Can flowers affect land surface albedo and soil microclimates?

The phenology of vegetation, namely leaf-out and senescence, can influence the Earth's climate over regional spatial scales and long time periods (e.g., over 30 years or more), in addition to microclimates over local spatial scales and shorter time periods (weeks to months). However, the effects of flowers on climate and microclimate are unknown. We investigate whether flowers can influence light reflected by the land surface and soil microclimate in a subalpine meadow. We conducted a flower removal experiment with a common sunflower species, Helianthella quinquenervis, for 3 years (2015, 2017, and 2019). The flower removal treatment simulates the appearance of the meadow when Helianthella flowers earlier under climate change and loses its flowers to frost (other plant structures are not damaged by frost). We test the hypotheses that a reduction in cover of yellow flowers leads to a greener land surface, lower reflectance, warmer and drier soils, and increased plant water stress. Flower removal plots are greener, reflect less light, exhibit up to 1.2 °C warmer soil temperatures during the warmest daylight hours, and contain ca. 1% less soil moisture compared to controls. However, soils were warmer in only 2 of the 3 years, when flower abundance was high. Helianthella water use efficiency did not differ between removal and control plots. Our study provides evidence for a previously undocumented effect of flowers on soil microclimate, an effect that is likely mediated by climate change and flowering phenology. Many anthropogenic environmental changes alter landscape albedo, all of which could be mediated by flowers: climate change, plant invasions, and agriculture. This study highlights how further consideration of the effects of flowers on land surface albedo could improve our understanding of the effects of vegetation on microclimate.

Diverse values, philosophies and ideas beget innovation and resilience in ecology and for our world.

Intellectual diversity, which is often assessed through social diversity proxies, such as race, ethnicity, and gender, is essential to innovation in ecology. There are many diverse ways of knowing, valuing, and discerning alternatives within ecology and as it is applied to solve global-change issues. However, science is a cultural activity, one that is affected by colonialism, racism, and White supremacy and, like other fields of science, the pursuit of ecological careers has historically been a space of limited opportunities for participation for Black, Indigenous, and People of Color (BIPOC)-narrowing the diverse ways of knowing, valuing, and engaging in ecological work. We seek to debunk the view that such limited participation is a result of BIPOC lacking interest in or proficiency in science, and instead offer that BIPOC communities are places of deep scientific understanding, sociocultural resilience, and cultural wealth. Scientists must broaden their consideration of the ecological sciences and work with BIPOC to establish community-based scientific partnerships that will foster increased ecological career pathways for BIPOC youth.

Experimental warming differentially affects vegetative and reproductive phenology of tundra plants.

Rapid climate warming is altering Arctic and alpine tundra ecosystem structure and function, including shifts in plant phenology. While the advancement of green up and flowering are well-documented, it remains unclear whether all phenophases, particularly those later in the season, will shift in unison or respond divergently to warming. Here, we present the largest synthesis to our knowledge of experimental warming effects on tundra plant phenology from the International Tundra Experiment. We examine the effect of warming on a suite of season-wide plant phenophases. Results challenge the expectation that all phenophases will advance in unison to warming. Instead, we find that experimental warming caused: (1) larger phenological shifts in reproductive versus vegetative phenophases and (2) advanced reproductive phenophases and green up but delayed leaf senescence which translated to a lengthening of the growing season by approximately 3%. Patterns were consistent across sites, plant species and over time. The advancement of reproductive seasons and lengthening of growing seasons may have significant consequences for trophic interactions and ecosystem function across the tundra.

The Snowmelt Niche Differentiates Three Microbial Life Strategies That Influence Soil Nitrogen Availability During and After Winter.

