Agassiz’s desert tortoises (Gopherus agassizii) approach the southern edge of their mostly Mojave Desert range near Joshua Tree National Park. Modern desert tortoise research started in the Park in 1978 when the first tortoise population census was conducted on a one square mile area in the Pinto Basin known as the “Barrow Plot.” U.S. Geological Survey research began at the plot in 1997 and continued intermittently until 2012, providing important long-term data on variation in population size and survivorship of long-lived tortoises during a period of widely variable environmental conditions. As a threatened species under the Endangered Species Act, resource managers need information on how best to manage for healthy tortoise populations. Other long-term research is focused on tortoise populations living near the mouth of Cottonwood Canyon in the Sonoran Desert portion of the Park. Research at Cottonwood examined the behavior, movements and reproductive ecology of tortoises using radio-telemetry and X-radiography. Scientists from the U.S. Geological Survey and the National Park Service have a long history of collaboration on tortoise research at Joshua Tree National Park.
Background & Importance
Agassiz’s desert tortoise populations are declining across their vast range in the Mojave and Sonoran deserts of Arizona, California, Nevada and Utah. Research is needed by resource management agencies to help recover populations and ultimately delist them from their status as threatened species under the Endangered Species Act.
Research is also needed to understand how populations will respond to utility-scale solar and wind energy development in desert tortoise habitat and to understand how they will respond to climate change. Large areas of the Southwestern Desert are developed or slated for development of large renewable energy projects, often requiring relocation of tortoises with results that are not always predictable. This research also supports the monitoring requirements for desert tortoises under the Coachella Valley Multiple Species Habitat Conservation Plan.
General Methods
Population censuses for tortoises often used plot-based surveys of one square mile study plots and this is how work started at the Barrow Plot in Joshua Tree National Park. Teams of scientists and technicians walk closely-spaced transects across the plot, noting all tortoises and tortoise sign (tortoise burrows, shells, and scat). Live tortoises are marked with unique and permanent notches on their shell that allow researchers to identify individuals at subsequent captures to measure growth, movements, etc. An advantage of plot-based surveys is that detailed recapture data can be analyzed over time for a cohort of marked tortoises since they generally don’t move great distances but they can live a long time. A disadvantage is that extrapolation of density results to surrounding areas can be flawed due to the patchy distribution of habitats and tortoises in the larger desert landscape.
Individual tortoises are measured and weighed, and sex is determined by examining shell shape and tail size. A small sample of tortoises are outfitted with radio transmitters that allow scientists to find them on subsequent visits to the field site. Adult female tortoises are X-rayed to determine the clutch size, clutch frequency (they produce up to two clutches per female per year in Joshua Tree National Park), and egg width. Previously published research by the Principal Investigator determined that this technique poses minimal risk to females and embryos. Small blood samples are occasionally taken to obtain DNA used to compare genetic differences between populations. In addition, wildlife “trail cameras” were used in 2015 to detect tortoise predators.
Important Results
To date, research has demonstrated that National Parks provide good protection to desert tortoise populations but they cannot protect them from all the negative impacts of drought, fire, disease, climate change, and road mortality. For example, analysis of long-term data from the Barrow Plot found a strong relationship between average rainfall amounts and survival of tortoises. Simply put, tortoise populations decreased in size during droughts. What was once a robust and large population of tortoises in the early 1990s declined precipitously by 2012. The effects of the drought also caused predators like coyotes to eat more tortoises instead of their normal prey: rabbits and rodents. Research at Cottonwood demonstrates that tortoises often spend most of the year in steep boulder piles and small mountains, even nesting there. Elsewhere in California, tortoises appear to prefer flatter ground with more sandy soils for digging burrows. Steep terrain may be underappreciated by resource managers as habitat for desert tortoises in some areas.

Below are other science projects related to this project.
The Gemini Solar Project
Turtle Ecology
Desert Tortoise Ecology and Renewable Energy Development
Below are selected publications associated with this project.
