Roadside habitat: Boon or bane for pollinating insects?

Pollinators, which provide vital services to wild ecosystems and agricultural crops, are facing global declines and habitat loss. As undeveloped land becomes increasingly scarce, much focus has been directed recently to roadsides as potential target zones for providing floral resources to pollinators. Roadsides, however, are risky places for pollinators, with threats from vehicle collisions, toxic pollutants, mowing, herbicides, and more. Although these threats have been investigated, most studies have yet to quantify the costs and benefits of roadsides to pollinators and, therefore, do not address whether the costs outweigh the benefits for pollinator populations using roadside habitats. In this article, we address how, when, and under what conditions roadside habitats may benefit or harm pollinators, reviewing existing knowledge and recommending practical questions that managers and policymakers should consider when planning pollinator-focused roadside management.

Using Regional Climate Projections to Guide Grassland Community Restoration in the Face of Climate Change.

Grassland loss has been extensive worldwide, endangering the associated biodiversity and human well-being that are both dependent on these ecosystems. Ecologists have developed approaches to restore grassland communities and many have been successful, particularly where soils are rich, precipitation is abundant, and seeds of native plant species can be obtained. However, climate change adds a new filter needed in planning grassland restoration efforts. Potential responses of species to future climate conditions must also be considered in planning for long-term resilience. We demonstrate this methodology using a site-specific model and a maximum entropy approach to predict changes in habitat suitability for 33 grassland plant species in the tallgrass prairie region of the U.S. using the Intergovernmental Panel on Climate Change scenarios A1B and A2. The A1B scenario predicts an increase in temperature from 1.4 to 6.4°C, whereas the A2 scenario predicts temperature increases from 2 to 5.4°C and much greater CO2 emissions than the A1B scenario. Both scenarios predict these changes to occur by the year 2100. Model projections for 2040 under the A1B scenario predict that all but three modeled species will lose ~90% of their suitable habitat. Then by 2080, all species except for one will lose ~90% of their suitable habitat. Models run using the A2 scenario predict declines in habitat for just four species by 2040, but models predict that by 2080, habitat suitability will decline for all species. The A2 scenario appears based on our results to be the less severe climate change scenario for our species. Our results demonstrate that many common species, including grasses, forbs, and shrubs, are sensitive to climate change. Thus, grassland restoration alternatives should be evaluated based upon the long-term viability in the context of climate change projections and risk of plant species loss.

Climate Extremes, Vegetation Change, and Decoupling of Interactive Fire-Grazing Processes Exacerbate Fly Parasitism of Cattle.

We assessed local horn fly (Haematobia irritans L.) and face fly (Musca autumnalis De Geer) communities on cattle in 2012 and 2013 relative to vegetation and climate data to understand how parasitism of cattle is influenced by change in climate and vegetation structure. We compared heterogeneity management using spatially and temporally discrete fires (i.e., patch-burning one-third of a pasture annually) to homogeneity management (i.e., burning entire pasture in 2012 then no burning in 2013), with cattle grazing all years in both treatments. Predicted emergence of horn flies and face flies was 24 and 34 d earlier in 2012 associated with earlier spring warming, a significant deviation from the five-year mean. Intraannual horn fly dynamics were explained by concurrent high ambient air temperature the day of observations, but face flies were explained by low ambient air temperatures and dry conditions 3 wk before observations. Importance values of information for the theoretic models including fire treatments ranged from 0.89 to 1, indicating that both horn flies and face flies are sensitive to habitat alterations and fire-driven animal movements. Ordination indicates herds on unburned pastures were dissimilar to herds on pastures burned with patchy fires or pastures burned completely and species-specific fly responses to different vegetation structure metrics. For example, horn flies were correlated with vegetation visual obstruction, and face flies were correlated with woody plant cover. Vegetation structure may be as important as climate in driving the dynamics of fly parasites of cattle.

Exotic-Dominated Grasslands Show Signs of Recovery with Cattle Grazing and Fire.

