Legacies of more frequent drought in ponderosa pine across the western United States

Despite widespread interest in drought legacies—multiyear impacts of drought on tree growth—the key implication of reported drought legacies remains unaddressed: as impaired growth and slow recovery associated with drought legacies are pervasive across forest ecosystems, what is the impact of more frequent drought conditions? We investigated the assumption that either multiple drought years occurring during a short period (multiyear droughts), or droughts occurring during the recovery period from previous drought (compounded droughts), are detrimental to subsequent growth. There is evidence that drought responses may vary among populations of widespread species, leading us to examine regional differences in responses of the conifer Pinus ponderosa to historic drought frequency in the western United States. More frequent drought conditions incurred additional growth declines and shifts in growth–climate sensitivities in the years following drought relative to single‐drought events, with ‘triple‐droughts' being worse than ‘double‐droughts'. Notably, prediction skill was not strongly reduced when ignoring compounded droughts, a consequence of the temporally comprehensive formulation of our stochastic antecedent model that accounts for the climatic memory of tree growth. We argue that incorporating drought‐induced temporal variability in tree growth sensitivities can aid inference gained from statistical models, where more simplistic models could overestimate the severity of drought legacies. We also found regional differences in response to repeated drought, and suggest plastic post‐drought sensitivities and climatic memory may represent beneficial physiological adjustments in interior regions. Within‐species variability may thus mediate forest responses to increasing drought frequency under future climate change, but experimental approaches using more species are necessary to improve our understanding of the mechanisms that underlie drought legacy effects on tree growth.     

Soil legacy effects of plants and drought on aboveground insects in native and range-expanding plant communities

Soils contain biotic and abiotic legacies of previous conditions that may influence plant community biomass and associated aboveground biodiversity. However, little is known about the relative strengths and interactions of the various belowground legacies on aboveground plant–insect interactions. We used an outdoor mesocosm experiment to investigate the belowground legacy effects of range-expanding versus native plants, extreme drought and their interactions on plants, aphids and pollinators. We show that plant biomass was influenced more strongly by the previous plant community than by the previous summer drought. Plant communities consisted of four congeneric pairs of natives and range expanders, and their responses were not unanimous. Legacy effects affected the abundance of aphids more strongly than pollinators. We conclude that legacies can be contained as soil ‘memories’ that influence aboveground plant community interactions in the next growing season. These soil-borne ‘memories’ can be altered by climate warming-induced plant range shifts and extreme drought.     

Drought legacies are dependent on water table depth,wood anatomy and drought timing across the eastern US

Severe droughts can impart long‐lasting legacies on forest ecosystems through lagged effects that hinder tree recovery and suppress whole‐forest carbon uptake. However, the local climatic and edaphic factors that interact to affect drought legacies in temperate forests remain unknown. Here, we pair a dataset of 143 tree ring chronologies across the mesic forests of the eastern US with historical climate and local soil properties. We found legacy effects to be widespread, the magnitude of which increased markedly in diffuse porous species, sites with deep water tables, and in response to late‐season droughts (August–September). Using an ensemble of downscaled climate projections, we additionally show that our sites are projected to drastically increase in water deficit and drought frequency by the end of the century, potentially increasing the size of legacy effects by up to 65% and acting as a significant process shaping forest composition, carbon uptake and mortality.     

Context-Dependency of Agricultural Legacies in Temperate Forest Soils

Ecosystems - Anthropogenic activities have affected forests for centuries, leading to persistent legacies. Observations of agricultural legacies on forest soil properties have been site specific...     

