Chapter I: Part 1

saguaro
national
monument
ARIZONA

Napier Shelton
_based on an earlier work by Natt Dodge_

NATURAL HISTORY SERIES

1972
National Park Service
U.S. DEPARTMENT of the INTERIOR

For sale by the Superintendent of Documents, U.S. Government Printing
Office
Washington, D.C. 20402—Price $1.25
Stock Number 2405-0292

preface

This book is a simple account of the natural history of Saguaro National Monument. It is intended to help you understand the relationships between land, climate, plants, wild animals, and man in the environment of a hot desert. While it includes brief profiles of many representative species, it is not intended to serve as a guide to the monument. It does indicate where the several distinctive natural communities exist, and when and where to look for certain plants and animals. For identification purposes, you will need field guides.

The present edition is my revision of the 1957 book by Natt Dodge, then naturalist for the Southwest Region of the National Park Service. The first five and the last two chapters are essentially new; the central chapters on plants and animals remain largely as written in the first edition.

The authors wish to thank former and present members of the monument staff for their help and companionship, in both field and office. We are particularly grateful to Chief Naturalist Harold T. Coss, Jr., who devoted much time and effort to obtaining many of the photographs and, with Park Biologist Warren F. Steenbergh, gave the manuscript a thorough scrutiny. The cooperation and hospitality of Superintendent Harold R. Jones have created the best possible climate for work on the revised edition. National Park Service geologist Robert H. Rose contributed in very great measure to the geological content. Finally, our thanks go to the many students of desert life on whose knowledge this book has been built, and to monument visitors who ask questions—for their concern gives hope for better relations between man and nature.

—N.S.

contents

The Desert Scene, 1
The Sonoran Desert and the Monument, 3
Climate: The Vital Factor, 9
Rocks: Foundation and Soilmakers, 13
Plant and Animal Zones, 19
Desert Plants, 23
Succulents, 25
The Saguaro—Monarch of the Monument, 27
Other Common Cactuses, 33
Non-Succulents, 36
Perennials, 36
Ephemerals, 42
Major Plant Communities of Saguaro, 49
Plants of the Hills and Mountains, 55
Oak-Pine Woodland, 55
Ponderosa Pine Forest, 58
Animals and How They Survive, 61
Invertebrates, 62
Amphibians, 64
Reptiles, 64
Birds, 66
Mammals, 72
The Rhythms of Nature, 79
The Impact of Man, 83
Appendix, 87
Suggested Reading, 87
Common and Scientific Names of Plants, 89
Reptiles and Amphibians of the monument, 92
Birds of the monument, 94
Mammals of the monument, 97

the desert scene

Scattered through the wide, lonely Sonoran Desert, isolated mountain ranges raise jagged blue silhouettes against the sky. The high ones wear a crown of dark pines and a speckled mantle of oaks. Lapping against their feet is the desert sea, studded with the green masts of giant saguaro cactuses rising above a motley crew of tough, strange, often handsome subordinates.

On either side of the Santa Cruz Valley in southeastern Arizona, Saguaro National Monument embraces two of these ranges, with the desert lands at their feet. Although it was established primarily to preserve impressive stands of saguaros, it gains wholeness by including the mountains that shed the rocky soil on which the saguaros grow. The monument is in two sections, some 30 miles apart. The _Rincon Mountain Section_, east of Tucson, includes in its 99 square miles a spectrum of plantlife ranging from saguaro communities at the low elevations to a wet fir forest on the north slope of 8,666-foot Mica Mountain. The 24-square-mile _Tucson Mountain Section_, about 12 miles west of the city, has a denser stand of saguaros, on the lower slopes of a wild jumble of volcanic mountains dominated by 4,687-foot Wasson Peak.

Within the cactus forests and upon the mountain slopes, life in myriad forms goes on in a delicate and continuous adjustment to changing environments. Over millennia, mountains rise and crumble, species evolve and fade from the scene. Over days and weeks and years, populations of plants and animals rise and fall, in response to thousands of interactions which we are only beginning to understand. In the following pages, we will meet the main characters in this natural drama (including some of the most improbable in all of nature); we will investigate the environmental stage upon which they perform; and we will try to understand something of the play itself, including our own role in it.

the sonoran desert
and the monument

Saguaro National Monument is at the northeastern edge of the Sonoran Desert. Named for the state of Sonora, Mexico, in which the greater part of it lies, this, one of four major deserts in North America, is distinguished by differences in climate and vegetation. The Great Basin Desert, mainly in Nevada and Utah between the Rockies and the Sierra Nevada, has cold winters, sparse precipitation distributed fairly evenly throughout the year, and rather simple vegetation dominated by the low shrubs, sagebrush and saltbrush. Immediately to the south, in southern Nevada and southeastern California, is the Mohave Desert, with cool winters during which most of the year’s precipitation comes, and with plant cover consisting mostly of shrubs such as creosotebush. At higher elevations grows the Joshua-tree, a giant yucca. The Chihuahuan Desert, with cool winters and summer rainfall, covers the broad plateau of north-central Mexico, extending into southern New Mexico and west Texas. Its vegetation consists mostly of small cactuses, spiny shrubs, and succulent-leaved plants such as yuccas.

Lying between the Mohave and Chihuahuan deserts, the Sonoran Desert has both winter and summer rainfall, with spring and autumn droughts. Its mild winters and bi-seasonal rainfall encourage a vegetation far surpassing in lushness and variety that of the other deserts. From west to east, the land rises and precipitation increases. Yuma, in southwestern Arizona, lying at 141 feet above sea level, gets about 3 inches a year. Tucson, in southeastern Arizona, is at 2,400 feet elevation and averages 11 inches a year. In the western part, where plains are extensive and mountain ranges low and far apart, a very few small-leaved species such as creosotebush dominate. Toward the east, mountain ranges become more numerous, shedding material on which more kinds of plants, particularly paloverde, mesquite, and cactuses, assume leading roles in the vegetative cover. Both sections of the monument lie within the eastern, wetter, more diversified part of the Sonoran Desert known as the Arizona Upland. Organ Pipe Cactus National Monument, to the southwest, is in an area transitional between the Arizona upland and Colorado River lowland phases of the Sonoran Desert.

