Brenda J. Grewell

PARASITE FACILITATES PLANT SPECIES COEXISTENCE IN A COASTAL WETLAND

Outbreaks of infectious agents in natural ecosystems are on the rise. Understanding host-pathogen interactions and their impact on community composition may be central to the conservation of biological diversity. Infectious agents can convey both exploitive and facilitative effects that regulate host populations and community structure. Parasitic angiosperms are highly conspicuous in many plant communities, and they provide a tractable model for understanding parasite effects in multispecies communities. I examined host identity and variation in host infectivity of a holoparasitic vine (Cuscuta salina) within a California salt marsh. In a two-year parasite removal experiment, I measured the effect of C. salina on its most frequent host, a rare hemiparasite, and the plant community. C. salina clearly suppressed the dominant host, but rare plant fitness and plant species diversity were enhanced through indirect effects. Priority effects played a role in the strength of the outcome due to the timing of life history characteristics. The differential influence of parasites on the fecundity of multiple hosts can change population dynamics, benefit rare species, and alter community structure. The continuum of negative to positive consequences of parasitic interactions deserves more attention if we are to understand community dynamics and successfully restore tidal wetlands.

Phenotypic plasticity of invasive Spartina densiflora (Poaceae) along a broad latitudinal gradient on the Pacific Coast of North America

PREMISE OF THE STUDY Phenotypic acclimation of individual plants and genetic differentiation by natural selection within invasive populations are two potential mechanisms that may confer fitness advantages and allow plants to cope with environmental variation. The invasion of Spartina densiflora across a wide latitudinal gradient from California (USA) to British Columbia (Canada) provides a natural model system to study the potential mechanisms underlying the response of invasive populations to substantial variation in climate and other environmental variables. METHODS We examined morphological and physiological leaf traits of Spartina densiflora plants in populations from invaded estuarine sites across broad latitudinal and climate gradients along the Pacific west coast of North America and in favorable conditions in a common garden experiment. KEY RESULTS Our results show that key foliar traits varied widely among populations. Most foliar traits measured in the field were lower than would be expected under ideal growing conditions. Photosynthetic pigment concentrations at higher latitudes were lower than those observed at lower latitudes. Greater leaf rolling, reduced leaf lengths, and lower chlorophyll and higher carbon concentrations were observed with anoxic sediments. Lower chlorophyll to carotenoids ratios and reduced nitrogen concentrations were correlated with sediment salinity. Our results suggest that the variations of foliar traits recorded in the field are a plastic phenotypic response that was not sustained under common garden conditions. CONCLUSIONS SPARTINA DENSIFLORA shows wide differences in its foliar traits in response to environmental heterogeneity in salt marshes, which appears to be the result of phenotypic plasticity rather than genetic differentiation.

Rare plants in the Golden Gate Estuary (California): the relationship between scale and understanding

We analysed 10 rare-plant projects conducted from 1982 to 2005 for trends in scientific findings, project costs, effort and efficacy. Our purpose was to determine whether generalisations about these factors can be found, and if so, whether they might be useful for designing and implementing successful future rare-plant endeavours. Analysis results revealed that rare plant species persist despite their restriction to a highly fragmented and degraded urbanised estuary of more than seven million people. Also important were the findings that with sufficient funding, successful rare-plant reintroduction is possible in the short-term at minimum. Habitat considerations, however, are paramount—i.e. habitat requirements of a rare species should be known before reintroduction for the restoration effort to be likely to be successful. Understanding ecosystem functions that support rare species, therefore, remains the highest priority for rare-plant restorationists. Project costs varied significantly, as expected, but a ‘middle ground’ provides necessary and sufficient funding to conduct most rare-plant work for one or a few species. Costs rise, but not linearly, when additional rare taxa are included. Given our experience is applicable elsewhere, taking an ecosystem approach to protection of rare flora is most successful and cost effective.

