Educational Bulletin #09-2
A publication of the Desert Protective Council www.dpcinc.org
Are Joshua trees facing extinction?
by Chris Clarke
At California’s Saddleback Butte State Park in the eastern Antelope Valley, no new Joshua trees
have grown in the last 20 years. Chris Clarke photo
T
he emblematic tree of the Mojave Desert is in big
trouble. That’s probably not news to you if you read
the papers. News organizations from the Riverside
Press-Enterprise to National Public Radio are cover-
ing the increasing threat to the Joshua tree, Yucca
brevifolia, from human-caused climate change. At a February,
2009 climate symposium in Joshua Tree sponsored by the Nation-
al Parks Conservation Association, environmental activists and
biologists and Park staff gathered to discuss the fate of the North
American deserts in a world with increased global temperatures.
Those in attendance spoke with gallows humor about the eventual
need for Joshua Tree National Park (JTNP) to change its name.
The way things are going, it looks as if there will be no Joshua trees
left in the park before too many more decades go by.
Politically motivated misinformation on climate change is still
ubiquitous in mainstream media, giving the false impression that
the jury is still out as to whether human activity is changing the
global climate. Despite this misinformation the scientific com-
munity is as near-unanimous as it ever gets: our releasing of car-
bon and other greenhouse gases into the atmosphere has already
changed our climate, and widespread catastrophe is assured un-
less we make significant changes in the way we run our society.
Just how climate change will play out in a particular place in
the desert is far harder to determine. Future temperatures are
hard enough to predict. Though human-caused climate change
is popularly called “global warming,” some places in the desert
southwest may actually see cooler average temperatures in years
to come due to changing cloud cover or prevailing winds. Trying
to foresee the effects of global climate change on a particular spe-
cies is even more difficult. In addition to temperature, one must
anticipate changes in available precipitation, the effect of higher
concentrations of atmospheric CO2, and the secondary and ter-
tiary effects of those changes on a thousand other species that
compete with, predate on, or assist the species in question.
Still, we do know a great deal about the factors now threatening
to weaken the tenuous hold Joshua trees have on survival.
Desert plants, with their hard-won adaptations to heat and
drought, would seem well-suited to survive a warmer world. But
desert plants are adapted to thrive in the range of conditions
in which they have evolved. Change those conditions, and you
change the plants’ chances of survival, of successful reproduction.
Those conditions aren’t limited to temperature and precipitation,
but also include the web of relationships the plant species has with
a hundred other species, plant and animal.
On the wings of a moth
The welfare of the species Yucca brevifolia depends utterly on the
welfare of a small and inconspicuous insect. Like all their fellow
yuccas, Yucca brevifolia depend for their reproductive success on
small moths. The moths and the yuccas have evolved a remark-
able partnership, without which neither could reproduce. The
moths visit male yucca flowers and gather pollen, using special
prehensile tentacles found in no other type of moth. They then
use that pollen to fertilize female blossoms, and lay eggs in the
fertilized ovary. The eggs hatch out and the larvae eat the develop-
ing seeds until they are large enough to pupate. Enough seeds are
produced that each ripe fruit generally has several dozen viable
and undamaged seeds remaining when the larvae emerge, drop
to the ground, burrow an undetermined depth into the soil, and
pupate. Joshua trees are pollinated by one of two species of yucca
moth; the western populations by Tegeticula synthetica, and the
populations within the Colorado River drainage —the subspecies
Y. brevifolia jaegeriana – by T. antithetica. The moths emerge from
the ground after a long pupation – “diapause” – during the trees’
season of bloom, the fine timing due to unknown environmental
triggers. They pollinate actively for about three days, then die. If
changes in spring temperatures prompt moths to emerge when
Joshua trees have no open blooms, then both species could suffer
catastrophic reproductive failure.

Drought
Increasing the global average temperature may bring an increase
in overall precipitation to some desert areas, as prevailing winds
change and storms intensify. Still, it’s a safe bet that most of the
Southwest will see more frequent and deeper droughts. As re-
search on the massive Joshua tree forests of Joshua Tree National
Park indicates, the effect of drought on those forests may extend
beyond merely forcing the drought-tolerant trees to cope with
even less rain than usual.
In October 2001, National Park staff observed large missing
patches of the bark-like layer on Joshua trees throughout JTNP.