In grasslands, overgrazing by domestic livestock, fertilization, and introduction of exotic forage species leads to plant communities consisting of a mixture of native and exotic species. These degraded grasslands present a problem for land managers, farmers, and restoration ecologists concerned with improving biodiversity while continuing to use the land for livestock production. Here we assessed the response of butterfly and plant community composition to the use of fire and moderate grazing by domestic cattle on degraded grasslands dominated by exotic plants. We evaluated change by comparing experimental pastures to two reference sites that were grasslands dominated by native plants. We used two burning and grazing treatments: 1) patch-burn graze, a heterogeneously managed treatment, where one third of the pasture is burned each year and cattle have free access to the entire pasture, and 2) graze-and-burn, a homogenously managed treatment, where the entire pasture is grazed each year and burned in its entirety every three years. We tested for change in the butterfly and plant community composition over seven years using Bray-Curtis dissimilarity measures. Over the course of seven years, degraded pastures in both treatments became more similar to reference sites with respect to the butterfly and plant communities. Only two butterfly species and two plant functional guilds exhibited significant linear trends over time, with varying responses. Compositional changes in both the butterfly and plant communities indicate that the use of moderate grazing and fire may shift butterfly and plant communities of exotic-dominated grasslands to be more similar to reference tallgrass prairies over time.

Temporal variability in aboveground plant biomass decreases as spatial variability increases.

Ecological theory predicts that diversity decreases variability in ecosystem function. We predict that, at the landscape scale, spatial variability created by a mosaic of contrasting patches that differ in time since disturbance will decrease temporal variability in aboveground plant biomass. Using data from a multi-year study of seven grazed tallgrass prairie landscapes, each experimentally managed for one to eight patches, we show that increased spatial variability driven by spatially patchy fire and herbivory reduces temporal variability in aboveground plant biomass. This pattern is associated with statistical evidence for the portfolio effect and a positive relationship between temporal variability and functional group synchrony as predicted by metacommunity variability theory. As disturbance from fire and grazing interact to create a shifting mosaic of spatially heterogeneous patches within a landscape, temporal variability in aboveground plant biomass can be dampened. These results suggest that spatially heterogeneous disturbance regimes contribute to a portfolio of ecosystem functions provided by biodiversity, including wildlife habitat, fuel, and forage. We discuss how spatial patterns of disturbance drive variability within and among patches.

Bee Abundance and Nutritional Status in Relation to Grassland Management Practices in an Agricultural Landscape.

Grasslands provide important resources for pollinators in agricultural landscapes. Managing grasslands with fire and grazing has the potential to benefit plant and pollinator communities, though there is uncertainty about the ideal approach. We examined the relationships among burning and grazing regimes, plant communities, and Bombus species and Apis mellifera L. abundance and nutritional indicators at the Grand River Grasslands in southern Iowa and northern Missouri. Treatment regimes included burn-only, grazed-and-burned, and patch-burn graze (pastures subdivided into three temporally distinct fire patches with free access by cattle). The premise of the experimental design was that patch-burn grazing would increase habitat heterogeneity, thereby providing more diverse and abundant floral resources for pollinators. We predicted that both bee abundance and individual bee nutritional indicators (bee size and lipid content) would be positively correlated with floral resource abundance. There were no significant differences among treatments with respect to bee abundance. However, some of the specific characteristics of the plant community showed significant relationships with bee response variables. Pastures with greater abundance of floral resources had greater bee abundance but lower bee nutritional indicators. Bee nutritional variables were positively correlated with vegetation height, but, in some cases, negatively correlated with stocking rate. These results suggest grassland site characteristics such as floral resource abundance and stocking rate are of potential importance to bee pollinators and suggest avenues for further research to untangle the complex interactions between grassland management, plant responses, and bee health.

Effects of Tall Fescue and Its Fungal Endophyte on the Development and Survival of Tawny-Edged Skippers (Lepidoptera: Hesperiidae).

