Unpalatable Plants Protect Neighbors from Grazing and Increase Plant Community Diversity

Tasty plants can be protected from herbivores by unpalatable neighbors. We used experimental exclosures, removal of unpalatable species, and transplants of palatable and unpalatable species in subalpine meadows of the Caucasus Mountains of Georgia to study the effects of two unpalatable species on plant communities. We found that Cirsium obalatum and Veratrum lobelianum, two large native perennial herbs that invade after heavy grazing, had strong facilitative effects on communities through their indirect effects on livestock herbivores. These unpalatable invaders had different effects on community composition when livestock were present than when livestock were excluded. Furthermore, removing Cirsium and Veratrum where herbivory was permitted decreased the richness of associated communities, but inside a livestock exclosure removal of these species increased community richness. Transplanted palatable species (Anthoxanthum odoratum and Phleum alpinum) grew larger inside the exclosure, and in the exclosure Cirsium and Veratrum had no effect on their growth. However, outside of the exclosure, Cirsium and Veratrum had strong positive effects on the growth of A. odoratum and P. alpinum. Excluding livestock decreased the growth of Luzula pseudosudetica, another unpalatable species, and Cirsium and Veratrum had no effect on L. pseudosudetica outside the exclosure. In contrast, inside exclosures Cirsium and Veratrum had competitive effects on L. pseudosudetica. Our results indicate that Cirsium and Veratrum, which are in some ways undesirable rangeland weeds, may also play an important role in maintaining species and functional diversity of overgrazed plant communities in the Caucasus

Range-Expanding Populations of a Globally Introduced Weed Experience Negative Plant-Soil Feedbacks

BACKGROUND: Biological invasions are fundamentally biogeographic processes that occur over large spatial scales. Interactions with soil microbes can have strong impacts on plant invasions, but how these interactions vary among areas where introduced species are highly invasive vs. naturalized is still unknown. In this study, we examined biogeographic variation in plant-soil microbe interactions of a globally invasive weed, Centaurea solstitialis (yellow starthistle). We addressed the following questions (1) Is Centaurea released from natural enemy pressure from soil microbes in introduced regions? and (2) Is variation in plant-soil feedbacks associated with variation in Centaurea's invasive success? METHODOLOGY/PRINCIPAL FINDINGS: We conducted greenhouse experiments using soils and seeds collected from native Eurasian populations and introduced populations spanning North and South America where Centaurea is highly invasive and noninvasive. Soil microbes had pervasive negative effects in all regions, although the magnitude of their effect varied among regions. These patterns were not unequivocally congruent with the enemy release hypothesis. Surprisingly, we also found that Centaurea generated strong negative feedbacks in regions where it is the most invasive, while it generated neutral plant-soil feedbacks where it is noninvasive. CONCLUSIONS/SIGNIFICANCE: Recent studies have found reduced below-ground enemy attack and more positive plant-soil feedbacks in range-expanding plant populations, but we found increased negative effects of soil microbes in range-expanding Centaurea populations. While such negative feedbacks may limit the long-term persistence of invasive plants, such feedbacks may also contribute to the success of invasions, either by having disproportionately negative impacts on competing species, or by yielding relatively better growth in uncolonized areas that would encourage lateral spread. Enemy release from soil-borne pathogens is not sufficient to explain the success of this weed in such different regions. The biogeographic variation in soil-microbe effects indicates that different mechanisms may operate on this species in different regions, thus establishing geographic mosaics of species interactions that contribute to variation in invasion success

Data from: The importance of factors controlling species abundance and distribution varies in native and non-native species.

How variation in factors controlling species abundance and distribution between native and non-native ranges compares to that within ranges remains poorly understood. We used a globally distributed ruderal, Centaurea solstitialis (Centaurea), to explore the possibility that the importance of those factors exhibits great variation between and within ranges. To test our hypothesis, we established seed addition experiments with soil disturbance (turnover and control) and biocide (fungicides, insecticide, and control) treatments in two regions within native (the Caucasus and south-western Turkey) and non-native (the western United States – US – and central Argentina) distributions. Also, we estimated the rate of vegetation recovery after disturbance (resilience) and related it to Centaurea density in experimental plots. Disturbance strongly increased Centaurea density in all regions. Density was similar between the native Caucasus and non-native Argentina and much greater in those regions than in the native Turkey and non-native US in biocide-free plots. Fungicides had positive effects on density in the US and negative ones in the Caucasus and Argentina, resulting in no differences between those three regions and greater density in the US than Turkey. Insecticide applications promoted Centaurea density in Turkey and Argentina, but inter-regional comparisons of density in treated plots were comparable to those in biocide-free plots. Overall, plants were smaller and less fecund in Turkey than the other regions, except the US. The greatest fungal attack was documented in Turkey, and herbivory was stronger there and in Argentina than in the Caucasus and US. The resilience of the local community explained a large proportion of variation in Centaurea density. These results support our hypothesis, and reveal that the speed at which competition is re-gained after disturbance may influence global variation in Centaurea abundance. Because many ruderals exhibit native and non-native distributions, our results are likely to be generalized to other systems

