Global change impacts on cacti (Cactaceae): current threats, challenges and conservation solutions.

BACKGROUND: The plant family Cactaceae provides some of the most striking examples of adaptive evolution, expressing undeniably the most spectacular New World radiation of succulent plants distributed across arid and semi-arid regions of the Americas. Cacti are widely regarded for their cultural, economic and ecological value, yet they are also recognized as one of the most threatened and endangered taxonomic groups on the planet. SCOPE: This paper reviews current threats to species of cacti that have distributions in arid to semi-arid subtropical regions. Our review focuses primarily on four global change forces: (1) increases in atmospheric CO2 concentrations; (2) increases in mean annual temperatures and heat waves; (3) increases in the duration, frequency and intensity of droughts; and (4) and increases in competition and wildfire frequency from invasion by non-native species. We provide a broad range of potential priorities and solutions for stemming the extinction risk of cacti species and populations. CONCLUSIONS: Mitigating ongoing and emerging threats to cacti will require not only strong policy initiatives and international cooperation, but also new and creative approaches to conservation. These approaches include determining species at risk from climate extremes, enhancing habitat quality after disturbance, approaches and opportunities for ex situ conservation and restoration, and the potential use of forensic tools for identifying plants that have been removed illegally from the wild and sold on open markets.

Quantifying the immediate response of the soil microbial community to different grazing intensities on irrigated pastures.

Grazing is known to affect soil microbial communities, nutrient cycling, and forage quantity and quality over time. However, a paucity of information exists for the immediate changes in the soil physicochemical and microbial environment in response to different grazing strategies. Soil microbes drive nutrient cycling and are involved in plant-soil-microbe relationships, making them potentially vulnerable to plant-driven changes in the soil environment caused by grazing. To test the hypothesis that variable grazing intensities modulate immediate effects on the soil microbial community, we conducted a grazing trial of three management approaches; high-intensity, short-duration grazing (HDG), low-intensity, medium-duration grazing (LDG), and no grazing (NG). Soil and vegetation samples were collected before grazing and 24 hours, 1 week, and 4 weeks after HDG grazing ended. Soil labile carbon (C) and nitrogen (N) pools, vegetation biomass, and soil microbial diversity and functional traits were determined, including extracellular enzymatic assays and high-throughput sequencing of the bacterial 16S rRNA and fungal ITS2 regions. We found that labile soil C and inorganic N increased following the LDG grazing while C-cycling extracellular enzymatic activities increased in response to HDG grazing but both total extracellular enzymatic activity profiles and soil abiotic profiles were mostly affected by temporal fluxes. The soil fungal community composition was strongly affected by the interaction of sampling time and grazing treatment, while the soil bacterial community composition was largely affected by sampling time with a lesser impact from grazing treatment. We identified several key fungal taxa that may influence immediate responses to grazing and modulate plant-soil-microbe interactions. There was strong evidence of temporal influences on soil biogeochemical variables and the soil microbiome, even within our narrow sampling scheme. Our results indicate that the soil ecosystem is dynamic and responsive to different grazing strategies within very short time scales, showing the need for further research to understand plant-soil-microbe interactions and how these feedback mechanisms can inform sustainable land management.

Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

rKIN: Kernel-based method for estimating isotopic niche size and overlap.

The isotopic niche of consumers represents biologically relevant information on resource and habitat use. Several tools have been developed to quantify niche size and overlap. Nonetheless, methods adapted by spatial ecologists to quantify animal home ranges can be modified for use in stable isotope ecology when data are not normally distributed in bivariate space. We offer a tool that draws on existing spatial metrics, such as minimum convex polygon (MCP) and standard ellipse area (SEA), and add novel metrics using kernel utilization density (KUD) estimators to measure isotopic niche size and overlap. We present examples using empirical and simulated data to demonstrate the performance of the package kernel isotopic niches in r (rKIN) under various scenarios. Results of niche size from MCP, SEA and KUD were highly correlated but divergent among datasets. Overall, the KUD method produced the largest niche sizes and was more sensitive to the distribution of the isotopic data. Pairwise estimates of overlap were highly variable, likely because MCP and SEA inherently include or exclude unused areas in the resulting niche estimate. Four bandwidth methods (reference, normal scale, plug-in and biased cross-validation) produced comparable estimates of niche size and overlap at various sample sizes (10-40). Niche size and overlap were consistent across sample sizes >15. Use of rKIN will allow isotope ecologists to quantify niche shifts, expansions or contractions, as well as assess the performance of several estimation methods. The package also can be applied to other data types (e.g. principal component analysis, multi-dimensional scaling) so long as axes and measurement units are identical and can be converted to Cartesian coordinates.

Nitrogen deposition sources and patterns in the Greater Yellowstone Ecosystem determined from ion exchange resin collectors, lichens, and isotopes.