Soil microbial biomass can reach its annual maximum pool size beneath the winter snowpack and is known to decline abruptly following snowmelt in seasonally snow-covered ecosystems. Observed differences in winter versus summer microbial taxonomic composition also suggests that phylogenetically conserved traits may permit winter- versus summer-adapted microorganisms to occupy distinct niches. In this study, we sought to identify archaea, bacteria, and fungi that are associated with the soil microbial bloom overwinter and the subsequent biomass collapse following snowmelt at a high-altitude watershed in central Colorado, United States. Archaea, bacteria, and fungi were categorized into three life strategies (Winter-Adapted, Snowmelt-Specialist, Spring-Adapted) based upon changes in abundance during winter, the snowmelt period, and after snowmelt in spring. We calculated indices of phylogenetic relatedness (archaea and bacteria) or assigned functional attributes (fungi) to organisms within life strategies to infer whether phylogenetically conserved traits differentiate Winter-Adapted, Snowmelt-Specialist, and Spring-Adapted groups. We observed that the soil microbial bloom was correlated in time with a pulse of snowmelt infiltration, which commenced 65 days prior to soils becoming snow-free. A pulse of nitrogen (N, as nitrate) occurred after snowmelt, along with a collapse in the microbial biomass pool size, and an increased abundance of nitrifying archaea and bacteria (e.g., Thaumarchaeota, Nitrospirae). Winter- and Spring-Adapted archaea and bacteria were phylogenetically clustered, suggesting that phylogenetically conserved traits allow Winter- and Spring-Adapted archaea and bacteria to occupy distinct niches. In contrast, Snowmelt-Specialist archaea and bacteria were phylogenetically overdispersed, suggesting that the key mechanism(s) of the microbial biomass crash are likely to be density-dependent (e.g., trophic interactions, competitive exclusion) and affect organisms across a broad phylogenetic spectrum. Saprotrophic fungi were the dominant functional group across fungal life strategies, however, ectomycorrhizal fungi experienced a large increase in abundance in spring. If well-coupled plant-mycorrhizal phenology currently buffers ecosystem N losses in spring, then changes in snowmelt timing may alter ecosystem N retention potential. Overall, we observed that snowmelt separates three distinct soil niches that are occupied by ecologically distinct groups of microorganisms. This ecological differentiation is of biogeochemical importance, particularly with respect to the mobilization of nitrogen during winter, before and after snowmelt.

Accelerated Snowmelt Protocol to Simulate Climate Change Induced Impacts on Snowpack Dependent Ecosystems.

Field studies that simulate the effects of climate change are important for a predictive understanding of ecosystem responses to a changing environment. Among many concerns, regional warming can result in advanced timing of spring snowmelt in snowpack dependent ecosystems, which could lead to longer snow-free periods and drier summer soils. Past studies investigating these impacts of climate change have manipulated snowmelt with a variety of techniques that include manual snowpack alteration with a shovel, infrared radiation, black sand and fabric covers. Within these studies however, sufficient documentation of methods is limited, which can make experimental reproduction difficult. Here, we outline a detailed plot-scale protocol that utilizes a permeable black geotextile fabric deployed on top of an isothermal spring snowpack to induce advanced snowmelt. The method offers a reliable and cost-effective approach to induce snowmelt by passively increasing solar radiation absorption at the snow surface. In addition, control configurations with no snowpack manipulation are paired adjacent to the induced snowmelt plot for experimental comparison. Past and ongoing deployments in Colorado subalpine ecosystems indicate that this approach can accelerate snowmelt by 14-23 days, effectively mimicking snowmelt timing at lower elevations. This protocol can be applied to a variety of studies to understand the hydrological, ecological, and geochemical impacts of regional warming in snowpack dependent ecosystems.

Limited effects of early snowmelt on plants, decomposers, and soil nutrients in Arctic tundra soils.

In addition to warming temperatures, Arctic ecosystems are responding to climate change with earlier snowmelt and soil thaw. Earlier snowmelt has been examined infrequently in field experiments, and we lack a comprehensive look at belowground responses of the soil biogeochemical system that includes plant roots, decomposers, and soil nutrients. We experimentally advanced the timing of snowmelt in factorial combination with an open-top chamber warming treatment over a 3-year period and evaluated the responses of decomposers and nutrient cycling processes. We tested two alternative hypotheses: (a) Early snowmelt and warming advance the timing of root growth and nutrient uptake, altering the timing of microbial and invertebrate activity and key nutrient cycling events; and (b) loss of insulating snow cover damages plants, leading to reductions in root growth and altered biological activity. During the 3 years of our study (2010-2012), we advanced snowmelt by 4, 15, and 10 days, respectively. Despite advancing aboveground plant phenology, particularly in the year with the warmest early-season temperatures (2012), belowground effects were primarily seen only on the first sampling date of the season or restricted to particular years or soil type. Overall, consistent and substantial responses to early snowmelt were not observed, counter to both of our hypotheses. The data on soil physical conditions, as well interannual comparisons of our results, suggest that this limited response was because of the earlier date of snowmelt that did not coincide with substantially warmer air and soil temperatures as they might in response to a natural climate event. We conclude that the interaction of snowmelt timing with soil temperatures is important to how the ecosystem will respond, but that 1- to 2-week changes in timing of snowmelt alone are not enough to drive season-long changes in soil microbial and nutrient cycling processes.

