EcoMem: An R package for quantifying ecological memory

Ecological processes may exhibit memory to past disturbances affecting the resilience of ecosystems to future disturbance. Understanding the role of ecological memory in shaping ecosystem responses to disturbance under global change is a critical step toward developing effective adaptive management strategies to maintain ecosystem function and biodiversity. We developed EcoMem, an R package for quantifying ecological memory functions using common environmental time series data (continuous, count, proportional) applying a Bayesian hierarchical framework. The package estimates memory functions for continuous and binary (e.g., disturbance chronology) variables making no a priori assumption on the form of the functions. EcoMem allows users to quantify ecological memory for a wide range of ecosystem processes and responses. The utility of the package to advance understanding of the memory of ecosystems to environmental drivers is demonstrated using a simulated dataset and a case study assessing the memory of boreal tree growth to insect defoliation.Peer reviewe

Identifying and Separating the Processes Underlying Boreal Forest Understory Community Assembly

Identifying the ecological processes underlying community assembly remains an elusive goal in community ecology. We formalize assembly hypotheses as alternative models and apply each to predict 1,918 out-of-sample boreal forest understory communities to identify and separate the processes driving community assembly. Models are specified within a Bayesian joint species distribution framework that allows for the inclusion and separation of stochastic processes, environmental filtering, and two different species dependence structures. We found clear evidence that study communities are structured by both environmental filtering and compositional dependence highlighting the importance of selection in community assembly. The relative importance of environmental filtering was greater than compositional dependence in predicting both understory communities and the abundance of constituent species across broad suc-cessional and bioclimatic gradients. Contrary to ecological expectations, the inclusion of a flexible residual species dependence structure (accounting for more than compositional dependence) did not improve model predictions after accounting for the strong role of environmental filtering. Our results provide novel inference on the processes underlying community assembly facilitated by applying empirical approximations of alternative assembly processes to predict communities across a range of environmental conditions

Shifts in timing and duration of breeding for 73 boreal bird species over four decades

Breeding timed to match optimal resource abundance is vital for the successful reproduction of species, and breeding is therefore sensitive to environmental cues. As the timing of breeding shifts with a changing climate, this may not only affect the onset of breeding but also its termination, and thus the length of the breeding period. We use an extensive dataset of over 820K nesting records of 73 bird species across the boreal region in Finland to probe for changes in the beginning, end, and duration of the breeding period over four decades (1975 to 2017). We uncover a general advance of breeding with a strong phylogenetic signal but no systematic variation over space. Additionally, 31% of species contracted their breeding period in at least one bioclimatic zone, as the end of the breeding period advanced more than the beginning. We did not detect a statistical difference in phenological responses of species with combinations of different migratory strategy or number of broods. Nonetheless, we find systematic differences in species responses, as the contraction in the breeding period was found almost exclusively in resident and short-distance migrating species, which generally breed early in the season. Overall, changes in the timing and duration of reproduction may potentially lead to more broods co-occurring in the early breeding season-a critical time for species' reproductive success. Our findings highlight the importance of quantifying phenological change across species and over the entire season to reveal shifts in the community-level distribution of bird reproduction.Peer reviewe

Shifts in timing and duration of breeding for 73 boreal bird species over four decades

Breeding timed to match optimal resource abundance is vital for the successful reproduction of species, and breeding is therefore sensitive to environmental cues. As the timing of breeding shifts with a changing climate, this may not only affect the onset of breeding but also its termination, and thus the length of the breeding period. We use an extensive dataset of over 820K nesting records of 73 bird species across the boreal region in Finland to probe for changes in the beginning, end, and duration of the breeding period over four decades (1975 to 2017). We uncover a general advance of breeding with a strong phylogenetic signal but no systematic variation over space. Additionally, 31% of species contracted their breeding period in at least one bioclimatic zone, as the end of the breeding period advanced more than the beginning. We did not detect a statistical difference in phenological responses of species with combinations of different migratory strategy or number of broods. Nonetheless, we find systematic differences in species responses, as the contraction in the breeding period was found almost exclusively in resident and short-distance migrating species, which generally breed early in the season. Overall, changes in the timing and duration of reproduction may potentially lead to more broods co-occurring in the early breeding season—a critical time for species’ reproductive success. Our findings highlight the importance of quantifying phenological change across species and over the entire season to reveal shifts in the community-level distribution of bird reproduction

Peak radial growth of diffuse-porous species occurs during periods of lower water availability than for ring-porous and coniferous trees

