Trait-based filtering mediates the effects of realistic biodiversity losses on ecosystem functioning.

Biodiversity losses are a major driver of global changes in ecosystem functioning. While most studies of the relationship between biodiversity and ecosystem functioning have examined randomized species losses, trait-based filtering associated with species-specific vulnerability to drivers of diversity loss can strongly influence how ecosystem functioning responds to declining biodiversity. Moreover, the responses of ecosystem functioning to diversity loss may be mediated by environmental variability interacting with the suite of traits remaining in depauperate communities. We do not yet understand how communities resulting from realistic diversity losses (filtered by response traits) influence ecosystem functioning (via effect traits of the remaining community), especially under variable environmental conditions. Here, we directly test how realistic and randomized plant diversity losses influence productivity and invasion resistance across multiple years in a California grassland. Compared with communities based on randomized diversity losses, communities resulting from realistic (drought-driven) species losses had higher invasion resistance under climatic conditions that matched the trait-based filtering they experienced. However, productivity declined more with realistic than with randomized species losses across all years, regardless of climatic conditions. Functional response traits aligned with effect traits for productivity but not for invasion resistance. Our findings illustrate that the effects of biodiversity losses depend not only on the identities of lost species but also on how the traits of remaining species interact with varying environmental conditions. Understanding the consequences of biodiversity change requires studies that evaluate trait-mediated effects of species losses and incorporate the increasingly variable climatic conditions that future communities are expected to experience.

Phylogenomic Data Reveal Widespread Introgression Across the Range of an Alpine and Arctic Specialist.

Understanding how gene flow affects population divergence and speciation remains challenging. Differentiating one evolutionary process from another can be difficult because multiple processes can produce similar patterns, and more than one process can occur simultaneously. Although simple population models produce predictable results, how these processes balance in taxa with patchy distributions and complicated natural histories is less certain. These types of populations might be highly connected through migration (gene flow), but can experience stronger effects of genetic drift and inbreeding, or localized selection. Although different signals can be difficult to separate, the application of high-throughput sequence data can provide the resolution necessary to distinguish many of these processes. We present whole-genome sequence data for an avian species group with an alpine and arctic tundra distribution to examine the role that different population genetic processes have played in their evolutionary history. Rosy-finches inhabit high elevation mountaintop sky islands and high-latitude island and continental tundra. They exhibit extensive plumage variation coupled with low levels of genetic variation. Additionally, the number of species within the complex is debated, making them excellent for studying the forces involved in the process of diversification, as well as an important species group in which to investigate species boundaries. Total genomic variation suggests a broadly continuous pattern of allele frequency changes across the mainland taxa of this group in North America. However, phylogenomic analyses recover multiple distinct, well supported, groups that coincide with previously described morphological variation and current species-level taxonomy. Tests of introgression using D-statistics and approximate Bayesian computation reveal significant levels of introgression between multiple North American taxa. These results provide insight into the balance between divergent and homogenizing population genetic processes and highlight remaining challenges in interpreting conflict between different types of analytical approaches with whole-genome sequence data. [ABBA-BABA; approximate Bayesian computation; gene flow; phylogenomics; speciation; whole-genome sequencing.].

Flowering phenology shifts in response to biodiversity loss.

Observational studies and experimental evidence agree that rising global temperatures have altered plant phenology-the timing of life events, such as flowering, germination, and leaf-out. Other large-scale global environmental changes, such as nitrogen deposition and altered precipitation regimes, have also been linked to changes in flowering times. Despite our increased understanding of how abiotic factors influence plant phenology, we know very little about how biotic interactions can affect flowering times, a significant knowledge gap given ongoing human-caused alteration of biodiversity and plant community structure at the global scale. We experimentally manipulated plant diversity in a California serpentine grassland and found that many plant species flowered earlier in response to reductions in diversity, with peak flowering date advancing an average of 0.6 days per species lost. These changes in phenology were mediated by the effects of plant diversity on soil surface temperature, available soil N, and soil moisture. Peak flowering dates were also more dispersed among species in high-diversity plots than expected based on monocultures. Our findings illustrate that shifts in plant species composition and diversity can alter the timing and distribution of flowering events, and that these changes to phenology are similar in magnitude to effects induced by climate change. Declining diversity could thus contribute to or exacerbate phenological changes attributed to rising global temperatures.