Historical forest disturbance mediates soil microbial community responses to drought

Despite the abundance of studies demonstrating the effects of drought on soil microbial communities, the role of land use legacies in mediating these drought effects is unclear. To assess historical land use influences on microbial drought responses, we conducted a drought-rewetting experiment in soils from two adjacent and currently forested watersheds with distinct land use histories: an undisturbed ‘reference’ site and a ‘disturbed’ site that was clear-cut and converted to agriculture ~60 years prior. We incubated intact soil cores at either constant moisture or under a drought-rewet treatment and characterized bacterial and fungal communities using amplicon sequencing throughout the experiment. Bacterial alpha diversity decreased following drought-rewetting while fungal diversity increased. Bacterial beta diversity also changed markedly following drought-rewetting, especially in historically disturbed soils, while fungal beta diversity exhibited little response. Additionally, bacterial beta diversity in disturbed soils recovered less from drought-rewetting compared with reference soils. Disturbed soil communities also exhibited notable reductions in nitrifying taxa, increases in putative r-selected bacteria, and reductions in network connectivity following drought-rewetting. Overall, our study reveals historical land use to be important in mediating responses of soil bacterial communities to drought, which will influence the ecosystem-scale trajectories of these environments under ongoing and future climate change.     

Effects of climate legacies on above‐ and belowground community assembly

The role of climatic legacies in regulating community assembly of above‐ and belowground species in terrestrial ecosystems remains largely unexplored and poorly understood. Here, we report on two separate regional and continental empirical studies, including >500 locations, aiming to identify the relative importance of climatic legacies (climatic anomaly over the last 20,000 years) compared to current climates in predicting the relative abundance of ecological clusters formed by species strongly co‐occurring within two independent above‐ and belowground networks. Climatic legacies explained a significant portion of the variation in the current community assembly of terrestrial ecosystems (up to 15.4%) that could not be accounted for by current climate, soil properties, and management. Changes in the relative abundance of ecological clusters linked to climatic legacies (e.g., past temperature) showed the potential to indirectly alter other clusters, suggesting cascading effects. Our work illustrates the role of climatic legacies in regulating ecosystem community assembly and provides further insights into possible winner and loser community assemblies under global change scenarios.     

Legacies in life histories

Complex life-histories are common in nature, have many importantbiological consequences, and are an important focal area forintegrative biology. For organisms with complex life-histories,a legacy is something handed down from an ancestor or previousstage, and can be genetic, nutritional/provisional, experiential,as well as the result of random chance and natural variationin the environment. As we learn more about complex life-histories,it becomes clear that legacies are inexorably linked in theshort- and long-term through ecology and evolution. Understandingthe consequences and drivers of life-history patterns can thereforeonly be understood by considering all types of legacies andintegrating legacies across the entire life cycle. Larry McEdwardwas a leader in the field of ecological physiology, and evolutionaryecology of marine invertebrate larvae with complex life-histories.Through his scientific work and publications, devotion to students,colleagues, family, and friends, Larry has left a lasting legacythat will impact the future development of the field of larvalecology and complex life-histories.     

Within‐Year Soil Legacies Contribute to Strong Priority Effects of Exotics on Native California Grassland Communities

Understanding priority effects, in which one species in a habitat decreases the success of later species, may be essential for restoring native communities. Priority effects can operate in two ways: size‐asymmetric competition and creation of “soil legacies,” effects on soil that may last long after the competitive effect. We examined how these two types of priority effects, competition and soil legacies, drive interactions between seedlings of native and exotic California grassland plants. We established native and exotic communities in a mesocosm experiment. After 5 weeks, we removed the plants from half the treatments (soil legacy treatment) and retained the plants in the other half (priority effect treatment, which we interpret to include both competition and soil legacies). We then added native or exotic seed as the colonizing community. After 2 months, we measured the biomass of the colonizing community. When germinating first, both natives and exotics established priority effects, reducing colonist biomass by 86 and 92%, respectively. These priority effects were predominantly due to size‐asymmetric competition. Only exotics created soil legacies, and these legacies only affected native colonizers, reducing biomass by 74%. These results imply that exotic species priority effects can affect native grassland restorations. Although most restorations focus on removing exotic seedlings, amending soil to address soil legacies may also be critical. Additionally, because native species can exclude exotics if given a head start, ensuring that natives germinate first may be a cost‐effective restoration technique.     

