Native plant diversity resists invasion at both low and high resource levels

Human modification of the environment is causing both loss of species and changes in resource availability. While studies have examined how species loss at the local level can influence invasion resistance, interactions between species loss and other components of environmental change remain poorly studied. In particular, the manner in which native diversity interacts with resource availability to influence invasion resistance is not well understood. We created experimental plant assemblages that varied in native species (1-16 species) and/or functional richness (defined by rooting morphology and phenology; one to five functional groups). We crossed these diversity treatments with resource (water) addition to determine their interactive effects on invasion resistance to spotted knapweed (Centaurea maculosa), a potent exotic invader in the intermountain West of the United States. We also determined how native diversity and resource addition influenced plant-available soil nitrogen, soil moisture, and light. Assemblages with lower species and functional diversity were more heavily invaded than assemblages with greater species and functional diversity. In uninvaded assemblages, experimental addition of water increased soil moisture and plant-available nitrogen and decreased light availability. The availability of these resources generally declined with increasing native plant diversity. Although water addition increased susceptibility to invasion, it did not fundamentally change the negative relationship between diversity and invasibility. Thus, native diversity provided strong invasion resistance even under high resource availability. These results suggest that the effects of local diversity can remain robust despite enhanced resource levels that are predicted under scenarios of global change.     

Geographic variation in resource dominance-discovery in Brazilian ant communities

A predictive framework for the ecology of species invasions requires that we learn what limits successful invaders in their native range. The red imported fire ant (Solenopsis invicta) is invasive in the United States, Puerto Rico, Australia, New Zealand, and China. Solenopsis invicta appears to be a superior competitor in its introduced range, where it can cause the local extirpation of native species, but little is known about its competitive ability in its native range in South America. Here we examine the competitive ability of S. invicta for food resources in three widely separated Brazilian ant communities. Each of these communities contains 20-40 ant species, 8-10 of which were common and frequently interacted with S. invicta. S. invicta at all three sites was attacked by several species-specific phorid parasitoids, and at one site, two other species were attacked by their own specialized parasitoids. We examined interactions in these local communities for evidence that trade-offs among ant species between resource dominance and resource discovery, and between resource dominance and parasitoid vulnerability facilitate local coexistence. The trade-off between resource dominance and resource discovery was strong and significant only at Santa Genebra, where parasitoids had no effect on the outcome of confrontations at resources. At Bonito, parasitoids significantly reduced the ability of S. invicta, which was the top-ranked behavioral dominant, from defending and usurping food resources from subordinate species. In the Pantanal, S. invicta ranked behind three other ant species in a linear hierarchy of behavioral dominance, and lost the majority of its interactions with a fourth more subordinate species, Paratrechina fulva, another invasive species. Parasitoids of S. invicta were uncommon in the Pantanal, and did not affect its low position in the hierarchy relative to the other two sites. Parasitoids, however, did affect the ability of Linepithema angulatum, the top-ranked behavioral dominant in this community, from defending and usurping resources from behavioral subordinates. These results indicate that both interspecific competition and trait-mediated indirect effects of phorid parasitoids affect the ecological success of the red imported fire ant in its native range, but that the relative importance of these factors varies geographically.     

Differing Effects of Cattle Grazing on Native and Alien Plants

Abstract:   Habitat managers use cattle grazing to reduce alien plant cover and promote native species in California grasslands and elsewhere in the western United States. We tested the effectiveness of grazing as a restoration method by examining the effects of herbivory on native and alien plants. At Carrizo Plain National Monument, California, we surveyed native and alien species cover in adjacent grazed and ungrazed areas. We also established experimental plots in which plants were clipped or mulch (dead biomass) was removed. In addition, we clipped plants grown in pots and plants in the field that grew with and without competitors. Native species were negatively affected by clipping in 1999, 2000, and 2001, whereas alien species were unaffected. In the experimental field plots, the European annual forb Erodium cicutarium compensated in growth and reproduction following simulated herbivory. In contrast, growth and reproduction of the native perennial bunchgrass Poa secunda were reduced 1 year after clipping. In pots, E. cicutarium overcompensated and grasses undercompensated. In the field, European grasses were unaffected by the removal of competitors. It is unclear by what mechanism E. cicutarium was able to compensate, but the ability may be related to its basal rosette growth form and indeterminately growing inflorescences. The native California grassland community assembled in the absence of grazing herds, whereas invasive European species have been exposed to grazing for centuries. It may be that these invaders have adaptations that better enable them to recover from grazing. In the grassland we studied, the strategy of livestock grazing for restoration is counterproductive. It harms native species and promotes alien plant growth.     

