Craig Guyer’s research while affiliated with Auburn University and other places

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Publications (160)


Figure 2. Bivariate plot of radiotelemetry fixes (x axis) and number of long-term captures (y axis) of gopher tortoises (Gopherus polyphemus) from a population in southern Alabama. Points are: females (open circles), males (filled circles), and an individual of unidentified sex (X). Dashed line separates individuals defined to be residents (above line) from those defined to be emigrants (below line). Presumed emigrants during year 2000 are indicated in red.
Figure 4. Annual home ranges of resident female (A) and male (B) gopher tortoises (Gopherus polyphemus) during 2000-2001 and lifetime home ranges of those same resident female (C) and male (D) tortoises during 1992-2020.
Figure 7. Regression of how local population density influences emigration rate for 6 sites described in Guyer et al. (2012). Dashed line represents ordinary least-squares regression line (y ¼ À0.20x þ 0.33; R 2 ¼ 0.91). Lower-density sites have higher emigration rates. Site abbreviations and sample sizes (total number of individuals transmittered) are Camp Shelby, MS (CS; 38), Conecuh National Forest, AL (CNF; 41), Dixon Center, AL (DC; 13), Mobile County, AL (MC; 40), Green Grove, AL (GG; 132), and Wade Tract, GA (WT; 66).
Annual and Lifetime Home Ranges Reveal Movement Patterns Within and Among Local Populations of Gopher Tortoises (Gopherus polyphemus)
  • Article
  • Full-text available

July 2024

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56 Reads

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1 Citation

Chelonian Conservation and Biology

Craig Guyer

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Jeffrey M. Goessling

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Download

Map of the study area and sampling locations. a The southeastern United States, with the study area indicated by the black rectangle. b Sampling locations (black dots) in southern Alabama, including Fred T. Stimpson State Game Sanctuary, Perdido Wildlife Management Area, Solon Dixon Forestry Education Center, Conecuh National Forest (administrative boundary in gray), Rayonier Tract, and Geneva State Forest. c Samples from six sites (colored dots) within Conecuh National Forest and Solon Dixon Forestry Education Center (Solon Dixon); opportunistic samples outside of these sites are indicated in red as ‘opportunistic samples’. Satellite map is provided by Google through the R package ggmap (Kahle and Wickham 2013)
Pairwise genetic distance by geographic distance for Gopher Tortoise (Gopherus polyphemus) samples (n = 336; 56,280 total comparisons). Pairwise genetic distance increases significantly with geographic distance (Mantel r = 0.55, P = 0.001). Regression line is shown for heuristic purposes
Cluster-based population assignment analyses for various subsets of Gopher Tortoise (Gopherus polyphemus) samples from sites in southern Alabama. Results from analyses are shown both as a map (top) and as a bar plot (bottom) for a all locations using non-spatial fastSTRUCTURE, b all locations using spatially explicit conStruct analyses. c all locations excluding Perdido and Stimpson using non-spatial fastSTRUCTURE and d all locations excluding Perdido and Stimpson using spatially explicit conStruct (d). Each pie chart or bar represents a single individual composed of admixture from differently colored inferred populations. Sites are: Conecuh National Forest (Conecuh), Fred T. Stimpson State Game Sanctuary (Stimpson), Geneva State Forest (Geneva), Perdido River Wildlife Management Area (Perdido), Rayonier Tract (Rayonier), and Solon Dixon Forestry Education Center (Solon Dixon). Samples from Conecuh National Forest were further divided into six sites (‘Sites 1–6’) and other opportunistic samples (‘Other’)
Principal components analyses (PCA) and spatial principal component analyses (sPCA) for all Gopher Tortoises (a and b, respectively), and all Gopher Tortoises excluding individuals from Perdido and Stimpson (c and d, respectively). Dots represent one individual colored by their sampling location. Conecuh in a and b includes Sites 1 through 6, and other as shown in plots ( c , d)
Using genomic data to estimate population structure of Gopher Tortoise (Gopherus polyphemus) populations in Southern Alabama