Higher susceptibility of beech to drought in comparison to oak

The expected increase in drought severity and frequency as a result of anthropogenic climate change leads to concerns about the ability of native tree species to cope with these changes. To determine the susceptibility of Fagus sylvatica (European beech) and Quercus robur (pedunculate oak) – the two dominant deciduous tree species in Central Europe – to drought, we quantified the climate sensitivity and drought-response of radial growth for both species using an array of dendroecological methods. Tree-ring data were collected from a site east of Coburg, Bavaria which had shown pronounced stress-symptoms (early leaf coloration) during the record drought of 2018. Climate-growth relationships were used to establish the sensitivity of radial growth to multiple climatic variables. The impact of specific drought events on tree growth was quantified using tolerance indices. In addition, we employed a Principal Component Gradient Analysis (PCGA) and remote sensing data (MODIS Normalized Difference Vegetation Index (NDVI)) to delineate the species specific drought responses. Using these methods we were able to show a clear difference in drought susceptibility between beech and oak. Beech displayed a higher sensitivity to temperature and the standardized precipitation evapotranspiration index (SPEI) and showed lower resistance and resilience to drought events than oak. In particular, beech was unable to fully recover from the 2003 drought, after which it expressed a stark growth decline, i.e. drought legacies, which was not observed for oak. The PCGA revealed a clear differentiation in the grouping of drought responses between beech and oak, supporting the findings of the climate-growth analysis and the tolerance indices. Correlations of NDVI and ring-width indices (RWI) indicated that under normal climatic conditions NDVI variability is linked to the start of the growing season. This is in contrast to drought years, such as 2003, where summer NDVI mirrored the drought response of beech and oak. These results reveal beech to have both a higher sensitivity to summer temperature and SPEI and a higher susceptibility to drought events. Although, in the past high plasticity and adaptability to drought have been attributed to both beech and oak, our study assigns beech a higher risk than oak to suffer from anticipated increases in drought frequency and intensity as a consequence of climate change.     

Quaternary and pre‐Quaternary historical legacies in the global distribution of a major tropical plant lineage

Aim The relative importance of current climate and past historical legacies is hotly debated. Here, we assess their role in determining the global distribution and diversity patterns of palms (Arecaceae), a widespread, species‐rich group of keystone ecological importance in tropical ecosystems. Location Global. Methods We assembled country‐level species lists world‐wide and compiled associated data on potential contemporary environmental drivers (current climate, habitat heterogeneity, area, and insularity), Quaternary glacial–interglacial climate change and major biogeographic regions to evaluate to what extent the global distribution and species richness patterns in palms reflect Quaternary climatic oscillations or regional effects reflecting pre‐Quaternary legacies. We also assessed for the first time if historical legacies differ between continents and islands, providing novel insights into determinants of insular species richness. Results Palm species richness was significantly affected by Quaternary climate changes and further differed between biogeographic regions even when both current environmental conditions and Quaternary climate changes were accounted for. In contrast, global limits to the distribution of the palm family were best explained by current temperature while biogeographic regional differences were unimportant and Quaternary climate change caused only a small constraint. Historical legacies were weak on islands, with only a small regional effect and no effect of Quaternary climate changes. Main conclusions Strong historical legacies supplement current environment as determinants of palm species richness. These primarily comprise pre‐Quaternary historical effects, reflected in low African species richness (possibly linked to pre‐Quaternary extinctions) and outstandingly high Neotropical and Indomalayan palm species richness (possibly linked to these regions' long‐term climatic suitability for palms). In contrast to species richness, the global distribution of the family range is largely in equilibrium with current climate. The small historical effects on islands are consistent with climatic buffering from their oceanic environment.     

Legacy effects of a regional drought on aboveground net primary production in six central US grasslands

