overlooked the previous year. Many of the<br />

plants with abnormal fruits were found to be the hosts of a species<br />

of Lepidoptera. At fruiting time larvae approximately 3/4 of<br />

an inch long were found in the hollowed-out centers of some of<br />

the enlarged fruits, one per fruit. While these fruits were under<br />

observation the larvae became ready for the pupal stage and<br />

burrowed out through the sides of the fruits, left the cacti, and<br />

climbed nearby perennial stems, where they pupated. The insects<br />

entered the winter in the pupal stage, but we were unable<br />

to rear any adults from the pupae we had collected, as all of<br />

our specimens died during the winter. These larvae left round<br />

holes in the sides of the fruits as they burrowed out of them,<br />

and holes of this type were found in some of the pads as well.<br />

Dissection showed that each larva had entered its fruit by tunneling<br />

into it from the pad, through the narrow base of the fruit.<br />

Some of the tunnels were traced through two pads before en-<br />

tering the fruits.<br />

This Lepidoptera has not been encountered by us in any other<br />

instance in the cacti of Texas, and we find no record of it. It is<br />

impossible to find it in the cacti on any side of this thicket.<br />

It was thought that this Lepidoptera might be the cause of<br />

the abnormality in the cactus, since the larvae were observed<br />

only in the pads and the elongated fruits of these abnormal<br />

specimens, but exactly how this may be is not clear, since only<br />

a small per cent of the elongated fruits contained any larvae.<br />

Many of them showed no sign of the insect. Furthermore, the<br />

unusual growth originates at least as early as the flower stage,<br />

affecting the ovary when still in the bud stage, and the larvae<br />

were not observed to enter any fruit until much later than that.<br />

The presence of the larvae in the lower parts of the plant causing<br />

the abnormal growth of ovary is however a possibility, and the<br />

fungi may enter the plants with the insects.<br />

Be that as it may, O. macateei seems clearly only a diseased<br />

form of variety fusco-atra, representing one of the few cases<br />

where such a diseased form in cacti has been called a species.<br />

Opuntia compressa var. grandiflora (Eng.)<br />

“Large-Flowered Opuntia”<br />

Description Plate 55<br />

roots: Entirely fibrous in all specimens observed, with no<br />

tubers present, but in a few specimens the main roots were<br />

found to 3/8 of an inch in diameter and somewhat fleshy before<br />

branching.<br />

stems: The plant is spreading and semiprostrate, but with new<br />

pads standing perfectly erect their first year or sometimes<br />

until there are two pads upright, before falling over, giving<br />

a maximum height of sometimes 12 inches. The pads are eggshaped<br />

to rather spindle-shaped, narrowing at the base or<br />

both above and below, 4 to 6 inches long by 21/2 to 4 inches<br />

wide. They are medium in thickness, the surface conspicuously<br />

tubercled by elevated areoles, and except in very wet seasons<br />

usually markedly wrinkled. In color the surface is bright<br />

green.<br />

areoles: Small and elongated, to only 1/8 of an inch long<br />

on young pads, becoming round and to 1/4 of an inch on old<br />

pads. In position they are 1/2 to 1 inch apart.<br />

spines None.<br />

glochids: Straw-colored, many and conspicuous, although<br />

short and in a tight bunch at first, becoming to 1/4 of an inch<br />

long in a spreading cluster on old pads.<br />

flowers: Normally very large and beautiful, 4 to 5 inches in<br />

diameter and 3 to 31/2 inches tall, but they can be stunted<br />

and reduced to 2 inches in diameter when denied good growing<br />

conditions. In color they are yellow streaked at random<br />

with reddish markings or yellow with reddish centers. The inner<br />

petals are about 8 in number and 2 inches long by 11/2<br />

inches wide on normal sized flowers. The ovary is 2 inches or<br />

more long and slender clubshaped. The stigma has 5 white,<br />

thick, grooved lobes.<br />

fruits: About 21/2 inches long, very elongated club-shaped,<br />

being only 1/2 to 3/4 of an inch wide at the thickest upper part,<br />

with a prolonged constriction below. The top is somewhat<br />

pitted.<br />

range. Given originally as “on the Brazos” in Texas. The only<br />

specific location known seems to be that of our collection in<br />

Bastrop State Park, Bastrop County. This is on the Colorado<br />

River about 30 miles southeast of Austin, Texas. This is about<br />

60 miles southwest of the Brazos River where it passes near<br />

Bryan, Texas, and, although we have not been able to collect it<br />

on the Brazos River, the range could be assumed to be from the<br />

Colorado River at Bastrop to the Brazos near Bryan. If the<br />

speading -> spreading


210 cacti of the southwest<br />

debatable assumption is followed that Engelmann’s O. intermedia<br />

from Industry, Texas, is the same plant, this would extend<br />

the range only slightly farther southwest between the two<br />

rivers.<br />

Remarks. This is a truly beautiful cactus when its huge blooms<br />

appear. It has the largest flowers of any Opuntia in the south-<br />

west except O. compressa var. allairei, which sometimes equals<br />

it in cultivation but seems always to have much smaller flowers<br />

in the wild. It would seem it would be better known than it is.<br />

However, except for that week or two when it flowers each<br />

spring, it is an uninteresting, sprawling prickly pear and so is<br />

usually overlooked or even avoided. It is also far from wide-<br />

spread, and no doubt few people have ever seen its flowers.<br />

We have been able to discover it growing only in the Bastrop<br />

State Park, where there are, however, numerous specimens. Accepting<br />

all we can of the early, general statements about its<br />

locations, we come up with a total range for it consisting of a<br />

triangle about 60 miles or less on each side between the Colorado<br />

and Brazos Rivers just east of Austin, Texas.<br />

This is, no doubt, a very local form of the O. compressa complex,<br />

more robust in size of pads and flowers, but totally without<br />

spines. It is interesting to note that no description has been<br />

given by anyone of the seeds of this plant, although several have<br />

described the fruits. In 5 years of cultivation in my garden,<br />

during which time it has bloomed repeatedly, no fruit has<br />

ripened and no seed has been set. We wonder if it may be some<br />

sort of sterile hybrid propagated entirely by separated pads.<br />

Engelmann’s O. intermedia was very incompletely described,<br />

but had similar flowers and fruits, and everyone since his time<br />

has assumed it to be the same plant as his O. grandiflora. However,<br />

he said that O. intermedia was erect and ascending to several<br />

feet high, which has always cast a doubt upon the combination.<br />

No erect plant with this flower and fruit is known in<br />

the area. However, noticing that Engelmann says he thinks he<br />

saw his O. intermedia also near Natchitoches, which is in western<br />

Louisiana, in the area of the larger and more ascending O.<br />

compressa var. allairei, and finding that that plant can produce<br />

flowers and fruits identical to those of variety grandiflora, I<br />

would advance the theory that Engelmann’s O. intermedia from<br />

Industry, Texas, was variety grandiflora, but that he tempered<br />

the description of it from his memory of the similar but larger<br />

and more ascending plant which grows in western Louisiana<br />

and which was named much later by Griffiths as O. allairei.<br />

Opuntia compressa var. allairei (Gr.)<br />

Description Plate 55<br />

roots: Tuberous, with clusters of spindle-shaped enlargements<br />

on the fibrous roots of all specimens we have examined.<br />

stems: A spreading plant, the joints of which only recline on<br />

one edge when very old and send up sprawling branches of<br />

2 to 4 pads to a height of 12 or even 18 inches. Pads elongated<br />

egg-shaped or spindle-shaped, thick and firm, not becoming<br />

flabby or wrinkled even when desiccated, the surface<br />

often somewhat tuberculate when growing but when older<br />

always becoming smooth and flat. The color is blue-green to<br />

sometimes rather yellow-green. The size of the pads is 4 to<br />

sometimes 8 inches long by 2 to 33/4 inches wide.<br />

areoles: Small and oval to round on young pads, increasing<br />

to round and 1/4 of an inch across on old pads. They are<br />

spaced 1/2 to 11/4 inches apart.<br />

spines: Often spineless, but some specimens having 1 spine<br />

each on perhaps half a dozen of the uppermost edge areoles.<br />

This spine is straight, 1/2 to 11/4 inches long, medium in thickness,<br />

round or slightly flattened, gray or whitish, and some-<br />

times slightly annulate above, with a brownish base.<br />

glochids: Yellow, many, and conspicuous, to 1/4 of an inch<br />

long, in a compact tuft, as is typical in this complex.<br />

flowers: Entirely yellow, yellow irregularly streaked with<br />

red, or yellow with red centers. Plants blooming in the field<br />

presented flowers about 3 inches across by 21/2 inches tall, but<br />

the same plants after 2 years of cultivation produced identical<br />

flowers except 41/2 to 5 inches in diameter and 3 inches<br />

tall. There were always 9 inner petals up to 21/2 inches long<br />

and 11/2 inches wide. The ovary was slender and elongated<br />

to 2 inches on the larger flowers. Stigma lobes varied from 4<br />

to 7 in number, and were white or cream in color.<br />

fruits: 1/4 to 21/2 inches long by 3/4 to 7/8 of an inch thick,<br />

elongated club-shaped, constricted at both top and bottom.<br />

At first they have some glochids on them, but these soon<br />

drop off. When ripe they are bright rose-plum in color, the<br />

pulp light red. The seeds are about 3/16 of an inch (4–5 millimeters)<br />

in diameter, thick, with narrow rims.<br />

Range. The extreme southeast corner of Texas and southwest<br />

corner of Louisiana. Specifically, along the east side of the Trinity<br />

River from the mouth of that stream to near Livingston,<br />

Texas, and east into Louisiana, perhaps, if Engelmann’s remark<br />

is taken to indicate this plant, as far north as Natchitoches, and<br />

known to grow as far east as Marksville, Louisiana.<br />

Remarks. It is obvious that this cactus is closely related to variety<br />

grandiflora, but I was greatly surprised to find the flow-<br />

ers, which have never been described before, becoming identical<br />

with those huge flowers of that form when the plant was grown<br />

in the most favorable conditions in my garden. It is different<br />

from variety grandiflora only in details of the pads and in<br />

sometimes having a few spines, and for the fact that it grows<br />

more robust and more upright and does not have the same difficulty<br />

with its fruits, these ripening and forming seeds in profusion.<br />

But the two forms respond differently to the same<br />

growing conditions in my garden in San Antonio. While variety<br />

allairei prospers and sprawls all over the place, variety grandi-


genus Opuntia 211<br />

flora in the same bed does not grow well and only the most<br />

careful treatment will keep it alive here.<br />

I consider variety allairei to be the largest, most robust form<br />

of the O. compressa complex, at least in our area. It is very<br />

much like a large edition of the small eastern form in nearly<br />

every character, although its pads are always elongated and<br />

never so broad as the eastern form shows.<br />

Variety allairei grows in that unique and beautiful southeast<br />

corner of Texas called the Big Thicket. Most people are very<br />

surprised to find a cactus growing in this densely wooded and<br />

humid area, but this cactus thrives there. It accomplishes this<br />

by growing in alkali spots which occur naturally in that area,<br />

where the trees and brush are held back by the alkaline, almost<br />

sterile soil. Here, in these natural clearings, one will often find<br />

this cactus happily growing among the coarse grasses which<br />

share these special habitats. While these special natural situations<br />

are usually widely scattered, the oil fields of the area are<br />

among the oldest in existence and have for many years poured<br />

out salty water upon the ground. The cactus has quickly taken<br />

advantage of the alkali tracts so produced by man, and among<br />

the ancient oil tools and rusting pipes of these old fields, as<br />

for instance around Batson, Texas, in Hardin County, one can<br />

find variety allairei growing in some profusion.<br />

Going east into Louisiana there are few records of the cactus,<br />

but it has been seen near Leesville. McAtee’s collection of it near<br />

Marksville is by far the most easterly, and establishes the presently<br />

known eastern limit of its range. As has been mentioned<br />

in the remarks on variety grandiflora, if we explain Engelmann’s<br />

statement about remembering having seen his O. intermedia<br />

at Natchitoches, Louisiana, by assuming that he saw this<br />

plant, that would push variety allairei’s known range some 60<br />

miles or so north of any other record.<br />

Rose and McAtee both collected plants on the lower Texas<br />

coast which they called O. allairei, Rose’s from Brownsville and<br />

McAtee’s from Matagorda Island, but having examined their<br />

herbarium specimens of these plants, I do not believe we can<br />

consider them this variety. They seem rather to be robust specimens<br />

of variety fusco-atra as are found all along the lower<br />

Texas coast. There is still no record of variety allairei west of<br />

the Trinity River.<br />

Opuntia compressa var. stenochila (Eng.)<br />

Description Plate 56<br />

roots: Usually entirely fibrous, but sometimes with some<br />

spindle-shaped to oval tuberous thickenings on them.<br />

stems: Low and prostrate, with older pads usually resting on<br />

the ground, but young pads temporarily ascending. In the<br />

spring this often results in a small clump of the young, ascending<br />

pads standing to 8 or even 10 inches tall, but during<br />

the winter these pads become flaccid and the whole thing is<br />

entirely prostrate. The pads are usually 21/2 to 4 but sometimes<br />

to 6 inches long, nearly round with only a slight constriction<br />

below to elongated oval or egg-shaped with very<br />

pronounced constriction below. These pads are of medium<br />

thickness to very thick, often 5/8 of an inch thick, tuberculate<br />

when growing but then nearly smooth, except often very<br />

shrunken and wrinkled from lack of water or from winter<br />

cold. The color is a light grayish-green or yellowish-green<br />

with a dull surface.<br />

areoles: Small and inconspicuous on young pads, becoming<br />

to almost 1/4 of an inch in diameter on old pads. They are<br />

spaced 1/2 to 1 inch apart.<br />

spines: 1 to 4 in number in the areoles on the upper edge to<br />

the upper one-half of the pad only. There is in these upper,<br />

armed areoles one main central spine which is porrect or<br />

deflexed a little, 1 to 21/4 inches long, slender to medium<br />

thickness, and round to somewhat flattened. Besides this there<br />

are usually 1 or 2 upper spines porrect or spreading upward,<br />

3/4 to 21/4 inches long, of medium thickness, and always round,<br />

plus rarely 1 lower spine 3/8 to 11/8 inches long, deflexed,<br />

slender, and flattened. All spines are entirely white or gray,<br />

or sometimes gray with bases light brown or yellowish. Rarely<br />

the upper spines have darker brown bases or brown mot-<br />

tling.<br />

glochids: Yellow, greenish-yellow, or straw-colored, few to<br />

medium in number and short at first, increasing somewhat in<br />

number and length until rather conspicuous and to 3/8 of an<br />

inch long on old pads.<br />

flowers: Light sulphur or greenish-yellow in color, 21/2 to 4<br />

inches in diameter. The ovary is a slender club about 11/4 to<br />

2 inches long. There are 10 inner petals. The stigma has 6<br />

small white lobes.<br />

fruits: Club-shaped with constricted and more or less elongated<br />

bases. The umbilicus is deeply pitted. In size they measure<br />

11/2 to 21/2 inches long. They seeds are approximately 3/16<br />

of an inch (4–5 millimeters) in diameter, thick to very thick,<br />

with narrow rims.<br />

Range. Common only in northwestern New Mexico and adjacent<br />

Arizona, but found in very scattered locations over most<br />

of the northern two-thirds of New Mexico and also collected<br />

a few times in the upper Texas Panhandle.<br />

Remarks. I have found in all herbarium collections I have examined<br />

some small, prostrate New Mexico Opuntias which do<br />

not quite fit the descriptions of O. cymochila or O. compressa<br />

var. macrorhiza, but which are usually placed in the folder of<br />

one or the other of these species that everyone expects to find<br />

in New Mexico. Many of these specimens are so incomplete that<br />

there is little way to prove what they are, but taking those<br />

which present fruits and seeds as well as pads, and lumping<br />

them together, one gets a rather homogeneous group of speci-<br />

mens. In living collections I have found the same plants grow-


212 cacti of the southwest<br />

ing, and finally, we have collected specimens with the same<br />

characters in a number of places in New Mexico, Arizona, and<br />

Texas.<br />

I soon noticed that when I compiled the description of this<br />

body of plants I had thus brought together, it paralleled remarkably<br />

well Engelmann’s old description of O. stenochila.<br />

The only aspects in which it exceeds the limits of that description<br />

are in the maximum length of spines found on some of my<br />

specimens and in the slightly smaller seeds on some than he gave<br />

for the species. When it is noticed that the majority of these<br />

maverick plants are from northwestern New Mexico, including<br />

some from just east of Zuni, which must be from very near the<br />

type locality of O. stenochila (it having been given as “canyon<br />

of Zuni, western New Mexico”), it becomes very convincing<br />

that here we have a collection of that long-lost cactus.<br />

But if so, then why has the cactus been so long overlooked,<br />

and why are there no specimens called by this name in any collection<br />

except the type specimens? I think this goes back to<br />

Wooton and Standley. These men were the acknowledged authorities<br />

on New Mexico plants in the next generation after<br />

Coulter, and somehow they misunderstood this plant so much<br />

that they dismissed it by saying, “Known only from the original<br />

collection by Bigelow. We have seen no material of this species.”<br />

And almost everyone since has been all too quick to take them<br />

at their word; so, since we see what we expect to see, whatever<br />

specimens of this cactus have been found since that time have<br />

been quickly assigned to O. cymochila or O. compressa var.<br />

macrorhiza, where they almost, but not quite, fit. Being more<br />

careful, Peebles erected a new species, O. loomisii, to take care<br />

of the Arizona specimens of this same cactus, and so the more<br />

western examples of it were dispersed in other folders.<br />

But perhaps this is one place where we should pay attention<br />

to Britton and Rose. After and in spite of Wooton and Standley’s<br />

statement about O. stenochila, they said, “It is the common<br />

low, spreading Opuntia of northwestern New Mexico and<br />

Arizona.” I agree with them, and think that many herbarium<br />

specimens would be shifted out of folders where they don’t quite<br />

look comfortable into the folder of this form if we regarded it<br />

as a plant we might expect to find and if the specimens collected<br />

were complete enough and well-enough preserved to show its<br />

differences from the other forms.<br />

The cactus is obviously one of the O. compressa complex. Engelmann<br />

himself reduced it in a later writing from a separate<br />

species to a variety, and Coulter followed him. Its fruits are<br />

almost exactly like those of variety macrorhiza, both in shape<br />

and in the limits of their sizes. The seeds of the two forms are<br />

very nearly alike, and both may have either fibrous or tuberous<br />

roots. But variety stenochila presents yellow glochids and longer<br />

spines than are typical of variety macrorhiza, and its pads are<br />

gray or yellowish-green and dull, while those of variety macrorhiza<br />

are deep and shining green. The flowers are also dif-<br />

ferent.<br />

When the fruits are considered, it is immediately obvious that<br />

variety stenochila is not just a poorly armed O. cymochila, as<br />

its pads have led many to believe. Its long fruits with constricted<br />

bases and its thick, narrow-margined seeds show that it<br />

belongs with the O. compressa group.<br />

If this is truly variety stenochila, then it is, as originally<br />

stated, a plant of northwest New Mexico. One may actually<br />

find it fairly easily in that area. It is my opinion that it spreads<br />

into Arizona, where it was described as O. loomisii by Peebles.<br />

He states that that plant comes into New Mexico, and I know<br />

of nothing else in this state to equal his description. However,<br />

the eastern limit of its range is very hard to define. I have the<br />

cactus collected in the Mescalero Sands. This is east of Roswell,<br />

and getting near the southeastern corner of New Mexico. I have<br />

also collected what must be this plant in Gray County, east of<br />

Pampa in the Texas Panhandle, which marks by far the easternmost<br />

known record of it.<br />

Opuntia pottsii SD<br />

Description Plate 56<br />

roots: Always having a large central taproot 3/4 to 11/2<br />

inches in diameter and 6 inches to more than a foot long. This<br />

may be smooth and carrot-like or, in rocky ground, is often<br />

contorted and constricted here and there by the edges of<br />

rocks. Sometimes it subdivides into 2 or 3 fleshy branches, but<br />

it is not ever a tuberous, spherical, or spindle-shaped, potato-<br />

like enlargement on otherwise fibrous roots. When damaged<br />

this root slowly exudes a milky sap.<br />

stems: Although small, this is basically an ascending plant<br />

with pads only touching the ground when extremely dehydrated<br />

or in their winter-shrunken condition. It appears not<br />

to root from the pads, so it is not mat-forming, but stands as<br />

a small clump of upright pads 6 to 12 inches tall from a central<br />

base which is often a short, cylindrical trunklet. The pads<br />

are 11/2 to 5 inches long by 1 to 4 inches wide, almost round<br />

to elongated egg-shaped, with some constriction at the bases.<br />

They are thin or medium thickness, the surface flat without<br />

raised areoles. In color it is glaucous blue-green, often purplish<br />

around the areoles when young and healthy, but fading<br />

to yellow-green when old or unhealthy. The leaves of this<br />

plant are extremely tiny, being around 1/8 of an inch long.<br />

areoles: To only 1/8 of an inch long on young pads, but becoming<br />

to 1/4 of an inch in diameter on old pads. They are<br />

spaced 3/8 to 1 inch apart.<br />

spines: Having spines on only the upper edge or at most the<br />

upper one-third of the pad. These upper areoles have 1 to 3<br />

spines which are straight although usually twisted and rather<br />

slender. They may be round or flattened or sometimes ridged,<br />

and are whitish, gray, or sometimes gray mottled with tan,<br />

or brownish. There will be 1 or 2 upper spines 1 to 21/2 inches


genus Opuntia 213<br />

long, deflexed or spreading, plus sometimes 1 lower, very<br />

deflexed spine 1/2 to 1 inch long. The plant has been said to<br />

be occasionally spineless, but I have not seen such a speci-<br />

men.<br />

glochids: Greenish-yellow to dirty straw-colored, usually<br />

rather numerous in compact clusters, 1/8 to sometimes 1/4 of<br />

an inch long on old pads.<br />

flowers: Brilliant purplish or rose-red and very beautiful. 2<br />

to 23/4 inches in diameter and 21/2 to 3 inches tall. The ovary<br />

is very slender and 1 to 2 inches long, with some glochids on<br />

it. There are about 7 or 8 inner petals. The filaments are green<br />

below and yellowish above, the anthers bright yellow. The<br />

style is pinkish and the stigma lobes cream-colored, fat and<br />

velvety, and 5 to 7 in number.<br />

fruits: 11/2 to 2 inches long, slender club-shaped, being only<br />

3/4 of an inch in diameter at the upper, widest part above the<br />

slender, attenuated base. The apex is deeply pitted. When<br />

ripening, they become light red. The seeds are about 3/16 of an<br />

inch or a little larger (4–51/2 millimeters) in diameter in my<br />

observation, although Coulter says 31/2 to 4 millimeters. These<br />

seeds are thick—Engelmann says they are the thickest of the<br />

Opuntias. The rims are of medium width, blunt instead of<br />

acute, and irregular.<br />

Range. From Chihuahua, Mexico, north past El Paso and<br />

through the western part of the Big Bend into the Davis Mountains<br />

of Texas and on into southeastern New Mexico. Known<br />

to be found in the southern Guadalupe Mountains and to extend<br />

in New Mexico at least as far north as Caprock and Roswell.<br />

Remarks. This cactus is very close to the O. compressa group<br />

and more study may show that it should be included in it, but<br />

the step seems premature with no more than our present knowledge.<br />

At any rate, it has very distinct differences from any of<br />

the varieties of that group. It was first described from Chihuahua<br />

specimens by Prince Salm-Dyck. Engelmann redescribed<br />

it from Texas collections, calling it O. filipendula. Once it was<br />

determined by Britton and Rose that these two were the same,<br />

there has been little confusion over this species.<br />

The cactus appears at a glance much like O. compressa var.<br />

macrorhiza, until one notices the more blue-green, glaucous<br />

coloring and the fact that, although it is no taller than the other<br />

plant, O. pottsii is not a fully prostrate, diffuse thing rooting<br />

and spreading from chains of pads on the ground, but a compact<br />

plant ascending from one center by many upright or<br />

sprawling pads. The spination is similar, each plant having either<br />

round or flattened spines on only the upper areoles, but O.<br />

pottsii has more slender and usually longer spines, although never<br />

reaching the maximum number found in some specimens of va-<br />

riety macrorhiza and always lacking the lower bristles often<br />

found on that other cactus. If one digs up the plants, he will<br />

find there an obvious difference between the two forms. The<br />

root of O. pottsii is always a central taproot extension of the<br />

thick, trunklike stem which often goes without interruption and<br />

with very little taper up to a foot into the ground. It is basically<br />

different from the globular to spindle-shaped tubers often found<br />

on the fibrous roots of the O. compressa forms. There is also a<br />

striking difference in the leaves of this plant, those of O. pottsii<br />

being very tiny, while those of all forms of O. compressa examined<br />

in this study are 3 or 4 times as large. But when the<br />

beautiful purplish or rose-red flowers of O. pottsii appear one<br />

really appreciates the difference between the two plants. These<br />

are certainly some of the most beautiful flowers found among<br />

the Opuntias.<br />

O. pottsii was reported by Anthony from western Brewster<br />

County in the Big Bend, although I have been unable to recollect<br />

it there. Dr. Rose collected it “in the valley of the Rio<br />

Grande below El Paso, Texas,” and Engelmann’s specimens were<br />

from near Dona Ana and San Elizario in the El Paso area, but I<br />

have seen no recent specimens from that locality. Engelmann<br />

said it was growing there on alluvial prairies, and in looking<br />

for likely spots, I found that almost everything there which<br />

might be described at all by the term of alluvial prairies is now<br />

a part of the city, in cultivation and irrigation, or otherwise<br />

developed, so I wonder if the cactus may not have been elimi-<br />

nated around El Paso by the developments.<br />

The cactus does seem to be strictly a resident of alluvial flats.<br />

It still grows along stream beds in the Davis Mountains of<br />

Texas. Engelmann’s statement that the eastern boundary of its<br />

range was the Limpia was correct. This is a creek in the eastern<br />

part of the Davis Mountains, and the plant has not been recorded<br />

from east of it in Texas. Nor does his additional statement<br />

of “El Paso and eastward on the Pecos,” contradict this.<br />

It does not mean on the lower Pecos, as many have thought.<br />

Limpia Creek runs north out of the mountains into the Pecos<br />

River just below the town of Pecos, Texas, and the cactus occurs<br />

to the Pecos only above this point. Farther north it enters<br />

New Mexico, to be found in the creek valleys of the southeasternmost<br />

part of the Guadalupe Mountains and east across the<br />

Pecos to at least as far north and east as Caprock, New Mexico.<br />

If the New Mexico herbarium material usually labeled O. ballii<br />

were considered as this plant, as some think it should be, then<br />

the range would extend over all of the southeastern corner of<br />

New Mexico and along the eastern boundary of that state to<br />

Logan on the Canadian River.<br />

Besides certain morphological differences, there is an ecological<br />

reason why I hesitate to lump these northern specimens<br />

with the Texas O. pottsii. There has been, in the past few years,<br />

a marked die-off of the O. pottsii population in the Davis<br />

Mountains, which has become so bad that in our old collecting<br />

spot north of Alpine, Texas (where a few years ago we could<br />

count dozens of healthy specimens), in 1966 I failed to find one<br />

remaining example. The preceding few winters had been very<br />

severe, setting new low temperature records for the area. Dr.<br />

Warnock, botanist at Sul Ross State College in Alpine, states<br />

that most specimens of this cactus in the area were killed by this<br />

unusual cold. Remembering that this is a Chihuahuan species


214 cacti of the southwest<br />

restricted to alluvial bottoms in the south of its range and noting<br />

this evidence of tenderness in the plant, I wonder if it can be<br />

the same plant which grows rather profusely in the very severe<br />

conditions of the eastern New Mexico mountains and plains.<br />

Statements by those who have observed many of the plants<br />

growing far up in eastern New Mexico that they bloom yellow<br />

as well as red and often have tuberous roots instead of the typical<br />

taproot add to my doubts about them. Further study of<br />

these plants is needed to determine whether we have in them a<br />

transitional form bridging the gap between O. pottsii and the<br />

O. compressa complex or whether they are something entirely<br />

different.<br />

The specimens definitely known to be O. pottsii show us a<br />

rather strange little Opuntia with beautiful flowers. While it<br />

grows over a rather good-sized area, it seems always to be far<br />

from common anywhere. It is an easy cactus to grow, in my<br />

experience, not being harmed by moisture as easily as are some<br />

others. Perhaps this is because it grows naturally in valleys in-<br />

stead of upon the more dry hills and ledges.<br />

Opuntia ballii Rose<br />

Description Plate 56<br />

roots: According to Rose, “somewhat tuberous.” In our observation<br />

a large specimen had small, spindle-shaped tubers<br />

upon the fibrous roots, but plants started as cuttings from the<br />

original had not developed tubers upon their roots in several<br />

years’ growth.<br />

stems: Prostrate and spreading. The pads are 23/8 to 4 inches<br />

long, almost circular to broadly egg-shaped, very thick for<br />

their size, pale yellowish-green and glaucous, the surface entirely<br />

smooth without raised areoles.<br />

areoles: Small, nearly round.<br />

spines: 2 to 4 in number on most or all areoles. There are 1<br />

to 3 main spines spreading upward or erect, 11/2 to 23/4 inches<br />

long, plus 1 lower, deflexed, and much shorter spine. All<br />

spines are straw or pale brownish in color, heavy, very rigid,<br />

straight, and a little flattened.<br />

glochids: Very conspicuous, each areole having a great, compact<br />

mass of bright yellow glochids 1/4 to 1/2 inch long.<br />

flowers: Small, 11/4 to about 2 inches across and about 2<br />

inches tall, deep rose-red. The ovary is about 11/4 inches long<br />

and very narrow club-shaped or almost linear, with very few<br />

yellow glochids in the areoles. The outer perianth segments<br />

have green midlines, the smaller ones shading to flesh-colored<br />

