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Geology topics

Joshua R. Ennen

Publications and source records attributed to Joshua R. Ennen.

36 records · Page 2Linked to original sources

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

Intersexual allometry differences and ontogenetic shifts of coloration patterns in two aquatic turtles, Graptemys oculifera and Graptemys flavimaculata

Coloration can play critical roles in a species' biology. The allometry of color patterns may be useful for elucidating the evolutionary mechanisms responsible for shaping the traits. We measured characteristics relating to eight aspects of color patterns from Graptemys oculifera and G. flavimaculata to investigate the allometric differences among male, female, and unsexed juvenile specimens. Additionally, we investigated ontogenetic shifts by incorporating the unsexed juveniles into the male and female datasets. In general, male color traits were isometric (i.e., color scaled with body size), while females and juvenile color traits were hypoallometric, growing in size more slowly than the increase in body size. When we included unsexed juveniles in our male and female datasets, our linear regression analyses found all relationships to be hypoallometric and our model selection analysis found support for nonlinear models describing the relationship between body size and color patterns, suggestive of an ontogenetic shift in coloration traits for both sexes at maturity. Although color is critical for many species' biology and therefore under strong selective pressure in many other species, our results are likely explained by an epiphenomenon related to the different selection pressures on body size and growth rates between juveniles and adults and less attributable to the evolution of color patterns themselves.

Ecology and Evolution

Turbines and terrestrial vertebrates: variation in tortoise survivorship between a wind energy facility and an adjacent undisturbed wildland area in the desert southwest (USA)

With the recent increase in utility-scale wind energy development, researchers have become increasingly concerned how this activity will affect wildlife and their habitat. To understand the potential impacts of wind energy facilities (WEF) post-construction (i.e., operation and maintenance) on wildlife, we compared differences in activity centers and survivorship of Agassiz's desert tortoises ( Gopherus agassizii ) inside or near a WEF to neighboring tortoises living near a wilderness area (NWA) and farther from the WEF. We found that the size of tortoise activity centers varied, but not significantly so, between the WEF (6.25 ± 2.13 ha) and adjacent NWA (4.13 ± 1.23 ha). However, apparent survival did differ significantly between the habitat types: over the 18 year study period apparent annual survival estimates were 0.96 ± 0.01 for WEF tortoises and 0.92 ± 0.02 for tortoises in the NWA. High annual survival suggests that operation and maintenance of the WEF has not caused considerable declines in the adult population over the past two decades. Low traffic volume, enhanced resource availability and decreased predator populations may influence annual survivorship at this WEF. Further research on these proximate mechanisms and population recruitment would be useful for mitigating and managing post-development impacts of utility scale wind energy on long-lived terrestrial vertebrates.

Environmental Management

Not putting all their eggs in one basket: bet-hedging despite extraordinary annual reproductive output of desert tortoises

Bet-hedging theory makes the counter-intuitive prediction that, if juvenile survival is low and unpredictable, organisms should consistently reduce short-term reproductive output to minimize the risk of reproductive failure in the long-term. We investigated the long-term reproductive output of an Agassiz's desert tortoise ( Gopherus agassizii ) population and conformance to a bet-hedging strategy of reproduction in an unpredictable but comparatively productive environment. Most females reproduced every year, even during periods of low precipitation and poor germination of food plants, and the mean percentage of reproducing females did not differ significantly on an annual basis. Although mean annual egg production (clutch size × clutch frequency) differed significantly among years, mean clutch size and mean clutch frequency remained relatively constant. During an El Niño year, mean annual egg production and mean annual clutch frequency were the highest ever reported for this species. Annual egg production was positively influenced by maternal body size but clutch size and clutch frequency were not. Our long-term results confirm earlier conclusions based on short-term research that desert tortoises have a bet-hedging strategy of producing small clutches almost every year. The risk of long-term reproductive failure is minimized in unpredictable environments, both through time by annually producing multiple small clutches over a long reproductive lifespan, even in years of low resource availability, and through space by depositing multiple annual clutches in different locations. The extraordinary annual reproductive output of this population appears to be the result of a typically high but unpredictable biomass of annual food plants at the site relative to tortoise habitat in dryer regions. Under the comparatively productive but unpredictable conditions, tortoises conform to predictions of a bet-hedging strategy of reproduction with relatively small but consistent clutch sizes.

