The Deep
History of the Sonoran Desert
Thomas R.
Van Devender, AZ/Sonora Desert Museum
The
Sonoran Desert is considered to be the most “tropical” of the North American
deserts. Its climate is virtually frost-free, and summer rainfall comes from
the tropical oceans. The Sonoran Desert’s structurally diverse vegetation,
which includes columnar cacti and leguminous trees, certainly differs from
those of the shrub-dominated Great Basin, Mohave and Chihuahuan deserts. It has
both geographic and biologic connections with more tropical communities. In a
single day’s travel, naturalists can begin in the oak woodlands and desert
grasslands in southern Arizona along Interstate 19, and travel through the
various desertscrub, thornscrub, and tropical deciduous forest habitats along
México 15 in Sonora, experiencing the remarkable transition from temperate zone
communities to the New World tropics. This vegetational gradient gives us a
sense of the Sonoran Desert’s connection with the tropics, but does not really
explain it. The explanation lies in the tropical roots of the Sonoran Desert,
deep in its evolutionary history.
A Walk
through Time
The
Sonoran Desert that we see today, with its characteristic assemblages of plants
and animals is quite recent, at least in terms of geologic time. In fact, it
and the other North American deserts are among the youngest biotic communities
on the continent. Although some Sonoran species evolved in ancestral
seasonally-dry tropical communities, the development of the unique regional
climates and the evolution of characteristic desert-adapted plants and animals
are thought to have combined to form the Sonoran Desert by about 8 million
years ago (mya) in the late Miocene. Similar conditions developed many times
subsequently as global climates changed, with the Sonoran Desert continually
expanding, contracting, and redefining itself. The most recent expansion of the
Sonoran Desert into its modern area in Arizona and California occurred only
9000 years ago, with the modern communities of plants and animals developing
4500 years later. This chapter is a walk through time examining the conditions
that led to the development of the Sonoran Desert and exploring what shaped its
dynamic history.
The
Paleocene
(66.4 to 57.8 mya)
In the
Paleocene epoch, soon after the extinction of the dinosaurs (65 mya), most of
North America was covered with temperate evergreen and tropical rainforests.
There was little regional variation. The warm climates promoted humid forests
with strong Asian affinities; primitive ferns (Anemia spp.), cycads (Diön, Zamia spp.), and palms grew as far north as Alaska. The flowering
plants (angiosperms), whose spectacular evolutionary radiation began in the
Late Cretaceous, became increasingly important in the forests, displacing
archaic cycads, conifers, and tree ferns. The earliest indisputably
recognizable fossil grasses were found in sediments dating to about 58 mya.
These were broad-leaved forest grasses ancestral to the modern bamboos.
The
Eocene
(57.8 to 36.6 mya)
In the
Eocene, deciduous trees became increasingly common, providing the first
evidence of a dry season. The landscape now included tropical deciduous
forests, in which trees dropped their leaves in response to drought. During
these periods, sunlight passed through the leafless canopy and heated and dried
the surface of the ground. Many new species of plants and animals evolved,
adapting to these new heat and moisture regimes. The origins of cacti and other
succulents likely occurred during the Eocene in dry tropical forests. Fossils
of an alligator (Allognathosuchus), a softshell turtle (Trionyx),
a primitive tortoise (Geochelone), a primitive monitor lizard (Varanidae), a ground boa (Boidae), and many small mammals from
Ellesmere Island in northeastern Canada, then at 78° latitude, indicate that
the world was very warm and that plants and animals freely traversed a land
bridge between North America and Europe. The results were dramatic shifts in
the biota as more advanced forms displaced archaic ones.
The
Oligocene
(36.6 to 23.7 mya)
By the
Oligocene, grasses, including the important taxonomic groups in arid lands, had
achieved relatively modern diversity. Unfortunately, most of this dramatic
radiation in the grasses, one of the most important plant families, was not
captured in the fossil record. Gopherus, the genus of the modern
desert and gopher tortoises (G. agassizii and G. polyphemus), appeared in the Oligocene. Modern genera of lizards in
the Oligocene fauna were skinks (Eumeces) and beaded lizard or Gila
monster (Heloderma). The snake fauna was
dominated by small ground boas related to the living rubber boa (Charina bottae) and desert rosy boa (Lichanura trivirgata) of western
North America and the sand boas (Eryx spp.) of Africa. The lizards
common today in the Sonoran Desert—Iguanidae and Teidae, (that is, iguanas and
their relatives, and whiptails)—and the common snakes of the Sonoran Desert
today—Colubridae and Viperidae (colubrid snakes, for example, bull snakes,
king-snakes; and pitvipers) were uncommon or absent at these early dates.
