Showing papers in "Ecological Monographs in 2016"

Dynamics of non-structural carbohydrates in terrestrial plants: a global synthesis

Abstract: Plants store large amounts of non-structural carbohydrates (NSC). While multiple functions of NSC have long been recognized, the interpretation of NSC seasonal dynamics is often based on the idea that stored NSC is a reservoir of carbon that fluctuates depending on the balance between supply via photosynthesis and demand for growth and respiration (the source–sink dynamics concept). Consequently, relatively high NSC concentrations in some plants have been interpreted to reflect excess supply relative to demand. An alternative view, however, is that NSC accumulation reflects the relatively high NSC levels required for plant survival; an important issue that remains highly controversial. Here, we assembled a new global database to examine broad patterns of seasonal NSC variation across organs (leaves, stems, and belowground), plant functional types (coniferous, drought-deciduous angiosperms, winter deciduous angiosperms, evergreen angiosperms, and herbaceous) and biomes (boreal, temperate, Mediterranean, and tropical). We compiled data from 121 studies, including seasonal measurements for 177 species under natural conditions. Our results showed that, on average, NSC account for ~10% of dry plant biomass and are highest in leaves and lowest in stems, whereas belowground organs show intermediate concentrations. Total NSC, starch, and soluble sugars (SS) varied seasonally, with a strong depletion of starch during the growing season and a general increase during winter months, particularly in boreal and temperate biomes. Across functional types, NSC concentrations were highest and most variable in herbaceous species and in conifer needles. Conifers showed the lowest stem and belowground NSC concentrations. Minimum NSC values were relatively high (46% of seasonal maximums on average for total NSC) and, in contrast to average values, were similar among biomes and functional types. Overall, although starch depletion was relatively common, seasonal depletion of total NSC or SS was rare. These results are consistent with a dual view of NSC function: whereas starch acts mostly as a reservoir for future use, soluble sugars perform immediate functions (e.g., osmoregulation) and are kept above some critical threshold. If confirmed, this dual function of NSC will have important implications for the way we understand and model plant carbon allocation and survival under stress.

The ecology of methane in streams and rivers: patterns, controls, and global significance

Abstract: "Support for this paper was provided by funding from the North Temperate Lakes LTER program, NSF DEB‐0822700."

Stoichiometry of microbial carbon use efficiency in soils

Abstract: The carbon use efficiency (CUE) of microbial communities partitions the flow of C from primary producers to the atmosphere, decomposer food webs, and soil C stores. CUE, usually defined as the ratio of growth to assimilation, is a critical parameter in ecosystem models, but is seldom measured directly in soils because of the methodological difficulty of measuring in situ rates of microbial growth and respiration. Alternatively, CUE can be estimated indirectly from the elemental stoichiometry of organic matter and microbial biomass, and the ratios of C to nutrient-acquiring ecoenzymatic activities. We used this approach to estimate and compare microbial CUE in >2000 soils from a broad range of ecosystems. Mean CUE based on C:N stoichiometry was 0.269 ± 0.110 (mean ± SD). A parallel calculation based on C:P stoichiometry yielded a mean CUE estimate of 0.252 ± 0.125. The mean values and frequency distributions were similar to those from aquatic ecosystems, also calculated from stoichiometric models, and to those calculated from direct measurements of bacterial and fungal growth and respiration. CUE was directly related to microbial biomass C with a scaling exponent of 0.304 (95% CI 0.237–0.371) and inversely related to microbial biomass P with a scaling exponent of −0.234 (95% CI −0.289 to −0.179). Relative to CUE, biomass specific turnover time increased with a scaling exponent of 0.509 (95% CI 0.467–0.551). CUE increased weakly with mean annual temperature. CUE declined with increasing soil pH reaching a minimum at pH 7.0, then increased again as soil pH approached 9.0, a pattern consistent with pH trends in the ratio of fungal : bacteria abundance and growth. Structural equation models that related geographic variables to CUE component variables showed the strongest connections for paths linking latitude and pH to β-glucosidase activity and soil C:N:P ratios. The integration of stoichiometric and metabolic models provides a quantitative description of the functional organization of soil microbial communities that can improve the representation of CUE in microbial process and ecosystem simulation models.

