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We examined overwintering behavior in gopher tortoises (Gopherus polyphemus) translocated to the northern periphery of their geographic range by using miniature temperature data loggers during 2 winters. All adult and juvenile tortoises monitored with temperature loggers survived overwintering; however, during the course of the study 2 translocated juvenile tortoises without temperature loggers died during winter months. Onset and termination of overwintering were not different between the 2 yrs and were not correlated with mean above-ground air temperature. Mean overwinter duration was 127 ± 9 d SD and 128 ± 13 d SD during 2002–2003 and 2004–2005, respectively. Tortoises experienced temperatures as low as 7°C and as high as 31°C while overwintering; however, most (12 of 15) tortoises experienced very little (< 1°C) mean daily temperature fluctuation despite air temperatures regularly dropping below 0°C and exceeding 20°C. The overall mean temperature of overwintering tortoises was 12.4° ± 0.8°C (2002–2003) and 12.6° ± 1.2°C (2004–2005). Large fluctuations in temperature occurred when tortoises actively basked, and half of the monitored tortoises did, particularly juveniles, which accounted for 42% of winter basking events. Our results suggest that, given timely access to suitable refugia at recipient sites, overwinter mortality of translocated adult individuals may be minimal.

Population viability analyses are useful tools to predict abundance and extinction risk for imperiled species. In southeastern North America, the federally threatened gopher tortoise (Gopherus polyphemus) is a keystone species in the diverse and imperiled longleaf pine (Pinus palustris) ecosystem, and researchers have suggested that tortoise populations are declining and characterized by high extinction risk. We report results from a 30-year demographic study of gopher tortoises in southern Alabama (1991–2020), where 3 populations have been stable and 3 others have declined. To better understand the demographic vital rates associated with stable and declining tortoise populations, we used a multi-state hierarchical markrecapture model to estimate sex- and stage-specific patterns of demographic vital rates at each population. We then built a predictive population model to project population dynamics and evaluate extinction risk in a population viability context. Population structure did not change significantly in stable populations, but juveniles became less abundant in declining populations over 30 years. Apparent survival varied by age, sex, and site; adults had higher survival than juveniles, but female survival was substantially lower in declining populations than in stable ones. Using simulations, we predicted that stable populations with high female survival would persist over the next 100 years but sites with lower female survival would decline, become male-biased, and be at high risk of extirpation. Stable populations were most sensitive to changes in apparent survival of adult females. Because local populations varied greatly in vital rates, our analysis improves upon previous demographic models for northern populations of gopher tortoises by accounting for population-level variation in demographic patterns and, counter to previous model predictions, suggests that small tortoise populations can persist when habitat is managed effectively.

Population manipulations such as translocation and head-starting are increasingly used as recovery tools for chelonians. However, evaluating success of individual projects can require decades of monitoring to detect population trends in these long-lived species. Furthermore, there are often few benchmarks from stable, unmanipulated populations against which to compare demographic rates, particularly for the immature stages. Here, we used 8 years of mark-recapture data to estimate apparent survival of immature gopher tortoises recruited into an introduced population of gopher tortoises (Gopherus polyphemus) first established on St. Catherines Island, Georgia, USA in 1987. During 2006 29 -2013, we conducted targeted trapping of immature gopher tortoises and compared survival of the hatchling, juvenile and subadult stages among treatments: 1) individuals released shortly after hatching from eggs obtained from gravid female founders (‘direct releases’); 2) individuals reared in captivity for 6-9 months following hatching (‘head-starts’); and 3) individuals first encountered as free-ranging, wild-recruited offspring (‘wild recruits’). Among the candidate models we examined, the best fit model included additive effects of tortoise stage and treatment, however, overlapping 95% credible intervals among treatments (CI) suggested that survival did not vary significantly among treatments. Annual apparent survival increased over the immature period, highlighting the importance of calculating separate estimates for the different immature stages. Across all treatments, the additive model estimated annual apparent survival probability to be 0.37 (CI: 0.25 40 – 0.48) for hatchlings, 0.71 (CI: 0.61 – 0.81) for juveniles, and 0.83 (CI: 0.74 – 0.94) for subadults. Our study, in combination with previous monitoring efforts at St. Catherines Island, provides strong evidence that the translocation and subsequent population augmentation efforts have been successful in establishing a robust population of gopher tortoises. Additionally, our results provide estimates of demographic rates for life stages that are poorly understood but critical to understanding population dynamics of this imperiled species.

