Fire Management Effects on Long-Term Gopher Tortoise Population Dynamics

Long-term datasets are required to understand the response of long-lived organisms (e.g., gopher tortoises [Gopherus polyphemus]) to management actions, such as prescribed burns. Our objective was to estimate the effects of prescribed burning on gopher tortoise population dynamics over decadal time frames at Fort Stewart Army Reserve, southeastern Georgia, USA. We captured and marked adult tortoises from 1994–2020. In addition, since the early 1990s, managers at Fort Stewart collected spatial records of prescribed burns; thus, we could compare demography of the population to prescribed burning. We used a Bayesian hierarchical model (open population Jolly-Seber model) to estimate population parameters (emigration and survival, immigration and recruitment, and adult abundance) and their relationships with years since burn. We observed opposing responses to years since burn at 2 sites: abundance and the probability of staying (survival plus not emigrating) increased within 1 site when it had been more recently burned (F zones), but abundance and probability of staying in a second site increased when it had been longer since the site was burned (E zones). Some of these effects were weak but indicative of different responses to burning between the sites. Although the sites experienced similar burning regimes, they differed substantially in other habitat features: the F zones had almost twice the tree cover and lower soil sand composition, indicating that tortoise population responses to burning depend on habitat context. We inferred that the primary mechanism for demographic responses to years since burn was likely emigrating adults, which indicates the need for more detailed movement data. Our results demonstrate that gopher tortoise population responses to prescribed burning are complex, context dependent, and primarily influenced by tortoise movements. Therefore, prescribed burn plans may best accommodate spatially dynamic tortoise populations when they create spatial heterogeneity in burn ages within the range of typical tortoise movements. © 2021 The Wildlife Society.     

Gopher Tortoise Demographic Responses to a Novel Disturbance Regime

The long-term viability of gopher tortoise (Gopherus polyphemus) populations is jeopardized by increased urbanization and habitat degradation owing to fire suppression. Because the species' remaining natural habitats in the southeastern United States exist within a mosaic of anthropogenic land uses, it is important to understand demographic responses to contrasting land uses and habitat management regimes. We examined differences in demographic parameters among fire-suppressed sandhill, restored sandhill, and former sandhill (i.e., ruderal) land use-land cover (LULC) types at Archbold Biological Station in south-central Florida, USA. Using Program MARK, we estimated population size, and sex-specific and LULC-specific survivorship based on 6 years of mark-recapture data. We also analyzed individual growth trajectories and clutch sizes to determine whether growth rates or reproductive output differed among LULC types. Tortoises in an open, ruderal field occurred at a higher density (7.79/ha) than in adjacent restored (1.43/ha) or fire-suppressed (0.40/ha) sandhill. Despite this higher density, both adult survivorship and body size were significantly higher in the ruderal field. Furthermore, the larger female body size in the ruderal field likely contributed to increased annual survivorship and slightly larger average clutch sizes. We did not detect offsetting negative demographic effects; in particular, we did not find significant biological or statistical differences in body condition, asymptotic body size, or growth rate among the 3 LULC types. Our results suggest that anthropogenic, grass-dominated land-cover types may be important components of the habitat mosaic currently available to this at-risk species. © 2019 The Wildlife Society.     

Waif Gopher Tortoise Survival and Site Fidelity Following Translocation

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 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.     

Population Genetic Assignment of Confiscated Gopher Tortoises

Abstract As gopher tortoises (Gopherus polyphemus) increasingly become threatened throughout their range in Florida, USA, the need for management and conservation will intensify. Here we evaluate the forensic applicability of genetic assignment tests based on microsatellite genotypic data to 1) accurately assign individuals in our genetic database to the sample location or population of origin and 2) determine the origin of 6 confiscated tortoises. Overall, we could correctly assign 90% of the individuals in the database to their population of origin, but we were unable to determine the source of the confiscated tortoises. However, these individuals are unlikely to have come from any of our sampled sites and all 6 may have come from the same population. This approach can be used by law enforcement personnel to identify the origin of confiscated tortoises as well as by developers and wildlife managers to determine the genetic appropriateness of potential recipient populations when it is necessary to relocate individuals.     