Carbon content and climate variability drive global soil bacterial diversity patterns

Abstract: Despite the vital role of microorganisms for ecosystem functioning and human welfare, our understanding of their global diversity and biogeographical patterns lags significantly behind that of plants and animals. We conducted a meta-analysis including ~600 soil samples from all continents to evaluate the biogeographical patterns and drivers of bacterial diversity in terrestrial ecosystems at the global scale. Similar to what has been found with plants and animals, the diversity of soil bacteria in the Southern Hemisphere decreased from the equator to Antarctica. However, soil bacteria showed similar levels of diversity across the Northern Hemisphere. The composition of bacterial communities followed dissimilar patterns between hemispheres, as the Southern and Northern Hemispheres were dominated by Actinobacteria and Acidobacteria, respectively. However, Proteobacteria was co-dominant in both hemispheres. Moreover, we found a decrease in soil bacterial diversity with altitude. Climatic features (e.g. high diurnal temperature range and low temperature) were correlated with the lower diversity found at high elevations, but geographical gradients in soil total carbon and species turnover were important drivers of the observed latitudinal patterns. We thus found both parallels and differences in the biogeographical patterns of above- versus soil bacterial diversity. Our findings support previous studies that highlighted soil pH, spatial influence and organic matter as important drivers of bacterial diversity and composition. Furthermore, our results provide a novel integrative view of how climate and soil factors influence soil bacterial diversity at the global scale, which is critical to improve ecosystem and earth system simulation models and for formulating sustainable ecosystem management and conservation policies. This article is protected by copyright. All rights reserved.

Characterizing the ecological trade‐offs throughout the early ontogeny of coral recruitment

Abstract: Drivers of recruitment in sessile marine organisms are often poorly understood, due to the rapidly changing requirements experienced during early ontogeny. The complex suite of physical, biological, and ecological interactions beginning at larval settlement involves a series of trade-offs that influence recruitment success. For example, while cryptic settlement within complex microhabitats is a commonly observed phenomenon in sessile marine organisms, it is unclear whether trade-offs between competition in cryptic refuges and predation on exposed surfaces leads to higher recruitment.To explore the trade-offs during the early ontogeny of scleractinian corals, we combined field observations with laboratory and field experiments to develop a mechanistic understanding of coral recruitment success. Multiple experiments conducted over 15 months in Palau (Micronesia) allowed a mechanistic approach to study the individual factors involved in recruitment: settlement behavior, growth, competition, and predation, as functions of microhabitat and ontogeny. We finally developed and tested a predictive recruitment model with the broader aim of testing whether our empirical insights explained patterns of coral recruitment and quantifying the relative importance of each trade-off.Coral settlement was higher in crevices than exposed microhabitats, but post-settlement bottlenecks differed markedly in the presence (uncaged) and absence (caged) of predators. Incidental predation by herbivores on exposed surfaces at early post-settlement (<3 mm) stages and targeted predation by corallivores at late post-settlement (3–10 mm) stages exceeded competition in crevices as major drivers of mortality. In contrast, when fish were excluded, competition with macroalgae and heterotrophic invertebrates intensified mortality, particularly in crevices. As a result, post-settlement trade-offs were reversed, and recruitment was more than twofold higher on exposed surfaces than crevices. Once post-settlement bottlenecks were overcome, survival was higher on exposed surfaces regardless of fish exclusion. However, maximum recruitment occurred in crevices of uncaged treatments, being ninefold higher than caged treatments. Overall, we characterize recruitment success throughout the earliest life-history stages of corals and uncover some intriguing trade-offs between growth, competition and predation, highlighting how these change and even reverse during ontogeny and under alternate disturbance regimes.

Legacy effects of drought in the southwestern United States: A multi‐species synthesis

Abstract: Understanding impacts of drought on tree growth and forest health is of major concern given projected climate change. Droughts may become more common in the Southwest due to extreme temperatures that will drive increased evapotranspiration and lower soil moisture, in combination with uncertain precipitation changes. Utilizing ~1.3 million tree-ring widths from the International Tree Ring Data Bank representing 10 species (eight conifers, two oaks) in the Southwest, we evaluated the effects of drought on tree growth. We categorized ring widths by formation year in relation to drought (pre-drought, drought year, and post-drought), and we used a mixed-effects model to estimate the effects of current and antecedent precipitation and temperature on tree growth during the post-drought recovery period. This allowed us to assess changes in sensitivity of tree growth to precipitation and temperature at multiple timescales following multiple droughts, and to evaluate drought resistance and recovery in these species. The effects of precipitation and temperature on ring widths following drought varied among species and time since drought. Across species, 16% of the climate effects (i.e., “sensitivities”) were significantly different from their pre-drought values. Species differed, with some showing increased sensitivities to precipitation and temperature following drought, and others showing decreased sensitivities. Furthermore, some species (e.g., Abies concolor and Pinus ponderosa) showed low resistance and slow recovery, with changes in growth sensitivities persisting up to 5 yr; others (e.g., Juniper spp.) showed high resistance, such that their climatic sensitivities did not change. Among species, the importance of different antecedent climate variables changed with time since drought. Though a majority of species responded positively to same-year precipitation pre-drought, all 10 species were positively affected by same-year precipitation the second year after drought. Our results demonstrate tree growth sensitivities vary among species and with time since drought, raising questions about physiological mechanisms and implications for forest health under future drought.