But even between the two sections of Saguaro National Monument there are differences. There is the obvious fact that the Tucson Mountains are much lower and smaller in mass than the Rincons. And there is the not-so-obvious fact that the Tucson Mountain Section extends to lower elevations (2,200 as opposed to 2,700 feet). These conditions are reflected in the somewhat warmer environment of the western section. This helps to explain why certain Sonoran Desert plants and animals here reach their northeastern limits; among them are ironwood, desert iguana, desert horned lizard, western shovel-nosed snake, sidewinder, desert kangaroo rat, and the Le Conte thrasher. The Rincon Mountain Section, by virtue of its higher elevations, has plants and animals—such as ponderosa pine, Steller’s jay, and whitetail deer—not found in the western section.

On the geologic time scale the Sonoran Desert, like all the earth’s present deserts, is a recent development. Some 50 million years ago tropical forests grew here, as they did over most of southern North America. As the Southwest gradually became drier, new species adapted to the new conditions evolved; and the general type of vegetation changed—with some retrogression during wetter periods—from forest to savanna (grasslands with scattered trees) to arid subtropical scrub (such as is now found in southern Sonora State), and finally to the plant communities of today’s Sonoran Desert. The last stage has occurred only during the past few million years.

The assemblage of species from which most of our present desert and lower mountain plants were derived is known as the Madro-Tertiary Flora, to denote its center (Sierra Madre in Mexico) and time (Tertiary period—one million to 65 million years ago) of early development. Reflecting these origins, a majority of the species growing below 6,000 feet elevation in the monument today are basically Mexican or Central or South American in their distribution.

On the higher mountains of the Sonoran Desert, however, exist species with an entirely different ancestry. Most of the plants of the pine and fir forests of these high places, like the plants of the Great Basin Desert, derive from the Arcto-Tertiary Flora, which dominated the northern part of our continent during Tertiary times. As climate changes, one or the other of these great plant assemblages will benefit at the expense of the other. If cooler, wetter periods return, as they did during the Pleistocene epoch, ponderosa pines, now found above 6,000 feet, may again grow nearly as low down as today’s cactus forests. But if the present long-term drying trend continues, the northern plants will eventually be squeezed off their mountaintops.

Animals, to some extent, reflect the same dichotomy of origins we have seen in the plants. Thus, on the mountaintops supporting pine and fir forests, a northern contingent of animals predominates, while on the lower slopes and desert the fauna has a Mexican character. The canyons and oak-pine forests of middle elevations in southeastern Arizona are particularly exciting, because in these biological islands, isolated by surrounding desert, live many “Mexican” species of animals found nowhere else in the United States. Naturally, mountain ranges nearest the border—such as the Huachucas and Chiricahuas—have the greatest numbers of Mexican specialties; but the Rincons have their share too.

The wonderful diversity of plant and animal life in Saguaro National Monument depends on a diversity of habitats. These in turn owe their existence to a widely varying triumvirate of environmental factors—climate, soil, and topography. To understand the biological interplay that goes on here, we first must know something of the environmental conditions that circumscribe it.

Geological Time Table

Era Period Epoch Years Major
before geological
present events
time
Cenozoic Quaternary Recent 10,000 Cascade Range
and Sierra
Nevada uplifted
Pleistocene
1,000,000
Tertiary Pliocene
12,000,000
Miocene
26,000,000
Oligocene
38,000,000
Eocene
54,000,000 Rocky Mountains
formed
Paleocene
65,000,000
Mesozoic Cretaceous
136,000,000
Jurassic
195,000,000
Triassic
250,000,000 Appalachian
folding
Paleozoic Permian
280,000,000
Pennsylvanian
320,000,000
Mississippian
345,000,000
Devonian
395,000,000
Silurian
440,000,000
Ordovician
500,000,000
Cambrian
570,000,000
Precambrian

climate:
the vital factor

Climate is the chief arbiter of life on earth. Each plant and animal, including man, has tolerance limits for heat and water below or above which it dies. The Sonoran desert, with its low and uncertain rainfall and high summer temperatures, thus presents one of earth’s most taxing environments. For most desert plants and animals, the main problem presented by this climatic combination is a scarcity of water. Why, we might ask, is this part of the globe so dry?

Essentially, the dryness of the Sonoran and other southwestern deserts results from their geographic situation. First, these lands lie east of a persistent high-pressure cell over the eastern Pacific—part of a belt of high-pressure cells which encircles the earth at this latitude. High-pressure systems in the Northern Hemisphere have descending, warming, dry air in their middles and along their eastern sides. The air that flows eastward from the “East Pacific high” thus contains little moisture, and dictates desert conditions over much of the Southwest. Furthermore, the mountains that more or less ring these deserts intercept much of the meager incoming moisture, by forcing the air to rise and cool so that its moisture condenses and falls to the ground.

But during two periods of the year, the desert does receive rain. In winter, the East Pacific high shifts southward with the sun, reducing its effects on the Southwest. Now the low pressure storm systems, which at this season move eastward across the continent, can sometimes dip far enough south to bring rain to the Sonoran Desert. These rains are usually gentle, and cover large areas. Their clouds may cover the sky for several days at a time.

