Legacy Effects Research Articles

Declined benefit of earlier spring greening on summer growth in northern ecosystems under future scenarios

Widespread spring warming and an earlier start of the growing season (SOS) compensated for spring vegetation productivity, simultaneously resulting in reduced retention of accessible nutrition and water resources in subsequent seasons. In this case, however, it is unknown whether the increased summer growth induced by earlier SOS will be sustained. Here, we analyzed the legacy effects of SOS on summer growth under historical and future scenarios, through combining remote sensing products, flux tower observations and model simulations. We observed that the background climate (summer temperature and summer precipitation) modulated the response of summer growth to SOS. In the past two decades, the advance of SOS promoted summer growth in most areas of the northern ecosystems, except for dry and hot regions. Moreover, the effects of SOS on plant growth reversed within the summer months, which may be related to the continued deterioration of moisture conditions. Models predicted a general shift from a negative to a positive correlation between SOS and summer growth under future warming and regionally uneven precipitation. In particular, under SSP370 and SSP585 scenarios, areas of positive correlation will exceed that of negative correlation by the end of the century. These results indicated the stimulation of summer growth by SOS advance may shift to inhibition in more regions in the future. Our findings highlighted the limited enhancement of plant carbon sink by altered spring phenology under future climate change.

Legacy effects of premature defoliation in response to an extreme drought event modulate phytochemical profiles with subtle consequences for leaf herbivory in European beech.

Extreme droughts can have long-lasting effects on forest community dynamics and species interactions. Yet, our understanding of how drought legacy modulates ecological relationships is just unfolding. We tested the hypothesis that leaf chemistry and herbivory show long-term responses to premature defoliation caused by an extreme drought event in European beech (Fagus sylvatica L.). For two consecutive years after the extreme European summer drought in 2018, we collected leaves from the upper and lower canopy of adjacently growing drought-stressed and unstressed trees. Leaf chemistry was analyzed and leaf damage by different herbivore-feeding guilds was quantified. We found thatdrought had lasting impacts on leaf nutrients and on specialized metabolomic profiles. However, drought did not affect the primary metabolome. Drought-related phytochemical changes affected damage of leaf-chewing herbivores whereas damage caused by other herbivore-feeding guilds was largely unaffected. Drought legacy effects on phytochemistry and herbivory were often weaker than between-year or between-canopy strata variability. Our findings suggest that a single extreme drought event bears the potential to long-lastingly affect tree-herbivore interactions. Drought legacy effects likely become more important in modulating tree-herbivore interactions since drought frequency and severity are projected to globally increase in the coming decades.

Grasses and shrubs have decoupled legacy effects related to plant nutrition: the interplay of symbiotic fungi and nutrients in soil

Grasses and shrubs have decoupled legacy effects related to plant nutrition: the interplay of symbiotic fungi and nutrients in soil

Water level drawdown induces a legacy effect on the seed bank and retains sediment chemistry in a eutrophic clay wetland

The lack of extreme water level fluctuations in managed, non-peat forming wetland ecosystems can result in decreased productivity through the loss of heterogeneity of these ecosystems. Stochastic disruption, such as a water level drawdown, can effectively reverse this effect and return the wetland to a more productive state, associated with higher biodiversity through new vegetation development. Yet, aside from the effect on vegetation dynamics, little is known about longer-term effects (30 years) of a water level drawdown, hereafter referred to as legacy effects, and how this may impact future water level drawdowns.Here, we aim to unravel the legacy effects of a water level drawdown, stand alone and along a water level gradient, on seed bank properties and nutrient availability in a eutrophic clay wetland. To identify these, we studied the hydrologically managed nature reserve Oostvaardersplassen in the Netherlands. Here, one section was subjected to a multi-year water level drawdown and another section was kept inundated. We determined seed bank properties in both areas, spatially and along a soil elevation gradient (20 cm). Nutrient availability was measured by taking sediment samples along the water level gradient and through experimental manipulation of the water level in an indoor mesocosm experiment.Germination was higher in locations with a water level drawdown history, especially at relatively high elevations. Additionally, the proportion of pioneer species in the seed bank was higher in the water level drawdown area. Overall, nutrient concentrations were higher compared to other aquatic systems. Nutrient availability was higher in the inundated area and did not respond to the water level gradient. We conclude that 30 years after an induced water level drawdown there is no depletion of nutrients, while we still observe a legacy effect in the number of viable seeds in the seed bank.