The adaptive capacity of lake food webs: from individuals to ecosystems

Abstract: Aquatic ecosystems support size structured food webs, wherein predator-prey body sizes span orders of magnitude. As such, these food webs are replete with extremely generalized feeding strategies, especially among the larger bodied, higher trophic position taxa. The movement scale of aquatic organisms also generally increases with body size and trophic position. Together, these body size, mobility, and foraging relationships suggest that organisms lower in the food web generate relatively distinct energetic pathways by feeding over smaller spatial areas. Concurrently, the potential capacity for generalist foraging and spatial coupling of these pathways often increases, on average, moving up the food web toward higher trophic levels. We argue that these attributes make for a food web architecture that is inherently ‘adaptive’ in its response to environmental conditions. This is because variation in lower trophic level dynamics is dampened by the capacity of predators to flexibly alter their foraging behavior. We argue that empirical, theoretical, and applied research needs to embrace this inherently adaptive architecture if we are to understand the relationship between structure and function in the face of ongoing environmental change. Toward this goal, we discuss empirical patterns in the structure of lake food webs to suggest that ecosystems change consistently, from individual traits to the structure of whole food webs, under changing environmental conditions. We then explore an empirical example to reveal that explicitly unfolding the mechanisms that drive these adaptive responses offers insight into how human-driven impacts, such as climate change, invasive species, and fisheries harvest, ought to influence ecosystem structure and function (e.g., stability, secondary productivity, maintenance of major energy pathways). We end by arguing that such a directed food web research program promises a powerful across-scale framework for more effective ecosystem monitoring and management.

Old-growth Neotropical forests are shifting in species and trait composition

Abstract: Tropical forests have long been thought to be in stable state, but recent insights indicate that global change is leading to shifts in forest dynamics and species composition. These shifts may be driven by environmental changes such as increased resource availability, increased drought stress, and/or recovery from past disturbances. The relative importance of these drivers can be inferred from analyzing changes in trait values of tree communities. Here, we evaluate a decade of change in species and trait composition across five old-growth Neotropical forests in Bolivia, Brazil, Guyana, and Costa Rica that cover large gradients in rainfall and soil fertility. To identify the drivers of compositional change, we used data from 29 permanent sample plots and measurements of 15 leaf, stem, and whole-plant traits that are important for plant performance and should respond to global change drivers. We found that forests differ strongly in their community-mean trait values, resulting from differences in soil fertility and annual rainfall seasonality. The abundance of deciduous species with high specific leaf area increases from wet to dry forests. The community-mean wood density is high in the driest forests to protect xylem vessels against drought cavitation, and is high in nutrient-poor forests to increase wood longevity and enhance nutrient residence time in the plant. Interestingly, the species composition changed over time in three of the forests, and the community-mean wood density increased and the specific leaf area decreased in all forests, indicating that these forests are changing toward later successional stages dominated by slow-growing, shade-tolerant species. We did not see changes in other traits that could reflect responses to increased drought stress, such as increased drought deciduousness or decreased maximum adult size, or that could reflect increased resource availability (CO2, rainfall, or nitrogen). Changes in species and trait composition in these forests are therefore most likely caused by recovery from past disturbances. These compositional changes may also lead to shifts in ecosystem processes, such as a lower carbon sequestration and “slower” forest dynamics.

Ecological effects of extreme drought on Californian herbaceous plant communities

Abstract: Understanding the consequences of extreme climatic events is a growing challenge in ecology Climatic extremes may differentially affect varying elements of biodiversity, and may not always produce ecological effects exceeding those of “normal” climatic variation in space and time We asked how the extreme drought years of 2013–2014 affected the cover, species richness, functional trait means, functional diversity, and phylogenetic diversity of herbaceous plant communities across the California Floristic Province We compared the directions and magnitudes of these drought effects with expectations from four “pre-drought” studies of variation in water availability: (1) a watering experiment, (2) a long-term (15-yr) monitoring of interannual variability, (3) a resampling of historic (57-yr-old) plots within a warming and drying region, and (4) natural variation in communities over a broad geographic gradient in precipitation We found that the drought was associated with consistent reductions in species richness and cover, especially for annual forbs and exotic annual grasses, but not with changes in functional or phylogenetic diversity Except for total cover and cover of exotic annual grasses, most drought effects did not exceed quantitative expectations based on the four pre-drought studies Qualitatively, plant community responses to the drought were most concordant with responses to pre-drought interannual rainfall variability in the 15-yr monitoring study, and least concordant with responses to the geographic gradient in precipitation Our results suggest that, at least in the short term, extreme drought may cause only a subset of community metrics to respond in ways that exceed normal background variability