The Miocene
(23.7
to 5.3 mya)
A series
of enormous volcanic eruptions from the middle Oligocene to the middle Miocene
(about 30 to 15 mya) changed the climates and established the modern
biogeographic provinces of North America. (See the chapter “The Geologic Origin
of the Sonoran Desert” for a more complete discussion of these processes and
the terminology used to describe them.) The Rocky Mountains were uplifted to
new heights by the accumulation of over a kilometer and a half of volcanic
rock. A kilometer (.6 mile) thick layer of rhyolitic ash fell in the Sierra
Madre Occidental in northwestern Mexico—on top of a kilometer of early Tertiary
(the geologic period that includes the Paleocene, Eocene, Oligocene, Miocene,
and Pliocene epochs) andesites. As regional uplift pushed them even higher, the
mountains interrupted the upper flow of the atmosphere for the first time.
Tropical moisture from both the Pacific Ocean and the Gulf of Mexico was
blocked from the mid-continent, drying out the modern Great Plains and the Mexican
Plateau. Harsher climates segregated drought- and cold-tolerant species into
new environmentally- limited biomes, including tundra, conifer forests and
grasslands, and restricted them along elevational and latitudinal environmental
gradients. The Miocene also was a time when major evolutionary radiations began
in many of today’s successful groups including composites (plants in the
sunflower family), grasses, toads, iguanid and teid lizards, colubrid snakes
and pitvipers.
The “Miocene
Revolution”
After the
rise of the Sierra Madres and the resulting changes in climate, tropical
forests were found only in the lowlands along the coasts of Mexico and Central
America. There, newly evolved species joined archaic ones in the biome known as
tropical deciduous forest. And, along the lower, drier edges of tropical
deciduous forest evolved a new biome—thornscrub. Thornscrub looks very much
like a transitional state between tropical deciduous forest and Sonoran Desert:
its vegetation is shorter and sparser than tropical forest and it does not
require much moisture. During the Miocene, thornscrub may well have been the
regional vegetation in drier areas to the north that are now Sonoran Desert.
Thornscrub, in fact, may be the ancestral biome of many Sonoran Desert plants
and animals.
The
Sonoran Desert itself came about during a drying trend in the middle Miocene
(15 to 8 mya). Much of the desert’s vegetation, however, predates the Sonoran
Desert itself, having evolved in climates that also required adaptations to aridity.
For example, guayacán (Guaiacum coulteri), organpipe cactus (Stenocereus thurberi), palo brea (Cercidium praecox), senita (Lophocereus schottii), and tree ocotillo (Fouquieria
macdougalii) likely
evolved in thornscrub. Other plants such as desert ironwood (Olneya tesota), foothills palo verde (Cercidium
microphyllum), and
saguaro (Carnegiea gigantea) evolved along with the
Sonoran Desert.
Another
important chapter in the history of the Sonoran Desert concerns the formation
of the Baja California peninsula. Before 12 million years ago, much of the land
which is now Baja California was part of the Mexican mainland. Activity along
the San Andreas fault caused the Gulf of California to open, and several large
chunks broke away from the mainland and drifted in splendid isolation
northwestward. The timing of the formation of Baja California is controversial,
with some estimates as recent as 5 to 6 million years ago. These islands were
populated with tropical plants and animals which soon evolved into regional
endemics, including the boojum tree or cirio (Fouquieria
columnaris).
Eventually the islands combined and joined California to form the Baja
peninsula. Today, the modern islands on both sides of the peninsula in the
Pacific Ocean and the Gulf of California, many of which are geologically young,
are the most active evolutionary arenas in the Sonoran Desert Region, and have
many endemic species.
