Phragmites australis as a model organism for studying plant invasions

© 2016, Springer International Publishing Switzerland. The cosmopolitan reed grass Phragmites australis (Poaceae) is an intensively studied species globally with a substantial focus in the last two decades on its invasive populations. Here we argue that P. australis meets the criteria to serve as a model organism for studying plant invasions. First, as a dominant species in globally important wetland habitats, it has generated significant pre-existing research, demonstrating a high potential for funding. Second, this plant is easy to grow and use in experiments. Third, it grows abundantly in a wide range of ecological systems and plant communities, allowing a broad range of research questions to be addressed. We formalize the designation of P. australis as a model organism for plant invasions in order to encourage and standardize collaborative research on multiple spatial scales that will help to integrate studies on the ecology and evolution of P. australis invasive populations, their response to global environmental change, and implications for biological security. Such an integrative framework can serve as guidance for studying invasive plant species at the population level and global spatial scale

Geographic variation in apparent competition between native and invasive \u3ci\u3ePhragmites australis\u3c/i\u3e

Apparent competition, the negative interaction between species mediated by shared natural enemies, is thought to play an important role in shaping the structure and dynamics of natural communities. However, its importance in driving species invasions, and whether the strength of this indirect interaction varies across the latitudinal range of the invasion, has not been fully explored. We performed replicated field experiments at four sites spanning 900 km along the Atlantic Coast of the United States to assess the presence and strength of apparent competition between sympatric native and invasive lineages of Phragmites australis. Four herbivore guilds were considered: stem‐feeders, leaf‐miners, leaf‐chewers and aphids. We also tested the hypothesis that the strength of this interaction declines with increasing latitude. Within each site, native and invasive plants of P. australis were cross‐transplanted between co‐occurring native and invasive patches in the same marsh habitat and herbivore damage was evaluated at the end of the growing season. Apparent competition was evident for both lineages and involved all but the leaf‐chewer guild. For native plants, total aphids per plant was 296% higher and the incidence of stem‐feeding and leaf‐mining herbivores was 34% and 221% higher, respectively, when transplanted into invasive than native patches. These data suggest that invasive P. australis has a negative effect on native P. australis via apparent competition. Averaged among herbivore types, the indirect effects of the invasive lineage on the native lineage was 57% higher than the reverse situation, suggesting that apparent competition was asymmetric. We also found that the strength of apparent competition acting against the native lineage was comparable to the benefits to the invasive lineage from enemy release (i.e., proportionately lower mean herbivory of the invasive relative to the native taxa). Finally, we found the first evidence that the strength of apparent competition acting against the native lineage (from stem‐feeders only) decreased with increasing latitude. These results suggest that not only could apparent competition be of tantamount importance to enemy release in enhancing the establishment and spread of invasive taxa, but also that these indirect and direct herbivore effects could vary over the invasion range

Biogeography of a Plant Invasion: Plant–Herbivore Interactions

Theory predicts that native plant species should exhibit latitudinal gradients in the strength of their interactions with herbivores. We hypothesize that if an invasive plant species exhibits a different latitudinal gradient in response to herbivores (e.g., a nonparallel gradient), it can create large-scale heterogeneities in community resistance/susceptibility to the invasive species. We conducted a study of latitudinal variation in the strength of herbivory and defenses of native genotypes of Phragmites australis in North America (NA) and Europe (EU) and European invasive genotypes in NA. Within NA, we tested whether (1) invasive genotypes are better defended and suffer less herbivory than co-occurring native genotypes, (2) herbivory and defenses of native P. australis decreases with increasing latitude; and (3) invasive genotypes exhibit either no latitudinal gradient, or a nonparallel latitudinal gradient in herbivory and defenses compared to native genotypes. For the European genotypes, we tested two additional hypotheses: (4) defenses, nutritional condition, and herbivory would differ between the native (EU) and invasive ranges (NA) and (5) latitudinal gradients in defenses and herbivory would be similar between ranges. Within NA, chewing damage, internal stem-feeding incidence, and aphid abundance were 650%, 300%, and 70% lower, respectively, on invasive than native P. australis genotypes. Genotypes in NA also differed in nutritional condition (percent N, C:N ratio), but there was little support for invasive genotypes being better defended than native genotypes. For the European genotypes, herbivory was significantly lower in the invaded than native range, supporting the enemy-release hypothesis. Defense levels (leaf toughness and total phenolics) and tissue percent C and percent N were higher in the invaded than native range for European genotypes. Overall, latitudinal gradients in P. australis nutritional condition, defenses, and herbivory were common. Interestingly, chewing damage and stem-feeder incidence decreased with latitude for native P. australis genotypes in NA and EU, but no latitudinal gradients in response to herbivores were evident for invasive genotypes in NA. Nonparallel latitudinal gradients in herbivory between invasive and native P. australis suggest that the community may be more susceptible to invasion at lower than at higher latitudes. Our study points to the need for invasion biology to include a biogeographic perspective

Biological control of invasive Phragmites australis will be detrimental to native P. australis

Phragmites australis biological control is intended to address a major invasion in North America and is likely to change the ecology of vast areas of coastal and inland wetlands. However, the real risks to the native North American genotypes of P. australis (as indicated by recent research summarized above) may not have been fully considered, particularly the extirpation of native populations or the eventual extinction of the native North American lineage altogether. The concerns we raise need to be considered in the process of developing and approving the release of biological control agents and the entire approval process would benefit from greater transparency and wider input from Phragmites researchers globally