Phosphorus metabolism of reef organisms with algal symbionts

Abstract: Phosphorus (P), an essential structural and functional component of all living organisms, is considered to be the ultimate limiting nutrient in marine ecosystems To optimize its acquisition, marine species such as protozoa, sponges, foraminifera, clams, and reef corals, among others, have entered symbiotic relationships with algae, which recycle waste products of the animal host and transform dissolved inorganic nutrients into organic molecules, making them bioavailable to their host Such associations provide a competitive edge in an environment where ambient nutrient availability is low The aim of this review is to summarize the current knowledge on the P sources available to reef organisms with algal symbionts, to discuss the means by which P is taken up and stored within the symbiosis, and to assess the effects of eutrophication-induced phosphate enrichment on the algal and host physiology Finally, we give an overview of knowledge gaps and open questions that should be addressed to better understand the role of phosphorus in reef symbioses functioning

Deterministic and stochastic processes lead to divergence in plant communities 25 years after the 1988 Yellowstone fires

Abstract: Young, recently burned forests are increasingly widespread throughout western North America, but forest development after large wildfires is not fully understood, especially regarding effects of variable burn severity, environmental heterogeneity, and changes in drivers over time. We followed development of subalpine forests after the 1988 Yellowstone fires by periodically resampling permanent plots established soon after the fires. We asked two questions about patterns and processes over the past 25 years: (1) Are plant species richness and community composition converging or diverging across variation in elevation, soils, burn severity, and post-fire lodgepole pine (Pinus contorta var. latifolia) density? (2) What are the major controls on post-fire species composition, and has the relative importance of controls changed over time? For question 1, we sampled 10-m2 plots (n = 552) distributed among three geographic areas that differ in elevation and substrate; plots spanned the spectrum of fire severities and were resampled periodically from 1991 to 2013. For question 2, we sampled 0.25-ha plots (n = 72), broadly distributed across areas that burned as stand-replacing fire, in 1999 and 2012. Richness and species composition diverged early on between infertile low-elevation areas (lower richness) and more fertile high-elevation areas (greater richness). Richness increased rapidly for the first 5 yr post-fire, then leveled off or increased only slowly thereafter. Only 6% of 227 recorded species were nonnative. Some annuals and species with heat-stimulated soil seed banks were associated with severely burned sites. However, most post-fire species had been present before the fire; many survived as roots or rhizomes and regenerated rapidly by sprouting. Among the 72 plots, substrate, temperature, and precipitation (the abiotic template) were consistently important drivers of community composition in 1999 and 2012. Post-fire lodgepole pine abundance was not significant in 1999 but was the most important driving variable by 2012, with a negative effect on presence of most understory species, especially annuals and shade-intolerant herbs. Burn severity was significant in 1999 but not in 2012, and distance to unburned forest had no influence in either year. The 1988 fires did not fundamentally alter subalpine forest community assemblages in Yellowstone, and ecological memory conferred resilience to high-severity fire.

Reorganization of taxonomic, functional, and phylogenetic ant biodiversity after conversion to rubber plantation

Abstract: Agricultural activity associated with habitat conversion is a major driver of biodiversity loss across the tropics. The decline of species richness is a common outcome of conversion to agricultural land use, but the associated changes in functional and phylogenetic diversities, and spatial patterns of beta diversity, are not well understood. These patterns may shed light on underlying ecological processes that are of both basic and applied interest; for example the relative roles of stochastic assembly, ecological filtering, and competition in structuring ecological communities as well as broader consequences for ecological functioning in agroecosystems. Here we investigate the effects of conversion to rubber plantation (Hevea brasiliensis), a rapidly spreading agroecosystem in Southeast Asia, on leaf-litter ant taxonomic, functional, and phylogenetic diversities at local and among-sites scales in Xishuangbanna, Southeast China. We found a sharp decline of ant species richness in rubber plantations compared with nearby forest habitat, with low beta diversity indicating spatial homogeneity of communities in rubber plantations. In addition, patterns of both functional alpha and beta diversities suggested the emergence of a functionally distinct ant community in the agroecosystem compared to the forest habitats. These results support a role of ecological filtering in structuring the taxonomic and functional composition of both rubber and forest habitats. In contrast, changes in phylogenetic diversity were modest and not significantly different from random expectations despite strong phylogenetic signal of functional traits. This study highlights the need for a pluralistic approach to characterizing the loss of biodiversity in agroecosystems, as well as understanding the underlying mechanisms of community assembly driving this biodiversity loss.

Environmental heterogeneity has a weak effect on diversity during community assembly in tallgrass prairie

Abstract: Citation: Baer, S. G., Blair, J. M., & Collins, S. L. (2016). Environmental heterogeneity has a weak effect on diversity during community assembly in tallgrass prairie. Ecological Monographs, 86(1), 94-106. doi:10.1890/15-0888.1

