Tortoise Survival Research Articles

Unexpected effect of geographic origin on post‐translocation survival in a long‐lived reptile, the gopher tortoise

AbstractMitigation translocations move wildlife from specific areas due to conflict with humans over land use at the site. A critical decision when carrying out mitigation translocation is the acceptable distance across which animals can be moved. This decision trades off logistical expediency of unrestricted translocation with the risk of reducing translocation success due to environmental mismatch between origin and translocation site conditions. In this study, we used a large dataset of 502 individually identifiable carcasses to examine the role of geographic origin and translocation distance in the relative survival of 2822 translocated subadult and adult gopher tortoises (Gopherus polyphemus), a species experiencing large‐scale mitigation translocation, at a recipient site in the Florida panhandle, USA. We hypothesized that if climate or habitat differences between the origin and translocation site influenced survival, tortoises translocated from within the Florida panhandle would have the highest survival. To the contrary, we found that survival slightly increased with increasing climatic difference between origin and recipient site, driven by higher survival of tortoises coming from central Florida sites compared to those from the panhandle and north Florida. This suggests that environmental mismatch due to long‐distance translocation is not a main driver of mortality. These models also indicated an effect of season, with a survival advantage to tortoises translocated in the spring and late fall, relative to summer translocations, and a negative effect of initial density on survival. Finally, we also estimated the upper bound on annual survival in three well‐monitored groups to be quite low (92–95%) for several years following release, suggesting caution when considering large translocated populations to be viable without first assessing adult survival. Our unexpected results highlight the importance of investigating species‐specific sensitivities to translocation distances and indicate the limitations of assumed linear effects of translocation distance on outcomes.

Survival and behavior of Mojave desert tortoises head‐started with and without outdoor rearing

AbstractMojave desert tortoise (Gopherus agassizii) populations in some regions have declined by >50% since 2004, prompting the need for more research on ways to recover populations. One possible recovery tool is head‐starting (i.e., the act of protecting and raising juvenile tortoises to sizes that increase survival upon release); however, head‐starting can have high start‐up and maintenance costs that can limit its feasibility. Strategies that reduce cost and rearing duration may foster broader and more effective use. We released and radio‐tracked 60 juvenile tortoises in the Mojave National Preserve in California, USA, that had been reared under 2 treatments: those reared 1 year indoors after hatching, then 1 year outdoors (combo) and those reared just 1 year indoors (indoor‐only). We tested whether indoor‐only rearing alone could be a more efficient means of producing robust head‐started tortoises. We examined the behavior, movement, and survival of tortoises after release into the wild from 2020 to 2021 to determine whether these outcomes differed between husbandry treatments. Combo tortoises tended to perform settling behaviors (mean ± 1 SE days to building first burrow = 6.7 ± 0.8, entering dormancy = 23.3 ± 2.1, and emerging from dormancy = 189.6 ± 4.4) earlier than indoor‐only tortoises (7.4 ± 0.9, 31.5 ± 2.6, and 193.9 ± 5.9, respectively), but this difference was not significant, suggesting the rearing method did not greatly alter settling behavior. Indoor‐only tortoises dispersed at least twice as far from their release site (156.2 ± 26.3 m compared with 77.3 ± 20.6 m for combo tortoises), had larger mean use areas (3.7 ± 0.1 ha compared with 2.8 ± 0.3 ha for combo tortoises for 95% Brownian bridge movement model estimates), and greater variability in their movements than combo tortoises (daily average step length post‐emergence: 4.3 ± 0.2 m compared with 2.8 ± 0.1 m for combo tortoises). Despite differences in their movements, indoor‐only and combo tortoises had similar survival rates over the study, 51% versus 42%, respectively, during a period of extreme drought in 2021. The similarity in survival between groups gives head‐starting practitioners freedom in their rearing methods. The indoor‐only group had lower site fidelity, which should be considered when this is an undesirable trait for released tortoises.

Addressing the raven food subsidy challenge by engaging restaurants to close their dumpsters.

Ravens have benefitted from resource subsidies provided by humans so much that their population has increased by over 800% in the western Mojave desert over the last 50 years. Our food waste is an especially large subsidy. Raven predation is one of the greatest threats to desert tortoise survival in the California desert. We sought to create and evaluate a behavioral change program among restaurants in Yucca Valley and Twentynine Palms, California in the western Mojave with elevated raven populations. Half of the 60 restaurants in these two communities received an intervention encouraging them to close their dumpsters while the other half served as controls with no intervention. Treatment restaurants received two in-person visits to discuss the manifold importance of dumpster closure and laminated information signs to display and extend communication to staff. We surveyed all dumpsters for open/closed status five times before the intervention and five times again 3 months after the intervention. We found a significant increase in closure rates among treatment restaurants due to the interventions, with an average of 9.5% increase toward maximum possible lid closure. Restaurants achieving 80%+ closure received "Gold Star Awards" to further stimulate community-wide behavior change by publicly recognizing and encouraging desired behaviors. We summarize dumpster closure rates from similar interventions in nearby Joshua Tree and across the Coachella Valley to illustrate how local social norms may influence behavior.

