Home Range and Movement of Desert Tortoises Gopherus Agassizii in the Mojave Desert of California, USA
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Vol. 13: 191–201, 2011 ENDANGERED SPECIES RESEARCH Published online March 9 doi: 10.3354/esr00313 Endang Species Res OPENPEN ACCESSCCESS Home range and movement of desert tortoises Gopherus agassizii in the Mojave Desert of California, USA Bryan R. Franks, Harold W. Avery*, James R. Spotila Department of Biology, Drexel University, 3141 Chestnut St., Philadelphia, Pennsylvania 19104, USA ABSTRACT: Radio telemetry of 29 desert tortoises Gopherus agassizii in 2000 and 60 in 2001 at 4 sites in the Mojave Desert of California, USA, revealed that males had larger home ranges and moved greater distances than females, but that tortoises at sites with more rainfall did not always have larger home ranges or move greater distances. The Ivanpah Valley received more rainfall than the Fort Irwin area, Superior Valley, and Fremont, but differences between home range size and rain- fall were not consistent. Within the Ivanpah Valley, the plot at the highest elevation received more rainfall than the lower plots. However, within the Ivanpah Valley, tortoises at the 3 locations did not differ in home range and movement. In the wetter year (2001), female tortoises moved greater dis- tances each day at Ivanpah than female tortoises at Superior and Fremont, which were drier sites. Males moved farther per day than females in both years. In 2001, female tortoises in the Ivanpah Val- ley used more burrows than tortoises at Fremont and Superior, but similar numbers as tortoises at Fort Irwin. There was no mortality of tortoises in 2000 and a 5% mortality in 2001. KEY WORDS: Desert tortoise · Radio telemetry · Rainfall · Ivanpah Valley · Fort Irwin · Minimum convex polygon · Climate change Resale or republication not permitted without written consent of the publisher INTRODUCTION more thorough and detailed knowledge of desert tortoise life history and ecology, including knowledge The desert tortoise Gopherus agassizii inhabits the of desert tortoise movements, home range, and activity Mojave, Colorado, and Sonoran Deserts, USA, (Ernst patterns, in order to establish a sound conservation et al. 1994) and is listed as a threatened species under or management strategy for the species (Berry 1986, the US Endangered Species Act in the western part of Gibbons 1986, US Fish & Wildlife Service 1994, its range (US Fish & Wildlife Service 1990). Field data Freilich et al. 2000). (Berry 1997) and demographic analyses by modeling Home range (White & Garrott 1990) and movements (Doak et al. 1994) have indicated that many popula- are determined in part by the biotic resources and tions of desert tortoises are in decline. However, accu- biophysical properties in an environment. The re- racy of historical density estimates have been chal- source environment (e.g. water, energy, protein) and lenged (Bury & Corn 1995) due to difficulties with the social environment (e.g. mating strategies, territo- desert tortoise surveys (Corn 1994, Freilich et al. 2000), riality) of an ectotherm influence its daily activity bud- large differences in local densities (Germano et al. get (Dunham et al. 1989). This daily activity budget in 1994a), inter-observer bias (Freilich & LaRue 1998), turn affects the movements and home range of that and changes in tortoise activity due to changes in rain- animal. Predator avoidance and demographics within fall and plant productivity (Duda et al. 1999). In addi- a population also affect home range and movements. tion, tortoises may be undercounted in dry years Availability of water varies both temporally and spa- (Freilich et al. 2000). Research is still needed to provide tially in the Mojave Desert (Shreve 1925, Beatley 1974, *Corresponding author. Email: [email protected] © Inter-Research 2011 · www.int-res.com 192 Endang Species Res 13: 191–201, 2011 Germano et al. 1994b). This variation has a direct ef- et al. 2007). The reasons for these differences are fect on the success of vegetation and the types of vege- unclear. Additional data are needed to determine tation present in an area. From year to year, rainfall is whether differences in home range and movement at highly variable, with years of drought followed by different locations represent a general phenomenon in years of high rainfall. In the western Mojave Desert, desert tortoises and to determine the mechanisms that rainfall primarily occurs in winter, with little or no sum- control home range size and movement of male and mer rain. Historically, rainfall in the eastern Mojave female tortoises at different sites in the Mojave Desert. Desert has been bimodal, with winter rains as well as a We used radio telemetry to examine effects of year, pulse of summer rain commonly seen in August and sex, location, and rainfall on desert tortoise home September (Shreve 1925). Since 1950, California has range size, movement, and activity at 4 sites