January 2024

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121 Reads

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2 Citations

Conservation Genetics

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Jeffrey M. Goessling

In the North American longleaf pine (Pinus palustris) ecosystem, the Gopher Tortoise (Gopherus polyphemus) is a keystone species that has declined significantly over the last century. Habitat degradation and fragmentation may have caused G. polyphemus to become separated into small, isolated local populations that suffer from decreased genetic diversity or inbreeding depression. Here we use genome-scale methods to sequence thousands of loci for 336 G. polyphemus individuals from 11 sites across southern Alabama to estimate population genetic structure and levels of genetic diversity. We found a pattern of isolation by distance among samples, where geographic distance predicted genetic difference. Principal components and structure analyses supported the existence of three weak genetic populations comprising individuals from (1) Fred T. Stimpson State Game Sanctuary and Perdido Wildlife Management Area, (2) Conecuh National Forest and Solon Dixon Forestry Education Center, and (3) Geneva State Forest Wildlife Management Area. We did not observe strong variation in genetic diversity or effective population size metrics among sampling locations or genetic populations identified by population structure analyses. Our results suggest that G. polyphemus historically operated on larger geographic scales than those considered by contemporary mark-recapture studies. Absence of variation in population genetic metrics suggests that either effects of fragmentation have not manifested themselves, or that the effects are similar across all locations. Given the common use of translocations in Gopher Tortoise management, we provide a framework for tortoise translocations based on our genomic data.


FIG. 3.-Growth curves based on the results of a von Bertalanffy growth model for Eastern Indigo Snake (Drymarchon couperi) SVL across wild (A), captive (B), and reintroduced (C) populations. Shaded regions equal 95% credible intervals, and curves were fit assuming a hatchling SVL of 40 cm based on data from captive individuals. The horizontal line represents the largest indigo snake recorded from wild populations in this study.
FIG. 4.-Distributions of daily change in body weight for headstarted Eastern Indigo Snakes (D. couperi) housed at four captive rearing facilities (Auburn, OCIC, WNFH, and Zoo Atlanta [ZooATL]). Values are unitless and represent the change in weight per day standardized by the initial weight during that growth increment.
Evaluating Growth Rates of Captive, Wild, and Reintroduced Populations of the Imperiled Eastern Indigo Snake (Drymarchon couperi)

December 2023

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310 Reads

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1 Citation

Herpetologica

Reintroduction of species at sites where populations have been extirpated has become a common technique in wildlife conservation. To track progress towards reintroduction success, effective postrelease monitoring is needed to document vital rates of individuals and the corresponding impact on population trajectories. We assessed growth and body size in Eastern Indigo Snakes (Drymarchon couperi) using a data set from multiple projects across the species' distribution, including free-ranging wild snakes, snakes reared in captive-breeding programs, and snakes released at two reintroduction sites. We used these data to fit a von Bertalanffy growth model in a Bayesian framework to quantify differences in growth among three broad categories of snakes (wild, captive, and reintroduced), while accounting for measurement error across various projects. We also compared changes in body mass of captive-born individuals from four captive rearing facilities. Asymptotic snout–vent length across all groups was 185 cm (95% credible interval = 177–194 cm) for males and 157 cm (95% credible interval = 153–161 cm) for females. Reintroduced snakes had a higher growth coefficient than either captive or wild snakes (e.g., captive females = 1.20 [1.06–1.35] d–1; wild females = 1.22 [0.95–1.49] d–1; reintroduced females = 1.62 [1.21–2.05] d–1), indicating that current captive-breeding and rearing efforts for indigo snakes produce similar or faster growth trends compared to wild populations. Furthermore, daily changes in juvenile body weight relative to body size were similar in three of the four captive rearing facilities (mean for females at Orianne Center for Indigo Conservation = 0.57 [0.48–0.65]; Zoo Atlanta = 0.55 [0.37–0.72]; Welaka National Fish Hatchery = 0.55, [0.36–0.73]; Auburn University = 0.39 [0.21–0.58]). Long-term project success for indigo snake reintroductions will depend on continuing to implement best practices in an adaptive management framework.



Distribution and Status of the Alligator Snapping Turtle (Macrochelys temminckii) in Alabama

June 2023

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152 Reads

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1 Citation

Southeastern Naturalist

Macrochelys temminckii (Alligator Snapping Turtle) has experienced historical harvesting pressures that impact a number of current populations. Filling distribution gaps and obtaining demographic data can better inform conservation and management. We summarized multi-year samples targeting the Alligator Snapping Turtle in large river systems and major tributaries of Alabama. Although catch per unit effort was lower than observed in other states, we filled distribution gaps in several river systems. Though studies from other states documented effects of historical harvesting pressure on Alligator Snapping Turtle populations, mean adult mass of both sexes, mean carapace length, and capture ratio of adults to immatures suggested reduced pressures in Alabama and possible population stability. Nevertheless, smaller tributaries and creeks need to be surveyed for previously undetected populations with mark–recapture and movement studies.