Meta-analysis of the effects of upstream land cover on stream recovery.

Unpredictable or variable ecosystem recovery from disturbance presents a challenge to conservation, particularly as the scale of human disturbance continues to increase. Theory suggests land-cover and disturbance characteristics affect recovery, but individual studies of disturbance and recovery frequently struggle to uncover generalizable patterns because of high levels of site-specific variation. To understand how land-cover, disturbance type, and disturbance duration influence ecosystem recovery, we used studies documenting recovery of 50 streams to perform a global meta-analysis of stream recovery from disturbances that affect water quality (e.g., oil spill, fire, wastewater). We extracted upstream natural and urban land-cover percentages for each site and performed model selection and averaging to identify influences on recovery completeness. Most streams improved following the end of a disturbance (median 240% of disturbed condition) but did not recover fully to baseline predisturbance condition within the studied period (median study period 2 years; median recovery 60% of baseline). Scale of disturbance in time and space did not predict recovery, but sites with higher percentages of upstream natural land cover had less complete recovery relative to sites with more urban or agricultural cover, possibly due to higher baseline conditions in these streams. Our findings suggest impacts to systems with low anthropogenic stress may be more irreversible than impacts to already modified systems. We call for more long-term evaluations of ecosystem response to disturbance and the inclusion of regional references and predisturbance reference conditions for comparison. A more thorough understanding of the role of the surrounding landscape in shaping stream response to disturbance can help managers calibrate expectations for recovery and prioritize protection.

Meta-Análisis de los Efectos de la Cobertura Río-Arriba sobre la Restauración de Arroyos Resumen La restauración impredecible o variable de un ecosistema después de una perturbación presenta un reto para la conservación, particularmente conforme la escala de perturbaciones humanas continúa incrementando. La teoría sugiere que la cobertura de suelo y las características de la perturbación afectan a la restauración pero los estudios individuales sobre las perturbaciones y las restauraciones constantemente luchan por descubrir patrones generalizables debido a los niveles altos de variación específica en el sitio. Usamos estudios que documentan la restauración de 50 arroyos para realizar un meta-análisis global de la restauración de arroyos después de perturbaciones que afectaron la calidad del agua (p. ej.: derrames de petróleo, incendios, aguas negras) y así entender cómo la cobertura de suelo, el tipo de perturbación, y la duración de la perturbación influyen sobre la restauración del ecosistema. Extrajimos porcentajes de la cobertura natural y urbana de suelo río-arriba para cada sitio y realizamos una selección y promedio de modelos para identificar las influencias sobre la completitud de la restauración. La mayoría de los arroyos mejoraron después de que terminó la perturbación (mediana del 240% de la condición perturbada) pero no se recuperó completamente hasta la línea base de condiciones previas a la perturbación dentro del periodo estudiado (mediana del periodo de estudio: dos años; mediana de la restauración 60% de la línea base). La escala de perturbación en el tiempo y en el espacio no pronosticó la restauración, pero los sitios con porcentajes más altos de cobertura natural de suelo río-arriba tuvieron una restauración menos completa en relación con los sitios con una cobertura más urbana o agrícola, posiblemente debido a las condiciones más altas de línea base en estos arroyos. Nuestros hallazgos sugieren que los impactos sobre los sistemas con un bajo estrés antropogénico pueden ser más irreversibles que los impactos sobre sistemas que ya han sido modificados. Hacemos un llamado por más evaluaciones a largo plazo de la respuesta de los ecosistemas ante las perturbaciones y por la inclusión de referencias regionales y condiciones previas a la perturbación como referencia para realizar comparaciones. Un entendimiento más a fondo del papel del paisaje circundante en la formación de la respuesta de los arroyos ante las perturbaciones puede ayudar a los administradores a calibrar expectativas para la restauración y a priorizar la protección.