Nutrient deficiency in dune slack pioneer vegetation: a review

A review of results of fertilization experiments in wet dune slacks is presented. In most cases the above-ground biomass appeared to be limited by nitrogen availability. Primary phosphorus limitation was assessed only once in a dune slack where sod cutting had been applied very recently. In most other case studies phosphorus limits biomass production after nitrogen deficiency was lifted. Potassium availability is of minor importance for biomass production in this type of ecosystem. Singular nitrogen additions led to increased dominance ofCarex andJuncus species as well as perennial grasses, such asAgrostis stolonifera andCalamagrostis epigejos. A combined addition of nitrogen and phosphorus led to total dominance of grasses, while the characteristic basiphilous pioneer species (including mosses) decreased or even disappeared. Certain mechanisms are considered which may maintain nutrient availability in slacks with basiphilous pioneer vegetation at a low level, despite of the accumulation of nutrients in the developing organic soil layer. Some implications for management and further research are discussed.     

Long-term population dynamics of seeded plants in invaded grasslands

In recent decades, dozens of studies have involved attempts to introduce native and desirable nonnative plant species into grasslands dominated by invasive weeds. The newly introduced plants have proved capable of establishing, but because they are rarely monitored for more than four years, it is unknown if they have a high likelihood of persisting and suppressing invaders for the long-term. Beyond invaded grasslands, this lack of long-term monitoring is a general problem plaguing efforts to reintroduce a range of taxa into a range of ecosystems. We introduced species from seed and then periodically measured plant abundances for nine years at one site and 15 years at a second site. To our knowledge, our 15-year data are the longest to date from a seeding experiment in invaded, never-cultivated grassland. At one site, three seeded grasses maintained high densities for three or more years, but then all or nearly all individuals died. At the second site, one grass performed similarly, but two other grasses proliferated and at least one greatly suppressed the dominant invader (Centaurea maculosa). In one study, our point estimate suggests that the seeded grass Thinopyrum intermedium reduced C. maculosa biomass by 93% 15 years after seeding. In some cases, data from three and fewer years after seeding falsely suggested that seeded species were capable of persisting within the invaded grassland. In other cases, data from as late as nine years after seeding falsely suggested seeded populations would not become large enough to suppress the invader. These results show that seeded species sometimes persist and suppress invaders for long periods, but short-term data cannot predict if, when, or where this will occur. Because short-term data are not predictive of long-term seeded species performances, additional long-term data are needed to identify effective practices, traits, and species for revegetating invaded grasslands.     

An invader differentially affects leaf physiology of two natives across a gradient in diversity

Little is known about how exotics influence the ecophysiology of co-occurring native plants or how invader impact on plant physiology may be mediated by community diversity or resource levels. We measured the effect of the widespread invasive forb spotted knapweed (Centaurea maculosa) on leaf traits (leaf dry matter content, specific leaf area, leaf nitrogen percentage, leaf C:N ratios, and delta13C as a proxy for water use efficiency) of two co-occurring native perennial grassland species, Monarda fistulosa (bee balm) and Koeleria macrantha (Junegrass). The impact of spotted knapweed was assessed across plots that varied in functional diversity and that either experienced ambient rainfall or received supplemental water. Impact was determined by comparing leaf traits between identical knapweed-invaded and noninvaded assemblages. Virtually all M. fistulosa leaf traits were affected by spotted knapweed. Knapweed impact, however, did not scale with its abundance; the impact of knapweed on M. fistulosa was similar across heavily invaded low-diversity assemblages and lightly invaded high-diversity assemblages. In uninvaded assemblages, M. fistulosa delta13C, leaf nitrogen, and C:N ratios were unaffected by native functional group richness, whereas leaf dry matter content significantly increased and specific leaf area significantly decreased across the diversity gradient. The effects of spotted knapweed on K. macrantha were weak; instead native functional group richness strongly affected K. macrantha leaf C:N ratio, delta13C, and specific leaf area, but not leaf dry matter content. Leaf traits for both species changed in response to spotted knapweed or functional richness, and in a manner that may promote slower biomass accumulation and efficient conservation of resources. Taken together, our results show that an invader can alter native plant physiology, but that these effects are not a simple function of how many invaders exist in the community.     