Assessing the influence of climate on the growth of green ash trees from five Plant Hardiness Zones growing in a range-wide provenance test near the species’ northern range limit

Green ash is threatened with extirpation from emerald ash borer (EAB) attack. Because green ash has an extensive range, its restoration could require both breeding for EAB resistance and possible adaptation to a variety of environmental gradients. We assessed the growth and climate sensitivity of green ash from five Plant Hardiness Zones (PHZs) growing in a range-wide provenance test in Vermont. Although there tended to be greater growth among trees from the warmest PHZ (7), differences were rarely statistically distinguishable. For trees from all PHZs, growth was positively correlated with precipitation in the current year and negatively correlated with precipitation the year before (a possible legacy effect). Growth was negatively associated with temperature the year of ring formation but positively associated with temperatures the year before. Growth was often positively correlated with winter snow but was negatively associated with spring or fall snow for the warmest PHZs. Climate correlations for PHZ 3 were unusual in that (1) only positive correlations were detected, (2) no legacy effects were noted, and (3) despite being from the coldest region, no correlations with snow were found. Growth increased over time for the warmest PHZs during a period of simultaneous increases in temperature and precipitation.

Seasonal aridity regulates drivers and temporal variability of wood phenology: A meta-analysis of dendrometer monitoring data across the Northern Hemisphere

The secondary growth phenology of trees, also known as wood phenology, plays a vital role in assessing the timing of carbon sequestration in forests. However, its spatio-temporal variability and determinants in the Northern Hemisphere remain debated. Here, we presented a meta-analysis using secondary growth phenology data based on dendrometer monitoring of 59 tree species and 84 sites to explore the drivers and temporal connections of phenology across seasonal arid and humid regions in the Northern Hemisphere. We found spring temperatures and precipitation co-regulated the start of secondary growth (SOG) in spring arid regions, and SOG advanced with the increase in spring temperatures and precipitation. However, SOG was only influenced by spring temperatures in spring humid regions. The end of secondary growth (EOG) indicated strong correlations with autumn temperatures only in autumn arid regions but not in humid regions. Interestingly, the earlier SOG promoted advancement of the time of maximum secondary growth rate (MOG) and EOG both in seasonal humid and arid regions, while the end of secondary growth in the previous year (prEOG) negatively affected SOG only in seasonal arid regions. The legacy effects of previous phenological events on subsequent ones should be integrated into phenological prediction models to help accurately assess global carbon, water, and energy cycles.

Tree post-drought recovery: scenarios, regulatory mechanisms and ways to improve.

Efficient post-drought recovery of growth and assimilation enables a plant to return to its undisturbed state and functioning. Unlike annual plants, trees suffer not only from the current drought, but also from cumulative impacts of consecutive water stresses which cause adverse legacy effects on survival and performance. This review provides an integrated assessment of ecological, physiological and molecular evidence on the recovery of growth and photosynthesis in trees, with a view to informing the breeding of trees with a better ability to recover from water stress. Suppression of recovery processes can result not only from stress damage but also from a controlled downshift of recovery as part of tree acclimation to water-limited conditions. In the latter case, recovery processes could potentially be activated by turning off the controlling mechanisms, but several obstacles make this unlikely. Tree phenology, and specifically photoperiodic constraints, can limit post-drought recovery of growth and photosynthesis, and targeting these constraints may represent a promising way to breed trees with an enhanced ability to recover post-drought. The mechanisms of photoperiod-dependent regulation of shoot, secondary and root growth and of assimilation processes are reviewed. Finally, the limitations and trade-offs of altering the photoperiodic regulation of growth and assimilation processes are discussed.