Highly individualistic rates of plant phenological advance associated with arctic sea ice dynamics.

We analysed 12 years of species-specific emergence dates of plants at a Low-Arctic site near Kangerlussuaq, Greenland to investigate associations with sea ice dynamics, a potential contributor to local temperature variation in near-coastal tundra. Species displayed highly variable rates of phenological advance, from a maximum of -2.55 ± 0.17 and -2.93 ± 0.51 d yr-1 among a graminoid and forb, respectively, to a minimum of -0.55 ± 0.19 d yr-1 or no advance at all in the two deciduous shrub species. Monthly Arctic-wide sea ice extent was a significant predictor of emergence timing in 10 of 14 species. Despite variation in rates of advance among species, these rates were generally greatest in the earliest emerging species, for which monthly sea ice extent was also the primary predictor of emergence. Variation among species in rates of phenological advance reshuffled the phenological community, with deciduous shrubs leafing out progressively later relative to forbs and graminoids. Because early species advanced more rapidly than late species, and because rates of advance were greatest in species for which emergence phenology was associated with sea ice dynamics, accelerating sea ice decline may contribute to further divergence between early- and late-emerging species in this community.

Five years of phenology observations from a mixed-grass prairie exposed to warming and elevated CO2.

Atmospheric CO2 concentrations have been steadily increasing since the Industrial Era and contribute to concurrent increases in global temperatures. Many observational studies suggest climate warming alone contributes to a longer growing season. To determine the relative effect of warming on plant phenology, we investigated the individual and joint effects of warming and CO2 enrichment on a mixed-grass prairie plant community by following the development of six common grassland species and recording four major life history events. Our data support that, in a semi-arid system, while warming advances leaf emergence and flower production, it also expedites seed maturation and senescence at the species level. However, the additive effect can be an overall lengthening of the growing and reproductive seasons since CO2 enrichment, particularly when combined with warming, contributed to a longer growing season by delaying plant maturation and senescence. Fostering synthesis across multiple phenology datasets and identifying key factors affecting plant phenology will be vital for understanding regional plant community responses to climate change.

Earlier snowmelt and warming lead to earlier but not necessarily more plant growth.

Climate change over the past ∼50 years has resulted in earlier occurrence of plant life-cycle events for many species. Across temperate, boreal and polar latitudes, earlier seasonal warming is considered the key mechanism leading to earlier leaf expansion and growth. Yet, in seasonally snow-covered ecosystems, the timing of spring plant growth may also be cued by snowmelt, which may occur earlier in a warmer climate. Multiple environmental cues protect plants from growing too early, but to understand how climate change will alter the timing and magnitude of plant growth, experiments need to independently manipulate temperature and snowmelt. Here, we demonstrate that altered seasonality through experimental warming and earlier snowmelt led to earlier plant growth, but the aboveground production response varied among plant functional groups. Earlier snowmelt without warming led to early leaf emergence, but often slowed the rate of leaf expansion and had limited effects on aboveground production. Experimental warming alone had small and inconsistent effects on aboveground phenology, while the effect of the combined treatment resembled that of early snowmelt alone. Experimental warming led to greater aboveground production among the graminoids, limited changes among deciduous shrubs and decreased production in one of the dominant evergreen shrubs. As a result, we predict that early onset of the growing season may favour early growing plant species, even those that do not shift the timing of leaf expansion.

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake.

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub-dominated and evergreen/graminoid-dominated community-level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant-community-specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra-specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10-day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.

Elevated CO2 further lengthens growing season under warming conditions.

Observations of a longer growing season through earlier plant growth in temperate to polar regions have been thought to be a response to climate warming. However, data from experimental warming studies indicate that many species that initiate leaf growth and flowering earlier also reach seed maturation and senesce earlier, shortening their active and reproductive periods. A conceptual model to explain this apparent contradiction, and an analysis of the effect of elevated CO2--which can delay annual life cycle events--on changing season length, have not been tested. Here we show that experimental warming in a temperate grassland led to a longer growing season through earlier leaf emergence by the first species to leaf, often a grass, and constant or delayed senescence by other species that were the last to senesce, supporting the conceptual model. Elevated CO2 further extended growing, but not reproductive, season length in the warmed grassland by conserving water, which enabled most species to remain active longer. Our results suggest that a longer growing season, especially in years or biomes where water is a limiting factor, is not due to warming alone, but also to higher atmospheric CO2 concentrations that extend the active period of plant annual life cycles.

