Tortoise Densities Research Articles

Simulation Modeling to Assess Line Transect Distance Sampling Under a Range of Translocation Scenarios

Abstract The accuracy of posttranslocation population monitoring methods is critical to assessing long-term success in translocation programs. Translocation can produce unique challenges to monitoring efforts; therefore, it is important to understand the flexibility and robustness of commonly used monitoring methods. In Florida, USA, thousands of gopher tortoises Gopherus polyphemus have been, and continue to be, translocated from development sites to permitted recipient sites. These recipient sites create a broad range of potential monitoring scenarios due to variability in soft-release strategies, habitat conditions, and population demographics. Line transect distance sampling is an effective method for monitoring natural tortoise populations, but it is currently untested for translocated populations. We therefore produced 3,024 individual-based, spatially explicit scenarios of translocated tortoise populations that differed in recipient site and tortoise population properties, based on real-world examples, literature review, and expert opinion. We virtually sampled simulated tortoise populations by using line transect distance sampling methods and built a Bayesian hierarchical model to estimate the population density for each simulation, which incorporated individual-level covariates (i.e., burrow width and burrow occupancy). Line transect distance sampling was largely appropriate for the conditions that typify gopher tortoise recipient sites, particularly when detection probability on the transect lines was greater than or equal to 0.85. Designing the layout of transects relative to the orientation of soft-release pens, to avoid possible sampling biases that lead to extreme outliers in estimates of tortoise densities, resulted in more accurate population estimates. We also suggest that use of individual-level covariates, applied using a Bayesian framework as demonstrated in our study, may improve the applicability of line transect distance sampling surveys in a variety of contexts and that simulation can be a powerful tool for assessing survey design in complex sampling situations.

A spatially explicit model for density that accounts for availability: a case study withMojavedesert tortoises

AbstractEstimating population density and identifying those areas where density is changing through time are central to prioritizing conservation and management strategies. Obtaining reliable estimates of density and trends can be challenging, however, especially for long‐lived species that are rare, have broad geographic distributions, and are difficult to detect reliably during field surveys. We developed a hierarchical model for distance‐sampling data that characterizes spatial variation in density at two scales and simultaneously estimates regional trends while accounting for variation in detection probability and availability across surveys. We applied the model to data collected over a 20‐year period (2001–2020) in an area that encompassed most of the geographic range of the Mojave desert tortoise (Gopherus agassizii). Density of adult tortoises varied with multiple biotic and abiotic features, including topography, aspect, geology, and seasonal precipitation and temperature regimes. Across the entire period and study area, the density of adult tortoises decreased by an average of 1.8% per year (95% CI = −3.5% to −0.2%). Trends varied geographically, however, with the steepest declines in the western part of the range (−4.1%, −6.9% to −1.3%). Accounting for habitat loss across our study area, the abundance of this threatened species declined by an estimated 129,000 adults (36%) between 2001 and 2020. Our modeling approach extends traditional distance‐sampling frameworks by accounting for ecological and observational processes that could mask spatiotemporal variation in density and, at the same time, provides spatially explicit estimates to guide conservation and management strategies for tortoises and other rare species.

Sharing land with giants: Habitat preferences of Galapagos tortoises on farms

One of the most pressing dilemmas of our time is determining how to satisfy the demands of a growing human population while still conserving biodiversity. Worldwide, land modification to accommodate human resource needs has caused significant declines in wildlife populations. To help minimize biodiversity loss, we must support wildlife on human-dominated land, such as farms and urban areas, but our knowledge of how to do so is lacking. Agriculture is a major driver of land modification; but also has the potential to play a role in conserving biodiversity. To support critically endangered ecosystem engineers that use farms, such as giant Galapagos tortoises, we need to understand the characteristics encouraging or hindering them. To quantify tortoise habitat preferences, we assessed the relationship between tortoise density, habitat structure, and land-use type, by recording tortoise density on farms on Santa Cruz Island, Galapagos, over two years. Tortoise density was lowest in abandoned farmland and highest in tourist areas and was most strongly positively correlated with abundant ground cover, short vegetation, and few shrubs. The habitat features favoured by tortoises could potentially be manipulated to help support tortoise conservation on farms. Measuring wildlife preferences in human-dominated areas is an important step towards balancing biodiversity conservation and human-enterprise. • Wildlife preferences should be measured on human-dominated land. • Habitat structure on farms impacts wildlife density. • Agricultural land can support wildlife. • Abandoned land supports the least tortoises. • Land-use type impacts the density of giant tortoises in farmland.