Drought legacies in mixed Mediterranean forests: Analysing the effects of structural overshoot, functional traits and site factors

Previous favorable climate conditions stimulate tree growth making some forests more vulnerable to hotter droughts. This so-called structural overshoot may contribute to forest dieback, but there is little evidence on its relative importance depending on site conditions and tree species because of limited field data. Here, we analyzed remote sensing (NDVI) and tree-ring width data to evaluate the impacts of the 2017 drought on canopy cover and growth in mixed Mediterranean forests (Fraxinus ornus, Quercus pubescens, Acer monspessulanum, Pinus pinaster) located in southern Italy. Legacy effects were assessed by calculating differences between observed and predicted basal area increment (BAI). Overall, the growth response of the study stands to the 2017 drought was contingent on site conditions and species characteristics. Most sites presented BAI and canopy cover reductions during the drought. Growth decline was followed by a quick recovery and positive legacy effects, particularly in the case of F. ornus. However, we found negative drought legacies in some species (e.g., Q. pubescens, A. monspessulanum) and sites. In those sites showing negative legacies, high growth rates prior to drought in response to previous wet winter-spring conditions may have predisposed trees to drought damage. Vice versa, the positive drought legacy found in some F. ornus site was linked to post-drought growth release due to Q. pubescens dieback and mortality. Therefore, we found evidences of structural drought overshoot, but it was restricted to specific sites and species. Our findings highlight the importance of considering site settings such as stand composition, pre-drought conditions and different tree species when studying structural overshoot. Droughts contribute to modify the composition and dynamics in mixed forests.

Diversified grain rotations can be highly and reliably productive in unstable climates

The crop rotation is a farm-scale management choice that dictates agronomic output, ecological impacts, and farm viability. Rotations have become less diverse recently. Re-diversifying may help agricultural systems meet the growing, dynamic demands from society and challenges from climate change. Using a long-term experiment, we tested whether diversified rotations could a) match the productivity (grain yield) of simplified rotations while b) stabilizing productivity against variable weather in the western U.S. Corn Belt. We identified rotation design choices that drive these productivity-stability dynamics. Although more productive rotations were more susceptible to weather (r = 0.63; p < 0.001), this tradeoff was surmountable (p < 0.001): the corn-oat-winter wheat-soybean (Zea mays L., Avena sativa L., Triticum aestivum L., Glycine max (L.) Merr.) rotation was a) no less productive than the highly unstable, corn-soybean rotation (p = 0.11); and b) only marginally less stable than the most stable rotation (p = 0.06). Crop selection and sequencing were critical to this outcome. High productivity was due to a) overyielding of individual crops in diversified rotations that increased with time; b) sequencing to allow higher-yielding winter crops; and c) beneficial crop legacy effects. Stability was highest in rotations that included crops belonging to more weather-niches (r2 = 0.67; p < 0.001). Surprisingly, these weather-niches groups did not correspond to traditional categories like cool- vs warm-season crops. These results suggest four principles for the design, study, and implementation of diversified grain rotations that are stable under erratic weather and are as productive as current standard practices.

Land use legacies affect early tropical forest succession in Mexico

AbstractQuestionsAgricultural expansion is one of the dominant drivers of forest and biodiversity loss, and shifting cultivation is the most widely used form of agriculture in many tropical forest regions. Where forests have been cleared, they have the potential to recover once the land is abandoned. However, legacies of land use are often overlooked in successional studies, and a deeper understanding of this legacy effect is needed to define efficient restoration practices using natural or assisted regeneration. Here, we analysed how land‐use history affects soil properties and early succession on abandoned agricultural fields in two contrasting Mexican socio‐ecological systems.LocationMexico, Oaxaca and Chiapas.MethodsWe sampled soil and monitored vegetation for 2 years after agricultural abandonment, and interviewed landowners about their land‐use practices.ResultsLand‐use practices were clearly influenced by landowners’ social context (residence time, rural or urban origin), and topography and soil type also constrained or facilitated land‐use practices. Soil characteristics were strongly affected by three land‐use practices: mechanical tillage decreased soil N and K; frequent herbicide and pesticide use increased N and K; and for pasture systems, stocking density increased soil bulk density and decreased pH and N. High‐intensity land management practices, specifically use of machinery, had the highest impact on early forest succession. When machinery was not used, the frequency of land‐use practices, particularly weeding frequency, is the main factor influencing tree cover and sapling diversity.ConclusionsTo facilitate post‐agricultural forest recovery, we recommend restoration efforts using natural regeneration in areas with low previous land‐use intensity and frequency.