The
Pliocene
(5.3 to 1.8 mya)
During
the latest Miocene and early Pliocene, geological forces again altered landscapes
and climate regimes, causing a reversal to more tropical climates. Sea level
rose enough that the Gulf of California expanded into the Los Angeles area of
southern California. A fossil skull of an iguana (Pumilia
novaceki), a
primitive relative of the tropical green iguana (Iguana
iguana) that
today occurs no farther north than southern Sinaloa, was found in sediments 2.5
to 4.3 million years old in southern California. With tropical circulation
patterns enhanced by warmer oceans, tropical forests of western Mexico likely
expanded, reaching farther north than they do today in Sonora; likewise, the
Sonoran Desert in Arizona and California extended further, perhaps as far as
southern Nevada.
The
Pleistocene
(1.8 mya to today)
The
warmth of the Pliocene ended abruptly at the beginning of the Pleistocene about
1.8 million years ago, as the Earth entered a new climatic era that far
surpassed the middle Miocene in cool, continental conditions. Traditionally,
four ice ages or glacial periods were recognized, based on terrestrial
sedimentary deposits in North America, and these were widely correlated with
conditions in Europe and South America. However, recent studies of sediment
cores from the ocean floors record fifteen to twenty glacial periods in the Pleistocene.
Ice ages were about ten times longer than interglacials (the warm periods
between ice ages) which lasted 10,000 to 20,000 years. Officially, the end of
the Pleistocene was defined as the beginning of the Holocene 10,000 years ago,
based on changes in sediments in European lakes. Today we understand that the
Holocene is the present interglacial period and that the cyclic environmental
fluctuations of the Pleistocene likely have not ended.
In the
last glacial period (the Wisconsin), the massive Laurentide continental glacier
covered most of Canada and extended as far south as New York and Ohio. As much
as two miles (3 km) of ice covered the Great Lakes and New York City. Boreal
forest with spruce and jack pine moved southward displacing the mixed deciduous
forests of the eastern United States. Mountain glaciers covered the tops of the
Rocky Mountains and the Sierra Nevada in the western United States and the
Sierra Madre del Sur in south-central Mexico. Now-dry playa lakes in the Great
Basin were full. Enough water was tied up in ice on land to lower sea level
about 425 feet (130 m).
During
the last half of this glacial period (from 45,000 to 11,000 years ago), plant
remains in ancient packrat (Neotoma spp.) middens document the
expansion of woodland trees and shrubs into areas that had been desert.
Woodlands with singleleaf pinyon (Pinus monophylla), junipers (Juniperus spp.), shrub live oak (Quercus
turbinella), and
joshua tree (Yucca brevifolia) were widespread in the
present Arizona Upland subdivision of the Sonoran Desert. Ice age climates with
greater winter rainfall from the Pacific Ocean and reduced summer monsoonal
rainfall from the tropical oceans favored woody cool-season shrubs related to
plants living farther north, rather than to the summer-rainfall trees, shrubs
and cacti of tropical forests and subtropical deserts. The isolated chaparral
communities in central Arizona, mostly in a northwest-southeast band below the
Mogollon Rim, are relicts of ice-age chaparral connections with California.
Many species are shared between California and Arizona chaparral, including
shrub live oak. The Arizona black rattlesnake (Crotalus
viridis cerberus) in
Arizona chaparral is essentially a dark form of the southern Pacific
rattlesnake (C. v. helleri); it is more distantly related
to the other four subspecies of western rattlesnakes in Arizona.
Warm
desert communities dominated by creosote bush (Larrea
tridentata) were
restricted to below 1100 feet (300 m) elevation in the Lower Colorado River
Valley in the Sonoran Desert and in the southern Chihuahuan Desert. Although
brittlebush and saguaro returned to Arizona soon after the beginning of the
present interglacial (the Holocene) about 11,000 years ago, the Sonoran Desert
did not re-form until about 9000 years ago, as the last displaced woodland
plants retreated upslope. Relatively modern community composition was not
achieved until about 4500 years ago when foothills palo verde, desert ironwood
and organpipe cactus arrived from their retreats to the south and to low
elevations. However, the modern assemblages that we recognize as the Sonoran
Desert communities must have recurred many times during the Pleistocene
interglacials, only to retreat to warmer climates as ice age climates returned.