High female desert tortoise mortality in the western Sonoran Desert during California’s epic 2012-2016 drought

We conducted population surveys for desert tortoisesGopherus agassiziiat 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 periods of resource limitation. At the higher-elevation, cooler, wetter Cottonwood site from 2015-2016, the sex ratio of live adult tortoises was biased toward males and the sex ratio of tortoises estimated to have died during the intensified drought conditions from 2012-2016 was essentially even. At the lower-elevation, warmer, drier Orocopia site from 2017-2018, the sex ratio of live adult tortoises was biased toward males and the sex ratio of tortoises with estimated times of death from 2012-2016 was biased toward females. High female mortality at the Orocopia site may have resulted from the interaction of drought effects and the bet-hedging reproductive strategy of tortoises wherein they continue to produce clutches of eggs in drought years. Annual reproductive output results in an estimated loss of up to 13.5% of female tortoise body mass including over 0.20 l of water. Combined with dehydration during severe droughts, these losses may compromise their ability to survive droughts lasting more than 2 yr. The low tortoise density and high mortality of females observed may reflect reduced survival of tortoises near the southern edge of their range due to climate change, including protracted and intensified droughts.

Influence of Habitat and Food Resource Availability on Common Raven Nest Site Selection and Reproductive Success in Mediterranean Forests

Bird nest selection in forests can be influenced by the composition of key structural elements and resources. This has important consequences in terms of species population dynamics since it can determine reproduction success. Here, we assessed Common raven nest-site selection and reproductive success, and how these might be determined by foraging behavior and habitat structure. A previously documented breeding raven population that exerts high predation pressure on young Spur-thighed tortoises (Testudo graeca) in a Mediterranean forest was monitored. Generalized linear mixed models were performed to determine the singularities of the trees with nests and the drivers of reproductive success of breeding pairs of ravens. The results showed a high density of breeding pairs in the study area (0.8 pairs/km2), which selected taller trees in areas with higher bare ground cover and a high density of tortoises for nesting. Nests were spatially aggregated; breeding pairs occupied smaller territories and intraspecific competition seemed relaxed, reflecting the abundance of food resources. Most breeding pairs occasionally predated on young tortoises. Tortoises seem to play a part in raven reproductive success in our study area, which might be associated with the availability/catchability of young tortoises. The study illustrates that Spur-thighed tortoise distribution and abundance plays a role in the breeding behavior of ravens and is mediated by habitat structure. Understanding the drivers of nest-site selection and the breeding behavior of ravens is pivotal to implementing appropriate habitat management and conservation strategies across their distribution range, particularly in areas where ravens potentially affect threatened species.

