Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H 2 O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong eVects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone signiWcantly aVected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a signiWcant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results conWrm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.

In a laboratory study, we examined growth and survival of the caddisfly, Lepidostoma unicolor, feeding on two types of native leaf litter (Alnus rhombifolia [Alder] and a Salicaceae mix [Salix spp. and Populus fremontii]) and two invasive, non-native species (Tamarix ramosissima [saltcedar] and Arundo donax [giant reed]). Larval survival was high ([85%), and did not differ, among those fed Alnus, Salicaceae, or Tamarix litter, but was much lower (20%) for larvae fed Arundo litter. Mean dry biomass of larvae fed Tamarix was 45% greater than that of larvae fed Alnus, and both were significantly greater than the biomasses of insects fed Salicaceae or Arundo. Although both Alnus and Tamarix increased in percent nitrogen with conditioning,

In stream ecosystems, the growth of aquatic primary producers is affected by spatial and temporal variations in the riparian canopy, which can influence the availability of light resources. Aquatic plants can acclimate to low light environments by employing a suite of morphological or physiological mechanisms to increase light capture or photosynthetic efficiency. Some species may also use alternate types of propagules to colonize environments with heterogeneous light environments. In a greenhouse experiment we examined the morphological and physiological response of watercress (Nasturtium officinale R. Br.) to a gradient of increasing light levels, which ranged from 7% ambient light to full sunlight. We also determined if watercress seedlings and vegetative fragments differed in their growth response to increasing light levels. Total biomass and root biomass of seedlings and vegetative fragments decreased with decreasing light levels. The difference in plant biomass across treatments was due to morphological changes in total canopy area and leaf area, both of which increased with decreasing light levels. Seedlings and vegetative fragments did not differ in their response to light availability, but vegetative fragments had higher final biomass as a result of higher initial biomass. Physiological acclimation to low light levels appears to be of secondary importance for watercress as the concentrations of total chlorophyll, chlorophyll a, chlorophyll b, and chlorophyll a:b did not differ among light levels or between seedlings and vegetative fragments. Seedlings and vegetative fragments grown under high light levels had a greater percentage of carbon and a lower percentage of nitrogen than plants grown under low light conditions. The results of this study indicate that watercress displays considerable morphological plasticity and acclimates to low light conditions primarily by increasing leaf area and canopy surface area. There is no evidence that the type of watercress propagule (seedling vs. vegetative fragment) imparts any growth advantage in low light environments and watercress grown from either type of propagule showed no differences in their morphological or physiological responses to varying light regimes.

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