During the next year, only 20 mm of rain fell in the park, less than
a fifth the historic average. USGS researcher Todd Esque and his
colleagues found that the drought was a root cause of the damage
to the trees’ periderm: the local fauna, struggling to obtain food in
a year when many desert plants failed to germinate or put out new
growth, were busily stripping the bark off the trees. Black-tailed
jackrabbits (Lepus californicus) were denuding the lower meter or
so of Joshua trees’ trunks. Antelope ground squirrels (Ammosper-
mophilus leucurus) were climbing the trees and stripping periderm
from the upper branches. Botta’s pocket gophers (Thomomys bot-
tae) tunnelled beneath the trees and hollowed them out from the
inside. More than 90 percent of trees with undamaged periderms
survived the drought year handily, but mortaility rose steeply for
those trees that had been stripped of even a little periderm. No
trees that lost a quarter or more of their periderm survived. By
2003, Esque and his colleagues observed that there were no areas
of JTNP in which the Joshua trees did not exhibit periderm dam-
age. In a 2003 report published by the USGS, Esque et.al. con-
cluded that thousands of the park’s trees had been killed by native
fauna, and said:
“Reduced Joshua tree survivorship due to the combined effects of
drought and herbivory has changed the population structure of im-
pacted Joshua tree stands by thinning out mature trees. The future of
Joshua tree stands depends on whether or not additional severe and
widespread disturbances occur in coming decades.”
Drought can kill off mature trees, but it can also interfere with
the production of new trees as well. In order to bloom abundantly
Since the 1999 Juniper Complex fire in Joshua Tree National
Park few trees have germinated or resprouted here, though
by the time this photo was taken in 2001 invasive grasses had
already recovered the entire area. Photo by Chris Clarke.
Joshua trees require significant precipitation in mid-winter, pref-
erably in February. The fewer flowers are produced the fewer seeds
there will be, and thus the fewer chances a drought-stricken forest
will have to recruit younger Joshua trees that may, one day, replace
the older trees lost to periderm damage.
Fire
The increase in desert wildfire in recent years may be, in part,
another climate-related change. Torrential storms in the 2004-
5 rainy season, for instance, prompted a flush of plant growth
throughout the desert southwest: much of that growth caught
fire in the summer of 2005. In that year, some desert landscapes
burned that had not done so in the previous 10,000 years.
Some factors contributing to the increase in wildfires aren’t
directly related to climate. Invasive exotic plants have spread
throughout the deserts, not only increasing fuel loads but also
filling in what were previously bare soil areas between shrubs that
functioned as natural firebreaks. Before the advent of the inva-
sives, lightning could strike a Joshua tree, and that tree could burn
to a crisp, without the conflagration spreading to other plants.
Now a carpet of introduced vegetation has filled in the barren soil
– primarily red brome (Bromus rubens), buffelgrass (Pennisetum
ciliare)and Sahara mustard (Brassica tournefortii), but also includ-
ing Mediterranean grass (Schismus spp.), tumbleweed (Salsola),
tamarisk and about a dozen other problem species. Each not only
allows the spread of wildfire but adds significantly to fuel load, by
as much as 2,500 pounds per acre in the case of Sahara mustard.
Joshua trees are damaged directly by fire, often with even ap-
parently minor damage resulting in mortality. Different popula-
tions of Joshua trees respond differently to fire damage. The west-
ernmost population of Joshua trees, visible from Interstate 5 near
Gorman, CA in the Tehachapi Mountains, suffered a catastrophic
fire in the early part of this decade: by 2006, almost every single
tree in the grove had regrown at least one healthy basal sprout.
These trees have thrived in the fire-prone coastal mountains for
some millennia, and it makes sense they’d have evolved some
adaptation to a regular fire regime. Farther east, where wildfires
have historically been less frequent or nonexistent, most trees do
not recover from such catastrophic fire damage. A grove of trees
near Mountain Pass, burned in the early 1990s, still shows almost
no signs of recovery. Similarly slow recovery can be observed in
burned groves in the Pakoon Springs area in the Arizona Strip,
the Bulldog Wash area in southwestern Utah, and throughout
the JTNP, where – for example – recovery from the huge Juniper
Complex fire a decade ago has been less than exuberant.
If the direct damage done Joshua trees by fire is devastating, the
indirect damage done by altering their environment is even more
so. Wildfires change the vegetative structure of desert shrub­land,
destroying the slow-growing shrubs and facilitating their replace-
ment with grasses. Even if a stand of Joshua trees mostly survives
an invasive-fueled fire, the shrubs that surround them almost cer-
tainly will not, and those shrubs will be very slow to reestablish,
if the come back at all. Blackbrush (Coleogyne ramosissima), an
important understory plant in most Joshua tree forests, seems not
to reestablish at all after wildfires: one 1985 study found almost
no regrowth of blackbrush in stands that had burned as long as 37
years previously.