The Catastrophic Decline of Tortoises at a Fenced Natural Area

ABSTRACTAgassiz’s desert tortoise (Gopherus agassizii), a threatened species of the southwestern United States, has severely declined to the point where 76% of populations in critical habitat (Tortoise Conservation Areas) are below viability. The potential for rapid recovery of wild populations is low because females require 12–20 years to reach reproductive maturity and produce few eggs annually. We report on a 34‐year mark‐recapture study of tortoises initiated in 1979 at the Desert Tortoise Research Natural Area in the western Mojave Desert, California, USA, and provide substantive data on challenges faced by the species. In 1980, the United States Congress designated the Research Natural Area and protected the land from recreational vehicles, livestock grazing, and mining with a wildlife‐permeable fence. The 7.77‐km2study area, centered on interpretive facilities, included land both within the Natural Area and outside the fence. We expected greater benefits to accrue to the tortoises and habitat inside compared to outside. Our objectives were to conduct a demographic study, analyze and model changes in the tortoise population and habitat, and compare the effectiveness of fencing to protect populations and habitat inside the fence versus outside, where populations and habitat were unprotected. We conducted surveys in spring in each of 7 survey years from 1979, when the fence was under construction, through 2012. We compared populations inside to those outside the fence by survey year for changes in distribution, structure by size and relative age, sex ratios, death rates of adults, and causes of death for all sizes of tortoises. We used a Bayesian implementation of a Jolly Seber model for mark‐recapture data. We modeled detection, density, growth and transition of tortoises to larger size‐age classes, movements from inside the protective fence to outside and vice versa, and survival. After the second and subsequent survey years, we added surveys to monitor vegetation and habitat changes, conduct health assessments, and collect data on counts of predators and predator sign. At the beginning of the study, counts and densities for all sizes of tortoises were high, but densities were approximately 24% higher inside the fence than outside. By 2002, the low point in densities, densities had declined 90% inside the fence and 95% outside. Between 2002 and 2012, the population inside the fence showed signs of improving with a 54% increase in density. Outside the fence, densities remained low. At the end of the study, when we considered the initial differences in location, densities inside the fence were roughly 2.5 times higher than outside. The pattern of densities was similar for male and female adults. When evaluating survival by blocks of years, survivorship was higher in 1979–1989 than in 1989–2002 (the low point) and highest from 2002 to 2012. Recruitment and survival of adult females into the population was important for growing the population, but survival of all sizes, including juveniles, was also critical.Major events and activities driving the decline in populations both inside and outside the fence included illegal collecting, upper respiratory tract disease, and hyperpredation by the common raven (Corvus corax) on juvenile tortoises. Other sources of death were gunshots, vehicles, and predation by mammals. Outside the fence, fragmentation and deterioration of habitat was a critical driver. Between the first and last surveys, 2 different ecosystem processes were underway: recovery of vegetation and soils from grazing and vehicles inside the fence and continued deterioration outside the fence. Habitat outside the fence became increasingly denuded of shrubs and fragmented by roads and trails, and habitat fragments increased 50‐fold. Outside the fence, biomass of non‐native annual plants was higher and the cover of shrubs was lower, a reflection of ongoing deterioration. These changes and losses of habitat resulted in loss of shrub cover and sites for burrows, reduction in preferred food plants, and greater exposure to predators and extremes in temperature. Overall, the tortoise population and habitat inside the fence appeared to benefit from protection and showed signs of recovery at the end of the study. Fencing, control of vehicular access, and removal of livestock grazing were among several recommended management actions for critical habitat in the first recovery plan in 1994. At the end of the study, the Natural Area remained as 1 of 2 fenced, official protected areas for the species in the geographic range. We attribute fencing to continuing higher densities of adults inside the fence compared with outside the fence and promising signs of recovery. Densities of adults at the Natural Area also were 2.3 to 5.5 times higher than in 16 of the 17 Tortoise Conservation Areas (critical habitat units) within the geographic range. © 2020 The Authors.Wildlife Monographspublished by Wiley Periodicals, LLC on behalf of The Wildlife Society.

Epidemiological Investigation of a Mortality Event in a Translocated Gopher Tortoise (Gopherus polyphemus) Population in Northwest Florida.

Nokuse Plantation, a 22,055 ha private conservation preserve in northwest Florida, is a recipient site for gopher tortoises translocated from development sites in Florida. Since 2006, Nokuse has received over 5,000 tortoises from multiple development sites. During 2013–2015, 52 tortoises were found sick (n = 14) or dead (n = 38) in multiple soft-release enclosures in which tortoises consistently exhibited clinical signs, with additional sick (n = 5) and dead (n = 5) tortoises presenting similarly during 2016–2017. When found alive, tortoises behaved abnormally (e.g., frequently out of burrows during cold weather, pacing along enclosure fencing), appeared emaciated, were lethargic, and had developed redness under plastron scutes. Similar numbers of male (n = 28) and female (n = 32) tortoises were recovered along with two of unidentified sex, including mainly adults (n = 59) and three subadults. Physical examination, blood analysis, and other diagnostics were indicative of starvation and dehydration. Most sick tortoises provided with supportive care recovered. Necropsy findings generally confirmed starvation, with no evidence of infectious pathogens or contaminants. There were no apparent differences in quality of habitat, plant community, or soil or water among affected and unaffected enclosures. Botanical surveys indicated adequate forage quality and quantity, with no poisonous exotic or native plants detected. No land management practices changed prior to this event. Analysis of epidemiological data and demographic factors from before and during this mortality event identified initial density of tortoises in the enclosures as exerting the strongest influence on detection of tortoise morbidity and mortality. We believe that the stress associated with mixing tortoises from different populations and at higher densities during translocation impacted an individual tortoise's ability to obtain or absorb adequate nutrients from foraging, ultimately leading to a wasting condition consistent with starvation. Based on our findings, we recommend a maximum of 3 gopher tortoises per ha in soft-release enclosures for translocation, but further research is warranted to investigate the complexity of stress and social pressures associated with translocation.