Line-transect distance sampling (LTDS) surveys are commonly used to estimate abundance of animals or objects. In terrestrial LTDS surveys of gopher tortoise (Gopherus polyphemus) burrows, the presence of ground-level vegetation substantially decreases detection of burrows of all sizes, but no field or analytical methods exist to control for spatially heterogeneous vegetation obstruction as a source of variation in detection. We propose the addition of a simple measurement of groundlevel vegetation that serves as a covariate for the detection function. We present a Bayesian hierarchical model in which covariates burrow width and nearby vegetation height help to account for detection bias and improve precision of estimated density. We investigate the performance of this covariate by simulation and by using real LTDS data collected before and after application of prescribed fire. We collected data in 2018 at the Jones Center at Ichauway in Newton, Georgia, USA. Across all simulations, our model including both covariates produced the most accurate density point estimates of any of the models tested. For our case study, our Bayesian model with vegetation covariates tended to produce similar estimates of density before and after burns. Our study indicates that any level of spatial variation in vegetation obstruction decreases detection of burrows and may lead to underestimation in population size (≤68%) and proportion of individuals with small burrow sizes (≤32%) when not considered during analysis. Our work is extensible to other terrestrial sampling efforts where systematic measurement of a spatially distributed obstructing feature is feasible during the LTDS survey.

Gopher Tortoise (Gopherus polyphemus) populations are declining throughout the southeastern United States. Range-wide conservation efforts include identifying populations that do not currently meet viability criteria but are suitable candidates for population enhancement. We investigated the potential role of head starting as a recovery tool by releasing hatchling and head-started yearling Gopher Tortoises as pairs (n = 28) into adult burrows in fall and comparing their movements and survival until winter dormancy. Head-started yearlings experienced higher predicted survival than did hatchlings (87.7% vs. 56.5%). Head-started individuals also tended to move greater distances between burrows and established dormancy burrows further from their release burrows, but the differences were not significant. Most individuals of both groups used a small number of closely spaced burrows, although hatchlings took longer than head-started individuals to establish their first burrow (11.3 vs. 4.4 d on average) and a higher proportion were depredated or censored before establishing a burrow (35.7% vs. 10.7%). The paired release design provides strong experimental evidence that head-started yearling Gopher Tortoises experience at least a short-term survival advantage over hatchlings, while exhibiting comparable fidelity to the release site. Soft-release pens were not necessary to promote high site fidelity in our study, but the decision as to whether or not to use them at other release sites may be dictated by the predator community and what is practical to implement. We contend that head starting shows promise as a recovery tool for Gopher Tortoises and that hard releases may be a worthwhile option for managers to consider.

Camera trap time-lapse recordings can collect vast amounts of data on wildlife in their natural settings. Transforming these data into information useful to ecologists is a major challenge. Machine learning techniques show promise for becoming important tools in the cost-effective analysis of camera trap data, but only if they become readily available to researchers without requiring advanced computing skills and resources. We present a new suite of software tools that reduce the amount of human effort needed to segment time-lapse, camera trap recordings in preparation for analysis. The tools incorporate a convolutional neural network trained to detect a focal species and to generate a draft video segmentation indicating the ranges of time when the focal species is present. We evaluated the utility of our neural network by comparing manual and automatic segmentations of 64 time-lapse recordings of gopher tortoise (Gopherus polyphemus) burrows, recorded in Pinellas County, Florida, USA between 25 November 2020 and 30 November 2020. The neural network correctly found 130 of the 145 segments containing tortoises (89.7%), whereas student graders found 135 segments (93.1%). A year of experience using the new software suite in an ongoing study of gopher tortoises deploying 12 camera traps indicates one person, assisted by machine learning algorithms, can segment a week's worth of time-lapse recordings—11.5 hours of standard-speed video—in under 3 hours. We concluded that the use of machine learning algorithms is practical and allows researchers to process large volumes of time-lapse data with minimal human effort.

Gopher tortoises (Gopherus polyphemus) are among the most commonly translocated reptiles. Waif tortoises are animals frequently of unknown origin that have been displaced from the wild and often held in human possession for various reasons and durations. Although there are risks associated with any translocation, waif tortoises are generally excluded from translocation projects because of heightened concerns of introducing pathogens and uncertainty about the post-release survival of these individuals. If these risks could be managed, waif tortoises could have conservation value because they can provide the needed numbers to stabilize populations. In the early 1990s, the discovery of an isolated population of gopher tortoises (≤15 individuals) near Aiken, South Carolina, USA, prioritized establishment of the Aiken Gopher Tortoise Heritage Preserve (AGTHP). Because of the population’s need for augmentation and the site’s isolation from other tortoise populations, the AGTHP provided the opportunity to evaluate the post-release survival of translocated waif tortoises without compromising a viable population. Since 2006, >260 waif tortoises have been introduced to the preserve. Using a Cormack-Jolly-Seber modeling framework to analyze release records and capture histories from trapping efforts in 2017 and 2018, we estimated the long-term apparent survival and site fidelity of this population composed largely of waif tortoises. We estimated annual apparent survival probabilities to be high (≥0.90) for subadult, adult male, and adult female tortoises, and these rates were similar to those reported for wild-to-wild translocated gopher tortoises and those from unmanipulated populations. Of the tortoises recaptured within the boundaries of the preserve, 75% were located within 400m of their release location. These results suggest that waif tortoises could be an important resource in reducing the extirpation risk of isolated populations.

