Seed classification with random forest models.

Premise: To improve forest conservation monitoring, we developed a protocol to automatically count and identify the seeds of plant species with minimal resource requirements, making the process more efficient and less dependent on human operators. Methods and Results: Seeds from six North American conifer tree species were separated from leaf litter and imaged on a flatbed scanner. In the most successful species-classification approach, an ImageJ macro automatically extracted measurements for random forest classification in the software R. The method allows for good classification accuracy, and the same process can be used to train the model on other species. Conclusions: This protocol is an adaptable tool for efficient and consistent identification of seed species or potentially other objects. Automated seed classification is efficient and inexpensive, making it a practical solution that enhances the feasibility of large-scale monitoring projects in conservation biology.

Fire sparks upslope range shifts of North Cascades plant species.

As ongoing climate change drives suitable habitats to higher elevations, species ranges are predicted to follow. However, observed range shifts have been surprisingly variable, with most species differing in rates of upward shift and others failing to shift at all. Disturbances such as fires could play an important role in accelerating range shifts by facilitating recruitment in newly suitable habitats (leading edges) and removing adults from areas no longer suited for regeneration (trailing edges). To date, empirical evidence that fires interact with climate change to mediate elevational range shifts is scarce. Resurveying historical plots in areas that experienced climate change and fire disturbance between surveys provides an exciting opportunity to fill this gap. To investigate whether species have tended to shift upslope and if shifts depend on fires, we resurveyed historical vegetation plots in North Cascades National Park, Washington, USA, an area that has experienced warming, drying, and multiple fires since the original surveys in 1983. We quantified range shifts by synthesizing across two lines of evidence: (1) displacement at range edges and the median elevation of species occurrences, and (2) support for the inclusion of interactions among time, fire and elevation in models of species presence with elevation. Among species that experienced fire since the original survey, a plurality expanded into new habitats at their upper edge. In contrast, a plurality of species not experiencing fire showed no evidence of shifts, with the remainder exhibiting responses that were variable in magnitude and direction. Our results suggest that fires can facilitate recruitment at leading edges, while species in areas free of disturbance are more likely to experience stasis.

Applying climate change refugia to forest management and old-growth restoration.

Recent studies highlight the potential of climate change refugia (CCR) to support the persistence of biodiversity in regions that may otherwise become unsuitable with climate change. However, a key challenge in using CCR for climate resilient management lies in how CCR may intersect with existing forest management strategies, and subsequently influence how landscapes buffer species from negative impacts of warming climate. We address this challenge in temperate coastal forests of the Pacific Northwestern United States, where declines in the extent of late-successional forests have prompted efforts to restore old-growth forest structure. One common approach for doing so involves selectively thinning forest stands to enhance structural complexity. However, dense canopy is a key forest feature moderating understory microclimate and potentially buffering organisms from climate change impacts, raising the possibility that approaches for managing forests for old-growth structure may reduce the extent and number of CCR. We used remotely sensed vegetation indices to identify CCR in an experimental forest with control and thinned (restoration) treatments, and explored the influence of biophysical variables on buffering capacity. We found that remotely sensed vegetation indices commonly used to identify CCR were associated with understory temperature and plant community composition, and thus captured aspects of landscape buffering that might instill climate resilience and be of interest to management. We then examined the interaction between current restoration strategies and CCR, and found that selective thinning for promoting old-growth structure had only very minor, if any, effects on climatic buffering. In all, our study demonstrates that forest management approaches aimed at restoring old-growth structure through targeted thinning do not greatly decrease buffering capacity, despite a known link between dense canopy and CCR. More broadly, this study illustrates the value of using remote sensing approaches to identify CCR, facilitating the integration of climate change adaptation with other forest management approaches.

Animal-mediated plant niche tracking in a changing climate.

Over half of plant species are animal-dispersed, and our understanding of how animals can help plants move in response to climate change - a process known as niche tracking - is limited, but advancing rapidly. Recent research efforts find evidence that animals are helping plants track their niches. They also identify key conditions needed for animal-mediated niche tracking to occur, including alignment of the timing of seed availability, the directionality of animal movements, and microhabitat conditions where seeds are deposited. A research framework that measures niche tracking effectiveness by considering all parts of the niche-tracking process, and links together data and models from multiple disciplines, will lead to further insight and inform actions to help ecosystems adapt to a changing world.

Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery.

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

MeadoWatch: a long-term community-science database of wildflower phenology in Mount Rainier National Park.

We present a long-term and high-resolution phenological dataset from 17 wildflower species collected in Mt. Rainier National Park, as part of the MeadoWatch (MW) community science project. Since 2013, 457 unique volunteers and scientists have gathered data on the timing of four key reproductive phenophases (budding, flowering, fruiting, and seeding) in 28 plots over two elevational gradients alongside popular park trails. Trained volunteers (87.2%) and University of  Washington scientists (12.8%) collected data 3-9 times/week during the growing season, using a standardized method. Taxonomic assessments were highly consistent between scientists and volunteers, with high accuracy and specificity across phenophases and species. Sensitivity, on the other hand, was lower than accuracy and specificity, suggesting that a few species might be challenging to reliably identify in community-science projects. Up to date, the MW database includes 42,000+ individual phenological observations from 17 species, between 2013 and 2019. However, MW is a living dataset that will be updated through continued contributions by volunteers, and made available for its use by the wider ecological community.

North American tree migration paced by climate in the West, lagging in the East.

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

Is there tree senescence? The fecundity evidence.