Population density of the spur-thighed tortoise Testudo graeca declines after fire in north-western Africa.

Fire is a key ecological process in several biomes worldwide. Over recent decades, human activities (e.g. rural abandonment, monoculture plantations) and global warming are magnifying the risk of fire, with changes in fire intensity and frequency. Here, we offer the first study that examines the impact of fire on the spur-thighed tortoise Testudo graeca living in a native cork oak forest and pine plantation in north-western Africa. A total of 44 transects (22 burnt and 22 unburnt) were sampled at 8 sites affected by fires of natural cork oak forest and pine plantation with 8 surveys per site in 2015–2017 (264 hours of sampling effort). Tortoise densities were estimated with line-transect distance sampling. The detection probability of tortoises was higher in burnt (0.915) than unburnt (0.474) transects. The density of tortoises was negatively associated with elevation and declined with fire by c. 50% in both forest types. The negative response of T. graeca to fire should be considered in conservation planning of this species in north-western Africa in a future scenario of changes in fire regime.

Coyote diet patterns in the Mojave Desert: implications for threatened desert tortoises

Coyotes (Canis latrans) are generalist predators and are ubiquitous in North America. Occasionally, predation by coyotes can pose a threat to populations of rare species. We assessed diet patterns of coyotes over a 5-year period (2009–14) in a region of the Mojave Desert where high predation rates on threatened desert tortoises (Gopherus agassizii) had been reported. Our goal was to identify primary food items for coyotes and to assess the importance of desert tortoises in the diet. Coyotes primarily consumed rabbits and rodents with rabbits being consumed preferentially and rodents, along with secondary foods including various birds, reptiles, arthropods, and fruits, being consumed more opportunistically. In response to low annual precipitation in the last three years of the study, dietary diversity increased, as did use of anthropogenic food items by coyotes. However, coyotes did not seem to be dependent upon anthropogenic items. Remains of desert tortoises occurred in coyote scats at low frequencies (<6%) in all years and seasons, and use of tortoises appeared to be opportunistic as use varied with tortoise abundance. In the portion of the study area where 571 translocated desert tortoises had been released in 2008, the frequencies of tortoise remains in coyote scats were markedly higher in the two years following the releases (7.5% and 8.8%, respectively). The high predation rates on tortoises reported in this area may have resulted from focussed coyote foraging efforts due to the availability of vulnerable individuals (e.g. disoriented and displaced tortoises) as well as higher tortoise densities.

Distance-Generated Field Density Estimates for the Threatened Sahel Tortoise <i>Centrochelys sulcata</i

In this paper, the results of the first density investigations (via Distance methodology on line transect surveys) for the threatened African spurred tortoise Centrochelys sulcata from West African Sahel (Niger and Burkina Faso) are reported. Spurred tortoises were encountered in 51.8% of Burkina Faso transects ( n = 27) and in 60.7% of Niger transects. In several transects we found evidence of the tortoise presence, but without actually seeing any individual. Pooling all the transects from which tortoise density was calculated, the mean density of tortoises in Burkina Faso was 0.0021 ind./ha, and in Niger was 0.167 ind./ha. Nonetheless, there was a considerable variability in density estimates by transect in both countries, and especially in Teneré-Termit (Niger), the density of tortoises was very high (over 2 ind./ha). Teneré-Termit region in Niger may therefore represent a potentially crucial area for the conservation of this declining species.

Distance-Generated Field Density Estimates for the Threatened Sahel Tortoise <i>Centrochelys sulcata</i

In this paper, the results of the first density investigations (via Distance methodology on line transect surveys) for the threatened African spurred tortoise Centrochelys sulcata from West African Sahel (Niger and Burkina Faso) are reported. Spurred tortoises were encountered in 51.8% of Burkina Faso transects ( n = 27) and in 60.7% of Niger transects. In several transects we found evidence of the tortoise presence, but without actually seeing any individual. Pooling all the transects from which tortoise density was calculated, the mean density of tortoises in Burkina Faso was 0.0021 ind./ha, and in Niger was 0.167 ind./ha. Nonetheless, there was a considerable variability in density estimates by transect in both countries, and especially in Teneré-Termit (Niger), the density of tortoises was very high (over 2 ind./ha). Teneré-Termit region in Niger may therefore represent a potentially crucial area for the conservation of this declining species.