Biological Journal of the Linnean Society

Using motion-sensor camera technology to infer seasonal activity and thermal niche of the desert tortoise ( Gopherus agassizii )

Understanding the relationships between environmental variables and wildlife activity is an important part of effective management. The desert tortoise ( Gopherus agassizii ), an imperiled species of arid environments in the southwest US, may have increasingly restricted windows for activity due to current warming trends. In summer 2013, we deployed 48 motion sensor cameras at the entrances of tortoise burrows to investigate the effects of temperature, sex, and day of the year on the activity of desert tortoises. Using generalized estimating equations, we found that the relative probability of activity was associated with temperature (linear and quadratic), sex, and day of the year. Sex effects showed that male tortoises are generally more active than female tortoises. Temperature had a quadratic effect, indicating that tortoise activity was heightened at a range of temperatures. In addition, we found significant support for interactions between sex and day of the year, and sex and temperature as predictors of the probability of activity. Using our models, we were able to estimate air temperatures and times (days and hours) that were associated with maximum activity during the study. Because tortoise activity is constrained by environmental conditions such as temperature, it is increasingly vital to conduct studies on how tortoises vary their activity throughout the Sonoran Desert to better understand the effects of a changing climate.

California

The effect of research activities and winter precipitation on voiding behaviour of Agassiz’s desert tortoises ( Gopherus agassizii )

Context: There is little information available on how research activities might cause stress responses in wildlife, especially responses of threatened species such as the desert tortoise ( Gopherus agassizii ). Aims: The present study aims to detect behavioural effects of researcher handling and winter precipitation on a natural population of desert tortoises in the desert of Southwestern United States, over the period 1997 to 2014, through extensive assessments of capture events during multiple research studies, and capture–mark–recapture survivorship analysis. Methods: Juvenile and adult desert tortoises were repeatedly handled with consistent methodology across 18 years during 10 study seasons. Using a generalised linear mixed-effects model, we assessed the effects of both research manipulation and abiotic conditions on probability of voiding. Additionally, we used a Cormack–Jolly–Seber model to assess the effects of winter precipitation and voiding on long-term apparent survivorship. Key results: Of 1008 total capture events, voiding was recorded on 83 (8.2%) occasions in 42 different individuals. Our top models indicated that increases in handling time led to significantly higher probabilities of voiding for juveniles, females and males. Similarly, increases in precipitation resulted in significantly higher probabilities of voiding for juveniles and females, but not for males. Tortoise capture frequency was negatively correlated with voiding occurrence. Cormack–Jolly–Seber models demonstrated a weak effect of winter precipitation on survivorship, but a negligible effect for both voiding behaviour and sex. Conclusions: Handling-induced voiding by desert tortoises may occur during common research activities and years of above average winter precipitation. Increased likelihood of voiding in individuals with relatively low numbers of recaptures suggested that tortoises may have perceived researchers initially as predators, and therefore voided as a defensive strategy. Voiding does not appear to impact long-term survivorship in desert tortoises at this site. Implications: This study has demonstrated that common handling practices on desert tortoise may cause voiding behaviour. These results suggest that in order to minimise undesirable behavioural responses in studied desert tortoise populations, defined procedures or protocols must be followed by the investigators to reduce contact period to the extent feasible.

Wildlife Research

Clinal variation or validation of a subspecies? A case study of the Graptemys nigrinoda complex (Testudines: Emydidae)