Conservation Development Practices,Extent, and Land‐Use Effects in the United States

Abstract: Conservation development projects combine real‐estate development with conservation of land and other natural resources. Thousands of such projects have been conducted in the United States and other countries through the involvement of private developers, landowners, land trusts, and government agencies. Previous research has demonstrated the potential value of conservation development for conserving species, ecological functions, and other resource values on private lands, especially when traditional sources of conservation funding are not available. Nevertheless, the aggregate extent and effects of conservation development were previously unknown. To address this gap, we estimated the extent and trends of conservation development in the United States and characterized its key attributes to understand its aggregate contribution to land‐conservation and growth‐management objectives. We interviewed representatives from land trusts, planning agencies, and development companies, searched the Internet for conservation development projects and programs, and compiled existing databases of conservation development projects. We collected data on 3884 projects encompassing 1.38 million ha. About 43% of the projects targeted the conservation of specific plant or animal species or ecological communities of conservation concern; 84% targeted the protection of native ecosystems representative of the project area; and 42% provided buffers to existing protected areas. The percentage of protected land in conservation development projects ranged from <40% to >99%, and the effects of these projects on natural resources differed widely. We estimate that conservation development projects have protected roughly 4 million ha of land in the United States and account for about 25% of private‐land conservation activity nationwide.     

Competition between Native Populus deltoides and Invasive Tamarix ramosissima and the Implications for Reestablishing Flooding Disturbance

Abstract: Changes in historical disturbance regimes have been shown to facilitate non-native plant invasions, but reinstatement of disturbance can be successful only if native colonizers are able to outcompete colonizing invasives. Reintroduction of flooding in the southwestern United States is being promoted as a means of reestablishing Populus deltoides subsp. wislizenii , but flooding can also promote establishment of an introduced, invasive species, Tamarix ramosissima . We investigated competition between Populus and Tamarix at the seedling stage to aid in characterizing the process by which Tamarix may invade and to determine the potential ability of Populus to establish itself with competitive pressure from Tamarix . We planted seedlings of Tamarix and Populus in five ratios at three densities for a total of 15 treatments. The growth response of each species was measured in terms of height, above-ground biomass, and tissue concentrations of nitrogen and phosphorous. These measurements across treatments were modeled as three-dimensional response surfaces. For both species, Populus density was more important than Tamarix density for determining growth response. Both species were negatively affected by increasing numbers of Populus seedlings. Due to the larger size of the native Populus , we predict that its superior competitive ability can lead to its dominance when conditions allow native establishment. Our results suggest that even in the presence of an invader that positively responds to disturbance, reestablishment of historical flooding regimes and post-flood hydrology can restore this ecosystem by promoting its dominant plant species.     

Postfire Management on Forested Public Lands of the Western United States

Abstract:  Forest ecosystems in the western United States evolved over many millennia in response to disturbances such as wildfires. Land use and management practices have altered these ecosystems, however, including fire regimes in some areas. Forest ecosystems are especially vulnerable to postfire management practices because such practices may influence forest dynamics and aquatic systems for decades to centuries. Thus, there is an increasing need to evaluate the effect of postfire treatments from the perspective of ecosystem recovery. We examined, via the published literature and our collective experience, the ecological effects of some common postfire treatments. Based on this examination, promising postfire restoration measures include retention of large trees, rehabilitation of firelines and roads, and, in some cases, planting of native species. The following practices are generally inconsistent with efforts to restore ecosystem functions after fire: seeding exotic species, livestock grazing, placement of physical structures in and near stream channels, ground-based postfire logging, removal of large trees, and road construction. Practices that adversely affect soil integrity, persistence or recovery of native species, riparian functions, or water quality generally impede ecological recovery after fire. Although research provides a basis for evaluating the efficacy of postfire treatments, there is a continuing need to increase our understanding of the effects of such treatments within the context of societal and ecological goals for forested public lands of the western United States.     

Forest succession suppressed by an introduced plant-fungal symbiosis

Microbial symbionts can affect plant nutrition, defensive chemistry, and biodiversity. Here we test the hypothesis that symbionts alter the speed and direction of plant succession in communities that are shifting from grasslands to forests. A widespread C3 grass introduced to the United States, Lolium arundinaceum (tall fescue), hosts a fungal endophyte that is toxic to herbivores. In replicated experimental grasslands, the presence of the endophyte in tall fescue reduced tree abundance and size, altered tree composition, and slowed plant species turnover. In addition, consumption of tree seedlings by voles (Microtus spp.) was 65% higher in plots with the endophyte at the one grassland site where these data were collected. Despite its negligible contribution to community biomass, a microbial symbiont suppressed tree establishment, posing an important constraint on the natural transition from grasslands to forests.     

