Habitat Use by Sonoran Desert Tortoises

ABSTRACT The distribution of desert tortoises (Gopherus agassizii) spans a wide range of biotic and abiotic conditions in the southwestern United States and northwestern Mexico, with physical and behavioral differences distinguishing tortoises inhabiting the Mojave Desert from those inhabiting the Sonoran Desert. Relative to tortoise populations in the Mojave Desert, populations in the Sonoran Desert have not been well-studied. To assess how habitat use of desert tortoises in the Sonoran Desert was influenced by topography, vegetation, geomorphology, and soil, we surveyed 40 randomly located 3-ha sites for presence of adult tortoises within a site-occupancy framework. We modeled both occupancy and detection probability as a function of environmental features, and compared those results with a logistic regression model that assumed detection probability was equal to 1. Results from both approaches agreed, suggesting that habitat selection of tortoises in the Sonoran Desert was influenced primarily by topographic and geomorphologic features rather than by vegetation. Specifically, tortoises were more likely to occupy sites that were steep (we detected tortoises on 29% of sites with mean slope <5° and 92% of sites with mean slope >15°) and predominantly east-facing (53% of sites with <5% of site facing E and 92% of sites with >20% facing E), and less likely to occupy north-facing slopes (100% of sites with <10% of site facing N and 14% of sites with >60% facing N). Our results contrast with patterns of habitat use in the Mojave Desert where tortoises primarily occupy valley bottoms. Habitat use of tortoises in Sonoran and Mojave Desert populations differ considerably, contributing to the mounting body of evidence suggesting that these geographically distinct populations may represent separate species.     

Estimating the probability of illegal desert tortoise collection in the Sonoran Desert

The expansion of road networks in desert tortoise (Gopherus agassizii) habitat in the Sonoran Desert has raised questions concerning appropriate mitigation to reduce impacts at the population level. Although some effects, namely road-kill and habitat loss, have been well documented, illegal tortoise collection has been insufficiently addressed. It has become increasingly important for wildlife and land-use managers to understand the cumulative impacts of roads on tortoises and the effect that those impacts have on population persistence. We estimated the probability of desert tortoise detection and collection along 2-lane paved, maintained gravel, and non-maintained gravel roads to evaluate whether collection probabilities were related to road type. Although collection probability did not vary by road type, the probability of desert tortoise detection by passing motorists was greatest on maintained gravel roads and fewest on non-maintained gravel and paved roads. These results have implications for effectively mitigating the impacts of roads on desert tortoises. Published 2011. This article is a U.S. Government work and is in the public domain in the USA.     

Implications of Anthropogenic Landscape Change on Inter-Population Movements of the Desert Tortoise (Gopherus agassizii)

In the Sonoran Desert of North America, populations of the desert tortoise (Gopherus agassizii) occur in rocky foothills throughout southwestern Arizona and northwestern Mexico. Although tortoise populations appear to be isolated from each other by low desert valleys, individuals occasionally move long distances between populations. Increasingly, these movements are hindered by habitat fragmentation due to anthropogenic landscape changes. We used molecular techniques and radiotelemetry to examine movement patterns of desert tortoises in southern Arizona. We collected blood samples from 170 individuals in nine mountain ranges and analyzed variability in seven microsatellite loci to determine genetic differentiation among populations. Gene flow estimates between populations indicate that populations exchanged individuals historically at a rate greater than one migrant per generation, and positive correlation between genetic and geographic distance of population pairs suggests that the limiting factor for gene flow among populations is isolation by distance. Life history traits of the desert tortoise, a long-lived species with delayed sexual maturity, may severely constrain the ability of small populations to respond to disturbances that increase adult mortality. Historic gene flow estimates among populations suggests that recovery of declining populations may rely heavily on the immigration of new individuals from adjacent mountain ranges. Management strategies compatible with the evolutionary history of gene flow among disjunct populations will help ensure the long-term persistence of Sonoran desert tortoise populations.     