on the edges, the larger ones to burnt-orange pinkish on the<br />

edges. There are about 6 inner segments about 1 inch long and<br />

wide, with blunt ends notched at the apex. The base and<br />

most of each inner segment shade from old red through deep<br />

rose-red, the edges and tips cerise. The filaments are green at<br />

the base, then cream-colored, the anthers cream. The style is<br />

short and whitish. The stigma has 5 cream-colored, fat, and<br />

thick lobes.<br />

fruits: Small and very slender, 3/4 to 11/4 inches long, very<br />

narrow club-shaped, being only 1/4 of an inch thick at the<br />

center of the widest part, with greatly constricted bases and<br />

rather narrowed, pitted umbilici. They are spineless. The seeds<br />

are slightly over 1/8 of an inch (about 31/2 millimeters) broad,<br />

and thick for their size.<br />

Range. According to Dr. Rose, “the dry mesa beyond Pecos,<br />

Texas.”<br />

Remarks. This remarkable little cactus was found and described<br />

by Dr. Rose in 1911. He gave for its type location Pecos, Reeves<br />

County, Texas.<br />

Almost immediately after Rose described the species, Wooton<br />

said it was common in New Mexico, and later Wooton and<br />

Standley referred it to O. filipendula (O. pottsii) as a synonym.<br />

Dr. Rose had opportunity to publish a rebuttal and did so,<br />

stressing that O. ballii grows in a different habitat, has smaller<br />

fruits, stouter and more erect spines, and different areoles than<br />

that plant.<br />

There thus arose a disagreement about this cactus which has<br />

continued until this day. Some students, influenced by Wooton,<br />

have expected to find it in New Mexico. We often convince our-<br />

selves that we have found what we expect to find, at least to<br />

our own satisfaction, and the New Mexico herbarium contains<br />

a whole series of plants labeled O. ballii, specifically from<br />

Eunice, Tatum, Caprock, Texico, near Tucumcari, and near Logan,<br />

New Mexico. If these are correctly assigned, the cactus<br />

extends its range all up along the eastern edge of the state.<br />

I have studied the cacti growing in all of these areas as much<br />

as possible during a long search for the true O. ballii, collecting<br />

most of them in the field rather than relying on the herbarium<br />

specimens. The plants growing west of Tucumcari seem<br />

clearly to be O. compressa var. macrorhiza, which might be expected<br />

there, since I have collected it within less than one hundred<br />

miles both southeast and northwest of this location. The<br />

Logan specimens are too incomplete to identify certainly, and I<br />

was unable to find the plant growing in that area today. The<br />

Texico plants are quite certainly O. cymochila, which grows in<br />

profusion there. The Caprock specimens appear to be O. pottsii,<br />

which I have collected myself near Caprock. In short, I have<br />

found no New Mexican specimens under this name which<br />

seemed to me to be actually O. ballii, and most of them seemed<br />

clearly other forms.<br />

But the plant was originally described from Texas, and Britton<br />

and Rose long ago denied New Mexico the plant in restating<br />

its range to be west Texas. What was the plant they actually<br />

collected near Pecos?<br />

When I examined Rose’s own specimens it immediately seemed<br />

clear to me that his was a different cactus, not only from<br />

O. pottsii, but from all those collected and labeled O. ballii in


genus Opuntia 215<br />

New Mexico. His specimens are easy to recognize by their long,<br />

heavy spines, their yellow glochids mostly to 1/2 inch long, and<br />

very nearly the longest found on any Opuntia, as well as by<br />

their very small fruits. None of these characters are shared by<br />

O. pottsii or by any New Mexico plants I have seen, to say<br />

nothing of the difference in roots between O. pottsii and this<br />

plant. I became persuaded that we have here a unique small<br />

cactus perhaps not found outside of the Pecos area of west<br />

Texas. And there is only one sign of its ever having been seen<br />

again since Rose’s original collection. ‘There is in the Southern<br />

Methodist University herbarium a single unlabeled specimen<br />

which seems clearly to be this plant. It was collected by Warnock<br />

22 miles northwest of Pecos, Texas. What was needed was the<br />

rediscovery of the thing itself so we could know more about it.<br />

I made repeated trips to Reeves County and spent days<br />

scouring all of the region I could, but perhaps the most frustrating<br />

experience of this study has been the failure to relocate<br />

O. ballii. There is a large amount of dry mesa, and it must still<br />

be there, but also much of the area is now irrigated and farmed,<br />

and it may have been largely eradicated.<br />

I had given up hope of ever seeing the cactus alive, when one<br />

day I was looking over the very interesting cactus gardens of<br />

Mr. Clark Champie, the widely known cactus student and dealer<br />

of Anthony, New Mexico-Texas. Mr. Champie began showing<br />

me some plants he could not identify, and there it was! Its small,<br />

thick, very light green pads with their very heavy spines often<br />

as long as the pads themselves and the brilliant yellow glochids<br />

so profuse and so long in each areole were unlike anything I<br />

had seen before. The tiny fruits less than an inch long in many<br />

cases and no thicker than a lead pencil were complete confirmation.<br />

Each fruit contained a dozen or so seeds hardly over<br />

1/8 of an inch in diameter, but very thick.<br />

So I have seen O. ballii alive, have my own cutting of it, and,<br />

thanks to Mr. Champie, am able to present a photograph of it,<br />

but still I can tell no more about where it grows than Rose’s<br />

statement. Mr. Champie says that this plant was given to him<br />

some 8 or 10 years ago by a fellow cactophile who didn’t remember<br />

where it was collected beyond the fact that it was in<br />

west Texas. This may very well be the only specimen of this interesting<br />

species living today. To those who might be intrigued<br />

and want to try to find it, I can say nothing more than for them<br />

to try the dry mesa beyond Pecos, Texas, and good luck! It must<br />

be there somewhere.<br />

Opuntia plumbea Rose<br />

Description Plate 56<br />

roots: Having a very unusual underground structure consisting<br />

of what appears to be a fleshy rhizome rather than a<br />

root. This structure is 1/2 inch or a little more in diameter,<br />

and runs horizontally just under the ground, giving off pads<br />

at intervals of 4 to 8 inches along its length. It has been observed<br />

with a total length of about 4 feet. Fibrous roots are<br />

produced from this apparent rhizome.<br />

stems: Plants ascending at the point of each sprouting from<br />

the underground stem, but not standing over 1 or 2 pads high.<br />

These pads are almost circular to broadly egg-shaped and 11/4<br />

to 21/2 inches long. They soon develop into a cluster of small<br />

pads standing to only 4 inches or so high but in time becoming<br />

to 8 or 12 inches across at each growing point. These<br />

separate clusters finally meet at their edges to form a more<br />

or less solid mat of pads up to several feet across. The pads<br />

are rather thin and the surface somewhat tuberculate by<br />

raised areoles. The color of the surface is very glaucous, dull<br />

blue-gray, sometimes described as lead-colored.<br />

areoles: Large for the size of the pads, being oval and 3/16 of<br />

an inch or so long, spaced 3/8 to 1/2 inch apart.<br />

spines: 1 to 4, but usually 2 in number, found in upper areoles<br />

of the pads only. They are gray in color, more or less mottled<br />

with pale brown. There is one main spine porrect or slightly<br />

deflexed, which is 1 to 17/8 inches long, slender, straight, and<br />

round, or nearly so, although sometimes slightly flattened at<br />

the base. Sometimes also present will be 1 or 2 upper spines<br />

standing porrect, or practically so, 1 to 2 inches long, slender,<br />

straight, and round. There may be also 1 lower deflexed spine<br />

1/2 to 11/4 inches long.<br />

glochids: There are conspicuous, bright, red-brown glochids<br />

in all areoles. They are in compact clusters and are 1/8 to 3/16<br />

of an inch long.<br />

flowers: Small, being 11/2 inches or so long and wide, purplish<br />

in color. The ovary is less than 1 inch long, but fairly<br />

broad. The stigmas are white.<br />

fruits: Small, being 5/8 to 7/8 of an inch long and broadly<br />

pear-shaped with pitted umbilici. The seeds are slightly over<br />

1/8 of an inch (31/2–4 millimeters) in diameter.<br />

Range. The San Carlos Indian Reservation in Arizona and the<br />

edge of New Mexico west of Silver City.<br />

Remarks. Dr. Rose described this strange little cactus from a<br />

collection made on the San Carlos Indian Reservation of Arizona<br />

in 1904. The plant must be very rare, as there is no record<br />

of its ever having been seen again in Arizona. More recent<br />

works have generally ignored it or else submerged it as a synonym<br />

of O. compressa var. macrorhiza. It has never been con-<br />

sidered a New Mexico plant.<br />

Mr. Clark Champie of Anthony, New Mexico-Texas, who has<br />

a fine cactus collection, has a strange Opuntia which he has kept<br />

because of its uniqueness. He had had it for about 5 years when<br />

I first saw it, during which time it had spread in a most unusual,<br />

linear manner. When he transplanted it he was surprised to find<br />

that it had in this time developed a long, rhizome-like structure<br />

which, growing horizontally, was at that time nearly 3 feet<br />

long. It was observed that this structure had sprouted a series


216 cacti of the southwest<br />

of pads at intervals of about 6 inches all along its length, each<br />

of these points in time forming a new growing center with numerous<br />

pads in a cluster. Since planted in the ground this cactus<br />

has continued the elongation of this underground structure, with<br />

new growth centers from it.<br />

The manner of growth which results is rather well described<br />

by Rose’s word, “creeping,” in his description of O. plumbea.<br />

This is a term which Rose used concerning no other prostrate<br />

cactus and which I take to mean specifically the creeping,<br />

spreading growth outward which results in this cactus from<br />

these peculiar underground growths. I know of only one other<br />

cactus in the U. S. Southwest with a similar creeping and sprouting<br />

underground structure. This is O. arenaria, whose underground<br />

part has areoles with many glochids all over it. These<br />

are lacking on this plant, and the two cacti are different in al-<br />

most all other respects.<br />

Mr. Champie’s strange cactus has small pads of a dull, bluish-<br />

gray color, long slender spines, and conspicuous, bright, red-<br />

brown glochids. It blooms with a small purplish flower and has<br />

small though broad fruits less than 1 inch long. It fits perfectly<br />

Rose’s description of O. plumbea, and we are certain it is that<br />

long-lost plant. The specimen in Mr. Champie’s garden was presented<br />

to him by a collector who gave little detail of the location<br />

of its collection, saying only that he found it in New Mex-<br />

ico somewhere west of Silver City. Although we have been unsuccessful<br />

in collecting it ourselves, I feel obliged to list it here as<br />

a species probably growing in New Mexico. Since I cannot give<br />

the exact location of it, I realize that this does not constitute an<br />

actual record in the strict scientific sense; however, I feel it<br />

is important to tell of this apparent instance of the plant in our<br />

area in the hope that it will stimulate someone to re-collect it<br />

and give us an exact location for it.<br />

Opuntia drummondii Graham<br />

“Crow-Foot Prickly Pear,” “Cock-Spur Cactus,” “Cockle-<br />

Burr Cactus,” “Sand-Burr Cactus”<br />

Description Plate 57<br />

roots: Either entirely fibrous or often having several oval<br />

tubers up to 21/2 inches long and 11/2 inches thick upon these<br />

roots.<br />

stems: A very prostrate and diffuse cactus, forming mats<br />

from 1 to sometimes at least 15 feet across, but hardly over<br />

4 or 5 inches high, as all older pads lie on the ground. The<br />

pads are broadly oblong to elongated club-shaped, 1 to 41/2<br />

inches long by 1 to 21/4 inches wide. They are often 3/4 of an<br />

inch and sometimes even 1 inch thick, young joints being almost<br />

globular. The surface is smooth, without raised areoles<br />

when well-watered. They shrink and wrinkle when dry, but<br />

can never be described as tuberculate. They are deep green<br />

with darker, bluish coloring around the areoles, becoming<br />

lighter green when old. These pads are very loosely articulat-<br />

ed, coming off the plant at a touch.<br />

areoles: Round or oval, 1/8 to 3/16 of an inch in diameter, 3/8<br />

to 3/4 of an inch apart.<br />

spines: There are 1 to 4 straight spines on almost all areoles<br />

or often on upper edge areoles only. There is 1 porrect spine<br />

1/2 to 11/2 inches long, flattened, twisted, and of medium<br />

thickness. There may be 1 or 2 upper spines 1 to 15/8 inches<br />

long and round in shape. Sometimes there is 1 lower spine 1/4<br />

to 3/4 of an inch long, slender, round or flattened, deflexed<br />

or porrect. At first these spines are light brown, but they soon<br />

fade to gray with the tips remaining clear yellow or else becoming<br />

blackish.<br />

glochids: Greenish-yellow or bright straw-colored, 1/8 to 1/4<br />

of an inch long, average number to many, in compact clusters.<br />

flowers: Clear lemon to greenish-yellow, sometimes with a<br />

slightly deeper yellow coloring in the center. They are 21/2 to<br />

3 inches in diameter. The ovary is 7/8 to 1 inch long, with a<br />

few short, red glochids on it. There are 8 inner petals. The<br />

filaments are orange, the anthers yellow. The stamens are very<br />

sensitive, moving at a touch. The stigma has 5 or 6 thick, yel-<br />

low or slightly greenish-yellow lobes.<br />

fruits: Light red when ripe, club-shaped, with some constriction<br />

below and a shallow to rather deeply pitted umbilicus at<br />

the top. These fruits are 1 to 11/2 inches long by 1/2 to 5/8 of an<br />

inch thick at the thickest point. The seeds are not quite 3/16 of<br />

an inch (about 4 millimeters) in diameter, regular, and rather<br />

thick, with rather narrow and blunt rims.<br />

Range. Previously known from the beach areas of North Carolina<br />

through Florida to Alabama. It is known to grow in our<br />

area only in a strip a mile or so long near the tip of Bolivar<br />

Peninsula, in front of Galveston Bay. Perhaps it also grows in<br />

Louisiana.<br />

Remarks. I had been told by Mr. H. C. Lawson, the San Antonio<br />

cactus dealer so well known during more than 30 years of business,<br />

about the strange little prickly pear which grew on the<br />

Bolivar Peninsula directly across the pass from Galveston. I had<br />

repeatedly failed to locate it myself, and had about concluded<br />

that it had been eliminated by the developments which that area<br />

has undergone through the years. Then Mr. Glenn Spraker, a<br />

collector in Houston, told me of such a cactus which his father-<br />

in-law, Mr. Herman McGowan, had noticed during fishing trips<br />

on the peninsula. These men were so kind as to take me to the<br />

cactus, and it turned out to be O. drummondii, previously unknown<br />

in Texas.<br />

The reason I had not found it earlier is that it grows only on<br />

the bay side of the peninsula from about Baffle Point toward the<br />

tip of the peninsula. This narrow strip is perhaps the highest part<br />

of the peninsula, but since the Intracoastal Canal was dredged<br />

down the length of the peninsula, it is entirely separated by this<br />

canal from the major part of the peninsula. It now forms a long,


genus Opuntia 217<br />

slender island not over 1/4 mile wide and totally uninhabited<br />

except by cattle, so anyone going to hunt this cactus has to have<br />

the interesting experience of hunting cactus by boat, since there<br />

is no other way to get to this island.<br />

Once having gotten there, the collector will find the highest<br />

part of this strip to be almost covered by mats of O. drummondii.<br />

The small pads are flat on the very sandy ground or<br />

even partly covered by blowing sand. With the sparse grass<br />

growing up even higher through the cactus mats, they are almost<br />

impossible to see until one is within a few feet. And if a<br />

person fails entirely to notice them and makes the mistake of<br />

stepping into the mat of them, his foot comes up with sometimes<br />

half a dozen pads pulled loose and firmly stuck by their spines<br />

into his shoes, his trousers and, if he is little less fortunate, into<br />

his ankles. With its pads so loosely held, this cactus must be a<br />

constant hitchhiker on the legs of animals. It is well named<br />

the sand-burr cactus. Its adaptation for animal dispersal is probably<br />

even more cruel than that of the sand-burr or cockle-burr.<br />

However, no doubt the pads are broken off and distributed<br />

wholesale by storm tides as well, as there were unmistakable<br />

signs that the last hurricane had swept large logs across this<br />

ridge.<br />

Although very common on and near beaches of the southeastern<br />

part of the U. S., O. drummondii has not been reported<br />

before in Texas. It is interesting, but probably fruitless to speculate<br />

upon how it happens to be growing here, so far west of<br />

Alabama, its previous definitely known western limit. This location<br />

in Texas is at the point where ships enter Galveston Bay<br />

and where more recently barges and boats come by constantly<br />

after having traveled west along the Intracoastal Waterway. It<br />

is not hard to imagine pads of this cactus moving west with<br />

them, but there is no evidence of this, and if it happened, it was<br />

long ago, as a large area is covered with this cactus now.<br />

A specimen which appears to be O. drummondii is in the U. S.<br />

National Museum, collected, according to the almost illegible<br />

label, in 1910, by McAtee at Caudwerdee (?), Louisiana. I have<br />

not been able to find this location, but this raises the possibility<br />

that the plant grows somewhere along the coast of Louisiana.<br />

This cactus was regarded by Coulter as very close to the eastern,<br />

typical form of Opuntia compressa, and it does have similarities,<br />

particularly of flower, fruit, and seed to that cactus. It<br />

is like the O. compressa group also in often having tuberous<br />

roots, but in thickness of pads it surpasses all of those forms.<br />

Britton and Rose, on the other hand, in devising their series<br />

within the genus Opuntia, set up the series Curassavicae, made<br />

up of a number of small cacti from the West Indies characterized<br />

by fragile branches which separate at a touch, and naturally put<br />

this cactus into that series, thus separating it entirely from the<br />

O. compressa complex, whose pads have to be twisted off. However,<br />

this character seems a doubtful one upon which to erect a<br />

series. The looseness of joints is shared by a number of other<br />

cacti described for the U. S. One is O. tracyi Britt., which grows<br />

on the coast from Florida to Mississippi, and which seems very<br />

close to O. drummondii. Another is O. nemoralis Gr., from northeast<br />

Texas, which would seem to be a typical O. compressa var.<br />

humifusa except for this feature. It has not been seen since its<br />

first description and nothing appears today in its area except<br />

variety humifusa, which makes it, in our estimation, of doubtful<br />

validity. A third is O. fragilis of northwest Texas and New<br />

Mexico and far into the north. However, even though this cactus<br />

has similar loose pads, it has dry fruits, and so Britton and<br />

Rose place it into another series entirely. O. schottii, a Corynopuntia<br />

of south Texas has this character, and of course there<br />

are various of the large chollas, such as O. tunicata and O. fulgida<br />

featuring this. It seems to be a feature running through<br />

many major segments of the genus and not one upon which any<br />

major divisions of the group should be made.<br />

Opuntia fragilis (Nutt.) Haw.<br />

“Brittle Cactus,” “Fragile Prickly Pear”<br />

Description Plate 57<br />

roots: Fibrous.<br />

stems: A low-growing cactus almost completely prostrate or<br />

with spreading branches rising to only 6 or 8 inches high, but<br />

forming very dense mats up to a foot or two in diameter. The<br />

joints begin as almost globular outgrowths up to 1/2 or even<br />

5/8 of an inch in diameter. In growing, they first elongate,<br />

reaching to 1 or 11/2 inches long without broadening, so they<br />

are at this stage almost cylindrical. If the plant is not robust<br />

its whole growth may remain like this, but if it is situated so<br />

that it can proceed fully, these joints then begin broadening<br />

until they become small, very thick pads 11/2 to 2 inches long<br />

by 3/4 to 11/4 inches wide by about 5/8 of an inch thick. These<br />

joints are very loosely attached and separate at a touch. They<br />

are often very wrinkled and flaccid.<br />

areoles: Small to medium size, with some white wool when<br />

young, situated about 3/8 to 1/2 inch apart.<br />

spines: 1 to sometimes at least 7, spreading, fairly stout, round<br />

or nearly so, and 1/4 to 1 inch long. They are whitish with<br />

darker tips or straw-colored when matured, but dark brown<br />

when growing.<br />

glochids: Very few and short, and yellowish in color.<br />

flowers: Clear yellow, sometimes with orange centers. They<br />

are up to about 2 inches in diameter. The stamens are yellowish.<br />

The stigma has 4 to 6 green stigma lobes. The ovary is<br />

small and almost spherical.<br />

fruits: Oval or egg-shaped, 1/2 to 1 inch long, with pitted<br />

flower scars. They have some short spines on the upper areoles<br />

and become dry when ripe. The seeds are large, around 1/4 of<br />

an inch (5–7 millimeters) in diameter, flat, with broad, ir-<br />

regular rims.<br />

Range. Extreme northern New Mexico into Colorado and Utah,<br />

Corny- -> Coryn-


218 cacti of the southwest<br />

and extreme northwestern Texas across the Oklahoma Pan-<br />

handle and on into Kansas. It is found at various scattered locations<br />

throughout the northwestern U. S. from Wisconsin to Washington<br />

and on far into Canada.<br />

Remarks. O. fragilis is the first of our dry-fruited Opuntias—<br />

that is, those in which the fruits become dry and their walls<br />

papery instead of juicy and pulpy when ripe—for us to study.<br />

These cacti have another character in common, the possession of<br />

green stigma lobes in the flowers. If one is fortunate enough to<br />

see both their flowers and fruits these two characters make them<br />

easy to know, but as some of them bloom and fruit very sparingly,<br />

most people see their vegetative parts only, and some of<br />

them are rather hard to recognize from just these parts.<br />

O. fragilis grows in very sandy soil, and anyone who has tried<br />

to treat it to the same soil and extreme heat as he does most<br />

other cacti will find it is fragile indeed. It just will not grow in<br />

the typical hot desert situation or in heavy soil. But it is not<br />

fragile in regard to cold. It is one of our most northern cacti,<br />

said to grow in Canada almost to the arctic circle. Of course, its<br />

name comes from its loosely articulated joints, which are dislodged<br />

by a touch and which some say can even be loosened and<br />

distributed by the wind. A person or an animal, not seeing the<br />

low mat of these little pads, often steps away with pads securely<br />

stuck to clothing or fur—or flesh—to be shaken off later and to<br />

root wherever they fall.<br />

One seeing the cactus only in the form it takes when less than<br />

robust might never think of it as a prickly pear, but as some sort<br />

of tiny cylindrical jointed cactus. However, when growing well,<br />

the joints broaden into thick, but definitely flattened little pads<br />

up to 2 inches long. The flowers and fruits are not well known<br />

because it does not flower at all unless very well situated, often<br />

relying upon the scattering of the little joints to propagate it for<br />

years at a time before every factor of season and soil pleases it<br />

and it blooms. For this reason it rarely blooms in cultivation.<br />

This strange little northern cactus just enters the edge of our<br />

southwestern area, living only in extreme northern New Mexico<br />

and the panhandles of Oklahoma and Texas. Here its locations<br />

are not numerous and are very scattered. In Texas it can occasionally<br />

be found in the sandy breaks along the Canadian River<br />

north of Amarillo. There once was a good population of it along<br />

the railroad right-of-way northeast of Amarillo, but this is apparently<br />

all gone now, wiped out in the last few years, perhaps<br />

by the herbicides now used on such right-of-ways. Although I<br />

lived two years in the Oklahoma Panhandle and was in the field<br />

over much of that area, I know of no place where it grows there<br />

except on the Black Mesa, that remarkable formation in the extreme<br />

northwest corner of the state. It appears from old accounts<br />

that the cactus may have been more widespread in the<br />

past in the Oklahoma and Texas panhandles, but it is my theory<br />

that it has been practically eliminated by the farming which has<br />

been practiced on almost all of that area, or where there has not<br />

been farming, by the drifting sand of the dust-bowl days. It<br />

would have been limited to the most sandy areas, the very areas<br />

where drifts sometimes covered automobiles and reached the<br />

eaves of buildings, and these would surely have covered and<br />

smothered this tiny cactus. Wherever we find this northern cactus<br />

in our area it is just the extreme southern extension of its<br />

range, and we get the feeling that it is not really one of ours, but<br />

merely one whose little pads have been carried in on who knows<br />

what animal’s pelt and which is at best just hanging on with us.<br />

As treacherous as it is, we are not sure whether we wish it luck<br />

or not.<br />

Engelmann received specimens from near Inscription Rock in<br />

northwestern New Mexico which he thought a different plant<br />

and named O. brachyarthra. Coulter made this a variety of the<br />

species, although saying that whether it should even stand as a<br />

distinct variety seemed doubtful to him. Britton and Rose considered<br />

it merely a synonym of the species. Boissevain, in his<br />

study of Colorado cacti, says that all of the specimens on the<br />

western slopes of the Continental Divide are more robust than<br />

those on the eastern side and that these western plants fit the<br />

description of Engelmann’s plant, so he calls the whole western<br />

population O. fragilis var. brachyarthra. But upon examining<br />

specimens from Inscription Rock, we do not find constant<br />

enough differences from the more eastern specimens, in our opinion,<br />

to justify the variety.<br />

Opuntia Sphaerocarpa Eng.<br />

Description Plate 57<br />

roots: Mostly tuberous, with small, oblong or spindle-shaped<br />

tubers on the fibrous roots, but sometimes lacking these and<br />

remaining entirely fibrous.<br />

stems: A low, diffuse, and mostly prostrate plant with only<br />

the new pads temporarily upright. The pads are round or even<br />

wider than they are long to sometimes very broadly egg-<br />

shaped, 21/2 to 4 inches long and 21/2 to 31/2 inches wide. These<br />

pads are average thickness to thick, tuberculate by raised<br />

areoles, Soft, shrinking to become very wrinkled when water<br />

is scarce or especially in the winter. They are bright green<br />

when young, fading to a somewhat lighter green when old,<br />

and often suffused with purplish coloring in the winter.<br />

areoles: Elongated in shape, tiny or very small, 1/8 of an inch<br />

or less long on the sides of the pads, becoming oval, but hard-<br />

ly larger on the edges. They are 1/4 to 3/4 of an inch apart.<br />

spines: Only upper edge areoles are armed. These have typically<br />

1 to 3 spines consisting of 1 central which is deflexed,<br />

1/4 to 11/2 inches long, slender to medium thickness, and flattened;<br />

sometimes 1 upper spine which is porrect, 1/2 to 11/2<br />

inches long, slender to medium thickness, and round; plus oc-


genus Opuntia 219<br />

casionally 1 or 2 lower, much deflexed, slender spines, 3/16 to<br />

1/2 inch long. The larger spines are brown or reddish-brown<br />

mottled above with tan or whitish. The lower, slender spines<br />

are gray with dark tips.<br />

glochids: Very few and short in a compact clump in each<br />

areole on the sides of the pads, becoming somewhat more<br />

numerous and longer to 3/16 of an inch on the edges. They are<br />

yellow to dirty straw-colored.<br />

flowers: Greenish-yellow, sometimes with some brownish<br />

coloring in their centers, 2 inches in diameter by 11/2 to 21/4<br />

inches tall. The ovary is very broad club-shaped, 3/4 to 11/2<br />

inches long, with some tan glochids on it. There are 7 or 8<br />

inner petals. The stamens are cream or yellowish. The style is<br />

long and greenish, with 5 fat, bright green lobes making up<br />

the stigma.<br />

fruits: Perfectly spherical to somewhat oval, 3/4 to 13/8 inches<br />

long and wide. The umbilicus is very broad and either flat or<br />

only very shallowly pitted. Having short white wool and<br />

many tan glochids up to 1/8 of an inch long in the areoles, but<br />

spineless. This fruit becomes dry, brown, and papery when<br />

ripe, but does this very slowly. It typically remains yellowish-<br />

green all summer and only dries the following winter. The<br />

seeds are just over 3/16 of an inch (5 millimeters) in diameter,<br />

irregular in shape, rather thick with narrow, acute rims.<br />

Range. The Sandia Mountains east of Albuquerque, New Mex-<br />

ico, through the Jicarilla Mountains west of Capitan, New<br />

Mexico, to the Sierra Blanca Mountains north of Ruidosa, New<br />

Mexico.<br />

Remarks. This seems to have been one of the most sadly misunderstood<br />

of all cacti, and as a result it is almost unknown today.<br />

Being impressed by the fact that it is supposed to have dry,<br />

spherical fruits, many have seized upon any plant from the Sandia<br />

Mountains with such fruits as representing this species. But<br />

they forget that O. polyacantha is specifically stated by the<br />

original description to sometimes have spherical fruits, and that<br />

these can sometimes be almost spineless. As a result I have seen<br />

numerous herbarium specimens of O. polyacantha with round<br />

fruits called O. sphaerocarpa. Taking this one step farther, there<br />

have even been attempts to call this a variety of O. polyacantha.<br />

Such a mistake should not have been made, in view of Engelmann’s<br />

very specific statement to forestall the confusion of these<br />

two. In his original description of O. sphaerocarpa he states that<br />

this form lacks the numerous small, bristle-like spines of O. polyacantha,<br />

which distinction will swiftly cull out the examples of<br />

O. polyacantha erroneously called this species. We have here a<br />

cactus with 1 to at most 4 spines, while O. polyacantha has al-<br />

ways a total of 5 to 18 spines per fully armed areole.<br />

The above error has so completely persuaded most students to<br />

regard O. sphaerocarpa as some sort of variation of O. polyacantha<br />

that Britton and Rose, for instance, describe it as light<br />

green in color like that other species. This has blinded most to the<br />

idea that there might be another cactus in the Sandias and nearby<br />

mountains having round, dry fruits, but being by comparison<br />

almost spineless and bright green when in healthy color. Add to<br />

this the lamentable tendency to call any bright green, prostrate,<br />

sparsely spined, tuberous-rooted cactus found anywhere O. com-<br />

pressa var. macrorhiza, and the true O. sphaerocarpa has remained<br />

unknown for most of the time since its original description.<br />

Nor is it any wonder that it is often mistaken for O. compressa<br />

var. macrorhiza, which is also found within its range.<br />

Stumbling upon a cactus in central New Mexico having prostrate,<br />

bright green, fleshy, tuberculate, and often wrinkled pads,<br />

1 to 4 spines more or less flattened on upper areoles only, and<br />

usually tuberous roots, almost anyone would call it that cactus.<br />

It keys out to that in any key I have seen that uses vegetative<br />

characters only. I called it that myself when I first collected it.<br />

But then I saw it bloom, and I got a distinct surprise, for it<br />

had too short an ovary, and standing there in plain sight were<br />