Native fish diversity alters the effects of an invasive species on food webs

Aquatic communities have been altered by invasive species, with impacts on native biodiversity and ecosystem function. At the same time, native biodiversity may mitigate the effects of an invader. Common carp (Cyprinus carpio) is a ubiquitous, invasive fish species that strongly influences community and ecosystem processes. We used common carp to test whether the potential effects of an invasive species are altered across a range of species diversity in native communities. In mesocosms, treatments of zero, one, three, and six native fish species were used to represent the nested subset patterns observed in fish communities of lakes in Illinois, USA. The effect of the invader was tested across fish richness treatments by adding common carp to the native community and substituting native biomass with common carp. Native species and intraspecific effects reduced invader growth. The invader reduced native fish growth; however, the negative effect was minimized with increasing native richness. The zooplankton grazer community was modified by a top-down effect from the invader that increased the amount of phytoplankton. Neither the invader nor richness treatments influenced total phosphorus or community metabolism. Overall, the invader reduced resources for native species; and the effect scaled with how the invader was incorporated into the community. Higher native diversity mitigated the impact of the invader, confirming the need to consider biodiversity when predicting the impacts of invasive species.     

Forest Restoration and Fire: Principles in the Context of Place

Abstract:  There is broad consensus that active management through thinning and fire is urgently needed in many forests of the western United States. This consensus stems from physically based models of fire behavior and substantial empirical evidence. But the types of thinning and fire and where they are applied are the subjects of much debate. We propose that low thinning is the most appropriate type of thinning practice. Treating surface fuels, reducing ladder fuels, and opening overstory canopies generally produce fire-safe forest conditions, but large, fire-resistant trees are also important components of fire-safe forests. The context of place is critical in assigning priority for the limited resources that will be available for restoration treatments. Historical low-severity fire regimes, because of their current high hazards and dominance by fire-resistant species, are the highest priority for treatment. Mixed-severity fire regimes are of intermediate priority, and high-severity fire regimes are of lowest priority. Classification systems based on potential vegetation will help identify these fire regimes at a local scale.     

Weed-biocontrol insects reduce native-plant recruitment through second-order apparent competition

Small-mammal seed predation is an important force structuring native-plant communities that may also influence exotic-plant invasions. In the intermountain West, deer mice (Peromyscus maniculatus) are prominent predators of native-plant seeds, but they avoid consuming seeds of certain widespread invasives like spotted knapweed (Centaurea maculosa). These mice also consume the biological-control insects Urophora spp. introduced to control C. maculosa, and this food resource substantially increases deer mouse populations. Thus, mice may play an important role in the invasion and management of C. maculosa through food-web interactions. We examined deer mouse seed predation and its effects on seedling emergence and establishment of a dominant native grass, Pseudoroegneria spicata, and forb, Balsamorhiza sagittata, in C. maculosa-invaded grasslands that were treated with herbicide to suppress C. maculosa or left untreated as controls. Deer mice readily took seeds of both native plants but removed 2-20 times more of the larger B. sagittata seeds than the smaller P. spicata seeds. Seed predation reduced emergence and establishment of both species but had greater impacts on B. sagittata. The intensity of seed predation corresponded with annual and seasonal changes in deer mouse abundance, suggesting that abundance largely determined mouse impacts on native-plant seeds. Accordingly, herbicide treatments that reduced mouse abundance by suppressing C. maculosa and its associated biocontrol food subsidies to mice also reduced seed predation and decreased the impact of deer mice on B. sagittata establishment. These results provide evidence that Urophora biocontrol agents may exacerbate the negative effects of C. maculosa on native plants through a form of second-order apparent competition-a biocontrol indirect effect that has not been previously documented. Herbicide suppressed C. maculosa and Urophora, reducing mouse populations and moderating seed predation on native plants, but the herbicide's direct negative effects on native forb seedlings overwhelmed the indirect positive effect of reducing deer mouse seed predation. By manipulating this four-level food chain, we illustrate that host-specific biological control agents may impact nontarget plant species through food-web interactions, and herbicides may influence management outcomes through indirect trophic interactions in addition to their direct effects on plants.     