The importance of factors controlling species abundance and distribution varies in native and non-native ranges

How variation in factors controlling species abundance and distribution between native and non-native ranges compares to that within ranges remains poorly understood. We used a globally distributed ruderal, Centaurea solstitialis (Centaurea), to explore the possibility that the importance of those factors exhibits great variation between and within ranges. To test our hypothesis, we established seed addition experiments with soil disturbance (turnover and control) and biocide (fungicides, insecticide, and control) treatments in two regions within native (the Caucasus and south-western Turkey) and non-native (the western United States -US- and central Argentina) distributions. Also, we estimated the rate of vegetation recovery after disturbance (resilience) and related it to Centaurea density in experimental plots. Disturbance strongly increased Centaurea density in all regions. Density was similar between the native Caucasus and non-native Argentina and much greater in those regions than in the native Turkey and non-native US in biocide-free plots. Fungicides had positive effects on density in the US and negative ones in the Caucasus and Argentina, resulting in no differences between those three regions and greater density in the US than Turkey. Insecticide applications promoted Centaurea density in Turkey and Argentina, but inter-regional comparisons of density in treated plots were comparable to those in biocide-free plots. Overall, plants were smaller and less fecund in Turkey than the other regions, except the US. The greatest fungal attack was documented in Turkey, and herbivory was stronger there and in Argentina than in the Caucasus and US. The resilience of the local community explained a large proportion of variation in Centaurea density. These results support our hypothesis, and reveal that the speed at which competition is re-gained after disturbance may influence global variation in Centaurea abundance. Because many ruderals exhibit native and non-native distributions, our results are likely to be generalized to other systems.Fil: Hierro, Jose Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Ciencias de la Tierra y Ambientales de La Pampa. Universidad Nacional de La Pampa. Facultad de Ciencias Exactas y Naturales. Instituto de Ciencias de la Tierra y Ambientales de La Pampa; ArgentinaFil: Khetsuriani, Liana. Ilia State University; GeorgiaFil: Andonian, Krikor. De Anza College; Estados UnidosFil: Eren, Ozkan. Adnan Menderes University; TurquíaFil: Villarreal, Diego. Universidad Nacional de La Pampa. Facultad de Ciencias Exactas y Naturales; ArgentinaFil: Janoian, Grigor. American University of Armenia; ArmeniaFil: Reinhart, Kurt O.. United States Department of Agriculture; Estados UnidosFil: Callaway, Ragan M.. University of Montana; Estados Unido

Geographic mosaics of plant-soil microbe interactions in a global plant invasion

Aim Our aim in this study was to document the global biogeographic variation in the effects of soil microbes on the growth of Centaurea solstitialis (yellow starthistle; Asteraceae), a species that has been introduced throughout the world, but has become highly invasive only in some introduced regions. Location To assess biogeographic variation in plant-soil microbe interactions, we collected seeds and soils from native Eurasian C. solstitialis populations and introduced populations in California, Argentina and Chile. Methods To test whether escape from soil-borne natural enemies may contribute to the success of C. solstitialis, we compared the performance of plants using seeds and soils collected from each of the biogeographic regions in greenhouse inoculation/sterilization experiments. Results We found that soil microbes had pervasive negative effects on plants from all regions, but these negative effects were significantly weaker in soils from non-native ranges in Chile and California than in those from the non-native range in Argentina and the native range in Eurasia. Main conclusions The biogeographic differences in negative effects of microbes in this study conformed to the enemy-release hypothesis (ERH) overall, but the strong negative effect of soil biota in Argentina, where C. solstitialis is invasive, and weaker effects in Chile where it is not, indicated that different factors influencing invasion are likely to occur in large scale biogeographic mosaics of interaction strengths.Fil: Andonian, Krikor. University of California; Estados UnidosFil: Hierro, Jose Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Ciencias de la Tierra y Ambientales de La Pampa. Universidad Nacional de La Pampa. Facultad de Ciencias Exactas y Naturales. Instituto de Ciencias de la Tierra y Ambientales de La Pampa; ArgentinaFil: Khetsuriani, Liana. Institute of Botany of the Georgian Academy of Sciences; Estados UnidosFil: Becerra, Pablo I.. Universidad Católica de Chile; ChileFil: Janoyan, Grigor. American University of Armenia; ArmeniaFil: Villareal, Diego. Universidad Nacional de La Pampa. Facultad de Ciencias Exactas y Naturales; ArgentinaFil: Cavieres, Lohengrin A.. Universidad de Concepción; ChileFil: Fox, Laurel R.. University of California; Estados UnidosFil: Callaway, Ragan M.. University of Montana; Estados Unido

Dispersal Pathways and Genetic Differentiation among Worldwide Populations of the Invasive Weed Centaurea solstitialis L. (Asteraceae)