By April, the East Pacific high has returned far enough northward to resume its control of southwestern weather. Now the fore-summer drought sets in, intensified by gradually rising temperatures. Day after day the sun climbs higher in the sky, baking the earth and testing the endurance of desert life. Then, sometime in late June or early July, clouds begin forming in the afternoons over the mountains; finally, one day they build up to great thunderheads. The drought ends suddenly as rain pours down and lightning cracks the sky. Down the canyons and along the desert washes rages the water, brown with soil. In a short time, the storm is over, and the water disappears into the ground. Just 1 mile away, rain may not have fallen.

These intense, local summer rains, so different from the winter ones, have a different origin as well. They are spawned by the Atlantic’s Bermuda high, which has shifted northwestward and sent moist air from the Atlantic and Gulf of Mexico flowing off its western edge toward the deserts. This air rises quickly when it strikes hot, high mountains such as the Rincons, and its moisture condenses to fall as rain.

Some time in September the rains of summer stop, as the Bermuda High moves south again, removing its influence and returning the East Pacific high to a position of dominance. The sun and wind again wring moisture from the soil, but with diminishing effect. The thermometer no longer reaches the 100’s as it did in June, July, and August. In November or December, the winter rains begin. Water stays longer in the soil, because temperatures now range between 30° and 80°. Life in the desert begins a new cycle. This rainy period usually lasts into March.

On the mountain slopes, of course, the same seasonal patterns of heat and precipitation occur, but they are tempered or enhanced by elevation. With increasing elevation, temperatures decline and precipitation increases. When July temperatures at headquarters are averaging in the nineties, they will be in the sixties on top of the Rincons. In the cactus forest at the foot of Tanque Verde Ridge only about 10 to 12 inches of rain fall in a year, while Manning Camp, at 8,000 feet, gets about 18 to 25 inches. These up-slope gradients of temperature and precipitation, as we shall see, have pronounced effects on plant and animal life.

rocks:
foundation and soilmakers

Soils are derived from the fragmentation and decomposition of rocks. Combinations of soils and climate, varying from place to place, create an almost infinite number of environments with differences so subtle and small as to make it appropriate to refer to them as microenvironments. Though these differences be minor, they are often sufficient to create niches each of which becomes a habitat for a particular group of plants and animals. Thus it is that the saguaros and other desert vegetation in Saguaro National Monument and vicinity are found in particular environmental or ecological niches where just the right combinations of soils, moisture, sunlight, temperature and other factors are present.

Since the rocks provide the foundation and the source of soils that support the plant and animal life, it is helpful to understand something about the origin and evolution of the rocks and the landscape of which they are a part. Clues to this story are to be found in the composition of the rocks and their relationships to one another.

Rocks of the three major classes—igneous, metamorphic, and sedimentary—are found within the monument. The igneous rocks include granites and various kinds of lava flows, together with some intrusive dikes and veins. The metamorphic rocks are represented by gneiss and schist. Lastly, sedimentary rocks occur as limestone, sandstone, and alluvial fill material. Each of these great classes of rocks was formed in a different manner, and these differences reveal the nature of the events which are a part of the geologic history of the monument and surrounding area.

Catalina Gneiss (pronounced NICE) is the predominant rock that visitors see in the Rincon Mountains and the Tanque Verde Ridge; exposures of granite and schist are also found. Gneiss is a coarse-grained metamorphic rock resembling granite, having a banded appearance, and consisting of alternating layers of different minerals such as feldspar, quartz, mica, and hornblende. The banding and texture reveal that the gneiss, now exposed by uplift and erosion, was formed from parent rocks deep below the earth’s surface during the Precambrian Era more than a half billion years ago. There the parent rocks, subjected to pressure and heat, melted, flowed, and crystallized, before resolidification.

Though the gneiss of the Rincon Mountains is ancient, the uplift that raised it to its present height is thought to have occurred rather recently (within the past 24 million years), during the period when most of the Sonoran Desert ranges apparently were formed. In age, these mountains thus would fall somewhere between the older Rockies and the younger Sierra Nevada.

The schist, which underlies much of the Cactus Forest, where it is exposed along washes, was probably formed during the Cretaceous period more than 65 million years ago. Like the gneiss, the schist is a metamorphic rock, having been formed by the transformation of parent rocks at depth under great pressure and heat. Due mainly to differences in the composition of the parent rock, together with its mica content, the rock that resulted was platy and cleavable (accounting for its classification as schist).

Granite, which forms Wasson and Amole peaks in the Tucson Mountain Section, is an igneous rock that (like the metamorphic gneiss and schist) originated at depths below the earth’s surface. The granite was formed by the solidification of molten rock material that moved upward en masse from greater depths, rather than by the alteration of ancestral rocks. Erosion of the uplifted land mass has not only stripped away the overburden but has developed valleys deeply incised into the granite itself.

Volcanic rocks in the form of rhyolite, andesite, and basalt flows also are exposed, chiefly in the Tucson Mountain section of the monument. These are all extrusive igneous rocks composed of magma that solidified after reaching the earth’s surface through vents or fissures. There are differences in the lava flows, reflecting differences in composition, temperature, and other conditions of the magma from which they were formed. The rhyolite is lighter in color than andesite, and it is somewhat richer in feldspar. Basalt, on the other hand, is dark, is deficient in feldspar and quartz, and contains relatively large amounts of the darker minerals such as hornblende, pyroxene, and olivene. Lava flows of the rhyolite and andesite variety occur in the Cactus Forest locality of the Rincon Mountain Section. Cat Mountain Rhyolite is the name given to a rhyolite flow of Tertiary age that forms the topmost layer in much of the Tucson Mountain Section. A small exposure of basalt is also found here.

Limestone, sandstone, and shale formations are also exposed at various places within the monument. They are reliable indicators of seas that covered the area during one or more times of the geologic past. Comparisons with the limestone formations farther to the south in the Colossal Cave region indicate that the limestone in the monument was formed as far back as late in the Paleozoic Era (345 million years or more ago). This age is suggested by types of fossils, including fragments of crablike trilobites and crinoid (sea lily) stems, found in limestones of the same age near Colossal Cave.