HEMIPARASITES GENERATE ENVIRONMENTAL HETEROGENEITY AND ENHANCE SPECIES COEXISTENCE IN SALT MARSHES

Tidal inundation and salinity are considered to be controlling factors in salt marsh species distributions. Parasitic plants may also influence community organization as parasite-host interactions may play a functional role in stress amelioration due to physiological mechanisms for salinity tolerance and resource acquisition. Endangered root hemiparasites (Cordylanthus maritimus ssp. palustris and Cordylanthus mollis ssp. mollis) occupy unique habitat within fragmented northern California tidal wetlands. My objective was to examine the effects of these root hemiparasites on soil salinity, aeration, and community composition. I compared experimentally established bare patches, shaded and unshaded, and parasite removal patches to controls with hemiparasites across intertidal elevation gradients. Plant community composition, soil salinity, and redox potential were measured as response variables. In this field removal experiment, I demonstrated that parasite-host associations can enhance the amelioration of physical stress conditions in the salt marsh exceeding the passive role of shading by vegetation. Consumer-driven reduction of physical stress resulted in increased plant species richness, and the effect was most pronounced with elevated salinity and hypoxia stress. Although previous studies have demonstrated that removal of dominant plant biomass by herbivores can increase physical stress in salt marshes, this is one of the first examples of a positive indirect effect of a consumer on community diversity through physical stress relief. Greater understanding of biological interactions coupled with abiotic factors may improve rare plant conservation and salt marsh restoration success.

Variation in tussock architecture of the invasive cordgrass Spartina densiflora along the Pacific Coast of North America

Some introduced species spread rapidly beyond their native range and into novel habitats mediated by a high degree of phenotypic plasticity and/or rapid evolutionary responses. In this context, clonality has been described as a significant factor contributing to invasiveness. We studied the abiotic environment and the responses of different tussock architecture traits of the invasive cordgrass Spartina densiflora Brongn. (Poaceae). A common garden experiment and field studies of S. densiflora in salt marshes across a wide latitudinal gradient from California (USA) to British Columbia (Canada) provided a model system for an integrated study of the potential mechanisms underlying the response of invasive S. densiflora populations to changes in environmental conditions. Our results showed that S. densiflora is able to adjust to widely variable climate (specifically, air temperature and the duration of the growing season) and sediment conditions (specifically, texture and hypoxia) through phenotypical plastic key functional tussock traits (e.g. shoot density, height, above- and below-ground biomass allocation patterns). Root biomass increased in coarser sediments in contrast to rhizomes, which were more abundant in finer sediments. Above-ground biomass and leaf area index increased mainly with air temperature during summer, and more robust (taller and wider) shoots were associated with more oxygenated sediments. In view of our results, S. densiflora appears to be a halophyte with a high degree of phenotypic plasticity that would enable it to respond successfully to changes in the abiotic conditions of salt marshes driven by global climate change, such as increasing salinity and temperatures.

Phenotypic plasticity and population differentiation in response to salinity in the invasive cordgrass Spartina densiflora

Salinity and tidal inundation induce physiological stress in vascular plant species and influence their distribution and productivity in estuarine wetlands. Climate change-induced sea level rise may magnify these abiotic stressors and the physiological stresses they can cause. Understanding the potential of invasive plants to respond to predicted salinity increases will elucidate their potential niche breadth. To examine potential phenotypic plasticity and functional trait responses to salinity stress in the invasive cordgrass Spartina densiflora, we collected rhizomes from four invasive populations occurring from California to Vancouver Island, British Columbia on the Pacific Coast of North America. In a glasshouse common garden experiment, we measured plant traits associated with growth and allocation, photosynthesis, leaf pigments, and leaf chemistry and calculated plasticity indices across imposed salinity treatments. Fifteen of 21 leaf chemistry, pigment, morphological and physiological traits expressed plastic responses to salinity. When averaged across all measured traits, degree of plasticity did not vary among sampled populations. However, differences in plasticity among populations in response to salinity were observed for 9 of 21 measured plant traits. Leaf chemistry and adaxial leaf rolling trait responses demonstrated the highest degree of plasticity, while growth and allocation measures were less plastic. Phenotypic plasticity of leaf functional traits to salinity indicates the potential of S. densiflora to maintain invasive growth in response to rising estuarine salinity with climate change.

Trait responses of invasive aquatic macrophyte congeners: colonizing diploid outperforms polyploid

Polyploidy (multiple copies of whole genomes) is over-represented in invasive plants and thought to promote their success in novel environments. Understanding functional traits supporting colonization can provide a foundation for development of effective management strategies. We compared how two aquatic invasive congeners differing in ploidy (diploid, decaploid) respond to resource availability (light, nutrients). Counter to our predictions, the diploid congener out-performed the decaploid with nutrient enrichment. Our results suggest the congeners have alternate colonization strategies, and trait responses underlying their success may change with ontogeny. Management strategies for invasive Ludwigia species should therefore be tailored for specific cytotypes and unique characteristics of their life stages.

Wastewater polishing by a channelized macrophyte-dominated wetland and anaerobic digestion of the harvested phytomass.