bright green stigma lobes! No O. compressa var. macrorhiza of<br />

the hundreds I had seen flower ever had other than yellowish<br />

or whitish stigmas. So I watched this cactus closely, and when<br />

the fruits formed they were as round as marbles or at least very<br />

broadly oval with no constriction below and with the umbilici<br />

very broad and flat. I now knew that this could not be variety<br />

macrorhiza, which always has extremely elongated, club-shaped,<br />

constricted fruits, with deeply pitted umbilici.<br />

What could this cactus be? All summer, as its fruits remained<br />

greenish way past the usual ripening time, I couldn’t place it,<br />

but when, in the following winter, they dried to become brittle<br />

balls, I first thought of it as a dry-fruited species, and once I<br />

did that, it fell into place perfectly as O. sphaerocarpa.<br />

Because this is apparently the first description of the flowers<br />

or the roots, and because I find no herbarium specimens since<br />

the type with the spineless, round fruits included, it is my opinion<br />

that any previously collected specimens, showing only vegetative<br />

parts, are buried in the O. compressa var. macrorhiza<br />

folders, from which they cannot be separated without reproductive<br />

parts. Nor do I think any collector will be able to tell<br />

when he has O. sphaerocarpa unless he is willing to wait in the<br />

New Mexico mountains until his specimens bloom and fruit, or<br />

unless he goes to the trouble to grow his specimens to these stages<br />

after he collects them.<br />

As far as relationships are concerned, this plant is a remarkable<br />

intermediate between the dry-fruited species and the fleshy-<br />

fruited O. compressa group. It has the green stigmas of the dry-<br />

fruited, but its fruits dry tardily. By character of pads, spines,<br />

roots, and seeds it is much more closely related to the O. com-<br />

pressa group than to O. polyacantha.<br />

I have collected this cactus from several locations in the Sandia<br />

Mountains, the type locality. I also have it from locations<br />

west of Capitan and from near Ruidosa, which enlarges its<br />

known range to include the mountains almost continuous with<br />

the Sandias on their southeast. It has been found only at com-


220 cacti of the southwest<br />

paratively high altitudes and usually in association with the<br />

forests of these mountains.<br />

Opuntia rhodantha Schumann<br />

“’Wide Cactus,” “Cliff Prickly Pear”<br />

Description Plate 58<br />

roots: Fibrous in all specimens seen.<br />

stems: Low-growing and spreading, only 6 inches to rarely<br />

18 inches high, with the pads upright at first, later reclining<br />

and rooting. These pads are almost circular, egg-shaped, or<br />

oblong, 2 to 9 inches long by 2 to 6 inches wide, 1/2 to 1 inch<br />

thick, and often slightly tuberculate. In color they are deep<br />

green or glaucous and gray-green.<br />

areoles: Oval and small to medium in size, being only about<br />

1/8 to 3/16 of an inch long. They are spaced 1/2 to 7/8 of an<br />

inch apart.<br />

spines: Usually 1 to 6 per areole in number, on only the upper<br />

one-half or less of the pad, except in one variety on which<br />

there may be up to 10 on most areoles. There are 1 to 4 main,<br />

spreading spines 3/4 to 2 inches long, medium to heavy, and<br />

somewhat to much flattened. These spines are whitish, yellowish,<br />

or variegated with brown. There are also 1 to 6 slender,<br />

round to slightly flattened, deflexed, whitish spines 1/4 to<br />

3/4 of an inch long.<br />

glochids: Brown in color, very few in one variety, quite<br />

numerous in another.<br />

flowers: Yellow or pink or reddish in color, and 2 to 3<br />

inches in diameter and 2 to 23/4 inches tall. The ovary is about<br />

11/2 inches long, tuberculate, with white wool and often some<br />

glochids, but with no spines on it. There are about 12 inner<br />

petals which are to 1 inch wide at the top. The filaments are<br />

greenish, yellowish, or reddish; the anthers cream. There are<br />

8 to 12 long, slender, deep green stigma lobes.<br />

fruits: Broadly club-shaped, tapering below, 11/2 to 13/4<br />

inches long. They usually have from a few areoles at the top<br />

to sometimes all areoles with clusters of short, whitish spines,<br />

but are occasionally spineless. They become dry and brittle<br />

when ripe. The seeds are between 3/16 and 1/4 of an inch (5–6<br />

millimeters), flat, with wide rims.<br />

Range. Southwestern Colorado and southern Utah into northern<br />

New Mexico and Arizona. In New Mexico found northwest<br />

of a line approximately from Taos to Albuquerque to Gallup.<br />

Remarks. This is perhaps the most beautiful of the small, dryfruited<br />

Opuntias. Although not very big, it is the largest of<br />

them, with thick, fleshy, deep-colored pads having not too many<br />

spines but those it has being very rigid and heavy. Its beauty,<br />

however, lies in its flowers. Although many specimens have clear<br />

yellow blooms, others have the only definitely pink blossoms<br />

I have seen in this genus. There are also specimens presenting<br />

shades intermediate between these colors, these often turning<br />

out to be salmon or reddish hues. The New Mexican plants tend<br />

to be mostly yellow, while in Colorado pink flowers are more<br />

commonly encountered.<br />

O. rhodantha is a cactus of the higher elevations, usually<br />

found at 5,000 or more feet above sea level. Within our area it<br />

is limited to the mountains of northwestern New Mexico. While<br />

it is immune to cold, I find that it does not grow well or bloom<br />

very successfully in the extremes of heat that most of our south-<br />

western cacti thrive in. In growing it, one should remember to<br />

provide all he can of the conditions of high altitudes.<br />

Opuntia rhodantha var. rhodantha (Schumann)<br />

Description<br />

roots: As the species.<br />

stems: As the species, except pads to only 7 inches long by 4<br />

inches wide.<br />

areoles: As the species, except to only 7/8 of an inch apart.<br />

spines: As the species, except to only 6 per areole, with the<br />

lower half or more of the areoles on each pad spineless, with<br />

the largest spines to only 15/8 inches long, and with only 1 to<br />

4 smaller, bristle-like spines per areole.<br />

glochids: Very few and inconspicuous.<br />

flowers: As the species.<br />

fruits: As the species.<br />

Range. As the species.<br />

Remarks. Originally, in describing this cactus, K. Schumann<br />

thought the red-flowered and the yellow-flowered specimens<br />

were different species, and he called them O. rhodantha and O.<br />

xanthostemma [sic] respectively. All students since then have<br />

realized that these two are one, and combined them under the<br />

name of O. rhodantha, with the exception of L. Benson, who has<br />

chosen to consider this a variety of the more westerly grizzly<br />

bear cactus, O. erinacea Eng., and so calls it O. erinacea var. xanthostema.<br />

This combination of our moderately spined cactus with<br />

the extremely spiny and hairy Mojave cactus does not seem too<br />

logical to us, and is not followed even by Earle in the most recent<br />

cactus book out of Arizona, so I continue with the original<br />

name for the plant.<br />

There have been numerous names given to various garden varieties<br />

of this cactus by horticulturists, particularly in Europe.<br />

Examples of some of them are variety pisciformis and variety<br />

schumanniana Spath; variety pallida, variety salmonea, variety<br />

rosea, variety rubra, variety gracilis, variety elegans, variety<br />

fulgens, variety orbicularis, variety brevispina, variety flavispina,<br />

etc. Hort. These refer to the various flower colors and<br />

other characters which have appeared in cultivation, and have


genus Opuntia 221<br />

no significance except to show the extent to which this cactus<br />

has been grown and the degree of its popularity among col-<br />

lectors.<br />

Opuntia rhodantha var. spinosior Boissevain<br />

Description Plate 58<br />

roots: As the species.<br />

stems: As the species, except pads averaging larger. When<br />

growing wild the pads are 4 to 8 inches long and typical in<br />

shape, but cultivated plants produced pads more spindleshaped<br />

and to 9 inches long by 6 inches wide, the plants then<br />

standing to 18 inches tall in the most robust cases.<br />

areoles: As the species, except a little larger and to 1 inch<br />

apart.<br />

spines: 1 to 10 on almost all areoles of each pad. ‘There are<br />

the same 3 or 4 main spines as on the species, except they are<br />

here up to 2 inches long. ‘There are 1 to 6 smaller, bristle-like<br />

spines below these.<br />

glochids: As the species, except quite numerous and sometimes<br />

to 1/4 of an inch long.<br />

flowers: As the species.<br />

fruits: As the species, except not so spiny. About half of the<br />

specimens observed had spineless fruits and the rest had spines<br />

on only the upper edges of the fruits.<br />

Range. From southwestern Colorado to near Albuquerque, New<br />

Mexico.<br />

Remarks. Boissevain and Davidson described this variety from<br />

the arid southwestern corner of Colorado. It has seldom been<br />

studied since, but in re-evaluating the Species in 1944, Croizat<br />

concluded that this was a “geographic well established form peculiar<br />

of the species in the southwestern Colorado desert,” and<br />

that it therefore deserves a position much higher than all of the<br />

garden varieties, which have been mentioned for the species. No<br />

one has reported it outside of Colorado until this time, however.<br />

We were very surprised to find this plant growing in the<br />

yard of friends who live in one of the new subdivisions across<br />

the Rio Grande northwest of downtown Albuquerque, New<br />

Mexico. They had a whole row of the plants grown very large<br />

and beautiful with the care they had given them. There were<br />

both yellow and pink-flowering individuals in the row, and<br />

when they put on their fruits, the identification was unmistakable.<br />

I visited the location where they had gathered their plants,<br />

only a few miles away on the flat, sandy mesa below the lava<br />

outcrops on the west and above the river valley a short distance<br />

to the east. The location is beside state road 448, in an area fast<br />

being developed, and the wild population is in real danger. I<br />

could not find the form growing anywhere else in the area. O.<br />

polyacantha, O. hystricina, and O. phaeacantha var. camanchica<br />

grew in profusion on the hills just to the west of this, but<br />

variety spinosior did not seem to be up there with them.<br />

On the basis of this collection, and after observing the variety<br />

growing in Colorado, I feel it necessary to include this variety<br />

as another New Mexico cactus. One would assume that the form<br />

must exist in similar situations between this point and south-<br />

western Colorado, and it will be interesting to see whether this<br />

supposition is borne out.<br />

I have included in my description of the variety the larger<br />

measurements which the cactus has achieved in the abovementioned<br />

garden. Although the specimens seen growing wild<br />

were somewhat smaller, they still averaged larger than the typical<br />

O. rhodantha, and since this garden was not over 4 miles<br />

or so from the collection point in the same soil and the larger<br />

growth could be the result only of some extra feeding and<br />

watering, I feel that the plant might attain the size in a favored<br />

natural situation also.<br />

The cactus is clearly very close to O. rhodantha, and should<br />

probably be left as a variety of this species. Other relationships<br />

have not been worked Out. I find that plants sent me from<br />

Arizona for O. nicholii are rather similar, and this similarity<br />

should be studied.<br />

Opuntia polyacantha Haw.<br />

“Hunger Cactus,” “Starvation Cactus”<br />

Description Plates 58, 59<br />

roots: Fibrous.<br />

stems: A prostrate, spreading cactus rarely rising over 6<br />

inches high, preferring to string out its joints along the<br />

ground, rooting on the edges of them, and thus often forming<br />

dense mats of growth. The pads are circular to oval, spindle-<br />

shaped, or broadly egg-shaped, but without any real constriction<br />

at the bases. They are average thickness to thick<br />

for their size, tuberculate by raised areoles, and also usually<br />

wrinkled. In color they are pale green or yellow-green, becoming<br />

reddish when suffering from extreme heat or cold.<br />

They vary from 11/2 to 5 inches long and from 11/2 to 4<br />

inches wide.<br />

areoles: Oval or elongated and small, 1/16 to 1/8 of an inch<br />

long on young pads, sometimes increasing in size on old pads<br />

and sometimes not. They contain much white wool when<br />

young. They are spaced 1/4 to 5/8 of an inch apart.<br />

spines: Very variable. On typical specimens there are 1 to 15<br />

spines present on all or almost all areoles. The main, interior<br />

spines may be missing or up to 5 in number. When present<br />

they consist of 1 main spine 1/4 to 2 inches long, very slender<br />

to medium in thickness, round to somewhat flattened, porrect


222 cacti of the southwest<br />

or deflexed; plus 0 to 2 upper spines 1/4 to 3 inches long, upright<br />

or spreading, slender to medium thickness, and round;<br />

and 0 to 2 laterals 1/4 to 11/4 inches long, slender, and round<br />

or slightly flattened. These spines may be white, yellowish,<br />

brownish, red-brown, or variegated. Besides this there are<br />

from 1 to at least 10 outer, radiating spines 1/8 to 3/4 of an<br />

inch long, slender to very slender and bristle-like, white or<br />

gray, often with darker tips. On some forms of the species the<br />

larger, main spines may be missing on all but the upper edge<br />

areoles, the spination of the pad thus being made up almost<br />

entirely of just the lower, bristle-like spines radiating downward.<br />

On the other hand, specimens are occasionally seen on<br />

which only the upper areoles have spines and these only 1 to<br />

3 of the main upper spines, with the lower bristles only 1 or<br />

2 in number and very short. Quite often the radial spines of<br />

old pads increase greatly in number, especially toward the<br />

bases of the pads, and become flexible, hairlike, and elon-<br />

gated to 2 to 8 inches.<br />

glochids: Few and short on young pads, being a very compact<br />

cluster only 1/16 of an inch or so long, but becoming<br />

sometimes to 3/16, of an inch long and fairly numerous on old<br />

pads. In color, yellow to bright brown.<br />

flowers: 2 to 31/2 inches in diameter, 11/2 to 21/2 inches tall.<br />

They are almost always—if not always—yellow in our area,<br />

but often pink, rose, or even reddish in color in Colorado and<br />

on north. The ovary is broad club-shaped to almost spherical,<br />

3/4 to 15/8 inches long and 3/4 of an inch or so wide. It is tuberculate<br />

by many elevated areoles containing much white<br />

wool and many yellow glochids plus some very slender spines<br />

sometimes up to 1/2 inch long. There are 5 to 10 bright green<br />

stigma lobes.<br />

fruits: Very variable. They are spherical to oval or eggshaped,<br />

more or less tuberculate when growing, but without<br />

tubercles when ripe and dry. They measure 3/4 to 2 inches<br />

long by 1/2 to 2 inches thick. The top is usually somewhat<br />

pitted, but may be entirely flat. When completely ripe the<br />

fruit becomes dry with a thin, papery skin over the tightly<br />

packed mass of seeds. On typical specimens there are 2 to 12<br />

slender spines 3/16 to 5/8 of an inch long on each areole of the<br />

upper third to three-fourths of the fruit, but fruits wholly<br />

spiny to wholly spineless may be found. The seeds are also<br />

extremely variable, ranging in size from 1/8 to more than 1/4<br />

of an inch (3–7 millimeters), being however over 3/16 of an<br />

inch (5–7 millimeters) in diameter on typical plants. The rim<br />

of the seed can be from narrow to very wide, this giving most<br />

of the variation in diameter which occurs.<br />

Range. The northwestern United States far into Canada, and<br />

south through western Nebraska and Kansas across the western<br />

tip of the Oklahoma Panhandle into the upper part of the<br />

Texas Panhandle, as well as through Colorado, over all of New<br />

Mexico, south into the Hueco and Davis mountains of southwestern<br />

Texas, and west into Arizona.<br />

remarks. This is one of the widest ranging of all cacti. It is<br />

probably the most northern cactus, growing practically up to<br />

the Arctic Circle, and followed north only by Opuntia fragilis.<br />

It is, however, not a conspicuous plant, perhaps surviving so<br />

successfully because it lies half buried in the sand or lost in the<br />

grass. The mats it forms effectively hold the soil from erosion<br />

and even stabilize the sand which drifts into them on the wind.<br />

As a result there is usually grass growing up through these carpets<br />

of cacti, and the very pale coloring of the pads plus the<br />

many spines which blend with the dead grass and trash caught<br />

in them render the plant so nearly invisible that it is much<br />

easier to walk into a stand of it before seeing it than it is to get<br />

out again. It is an aid to soil conservation by this holding action<br />

against erosion, but it is an ally few people can appreciate prop-<br />

erly.<br />

The cactus was apparently first referred to by Nuttall as<br />

Cactus ferox, but his description was so inadequate that authorities<br />

take as the first valid name Haworth’s Opuntia polyacantha.<br />

For a long period it was known by the name, O. missouriensis<br />

DC, but there is agreement now that this is merely<br />

a synonym.<br />

Having such a huge range and living in such different climates,<br />

this cactus shows many variations—witness the spines<br />

numbering from only a few to 15 and the color from white to<br />

red-brown, the fruit from spherical to egg-shaped and its size<br />

from 3/4 to 2 inches long, and most obvious of all, the seed size<br />

from about 1/8 to more than 1/4 of an inch in diameter. It has<br />

always been difficult to understand these differences. Engelmann<br />

found it necessary to list 5 varieties in the species, and<br />

Coulter increased that. More recently most of these have been<br />

ignored as mere growth forms, as for instance, variety rufispina<br />

Eng., and variety albispina Eng., based on spine color almost<br />

entirely, but some of them have, on the other hand, been elevated<br />

by various students to separate species, as O. trichophora<br />

(Eng.) B. & R. Then, some new species have been described, as<br />

for example O. juniperina Rose and O. schweriniana Schumann.<br />

Other students think these should hardly be carried even as<br />

separate varieties of the species.<br />

Here is a place where classical taxonomy based on discernible<br />

structural characters has as bad a time and gives us as little hope<br />

of constructing a logical order of relationships as anywhere.<br />

Here is where we need help. And right here is about the first place<br />

in the study of cacti where the results of modern biosystematics<br />

comes forward with some very limited but suggestive data for<br />

us.<br />

Chromosome counts have been made so far on only a few<br />

cacti, and most results published have been numbers from only<br />

single plants or plants from a single location. It happens, however,<br />

that chromosome counts of specimens of O. polyacantha<br />

from widely separate locations have been made and the results<br />

published by Stockwell. Here we find that there is variation in<br />

the chromosome number within the species. Where the typical<br />

2N chromosome number for cacti is 22, specimens of this cactus


genus Opuntia 223<br />

checked by Stockwell had either 44 or 66 chromosomes, with<br />

the specimens having 44 chromosomes appearing typical and<br />

those with 66 chromosomes being Canadian specimens which<br />

were smaller plants with smoother pads, fewer spines, and the<br />

small-sized seeds.<br />

What the modern technique seems to have given us thus far is<br />

a possible physical basis in the variation of chromosome numbers<br />

and a mechanism in the process of ploidy for some of the<br />

wide variation we have already seen in structural characters in<br />

the species. The only pattern we can see so far is that the chromosome<br />

number rises as we go north, which is as we would expect,<br />

ploidy often enhancing vigor and resistance to environmental<br />

extremes. All of the 66 chromosome counts reported<br />

were in the tiny-seeded Canadian form once described by Engelmann<br />

as O. missouriensis var. microsperma, which hardly<br />

enters the United States at all. This would seem to establish it as a<br />

definite variety based upon ploidy. But we are left with little<br />

aid in trying to logically separate the forms in our own area,<br />

since no variation in chromosome numbers among them has so<br />

far been shown. Therefore, we will leave the species as it stands,<br />

with our description wide enough to cover the usual forms, and<br />

deal individually with the most often separated of them.<br />

A list of the named varieties which we do not see as distinct<br />

and consistent enough to be regarded separately is included<br />

here in order to try to eliminate as much confusion as possible.<br />

While we have seen specimens which fit the description of each<br />

one of these, we have not been able to find them making up<br />

any separate populations in any definite geographical range.<br />

Always we have found them existing as the extremes within a<br />

population of varying individuals on the same hillside with<br />

typical individuals, and more often than not 2 or 3 different<br />

ones of these variations have been found side by side with the<br />

typical. This is how they differ from what we consider the defi-<br />

nite varieties of, for instance, O. engelmannii.<br />

Variety rufispina Eng. is only the typical plant as it sometimes<br />

appears with its main spines brown instead of the more<br />

common white.<br />

Variety albispina Eng. is the more common, pure white-spined<br />

form.<br />

Variety platycarpa (Eng.) Coult. is the form as it often appears<br />

at high altitudes in the northern New Mexico mountains<br />

and on into Colorado. One will often find plants fitting this<br />

description growing under the rather dense shade of junipers,<br />

while typical specimens are usually only a few feet away in the<br />

open. Nor is this form distinct for having the fruits globose with<br />

broad, flat umbilici, as some have supposed. In Taos Canyon<br />

and in several other high mountain locations in northern New<br />

Mexico, I have seen O. polyacantha fruits from almost club-<br />

shaped to perfectly spherical and from 3/4 to 2 inches long, as<br />

well as from very spiny to spineless, all on the same mountain-<br />

side. Many of them fitted Engelmann’s description of this supposed<br />

variety, but they were only part of the one large, greatly<br />

varying population. It was noted that all of those fruits over<br />

about 11/4 inches long were spherical or nearly so and spineless.<br />

Every one of these very large fruits which was opened was found<br />

to be sterile and apparently parasitized. We feel these exaggerated<br />

round fruits may be this species’ reaction to fruit parasites<br />

just as the extreme enlargement by elongation is the response of<br />

O. compressa var. fusco-atra to something similar on the Texas<br />

coast. These abnormal fruits we have heard called both variety<br />

platycarpa and O. sphaerocarpa, as we have those more normal<br />

and smaller fruits which happen to be round and nearly spineless.<br />

Many otherwise typical O. polyacantha specimens have<br />

normally round fruits, and to separate those with fewer than<br />

typical spines on the pads from a mixed population and call<br />

them another species or even another variety does not seem<br />

proper.<br />

Variety subinermis Eng. is yet another supposed variety.<br />

Plants fitting its description are occasionally found in the large,<br />

mixed populations of northern New Mexico. They have more<br />

elongated pads with areoles a little farther apart and only a<br />

very few short and slender spines on the upper edges. ‘These<br />

plants often make up part of the same large population, with 2<br />

or 3 others of these supposed varieties present as well as typical<br />

specimens. In each observed case, however, all of the specimens<br />

of this description were the result of environment, all of them<br />

growing far under junipers in deep accumulations of leaf mold.<br />

The elongated pads and poor armament are clear signs of etiolation.<br />

This form is very similar to O. schweriniana Schumann,<br />

which is not treated here because it is a Colorado form, but the<br />

relation between the two should be given careful study. I have<br />

seen a series of specimens from one location in Colorado which<br />

seemed to run all the way from fairly typical O. polyacantha<br />

through something like this form and on to something very close<br />

to O. schweriniana.<br />

O. juniperina Rose has been considered a separate species.<br />

Early in this study, on a trip through the northern mountains<br />

of New Mexico during the dead of winter, I hurriedly dug out<br />

of the snow some specimens which exactly fitted Rose’s description<br />

of this form. When I also saw similar specimens in the University<br />

of Colorado herbarium under this name, I was very sure<br />

of the species, and held out for it in some arguments with friends<br />

who said this was merely another form occurring in its type<br />

locality as part of the same sort of widely varying O. polyacantha<br />

population. More recently, when I studied the locality<br />

of my original collection of these specimens during the summer,<br />

I found the same thing there. There are typical O. polyacantha<br />

and all gradations between that and this supposed species wherever<br />

I have seen this form, including in its type locality at Cedar<br />

Hill, New Mexico. I therefore find it impossible to maintain it<br />

as even a consistent variety of the species.<br />

Variety trichophora (Eng.) Coult. has been one of the most<br />

persistent varieties. This is the form of the species in which the<br />

radial spines of at least the lower areoles and sometimes of all<br />

areoles on older pads elongate greatly and become flexible and<br />

hairlike. They often grow to 4 inches long, and Boissevain says


224 cacti of the southwest<br />

they sometimes reach 8 inches. At the same time they often increase<br />

in number to 25 or 30 per areole. Such a plant is surely<br />

a remarkable sight. This is naturally the most commonly noticed<br />

variant from the typical form of the species. Engelmann and<br />

Coulter called it a variety, Britton and Rose elevated it to a<br />

separate species, but most authors since have left it a variety.<br />

We started out considering it as separate from the typical<br />

form, but we soon ran into the same trouble as with the rest of<br />

these forms. We found all degrees of this hairiness, sometimes in<br />

the same population. Where could we draw the line in separating<br />

them? We have wearied at trying to be consistent about this,<br />

and tired of reading the contradictory explanations of the variety<br />

by various authors. The condition intergrades all the way<br />

with the typical forms which fail to elongate and proliferate the<br />

spines. In our northern New Mexico locations mentioned above<br />

we have found this hairy form growing only yards away from<br />

very good variety platycarpa, variety subinermis, and O. juniperina<br />

specimens having their less than typical armaments. In<br />

Taos Canyon I once collected a specimen fitting the description<br />

of O. juniperina in all pads except one, this one aberrant pad<br />

having 20 to 30 hairlike, flexible spines to 3 inches long in each<br />

areole of its surface. It was literally a pad of variety tricho-<br />

phora growing from an O. juniperina specimen. I grew this specimen<br />

carefully for 4 years, and was very disappointed to find<br />

the two pads which sprouted from the hairy pad developing<br />

into typical O. juniperina, sparsely spined pads in the greenhouse.<br />

So we have begun disappointing all those who constantly<br />

ask, “Which do I have, O. polyacantha or O. trichophora?” by<br />

refusing to separate them. In this we claim the support of Wooton,<br />

who said long ago that variety trichophora may not be dis-<br />

tinct from O. polyacantha.<br />

The species, O. polyacantha, blankets all of the state of New<br />

Mexico. There are records from within 50 miles of each of the<br />

four corners of that state. However, it does not go far east or<br />

south beyond that. In Oklahoma it has been recorded only from<br />

Cimarron County, the end county of the Panhandle, and rarely<br />

seen east of the Black Mesa, which is the extreme corner of that<br />

county. It enters Texas in two separate areas, coming into the<br />

Texas Panhandle where it seems to grow only as far as the south<br />

breaks of the Canadian River. Not being seen in all of the Texas<br />

high plains south of that, it enters the state again from southeastern<br />

New Mexico along the Hueco and Guadalupe mountains,<br />

growing as far south as the Davis Mountains, but appar-<br />

ently no farther into the lower Big Bend. It is definitely a north-<br />

ern cactus.<br />

Opuntia hystricina Eng.<br />

“Porcupine Prickly Pear”<br />

Description Plate 59<br />

roots: Fibrous.<br />

stems: Low, diffuse, and spreading, but not prostrate. They<br />

form small clumps of upright pads standing 6 inches to sometimes<br />

1 foot high. The pads are nearly circular to more often<br />

elongated obovate or even oblong, 3 to 5 inches long and 21/2<br />

to 3 inches wide, medium thickness, not tuberculate, and not<br />

becoming noticeably wrinkled from lack of water. They are<br />

medium or bright green in color.<br />

areoles: Large and conspicuous, oval, and about 3/16 of an<br />

inch long on young pads, becoming round and 1/4 of an inch<br />

or more in diameter on old pads. They are situated 1/4 to 1/2<br />

inch apart.<br />

spines: A total of 6 to 15 spines in each areole of the pad.<br />

There are in each areole 1 to 8, and in fully armed upper<br />

areoles always 5 to 8, main spines which are 11/2 to 4 inches<br />

long. These spines are irregularly arranged and spread in all<br />

directions, including upward. They are brownish or gray or<br />

variegated in color. They are of slender to average thickness,<br />

somewhat flattened, often twisted or bent, and somewhat<br />

flexible. There are also up to 7 lower, very slender, white,<br />

radiating spines 3/4 to 1 inch long in each areole. There is<br />

sometimes a tendency for a few very slender, white, hairlike<br />

spines 2 to 3 inches long to grow at the bases of the pads.<br />

glochids: Straw-colored or brown, often both in the same<br />

areole due to the existence of an outer ring of older, longer,<br />

darkened glochids and an inner cluster of younger, shorter,<br />

brighter, straw-colored ones.<br />

flowers: Ordinarily clear, pale yellow, but said to sometimes<br />

have various hues such as orange, rose, red, or even purplish.<br />

They are 2 to 3 inches in diameter. The ovary has white<br />

wool and bristle-like spines. The stigma lobes number 8 to 10<br />

and are green in color.<br />

fruits: Egg-shaped to broadly club-shaped and 7/8 to 11/4<br />

inches long by about 1/2 inch thick. The areoles of the upper<br />

half of the fruit possess 4 to 12 bristly spines 1/2 to 3/4 of an<br />

inch long per areole. The umbilicus is flat, or nearly so. The<br />

seeds are very large, between 1/4 and 5/16 of an inch (7 millimeters<br />

or more) in diameter, with broad rims.<br />

range. From the Rio Grande in New Mexico west to Nevada<br />

and California.<br />

remarks. Over 100 years ago Engelmann felt it necessary to<br />

distinguish a cactus closely allied to O. polyacantha from that<br />

species, calling it O. hystricina. He admitted that he himself had<br />

difficulty in keeping the two separate, and even conjectured<br />

that one might be only a form of the other. Everyone since then<br />

has had trouble with these two, but no one has actually combined<br />

them. When a combination involving this cactus did<br />

finally come, it was Benson’s attempt to combine it instead with<br />

erinaceae -><br />

an entirely different California cactus, O. erinacea Eng., giving erinacea<br />

us O. erinacea var. hystricina (Eng.) Benson.<br />

Let us see first how we distinguish O. hystricina from O. polyacantha<br />

and what characters keep it from being united with that<br />

cactus. We find that it differs from O. polyacantha as follows:


genus Opuntia 225<br />

it is diffuse and spreading with more upright growth than that<br />

cactus; its pads are darker green; it lacks distinct tubercles by<br />

raised areoles or marked wrinkling; it has more and longer and<br />

more erect main spines, which are also usually bent and somewhat<br />

flexible; and it has many more and longer glochids. I<br />

should state emphatically that it cannot be told from O. polyacantha<br />

by having flattened spines, as some have argued, since<br />

some specimens of that cactus have more definitely flattened<br />

spines than this does; nor by having more stout spines, since<br />

specimens could be matched with this character just reversed;<br />

nor larger pads; nor more widely spaced areoles. It is not just<br />

a robust version of the hunger cactus, as some articles about it<br />

would lead one to believe. Its whole manner of growth is different.<br />

But in many collections, both herbarium collections and<br />

fanciers’ gardens, I see smaller specimens of O. polyacantha<br />

labeled correctly, but plants of that species merely more robust<br />

in size or spines wrongly called O. hystricina. This has made the<br />

tracing of this species very difficult.<br />

Part of the trouble may be that although Engelmann says correctly<br />

that it grows all the way east to the Rio Grande, it is not<br />

at all common in New Mexico. It should be noticed that his<br />

descriptions were all compiled from specimens of Arizona and<br />

Nevada, and the only examples he actually mentions having<br />

seen from New Mexico were Fendler’s specimens from near<br />

Santa Fe—these Engelmann says, “seem to be intermediate [between<br />

O. hystricina and O. polyacantha] and may make it necessary<br />

to combine them.” These may have actually been only<br />

robust O. polyacantha specimens. If so, Engelmann himself<br />

would have been the first to make this often repeated mistake.<br />

It is interesting that Coulter found no New Mexico records of<br />

O. hystricina as late as 1896, and that in 1915, Wooton and<br />

Standley, those very complete students of New Mexico plants,<br />

said, “We have seen only one doubtful specimen [of O. hystridna].<br />

It is reported here… on the authority of the first collector,<br />

Dr. Bigelow.” So we should get over the idea which I find<br />

in some collectors that every New Mexico hill west of the Rio<br />

Grande can be expected to have this cactus on it, and stop call-<br />

ing all of our big specimens of O. polyacantha this.<br />

This is not to say that it does not exist in New Mexico, but<br />

I believe it is rare anywhere in that state. I have seen only two<br />

definite specimens from New Mexico, and the one which I have<br />

growing before me is from only a short distance west of Las<br />

Cruces, so far southeast that it confirms nicely the fact that, although<br />

it is rare, it must range the whole state west of the Rio<br />

Grande.<br />

These plants also show clearly why Benson combined this<br />

form with the more western form he calls O. erinacea. They are<br />

upright in growth, with ovate or oblong pads, having spreading,<br />

often erect spines up to 4 inches long and only moderately rigid,<br />

and with many conspicuous glochids. All of these are characters in<br />

common with the more western plant. In fact, it is easy to agree<br />

with Backeberg, who maintains that the two are actually the<br />

same. But Backeberg also points out that Engelmann himself<br />

withdrew his name, O. erinacea, and so it would not be technically<br />

correct to combine anything under that name. We therefore<br />

follow him in leaving the name of our rare New Mexico<br />

plant O. hystricina. Whether it is the same as the California<br />

plant is outside the scope of our study. It is the same plant that<br />

grows at least to the Colorado River and Nevada, and it is<br />

more closely related to the equally upright O. ursina Weber, the<br />

extremely hairy grizzly bear cactus of the Mojave, than it is to<br />

O. polyacantha.<br />

Opuntia arenaria Eng.<br />

Description Plate 59<br />

roots: Unique among Opuntias in our area. The roots are<br />

not tuberous, but have large, rhizome-like structures up to 1/2<br />

inch thick which sometimes run 3 to 6 feet horizontally just<br />

under the surface of the almost pure sand in which this plant<br />

grows. These underground structures are covered with many<br />

large areoles and long glochids like old stems, and they give<br />

off pads and fibrous roots all along them.<br />

stems: If these underground structures are considered as rhizomes<br />

or stem structures, the main stems of the plant are prostrate<br />

strings up to 6 feet long of old pads which are buried in<br />

the drifting sand. However, if these underground structures<br />

are made up from old pads, the separate joints are not clearly<br />

observable; therefore some students have interpreted them as<br />

roots. But the presence of areoles and glochids upon them<br />

makes it difficult for us to consider them roots. From these<br />

structures there are ascending and spreading young pads which<br />

stand 6 inches or rarely a little higher. The pads are 11/4 to 4<br />

inches long by 3/4 to 2 inches wide, and often are 3/4 of an<br />

inch thick. When small these pads are almost cylindrical, but<br />

as they grow larger they widen and flatten to become elongated<br />

oval to very elongated egg-shaped or even clavate.<br />

Young pads are light, shiny green in color, tuberculate, but<br />

not becoming wrinkled when dehydrated. Old pads, on the<br />

prostrate, more or less covered stems become brown and<br />

woody.<br />

areoles: Small, with a little white wool when young, enlarging<br />

somewhat on old stems. They are spaced 3/16 to 3/8 of<br />

an inch apart.<br />

spines: Almost all areoles are armed with 3 to 10 spines white<br />

or brownish or white with brownish bases and tips. There are<br />

1 to 4 main spines. The central one of these is 3/4 to 2 inches<br />

long, rigid, of average thickness, flattened slightly, and standing<br />

porrect or turned upward. There are besides this 0 to 3<br />

others 3/8 to 1 inch long, round or slightly flattened, and<br />

spreading downward. Below these there are 2 to 6 bristle-like<br />

spines 1/8 to 1/2 inch long, radiating downward.<br />

glochids: Straw or brownish in color, few in number but


226 cacti of the southwest<br />

rather long on young pads, becoming very many and to 1/4 of<br />

an inch long in large, spreading clusters which almost completely<br />

cover old pads.<br />

flowers: Yellow, 2 to 23/4 inches in diameter. The ovary is<br />

egg-shaped and 7/8 to 11/4 inches long. There are about 8 inner<br />

petals and 5 green stigma lobes.<br />

fruits: Oblong or club-shaped, usually constricted at both<br />

the bottom and the top, and deeply pitted above. They are 1<br />

to 11/2 inches long by 1/4 to 3/8 of an inch thick. They have 1<br />

to 5 slender spines 1/4 to 1/2 inch long on each areole of the<br />

broader central zone. The seeds are from nearly 3/16 to almost<br />

5/16 of an inch (5–7 millimeters) in diameter, very thin, irreg-<br />

ular, and rather elongated in shape, with broad margins.<br />

Range. Known only from sandy areas along the Rio Grande<br />

just a few miles above El Paso, Texas. Those on the east side of<br />

the river would be in Texas and those on the west side in New<br />

Mexico.<br />

Remarks. This is a unique little cactus equipped with the extremely<br />

long, stolon-like root or stem structure of a plant adapted<br />

to survive in deep and shifting sand. And it is found only in<br />

such sand, where often three-fourths of the plant is entirely<br />

buried, with only the current year’s new pads standing upright<br />

out of the ground.<br />

This is an almost vanished species. Engelmann said it grew in<br />

the sandy bottoms of the Rio Grande near El Paso. Most of<br />

these have long since had buildings or irrigated farming upon<br />

them. Dr. Rose said he found the plant about 8 miles above El<br />

Paso on the New Mexico side of the river. I had spent some time<br />

slogging through the sand over there to no avail, and would no<br />

doubt never have seen the cactus but for Mr. Clark Champie,<br />

well-known cactus grower and dealer of the area. When I told<br />

him of my problem, I told the right person, for he has a definite<br />

corner on this cactus. He immediately took me to one of the<br />

two known living stands just a few hundred yards from his own<br />

cactus farm on the outskirts of Anthony, east of the river at the<br />

Texas-New Mexico border. The stand is only a few dozen yards<br />

in extent, and a road is now bull-dozed through the center of it,<br />

so let us hope Mr. Champie is a good custodian of the cactus, or<br />

it may be completely extinct very soon.<br />

Coulter’s opinion was that O. arenaria was allied to O. fragilis,<br />

because of the almost cylindrical form of its early joints, but<br />

O. arenaria’s pads are tightly joined to each other instead of<br />

loose, and there are other differences.<br />

Britton and Rose say that its flowers are red, which is definitely<br />

not true of the stand near Anthony. The blooms are very<br />

yellow there. However, Britton and Rose had plants from another<br />

location on the west side of the river, and knowing the<br />

tendency to variability of flower color in this whole group, we<br />

must admit that their plants might have bloomed with a differ-<br />

ent color.<br />

In cultivation this cactus does not usually do well. It will not<br />

grow successfully in regular soil, and it is hard to provide a big<br />

enough and deep enough plot of sand containing the proper elements<br />

so that it can spread out and feel at home. While I have<br />

succeeded in keeping it alive for 4 years now, and in getting<br />

some growth, it has never flowered for me. So far as I know, to<br />

see it growing successfully one must make a pilgrimage to Mr.<br />

Champie’s natural stand. We wish such a rare stand as this could<br />

have some sort of official protection.<br />

Opuntia grahamii Eng.<br />

“Mounded Dwarf Cholla”<br />

Description Plate 59<br />

stems: Joints elongated oblong, club-shaped, or sometimes<br />

spindle-shaped, 11/2 to 21/2 inches long, covered with low, oblong<br />

tubercles 1/2 inch or slightly more in length. Sometimes<br />

these tubercles are very indistinct. These joints form very<br />

dense mats up to about a foot across, those on the edges often<br />

lying prostrate, but those in the center ascending. The plant<br />

may round up into quite a mound, but this is due to sand<br />

blowing into it. The joints only branch once above the surface,<br />

and, so stand to only 4 inches or so above the accumulated<br />

level of the sand. The joints are firmly attached.<br />

areoles: Round, 1/8 to 3/16 of an inch across, with much white<br />

wool in them at first. They are located on the ends of the<br />

tubercles, and are about 1/2 inch apart.<br />

spines: 8 to 14 per areole. There are 4 to 8 main inner spines<br />

spreading in all directions. They are 11/4 to 21/2 inches long,<br />

straight, medium in thickness, round or nearly so, brown or<br />

red-brown and smooth at first, becoming grayish and rough<br />

when old. They are not cross-striated or edged. Besides these<br />

there are 4 to 6 Outer spines spreading below, slender, round,<br />

whitish, 1/2 to 1 inch long.<br />

glochids: Few at first, becoming quite numerous and to 1/4<br />

of an inch long on old joints. They are brown in color.<br />

flowers: Yellow, 2 to 21/2 inches across. The stigmas are<br />

whitish.<br />

fruits: Elongated egg-shaped or oblong, 11/4 to 1/4 inches<br />

long, and yellow when ripe. They are tuberculate, with many<br />

areoles, each having white wool and several slender white<br />

spines, as well as 20 to 30 white glochids. The seeds are slightly<br />

over 3/16 of an inch (5–51/2 millimeters) in diameter.<br />

Range. As far as is definitely known, a rather small area of<br />

west Texas consisting of parts of El Paso and Hudspeth counties<br />

from the foothills of the Franklin Mountains through the Hueco<br />

Mountains and hills east and southeast of El Paso to near Sierra<br />

Blanca, Texas. It is known to grow in adjacent Chihuahua and<br />

is assumed to enter slightly into adjacent New Mexico, but I<br />

have not been able to confirm this. It is reported by Anthony<br />

from the Texas Big Bend, but this we also failed to confirm.<br />

Remarks. This species introduces us to the cylindrical-stemmed


Cornyopuntia -><br />

Corynopuntia<br />

genus Opuntia 227<br />

Opuntias, sometimes called Cylindropuntia in technical language,<br />

and also almost universally known by the common name<br />

of chollas. It is also one of the low-growing chollas whose stems<br />

are more or less club-shaped, and so it is part of the group<br />

sometimes separated out of the Cylindropuntia as Clavat-<br />

opuntia or Corynopuntia.<br />

O. grahamii is an interesting small cactus of the sand hills<br />

east of El Paso, Texas. It is very common on some of those<br />

hills, particularly near the Hueco Mountains. It grows more<br />

compactly and with its joints more ascending than those of any<br />

of its relatives except O. clavata. A typical specimen usually<br />

forms a little mound 3 to 6 inches high and up to about a foot or<br />

so in diameter. But the sand drifts with the wind into this compact<br />

mass, and soon covers the lower stems. As this happens,<br />

new ones grow above, and although no joint is over 4 inches or<br />

so above the surface, the accumulation often results in a mound<br />

a number of inches high. Covered as it is by its very many long,<br />

rather slender spines, the appearance from a distance is exactly<br />

that of a clump of short grass partly filled with sand, and the<br />

cactus is very hard to see.<br />

O. grahamii can be told from all of its close relatives by the<br />

fact that its main spines are comparatively slender, smooth and<br />

round, or practically so, while all of the others have their main<br />

spines very heavy, very rough, cross-striated, and greatly flattened.<br />

Noticing this difference will enable anyone to distinguish<br />

it from O. schottii, with which it is most often confused.<br />

Opuntia schottii Eng.<br />

“Devil Cactus,” “Dog Cholla,” “Clavellina”<br />

Description Plate 60<br />

stems: Joints elongated, cylindrical, or more often clubshaped,<br />

lying prostrate or sometimes with the upper ends<br />

turned upward. The bases of these joints are constricted and<br />

around 1/4 of an inch thick, from which they gradually broaden<br />

to almost 1 inch thick at the upper end. The over-all<br />

length of a joint is 1 to 3 inches. The joint is covered by<br />

broad, elongated tubercles about 1/2 to 3/4 of an inch long.<br />

These joints are rather loosely attached, coming off the plant<br />

with only a small tug.<br />

areoles: Round, or nearly so, located on the upper ends of<br />

the tubercles, small below to over 1/4 of an inch in diameter<br />

toward the end of the joint.<br />

spines: There are 8 to 14 spines per areole. Of these, 1 main<br />

central, which is very heavy and very flat, stands porrect or<br />

a little deflexed. It is 1 to 21/2 inches long and 1/16 to 1/8 of<br />

an inch wide for most of its length. Its surface is very rough<br />

and usually distinctly cross-striated. In color it is at first<br />

straw to light brown with the tip translucent yellow and<br />

edged with light yellow or whitish the length of the spine. It<br />

fades to rough gray when old. There are two lower centrals<br />

spreading downward, to 11/2 inches long and as the main central<br />

in character. There are 1 to 4 centrals spreading upward,<br />

1/4 to 11/2 inches long, heavy but triangular or completely<br />

round, darker brown or sometimes red-brown in color with<br />

the tips translucent yellow. Besides these there are 5 to 7<br />

small, slender, round, whitish radial spines 1/4 to 3/4 of an inch<br />

long spreading below. There are no sheaths on matured spines<br />

of this plant.<br />

glochids: Few. In the upper areoles there will be up to a<br />

dozen straw-colored bristles to 3/16 of an inch long at the top<br />

of the areole.<br />

flowers: Yellow, about 2 to 21/2 inches across and 2 inches<br />

tall. The stamens are whitish. The stigmas are whitish or very<br />

pale greenish.<br />

fruits: 1 to 11/2 inches long by 3/8 to 1/2 inch thick, elongated<br />

club-shaped, with perianth persistent upon it, and very light<br />

yellow when ripe. It is covered with narrow tubercles 1/4 to<br />

3/8 of an inch long, each ending in an areole above. These<br />

areoles are small, each containing white wool, 4 main central<br />

spines which are slender, round, 3/16 to 5/16 of an inch long,<br />

and arranged cross-wise, plus 25 to 35 radial spines which<br />

are very slender and bristle-like. The seeds are not quite 3/16<br />

of an inch (about 4 millimeters) in diameter.<br />

Range. Along the Rio Grande from near Brownsville, Texas,<br />

to Lajitas in the Big Bend. Becoming common from the area of<br />

Zapata, Texas, to the Big Bend, but never seen over 10 miles or<br />

so north of the river.<br />

Remarks. This is a very low, inconspicuous little cactus growing<br />

practically invisibly in the grass, but if stepped upon, it can become<br />

very painfully evident. The spines are strong enough to<br />

penetrate shoe leather sometimes, and being very barbed, they<br />

stick to shoes or clothing. The next step often pulls joints loose,<br />

and one has unwanted riders which it is a major chore to disengage<br />

without injury to oneself.<br />

O. schottii grows on the gravelly hillsides overlooking the Rio<br />

Grande and the last few miles of its tributaries such as the<br />

Devil’s River and the Pecos. Below the Pecos the plant usually<br />

grows as single chains of prostrate joints blooming and fruiting<br />

sparingly and apparently propagated mostly by separated pads.<br />

These can be pulled apart rather easily, but are not nearly so<br />

loosely attached as those of O. fragilis or O. drummondii. From<br />

the Pecos west the cactus grows more often in low, compact<br />

mats up to 2 feet or so across, and in season these are covered<br />

with very many flowers and fruits.<br />

O. schottii is a Texas cactus, growing along an immense length<br />

of the Rio Grande, but never leaving the hills overlooking the<br />

river. Its westernmost range does not seem to quite reach the<br />

range of its nearest relatives, but it apparently has grown, at<br />

least in the past, almost to the mouth of the Rio Grande. Runyon’s<br />

record of it, “near Brownsville,” attests to that. We have


228 cacti of the southwest<br />

not been able to find it below Zapata, and surmise that the great<br />

development of the area below there may have reduced its range<br />

considerably.<br />

Opuntia stanlyi Eng.<br />

“Stanley’s Cholla,” “Devil Cholla,” “Creeping Cholla”<br />

Description Plate 60<br />

stems: Consisting of cylindrical, club-shaped joints, 31/2 to 6<br />

inches long, slender at the base, but enlarging to from 1 to 2<br />

inches in diameter at the outer end. The plant consists of many<br />

of these joints lying prostrate or with the ends curving upward,<br />

or sometimes with the whole joint ascending, thus<br />

forming dense mats standing from 6 to sometimes 12 inches<br />

high and extending from several feet to sometimes 15 or 20<br />

feet in diameter. The joints are covered with tubercles appearing<br />

as ridges 1 to 13/4 inches long by 1/4 to 3/4 of an inch<br />

tall by 1/4 to 1/2 inch wide.<br />

areoles: Round, or nearly so, located on the upper slopes of<br />

the tubercles, 1/4 of an inch or slightly more in diameter, with<br />

some white wool at first.<br />

spines: 10 to at least 20 in number. There are 5 to 9 large inner<br />

spines 11/8 to 23/8 inches long, spreading, with the largest<br />

one porrect or deflexed, very flat, and 1/16 to sometimes 1/8<br />

of an inch wide, with the upper surface roughened by tiny<br />

pits and irregular cross-striation. The other main spines are<br />

not so heavy nor so flattened. All these are yellowish or reddish-brown<br />

at first, becoming ashy gray when old. There are<br />

also 7 to at least 15 outer spines 3/8 to 3/4 of an inch long,<br />

whitish, round to flattened.<br />

glochids: Few in number, but to 1/4 of an inch long and ro-<br />

bust. They are yellow in color.<br />

flowers: Pale yellow, 11/2 to 23/4 inches in diameter. The<br />

stigma lobes are white.<br />

fruits: Broadly club-shaped or egg-shaped, 2 to 23/4 inches<br />

long, and very spiny. The seeds are about 3/16 of an inch<br />

(about 5 millimeters) in diameter.<br />

Range. Entering our area from Arizona and Mexico, and ranging<br />

over southwestern New Mexico at least as far as the upper<br />

edge of Grant County, and east to Socorro and near El Paso. It<br />

is found in Texas along the Rio Grande from near El Paso to<br />

near Candelaria in Presidio County.<br />

Remarks. This is one of the worst cactus pests in existence, earning<br />

the common name, devil cholla. Fortunately it is not common<br />

in our area, but where it is met with it often renders whole<br />

hillsides impassible not only to men but even to goats, by covering<br />

many square feet with thickets only a foot or less high but<br />

so compact as to be impenetrable by all except small creeping<br />

things. Such stands are to be seen in western Grant County,<br />

New Mexico, and along the Rio Grande in western Presidio<br />

County in Texas. Other than in these two locations in our area,<br />

we are not at all sorry to Say, we encounter only isolated small<br />

stands of the cactus in widely scattered places. These locations<br />

are so scattered that it is hard to delimit the eastern edge of its<br />

range in New Mexico. The most northeastern record of it is an<br />

old one of Vasey’s given as Socorro, New Mexico. There are old<br />

records from near El Paso as well, but the plant does not seem<br />

to have been collected nearly so far east in recent times.<br />

O. stanlyi has had several other similar forms attached to it<br />

as varieties, but these occur only in Arizona, California, and<br />

Nevada. The cactus as we find it in our area is the typical form.<br />

It is most closely related to O. schottii. The main characters of<br />

the two are almost all the same, O. schottii presenting most of<br />

the features of the other in miniature. However, the joints of<br />

O. stanlyi are not easily separated, as are those of the smaller<br />

cactus. The ranges of the two are separated by only a short distance<br />

along the Rio Grande in Texas, and the Candelaria population<br />

of O. stanlyi has individuals averaging a little smaller<br />

than those farther west, but there is no actual intergrading in<br />

size between the two. Hester, who studied them both in the Big<br />

Bend, did not consider any combination of them, but a fair case<br />

could be made perhaps for considering one a variety of the<br />

other.<br />

Engelmann apparently described this cactus under two names,<br />

first as O. stanlyi, and then later as O. emoryi. The two names<br />

are, therefore, considered synonyms.<br />

Opuntia clavata Eng.<br />

“Club Cholla,” “Dagger Cholla”<br />

Description Plate 60<br />

stems: Each plant forming a low mat often 3 to at least 6<br />

feet across but only 3 to 6 inches high. The joints are upright<br />

or nearly so, short club-shaped to egg-shaped, with some narrowing<br />

below but not markedly constricted below, 1 to 21/2<br />

inches long and to 1 inch thick above. These joints are covered<br />

with low, broad, rather indistinct tubercles about 1/4 to 1/2<br />

inch long.<br />

areoles: Large, usually 3/16 to 1/4 of an inch across, and<br />

round.<br />

spines: There are 10 to 20 spines, all white and rough when<br />

mature, but bright pink when growing. 4 to 7 of these are<br />

main inner spines 1/2 to 11/4 inches long. The upper ones of<br />

these are more slender, shorter, only somewhat flattened, and<br />

stand erect. The lower 3 or 4 of them are spreading downward,<br />

larger, and flatter. The main central spine is deflexed<br />

and conspicuous for its flatness and thickness, to only 11/4<br />

inches long, but about 1/8 of an inch thick at its base and<br />

tapering evenly to the point. The upper surface of this spine<br />

is conspicuously cross-striated and the lower surface is keeled.