The Recreation Redbeds of the Tucson Mountain Section is one of the important sandstone members exposed in the monument. Geologists believe that its deposition occurred early in the Cretaceous Period more than 65 million years ago. Above the Recreation Redbeds lies a formation consisting mainly of limestone and commonly known as the Tucson Mountain “Chaos”—an appropriate name, in view of the geologists’ meager knowledge of it. Within the Chaos formation, believed to be lower to Middle Tertiary in age, are limestone blocks of greater age than the rocks within which they are entombed. No generally accepted explanations have been advanced as to how these relationships developed.

The processes that destroy mountains continue concurrently with those that build them. Temperature changes, weathering, and downslope creep under the influence of gravity are among the agents which destroy the rocks and eventually convert them into soil. Though these processes work rapidly from a geological standpoint, in terms of the average human life span, they progress at an imperceptible rate. Violent thunderstorms and cloudbursts, however, cause a massive, often spectacular movement of boulders, gravel, sand, and silt by torrential streams.

In desert regions, most of the runoff from storms sinks into the slopes, dropping its burden along the way. Big rocks are dropped early as the carrying power of the water diminishes with the speed of flow; smaller fragments travel farther. The finest material is carried far out into the basins between mountain ranges, gradually filling them. (The alluvial material in the Tucson Basin is estimated to be 2,000 to 5,000 feet thick.) Thus desert mountains tend to bury themselves in their own debris.

Alluvial fans (fan-shaped deposits built by rivers flowing from mountains into lowlands) form at the mouths of canyons. Sometimes the alluvial fans of adjacent canyons coalesce, forming the long, sweeping slope known as a bajada (pronounced ba-HA-da). In other places the eroded bedrock extends outward from the bases of desert mountains, forming “pediments,” which are usually covered by a veneer of alluvial material. Pediments of this kind stretch from the lower slopes of the Rincon Mountains. In the Tucson Mountains the alluvial material deepens rapidly toward the Avra Valley because the bedrock at the base of the mountain dips steeply downward.

Serious gaps remain in the story of the origin and evolution of the landscape in Saguaro National Monument. Moreover, it is difficult to establish the exact sequence in which the various events occurred. The composition, texture, and relationships of the rocks, however, do reveal much about the nature of the events and the processes that were involved.

Faults are hypothetical

Showing:

Valley Fill
Wasson Peak
_Granite_
_Folded and tilted Sedimentary and Volcanic Rock_
_Santa Cruz River_
_Flood Plain_
_Valley Fill_
Tucson
_Pantano Wash_
Pediment
Tanque Verde Ridge
Mica Mountain
_Gneiss_

The presence of the limestones and sandstones indicates that that area was submerged below the water of ancient seas one or more times in the geologic past. The gneiss, schist, and granite bespeak deep-seated metamorphism and magmatic intrusion, which gave these rocks the form, composition, and texture they possess today. The lava flows are indicative of volcanic activity that was a part of the extensive volcanism that occurred in this part of southern Arizona. And, finally, the great alluvial fans and bajadas suggest that these mountain ranges could bury themselves in their own products of erosion unless the mountain-building processes in future eons continue at a faster pace than the wearing-down processes. This evidence enables us to perceive today’s landscape as but a transitional phase in the drama of change that will continue for milleniums.

plant and animal zones

Mountains are biologically exciting because their elevation produces, in a short vertical distance, the same climatic changes that occur over long latitudinal distances. In Arizona, average temperatures drop about 4° with each 1,000-foot rise in elevation, and precipitation increases about 4 to 5 inches. At sea level, you would have to travel about 900 miles north to experience the climatic change found in going from the lowest part of the monument (2,200 feet) to the top of Mica Mountain (8,600 feet). Plant-and-animal communities change along these climatic gradients.

In southern Arizona, the sequence of vegetation types begins with desert scrub at the lowest elevations, and ranges through desert grassland, oak woodland and chaparral, oak-pine woodland, ponderosa pine forest, and Douglas-fir forest, to Engelmann spruce forest on top of the highest mountains. In the national monument, we have all these types except the last—although desert grassland is poorly represented here because the steep slopes squeeze it into a narrow altitudinal band; and Douglas-fir forest is restricted to small areas in canyons and on north-facing slopes at high elevations. A few drainageways (notably Chiminea Canyon, on the south side of the monument) support patches of deciduous trees classified as riparian woodland.

These vegetation zones should not be visualized as nice neat bands on the mountainsides, however, for roughness of topography and differing tolerances of plant species usually lead to gradual changes from one zone to the next. Because of the greater exposure to drying heat and sunlight, each belt occupies a higher range of elevation on south-facing slopes than on north-facing ones. And along draws, where conditions are wetter and cooler than on ridges at the same elevations, the vegetation zones finger down to lower elevations; on ridges the opposite is true.

Animals are less restricted than plants to particular vegetation belts, but they, too, show a zonation with altitude. You must look, for instance, in the paloverde-saguaro community (part of the desert scrub type) for kangaroo rats and cactus wrens; in the oak-pine woodland for Mexican jays; and in the ponderosa pine forest for whitetail deer and tassel-eared squirrels.

In the Tucson Mountains, only the desert scrub type is well represented, although the highest ridges support a suggestion of desert grassland. A small patch of shrub live oaks, relicts of a wetter period, huddles on the north side of Wasson Peak. Zonation is poorly developed here because the mountains are low and small in mass. (Studies have shown that mountains of smaller mass tend to have warmer and drier climates than bulkier mountains of the same height.)

