Showing papers in "Ecology in 2013"

Responses of ecosystem carbon cycle to experimental warming: a meta‐analysis

Abstract: Global warming potentially alters the terrestrial carbon (C) cycle, likely feeding back to further climate warming. However, how the ecosystem C cycle responds and feeds back to warming remains unclear. Here we used a meta-analysis approach to quantify the response ratios of 18 variables of the ecosystem C cycle to experimental warming and evaluated ecosystem C-cycle feedback to climate warming. Our results showed that warming stimulated gross ecosystem photosynthesis (GEP) by 15.7%, net primary production (NPP) by 4.4%, and plant C pools from above- and belowground parts by 6.8% and 7.0%, respectively. Experimental warming accelerated litter mass loss by 6.8%, soil respiration by 9.0%, and dissolved organic C leaching by 12.1%. In addition, the responses of some of those variables to experimental warming differed among the ecosystem types. Our results demonstrated that the stimulation of plant-derived C influx basically offset the increase in warming-induced efflux and resulted in insignificant changes in litter and soil C content, indicating that climate warming may not trigger strong positive C-climate feedback from terrestrial ecosystems. Moreover, the increase in plant C storage together with the slight but not statistically significant decrease of net ecosystem exchange (NEE) across ecosystems suggests that terrestrial ecosystems might be a weak C sink rather than a C source under global climate warming. Our results are also potentially useful for parameterizing and benchmarking land surface models in terms of C cycle responses to climate warming.

The AIC model selection method applied to path analytic models compared using a d-separation test.

Abstract: Classical path analysis is a statistical technique used to test causal hypotheses involving multiple variables without latent variables, assuming linearity, multivariate normality, and a sufficient sample size. The d-separation (d-sep) test is a generalization of path analysis that relaxes these assumptions. Although model selection using Akaike's information criterion (AIC) is well established for classical path analysis, this model selection technique has not yet been developed for d-sep tests. In this paper, I explain how to use the AIC statistic for d-sep tests, give a worked example, and include instructions (supplemental material) to implement the analysis in the R computing language.

Reconstructing patterns of temperature, phenology, and frost damage over 124 years: Spring damage risk is increasing

Abstract: Climate change, with both warmer spring temperatures and greater temperature fluctuations, has altered phenologies, possibly leading to greater risk of spring frost damage to temperate deciduous woody plants. Phenological observations of 20 woody species from 1993 to 2012 in Trelease Woods, Champaign County, Illinois, USA, were used to identify years with frost damage to vegetative and reproductive phases. Local temperature records were used in combination with the phenological observations to determine what combinations of the two were associated with damage. Finally, a long-term temperature record (1889-1992) was evaluated to determine if the frequency of frost damage has risen in recent decades. Frost < or = -1.7 degrees C occurred after bud-break in 14 of the 20 years of observation. Frost damage occurred in five years in the interior and in three additional years at only the forest edge. The degree of damage varied with species, life stage, tissue (vegetative or reproductive), and phenological phase. Common features associated with the occurrence of damage to interior plants were (1) a period of unusual warm temperatures in March, followed by (2) a frost event in April with a minimum temperature < or = -6.1 degrees C with (3) a period of 16-33 days between the extremes. In the long-term record, 10 of 124 years met these conditions, but the yearly probability of frost damage increased significantly, from 0.03 during 1889-1979 to 0.21 during 1980-2012. When the criteria were "softened" to < or = -1.7 degrees C in April and an interval of 16-37 days, 31 of 124 years met the conditions, and the yearly damage probability increased significantly to 0.19 for 1889-1979 and 0.42 for 1980-2012. In this forest, the combination of warming trends and temperature variability (extremes) associated with climate change is having ecologically important effects, making previously rare frost damage events more common.

Microbial abundance and composition influence litter decomposition response to environmental change

Abstract: Rates of ecosystem processes such as decomposition are likely to change as a result of human impacts on the environment. In southern California, climate change and nitrogen (N) deposition in particular may alter biological communities and ecosystem processes. These drivers may affect decomposition directly, through changes in abiotic conditions, and indirectly through changes in plant and decomposer communities. To assess indirect effects on litter decomposition, we reciprocally transplanted microbial communities and plant litter among control and treatment plots (either drought or N addition) in a grassland ecosystem. We hypothesized that drought would reduce decomposition rates through moisture limitation of decomposers and reductions in plant litter quality before and during decomposition. In contrast, we predicted that N deposition would stimulate decomposition by relieving N limitation of decomposers and improving plant litter quality. We also hypothesized that adaptive mechanisms would allow microbes to decompose litter more effectively in their native plot and litter environments. Consistent with our first hypothesis, we found that drought treatment reduced litter mass loss from 20.9% to 15.3% after six months. There was a similar decline in mass loss of litter inoculated with microbes transplanted from the drought treatment, suggesting a legacy effect of drought driven by declines in microbial abundance and possible changes in microbial community composition. Bacterial cell densities were up to 86% lower in drought plots and at least 50% lower on litter derived from the drought treatment, whereas fungal hyphal lengths increased by 13–14% in the drought treatment. Nitrogen effects on decomposition rates and microbial abundances were weaker than drought effects, although N addition significantly altered initial plant litter chemistry and litter chemistry during decomposition. However, we did find support for microbial adaptation to N addition with N-derived microbes facilitating greater mass loss in N plots than in control plots. Our results show that environmental changes can affect rates of ecosystem processes directly through abiotic changes and indirectly through microbial abundances and communities. Therefore models of ecosystem response to global change may need to represent microbial biomass and community composition to make accurate predictions.