Cohort variation in individual body mass dissipates with age in large herbivores

Abstract: Environmental conditions experienced during early growth and development markedly shape phenotypic traits. Consequently, individuals of the same cohort may show similar life-history tactics throughout life. Conditions experienced later in life, however, could fine-tune these initial differences, either increasing (cumulative effect) or decreasing (compensatory effect) the magnitude of cohort variation with increasing age. Our novel comparative analysis that quantifies cohort variation in individual body size trajectories shows that initial cohort variation dissipates throughout life, and that lifetime patterns change both across species with different paces of life and between sexes. We used longitudinal data on body size (mostly assessed using mass) from 11 populations of large herbivores spread along the “slow-fast” continuum of life histories. We first quantified cohort variation using mixture models to identify clusters of cohorts with similar initial size. We identified clear cohort clusters in all species except the one with the slowest pace of life, revealing that variation in early size is structured among cohorts and highlighting typological differences among cohorts. Growth trajectories differed among cohort clusters, highlighting how early size is a fundamental determinant of lifetime growth patterns. In all species, among-cohort variation in size peaked at the start of life, then quickly decreased with age and stabilized around mid-life. Cohort variation was lower in species with a slower than a faster pace of life, and vanished at prime age in species with the slowest pace of life. After accounting for viability selection, compensatory/catch-up growth in early life explained much of the decrease in cohort variation. Females showed less phenotypic variability and stronger compensatory/catch-up growth than males early in life, whereas males showed more progressive changes throughout life. These results confirm that stronger selective pressures for rapid growth make males more vulnerable to poor environmental conditions early in life and less able to recover after a poor start. Our comparative analysis illustrates how variability in growth changes over time in closely related species that span a wide range on the “slow-fast” continuum, the main axis of variation in life-history strategies of vertebrates. This article is protected by copyright. All rights reserved.

Habitat, predators, and hosts regulate disease in Daphnia through direct and indirect pathways

Abstract: Community ecology can link habitat to disease via interactions among habitat, focal hosts, other hosts, their parasites, and predators. However, complicated food web interactions (i.e., trophic interactions among predators and their impacts on host density and diversity) often obscure the important pathways regulating disease. Here, we disentangle community drivers in a case study of planktonic disease, using a two-step approach. In step one, we tested univariate field patterns linking community interactions directly to two disease metrics. Density of focal hosts (Daphnia dentifera) was related to density but not prevalence of fungal (Metschnikowia bicuspidata) infections. Both disease metrics appeared to be driven by selective predators that cull infected hosts (fish, e.g., Lepomis macrochirus), sloppy predators that spread parasites while feeding (midges, Chaoborus punctipennis), and spore predators that reduce contact between focal hosts and parasites (other zooplankton, especially small-bodied Ceriodaphnia sp.). Host diversity also negatively correlated with disease, suggesting a dilution effect. However, several of these univariate patterns were initially misleading, due to confounding ecological links among habitat, predators, host density, and host diversity. In step two, path models uncovered and explained these misleading patterns, and grounded them in habitat structure (refuge size). First, rather than directly reducing infection prevalence, fish predation drove disease indirectly through changes in density of midges and frequency of small spore predators (which became more frequent in lakes with small refuges). Second, small spore predators drove the two disease metrics through fundamentally different pathways: they directly reduced infection prevalence, but indirectly reduced density of infected hosts by lowering density of focal hosts (likely via competition). Third, the univariate diversity–disease pattern (signaling a dilution effect) merely reflected the confounding direct effects of these small spore predators. Diversity per se had no effect on disease, after accounting for the links between small spore predators, diversity, and infection prevalence. In turn, these small spore predators were regulated by both size-selective fish predation and refuge size. Thus, path models not only explain each of these surprising results, but also trace their origins back to habitat structure.

The effect of demographic correlations on the stochastic population dynamics of perennial plants

Abstract: Understanding the influence of environmental variability on population dynamics is a fundamental goal of ecology. Theory suggests that, for populations in variable environments, temporal correlations between demographic vital rates (e.g., growth, survival, reproduction) can increase (if positive) or decrease (if negative) the variability of year-to-year population growth. Because this variability generally decreases long-term population viability, vital rate correlations may importantly affect population dynamics in stochastic environments. Despite long-standing theoretical interest, it is unclear whether vital rate correlations are common in nature, whether their directions are predominantly negative or positive, and whether they are of sufficient magnitude to warrant broad consideration in studies of stochastic population dynamics. We used long-term demographic data for three perennial plant species, hierarchical Bayesian parameterization of population projection models, and stochastic simulations to address the following questions: (1) What are the sign, magnitude, and uncertainty of temporal correlations between vital rates? (2) How do specific pairwise correlations affect the year-to-year variability of population growth? (3) Does the net effect of all vital rate correlations increase or decrease year-to-year variability? (4) What is the net effect of vital rate correlations on the long-term stochastic population growth rate (λS)? We found only four moderate to strong correlations, both positive and negative in sign, across all species and vital rate pairs; otherwise, correlations were generally weak in magnitude and variable in sign. The net effect of vital rate correlations ranged from a slight decrease to an increase in the year-to-year variability of population growth, with average changes in variance ranging from -1% to +22%. However, vital rate correlations caused virtually no change in the estimates of λS (mean effects ranging from -0.01% to +0.17%). Therefore, the proportional changes in the variance of population growth caused by demographic correlations were too small on an absolute scale to importantly affect population growth and viability. We conclude that in our three focal populations and perhaps more generally, vital rate correlations have little effect on stochastic population dynamics. This may be good news for population ecologists, because estimating vital rate correlations and incorporating them into population models can be data-intensive and technically challenging. This article is protected by copyright. All rights reserved.