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.

Waif Gopher Tortoise Survival and Site Fidelity Following Translocation

ABSTRACTGopher 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 400 m of their release location. These results suggest that waif tortoises could be an important resource in reducing the extirpation risk of isolated populations. © 2021 The Wildlife Society.

Integrating Gene Transcription-Based Biomarkers to Understand Desert Tortoise and Ecosystem Health.

Tortoises are susceptible to a wide variety of environmental stressors, and the influence of human disturbances on health and survival of tortoises is difficult to detect. As an addition to current diagnostic methods for desert tortoises, we have developed the first leukocyte gene transcription biomarker panel for the desert tortoise (Gopherus agassizii), enhancing the ability to identify specific environmental conditions potentially linked to declining animal health. Blood leukocyte transcript profiles have the potential to identify physiologically stressed animals in lieu of clinical signs. For desert tortoises, the gene transcript profile included a combination of immune or detoxification response genes with the potential to be modified by biological or physical injury and consequently provide information on the type and magnitude of stressors present in the animal's habitat. Blood from 64 wild adult tortoises at three sites in Clark County, NV, and San Bernardino, CA, and from 19 captive tortoises in Clark County, NV, was collected and evaluated for genes indicative of physiological status. Statistical analysis using a priori groupings indicated significant differences among groups for several genes, while multidimensional scaling and cluster analyses of transcription C T values indicated strong differentiation of a large cluster and multiple outlying individual tortoises or small clusters in multidimensional space. These analyses highlight the effectiveness of the gene panel at detecting environmental perturbations as well as providing guidance in determining the health of the desert tortoise.

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.

Effects of predator exclusion on nest and hatchling survival in the gopher tortoise

AbstractGopher tortoise (Gopherus polyphemus) populations are declining throughout the Southeast, and high levels of predation on nests and juveniles have been suggested as a potential contributor to this decline. Therefore, we documented gopher tortoise nest success and hatchling survival relative to mammalian predator control. We used 4, large (approx. 40‐ha) fenced, predator exclosures to exclude mid‐sized mammalian predators: bobcat (Lynx rufus), raccoon (Procyon lotor), Virginia opossum (Didelphis virginianus), fox (Urocyon cinereoargenteusandVulpes vulpes), coyote (Canis latrans), nine‐banded armadillo (Dasypus novemcinctus), and skunk (Mephitis mephitis); 4 unfenced plots served as controls. We monitored nests for survival through hatching and used radio‐telemetry to examine hatchling survival. We radio‐tracked 40 hatchlings for up to 329 days, but we were only able to track 8 individuals from a single nest at an unfenced plot because of high nest predation. Mean nest survival was greater at exclosures than at unfenced controls (F1, 2 = 45.80,P = 0.0001). Hatchling survival differed (χ2 = 5.839,P = 0.016) between unfenced plots (37.5%) and exclosures (74.4%), suggesting that mammals also were significant predators of hatchlings. The number of juvenile (<13 cm in diameter) and subadult tortoise burrows (13–21.9 cm) increased over a 6‐year period in exclosures, providing further support for an effect of excluding mammalian predators on nest and juvenile tortoise survival. © 2012 The Wildlife Society.

Long-Term Post-Fire Effects on Spatial Ecology and Reproductive Output of Female Agassiz’s Desert Tortoises (Gopherus agassizii) at a Wind Energy Facility near Palm Springs, California, USA

We studied the long-term response of a cohort of eight female Agassiz’s desert tortoises (Gopherus agassizii) during the first 15 years following a large fire at a wind energy generation facility near Palm Springs, California, USA. The fire burned a significant portion of the study site in 1995. Tortoise activity areas were mapped using minimum convex polygons for a proximate post-fire interval from 1997 to 2000, and a long-term post-fire interval from 2009 to 2010. In addition, we measured the annual reproductive output of eggs each year and monitored the body condition of tortoises over time. One adult female tortoise was killed by the fire and five tortoises bore exposure scars that were not fatal. Despite predictions that tortoises would make the short-distance movements from burned to nearby unburned habitats, most activity areas and their centroids remained in burned areas for the duration of the study. The percentage of activity area burned did not differ significantly between the two monitoring periods. Annual reproductive output and measures of body condition remained statistically similar throughout the monitoring period. Despite changes in plant composition, conditions at this site appeared to be suitable for survival of tortoises following a major fire. High productivity at the site may have buffered tortoises from the adverse impacts of fire if they were not killed outright. Tortoise populations at less productive desert sites may not have adequate resources to sustain normal activity areas, reproductive output, and body conditions following fire.