Practical guidance on characterizing availability in resource selection functions under a use–availability design

Abstract: Habitat selection is a fundamental aspect of animal ecology, the understanding of which is critical to management and conservation. Global positioning system data from animals allow fine-scale assessments of habitat selection and typically are analyzed in a use–availability framework, whereby animal locations are contrasted with random locations (the availability sample). Although most use–availability methods are in fact spatial point process models, they often are fit using logistic regression. This framework offers numerous methodological challenges, for which the literature provides little guidance. Specifically, the size and spatial extent of the availability sample influences coefficient estimates potentially causing interpretational bias. We examined the influence of availability on statistical inference through simulations and analysis of serially correlated mule deer GPS data. Bias in estimates arose from incorrectly assessing and sampling the spatial extent of availability. Spatial autocorrelati...

Early onset of spring increases the phenological mismatch between plants and pollinators

Abstract: Climate warming accelerates the timing of flowering and insect pollinator emergence, especially in spring. If these phenological shifts progress independently between species, features of plant-pollinator mutualisms may be modified. However, evidence of phenological mismatch in pollination systems is limited. We investigated the phenologies of a spring ephemeral, Corydalis ambigua, and its pollinators (bumble bees), and seed-set success over 10-14 years in three populations. Although both flowering onset and first detection of overwintered queen bees in the C. ambigua populations were closely related to snowmelt time and/or spring temperature, flowering tended to be ahead of first pollinator detection when spring came early, resulting in lower seed production owing to low pollination service. Relationships between flowering onset time, phenological mismatch, and seed-set success strongly suggest that phenological mismatch is a major limiting factor for reproduction of spring ephemerals. This report demonstrates the mechanism of phenological mismatch and its ecological impact on plant-pollinator interactions based on long-term monitoring. Frequent occurrence of mismatch can decrease seed production and may affect the population dynamics of spring ephemerals.

MARSPEC: ocean climate layers for marine spatial ecology

Abstract: Ecological niche models are widely used in terrestrial studies to address critical ecological and evolutionary questions related to past and future climate change, local adaptation and speciation, the discovery of rare endemics, and biological invasions. However the application of niche models to similar questions in marine ecosystems has lagged behind, in part due to the lack of a centralized high-resolution spatial data set representing both benthic and pelagic marine environments. Here we describe the creation of MARSPEC, a high-resolution GIS database of ocean climate layers intended for marine ecological niche modeling and other applications in marine spatial ecology. MARSPEC combines information related to topographic complexity of the seafloor with bioclimatic measures of sea surface temperature and salinity for the world ocean. We derived seven geophysical variables from a high-resolution raster grid representing depth of the seafloor (bathymetry) to characterize six facets of topographic...

Bee diversity effects on pollination depend on functional complementarity and niche shifts.

Abstract: Biodiversity is important for many ecosystem processes Global declines in pollinator diversity and abundance have been recognized, raising concerns about a pollination crisis of crops and wild plants However, experimental evidence for effects of pollinator species diversity on plant reproduction is extremely scarce We established communities with 1-5 bee species to test how seed production of a plant community is determined by bee diversity Higher bee diversity resulted in higher seed production, but the strongest difference was observed for one compared to more than one bee species Functional complementarity among bee species had a far higher explanatory power than bee diversity, suggesting that additional bee species only benefit pollination when they increase coverage of functional niches In our experiment, complementarity was driven by differences in flower and temperature preferences Interspecific interactions among bee species contributed to realized functional complementarity, as bees reduced interspecific overlap by shifting to alternative flowers in the presence of other species This increased the number of plant species visited by a bee community and demonstrates a new mechanism for a biodiversity-function relationship ("interactive complementarity") In conclusion, our results highlight both the importance of bee functional diversity for the reproduction of plant communities and the need to identify complementarity traits for accurately predicting pollination services by different bee communities

Moving to stay in place: behavioral mechanisms for coexistence of African large carnivores.

