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Brian Folt

Publications and source records attributed to Brian Folt.

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Strengths and opportunities in gopher tortoise population modeling: Reply to Loope et al.

We read the letter written by Loope et al. (2024) that evaluated the predictive population model for gopher tortoises from our previous paper ( Folt et al., 2022 ), and, despite criticism raised, we contend that our effort remains a valuable contribution. In our previous paper, we performed an unprecedented and exhaustive review of gopher tortoise population demography, which synthesized a large volume of literature about the current state of knowledge on gopher tortoise population dynamics and factors that influence demographic rates, including anthropogenic threats ( Folt et al., 2022 ). We then built a predictive population model that examined the relative influence of multiple anthropogenic stressors on tortoise populations into the future, while accounting for important geographic variation ( Folt et al., 2022 ).

Global Ecology and Conservation

Using predictions from multiple anthropogenic threats to estimate future population persistence of an imperiled species

Imperiled species face numerous and diverse anthropogenic threats to their persistence, and wildlife managers charged with making conservation decisions benefit from a sound understanding of how populations, species, and ecosystems will respond to future changes in threats to biodiversity. In southeastern North America, the gopher tortoise ( Gopherus polyphemus ) is a keystone species in upland ecosystems; however, tortoise populations have declined strongly over the last century, and the species is a candidate for increased protection by the United States federal government under the Endangered Species Act (ESA). Here, we sought to support conservation decision making for G. polyphemus by developing a spatially-explicit predictive population model that linked four anthropogenic threats (climate warming, sea-level rise, urbanization, habitat degradation) to demographic vital rates and used the model to estimate future changes in the number of individuals, populations, and metapopulations across the species’ range. Using recent survey data, we projected 457 populations for 80 years into the future under scenarios varying in threat magnitude, management magnitude, and demographic uncertainty. Population projections predicted that the number of individuals, populations, and metapopulations would decline among all simulated scenarios in the next 80 years. Model predictions were more sensitive to variation in adult survival and immigration rates than to variation in threat magnitude. A scenario with decreased habitat management and threat effects from climate warming, sea-level rise, and urbanization predicted geographic variation in persistence probabilities for populations that might result in decreased genetic representation across the species' range. Our results can be used to support conservation listing decisions for the gopher tortoise as part of its federal Species Status Assessment and provide an analytical framework for how to link diverse threats to geographically-varying demographic rates during population viability analyses for wide-ranging imperiled species around the world.

Global Ecology and Conservation

Decision context as an essential component of population viability analysis

Population viability analysis (PVA) is a widely used tool that applies demographic data in simulation frameworks to assess extinction risk for species or populations. It is used in diverse conservation applications, including evaluating management effectiveness, relative risk of threats, and potential changes to protective status (Beissinger & McCullough, 2002 ), and can be a critical tool for making decisions with imperfect knowledge of the system state, often on limited timelines (Meine et al., 2006 ). Chaudhary and Oli ( 2020 ) recently developed a framework to appraise the quality of PVAs based on the presence of essential background, model, and analysis components. They evaluated 160 published PVAs and reported a decline in the quality of PVAs over time (1990−2017). We agree PVA studies should report unambiguous descriptions of their essential components (Table 1 in Chaudhary and Oli) and explicitly state the model's biological and statistical assumptions. The need for increased transparency in PVAs is evident. Morrison et al. ( 2016 ) reported that only 50% of PVAs published in peer-reviewed and gray literature were both reproducible and repeatable. Further, in an examination of 67 studies that used matrix population models (widely used in PVAs), Kendall et al. ( 2019 ) reported that models frequently contained misspecification errors. Given the rapid advancement of simulation techniques, updated guidance for PVA construction is warranted. However, we believe the essential PVA components identified by Chaudhary and Oli contain a critical omission: the decision context in which the PVA was created and its usefulness in that context. Quality and utility are not mutually exclusive; however, some models that do not meet idealized quality standards might still be valuable because they are useful and represent the best available science for a given decision context (hereafter, decision-support models). The definition of quality for decision-support models should be different than models developed for the purpose of learning (hereafter, heuristic models) and should incorporate how useful the model was, despite information gaps. We further argue that assessment questions should be used prospectively to guide modeling projects, rather than for retrospective comparison of model quality.