Georgia Distribution and Characterization of Species within the Eurycea quadridigitata Complex

July 2022

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83 Reads

Southeastern Naturalist

We performed systematic surveys for members of the Eurycea quadridigitata (Dwarf Salamander) complex in Georgia to determine the distribution of member species. Over 409 person-hours of search effort, we sampled 211 locations in 38 counties, detecting 108 individuals, including many that bridge substantial distribution gaps. Molecular data document Georgia specimens belong to 2 species, E. hillisi and E. quadridigitata. No specimens attributable to E. chamberlaini were found in Georgia, indicating this species has a distribution restricted to South and North Carolina. We used color patterns and measures of body size and shape of specimens identified by molecular methods to separate 4 species of 4-toed Eurycea from the Coastal Plain of Alabama, Florida, Georgia, South Carolina, and North Carolina. Large body size and the presence of melanophores across the chin and surrounding the cloaca separated E. quadridigitata from all others. Additionally, E. sphagnicola and E. quadridigitata possessed dark sides to the body and tail, with these features being disrupted by irregular white stripes or spots. In contrast, E. chamberlaini and E. hillisi possessed light sides to the body and tail that lacked irregular white stripes. Small body size distinguished E. sphagnicola from E. quadridigitata, but no feature distinguished E. chamberlaini from E. hillisi.



Figure 1. Two species of abundant terrestrial frogs and a predatory spider (Ctenidae) from lowland Caribbean forests of Costa Rica: (A) Craugastor bransfordii, (B) Oophaga pumilio and (C) Cupiennius sp. consuming a rain frog (Pristimantis ridens). See Folt and Lapinski (2017) for other examples of ctenid spiders consuming various frog species from the Caribbean lowlands of Costa Rica. Photographs by J. Folt (A, B) and W. Lapinski (C).
Six models describing hypotheses for co-occurrence patterns between predatory spiders and prey frogs in forest plots at La Selva, Costa Rica. Species A is assumed to be dominant over species B. All models include terms describing species A and B differing in probability of occupancy and detection.
Habitat-dependent effects of predatory spiders on prey frogs in a Neotropical wet forest

August 2021

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129 Reads

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5 Citations

Journal of Tropical Ecology

In seasonal wet Neotropical forests, many studies have suggested that species-rich terrestrial frog assemblages are regulated bottom-up by the abundance of leaf litter. However, terrestrial frogs are prey to a diverse community of predators, and no studies have tested for top-down effects of predators on this or other anuran assemblages. Here, we used an extensive field dataset to model the relative contribution of food resources, microhabitat resources and predators towards the occupancy and detection of two frog species ( Craugastor bransfordii and Oophaga pumilio ) at La Selva, Costa Rica. Frog occupancy was most strongly influenced by predatory spiders and secondarily influenced by the abundance of leaf litter. Predators exerted stronger effects on frogs than food resources, and frogs avoided predators more as leaf litter decreased. Detection probability was elevated when predators were present. We found support for bottom-up effects of leaf litter on the terrestrial frog assemblage, but top-down effects by predators exerted stronger effects on frog occupancy and detection. Because predator avoidance varied along a resource gradient, predator and resource effects appear to be dependent, supporting interactions between top-down and bottom-up mechanisms. Climate-driven decreases in leaf litter may drive decreased availability of frog refugia and increased interactions between frogs and predators.