Abstract: Most ecosystems have multiple predator species that not only compete for shared prey, but also pose direct threats to each other. These intraguild interactions are key drivers of carnivore community structure, with ecosystem-wide cascading effects. Yet, behavioral mechanisms for coexistence of multiple carnivore species remain poorly understood. The challenges of studying large, free-ranging carnivores have resulted in mainly coarse-scale examination of behavioral strategies without information about all interacting competitors. We overcame some of these challenges by examining the concurrent fine-scale movement decisions of almost all individuals of four large mammalian carnivore species in a closed terrestrial system. We found that the intensity of intraguild interactions did not follow a simple hierarchical allometric pattern, because spatial and behavioral tactics of subordinate species changed with threat and resource levels across seasons. Lions (Panthera leo) were generally unrestricted and anchored themselves in areas rich in not only their principal prey, but also, during periods of resource limitation (dry season), rich in the main prey for other carnivores. Because of this, the greatest cost (potential intraguild predation) for subordinate carnivores was spatially coupled with the highest potential benefit of resource acquisition (prey-rich areas), especially in the dry season. Leopard (P. pardus) and cheetah (Acinonyx jubatus) overlapped with the home range of lions but minimized their risk using fine-scaled avoidance behaviors and restricted resource acquisition tactics. The cost of intraguild competition was most apparent for cheetahs, especially during the wet season, as areas with energetically rewarding large prey (wildebeest) were avoided when they overlapped highly with the activity areas of lions. Contrary to expectation, the smallest species (African wild dog, Lycaon pictus) did not avoid only lions, but also used multiple tactics to minimize encountering all other competitors. Intraguild competition thus forced wild dogs into areas with the lowest resource availability year round. Coexistence of multiple carnivore species has typically been explained by dietary niche separation, but our multi-scaled movement results suggest that differences in resource acquisition may instead be a consequence of avoiding intraguild competition. We generate a more realistic representation of hierarchical behavioral interactions that may ultimately drive spatially explicit trophic structures of multi-predator communities.

Root depth distribution and the diversity–productivity relationship in a long-term grassland experiment

Abstract: The relationship between plant diversity and productivity in grasslands could depend, partly, on how diversity affects vertical distributions of root biomass in soil; yet, no prior study has evaluated the links among diversity, root depth distributions, and productivity in a long-term experiment. We use data from a 12-year experiment to ask how plant species richness and composition influenced both observed and expected root depth distributions of plant communities. Expected root depth distributions were based on the abundance of species in each community and two traits of species that were measured in monocultures: root depth distributions and root to shoot ratios. The observed proportion of deep root biomass increased more than expected with species richness and was positively correlated with aboveground productivity. Indeed, the proportion of deep root biomass explained variation in productivity even after accounting for legume presence/abundance, and greater nitrogen availability in diverse plots. Diverse plots had root depth distributions that were twice as deep as expected from their species composition and corresponding monoculture traits, partly due to interactions between C4 grasses and legumes. These results suggest the productivity of diverse plant communities was partly dependent on belowground plant interactions that caused roots to be distributed more deeply in soil.

Successional changes in functional composition contrast for dry and wet tropical forest

Abstract: We tested whether and how functional composition changes with succession in dry deciduous and wet evergreen forests of Mexico. We hypothesized that compositional changes during succession in dry forest were mainly determined by increasing water availability leading to community functional changes from conservative to acquisitive strategies, and in wet forest by decreasing light availability leading to changes from acquisitive to conservative strategies. Research was carried out in 15 dry secondary forest plots (5–63 years after abandonment) and 17 wet secondary forest plots (<1–25 years after abandonment). Community-level functional traits were represented by community-weighted means based on 11 functional traits measured on 132 species. Successional changes in functional composition are more marked in dry forest than in wet forest and largely characterized by different traits. During dry forest succession, conservative traits related to drought tolerance and drought avoidance decreased, as predicted. Une...

Consumer diversity interacts with prey defenses to drive ecosystem function.

Abstract: Prey traits linking consumer diversity to ecosystem function remain poorly understood. On tropical coral reefs, herbivores promote coral dominance by suppressing competing macroalgae, but the roles of herbivore identity and diversity, macroalgal defenses, and their interactions in affecting reef resilience and function are unclear. We studied adjacent pairs of no-take marine reserves and fished areas on reefs in Fiji and found that protected reefs supported 7–17× greater biomass, 2–3× higher species richness of herbivorous fishes, and 3–11× more live coral cover than did fished reefs. In contrast, macroalgae were 27–61× more abundant and 3–4× more species-rich on fished reefs. When we transplanted seven common macroalgae from fished reefs into reserves they were rapidly consumed, suggesting that rates of herbivory (ecosystem functioning) differed inside vs. outside reserves. We then video-recorded feeding activity on the same seven macroalgae when transplanted into reserves, and assessed the functional redundancy vs. complementarity of herbivorous fishes consuming these macroalgae. Of 29 species of larger herbivorous fishes on these reefs, only four species accounted for 97% of macroalgal consumption. Two unicornfish consumed a range of brown macroalgae, a parrotfish consumed multiple red algae, and a rabbitfish consumed a green alga, with almost no diet overlap among these groups. The two most chemically rich, allelopathic algae were each consumed by a single, but different, fish species. This striking complementarity resulted from herbivore species differing in their tolerances to macroalgal chemical and structural defenses. A model of assemblage diet breadth based on our feeding observations predicted that high browser diversity would be required for effective control of macroalgae on Fijian reefs. In support of this model, we observed strong negative relationships between herbivore diversity and macroalgal abundance and diversity across the six study reefs. Our findings indicate that the total diet breadth of the herbivore community and the probability of all macroalgae being removed from reefs by herbivores increases with increasing herbivore diversity, but that a few critical species drive this relationship. Therefore, interactions between algal defenses and herbivore tolerances create an essential role for consumer diversity in the functioning and resilience of coral reefs.