Major Vegetation Types in Saguaro National Monument

Vegetation type Average Elevations Annual Life zone
July (feet) rainfall
temp. (inches)
Prominent species
Southwestern Desert 94 2,200 to 4,000 7 to 13 Lower Sonoran
Scrub
Paloverde
Saguaro
Pricklypear
Cholla
Catclaw
Ocotillo
Creosotebush
Mesquite
Ironwood[1]
Grassland Transition 85 3,500 to 4,500 10 to 15 Upper Sonoran
Mesquite
Beargrass
Gramagrasses
Amole
Sotol
Oak-pine-juniper 74 4,500 to 7,000 12 to 22 Upper Sonoran
Woodland and
Chaparral
Emory Oak
Mexican Blue Oak
Shrub Live Oak
Pinyon Pine
Mountain-mahogany
Manzanita
Skunkbush
Coniferous Forest 68 above 6,000 18 to 30 Transition and
start of Canadian
Ponderosa Pine
Gambel Oak
Buckbrush
Mountain Muhly
Douglas-fir
White Fir
Aspen
Snowberry
Mexican White Pine

[1]TUCSON MOUNTAIN SECTION

Biologists have developed several systems for classifying assemblages of plants and animals over broad regions, and one of the most widely known is that of C. Hart Merriam. Around the turn of the century, he conceived a system of “life zones,” named for the parts of the continent where they are best developed: Tropical, Lower Sonoran, Upper Sonoran, Transition, Canadian, Hudsonian, and Arctic. In Saguaro National Monument, the Lower Sonoran Zone corresponds to the desert scrub type; the Upper Sonoran includes desert grassland transition, oak woodland, and oak-pine woodland; the Transition Zone is equivalent to the ponderosa pine forest; and the poorly represented Canadian Zone has Douglas-fir and white fir. Because Merriam’s system is so widely used, his terms are included in the tabulation on page 20, which summarizes the main characteristics of plant zones in the monument.

The best way to appreciate the biotic changes that occur with elevation is to walk or ride a horse to the top of the Rincons. But if your time or energy is limited you can get a quick view of these sequences by driving up nearby Mount Lemmon, in the Santa Catalina Mountains.

desert plants

When you look out over the cactus forest, in either part of Saguaro National Monument, you may think there’s a sameness to it in all directions—saguaros standing amid scattered shrubs. But look harder, or walk about, and you will discover variations in the scene. First, there is a gradual change in the vegetation from the mountain foot down the bajada or pediment to the valley floor, as saguaros and paloverdes (green) become sparser and creosotebushes (smaller and brownish) take over. (The monument itself does not extend far enough from the mountain bases to include extensive creosotebush communities, but these cover large areas in the lowest parts of the valleys.) Then there is the luxuriant growth along washes, where mesquites and paloverdes grow to tree size and there are more kinds of plants. And if you are observant you may notice slight variations with each change in slope on the rolling hills—for example, more grass growing on their north sides. On another scale, you can see separate little communities of plants in special situations, as under shrubs or on rocks.

These patterns are due to variations in the environment. For the desert is hotter and drier in some places than in others. The gradual downslope changes in vegetation reflect the decrease in the amount of soil moisture available to plants—a condition caused by the decreasing size of soil particles and consequent shrinkage of water-holding space between particles. Desert washes encourage plant growth because they channel water and cold air. Strangely enough, night temperatures are often lower in a desert valley than farther up the slopes. This “inversion” is due to drainage of cold air down mountainsides, forming cold “pools” in valley bottoms, especially in winter. Cold air is heavier than warm, and (since air flows much like water) it is channeled down the draws—a fact that will strike you if you walk into a wash near the mountains at night or early in the morning. Add this phenomenon to the great amounts of moisture that lie beneath the surface of washes, and you can see that desert drainageways are really linear oases. More subtly, desert hills reflect in their vegetation the differences in soil moisture from north slopes to south slopes caused by increasing exposure to sunlight.

One controlling factor, then, is dominant in the desert: the scarcity of water. This results not only in the unusual forms and adaptations of desert plants, but also in a distinctive type of plant community. In wetter climes, plants compete mostly for sunlight. They can grow close together, as long as they receive adequate illumination. During the regrowth of forests, several distinct sets of plants appear, each succeeding group more tolerant of shade than the last, as the forest canopy closes. In the desert, there is no such succession. Clear a patch of desert vegetation, and the same species will reappear—spaced out, with bare ground between them as before. For here sunlight is abundant (we might say overabundant), but there is not enough water to allow plants to cover the ground.

Dr. Forrest Shreve, who was a botanist at the Carnegie Institution’s Desert Laboratory near Tucson and a master student of deserts, defined a desert as “an area in which deficient and uncertain rainfall ... has made a strong impression on the structure, functions, and behavior of living things.” The distinctive characteristics of desert plants and animals have evolved through millions of years, in a trial-and-error process in which only the better-fitted forms have survived. It would be enlightening to know how many of the species and varieties of plants that developed during the past 60 million years or so have failed to adapt to Sonoran Desert conditions. It is fascinating to study the hundreds of forms that have succeeded and to try to determine what structures they have perfected and what methods they have originated that enabled them to maintain themselves in such a harsh environment.

Desert plants can be classified as “escapers,” “evaders,” and “resisters,” according to their means of adaptation to water shortage. Escapers, such as the annuals, avoid the problem entirely by waiting out the dry periods as seeds, to sprout and reproduce only when the rains come. Evaders, such as the ocotillo, reduce their water loss during droughts by such methods as dropping their leaves or going into a state of dormancy. Resisters, however, “hang in there” all year, taking the desert’s worst. The cactuses, prime examples of this group, rapidly soak up water from each rain and store it for use during drought; the mesquite’s deep roots tap a more constant source of moisture.

Succulents

A large group of desert plants conserve water for use in periods of drought by storing it in specialized tissues during the wet season. Some of these “succulents,” principally the yuccas, eschevarias, and agaves, have developed water-storage tissues in their leaves. A few, notably the NIGHTBLOOMING CEREUS (see appendix for scientific names of plants), have slender stems but an enormous, carrot-shaped root in which the moisture-storage tissue is located. The GOURDS also have large, thick, moisture-retaining roots, as does the WILD-CUCUMBER.