Spatial heterogeneity in the effects of climate change on the population dynamics of a Mediterranean tortoise

AbstractClimatic shifts may increase the extinction risk of populations, especially when they are already suffering from other anthropogenic impacts. Our ability to predict the consequences of climate change on endangered species is limited by our scarce knowledge of the effects of climate variability on the population dynamics of most organisms and by the uncertainty of climate projections, which depend strongly on the region of the earth being considered. In this study, we analysed a long‐term monitoring programme (1988–2009) of Hermann's tortoise (Testudo hermanni) aimed at evaluating the consequences of the drastic changes in temperature and precipitation patterns predicted for the Mediterranean region on the demography of a long‐lived species with low dispersal capability and already suffering a large number of threats. Capture–recapture modelling of a population in the Ebro Delta (NE Spain) allowed us to assess the effect of climate variability on the survival of tortoises. Winter rainfall was found to be the major driver of juvenile and immature survival, whereas that of adults remained high and constant across the study. Furthermore, local climate series obtained ad hoc from regional climate simulations, for this and 10 additional Mediterranean locations where tortoises occurred, provided us with reliable future climate forecasts, which were used to simulate the fate of these populations under three precipitation scenarios (mean, wet and dry) using stochastic population modelling. We show that a shift to a more arid climate would have negative consequences for population persistence, enhancing juvenile mortality and increasing quasiextinction risk because of a decrease in recruitment. These processes varied depending on the population and the climate scenario we considered, but our results suggest that unless other human‐induced causes of mortality are suppressed (e.g. poaching, fire, habitat fragmentation), climate variability will increase extinction risk within most of the species’ current range.

Upper respiratory tract disease, force of infection, and effects on survival of gopher tortoises

Upper respiratory tract disease (URTD) caused by Mycoplasma agassizii has been hypothesized to contribute to the decline of some wild populations of gopher tortoises (Gopherus polyphemus). However, the force of infection (FOI) and the effect of URTD on survival in free-ranging tortoise populations remain unknown. Using four years (2003-2006) of mark-recapture and epidemiological data collected from 10 populations of gopher tortoises in central Florida, USA, we estimated the FOI (probability per year of a susceptible tortoise becoming infected) and the effect of URTD (i.e., seropositivity to M. agassizii) on apparent survival rates. Sites with high (> or = 25%) seroprevalence had substantially higher FOI (0.22 +/- 0.03; mean +/- SE) than low (< 25%) seroprevalence sites (0.04 +/- 0.01). Our results provide the first quantitative evidence that the rate of transmission of M. agassizii is directly related to the seroprevalence of the population. Seropositive tortoises had higher apparent survival (0.99 +/- 0.0001) than seronegatives (0.88 +/- 0.03), possibly because seropositive tortoises represent individuals that survived the initial infection, developed chronic disease, and experienced lower mortality during the four-year span of our study. However, two lines of evidence suggested possible effects of mycoplasmal URTD on tortoise survival. First, one plausible model suggested that susceptible (seronegative) tortoises in high seroprevalence sites had lower apparent survival rates than did susceptible tortoises in low seroprevalence sites, indicating a possible acute effect of infection. Second, the number of dead tortoise remains detected during annual site surveys increased significantly with increasing site seroprevalence, from approximately 1 to approximately 5 shell remains per 100 individuals. If (as our results suggest) URTD in fact reduces adult survival, it could adversely influence the population dynamics and persistence of this late- maturing, long-lived species.

Assessing the efficacy of reintroduction programmes by modelling adult survival: the example of Hermann's tortoise

AbstractAlthough reintroduction programmes are often implemented for recovering animal populations, projects seldom monitor the long‐term survival of released animals. In addition, although many releases may occur in the same area, little is known about how the survival of successive release batches is affected by the presence of conspecifics and density dependence. Here, we use multi‐state capture–recapture modelling (combining information from recaptures and recoveries) to analyse the survival of two batches of Hermann's tortoise Testudo hermanni hermanni released in a 10‐year interval and monitored for 18 years at the Ebro Delta (western Mediterranean). We also tested whether the released animals experienced lower survival (i.e. a release cost) before becoming familiar with the new environment. Although we used a hard‐release method, neither group experienced a short‐term release cost. Annual survival of both groups differed and was not negatively affected by density‐dependent factors. Annual survivorship of the first group of released tortoises was constant and very high (0.945, se=0.011), and similar to that estimated from several natural populations. The presence of a terrestrial predator in 1 year (before the release of the second group) significantly decreased the survival of tortoises (0.819, se=0.073). Strikingly, survival of the second batch was significantly lower than that of the first group after the first years of release (0.775, se=0.049). Although survivorship for the first group suggested that habitat quality was high, the second group seemed not to acclimate well to the new environment, possibly due to the presence of resident tortoises. From a management perspective, reintroduction programmes of the Hermann's tortoise are a successful strategy for its recovery. Nevertheless, it seems advisable to avoid: (1) the release of tortoises at the core of well‐established populations and (2) areas with a high density of predators, which can jeopardize the reintroduction success, especially when the number of released individuals is small.

