Bitty A. Roy

RESPONSE OF PLANT PATHOGENS AND HERBIVORES TO A WARMING EXPERIMENT

Pathogens and herbivores can severely reduce host fitness, potentially leading to altered succession rates and changes in plant community composition. Thus, to predict vegetation dynamics under climate change, it is necessary to understand how plant pathogens and herbivores will respond. Pathogens and herbivores are predicted to increase under climate warming because the amount of time available for growth and reproduction will increase. To test this prediction, we used a warming experiment in which heaters were suspended over a natural montane meadow for 12 years. In the summer of 2002, we quantified damage by all the observable (aboveground) pathogens and herbivores on six of the most common plant species (Artemisia tridentata, Helianthella quinquenervis, Erigeron speciosus, Potentilla gracilis, Potentilla hippiana, and Lathyrus leucanthus). We found that plants in the earlier melting plots generally had the most damage and were attacked by a larger number of species, which is consistent with predictions. However, although the overall trend was an increase in damage with warmer temperatures and earlier snowmelt, some pathogens and herbivores performed better in cooler or later melting plots. The idiosyncratic response of each species to environmental conditions suggests that there are likely to be changes in community composition as the planet warms.

Increasing forest loss worldwide from invasive pests requires new trade regulations

Loss of forests due to non-native invasive pests (including insects, nematodes, and pathogens) is a global phenomenon with profound population, community, ecosystem, and economic impacts. We review the magnitude of pest-associated forest loss worldwide and discuss the major ecological and evolutionary causes and consequences of these invasions. After compiling and analyzing a dataset of pest invasions from 21 countries, we show that the number of forest pest invasions recorded for a given country has a significant positive relationship with trade (as indicated by gross domestic product) and is not associated with the amount of forested land within that country. We recommend revisions to existing international protocols for preventing pest entry and proliferation, including prohibiting shipments of non-essential plants and plant products unless quarantined. Because invasions often originate from taxa that are scientifically described only after their introduction, current phytosanitary regulations – which ...

CONTEXT-DEPENDENT POLLINATOR BEHAVIOR: AN EXPLANATION FOR PATTERNS OF HYBRIDIZATION AMONG THREE SPECIES OF INDIAN PAINTBRUSH

Abstract In some areas of sympatry, reproductively compatible plant species hybridize, but in other areas of sympatry, they do not and they remain reproductively isolated from one another. Explanations offered to explain patterns of hybridization that vary by population have usually focused on genetic or environmental factors. Instead, we examined whether different community contexts might change pollinator preference and constancy and thus influence the likelihood of hybridization among three Indian paintbrush species (Castilleja miniata, C. rhexifolia, and C. sulphurea). To determine whether visitation was context-dependent, we observed pollinator behavior in experimental arrays (constructed using flowering stems of the three Indian paintbrush species) in different contexts. Contexts were defined by which Castilleja species occurred in the immediate neighborhood of the arrays. Specifically, we asked, does visitation to particular species in the arrays depend on context? In general, each Castilleja species was preferred when it matched the surrounding community context, as is predicted by optimal foraging theory. More interestingly, pollinator constancy was weakened in the hybrid context (an area where the three species co-occurred with morphologically intermediate plants), which is likely to increase pollen flow among the species. Reduced pollinator constancy in hybrid zones could set up a positive feedback loop in which more flower diversity is created through hybridization, decreasing pollinator constancy, and leading to more hybridization. This self-reinforcing mechanism could lead to “hybridization hot spots” and to a patchy distribution of hybrid populations. We expect that this mechanism may be important in other animal-pollinated plant hybrid zones.

Abiotic constraints on the competitive ability of exotic and native grasses in a Pacific Northwest prairie

