Kenneth E. Nussear

Making molehills out of mountains: landscape genetics of the Mojave desert tortoise

Heterogeneity in habitat often influences how organisms traverse the landscape matrix that connects populations. Understanding landscape connectivity is important to determine the ecological processes that influence those movements, which lead to evolutionary change due to gene flow. Here, we used landscape genetics and statistical models to evaluate hypotheses that could explain isolation among locations of the threatened Mojave desert tortoise (Gopherus agassizii). Within a causal modeling framework, we investigated three factors that can influence landscape connectivity: geographic distance, barriers to dispersal, and landscape friction. A statistical model of habitat suitability for the Mojave desert tortoise, based on topography, vegetation, and climate variables, was used as a proxy for landscape friction and barriers to dispersal. We quantified landscape friction with least-cost distances and with resistance distances among sampling locations. A set of diagnostic partial Mantel tests statistically separated the hypotheses of potential causes of genetic isolation. The best-supported model varied depending upon how landscape friction was quantified. Patterns of genetic structure were related to a combination of geographic distance and barriers as defined by least-cost distances, suggesting that mountain ranges and extremely low-elevation valleys influence connectivity at the regional scale beyond the tortoises’ ability to disperse. However, geographic distance was the only influence detected using resistance distances, which we attributed to fundamental differences between the two ways of quantifying friction. Landscape friction, as we measured it, did not influence the observed patterns of genetic distances using either quantification. Barriers and distance may be more valuable predictors of observed population structure for species like the desert tortoise, which has high dispersal capability and a long generation time.

Lizards, Lipids, and Dietary Links to Animal Function

Our experiments were designed to test the hypotheses that dietary lipids can affect whole‐animal physiological processes in a manner concordant with changes in the fluidity of cell membranes. We measured (1) the lipid composition of five tissues, (2) body temperatures selected in a thermal gradient (Tsel), (3) the body temperature at which the righting reflex was lost (critical thermal minimal [CTMin]), and (4) resting metabolic rate (RMR) at three body temperatures in desert iguanas (Dipsosaurus dorsalis) fed diets enriched with either saturated or unsaturated fatty acids. The composition of lipids in tissues of the lizards generally reflected the lipids in their diets, but the particular classes and ratios of fatty acids varied among sampled organs, indicating the conservative nature of some tissues (e.g., brain) relative to others (e.g., depot fat). Lizards fed the diet enriched with saturated fatty acids selected warmer nighttime body temperatures than did lizards fed a diet enriched with unsaturated fatty acids. This difference is concordant with the hypothesis that the composition of dietary fats influences membrane fluidity and that ectotherms may compensate for such changes in fluidity by selecting different body temperatures. The CTMin of the two treatment groups was indistinguishable. This may reflect the conservatism of some tissues (e.g., brain) irrespective of diet treatment. The RMR of the saturated treatment group nearly doubled between 30° and 40°C. Here, some discrete membrane domains in the lizards fed the saturated diet may have been in a more‐ordered phase at 30°C and then transformed to a less‐ordered phase at 40°C. In contrast, the RMR of the unsaturated treatment group exhibited temperature independence in metabolic rate from 30° to 40°C. Perhaps the unsaturated diet resulted in membranes that developed a higher degree of disorder (i.e., a certain phase) at a lower temperature than were membranes of lizards fed the saturated diet. Our study demonstrates links between dietary fats and whole‐animal physiology; however, the mechanistic basis of these links, and the general knowledge of lipid metabolism in squamate reptiles, remain poorly understood and warrant further study.

Microclimate and limits to photosynthesis in a diverse community of hypolithic cyanobacteria in northern Australia.

Hypolithic microbes, primarily cyanobacteria, inhabit the highly specialized microhabitats under translucent rocks in extreme environments. Here we report findings from hypolithic cyanobacteria found under three types of translucent rocks (quartz, prehnite, agate) in a semiarid region of tropical Australia. We investigated the photosynthetic responses of the cyanobacterial communities to light, temperature and moisture in the laboratory, and we measured the microclimatic variables of temperature and soil moisture under rocks in the field over an annual cycle. We also used molecular techniques to explore the diversity of hypolithic cyanobacteria in this community and their phylogenetic relationships within the context of hypolithic cyanobacteria from other continents. Based on the laboratory experiments, photosynthetic activity required a minimum soil moisture of 15% (by mass). Peak photosynthetic activity occurred between approximately 8 degrees C and 42 degrees C, though some photosynthesis occurred between -1 degrees C and 51 degrees C. Maximum photosynthesis rates also occurred at light levels of approximately 150-550 micromol m(-2) s(-1). We used the field microclimatic data in conjunction with these measurements of photosynthetic efficiency to estimate the amount of time the hypolithic cyanobacteria could be photosynthetically active in the field. Based on these data, we estimated that conditions were appropriate for photosynthetic activity for approximately 942 h (approximately 75 days) during the year. The hypolithic cyanobacteria community under quartz, prehnite and agate rocks was quite diverse both within and between rock types. We identified 115 operational taxonomic units (OTUs), with each rock hosting 8-24 OTUs. A third of the cyanobacteria OTUs from northern Australia grouped with Chroococcidiopsis, a genus that has been identified from hypolithic and endolithic communities from the Gobi, Mojave, Atacama and Antarctic deserts. Several OTUs identified from northern Australia have not been reported to be associated with hypolithic communities previously.