genus Opuntia 229<br />

There are also 6 to 13 outer spines radiating in every direction.<br />

They are only 3/16 to 5/8 of an inch long, slender, and<br />

round.<br />

glochids: Few to rather many, white or straw-colored, about<br />

1/8 to 3/16 of an inch long.<br />

flowers: Yellow and small, to only 2 inches in diameter and<br />

even less in length.<br />

fruits: Elongated club-shaped or almost spindle-shaped, with<br />

a deep umbilicus. They are 11/4 to 2 inches long and to 1<br />

inch thick. The light yellow surface of the ripe fruit is almost<br />

completely covered with a large number of very slender,<br />

white or straw-colored bristles which grow from the many<br />

areoles. The seeds are slightly over 3/16 to 1/4 of an inch (5–6<br />

millimeters) in diameter.<br />

Range. Central and a portion of northwestern New Mexico,<br />

sometimes said to extend into Arizona. From Tularosa and near<br />

Truth-or-Consequences, New Mexico, on the south to near Las<br />

Vegas on the northeast, to El Rito on the north, then back<br />

southwest to near Cubero. Said to enter Arizona north of the<br />

Zuni River, but I find no New Mexico record of it west of a<br />

line from El Rito to Cubero to Socorro to Truth-or-Consequences.<br />

Remarks. On all sides of Albuquerque, New Mexico, one can<br />

find large areas of almost barren flats and slopes where the<br />

short grass alternates with the often huge mats of O. clavata.<br />

At first glance the cactus appears much like grass, but one had<br />

better learn to distinguish the one from the other before he<br />

walks in the area. While some point out that it is a great pest<br />

to livestock, an unprejudiced observation reveals that it doesn’t<br />

often compete with the grass, but usually occupies otherwise<br />

barren ground. The soil is usually bare and eroded away on all<br />

sides of these thick mats of cactus, leaving the stand elevated<br />

several inches on the soil it has held and showing clearly how<br />

valuable a soil holder it is where little else seems able to do the<br />

job. It is thus another “pest” which we should consider carefully<br />

before condemning entirely, and we should know very<br />

certainly what can be introduced to hold the soil in its place<br />

before the herbicide gunners are turned loose on it.<br />

The cactus is well named the dagger cholla. The name comes<br />

from the shape and appearance of the largest spine, which is so<br />

short, but so broad and flattened that it does resemble a tiny<br />

dagger. The other common name sometimes applied to it, the<br />

club cholla, is probably not so apt, since the joints are at most<br />

only very broadly club-shaped without much constriction at the<br />

base and usually better described as egg-shaped. Although they<br />

stand upright, they rarely give off new joints except near their<br />

bases, and so the plant is seldom over 3 or 4 inches tall.<br />

This is a cactus of central New Mexico. While it ranges most<br />

of the length of that state, it does not reach either the north or<br />

south border, and there are no known records of it near either<br />

the eastern or western boundaries. Records of it place it in a<br />

strip only up to 100 or so miles wide. If it does in fact enter<br />

Arizona, as has been said, it must be very rare in the intervening<br />

western part of New Mexico.<br />

This cactus is most closely related to O. stanlyi var. parishii<br />

Benson, which, however, does not seem to come east of western<br />

Arizona.<br />

Opuntia imbricata var. arborescens (Eng.)<br />

“Tree Cactus,” “Cane Cactus,” “Candelabrum Cactus,”<br />

“Cholla,” “Velas de Coyote (Coyote Candles)”<br />

Description Plate 61<br />

stems: Growing as an upright, bushy, or treelike plant, 3 to<br />

at least 8 feet tall. There is a round trunk quite often becoming<br />

up to 3 or 4 inches in diameter and said to have been<br />

seen 5 to 10 inches thick. This trunk is more or less covered<br />

with rough bark and enlarged areoles. The current year’s<br />

joints are cylindrical, usually 2 to 6 inches, but occasionally<br />

to 8 inches long, and to about 1 inch in diameter. They are<br />

covered with elongated tubercles 3/4 to 11/4 inches long and<br />

to about 1/4 of an inch high, their upper and lower slopes<br />

about equal. The old stems have a woody skeleton.<br />

areoles: Oval or oblong, about 1/4 of an inch long, enlarging<br />

somewhat with age. They possess some short, whitish, or yellowish<br />

wool.<br />

spines: Very variable in number, from 2 to 10 on the current<br />

year’s growth, increasing with age to 20 or 30. There are 1 to<br />

8 central spines spreading in all directions. These are 1/2 to 11/4<br />

inches in length, slender to medium strength, round, yellow,<br />

brownish, or variegated in color. They are covered by loose<br />

sheaths. The remainder of the spines are radiating exterior<br />

spines, 1/4 to 5/8 of an inch long, slender, white to brownish,<br />

at least partly covered with tight, whitish sheaths.<br />

glochids: Very few, sometimes apparently missing. Those<br />

present are found in the upper part of the areole and are so<br />

short as to be inconspicuous in the wool.<br />

flowers: To 3 inches in diameter, 11/2 to 2 inches tall, purplish,<br />

lavender, or rose-pink in color. The ovary is 1 inch long<br />

or slightly more, and nearly as thick. It is very tuberculate<br />

and often has perianth segments and 2 or 3 long, white bristles<br />

1/4 to 5/8 of an inch long on its upper areoles. The fila-<br />

ments are reddish toward their bases, becoming greenish-pink<br />

above. The anthers are cream-colored. The style is reddish and<br />

slightly longer than the stamens. There are 6 to 8 long, fat,<br />

pinkish-white or tan stigma lobes.<br />

fruits: Spherical or even hemispherical in shape, about 1 inch<br />

long, but often a little wider than they are long. They are unarmed,<br />

whatever bristles were upon the ovary being early<br />

deciduous, and have very deeply pitted umbilici, and the rest<br />

of the surface covered with pronounced tubercles around 1/2


230 cacti of the southwest<br />

inch long and 1/8 to 3/16 of an inch high. They become yellow<br />

when ripe, but not juicy. The fruits usually remain on the<br />

branches for at least a year, and sometimes even turn yellow<br />

and then turn brown and dry on the branches, but all this<br />

without ever losing their conspicuous tubercles. The seeds are<br />

regular, smooth, and 1/8 of an inch or a little more (3–4 millimeters)<br />

in diameter.<br />

Range. From southeastern Colorado into New Mexico east of a<br />

line approximately from the upper Rio Grande to the vicinity<br />

of Grants, New Mexico, to Silver City, and on into Chihuahua.<br />

The eastern boundary of its range runs from the extreme southwestern<br />

corner of Kansas across the western part of the Oklahoma<br />

and Texas panhandles, then southeast to near Abilene,<br />

Texas, then back southwest to near Fort Stockton, and into<br />

Mexico between Sanderson and the Big Bend. It is encountered<br />

in northwestern Coahuila.<br />

Remarks. This is the large cholla common over so much of the<br />

western part of our area, and seen almost anywhere else reduced<br />

to the canes, lamp bases, and other trinkets made of reticulated<br />

cactus wood and brought home from western trips. However,<br />

the larger and more firm cholla wood comes from other species<br />

growing even farther west than this one.<br />

On the high plains of northwest Texas and at the tip of the<br />

Oklahoma Panhandle one may see proud old individuals of this<br />

cactus standing defiant of all the extreme elements, often miles<br />

apart, but clearly visible where no trees grow on the vacant<br />

prairies. Then again, some 20 miles northwest of Amarillo, the<br />

tree cactus somehow manages to take over whole stretches of<br />

the prairie and there are forests of individuals often 6 to 8<br />

feet tall. And farther southeast toward Big Spring, Texas, this<br />

cactus alternates with the mesquite to make a spiny mixed forest.<br />

The cactus also grows on mountain slopes farther south and<br />

west, into Mexico and almost to the Arizona border. Sometimes<br />

in the mountains it becomes very numerous, but the individuals<br />

rarely seem to become as robust and proud as on the prairies,<br />

usually remaining low bushes 3 or 4 feet high and showing signs<br />

of dying back almost as fast as they grow. The best stands of<br />

robust, healthy individuals I have seen in a mountain habitat<br />

are in the southwestern parts of the Davis Mountains.<br />

There is some variation in the color of the flowers of this cactus—the<br />

exact extent being hard to discover. Typically in Texas,<br />

the flowers are pale purple to lavender, with the outside of the<br />

segments often distinctly green and the center of the flower<br />

greenish. Almost everyone in the area of the Davis Mountains<br />

insists that occasional plants are seen there with white flowers,<br />

and that very rarely there have been seen flowers which were<br />

pale yellowish. These reports are so numerous as to be intriguing<br />

and, remembering the same sort of reports which, when finally<br />

investigated by an organized hunt during flowering season in<br />

the Arbuckle Mountains of Oklahoma, actually produced these<br />

flower colors in the also purple-flowered Echinocereus caespitosus,<br />

I have spent some time trying to substantiate them. So far,<br />

however, I have not seen these flower colors myself. On the<br />

other hand, the flowers on the cactus as it grows in the mountains<br />

of northern New Mexico, as for instance in Taos Canyon,<br />

are a very bright rose-purple or almost red, with no green at all.<br />

This is unlike anything I have seen on this cactus elsewhere.<br />

Boissevain gives purple to rose-pink for the Colorado specimens.<br />

The fruits of variety arborescens are entirely spherical or even<br />

broader than they are long, but always covered with large tubercles<br />

and having very large, very deep umbilici caving in the<br />

whole top of each. They ripen slowly, but by late summer are<br />

turning bright yellow. They remain on the plants most or all of<br />

the winter, usually falling in the spring. They do not change<br />

shape, however, remaining extremely tuberculate and pitted to<br />

the end. This shape distinguishes this plant most clearly from the<br />

following cactus which is probably another variety of the species.<br />

I am using Engelmann’s name, arborescens, for this cactus because<br />

we know this is exactly the form he was describing under<br />

that name. I have examined his types for his cactus, and this is<br />

his form.<br />

An admittedly older name, O. imbricata (Haw.) DC, exists,<br />

but this was the name given for a cactus whose type specimens<br />

or even whose type locality are unknown (beyond that it was<br />

from Mexico) and whose original description is so vague that it<br />

would fit almost any large cholla we have. Many people have<br />

thought that our U.S. cholla is the same as the Mexican one,<br />

and so have used the name, O. imbricata for our cactus. Why,<br />

then, am I returning to the use of Engelmann’s later name?<br />

In studying this cactus, I have followed Engelmann’s cactus<br />

which he called O. arborescens down from its southernmost U.S.<br />

range in the Big Bend of Texas into the mountains of north-<br />

western Coahuila, Mexico. All along I saw the same plant with<br />

its greatly tuberculate, but small fruits, and no other, until it<br />

reached the end of its range and disappeared near Monclova,<br />

Coahuila.<br />

Somewhat farther on, in the hills north of Saltillo, I came<br />

upon a similar but surely different cholla. This one grows with<br />

joints larger in both diameter and maximum length, and with its<br />

more or less ovate fruits becoming twice as big as those of the<br />

U.S. form, as well as becoming perfectly smooth as they ripen,<br />

all tubercles vanishing and the umbilici becoming flat. The resulting<br />

ripe fruit is quite like the shape of a fig. This cactus begins<br />

to appear near Saltillo, and is very common, becoming one<br />

of the dominants in much of the brush country all the way<br />

through San Luis Potosi, while our northern form does not ap-<br />

pear anywhere at all in this whole area.<br />

Both Coulter and Griffiths state clearly the differences between<br />

these two forms, as well as their proper ranges. Both use<br />

Engelmann’s name O. arborescens, for the northern one, while<br />

assuming that the southern one is the original O. imbricata (as<br />

did Engelmann) and using that name for it.<br />

Most later writers, following Schumann and Weber, ignored<br />

the differences between these two forms and united the two<br />

names as synonyms. While we feel that they may be too close


genus Opuntia 231<br />

to each other to be considered entirely separate species, we do<br />

not believe they are identical. They even show differing chromatograph<br />

maps, as we discovered in our own laboratory, which<br />

is added evidence for keeping them separate. While there is no<br />

certainty possible from only the very poor early descriptions of<br />

Haworth and De Candolle as to what the typical form of O. imbricata<br />

was, it is clear that it was a Mexican form, and that the<br />

much later discovered northern form Engelmann described was<br />

not for many years considered to be the same, although Engelmann<br />

himself speculated that his might not be a totally different<br />

species. We believe that, all evidence considered, our northern<br />

form can best be regarded as a variety of the larger species<br />

which all agree now must bear the original name, O. imbricata,<br />

and so we call it O. imbricata var. arborescens.<br />

Opuntia imbricata var. viridiflora (B. & R.)<br />

Description Plate 61<br />

stems: Growing as an upright bush 1 to 3 feet tall. It is much<br />

branched and with no enlarging trunk, but the old stems are<br />

more or less bark-covered and with a woody support. The<br />

current year’s joints are usually 1/2 to 3/4 of an inch in diameter,<br />

but occasionally in cultivated examples reach 1 inch<br />

thick. The surface is covered with prominent tubercles 3/4 of<br />

an inch or a little longer.<br />

areoles: Circular or oval, with short gray or yellowish wool.<br />

spines: 2 to 8 per areole on current year’s growth, not increasing<br />

much with age. They are 3/4 to 1 inch long, dark<br />

brown, with brownish sheaths.<br />

glochids: Fairly numerous, but short.<br />

flowers: 1 to 2 inches in diameter, not opening widely. They<br />

are coral-pink within, yellowish to pale green on the outside.<br />

The filaments are green, the anthers yellow. The pistil is long<br />

and reddish, and the stigma has 8 or 9 reddish lobes.<br />

fruits: 1 inch or less in diameter, as those of variety arborescens<br />

in shape, with very prominent, persisting tubercles and<br />

deep umbilici. They have long, deciduous bristles at first, but<br />

are usually naked when ripe.<br />

Range. Known only from the type locality, which is the hills<br />

just north of Santa Fe, New Mexico.<br />

Remarks. Although described by Britton and Rose as a separate<br />

species, I find it only possible to think of this form as at best a<br />

doubtful variety of the O. imbricata complex very close to the<br />

other northern form, variety arborescens. Growing wild, it appears<br />

in no essential character different from a small, stunted<br />

version of that cactus, all its elements present but reduced quantitatively.<br />