Rainfall and soils modify plant community response to grazing in Serengeti National Park

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.     

Forecasting the Expansion of Zebra Mussels in the United States

Abstract:  Because zebra mussels spread rapidly throughout the eastern United States in the late 1980s and early 1990s, their spread to the western United States has been expected. Overland dispersal into inland lakes and reservoirs, however, has occurred at a much slower rate than earlier spread via connected, navigable waterways. We forecasted the potential western spread of zebra mussels by predicting the overland movement of recreational boaters with a production-constrained gravity model. We also predicted the potential abundance of zebra mussels in two western reservoirs by comparing their water chemistry characteristics with those of water bodies with known abundances of zebra mussels. Most boats coming from waters infested with zebra mussels were taken to areas that already had zebra mussels, but a small proportion of such boats did travel west of the 100th meridian. If zebra mussels do establish in western U.S. water bodies, we predict that population densities could achieve similar levels to those in the Midwestern United States, where zebra mussels have caused considerable economic and ecological impacts. Our analyses suggest that the dispersal of zebra mussels to the western United States is an event of low probability but potentially high impact on native biodiversity and human infrastructure. Combining these results with economic analyses could help determine appropriate investment levels in prevention and control strategies .     

Provenance, life span, and phylogeny do not affect grass species' responses to nitrogen and phosphorus

Successful conservation management requires an understanding of how species respond to intervention. Native and exotic species may respond differently to management interventions due to differences arising directly from their origin (i.e., provenance) or indirectly due to biased representations of different life history types (e.g., annual vs. perennial life span) or phylogenetic lineages among provenance (i.e., native or exotic origin) groups. Thus, selection of a successful management regime requires knowledge of the life history and provenance-bias in the local flora and an understanding of the interplay between species characteristics across existing environmental gradients in the landscape. Here we tested whether provenance, phylogeny, and life span interact to determine species distributions along natural gradients of soil chemistry (e.g., soil nitrogen and phosphorus) in 10 upland prairie sites along a 600-km latitudinal transect running from southern Vancouver Island in British Columbia, Canada, to the Willamette Valley in Oregon, USA. We found that soil nitrate, phosphorus, and pH exerted strong control over community composition. However, species distributions along environmental gradients were unrelated to provenance, life span, or phylogenetic groupings. We then used a greenhouse experiment to more precisely measure the response of common grass species to nitrogen and phosphorus supply. As with the field data, species responses to nutrient additions did not vary as a function of provenance, life span, or phylogeny. Native and exotic species differed strongly in the relationship between greenhouse-measured tolerance of low nutrients and field abundance. Native species with the greatest ability to maintain biomass production at low nutrient supply rates were most abundant in field surveys, as predicted by resource competition theory. In contrast, there was no relationship between exotic-species biomass at low nutrient levels and field abundance. The implications of these findings for management of invasive species are substantial in that they overturn a general belief that reduction of nutrient supplies favors native species. The idiosyncratic nature of species response to nutrients in this study suggests that manipulation of nutrient supplies is unlikely to alter the overall balance between native and exotic species, although it may well be useful to control specific exotic species.     

Nitrogen inputs promote the spread of an invasive marsh grass.

Excess nutrient loading and large-scale invasion by nonnatives are two of the most pervasive and damaging threats to the biotic and economic integrity of our estuaries. Individually, these are potent forces, but it is important to consider their interactive impacts as well. In this study we investigated the potential limitation of a nonnative intertidal grass, Spartina alterniflora, by nitrogen (N) in estuaries of the western United States. Nitrogen fertilization experiments were conducted in three mud-flat habitats invaded by S. alterniflora in Willapa Bay, Washington, USA, that differed in sediment N. We carried out parallel experiments in San Francisco Bay, California, USA, in three habitats invaded by hybrid Spartina (S. alterniflora x S. foliosa), in previously unvegetated mud flat, and in native S. foliosa or Salicornia virginica marshes. We found similar aboveground biomass and growth rates between habitats and estuaries, but end-of-season belowground biomass was nearly five times greater in San Francisco Bay than in Willapa Bay. In Willapa Bay, aboveground biomass was significantly correlated with sediment N content. Addition of N significantly increased aboveground biomass, stem density, and the rate of spread into uninvaded habitat (as new stems per day) in virtually all habitats in both estuaries. Belowground biomass increased in Willapa Bay only, suggesting that belowground biomass is not N limited in San Francisco Bay due to species differences, N availability, or a latitudinal difference in the response of Spartina to N additions. The relative impact of added N was greater in Willapa Bay, the estuary with lower N inputs from the watershed, than in San Francisco Bay, a highly eutrophic estuary. Nitrogen fertilization also altered the competitive interaction between hybrid Spartina and Salicornia virginica in San Francisco Bay by increasing the density and biomass of the invader and decreasing the density of the native. There was no significant effect of N on the native, Spartina foliosa. Our results indicate that excess N loading to these ecosystems enhances the vulnerability of intertidal habitats to rapid invasion by nonnative Spartina sp.     