Do invasive species perform better in their new ranges

Abstract: A fundamental assumption in invasion biology is that most invasive species exhibit enhanced performance in their introduced range relative to their home ranges. This idea has given rise to numerous hypotheses explaining ''invasion success'' by virtue of altered ecological and evolutionary pressures. There are surprisingly few data, however, testing the underlying assumption that the performance of introduced populations, including organism size, reproductive output, and abundance, is enhanced in their introduced compared to their native range. Here, we combined data from published studies to test this hypothesis for 26 plant and 27 animal species that are considered to be invasive. On average, individuals of these 53 species were indeed larger, more fecund, and more abundant in their introduced ranges. The overall mean, however, belied significant variability among species, as roughly half of the investigated species (N ¼ 27) performed similarly when compared to conspecific populations in their native range. Thus, although some invasive species are performing better in their new ranges, the pattern is not universal, and just as many are performing largely the same across ranges.

Animal migration amid shifting patterns of phenology and predation: lessons from a Yellowstone elk herd

Abstract: Migration is a striking behavioral strategy by which many animals enhance resource acquisition while reducing predation risk Historically, the demographic benefits of such movements made migration common, but in many taxa the phenomenon is considered globally threatened Here we describe a long-term decline in the productivity of elk (Cervus elaphus) that migrate through intact wilderness areas to protected summer ranges inside Yellowstone National Park, USA We attribute this decline to a long-term reduction in the demographic benefits that ungulates typically gain from migration Among migratory elk, we observed a 21-year, 70% reduction in recruitment and a 4-year, 19% depression in their pregnancy rate largely caused by infrequent reproduction of females that were young or lactating In contrast, among resident elk, we have recently observed increasing recruitment and a high rate of pregnancy Landscape-level changes in habitat quality and predation appear to be responsible for the declining productivity of Yellowstone migrants From 1989 to 2009, migratory elk experienced an increasing rate and shorter duration of green-up coincident with warmer spring-summer temperatures and reduced spring precipitation, also consistent with observations of an unusually severe drought in the region Migrants are also now exposed to four times as many grizzly bears (Ursus arctos) and wolves (Canis lupus) as resident elk Both of these restored predators consume migratory elk calves at high rates in the Yellowstone wilderness but are maintained at low densities via lethal management and human disturbance in the year-round habitats of resident elk Our findings suggest that large-carnivore recovery and drought, operating simultaneously along an elevation gradient, have disproportionately influenced the demography of migratory elk Many migratory animals travel large geographic distances between their seasonal ranges Changes in land use and climate that disparately influence such seasonal ranges may alter the ecological basis of migratory behavior, representing an important challenge for, and a powerful lens into, the ecology and conservation of migratory taxa

Sensitivity of grassland plant community composition to spatial vs. temporal variation in precipitation

Abstract: Climate gradients shape spatial variation in the richness and composition of plant communities. Given future predicted changes in climate means and variability, and likely regional variation in the magnitudes of these changes, it is important to determine how temporal variation in climate influences temporal variation in plant community structure. Here, we evaluated how species richness, turnover, and composition of grassland plant communities responded to interannual variation in precipitation by synthesizing long-term data from grasslands across the United States. We found that mean annual precipitation,(MAP) was a positive predictor of species richness across sites, but a positive temporal relationship between annual precipitation and richness was only evident within two sites with low MAP. We also found higher average rates of species turnover in dry sites that in turn had a high proportion of annual species, although interannual rates of species turnover were surprisingly high across all locations. Annual species were less abundant than perennial species at nearly all sites, and our analysis showed that the probability of a species being lost or gained from one year to the next increased with decreasing species abundance. Bray-Curtis dissimilarity from one year to the next, a measure of species composition change that is influenced mainly by abundant species, was insensitive to precipitation at all sites. These results suggest that the richness and turnover patterns we observed were driven primarily by rare species, which comprise the majority of the local species pools at these grassland sites. These findings are consistent with the idea that short-lived and less abundant species are more sensitive to interannual climate variability than longer-lived and more abundant species. We conclude that, among grassland ecosystems, xeric grasslands are likely to exhibit the greatest responsiveness of community composition (richness and turnover) to predicted future increases in interannual precipitation variability. Over the long-term, species composition may shift to reflect spatial patterns of mean precipitation; however, perennial-dominated systems will be buffered against rising interannual variation, while systems that have a large number of rare, annual species will show the greatest temporal variability in species composition in response to rising interannual variability in precipitation.