Over the past century, atmospheric nitrogen deposition (Ndep) has increased across the western United States due to agricultural and urban development, resulting in degraded ecosystem quality. Regional patterns of Ndep are often estimated by coupling direct measurements from large-scale monitoring networks and atmospheric chemistry models, but such efforts can be problematic in the western US because of complex terrain and sparse sampling. This study aimed not only to understand Ndep patterns in mountainous ecosystems but also to investigate whether isotope values of lichens and throughfall deposition can be used to determine Ndep sources, and serve as an additional tool in ecosystem health assessments. We measured Ndep amounts and δ15N in montane conifer forests of the Greater Yellowstone Ecosystem using canopy throughfall and bulk monitors and lichens. In addition, we examined patterns of C:N ratios in lichens as a possible indicator of lichen physiological condition. The isotopic signature of δ15N of Ndep helps to discern emission sources, because δ15N of NOx from combustion tends to be high (-5 to +25‰) while NHx from agricultural sources tends to be comparatively low (-40 to -10‰). Summertime Ndep increased with elevation and ranged from 0.26 to 1.66 kg ha-1. Ndep was higher than expected in remote areas. The δ15N values of lichens were typically -15.3 to -10‰ suggesting agriculture as a primary emission source of deposition. Lichen %N, δ15N and C:N ratios can provide important information about Ndep sources and patterns over small spatial scales in complex terrain.

Hydraulic and photosynthetic responses of big sagebrush to the 2017 total solar eclipse.

The total solar eclipse of August 21, 2017 created a path of totality ~115 km in width across the United States. While eclipse observations have shown distinct responses in animal behavior often emulating nocturnal behavior, the influence of eclipses on plant physiology are less understood. We investigated physiological perturbations due to rapid changes of sunlight and air temperature in big sagebrush (Artemisia tridentata ssp. vaseyana), a desert shrub common within the path of eclipse totality. Leaf gas exchange, water potential, and chlorophyll a fluorescence were monitored during the eclipse and compared to responses obtained the day before in absence of the eclipse. On the day of the eclipse, air temperature decreased by 6.4 °C, coupled with a 1.0 kPa drop in vapor pressure deficit having a 9-minute lag following totality. Using chlorophyll a fluorescence measurements, we found photosynthetic efficiency of photosystem II (Fv'/Fm') recovered to near dark acclimated state (i.e., 87%), but the short duration of darkness did not allow for complete recovery. Gas exchange data and a simple light response model were used to estimate a 14% reduction in carbon assimilation for one day over sagebrush dominated areas within the path of totality for the Western United States.

Microbiome Composition in Both Wild-Type and Disease Model Mice Is Heavily Influenced by Mouse Facility.

Murine models have become essential tools for understanding the complex interactions between gut microbes, their hosts, and disease. While many intra-facility factors are known to influence the structure of mouse microbiomes, the contribution of inter-facility variation to mouse microbiome composition, especially in the context of disease, remains under-investigated. We replicated microbiome experiments using identical mouse lines housed in two separate animal facilities and report drastic differences in composition of microbiomes based upon animal facility of origin. We observed facility-specific microbiome signatures in the context of a disease model [the Ednrb (endothelin receptor type B) Hirschsprung disease mouse] and in normal C57BL/6J mice. Importantly, these facility differences were independent of cage, sex, or sequencing-related influence. In addition, we investigated the reproducibility of microbiome dysbiosis previously associated with Ednrb-/- (knock-out; KO) mice. While we observed genotype-based differences in composition between wild-type (WT) and KO mice, these differences were inconsistent with the previously reported conclusions. Furthermore, the genotype-based differences were not identical across animal facilities. Despite this, through differential abundance testing, we identified several conserved candidate taxa and candidate operational taxonomic units that may play a role in disease promotion or protection. Overall, our findings raise the possibility that previously reported microbiome-disease associations from murine studies conducted in a single facility may be heavily influenced by facility-specific effects. More generally, these results provide a strong rationale for replication of mouse microbiome studies at multiple facilities, and for the meticulous collection of metadata that will allow the confounding effects of facility to be more specifically identified.

Identifying Holes in the Greater Sage-Grouse Conservation Umbrella.

The umbrella species concept, wherein multiple species are indirectly protected under the umbrella of a reserve created for one, is intended to enhance conservation efficiency. Although appealing in theory and common in practice, empirical tests of the concept have been scarce. We used a real-world, semi-protected reserve established to protect a high-profile umbrella species (greater sage-grouse [Centrocercus urophasianus]) to investigate 2 potential mechanisms underlying the concept's successful application: reserve size and species similarity. We estimated how much habitat protection the established reserve provided to 52 species of conservation concern associated with vegetation communities where greater sage-grouse occur. To illustrate the importance of reserve size, we compared the effectiveness of the established reserve to alternative greater sage-grouse reserves of various sizes and to simulated reserves of equal size but sited with no regard for greater sage-grouse. We further assessed whether key species' traits were associated with different levels of protection under the umbrella reserve. The established umbrella reserve protected 82% of the state's greater sage-grouse population and 0-63% of the habitat of the background species examined. The reserve outperformed equally sized, simulated reserves for only 12 of 52 background species. As expected, larger alternative reserves served as better umbrellas, but regardless of reserve size, not all species received equal protection. The established reserve was most effective at protecting the habitat of species that were most similar to the umbrella species (i.e., avian species, those highly associated with sagebrush plant communities, and those with widespread habitat). In contrast, the habitat of species with restricted distributions, particularly when combined with vegetation associations not closely matching the umbrella species, was not protected as well by the umbrella reserve. Such species require additional, targeted attention to achieve conservation objectives. Successful application of the umbrella species concept requires careful consideration of the characteristics of the umbrella species, the reserve delineated on its behalf, and the similarity of the umbrella species to its purported background species.