The natural history of introduced species is often unclear due to a lack of historical records. Even when historical information is readily available, important factors of the invasions such as genetic bottlenecks, hybridization, historical relationships among populations and adaptive changes are left unknown. In this study, we developed a set of nuclear, simple sequence repeat markers and used these to characterize the genetic diversity and population structure among native (Eurasian) and non-native (North and South American) populations of Centaurea solstitialis L., (yellow starthistle). We used these data to test hypotheses about the invasion pathways of the species that were based on historical and geographical records, and we make inferences about historical relationships among populations and demographic processes following invasion. We confirm that the center of diversity and the native range of the species is likely the eastern Mediterranean region in the vicinity of Turkey. From this region, the species likely proceeded to colonize other parts of Europe and Asia via a slow, stepwise range expansion. Spanish populations were the primary source of seed to invade South America via human-mediated events, as was evident from historical records, but populations from the eastern Mediterranean region were also important. North American populations were largely derived from South America, but had secondary contributors. We suggest that the introduction history of non-native populations from disparate parts of the native range have allowed not just one, but multiple opportunities first in South America then again in North America for the creation of novel genotypes via intraspecific hybridization. We propose that multiple intraspecific hybridization events may have created especially potent conditions for the selection of a noxious invader, and may explain differences in genetic patterns among North and South America populations, inferred differences in demographic processes, as well as morphological differences previously reported from common garden experiments

Data from Hierro et al. 2016 Ecography

This is an Excel file, where data from our study is organized in different sheets

Germination responses of an invasive species in native and non-native ranges

Studying germination in the native and non-native range of a species can provide unique insights into processes of range expansion and adaptation; however, traits related to germination have rarely been compared between native and nonnative populations. In a series of common garden experiments, we explored whether differences in the seasonality of precipitation, specifically, summer drought vs summer rain, and the amount and variation of annual and seasonal precipitation affect the germination responses of populations of an annual ruderal plant, Centaurea solstitialis, from its native range and from two non-native regions with different climates. We found that seeds from all native populations, irrespective of the precipitation seasonality of the region in which they occurred, and non-native populations from regions with dry summers displayed similarly high germination proportions and rates. In contrast, genotypes from the non-native region with predominantly summer rain exhibited much lower germination fractions and rates. Also, percent germination was strongly correlated with variation in precipitation in winter, the season that follows germination for C. solstitialis. Specifically, germination was lower for native and non-native populations experiencing greater variation in winter precipitation. This correlation, however, was greatly influenced by the non-native region with summer rain, which also exhibited the greatest variation in winter precipitation among studied regions. These results suggest that rather than general climatic patterns, the degree of risk experienced at early developmental stages could exert an important control over the germination strategy of C. solstitialis populations in both native and non-native ranges. Also, these findings reveal a largely unique germination response in C. solstitialis genotypes growing in the non-native region with summer rain and high variation in winter precipitation. Our work raises the possibility that rapid adaptive changes in germination strategies may contribute to the success of globally distributed invaders.Centaurea solstitialis, from its native range and from two non-native regions with different climates. We found that seeds from all native populations, irrespective of the precipitation seasonality of the region in which they occurred, and non-native populations from regions with dry summers displayed similarly high germination proportions and rates. In contrast, genotypes from the non-native region with predominantly summer rain exhibited much lower germination fractions and rates. Also, percent germination was strongly correlated with variation in precipitation in winter, the season that follows germination for C. solstitialis. Specifically, germination was lower for native and non-native populations experiencing greater variation in winter precipitation. This correlation, however, was greatly influenced by the non-native region with summer rain, which also exhibited the greatest variation in winter precipitation among studied regions. These results suggest that rather than general climatic patterns, the degree of risk experienced at early developmental stages could exert an important control over the germination strategy of C. solstitialis populations in both native and non-native ranges. Also, these findings reveal a largely unique germination response in C. solstitialis genotypes growing in the non-native region with summer rain and high variation in winter precipitation. Our work raises the possibility that rapid adaptive changes in germination strategies may contribute to the success of globally distributed invaders.Fil: Hierro, Jose Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Ciencias de la Tierra y Ambientales de La Pampa. Universidad Nacional de La Pampa. Facultad de Ciencias Exactas y Naturales. Instituto de Ciencias de la Tierra y Ambientales de La Pampa; Argentina. University of Montana; Estados UnidosFil: Eren, Ozkan. Adnan Menderes Universitesi; TurquíaFil: Khetsuriani, Liana. Academy of Sciences; GeorgiaFil: Diaconu, Alecu. Institute of Biological Research; RumaniaFil: Török, Katalin. Institute of Ecology and Botany of HAS; HungríaFil: Montesinos, Daniel. Centro de Investigaciones sobre Desertificación; EspañaFil: Andonian, Krikor. University of California; Estados UnidosFil: Kikodze, David. Academy of Sciences; GeorgiaFil: Janoian, Levan. University of Armenia; ArmeniaFil: Villarreal, Diego. Universidad Nacional de La Pampa. Facultad de Ciencias Exactas y Naturales; ArgentinaFil: Estanga Mollica, María E.. Universidad Nacional de La Pampa. Facultad de Ciencias Exactas y Naturales; ArgentinaFil: Callaway, Ragan M.. University of Montana; Estados Unido