Desert Tortoise Hibernation: Temperatures, Timing, and Environment

Abstract This research examined the onset, duration, and termination of hibernation in Desert Tortoises (Gopherus agassizii) over several years at multiple sites in the northeastern part of their geographic range, and recorded the temperatures experienced by tortoises during winter hibernation. The timing of hibernation by Desert Tortoises differed among sites and years. Environmental cues acting over the short-term did not appear to influence the timing of the hibernation period. Different individual tortoises entered hibernation over as many as 44 days in the fall and emerged from hibernation over as many as 49 days in the spring. This range of variation in the timing of hibernation indicates a weak influence at best of exogenous cues hypothesized to trigger and terminate hibernation. There do appear to be regional trends in hibernation behavior as hibernation tended to begin earlier and continue longer at sites that were higher in elevation and generally cooler. The emergence date was generally more similar among study sites than the date of onset. While the climate and the subsequent timing of hibernation differed among sites, the average temperatures experienced by tortoises while hibernating differed by only about five degrees from the coldest site to the warmest site.

Long‐term plant responses to climate are moderated by biophysical attributes in a North American desert

Summary 1. Recent elevated temperatures and prolonged droughts in many already water-limited regions throughout the world, including the southwestern United States, are likely to intensify according to future climate-model projections. This warming and drying can negatively affect perennial vegetation and lead to the degradation of ecosystem properties. 2. To better understand these detrimental effects, we formulate a conceptual model of dryland ecosystem vulnerability to climate change that integrates hypotheses on how plant species will respond to increases in temperature and drought, including how plant responses to climate are modified by landscape, soil and plant attributes that are integral to water availability and use. We test the model through a synthesis of fifty years of repeat measurements of perennial plant species cover in large permanent plots across the Mojave Desert, one of the most water-limited ecosystems in North America. 3. Plant species ranged in their sensitivity to precipitation in different seasons, capacity to increase in cover with high precipitation and resistance to decrease in cover with low precipitation. 4. Our model successfully explains how plant responses to climate are modified by biophysical attributes in the Mojave Desert. For example, deep-rooted plants were not as vulnerable to drought on soils that allowed for deep-water percolation, whereas shallow-rooted plants were better buffered from drought on soils that promoted water retention near the surface. 5. Synthesis. Our results emphasize the importance of understanding climate–vegetation relationships in the context of biophysical attributes that influence water availability and provide an important forecast of climate-change effects, including plant mortality and land degradation in dryland regions throughout the world.

CAN MODELING IMPROVE ESTIMATION OF DESERT TORTOISE POPULATION DENSITIES

The federally listed desert tortoise (Gopherus agassizii) is currently monitored using distance sampling to estimate population densities. Distance sampling, as with many other techniques for estimating population density, assumes that it is possible to quantify the proportion of animals available to be counted in any census. Because desert tortoises spend much of their life in burrows, and the proportion of tortoises in burrows at any time can be extremely variable, this assumption is difficult to meet. This proportion of animals available to be counted is used as a correction factor (g0) in distance sampling and has been estimated from daily censuses of small populations of tortoises (6-12 individuals). These censuses are costly and produce imprecise estimates of go due to small sample sizes. We used data on tortoise activity from a large (N = 150) experimental population to model activity as a function of the biophysical attributes of the environment, but these models did not improve the precision of estimates from the focal populations. Thus, to evaluate how much of the variance in tortoise activity is apparently not predictable, we assessed whether activity on any particular day can predict activity on subsequent days with essentially identical environmental conditions. Tortoise activity was only weakly correlated on consecutive days, indicating that behavior was not repeatable or consistent among days with similar physical environments.