over a 2 yr experienced increased effects of global warming, with period (2000 to 2001) in the California Mojave Desert. warmer winter and spring temperatures and earlier Radio telemetry continues to be a useful method to spring flowering (Cayan et al. 2006). Climate models determine the home ranges of endangered species as project that there will be an increase of 2.8 to 5.5°C in varied as Mt. Graham red squirrels (Koprowski et al. June to August temperatures in the Mojave Desert in 2008) and pangolins (Lim & Ng 2008). The objectives of 2070 to 2099 and a change of 5 to 20% in total precipi- the study were to: (1) determine whether rainfall tation during the 21st century due to an increase in affected home range size, movements, and activity, (2) CO2 that will cause global warming (Cayan et al. determine whether home ranges, movements, and 2008). We already know that seasonal, annual, and ge- activity differed in male and female tortoises, and (3) ographic differences in rainfall affect the physiology, determine whether home range size and location behavior, and ecology of desert tortoises (Nagy & Med- differed seasonally. ica 1986, Peterson 1996b, Henen et al. 1998). There- fore, it is important to determine how desert tortoises change behavior in response to differences in climate MATERIALS AND METHODS variables so that we can anticipate how tortoises will react to changes in climate over the next 50 to 100 yr. We conducted this study at 4 locations in the Cali- Home range size, movements, and activity patterns fornia Mojave Desert (Fig. 1). We collected data from 2 of tortoises vary in time and space. Annual home sites in 2000 and from all 4 sites during 2001. Sites range sizes of desert tortoises in the Mojave Desert ranged from the western Mojave Desert near Helen- are highly variable, ranging from 1 to 89 ha both dale, California, to the eastern Mojave Desert near the between and within populations (Berry 1986, Duda et Nevada–California border within the Mojave National al. 1999, Field et al. 2007, Harless et al. 2009). Home Preserve (Fig. 1). Sites differed in human impact, rang- range sizes and movements in 2 populations of desert ing from heavy activity such as dumping and off-road tortoises differed between 1995 and 1996 in the south- vehicle (OHV) use (west Mojave) to little visible dis- central Mojave Desert (Duda et al. 1999). The first year turbance (east Mojave). Sites varied in elevation, rain- was wet and had more plant productivity, and tortoises fall amounts and frequency, and vegetation communi- had larger home ranges and traveled higher average ties. We monitored tortoises of both sexes at 3 sites but distances per day than in the subsequent drought year. only monitored female tortoises at 1 site due to logisti- Tortoises at 1 site had significantly smaller home range cal limitations. Sample sizes are given in Table 1. We sizes than tortoises at the second site, correlated with were seriously limited in our ability to acquire equal plant productivity differences between sites (Duda sample sizes and times due to delays in permitting et al. 1999). from United States Fish and Wildlife Service (USFWS) The effect of sex on the home range and movements and the Mojave National Preserve of the National of tortoises varies. Home range size was the same for Parks Service. male and female tortoises in the Sonoran population in Ivanpah Valley site. The Ivanpah Valley site was the Picacho Mountains (Barrett 1990) and in Saguaro within the Mojave National Preserve approximately National Park in Arizona (Stitt et al. 2003), and for the 10 km south and west of Nipton, California (Fig. 1). Mojave population near the Desert Tortoise Conserva- The Mojave National Preserve is a 650 000 ha pro- tion Center in the Las Vegas Valley in Nevada (O’Con- tected area created in 1994. The valley is located nor et al. 1994). However, male desert tortoises had between the Ivanpah and New York Mountains. Eleva- larger home ranges than females near Lake Mead, tion ranges from 870 to 1100 m and increases from Nevada (Duda et al. 1999), in Joshua Tree National northeast to southwest. A creosote scrub-dominated Park (Freilich et al. 2000), and in Fort Irwin, California community occurs at lower elevations, with Mojave (Harless et al. 2009), and translocated males had larger yucca and Joshua trees occurring at higher elevations. home ranges than females in southern Nevada (Field Human impacts included a powerline corridor, cattle Franks et al.: Desert tortoise home range and movement 193 fall amounts along a 10 km gradient. Areas at higher el- evations in the valley received more rain than areas at lower elevations (Avery 1998, Avery et al. 2002). We monitored tortoises from 3 study plots of 4 km2 each within the valley. Plot 1 (Lower Powerline) was in the lower portion of the valley, with elevations ranging from 850 to 950 m. Plot 2 (Lower Water Tank) was in the middle to lower portion of the valley, with eleva- tions ranging from 980 to 1050 m.