In prairie ecosystems, abiotic constraints on competition can structure plant communities; however, the extent to which competition between native and exotic plant species is constrained by environmental factors is still debated. The objective of our study was to use paired field and greenhouse experiments to evaluate the competitive dynamics between two native (Danthonia californica and Deschampsia cespitosa) and two exotic (Schedonorus arundinaceus and Lolium multiflorum) grass species under varying nutrient and moisture conditions in an upland prairie in the Willamette Valley, Oregon. We hypothesized the two invasive, exotic grasses would be more competitive under high-nutrient, moderate-moisture conditions, resulting in the displacement of native grasses from these environments. In the field, the experimental reduction of competition resulted in shorter, wider plants, but only the annual grass, Lolium multiflorum, produced more aboveground biomass when competition was reduced. In the greenhouse, the two exotic grasses produced more total biomass than the two native grasses. Competitive hierarchies were influenced by nutrient and/or moisture treatments for the two exotic grasses, but not for the two native grasses. L. multiflorum dominated competitive interactions with all other grasses across treatments. In general, S. arundinaceus dominated when in competition with native grasses, and D. cespitosa produced the most biomass in monoculture or under interspecific competition with the other native grass, D. californica. D. californica, D. cespitosa, and S. arundinaceus all produced more biomass in high-moisture, high-nutrient environments, and D. cespitosa, L. multiflorum, and S. arundinaceus allocated more biomass belowground in the low nutrient treatment. Taken together, these experiments suggest the competitive superiority of the exotic grasses, especially L. multiflorum, but, contrary to our hypothesis, the native grasses were not preferentially excluded from nutrient-rich, moderately wet environments.

Tests for Parasite-mediated Frequency-dependent Selection in Natural Populations of an Asexual Plant Species

Genetic variation in plant populations for resistance to pathogens and herbivores might be maintained by parasite-mediated negative frequency-dependent selection (FDS). But it is difficult to observe the time-lagged oscillations between host and parasite genotypes that should result from FDS. To evaluate the potential for FDS, we tested for local adaptation of parasites to common clones, the role of host genetic diversity in resistance to parasites, and genetic correlations among fitness, parasitism, and the frequency of host clones. We studied three populations of Arabis holboellii, a short-lived apomictic (asexual by seed) plant attacked by rust fungi and insect herbivores. To estimate clone frequency, we used polymorphic allozyme markers on 200 individuals in each population in 1990 and in 2000. We also recorded levels of parasitism and host fitness (fruit production). Only the rust fungi showed evidence for local host adaptation; they usually increased in incidence as a function of clone frequency, and they tracked temporal change in clone frequency. In further support of FDS, parasitism was lower in populations with higher genetic diversity. However, total parasitism (herbivory and disease combined) decreased as host clone frequency and fitness increased. Thus, although the highly virulent rust pathogen showed potential for driving the cycles that result from FDS, this apparently does not occur in the populations studied because the host clones were also attacked by herbivores.

Testing assumptions of the enemy release hypothesis: generalist versus specialist enemies of the grass Brachypodium sylvaticum

The enemy release hypothesis (ERH) suggests greater success of species in an invaded range due to release from natural enemies. The ERH assumes there will be more specialist enemies in the native range and that generalists will have an equal effect in both ranges. We tested these assumptions with the grass Brachypodium sylvaticum in the native range (Switzerland) and invaded range (Oregon, USA). We assessed all the kinds of damage present (caused by fungi, insects, mollusk and deer) on both leaves and seeds at 10 sites in each range and correlated damage with host fitness. Only two of the 20 fungi found on leaves were specialist pathogens, and these were more frequent in the native range. Conversely there was more insect herbivory on leaves in the invaded range. All fungi and insects found on seeds were generalists. More species of fungi were found on seeds in the native range, and a higher proportion of them were pathogenic than in the invaded range. There were more kinds of enemies in the native range, where the plants had lower fitness, in accordance with the ERH. However, contrary to assumptions of the ERH, generalists appear to be equally or more important than specialists in reducing host fitness.

Context-dependent pattern and process: the distribution and competitive dynamics of an invasive grass, Brachypodium sylvaticum

Brachypodium sylvaticum, or false brome, a recent invader of North America, has spread rapidly following establishment in Oregon in 1939 and is now colonizing at least five US states. To understand how ecological processes and environmental context interact to facilitate invasion, we combined a GIS analysis of spatial patterns of invasion with a greenhouse competition experiment under two light and two nutrient levels, and a field competition-removal experiment. Brachypodium sylvaticum was concentrated along human-use corridors and in forests, where our experiments indicate it is a better competitor than co-occurring grasses. Forested areas near trails require the greatest focus for detection and control because the species’ dual propensities to establish and thrive in these locations may make them key source areas for further invasion. Our methodology demonstrates how understanding both the patterns and processes of invasion enables the prescription of context-specific remedial actions.