High female desert tortoise mortality in the western Sonoran Desert during California’s epic 2012–2016 drought
Refining genetic boundaries for Agassiz’s desert tortoise (Gopherus agassizii) in the western Sonoran Desert: The influence of the Coachella Valley on gene flow among populations in southern California
Turtles and tortoises are in trouble
Where have all the turtles gone, and why does it matter?
Agassiz’s desert tortoise (Gopherus agassizii) activity areas are little changed after wind turbine-induced fires in California
Where have all the turtles gone, and why does it matter?
Reproductive output and clutch phenology of female Agassiz's desert tortoises (Gopherus agassizii) in the Sonoran Desert region of Joshua Tree National Park
Global conservation status of turtles and tortoises (order Testudines)
Variation in annual clutch phenology of desert tortoises (Gopherus morafkai) in the Sonoran Desert of Arizona
Not putting all their eggs in one basket: bet-hedging despite extraordinary annual reproductive output of desert tortoises
Studies of reproductive output of the desert tortoise at Joshua Tree National Park, the Mojave National Preserve, and comparative sites
Below are news features about desert tortoise and turtle research led by the Southwest Biological Science Center.
Below are partners associated with this project.
- Overview
Agassiz’s desert tortoises (Gopherus agassizii) approach the southern edge of their mostly Mojave Desert range near Joshua Tree National Park. Modern desert tortoise research started in the Park in 1978 when the first tortoise population census was conducted on a one square mile area in the Pinto Basin known as the “Barrow Plot.” U.S. Geological Survey research began at the plot in 1997 and continued intermittently until 2012, providing important long-term data on variation in population size and survivorship of long-lived tortoises during a period of widely variable environmental conditions. As a threatened species under the Endangered Species Act, resource managers need information on how best to manage for healthy tortoise populations. Other long-term research is focused on tortoise populations living near the mouth of Cottonwood Canyon in the Sonoran Desert portion of the Park. Research at Cottonwood examined the behavior, movements and reproductive ecology of tortoises using radio-telemetry and X-radiography. Scientists from the U.S. Geological Survey and the National Park Service have a long history of collaboration on tortoise research at Joshua Tree National Park.
USGS Research Ecologist Jeff Lovich gets ready to release a male desert tortoise into its cave-like shelter in southern Joshua Tree National Park after collecting data. Photo by Shellie Puffer, USGS, SBSC. Background & Importance
Agassiz’s desert tortoise populations are declining across their vast range in the Mojave and Sonoran deserts of Arizona, California, Nevada and Utah. Research is needed by resource management agencies to help recover populations and ultimately delist them from their status as threatened species under the Endangered Species Act.
A female Agassiz's desert tortoise at Joshua Tree National Park lounges in the entrance of her burrow, wearing a USGS radio. Research is also needed to understand how populations will respond to utility-scale solar and wind energy development in desert tortoise habitat and to understand how they will respond to climate change. Large areas of the Southwestern Desert are developed or slated for development of large renewable energy projects, often requiring relocation of tortoises with results that are not always predictable. This research also supports the monitoring requirements for desert tortoises under the Coachella Valley Multiple Species Habitat Conservation Plan.
General Methods
Population censuses for tortoises often used plot-based surveys of one square mile study plots and this is how work started at the Barrow Plot in Joshua Tree National Park. Teams of scientists and technicians walk closely-spaced transects across the plot, noting all tortoises and tortoise sign (tortoise burrows, shells, and scat). Live tortoises are marked with unique and permanent notches on their shell that allow researchers to identify individuals at subsequent captures to measure growth, movements, etc. An advantage of plot-based surveys is that detailed recapture data can be analyzed over time for a cohort of marked tortoises since they generally don’t move great distances but they can live a long time. A disadvantage is that extrapolation of density results to surrounding areas can be flawed due to the patchy distribution of habitats and tortoises in the larger desert landscape.