Precipitation legacies in desert grassland primary production occur through previous‐year tiller density

Abstract: In arid ecosystems, current-year precipitation often explains only a small proportion of annual aboveground net primary production (ANPP). We hypothesized that lags in the response of ecosystems to changes in water availability explain this low explanatory power, and that lags result from legacies from transitions from dry to wet years or the reverse. We explored five hypotheses regarding the magnitude of legacies, two possible mechanisms, and the differential effect of previous dry or wet years on the legacy magnitude. We used a three-year manipulative experiment with five levels of rainfall in the first two years (� 80% and � 50% reduced annual precipitation (PPT), ambient, þ50% and þ80% increased PPT), and reversed treatments in year 3. Legacies of previous two years, which were dry or wet, accounted for a large fraction (20%) of interannual variability in production on year 3. Legacies in ANPP were similar in absolute value for both types of precipitation transitions, and their magnitude was a function of the difference between previous and current-year precipitation. Tiller density accounted for 40% of legacy variability, while nitrogen and carry- over water availability showed no effect. Understanding responses to changes in interannual precipitation will assist in assessing ecosystem responses to climate change-induced increases in precipitation variability.

Influence of competition and rainfall manipulation on the growth responses of savanna trees and grasses

Abstract: In this study, we explored how rainfall manipulation influenced competitive interactions between grasses and juvenile trees (small nonreproductive trees capable of resprouting) in savanna. To do this, we manipulated rainfall amount in the field using an incomplete factorial experiment that determined the effects of rainfall reduction, no manipulation, rainfall addition, and competition between grasses and trees on grass and tree growth. As response variables, we focused on several measures of tree growth and Disc Pasture Meter settling height as an estimate of grass aboveground biomass. We conducted the study over four years, at two sites in the Kruger National Park, South Africa. Our results show that rainfall manipulation did not have substantial effects on any of the measures of tree growth we considered. However, trees at plots where grasses had been removed grew on average 15 cm more in height and 1.3-1.7 times more in basal area per year than those in plots with grasses. Grass biomass was not influenced by the presence of trees but was significantly and positively influenced by rainfall addition. These findings were not fundamentally influenced by soil type or by prevailing precipitation, suggesting applicability of our results to a wide range of savannas. Our results suggest that, in savannas, increasing rainfall serves to increase the competitive pressure exerted by grasses on trees. The implication is that recruitment into the adult tree stage from the juvenile stage is most likely in drought years when there is little competition from grass for resources and grass fuel loads are low.

Benthic algae stimulate leaf litter decomposition in detritus‐based headwater streams: a case of aquatic priming effect?

Abstract: In detritus-based ecosystems, autochthonous primary production contributes very little to the detritus pool. Yet primary producers may still influence the functioning of these ecosystems through complex interactions with decomposers and detritivores. Recent studies have suggested that, in aquatic systems, small amounts of labile carbon (C) (e.g., producer exudates), could increase the mineralization of more recalcitrant organic-matter pools (e.g., leaf litter). This process, called priming effect, should be exacerbated under low- nutrient conditions and may alter the nature of interactions among microbial groups, from competition under low-nutrient conditions to indirect mutualism under high-nutrient conditions. Theoretical models further predict that primary producers may be competitively excluded when allochthonous C sources enter an ecosystem. In this study, the effects of a benthic diatom on aquatic hyphomycetes, bacteria, and leaf litter decomposition were investigated under two nutrient levels in a factorial microcosm experiment simulating detritus- based, headwater stream ecosystems. Contrary to theoretical expectations, diatoms and decomposers were able to coexist under both nutrient conditions. Under low-nutrient conditions, diatoms increased leaf litter decomposition rate by 20% compared to treatments where they were absent. No effect was observed under high-nutrient conditions. The increase in leaf litter mineralization rate induced a positive feedback on diatom densities. We attribute these results to the priming effect of labile C exudates from primary producers. The presence of diatoms in combination with fungal decomposers also promoted decomposer diversity and, under low-nutrient conditions, led to a significant decrease in leaf litter C:P ratio that could improve secondary production. Results from our microcosm experiment suggest new mechanisms by which primary producers may influence organic matter dynamics even in ecosystems where autochthonous primary production is low.

Effects of predator richness on prey suppression: a meta-analysis

Abstract: It is well established that species richness of primary producers and primary consumers can enhance efficiency of resource uptake and biomass production of respective trophic levels. At the level of secondary consumers (predators), however, conclusions about the functional role of biodiversity have been mixed. We take advantage of a recent surge of published experiments (totaling 46 since 2005) to both evaluate general effects of predator richness on aggregate prey suppression (top-down control) and explore sources of variability among experiments. Our results show that, across experiments, predator richness enhances prey suppression relative to the average single predator species (mean richness effect), but not the best-performing species. Mean richness effects in predator experiments were stronger than those for primary producers and detritivores, suggesting that relationships between richness and function may increase with trophic height in food webs. The strength of mean predator richness effects increased with the spatial and temporal scale of experiments, and the taxonomic distinctness (TD, used as a proxy of phylogenetic diversity) of species present. This latter result suggests that TD captures important aspects of functional differentiation among predators and that measures of biodiversity that go beyond species richness may help to better predict the effects of predator species loss.