High plant diversity is needed to maintain ecosystem services.

Biodiversity is rapidly declining worldwide, and there is consensus that this can decrease ecosystem functioning and services. It remains unclear, though, whether few or many of the species in an ecosystem are needed to sustain the provisioning of ecosystem services. It has been hypothesized that most species would promote ecosystem services if many times, places, functions and environmental changes were considered; however, no previous study has considered all of these factors together. Here we show that 84% of the 147 grassland plant species studied in 17 biodiversity experiments promoted ecosystem functioning at least once. Different species promoted ecosystem functioning during different years, at different places, for different functions and under different environmental change scenarios. Furthermore, the species needed to provide one function during multiple years were not the same as those needed to provide multiple functions within one year. Our results indicate that even more species will be needed to maintain ecosystem functioning and services than previously suggested by studies that have either (1) considered only the number of species needed to promote one function under one set of environmental conditions, or (2) separately considered the importance of biodiversity for providing ecosystem functioning across multiple years, places, functions or environmental change scenarios. Therefore, although species may appear functionally redundant when one function is considered under one set of environmental conditions, many species are needed to maintain multiple functions at multiple times and places in a changing world.

Several scales of biodiversity affect ecosystem multifunctionality.

Society values landscapes that reliably provide many ecosystem functions. As the study of ecosystem functioning expands to include more locations, time spans, and functions, the functional importance of individual species is becoming more apparent. However, the functional importance of individual species does not necessarily translate to the functional importance of biodiversity measured in whole communities of interacting species. Furthermore, ecological diversity at scales larger than neighborhood species richness could also influence the provision of multiple functions over extended time scales. We created experimental landscapes based on whole communities from the world's longest running biodiversity-functioning field experiment to investigate how local species richness (α diversity), distinctness among communities (ß diversity), and larger scale species richness (γ diversity) affected eight ecosystem functions over 10 y. Using both threshold-based and unique multifunctionality metrics, we found that α diversity had strong positive effects on most individual functions and multifunctionality, and that positive effects of ß and γ diversity emerged only when multiple functions were considered simultaneously. Higher ß diversity also reduced the variability in multifunctionality. Thus, in addition to conserving important species, maintaining ecosystem multifunctionality will require diverse landscape mosaics of diverse communities.

Species traits outweigh nested structure in driving the effects of realistic biodiversity loss on productivity.

While most studies of the relationship between biodiversity and ecosystem functioning have examined randomized diversity losses, several recent experiments have employed nested, realistic designs and found that realistic species losses had larger consequences than random losses for ecosystem functioning. Progressive, realistic, biodiversity losses are generally strongly nested, but this nestedness is a potentially confounding effect. Here, we address whether nonrandom trait loss or degree of nestedness drives the relationship between diversity and productivity in a realistic biodiversity-loss experiment. We isolated the effect of nestedness through post hoc analyses of data from an experimental biodiversity manipulation in a California serpentine grassland. We found that the order in which plant traits are lost as diversity declines influences the diversity-productivity relationship more than the degree of nestedness does. Understanding the relationship between the expected order of species loss and functional traits is becoming increasingly important in the face of ongoing biodiversity loss worldwide. Our findings illustrate the importance of species composition and the order of species loss, rather than nestedness per se, for understanding the mechanisms underlying the effects of realistic species losses on ecosystem functioning.

Grazing maintains native plant diversity and promotes community stability in an annual grassland.