The cactuses store water in their stems and thus are called stem succulents. In the course of its evolution the cactus has eliminated leaves, and their function has been taken over by the green outer covering of the stems. Thus the amount of transpiration (moisture loss through plant breathing pores) has been greatly reduced.

The cactuses are thought to have evolved from relatives of the rose family in the West Indies, beginning some 18,000 to 20,000 years ago. From there they spread to most parts of the Western Hemisphere, but particularly to the drier regions, changing their forms to meet new conditions. One of the youngest of plant families, the cactuses are still evolving rapidly. This doesn’t make the task of classifying them easier for the taxonomists.

In varied forms, cactuses enliven the paloverde-saguaro community. In size they range from tiny button and pincushion types, some of which weigh only a few ounces, to the giant saguaro, the bulkiest of which may weigh several tons.

Cactuses as a group are easily recognized, although many people mistakenly believe that any desert plant with spines or thorns is a cactus. Shreve describes their main characteristics thus:

Several structural features have served to give the cacti their
outstanding appearance, so unlike that of other plants. Most general
have been the loss of the leaf as a permanent organ, the enlargement
of the stem to accommodate water-storing tissue, and the development
of local spinebearing structures known as “areoles.” In several
genera, the stem is segmented into sections which are flat and
somewhat leaflike; in others the stem is round, much branched, and the
surface occupied by close-set tubercles. In a large group, including
massive erect forms, as well as slender climbing ones, the stem is
grooved or fluted and thus able readily to accommodate its surface to
great fluctuations in the water content of the tissues.

If you are trying to identify species, however, cactuses can be annoying, since they often hybridize. You must expect to find some individuals that don’t fit the book descriptions.

_The Saguaro—Monarch of the Monument_

Of all the species of cactus recorded in Saguaro National Monument, the giant SAGUARO (pronounced sah-WAH-roe) holds the center of interest. From the visitor’s standpoint, all other plants, no matter how bizarre in appearance or peculiar in living habits, are merely stage scenery for setting off the star of the desert drama. For size, this vegetable mammoth tops all other succulents of this country; heights of more than 50 feet and weights of more than 5 tons have been reported. There have been specimens with more than 50 arms, or branches. Although no accurate method of determining saguaro age has been devised, it is estimated that an occasional veteran may reach the two-century mark.

Structurally, the giant cactus is well adapted to meet the stern requirements of its habitat. Its widespread root system, as much as 70 feet in diameter, lying close to the surface of the ground, anchors and holds the heavy plant erect. The shallow root system quickly and efficiently collects and channels to the main stem any moisture that may penetrate the topsoil. The trunk and branches have a cylindrical framework of long slender poles or ribs fused at the constricted base. This woody skeleton supports the mass of pulpy tissue, the whole being covered with tough, waxy, spine-bearing “skin.” Numerous vertical ridges, like the pleats of a huge accordion, permit the stems and branches to expand in girth as the tissues swell with water during wet weather and to shrink during times of drought.

So efficient is the saguaro’s water-storage system that, even after years of extreme drought, the plant retains enough moisture in reserve to enable it to produce flower buds. The buds appear in vertical rows at the tips of the main stem and branches, a few opening each evening over a period of several weeks in May and June. The flowers, up to 4 inches in diameter, have waxy white petals. This beautiful blossom is the State flower of Arizona. The egg-shaped fruits mature in late June and July, splitting open when ripe to reveal masses of juicy, deep-red pulp filled with tiny black seeds. Pulp and seeds are consumed by several kinds of birds, especially white-winged doves. Many fruits that fall to the ground are promptly eaten by rodents and other animals.

Indians, too, eat the fruits. European explorers who followed Coronado’s expedition into this region found peaceful Papagos and Opatas living here, hunting animals and utilizing many native plants. Among the most dependable of Papago food sources was the fruit of the giant cactus. So important was this fruit harvest in their economy that they designated this season as the start of the new year. Today, in some parts of the desert, Pima and Papago Indians still harvest the fruits. The word “saguaro” is believed to derive from a Spanish corruption of a Papago word for the big cactus.

Saguaros provide not only food for man and beast, but homes for animals. Walk through a giant cactus forest and you will be amazed at the number of holes drilled in these plants. The holes are made by Gila woodpeckers and related gilded flickers, which often relinquish them after one nesting season. The next occupant may be any of a host of desert dwellers, including screech and elf owls, purple martins, and invertebrates. Some birds use the plant as a foundation for their homes. White-winged doves, for instance, often build flimsy stick platforms on the tips of saguaro arms; red-tailed hawks and horned owls construct more substantial nests in the forks.

Although billions of saguaro seeds are produced yearly in the extensive stands of the monument, only a very few find favorable locations for germination and growth. Trees, rocks, dead limbs, pebbles—anything that reduces evaporation in the immediate vicinity of the seed—improve the chance of germination. These kinds of shelter not only provide the necessary moisture conditions, but also hide the seed from armies of ants, rodents, and other animals searching for food.

Early growth is extremely slow. A 2-year-old saguaro may be only one-quarter of an inch in diameter, and a 9-year-old plant may be 6 inches high. These years are the most hazardous. Insect larvae devour the tiny cactuses. Woodrats and other rodents chew the succulent tissue for its water, and ground squirrels uproot the young plants with their digging. In later life, the saguaro must contend with uprooting wind and human vandalism, as well as the earlier foes—drought, frost, erosion, and animals.

In a century of maturity, a saguaro may produce 50 million seeds; replacement of the parent plant would require only that one of these germinate and grow. But in the cactus forest of the Rincon Mountain Section, the rate of survival has been even lower, so that over the last few decades the stand has been dwindling. What is wrong?