Fire drives functional thresholds on the savanna–forest transition

Abstract: In tropical landscapes, vegetation patches with contrasting tree densities are distributed as mosaics. However, the locations of patches and densities of trees within them cannot be predicted by climate models alone. It has been proposed that plant-fire feedbacks drive functional thresholds at a landscape scale, thereby maintaining open (savanna) and closed (forest) communities as two distinct stable states. However, there is little rigorous field evidence for this threshold model. Here we aim to provide support for such a model from a field perspective and to analyze the functional and phylogenetic consequences of fire in a Brazilian savanna landscape (Cerrado). We hypothesize that, in tropical landscapes, savanna and forest are two stable states maintained by plant-fire feedbacks. If so, their functional and diversity attributes should change abruptly along a community closure gradient. We set 98 plots along a gradient from open savanna to closed forest in the Brazilian Cerrado and tested for a threshold pattern in nine functional traits, five soil features, and seven diversity indicators. We then tested whether the threshold pattern was associated with different fire regimes. Most community attributes presented a threshold pattern on the savanna-forest transition with coinciding breakpoints. The thresholds separated two community states: (1) open environments with low-diversity communities growing in poor soils and dominated by plants that are highly resistant to high-intensity fires; and (2) closed environments with highly diverse plant communities growing in more fertile soils and dominated by shade-tolerant species that efficiently prevent light from reaching the understory. In addition, each state was associated with contrasting fire regimes. Our results are consistent with the hypothesis that forests and savannas are two coexisting stable states with contrasting patterns of function and diversity that are regulated by fire- plant feedbacks; our results also shed light on the mechanism driving each state. Overall, our results support the idea that fire plays an important role in regulating the distribution of savanna and forest biomes in tropical landscapes.

A direct test of nitrogen and phosphorus limitation to net primary productivity in a lowland tropical wet forest

Abstract: Experimental evidence for limitation of net primary productivity (NPP) by nitrogen (N) or phosphorus (P) in lowland tropical forests is rare, and the results from the few existing studies have been inconclusive. To directly test if N or P limit NPP in a lowland tropical wet forest in Costa Rica, we conducted a full factorial fertilization experiment (4 treatments × 6 replicates in 30 × 30 m plots). We focused on the influence of tree size and taxa on nutrient limitation, because in these forests a wide variety of tree functional traits related to nutrient acquisition and use are likely to regulate biogeochemical processes. After 2.7 years, a higher percentage of trees per plot increased basal area (BA) with P additions (66.45% ± 3.28% without P vs. 76.88% ± 3.28% with P), but there were no other community-level responses to N or P additions on BA increase, litterfall productivity, or root growth. Phosphorus additions resulted in doubled stem growth rates in small trees (5–10 cm diameter at breast height (dbh); [P ≤ 0.01]) but had no effect on intermediate (10–30 cm dbh) or large trees (>30 cm dbh). Phosphorus additions also increased the percentage of seedling survival from 59% to 78% (P < 0.01), as well as the percentage of seedlings that grew (P = 0.03), and increased leaf number (P = 0.02). Trees from Pentaclethra macroloba, the most abundant species, did not increase growth rates with fertilization (P = 0.40). In contrast, the most abundant palms (Socratea exorrhiza) had more than two times higher stem growth rates with P additions (P = 0.01). Our experiment reiterates that P availability is a significant driver of plant processes in these systems, but highlights the importance of considering different aspects of the plant community when making predictions concerning nutrient limitation. We postulate that in diverse, lowland tropical forests “heterogeneous nutrient limitation” occurs, not only driven by variability in nutrient responses among taxa, but also among size classes and potential functional groups. Heterogeneous responses to nutrient additions could lead to changes in forest structure or even diversity in the long term, affecting rates of NPP and thus carbon cycling.

Mycorrhizas in the Central European flora: relationships with plant life history traits and ecology

Abstract: Plant traits have been widely used to characterize different aspects of the ecology of plant species. Despite its wide distribution and its proven significance at the level of individuals, communities, and populations, the ability to form mycorrhizal associations has been largely neglected in these studies so far. Analyzing plant traits associated with the occurrence of mycorrhizas in plants can therefore enhance our understanding of plant strategies and distributions. Using a comparative approach, we tested for associations between mycorrhizal status and habitat characteristics, life history traits, and plant distribution patterns in 1752 species of the German flora (a major part of the Central European flora). Data were analyzed using log-linear models or generalized linear models, both accounting for phylogenetic relationships. Obligatorily mycorrhizal (OM) species tended to be positively associated with higher temperature, drier habitats, and higher pH; and negatively associated with moist, acidic, an...

Diel horizontal migration in streams: juvenile fish exploit spatial heterogeneity in thermal and trophic resources.

Abstract: Vertical heterogeneity in the physical characteristics of lakes and oceans is ecologically salient and exploited by a wide range of taxa through diel vertical migration to enhance their growth and survival. Whether analogous behaviors exploit horizontal habitat heterogeneity in streams is largely unknown. We investigated fish movement behavior at daily timescales to explore how individuals integrated across spatial variation in food abundance and water temperature. Juvenile coho salmon made feeding forays into cold habitats with abundant food, and then moved long distances (350–1300 m) to warmer habitats that accelerated their metabolism and increased their assimilative capacity. This behavioral thermoregulation enabled fish to mitigate trade-offs between trophic and thermal resources by exploiting thermal heterogeneity. Fish that exploited thermal heterogeneity grew at substantially faster rates than did individuals that assumed other behaviors. Our results provide empirical support for the importance of...