Constructed wetlands (CW) offer a mechanism to meet increasingly stringent regulatory standards for wastewater treatment while minimizing energy inputs. Additionally, harvested wetland phytomass subjected to anaerobic digestion can serve as a source of biogas methane. To investigate CW wastewater polishing activities and potential energy yield we constructed a pair of secondary wastewater-fed channelized CW modules designed to retain easily harvestable floating aquatic vegetation and maximize exposure of water to roots and sediment. Modules that were regularly harvested averaged a nitrate removal rate of 1.1 g N m−2 d−1; harvesting, sedimentation and gasification were responsible for 30.5%, 8.0% and 61.5% of the N losses, respectively. Selective harvesting of a module to maintain dominance of filamentous algae had no effect on nitrate removal rate but lowered productivity by one-half. The average monthly productivity for unselectively harvested modules was 9.3 ± 1.7 g dry wt. m−2 d−1 (±SE). Cessation of harvesting in one module resulted in a significant increase in nitrate removal rate and decrease in phosphate removal rate. Compared to the influent, the effluent of the harvested module had significantly lower levels of estrogenic activity, as determined by a quantitative PCR-based juvenile trout bioassay, and significantly lower densities of E. coli. In mixed vertical-flow reactors anaerobic co-digestion of equal dry weight proportions of harvested aquatic vegetation, wine yeast lees and dairy manure was greatly improved when the manure was replaced with the crude glycerol by-product of biodiesel production. Remaining solids were vermicomposted for use as a soil amendment. Our results indicate that incorporation of constructed wetlands into an integrated treatment system can simultaneously enhance the economic and energetic feasibility of wastewater and organic waste treatment processes.

Low genetic diversity contrasts with high phenotypic variability in heptaploid Spartina densiflora populations invading the Pacific coast of North America

Abstract Species can respond to environmental pressures through genetic and epigenetic changes and through phenotypic plasticity, but few studies have evaluated the relationships between genetic differentiation and phenotypic plasticity of plant species along changing environmental conditions throughout wide latitudinal ranges. We studied inter‐ and intrapopulation genetic diversity (using simple sequence repeats and chloroplast DNA sequencing) and inter‐ and intrapopulation phenotypic variability of 33 plant traits (using field and common‐garden measurements) for five populations of the invasive cordgrass Spartina densiflora Brongn. along the Pacific coast of North America from San Francisco Bay to Vancouver Island. Studied populations showed very low genetic diversity, high levels of phenotypic variability when growing in contrasted environments and high intrapopulation phenotypic variability for many plant traits. This intrapopulation phenotypic variability was especially high, irrespective of environmental conditions, for those traits showing also high phenotypic plasticity. Within‐population variation represented 84% of the total genetic variation coinciding with certain individual plants keeping consistent responses for three plant traits (chlorophyll b and carotenoid contents, and dead shoot biomass) in the field and in common‐garden conditions. These populations have most likely undergone genetic bottleneck since their introduction from South America; multiple introductions are unknown but possible as the population from Vancouver Island was the most recent and one of the most genetically diverse. S. densiflora appears as a species that would not be very affected itself by climate change and sea‐level rise as it can disperse, establish, and acclimate to contrasted environments along wide latitudinal ranges.

Temporal and nonlinear dispersal patterns of Ludwigia hexapetala in a regulated river

Rivers are vulnerable to biological invasion due to hydrologic connectivity, which facilitates post-entry movement of aquatic plant propagules by water currents. Ecological and watershed factors may influence spatial and temporal dispersal patterns. Field-based data on dispersal could improve risk assessment models and management responses. Ludwigia hexapetala, an invasive emergent macrophyte, provides a case study for understanding dispersal patterns throughout a watershed. The species spreads via hydrochory and is increasingly imposing detrimental ecological and economic impacts within watersheds of the United States and Europe. We investigated morphology of shoot fragments and their dispersal in the Russian River watershed of California, capturing shoot fragments of L. hexapetala during repeated summer surveys at five locations in the river and quantifying their morphological traits that predict establishment success. Highly variable capture counts suggest the importance of pulse disturbance events in local dispersal of L. hexapetala. Unexpectedly, dispersing propagule pressure was nonlinear, with more shoot fragments captured in the middle rather than lower river. Captured fragments in the middle river were twice the length of fragments captured in the lower river and bore 83% more stem nodes, characteristics associated with greater establishment success. Our results support development of spatially targeted management, outreach, and prevention efforts that could lead to decreased propagule pressure in the watershed.