Techniques for Estimating the Size of Low-Density Gopher Tortoise Populations

AbstractGopher tortoises (Gopherus polyphemus) are candidates for range-wide listing as threatened under the U.S. Endangered Species Act. Reliable population estimates are important to inform policy and management for recovery of the species. Line transect distance sampling has been adopted as the preferred method to estimate population size. However, when tortoise density is low, it can be challenging to obtain enough tortoise observations to reliably estimate the probability of detection, a vital component of the method. We suggest a modification to the method based on counting usable tortoise burrows (more abundant than tortoises) and separately accounting for the proportion of burrows occupied by tortoises. The increased sample size of burrows can outweigh the additional uncertainty induced by the need to account for the proportion of burrows occupied. We demonstrate the method using surveys conducted within a 13,118-ha portion of the Gopher Tortoise Habitat Management Unit at Fort Gordon Army Installation, Georgia. We used a systematic random design to obtain more precise estimates, using a newly developed systematic variance estimator. Individual transects had a spatially efficient design (pseudocircuits), which greatly improved sampling efficiency on this large site. Estimated burrow density was 0.091 ± 0.011 burrows/ha (CV = 12.6%, 95% CI = 0.071–0.116), with 25% of burrows occupied by a tortoise (CV = 14.4%), yielding a tortoise density of 0.023 ± 0.004 tortoise/ha (CV = 19.0%, 95% CI = 0.016–0.033) and a population estimate of 297 tortoises (95% CI = 210–433). These techniques are applicable to other studies and species. Surveying burrows or nests, rather than animals, can produce more reliable estimates when it leads to a significantly larger sample of detections and when the occupancy status can reliably be ascertained. Systematic line transect survey designs give better precision and are practical to implement and analyze.

Adapting coastal management to climate change: Mitigating our shrinking shorelines

ABSTRACTSea‐level rise due to climate change is a major threat to coastal ecosystems worldwide. Current management to reduce beach erosion often focuses on protecting human structures and research on effects on wildlife is lacking. Using a combination of hierarchical models and generalized linear models, we evaluated how the gopher tortoise (Gopherus polyphemus) colonized constructed dunes along coastal scrub at the Merritt Island National Wildlife Refuge, central Florida, USA. Over 2 years, we surveyed tortoise populations along natural dunes and 2 constructed dunes (completed in 2012 and in 2014) and estimated tortoise density each summer and winter. Our models indicated that tortoise density along the 2014 dune was comparable to that of natural dunes ( = 0–8 tortoises/ha), and density peaked at a mean of 21 tortoises/ha along the 2012 constructed dune. Gopher tortoises rapidly colonized constructed dunes, and dune construction may represent effective management against habitat loss for this species. © 2017 The Wildlife Society.

Low tortoise abundances in pine forest plantations in forest-shrubland transition areas.

In the transition between Mediterranean forest and the arid subtropical shrublands of the southeastern Iberian Peninsula, humans have transformed habitat since ancient times. Understanding the role of the original mosaic landscapes in wildlife species and the effects of the current changes as pine forest plantations, performed even outside the forest ecological boundaries, are important conservation issues. We studied variation in the density of the endangered spur-thighed tortoise (Testudo graeca) in three areas that include the four most common land types within the species’ range (pine forests, natural shrubs, dryland crop fields, and abandoned crop fields). Tortoise densities were estimated using a two-stage modeling approach with line transect distance sampling. Densities in dryland crop fields, abandoned crop fields and natural shrubs were higher (>6 individuals/ha) than in pine forests (1.25 individuals/ha). We also found large variation in density in the pine forests. Recent pine plantations showed higher densities than mature pine forests where shrub and herbaceous cover was taller and thicker. We hypothesize that mature pine forest might constrain tortoise activity by acting as partial barriers to movements. This issue is relevant for management purposes given that large areas in the tortoise’s range have recently been converted to pine plantations.