Increased Arctic sea ice drift alters adult female polar bear movements and energetics.

Recent reductions in thickness and extent have increased drift rates of Arctic sea ice. Increased ice drift could significantly affect the movements and the energy balance of polar bears (Ursus maritimus) which forage, nearly exclusively, on this substrate. We used radio-tracking and ice drift data to quantify the influence of increased drift on bear movements, and we modeled the consequences for energy demands of adult females in the Beaufort and Chukchi seas during two periods with different sea ice characteristics. Westward and northward drift of the sea ice used by polar bears in both regions increased between 1987-1998 and 1999-2013. To remain within their home ranges, polar bears responded to the higher westward ice drift with greater eastward movements, while their movements north in the spring and south in fall were frequently aided by ice motion. To compensate for more rapid westward ice drift in recent years, polar bears covered greater daily distances either by increasing their time spent active (7.6%-9.6%) or by increasing their travel speed (8.5%-8.9%). This increased their calculated annual energy expenditure by 1.8%-3.6% (depending on region and reproductive status), a cost that could be met by capturing an additional 1-3 seals/year. Polar bears selected similar habitats in both periods, indicating that faster drift did not alter habitat preferences. Compounding reduced foraging opportunities that result from habitat loss; changes in ice drift, and associated activity increases, likely exacerbate the physiological stress experienced by polar bears in a warming Arctic.

Modeling Behavior by Coastal River Otter (Lontra Canadensis) in Response to Prey Availability in Prince William Sound, Alaska: A Spatially-Explicit Individual-Based Approach.

Effects of climate change on animal behavior and cascading ecosystem responses are rarely evaluated. In coastal Alaska, social river otters (Lontra Canadensis), largely males, cooperatively forage on schooling fish and use latrine sites to communicate group associations and dominance. Conversely, solitary otters, mainly females, feed on intertidal-demersal fish and display mutual avoidance via scent marking. This behavioral variability creates "hotspots" of nutrient deposition and affects plant productivity and diversity on the terrestrial landscape. Because the abundance of schooling pelagic fish is predicted to decline with climate change, we developed a spatially-explicit individual-based model (IBM) of otter behavior and tested six scenarios based on potential shifts to distribution patterns of schooling fish. Emergent patterns from the IBM closely mimicked observed otter behavior and landscape use in the absence of explicit rules of intraspecific attraction or repulsion. Model results were most sensitive to rules regarding spatial memory and activity state following an encounter with a fish school. With declining availability of schooling fish, the number of social groups and the time simulated otters spent in the company of conspecifics declined. Concurrently, model results suggested an elevation of defecation rate, a 25% increase in nitrogen transport to the terrestrial landscape, and significant changes to the spatial distribution of "hotspots" with declines in schooling fish availability. However, reductions in availability of schooling fish could lead to declines in otter density over time.

Supplemental feeding alters migration of a temperate ungulate.

Conservation of migration requires information on behavior and environmental determinants. The spatial distribution of forage resources, which migration exploits, often are altered and may have subtle, unintended consequences. Supplemental feeding is a common management practice, particularly for ungulates in North America and Europe, and carryover effects on behavior of this anthropogenic manipulation of forage are expected in theory, but have received limited empirical evaluation, particularly regarding effects on migration. We used global positioning system (GPS) data to evaluate the influence of winter feeding on migration behavior of 219 adult female elk (Cervus elaphus) from 18 fed ranges and 4 unfed ranges in western Wyoming. Principal component analysis revealed that the migratory behavior of fed and unfed elk differed in distance migrated, and the timing of arrival to, duration on, and departure from summer range. Fed elk migrated 19.2 km less, spent 11 more days on stopover sites, arrived to summer range 5 days later, resided on summer range 26 fewer days, and departed in the autumn 10 days earlier than unfed elk. Time-to-event models indicated that differences in migratory behavior between fed and unfed elk were caused by altered sensitivity to the environmental drivers of migration. In spring, unfed elk migrated following plant green-up closely, whereas fed elk departed the feedground but lingered on transitional range, thereby delaying their arrival to summer range. In autumn, fed elk were more responsive to low temperatures and precipitation events, causing earlier departure from summer range than unfed elk. Overall, supplemental feeding disconnected migration by fed elk from spring green-up and decreased time spent on summer range, thereby reducing access to quality forage. Our findings suggest that ungulate migration can be substantially altered by changes to the spatial distribution of resources, including those of anthropogenic origin, and that management practices applied in one season may have unintended behavioral consequences in subsequent seasons.

Linking anti-predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore.

Ecological theory predicts that the diffuse risk cues generated by wide-ranging, active predators should induce prey behavioural responses but not major, population- or community-level consequences. We evaluated the non-consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high-risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20-fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.