Revealing legacy effects of extreme droughts on tree growth of oaks across the Northern Hemisphere

Forests are undergoing increasing risks of drought-induced tree mortality. Species replacement patterns following mortality may have a significant impact on the global carbon cycle. Among major hardwoods, deciduous oaks (Quercus spp.) are increasingly reported as replacing dying conifers across the Northern Hemisphere. Yet, our knowledge on the growth responses of these oaks to drought is incomplete, especially regarding post-drought legacy effects. The objectives of this study were to determine the occurrence, duration, and magnitude of legacy effects of extreme droughts and how that vary across species, sites, and drought characteristics. The legacy effects were quantified by the deviation of observed from expected radial growth indices in the period 1940–2016. We used stand-level chronologies from 458 sites and 21 oak species primarily from Europe, north-eastern America, and eastern Asia. We found that legacy effects of droughts could last from 1 to 5 years after the drought and were more prolonged in dry sites. Negative legacy effects (i.e., lower growth than expected) were more prevalent after repetitive droughts in dry sites. The effect of repetitive drought was stronger in Mediterranean oaks especially in Quercus faginea. Species-specific analyses revealed that Q. petraea and Q. macrocarpa from dry sites were more negatively affected by the droughts while growth of several oak species from mesic sites increased during post-drought years. Sites showing positive correlations to winter temperature showed little to no growth depression after drought, whereas sites with a positive correlation to previous summer water balance showed decreased growth. This may indicate that although winter warming favors tree growth during droughts, previous-year summer precipitation may predispose oak trees to current-year extreme droughts. Our results revealed a massive role of repetitive droughts in determining legacy effects and highlighted how growth sensitivity to climate, drought seasonality and species-specific traits drive the legacy effects in deciduous oak species.

Assessing the response of an urban stream ecosystem to salinization under different flow regimes

Urban streams are exposed to a variety of anthropogenic stressors. Freshwater salinization is a key stressor in these ecosystems that is predicted to be further exacerbated by climate change, which causes simultaneous changes in flow parameters, potentially resulting in non-additive effects on aquatic ecosystems. However, the effects of salinization and flow velocity on urban streams are still poorly understood as multiple-stressor experiments are often conducted at pristine rather than urban sites. Therefore, we conducted a mesocosm experiment at the Boye River, a recently restored stream located in a highly urbanized area in Western Germany, and applied recurrent pulses of salinity along a gradient (NaCl, 9 h daily of +0 to +2.5 mS/cm) in combination with normal and reduced current velocities (20 cm/s vs. 10 cm/s). Using a comprehensive assessment across multiple organism groups (macroinvertebrates, eukaryotic algae, fungi, parasites) and ecosystem functions (primary production, organic-matter decomposition), we show that flow velocity reduction has a pervasive impact, causing community shifts for almost all assessed organism groups (except fungi) and inhibiting organic-matter decomposition. Salinization affected only dynamic components of community assembly by enhancing invertebrate emigration via drift and reducing fungal reproduction. We caution that the comparatively small impact of salt in our study can be due to legacy effects from past salt pollution by coal mining activities >30 years ago. Nevertheless, our results suggest that urban stream management should prioritize the continuity of a minimum discharge to maintain ecosystem integrity. Our study exemplifies a holistic approach for the assessment of multiple-stressor impacts on streams, which is needed to inform the establishment of a salinity threshold above which mitigation actions must be taken.