Despite its importance for forest regeneration, food webs, and human economies, changes in tree fecundity with tree size and age remain largely unknown. The allometric increase with tree diameter assumed in ecological models would substantially overestimate seed contributions from large trees if fecundity eventually declines with size. Current estimates are dominated by overrepresentation of small trees in regression models. We combined global fecundity data, including a substantial representation of large trees. We compared size-fecundity relationships against traditional allometric scaling with diameter and two models based on crown architecture. All allometric models fail to describe the declining rate of increase in fecundity with diameter found for 80% of 597 species in our analysis. The strong evidence of declining fecundity, beyond what can be explained by crown architectural change, is consistent with physiological decline. A downward revision of projected fecundity of large trees can improve the next generation of forest dynamic models.

Early snowmelt and warmer, drier summers shrink postflowering transition times in subalpine wildflowers.

Plant reproductive phenology-the timing of reproduction-is shifting rapidly with global climate change. Many studies focus on flowering responses to climate, but few investigate how postflowering processes, such as how quickly plants develop from flowering to seed dispersal, respond to environmental factors. We examined the climatic drivers of postflowering phenology in 28 species of western North American subalpine meadow plants over large spatial and temporal climate gradients. We took a Bayesian hierarchical approach to address whether and how climate influences the time it takes for wildflower populations to transition from flower to seed. Our previous work on the same species demonstrated that the initiation of flowering depends on snowmelt timing, with warmer temperatures and soil moisture also playing a role. Here, we found that for the majority of the flowering community, the same climate drivers also affected the time it takes to move from flowering to seed dispersal. Climate-sensitive species shortened flower-seed transitions when snow melted earlier, temperatures were warmer, and/or soil dried down more quickly-conditions we expect with higher frequency under climate change. Our work underscores the fact that predicting the impact of climate change on plant reproductive phenology demands empirical data on phases beyond flowering. Additionally, it suggests that some species face a future in which multiple environmental factors will push them towards more rapid transitions from flowering to postflowering phases, with potential effects on plants themselves and the many animal associates that rely on them, including frugivores and seed predators.

Spatial heterogeneity in ecologically important climate variables at coarse and fine scales in a high-snow mountain landscape.

Climate plays an important role in determining the geographic ranges of species. With rapid climate change expected in the coming decades, ecologists have predicted that species ranges will shift large distances in elevation and latitude. However, most range shift assessments are based on coarse-scale climate models that ignore fine-scale heterogeneity and could fail to capture important range shift dynamics. Moreover, if climate varies dramatically over short distances, some populations of certain species may only need to migrate tens of meters between microhabitats to track their climate as opposed to hundreds of meters upward or hundreds of kilometers poleward. To address these issues, we measured climate variables that are likely important determinants of plant species distributions and abundances (snow disappearance date and soil temperature) at coarse and fine scales at Mount Rainier National Park in Washington State, USA. Coarse-scale differences across the landscape such as large changes in elevation had expected effects on climatic variables, with later snow disappearance dates and lower temperatures at higher elevations. However, locations separated by small distances (∼20 m), but differing by vegetation structure or topographic position, often experienced differences in snow disappearance date and soil temperature as great as locations separated by large distances (>1 km). Tree canopy gaps and topographic depressions experienced later snow disappearance dates than corresponding locations under intact canopy and on ridges. Additionally, locations under vegetation and on topographic ridges experienced lower maximum and higher minimum soil temperatures. The large differences in climate we observed over small distances will likely lead to complex range shift dynamics and could buffer species from the negative effects of climate change.

'Natural experiment' demonstrates top-down control of spiders by birds on a landscape level.

The combination of small-scale manipulative experiments and large-scale natural experiments provides a powerful approach for demonstrating the importance of top-down trophic control on the ecosystem scale. The most compelling natural experiments have come from studies examining the landscape-scale loss of apex predators like sea otters, wolves, fish and land crabs. Birds are dominant apex predators in terrestrial systems around the world, yet all studies on their role as predators have come from small-scale experiments; the top-down impact of bird loss on their arthropod prey has yet to be examined at a landscape scale. Here, we use a unique natural experiment, the extirpation of insectivorous birds from nearly all forests on the island of Guam by the invasive brown tree snake, to produce the first assessment of the impacts of bird loss on their prey. We focused on spiders because experimental studies showed a consistent top-down effect of birds on spiders. We conducted spider web surveys in native forest on Guam and three nearby islands with healthy bird populations. Spider web densities on the island of Guam were 40 times greater than densities on islands with birds during the wet season, and 2.3 times greater during the dry season. These results confirm the general trend from manipulative experiments conducted in other systems however, the effect size was much greater in this natural experiment than in most manipulative experiments. In addition, bird loss appears to have removed the seasonality of spider webs and led to larger webs in at least one spider species in the forests of Guam than on nearby islands with birds. We discuss several possible mechanisms for the observed changes. Overall, our results suggest that effect sizes from smaller-scale experimental studies may significantly underestimate the impact of bird loss on spider density as demonstrated by this large-scale natural experiment.

Productivity is a poor predictor of plant species richness.

For more than 30 years, the relationship between net primary productivity and species richness has generated intense debate in ecology about the processes regulating local diversity. The original view, which is still widely accepted, holds that the relationship is hump-shaped, with richness first rising and then declining with increasing productivity. Although recent meta-analyses questioned the generality of hump-shaped patterns, these syntheses have been criticized for failing to account for methodological differences among studies. We addressed such concerns by conducting standardized sampling in 48 herbaceous-dominated plant communities on five continents. We found no clear relationship between productivity and fine-scale (meters(-2)) richness within sites, within regions, or across the globe. Ecologists should focus on fresh, mechanistic approaches to understanding the multivariate links between productivity and richness.