Global climate models predict increases in the frequency and severity of drought worldwide, directly affecting most ecosystem types. Consequently, drought legacy effects (drought-induced alterations in ecosystem function postdrought) are expected to become more common in ecosystems varying from deserts to grasslands to forests. Drought legacies in grasslands are usually negative and reduce ecosystem function, particularly after extended drought. Moreover, ecosystems that respond strongly to drought (high sensitivity) might be expected to exhibit the largest legacy effects the next year, but this relationship has not been established. We quantified legacy effects of a severe regional drought in 2012 on postdrought (2013) aboveground net primary productivity (ANPP) in six central US grasslands. We predicted that (1) the magnitude of drought legacy effects measured in 2013 would be positively related to the sensitivity of ANPP to the 2012 drought, and (2) drought legacy effects would be negative (reducing 2013 ANPP relative to that expected given normal precipitation amounts). The magnitude of legacy effects measured in 2013 was strongly related (r² = 0.88) to the sensitivity of ANPP to the 2012 drought across these six grasslands. However, contrary to expectations, positive legacy effects (greater than expected ANPP) were more commonly observed than negative legacy effects. Thus, while the sensitivity of ANPP to drought may be a useful predictor of the magnitude of legacy effects, short-term (1-year) severe droughts may cause legacy effects that are more variable than those observed after multiyear droughts.     

Persistent land-use legacies increase small-scale diversity and strengthen vegetation–soil relationships on an unmanaged heathland

Land-use legacies are recognized determinants of vegetation dynamics and plant community assembly. The duration of these legacies and how they influence the structure of vegetation communities developing naturally in nutrient-poor ecosystems is not well understood. Here, we focus on the effects of increased nutrient availability from previous agricultural practices on multiple vegetation properties in a heathland where agriculture and domestic grazing ceased near 1870 and 1895, respectively. We compared diversity, compositional and functional properties of the vegetation responses to land-use legacies in the soil between areas with different agricultural histories (previously cultivated vs. uncultivated). Diversity measures were found to be higher in the previously cultivated soils. β-diversity was mainly driven by changes in species relative cover and increased with increasing nutrient availability in the cultivated area. Furthermore, functional traits related to nutrient acquisition (SLA and Leaf Nitrogen content) and the changes in vegetation composition were directly linked to soil properties only in the previously cultivated part of the heathland. Together these results revealed a shift to a deterministic control of the plant community, where increased nutrient availability leads to stronger associations between soil and vegetation properties. This suggests that as nutrients become available, niche differentiation and competitive interactions become the predominant underlying mechanisms structuring the community. Our study shows that land-use legacies of moderate intensity can alter the assembly mechanisms and diversity patterns in unmanaged vegetation that can be maintained after more than a century since cessation of agricultural practice. Identifying land-use legacies and understanding how they structure heathland communities can thus lead to management decisions adapted to the specific assembly mechanisms and result in a more effective management.     

Plant community composition steers grassland vegetation via soil legacy effects

Soil legacy effects are commonly highlighted as drivers of plant community dynamics and species co‐existence. However, experimental evidence for soil legacy effects of conditioning plant communities on responding plant communities under natural conditions is lacking. We conditioned 192 grassland plots using six different plant communities with different ratios of grasses and forbs and for different durations. Soil microbial legacies were evident for soil fungi, but not for soil bacteria, while soil abiotic parameters did not significantly change in response to conditioning. The soil legacies affected the composition of the succeeding vegetation. Plant communities with different ratios of grasses and forbs left soil legacies that negatively affected succeeding plants of the same functional type. We conclude that fungal‐mediated soil legacy effects play a significant role in vegetation assembly of natural plant communities.     

Global environmental change effects on plant community composition trajectories depend upon management legacies

The contemporary state of functional traits and species richness in plant communities depends on legacy effects of past disturbances. Whether temporal responses of community properties to current environmental changes are altered by such legacies is, however, unknown. We expect global environmental changes to interact with land‐use legacies given different community trajectories initiated by prior management, and subsequent responses to altered resources and conditions. We tested this expectation for species richness and functional traits using 1814 survey‐resurvey plot pairs of understorey communities from 40 European temperate forest datasets, syntheses of management transitions since the year 1800, and a trait database. We also examined how plant community indicators of resources and conditions changed in response to management legacies and environmental change. Community trajectories were clearly influenced by interactions between management legacies from over 200 years ago and environmental change. Importantly, higher rates of nitrogen deposition led to increased species richness and plant height in forests managed less intensively in 1800 (i.e., high forests), and to decreases in forests with a more intensive historical management in 1800 (i.e., coppiced forests). There was evidence that these declines in community variables in formerly coppiced forests were ameliorated by increased rates of temperature change between surveys. Responses were generally apparent regardless of sites’ contemporary management classifications, although sometimes the management transition itself, rather than historic or contemporary management types, better explained understorey responses. Main effects of environmental change were rare, although higher rates of precipitation change increased plant height, accompanied by increases in fertility indicator values. Analysis of indicator values suggested the importance of directly characterising resources and conditions to better understand legacy and environmental change effects. Accounting for legacies of past disturbance can reconcile contradictory literature results and appears crucial to anticipating future responses to global environmental change.     