This is a serious long-term problem for Joshua trees. Black-
brush is a stout, sturdy, nearly impenetrable small shrub, and thus
is an effective “nurse plant” for Joshua tree seedlings. Joshua trees
are remarkably vulnerable to herbivores, especially Audubon’s
cottontails and black-tailed jackrabbits, for their first few years
after germination. Seeds that germinate within a blackbrush can-
opy stand a far better chance of escaping predation until they are
older and less palatable. Wiping out the blackbrush understory of
a Joshua tree forest may ensure that no new trees grow to replace
mature trees lost to fire, periderm damage, lightning or simple
old age. And invasive plants fill in where the blackbrush doesn’t,
ensuring more frequent future fires.
Ranges past
To understand the threat climate change poses
Joshua trees in the near future, it helps
to consider the trees’ distant past. It’s
tempting to think of pre-industrial
wild landscapes as primeval, un-
changing in the long term. But
all species’ ranges shift, usually
in response to conditions that
change over time. The current-day range of the
Joshua tree coincides more or less with the bound-
aries of the Mojave Desert in eastern California,
southern Nevada, northwestern Arizona, and
The Shasta ground sloth
(­Nothrotheriops shastense) was
most likely a significant factor in the
dispersal of Joshua tree seeds.
Painting by Carl S. Buell.
the extreme southwestern corner of Utah. Within that area, the
trees are generally found growing on well-drained slopes between
2,000 and 6,000 feet in elevation.
Thirty thousand years ago or so, the tree’s range was quite dif-
ferent. Paleontological evidence indicates that Joshua trees grew
significantly farther south, in southern Arizona and – most likely
– northern Mexico. The trees also seem to have grown at signifi-
cantly lower elevations, even that far south of their current range.
It’s safe to assume that conditions in those days were significantly
cooler, and perhaps a bit wetter, than those to found in the ­Arizona
Upland biome that occupies that landscape today.
Fast forward about 15,000 years and head back north to the
Mojave Desert. Joshua trees grew there, but the desert itself was
very different than it would be in the age of off-road vehicles.
Huge freshwater lakes filled the valleys, some of them hundreds of
feet deep, fed by a broad Mojave River. On the plains above those
lakes were forests of piñon and juniper – what desert biogeogra-
phers call “the P-J” – with Joshua trees scattered among them.
Over the next few thousand years the Mojave’s climate changed
radically yet again. Rainfall slackened. The Mojave River dwindled
and dried up, for the most part. The lakes became alkaline playas
and salinas. The verdant piñon-juniper forests retreated upslope
above the 6,000-foot line, at least where they could. Joshua trees
moved upslope as well, though not by as much. Joshua trees had
been a minor component of the piñon-juniper forest, and in a few
places, like the westernmost part of the Antelope Valley, they still
grow intermingled with piñon and juniper. For the most part,
however, Joshua trees became the dominant tree in a distinct for-
est type of their own, generally growing just beneath the lower
limits of the P-J.
And so things stand today, more or less. It’s clear that the Joshua
tree has responded to climate change in the past by moving into
new areas with more favorable climates. Why can’t the trees do
this now? Why does our current climate crisis spur talk of extinc-
tion when the species has survived past changes that were likely
even more profound?
Migrating trees
When we speak of “migration” in tree populations, we obviously
aren’t talking about individual trees moving. For plant species to
migrate, they must disperse their seeds to new locations, and then
– with luck – those seeds will
germinate, survive to re-
productive maturity, and
produce new genera-
tions that can then
disperse their seed
to even more
new habitats.
But Joshua
trees aren’t
very good
at seed
dispersal.
Pack rats will pick up seeds and fruit and carry them 50 feet from
the tree or so, and there are a few other birds and rodents that play
a role in moving Joshua tree seeds around. Ladder-backed wood-
peckers will carry ripe Joshua tree fruit to a safe spot and hammer
at fallen fruit to get the larvae that live inside, spilling seeds all over.
Every once in a while, a very hungry coyote might eat a fruit and ex-
crete it 10 miles away. But unless the tree is on a long, steep hill, its
golf-ball-sized fruit tend to stay within a few meters of the parent
tree. The seeds are large, and have none of the adaptations found
in many other desert plants to enable birds to carry them long dis-
tances: no sticky burrs, no sweet pulp surrounding a gut-proof seed
coat, nothing but a delicate little black flake. Usually when some-
thing eats a Joshua tree fruit, it kills the seed.