If winter rains are adequate, the Sonoran Desert in Joshua Tree National Park springs to life with wildflowers like these desert dandelions. Wildflowers are an important source of food and moisture for desert tortoises and may only be available for a few weeks of the year before they dry out and lose much of their nutritional value. (Credit: Shellie Puffer, USGS. Public domain.) Individual tortoises are measured and weighed, and sex is determined by examining shell shape and tail size. A small sample of tortoises are outfitted with radio transmitters that allow scientists to find them on subsequent visits to the field site. Adult female tortoises are X-rayed to determine the clutch size, clutch frequency (they produce up to two clutches per female per year in Joshua Tree National Park), and egg width. Previously published research by the Principal Investigator determined that this technique poses minimal risk to females and embryos. Small blood samples are occasionally taken to obtain DNA used to compare genetic differences between populations. In addition, wildlife “trail cameras” were used in 2015 to detect tortoise predators.
Sunset on the Barrow Plot in eastern Joshua Tree National Park. This area used to be inhabited by a healthy population of desert tortoises, but by 2012 many were dead due to the effects of protracted drought and predation by coyotes. Photo taken May 4, 2012.(Credit: Jeff Lovich, USGS. Public domain.) Important Results
To date, research has demonstrated that National Parks provide good protection to desert tortoise populations but they cannot protect them from all the negative impacts of drought, fire, disease, climate change, and road mortality. For example, analysis of long-term data from the Barrow Plot found a strong relationship between average rainfall amounts and survival of tortoises. Simply put, tortoise populations decreased in size during droughts. What was once a robust and large population of tortoises in the early 1990s declined precipitously by 2012. The effects of the drought also caused predators like coyotes to eat more tortoises instead of their normal prey: rabbits and rodents. Research at Cottonwood demonstrates that tortoises often spend most of the year in steep boulder piles and small mountains, even nesting there. Elsewhere in California, tortoises appear to prefer flatter ground with more sandy soils for digging burrows. Steep terrain may be underappreciated by resource managers as habitat for desert tortoises in some areas.
Sources/Usage: Public Domain. Visit Media to see details.A radioed male tortoise in the Santa Rosa Mountains, CA eating a beavertail prickly pear cactus. During the extreme drought conditions we are experiencing, that is the only food available for them with any moisture. They have to cope with all the little spiny gloccids (hair-like spines) that cover the pads of the cactus. Video provided by Dr. Bill Hoese at California State University, Fullerton provided to Jeff Lovich, SBSC, USGS with permission to use. Scattered mature Joshua trees in this northernmost stand of Joshua trees are surrounded by abundant seedlings and saplings. USGS, public domain. - Science
Below are other science projects related to this project.
The Gemini Solar Project
The United States is developing renewable energy resources, especially solar, at a rapid rate. Although renewable energy development is widely perceived by the public as “green technology,” construction, operation, maintenance, and eventual decommissioning of facilities all have known and potential negative impacts to natural resources, including plant communities and wildlife. This is especially...Turtle Ecology
Turtles are among the most recognizable and iconic of animals. Any animal with a shell and a backbone is a turtle whether they are called turtles, tortoises, or terrapins. In fact, terrapin is an Algonquian Native American name for turtle. Worldwide there are 356 turtle species on all continents except for Antarctica. The United States has more species than any other country with about 62...Desert Tortoise Ecology and Renewable Energy Development
The desert Southwest is experiencing rapid development of utility-scale solar and wind energy facilities. Although clean renewable energy has environmental benefits, it can also have negative impacts on wildlife and their habitats. Understanding those impacts and effectively mitigating them is a major goal of industry and resource managers. One species of particular concern is Agassiz’s desert... - Publications
Below are selected publications associated with this project.