Maintaining native biodiversity in grasslands requires management and mitigation of anthropogenic changes that have altered resource availability, grazing regimes, and community composition. In California (USA), high levels of atmospheric nitrogen (N) deposition have facilitated the invasion of exotic grasses, posing a threat to the diverse plant and insect communities endemic to serpentine grasslands. Cattle grazing has been employed to mitigate the consequences of exotic grass invasion, but the ecological effects of grazing in this system are not fully understood. To characterize the effects of realistic N deposition on serpentine plant communities and to evaluate the efficacy of grazing as a management tool, we performed a factorial experiment adding N and excluding large herbivores in California's largest serpentine grassland. Although we observed significant interannual variation in community composition related to climate in our six-year study, exotic cover was consistently and negatively correlated with native plant richness. Sustained low-level N addition did not influence plant community composition, but grazing reduced grass abundance while maintaining greater native forb cover, native plant diversity, and species richness in comparison to plots excluding large herbivores. Furthermore, grazing increased the temporal stability of plant communities by decreasing year-to-year variation in native forb cover, native plant diversity, and native species richness. Taken together, our findings demonstrate that moderate-intensity cattle grazing can be used to restrict the invasive potential of exotic grasses and maintain native plant communities in serpentine grasslands. We hypothesize that the reduced temporal variability in serpentine plant communities managed by grazing may directly benefit populations of the threatened Edith's Bay checkerspot butterfly (Euphydryas editha bayensis).

Realistic diversity loss and variation in soil depth independently affect community-level plant nitrogen use.

Numerous experiments have demonstrated that diverse plant communities use nitrogen (N) more completely and efficiently, with implications for how species conservation efforts might influence N cycling and retention in terrestrial ecosystems. However, most such experiments have randomly manipulated species richness and minimized environmental heterogeneity, two design aspects that may reduce applicability to real ecosystems. Here we present results from an experiment directly comparing how realistic and randomized plant species losses affect plant N use across a gradient of soil depth in a native-dominated serpentine grassland in California. We found that the strength of the species richness effect on plant N use did not increase with soil depth in either the realistic or randomized species loss scenarios, indicating that the increased vertical heterogeneity conferred by deeper soils did not lead to greater complementarity among species in this ecosystem. Realistic species losses significantly reduced plant N uptake and altered N-use efficiency, while randomized species losses had no effect on plant N use. Increasing soil depth positively affected plant N uptake in both loss order scenarios but had a weaker effect on plant N use than did realistic species losses. Our results illustrate that realistic species losses can have functional consequences that differ distinctly from randomized losses, and that species diversity effects can be independent of and outweigh those of environmental heterogeneity on ecosystem functioning. Our findings also support the value of conservation efforts aimed at maintaining biodiversity to help buffer ecosystems against increasing anthropogenic N loading.

Imperfect replacement of native species by non-native species as pollinators of endemic Hawaiian plants.

Native plant species that have lost their mutualist partners may require non-native pollinators or seed dispersers to maintain reproduction. When natives are highly specialized, however, it appears doubtful that introduced generalists will partner effectively with them. We used visitation observations and pollination treatments (experimental manipulations of pollen transfer) to examine relationships between the introduced, generalist Japanese White-eye (Zosterops japonicus) and 3 endemic Hawaiian plant species (Clermontia parviflora, C. montis-loa, and C. hawaiiensis). These plants are characterized by curved, tubular flowers, apparently adapted for pollination by curve-billed Hawaiian honeycreepers. Z. japonicus were responsible for over 80% of visits to flowers of the small-flowered C. parviflora and the midsize-flowered C. montis-loa. Z. japonicus-visited flowers set significantly more seed than did bagged flowers. Z. japonicus also demonstrated the potential to act as an occasional Clermontia seed disperser, although ground-based frugivory by non-native mammals likely dominates seed dispersal. The large-flowered C. hawaiiensis received no visitation by any birds during observations. Unmanipulated and bagged C. hawaiiensis flowers set similar numbers of seeds. Direct examination of Z. japonicus and Clermontia morphologies suggests a mismatch between Z. japonicus bill morphology and C. hawaiiensis flower morphology. In combination, our results suggest that Z. japonicus has established an effective pollination relationship with C. parviflora and C. montis-loa and that the large flowers of C. hawaiiensis preclude effective visitation by Z. japonicus.

Beyond reporting: proactive strategies for safer scientific fieldwork.