Population manipulations such as translocation and head-starting are increasingly used as recovery tools for chelonians. But evaluating success of individual projects can require decades of monitoring to detect population trends in these long-lived species. Furthermore, there are often few benchmarks from stable, unmanipulated populations against which to compare demographic rates, particularly for the immature stages. We used 8 years of mark-recapture data to estimate apparent survival of immature gopher tortoises (Gopherus polyphemus) recruited into an introduced population of gopher tortoises first established on St. Catherines Island, Georgia, USA, in 1987. During 2006–2013, we conducted targeted trapping of immature gopher tortoises and compared survival of the hatchling, juvenile and subadult stages among treatments: individuals released shortly after hatching from eggs obtained from gravid female founders (direct releases), individuals reared in captivity for 6–9 months following hatching (head-starts), and individuals first encountered as free-ranging, wild-recruited offspring (wild recruits). Among the candidate models we examined, the best fit model included additive effects of tortoise stage and treatment; however, overlapping 95% credible intervals among treatments (CrI) suggested that survival did not vary significantly among treatments. Annual apparent survival increased over the immature period, highlighting the importance of calculating separate estimates for the different immature stages. Across all treatments, the additive model estimated annual apparent survival probability to be 0.37 (CrI=0.25–0.48) for hatchlings, 0.71 (CrI=0.61–0.81) for juveniles, and 0.83 (CrI=0.74–0.94) for subadults. Our study, in combination with previous monitoring efforts at St. Catherines Island, provides strong evidence that the translocation and subsequent population augmentation efforts have been successful in establishing a robust population of gopher tortoises. Additionally, our results provide estimates of demographic rates for life stages that are poorly understood but critical to understanding population dynamics of this imperiled species.

An inherent challenge in managing rare or cryptic species is data deficiency. For this reason, ancillary data is a potentially valuable resource for generating key population estimates for priority species. We compiled ancillary commensal data collected between 1982 and 2020 during surveys of gopher tortoise (Gopherus polyphemus) burrows to estimate the terrestrial distribution of gopher frogs (Rana capito) from potential breeding wetlands. Gopher frogs were detected in gopher tortoise burrows 30‒3,879 m from identified wetlands. A global model of all records from all sites indicated that the probability of a gopher frog residing in a gopher tortoise burrow declined with increasing distance from a wetland. This pattern also held for 4 of 5 sites with a sufficient number of gopher frog detections to model independently. Based on the full data set, we estimated that 50%, 90%, and 99% of gopher frog observations occurred within 392 m, 1,019 m, and 2,752 m of the nearest wetland, respectively. Our results indicate a higher proportion of gopher frogs emigrate longer distances from wetlands than was previously reported using other methods, such as radio-telemetry. This information can directly assist with management decisions, notably the spatial extent for application of habitat management surrounding breeding wetlands. More generally, this study illustrates the capacity of ancillary data to fill data deficiencies for a rare and cryptic species and highlights the importance of these data.

Conservation biologists need to effectively monitor species given resource limitations and the inherent challenges of assessing long-term demographic processes. We assessed gopher tortoise (Gopherus polyphemus) abundance at a landscape scale and at the scale of 3 local populations within the Conecuh National Forest (CNF), Alabama, USA, between 1991 and 2017. We collected landscape-level data from line transect distance sampling arranged uniformly across the CNF during a single season (2011); we obtained data for local populations from long-term mark-recapture of individuals at 3 sites selected based on prior knowledge of high density at each. At a landscape scale, we estimated 5,242 (95% CI=3,538–7,768) tortoises occurred across the approximately 34,000-ha forest, yielding a density of 0.14–0.32 tortoises/ha. These low densities across the landscape suggest that, on average, management activities across the property have not allowed tortoise populations to retain the social structure needed for long-term persistence. The 3 local populations, however, contained 25–60 individuals and densities of 1.9–6.9 tortoises/ha. Over the study period, populations at 2 sites were stable and the third experienced significant population growth. Mean annual survival of individuals was 0.89 and invariant across size classes. Overall, line transect distance sampling is important for assessing landscape-scale abundance of tortoises but may fail to detect local clusters of high-density sites important for population persistence. Our mark-recapture efforts at the local scale revealed that small populations on these high-density sites can exhibit long-term stability or growth even though they do not meet current established criteria for viability. Improved models that incorporate immigration and emigration and better reflect the dynamics of peripheral populations would assist in determining how such populations best contribute to species recovery and regional conservation targets.