Long-Term Retention of a Relocated Population of Gopher Tortoises

Abstract: Relocations of gopher tortoises (Gopherus polyphemus) in Florida, USA, are frequently employed as mitigation tools when tortoises occupy land desired for development. Here we present information about retention and health of a relocated population of gopher tortoises 17 years after relocation. We combine our 17-year postrelocation data with earlier surveys 1 year and 2 years postrelocation to examine whether retention rates change over time. We also evaluate whether retention rates vary by age and gender. Of 74 gopher tortoises relocated in 1985, 31 were present in 2002. We found a 1-year retention rate of 42%, with retention rates of 100% each year thereafter, when we used the percentage of relocated individuals captured at each survey. We found a 1-year retention rate of 73%, a retention rate of 92% from year 1 to year 2, and an overall retention rate of 98.5% from year 2 through year 17, based on the assumption that all individuals present in later surveys were present in earlier surveys. We found no significant difference in retention rate over the 17-year period for adults and juveniles and for adult males and females. Relocated gopher tortoises showed natural growth patterns, indicating good health, but 35% of these gopher tortoises had ≥1 symptom of upper respiratory tract disease, a disease associated elsewhere with population declines of tortoises. Thus, retention rates of relocated gopher tortoises change over time, with relatively low retention during the first year postrelocation but nearly 100% retention in subsequent years. In general, our study shows that relocations can successfully lead to long-term retention of gopher tortoises, but we predict that without management (e.g., fire management and predator control) this relocated population is not viable.     

Northern Bobwhite Non-Breeding Habitat Selection in a Longleaf Pine Woodland

Efforts to halt the decline of the northern bobwhite (Colinus virginianus; bobwhite) across its distribution have had limited success. Understanding bobwhite habitat requirements across the annual cycle and at varying scales is essential to aid efforts to conserve bobwhites. We monitored radio-tagged bobwhites from 2016 to 2018 on a 165-km² portion of Fort Bragg Military Installation in the Sandhills physiographic region of North Carolina, USA, to determine factors influencing non-breeding bobwhite habitat selection at multiple scales. We used generalized linear models (GLM) and generalized linear mixed models to assess bobwhite habitat selection at the microsite scale (the immediate vicinity of an animal) and the macrosite scale (across the study area), respectively, by comparing used points to available random points. At the microsite scale, bobwhites strongly selected areas with greater woody understory cover. Also, bobwhite selection increased with greater forb and switchcane (Arundinaria tecta) cover, but this effect plateaued at 65% forb cover and 50% switchcane cover. At the macrosite scale, bobwhites generally selected areas with greater understory cover within a 200-m radius but avoided areas with >55% understory cover; these areas primarily were located in the core areas of drainages with extensive ericaceous vegetation. Bobwhites selected areas with 3–6 m²/ha hardwood basal area in uplands, potentially because of the availability of mast, but avoided uplands when pine (Pinus spp.) or hardwood basal area exceeded 20 m²/ha or 12 m²/ha, respectively, likely because high basal area is associated with increased shading and subsequent loss of understory cover. In addition, bobwhites selected uplands 1 growing season (≥2-month period falling entirely between 1 Apr and 1 Oct) post-fire regardless of burn season. Overall, managers seeking to improve habitat quality for bobwhites in longleaf pine (Pinus palustris) woodlands should employ management practices that maintain available woody understory across the landscape to provide cover during the non-breeding season. © 2020 The Wildlife Society.     