Many answers to this question have been advanced, but like all interrelationships in nature, the saguaro’s role in the desert web of life is very complex, and involves past events as well as present ones; a partial answer to the problem may be all we can hope for. The following reasons for the decline of the saguaros have been suggested by researchers.

There is some evidence to suggest that the Southwest has been getting drier since at least the late 19th century, and while the saguaro is adapted to extreme aridity, some of the “nurse” plants that shelter it during infancy are not. If such shrubs as paloverdes and mesquites dwindle, it is argued, so must the saguaro, which in its early years depends on them for shade.

Other culprits in the saguaro problem are man himself and his livestock. Around 1880, soon after the first railroad reached Tucson, a cattle boom began in southern Arizona. The valleys were soon overstocked, and cattle scoured the mountainsides in search of food. By 1893, when drought and starvation decimated the herds, the land had been severely overgrazed. Though the monument was established in 1933, grazing in the Rincon Mountain Section’s main cactus forest continued until 1958. (Elsewhere in the monument, it still goes on.) Compounding the problem, woodcutters removed acres of mesquite and other trees. In the center of the present Cactus Forest Loop Drive, lime kilns devoured quantities of woody fuel. Further upsetting the desert’s natural balance, ranchers and Government agents poisoned coyotes and shot hawks and other predators—in the belief that this would benefit the owners of livestock.

This unrestrained assault on the environment had unfortunate effects on saguaros as well as on the human economy. Overgrazing may have resulted in an increase in kangaroo rats (which benefit from bare ground on which to hunt seeds) and certain other rodents adapted to an open sort of ground cover. Man’s killing of predators, their natural enemies, further encouraged proliferation of these rodents, which some people say are especially destructive of saguaro seeds and young plants. Whatever the effect these rodents have on the saguaros, the removal of ground cover intensified erosion and reduced the chances for the seeds to germinate and grow. And certainly the cutting of desert trees removed shade that would have benefited young saguaros. In the Tucson Mountain Section, which is near the northeastern edge of the Sonoran Desert, freezing temperatures are perhaps the most important environmental factor in saguaro mortality.

Although the causes of decline of the cactus forest lying northwest of Tanque Verde Ridge are still something of a puzzle, several facts are clear: the saguaro is not becoming extinct; in rocky habitats many young saguaros are surviving, promising continued stands for the future; in non-rocky habitats, some young saguaros are surviving, ensuring that at least thin stands will endure in these areas. Furthermore, since grazing was stopped here, ground cover has improved—a plus factor for the saguaro’s welfare. On the negative side, it is possible that, in addition to suffering from climatic, biotic, and human pressures, the once-dense mature stands of the monument are in the down-phase of a natural fluctuation. It is possible, too, that these stands owed their exceptional richness to an unusually favorable past environment which may not occur again. We can hope, however, that sometime in the not-distant future the total environmental balance will shift once again in favor of the giant cactus.

_Other Common Cactuses_

Many other cactuses share the saguaro’s environment. The BARREL CACTUS is sometimes mistaken for a young saguaro, but can easily be distinguished by its curved red spines. Stocky and unbranching, this cactus rarely attains a height of more than 5 or 6 feet. It bears clusters of sharp spines, called “areoles,” with the stout central spine flattened and curved like a fishhook. In bloom, in late summer or early autumn, this succulent plant produces clusters of yellow or orange flowers on its crown. The widely circulated story that water can be obtained by tapping the barrel cactus has little basis in fact, although it is possible that the thick, bitter juice squeezed from the plant’s moist tissues might, under extreme conditions, prevent death from thirst. Desert rats, mice, and rabbits, carefully avoiding the spines, sometimes gnaw into the plant’s tissues to obtain moisture.

The group of cactuses called opuntias (oh-POON-cha) have jointed stems and branches. They are common and widespread throughout the desert and are well represented in the monument.

Those having cylindrical joints are known as chollas (CHO-yah), while those with flat or padlike joints are called pricklypears.

Chollas range in size and form from low mats to small trees, but most of those in the monument are shrublike. TEDDY BEAR CHOLLA, infamous for its barbed, hard-to-remove-from-your-skin spines, forms thick stands on warm south- or west-facing slopes. Its dense armor of straw-colored spines and its black trunk identify it. Because its joints break off easily when in contact with man or animal, this uncuddly customer is popularly called “jumping cactus.” A similar species is CHAIN FRUIT CHOLLA, notable for its long, branched chains of fruit, which sometimes extend to the ground. Each year, the new flowers blossom from the persistent fruit of the previous year. There is a common variety of this species that is almost spineless. STAGHORN CHOLLA, an inhabitant primarily of washes and other damp places, is named from its antler-shaped stems. This cactus’ scientific name—_Opuntia versicolor_—refers to the fact that its flowers, which appear in April and May, may be yellow, red, green, or brown. (Each plant sticks with one color through its lifetime.) Among the smaller chollas, thin-stemmed PENCIL CHOLLA grows from 2 to 4 feet high on plains and sandy washes. DESERT CHRISTMAS CACTUS, almost mat-like in form, blooms in late spring but develops brilliant red fruits which last through the winter.

PRICKLYPEARS, like many of the chollas, produce large blossoms in late spring. Those on the monument are principally the yellow-flowered species. The reddish brown-to-mahogany colored edible fruits, called tunas, attain the size of large strawberries. When mature in autumn, they are consumed by many desert animals.

Some of the smaller cactuses are so tiny as to be unnoticeable except when in bloom; examples are the HEDGEHOGS, the FISHHOOKS, and the PINCUSHIONS. Blossoms of some of these ground-hugging species are large, in some cases larger than the rest of the plant, and spectacular in form and color. All add to the monument’s spring and early summer display of floral beauty.