Temporal monitoring of free-living nematode communities for evaluation of soil health in an arable crop rotation

There is a growing interest in finding reliable methods for monitoring soil health using bioindicators. Free-living nematodes are an ideal indicator group because of their rapid response to changes in soil conditions. This UK study aims to assess their efficacy as bioindicators using two field experiments. In Experiment-1, the treatments included Farmyard Manure, Green Manure consisting of a mix of Raphanus sativus and Vicia sp., and Standard Practice serving as the control receiving N-fertiliser only. The same treatments, except Farmyard Manure, were compared in Experiment-2, which was on a sloping site with a different textured soil. Soil samples were collected twice during each crop season, in Spring and Autumn, for Experiment-1, and only in Autumn for Experiment-2. Ecological indices that categorise nematodes by feeding preference using morphological differences and life strategies (i.e. functional guilds) were calculated. Indices were compared with the abundance of nematode trophic groups to evaluate their use as soil indicators for understanding crop management practices and their legacy effects. Results showed that identification to trophic groups alone was not a sufficiently sensitive approach for assessing changes in the selected management practices. The variations among trophic groups and treatments within the same sampling period were significantly different for bacterivores, fungivores, predators, omnivores, and herbivores. These differences did not always cooccur within the same sampling period, with bacterivores and plant-parasites of economic importance showing greater responses. The food web analyses, calculated by applying the Enrichment Index and Structure Index, and Plant Parasite Index, provided a more sensitive indicator and allowed more effective diachronic monitoring. While using the composition of trophic groups appears to be an attractive solution, their application is best linked to quantifying short-term changes in soil condition and were not as well suited to longer-term soil health monitoring.

The Short-Term Responses of Forest Soil Invertebrate Communities to Typhoon Disturbances

Knowledge regarding the response of soil invertebrate communities to typhoon disturbance is limited, although it is known that soil invertebrates are sensitive to forest disturbances and that tropical cyclones (typhoons/hurricanes) are the most destructive natural disasters affecting the structure and function of forest ecosystems. To fill this knowledge gap, soil invertebrates in both litter and topsoil layers were investigated in four representative subtropical coastal forests of eastern China one week before the first typhoon (Hinnamnor) (T1), one day after the first typhoon (Hinnamnor) (T2), one day after the second typhoon (Muifa) (T3), and one week after the second typhoon (Muifa) (T4) in September 2022. Typhoon disturbances decreased the density and taxa abundance of soil invertebrate communities in litter layer, but the first typhoon disturbance increased these values in the topsoil layer. One week after the second typhoon disturbance, soil invertebrate communities in the litter layer showed a gradual recovery trend. Meanwhile, the soil invertebrate communities in the litter layer were more sensitive to typhoon disturbances than those in the topsoil layer. Furthermore, the responses of the soil invertebrate communities to the typhoon disturbances varied greatly with the forest types. The invertebrate densities in the litter layer decreased by 62.1%, 63.53%, 47.01%, and 46.92% in Chinese fir, second broad-leaved, mixed, and bamboo forests, respectively. Particularly, these two non-catastrophic typhoons significantly altered the functional group composition of detrital food webs in the short term, and the proportion of phytophages in detrital food webs in the litter layer increased after the typhoon disturbances. In conclusion, the effects of typhoon disturbances on soil invertebrate communities vary greatly with forest type and soil layer, and soil invertebrate communities can gradually recover after typhoon disturbances. The legacy effects of typhoon disturbances on the functional group composition of detrital food webs may influence carbon and nutrient cycling in forest ecosystems.