The Quagga Mussel Crisis at Lake Mead National Recreation Area,Nevada (U.S.A.)

Abstract: Parks are cornerstones of conservation; and non‐native invasive species drive extensive changes to biological diversity in parks. Knowing this, national park staff at Lake Mead National Recreation Area in the southwestern United States had a program in place for early detection of the non‐native, invasive quagga mussel (Dreissena rostriformis bugensis). Upon finding the mussel in January 2007, managers moved quickly to access funding and the best available science to implement a response. Managers considered four options—doing nothing, closing the park, restricting movement on the lakes, and educating and enforcing park visitors—and decided to focus on education and enforcing existing laws. Nonetheless, quagga spread throughout the park and soon began to appear throughout the western United States. I examined why efforts to control the expansion failed and determined the general lessons to be learned from this case. Concentrating human visitation on the lakes through land‐use zoning opened a pathway for invasion, reduced management options, and led to the rapid spread of quagga. To reconcile competing mandates to protect nature and provide recreation, zoning in parks has become a common practice worldwide. It reduces stress on some areas of a park by restricting and thus concentrating human activity in particular areas. Concentrating the human activity in one area does three things: cements pathways that repeatedly import and export vectors of non‐native invasive species; creates the disturbed area necessary to enable non‐native invasive species to gain a foothold; and, establishes a source of invasions that, without appropriate controls, can quickly spread to a park's wilderness areas.     

Grazers, producer stoichiometry, and the light : nutrient hypothesis revisited

The stoichiometric light:nutrient hypothesis (LNH) links the relative supplies of key resources with the nutrient content of tissues of producers. This resource-driven variation in producer stoichiometry, in turn, can mediate the efficiency of grazing. Typically, discussions of the LNH attribute this resource-stoichiometry link to bottom-up effects of light and phosphorus, which are mediated through producer physiology. Emphasis on bottom-up effects implies that grazers must consume food of quality solely determined by resource supply to ecosystems (i.e., they eat what they are served). Here, we expand upon this largely bottom-up interpretation with evidence from pond surveys, a mesocosm experiment, and a model. Data from shallow ponds showed the "LNH pattern" (positive correlation of an index of light : phosphorus supply with algal carbon : phosphorus content). However, algal carbon : phosphorus content also declined as zooplankton biomass increased in the ponds. The experiment and model confirmed that this latter correlation was partially caused by the various bottom-up and top-down roles of grazers: the LNH pattern emerged only in treatments with crustacean grazers, not those without them. Furthermore, model and experiment clarified that another bottom-up factor, natural covariation of nitrogen : phosphorus ratios with light : phosphorus supply (as seen in ponds), does not likely contribute to the LNH pattern. Finally, the experiment produced correlations between shifts in species composition of algae, partially driven by grazing effects of crustaceans, and algal stoichiometry. These shifts in species composition might shape stoichiometric response of producer assemblages to resource supply and grazing, but their consequences remain largely unexplored. Thus, this study accentuated the importance of grazing for the LNH; de-emphasized a potentially confounding, bottom-up factor (covarying nitrogen : phosphorus supply); and highlighted an avenue for future research for the LNH (grazer-mediated shifts in producer composition).     

Variational model of microorganism polyculture development without re-supply of mutually irreplaceable resources

Stationary stage of accumulating cultures of Pseudomonas aeruginosa dissociants is described by variational model of consumption and growth. The “generalized entropy” as a goal function is used. Model's parameters are the dissociants requirements for basic nutrients: carbon, nitrogen and phosphorus. Using this model we calculate the limitation ranges for arbitrary combinations of environmental resource factors and the community population sizes at a stationary stage of growth as functions of the resources that constrain the growth. The requirements values obtained from experimental data allowed us to predict the limiting resources and dissociant's abundances. Our estimations match experimental results. The possibility of composition control is discussed in paper.