Invasive, exotic grasses are increasing in tallgrass prairie and their dominance may be contributing to the decline of grassland butterflies through alterations in forage quality. Tall fescue (Schedonorus arundinaceus (Schreb.) Dumort.), an exotic grass covering millions of acres in the United States, can host a fungal endophyte, Epichloë coenophiala (Morgan-Jones & Gams). Alkaloids produced by the endophyte are known to be toxic to some foliar-feeding pest insects. Endophyte-infected tall fescue is commonly planted in hayfields, pastures, lawns, and is invading natural areas, but effects of the endophyte on nonpest insects such as butterflies are relatively unknown. Our objective was to investigate the role that tall fescue and its endophyte might play in the decline of grass skippers (Hesperiidae). We examined growth and survival parameters of tawny-edged skippers (Polites themistocles (Latreille)) that were reared on endophyte-infected tall fescue (E+), endophyte-free tall fescue (E-), and Kentucky bluegrass (KBG). KBG was included as a comparison because it is a cool season grass known to be palatable to P. themistocles larvae. Interestingly, results showed that the endophyte did not affect growth and survival of larvae compared to uninfected tall fescue, even though significant amounts of loline alkaloids (average 740 ppm) were measured in endophyte-infected plant material. Larvae feeding on KBG grew faster with greater survival rates than larvae on both tall fescue treatments. These results confirm that tall fescue invasion and dominance may be deteriorating the quality of grassland habitats for native pollinators; however, this effect does not appear to be linked to endophyte infection.

Predator identity influences the effect of habitat management on nest predation.

Predation is the leading cause of nest failure for many passerines and considerable effort is devoted to identifying the habitat characteristics and management practices that influence nest loss. The habitat components associated with nest loss are strongly influenced by the ecology of nest predators and differ among predator species as a result. Nevertheless, there is a tendency to generalize about the effects of habitat features and management on nest failure without considering how resulting patterns are influenced by nest predators. We examined how predator-specific patterns of nest loss differed among predators and in response to grassland management with fire and grazing by cattle (Bos taurus). We used video cameras to monitor and identify predators at nests of the Grasshopper Sparrow (Ammodramus savannarum), a species of conservation concern throughout its range. We observed predation by 15 different species that differed in their response to management and the habitat characteristics associated with nests they preyed on. Losses to mammals and snakes were more likely at nests with greater amounts of litter cover and tall fescue (Schedonorus phoenix). Mammals were less likely to prey on nests surrounded by greater forb cover. Nest predation by snakes was lower in burned areas, whereas predation by mammals and Brown-headed Cowbirds (Molothrus ater) was unaffected by the use of fire. Neither vegetation density at the nest, nor landscape context was related to nest loss by any predator taxon. Although there were many similarities, we identified important differences in the species composing the nest predator community between our. study and other published research. These differences are likely to be responsible for geographic variation in the influence of habitat features and management actions on nest success. Our results demonstrate the need for natural resource managers to incorporate knowledge of local nest predators and their ecology when developing management prescriptions aimed at enhancing the reproductive success of songbirds.

Connecting soil organic carbon and root biomass with land-use and vegetation in temperate grassland.

Soils contain much of Earth's terrestrial organic carbon but are sensitive to land-use. Rangelands are important to carbon dynamics and are among ecosystems most widely impacted by land-use. While common practices like grazing, fire, and tillage affect soil properties directly related to soil carbon dynamics, their magnitude and direction of change vary among ecosystems and with intensity of disturbance. We describe variability in soil organic carbon (SOC) and root biomass--sampled from 0-170 cm and 0-100 cm, respectively--in terms of soil properties, land-use history, current management, and plant community composition using linear regression and multivariate ordination. Despite consistency in average values of SOC and root biomass between our data and data from rangelands worldwide, broad ranges in root biomass and SOC in our data suggest these variables are affected by other site-specific factors. Pastures with a recent history of severe grazing had reduced root biomass and greater bulk density. Ordination suggests greater exotic species richness is associated with lower root biomass but the relationship was not apparent when an invasive species of management concern was specifically tested. We discuss how unexplained variability in belowground properties can complicate measurement and prediction of ecosystem processes such as carbon sequestration.

Fire induced reproductive mechanisms of a Symphoricarpos (Caprifoliaceae) shrub after dormant season burning.