Argentine Black and White Tegu (Salvator merianae) can survive the winter under semi-natural conditions well beyond their current invasive range

March 2021

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890 Reads

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21 Citations

The Argentine Black and White Tegu (Salvator merianae, formerly Tupinambis merianae) is a large lizard from South America. Now established and invasive in southern Florida, and it poses threats to populations of many native species. Models suggest much of the southern United States may contain suitable temperature regimes for this species, yet there is considerable uncertainty regarding either the potential for range expansion northward out of tropical and subtropical zones or the potential for the species establishing elsewhere following additional independent introductions. We evaluated survival, body temperature, duration and timing of winter dormancy, and health of wild-caught tegus from southern Florida held in semi-natural enclosures for over a year in Auburn, Alabama (> 900 km northwest of capture location). Nine of twelve lizards emerged from winter dormancy and seven survived the greater-than-one-year duration of the study. Average length of dormancy (176 d) was greater than that reported in the native range or for invasive populations in southern Florida and females remained dormant longer than males. Tegus grew rapidly throughout the study and the presence of sperm in the testes of males and previtellogenic or early vitellogenic follicles in female ovaries at the end of our study suggest the animals would have been capable of reproduction the following spring. The survival and overall health of the majority of adult tegus in our study suggests weather and climate patterns are unlikely to prevent survival following introduction in many areas of the United States far from their current invasive range.


Figure 2. Captures of gopher tortoises from 6 study populations (inset numbers) in Conecuh National Forest, Alabama, USA, 1991-2020. Black lines indicate the spatial extent searched for tortoise burrows during each sampling session at each study site; we did not include captures outside of the study area in analyses. Maps courtesy of Google (Mountain View, CA, USA) and Maxar Technologies (Westminster, CO, USA).
Demographic parameters used to predict recruitment in demographic projection models for populations of the gopher tortoise in Conecuh National Forest, Alabama, USA during simulated population projections.
Contrasting Patterns of Demography and Population Viability Among Gopher Tortoise Populations in Alabama

February 2021

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519 Reads

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21 Citations

Journal of Wildlife Management

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. Small gopher tortoise populations (10–50 individuals) may be stable and persist with low extinction risk if adult female survival is sufficiently high (≥0.95).


Citations (70)


... Tortoises sometimes move long distances between suitable patches of habitat, complicating multi-year studies of survival because mortality is often confounded with dispersal (Eubanks et al. 2003, Hunter and Rostal 2021, Guyer et al. 2024. A strength of this study is that we observed mortality directly via carcass encounter, rather than inferring it from absences of marked individuals that could be caused by dispersal. ...

Reference:

Social composition of soft-release groups is correlated with survival of translocated gopher tortoises
Annual and Lifetime Home Ranges Reveal Movement Patterns Within and Among Local Populations of Gopher Tortoises (Gopherus polyphemus)

Chelonian Conservation and Biology

... Here, we use a major recipient site for mitigation translocations to test for site-of-origin effects and impacts of genetic admixture on reproductive behavior and success in the gopher tortoise (Gopherus polyphemus), a keystone species and ecosystem engineer of the coastal plain of the south-eastern United States. This species exhibits substantial population structure across its range (Gaillard et al., 2017), including neutral (Krohn et al., 2024) and possibly adaptive (Elbers, Brown, & Taylor, 2018) genetic differentiation on relatively small scales. The gopher tortoise inhabits fire-maintained habitats, and its burrows promote diversity by serving as critical thermal and fire refugia for hundreds of species of commensals (Catano & Stout, 2015). ...

Using genomic data to estimate population structure of Gopher Tortoise (Gopherus polyphemus) populations in Southern Alabama

Conservation Genetics

... Corytophanes cristatus is a member of the family Corytophanidae, a group of New World lizards including the genera Basiliscus, Corytophanes, and Laemanctus called casquehead lizards or helmeted lizards due to their prominent parietal crest. Basiliscus, Corytophanes, and Laemanctus are sympatric in Costa Rica (Leenders, 2019); however, only Basiliscus and Corytophanes are present at La Selva Biological Station (Guyer and Donnelly, 2005). ...

Amphibians and Reptiles of La Selva, Costa Rica, and the Caribbean Slope: A Comprehensive Guide
  • Citing Book
  • November 2023

... As currently understood, the habitat requirements of Gopher Tortoises include herbaceous ground cover for foraging, ground-level sunlight for thermoregulating, and soils appropriate for burrowing and nesting (Auffenberg and Iverson 1979;Auffenberg and Franz 1982;Diemer 1986;Mushinsky et al. 2006;Pudner et al. 2021). The Gopher Tortoise is typically associated with sandy, well-drained soils characteristic of Longleaf Pine (Pinus palustris) savanna and sandhill ecosystems. ...