Many of the weaker specimens of variety arborescens<br />

found in the mountains of northern New Mexico which I have<br />

examined I could not distinguish from this form by their vege-<br />

tative characters. Only when they bloomed with their bright<br />

purplish flowers could I be Sure they weren’t this form, and<br />

only when my specimens bloomed with their pale greenish flow-<br />

ers could I be Sure that they were variety viridiflora.<br />

It does not seem that flower color can be a basis in the cacti,<br />

at least, for separating species, and anyway, most of the very<br />

large, robust variety arborescens specimens in northwest Texas<br />

bloom with pale lavender flowers having traces of green in<br />

them. So if there were no other factors I would think the two<br />

should be combined. However, the cactus described as O. viridiflora,<br />

even in cultivation in New Mexico where it has the best<br />

situation, does not seem to grow large like the other forms of<br />

the species. The quantitative difference remains. Yet this is still<br />

little basis for separating the forms. Figuring even more importantly<br />

in our evaluation, the chromatograph map of the cactus<br />

is quite essentially different from that of variety arborescens.<br />

For those reasons mentioned, I continue to keep this form<br />

separate, but I definitely do not feel that the differences are<br />

enough to warrant considering this more than a local variety<br />

within the O. imbricata complex.<br />

Opuntia imbricata var. vexans (Gr.)<br />

Description Plate 61<br />

stems: A treelike cholla with cylindrical branches from an<br />

upright, bark-covered trunk. Old specimens stand 6 to 12 feet<br />

tall with a compact crown of many branches from a cylindrical<br />

trunk becoming at least 8 inches in diameter. The current<br />

years’s branches are 4 to 16 inches long, the typical length<br />

being 6 to 12 inches. They are cylindrical, 3/4 to 1/4 inches<br />

thick, usually narrowing gradually at their bases. They are<br />

tuberculate, the tubercles about 11/4 inches long and 3/16 to 3/8<br />

of an inch tall, with the upper slope abrupt and the lower<br />

slope much longer and more gradual.<br />

areoles: Oval, 1/4 of an inch or slightly more in length, with<br />

gray wool at first, later enlarging and bulging outward.<br />

spines: 1 to 10 in the areoles of current growth, increasing to<br />

20 or 30 on old stems. Spreading in all directions, short, being<br />

only 1/4 to 5/8 of an inch long, and very slender to medium in<br />

thickness. They are brown, more or less annulate, with lighter<br />

tips, and are covered with rather tight, yellowish, gray, or<br />

whitish sheaths.<br />

glochids: Yellowish in color. They grow from the upper part<br />

of the areole, and are always very few, and from 1/16 to 3/8<br />

of an inch long.<br />

flowers: 2 to 23/4 inches in diameter by 11/2 to 2 inches tall.<br />

The ovary is 1 to 11/8 inches long by about 1/4 of an inch thick,<br />

very tuberculate, with a number of long perianth segments<br />

upon it. The outside of the flower is greenish, the inside light<br />

pinkish-purple. The filaments are pink suffused with green or


232 cacti of the southwest<br />

brown. The anthers are yellow or cream; the style is short;<br />

and the stigma lobes number 6 to 9 and are brownish, tan, or<br />

whitish. This form tends to bloom repeatedly during the summer,<br />

whenever there is moisture.<br />

fruits: When young, egg-shaped, about 13/4 inches long by 1<br />

inch wide, very tuberculate with tubercles 5/8 to 3/4 of an inch<br />

long and about 1/8 of an inch tall, and with the umbilici deeply<br />

pitted. At this stage they have some short glochids and around<br />

the upper edge half a dozen or so very slender, white spines<br />

3/8 to 13/8 inches long. When ripening later in the summer,<br />

they become yellowish-green, egg-shaped to almost spherical<br />

or sometimes even wider than they are long, being then 11/4<br />

to 2 inches long and 11/2 to 2 inches wide. These ripe fruits<br />

are spineless, wholly smooth, or with only the traces of a few<br />

tubercles at the base, and with the umbilici practically or completely<br />

flattened.<br />

Range. Said in the original description to be Webb County,<br />

Texas. It has not been seen growing wild since, but is found as a<br />

cultivated plant in gardens in Laredo and Del Rio, Texas.<br />

Remarks. Dr. Griffiths described O. vexans as a new species in<br />

1911. He said then: “The type specimen is… prepared… from<br />

plants cultivated from cuttings collected… in Webb County,<br />

Texas, March 13, 1908. Mature plants have not been seen elsewhere,<br />

but the species is frequently cultivated. The description<br />

is a compilation of several sets of notes taken from native and<br />

cultivated plants.”<br />

We carried on a long search for a cholla native to Webb County,<br />

covering as much of the area as possible ourselves, and asking<br />

everyone we could find who had reason to be abroad in the<br />

open, from ranch owners to cowboys, and particularly oldtimers,<br />

where the cholla grew wild in Webb County. Everywhere we<br />

met with complete surprise that anyone thought a cholla ever<br />

grew wild anywhere in that county. We were stymied entirely<br />

in trying to locate Griffiths’ cholla this way.<br />

But Laredo, in Webb County, is the site of one of the oldest<br />

and most famous cactus dealerships. In business continually for<br />

well over 30 years, first as the Shiner Cactus Nursery, and more<br />

recently as the Cactus Garden and Cafe, this establishment has<br />

sold literally millions of cacti from the local region, as well as<br />

imported cacti from all over Mexico. It still has a large garden<br />

with hundreds of huge old specimen plants which have stood<br />

there for decades. Mrs. Jones, the present proprietor, has been<br />

involved with the business almost since the beginning.<br />

Naturally, since I believe that she knows the most about the<br />

cacti of Webb County of anyone living, I asked her about Griffiths’<br />

cholla. She says that she has never known of any such<br />

native plant. She has had crews of men in the field at various<br />

times during the years, digging cacti for her business, and says<br />

she was never shown a cholla from the region.<br />

Although one hates to write off the statements of one so eminent<br />

as Dr. Griffiths, I was faced with having to do this, since<br />

I couldn’t even find a clue to his plant. Then one day I walked<br />

out behind the Cactus Garden Cafe in Laredo, and there on the<br />

edge of Mrs. Jones’s garden, fully 12 feet tall and vying successfully<br />

with the retama trees, stood a giant specimen of Griffiths’<br />

cholla. It had a trunk more than 6 inches in diameter, with its<br />

first branches 4 feet or more above the ground, and with a<br />

crown spreading 10 or 12 feet. On the branches were young,<br />

tubercled, spiny fruits, and also large, spherical, perfectly smooth<br />

and spineless, ripe fruits of the season’s several blooming periods.<br />

When I told Mrs. Jones that this was the cholla from Webb<br />

County, she disagreed, saying that it was instead an import from<br />

Mexico which had stood in that spot for very many years. There<br />

is a chance that this very specimen was growing there when Dr.<br />

Griffiths was collecting in Webb County, and it is entirely possible<br />

that he used it in writing his description. I have since seen<br />

several other specimens of this cactus in Laredo, but there are<br />

not many, and they are all in very old yards.<br />

There was another old business dealing in rocks, cacti, and<br />

curios along the highway in Del Rio, Texas. This was run for<br />

many years by Mr. and Mrs. Basket. In the yard around this<br />

establishment was to be seen a whole grove of 20 or more specimens<br />

of this cholla 8 to 12 feet tall. I was privileged to know<br />

Mrs. Basket, and she said these cacti were not native to Texas,<br />

but had their source somewhere in Mexico. Mrs. Basket sold her<br />

property recently, and within the past year all of these giant<br />

specimens were grubbed out and destroyed. I do not know that<br />

the form exists any more in Del Rio.<br />

There were also two fine specimens of this cholla beside a<br />

small roadside cafe and station some few miles west of Del Rio<br />

on U. S. 90, but this business has been razed and the plants went<br />

with it. So far as I know, the few Laredo specimens and some<br />

starts in my own collection are all that survive of the form.<br />

It is impossible with only the present knowledge of this cholla<br />

group to know what to do with this cactus. It seems doubtful,<br />

from all accounts, that the plant ever grew wild in the U.S.,<br />

and more likely that Dr. Griffiths saw examples escaped from<br />

very early cultivation, if he did see the cactus growing outside<br />

of yards. Neither is there any sign of the cactus growing wild<br />

anywhere near the Rio Grande in north Mexico. The nearest<br />

location to Webb County where I find chollas with fruits becoming<br />

smooth like this is in the hills around Saltillo, Coahuila,<br />

Mexico, and this might be a location from which it was introduced<br />

into Texas long ago; but the form with smooth fruits<br />

growing from Saltillo down through San Luis Potosi does not<br />

seem to be entirely identical with the form in Webb County. It<br />

does not grow so large anywhere that I have observed it, and<br />

there are differences of tubercle shape, spine development, and<br />

fruit details, which do not seem that they could be environmen-<br />

tal.<br />

What does seem very clear is that the cactus called O. vexans<br />

by Griffiths is different from the U. S. cholla which we are calling<br />

O. imbricata var. arborescens, having a different growth<br />

form when large, thicker joints, shorter spines, and different<br />

fruits. Since it is going to be noticed by some observant cacto-


genus Opuntia 233<br />

philes in Laredo yards, if nowhere else, I feel we must list it<br />

here if we are to have a complete account. That it is definitely<br />

one of the O. imbricata group is also clear. But the problem of<br />

how it relates to the typical O. imbricata is a hard one, mainly<br />

because, the early descriptions of that cactus being in only the<br />

most general terms and no type or type locality being given, we<br />

do not really know the exact form of the typical species type.<br />

It is assumed by all that the original collection was in Mexico,<br />

and a logical choice for the typical species form would be the<br />

form which is so common as to be one of the dominant plants<br />

from southern Coahuila at least to San Luis Potosi. Griffiths<br />

himself flatly calls this form O. imbricata, and yet he never<br />

seems to have thought of the plant he saw in Webb County as<br />

this one. I have studied this cactus widely in Mexico, and Griffiths’<br />

O. vexans is very close to it, but I can see differences also.<br />

Yet even if I were to conclude that they are the same, I would<br />

still hesitate to state that the form was the typical O. imbricata,<br />

since Griffiths also describes an O. spinotecta from Durango,<br />

which I have seen, and which is still different from any of the<br />

above, with only 3 to 6 spines, even on old stems, but with these<br />

much longer than on the other Mexican forms. This form is the<br />

one which fits Coulter’s description for O. imbricata, and could<br />

just as easily be the one which should bear the name.<br />

With nothing more definitely known than all this, it seems<br />

most prudent to leave our cactus growing, probably as a visitor,<br />

in Webb County, Texas, as a variety of this complex under the<br />

name O. imbricata var. vexans. With all the lack of certainty<br />

about it, the name is certainly well enough chosen.<br />

Opuntia spinosior (Eng.) Toumey<br />

“Cane Cholla”<br />

Description Plate 62<br />

stems: A large, open-branching shrub or treelike plant 3 to 8<br />

feet, and said to sometimes reach 12 feet tall. When large it<br />

produces a black, scaly trunk 2 to 4 inches in diameter, although<br />

smaller, shrubby examples will lack this. The young<br />

joints are 4 to 12 inches long, cylindrical, 5/8 to 11/4 inches in<br />

diameter, covered with conspicuous tubercles 1/4 to 3/4 of an<br />

inch long. The surface is grayish-green, often becoming pur-<br />

plish in the winter.<br />

areoles: Oval or nearly so, 1/8 of an inch in diameter when<br />

young, becoming larger and bulging outward with age.<br />

spines: There are 6 to 12 Spines per areole on the current<br />

year’s growth, the number increasing to at least 20 or 30 when<br />

very old. They are short, mostly 1/4 to 1/2 inch long, but rarely<br />

to 3/4 of an inch, and spread in all directions. They are white,<br />

gray, or brownish, all fading to gray. At first they are covered<br />

entirely, or the smaller ones on only the upper parts,<br />

with thin, comparatively inconspicuous sheaths which usually<br />

fall off after a year. The sheaths are gray, sometimes with a<br />

pinkish tinge.<br />

glochids: There are a few short, inconspicuous, white or yellowish<br />

glochids at the top of the areole.<br />

flowers: 1/2 to 21/4 inches in diameter, very beautiful, but<br />

also variable in color. The most common color seems to be<br />

purplish, but it is not uncommon to find various shades of<br />

red, more or less yellowish, or even white blossoms. The ovary<br />

is tubercled and with very slender white spines on it. The<br />

stigma lobes are cream-colored or yellowish.<br />

fruits: Yellow when ripe, 1 to 11/2 inches long, almost spherical<br />

to oblong or rather egg-shaped. They are very tuberculate,<br />

with also deeply pitted apexes. At first the fruits have<br />

the very slender white spines of the ovary upon them, but<br />

these fall off long before they ripen. The seeds are a little over<br />

1/8 of an inch (about 4 millimeters) in diameter.<br />

Range. Southwestern New Mexico from about Silver City to<br />

Hermanas and the Carrizalillo Hills, and on into adjacent Ari-<br />

zona and Mexico.<br />

Remarks. This is thought by many to be the most beautiful<br />

cholla of our area. Its treelike, open branching, and tidy manner<br />

of growth, its even covering of short, whitish spines, and its<br />

brilliant flowers make it a favorite. It is so often featured in the<br />

bright color photos of the southwestern desert which have become<br />

a fad these days that one might think he could find it anywhere<br />

he would look in that desert. The cactus really has a comparatively<br />

small range in the U.S., however, being found only<br />

in the southwest corner of New Mexico and over about the<br />

southeast quarter of Arizona. Other large chollas take its place<br />

farther west, and the casual traveler doesn’t stop to notice the<br />

difference.<br />

This cactus does have superficial similarities to the more<br />

northern and much smaller O. whipplei, and Engelmann confused<br />

them in all of his accounts, calling this cactus O. whipplei<br />

var. spinosior. There are various differences between the two,<br />

however, which will be noted in the following account of that<br />

other species.<br />

O. spinosior is another of the large chollas having a woody<br />

skeleton of beautiful open pattern from which everything from<br />

canes to lamp bases has been made, hence the common name. It<br />

is a favorite especially for cane work, because of the straight-<br />

ness of its stems.<br />

Opuntia whipplei Eng. & Big.<br />

“Whipple’s Cholla,” “Rat-Tail Cactus,” “Sticker Cactus”<br />

Description Plate 62<br />

stems: Ordinarily, and apparently always in our area, an<br />

erect but low-growing cholla 6 to about 24 inches tall. Plants


234 cacti of the southwest<br />

may stand alone and be rather openly branched bushes, the<br />

joints 6 to 10 inches long, but more often they are very<br />

crowded with their branches extremely numerous and their<br />

joints only 2 or 3 inches long, these forming very compact,<br />

matlike thickets of sometimes up to 8 feet across while still<br />

only a foot or so high. In some locations in Arizona the plant<br />

is reported to grow much taller, but this has not been demonstrated<br />

elsewhere. The current year’s joints are usually 3/8<br />

to 3/4 of an inch in diameter, but occasionally to 1 inch. They<br />

are covered with conspicuous, broad, short tubercles 3/8 to 5/8<br />

of an inch long. The surface is light or yellowish-green. The<br />

joints are rather easily detached.<br />

areoles: Egg-shaped or elliptical, about 3/16 of an inch long<br />

when young. They have some white wool at first, but very<br />

soon lose it.<br />

spines: There are 3 to 12 spines per areole. 1 to 4 of these are<br />

main spines spreading from the center of the areole. The<br />

lower, rather deflexed one is usually the longer of these, and<br />

is often somewhat flattened. The uppermost one is flattened<br />

and the heaviest, but while all of these are rigid, none of<br />

them are very stout. They are from 1/4 to 11/4 inches long.<br />

They are whitish to brown and covered with conspicuous,<br />

loose, straw-colored or whitish sheaths. There are 2 to 8 smaller,<br />

very slender, unsheathed spines 1/16 to 3/8 of an inch long,<br />

radiating or often recurving against the surface of the plant<br />

around the areole.<br />

glochids: There are several white or pale yellowish glochids<br />

up to 1/8 of an inch long near the upper edge of the areole.<br />

flowers: Pale yellow, not opening widely. They usually stand<br />

3/4 to 11/4 inches wide. The ovary tube is very tuberculate,<br />

with large white areoles and a few slender, white, soon-falling<br />

spines upon it. The filaments are greenish. The anthers are yel-<br />

lowish. The stigma lobes are about 5 or 6 in number and<br />

greenish or white in color.<br />

fruits: Almost round to somewhat obovate, 3/4 to 11/4 inches<br />

long. They are very tuberculate, with deep umbilici, and without<br />

spines. They remain green a long time, and then become<br />

yellowish when ripe. The seeds are 1/8 of an inch or slightly<br />

more (3–4 millimeters) in diameter.<br />

Range. Northwestern New Mexico from south of Ojo Caliente<br />

in Valencia County to western Socorro County around Puertecito,<br />

to near Farmington. Extending into southwestern Colorado<br />

and over almost the whole upper half of Arizona.<br />

Remarks. There has been much misinformation about this cactus<br />

because it has from the first been confused with two other chollas.<br />

Engelmann apparently never did distinguish clearly between<br />

this and O. spinosior. He began by having a species, O.<br />

whipplei, which encompassed both of these forms, and then<br />

describing under this species a variety laevior, which sounded<br />

much like this cactus, but for which he described the flowers as<br />

being red, and a variety spinosior, which was the larger plant.<br />

Later, in his Whipple’s Report, he stated his earlier described<br />

red flowers were actually those of variety spinosior, but he did<br />

not redescribe the flowers of the smaller plant.<br />

Coulter restricted the name of the species to the small northern<br />

cholla, and it has been called O. whipplei constantly since<br />

that time, but he repeated the error of stating that it had red<br />

flowers, and still called the larger southern cactus O. whipplei<br />

var. spinosior.<br />

The two cacti are so close to each other in many respects of<br />

stems and spines that it would be very easy to think of them as<br />

a large growth form in the south and a smaller one in the north<br />

due to more severe climate there, especially if one were thinking<br />

that the flowers were identical on both. Only later did Tourney<br />

separate the two as different species, and by Wooton’s time it<br />

was clear that in New Mexico at least, which is the type locality<br />

of O. whipplei, the flowers of the smaller northern form are always<br />

yellow, and the ranges of the two are separated by a band<br />

of at least 100 miles where neither grows. Since then the two<br />

have always been considered separate species, although Arizona<br />

students have mentioned what they think may be intermediates<br />

in their state. These have not been demonstrated in New Mexico.<br />

After all this had been pretty well settled, a new confusion<br />

arose between O. whipplei and another cholla of northeastern<br />

New Mexico, Oklahoma, and Texas, O. davisii. Here is another<br />

form surely very close to O. whipplei. It was only incompletely<br />

described by either Engelmann or Coulter, and Wooton seems<br />

the first one actually to try to distinguish between these two<br />

forms. Unfortunately, the only character he could find with<br />

which to try to tell them apart was the spine color. He said that<br />

O. whipplei has “spine sheaths pale yellowish to white,” while<br />

O. davisii has “spine sheaths yellowish brown,” or, as he put it<br />

again, “O. whipplei always looks whitish or very pale yellow,<br />

while O. davisii is a golden brown, the colors being due to the<br />

sheaths.”<br />

Now this distinction pointed out by Wooton is essentially accurate,<br />

but few things are more relative with us than color, both<br />

in the naming of the various shades and no doubt in the perception<br />

of them. The result was that other characteristics in<br />

which these two differed were not properly considered while<br />

people began to call their specimens whichever one of these they<br />

were more familiar with. Due to this confusion we have reports<br />

of O. whipplei in northeastern New Mexico and even in Oklahoma.<br />

All of these specimens collected outside of the above given<br />

range of O. whipplei which I have been able to investigate personally<br />

have proved to be O. davisii. A typical example is a<br />

collection made by Lahman in Greer County, Oklahoma, on the<br />

basis of which O. whipplei is often included in Oklahoma plant<br />

lists. I have examined Lahman’s preserved specimen, and it is<br />

clearly O. davisii.<br />

We need much more reliable characters with which to distinguish<br />

these two species, and I would mention that the length


genus Opuntia 235<br />

of spines of O. whipplei is usually no more than 3/4 of an inch<br />

and never over 11/4 inches, and the length of its tubercles is 3/8<br />

to 5/8 of an inch, while the spines of O. davisii are to 2 inches<br />

long and its tubercles 5/8 to 1 inch long. The flowers of the former<br />

are yellow and it produces seeds, while the latter has green<br />

flowers and is apparently sterile.<br />

O. whipplei, when definitely recognized, is the small pest<br />

cholla of northwestern New Mexico and northern Arizona. It<br />

has little to commend it to us, and its impenetrable thickets,<br />

with its severe spines and joints which easily attach themselves<br />

to us and hate to let go again, make it less than appreciated by<br />

most who encounter it. Although it is one of the forms which<br />

give cacti little besides a bad name, we must acknowledge it as<br />

one of ours.<br />

Opuntia davisii Eng.<br />

Description Plate 63<br />

stems: A low-growing, very much branched bush usually<br />

standing 12 to 18 inches tall, but occasionally reaching 30<br />

inches. The main stem is very short because of the immediate<br />

branching and it hardly increases in diameter, but it does become<br />

more or less covered with gray, scaly bark. The current<br />

year’s joints are cylindrical, often bordering upon being clubshaped<br />

because of the narrowing of their bases. They are 3 to<br />

6 inches long, 1/4 to 3/4 of an inch in diameter, very tuberculate,<br />

with the tubercles laterally compressed and 5/8 to at<br />

least 1 inch long. The color of the surface is light green. The<br />

joints are easily detached.<br />

areoles: Elliptical, about 3/16 is of an inch long, with some yellowish<br />

wool.<br />

spines: Producing 6 to 13 spines per areole. 4 to 7 of these are<br />

main spines spreading in all directions from the center of the<br />

areole. They are from 3/4 to 2 inches long, round, and heavy.<br />

They are bright brown or red-brown and somewhat annulate<br />

in color, but this coloring is not seen until their very large,<br />

very loose, bright, glistening, straw-colored, or light brown<br />

sheaths are removed. There are also 2 to 5 small radial spines<br />

which are 1/4 to 1/2 inch long, slender, brownish, and do not<br />

appear to be sheathed.<br />

glochids: There is a compact cluster of yellow glochids 1/16<br />

to 1/8 of an inch long at the top of the areole.<br />

flowers: About 2 inches tall, but not opening widely, and so<br />

only about 11/2 inches in diameter. They are deep green to<br />

pale green in color, the centers having yellowish coloring and<br />

the upper edges and outside surfaces of the petals tinted with<br />

brown or reddish. The filaments are greenish-red and very<br />

coarse, and the anthers yellow. The style is short, with 4 to 7<br />

very large, cream-white to sometimes pale purplish lobes. The<br />

ovary is conic, about 11/4 inches long by 1 inch wide, very<br />

tuberculate, with some yellow areoles and on the upper part<br />

some very slender, white, deciduous spines to 1 inch long.<br />

fruits: Egg-shaped to clavate, 1 to 11/2 inches long by 5/8 to<br />

3/4 of an inch thick at the top. They are very tuberculate, with<br />

deeply pitted umbilici. There are short yellow glochids upon<br />

them, but they are otherwise naked. They become greenish-<br />

yellow and then dry up. All examples examined were<br />

sterile.<br />

Range. From Greer and Harmon counties in extreme south-<br />

western Oklahoma, west across the Texas Panhandle and eastern<br />

New Mexico to near the mountains, also south across Texas<br />

to Gillespie County in the south-central part of that state, and<br />

to the Rio Grande in the Big Bend.<br />

Remarks. Opuntia davisii is the eastern counterpart of the more<br />

western O. whipplei. The two have many characters in common,<br />

and since, in most keys, as for instance those of Wooton and of<br />

Britton and Rose, the only way mentioned to distinguish them<br />

is by spine-sheath color, they have often been confused. Frequently<br />

this has worked to the advantage of O. davisii, in terms<br />

of the range given to it. In Coulter’s time, records were so confused<br />

between these two and even other small chollas that he<br />

gives the range of O. davisii as going all the way to California.<br />

Wooton seemed to be able to distinguish these plants well for<br />

himself, and put this cactus back in eastern New Mexico where<br />

it belonged, but his key led many others farther astray. There<br />

is just not enough difference between “sheaths pale yellowish to<br />

white—O. whipplei” and “sheaths yellowish brown—O. da-<br />

visii.”<br />

Perhaps the worst confusion was that of Boissevain and Davidson.<br />

Partly because of the above trouble with spine colors<br />

and partly because they repeat Engelmann’s erroneous statement<br />

that O. whipplei has red flowers, they call their small, yellow-<br />

flowered, whitish-spined cholla of southwestern Colorado O.<br />

davisii, when it is in reality O. whipplei. To date there are no<br />

actual records of O. davisii in Colorado, much less in extreme<br />

southwestern Colorado as they would have it.<br />

Conversely, Mrs. Lahman collected a cholla which she called<br />

O. whipplei in Green and Harmon counties, Oklahoma. Examination<br />

of her preserved specimens and re-collection of the living<br />

cacti show that they are clearly O. davisii.<br />

How can this sort of confusion between these two interesting<br />

plants be avoided? Only by noting that beyond the glistening<br />

straw or golden-brown sheaths and the more or less red-brown<br />

color of the main spines, O. davisii has 4 to 7 of these main<br />

spines 3/4 to 2 inches long, whereas O. whipplei shows no more<br />

than 1 to 4 main spines of which none are over 11/4 inches long.<br />

Also, O. davisii has tubercles compressed and 5/8 to 1 inch long<br />

instead of broader and shorter ones such as are found on the<br />

other cactus, and its flowers are at best yellowish-green while


236 cacti of the southwest<br />

the flowers of the other are clear yellow. With this sort of careful<br />

observation the two can be distinguished with certainty unless,<br />

of course, one is looking at stunted, abnormal growths, and<br />

in this condition one can hardly separate any of the Opuntias.<br />

In extremely stunted conditions, purposely induced in my garden,<br />

O. davisii takes on much of the aspect of O. kleiniae, with<br />

long, very slender joints, very indistinct tubercles and down to<br />

1 slender spine, while O. whipplei reacts in just the opposite<br />

manner, shortening its joints which remain very tuberculate and<br />

come to appear much like poorly armed, over-elongated exam-<br />

ples of O. grahamii.<br />

Hester reported O. davisii from the Big Bend of Texas, a very<br />

long way from any other report of it. Anthony later confirmed<br />

this, and I have re-collected it there myself. Some of Hester’s<br />

plants had very long roots with tuberous thickenings strung<br />

along them. I have not been able to find this sort of root forma-<br />

tion again.<br />

Not only is the occurrence of the cactus in the Big Bend, so<br />

far from any other report of it, not surprising, but such isolated<br />

records seem to be the rule with this species. It occurs over a<br />

very large range, but is found in most of this huge area at only<br />

widely scattered locations. The center of its range is clearly a<br />

strip extending about 50 miles either side of the Texas, New<br />

Mexico boundary from about Nara Visa, New Mexico, on the<br />

north to about Tatum, New Mexico, on the south. In this strip<br />

of territory one can find a small stand every 30 to 50 miles<br />

wherever he goes, and there are some expanses of the prairie<br />

where the individuals are up to a dozen or so to the square mile.<br />

But when one goes any direction from this strip, the occurrences<br />

are very widely spaced. Going west we find records from the<br />

Tucumcari Hills, and then no sign of it again until between<br />

Chimayo and Espanola, New Mexico, we again find a real<br />

stand. Going southwest or south there has been no known sign<br />

of it past Tatum, New Mexico, except Hester’s collections of it<br />

near Marfa, Texas, and at some unspecified place on the Rio<br />

Grande, and Anthony’s and my own collections of it, both a<br />

few miles from Marathon. Going east from the New Mexico-<br />

Texas border one can quickly list the known locations. They are<br />

Abernathy in Hale County, Texas, north of Lubbock; in Kent<br />

County, southeast of Lubbock some 60 miles; and then not again<br />

until a location in the southwest corner of Beckman County,<br />

one in Harmon County, and one in Greer County, all in Oklahoma;<br />

and yet one more at Seymour in Baylor County, Texas.<br />

Going southeast from the original area one finds it again around<br />

Colorado City, Texas, once again 60 miles or so from any other<br />

record, and then not again until a small population is found in<br />

northern Gillespie County, Texas, fully 160 miles from the near-<br />

est location and well down into the Texas hill country.<br />

This sort of widespread range with very scattered locations<br />

within it seems typical of many chollas. It may indicate, as some<br />

have conjectured, that these plants were previously much more<br />

common and that we have only islands of survival left, or it<br />

may be the result of the highly developed ability in these plants<br />

to reproduce themselves from detached joints. These joints cling<br />

tightly to flesh, fur, or clothing, and perhaps these plants were<br />

spread to their widely separated locations by the migrating ani-<br />

mals and wandering tribes of the prairies.<br />

The flower of O. davisii is among the most unusual and remarkable<br />

I have seen in the cacti. Its petals are very firm and<br />

waxy, unlike those of any other Opuntias I know, and similar<br />

to the stiff, persistent flower petals of the Echinocereus triglochidiatus<br />

group. It is the largest green flower with which I<br />

am acquainted, except for some orchids.<br />

Another peculiarity of this species is the fact that all fruits<br />

seem perfectly formed and developing normally, with many<br />

plants producing fruits in season, but seeds have never been described.<br />

While living in the Oklahoma Panhandle and traveling<br />

through the central range of this species in all seasons, I have<br />

opened many fruits, and all examples I have opened have been<br />

sterile. There did not seem to be any sign of parasites, and sterility<br />

appears to be a characteristic of the cactus, its reproduction<br />

apparently being solely by separated joints. What the sig-<br />

nificance of this is has not been determined.<br />

The sight of a large, healthy specimen of this cactus standing<br />

as a glistening, golden bush on the prairie makes one look<br />

around for the Midas responsible. Its spine sheaths in shining<br />

gold are only equaled in brilliance by those of the next species<br />

which glisten as brightly in silver.<br />

Opuntia tunicata (Lem.) Link & Otto in Pfeiffer, 1837<br />

“Abrojo,” “Clavelina”<br />

Description Plate 63<br />

stems: Erect, with a more or less definite woody stem, but<br />

with very many crowded, lateral branches, and, therefore, low<br />

and spreading in aspect. It grows from less than 1 to about<br />

2 feet tall in our area, but is said to grow taller sometimes in<br />

Mexico. The current year’s joints are 2 to 6 inches long, narrowly<br />

oblong or somewhat club-shaped when short, but almost<br />

cylindrical when longer. They are covered with prominent<br />

tubercles 3/4 to 11/2 inches long. The color is medium to<br />

light green. All joints are very easily detached.<br />

areoles: Oblong in shape, 3/16 of an inch long at first, enlarging<br />

considerably with age. They bulge outward with very<br />

white wool.<br />

spines: There are 6 to 10 Spines, all white, yellow, or rarely<br />

reddish, but all covered with very loose, very thin and papery,<br />

translucent, silvery white sheaths. There are 3 to 6<br />

spreading central spines 1 to 21/2 inches long. Underneath<br />

their sheaths these are more or less angular, and very heavy<br />

spines. There are also 2 to 4 radial spines from the lower part


genus Opuntia 237<br />

of the areole, these more slender and from bristle-like ones<br />

only 3/8 of an inch long to sometimes fairly stout ones, to 11/4<br />

inches long.<br />

glochids: There is a small cluster of very short whitish or<br />

pale yellow glochids in the upper edge of the areole.<br />

flowers: Pale greenish-yellow. About 2 inches in diameter.<br />

fruits: Spherical to broad club-shaped, tuberculate, yellow-<br />

ish-green. The seeds are apparently undescribed.<br />

Range. Found in the U. S. only on the southeast slopes of the<br />

Glass Mountains just within the southwest edge of Pecos County,<br />

Texas. Ranging throughout central Mexico, and reported also in<br />

Ecuador, Peru, and northern Chile.<br />

Remarks. Miss Anthony reported this species not many years<br />

ago from the Big Bend of Texas, this being the first knowledge<br />

that it grew in the United States. Dr. Warnock, of Sul Ross College<br />

in Alpine, Texas, kindly sent me some specimens collected<br />

on the Glass Mountains, the same location where Miss Anthony<br />

found it. We seem to have in that small area a very isolated<br />

population of the species, as it has never been reported from any<br />

of the Big Bend south of there to the Mexican border. This population<br />

must be the northernmost point it attains, a remarkable<br />

outpost since, although I have watched for it on several trips<br />

through the mountains of northwest Coahuila, the first place I<br />

have seen it again was at the latitude of Saltillo, hundreds of<br />

miles into Mexico.<br />

This gives the cactus ample reason to be considered for the<br />

title of the most widely ranging of all, since this means a range<br />

for it extending from the southern U. S. all the way into northern<br />

Chile. It is also said to have been found, probably as an<br />

escape, in Cuba.<br />

This immense range is all the more remarkable since this is<br />

probably the most difficult cholla to grow in cultivation. Even<br />

here in Texas, when trying one after another of the treatments<br />

which enable me to grow and flower most of the other cacti<br />

successfully, I find my specimens of O. tunicata, both from Texas<br />

and from Mexico, which were large and beautiful when I got<br />

them, refusing to bloom at all and reverting to the curious<br />

dwarf form, 8 to 10 inches high, with very short branches and<br />

less than typical spination, which one sees here and there even<br />

among the robust examples in the mountains of Mexico. The lack<br />

of published details about the flowers and fruits of this cactus<br />

makes me think that others have had the same trouble with it,<br />

and that perhaps it blooms and fruits sparingly even in the wild.<br />

At any rate, it has managed to survive in many places. However,<br />

the dwarf form of it is widespread even in the best loca-<br />

tions, as Britton and Rose mention.<br />

When growing robustly, O. tunicata is an amazingly beautiful<br />

plant. A good specimen is a small bush so compact in its branching<br />

and so covered with its shining silver-sheathed spines that it<br />

glistens, even from across a canyon, like an ice-covered bush in<br />

the sunlight of a northern winter, or like a globe of the finest<br />

Spanish silver filigree. But for all the magic of its appearance,<br />

it is a cactus which must be skirted by man or beast with the<br />

proper respect.<br />

Opuntia kleiniae DC<br />

“Klein Cholla,” “Candle Cholla”<br />

Description Plate 64<br />

stems: Erect, shrubby stems 3 to 6 feet tall, openly but rather<br />

sparingly branching from woody trunks 1 to 11/2 inches in<br />

diameter and covered with brown, scaly bark. The current<br />

year’s joints are 4 to at least 12 inches long, cylindrical, 5/16<br />

to 1/2 inch in diameter. They have low, broad, more or less indistinct<br />

tubercles 1/2 to as much as 13/8 inches long. The lateral<br />

branches are fairly easily detached, but are not as loosely held<br />

as on some chollas.<br />

areoles: Round to egg-shaped or even practically triangular,<br />

and about 3/16 of an inch long. They have white wool in them.<br />

spines: 1 to 4 per areole, with most often only 1 present. This<br />

main spine is 1/2 to at least 11/2 inches long, and said to reach<br />

2 inches; reddish to gray, with a loose yellowish sheath which<br />

usually falls off soon after the spine matures. There may be 1<br />

to 3 additional shorter Spines.<br />

glochids: A small, compact cluster of yellowish or brown<br />

glochids about 1/16 of an inch long is found at the upper edge<br />

of the areole.<br />

flowers: 1 to 11/4 inches in diameter, pale greenish-purple,<br />

lavender, or pinkish in color. The ovary is egg-shaped, 1/2 to<br />

3/4 of an inch long, with woolly areoles on indistinct tubercles.<br />

The filaments are pinkish, the stigma with 6 or 7 small, whit-<br />

ish lobes.<br />

fruits: Egg-shaped or sometimes almost club-shaped, 3/4 to<br />

11/2 inches long, red or bright orange, naked or with clusters<br />

of brown glochids 1/8 of an inch long in the areoles when ripe.<br />

They are more or less tuberculate when growing, and sometimes<br />

remain strongly tuberculate but more often become almost<br />

completely smooth when ripe. The amount of growth<br />

influences the degree of smoothness attained, since smaller<br />

fruits are consistently more tuberculate. The seeds are between<br />

1/8 and 3/16 of an inch (about 4 millimeters) in diameter.<br />

Range. From deep in central Mexico far north into the United<br />

States, having been reported from Texas, New Mexico, Oklahoma,<br />

and Arizona. The most northerly records from within our<br />

area, going from west to east, are as follows: Columbus, in<br />

southwest New Mexico; the hills west of San Antonio, New<br />

Mexico; Santa Rosa, New Mexico; Clayton, in the extreme<br />

northeast corner of New Mexico; then back to the Davis Moun-


238 cacti of the southwest<br />

tains of Texas and the lower Big Bend, except for one remarkable<br />

report of its having been found growing wild near Kingfisher<br />

in north-central Oklahoma.<br />

Remarks. O. kleiniae, while a tall-growing cholla, is an inconspicuous<br />

one. Its branches and stems are so long and slender and<br />

so weakly spined that from a distance it looks more like some<br />

leafless bush than a cactus. Many collectors no doubt pass right<br />

by it, thinking it is just another of the assorted desert shrubs<br />

with which it usually mingles in thickets. Few of them would<br />

care to take this cactus home with them anyway, since by either<br />

form or flower it has little to offer to a garden.<br />

If it grew over large areas of our country, this would certainly<br />

be one of the worst of the pest cacti, as it is in parts of Mexico,<br />

since, where it is found, it often makes large thickets of tall<br />

bushes which man or beast must avoid. Some of the creek flats<br />

in the Davis Mountains of west Texas are fairly choked with the<br />

cactus, and a large thicket grew, when I was last there, in a<br />

ditch in almost the center of Fort Davis, Texas. Here it was obvious<br />

that it was scraped away with each cleaning of the ditch,<br />

but it returned immediately from the roots with stems reaching<br />

4 or 5 feet tall in one season.<br />

It is fortunate that in the U. S. this potentially dangerous cactus<br />

is like so many of the chollas in occurring only here and<br />

there within its wide range. Although its range includes about<br />

half of New Mexico, in this huge area it has been found at only<br />

a dozen or so places, all of these more or less localized populations<br />

covering small areas. In Texas the same is true, the population<br />

in the Davis Mountains being the largest, with occasional<br />

thickets being found down around Presidio and up in the foothills<br />

of the Guadalupe Mountains.<br />

It is hundreds of miles from any of these locations to the<br />

location mentioned for the plant near Kingfisher, Oklahoma.<br />

There have never been any other reports of this species from<br />

Oklahoma. Were it not that this collection was made by Mr.<br />

Charles Polaski, the well-known cactus collector of Oklahoma<br />

City, who is a very careful observer and who has provided most<br />

of the recent specimens for European studies of Oklahoma forms,<br />

and that I have seen specimens collected at that site growing 6<br />

feet and more tall in Mr. Polaski’s garden, I would be skeptical,<br />

but apparently there is this isolated colony of the cactus far in<br />

north-central Oklahoma.<br />

There has not been much confusion over this cactus. Engelmann,<br />

overlooking De Candolle’s earlier description of it, renamed<br />

the species O. wrightii—this not to be confused with O.<br />

wrightiana (Baxter) Peebles, which is a low-growing, club cholla<br />

closely related to or a variety of O. stanlyi. Almost immediately<br />

the earlier name was recognized as the correct one, and I believe<br />

no one else has used Engelmann’s unfortunate name for the plant<br />

since.<br />

O. kleiniae is probably most closely related to O. arbuscula<br />

Eng., a cholla with similar slender, though much shorter branches<br />

from a much thicker trunk, which is therefore a much more<br />

compact, bushy shape when grown. The two have similar spines,<br />

but, so far as I know, O. arbuscula, which has yellowish flowers<br />

and green fruits, is an Arizona plant which has never been found<br />

in New Mexico, so we should have no problem with the two in<br />

our area.<br />

It may be noticed that this description gives a longer maximum<br />

spine length and larger maximum fruit size than is stated<br />

for O. kleiniae in most of the major works on cacti. The first<br />

few descriptions of the cactus do not give us either the length of<br />

the spines or the size of the fruits. Coulter seems to have been the<br />

first to state sizes for these, and his maximums are the smallest<br />

given by anyone—3/4 inch for the spines and about 5/8 inch for<br />

the fruits. Most others have increased his figures only slightly.<br />

Most of them give the maximum spine length as between 3/4 and<br />

1 inch, and the largest fruit size is stated variously as from 5/8 to<br />

1 inch long. Only Borg differs from the rest in this, and he gives<br />

the spine length as to 2 inches—a startling maximum, double<br />

that of any other authority, which at first looks doubtful.<br />

It is our conviction that stopping the study of cacti which<br />

enter the U. S. from Mexico at the north bank of the Rio Grande<br />

and drawing up our descriptions of such species from only the<br />

U.S. segment of the population, which may be only a northern<br />

clone, as is so often done, is not good practice and may be the<br />

cause for some of our confusing problems in cactus study. Here,<br />

in this cactus which ranges hundreds of miles into Mexico and<br />

occurs widely there as huge, dominant populations rather than<br />

in scattered small numbers as in the U. S. seems to be one place<br />

where more attention should have been paid to the Mexican<br />

specimens. It is significant that Borg, who was much more a<br />

student of Central and South American cacti than of the U.S.<br />

forms, gives a spine length for the species double that given by<br />

our students.<br />

With this in mind, I have observed this cactus carefully on<br />

numerous trips throughout Coahuila and as far south as San Luis<br />

Potosi, where it grows in great profusion. In Mexico I find it<br />

identical to our northern specimens in all respects except that<br />

the spine length is quite commonly to 11/2 inches and the fruit<br />

size often to 11/2 inches also. I have never been able to measure a<br />

spine on it 2 inches long, as Borg must have done, but must presume<br />

that since he is right in that the usually stated limits are<br />

too short, he may well be right in his new maximum also. The<br />

value of considering the Mexican population is shown in that it<br />

made me return to the U. S. situation with new questions about<br />

our long-accepted limits. I measured plants wholesale, and in the<br />

Davis Mountains, where the cactus grows most profusely in the<br />

U. S., I finally found a few examples equaling the best Mexican<br />

specimens I had seen in both spine and fruit size. I have not been<br />

able to find equally developed plants in New Mexico, where it<br />

seems the cactus is less successful and, therefore, less robust.<br />

While these are small and technical points, their importance<br />

may be greater than at first appears. Engelmann described another<br />

slender-stemmed cholla from “in mountains near El Paso.”<br />

He said it had a main spine 1 to 21/2 inches long. He at first


genus Opuntia 239<br />

thought it another new species and called it O. vaginata. In his<br />

earliest report of it he said it had pale yellowish flowers with a<br />

green tinge, which were 1/2 to 3/4 of an inch in diameter. Now<br />

these flowers would be exactly like those of O. leptocaulis, and<br />

perhaps because of this he said later that it might be a stout form<br />

of O. frutescens (his name for O. leptocaulis). Watson took him<br />

seriously on this and made the combination, O. leptocaulis var.<br />

vaginata.<br />

Whether or not Engelmann ever actually meant to submerge<br />

the form into the species O. leptocaulis is not clear, but seems<br />

unimportant in the light of later developments. What is important<br />

is that in later reports he retracts his description of the<br />

flower and says that the flower of O. vaginata is unknown. It is<br />

impossible to tell whether, without the earlier, erroneous concept<br />

of the yellow flower it would ever have occurred to him<br />

that there could be a stout form of O. leptocaulis, but what has<br />

happened is that all students since have either made this a variety<br />

of that species or considered it a synonym and ignored it.<br />

But what if the flower, which is unknown still, were purplish?<br />

Wouldn’t we then think of the cactus as a long-spined form of<br />

O. kleiniae instead? It is clear that while in several characters<br />

this cactus exceeds the usual limits of O. leptocaulis so that to<br />

include it in that species would require a redrawing of the species<br />

description, vegetatively, except for the spine length, the<br />

description of O. vaginata falls entirely within the limits of<br />

O. kleiniae. Even the longer spines, with our new knowledge of<br />

the length of spine sometimes attained in O. kleiniae, is no more<br />

of a problem here than it would be in combining this form with<br />

O. leptocaulis. The fruits were said to be small, yellowish, and<br />

strongly tuberculate, but they were within the range of size<br />

found easily on plants of both the other species. Both species<br />

sometimes have orange or yellow-orange fruits, but the smaller,<br />

less well developed fruits of O. kleiniae sometimes remain tuberculate,<br />

as was its ovary, while neither the fruits nor the ovaries<br />

of O. leptocaulis are ever seen distinctly tuberculate.<br />

This gives the idea of O. vaginata being an extra long-spined<br />

form of O. kleiniae somewhat the advantage, it seems to me.<br />

And here the fact that we now know O. kleiniae grows longer<br />

spines than before supposed becomes important. Now who can<br />

tell—may it not be that Engelmann merely had before him,<br />

from near El Paso, an O. kleiniae with spines yet a little longer<br />

than Borg’s?<br />

Attractive as this possibility is, it is impossible to know. It<br />

would seem to be essential to have living specimens to which to<br />

refer and from which especially to learn the characteristics of<br />

the flower, yet I know of no specimens which fit the description<br />

of this form collected since the type, and admit that it has eluded<br />

me. All of the herbarium specimens so labeled which I have examined,<br />

other than the type, have proved to be, like Lindheimer’s<br />

O. vaginata from New Braunfels, Texas, ordinary O. leptocaulis<br />

and have lacked the dimensions given for the species by Engelmann.<br />

In this situation it would be almost as great an assumption<br />

to call it a synonym or a variety of O. kleiniae as it has<br />

been on the part of Watson and others to place it in O. leptocaulis.<br />

Here is a place where there is real need of further work,<br />

and we hope by this discussion to clarify the problem which has<br />

been glossed over for too long and to stimulate someone to find<br />

the plant and place it properly.<br />

Opuntia leptocaulis DC<br />

“Desert Christmas Cactus,” “Slender Stem Cactus,” “Tasa-<br />

jillo,” “Aguijilla,” “Garrambullo”<br />

Description Plate 64<br />

stems: A small, upright bush 2 to sometimes 5 feet tall. It is<br />

usually compactly and extensively branched from a main<br />

trunk which, on old plants, becomes covered with scaly bark<br />

and grows to 1 to 11/4 inches in diameter. The current year’s<br />

joints are cylindrical and 1 to 12 inches long, but only 1/8 to<br />

1/4 of an inch thick. The surface of these joints may have indistinct<br />

tubercles 1/4 to 1/2 inch long, but are more often completely<br />

smooth. The color is deep green, often with purplish<br />

spots around the areoles. The lateral joints detach very easily.<br />

areoles: Ovate or often almost diamond-shaped. They are<br />

about 1/8 of an inch long, with short, white wool.<br />

spines: 0 to 3 per areole, but most commonly 1. The main<br />

spine is porrect, gray, and very variable. It may be 1/8 to 2<br />

inches long. When short, it is usually very slender with a<br />

close-fitting sheath. When long and well-developed, it is stout,<br />

more or less flattened, and covered with a loose, papery,<br />

white, yellow, or tan sheath. There may be 1 or 2 additional<br />

short, slender, bristle-like spines.<br />

glochids: Few and very short in 1 to 3 small bunches in the<br />

upper part of the areole. They are yellowish to brown in color.<br />

flowers: 1/2 to 7/8 of an inch wide by 3/4 to 1 inch tall, opening<br />

very wide. They are greenish-yellow in color. The ovary<br />

is about 5/8 of an inch long, egg-shaped to conical, slightly if<br />

at all tuberculate, with brown glochids and elongated perianth<br />

segments upon it. The outer segments are greenish-yellow with<br />

soft green spines at their summits. The inner segments are oblong<br />

and pointed. The stamens are greenish-yellow. The style<br />

is long, with 3 to 6 short, thick, greenish-yellow stigma lobes.<br />

fruits: Bright scarlet, orange-red, or yellowish when ripe.<br />

Almost globular, pear-shaped, or club-shaped, and 3/8 to 1<br />

inch long. The surface is smooth, often with brown glochids in<br />

the areoles and with a deeply pitted umbilicus. These fruits<br />

are persistent on the plant and often proliferous, sometimes<br />

having shoots 2 or 3 inches long from them while still in place<br />

on the branch. There are usually a dozen or less seeds per<br />

fruit, these 1/8 of an inch or a little more (3–4 millimeters) in<br />

diameter.<br />

Range. From south-central Mexico north into Arizona, over all<br />

but approximately the northwestern quarter of New Mexico,


240 cacti of the southwest<br />

over all of Texas south of the Canadian River east to the Dallas<br />

area and the lower Brazos River, and entering Oklahoma as far<br />

as Harmon and Greer counties in the extreme southwestern corner<br />

and the Arbuckle Mountains in the south-central part of the<br />

state.<br />

Remarks. This, probably the most hated cactus of all in our<br />

area, and surely the one with the least to endear it to us, is a<br />

very wide-ranging species and one which unfortunately grows<br />

in great numbers in much of its territory. No one who has ever<br />

carelessly brushed up against what he ignored as just another<br />

scrubby little bush, but which was really this cactus, will ever<br />

forget it. Its nefariousness lies not in its being so spiny—many<br />

other cacti far surpass it in that—but in that it is so uncactus-<br />

like that we often fail to give it the proper respect and so it<br />

pricks us worst of all. It likes to grow within other bushes, and<br />

reaches out stray branches to snare us as we pass. The Spanish<br />

names for it, which anyone who understands them will realize<br />

are not exactly terms of endearment, are most appropriate. The<br />

person who had charity enough to coin the name, desert christmas<br />

cactus, for it, after the brilliant fruits which do remain on<br />

the bushes and beautify the otherwise dull brush all winter, was<br />

a rare soul or else only saw it from the highway and never tried<br />

to walk between plants of the garrambullo.<br />

The cactus varies greatly in size in different situations, being<br />

often only 2 feet high, but in the southern part of our area, in<br />

good alluvial soil it sometimes becomes 5 feet tall. However,<br />

robust as it may grow, it does not increase the diameter of its<br />

terminal branches over about 1/4 of an inch nor of its trunk over<br />

about 11/2 inches. I believe this distinguishes it from O. arbuscula,<br />

a close relative found in Arizona, upon which I have seen<br />

old trunks over 4 inches in diameter. Fruit color also distinguishes<br />

these two. Another close relative sometimes found in the<br />

same thicket with this species is O. kleiniae. Although the trunks<br />

of that species do not get much larger, its current year’s stems<br />

do, and it also has larger, purplish flowers.<br />

The garrambullo was noticed early and has had its share of<br />

names assigned to it. The first and valid one is O. leptocaulis,<br />

given it by De Candolle in 1828. Afterward there followed various<br />

others, among them O. ramulifera SD and O. virgata Link,<br />

but the ones causing the most confusion were the array of different<br />

names used for it by Engelmann. At first he thought it was<br />

a variety of another cactus we have recently looked at, and so<br />

called it O. fragilis var. frutescens. He himself later realized the<br />

error in this, and so raised it to a separate species, but at the<br />

time he apparently did not know of De Candolle’s previous<br />

name, and so he burdened us with O. frutescens as a species<br />

name, which is of course superseded by the earlier one.<br />

But then Engelmann thought he saw consistent varieties in the<br />

species. He therefore described variety longispina and variety<br />

brevispina, the former with its spines 1 to 2 inches long and with<br />

a loose sheath, and the latter with its spines short, slender, and<br />

tightly sheathed. Coulter, later, when trying to get the names<br />

correct, only compounded the confusion by rechristening the<br />

long-spined variety O. leptocaulis var. stipata because of a supposedly<br />

consistent difference from the other in joint shape. Still<br />

later, Britton and Rose described what seems to be nothing but<br />

this same long-spined form as a new species, Opuntia mortolensis.<br />

There was also commonly listed an O. leptocaulis var. vaginata.<br />

This supposed taxon and the almost complete uncertainty<br />

concerning it is discussed in the remarks upon O. kleiniae, to<br />

which it seems to us much more closely linked. There followed<br />

variety badia, variety robustior, and variety pluriseta, all by<br />

Berger, and all for minor differences of spines, joints, or areoles.<br />

Most authors have long since dropped any attempt to maintain<br />

these varieties, and I agree that they seem only minor<br />

growth forms, probably environmental or else mere phenotypes.<br />

They are often found growing together, although in some areas<br />

one form does predominate and often only the weak-spined<br />

type is present at all. In numerous cases where I have watched<br />

over a period of several years, long-spined specimens have reverted<br />

to the short-spined, making it appear that we have here<br />

again the robust versus the stunted forms of the same plant, as<br />

we have seen so commonly in the Opuntias. At least there seem<br />

no separate ranges for these forms.<br />

O. leptocaulis is placed last in our account of the cacti. This<br />

has no significance except to indicate that it is probably last in<br />

the interest of cactophiles. Many would say it should have been<br />

first in our account, since it seems probably the most primitive<br />

of the cacti in our area. Be that as it may, the cactus is a major<br />

pest cholla, and hardly the form to use in introducing the cacti<br />

to anyone. So we place it here at the end of our account. Whether<br />

it is thought of as the unpleasant thing from which others so<br />

beautiful and fascinating arose or as an example of how degenerate<br />

a cactus can become, it is the poor relation of all the beau-<br />

tiful ones, and it cannot be left out.