BACKGROUND: Symphoricarpos, a genus of the Caprifoliaceae family, consists of about 15 species of clonal deciduous shrubs in North America and 1 species endemic to China. In North American tallgrass prairie, Symphoricarpos orbiculatus (buckbrush) is the dominant shrub often forming large colonies via sexual and asexual reproductive mechanisms. Symphoricarpos shrubs, in particular S. orbiculatus, use a unique sexual reproductive mechanism known as layering where vertical stems droop and the tips root upon contact with the soil. Because of conflicting societal values of S. orbiculatus for conservation and agriculture and the current attempt to restore historical fire regimes, there is a need for basic research on the biological response of S. orbiculatus to anthropogenic burning regimes. RESULTS: From 2007 through 2013 we applied prescribed fires in the late dormant season on grazed pastures in the Grand River Grasslands of Iowa. From 2011 to 2013, we measured how S. orbiculatus basal resprouting and layering stems were affected by patchy fires on grazed pastures, complete pasture fires on grazed pastures or fire exclusion without grazing for more than three years. We measured ramet height, ramet canopy diameter, stems per ramet, ramets per 100 m2, and probability of new layering stems 120 days after fire. Height in burned plots was lower than unburned plots but S. orbiculatus reached ~ 84% of pre-burn height 120 days after fire. Stems per ramet were 2x greater in the most recently burned plots due to basal re-sprouting. Canopy diameter and density of ramets was not affected by time since fire, but burned pastures had marginally lower densities than plots excluded from fire (P = 0.07). Fire triggered new layering stems and no new layering stems were found in plots excluded from fire. CONCLUSIONS: The mechanisms of both basal sprouting and aerial layering after fire suggest S. orbiculatus is tolerant to dormant season fires. Furthermore, dormant season fires, regardless if they were patchy fires or complete pasture fires, did not result in mortality of S. orbiculatus. Dormant season fires can reduce S. orbiculatus structural dominance and maintain lower ramet densities but also trigger basal resprouting and layering.

Gradient-based habitat affinities predict species vulnerability to drought.

Ecological fingerprints of climate change are becoming increasingly evident at broad geographical scales as measured by species range shifts and changes in phenology. However, finer-scale species-level responses to environmental fluctuations may also provide an important bellwether of impending future community responses. Here we examined changes in abundance of butterfly species along a hydrological gradient of six montane meadow habitat types in response to drought. Our data collection began prior to the drought, and we were able to track changes for 11 years, of which eight were considered mild to extreme drought conditions. We separated the species into those that had an affinity for hydric vs. xeric habitats. We suspected that drought would favor species with xeric habitat affinities, but that there could be variations in species-level responses along the hydrological gradient. We also suspected that mesic meadows would be most sensitive to drought conditions. Temporal trajectories were modeled for both species groups (hydric vs. xeric affinity) and individual species. Abundances of species with affinity for xeric habitats increased in virtually all meadow types. Conversely, abundances of species with affinity for hydric habitats decreased, particularly in mesic and xeric meadows. Mesic meadows showed the most striking temporal abundance trajectory: Increasing abundances of species with xeric habitat affinity were offset by decreasing or stable abundances of species with hydric habitat affinity. The one counterintuitive finding was that, in some hydric meadows, species with affinity for hydric habitats increased. In these cases, we suspect that decreasing moisture conditions in hydric meadows actually increased habitat suitability because sites near the limit of moisture extremes for some species became more acceptable. Thus, species responses were relatively predictable based upon habitat affinity and habitat location along the hydrological gradient, and mesic meadows showed the highest potential for changes in community composition. The implications of these results are that longer-term changes due to drought could simplify community composition, resulting in prevalence of species tolerant to drying conditions and a loss of species associated with wetter conditions. We contend that this application of gradient analysis could be valuable in assessing species vulnerability of other taxa and ecosystems.

Occurrence of sarmentosin and other hydroxynitrile glucosides in Parnassius (papilionidae) butterflies and their food plants.