Herbivores and pathogens on Alnus viridis subsp. fruticosa in Interior Alaska: effects of leaf, tree, and neighbour characteristics on damage levels

Parasite damage strongly affects dynamics of boreal forests. Damage levels may be affected by climate change, either directly or indirectly through changes in properties of host trees. We examined how herbivore and pathogen damage in Alnus viridis subsp. fruticosa (Rupr.) Nym. depend on leaf morphology and chemistry, tree size, and tree neighborhood. Damage and tree properties were measured in 2003 and 2004 on eight trees at each of 20 sites in Interior Alaska. Damage varied significantly among sites and among trees within sites, but Cartesian distances between sites were not correlated with similarity in damage levels. Compared with middle leaves, terminal leaves experienced less damage from phloem-feeding insects and pathogens, whereas leaf-roller damage was largely confined to terminal leaves. Summer drought in 2004 strongly reduced damage from phloem-feeding insects, while damage from chewing insects increased. Overall, herbivore damage was best explained by leaf morphology and chemistry, and pathogen...

The herbaceous landlord: integrating the effects of symbiont consortia within a single host

Plants are typically infected by a consortium of internal fungal associates, including endophytes in their leaves, as well as arbuscular mycorrhizal fungi (AMF) and dark septate endophytes (DSE) in their roots. It is logical that these organisms will interact with each other and the abiotic environment in addition to their host, but there has been little work to date examining the interactions of multiple symbionts within single plant hosts, or how the relationships among symbionts and their host change across environmental conditions. We examined the grass Agrostis capillaris in the context of a climate manipulation experiment in prairies in the Pacific Northwest, USA. Each plant was tested for presence of foliar endophytes in the genus Epichloë, and we measured percent root length colonized (PRLC) by AMF and DSE. We hypothesized that the symbionts in our system would be in competition for host resources, that the outcome of that competition could be driven by the benefit to the host, and that the host plants would be able to allocate carbon to the symbionts in such a way as to maximize fitness benefit within a particular environmental context. We found a correlation between DSE and AMF PRLC across climatic conditions; we also found a fitness cost to increasing DSE colonization, which was negated by presence of Epichloë endophytes. These results suggest that selective pressure on the host is likely to favor host/symbiont relationships that structure the community of symbionts in the most beneficial way possible for the host, not necessarily favoring the individual symbiont that is most beneficial to the host in isolation. These results highlight the need for a more integrative, systems approach to the study of host/symbiont consortia.

Grassland fires may favor native over introduced plants by reducing pathogen loads

Grasslands have been lost and degraded in the United States since Euro-American settlement due to agriculture, development, introduced invasive species, and changes in fire regimes. Fire is frequently used in prairie restoration to control invasion by trees and shrubs, but may have additional consequences. For example, fire might reduce damage by herbivore and pathogen enemies by eliminating litter, which harbors eggs and spores. Less obviously, fire might influence enemy loads differently for native and introduced plant hosts. We used a controlled burn in a Willamette Valley (Oregon) prairie to examine these questions. We expected that, without fire, introduced host plants should have less damage than native host plants because the introduced species are likely to have left many of their enemies behind when they were transported to their new range (the enemy release hypothesis, or ERH). If the ERH holds, then fire, which should temporarily reduce enemies on all species, should give an advantage to the natives because they should see greater total reduction in damage by enemies. Prior to the burn, we censused herbivore and pathogen attack on eight plant species (five of nonnative origin: Bromus hordaceous, Cynosuros echinatus, Galium divaricatum, Schedonorus arundinaceus (= Festuca arundinacea), and Sherardia arvensis; and three natives: Danthonia californica, Epilobium minutum, and Lomatium nudicale). The same plots were monitored for two years post-fire. Prior to the burn, native plants had more kinds of damage and more pathogen damage than introduced plants, consistent with the ERH. Fire reduced pathogen damage relative to the controls more for the native than the introduced species, but the effects on herbivory were negligible. Pathogen attack was correlated with plant reproductive fitness, whereas herbivory was not. These results suggest that fire may be useful for promoting some native plants in prairies due to its negative effects on their pathogens.