Alternative vegetation trajectories through passive habitat rewilding: opposite effects for animal conservation

ContextPassive habitat rewilding after rural abandonment can affect wildlife differently depending on the type of habitats that it generates.ObjectiveEvaluate and compare the effects of two alternative vegetation trajectories that occur through passive habitat rewilding in Mediterranean ecotone areas (crop-scrub and crop-pine forest transitions) on the long-term population dynamics of animal species.MethodsWe used the spur-thighed tortoise (Testudo graeca), a characteristic long-lived species of cultural landscapes, as study species. We applied a spatially explicit and individual-based model (STEPLAND) to simulate the movement and demographic processes in a long-term period, by comparing an “impact scenario” (i.e., historical land-use changes) to a “control scenario” (no land-use changes).ResultsThe two landscape scenarios resulted in different population trends. In the crop-scrub scenarios (control and impact), population densities increased similarly over time. However, the crop-pine forest scenario negatively affected population density throughout the simulation period, and showed a time-lag response of three decades. The extinction risk was 55% with a time-lag response of approximately 110 years.ConclusionsOur study highlights the need to analyse the legacy effects on long-lived ectotherms, using them as a proxy to understand the future effects of dynamic landscapes created by “passive habitat rewilding”. Our results showed how traditional agriculture in Mediterranean ecotone areas may generate “ecotone effects” (i.e. increase in demographical parameters), but also population extinction on long-lived ectotherms. Therefore, we consider it relevant to maintain traditional agricultural areas in Mediterranean landscapes, especially in ecotone areas associated with pine forests (generating mosaics with open habitats).

A historical contingency hypothesis for population ecology

Assessments of historical contingency have advanced our understanding of adaptive radiation and community ecology, but little attention has been given to assessing the importance of historical contingency in population ecology. An obstacle has been the unmet need to conceptualize historical contingencies for populations in a manner that allows for their explanatory power to be assessed and quantified so that it can be directly compared with the explanatory power of statistical models representing other hypotheses or theory-based explanations. Here we conceptualize historical contingencies as a series of random events characterized by (1) significant legacy effects that are comparable in length to the waiting time between such events, and (2) the disparate nature of individual events in the series. From that conceptualization, we present a simple quantitative framework for assessing the explanatory power of historical contingencies in population ecology and apply it to an existing long-term dataset on the predator-prey system in Isle Royale National Park. The population-level phenomenon that we focused on was predation rate because it is a synthesis of three basic elements in population ecology (predator abundance, prey abundance and kill rate). We also compared the explanatory power of models of the historical contingency hypothesis to a wide-range of alternative, theory-based, statistical models used to assess underlying mechanisms or forecast future dynamics. Models of the historical contingency hypothesis explained over half of the interannual variation in predation rate and performed similarly, or better than, the vast majority of alternative, theory-based, models. Those findings highlight the potential value of reconsidering the way that population ecologists traditionally attempt to explain phenomena. We also discuss how this new conceptualization of the historical contingency hypothesis can also be valuable for synthesizing several other important ecological concepts of broad significance, especially reddened spectra, tipping points, alternative stable states, and ecological surprises. If the historical contingency hypothesis were found to be broadly applicable, then it would likely explain why ecologists are conspicuously poor at forecasting future dynamics.

China’s urban green growth from 2000 to 2020 is influenced by legacy effects

China’s urban green growth from 2000 to 2020 is influenced by legacy effects

Spatio-temporal microbial regulation of aggregate-associated priming effects under contrasting tillage practices