Modern desert communities have been present for only about five percent of the
2.4 million years of the Pleistocene, while ice age woodlands in the desert
lowlands persisted for about ninety percent of this period.
Tropical
Interglacials
Surprisingly,
the vertebrate fossil record suggests that some interglacial climates were more
tropical during the Holocene. El Golfo de Santa Clara is near the mouth of the
Colorado River in northwestern Sonora. Early Pleistocene (1.8 mya) fossils
reflect a climate that was frost free, with much greater rainfall in the warm
season, and with higher humidity than today. Greater summer rainfall would
suggest that tropical oceans were warmer than they are today, in contrast to
most of the Pleistocene when ocean waters were colder. The fauna included such
mammals as antelope, bear, camels (dromedaries and llamas), cats, horses,
proboscidians, and a tapir (Tapirus). The giant anteater, capybara
(Neochoerus), and ground sloths in the
fauna were members of ten families of mammals that immigrated into North
America in the late Pliocene or early Pleistocene after the opening of the
Panamanian land bridge during the Great American Interchange. In contrast, the
imperial mammoth (Mammuthus imperator), a hyena (Chasmoporthetes johnstoni), and jaguar (Felis onca), were Eurasian immigrants. The nearest populations of
giant anteater are 1800 miles (3000 km) to the southeast in the humid, tropical
lowlands of Central America! As for many large mammals, the modern distribution
may not accurately reflect their physiological range limits because of human
predation in the last 11,000 years. Other fossils in the fauna include the
Sonoran Desert toad (Bufo alvarius), slider “turtle”(Trachemys scripta), boa constrictor (Constrictor
constrictor), and
the large extinct California beaver (Castor cf. C. californicus). The Sonoran Desert toad is a regional endemic, while the
slider and boa constrictor occur today in Sonora in wetter, more tropical areas
to the southeast. Although the El Golfo area is today part of the hyperarid
Gran Desierto, the delta of the Colorado River was historically a very wet area
that supported extensive cottonwood (Populus fremontii) gallery forests with abundant
beaver. There is even a December 1827 account of a large spotted cat (likely a
jaguar) that entered James Ohio Pattie’s camp on the Colorado River south of
Yuma to feed on drying beaver skins.
Rancho La
Brisca is in a riparian stream canyon north of Cucurpé, fifty-four miles (90
km) south of the Arizona boundary in Sonora. A pocket of 150,000-year-old
ciénega sediments yielded abundant Sonoran mudturtle (Kinosternon sonoriense), fish, and other small vertebrate
fossils associated with bison (Bison). The presence of the sabinal frog (Leptodactylus melanonotus) 144 miles (240 km) north of the northernmost
extant population on the Río Yaqui indicates that the climate of a late
Pleistocene interglacial was also more tropical than at the site today.
Sonoran
Desert Mammoths?
A few
years ago, a Mayo Indian found a very large bone in the bank of the arroyo
behind his house in Teachive, a village in coastal thornscrub in southern
Sonora. For him, as it has been for others who have discovered fossil mammoth
bones throughout North America and Europe for centuries, it was puzzling and
perhaps frightening. What animal could be so much larger than a deer or a cow?
Why has no one seen these monsters? Many a legend was born to explain them and
their disappearance.
Today we
know that about 11,000 years ago, nearly two-thirds of the large mammals of
North America went extinct. Common, widespread grazers, including horses and
mammoths, disappeared at the very time that spruce and pine retreated and
grasslands expanded from Arizona to Canada. Paul S. Martin of the University of
Arizona forcefully presented the case that big game hunters caused widespread
extinctions within a few hundred years after their entry into North America
from Siberia via the Bering Strait. The theory of “overkill” of “naive” large
mammals is controversial, and some suggest changes in climate may have caused
the extinctions. However, the paleobotanic record gives no evidence of climatic
changes severe enough to have resulted in the extinction of so many large
animals over such a broad, diverse area. The well-preserved plant remains in
packrat middens provide additional insights. A species could respond to a major
climatic change by (1) adapting genetically (speciation), (2) becoming extinct,
or (3) adjusting its geographical distribution. At the beginning of the
Holocene, the last glacial/interglacialclimatic shift, there are essentially no
records of speciation or extinction in plants or small animals. Most simply
shifted their geographic and elevational ranges. Moreover, woodland plants
survived in desert lowlands for several thousand years after the megafaunal
extinctions and before the expansion of the Sonoran Desert. The biotic
communities of North America have had fewer large herbivores in the last 11,000
years than at any time in the last 20 million years! The impacts of these
herbivores on tropical deciduous forest, thornscrub, and the Sonoran Desert
flora were undoubtedly profound—but we will never fully understand the
ecological roles of these missing animals.