There are many plants in the Americas that, like the Joshua
tree, seem ill-adapted to seed dispersal because their fruit isn’t de-
signed for any living animal to disperse it efficiently: the Osage or-
ange, pawpaw, and avocado being examples. Many of these plants
are now thought to have been dispersed by giant extinct animals
of the Pleistocene, big beasts who could denude a tree of its fruit,
walk a distance away and excrete the tree seeds. Without those
animals, the plants have trouble reproducing. One of those beasts,
the Shasta giant ground sloth (Nothrotheriops shastense), lived
throughout the range of the Joshua tree until about 12,000 years
ago. That’s recently enough that you can still find its mummified
dung in caves in the southwest. That dung often contains the re-
mains of Joshua tree fruit, with seeds that could have been viable
when they were excreted. It’s a sensible conclusion that the Joshua
tree’s mass migration from Mexico to the Mojave was made pos-
sible by the giant ground sloth. Now that the ground sloth is ex-
tinct, Joshua trees disperse their seed much less effectively.
The Joshua tree’s current range, in other words, seems to be an
artifact of a relationship with a species that has been extinct for
thousands of years. That partnership now extinct, the trees’ ability
to disperse its descendants to new habitats is severely curtailed.
There are mature Joshua trees growing in Ione and Fallon, Ne-
vada, more than a hundred miles north of their current range.
It’s clear the trees themselves could grow to maturity significantly
northward of their current distribution. But little is known of
what climatic conditions each of the two Joshua tree moth spe-
cies can withstand. Some researchers talk of the possibility of “as-
sisted migration” – the deliberate planting of new populations of
Joshua trees in habitats more likely to be hospitable to the trees’
long-term survival. But until we learn much more about the eco-
logical needs of the trees’ partner moth species, we have little way
of knowing whether those new forests would be able to sustain
themselves in the long-term.
A bleak forecast
In a recent study, Dr. Kenneth Cole and his colleagues at the Colo-
rado Plateau Research Station in Flagstaff took the predictions of a
number of climate change models and applied them to determine
the likely effects of global warming on the Joshua tree. The results
were bleak. To quote from the report:
“The future potential range of [the] Joshua tree (Yucca brevifolia)
is not only reduced and shifted northward by climate change, but the
plant’s lack of dispersal mechanisms should reduce its actual extent
by at least 90%.”
Cole et al released their preliminary data in poster form at a
2005 meeting of the US Climate Change Science Program in Ar-
lington, Virginia. Cole and colleagues mapped what they knew
about current Joshua tree distribution. To that data, the research-
ers applied projections of future climate in a world with augment-
ed atmospheric CO2, calculated the likelihood of tree survival
and the rate at which the trees disperse new seedlings to potential-
ly more welcoming habitat, and mapped the species’ consequent
projected range late in this century. The resulting map was empty.
Only when they adjusted their models to assume that the trees
disperse their seeds 10 times more efficiently than they actually
do, did the model predict that Joshua trees would still exist in the
wild in the late 21st century.
Cole’s starter map received some quiet criticism. It was a mark-
edly optimistic estimate of the Joshua trees’ current range, show-
ing spurious populations well away from known stands of the
trees, especially in the extreme south part of the species’ range
in California and Arizona. But adjusting the erroneous data only
makes the results even bleaker.
Even in Cole’s optimistic scenario, the Joshua tree would be-
come extinct in Arizona and Utah. No Joshua trees would sur-
vive in Joshua Tree National Park, nor in the Mojave National
Preserve. The species would be forced into the northernmost part
of its range in eastern California and central Nevada. And all the
web of life that depends on the trees elsewhere – the woodrats who
survive the summer on a modest diet of Joshua tree leaves, the
night lizards who use fallen limbs as shelter and the night snakes
that prey on them, the flickers and woodpeckers who nest in hol-
lows in the trees’ trunks – all that will unravel.
There are still some thriving Joshua tree forests in the center
and north of the species’ range, where a mix of mature and “seed-
ling” trees seem to offer promise for the future. How long those
forests will remain thriving depends in part on luck, and in part
on our willingness to change our lifestyles so that we stop adding
CO2 to the atmosphere – and soon.
Supplemental Information
Mortality of Adult Joshua Trees (Yucca brevifolia) Due to Small
Mammal Herbivory at Joshua Tree National Park, California
Todd C. Esque, Dustin F. Haines, Lesley A. DeFalco, Jane
E. Rodgers, Kimberley A. Goodwin and Sara J. Scoles
USGS, 2003
The Effects of Fire on the Blackbrush (Coleogyne ramosissima)
Community of Southwestern Utah
Jim Callison, Jack D. Brotherson and James E. Bowns
Journal of Range Management, Vol. 38, No. 6 (Nov., 1985), pp.
535-538
Transient Dynamics of Vegetation Response to Past and Future
­Climatic Changes in the Southwestern United States. (Poster)
Kenneth L. Cole, Kirsten Ironside, Phillip Duffy, and Samantha
Arundel
USGS, 2005
Yucca moths
Chris Clarke
http://qarrtsiluni.com/2007/12/19/yucca-moths/