GLOSSARY<br />

The terms below are defined as they are used in this work<br />

to refer to the cacti described, and the definitions are not<br />

intended to be so broad as to cover their usage for all other<br />

plant groups.<br />

Anther. The enlarged, pollen-bearing sac at the tip of a stamen.<br />

Apex. The tip or summit of any structure.<br />

Apical. Referring to the apex.<br />

Areole. A spot in the form of a pit or a raised area marking an<br />

opening through the epidermis from which leaves, spines, or other<br />

structures grow.<br />

Ascending. Not standing perfectly upright, but growing upward.<br />

Axil. The angle between a leaf, branch, tubercle, or other outgrowth<br />

and the stem.<br />

Basal. At or referring to the base or lower part of any structure.<br />

Berry. A pulpy or fleshy fruit with numerous seeds embedded in the<br />

flesh.<br />

Bud. An unopened flower; a growing tip surrounded by its immature<br />

perianth segments or leaves.<br />

Caespitose. Forming a cluster or clump of stems by repeated branch-<br />

ing of the stem at or near the base.<br />

Calyx. The outer series of perianth segments, whenever these are distinct<br />

from the inner series.<br />

Central. Positioned at or near the center of an area, as opposed to<br />

being peripheral in position. A spine originating in the center of the<br />

areole as opposed to those growing around the edge of the areole.<br />

Character. A characteristic or feature unique enough to have value<br />

in distinguishing forms and setting up relationships.<br />

Cholla. Any cylindrical-stemmed member of the genus Opuntia.<br />

Cilium (pl.: cilia). Very fine, hairlike filaments sometimes forming<br />

fringes on the margins of perianth segments.<br />

Clone. A local population usually propagated from one individual by<br />

vegetative means and therefore uniform genetically and in appear-<br />

ance.<br />

Confluent. Running together or more or less coalescing.<br />

Corolla. The inner series of perianth segments, when these are distinct<br />

from the outer ones; the petals collectively.<br />

Decumbent. Lying prostrate on the ground with the tip turning upward.<br />

Deflexed. Curved or bent back, down upon itself, or toward the surface<br />

of the plant. Recurved.<br />

Dehiscent. Splitting open at maturity.<br />

Dimorphic. Having two forms.<br />

Distal. Situated opposite the point of attachment or origin.<br />

Divergent. Spreading apart so as to form opposites.<br />

Ecotype. A population which is recognizable by distinct morphological<br />

and physiological features and which is the result of, but kept<br />

separate from its near relatives by environmental barriers; an ecological<br />

“race.”<br />

Entire. Having the margin continuous and not toothed, lobed, indent-<br />

ed or interrupted.<br />

Epidermis. The outer layer of cells on a plant, forming a protective<br />

covering. In ordinary usage thought of as including the waxy non-<br />

living layer that overlies the living cells.<br />

Erose. Ragged, with irregular indentations, as though bits were randomly<br />

chewed away.<br />

Felt. A very thick covering of hairs, filaments, or fibers.<br />

Fibrous roots. Finely subdivided roots with no obvious thickening or<br />

enlarged central root beyond the base of the plant.<br />

Filament. The stalk of a stamen; the threadlike part of the stamen<br />

that supports the anther.<br />

Genotype. A group of organisms having a common genetic makeup.<br />

Genus. A grouping of species possessing common characters unique<br />

enough to be treated as a unit distinct from others.<br />

Glabrous. Smooth and shiny; not pubescent, rough, or hairy.<br />

Gland. A secreting structure, usually in the form of a protuberance or<br />

appendage, but sometimes a surface.<br />

Glaucescence. A thin layer of whitish substance, often called the<br />

“bloom” and usually made up of tiny particles of wax, which rub<br />

off.


242 cacti of the southwest<br />

Globose. Spherical or spheroidal in shape.<br />

Glochid. A sharp, hairlike or bristlelike outgrowth equipped with<br />

minute and usually invisible barbs so that it resists withdrawal from<br />

any tissue.<br />

Hilum. The scar on the seed that marks the point at which the seed<br />

was attached during growth.<br />

Inferior. Beneath or below; in the flower, descriptive of having the<br />

ovary appearing below such flower appendages as the perianth and<br />

stamens.<br />

Intergrade. Not separated by any division into sets, but merging by<br />

having an unbroken series of intermediate forms.<br />

Lanceolate. Lance-shaped; much longer than broad, widest just above<br />

the base, with a gradual taper from there to the tip.<br />

Lateral. Growing or positioned at the sides.<br />

Linear. Long and narrow, with sides parallel or nearly so, as a blade<br />

of grass.<br />

Meristem. A body of tissue with the power to divide and differentiate.<br />

Microgenus. A nonofficial term used here for genera that have been<br />

officially described, but seem distinguished by less obvious or significant<br />

characters than the major genera.<br />

Midrib. The main or center rib of a leaf or perianth segment.<br />

Monomorphic. Having but a single form and not subdividing.<br />

Mucro. A short, abrupt, and more or less sharp point on the tip of a<br />

leaf or flower structure.<br />

Oblanceolate. Reversed lanceolate in shape, with the widest part near<br />

the tip.<br />

Obovate. Reversed ovate; the outline of a hen’s egg with the broader<br />

part above the middle.<br />

Ovary. The enlarged lower part of the pistil containing the ovules.<br />

Ovate. Having the outline of a hen’s egg, and with the broader part<br />

below the middle.<br />

Pectinate. Comblike; used in referring to spines or other structures<br />

spread flat like the teeth of a comb.<br />

Perianth segment. One of the parts making up the perianth; a petal<br />

or sepal.<br />

Perianth tube. A tubelike arrangement of the perianth segments extending<br />

in some forms to some distance above the ovary.<br />

Petal. One of the inner set of perianth segments or corolla.<br />

Phenotype. A group of organisms recognized by their common visible<br />

characters irrespective of their genetic composition.<br />

Pistil. The central, female part of the flower, made up of the ovary,<br />

style, and stigma.<br />

Porrect. Positioned outward; standing perpendicular to the surface<br />

of the plant.<br />

Prostrate. Lying completely flat upon the ground.<br />

Pubescent. Covered with short, soft, fine, hairlike outgrowths; downy.<br />

Ramose. Branching or having branches.<br />

Radial. Positioned around the edges of an area; peripheral, as opposed<br />

to central. A spine positioned somewhere upon the periphery<br />

of an areole.<br />

Radiating. Positioned outward like radii.<br />

Reclining. Sprawling or leaning against something.<br />

Recurved. Curving back upon itself; deflexed.<br />

Rib. A ridge; a raised surface running vertically or sometimes spiraling,<br />

and bearing areoles in a row along its summit. Often<br />

thought of as being composed of more or less coalescent tubercles<br />

which may be evident as bulging masses along it.<br />

Rotate flower. Spreading widely rather than remaining bell-shaped<br />

or funnel-shaped; wheel-shaped.<br />

Scale. A small, scarcely expanded leaflike structure; also a narrow,<br />

triangular, or sometimes spinelike lower perianth segment often<br />

found on the ovary or flower-tube surface.<br />

Sepal. One of the outer perianth segments of the calyx.<br />

Spatulate. Spoon-shaped; oblong or rounded above, the base long and<br />

narrow.<br />

Species. A population recognizable by characteristics of form and<br />

genetic relationship as a unit, often capable of subdivision into<br />

varieties all genetically close enough for interbreeding.<br />

Spicule. A very small, fine spine. A glochid.<br />

Spine. A sharp outgrowth, either rigid and woody or sometimes flexible<br />

and hairlike. In cacti always arising from an areole and thought<br />

of as a modified leaf.<br />

Spreading. Growing outwardly. Used in referring to a series of radial<br />

spines projecting obliquely outward around the areole, as opposed<br />

to lying flat against the surface of the plant. Used also for plant<br />

growth advancing outward by new shoots or by rooting from old,<br />

reclining stems.<br />

Stamen. The male part of the flower consisting of the filament and<br />

the anther bearing the pollen.<br />

Stem. The main upward axis of a plant.<br />

Stigma. The uppermost part of the pistil, at the tip of the style; the<br />

part that receives the pollen; in cacti usually divided into lobes.<br />

Stigma lobe. One of the expanded sections of the stigma.<br />

Stoma (pl.: stomata). A microscopic opening through the epidermis<br />

of the plant allowing for respiration and transpiration.<br />

Style. The central portion of the pistil, connecting ovary and stigma.<br />

Subgenus. A taxonomic rank sometimes used to divide a large genus<br />

into subdivisions above the species level.<br />

Subspecies. A subdivision of the species unit recognizable by certain<br />

morphological characters, but not isolated by genetic barriers from<br />

others within the species. (No attempt is made in this work to distinguish<br />

between the subspecies and the variety.)<br />

Taproot. The primary root axis, when larger and longer than the<br />

branch roots, and often thick and used for storage, as a carrot.<br />

Taxon. A formally described category in classification; a series of individuals<br />

distinct by some visible characteristics.<br />

Terminal. At the tip or end.<br />

Transpiration. ‘The giving off of water vapor through the stomata by<br />

a plant.<br />

Trichome. A hairlike structure found on plants; a slender filament<br />

growing from the plant’s epidermis by one end.<br />

Tuber. A short, thick, fleshy underground stem or stem branch for<br />

storage and having buds. The potato is an example.<br />

Tubercle. A knoblike protrusion from the surface of any structure;<br />

a more or less pyramidal knob rising from the stem surface of a<br />

cactus and having an areole on or near its summit.<br />

Tuberous root. A root of undistended size overall, a generally fibrous<br />

root, but having thick, fleshy sections like tubers scattered upon it.<br />

Not a taproot.<br />

Tuberculate. Having tubercles.<br />

Umbilicus. On those species which drop the perianth and upper parts<br />

of the flower, the scar left at the summit of the fruit after the<br />

floral parts are shed.<br />

Variety. See Subspecies.<br />

Woolly. Covered with long and very thick hairs.


princips -><br />

princeps<br />

Cornyopuntia<br />

Cf. Corynopuntia<br />

leptocanthus -><br />

leptacanthus<br />

INDEX <strong>OF</strong> SCIENTIFIC NAMES<br />

The number of the page on which the principal description of a form begins is in<br />

italics. The number of the plate where the illustration of the plant appears is in<br />

parentheses. All other numbers refer to pages on which there is incidental reference<br />