Canopy Closure and Emigration by Juvenile Gopher Frogs

ABSTRACT Although studies have addressed effects of abrupt transitions in habitat type (e.g., forest-clear-cut or forest-field edges) on amphibian movements, little is known about effects of more subtle habitat transitions on patterns of migration and habitat use in amphibians. We used radiotelemetry to study movement patterns of juvenile gopher frogs (Rana capito) emigrating from ponds that were surrounded by longleaf pine (Pinus palustris) forest that varied in structure as a result of fire suppression. Our primary purpose was to determine if frogs emigrate directionally from their natal ponds and select habitat at random during their first month following metamorphosis. We found that frogs emigrated in nonrandom directions from ponds that were surrounded by heterogeneous habitat and selected fire-maintained habitat that was associated with an open canopy, few hardwood trees, small amounts of leaf litter, and large amounts of wiregrass (Aristida beyrichiana). Fire-maintained habitat contained higher densities of burrows excavated by gopher tortoises (Gopherus polyphemus) and small mammals, which are the priamry refuge sites for both juvenile and adult gopher frogs. Frogs moved up to 691 m from their natal ponds, frequently crossed dirt roads, and even seemed to use these roads as migration corridors. To maintain suitable terrestrial habitat for gopher frogs, including habitat used by migrating individuals, it is important to apply frequent prescribed fire to uplands surrounding breeding ponds that lead all the way to the edges of breeding ponds, as well as through ponds during periodic droughts.     

Effects of longleaf pine planting density and other factors on stand structure and associated wildlife habitat

The primary objective of many longleaf pine (Pinus palustris) restoration programs is to enhance or restore habitat for wildlife dependent on herbaceous plant communities. Because herbaceous cover is inversely related to canopy cover, restoration programs often place restrictions on longleaf pine planting density. However, the influence of planting density on understory plant communities has been inadequately evaluated. Therefore, we initiated a study to examine the relative influences of planting density and other factors on overall understory composition and forage availability for white‐tailed deer (Odocoileus virginianus) and northern bobwhite (Colinus virginianus) in nine longleaf pine stands throughout the Coastal Plain of Alabama during 2017–2018. We found that coverage of herbaceous plants decreased 3.5%, coverage of woody plants decreased 2.4%, and coverage of northern bobwhite forage plants decreased 1.9% for each 1 m²/ha increase in longleaf pine basal area. However, planting density was not a significant predictor of current basal area, nor coverage of any functional group of plants we examined, likely because current longleaf pine density averaged only 46% (range = 30–64%) of seedling planting density. We did not detect an effect of prescribed fire on stand condition or understory plant communities, likely due to variability in fire timing and frequency. Our findings related to planting density were likely a function of low longleaf pine survival, which is not uncommon. Because of this and the inherent variability in growth rates for young longleaf pine stands, restoration programs should consider placing greater emphasis on post‐planting monitoring and management than planting density.     

Response of Six‐Lined Racerunner (Aspidoscelis sexlineata) to Habitat Restoration in Fire‐Suppressed Longleaf Pine (Pinus palustris) Sandhills

Six‐lined racerunner (Aspidoscelis sexlineata) is an indicator species of frequently burned Longleaf pine (Pinus palustris) forests. To evaluate how the species responded to forest restoration, we conducted a mark‐recapture study in formerly fire‐suppressed Longleaf pine forests exposed to prescribed fire or fire surrogates (i.e. mechanical or herbicide‐facilitated hardwood removal) as well as in fire‐suppressed control sites and reference sites, which represented the historic condition. After initial treatment, all sites were exposed to over a decade of prescribed burning with an average return interval of approximately 2 years. We used population‐level response of A. sexlineata as an indicator of the effectiveness of the different treatments in restoring habitat. Specifically, we compared mean numbers of marked adults and juveniles at treatment sites to that of reference sites. After 4 years, restoration objectives were met at sites treated with burning alone and at sites treated with mechanical removal of hardwoods followed by fire. After over 10 years of prescribed burning, restoration objectives were met at all treatments. We conclude that prescribed burning alone was sufficient to restore fire‐suppressed Longleaf pine sandhills for A. sexlineata populations.     

