Local groundwater decline exacerbates response of dryland riparian woodlands to climatic drought.

Dryland riparian woodlands are considered to be locally buffered from droughts by shallow and stable groundwater levels. However, climate change is causing more frequent and severe drought events, accompanied by warmer temperatures, collectively threatening the persistence of these groundwater dependent ecosystems through a combination of increasing evaporative demand and decreasing groundwater supply. We conducted a dendro-isotopic analysis of radial growth and seasonal (semi-annual) carbon isotope discrimination (Δ13 C) to investigate the response of riparian cottonwood stands to the unprecedented California-wide drought from 2012 to 2019, along the largest remaining free-flowing river in Southern California. Our goals were to identify principal drivers and indicators of drought stress for dryland riparian woodlands, determine their thresholds of tolerance to hydroclimatic stressors, and ultimately assess their vulnerability to climate change. Riparian trees were highly responsive to drought conditions along the river, exhibiting suppressed growth and strong stomatal closure (inferred from reduced Δ13 C) during peak drought years. However, patterns of radial growth and Δ13 C were quite variable among sites that differed in climatic conditions and rate of groundwater decline. We show that the rate of groundwater decline, as opposed to climate factors, was the primary driver of site differences in drought stress, and trees showed greater sensitivity to temperature at sites subjected to faster groundwater decline. Across sites, higher correlation between radial growth and Δ13 C for individual trees, and higher inter-correlation of Δ13 C among trees were indicative of greater drought stress. Trees showed a threshold of tolerance to groundwater decline at 0.5 m year-1 beyond which drought stress became increasingly evident and severe. For sites that exceeded this threshold, peak physiological stress occurred when total groundwater recession exceeded ~3 m. These findings indicate that drought-induced groundwater decline associated with more extreme droughts is a primary threat to dryland riparian woodlands and increases their susceptibility to projected warmer temperatures.

Performance of a Native Butterfly and Introduced Moth on Native and Introduced Lineages of Phragmites australis.

This study examined the performance of Poanes viator (Edwards) (Hesperiidae), a native North American skipper, and Rhizedra lutosa (Hübner) (Noctuidae), an introduced moth, reared on native and non-native, invasive lineages of Phragmites australis. Poanes viator is a generalist on monocots and larvae were also fed leaves of Zizania aquatica, a native macrophyte that the skipper commonly uses as a host plant. Larval survival and duration, pupal weight, and pupation time were compared for P. viator feeding on leaf tissue and R. lutosa feeding on rhizomes of either native or introduced plants. We also tested an artificial diet supplemented with P. australis rhizome powder as a potential food for rearing other stalk and rhizome boring Lepidoptera. In experiments using excised plant tissues, some individuals of both species fed and developed to the pupal stage on native and introduced plants, but overall, larval survival rates were low. Plant species/haplotype identity did not cause strong differences in larval survival for either species. However, P. viator larvae only pupated when feeding on native plants (Zizania aquatica and native P. australis haplotypes), whereas R. lutosa successfully pupated on both native and introduced P. australis. Although larval survival was low, 100% of P. viator and 95% of R. lutosa that reached the pupal stage emerged as adults. Rhizedra lutosa larvae fed an artificial diet supplemented with P. australis rhizome powder had significantly greater survival and pupal weights, and shorter pupation times than larvae fed rhizomes only. Several specialist Lepidopteran species are being considered for approval as biological control agents for the non-native P. australis haplotype, and the convenience and increased larval performance make this artificial diet a good alternative for rearing organisms.

Biology of an Adventive Population of the Armored Scale Rhizaspidiotus donacis, a Biological Control Agent of Arundo donax in California.

Arundo donax (giant reed) is invasive in Mediterranean, sub-, and tropical riparian systems worldwide. The armored scale Rhizaspidiotus donacis is approved for biocontrol in North America, but an adventive population was recently discovered in southern California. We documented this population's distribution, phylogeny, phenology, potential host spillover to Phragmites spp., and potential for parasitism by a common biocontrol parasitoid of citrus scale. The adventive scale was found within a single watershed and is genetically closest to Iberian scale genotypes. Rhizaspidiotus donacis developed on Phragmites haplotypes but at much lower densities than Arundo. The adventive population is univoltine, producing crawlers from March-June. Aphytis melinus parasitoids exhibited sustained interest in R. donacis during choice and no-choice trials and oviposition resulted in a small second generation. Rhizaspidiotus donacis appears limited in distribution by its univoltinism and sessile adult females. This presents challenges for broad biocontrol implementation but allows for targeted application. The genetic differentiation between imported biocontrol samples and adventive populations presents an opportunity for exploring benefits of hybrids and/or alternative genotypes where establishment has been difficult. While unlikely to occur in situ, spillover to vulnerable endemic Phragmites or deleterious parasitoid effects on scale biocontrol agents warrants consideration when planning use of R. donacis.