Widely distributed species often display intraspecific morphological variation due to the abiotic and biotic gradients experienced across their ranges. Historically, in many vertebrate taxa, such as birds and reptiles, these morphological differences within a species were used to delimit subspecies. Graptemys nigrinoda is an aquatic turtle species endemic to the Mobile Bay Basin. Colour pattern and morphological variability were used to describe a subspecies ( G. n. delticola ) from the lower reaches of the system, although it and the nominate subspecies also reportedly intergrade over a large portion of the range. Other researchers have suggested that these morphological differences merely reflect clinal variation. Our molecular data (mtDNA) did not support the existence of the subspecies, as the haplotypes were differentiated by only a few base pairs and one haplotype was shared between the putative subspecies. While there were significant morphological and pattern differences among putative specimens of G. n. nigrinoda, G. n. delticola and G. n. nigrinoda × delticola , these differences probably represent clinal variation as they were also related to environmental variables [i.e. cumulative drainage area and drainage (categorical)]. Specimens occupying slow-current, high-turbidity river reaches (e.g. the Tensaw River) exhibited greater relative carapace heights and more dark pigmentation, while specimens occupying fast-current, clearer rivers (e.g. the upper Alabama, Cahaba and Tallapoosa rivers) exhibited lower carapace heights and more yellow pigmentation. Given the absence of clear molecular and morphological differences that are related to drainage characteristics, we suggest that there is not sufficient evidence for the recognition of G. n. delticola as a distinct subspecies.

Alabama;Mississippi

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

Nest site characteristics, nesting movements, and lack of long-term nest site fidelity in Agassiz's desert tortoises at a wind energy facility in southern California

Nest site selection has important consequences for maternal and offspring survival and fitness. Females of some species return to the same nesting areas year after year. We studied nest site characteristics, fidelity, and daily pre-nesting movements in a population of Agassiz’s desert tortoises (Gopherus agassizii) at a wind energy facility in southern California during two field seasons separated by over a decade. No females returned to the same exact nest site within or between years but several nested in the same general area. However, distances between first and second clutches within a year (2000) were not significantly different from distances between nests among years (2000 and 2011) for a small sample of females, suggesting some degree of fidelity within their normal activity areas. Environmental attributes of nest sites did not differ significantly among females but did among years due largely to changes in perennial plant structure as a result of multiple fires. Daily pre-nesting distances moved by females decreased consistently from the time shelled eggs were first visible in X-radiographs until oviposition, again suggesting some degree of nest site selection. Tortoises appear to select nest sites that are within their long-term activity areas, inside the climate-moderated confines of one of their self-constructed burrows, and specifically, at a depth in the burrow that minimizes exposure of eggs and embryos to lethal incubation temperatures. Nesting in “climate-controlled” burrows and nest guarding by females relaxes some of the constraints that drive nest site selection in other oviparous species.

California

Graptemys gibbonsi Lovich and McCoy -- Pascagoula Map Turtle

DEFINITION. The Pascagoula Map Turtle, Graptemys gibbonsi , is a large riverine species that exhibits pronounced sexual dimorphism, where females attain a maximum carapace length (CL) of 295 mm and males a maximum of 141 mm (Lovich et al. 2009). Mean adult female CL (248 mm) can be well over twice the mean CL of adult males (104 mm; Gibbons and Lovich 1990, Lovich et al. 2009). In addition, females have conspicuously enlarged heads (37.9 mm, SD = 14.0 mm) with broad alveolar surfaces (12.1 mm, SD = 4.9) compared to males (head width – 16.4 mm, SD = 1.1 mm; alveolar width – 4.3 mm, SD = 0.40 mm; Lindeman, unpublished data). Males have longer tails with the vent posterior to the edge of the carapace. Both sexes have relatively flat plastrons. Similar to other species within the pulchra clade, Graptemys gibbonsi possess a high-domed shell with a median keel. The median carapace keel is composed of prominent spines on the posterior portions of the second and third vertebrals. A broken black stripe, most pronounced anteriorly, marks the median keel of the vertebrals, and pleural scutes 1– 3 have a network of intersecting yellow lines or circular yellow markings on the distal parts. The plastron is pale yellow with dark pigment on some seams. Ground color of the head and limbs is brown to olive with light yellow or yellowish-green stripes and blotches. The yellow pigment on the upper marginal scutes is wide in comparison to other members of the pulchra clade. Hatchling pigmentation patterns resemble those of adults, but with more conspicuous patterns on the pleural scutes. Similarly, the plastron of hatchlings commonly has more dark pigmentation along the seams than adults. The shell is highly serrated along the edge of the carapace and the vertebral keel is more pronounced than in adults.

