Plants simultaneously maintain mutualistic relationships with different partners that are connected through the same host, but do not interact directly. One or more participating mutualists may alter their host's phenotype, resulting in a shift in the host's ecological interactions with all other mutualists involved. Understanding the functional interplay of mutualists associated with the same host remains an important challenge in biology. Here, we show belowground nitrogen-fixing rhizobia on lima bean (Phaseolus lunatus) alter their host plant's defensive mutualism with aboveground ants. We induced extrafloral nectar (EFN), an indirect defense acting through ant attraction. We also measured various nutritive and defensive plant traits, biomass, and counted ants on rhizobial and rhizobia-free plants. Rhizobia increased plant protein as well as cyanogenesis, a direct chemical defense against herbivores, but decreased EFN. Ants were significantly more attracted to rhizobia-free plants, and our structural equation model shows a strong link between rhizobia and reduced EFN as well as between EFN and ants: the sole path to ant recruitment. The rhizobia-mediated effects on simultaneously expressed defensive plant traits indicate rhizobia can have significant bottom-up effects on higher trophic levels. Our results show belowground symbionts play a critical and underestimated role in determining aboveground mutualistic interactions.

Polar systems are experiencing rapid climate change and the high sensitivity of these Arctic and Antarctic ecosystems make them especially vulnerable to accelerated ecological transformation. In Antarctica, warming results in a mosaic of ice-free terrestrial habitats dominated by a diverse assemblage of cryptogamic plants (i.e. mosses and lichens). Although these plants provide key habitat for a wide array of microorganisms and invertebrates, we have little understanding of the interaction between trophic levels in this terrestrial ecosystem and whether there are functional effects of plant species on higher trophic levels that may alter with warming. Here, we used open top chambers on Fildes Peninsula, King George Island, Antarctica, to examine the effects of passive warming and moss species on the abiotic environment and ultimately on higher trophic levels. For the dominant mosses, Polytrichastrum alpinum and Sanionia georgicouncinata, we found species-specific effects on the abiotic environment, including moss canopy temperature and soil moisture. In addition, we found distinct shifts in sexual expression in P. alpinum plants under warming compared to mosses without warming, and invertebrate communities in this moss species were strongly correlated with plant reproduction. Mosses under warming had substantially larger total invertebrate communities, and some invertebrate taxa were influenced differentially by moss species. However, warmed moss plants showed lower fungal biomass than control moss plants, and fungal biomass differed between moss species. Our results indicate that continued warming may impact the reproductive output of Antarctic moss species, potentially altering terrestrial ecosystems dynamics from the bottom up. Understanding these effects requires clarifying the foundational, mechanistic role that individual plant species play in mediating complex interactions in Antarctica's terrestrial food webs.

Carbon allocation demands from root-nodulating nitrogen-fixing bacteria (NFB) can modulate the host plant's chemical phenotype, with strong bottom-up effects on herbivores. In contrast to well-studied rhizobia, the effects of other important NFB on plant chemistry and herbivory are much less understood. Here, combining field surveys in the Oregon Coast Range, USA with laboratory experiments, we analyzed how N-fixing Frankia bacteria influenced plant growth, chemistry, and herbivory on Alnus rubra (red alder) seedlings. In the field, we quantified Frankia nodulation, herbivore damage, and plant size. In the laboratory, we grew seedlings with Frankia (F+), Frankia-free but nitrogen-fertilized (N+), or both uncolonized and unfertilized (F-N-) and assessed growth and leaf chemistry. We further conducted choice trials with black slugs, Arion rufus, a natural red alder herbivore. In the field, Frankia nodulation was significantly positively correlated with herbivory and negatively with seedling height. In contrast, in the lab, F+ as well as N+ seedlings were significantly taller than the F-N- controls. Seedlings from both treatments also had significantly increased leaf protein concentration compared to controls, whereas carbon-based nutritive compounds (carbohydrates) as well as leaf palatability-decreasing condensed tannins, lignin, and fiber were decreased in F+ but not in N+ treatments. In the choice assays, slugs preferred leaf material from F+ seedlings, but the effects were only significant in young leaves. Our study indicates that colonization by Frankia causes short-term ecological costs in terms of susceptibility to herbivory. However, the ubiquity of this symbiosis in natural settings suggests that these costs are outweighed by benefits beyond the seedling stage.

Mosses display a number of hallmark life history traits that influence their ecology at the population and community level. The long lived separation of sexes observed in the haploid gametophyte (dioicy) is one such feature of particular importance, as it is observed in the majority of bryophytes and creates intraspecific specialization of male and female individuals.The prevalence of sexually dimorphic mosses raises the possibility of sex-specific interactions with fungi as observed in some vascular plants. Here I investigated how moss sex shapes fungal communities associated with gametophytic tissues of the ubiquitous moss, Ceratodon purpureus. Using greenhouse populations of C. purpureus grown in a common garden, I examined fungal community structure and overall abundance of fungal biomass associated with male and female individuals from multiple populations. I hypothesized that individual mosses would harbor unique fungal communities based on their sex, and that overall fungal biomass associated with host tissues would differ significantly due to differences in morphological and physiological characteristics between the sexes. I found that fungal community composition and overall abundance (i.e. biomass) differ between male and female individuals of C. purpureus, and that sex-specific patterns are retained across individuals from three different populations. This work provides a first glance at how genetically based sexual systems in early land plants influence affiliated fungal community composition.ii Acknowledgments

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