Climate models project warmer summer temperatures will increase the frequency and heat severity of droughts in temperate forests of Eastern North America. Hotter droughts are increasingly documented to affect tree growth and forest dynamics, with critical impacts on tree mortality, carbon sequestration and timber provision. The growing acknowledgement of the dominant role of drought timing on tree vulnerability to water deficit raises the issue of our limited understanding of radial growth phenology for most temperate tree species. Here, we use well-replicated dendrometer band data sampled frequently during the growing season to assess the growth phenology of 610 trees from 15 temperate species over 6 years. Patterns of diameter growth follow a typical logistic shape, with growth rates reaching a maximum in June, and then decreasing until process termination. On average, we find that diffuse-porous species take 16-18 days less than other wood-structure types to put on 50% of their annual diameter growth. However, their peak growth rate occurs almost a full month later than ring-porous and conifer species (ca. 24 +/- 4 days; mean +/- 95% credible interval). Unlike other species, the growth phenology of diffuse-porous species in our dataset is highly correlated with their spring foliar phenology. We also find that the later window of growth in diffuse-porous species, coinciding with peak evapotranspiration and lower water availability, exposes them to a higher water deficit of 88 +/- 19 mm (mean +/- SE) during their peak growth than ring-porous and coniferous species (15 +/- 35 mm and 30 +/- 30 mm, respectively). Given the high climatic sensitivity of wood formation, our findings highlight the importance of wood porosity as one predictor of species climatic sensitivity to the projected intensification of the drought regime in the coming decades.Peer reviewe

Shifts in timing and duration of breeding for 73 boreal bird species over four decades

Breeding timed to match optimal resource abundance is vital for the successful reproduction of species, and breeding is therefore sensitive to environmental cues. As the timing of breeding shifts with a changing climate, this may not only affect the onset of breeding but also its termination, and thus the length of the breeding period. We use an extensive dataset of over 820K nesting records of 73 bird species across the boreal region in Finland to probe for changes in the beginning, end, and duration of the breeding period over four decades (1975 to 2017). We uncover a general advance of breeding with a strong phylogenetic signal but no systematic variation over space. Additionally, 31% of species contracted their breeding period in at least one bioclimatic zone, as the end of the breeding period advanced more than the beginning. We did not detect a statistical difference in phenological responses of species with combinations of different migratory strategy or number of broods. Nonetheless, we find systematic differences in species responses, as the contraction in the breeding period was found almost exclusively in resident and short-distance migrating species, which generally breed early in the season. Overall, changes in the timing and duration of reproduction may potentially lead to more broods co-occurring in the early breeding season-a critical time for species' reproductive success. Our findings highlight the importance of quantifying phenological change across species and over the entire season to reveal shifts in the community-level distribution of bird reproduction.Peer reviewe

Data from: Boreal tree growth exhibits decadal-scale ecological memory to drought and insect defoliation, but no negative response to their interaction

1. Interactions between drought and insect defoliation may dramatically alter forest function under novel climate and disturbance regimes, but remain poorly understood. We empirically tested two important hypotheses regarding tree responses to drought and insect defoliation: 1) trees exhibit delayed, persistent, and cumulative growth responses to these stressors; 2) physiological feedbacks in tree responses to these stressors exacerbate their impacts on tree growth. These hypotheses remain largely untested at a landscape scale, yet are critical for predicting forest function under novel future conditions given the connection between tree growth and demographic processes such as mortality and regeneration. 2. We developed a Bayesian hierarchical model to quantify the ecological memory of tree growth to past water deficits and insect defoliation events, derive antecedent variables reflecting the persistent and cumulative effects of these stressors on current growth, and test for their interactive effects. The model was applied to extensive tree growth, weather, and defoliation survey data from western and eastern regions of the Canadian boreal forest impacted by recent drought and defoliation events and characterized by contrasting tree compositions, climates, and insect defoliators. 3. Results revealed persistent negative tree growth responses to past water (all trees) and defoliation (host trees) stress lasting 3-6 and 10-12 years, respectively, depending on study region. Accounting for the ecological memory of tree growth to water and defoliation stress allowed for detection of interactions not previously demonstrated. Contrary to expectations, we found evidence for positive interactions among non-host trees likely due to reduced water stress following defoliation events. Regional differences in ecological memory to water stress highlight the role of climate in shaping forest responses to drought. 4. Synthesis. Study results suggest negative feedbacks in tree responses to drought and insect attack may be weaker than predicted for defoliator-dominated boreal forest systems. Instead, insect defoliation may offset the impacts of water deficit on boreal tree growth by reducing transpirational water demand. This offset mimics increased resistance to drought following forest thinning and may lessen growth and mortality losses due to increased aridity and more severe insect damage forecast for the boreal forest under global change

Tree growth, defoliation, and climatic water deficit data

Individual tree growth along with stand-level defoliation and weather data are provided for each study region. The individual tree growth data includes ring-width observations as well as tree-level variables including age, diameter, and species. Detailed information on the growth, defoliation, and weather data can be found in the methods section of the accompanying article

Bayesian joint species distribution model selection for community-level prediction

Peer reviewe