Fieldwork is crucial for science but poses heightened risks of gender-based harassment and assault. Current practices prioritize post-incident reporting, despite the demonstrated potential of preventive approaches. We recommend proactive practices, training strategies, and systemic policy changes to build safe and inclusive fieldwork settings from the outset.

Testing the effectiveness of interactive training on sexual harassment and assault in field science.

Fieldwork is a critical tool for scientific research, particularly in applied disciplines. Yet fieldwork is often unsafe, especially for members of historically marginalized groups and people whose presence in scientific spaces threatens traditional hierarchies of power, authority, and legitimacy. Research is needed to identify interventions that prevent sexual harassment and assault from occurring in the first place. We conducted a quasi-experiment assessing the impacts of a 90-min interactive training on field-based staff in a United States state government agency. We hypothesized that the knowledge-based interventions, social modeling, and mastery experiences included in the training would increase participants' sexual harassment and assault prevention knowledge, self-efficacy, behavioural intention, and behaviour after the training compared to a control group of their peers. Treatment-control and pre-post training survey data indicate that the training increased participants' sexual harassment and assault prevention knowledge and prevention self-efficacy, and, to a lesser extent, behavioural intention. These increases persisted several months after the training for knowledge and self-efficacy. While we did not detect differences in the effect of the training for different groups, interestingly, post-hoc tests indicated that women and members of underrepresented racial groups generally scored lower compared to male and white respondents, suggesting that these groups self-assess their own capabilities differently. Finally, participants' likelihood to report incidents increased after the training but institutional reports remained low, emphasizing the importance of efforts to transform reporting systems and develop better methods to measure bystander actions. These results support the utility of a peer-led interactive intervention for improving workplace culture and safety in scientific fieldwork settings. PROTOCOL REGISTRATION: "The stage 1 protocol for this Registered Report was accepted in principle on August 24, 2022. The protocol, as accepted by the journal, can be found at: https://doi.org/10.6084/m9.figshare.21770165 .

Sustaining multiple ecosystem functions in grassland communities requires higher biodiversity.

Society places value on the multiple functions of ecosystems from soil fertility to erosion control to wildlife-carrying capacity, and these functions are potentially threatened by ongoing biodiversity losses. Recent empirically based models using individual species' traits suggest that higher species richness is required to provide multiple ecosystem functions. However, no study to date has analyzed the observed functionality of communities of interacting species over multiple temporal scales to assess the relationship between biodiversity and multifunctionality. We use data from the longest-running biodiversity-functioning field experiment to date to test how species diversity affects the ability of grassland ecosystems to provide threshold levels of up to eight ecosystem functions simultaneously. Across years and every combination of ecosystem functions, minimum-required species richness consistently increases with the number of functions considered. Moreover, tradeoffs between functions and variability among years prevent any one community type from providing high levels of multiple functions, regardless of its diversity. Sustained multifunctionality, therefore, likely requires both higher species richness than single ecosystem functionality and a diversity of species assemblages across the landscape.

Abundance declines of a native forb have nonlinear impacts on grassland invasion resistance.

The effects of declining plant biodiversity on ecosystem processes are well studied, with most investigations examining the role of species richness declines rather than declines of species abundance. Using grassland mesocosms, we examined how the abundance of a native, resident species, Hemizonia congesta (hayfield tarweed), affected exotic Centaurea solstitialis (yellow starthistle) invasion. We found that progressive H. congesta abundance declines had threshold effects on invasion resistance, with initial declines resulting in minor increases in invasion and subsequent declines leading to accelerating increases in invader performance. Reduced invasion resistance was explained by increased resource availability as H. congesta declined. We also found evidence that resident abundance might indirectly affect invasion by mediating invader impact on resident competitors; C. solstitialis disproportionately reduced H. congesta biomass in low-abundance rather than high-abundance populations. H. congesta's direct and indirect effects on invasion resistance illustrate that an individual species' declining abundance can have accelerating, deleterious effects on ecosystem functions of conservation value.

Phenological tracking enables positive species responses to climate change.