Polymorphic microsatellite markers for the Mojave desert tortoise, Gopherus agassizii

We describe primers and polymerase chain reaction (PCR) conditions to amplify 14 tri- and tetranucleotide microsatellite loci for the Mojave desert tortoise (Gopherus agassizii). Across three populations (87 individuals) located in the Mojave Desert, USA, the markers yielded a range of four to 33 alleles and an average observed heterozygosity of 0.733 (range 0.433 to 0.933). We neither detected linkage disequilibrium between any pair of loci nor did we find a consistent pattern of deviation from Hardy-Weinberg equilibrium. These microsatellites are designed for PCR multiplexing, and provide higher throughput capacity to aid in conservation genetics studies for this threatened species.     

Translocation as a conservation tool for Agassiz's desert tortoises: Survivorship,reproduction, and movements

We translocated 120 Agassiz's desert tortoises to 5 sites in Nevada and Utah to evaluate the effects of translocation on tortoise survivorship, reproduction, and habitat use. Translocation sites included several elevations, and extended to sites with vegetation assemblages not typically associated with desert tortoises in order to explore the possibility of moving animals to upper elevation areas. We measured survivorship, reproduction, and movements of translocated and resident animals at each site. Survivorship was not significantly different between translocated and resident animals within and among sites, and survivorship was greater overall during non-drought years. The number of eggs produced by tortoises was similar for translocated and resident females, but differed among sites. Animals translocated to atypical habitat generally moved until they reached vegetation communities more typical of desert tortoise habitat. Even within typical tortoise habitat, tortoises tended to move greater distances in the first year after translocation than did residents, but their movements in the second or third year after translocation were indistinguishable from those of resident tortoises. Our data show that tortoises translocated into typical Mojave desert scrub habitats perform well; however, the large first-year movements of translocated tortoises have important management implications. Projects that employ translocations must consider how much area will be needed to contain translocated tortoises and whether roads need fencing to prevent the loss of animals. © 2012 The Wildlife Society.     

Reproductive Consequences of Clonal Growth in Stenocereus eruca, a Rare Clonal Cactus of the Sonoran Desert

Stenocereus eruca is a prostrated, self-incompatible cactus endemic to the Sonoran Desert that regenerates primarily through clonal propagation. Clonal growth is expected to affect mate availability by influencing the number and spatial distribution of mating types. In this paper we examine the role of clonal growth on female fecundity through a series of pollination experiments in a population of S. eruca. We set up a pollen supplementation experiment using five distance treatments with pollen collected at 1, 10, 100, 1000 and 25000 m from receptor flowers during the years 2001 and 2002 and evaluated genetic sifmilarities between pairs of receptor-donor ramets through RAPD markers. Our data on fruit set, number of seeds/fruit, germination and overall fecundity revealed that S. eruca show a significant reduction in female fecundity when pollination occurs between ramets located at short distances (1 and 10 m), while genetic data showed high levels of similarity at those distances. The reduction in female fecundity is apparently a consequence of geitonogamy and inbreeding depression. Our data suggest that clonal growth and geitonogamy are likely to be partially responsible for the low levels of sexual reproduction and seedling recruitment observed in populations of S. eruca. Co-ordinating editor: H. Kudoh     

Root distribution and seasonal production in the northwestern Sonoran Desert for a C3 subshrub, a C4 bunchgrass, and a CAM leaf succulent

