The history of climate and vegetation for the last 60 million
years provide insight into the ecology and physiology of the modern
Gopherus agassizii. Both Gopherus and evidence for tropical deciduous
forest appeared in the fossil record in the Eocene (55 mya, million
years ago). Climates were tropical with only wet and dry seasons
and little temperature difference. Tortoises were likely active
in warm-wet periods, inactive in warm-dry periods, ate fresh and
dried understory warm-season herbs and grasses, and primarily
used sheltersites to avoid predators. The uplift of the Rocky
Mountains and the Sierra Madre Occidental in the late Oligocene-middle
Miocene (25-15 mya) established the modern biotic and climatic
provinces in North America. New climate limited biomes (thornscrub,
grassland, conifer forests, tundra, etc.) formed along environmental
gradients. The modern subgenera (genera) diverged primarily involving
enhanced burrowing behavior in Gopherus compared to the more conservative
Xerobates. With the development colder temperatures at higher
latitudes and elevations, temperate winter and a four-season climate
developed. In these areas, tortoise seasonal activity patterns,
diet and behavior were little changed except that part of the
extended inactive period due to cold (hibernation) and sheltersite
were increasingly important environmental buffers. The drying
trend that resulted in the formation of the Sonoran Desert (by
8 mya) resulted in the modern five-season climate (winter, spring,
foresummer, summer, fall). Selection likely intensified spring
inactivity and foresummer estivation. The reversal to more tropical
climates in the Pliocene (5-2.4 mya) and the northward expansion
of the Gulf of California (Bouse Formation) would not have resulted
in major life history evolution. In the Pleistocene, new climatic
regimes developed with much cooler summers and the cool season
greatly expanded in spring and fall resulting in a general contraction
(for the first time) of monsoonal rainfall from the tropical oceans
and widespread expansion of Mediterranean climates and biotas.
For 85-90% of the last 2.4 mya, glacial climates allowed woodland
and chaparral to expanded into desert elevations; Mohave Desert
species such as Yucca brevifolia (Joshua tree) were widespread.
The Mohave Desert climate can be viewed as a reversal to the original
Eocene two-season regime (wet, dry) with two devastating differences:
(1) the rains fall in the cool season instead of the warm season
leaving the summers very hot and dry and (2) winter temperatures
are cold. Tortoises adapted to this new environment in several
important ways: (1) building more extensive burrows to buffer
extreme cold, heat, and aridity, (2) shifting primary activity
period from hot-wet summer to relatively-cool spring rainy season,
(3) shift of diet from tropical subshrubs, herbs and grasses to
spring annuals, mostly of temperate origins, and (4) a related
shift from general consumption to general avoidance of dried plant
foods. In this deep historical context, it is clear that the Mohave
desert tortoise is the most recently evolved North American tortoise
and likely the only one to have adapted to a winter rainfall climate.