Predicted responses of arctic and alpine ecosystems to altered seasonality under climate change.

Global climate change is already having significant impacts on arctic and alpine ecosystems, and ongoing increases in temperature and altered precipitation patterns will affect the strong seasonal patterns that characterize these temperature-limited systems. The length of the potential growing season in these tundra environments is increasing due to warmer temperatures and earlier spring snow melt. Here, we compare current and projected climate and ecological data from 20 Northern Hemisphere sites to identify how seasonal changes in the physical environment due to climate change will alter the seasonality of arctic and alpine ecosystems. We find that although arctic and alpine ecosystems appear similar under historical climate conditions, climate change will lead to divergent responses, particularly in the spring and fall shoulder seasons. As seasonality changes in the Arctic, plants will advance the timing of spring phenological events, which could increase plant nutrient uptake, production, and ecosystem carbon (C) gain. In alpine regions, photoperiod will constrain spring plant phenology, limiting the extent to which the growing season can lengthen, especially if decreased water availability from earlier snow melt and warmer summer temperatures lead to earlier senescence. The result could be a shorter growing season with decreased production and increased nutrient loss. These contrasting alpine and arctic ecosystem responses will have cascading effects on ecosystems, affecting community structure, biotic interactions, and biogeochemistry.

Complex carbon cycle responses to multi-level warming and supplemental summer rain in the high Arctic.

The Arctic has experienced rapid warming and, although there are uncertainties, increases in precipitation are projected to accompany future warming. Climate changes are expected to affect magnitudes of gross ecosystem photosynthesis (GEP), ecosystem respiration (ER) and the net ecosystem exchange of CO2 (NEE). Furthermore, ecosystem responses to climate change are likely to be characterized by nonlinearities, thresholds and interactions among system components and the driving variables. These complex interactions increase the difficulty of predicting responses to climate change and necessitate the use of manipulative experiments. In 2003, we established a long-term, multi-level and multi-factor climate change experiment in a polar semidesert in northwest Greenland. Two levels of heating (30 and 60 W m(-2) ) were applied and the higher level was combined with supplemental summer rain. We made plot-level measurements of CO2 exchange, plant community composition, foliar nitrogen concentrations, leaf δ(13) C and NDVI to examine responses to our treatments at ecosystem- and leaf-levels. We confronted simple models of GEP and ER with our data to test hypotheses regarding key drivers of CO2 exchange and to estimate growing season CO2 -C budgets. Low-level warming increased the magnitude of the ecosystem C sink. Meanwhile, high-level warming made the ecosystem a source of C to the atmosphere. When high-level warming was combined with increased summer rain, the ecosystem became a C sink of magnitude similar to that observed under low-level warming. Competition among our ER models revealed the importance of soil moisture as a driving variable, likely through its effects on microbial activity and nutrient cycling. Measurements of community composition and proxies for leaf-level physiology suggest GEP responses largely reflect changes in leaf area of Salix arctica, rather than changes in leaf-level physiology. Our findings indicate that the sign and magnitude of the future High Arctic C budget may depend upon changes in summer rain.

Biological consequences of earlier snowmelt from desert dust deposition in alpine landscapes.

Dust deposition to mountain snow cover, which has increased since the late 19(th) century, accelerates the rate of snowmelt by increasing the solar radiation absorbed by the snowpack. Snowmelt occurs earlier, but is decoupled from seasonal warming. Climate warming advances the timing of snowmelt and early season phenological events (e.g., the onset of greening and flowering); however, earlier snowmelt without warmer temperatures may have a different effect on phenology. Here, we report the results of a set of snowmelt manipulations in which radiation-absorbing fabric and the addition and removal of dust from the surface of the snowpack advanced or delayed snowmelt in the alpine tundra. These changes in the timing of snowmelt were superimposed on a system where the timing of snowmelt varies with topography and has been affected by increased dust loading. At the community level, phenology exhibited a threshold response to the timing of snowmelt. Greening and flowering were delayed before seasonal warming, after which there was a linear relationship between the date of snowmelt and the timing of phenological events. Consequently, the effects of earlier snowmelt on phenology differed in relation to topography, which resulted in increasing synchronicity in phenology across the alpine landscape with increasingly earlier snowmelt. The consequences of earlier snowmelt from increased dust deposition differ from climate warming and include delayed phenology, leading to synchronized growth and flowering across the landscape and the opportunity for altered species interactions, landscape-scale gene flow via pollination, and nutrient cycling.