Changes in Vegetation Structure and Gopher Tortoise Population Structure after 17 Years of Restoration Management
  • Citing Article
  • October 2021

Natural Areas Journal

... Spiders are opportunistic generalist predators found in all terrestrial environments (Turnbull 1973) and are one of the largest groups of obligate carnivores (Li et al. 2021). Diets of spiders consist primarily of invertebrate prey, although species from dozens of families (e.g., Pisauridae, Theraphosidae, Dictynidae, Lycosidae) have been shown to consume vertebrate prey (Menin et al. 2005;Toledo 2005;Murray et al. 2016;Nyffeler et al. 2017;Baba et al. 2019;Badjedjea Babangenge et al. 2019;Nyffeler and Altig 2020;Folt and Guyer 2021;Nyffeler and Gibbons 2022;Wilder and Simpson 2022;Michalko and Pekár 2024). Indeed, many spider species can capture and secure vertebrate prey; in some cases, prey that is much larger than the spider itself (Menin et al. 2005;Nyffeler et al. 2017;Baba et al. 2019;Badjedjea Babangenge et al. 2019;Nyffeler and Gibbons 2022;Wilder and Simpson 2022;Michalko and Pekár 2024). ...

Habitat-dependent effects of predatory spiders on prey frogs in a Neotropical wet forest

Journal of Tropical Ecology

... Both of these reptiles are highly successful and ecologically destructive invasive species in southern Florida, USA Offner et al., 2021). Though difficult to document in situ, these species' success within their novel environment is thought to be attributable to behavioral and thermal plasticity supporting individual survival and increased fecundity (Goetz et al., 2021;Shine et al., 1997;Stahlschmidt & DeNardo, 2010). For example, unlike most reptiles, Burmese pythons provide maternal care via nest brooding and clutch defense (e.g., Currylow, McCollister, et al., 2022) and may produce heat for eggs via shivering thermogenesis (Snow et al., 2010). ...

Argentine Black and White Tegu (Salvator merianae) can survive the winter under semi-natural conditions well beyond their current invasive range

... Much of EAFB is upland sandhill habitat, dominated by pine savanna that is managed by thinning and prescribed burning. Native populations of Gopher Tortoises in this region are historically depleted because of harvesting by humans for food (Auffenberg and Franz 1982) and are predicted to experience continued declines and possible local extinction owed to excessively low densities (Chandler et al. 2020;Folt et al. 2021). Under the Florida Fish and Wildlife Conservation Commission (FFWCC) permits, over 10,000 tortoises have been translocated onto EAFB into areas of existing habitat of Gopher Tortoises since 2015. ...

Contrasting Patterns of Demography and Population Viability Among Gopher Tortoise Populations in Alabama

Journal of Wildlife Management

... Existing research suggests ways for identifying such communities (Becerra, 2007) and for how they differ from communities where member species have not co-evolved together (Semchenko et al., 2019). Our plea for identifying ESS communities aligns with Steen and colleagues' (2017) Evolutionary Community Concept (for critiques, see Sagoff, 2019;Barrett et al., 2020). ...

The evolutionary community concept is fully armed and operational: a reply to Sagoff

Biology & Philosophy

... The consistency of these trends leads us to speculate that scrub may act as a source habitat, despite the lack of a significant difference in our tortoise density estimates for scrub and pine flatwoods. Goessling et al. (2021) found that periodic emigration and immigration among local aggregations of tortoises were important aspects of population viability in the low-density populations of Alabama. Telemetry data from APAFR, on the other hand, revealed frequent forays by males to adjacent habitats, presumably to court females, but high site fidelity and little movement between habitats for female tortoises (Castellón et al. 2018). ...

Implications from Monitoring Gopher Tortoises at Two Spatial Scales
  • Citing Article
  • October 2020

Journal of Wildlife Management

... These EIDs outbreaks affect both captive and wild animals (Williams et al., 2002;Latney and Wellehan, 2013;Schilliger et al., 2023). The rapid spread of R. orientalis as an invasive parasite in native reptile species and its infection intensity in southern Florida (Miller et al., 2020) highlights the potential threat which may pose to free-ranging wildlife in Europe and potentially worldwide. A major contributing factor is the growing animal trade and the increase in invasive species resulting from it (Keller et al., 2011;Lockwood et al., 2019). ...

Highly Competent Native Snake Hosts Extend the Range of An Introduced Parasite Beyond Its Invasive Burmese Python Host

The Bulletin of the Ecological Society of America