Non-Succulents

For the diversity of devices for adaptation to an inhospitable environment, the many species making up the non-succulent desert vegetation provide an absorbing field for study. As we have seen, there are two ways to survive the harsh desert climate; one is to avoid the periods of excessive heat and drought (“escapers”); the other is to adopt various protective devices (“evaders” and “resisters”). Short-lived plants follow the first method; perennials, the second.

_Perennials_

Chief among the requirements for year-round survival in the desert is a plant’s ability to control transpiration and thus maintain a balance between water loss and water supply. In this struggle, the hours of darkness are a great aid because in the cool of the night the air is unable to take up as much moisture as it does under the influence of the sun’s evaporating heat. Therefore, less exhaling and evaporating of water occurs from plants, and both the rate and the amount of water loss are reduced. This reduction in transpiration at night allows the plants to recover from the severe drying effects of the day. One biologist may have been close to the truth when he stated, “If the celestial machinery should break down so that just one night were omitted in the midst of a dry season, it would spell the doom of half the nonsucculent plants in the desert.”

One of the common trees in the desert part of the monument is the MESQUITE (mess-KEET). In general appearance it resembles a small, spiny apple or peach tree with finely divided leaves. Its roots sometimes penetrate to a depth of 40 or more feet, thus securing moisture at the deeper, cooler soil levels, from a supply that remains nearly constant throughout the year. This enables the tree to expose a rather large expanse of leaf surface without losing more water than it can replace. A number of mechanical devices help the tree reduce its water loss during the driest part of the day (10 a.m. to 4 p.m.). Among these are its ability to fold its leaves and close the stomata (breathing pores), thereby greatly reducing the surface area exposed to exhaling and evaporating influences. In April and May, mesquite trees are covered with pale-yellow, catkinlike flowers which attract swarms of insects. These flowers develop to stringbeanlike pods rich in sugar and important as food for deer and other animals. In earlier days, the mesquite was also a valuable source of food and firewood for Indians and pioneers.

Another desert tree abundant in the monument is the YELLOW PALOVERDE. It is somewhat similar in size and general shape to the mesquite. Lacking the deeply penetrating root system of the mesquite, the paloverde (Spanish word meaning “green stick”) has no dependable moisture source; but it has made unusual adaptations that enable it to retain as much as possible of the water collected by its roots. In early spring the tree leafs out in dense foliage, which is followed closely by a blanket of yellow blossoms. At this season the paloverdes provide one of the most spectacular displays of the desert, particularly along washes, where they grow especially well. Blue paloverde, growing in the arroyos, blooms well every year. Yellow, or foothill, paloverde, a separate species, blooms only if the soil moisture is high following winter rains.

With the coming of the hot, drying weather of late spring, the trees need to reduce their moisture losses. They gradually drop their leaves until, by early summer, each tree has become practically bare. The trees do not enter a period of dormancy, but are able to remain active because their green bark contains chlorophyll. Thus, the bark takes over some of the food-manufacturing function normally performed by leaves, but without the high rate of water loss.

Carrying the drought-evasion habits of the paloverde a step further, the OCOTILLO (oh-koh-TEE-yoh) comes into full leaf following each rainy spell during the warmer months. During the intervening dry periods it sheds its foliage. The ocotillo, a common and conspicuous desert dweller, is a shrub of striking appearance, with thorny, whiplike, unbranching stems 8 to 12 feet long growing upward in a funnel-shaped cluster. In spring, showy scarlet flower clusters appear at the tips of the stems, making each plant a glowing splash of color.

A number of desert shrubs fail to display as much ingenuity as the paloverde. Some of these evade the dry season simply by going into a state of dormancy. The WOLFBERRY bursts into full leaf soon after the first winter rains and blossoms as early as January. Its small, tomato-red, juicy fruits are sought by birds, which also find protective cover for their nests and for overnight perches in the stiff, thorny shrubs. In the past, the berrylike fruits were important to the Indians, who ate them raw or made them into a sauce.

Commonest of the conspicuous desert non-succulent shrubs is the wispy-looking but tough CREOSOTEBUSH, found principally on poor soils and on the desert flats between mountain ranges. It is also sprinkled throughout the paloverde-saguaro community in the monument. A new crop of wax-coated, musty-smelling leaves, giving the plant the local (but mistaken) name “grease-wood,” appears as early as January. The leaves are followed by a profuse blooming of small yellow flowers and cottony seed balls. During abnormally moist summers or in damp locations, the leaves and flowers persist the year round; but usually the coming of dry weather brings an end to the blossoming period. If the dry spell is exceptionally long, the leaves turn brown, and the plants remain dormant until awakened by the next winter’s rainfall. Pima Indians formerly gathered a resinous material, known as lac, which accumulates on the bark of its branches, and used it to mend pottery and fasten arrow points. They also steeped the leaves to obtain an antiseptic medicine. Ground squirrels commonly feed on the seeds.

A large shrub of open, sprawling growth usually found along desert washes in company with mesquite is CATCLAW. Its name refers to the small curved thorns that hide on its branches. In April and May, the small trees are covered with fragrant, pale-yellow, catkinlike flower clusters that attract swarms of insects. The seed pods were ground into meal by the Indians and eaten as mush and cakes.

In lower elevations of the Tucson Mountain Section, the gray-blue foliage of IRONWOOD is a common sight, but the species does not range farther eastward. Its wisterialike lavender-and-white flowers blossom in May and June. The nutritious seeds are harvested by rodents and formerly were parched and eaten by Indians. The wood is so dense that it sinks in water; Indians used it for making arrowheads and tool handles.

Ferns—commonly, plants of dank woods and other moist habitats—seem entirely out of place in the desert; nevertheless, some members of the fern family have overcome drought conditions. The GOLDFERN is common on rocky ledges, where it persists by means of special drought-resistant cells.