To investigate root distribution with depth, which can affect competition for water, surface areas of young and old roots were determined in 4-cm-thick soil layers for the C3 subshrub Encelia farinosa Torrey and A. Gray, the C4 bunchgrass Pleuraphis rigida Thurber, and the CAM (crassulacean acid metabolism) leaf succulent Agave deserti Engelm. At a site in the northwestern Sonoran Desert these codominant perennials had mean rooting depths of only 9-10 cm for isolated plants. Young roots had mean depths of 5-6 cm after a winter wet period, but 11-13 cm after a summer wet period. Young roots were most profuse in the winter for E. farinosa, which has the lowest optimum temperature for root growth, and in the summer for P. rigida, which has the highest optimum temperature. Roots for interspecific pairs in close proximity averaged 2-3 cm shallower for A. deserti and a similar distance deeper for the other two species compared with isolated plants, suggesting partial spatial separation of their root niches when the plants are in a competitive situation. For plants with a similar root surface area, the twofold greater leaf area and twofold higher maximal transpiration rate of E. farinosa were consistent with its higher root hydraulic conductivity, leading to a fourfold higher estimated maximal water uptake rate than for P. rigida. Continuous water uptake accounted for the shoot water loss by A. deserti, which has a high shoot water-storage capacity. A lower minimum leaf water potential for P. rigida than for A. deserti indicates greater ability to extract water from a drying soil, suggesting that temporal niche separation for water uptake also occurs.     

Monitoring of the Flat-Tailed Horned Lizard With Methods Incorporating Detection Probability

Abstract: Difficulty in monitoring the flat-tailed horned lizard (Phrynosoma mcallii) has led to controversy over its conservation status. The difficulty in detecting this species has discouraged large-scale estimates of abundance and led to uncertainty over whether the species exists in population sizes of sufficient size for long-term persistence. We incorporated detection probability into monitoring of this species using closed mark—recapture and distance-sampling methods. Density estimation from mark—recapture abundance estimates was improved using an estimate of the proportion of time lizards were on the plot. We estimated the probability of detection on the line for distance sampling and adjusted density estimates accordingly. We estimated the populations of the Yuha Basin Management Area in 2002 and the East Mesa Management Area, Imperial County, California, USA, in 2003 to be 25,514 (95% CI 14,444-38,970) and 42,619 (95% CI 23,161-67,639), respectively. Two estimates of detection probability on the line in distance sampling by different methods were 0.45 and 0.65. Density estimates derived from distance analyses for 3 East Mesa Management Area plots and the Yuha Basin Management Area were 1.55 per ha (95% CI 0.64-3.76) and 0.41 per ha (95% CI 0.22-0.7), respectively. These are the first large-scale estimates of abundance and density for P. mcallii.     

The dazed and confused identity of Agassiz's land tortoise, Gopherus agassizii (Testudines, Testudinidae) with the description of a new species, and its consequences for conservation

We investigate a cornucopia of problems associated with the identity of the desert tortoise, Gopherus agassizii (Cooper). The date of publication is found to be 1861, rather than 1863. Only one of the three original cotypes exists, and it is designated as the lectotype of the species. Another cotype is found to have been destroyed in the 1906 San Francisco earthquake and subsequent fire. The third is lost. The lectotype is genetically confirmed to be from California, and not Arizona, USA as sometimes reported. Maternally, the holotype of Gopherus lepidocephalus (Ottley & Velázques Solis. 1989) from the Cape Region of Baja California Sur, Mexico is also from the Mojavian population of the desert tortoise, and not from Tiburon Island, Sonora, Mexico as previously proposed. A suite of characters serve to diagnose tortoises west and north of the Colorado River, the Mojavian population, from those east and south of the river in Arizona, USA, and Sonora and Sinaloa, Mexico, the Sonoran population. Species recognition is warranted and because Gopherus lepidocephalus is from the Mojavian population, no names are available for the Sonoran species. Thus, a new species, Gopherus morafkaisp. n., is named and this action reduces the distribution of Gopherus agassizii to only 30% of its former range. This reduction has important implications for the conservation and protection of Gopherus agassizii, which may deserve a higher level of protection.     