Mississippi

Assessing the state of knowledge of utility-scale wind energy development and operation on non-volant terrestrial and marine wildlife

A great deal has been published in the scientific literature regarding the effects of wind energy development and operation on volant (flying) wildlife including birds and bats, although knowledge of how to mitigate negative impacts is still imperfect. We reviewed the peer-reviewed scientific literature for information on the known and potential effects of utility-scale wind energy development and operation (USWEDO) on terrestrial and marine non-volant wildlife and found that very little has been published on the topic. Following a similar review for solar energy we identified known and potential effects due to construction and eventual decommissioning of wind energy facilities. Many of the effects are similar and include direct mortality, environmental impacts of destruction and modification of habitat including impacts of roads, and offsite impacts related to construction material acquisition, processing and transportation. Known and potential effects due to operation and maintenance of facilities include habitat fragmentation and barriers to gene flow, as well as effects due to noise, vibration and shadow flicker, electromagnetic field generation, macro- and micro-climate change, predator attraction, and increased fire risk. The scarcity of before-after-control-impact studies hinders the ability to rigorously quantify the effects of USWEDO on non-volant wildlife. We conclude that more empirical data are currently needed to fully assess the impact of USWEDO on non-volant wildlife.

Applied Energy

A quantitative analysis of the state of knowledge of turtles of the United States and Canada

The “information age” ushered in an explosion of knowledge and access to knowledge that continues to revolutionize society. Knowledge about turtles, as measured by number of published papers, has been growing at an exponential rate since the early 1970s, a phenomenon mirrored in all scientific disciplines. Although knowledge about turtles, as measured by number of citations for papers in scientific journals, has been growing rapidly, this taxonomic group remains highly imperiled suggesting that knowledge is not always successfully translated into effective conservation of turtles. We reviewed the body of literature on turtles of the United States and Canada and found that: 1) the number of citations is biased toward large-bodied species, 2) the number of citations is biased toward wide-ranging species, and 3) conservation status has little effect on the accumulation of knowledge for a species, especially after removing the effects of body size or range size. The dispersion of knowledge, measured by Shannon Weiner diversity and evenness indices across species, was identical from 1994 to 2009 suggesting that poorly studied species remained poorly-studied species while well-studied species remained well studied. Several species listed as threatened or endangered under the U.S. Endangered Species Act (e.g., Pseudemys alabamensis, Sternotherus depressus, and Graptemys oculifera) remain poorly studied with the estimated number of citations for each ranging from only 13-24. The low number of citations for these species could best be explained by their restricted distribution and/or their smaller size. Despite the exponential increase in knowledge of turtles in the United States and Canada, no species of turtle listed under the Endangered Species Act has ever been delisted for reason of recovery. Therefore, increased knowledge does not necessarily contribute appreciably to recovery of threatened turtles.

Amphibia-Reptilia

Nest guarding by female Agassiz's desert tortoise (Gopherus agassizii) at a wind-energy facility near Palm Springs, California

We observed behavior consistent with nest-guarding in Agassiz's desert tortoise (Gopherus agassizii) at two nests in a large wind-energy-generation facility near Palm Springs, California, locally known as the Mesa Wind Farm. As researchers approached the nests, female desert tortoises moved to the entrance of their burrows and positioned themselves sideways, directly over their nests. One female stretched her limbs outward and wedged herself into the burrow (her plastron directly above the nest). Guarding of nests is rarely observed in Agassiz's desert tortoise but can occur as a result of attempted predation on eggs by Gila monsters (Heloderma suspectum) or in direct response to the perceived threat posed by researchers. This is the first report of nest-guarding for G. agassizii in the Sonoran Desert ecosystem of California.

California

Home range characteristics and overwintering ecology of the stripe-necked musk turtle ( Sternotherus minor peltifer ) in middle Tennessee

Little is known about the movement behavior of the stripe-necked musk turtle, Sternotherus minor peltifer . Using radiotelemetry, we calculated mean (± SD) home range length, which was 341.4 ± 90.3 m, with home range length not differing between the sexes (males, 335 ± 194 m; females, 346 ± 79.5 m). Sternotherus m. peltifer were active in every month of the year but decreased their movement distance and frequency between December and March; during nonwinter and winter periods, individuals used limestone bluffs most often.

