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  1. Plants and herbivores are remarkably variable in space and time, and variability has been considered a defining feature of their interactions. Empirical research, however, has traditionally focused on understanding differences in means and overlooked the theoretically significant ecological and evolutionary roles of variability itself. We review the literature with the goal of showing how variability-explicit research expands our perspective on plant–herbivore ecology and evolution. We first clarify terminology for describing variation and then review patterns, causes, and consequences of variation in herbivory across scales of space, time, and biological organization. We consider how incorporating variability improves existing hypotheses and leads to new ones. We conclude by suggesting future work that reports full distributions, integrates effects of variation across scales, describes nonlinearities, and considers how stochastic and deterministic variation combine to determine herbivory distributions. Expected final online publication date for the Annual Review of Ecology, Evolution, and Systematics, Volume 54 is November 2023. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates. 
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    Free, publicly-accessible full text available November 2, 2024
  2. Background and Aims Human-driven nitrogen (N) deposition can alter soil biogeochemistry and plant communities, both critical to soil biota. However, understanding the relative impact of the relationship between nutrient resources and plants on soil communities has been hindered by a lack of experimental manipulations of both factors. We hypothesized that soil nematode communities would be structured predominantly by N addition via overall increased abundance, decreased diversity, and compositional shifts to dominance of r-selected bacterial-feeding nematodes. In contrast, we expected plant efects to be less evident and restricted to nematodes directly associated with plants. Methods We used a long-term (18-yrs) experiment in moist meadow alpine tundra involving N addition and codominant plant (nitrophilic Deschampsia cespitosa and nitrogen sensitive Geum rossii) removal. We characterized nematode communities via 18S rRNA metabarcoding and used soil biogeochemistry, plant, and microbial variables to determine factors shaping their communities. Results The N addition treatment increased overall nematode abundance, decreased diversity, and afected the composition of all nematode trophic groups. Overall, nematode communities shifted to dominance of bacterial feeding nematode taxa adapted to N-enriched environments. The likely drivers of this shift were increased soil nitrate and lower pH. The direct efects of codominant plants were more limited, with only changes in Geum rossii appearing to afect nematode responses. Conclusion Overall, nematode communities in N-limited alpine ecosystems are highly sensitive to increases in N availability, irrespective of the nature of N preferences of codominant plants. The resulting nematode community restructuring could signify future shifts in soil functioning throughout alpine landscapes. 
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    Free, publicly-accessible full text available October 13, 2024
  3. Abstract

    Crop diversification offers a promising solution to meet expanding global food demands while maintaining ecosystem services. Diversification strategies that use mixed planting to reduce pest damage (e.g. intercropping), termed ‘associational effects’ (AE) in the ecological literature, can decrease (associational resistance) or increase (associational susceptibility) herbivore abundance on a focal plant. While application of AE to agroecosystems typically reduces pest abundance, the range of outcomes varies widely.

    We conducted a meta‐analysis using 272 estimates of insect herbivore abundance on crops neighbored by a conspecific or heterospecific from 44 studies undertaken on six continents. We focus on four agricultural crops well represented from sites across the globe to test hypotheses related to understanding how herbivore traits (diet breadth, feeding guild, origin), plant traits (crop type, phylogenetic distance to neighbour) and environmental context (climate, experimental design) contribute to variation in the outcomes of AE.

    Overall, bicultures provided a strong reduction of insect abundance on the focal crop. Climate and interactions between herbivore traits, particularly diet breadth and origin, and plant traits or environmental context mediated the strength of AE.

    Bicultures provided the strongest reductions in insect abundance at low latitudes, and this effect decreased at higher latitudes but only for insects with certain traits. Abundance of generalist herbivores and globally distributed pests tended to be most strongly negatively affected by bicultures, under certain contexts, whereas specialist herbivores and native pests were less affected by neighbours.

    Synthesis and application. This meta‐analysis highlights that crop diversification schemes have an overall strongly beneficial effect of reducing pest abundance. However, there was also variability in the outcomes that is determined in part by the interactive effects of herbivore traits and environmental context. The results provide guidance for incorporating beneficial ecological interactions into integrated pest management strategies.