Estimating abundance and population trends when detection is low and highly variable: A comparison of three methods for the Hermann's tortoise

Assessing population trends is a basic prerequisite to carrying out adequate conservation strategies. Selecting an appropriate method to monitor animal populations can be challenging, particularly for low-detection species such as reptiles. This study compares 3 detection-corrected abundance methods (capture–recapture, distance sampling, and N-mixture) used to assess population size of the threatened Hermann's tortoise. We used a single dataset of 432 adult tortoise observations collected at 118 sampling sites in the Plaine des Maures, southeastern France. We also used a dataset of 520 tortoise observations based on radiotelemetry data collected from 10 adult females to estimate and model the availability (g0) needed for distance sampling. We evaluated bias for N-mixture and capture–recapture, by using simulations based on different values of detection probabilities. Finally, we conducted a power analysis to estimate the ability of the 3 methods to detect changes in Hermann's tortoise abundances. The abundance estimations we obtained using distance sampling and N-mixture models were respectively 1.75 and 2.19 times less than those obtained using the capture–recapture method. Our results indicated that g0 was influenced by temperature variations and can differ for the same temperature on different days. Simulations showed that the N-mixture models provide unstable estimations for species with detection probabilities <0.5, whereas capture–recapture estimations were unbiased. Power analysis showed that none of the 3 methods were precise enough to detect slow population changes. We recommend that great care should be taken when implementing monitoring designs for species with large variation in activity rates and low detection probabilities. Although N-mixture models are easy to implement, we would not recommend using them in situations where the detection probability is very low at the risk of providing biased estimates. Among the 3 methods allowing estimation of tortoise abundances, capture–recapture should be preferred to assess population trends. © 2013 The Wildlife Society.     

Methods for lion monitoring: a comparison from the Selous Game Reserve,Tanzania

The Selous Game Reserve in Tanzania is believed to contain Africa's largest population of lions (Panthera leo), making it a popular destination for trophy hunters and photographic tourists. However, a lack of recent data has raised concerns about the conservation status of this iconic population, so we collected two types of population data between 2006 and 2009. First, we identified 112 individual animals in an 800 km² study area in the photographic tourism part of Selous, giving a density of 0.14 individuals km^?2. This density estimate was similar to results using the same method from 1997 to 1999, but the adult sex ratio has decreased from 1 male : 1.3 female in 1997 to 1 male : 3 females in 2009. Second, using buffalo calf distress calls, we conducted call‐up surveys to census lions in three hunting sectors in the west, east and south of Selous and in the northern photographic area. Estimated adult lion densities varied from 0.02 to 0.10 km^?2, allowing an overall population estimate of 4,300 (range: 1,700–6,900). Our results highlight the value of call‐ups in surveying cryptic hunted carnivores but stress the importance of long‐term projects for calibrating the responses to call‐ups and for measuring trends in demography and population size.     

Comparison of Day and Night Fyke Netting, Electrofishing and Snorkelling for Monitoring a Population of the Threatened Golden Galaxias (Galaxias auratus)

The littoral zone of small off-stream water storage containing a translocated population of Galaxias auratus was sampled fortnightly at day and night with fyke nets, electrofishing and snorkelling over 3 months. Variation in population data provided by each method, including relative abundance indices, size structure, and habitat preferences, were examined. Aspects of behaviour and activity patterns were also investigated. Night sampling using all methods consistently yielded larger catches than day sampling. The size structure of catches varied, with electrofishing at night and fyke netting during the day having higher proportions of juveniles, whilst snorkelling at night and electrofishing during the day had higher proportions of adults. Fyke netting at night yielded by far the largest catches (~3-fold more than other methods) and also captured balanced proportions of juveniles and adults. Galaxias auratus had a strong diel activity pattern and were most active at night. The majority of the population migrated into the littoral zone during the night and back into deeper water during the day. A small number of juveniles remained in the littoral zone and some adults sheltered in the dense cover of species-rich littoral vegetation during the day. Shores with shallow depth profiles appeared to be preferred due to higher catches in these areas using all methods. Based on the results of this study, fyke netting at night in littoral habitats is recommended for monitoring populations of G. auratus. Fyke netting is likely to be an effective method for monitoring other lacustrine galaxiid species; however, further work is required to investigate the effects of habitat variables and fish community structure on activity patterns of galaxiids, and hence their catchability with various methods, in more extensive lentic environments.