Pyrogenic fuels produced by savanna trees can engineer humid savannas

Abstract: Natural fires ignited by lightning strikes following droughts frequently are posited as the ecological mechanism maintaining discontinuous tree cover and grass‐dominated ground layers in savannas. Such fires, however, may not reliably maintain humid savannas. We propose that savanna trees producing pyrogenic shed leaves might engineer fire characteristics, affecting ground‐layer plants in ways that maintain humid savannas. We explored our hypothesis in a high‐rainfall, frequently burned pine savanna in which the dominant tree, longleaf pine (Pinus palustris), produces resinous needles that become highly flammable when shed and dried. We postulated that pyrogenic needles should have much greater influence on fire characteristics at ground level, and hence post‐fire responses of dominant shrubs and grasses, than other abundant fine fuels (shed oak leaves and grass culms). We further reasoned that these effects should increase with amounts of needles. We managed site conditions that affect fuels (time since fire, dominant vegetation), manipulated amounts of needles in ground‐layer plots, prescribed burned the plots, and measured fire characteristics at ground level. We also measured characteristics of ground‐layer oaks and grasses before, then 2 and 8 months after fires. We tested our hypotheses regarding effects of pyrogenic pine fuels on fire characteristics and vegetation regrowth and explored direct and indirect effects of fuels on fire characteristics and vegetation using a structural equation model. Pine needles influenced fire characteristics, elevating maximum temperature increases, durations of heating above 60°C, and fine fuel consumption considerably above measurements when fuels only included other savanna plants. Presence of pine needles depressed post‐fire numbers of oak stems and grass culms, especially in the interior of grass genets, as well as post‐fire flowering of grasses. The structural equation model indicated strong direct and indirect pathways from pine needles to post‐fire responses of oaks and grasses. The experimental field tests of hypotheses, bolstered by structural equation modeling, indicate pyrogenic fine fuels modify characteristics of prescribed fires at ground level, negatively affecting dominant ground‐layer oaks and grasses. Frequent fires fueled by pyrogenic needles should maintain humid savannas and generate spatial pyrodiversity that affects composition and dynamics of pine savanna ground‐layer vegetation.

Pivotal effect of early-winter temperatures and snowfall on population growth of alpine Parnassius smintheus butterflies

Abstract: Geographic range shifts in species’ distributions, due to climate change, imply altered dynamics at both their northern and southern range limits, or at upper and lower elevational limits. There is therefore a need to identify specific weather or climate variable(s), and life stages or cohorts on which they act, and how these affect population growth. Identifying such variables permits prediction of population increase or decline under a changing climate, and shifts in a species’ geographic range. For relatively well studied groups, such as butterflies, geographic range shifts are well documented, but weather variables and mechanisms causing those shifts are not well known. The Holarctic butterfly genus Parnassius (Papilionidae) inhabits northern and alpine environments subject to variable and extreme weather. As such, Parnassius species are vulnerable not only to long-term changes in average conditions but especially to short-term extreme weather events. We use population growth estimates for the alpine butterfly, Parnassius smintheus, from 21 populations in the Rocky Mountains of Canada over a 20-yr interval combined with techniques of machine learning (randomForests) and parametric modeling to identify the important weather variables determining population growth. We do this to determine the seasons and life stages of P. smintheus most affected by climate change. Extreme minimum and maximum temperatures in November, in combination with November snowfall, affect annual population growth most, more so than do mean temperatures in November, and more so than weather at any other time of year. Populations decline both in years with low extreme minimum temperatures in November and especially in years with high extreme maximum temperatures in November, indicating that overwintering eggs are particularly vulnerable to early-winter weather. Snowfall ameliorates the negative effects of extreme temperatures, particularly for extreme warm events. Results provide insight into biological mechanisms by which overwintering eggs might be affected by early winter weather. Short-term extreme weather in November, acting on a single pivotal life stage (egg) is a far better predictor of population change of alpine P. smintheus butterflies than is the general index of climate, the Pacific Decadal Oscillation.