     
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  4. Abstract

    Herbivores often have highly variable impacts on plant fecundity. The relative contribution of different environmental factors operating at varying spatial scales in affecting this variability is often unclear. We examined how density‐dependent seed predation at local scales and regional differences in primary productivity are associated with variation in the magnitude of pre‐dispersal seed predation onMonarda fistulosa(Lamiaceae). WithinM. fistulosapopulations growing in a low‐productivity region (LPR), Montana, USA, and a high‐productivity region (HPR), Wisconsin, USA, we quantified the magnitude of pre‐dispersal seed predation among individual plants differing in seed head densities. Out of a total of 303M. fistulosaplants that were surveyed, we found half as many herbivores in seed heads in the LPR (n = 133 herbivores) compared to the HPR (n = 316). In the LPR, 30% of the seed heads were damaged in plants with low seed head density, while 61% of seed heads were damaged in plants with high seed head density. Seed head damage was consistently high in the HPR (about 49% across the range of seed head density) compared to the LPR (45% across a range of seed head density). However, the proportion of seeds per seed head that were destroyed by herbivores was nearly two times higher (~38% loss) in the LPR compared to HPR (22% loss). Considering the combined effects of probability of damage and seed loss per seed head, the proportion seed loss per plant was consistently higher in the HPR regardless of seed head density. Nevertheless, because of greater seed head production, the total number of viable seeds produced per plant was higher in HPR and high‐density plants, despite being exposed to greater herbivore pressure. These findings show how large‐scale factors can interact with local‐scale factors to influence how strongly herbivores suppress plant fecundity.

     
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  5. Abstract

    Recent declines in wild bee populations have led to increases in conservation actions and monitoring of bee communities. Pan traps are a commonly used sampling method for monitoring bee populations due to their efficiency and low cost. However, potential biases inherent in different sampling techniques may result in misleading characterizations of bee communities across space and time.

    In this paper, we examined how bee communities sampled using pan traps and aerial nets changed seasonally, and if they were affected by the availability of floral resources.

    We found strong seasonal changes in the abundance, but not the richness, of bees captured in pan traps. Notably, we captured the fewest bees during weeks in spring when most flowering plant species were in bloom, suggesting that floral resource availability influences pan trap captures. We also compared patterns of bee abundance in pan traps to those captured by aerial netting. Bee richness in pans and nets was positively correlated, but relative abundances in pan and net samples were dominated by different bee genera. Furthermore, most genera decreased in pans with increasing floral richness, but patterns were mixed for nets. When using presence/absence data, rather than abundance, community composition was more similar between netted and pan‐trapped bee communities and changed less substantially across the floral richness gradient.

    Overall, these differences led to sampling substantially different bee community compositions in pan traps versus nets, especially when using abundance‐based methods to characterize the bee community. By examining multiple years of intensive seasonal sampling of plant and bee communities, we document potential pitfalls with methods commonly used to sample bee communities.

    We suggest that pan trapping and aerial netting provide similar estimates of bee species richness and community composition when using presence/absence data, but that practitioners should interpret pan‐trapped bee abundance data with caution especially when comparing bee communities between sites where plant communities may differ.

     
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  6. Abstract

    A paradigm in the plant defence literature is that defending against herbivores comes at a cost to growth, resulting in a growth–defence trade‐off. However, while there is strong evidence for growth–defence trade‐offs across species, evidence is mixed within species.

    Several mechanisms can account for this equivocal support within species, but teasing them apart requires examining growth–defence relationships both within and among populations, an approach seldom employed.

    We examined correlations between plant biomass (growth) and terpene production (defence) within and among populations ofMonarda fistulosa, a perennial herb. We sampled populations from Montana and Wisconsin, regions that differ in resource availability characterized by different summer precipitation and associated abiotic conditions that influence plant productivity.

    We found negative, neutral and positive growth–defence correlations, depending on the scale examined. Negative correlations occurred across populations originating from divergent regions, positive correlations occurred across populations originating from within the high‐resource region and neutral correlations were found within single populations.

    Collectively, these results challenge the general expectation of ubiquitous trade‐offs and support emerging views that resource availability (as it affects productivity) shapes the evolution of defence at different scales.

    A freePlain Language Summarycan be found within the Supporting Information of this article.

     
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  7. Abstract

    Increasingly intensive strategies to maintain biodiversity and ecosystem function are being deployed in response to global anthropogenic threats, including intentionally introducing and eradicating species via assisted migration, rewilding, biological control, invasive species eradications, and gene drives. These actions are highly contentious because of their potential for unintended consequences. We conducted a global literature review of these conservation actions to quantify how often unintended outcomes occur and to elucidate their underlying causes. To evaluate conservation outcomes, we developed a community assessment framework for systematically mapping the range of possible interaction types for 111 case studies. Applying this tool, we quantified the number of interaction types considered in each study and documented the nature and strength of intended and unintended outcomes. Intended outcomes were reported in 51% of cases, a combination of intended outcomes and unintended outcomes in 26%, and strictly unintended outcomes in 10%. Hence, unintended outcomes were reported in 36% of all cases evaluated. In evaluating overall conservations outcomes (weighing intended vs. unintended effects), some unintended effects were fairly innocuous relative to the conservation objective, whereas others resulted in serious unintended consequences in recipient communities. Studies that assessed a greater number of community interactions with the target species reported unintended outcomes more often, suggesting that unintended consequences may be underreported due to insufficient vetting. Most reported unintended outcomes arose from direct effects (68%) or simple density‐mediated or indirect effects (25%) linked to the target species. Only a few documented cases arose from more complex interaction pathways (7%). Therefore, most unintended outcomes involved simple interactions that could be predicted and mitigated through more formal vetting. Our community assessment framework provides a tool for screening future conservation actions by mapping the recipient community interaction web to identify and mitigate unintended outcomes from intentional species introductions and eradications for conservation.

