Jae R. Pasari

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.

Structure-function analysis of photosystem II subunit S (PsbS) in vivo

In land plants, photosystem II subunit S (PsbS) plays a key role in xanthophyll- and pH-dependent non-photochemical quenching (qE) of excess absorbed light energy. Arabidopsis thaliana (L.) Heynh. npq4 mutants are defective in the psbS gene and have impaired qE. Exactly how the PsbS protein is involved in qE is unclear, but it has been proposed that PsbS binds H+ and/or de-epoxidized xanthophylls in excess light as part of the qE mechanism. To identify amino acid residues that are important for PsbS function, we sequenced the psbS gene from eight npq4 point mutant alleles isolated by forward genetics screening, including two new alleles. In the four transmembrane helices of PsbS, several amino acid residues were found to affect the stability and/or function of the protein. By comparing the predicted amino acid sequences of PsbS from several plant species and studying the proposed topological structure of PsbS, eight possible H+-binding amino acid residues on the lumenal side of the protein were identified and then altered by site-directed mutagenesis in vitro. The mutant psbS genes were transformed into npq4-1, a psbS deletion mutant, to test the stability and function of the mutant PsbS proteins invivo. The results demonstrate that two conserved, protonatable amino acids, E122 and E226, are especially critical for the function of PsbS.

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 (β 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 β and γ diversity emerged only when multiple functions were considered simultaneously. Higher β 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.

Ecosystem Responses to Community Disassembly

Ecosystems around the world are experiencing unprecedented rates of extinction and species decline. The question of how community disassembly—the ongoing process of nonrandom species losses and declines—affects ecosystem functions, including those that influence persistence of other species, is addressed. The order in which species disappear from a community depends on their vulnerability to specific stressors and on traits associated with inherent susceptibility to decline. Information on species characteristics associated with vulnerability (response traits) is synthesized, and it is asked whether they are associated with characteristics that underpin significant contributions to ecosystem functioning (effect traits). Direct evidence that community disassembly affects ecosystem functioning comes from a variety of sources, ranging from documentation of long‐term changes following the loss of an initial species or fragmentation of a landscape, to modeling and manipulative experiments that simulate species losses and observe their consequences. The usefulness to conservation and restoration practice of community disassembly as a concept is evaluated, and it is asked whether and how community disassembly can provide guidance about species loss order, its consequences, what each of these depends on, and whether a positive link exists between vulnerability and contribution to function—a link that would exacerbate the consequences of the ongoing extinction crisis.

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 Californias 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 Ediths Bay checkerspot butterfly (Euphydryas editha bayensis).

Interactive Effects of Nitrogen Deposition and Grazing on Plant Species Composition in a Serpentine Grassland

Abstract The interaction of resource availability and disturbance can strongly affect plant species richness and the spread of exotic plants. Several ecological theories posit that disturbance mediates the richness-reducing effects of increased competition as resource levels rise. In the low-nutrient serpentine grasslands of the San Francisco Bay Area, the fertilizing effects of atmospheric nitrogen (N) deposition may threaten native species by promoting nitrophilic exotic grasses. Attempts to mitigate these N deposition effects have focused on cattle grazing as a strategy to reduce exotic grass cover. We simulated realistic N deposition increases with low-level fertilization, manipulated grazing with fencing, and monitored grazing intensity using camera traps in a 4 yr factorial experiment to assess the effects of grazing and N deposition on several measures of native and exotic species dynamics in Californias largest serpentine grassland. Our results suggest that native species diversity may increase slightly under low-level N deposition with moderate grazing in this system. However, grazing may not be effective at limiting exotic cover as N accumulates in the future. Examination of treatment trajectories using principal response curves indicated that responses to grazing might be determined more by functional group (forb or grass) than origin (native or exotic). Grazing intensity varied dramatically within the single stocking rate used to manage this ecosystem. Given this variation and the contrasting effects of grazing on different functional groups, more targeted management may be required to improve conservation outcomes.

Reply to Mokany et al: Comprehensive measures of biodiversity are critical to investigations of ecosystem multifunctionality.

Mokany et al. (1) reiterate an important consideration about the interpretation of results in our PNAS report (2). The authors suggest that the use of a different metric of β diversity might reveal a larger effect of β diversity on ecosystem multifunctionality. We agree, and reiterate that our reported effects of β diversity on ecosystem multifunctionality are likely conservative for a number of reasons. We also emphasize the importance of our findings despite these conservative results, and encourage future studies to explore creative methods for capturing more biodiversity effects.