to the plant.<br />

Acanthocereus: 8, 57, 60<br />

Acanthocereus pentagonus: 60, 61, 62, (18)<br />

Acanthocereus tetragonus: 62<br />

Ancistrocactus: 64, 76, 77, 78, 79, 80<br />

Ancistrocactus brevihamatus: 76<br />

Ancistrocactus scheeri: 77<br />

Ancistrocactus tobuschii: 78, 79<br />

Anhalonium: 100<br />

Anhalonium engelmannii: 102<br />

Anhalonium lewinii: 98<br />

Ariocarpus: 9, 63, 100, 101, 102, 107, 108<br />

Ariocarpus fissuratus: 101, 102, (26)<br />

Ariocarpus fissuratus var. lloydii: 102<br />

Ariocarpus retusus: 101<br />

Ariocarpus trigonus: 101<br />

Astrophytum: 64, 70<br />

Astrophytum asterias: 70<br />

Brittonia: 85<br />

Brittonia davisii: 85<br />

Cactus: 110<br />

Cactus compressus: 202<br />

Cactus conoideus: 93<br />

Cactus ferox: 222<br />

Cactus glomeratus: 148<br />

Cactus humifusus: 202<br />

Cactus mammillaris: 123<br />

Cactus missouriensis var. similis: 123<br />

Cactus neo-mexicanus: 129<br />

Cactus opuntia: 202<br />

Cactus opuntia var. inermis: 167<br />

Cactus pentagonus: 61, 62<br />

Cactus pitajaya: 61<br />

Cactus proliferus: 148<br />

Cactus pusillus: 148<br />

Cactus similis: 123<br />

Cactus strictus: 167<br />

Cactus tetragonus: 62<br />

Cactus viviparus: 126<br />

Cereus: 55, 57, 60, 110<br />

Cereus acutangulus: 61<br />

Cereus aggregatus: 41<br />

Cereus baxaniensis: 61<br />

Cereus conoideus: 42<br />

Cereus dussii: 61<br />

Cereus leptacanthus: 51<br />

Cereus nitidus: 61<br />

Cereus octacanthus: 39<br />

Cereus pectinatus centralis: 89<br />

Cereus pentalophus: 51<br />

Cereus phoeniceus: 41, 130<br />

Cereus phoeniceus var. conoideus: 42<br />

Cereus poselgeri: 56<br />

Cereus princeps: 61<br />

Cereus prismaticus: 61<br />

Cereus procumbens: 51<br />

Cereus roemeri: 39, 42<br />

Cereus roetteri: 34<br />

Cereus runyonii: 51<br />

Cereus sirul: 61<br />

Cereus tuberosus: 56<br />

Cereus undulosus: 61<br />

Cereus variabilis: 61<br />

Cereus viridiflorus var. tubulosus: 15<br />

Clavatae: 161<br />

Clavatopuntia: 161, 227<br />

Coloradoa: 64, 75<br />

Coloradoa mesae-verdae: 75<br />

Coryphantha: 103, 110, 111, 112, 140, 141, 144<br />

Coryphantha aggregata: 130<br />

Coryphantha arizonica: 131<br />

Coryphantha clava: 111, 112<br />

Coryphantha conoidea: 93<br />

Coryphantha cornifera: 116<br />

Coryphantha deserti: 131<br />

Coryphantha echinus: 116<br />

Coryphantha engelmannii: 115<br />

Coryphantha erecta: 111, 112<br />

Coryphantha fragrans: 132<br />

Coryphantha hesteri: 139<br />

Coryphantha macromeris: 120<br />

Coryphantha minima: 139,140<br />

Coryphantha muehlenpfordtii: 114, 115<br />

Coryphantha muehlenpfordtii<br />

var. robustispina: 115<br />

Coryphantha nellieae: 139, 140<br />

Coryphantha neo-mexicana: 128, 129, 130<br />

Coryphantha piercei: 141<br />

Coryphantha pirtlei: 121<br />

Coryphantha pottsii: 144<br />

Coryphantha radiosa: 127, 128, 130, 131<br />

Coryphantha ramillosa: 119<br />

Coryphantha roberti: 140, 141<br />

Coryphantha runyonii: 121, 141<br />

Coryphantha scheeri: 114, 115<br />

Coryphantha scolymoides: 116<br />

Coryphantha sulcata: 118<br />

Coryphantha varicolor: 138<br />

Coryphantha vivipara: 126<br />

Coryphantha vivipara var. aggregata: 130<br />

Coryphantha vivipara var. vivipara: 126<br />

Corynopuntia: 161, 184, 217, 227<br />

Curassavicae: 217<br />

Cylindropuntia: 161, 227<br />

Dolichothele: 110, 157<br />

Dolichothele sphaerica: 156<br />

Ebnerella: 145<br />

Echinocactus: 9, 63, 64, 65, 69, 72, 73, 74, 75,<br />

76, 77, 78, 79, 80, 85, 87, 95, 106, 107<br />

Echinocactus arizonicus: 72<br />

Echinocactus asterias: 66, 70, (19)<br />

Echinocactus bicolor: 86, 87<br />

Echinocactus bicolor var. schottii: 66, 85, 86,<br />

87, (23)<br />

Echinocactus bicolor var. texensis: 86<br />

Echinocactus bicolor var. tricolor: 86<br />

Echinocactus brevihamatus: 66, 76, 78, 79,<br />

(21)<br />

Echinocactus conoideus: 66, 93, 94, 135, (25)<br />

Echinocactus emoryi: 72<br />

Echinocactus equitans: 67<br />

Echinocactus erectocentrus: 91<br />

Echinocactus erectocentrus var. pallidus: 66,<br />

90, 91, (24)<br />

Echinocactus falconeri: 72<br />

Echinocactus flavidispinus: 66, 87, (23)<br />

Echinocactus flexispinus: 85<br />

Echinocactus grusonii: 65<br />

Echinocactus hamatacanthus: 66, 83, 84, 85,<br />

(23)<br />

Echinocactus hamatacanthus var.<br />

brevispinus: 85<br />

Echinocactus hamatacanthus var.<br />

crassispinus: 85<br />

Echinocactus hamatacanthus var.<br />

gracilispinus: 85<br />

Echinocactus horizonthalonius: 65, 66, 67,<br />

69, 115, (18), (19)<br />

Echinocactus horizonthalonius var.<br />

centrispinus: 67<br />

Echinocactus horizonthalonius var.<br />

curvispina: 65, 67, 68, (18)<br />

Echinocactus horizonthalonius var.<br />

moelleri: 65, 68, (19)<br />

Curassauicae -><br />

Curassavicae<br />

scheerii -><br />

scheeri


scheerii -> scheeri<br />

244 cacti of the southwest<br />

Echinocactus ingens: 6<br />

Echinocactus intertextus: 88, 89, (24)<br />

Echinocactus intertextus var. dasyacanthus:<br />

66, 90, 132, (24)<br />

Echinocactus intertextus var. intertextus: 66,<br />

89, 90, (24)<br />

Echinocactus lindheimeri: 69<br />

Echinocactus longihamatus: 84<br />

Echinocactus longihamatus var. sinuatus: 83<br />

Echinocactus mariposensis: 66, 92, (25)<br />

Echinocactus megarhizus: 78<br />

Echinocactus mesae-verdae: 65, 66, 75, (21)<br />

Echinocactus micromeris: 107<br />

Echinocactus muehlenpfordtii: 115<br />

Echinocactus platycephalus: 69<br />

Echinocactus pottsianus: 144<br />

Echinocactus reichenbachii: 20, 21<br />

Echinocactus scheeri: 66, 76, 77, 78, (21)<br />

Echinocactus setispinus: 79, 80, 83, 84, 85,<br />

(22)<br />

Echinocactus setispinus var. cachetianus: 82<br />

Echinocactus setispinus var. hamatus: 65, 80,<br />

81, 82, (22)<br />

Echinocactus setispinus var. micrensis: 82<br />

Echinocactus setispinus var. robustus: 83<br />

Echinocactus setispinus var. setaceus: 65, 81,<br />

82, (22)<br />

Echinocactus setispinus var. sinuatus: 83<br />

Echinocactus similis: 123<br />

Echinocactus simpsonii: 104<br />

Echinocactus sinuatus: 66, 80, 82, 83, 84, (23)<br />

Echinocactus texensis: 66, 68, 69, 125, (19)<br />

Echinocactus texensis var. gourgensii: 69<br />

Echinocactus texensis var. longispinus: 69<br />

Echinocactus tobuschii: 66, 78, 79, (22)<br />

Echinocactus uncinatus: 64, 73, 80<br />

Echinocactus uncinatus var. wrightii: 65, 72,<br />

73, (20)<br />

Echinocactus victoriensis: 70<br />

Echinocactus whipplei: 66, 74, (20)<br />

Echinocactus wislizeni: 66, 71, 85, (20)<br />

Echinocactus wislizeni var. decipiens: 72<br />

Echinocereus: 8, 10, 55<br />

Echinocereus albispinus: 11, 28, (6)<br />

Echinocereus baileyi: 11, 21, 25, 26, 27, 28,<br />

(6)<br />

Echinocereus baileyi var. albispinus: 27, 28<br />

Echinocereus baileyi var. brunispinus: 27<br />

Echinocereus baileyi var. caespitosus: 27<br />

Echinocereus baileyi var. flavispinus: 27<br />

Echinocereus baileyi var. roseispinus: 27<br />

Echinocereus berlandieri: 13, 19, 52, 53, 54,<br />

(17)<br />

Echinocereus blanckii: 13, 53, 54, (17)<br />

Echinocereus caespitosus: 19, 20, 21, 22, 23,<br />

24, 25, 26, 27, 28, 29, 30, 40, 149, 230, (3)<br />

Echinocereus caespitosus var. albiflora: 22<br />

Echinocereus caespitosus var. aureiflora: 22<br />

Echinocereus caespitosus var. caespitosus: 11,<br />

22, (3), (7)<br />

Echinocereus caespitosus var. castaneus: 21<br />

Echinocereus caespitosus var. minor: 11, 22,<br />

23, (4)<br />

Echinocereus caespitosus var. perbellus: 11,<br />

23, (4)<br />

Echinocereus caespitosus var. purpureus: 11,<br />

23, (5)<br />

Echinocereus chisoensis: 11, 29, (7)<br />

Echinocereus chloranthus: 14, 15, 16, 17, 18,<br />

19, (2)<br />

Echinocereus chloranthus var. chloranthus:<br />

11, 17, (2)<br />

Echinocereus chloranthus var. neocapillus:<br />

11, 17, 18, (2)<br />

Echinocereus coccineus: 39, 41, 42, 43, 47,<br />

130, (12), (13)<br />

Echinocereus longispinus: 27<br />

Echinocereus mariae: 27<br />

Echinocereus melanocentrus: 11, 24, 25, (5)<br />

Echinocereus mojaviensis: 12, 39<br />

Echinocereus neo-mexicanus: 32, 44<br />

Echinocereus octacanthus: 39<br />

Echinocereus oklahomensis: 27<br />

Echinocereus papillosus: 49, 50, (16)<br />

Echinocereus papillosus var. angusticeps: 13,<br />

50, (16)<br />

Echinocereus papillosus var. papillosus: 13,<br />

50, (16), (17)<br />

Echinocereus papillosus var. rubescens: 50<br />

Echinocereus paucispinus: 40, 41<br />

Echinocereus pectinatus: 21, 22, 25, 29, 30,<br />

31, 32, 33, 34<br />

Echinocereus pectinatus centralis: 89<br />

Echinocereus pectinatus var. ctenoides: 12,<br />

30, 31, (7), (8)<br />

Echinocereus pectinatus var. neo-mexicanus:<br />

33<br />

Echinocereus pectinatus var. rigidissimus: 12,<br />

30, 31, (7)<br />

Echinocereus pectinatus var. rubescens: 22<br />

Echinocereus pectinatus var. steereae: 32<br />

Echinocereus pectinatus var. texana: 30<br />

Echinocereus pectinatus var. wenigeri: 12,<br />

29, (7)<br />

Echinocereus pentalophus: 13, 51, 52, 54,<br />

(16)<br />

Echinocereus pentalophus var. leptacanthus:<br />

51<br />

Echinocereus pentalophus var. propinquus:<br />

51, 53<br />

Echinocereus pentalophus var. radicans: 51<br />

Echinocereus pentalophus var. simplex: 51,53<br />

Echinocereus pentalophus var. subarticulatus:<br />

51<br />

Echinocereus perbellus: 21<br />

Echinocereus polyacanthus: 43, 44<br />

Echinocereus polyacanthus var. neo-<br />

mexicanus: 12, 44, (13)<br />

Echinocereus polyacanthus var. polyacanthus:<br />

12<br />

Echinocereus polyacanthus var. rosei: 12, 43,<br />

44, (13)<br />

Echinocereus poselgeri: 56<br />

Echinocereus procumbens: 51<br />

Echinocereus purpureus: 28<br />

Echinocereus reichenbachii: 21, 27<br />

Echinocereus roemeri: 40<br />

Echinocereus roetteri: 12, 33, 34, (10)<br />

Echinocereus roetteri var. lloydii: 34, (11)<br />

Echinocereus rosei: 44<br />

Echinocereus russanthus: 11, 18, 19, (3)<br />

Echinocereus steereae: 32<br />

Echinocereus stramineus: 12, 46, 47, 48, (14)<br />

Echinocereus triglochidiatus: 35, 36, 37, 40,<br />

236, (11), (12)<br />

Echinocereus triglochidiatus var.<br />

gonacanthus: 12, 37, 39, 40, (12)<br />

Echinocereus triglochidiatus var. hexaedrus:<br />

12, 40<br />

Echinocereus triglochidiatus var.<br />

melanacanthus: 41<br />

Echinocereus coccineus var. coccineus: 12, 42,<br />

(12)<br />

Echinocereus coccineus var. conoideus: 12,<br />

42, 43, (13)<br />

Echinocereus coccineus var. octacanthus:<br />

38, 39<br />

Echinocereus conglomeratus: 47<br />

Echinocereus dasyacanthus: 25, 30, 31, 32, 33,<br />

34, (9)<br />

Echinocereus dasyacanthus var. ctenoides: 31<br />

Echinocereus dasyacanthus var.<br />

dasyacanthus: 12, 33, (9)<br />

Echinocereus dasyacanthus var. hildmanii:<br />

12, 33, (9)<br />

Echinocereus dasyacanthus var. minor: 33, 34<br />

Echinocereus dasyacanthus var.<br />

neo-mexicanus: 32, 33<br />

Echinocereus dasyacanthus var. steereae: 32<br />

Echinocereus davisii: 11, 15, 16, (2)<br />

Echinocereus dubius: 13, 47, 48, (15)<br />

Echinocereus engelmannii: 48<br />

Echinocereus enneacanthus: 40, 44, 45, 47,<br />

(14)<br />

Echinocereus enneacanthus var. carnosus: 12,<br />

46, 42, (14)<br />

Echinocereus enneacanthus var.<br />

enneacanthus: 12, 45, 46, (14)<br />

Echinocereus enneacanthus var. major: 46<br />

Echinocereus fendleri: 29, 48, 49, (15)<br />

Echinocereus fendleri var. bonkerae: 34<br />

Echinocereus fendleri var. fendleri: 12, 49,<br />

(15)<br />

Echinocereus fendleri var. pauperculus: 49<br />

Echinocereus fendleri var. rectispinus: 12, 49,<br />

(15)<br />

Echinocereus fitchii: 11, 25, (5)<br />

Echinocereus gonacanthus: 38<br />

Echinocereus lloydii: 12, 34<br />

Echinocereus longisetus: 19, 26<br />

Echinocereus triglochidiatus var.<br />

octacanthus: 12, 37, 39, 40, 41, 42, 43, (12)<br />

Echinocereus triglochidiatus var.<br />

polyacanthus: 44<br />

Echinocereus triglochidiatus var.<br />

triglochidiatus: 12, 37, 38, 40 (11)<br />

Echinocereus tuberosus: 56<br />

Echinocereus viridiflorus: 13, 14, 15, 16, 17,<br />

19, 31, (1)<br />

Echinocereus viridiflorus var. viridiflorus: 17<br />

Echinocereus viridiflorus var. cylindricus:<br />

11, 14, 15, 17, 18, 19, 31(1)<br />

Echinocereus viridiflorus var. davisii: 16<br />

Echinocereus viridiflorus var. minor: 14


index of scientific names 245<br />

Echinocereus viridiflorus var. standleyi: 11,<br />

15, (1)<br />

Echinocereus viridiflorus var. viridiflorus:<br />

11, 14, 15, 17, (1)<br />

Echinomastus: 64, 73, 89, 93<br />

Echinomastus dasyacanthus: 90<br />

Echinomastus intertextus: 88<br />

Echinomastus intertextus var. dasyacanthus:<br />

90<br />

Echinomastus johnsonii: 64, 72<br />

Echinomastus mariposensis: 92<br />

Echinomastus pallidus: 90, 91<br />

Echinopsis: 39<br />

Echinopsis octacanthus: 39<br />

Echinopsis pectinata var. reichenbachiana: 20<br />

Eopuntia: 3<br />

Eopuntia douglassii: 3<br />

Epithelantha: 9, 107, 108<br />

Epithelantha micromeris: 108, 109, (27)<br />

Epithelantha micromeris var. greggii: 108<br />

Escobaria: 110, 111, 112, 138, 140, 141, 144<br />

Escobaria albicolumnaria: 137<br />

Escobaria bella: 143<br />

Escobaria dasyacantha: 135, 137<br />

Escobaria leei: 142, 143<br />

Escobaria nellieae: 140<br />

Escobaria orcuttii: 135<br />

Escobaria runyonii: 140, 141<br />

Escobaria sneedii: 141<br />

Escobaria tuberculosa: 133, 134, 144<br />

Escobaria varicolor: 138<br />

Escobesseya: 111, 136, 137<br />

Escobesseya dasyacantha: 136<br />

Escobesseya duncanii: 136<br />

Eu-coryphantha: 110<br />

Eumammillaria: 110<br />

Euphorbia: 3, 111, 161<br />

Euphorbia grandicornis: 161<br />

Euphorbia obesa: 161<br />

Euphorbia prostrata: 161<br />

crassahamatus Ferocactus: 64, 72, 73, 80, 85<br />

-> crassihamatus Ferocactus crassihamatus: 72<br />

Ferocactus hamatacanthus: 72, 83<br />

Ferocactus johnsonii: 64, 72<br />

Ferocactus rafaelensis: 70<br />

Ferocactus uncinatus: 64, 72<br />

Ferocactus wislizeni: 71, 72<br />

Frailea: 71<br />

Glandulicactus: 64, 65, 72, 74<br />

Glandulicactus uncinatus: 74<br />

Glandulicactus uncinatus var. wrightii: 72<br />

Gymnocactus: 93<br />

Hamatocactus: 64, 72, 74, 80, 85<br />

Hamatocactus hamatacanthus: 83, 84<br />

Hamatocactus hamatacanthus var. davisii: 85<br />

Hamatocactus setispinus: 79, 80<br />

Hamatocactus setispinus var. hamatus: 80<br />

Hamatocactus setispinus var. setaceus: 81<br />

Hamatocactus sinuatus: 82<br />

Hamatocactus uncinatus: 74<br />

Homalocephala: 64, 65, 69<br />

Homalocephala horizonthalonius: 69<br />

Homalocephala texensis: 65, 69<br />

Krainzia: 157<br />

Lepidocoryphantha: 112, 119, 120<br />

Lepidoptera: 209<br />

Lophophora: 9, 63, 95, 97, 107, 108<br />

Lophophora williamsii: 97, 99, (15), (26)<br />

Lophophora williamsii var. caespitosa: 98<br />

Lophophora williamsii var. echinata: 98, (26)<br />

Lophophora williamsii var. lewinii: 98<br />

Lophophora williamsii var. lutea: 98<br />

Lophophora williamsii var. pluricostata: 98<br />

Lophophora williamsii var. texana: 98<br />

Lophophora williamsii var. typica: 98<br />

Lophophora williamsii var. williamsii: 98,<br />

(26)<br />

Mammillaria: 9, 93, 100, 101, 103, 107, 110,<br />

111, 112, 115, 121, 141, 157<br />

Mammillaria aggregata: 41, 130, 131<br />

Mammillaria albicolumnaria: 114, 137, 138,<br />

(36)<br />

Mammillaria alversonii: 132<br />

Mammillaria bella: 113, 143<br />

Mammillaria borealis: 105<br />

Mammillaria calcarata: 119<br />

Mammillaria celsiana: 115<br />

Mammillaria conoidea: 93<br />

Mammillaria cornifera: 116<br />

Mammillaria cornifera var. scolymoides: 116<br />

Mammillaria dactylothele: 121<br />

Mammillaria dasyacantha: 114, 134, 135,<br />

136, 137, 138, (34), (35)<br />

Mammillaria deserti: 132<br />

Mammillaria diaphanacantha: 93<br />

Mammillaria duncanii: 114, 136, (35)<br />

Mammillaria echinocactoides: 93<br />

Mammillaria echinus: 113, 116, 117, 118, (28)<br />

Mammillaria elongata: 140<br />

Mammillaria engelmannii: 115<br />

Mammillaria fissurata: 102<br />

Mammillaria fragrans: 114, 132, 133, 137,<br />

(33)<br />

Mammillaria grahami: 149,150<br />

Mammillaria gummifera: 155,156<br />

Mammillaria hesteri: 113, 139, (37)<br />

Mammillaria heyderi: 112,152,153, 154, 156,<br />

(42), (43)<br />

Mammillaria heyderi var. applanata: 112,<br />

153, 154, 155, (42)<br />

Mammillaria heyderi var. hemisphaerica:<br />

112, 154, 155, (43)<br />

Mammillaria heyderi var. heyderi: 112, 153,<br />

154, (42)<br />

Mammillaria heyderi var. waltheri: 153<br />

Mammillaria inconspicua; 93<br />

Mammillaria lasiacantha: 145, (39), (40)<br />

Mammillaria lasiacantha var. denudata: 114,<br />

145, 146, (39), (40)<br />

Mammillaria lasiacantha var. lasiacantha:<br />

114, 145, 146, (39)<br />

Mammillaria lasiacantha var. minor: 146<br />

Mammillaria leei: 114, 142, 143, (39)<br />

Mammillaria leona: 144<br />

Mammillaria longimamma: 157<br />

Mammillaria macdougalii: 155,156<br />

Mammillaria macromeris: 113, 119, 120, 121,<br />

143, (30)<br />

Mammillaria meiacantha: 112, 152, 155, 156,<br />

(43)<br />

Mammillaria melanocentra: 156<br />

Mammillaria melanocentra var. meiacantha:<br />

156<br />

Mammillaria microcarpa: 113, 149, 150, (40)<br />

Mammillaria micromeris: 107<br />

Mammillaria milleri: 150<br />

Mammillaria minima; 140<br />

Mammillaria missouriensis: 123<br />

Mammillaria missouriensis var. robustior: 123<br />

Mammillaria missouriensis var. similis: 123<br />

Mammillaria muehlenpfordtii: 115,119<br />

Mammillaria multiceps: 114, 147, 148, (40)<br />

Mammillaria nellieae: 15, 114, 139, 140, (37)<br />

Mammillaria nuttallii: 123<br />

Mammillaria nuttallii var. borealis: 123<br />

Mammillaria nuttallii var. caespitosa: 123<br />

Mammillaria nuttallii var. robustior: 123<br />

Mammillaria pectinata: 116, 117, 118<br />

Mammillaria phellosperma: 150<br />

Mammillaria pottsii: 114, 143, 144, (39)<br />

Mammillaria prolifera: 148<br />

Mammillaria prolifera var. multiceps: 148<br />

Mammillaria prolifera var. texana: 148<br />

Mammillaria pusilla var. texana: 148<br />

Mammillaria purpusii: 104<br />

Mammillaria radians: 116<br />

Mammillaria radiosa: 128<br />

Mammillaria ramillosa: 113, 119, (29)<br />

Mammillaria roberti: 114, 140, 141, (38)<br />

Mammillaria robustispina: 115,116<br />

Mammillaria rosiflora: 112, 124, 125<br />

Mammillaria runyonii: 113, 121, 122, (30)<br />

Mammillaria scheeri: 93, 113, 114, 115, 116,<br />

(28)<br />

Mammillaria scheeri var. valida: 115<br />

Mammillaria scolymoides: 113, 116, (28)<br />

Mammillaria sheldonii: 150<br />

Mammillaria similis: 112, 119, 122, 123, 124,<br />

125, (30), (31)<br />

Mammillaria similis var. robustior: 119, 123,<br />

124<br />

Mammillaria simpsonii: 104<br />

Mammillaria sneedii: 114, 141, 142, 143, (38)<br />

Mammillaria sphaerica: 112, 156, 157, (44)<br />

Mammillaria strobiliformis: 93, 134, 135<br />

Mammillaria sulcata: 113, 118, 119, 120, 124,<br />

135, (28)<br />

Mammillaria tobuschii: 79<br />

Mammillaria tuberculosa: 114, 133, 134, 136,<br />

137, 138, 144, (34), (35)<br />

Mammillaria tuberculosa var. durispina: 138<br />

Mammillaria varicolor: 113, 138, (37)<br />

Mammillaria vivipara: 125, 126, 127, 128,<br />

130, 132, 133, (31), (32), (33)<br />

Mammillaria vivipara var. aggregata: 130<br />

Mammillaria vivipara var. arizonica: 113,<br />

131, 132, (33)<br />

pellosperma -><br />

phellosperma<br />

scheerii -> scheeri<br />

scheerii -> scheeri


Mammillaria -><br />

Mammillaria<br />

246 cacti of the southwest<br />

Mammillaria vivipara var. borealis: 113, 127,<br />

128, 129, 130, 131, (32)<br />

Mammillaria vivipara var. deserti: 114, 131,<br />

132<br />

Mammillaria vivipara var. neo-mexicana:<br />

114, 127, 128, 129, 130, 132, 133, (32)<br />

Mammillaria vivipara var. radiosa: 113, 127,<br />

128, 129, 130, 131, 133, 137, (32)<br />

Mammillaria vivipara var. texana: 128, 129<br />

Mammillaria vivipara var. vivipara: 113,<br />

126, 127, 128, 129, 130, 131, (31)<br />

Mammillaria wilcoxii: 112, 151, 152, (42)<br />

Mammillaria wissmannii: 119, 123, 124<br />

Mammillaria wrightii: 112, 150, 151, 152,<br />

(41)<br />

Melocactus: 69<br />

Melocactus laciniatus: 69<br />

Navajoa: 103<br />

Neobesseya: 110, 112<br />

Neobesseya rosiflora: 124<br />

Neobesseya similis: 122<br />

Neobesseya wissmannii: 119, 124<br />

Neocoryphantha: 111, 112<br />

Neoevansia: 57<br />

Neolloydia: 64, 89, 93, 94<br />

Neolloydia conoidea: 93, 135<br />

Neolloydia intertextus: 89<br />

Neolloydia orcuttii: 135<br />

Neolloydia texensis: 93<br />

Neomammillaria: 110, 111, 112<br />

Neomammillaria leei: 143<br />

Opuntia: 9, 110, 159, 202, 217<br />

Opuntia aciculata: 179<br />

Opuntia aciculata var. orbiculata: 179<br />

Opuntia allairei: 210, 211<br />

Opuntia anahuacensis: 175<br />

Opuntia arborescens: 230<br />

Opuntia arbuscula: 238<br />

Opuntia arenaria: 165, 216, 225, 226, (59)<br />

Opuntia atrispina: 164, 180, 183, 188, 190,<br />

194, (50)<br />

Opuntia ballii: 165, 213, 214, 215, (56)<br />

Opuntia bentonii: 177<br />

Opuntia brachyarthra: 218<br />

Opuntia cacanapa: 178<br />

Opuntia caespitosa: 202<br />

Opuntia chihuahuensis: 195<br />

Opuntia chlorotica: 163, 182, 184, 187, (48)<br />

Opuntia chlorotica var. santa-rita: 187<br />

Opuntia clavata: 166, 227, 228, 229, (60)<br />

Opuntia compressa: 191, 197, 200, 201, 202,<br />

203, 205, 206, 207, 208, 210, 211, 212, 213,<br />

214, 217, 219, (54), (55)<br />

Opuntia compressa var. allairei: 164, 204,<br />

210, 211, (55)<br />

Opuntia compressa var. fusco-atra: 166, 175,<br />

204, 207, 208, 209, 211, 223, (54), (55)<br />

Opuntia compressa var. grandiflora: 166,<br />

209, 210, 211, (55)<br />

Opuntia compressa var. humifusa: 165, 202,<br />

203, 204, 205, 206, 207, (54)<br />

Opuntia compressa var. macrorhiza: 166, 204,<br />

205, 206, 207, 208, 211, 212, 213, 214, 215,<br />

219, (54)<br />

Opuntia compressa var. microsperma: 166,<br />

206, 207, (54)<br />

Opuntia compressa var. stenochila: 165, 211,<br />

212, (56)<br />

Opuntia convexa: 175<br />

Opuntia cyanella: 176<br />

Opuntia cymochila: 165, 199, 200, 201, 211,<br />

212, 214, (53)<br />

Opuntia davisii: 167, 234, 235, 236, (63)<br />

Opuntia deltica: 177<br />

Opuntia dillei: 172, 173, 181<br />

Opuntia dillenii: 177<br />

Opuntia drummondii: 166, 216, 217, 227,<br />

(57)<br />

Opuntia emoryi: 228<br />

Opuntia engelmannii: 161,163,168, 169, 170,<br />

171, 172, 173, 174, 175, 176, 177, 178, 179,<br />

180, 181, 182, 183, 184, 187, 188, 190, 191,<br />

192, 193, 194, 223, (44), (45), (46), (47),<br />

(48)<br />

Opuntia engelmannii var. aciculata: 163, 164,<br />

178, 179, (47)<br />

Opuntia engelmannii var. alta: 163, 168, 175,<br />

177, 187, (45), (46)<br />

Opuntia engelmannii var. cacanapa: 163, 177,<br />

178, (46)<br />

Opuntia engelmannii var. cyclodes: 163, 173,<br />

174, 178<br />

Opuntia engelmannii var. dulcis: 163, 179,<br />

187, (47)<br />

Opuntia engelmannii var. engelmannii: 162,<br />

171, 173, 174, (44)<br />

Opuntia engelmannii var. flexispina: 163,<br />

178, 179, 182, (46)<br />

Opuntia engelmannii var. linguiformis: 163,<br />

181, (48)<br />

Opuntia engelmannii var. subarmata: 164,<br />

180, 181, (47)<br />

Opuntia engelmannii var. texana: 162, 168,<br />

174, 175, 176, 177, 182, (45), (51)<br />

Opuntia engelmannii var. wootonii: 175<br />

Opuntia engelmannii X phaeacantha: 194<br />

Opuntia erinacea: 165, 220, 224, 225<br />

Opuntia erinacea var. hystricina: 224<br />

Opuntia erinacea var. xanthostemma: 220<br />

Opuntia expansa: 193<br />

Opuntia ferruginispina: 175<br />

Opuntia filipendula: 213, 214<br />

Opuntia fragilis: 166, 217, 218, 222, 226,<br />

227, (57)<br />

Opuntia fragilis var. brachyarthra: 218<br />

Opuntia fragilis var. frutescens: 240<br />

Opuntia frutescens: 239, 240<br />

Opuntia fulgida: 217<br />

Opuntia fusiliformis: 205<br />

Opuntia gilvoalba: 177<br />

Opuntia gomei: 172<br />

Opuntia gosseliniana: 187<br />

Opuntia gosseliniana var. santa-rita: 164,<br />

185, 186, 187, (50)<br />

Opuntia grahamii: 166, 226, 227, 236, (59)<br />

Opuntia grandiflora: 210<br />

Opuntia gregoriana: 172<br />

Opuntia griffithsiana: 175<br />

Opuntia humifusa: 202, 203, 206<br />

Opuntia hystricina: 164, 165, 221, 224, 225,<br />

(59)<br />

Opuntia imbricata: 230, 231, 233<br />

Opuntia imbricata var. arborescens: 166, 229,<br />

230, 231, 232, (61)<br />

Opuntia imbricata var. vexans: 166, 231,<br />

233, (61)<br />

Opuntia imbricata var. viridiflora: 166, 167,<br />

231, (61)<br />

Opuntia inermis: 167<br />

Opuntia intermedia: 210, 211<br />

Opuntia juniperina: 222, 223, 224<br />

Opuntia kleiniae: 167, 236, 237, 238, 239,<br />

240, (64)<br />

Opuntia laevis: 173<br />

Opuntia lawsonii: 177<br />

Opuntia laxiflora: 177<br />

Opuntia leptocarpa: 164, 170, 190, 191, 193,<br />

(51)<br />

Opuntia leptocaulis: 56, 167, 239, 240, (64)<br />

Opuntia leptocaulis var. badia: 240<br />

Opuntia leptocaulis var. brevispina: 240<br />

Opuntia leptocaulis var. longispina: 240<br />

Opuntia leptocaulis var. pluriseta: 240<br />

Opuntia leptocaulis var. robustior: 240<br />

Opuntia leptocaulis var. stipata: 240<br />

Opuntia leptocaulis var. vaginata: 239, 240<br />

Opuntia lindheimeri: 170, 171, 172, 174,<br />

175, 176, 180, 190, 191<br />

Opuntia lindheimeri var. chisoensis: 174<br />

Opuntia lindheimeri var. dulcis: 180<br />

Opuntia loomisii: 212<br />

Opuntia macateei: 175, 208, 209<br />

Opuntia mackensenii: 197<br />

Opuntia macrocalyx: 185<br />

Opuntia macrocentra: 164, 173, 185, 186,<br />

187, 195, (49)<br />

Opuntia macrocentra var. castetteri: 186<br />

Opuntia macrocentra var. minor: 186<br />

Opuntia macrorhiza: 190, 204, 205<br />

Opuntia mesacantha: 200, 202, 203, 206<br />

Opuntia mesacantha var. cymochila: 200<br />

Opuntia mesacantha var. oplocarpa: 193,<br />

195, 197<br />

Opuntia microdasys: 185<br />

Opuntia microdasys var. rufida: 185<br />

Opuntia missouriensis: 222<br />

Opuntia missouriensis var. microsperma: 223<br />

Opuntia mortolensis: 240<br />

Opuntia nemoralis: 203, 217<br />

Opuntia nicholii: 221<br />

Opuntia opuntia: 202<br />

Opuntia phaeacantha: 173, 179, 184, 190,<br />

191, 192, 193, 194, 195, 196, 197, 199,<br />

200, (51), (52), (53)<br />

Opuntia phaeacantha var. brunnea: 164, 186,<br />

194, 195, 198, (52)<br />

Opuntia phaeacantha var. camanchica: 163,<br />

164, 166, 183, 191, 193, 196, 197, 198,<br />

200, 201, 221, (53)<br />

Opuntia phaeacantha var. major: 163, 164,<br />

191, 192, 193, 197, (51)<br />

Opuntia phaeacantha var. nigricans: 163,<br />

188, 190, 193, 194, (51)


uba -> rubra<br />

index of scientific names 247<br />

Opuntia phaeacantha var. tenuispina: 165,<br />

197, (53)<br />

Opuntia plumbea: 165, 215, 216, (56)<br />

Opuntia polyacantha: 165, 200, 219, 221,<br />

222, 223, 224, 225, (58), (59)<br />

Opuntia polyacantha var. albispina: 222, 223<br />

Opuntia polyacantha var. platycarpa: 223,<br />

224<br />

Opuntia polyacantha var. rufispina: 222, 223<br />

Opuntia polyacantha var. subinermis: 223,<br />

224<br />

Opuntia polyacantha var. trichophora: 223,<br />

224<br />

Opuntia pottsii: 165, 212, 213, 214, 215, (56)<br />

Opuntia pyrocarpa: 175, 197<br />

Opuntia rafinesquii: 170, 191, 200, 202, 203,<br />

204,206<br />

Opuntia rafinesquii var. cymochila: 200<br />

Opuntia ramulifera: 240<br />

Opuntia reflexa: 175<br />

Opuntia rhodantha: 220, 221, (58)<br />

Opuntia rhodantha var. brevispina: 220<br />

Opuntia rhodantha var. elegans: 220<br />

Opuntia rhodantha var. flavispina: 220<br />

Opuntia rhodantha var. fulgens: 220<br />

Opuntia rhodantha var. gracilis: 220<br />

Opuntia rhodantha var. orbicularis: 220<br />

Opuntia rhodantha var. pallida: 220<br />

Opuntia rhodantha var. pisciformis: 220<br />

Opuntia rhodantha var. rhodantha: 164, 166,<br />

220<br />

Opuntia rhodantha var. rosea: 220<br />

Opuntia rhodantha var. rubra: 220<br />

Opuntia rhodantha var. salmonea: 220<br />

Opuntia rhodantha var. schumanniana: 220<br />

Opuntia rhodantha var. spinosior: 164, 221,<br />

(58)<br />

Opuntia roseana: 205<br />

Opuntia rufida: 164, 184, 185, 187, (49)<br />

Opuntia rufida var. tortiflora: 185<br />

Opuntia sanguinocula: 206<br />

Opuntia santa-rita: 187<br />

Opuntia schottii: 166, 217, 227, 228, (60)<br />

Opuntia schweriniana: 222, 223<br />

Opuntia sinclairii: 175<br />

Opuntia sphaerocarpa: 166, 218, 219, 223,<br />

(57)<br />

Opuntia spinosibacca: 162, 183, 184, (49)<br />

Opuntia spinosior: 166, 233, 234, (62)<br />

Opuntia spinotecta: 233<br />

Opuntia stanlyi: 166, 228, 238, (60)<br />

Opuntia stanlyi var. parishii: 229<br />

Opuntia stenochila: 212<br />

Opuntia stricta: 162, 164, 167, 168, 177, (44)<br />

Opuntia strigil: 164, 187, 188, 190, 194, (50)<br />

Opuntia tardospina: 163, 182, 183, 190, (48)<br />

Opuntia tenuispina: 195, 198, 199<br />

Opuntia tortispina: 197, 198, 200, 201<br />

Opuntia tortispina var. cymochila: 200<br />

Opuntia tracyi: 217<br />

Opuntia trichophora: 222, 223<br />

Opuntia tricolor: 178<br />

Opuntia tunicata: 166, 217, 236, 237, (63)<br />

Opuntia ursina: 225<br />

Opuntia vaginata: 167, 239<br />

Opuntia valida: 172<br />

Opuntia vexans: 232, 233<br />

Opuntia virgata: 240<br />

Opuntia viridiflora: 231<br />

Opuntia vulgaris: 191, 202<br />

Opuntia whipplei: 166, 167, 233, 234, 235,<br />

236, (62)<br />

Opuntia whipplei var. laevior: 234<br />

Opuntia whipplei var. spinosior: 233, 234<br />

Opuntia wootonii: 175<br />

Opuntia wrightiana: 238<br />

Opuntia wrightii: 238<br />

Opuntia xanthostemma: 220<br />

Pediocactus: 9, 63, 64, 103, 106<br />

Pediocactus knowltonii: 105, (27)<br />

Pediocactus papyracanthus: 105, 106, (27)<br />

Pediocactus simpsonii: 104, 106<br />

Pediocactus simpsonii var. minor: 105<br />

Pediocactus simpsonii var. robustior: 105<br />

Pediocactus simpsonii var. simpsonii: 104,<br />

(27)<br />

Peniocereus: 8, 55, 57<br />

Peniocereus greggii: 4, 57, 58, (18)<br />

Peniocereus greggii var. cismontanus: 59<br />

Peniocereus greggii var. roseiflorus: 59<br />

Peniocereus greggii var. transmontanus: 59<br />

Pereskia: 3, 110<br />

Phellosperma: 157<br />

Platyopuntia: 161<br />

Pseudocoryphantha: 111, 112<br />

Roseocactus: 100, 101<br />

Roseocactus fissuratus: 101<br />

Sclerocactus: 64, 75<br />

Sclerocactus mesae-verdae: 75<br />

Sclerocactus whipplei: 74, 75<br />

Sclerocactus whipplei var. intermedius: 75<br />

Sclerocactus whipplei var. whipplei: 75<br />

Subgymnocarpae: 111<br />

Thelocactus: 64, 85, 86, 93<br />

Thelocactus bicolor var. flavidispinus: 87<br />

Thelocactus bicolor var. schottii: 85<br />

Thelocactus flavidispinus: 87<br />

Thelocactus uncinatus: 74, 80<br />

Thelocactus wagnerianus: 88<br />

Theloidei: 103<br />

Toumeya: 103, 106<br />

Toumeya papyracantha: 106<br />

Utahia: 103<br />

Wilcoxia: 8, 55<br />

Wilcoxia poselgeri: 4, 55, 56, (17)<br />

Pellosperma Cf.<br />

Phellosperma


248 cacti of the southwest<br />

INDEX <strong>OF</strong> COMMON NAMES<br />

The spelling of the Spanish names is that of local usage where the plants grow,<br />

without correction or standardization.<br />

Abrojo: 236<br />

Aggregate Cactus: 41<br />

Aguijilla: 239<br />

Alicoche: 51, 53<br />

Arizona Coryphantha: 131<br />

Arizona Queen of the Night: 57<br />

Arizona Rainbow Hedgehog: 30<br />

Ball Cactus: 125<br />

Barrel Cactus: 71<br />

Beehive Cactus: 42<br />

Berlandier’s Alicoche: 52<br />

Bisnaga de Dulce: 66<br />

Bisnaga Meloncillo: 66<br />

Bisnagre: 66<br />

Biznaga: 71<br />

Biznaga Costillona: 84<br />

Biznaga de Agua: 71<br />

Biznaga de Chilitos: 152, 153, 155<br />

Biznaga de Tuna: 84<br />

Biznaga es Pinosa: 84<br />

Biznaga Ganchuda: 84<br />

Biznaga Limilla: 84<br />

Black Lace: 23, 24<br />

Blind Pear: 184<br />

Brittle Cactus: 217<br />

Brown-Flowered Hedgehog: 72<br />

Brown-Spined Prickly Pear: 191<br />

Bunch-Ball Cactus: 41<br />

Button Cactus: 108<br />

Cabeza del Viejo: 30<br />

Candelabrum Cactus: 229<br />

Candle Cholla: 237<br />

Candy Barrel: 71<br />

Candy Cactus: 68<br />

Cane Cholla: 233<br />

Cat-Claw Cactus: 72<br />

Cave Cactus: 229, 233<br />

Chaparral Cactus: 57<br />

Chautle: 101<br />

Cholla: 229<br />

Claret-Cup Cactus: 35, 36, 37, 39<br />

Classen’s Cactus: 19<br />

Clavelina: 236<br />

Clavellina: 227<br />

Cliff Prickly Pear: 220<br />

Clock-Face Prickly Pear: 182<br />

Club Cholla: 228, 229<br />

Cockle-Burr Cactus: 216<br />

Cock-Spur Cactus: 216<br />

Comb Hedgehog: 29<br />

Cow’s Tongue Cactus: 181<br />

Coyote Candles: 229<br />

Creeping Cholla: 228<br />

Crow-Foot Prickly Pear: 216<br />

Dagger Cholla: 228, 229<br />

Dahlia Cactus: 55<br />

Deer-Horn Cactus: 57<br />

Desert Christmas Cactus: 239, 240<br />

Devil Cactus: 227<br />

Devil’s Cholla: 228<br />

Devil’s Claw Barrel: 74<br />

Devil’s Head: 66, 68<br />

Dog Cholla: 227<br />

Dry Whisky: 97<br />

Dumpling Cactus: 121<br />

Eagle Claws: 66, 67<br />

Early Bloomer: 88<br />

Engelmann’s Prickly Pear: 168<br />

Estria del Tarde: 128<br />

Fendler’s Hedgehog Cactus: 48<br />

Fendler’s Pitaya: 48<br />

Finger Cactus: 118<br />

Fishhook Barrel: 71<br />

Fishhook Cactus: 76, 77, 79, 80, 81, 149<br />

Flaming Prickly Pear: 168<br />

Flap-Jack Cactus: 180<br />

Fragile Prickly Pear: 217<br />

Garrambullo: 239, 240<br />

Glory of Texas: 85, 86<br />

Golden Rainbow Hedgehog: 32<br />

Grama-Grass Cactus: 105, 106<br />

Grape Cactus: 147<br />

Green-Flowered Pitaya: 13, 16<br />

Green-Flowered Torch Cactus: 13, 16<br />

Hair-Covered Cactus: 147<br />

Heart Twister: 41<br />

Hedgehog Cactus: 79, 80, 81<br />

Hicore: 95<br />

Horse Crippler: 68, 69<br />

Hunger Cactus: 221<br />

Jiculi: 95<br />

Junior Tom Thumb Cactus: 140<br />

King’s Cup Cactus: 35, 37<br />

Klein Cholla: 237<br />

Knowlton Cactus: 105<br />

Lace Cactus: 19, 20<br />

Lady-Finger Cactus: 51<br />

Large-Flowered Opuntia: 209<br />

Lengua de Vaca: 181<br />

Little Chilis: 152, 155<br />

Living Rock: 101, 102<br />

Long Mama: 120<br />

Long-Tubercled Coryphantha: 114<br />

Lower Rio Grande Valley Barrel: 82<br />

Low Prickly Pear: 201<br />

Manco Caballo: 68<br />

Mesa Verde Cactus: 75<br />

Mescal Button: 97<br />

Mounded Dwarf Cholla: 226<br />

Mountain Cactus: 104<br />

Mulato: 108<br />

Needle ‘Mulee’: 114<br />

New Mexico Coryphantha: 128<br />

New Mexico Prickly Pear: 191<br />

New Mexico Rainbow Cactus: 13<br />

Night-Blooming Cereus: 58, 60<br />

Nipple Cactus: 118, 122, 152<br />

Nopal: 168<br />

Nylon Cactus: 13<br />

Organo: 46, 60<br />

Paper-Spined Cactus: 105<br />

Pencil Cactus: 55<br />

Pest Pear: 167, 175<br />

Peyote: 95, 96, 97, 98<br />

Peyote Cimarron: 101<br />

Peyotl: 95<br />

Pincushion: 125, 128, 149<br />

Pineapple Cactus: 118<br />

Pink-Flowered Echinocereus: 48<br />

Pitahaya: 43, 60<br />

Pitaya: 44, 46, 47<br />

Porcupine Prickly Pear: 224<br />

Purple Candle: 19<br />

Purple Hedgehog: 48<br />

Purple Prickly Pear: 185<br />

Rat-Tail Cactus: 233<br />

Red-Flowered Hedgehog Cactus: 41<br />

Red-Goblet Cactus: 43<br />

Root Cactus: 77


index of scientific names 249<br />

Runyon’s Coryphantha: 121<br />

Runyon’s Escobaria: 140<br />

Sacasil: 55<br />

Sand-Burr Cactus: 216<br />

Sea-Urchin Cactus: 70<br />

Seni: 95<br />

Silverlace Cactus: 137<br />

Sitting Cactus: 48<br />

Slender Stem Cactus: 239<br />

Small Papillosus: 50<br />

Smooth Prickly Pear: 201<br />

Sour Cactus: 129, 131<br />

Spiny Star: 125, 128<br />

Standly’s Cholla: 228<br />

Star Cactus: 101<br />

Star Rock: 101<br />

Starvation Cactus: 221<br />

Sticker Cactus: 233<br />

Strawberry Cactus: 35, 39, 44, 45, 46, 47, 48<br />

Sunami: 101<br />

Sunset Cactus: 149<br />

Sweet-Potato Cactus: 57<br />

Tasajillo: 239<br />

Teonanacatl: 95<br />

Texas Hedgehog: 72<br />

Texas Night-Blooming Cereus: 57<br />

Texas Rainbow Cactus: 32<br />

Tlalcoyote: 95<br />

Torch Cactus: 48<br />

Toumeya: 105<br />

Tree Cactus: 229, 230<br />

Triangle Cactus: 60, 61, 62<br />

Tulip Prickly Pear: 191<br />

Tuna: 168<br />

Turk’s Head Cactus: 41, 66, 72, 84<br />

Twisted-Rib Cactus: 79, 80, 81<br />

Uocoui: 95<br />

Velas de Coyote: 229<br />

Visnaga: 71, 84<br />

Viznaga: 68<br />

Whipple’s Cholla: 233<br />

Whisky Cactus: 97<br />

White Column: 137<br />

White-Flowered Visnagita: 88<br />

Wide Cactus: 220<br />

Xicorl: 95<br />

Yellow-Flowered Alicoche: 49<br />

Yellow-Flowered Echinocereus: 49<br />

Yellow-Flowered Pitaya: 32


Trees, Shrubs, and Woody Vines of the Southwest<br />

A Guide for the States of Arkansas, Louisiana, New Mexico, Oklahoma, and Texas<br />

By Robert A. Vines<br />

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is an incredible achievement in the history of natural science publishing. It is an immense<br />

production, indispensable to naturalists of the region."—American Forests. $30.00<br />

Roadside Flowers of Texas<br />

Paintings by Mary Motz Wills<br />

Text by Howard S. Irwin<br />

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Adventures with a Texas Naturalist<br />

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UNIVERSITY <strong>OF</strong> TEXAS PRESS · AUSTIN & LONDON

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