Calcium carbonate deposition drives nutrient cycling in a calcareous headwater stream

Abstract: Calcium carbonate (CaCO3) deposition is common in aquatic ecosystems and may reduce phosphorus availability via coprecipitation of phosphate, an impact with broad implications for ecosystem processes. To determine if CaCO3 deposition in streams increases phosphorus (P) retention in minerals while reducing P availability to organisms, we studied paired streams (with and without active CaCO3 deposition) subjected to experimental shading and monitored changes in ecosystem attributes (e.g., periphyton biomass content, nutrient spiraling, periphyton nutrient limitation, and leaf litter decomposition). Shading reduced rates of CaCO3 deposition by over 50%, suggesting that a substantial proportion of CaCO3 deposition is supported by photosynthetically induced changes in alkalinity. Shading-induced reductions in CaCO3 deposition led to increases in epilithon biomass P content (P < 0.05) and periphyton growth (F2,12 = 5.79, P < 0.05). Reductions in CaCO3 deposition also relieved P limitation of periphyton growth (F3,16 = 59.32, P < 0.001), extended P uptake lengths at least an order of magnitude, and reduced both P mass transfer velocity and areal uptake rates by over 80% (F2,3 = 22.62, P < 0.05 and F2,3 = 13.19, P < 0.05, respectively). Finally, while shading caused reductions in leaf litter decomposition in the non-CaCO3 depositing stream (F5,7 = 22.45, P < 0.001), shading had no effect on leaf litter decomposition in the stream with active CaCO3 deposition. These results indicate that CaCO3 deposition can regulate P bioavailability and retention in streams and may drive streams to be P limited, as has been suggested in lake and wetland ecosystems.

Thermal sensitivity and the role of behavior in driving an intertidal predator‐prey interaction

Abstract: Environmental stress models (ESM) provide a useful framework to study the direct and indirect ecological drivers of community diversity and resilience. ESMs make predictions about the relative importance of structuring processes (e.g. predation) based on the relative stress suffered by consumers and prey. Their practical application, i.e. determining the conditions under which consumers and prey performance is more negatively affected, has been limited because the roles of behavior and physiology are not usually considered. We examined the role of thermal sensitivity and behavior on the thermal performance of the rocky intertidal predator Pisaster ochraceus and its main prey Mytilus californianus. We propose a novel framework that merges thermal performance curves (TPC) with observations of microhabitat use to provide a realistic perspective of the relative physiological conditions of predator and prey. First, by deriving aquatic and aerial TPCs for both species and from two sites, we found differences in parameter values that in some cases correspond to the individuals’ origins. Second, we calculated realized thermal performance in the field by combining TPCs with body temperatures recorded with biomimetic sensors. Notably, thermal performance of Pisaster was higher than that for Mytilus (i.e. prey-stress model), contrary to previous expectations based on caging experiments. Third, these estimates of thermal performance corresponded loosely with a measured indicator of overall physiological condition (body mass index, BMI) and a marker for extreme thermal stress (heat-shock proteins 70kDa), suggesting that environmental drivers other than temperature such as food supply must be considered. We found no evidence that Pisaster movement significantly influences thermal performance under typical conditions, suggesting instead that its preference for sheltered microhabitats provides a mechanism for avoiding exposure to extreme environmental conditions. Through the application of TPCs and ESMs, this study provides a unique perspective on the importance of physiology and behavior in driving the sensitivity of species interactions to environmental change. Crucially, this framework allowed clarifying that this system behaves as a prey- instead of consumer-stress model, which may also apply to many other ectotherm species interactions. This article is protected by copyright. All rights reserved.

A stochastic model for landscape patterns of biodiversity

Abstract: Many factors have been proposed to affect biodiversity patterns across landscapes, including patch area, patch isolation, edge distances, and matrix quality, but existing models emphasize only one or two of these factors at a time. Here we introduce a synthetic but simple individual-based model that generates realistic patterns of species richness and density as a function of landscape structure. In this model, we simulated the stochastic placement of home ranges in landscapes, thus combining features of existing random placement and mid-domain effect models. As such, the model allows investigation of whether and how geometric constraints on home range placement of individuals scale up to affect species abundance and richness in landscapes. The model encompassed a gradient of possible landscapes, from a strictly homogeneous landscape to an archipelago of discrete patches. The model incorporated only variation in home range size of individuals of different species, with a simple suitability index that controlled home range spread into areas of habitat and areas of inter-habitat matrix. Demographic details of birth, death, and migration, as well as effects of species interactions were not included. Nevertheless, this simple model generated realistic patterns of biodiversity, including species–area curves and increases in diversity and abundance with decreasing isolation and increasing distance from patch edges. Species–area slopes (z values) generated by the model fell within the range observed in empirical studies on both true islands and habitat patches. Isolation and edge effects were stronger when the matrix was unsuitable, and became progressively weaker as matrix suitability increased, again in accordance with many empirical studies. When applied to a real data set on the abundance of 20 small mammal species sampled in forest patches in the Atlantic Forest of Brazil, the model predicted increases in abundance and richness with increasing patch size, consistent with the general pattern observed with field data. The ability of this simple model to reproduce realistic qualitative patterns of biodiversity suggests that such patterns may be driven, at least in part, by geometric constraints acting on the placement of individual ranges, which ultimately affect biodiversity patterns at the landscape level.