Consequences of violating the recapture duration assumption of mark–recapture models: a test using simulated and empirical data from an endangered tortoise population

Summary Mark–recapture methods are frequently used to estimate survival rates, which inform wildlife management policies. Mark–recapture models assume zero mortality of marked animals during the period of recapture and therefore that this period is of negligible duration relative to the interval between recapture events. This ‘recapture duration assumption’ is frequently violated yet is rarely tested for its potential to bias survival estimates. We investigated whether violating the recapture duration assumption increased precision at the cost of increased bias in estimates of survival. Using both simulation and empirical approaches, annual survival and recapture rates were estimated under recapture period durations that either conformed to, or violated, the recapture duration assumption. Empirical data were derived from a population of endangered Madagascar ploughshare tortoises Geochelone yniphora. Estimates of ploughshare tortoise survival were higher and had smaller standard errors when the recapture duration assumption was violated. As the ‘true’ ploughshare tortoise survivorship was unknown, a simulation approach was used to assess whether short or long recapture periods gave the most reliable estimates. The simulation study showed that violating the recapture duration assumption increased precision but did not increase bias in parameter estimates, even if up to 50% of the annual mortality occurred during the recapture period. The most precise and least biased survival estimates were obtained when sample size (number of marked individuals) was greatest, i.e. when both survival and recapture rates were high. Bias in survival estimates was negligible unless the recapture rate during the recapture period was &lt; 0·2. We conclude that the most precise ploughshare tortoise parameter estimates were generated by using lengthy recapture occasions that violated the recapture duration assumption. Synthesis and applications. To achieve the most precise and least biased survival estimates we recommend violating the recapture duration assumption and using a recapture period that maximizes sample size and achieves a recapture rate of &gt; 0·2 during the recapture period. If both survival and recapture rates are relatively constant during the recapture period, survival estimates will then have negligible bias. If a recapture rate of &gt; 0·2 during the recapture period cannot be achieved, methods other than mark–recapture should be used to quantify demographics. These recommendations apply to any species with an annual survival rate &gt; 0·05.

Survival during Early Life Stages of the Desert Tortoise (Gopherus agassizii) in the South-Central Mojave Desert

Early age classes of the Desert Tortoise (Gopherus agassizii) are particularly vulnerable to predation by several mammal and bird species. We studied tortoise survival at the Sand Hill Training Area of the Marine Corps Air Ground Combat Center, Twentynine Palms, California, from 1998 to 1999. We radiographed and thread-spooled 25 females to determine reproductive and nesting ecology. Predators consumed 11 of 42 monitored nests during the first 70 days of incubation. Ninety-one of 132 eggs in nests fenced after 70 days developed into healthy neonates. We recorded 0.84 and 0.91 neonate survival probability during dispersal in 1998 and 1999, respectively. Applying neonatal survival to egg success, we predict that 40% of individuals survived from oviposition to hibernation. Our findings support other chelonian life-history models indicating relatively high mortality early in life. Our data also suggest that neonatal Desert Tortoises are less susceptible to predation than was previously thought, perhaps because of their cryptic coloration and secretive habits. The Common Raven (Corvus corax) was not found to be a source of neonate mortality during this study.

Desert Tortoise (Gopherus agassizii) Survival at Two Eastern Mojave Desert Sites: Death by Short-Term Drought?

Survival of adult Desert Tortoises (Gopherus agassizii) appears related to site-specific variation in precipitation and productivity of annual plants. We studied adult tortoise survival rates at two closely situated, but physiographically different, sites in the eastern Mojave Desert over a nine-year period (spring 1992 to spring 2001). Survival rates were initially derived from population surveys conducted over a three-year period and by radio-telemetry monitoring over a seven-year period beginning in 1994. After a period of initial stability, survival rates on the two sites diverged over the study period, and seven-year survival rates estimated from radio-telemetry monitoring were 0.900 and 0.269, respectively. A die-off in 1996 on the latter site appears to have been triggered by a period of drought, which began in the summer of 1995, coupled with a failure of annual vegetation production in 1996. Depressed survival rates on this site were associated with drought conditions during three of four years. Although the decline had the appearance of an epizootic, there were no clinical signs of disease. Relatively short-term drought, combined with little or no annual biomass, appears to have caused severe reductions in tortoise survival. If periods of drought-induced low survival are common over relatively small areas, then source-sink population dynamics may be an important factor determining tortoise population densities.