The Arctic
Connection
Tropical
communities with their great species diversity, outrageous morphological
adaptations, and mixtures of archaic and advanced species have been important
evolutionary arenas. However, the evolutionary mechanisms are not so clear,
considering that the climatic fluctuations that isolate populations and stimulate
the evolution of new species are more intense at high latitudes. Recent
paleomagnetic dating of fossil-bearing sediments in northern Canada indicate
that some plants appeared there as much as 18 million years earlier, and some
mammals two to four million years earlier than at lower latitudes. If the
relative ages are not the result of a fragmentary fossil record, they may
indicate that important biotic innovations evolved in the mild “tropical”
Arctic climate with its months-long polar day-night cycle, and then the species
moved southward into the tropics.
Fossil
records of tropical plants and animals from Arctic latitudes with
six-month-long nights not only reflect very warm global climates but raise
questions about how these organisms survived the dark. Today reptiles spend the
cold winters in hibernation and hot, dry periods in estivation. Deciduous
plants shed their leaves for long periods, either triggered by the onset of
cold temperatures in temperate latitudes or the beginning of the dry season in
tropical latitudes. The Arctic fossils suggest that deciduousness in plants and
hibernation and estivation in reptiles could have arisen as responses to the
polar night, and later shifted to these other stimuli.
Whose Child
Is This?
Although
the perception of the Sonoran Desert as “tropical” is based partly on the
presence of columnar cacti, there are climatic, physical, and deep historical
connections as well. As discussed above, the region likely supported tropical
deciduous forest from the Eocene to the early Miocene and then thornscrub later
in the Miocene. The desert biota of this region are rich in endemics, most of
which evolved in more tropical communities prior to the Sonoran Desert itself.
Many Sonoran species reach their southern limits in thornscrub—the structural,
biotic, and historical link to tropical deciduous forest—not in the tropical
deciduous forests of southern Sonora. With the exception of organpipe cacti and
a few others, the paucity of desert species shared with the tropical deciduous
forests challenges the popular idea that “tropical deciduous forest is the
mother of the Sonoran Desert.”
Thornscrub
could more accurately be called the “mother” of the Sonoran Desert and tropical
deciduous forest its “grandmother” or “great aunt”!
Additional
Readings
Bahre,
Conrad J. A Legacy of Change. Historic Human
Impact on Vegetation in the Arizona Borderlands. Tucson: University of Arizona
Press, 1991.
Betancourt,
Julio L., Thomas R. Van Devender, and Paul S. Martin. Packrat Middens. The Last 40,000 Years of Biotic Change. Tucson: University of Arizona
Press, 1990.
Davis,
Goode P., Jr. Man and Wildlife in
Arizona: The American Exploration Period 1824-1865. Phoenix: Arizona Game and
Fish Department, 1982.
Grayson,
Donald K. The Desert’s Past. A Natural Prehistory
of the Great Basin.
Washington, D.C.: Smithsonian Institution Press, 1993.
Hastings,
James R. and Raymond M. Turner. The Changing Mile. Tucson: University of Arizona
Press, 1965.
Imbrie,
John and Kathryn P. Imbrie. Ice Ages. Solving the
Mystery.
Cambridge: Harvard University Press, 1986.
Kurtén,
Bjorn and Elaine Anderson. Pleistocene Mammals of
North America. New
York: Columbia University Press, 1980.
Martin,
Paul S. and Richard G. Klein, eds., Quaternary Extinctions:
A Prehistoric Revolution.
Tucson: University of Arizona Press,
1989.
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