Conservation Biology

Contrasting patterns of demography and population viability among gopher tortoise (Gopherus polyphemus) populations at the species’ northern range edge

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 mark-recapture 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. © 2021 The Wildlife Society.

Alabama

Modeling strategies and evaluating success during repatriations of elusive and endangered species

Wildlife repatriation is an important tool to decrease extinction risk for imperiled species, but successful repatriations require significant time, resources and planning. Because repatriations can be long and expensive processes, clear release strategies and monitoring programs are essential to efficiently use resources and evaluate success. However, monitoring can be challenging and surrounded by significant uncertainty, particularly for secretive species with extremely low detection probability. Here, we simulated how alternative repatriation strategies influence repatriation success for the eastern indigo snake Drymarchon couperi , a federally-Threatened species that is currently being repatriated in Alabama and Florida. Critically, we demonstrate how observed population growth can differ from true population growth when detection probabilities are low and mark-recapture analyses are not an option. Specifically, we built a stochastic stage-based population model to predict population growth and extinction risk under different release strategies and use information from ongoing repatriations to predict success and guide future releases. Because D. couperi is difficult to monitor, we modeled how detection probability influenced perceptions of abundance and population growth by monitoring programs. Simulated repatriation strategies releasing older, head-started snakes in greater abundance and frequency created wild populations with decreased extinction risk relative to scenarios releasing fewer and younger snakes less frequently. Ongoing repatriations currently have a 0.23 (Alabama) and 0.61 (Florida) probability of quasi-extinction, but extinction risk decreased to 0.07 and 0.10 at sites upon achieving the targeted number of releases. Abundances observed under realistic detection thresholds for D. couperi did not always predict true population growth; specifically, we demonstrate that monitoring programs during repatriations of secretive species may indicate that efforts have been unsuccessful when populations are actually growing. Overall, our modeling framework informs release strategies to maximize repatriation success while demonstrating the need to consider how detection processes influence assessment of success during conservation interventions.

Florida

Interdrainage morphological and genetic differentiation in the Escambia Map Turtle, Graptemys ernsti

Graptemys ernsti, the Escambia Map Turtle, inhabits the Escambia/Conecuh River, the adjacent Yellow River, and the Pea River further to the east, all of which have been distinct drainage systems since the Pleistocene. We used continuous and meristic morphological and genetic data to compare populations of G. ernsti and found evidence of differences among the three drainages. Frequency of occurrence of a nasal trident differed among the three drainages. Yellow River specimens possessed unique mitochondrial haplotypes while the Conecuh and the Pea shared haplotypes. Five microsatellite loci identified the drainages as being distinct, with the strongest differentiation between the Yellow River and the other two drainages. While these differences do not appear great enough to warrant taxonomic recognition, they do suggest that each population has a distinct evolutionary and demographic history and that they should therefore be managed separately.

Herpetological Conservation and Biology

Hybridization of two megacephalic map turtles (testudines: emydidae: Graptemys ) in the Choctawhatchee River drainage of Alabama and Florida

Map turtles of the genus Graptemys are highly aquatic and rarely undergo terrestrial movements, and limited dispersal among drainages has been hypothesized to drive drainage-specific endemism and high species richness of this group in the southeastern United States. Until recently, two members of the megacephalic “ pulchra clade,” Graptemys barbouri and Graptemys ernsti , were presumed to be allopatric with a gap in both species' ranges in the Choctawhatchee River drainage. In this paper, we analyzed variation in morphology (head and shell patterns) and genetics (mitochondrial DNA and microsatellite loci) from G. barbouri , G. ernsti , and Graptemys sp. collected from the Choctawhatchee River drainage, and we document the syntopic occurrence of those species and back-crossed individuals of mixed ancestry in the Choctawhatchee River drainage. Our results provide a first counter-example to the pattern of drainage-specific endemism in megacephalic Graptemys . Geologic events associated with Pliocene and Pleistocene sea level fluctuations and the existence of paleo-river systems appear to have allowed the invasion of the Choctawhatchee system by these species, and the subsequent introgression likely predates any potential human-mediated introduction.

Alabama, Florida