Earlier spring phenology observed in many plant species in recent decades provides compelling evidence that species are already responding to the rising global temperatures associated with anthropogenic climate change. There is great variability among species, however, in their phenological sensitivity to temperature. Species that do not phenologically "track" climate change may be at a disadvantage if their growth becomes limited by missed interactions with mutualists, or a shorter growing season relative to earlier-active competitors. Here, we set out to test the hypothesis that phenological sensitivity could be used to predict species performance in a warming climate, by synthesizing results across terrestrial warming experiments. We assembled data for 57 species across 24 studies where flowering or vegetative phenology was matched with a measure of species performance. Performance metrics included biomass, percent cover, number of flowers, or individual growth. We found that species that advanced their phenology with warming also increased their performance, whereas those that did not advance tended to decline in performance with warming. This indicates that species that cannot phenologically "track" climate may be at increased risk with future climate change, and it suggests that phenological monitoring may provide an important tool for setting future conservation priorities.

Effects of native and non-native vertebrate mutualists on plants.

Extinctions can leave species without mutualist partners and thus potentially reduce their fitness. In cases where non-native species function as mutualists, mutualism disruption associated with species' extinction may be mitigated. To assess the effectiveness of mutualist species with different origins, we conducted a meta-analysis in which we compared the effectiveness of pollination and seed-dispersal functions of native and non-native vertebrates. We used data from 40 studies in which a total of 34 non-native vertebrate mutualists in 20 geographic locations were examined. For each plant species, opportunistic non-native vertebrate pollinators were generally less effective mutualists than native pollinators. When native mutualists had been extirpated, however, plant seed set and seedling performance appeared elevated in the presence of non-native mutualists, although non-native mutualists had a negative overall effect on seed germination. These results suggest native mutualists may not be easily replaced. In some systems researchers propose taxon substitution or the deliberate introduction of non-native vertebrate mutualists to reestablish mutualist functions such as pollination and seed dispersal and to rescue native species from extinction. Our results also suggest that in places where all native mutualists are extinct, careful taxon substitution may benefit native plants at some life stages.

The Adaptation for Conservation Targets (ACT) framework: a tool for incorporating climate change into natural resource management.

As natural resource management agencies and conservation organizations seek guidance on responding to climate change, myriad potential actions and strategies have been proposed for increasing the long-term viability of some attributes of natural systems. Managers need practical tools for selecting among these actions and strategies to develop a tailored management approach for specific targets at a given location. We developed and present one such tool, the participatory Adaptation for Conservation Targets (ACT) framework, which considers the effects of climate change in the development of management actions for particular species, ecosystems and ecological functions. Our framework is based on the premise that effective adaptation of management to climate change can rely on local knowledge of an ecosystem and does not necessarily require detailed projections of climate change or its effects. We illustrate the ACT framework by applying it to an ecological function in the Greater Yellowstone Ecosystem (Montana, Wyoming, and Idaho, USA)--water flows in the upper Yellowstone River. We suggest that the ACT framework is a practical tool for initiating adaptation planning, and for generating and communicating specific management interventions given an increasingly altered, yet uncertain, climate.

Climate-induced reversal of tree growth patterns at a tropical treeline.

Globally, cold-limited trees and forests are expected to experience growth acceleration as a direct response to warming temperatures. However, thresholds of temperature limitation may vary substantially with local environmental conditions, leading to heterogeneous responses in tree ecophysiology. We used dendroecological and isotopic methods to quantify shifting tree growth and resource use over the past 143 years across topographic aspects in a high-elevation forest of central Mexico. Trees on south-facing slopes (SFS) grew faster than those on north-facing slopes (NFS) until the mid-20th century, when this pattern reversed notably with marked growth rate declines on SFS and increases on NFS. Stable isotopes of carbon, oxygen, and carbon-to-nitrogen ratios suggest that this reversal is linked to interactions between CO2 stimulation of photosynthesis and water or nitrogen limitation. Our findings highlight the importance of incorporating landscape processes and habitat heterogeneity in predictions of tree growth responses to global environmental change.