High female desert tortoise mortality in the western Sonoran Desert during California’s epic 2012–2016 drought
We conducted population surveys for desert tortoises Gopherus agassizii at 2 nearby sites in the western Sonoran Desert of California, USA, from 2015-2018, during the driest ongoing 22 yr period (2000-2021) in the southwestern USA in over 1200 yr. We hypothesized that drought-induced mortality would be female-biased due to water and energy losses attributable to egg production during protracted peAuthorsJeffrey E. Lovich, Michele (Shellie) R. Puffer, Kristy L. Cummings, Terence R. Arundel, Michael S. Vamstad, Kathleen D. BrundigeRefining genetic boundaries for Agassiz’s desert tortoise (Gopherus agassizii) in the western Sonoran Desert: The influence of the Coachella Valley on gene flow among populations in southern California
Understanding the influence of geographic features on the evolutionary history and population structure of a species can assist wildlife managers in delimiting genetic units (GUs) for conservation and management. Landscape features including mountains, low elevation depressions, and even roads can influence connectivity and gene flow among Agassiz’s desert tortoise (Gopherus agassizii) populationsAuthorsJeffrey E. Lovich, Taylor Edwards, Kristin H. Berry, Shellie R. Puffer, Kristy L. Cummings, Ennen Joshua R., Mickey Agha, Rachel Woodard, Kathleen D. Brundige, Robert W. MurphyTurtles and tortoises are in trouble
Turtles and tortoises (chelonians) have been integral components of global ecosystems for about 220 million years and have played important roles in human culture for at least 400,000 years. The chelonian shell is a remarkable evolutionary adaptation, facilitating success in terrestrial, freshwater and marine ecosystems. Today, more than half of the 360 living species and 482 total taxa (species aAuthorsC.B. Stanford, J.B. Iverson, A.G.J. Rhodin, P.P. van Dijk, R.A. Mittermeier, G. Kuchling, Kristin H. Berry, A. Bertolero, K.A. Bjorndal, T.E.G. Blanck, K. A. Buhlmann, R.L. Burke, J.D. Congdon, T. Diagne, T. Edwards, C.C. Eisemberg, J.R. Ennen, G. Forero-Medina, M. Frankel, U. Fritz, N. Gallego-Garcia, A. Georges, J.W Gibbons, S. Gong, E.V. Goode, H.T. Shi, H. Hoang, M.D. Hofmeyr, B.D Horne, R. Hudson, J.O. Juvik, R.A. Kiester, P.V. Koval, M.-A. Lea, P.V. Lindeman, Jeffrey E. Lovich, L. Luiselli, T.E.M. McCormack, G.A. Meyer, V.P. Paez, K. Platt, S.G. Platt, P.C.H. Pritchard, H.R. Quinn, W.M. Roosenburg, J.A. Seminoff, H.B. Shaffer, R. Spencer, J.U. Van Dyke, R.C. Vogt, A.D. WaldeWhere have all the turtles gone, and why does it matter?
No abstract available.AuthorsJeffrey E. Lovich, Joshua R. Ennen, Mickey Agha, J. Whitfield GibbonsAgassiz’s desert tortoise (Gopherus agassizii) activity areas are little changed after wind turbine-induced fires in California
Wind turbine-induced fires at a wind energy facility in California, USA, provided an opportunity to study the before and after effects of fire on a population of protected Agassiz’s desert tortoises (Gopherus agassizii) in the Sonoran Desert, a species and ecosystem poorly adapted to fire. We compared annual activity areas (AAs) of tortoises in 2011 and 2013, before and after two 2012 fires, withAuthorsJeffrey E. Lovich, Mickey Agha, Joshua R. Ennen, Terence R. Arundel, Meaghan AustinWhere have all the turtles gone, and why does it matter?