Tennessee

Nesting ecology of a population of Gopherus agassizii at a utility-scale wind energy facility in southern California

We investigated the annual nesting ecology of a population of Desert Tortoises (Gopherus agassizii) inhabiting a utility-scale renewable energy (USRE) facility in southern California and compared our results with populations inhabiting relatively undisturbed sites. In 2000, 15 radio-tracked females produced 29 clutches, and 24 nests were monitored to examine nest-site selection, nest predation, hatching success, date of emergence of hatchlings, and hatchling mass and carapace length. Overall, the nesting ecology of the population inhabiting the USRE facility was very similar to other populations of Desert Tortoises inhabiting relatively undisturbed habitats. Oviposition occurred from 12 May to 8 July, which was similar to other sites. Nest depths (11.1 cm), nest predation (12%), hatchling emergence date (7 August and 29 September), and hatchling morphometrics (i.e., MCL: 44.5 mm; mass: 23 g) were all within ranges reported in other populations. Unlike within other populations, we observed no relationship between hatchling size and either maternal body size or egg width. We found no evidence of females selecting for a particular burrow for oviposition of eggs based on environmental or anthropogenic variables. Most nests were located in or near burrows, and nest depth was greater for nests near the entrance than those deeper in the burrow. Although this study suggests that the nesting ecology of the Desert Tortoise population we studied was not adversely affected by the USRE facility, this relationship is only correlative because our study was not a before-after-control-impact (BACI) study, which would establish a cause and effect relationship. As pointed out in a recent review, BACI studies are critically needed to address the wildlife impacts of utility-scale renewable energy development.

California

A comparison of artificial incubation and natural incubation hatching success of gopher tortoise (Gopherus polyphemus) eggs in southern Mississippi

Recent studies have found that Gopher Tortoise, Gopherus polyphemus, populations in southern Mississippi exhibit low recruitment, due in part to very low hatching success of their eggs. We sought to determine if the cause(s) of this low hatching success was related to egg quality (intrinsic factors), unsuitability of the nest environment (extrinsic factors), or a combination of the two. In 2003, hatching success was monitored simultaneously for eggs from the same clutches that were incubated in the laboratory and left to incubate in nests. A subset of randomly chosen eggs from each clutch was incubated in the laboratory under physical conditions that were known to be conducive to successful hatching to estimate the proportion of eggs that were capable of hatching in a controlled setting. Hatching success in the laboratory was compared with that of eggs incubated in natural nests to estimate the proportion of eggs that failed to hatch presumably from extrinsic factors. Laboratory hatching success was 58.8%, suggesting that roughly 40% of the eggs were intrinsically incapable of hatching even when incubated under controlled conditions. Hatching success in natural nests, 16.7%, was significantly lower than hatching success in the laboratory, suggesting that approximately 42.1% of eggs were capable of hatching but failed to hatch due to some extrinsic aspect(s) of the nest environment. Thus, the low hatching success of Gopher Tortoise eggs in southern Mississippi appears to be attributable to a combination of intrinsic (egg quality) and extrinsic (nest environment) factors.

Mississippi

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.

California

Distribution and habitat utilization of the gopher tortoise tick (Amblyomma tuberculatum) in southern Mississippi

The distribution of the gopher tortoise tick ( Amblyomma tuberculatum ) has been considered intrinsically linked to the distribution of its primary host, gopher tortoises ( Gopherus polyphemus ). However, the presence of G. polyphemus does not always equate to the presence of A. tuberculatum . There is a paucity of data on the ecology, habitat preferences, and distribution of A. tuberculatum . The goals of this study were to assess the distribution of A. tuberculatum in southern Mississippi and to determine which, if any, habitat parameters explain the distribution pattern of A. tuberculatum . During 2006–2007, we examined 13 G. polyphemus populations in southern Mississippi for the presence of A. tuberculatum , and we measured a suite of habitat parameters at each site. Only 23% of the G. polyphemus populations supported A. tuberculatum , suggesting a more restricted distribution than its host. The results of our multivariate analyses identified several habitat variables, e.g., depth of sand and percentage of sand in the topsoil and burrow apron, as being important in discriminating between sites with, and without, A. tuberculatum . Amblyomma tuberculatum was only found at sites with a mean sand depth of >100 cm and a mean percentage of topsoil and burrow apron sand composition >94.0 and 92.4, respectively. Thus, environmental factors, and not just its host's range, seem to influence the distribution of A. tuberculatum .

Mississippi