Effects of Chronic Human Activities on Invasion of Longleaf Pine Forests by Sand Pine

Chronic human activities may result in new and permanent successional trajectories in certain ecosystems. The invasion of longleaf pine ecosystems by sand pine in the Florida Panhandle is one such change in the landscape. This study examined the spatial pattern of sand pine expansion and explored the natural and anthropogenic disturbances that fostered this invasion. Aerial photographs (1949, 1994) and Geographic Information Systems analyses confirmed sand pine expansion at Eglin Air Force Base. In 1949, there were 8,982 ha of sand pine in the southern portion of the study area near riparian and coastal lowland forests. By 1994, sand pine had expanded further upland and inland, for a total of 17,147 ha in the study area. Sand pine age data showed that this expansion had started by 1920 but increased rapidly in the 1940s. Historical accounts and structural data from stands suggest that land-use activities associated with the extraction of turpentine promoted the invasion by sand pine. Fires were suppressed in longleaf pine forests to protect turpentine trees, resulting in increased vegetation cover and decreased regeneration of longleaf pine. In addition, stands were typically harvested after turpentining, and there was little or no advanced regeneration of longleaf pine. Sand pine age histograms showed that the onset of high establishment rates (1940s) coincided with changes in land ownership and widespread fire suppression. Sand pine is likely to persist in these ecosystems due to its abundant regeneration. Received 17 March 1999; accepted 28 January 2000.     

Effects of mesopredators and prescribed fire on hispid cotton rat survival and cause-specific mortality

Control of mid-sized mammalian predators (hereafter, mesopredators) is sometimes advocated in an attempt to reduce their impact on wildlife populations, particularly economically important (i.e., game) or endangered species. However, mesopredators may play a role in regulating small mammal populations; thus, lethal control of mesopredators may have unintended consequences. The hispid cotton rat (Sigmodon hispidus; hereafter, cotton rat) is one of the most common small mammals in the southeastern United States and is an important prey species for several of the region's predators. Within fire-maintained communities, such as the longleaf pine (Pinus palustris) forests of the Coastal Plain, cotton rat populations dramatically, yet temporarily, decline following prescribed fire. To evaluate the effects of mesopredator removal on cotton rat survival and cause-specific mortality, we conducted a large-scale mesopredator exclusion experiment that incorporated a prescribed fire during the winter of study. Between 18 May 2006 and 20 June 2007, we used radio-telemetry to monitor 252 cotton rats (131 in exclosures and 121 in controls) and documented 184 mortalities. During the 37-week period of monitoring prior to the prescribed fire event, weekly survival of cotton rats was greater in mesopredator exclusion plots. During the 19 weeks following the prescribed fire, there were no differences in weekly survival relative to mesopredator treatment, but fire caused a short-term reduction in weekly survival within both exclosures and controls. Of 36 cotton rats monitored on the date of prescribed fire, 18 were depredated within 1 month, 4 emigrated, and 5 were killed by the fire event. Overall, raptors preyed on cotton rats more in exclosures than in controls, but evidence for compensatory predation (raptor-caused morality greater in exclosures than in controls although survival rates were similar between treatments) only occurred following the prescribed fire event. Our results suggest that managing mesopredators may result in a temporary increase in cotton rat survival, but dormant season prescribed fire removes this effect until well into the following growing season. © 2011 The Wildlife Society.     

Fire in the Suburbs: Ecological Impacts of Prescribed Fire in Small Remnants of Longleaf Pine (Pinus palustris) Sandhill

Abstract Logging, fire suppression, and urbanization have all contributed to the serious decline and fragmentation of Pinus palustris (longleaf pine) ecosystems in the southeastern United States. Effective management of the remaining patches of these pyrogenic communities must incorporate periodic low‐intensity fires, even where they are located on private lands in populated urban and suburban areas. To explore the effects of fire and its potential use for restoration and management of small fragments surrounded by suburban development, we conducted growing season prescribed fires in remnant longleaf pine sandhill patches in the suburbs of Gainesville, Florida. Density and composition of hardwoods were surveyed pre‐burn and 1 and 9 months post‐burn. Woody stem density decreased in the burn plots, predominantly in the smaller size classes. Flowering responses of forbs and small shrubs were surveyed six times post‐burn for 1 year. Overall, the burns did not yield greater densities of flowering stems, but burn patches had higher species richness and diversity than control patches. In addition, there were consistently greater numbers of “showy flowered” sandhill species in flower in burn patches relative to controls. The results of this research demonstrate that prescribed fire can be used for restoration and management of small remnants of longleaf pine sandhill in suburban neighborhoods. It is also clear that although a single prescribed burn can be effective, it will take more than one burn to attain desired restoration goals in degraded longleaf remnants.     