Of the 356 species of turtles worldwide, approximately 61% are threatened or already extinct. Turtles are among the most threatened of the major groups of vertebrates, in general, more so than birds, mammals, fishes or even the much besieged amphibians. Reasons for the dire situation of turtles worldwide include the familiar list of impacts to other species including habitat destruction, unsustainAuthorsJeffrey E. Lovich, Joshua R. Ennen, Mickey Agha, J. Whitfield GibbonsReproductive output and clutch phenology of female Agassiz's desert tortoises (Gopherus agassizii) in the Sonoran Desert region of Joshua Tree National Park
Agassiz's desert tortoise (Gopherus agassizii) reaches the southern edge of its range in the Sonoran Desert of California. The reproductive ecology of this wideranging species is understudied here compared to populations in the adjacent Mojave Desert. Understanding potential geographic variation in reproductive ecology is important for effective management of conservation-reliant species like G. aAuthorsJeffrey E. Lovich, Shellie R. Puffer, Mickey Agha, Joshua R. Ennen, Kathie Meyer-Wilkins, Laura A. Tennant, Amanda L Smith, Terence R. Arundel, Kathleen D. Brundige, Michael S. VamstadGlobal conservation status of turtles and tortoises (order Testudines)
We present a review and analysis of the conservation status and International Union for Conservation of Nature (IUCN) threat categories of all 360 currently recognized species of extant and recently extinct turtles and tortoises (Order Testudines). Our analysis is based on the 2018 IUCN Red List status of 251 listed species, augmented by provisional Red List assessments by the IUCN Tortoise and FrAuthorsAnders G.J. Rhodin, Craig B. Stanford, Peter Paul van Dijk, Carla Eisemberg, Luca Luiselli, Russell A. Mittermeier, Rick Hudson, Brian D. Horne, Eric Goode, Gerald Kuchling, Andrew Walde, Ernst H. W. Baard, Kristin H. Berry, Albert Bertolero, Torsten E. G. Blanck, Roger Bour, Kurt A. Buhlmann, Linda J. Cayot, Sydney Collett, Andrea Currylow, Indraneil Das, Tomas Diagne, Joshua R. Ennen, Germán Forero-Medina, Matthew G. Frankel, Uwe Fritz, Gerardo García, J. Whitfield Gibbons, Paul M. Gibbons, Gong Shiping, Joko Guntoro, Margaretha D. Hofmeyr, John B. Iverson, A. Ross Kiester, Michael Lau, Dwight P. Lawson, Jeffrey E. Lovich, Edward O. Moll, Vivian P. Páez, Rosalinda Palomo-Ramos, Kalyar Platt, Steven G. Platt, Peter C. H. Pritchard, Hugh R. Quinn, Shahriar Caesar Rahman, Soary Tahafe Randrianjafizanaka, Jason Schaffer, Will Selman, H. Bradley Shaffer, Dionysius S. K. Sharma, Shi Haitao, Shailendra Singh, Ricky Spencer, Kahleana Stannard, Sarah Sutcliffe, Scott Thomson, Richard C. VogtVariation in annual clutch phenology of desert tortoises (Gopherus morafkai) in the Sonoran Desert of Arizona
The phenology of egg production and oviposition in organisms affects survival and development of neonates and thus, both offspring and maternal fitness. In addition, in organisms with environmental sex determination, clutch phenology can affect hatchling sex ratios with attendant effects on population demography. The rapid rate of contemporary climate change might disrupt reproductive phenologiesAuthorsJeffrey E. Lovich, Roy C. Averill-Murray, Mickey Agha, Joshua R. Ennen, Meaghan AustinNot putting all their eggs in one basket: bet-hedging despite extraordinary annual reproductive output of desert tortoises
Bet-hedging theory makes the counter-intuitive prediction that, if juvenile survival is low and unpredictable, organisms should consistently reduce short-term reproductive output to minimize the risk of reproductive failure in the long-term. We investigated the long-term reproductive output of an Agassiz's desert tortoise (Gopherus agassizii) population and conformance to a bet-hedging strategy ofAuthorsJeffrey E. Lovich, Joshua R. Ennen, Charles B. Yackulic, Kathie Meyer-Wilkins, Mickey Agha, Caleb L. Loughran, Curtis Bjurlin, Meaghan Austin, Sheila V. MadrakStudies of reproductive output of the desert tortoise at Joshua Tree National Park, the Mojave National Preserve, and comparative sites
The stability of any population is a function of how many young are produced and how many survive to reproduce. Populations with low reproductive output and high mortality will decline until such time as deaths and births are at least balanced. Monitoring populations of sensitive species is particularly important to ensure that conditions do not favor decline or extinction. Turtles, including tortAuthorsJ.E. Lovich, P. Medica, H. Avery, K. Meyer, G. Bowser, A. Brown - News
Below are news features about desert tortoise and turtle research led by the Southwest Biological Science Center.
- Partners
Below are partners associated with this project.