Sequestration of plant secondary metabolites is a widespread phenomenon among aposematic insects. Sarmentosin is an unsaturated γ-hydroxynitrile glucoside known from plants and some Lepidoptera. It is structurally and biosynthetically closely related to cyanogenic glucosides, which are commonly sequestered from food plants and/or de novo synthesized by lepidopteran species. Sarmentosin was found previously in Parnassius (Papilionidae) butterflies, but it was not known how the occurrence was related to food plants or whether Parnassius species could biosynthesize the compound. Here, we report on the occurrence of sarmentosin and related compounds in four different Parnassius species belonging to two different clades, as well as their known and suspected food plants. There were dramatic differences between the two clades, with P. apollo and P. smintheus from the Apollo group containing high amounts of sarmentosin, and P. clodius and P. mnemosyne from the Mnemosyne group containing low or no detectable amounts. This was reflected in the larval food plants; P. apollo and P. smintheus larvae feed on Sedum species (Crassulaceae), which all contained considerable amounts of sarmentosin, while the known food plants of the two other species, Dicentra and Corydalis (Fumariaceae), had no detectable levels of sarmentosin. All insects and plants containing sarmentosin also contained other biosynthetically related hydroxynitrile glucosides in patterns previously reported for plants, but not for insects. Not all findings could be explained by sequestration alone and we therefore hypothesize that Parnassius species are able to de novo synthesize sarmentosin.

Moving forward in global-change ecology: capitalizing on natural variability.

Natural resources managers are being asked to follow practices that accommodate for the impact of climate change on the ecosystems they manage, while global-ecosystems modelers aim to forecast future responses under different climate scenarios. However, the lack of scientific knowledge about short-term ecosystem responses to climate change has made it difficult to define set conservation practices or to realistically inform ecosystem models. Until recently, the main goal for ecologists was to study the composition and structure of communities and their implications for ecosystem function, but due to the probable magnitude and irreversibility of climate-change effects (species extinctions and loss of ecosystem function), a shorter term focus on responses of ecosystems to climate change is needed. We highlight several underutilized approaches for studying the ecological consequences of climate change that capitalize on the natural variability of the climate system at different temporal and spatial scales. For example, studying organismal responses to extreme climatic events can inform about the resilience of populations to global warming and contribute to the assessment of local extinctions. Translocation experiments and gene expression are particular useful to quantitate a species' acclimation potential to global warming. And studies along environmental gradients can guide habitat restoration and protection programs by identifying vulnerable species and sites. These approaches identify the processes and mechanisms underlying species acclimation to changing conditions, combine different analytical approaches, and can be used to improve forecasts of the short-term impacts of climate change and thus inform conservation practices and ecosystem models in a meaningful way.

Montane meadow change during drought varies with background hydrologic regime and plant functional group.

Climate change models for many ecosystems predict more extreme climatic events in the future, including exacerbated drought conditions. Here we assess the effects of drought by quantifying temporal variation in community composition of a complex montane meadow landscape characterized by a hydrological gradient. The meadows occur in two regions of the Greater Yellowstone Ecosystem (Gallatin and Teton) and were classified into six categories (M1-M6, designating hydric to xeric) based upon Satellite pour l'Observation de la Terre (SPOT) satellite imagery. Both regions have similar plant communities, but patch sizes of meadows are much smaller in the Gallatin region. We measured changes in the percent cover of bare ground and plants by species and functional groups during five years between 1997 and 2007. We hypothesized that drought effects would not be manifested evenly across the hydrological gradient, but rather would be observed as hotspots of change in some areas and minimally evident in others. We also expected varying responses by plant functional groups (forbs vs. woody plants). Forbs, which typically use water from relatively shallow s,oils compared to woody plants, were expected to decrease in cover in mesic meadows, but increase in hydric meadows. Woody plants, such as Artemisia, were expected to increase, especially in mesic meadows. We identified several important trends in our meadow plant communities during this period of drought: (1) bare ground increased significantly in xeric meadows of both regions (Gallatin M6 and Teton M5) and in mesic (M3) meadows of the Teton, (2) forbs decreased significantly in the mesic and xeric meadows in both regions, (3) forbs increased in hydric (M1) meadows of the Gallatin region, and (4) woody species showed increases in M2 and M5 meadows of the Teton region and in M3 meadows of the Gallatin region. The woody response was dominated by changes in Artemisia spp. and Chrysothamnus viscidiflorus. Thus, our results supported our expectations that community change was not uniform across the landscape, but instead could be predicted based upon functional group responses to the spatial and temporal patterns of water availability, which are largely a function of plant water use and the hydrological gradient.