Effects of Hardwood Reduction Techniques on Longleaf Pine Sandhill Vegetation in Northwest Florida

We tested whether the intensity of hardwood midstory reduction causes commensurate improvements of herbaceous groundcover in fire‐suppressed Pinus palustris (longleaf pine) sandhills. Using a complete randomized block design, we compared the effects of three hardwood reduction techniques (spring burning, application of the ULW^® form of the herbicide hexazinone, chainsaw felling/girdling) and a no‐treatment control on plant species richness, and on life form and common species densities at Eglin Air Force Base, Florida, U.S.A., from 1995 to 1998. ULW^® and felling/girdling plots were burned for fuel reduction two years after initial treatment application. We also sampled the same variables in frequently‐burned reference sandhills to establish targets for restoration. Spring burns achieved partial topkill of oaks (17.6–41.1% from 1995 to 1998) compared to reductions of 69.1–94% accomplished by ULW^® and of 93.2–67.8% by felling/girdling treatments. We predicted that plant species richness and densities of herbaceous groundcover life forms would increase according to the percent hardwood reductions. Predictions were not supported by treatment effects for species richness because positive responses to fire best explained increases in plant richness, whereas ULW^® effects accounted for the largest initial decreases. Legumes, non‐legume forbs, and graminoids did not respond to treatments as predicted by the hypothesis. Again, positive responses to fire dominated the results, which was supported by greater herbaceous densities observed in reference plots. Overall, we found that the least effective and least expensive hardwood midstory reduction method, fire, resulted in the greatest groundcover improvements as measured by species richness and herbaceous groundcover plant densities.     

Intraspecific trait variability shapes leaf trait response to altered fire regimes

Background and AimsUnderstanding impacts of altered disturbance regimes on community structure and function is a key goal for community ecology. Functional traits link species composition to ecosystem functioning. Changes in the distribution of functional traits at community scales in response to disturbance can be driven not only by shifts in species composition, but also by shifts in intraspecific trait values. Understanding the relative importance of these two processes has important implications for predicting community responses to altered disturbance regimes.MethodsWe experimentally manipulated fire return intervals in replicated blocks of a fire-adapted, longleaf pine (Pinus palustris) ecosystem in North Carolina, USA and measured specific leaf area (SLA), leaf dry matter content (LDMC) and compositional responses along a lowland to upland gradient over a 4 year period. Plots were burned between zero and four times. Using a trait-based approach, we simulate hypothetical scenarios which allow species presence, abundance or trait values to vary over time and compare these with observed traits to understand the relative contributions of each of these three processes to observed trait patterns at the study site. We addressed the following questions. (1) How do changes in the fire regime affect community composition, structure and community-level trait responses? (2) Are these effects consistent across a gradient of fire intensity? (3) What are the relative contributions of species turnover, changes in abundance and changes in intraspecific trait values to observed changes in community-weighted mean (CWM) traits in response to altered fire regime?Key ResultsWe found strong evidence that altered fire return interval impacted understorey plant communities. The number of fires a plot experienced significantly affected the magnitude of its compositional change and shifted the ecotone boundary separating shrub-dominated lowland areas from grass-dominated upland areas, with suppression sites (0 burns) experiencing an upland shift and annual burn sites a lowland shift. We found significant effects of burn regimes on the CWM of SLA, and that observed shifts in both SLA and LDMC were driven primarily by intraspecific changes in trait values.ConclusionsIn a fire-adapted ecosystem, increased fire frequency altered community composition and structure of the ecosystem through changes in the position of the shrub line. We also found that plant traits responded directionally to increased fire frequency, with SLA decreasing in response to fire frequency across the environmental gradient. For both SLA and LDMC, nearly all of the observed changes in CWM traits were driven by intraspecific variation.