Response to Blossey and Casagrande: ecological and evolutionary processes make host specificity at the subspecies level exceedingly unlikely

We agree with Blossey and Casagrande (2016) that absolute host-specificity is a necessity for successful biological control of invasive plants without unintended consequences for native species. However, inclusion of non-target native species in the diet of a biological control agent is a relatively common phenomenon with native congeners of the target plant species at greatest risk (Pemberton 2000)

Biogeography of a plant invasion: drivers of latitudinal variation in enemy release

© 2016 John Wiley & Sons Ltd Aim: The relationship between herbivory and latitude may differ between native and invasive plant taxa, which can generate biogeographical heterogeneity in the strength of enemy release. Our aim was to compare latitudinal gradients in herbivory between native and invasive plants and investigate whether gradients are driven by local adaptation or phenotypic plasticity. Location: North America. Methods: Using sympatric native and invasive lineages of the wetland grass Phragmites australis and the specialist gall-fly Lipara rufitarsis, we conducted a field survey to examine whether the relationship between herbivory (the proportion of stems galled) and latitude was parallel between lineages. In a subsequent common garden experiment, we assessed whether latitudinal gradients in herbivory were genetically based or driven by phenotypic plasticity. Results: In the field, L. rufitarsis herbivory on the native P. australis lineage increased from 27% of stems galled in southern populations (36.5°) to 37% in northern populations (43.6°), whereas there was no relationship for the invasive lineage. Similar relationships were evident in the common garden experiment, indicating a genetic basis to latitudinal variation in herbivory. Moreover, the invasive lineage suffered five times less herbivory than the native lineage on average, supporting the enemy release hypothesis. However, a genetic basis to this pattern was absent in the common garden experiment, suggesting that local environmental conditions were responsible for the enemy release observed in nature. Specifically, stem height, diameter and density during the L. rufitarsis oviposition period appeared to be important drivers of herbivory. Main conclusions: Non-parallel gradients in herbivory may help explain the equivocal results of other studies that examine enemy release and biotic resistance at local scales, and can be an important mechanism promoting biogeographical variation in invasion success. We suggest that these latitudinal patterns in herbivory and other species interactions are likely to be a common phenomenon across a range of invaded systems

DNA assays for genetic discrimination of three Phragmites australis subspecies in the United States

Premise: To genetically discriminate subspecies of the common reed (Phragmites australis), we developed real-time quantitative (qPCR) assays for identifying P. australis subsp. americanus, P. australis subsp. australis, and P. australis subsp. berlandieri. Methods and Results: Utilizing study-generated chloroplast DNA sequences, we developed three novel qPCR assays. Assays were verified on individuals of each subspecies and against two non-target species, Arundo donax and Phalaris arundinacea. One assay amplifies only P. australis subsp. americanus, one amplifies P. australis subsp. australis and/or P. australis subsp. berlandieri, and one amplifies P. australis subsp. americanus and/or P. australis subsp. australis. This protocol enhances currently available rapid identification methods by providing genetic discrimination of all three subspecies. Conclusions: The newly developed assays were validated using P. australis samples from across the United States. Application of these assays outside of this geographic range should be preceded by additional testing

Intraspecific and biogeographical variation in foliar fungal communities and pathogen damage of native and invasive Phragmites australis

AimRecent research has highlighted that the relationship between species interactions and latitude can differ between native and invasive plant taxa, generating biogeographical heterogeneity in community resistance to plant invasions. In the first study with foliar pathogens, we tested whether co‐occurring native and invasive lineages of common reed (Phragmites australis) exhibit non‐parallel latitudinal gradients in foliar fungal communities, pathogen susceptibility and damage, and whether these biogeographical patterns can influence the success of invasion.LocationNorth America.Time period2015–2017.Major taxa studiedPerennial grass P. australis.MethodsWe surveyed 35 P. australis field populations, spanning 17° latitude and comprising four phylogeographical lineages, including one endemic to North America and one invasive from Europe. For each population, we quantified the percentage of leaf pathogen damage and cultured fungi from diseased leaves, which we identified using molecular tools. To assess whether latitudinal gradients in pathogen damage had a genetic basis, we inoculated plants from 73 populations with four putative pathogens in a complementary common garden experiment and measured P. australis susceptibility (i.e., diseased leaf area).ResultsWe isolated 84 foliar fungal taxa. Phragmites australis lineage influenced fungal community composition but not diversity. Despite the invasive European P. australis lineage being the least susceptible to three of the four pathogens tested in the common garden experiment, pathogen damage in the field was similar between native and invasive lineages, providing no evidence that release from foliar pathogens contributes to the success of invasion. Genetically based latitudinal gradients in pathogen susceptibility observed in the common garden were isolate specific and obscured by local environmental conditions in the field, where pathogen damage was threefold higher for northern compared with southern populations, regardless of lineage.Main conclusionsOur results highlight that host plant lineage and genetically based biogeographical gradients strongly influence foliar fungal communities and pathogen susceptibility, but do not translate to patterns of pathogen damage observed in the field.Peer Reviewedhttps://deepblue.lib.umich.edu/bitstream/2027.42/155999/1/geb13097.pdfhttps://deepblue.lib.umich.edu/bitstream/2027.42/155999/2/geb13097_am.pd