Life-history traits predict species responses to habitat area and isolation: a cross-continental synthesis.

There is a lack of quantitative syntheses of fragmentation effects across species and biogeographic regions, especially with respect to species life-history traits. We used data from 24 independent studies of butterflies and moths from a wide range of habitats and landscapes in Europe and North America to test whether traits associated with dispersal capacity, niche breadth and reproductive rate modify the effect of habitat fragmentation on species richness. Overall, species richness increased with habitat patch area and connectivity. Life-history traits improved the explanatory power of the statistical models considerably and modified the butterfly species-area relationship. Species with low mobility, a narrow feeding niche and low reproduction were most strongly affected by habitat loss. This demonstrates the importance of considering life-history traits in fragmentation studies and implies that both species richness and composition change in a predictable manner with habitat loss and fragmentation.

Reproductive asynchrony in natural butterfly populations and its consequences for female matelessness.

1. Reproductive asynchrony, where individuals in a population are short-lived relative to the population-level reproductive period, has been identified recently as a theoretical mechanism of the Allee effect that could operate in diverse plant and insect species. The degree to which this effect impinges on the growth potential of natural populations is not yet well understood. 2. Building on previous models of reproductive timing, we develop a general framework that allows a detailed, quantitative examination of the reproductive potential lost to asynchrony in small natural populations. 3. Our framework includes a range of biologically plausible submodels that allow details of mating biology of different species to be incorporated into the basic reproductive timing model. 4. We tailor the parameter estimation methods of the full model (basic model plus mating biology submodels) to take full advantage of data from detailed field studies of two species of Parnassius butterflies whose mating status may be assessed easily in the field. 5. We demonstrate that for both species, a substantial portion of the female population (6.5-18.6%) is expected to die unmated. These analyses provide the first direct, quantitative evidence of female reproductive failure due to asynchrony in small natural populations, and suggest that reproductive asynchrony exerts a strong and largely unappreciated influence on the population dynamics of these butterflies and other species with similarly asynchronous reproductive phenology.

Quantifying relationships between bird and butterfly community shifts and environmental change.

Quantifying the manner in which ecological communities respond during a time of decreasing precipitation is a first step in understanding how they will respond to longer-term climate change. Here we coupled analysis of interannual variability in remotely sensed data with analyses of bird and butterfly community changes in montane meadow communities of the Greater Yellowstone Ecosystem. Landsat satellite imagery was used to classify these meadows into six types along a hydrological gradient. The northern portion of the ecosystem, or Gallatin region, has smaller mean patch sizes separated by ridges of mountains, whereas the southern portion of the ecosystem, or Teton region, has much larger patches within the Jackson Hole valley. Both support a similar suite of butterfly and bird species. The Gallatin region showed more overall among-year variation in the normalized difference vegetation index (NDVI) when meadow types were pooled within regions, perhaps because the patch sizes are smaller on average. Bird and butterfly communities showed significant relationships relative to meadow type and NDVI. We identified several key species that are tightly associated with specific meadow types along the hydrological gradient. Comparing taxonomic groups, fewer birds showed specific habitat affinities than butterflies, perhaps because birds are responding to differences in habitat structure among meadow types and using the landscape at a coarser scale than the butterflies. Comparing regions, the Teton region showed higher predictability of community assemblages as compared to the Gallatin region. The Gallatin region exhibited more significant temporal trends with respect to butterflies. Butterfly communities in wet meadows showed a distinctive shift along the hydrological gradient during a drought period (1997-2000). These results imply that the larger Teton meadows will show more predictable (i.e., static) species-habitat associations over the long term, but that the smaller Gallatin meadows may be an area that will exhibit the effects of global climate change faster.