Cultivation legacies alter soil nutrients and differentially affect plant species performance nearly a century after abandonment

Cultivation legacies affect native vegetation in old fields of the Great Basin, USA for nearly a century after these fields are abandoned. We hypothesized that cultivation lowered soil nutrients and that this legacy would differentially impact plant performance of four representative Great Basin species. To test these hypotheses, we compared soil nutrients (C, N, P, K, Mg and Ca) between two formerly cultivated and adjacent noncultivated sites in two soil series. We then compared the plant growth and foliar nutrient content of an exotic grass (Bromus tectorum L.), two native grasses (Elymus elymoides [Raf.] Swezey and Achnatherum hymenoides [Roem. and Schult.] Barkworth), and a native forb (Sphaeralcea grossulariifolia [Hook. and Arn.] Rydb) grown in these soils in the greenhouse and in the field. Only one sampling site had reduced soil nutrients associated with cultivation legacies, where most of the negative effects on plant performance were found. E. elymoides appeared to be less affected by cultivation legacies than did A. hymenoides, which had a reduced survivorship and 20 % less above-ground biomass in cultivated soils. No species, including B. tectorum, were favored by cultivation. Our findings suggest that cultivation legacies can affect plant performance of different species in different ways and that altered soil nutrients may interact with other abiotic and biotic cultivation legacies in complex ways.     

The influence of time,soil characteristics,and land‐use history on soil phosphorus legacies: a global meta‐analysis

Agriculturally driven changes in soil phosphorus (P) are known to have persistent effects on local ecosystem structure and function, but regional patterns of soil P recovery following cessation of agriculture are less well understood. We synthesized data from 94 published studies to assess evidence of these land‐use legacies throughout the world by comparing soil labile and total P content in abandoned agricultural areas to that of reference ecosystems or sites remaining in agriculture. Our meta‐analysis shows that soil P content was typically elevated after abandonment compared to reference levels, but reduced compared to soils that remained under agriculture. There were more pronounced differences in the legacies of past agriculture on soil P across regions than between the types of land use practiced prior to abandonment (cropland, pasture, or forage grassland). However, consistent patterns of soil P enrichment or depletion according to soil order and types of post‐agricultural vegetation suggest that these factors may mediate agricultural legacies on soil P. We also used mixed effects models to examine the role of multiple variables on soil P recovery following agriculture. Time since cessation of agriculture was highly influential on soil P legacies, with clear reductions in the degree of labile and total P enrichment relative to reference ecosystems over time. Soil characteristics (clay content and pH) were strongly related to changes in labile P compared to reference sites, but these were relatively unimportant for total P. The duration of past agricultural use and climate were weakly related to changes in total P only. Our finding of reductions in the degree of soil P alteration over time relative to reference conditions reveals the potential to mitigate these land‐use legacies in some soils. Better ability to predict dynamics of soil nutrient recovery after termination of agricultural use is essential to ecosystem management following land‐use change.     

The influence of climatic legacies on the distribution of dryland biocrust communities