Environmental controls on spatial patterns in the long-term persistence of giant kelp in central California

Abstract: As marine management measures increasingly protect static areas of the oceans, it is important to make sure protected areas capture and protect persistent populations. Rocky reefs in many temperate areas worldwide serve as habitat for canopy-forming macroalgae and these structure-forming species of kelps (order Laminariales) often serve as important habitat for a great diversity of species. Macrocystis pyrifera is the most common canopy-forming kelp species found along the coast of California, but the distribution and abundance of M. pyrifera varies in space and time. The purpose of this study is to determine what environmental parameters are correlated with and their relative contribution to the spatial and temporal persistence of M. pyrifera along the central coast of California and how well those environmental parameters can be used to predict areas where this species is more likely to persist. Nine environmental variables considered in this study included depth of the seafloor, structure of the rocky reef, proportion of rocky reef, size of kelp patch, biomass of kelp within a patch, distance from the edge of a kelp patch, sea surface temperature, wave orbital velocities, and population connectivity of individual kelp patches. Using a generalized linear mixed effects model (GLMM), the persistence of M. pyrifera was significantly associated with seven of the nine variables considered: depth, complexity of the rocky reef, proportion of rock, patch biomass, distance from the edge of a patch, population connectivity, and wave orbital velocities. These seven environmental variables were then used to predict the persistence of kelp across the central coast, and these predictions were compared to a reserved dataset of M. pyrifera persistence, which was not used in the creation of the GLMM. The environmental variables were shown to accurately predict the persistence of M. pyrifera within the central coast of California (r = 0.71, P < 0.001). Because persistence of giant kelp is important to the community structure of kelp forests, understanding those factors that support persistent populations of M. pyrifera will enable more effective management of these ecosystems.

A multiple-scale assessment of long-term aspen persistence and elevational range shifts in the Colorado Front Range

Abstract: Aspen forests and woodlands are some of the most speciesrich forest communities in the northern hemisphere. Changing climate, altered disturbance regimes, land use, and increased herbivore pressure threaten these forests both in Eurasia and North America. In addition, rapid mortality dubbed “Sudden Aspen Decline” is a concern for aspen’s longterm presence in the western United States, especially Colorado and Utah. Yet it is still unclear whether aspen is persistent or declining at the landscape scale. We assessed aspen persistence at different spatial scales in the Colorado Front Range by resampling 89 plots containing aspen from among 305 vegetation plots sampled by Robert Peet during 1972–1973. We hypothesized that aspen density and basal area had decreased at the landscape scale, with notable variability in change depending on the forest community type, and that this overall decrease has been more pronounced at lower elevations. We also assessed elevational range shifts of the major species in these forests. Aspen were no longer present in 22 of the 89 plots and aspen density for stems less than 2.5 cm diameter at breast height (DBH) had declined significantly overall, although density of medium (2.5–10 cm DBH) and large (>10 cm DBH) trees, as well as basal area, had not changed significantly. A comparison between montane (<2700 m elevation) and subalpine (2700–3500 m elevation) plots revealed that the decrease was more pronounced at higher elevations and was mostly the result of substantial decreases of stems in the eleven plots that were part of Peet’s aspendominated “Populus tremuloides series.” In these plots, aspen basal area decreased significantly whereas basal area of Abies bifolia, Picea engelmannii, Pinus contorta, and Pseudotsuga menziesii increased substantially. Upslope shifts were observed for most species, especially on northeast facing slopes, suggesting climaterelated responses. In summary, aspen have been resilient in mixed forests and may be beneficiaries of recent bark beetle epidemics, but have decreased and been subject to successional transitions in previously aspendominated stands. Our results confirm the importance of regionspecific, multiplescale assessments of species persistence to make best management recommendations.

A general parameterized mathematical food web model that predicts a stable green world in the terrestrial ecosystem

Abstract: Terrestrial ecosystems are generally green with vegetation and only a small part ( Pcc > 0); plants are less nutritive than animals (np < nh, nc); and carnivore searches huge volume daily while eating speed of herbivore is limited (S _ eh). These conditions are well-realized in aboveground terrestrial ecosystems where plant-rich “green worlds” are common, vs. animal-rich belowground and aquatic ecosystems where some conditions are not realized. The h and c calculated from our model agreed well with those from empirical observations in forest ecosystems, where both h and c are within an order of magnitude of 100 mg (fresh biomass/m2 forest), and in savannah ecosystems. The model predicts that plant nutritive value np, digestibility of plants by herbivores αh, and herbivore eating speed eh are positively correlated with h and the intensity of herbivory, which theoretically explains the outdoor defensive effects of plant anti-nutritive or quantitative defenses (e.g., tannins, protease inhibitors), and predicts that c and the carnivore/herbivore ratio c/h are positively correlated with the relative growth rate of herbivores Gh. The present model introduced parameterized realities into food web theory that previous models lack.