Multiscale connectivity and graph theory highlight critical areas for conservation under climate change

Conservation planning and biodiversity management require information on landscape connectivity across a range of spatial scales from individual home ranges to large regions. Reduction in landscape connectivity due changes in land use or development is expected to act synergistically with alterations to habitat mosaic configuration arising from climate change. We illustrate a multiscale connectivity framework to aid habitat conservation prioritization in the context of changing land use and climate. Our approach, which builds upon the strengths of multiple landscape connectivity methods, including graph theory, circuit theory, and least-cost path analysis, is here applied to the conservation planning requirements of the Mohave ground squirrel. The distribution of this threatened Californian species, as for numerous other desert species, overlaps with the proposed placement of several utility-scale renewable energy developments in the American southwest. Our approach uses information derived at three spatial scales to forecast potential changes in habitat connectivity under various scenarios of energy development and climate change. By disentangling the potential effects of habitat loss and fragmentation across multiple scales, we identify priority conservation areas for both core habitat and critical corridor or stepping stone habitats. This approach is a first step toward applying graph theory to analyze habitat connectivity for species with continuously distributed habitat and should be applicable across a broad range of taxa.

Diet quality does not affect resting metabolic rate or body temperatures selected by an herbivorous lizard

Abstract Diet quality can influence many aspects of digestion, but the links between diet quality and resting metabolism are poorly understood. In nature, it might be beneficial to reduce energy expenditure when only poor quality diets are available. Alternatively, animals might increase the processing capacity of the gut to more thoroughly extract energy. If maintaining the processing capacity of the gut is energetically expensive, then increasing gut size or function might result in higher resting metabolism. In ectotherms, most digestive functions are temperature dependent, thus another strategy to maintain energy balance might be to alter selected body temperatures. We tested whether differing concentrations of dietary fiber affected the resting metabolic rate or body temperatures selected by chuckwallas (Sauromalus obesus) – lizards that naturally experience marked variation in dietary fiber. Resting metabolic rates measured at two temperatures and over three time intervals did not differ between groups of lizards force-fed low- (30% neutral-detergent fiber; NDF) and high-fiber (45% NDF) diets, nor did these diet differences influence body temperatures selected in a thermal gradient. We conclude that ecologically relevant differences in diet quality may have negligible effects on resting metabolic rates and body temperatures selected by chuckwallas.

Habitat drives dispersal and survival of translocated juvenile desert tortoises

Summary 1.In spite of growing reliance on translocations in wildlife conservation, translocation efficacy remains inconsistent. One factor that can contribute to failed translocations is releasing animals into poor quality or otherwise inadequate habitat. 2.Here we used a targeted approach to test the relationship of habitat features to post-translocation dispersal and survival of juvenile Mojave desert tortoises Gopherus agassizii. 3.We selected three habitat characteristics—rodent burrows, substrate texture (prevalence and size of rocks), and washes (ephemeral river beds)–that are tied to desert tortoise ecology. At the point of release, we documented rodent burrow abundance, substrate texture, and wash presence and analysed their relationship to maximum dispersal. We also documented relative use by each individual for each habitat characteristic and analysed their relationships with survival and fatal encounters with a predator in the first year after release. 4.In general, the presence of refugia or other areas that enabled animals to avoid detection, such as burrows and substrate, decreased overall mortality as well as predator-mediated mortality. The presence of washes and substrate that enhanced the tortoises’ ability to avoid detection also associated with reduced dispersal away from the release site. These results indicate an important role for all three measured habitat characteristics in driving dispersal, survival, or fatal encounters with a predator in the first year after translocation. 5.Synthesis and applications. Resource managers using translocations as a conservation tool should prioritize acquiring data linking habitat to fitness. In particular, for species that depend on avoiding detection, refuges such as burrows and habitat that improved concealment had notable ability to improve survival and dispersal. Our study on juvenile Mojave desert tortoises showed that refuge availability or the distributions of habitat appropriate for concealment are important considerations for identifying translocation sites for species highly dependent on crypsis, camouflage, or other forms of habitat matching. This article is protected by copyright. All rights reserved.

Spatially explicit decision support for selecting translocation areas for Mojave desert tortoises

Spatially explicit decision support systems are assuming an increasing role in natural resource and conservation management. In order for these systems to be successful, however, they must address real-world management problems with input from both the scientific and management communities. The National Training Center at Fort Irwin, California, has expanded its training area, encroaching U.S. Fish and Wildlife Service critical habitat set aside for the Mojave desert tortoise (Gopherus agassizii), a federally threatened species. Of all the mitigation measures proposed to offset expansion, the most challenging to implement was the selection of areas most feasible for tortoise translocation. We developed an objective, open, scientifically defensible spatially explicit decision support system to evaluate translocation potential within the Western Mojave Recovery Unit for tortoise populations under imminent threat from military expansion. Using up to a total of 10 biological, anthropogenic, and/or logistical criteria, seven alternative translocation scenarios were developed. The final translocation model was a consensus model between the seven scenarios. Within the final model, six potential translocation areas were identified.