Tree species traits influence soil physical, chemical, and biological properties in high elevation forests.

BACKGROUND: Previous studies have shown that plants often have species-specific effects on soil properties. In high elevation forests in the Southern Rocky Mountains, North America, areas that are dominated by a single tree species are often adjacent to areas dominated by another tree species. Here, we assessed soil properties beneath adjacent stands of trembling aspen, lodgepole pine, and Engelmann spruce, which are dominant tree species in this region and are distributed widely in North America. We hypothesized that soil properties would differ among stands dominated by different tree species and expected that aspen stands would have higher soil temperatures due to their open structure, which, combined with higher quality litter, would result in increased soil respiration rates, nitrogen availability, and microbial biomass, and differences in soil faunal community composition. METHODOLOGY/PRINCIPAL FINDINGS: We assessed soil physical, chemical, and biological properties at four sites where stands of aspen, pine, and spruce occurred in close proximity to one-another in the San Juan Mountains, Colorado. Leaf litter quality differed among the tree species, with the highest nitrogen (N) concentration and lowest lignin:N in aspen litter. Nitrogen concentration was similar in pine and spruce litter, but lignin:N was highest in pine litter. Soil temperature and moisture were highest in aspen stands, which, in combination with higher litter quality, probably contributed to faster soil respiration rates from stands of aspen. Soil carbon and N content, ammonium concentration, and microbial biomass did not differ among tree species, but nitrate concentration was highest in aspen soil and lowest in spruce soil. In addition, soil fungal, bacterial, and nematode community composition and rotifer, collembolan, and mesostigmatid mite abundance differed among the tree species, while the total abundance of nematodes, tardigrades, oribatid mites, and prostigmatid mites did not. CONCLUSIONS/SIGNIFICANCE: Although some soil characteristics were unaffected by tree species identity, our results clearly demonstrate that these dominant tree species are associated with soils that differ in several physical, chemical, and biotic properties. Ongoing environmental changes in this region, e.g. changes in fire regime, frequency of insect outbreaks, changes in precipitation patterns and snowpack, and land-use change, may alter the relative abundance of these tree species over coming decades, which in turn will likely alter the soils.

Inference of allelopathy is complicated by effects of activated carbon on plant growth.

Allelopathy can play an important role in structuring plant communities, but allelopathic effects are often difficult to detect because many methods used to test for allelopathy can be confounded by experimental artifacts. The use of activated carbon, a technique for neutralizing allelopathic compounds, is now employed in tests for allelopathy; however, this technique also could produce large experimental artifacts. In three independent experiments, it was shown that adding activated carbon to potting media affected nutrient availability and plant growth. For most species tested, activated carbon increased plant biomass, even in the absence of the potentially allelopathic agent. The increased growth corresponded to increased plant nitrogen content, likely resulting from greater nitrogen availability. Activated carbon also affected nitrogen and other nutrient concentrations in soil media in the absence of plants. The observed effects of activated carbon on plant growth can confound its use to test for allelopathy. The detection of allelopathy relies on the difference between plant growth in medium with carbon and that in medium without carbon in the presence of the potentially allelopathic competitor; however, this difference may be biased if activated carbon alters soil nutrient availability and plant growth even in the absence of the focal allelopathic agent.

Modeling the effect of photosynthetic vegetation properties on the NDVI--LAI relationship.

Developing a relationship between the normalized difference vegetation index (NDVI) and the leaf area index (LAI) is essential to describe the pattern of spatial or temporal variation in LAI that controls carbon, water, and energy exchange in many ecosystem process models. Photosynthetic vegetation (PV) properties can affect the estimation of LAI, but no models integrate the effects of multiple species. We developed four alternative NDVI-LAI models, three of which integrate PV effects: no PV effects, leaf-level effects, canopy-level effects, and effects at both levels. The models were fit to data across the natural range of variation in NDVI for a widespread High Arctic ecosystem. The weight of evidence supported the canopy-level model (Akaike weight, wr = 0.98), which includes species-specific canopy coefficients that primarily scale fractional PV cover to LAI by accounting for the area of unexposed PV. Modeling the canopy-level effects improved prediction of LAI (R2 = 0.82) over the model with no PV effect (R2 = 0.71) across the natural range of variation in NDVI but did not affect the site-level estimate of LAI. Satellite-based methods to estimate species composition, a variable in the model, will need to be developed. We expect that including the effects of PV properties in NDVI-LAI models will improve prediction of LAI where species composition varies across space or changes over time.