Predicting the distribution of biocrust species, mosses, lichens and liverworts associated with surface soils is difficult, but climatic legacies (changes in climate over the last 20 k years) can improve our prediction of the distribution of biocrust species. To provide empirical support for this hypothesis, we used a combination of network analyses and structural equation modelling to identify the role of climatic legacies in predicting the distribution of ecological clusters formed by species of mosses, lichens and liverworts using data from 282 large sites distributed across 0.6 million km² of eastern Australia. Two ecological clusters contained 87% of the 120 moss, lichen and liverwort species. Both clusters contained lichen, moss and liverwort species, but were dominated by different families. Sites where the air temperature increased the most over 20k years (positive temperature legacies) were associated with reductions in the relative abundance of species from the lichen (Peltulaceae and Teloschistaceae) and moss (Bryaceae) families (Cluster A species), greater groundstorey plant cover and lower soil pH. Sites where precipitation has increased over the past 20k years (positive precipitation legacy) were associated with increases in the relative abundance of lichen (Cladoniaceae, Lecideaceae and Thelotremataceae) and moss (Pottiaceae) families (Cluster B species) and lower levels of soil pH. Sites where temperatures have increased the most in the past 20k years suppressed the negative effects of plant cover on Cluster B by reducing plant cover. Increased intensity of grazing suppressed the negative effect of soil pH and the positive effect of soil carbon, on the relative abundance of Cluster B taxa. Finally, increasing temperature and precipitation legacies reduced the negative effect of soil pH on Cluster B. Understanding of the importance of climatic legacies improves our ability to predict how biocrust assemblies might respond to ongoing global environmental change associated with increasing land use intensification, increasing temperature and reduced rainfall.     

The Effects of Spatial Legacies following Shifting Management Practices and Fire on Boreal Forest Age Structure

Forest age structure and its spatial arrangement are important elements of sustainable forestry because of their effects on biodiversity and timber availability. Forest management objectives that include specific forest age structure may not be easily attained due to constraints imposed by the legacies of historical management and natural disturbance. We used a spatially explicit stochastic model to explore the synergetic effects of forest management and fire on boreal forest age structure. Specifically, we examined (1) the duration of spatial legacies of different management practices in the boreal forest, (2) how multiple shifts in management practices affect legacy duration and the spatial trajectories of forest age structure, and (3) how fire influences legacy duration and pattern development in combination with harvesting. Results based on 30 replicates of 500 years for each scenario indicate that (1) spatial legacies persist over 200 years and the rate at which legacies are overcome depends on whether new management targets are in synchrony with existing spatial pattern; (2) age specific goals were met faster after multiple management shifts due to the similar spatial scale of the preceding management types; (3) because large fires can erase the spatial pattern created by smaller disturbances, scenarios with fire had shorter lags than scenarios without fire. These results suggest that forest management goals can be accelerated by applying management at a similar spatial scale as existing spatial patterns. Also, management planning should include careful consideration of historical management as well as current and likely future disturbances.     

Legacies of afforestation on soil nematode community composition,structure, and diversity in a northern Canadian prairie

Plant and Soil - We examined how legacies of afforestation affect soil food webs using the composition, structure, and diversity of soil nematode communities along a prairie restoration...     

Long-term fire and vegetation change in northwestern Amazonia

Amazonian forest plots are used to quantify biodiversity and carbon sequestration, and provide the foundation for much of what is known about tropical ecology. Many plots are assumed to be undisturbed, but recent work suggests that past fire, forest openings, and cultivation created vegetation changes that have persisted for decades to centuries (ecological legacies). The Yasuní Forest Dynamics plot is one of the most biodiverse places on earth, yet its human history remains unknown. Here, we use charcoal and phytolith analysis to investigate the fire and vegetation history of the Yasuní forest plot, and compare results with nearby forest plots in Colombia (Amacayacu) and Peru (Medio Putumayo-Algodón [MPA]) to explore the spatial variability of past disturbances and ecological legacies in northwestern Amazonia. Three ¹⁴C dated charcoal fragments provided evidence for a modern (1956 CE) and a past fire event ca. 750 years ago at Yasuní, compared with fire ages of 1000–1600 years ago documented at Amacayacu and MPA. Small-scale disturbances and localized canopy openings also occurred in the Yasuní plot. Phytolith assemblages from Yasuní and Amacayacu showed more variability in past vegetation change than MPA. Low-intensity, non-continuous disturbances occurred at all three plots in the past, and our results highlight the variability of past human activities both in space and time in northwestern Amazonia. Our data also suggest that post-Columbian human disturbances from the Rubber Boom (AD 1850–1920) and subsequent oil exploration have likely left stronger ecological legacies than those left by pre-Columbian peoples in our studied regions.