No abstract

Over the past several years, plantings of California native plant nursery stock in restoration areas have become recognized as a pathway for invasive species introductions, in particular Phytophthora pathogens, including first in the U.S. detections (Phytophthora tentaculata, Phytophthora quercina), new taxa, new hybrid species, and dozens of other soilborne species. Restoration plantings may be conducted in high-value and limited habitats to sustain or re-establish rare plant populations. Once established, Phytophthora pathogens infest the site and are very difficult to eradicate or manage—they degrade the natural resources the plantings were intended to enhance. To respond to unintended Phytophthora introductions, vegetation ecologists took a variety of measures to prevent pathogen introduction and spread, including treating infested areas by solarization, suspending plantings, switching to direct seeding, applying stringent phytosanitation requirements on contracted nursery stock, and building their own nursery for clean plant production. These individual or collective actions, loosely coordinated by the Phytophthoras in Native Habitats Work Group ensued as demands intensified for protection from the inadvertent purchase of infected plants from commercial native plant nurseries. Regulation and management of the dozens of Phytophthora species and scores of plant hosts present a challenge to the state, county, and federal agriculture officials and to the ornamental and restoration nursery industries. To rebuild confidence in the health of restoration nursery stock and prevent further Phytophthora introductions, a voluntary, statewide accreditation pilot project is underway which, upon completion of validation, is planned for statewide implementation.

The actinobacterial genus Frankia establishes nitrogen-fixing root nodule symbioses with specific hosts within the nitrogen-fixing plant clade. Of four genetically distinct subgroups of Frankia, cluster I, II, and III strains are capable of forming effective nitrogen-fixing symbiotic associations, while cluster IV strains generally do not. Cluster II Frankia strains have rarely been detected in soil devoid of host plants, unlike cluster I or III strains, suggesting a stronger association with their host. To investigate the degree of host influence, we characterized the cluster II Frankia strain distribution in rhizosphere soil in three locations in northern California. The presence/absence of cluster II Frankia strains at a given site correlated significantly with the presence/absence of host plants on the site, as determined by glutamine synthetase (glnA) gene sequence analysis, and by microbiome analysis (16S rRNA gene) of a subset of host/nonhost rhizosphere soils. However, the distribution of cluster II Frankia strains was not significantly affected by other potential determinants such as host-plant species, geographical location, climate, soil pH, or soil type. Rhizosphere soil microbiome analysis showed that cluster II Frankia strains occupied only a minute fraction of the microbiome even in the host-plant-present site and further revealed no statistically significant difference in the ␣-diversity or in the microbiome composition between the host-plantpresent or -absent sites. Taken together, these data suggest that host plants provide a factor that is specific for cluster II Frankia strains, not a general growthpromoting factor. Further, the factor accumulates or is transported at the site level, i.e., beyond the host rhizosphere.IMPORTANCE Biological nitrogen fixation is a bacterial process that accounts for a major fraction of net new nitrogen input in terrestrial ecosystems. Transfer of fixed nitrogen to plant biomass is especially efficient via root nodule symbioses, which represent evolutionarily and ecologically specialized mutualistic associations. Frankia spp. (Actinobacteria), especially cluster II Frankia spp., have an extremely broad host range, yet comparatively little is known about the soil ecology of these organisms in relation to the host plants and their rhizosphere microbiomes. This study reveals a strong influence of the host plant on soil distribution of cluster II Frankia spp.

Widespread Phytophthora infections have been discovered in nursery stock used in California restoration plantings. In response, nursery Phytophthora best management practices (NPBMPs) designed to exclude Phytophthora from nursery plants were developed to address the need for clean planting stock in restoration projects. A pilot program to implement the systematic use of the NPBMPs, Accreditation to Improve Restoration (AIR), was developed and started in 2018. As of 2020, 13 northern California restoration nurseries have been evaluated and five have met all the program requirements. In 564 tests conducted over four years with a sensitive leachate baiting protocol, no Phytophthora was detected from over 20,000 nursery plants produced in compliance with the NPBMPs. In comparison, Phytophthora was detected in 25% of tests conducted on partially-compliant stock, and in 71% of tests from nurseries following few or no NPBMPs. The AIR pilot program has demonstrated that container stock free of detectable Phytophthora can be reliably produced by adhering to an integrated program of clean nursery production practices. To obtain Phytophthora-free plants for habitat restoration, informed clients were willing to pay increased costs required to produce NPBMP-compliant nursery stock.

Phytophthora pathogens are damaging native wildland vegetation including plants in restoration areas and botanic gardens. The infestations threaten some plants already designated as endangered and degrade high-value habitats. Pathogens are being introduced primarily via container plant nursery stock and, once established, they can spread to adjacent areas where plant species not previously exposed to pathogens may become infected. We review epidemics in California – caused by the sudden oak death pathogen Phytophthoraramorum Werres, De Cock & Man in ‘t Veld and the first USA detections of P. tentaculata Kröber & Marwitz, which occurred in native plant nurseries and restoration areas – as examples to illustrate these threats to conservation plantings.