     
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  8. Abstract

    Biological organisms are increasingly being introduced and eradicated in an effort to maintain biodiversity and ecosystem function in the face of anthropogenic threats. However, these conservation actions can have unintended consequences to non‐target species. Careful vetting of these actions using ecological modelling tools could help predict and avoid unintended consequences.

    Qualitative modelling tools, such as fuzzy interaction webs (FIWs), allow for qualitative rankings of community properties (e.g. interaction strength = high, medium, low) in combination with quantitative information to predict management outcomes. These tools have lower data requirements than strictly quantitative models, facilitating their use for communities lacking comprehensive parameterization. However, no studies have evaluated the efficacy of FIWs for predicting unintended consequences against empirically documented outcomes. Moreover, there is no process for systematically identifying which species to incorporate in community‐level conservation assessments to overcome model structure uncertainty. Finally, there is a need to make qualitative modelling tools more accessible for conservation practitioners.

    We applied FIWs to the case study of lake trout introduction into Yellowstone Lake, Yellowstone National Park, to assess its ability to predict documented community‐level outcomes from an intentional species introduction. Next, we used the case study of the intentional red squirrel introduction to Newfoundland to show how a community assessment framework can help define the community interaction web needed for applying a FIW. Lastly, we introduced a user‐friendly web interface (https://matrix.mpgranch.com/#/) for applying FIWs to conservation questions.

    We found that the FIW predicted previously documented directional changes in the abundance of community components relatively well in the Yellowstone Lake case study, even with minimal knowledge of the system. The community assessment framework provided a formal process for identifying community components for the Newfoundland case study, and the resulting FIW predicted documented unintended consequences. The user interface predicts realistic outcomes in our study system and allows managers to build and apply FIWs for conservation planning.

    Synthesis and applications. Our community assessment framework and user interface can be used to apply FIWs to identify and avert potential unintended outcomes of species introductions and eradications for improved conservation management.

     
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  9. Abstract

    Despite theoretical advances, the ecological factors and functional traits that enable species varying in seed size and fecundity to coexist remain unclear. Given inherent fecundity advantages, why don't small‐seeded species dominate communities?

    In perennial grasslands, we evaluated whether small‐seeded species are less tolerant of competition from the community dominant bunchgrass than large‐seeded species, but also less vulnerable to seed predation by mice. We also explored whether trade‐offs involving competitive tolerance include two other functional traits, height and leaf mass per area (LMA). We added seeds of 17 forb species to plots where bunchgrass competition and rodent seed predation were manipulated across sites varying in bunchgrass productivity and thus competitive intensity. Seeds were added at densities mimicking interspecific variation in fecundity among target species.

    Standardizing for differences in fecundity (i.e. seed input, which enabled us to evaluate inherent interspecific differences in susceptibility to biotic interactions), bunchgrass competition more greatly reduced recruitment and establishment of small‐ versus large‐seeded species, whereas rodent seed predation more greatly reduced the recruitment of large‐ versus small‐seeded species. Plant height and LMA were unrelated to the competition effect size.

    Small‐seeded species abundance decreased across sites increasing in bunchgrass productivity, whereas this was not the case for large‐seeded species. For adult plants but not seedlings, community‐weighted functional trait means (CWM) for seed size, height and LMA increased in plots with versus without bunchgrass competition and the CWM for seed size and height also increased at sites with greater bunchgrass productivity (for adults only). In contrast, rodent seed predation had no significant effects on CWM seed size.

    At the end of the experiment, adult abundance positively correlated with plant fecundity in plots lacking bunchgrass, indicating the inherent advantages accrued to high fecundity small‐seeded species. However, with bunchgrass competition, abundances were equalized across species due to reduced competitive tolerance of high fecundity small‐seeded species.

    Synthesis. Our results suggest that coexistence among subordinate forb species varying in seed size and fecundity is in‐part due to a trade‐off involving competitive tolerance and fecundity, mediated by seed size and associated functional traits.

     
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