Historical thermal regimes define limits to coral acclimatization

Abstract: Knowledge of the degree to which corals undergo physiological acclimatization or genetic adaptation in response to changes in their thermal environment is crucial to the success of coral reef conservation strategies. The potential of corals to acclimatize to temperatures exceeding historical thermal regimes was investigated by reciprocal transplantation of Acropora millepora colonies between the warm central and cool southern regions of the Great Barrier Reef (GBR) for a duration of 14 months. Colony fragments retained at native sites remained healthy, whereas transplanted fragments, although healthy over initial months when temperatures remained within native thermal regimes, subsequently bleached and suffered mortality during seasonal temperature extremes. Corals hosting Symbiodinium D transplanted to the southern GBR bleached in winter and the majority suffered whole (40%; n=20 colonies) or partial (50%) mortality at temperatures 1.1 degrees C below their 15-year native minimum. In contrast, corals hosting Symbiodinium C2 transplanted to the central GBR bleached in summer and suffered whole (50%; n=10 colonies) or partial (42%) mortality at temperatures 2.5 degrees C above their 15-year native maximum. During summer bleaching, the dominant Symbiodinium type changed from C2 to D within corals transplanted to the central GBR. Corals transplanted to the cooler, southern GBR grew 74-80% slower than corals at their native site, and only 50% of surviving colonies reproduced, at least partially because of cold water bleaching of transplants. Despite the absence of any visual signs of stress, corals transplanted to the warmer, central GBR grew 52-59% more slowly than corals at their native site before the summer bleaching (i.e., from autumn to spring). Allocation of energy to initial acclimatization or reproduction may explain this pattern, as the majority (65%) of transplants reproduced one month earlier than portions of the same colonies retained at the southern native site. All parameters investigated (bleaching, mortality, Symbiodinium type fidelity, reproductive timing) demonstrated strong interactions between genotype and environment, indicating that the acclimatization potential of A. millepora populations may be limited by adaptation of the holobiont to native thermal regimes.

Yeasts in nectar of an early‐blooming herb: sought by bumble bees, detrimental to plant fecundity

Abstract: Through their effects on physicochemical features of floral nectar, nectar-dwelling yeasts can alter pollinator behavior, but the effect of such changes on pollination success and plant reproduction is unknown. We present results of experiments testing the effects of nectar yeasts on foraging patterns of captive and free-ranging bumble bees, and also on pollination success and fecundity of the early-blooming, bumble bee-pollinated Helleborus foetidus (Ranunculaceae). Under controlled experimental conditions, inexperienced Bombus terrestris workers responded positively to the presence of yeasts in artificial sugar solutions mimicking floral nectar by visiting proportionally more yeast-containing artificial flowers. Free-ranging bumble bees also preferred yeast-containing nectar in the field. Experiments conducted in two different years consistently showed that natural and artificial nectars containing yeasts were more thoroughly removed than nectars without yeasts. Experimental yeast inoculation of the nectar of H. foetidus flowers was significantly associated with reductions in number of pollen tubes in the style, fruit set, seed set, and mass of individual seeds produced. These results provide the first direct evidence to date that nectar yeasts can modify pollinator foraging patterns, pollination success, and the quantity and quality of seeds produced by insect-pollinated plants.

Biodiversity simultaneously enhances the production and stability of community biomass, but the effects are independent

Abstract: To predict the ecological consequences of biodiversity loss, researchers have spent much time and effort quantifying how biological variation affects the magnitude and stability of ecological processes that underlie the functioning of ecosystems. Here we add to this work by looking at how biodiversity jointly impacts two aspects of ecosystem functioning at once: (1) the production of biomass at any single point in time (biomass/area or biomass/ volume), and (2) the stability of biomass production through time (the CV of changes in total community biomass through time). While it is often assumed that biodiversity simultaneously enhances both of these aspects of ecosystem functioning, the joint distribution of data describing how species richness regulates productivity and stability has yet to be quantified. Furthermore, analyses have yet to examine how diversity effects on production covary with diversity effects on stability. To overcome these two gaps, we reanalyzed the data from 34 experiments that have manipulated the richness of terrestrial plants or aquatic algae and measured how this aspect of biodiversity affects community biomass at multiple time points. Our reanalysis confirms that biodiversity does indeed simultaneously enhance both the production and stability of biomass in experimental systems, and this is broadly true for terrestrial and aquatic primary producers. However, the strength of diversity effects on biomass production is independent of diversity effects on temporal stability. The independence of effect sizes leads to two important conclusions. First, while it may be generally true that biodiversity enhances both productivity and stability, it is also true that the highest levels of productivity in a diverse community are not associated with the highest levels of stability. Thus, on average, diversity does not maximize the various aspects of ecosystem functioning we might wish to achieve in conservation and management. Second, knowing how biodiversity affects productivity gives no information about how diversity affects stability (or vice versa). Therefore, to predict the ecological changes that occur in ecosystems after extinction, we will need to develop separate mechanistic models for each independent aspect of ecosystem functioning.