A comparison of line-transect distance sampling methods for estimating gopher tortoise population densities

Line-transect distance sampling (LTDS) is increasingly used to estimate gopher tortoise (Gopherus polyphemus) densities. The process requires detecting tortoise burrows, and then determining occupancy. We compared 3 LTDS approaches that differ in how burrow detection and occupancy data are integrated, and 2 search strategies. Surveys were conducted at Avon Park Air Force Range in South-central Florida, USA, from April through October 2009. These include 1) LTDS using data from occupied burrows only; 2) LTDS with burrows treated as “clusters” of size 1 or 0 depending on occupancy; 3) LTDS to estimate burrow densities, with occupancy modeling to estimate occupancy rates; and 4) 2 search strategies: standard versus expanded searches. The LTDS method produced reliable burrow density estimates, and tortoise densities were estimated most precisely and with least effort using data from occupied burrows only. However, any method could be biased if burrow occupancy is uncertain. Indeed, in the Florida scrub and pine flatwoods habitats, video-camera scoping was hindered by groundwater or obstructions, resulting in undetermined occupancy for 45% of burrows. Improving scope technologies and surveying during the dry season has improved subsequent results for xeric habitats; however, research is needed to improve detectability of tortoises in mesic habitats with flooded burrows. If occupancy is uncertain, we recommend collecting data on all burrows, irrespective of occupancy, so that burrow densities can be estimated even if occupancy cannot. We do not recommend expanding the search area with sigmoid search paths adjacent to transects owing to potential violations of assumptions, nor use of occupancy modeling, because model requirements may lead to unacceptable exclusion of data. © 2015 The Wildlife Society.

A plant invasion affects condition but not density or population structure of a vulnerable reptile

By altering ecosystem structure and function, invasions by nonnative plants have the potential to alter the quantity and quality of habitat for animals. We examined effects of buffelgrass (Pennisetum ciliare), a nonnative grass that is increasing in distribution markedly throughout the southwestern United States and northern Mexico, on demographic characteristics and condition of Sonoran desert tortoises (Gopherus morafkai). In 2010 and 2011, we established 50 4-ha plots that spanned the gradient of buffelgrass cover in areas with environmental features characteristic of high-quality habitat for tortoises in southern Arizona. We surveyed each plot four times per year to characterize density, population structure, and condition of tortoises. We detected tortoises on 45 of 50 plots (90 %) and ≥1 tortoise during 114 of 200 surveys (57 %). We used a hierarchical model to estimate density of tortoises while accounting for imperfect detection rates. Density of tortoises averaged 0.35 individuals/ha (SE = 0.04) and did not vary appreciably with the amount of buffelgrass cover; similarly, age and sex structure of tortoise populations did not vary with buffelgrass cover. Condition of adult tortoises, however, averaged 10 % lower in areas where cover of buffelgrass was high (>15 %) relative to areas where buffelgrass was absent or cover was low (<1 %). Although the demographic characteristics we measured on this long-lived species did not vary with current levels of buffelgrass cover, reduced condition of tortoises in areas invaded by this nonnative grass could manifest as population-level effects over longer time periods, especially because buffelgrass is predicted to expand its current distribution.

Protection Benefits Desert Tortoise (Gopherus agassizii) Abundance: The Influence of Three Management Strategies on a Threatened Species

We surveyed an area of ∼260 km2 in the western Mojave Desert to evaluate relationships between condition of Agassiz's Desert Tortoise populations (Gopherus agassizii) and habitat on lands that have experienced three different levels of management and protection. We established 240 1-ha plots using random sampling, with 80 plots on each of the three types of managed lands. We conducted surveys in spring 2011 and collected data on live tortoises, shell-skeletal remains, other signs of tortoises, perennial vegetation, predators, and evidence of human use. Throughout the study area and regardless of management area, tortoise abundance was positively associated with one of the more diverse associations of perennial vegetation. The management area with the longest history of protection, a fence, and legal exclusion of livestock and vehicles had significantly more live tortoises and lower death rates than the other two areas. Tortoise presence and abundance in this protected area had no significant positive or negative associations with predators or human-related impacts. In contrast, the management area with a more recent exclusion of livestock, limited vehicular traffic, and with a recent, partial fence had lower tortoise densities and high death rates. Tortoise abundance here was negatively associated with vehicle tracks and positively associated with mammalian predators and debris from firearms. The management area with the least protection—unfenced, with uncontrolled vehicle use, sheep grazing, and high trash counts—also had low tortoise densities and high death rates. Tortoise abundance was negatively associated with sheep grazing and positively associated with trash and mammalian predator scat.