Does hybrid Phragmites australis differ from native and introduced lineages in reproductive, genetic, and morphological traits?

PREMISE OF THE STUDY: Hybridization between previously isolated species or lineages can stimulate invasiveness because of increased genetic diversity and inherited traits facilitating competitive and reproductive potential. We evaluated differences in stand characteristics and sexual and vegetative reproduction among native, introduced, and hybrid Phragmites australis lineages in the southwestern United States. We also assessed the degree of hybridization among lineages and backcrossing of hybrids with parental lineages. METHODS: Growth and morphological characteristics were measured in native, introduced, and hybrid Phragmites stands to evaluate relative cover and dominance in associated plant communities. Panicles were collected from stands to evaluate germination, dormancy, and differences in seed traits. Seedlings from germination trials were genotyped to determine frequency of crossing and backcrossing among lineages. KEY RESULTS: Introduced and hybrid Phragmites stands had significantly greater stem and panicle densities than native stands and were more likely to be dominant members of their respective plant communities. Hybrid seed outputs were significantly greater, but hybrid seeds had lower germination rates than those from native and introduced lineages. We detected a novel hybridization event between native and introduced lineages, but found no strong evidence of hybrids backcrossing with parental lineages. CONCLUSIONS: Hybrid Phragmites in the Southwest exhibits reproductive, genetic, and morphological characteristics from both parental lineages that facilitate dispersal, establishment, and aggressive growth, including high reproductive output, rhizome viability, and aboveground biomass, with smaller seeds and greater genetic diversity than its progenitors. Our results show hybrids can inherit traits that confer invasiveness and provide insight for managing this species complex and other cryptic species with native and introduced variants with potential for intraspecific hybridization.

Competition from native hydrophytes reduces establishment and growth of invasive dense-flowered cordgrass (Spartina densiflora).

Experimental studies to determine the nature of ecological interactions between invasive and native species are necessary for conserving and restoring native species in impacted habitats. Theory predicts that species boundaries along environmental gradients are determined by physical factors in stressful environments and by competitive ability in benign environments, but little is known about the mechanisms by which hydrophytes exclude halophytes and the life history stage at which these mechanisms are able to operate. The ongoing invasion of the South American Spartina densiflora in European marshes is causing concern about potential impacts to native plants along the marsh salinity gradient, offering an opportunity to evaluate the mechanisms by which native hydrophytes may limit, or even prevent, the expansion of invasive halophytes. Our study compared S. densiflora seedling establishment with and without competition with Phragmites australis and Typha domingensis, two hydrophytes differing in clonal architecture. We hypothesized that seedlings of the stress tolerant S. densiflora would be out-competed by stands of P. australis and T. domingensis. Growth, survivorship, biomass patterns and foliar nutrient content were recorded in a common garden experiment to determine the effect of mature P. australis and T. domingensis on the growth and colonization of S. densiflora under fresh water conditions where invasion events are likely to occur. Mature P. australis stands prevented establishment of S. densiflora seedlings and T. domingensis reduced S. densiflora establishment by 38%. Seedlings grown with P. australis produced fewer than five short shoots and all plants died after ca. 2 yrs. Our results showed that direct competition, most likely for subterranean resources, was responsible for decreased growth rate and survivorship of S. densiflora. The presence of healthy stands of P. australis, and to some extent T. domingensis, along river channels and in brackish marshes may prevent the invasion of S. densiflora by stopping the establishment of its seedlings.

Susceptibility of native and non-native common reed to the non-native mealy plum aphid (Homoptera: Aphididae) in North America.

An aggressive, non-native haplotype (distinct genetic lineage within a species) of Phragmites australis is invading brackish and freshwater systems in the eastern United States, potentially displacing native haplotypes. We studied the differential susceptibility of native and non-native populations collected from sites throughout North America to the non-native aphid, Hyalopterus pruni. In a greenhouse study, we found significantly higher aphid populations on native haplotypes than on the non-native haplotype 2 mo after infestation. Aphid feeding caused chlorosis and death of native stems, and in some cases, killed whole native genets. The non-native plants remained relatively undamaged. In a field study, non-native plants had significantly lower aphid densities than native plants or remained aphid free. There was an interactive effect in which aphid populations increased on the native plants over the 1-mo study period but remained low on non-native plants over the same period. The susceptibility of native North American populations of P. australis to non-native aphid infestation may indirectly affect the ability of these native plants to compete with non-native plant populations, ultimately contributing to the decline of native haplotypes.