Aggregated filter-feeding consumers alter nutrient limitation: consequences for ecosystem and community dynamics.

Abstract: Nutrient cycling is a key process linking organisms in ecosystems. This is especially apparent in stream environments in which nutrients are taken up readily and cycled through the system in a downstream trajectory. Ecological stoichiometry predicts that biogeochemical cycles of different elements are interdependent because the organisms that drive these cycles require fixed ratios of nutrients. There is growing recognition that animals play an important role in biogeochemical cycling across ecosystems. In particular, dense aggregations of consumers can create biogeochemical hotspots in aquatic ecosystems via nutrient translocation. We predicted that filter-feeding freshwater mussels, which occur as speciose, high-biomass aggregates, would create biogeochemical hotspots in streams by altering nutrient limitation and algal dynamics. In a field study, we manipulated nitrogen and phosphorus using nutrient-diffusing substrates in areas with high and low mussel abundance, recorded algal growth and community composition, and determined in situ mussel excretion stoichiometry at 18 sites in three rivers (Kiamichi, Little, and Mountain Fork Rivers, south-central United States). Our results indicate that mussels greatly influence ecosystem processes by modifying the nutrients that limit primary productivity. Sites without mussels were N-limited with -26% higher relative abundances of N-fixing blue-green algae, while sites with high mussel densities were co-limited (N and P) and dominated by diatoms. These results corroborated the results of our excretion experiments; our path analysis indicated that mussel excretion has a strong influence on stream water column N:P. Due to the high N:P of mussel excretion, strict N-limitation was alleviated, and the system switched to being co-limited by both N and P. This shows that translocation of nutrients by mussel aggregations is important to nutrient dynamics and algal species composition in these rivers. Our study highlights the importance of consumers and this imperiled faunal group on nutrient cycling and community dynamics in aquatic ecosystems.

Spatial occupancy models for large data sets

Abstract: Since its development, occupancy modeling has become a popular and useful tool for ecologists wishing to learn about the dynamics of species occurrence over time and space. Such models require presence–absence data to be collected at spatially indexed survey units. However, only recently have researchers recognized the need to correct for spatially induced overdisperison by explicitly accounting for spatial autocorrelation in occupancy probability. Previous efforts to incorporate such autocorrelation have largely focused on logit-normal formulations for occupancy, with spatial autocorrelation induced by a random effect within a hierarchical modeling framework. Although useful, computational time generally limits such an approach to relatively small data sets, and there are often problems with algorithm instability, yielding unsatisfactory results. Further, recent research has revealed a hidden form of multicollinearity in such applications, which may lead to parameter bias if not explicitly addressed. Com...

Telemetry and random-walk models reveal complex patterns of partial migration in a large marine predator

Abstract: Animals are often faced with complex movement decisions, particularly those that involve long-distance dispersal. Partial migrations, ubiquitous among all groups of vertebrates, are a form of long-distance movement that occurs when only some of the animals in a population migrate. The decision to migrate or to be a resident can be dependent on many factors, but these factors are rarely quantified in fishes, particularly top predators, even though partial migrations may have important implications for ecosystem dynamics and conservation. We utilized passive acoustic telemetry, with a Brownian bridge movement model and generalized additive mixed models, to explore the factors regulating partial migration in a large marine predator, the tiger shark, throughout the Hawaiian Islands. Although sharks tended to utilize a particular “core” island, they also demonstrated inter-island movements, particularly mature females that would swim from the northwestern Hawaiian Islands to the main Hawaiian Islands (MHI). Im...

Life-history traits and landscape characteristics predict macro-moth responses to forest fragmentation

Abstract: How best to manage forest patches, mitigate the consequences of forest fragmentation, and enable landscape permeability are key questions facing conservation scientists and managers. Many temperate forests have become increasingly fragmented, resulting in reduced interior forest habitat, increased edge habitats, and reduced connectivity. Using a citizen science landscape-scale mark-release-recapture study on 87 macro-moth species, we investigated how both life-history traits and landscape characteristics predicted macro-moth responses to forest fragmentation. Wingspan, wing shape, adult feeding, and larval feeding guild predicted macro-moth mobility, although the predictive power of wingspan and wing shape depended on the species' affinity to the forest. Solitary trees and small fragments functioned as "stepping stones," especially when their landscape connectivity was increased, by being positioned within hedgerows or within a favorable matrix. Mobile forest specialists were most affected by forest fragmentation: despite their high intrinsic dispersal capability, these species were confined mostly to the largest of the forest patches due to their strong affinity for the forest habitat, and were also heavily dependent on forest connectivity in order to cross the agricultural matrix. Forest fragments need to be larger than five hectares and to have interior forest more than 100 m from the edge in order to sustain populations of forest specialists. Our study provides new insights into the movement patterns of a functionally important insect group, with implications for the landscape-scale management of forest patches within agricultural landscapes.