Tillage intensity significantly influences the heterogeneous distribution and dynamic changes of soil microorganisms, consequently shaping spatio-temporal patterns of SOC decomposition. However, little is known about the microbial mechanisms by which tillage intensity regulates the priming effect (PE) dynamics in heterogeneous spatial environments such as aggregates. Herein, a microcosm experiment was established by adding 13C-labeled straw residue to three distinct aggregate-size classes (i.e., mega-, macro-, and micro-aggregates) from two long-term contrasting tillage histories (no-till [NT] and conventional plow tillage [CT]) for 160 days to observe the spatio-temporal variations in PE. Metagenomic sequencing and Fourier transform mid-infrared techniques were used to assess the relative importance of C-degrading functional genes, microbial community succession, and SOC chemical composition in the aggregate-associated PE dynamics during straw decomposition. Spatially, straw addition induced a positive PE for all aggregates, with stronger PE occurring in larger aggregates, especially in CT soil compared to NT soil. Larger aggregates have more unique microbial communities enriched in genes for simple C degradation (e.g., E5.1.3.6, E2.4.1.7, pmm-pgm, and KduD in Nitrosospeera and Burkholderia), contributing to the higher short-term PE; however, CT soils harbored more genes for complex C degradation (e.g., TSTA3, fcl, pmm-pgm, and K06871 in Gammaproteobacteria and Phycicoccus), supporting a stronger long-term PE. Temporally, soil aggregates played a significant role in the early-stage PEs (i.e., < 59 days after residue addition) through co-metabolism and nitrogen (N) mining, as evidenced by the increased microbial biomass C and dissolved organic C (DOC) and reduced inorganic N with increasing aggregate-size class. At a later stage, however, the legacy effect of tillage histories controlled the PEs via microbial stoichiometry decomposition, as suggested by the higher DOC-to-inorganic N and DOC-to-available P stoichiometries in CT than NT. Our study underscores the importance of incorporating both spatial and temporal microbial dynamics for a comprehensive understanding of the mechanisms underlying SOC priming, especially in the context of long-term contrasting tillage practices.

Recovery of lakes from eutrophication: changes in nitrogen retention capacity and the role of nitrogen legacy in 10 Danish lakes studied over 30 years

AbstractUsing data on 23 Danish lakes, we conducted mass balances to develop total nitrogen (TN) models for predicting annual mean TN in lakes based on external TN loading and found high predictability when including lake hydraulic retention time and mean depth in the model. We further used a unique 30-year mass balance data series from 10 Danish lakes with contrasting mean depths and hydraulic retention times to elucidate the effect of external TN loading reduction and N legacy on lake TN. We found that the TN retention percentage during the 30 years was generally not sensitive to an often major reduction in the external TN loading; it overall followed the pattern of the above model predictions, suggesting a low TN legacy effect. Moreover, the TN retention percentage was not affected by changes in TP. Our results, therefore, show a fast response to TN loading reduction, indicating that we can expect an immediate effect on lake water quality in shallow lakes suffering from internal phosphorus loading during re-oligotrophication provided that inorganic N is low enough to become a growth-limiting nutrient.

The synergistic effects of a leaf mixture on decomposition change with a period of terrestrial exposure prior to immersion in a stream.

The effect of mixing litter on decomposition has received considerable attention in terrestrial and aquatic (but rarely in both) ecosystems, with a striking lack of consensus in the obtained results. We studied the decomposition of a mixture of poplar and alder in three terrestrial: aquatic exposures to determine (1) if the effect of mixing litter on mass loss, associated decomposers (fungal biomass, sporulation rates, and richness), and detritivores (abundance, biomass, and richness of invertebrate shredders) differs between the stream (fully aquatic exposure) and when litter is exposed to a period of terrestrial exposure prior to immersion and (2) the effect of the mixture across exposure scenarios. The effect of the mixture was additive on mass loss and synergistic on decomposers and detritivores across exposure scenarios. Within scenarios, mass loss and decomposers showed synergistic effects only in the fully aquatic exposure, detritivores showed synergistic effects only when the period of terrestrial was shorter than the period of aquatic exposure, and when the period of terrestrial was equal to the period of aquatic exposure the effect of the mixture was additive on mass loss, decomposers, and detritivores. The species-specific effects also differed among exposure scenarios. Alder affected poplar only when there was a period of terrestrial exposure, with increased sporulation rates and fungal richness in exposure 25:75, and increased mass loss in exposure 50:50. Poplar affected alder only under fully aquatic exposure, with increased mass loss. In conclusion, the synergistic effects of the mixture changed with a period of terrestrial exposure prior to immersion. These results provide a cross-boundary perspective on the effect of mixing litter, showing a legacy effect of exposure to terrestrial decomposition on the fate of plant litter in aquatic ecosystems and highlighting the importance of also assessing the effect of mixing litter on the associated biota and not only on mass loss.