Climatic variation and tortoise survival: Has a desert species met its match?

While demographic changes in short-lived species may be observed relatively quickly in response to climate changes, measuring population responses of long-lived species requires long-term studies that are not always available. We analyzed data from a population of threatened Agassiz’s desert tortoises (Gopherus agassizii) at a 2.59km2 study plot in the Sonoran Desert ecosystem of Joshua Tree National Park, California, USA from 1978 to 2012 to examine variation in apparent survival and demography in this long-lived species. Transect-based, mark-recapture surveys were conducted in 10 of those years to locate living and dead tortoises. Previous modeling suggested that this area would become unsuitable as tortoise habitat under a warming and drying climate scenario. Estimated adult population size declined greatly from 1996 to 2012. The population appeared to have high apparent survival from 1978 to 1996 but apparent survival decreased from 1997 to 2002, concurrent with persistent drought. The best model relating apparent survivorship of tortoises ⩾18cm over time was based on a three year moving average of estimated winter precipitation. The postures and positions of a majority of dead tortoises found in 2012 were consistent with death by dehydration and starvation. Some live and many dead tortoises found in 2012 showed signs of predation or scavenging by mammalian carnivores. Coyote (Canis latrans) scats and other evidence from the site confirmed their role as tortoise predators and scavengers. Predation rates may be exacerbated by drought if carnivores switch from preferred mammalian prey to tortoises during dry years. Climate modeling suggests that the region will be subjected to even longer duration droughts in the future and that the plot may become unsuitable for continued tortoise survival. Our results showing wide fluctuations in apparent survival and decreasing tortoise density over time may be early signals of that possible outcome.

Densities of Ecological Replacement Herbivores Required to Restore Plant Communities: A Case Study of Giant Tortoises on Pinta Island, Galápagos

AbstractLoss of native herbivores and introduction of livestock in many arid and semi‐arid ecosystems around the world has shifted the competitive balance from herbaceous to woody plants, leading to biodiversity loss, reduced plant productivity, and soil erosion. To restore functions of these ecosystems, ecological replacements have been proposed as substitutes for extinct native herbivores. Here we predict how an ecological replacement giant tortoise population (Chelonoidis spp.) would interact with woody plants on Pinta Island in the Galápagos Archipelago, where a small group of replacement tortoises was introduced in 2010 to initiate restoration of the island's plant community. We developed an individual‐based, spatially explicit simulation model that incorporated field‐derived tortoise behavior and tortoise–plant interaction data to test whether tortoise introductions could lead to broad‐scale changes in the plant community and, if so, at what tortoise densities. Tortoises reduced vegetation density in most (81%) 50‐year‐long simulations if the tortoise density was at least 0.7 per hectare, a value well below typical densities. In a smaller proportion of simulations (30%), tortoises increased local vegetation patchiness. Our results suggest that even moderate‐density tortoise populations can reverse woody plant encroachment. Deployment of ecological replacement giant tortoises may therefore be a viable approach for restoring other arid and semi‐arid ecosystems where a native herbivore that previously had strong interactions with the plant community has gone extinct.

Intestinal parasites of the gopher tortoise (Gopherus polyphemus) from eight populations in Georgia

The gopher tortoise (Gopherus polyphemus), one of five tortoise species endemic in the USA, was recently classified as a candidate for federal listing as a threatened species. Fecal samples collected from 117 tortoises from eight sites in Georgia were examined for endoparasites using a combination of sedimentation and flotation. Samples from an island population were examined for parasitic oocysts and ova only by flotation, protozoan cysts by trichrome-stained direct smear, and Cryptosporidium by direct immunofluorescence assay and ProSpecT rapid assay. A total of 99 tortoises (85, range 0-100%) was infected with pinworms (Alaeuris spp.), 47 (40, 0-86%) with cestodes (Oochorstica sp.), 34 (41, 0-74%) with Chapiniella spp., 2 (3, 0-33%) with Eimeria paynei, and a single tortoise each with a capillarid and ascarid (1%). On the island, Entamoeba was detected in one tortoise (2%) while Cryptosporidium oocysts were detected in eight (17%). In conclusion, at least eight species of parasites were detected including Cryptosporidium, a possible pathogen of tortoises. Interestingly, we detected spatial variation in the distribution of several parasites among populations suggesting additional work should be conducted across a gradient of tortoise densities, land use, and habitat characteristics.