Interactive effects of elevated CO<Subscript>2</Subscript>, N deposition and climate change on plant litter quality in a California annual grassland

Although global changes can alter ecosystem nutrient dynamics indirectly as a result of their effects on plant litter quality, the interactive effects of global changes on plant litter remain largely unexplored in natural communities. We investigated the effects of elevated CO₂, N deposition, warming and increased precipitation on the composition of organic compounds in plant litter in a fully-factorial experiment conducted in a California annual grassland. While lignin increased within functional groups under elevated CO₂, this effect was attenuated by warming in grasses and by water additions in forbs. CO₂-induced increases in lignin within functional groups also were counteracted by an increase in the relative biomass of forbs, which contained less lignin than grasses. Consequently, there was no net change in the overall lignin content of senesced tissue at the plot level under elevated CO₂. Nitrate additions increased N in both grass and forb litter, although this effect was attenuated by water additions. Relative to changes in N within functional groups, changes in functional group dominance had a minor effect on overall litter N at the plot level. Nitrate additions had the strongest effect on decomposition, increasing lignin losses from Avena litter and interacting with water additions to increase decomposition of litter of other grasses. Increases in lignin that resulted from elevated CO₂ had no effect on decomposition but elevated CO₂ increased N losses from Avena litter. Overall, the interactions among elements of global change were as important as single-factor effects in influencing plant litter chemistry. However, with the exception of variation in N, litter quality had little influence on decomposition over the short term.     

The Effects of Exotic Grasses on Litter Decomposition in a Hawaiian Woodland: The Importance of Indirect Effects

Exotic grasses and grass-fueled fires have altered plant species composition in the seasonal submontane woodlands of Hawaii Volcanoes National Park. These changes have altered both structural and functional aspects of the plant community, which could, in turn, have consequences for litter decomposition and nitrogen (N) dynamics. In grass-invaded unburned woodland, grass removal plots within the woodland, and woodland converted to grassland by fire, we compared whole-system fluxes and the contributions of individual species to annual aboveground fine litterfall and litterfall N, and litter mass and net N loss. We assessed the direct contribution of grass biomass to decomposition and N dynamics, and we determined how grasses affected decomposition processes indirectly via effects on native species and alteration of the litter layer microenvironment. Grasses contributed 35% of the total annual aboveground fine litterfall in the invaded woodland. However, total litterfall mass and N were not different between the invaded woodland and the grass removal treatment because of compensation by the native tree Metrosideros polymorpha, which increased litter production by 37% ± 5% when grasses were removed. The 0.3 g N m^–2/y^–1 contained in this production increase was equal to the N contained in grass litter. Litter production and litterfall N was lowest in the grassland due to the loss of native litter inputs. Decomposition of litterfall on an area basis was highest in the grass-invaded woodland. We attributed this effect to increased inherent decomposability of native litter in the presence of grasses because (a) the microenvironment of the three vegetation treatments had little effect on decomposition of common litter types and (b) M. polymorpha litter produced in the invaded woodland decomposed faster than that produced in the grass removal plots due to higher lignin concentrations in the latter than in the former. Area-weighted decomposition was lowest in the grassland due to the absence of native litter inputs. Across all treatments, most litter types immobilized N throughout the incubation, and litter net N loss on an area basis was not different among treatments. Our results support the idea that the effects of a plant species or growth form on decomposition cannot be determined in isolation from the rest of the community or from the direct effects of litter quality and quantity alone. In this dry woodland, exotic grasses significantly altered decomposition processes through indirect effects on the quantity and quality of litter produced by native species.     

Ecosystem engineering varies spatially: a test of the vegetation modification paradigm for prairie dogs

Colonial, burrowing herbivores can be engineers of grassland and shrubland ecosystems worldwide. Spatial variation in landscapes suggests caution when extrapolating single‐place studies of single species, but lack of data and the need to generalize often leads to ‘model system’ thinking and application of results beyond appropriate statistical inference. Generalizations about the engineering effects of prairie dogs (Cynomys sp.) developed largely from intensive study at a single complex of black‐tailed prairie dogs C. ludovicianus in northern mixed prairie, but have been extrapolated to other ecoregions and prairie dog species in North America, and other colonial, burrowing herbivores. We tested the paradigm that prairie dogs decrease vegetation volume and the cover of grasses and tall shrubs, and increase bare ground and forb cover. We sampled vegetation on and off 279 colonies at 13 complexes of 3 prairie dog species widely distributed across 5 ecoregions in North America. The paradigm was generally supported at 7 black‐tailed prairie dog complexes in northern mixed prairie, where vegetation volume, grass cover, and tall shrub cover were lower, and bare ground and forb cover were higher, on colonies than at paired off‐colony sites. Outside the northern mixed prairie, all 3 prairie dog species consistently reduced vegetation volume, but their effects on cover of plant functional groups varied with prairie dog species and the grazing tolerance of dominant perennial grasses. White‐tailed prairie dogs C. leucurus in sagebrush steppe did not reduce shrub cover, whereas black‐tailed prairie dogs suppressed shrub cover at all complexes with tall shrubs in the surrounding habitat matrix. Black‐tailed prairie dogs in shortgrass steppe and Gunnison's prairie dogs C. gunnisoni in Colorado Plateau grassland both had relatively minor effects on grass cover, which may reflect the dominance of grazing‐tolerant shortgrasses at both complexes. Variation in modification of vegetation structure may be understood in terms of the responses of different dominant perennial grasses to intense defoliation and differences in foraging behavior among prairie dog species. Spatial variation in the engineering role of prairie dogs suggests spatial variation in their keystone role, and spatial variation in the roles of other ecosystem engineers. Thus, ecosystem engineering can have a spatial component not evident from single‐place studies.     

Mycorrhizal suppression alters plant productivity and forb establishment in a grass-dominated prairie restoration

A fundamental goal of restoration is the re-establishment of plant diversity representative of native vegetation. However, many prairie restorations or Conservation Reserve Program sites have been seeded with warm-season grasses, leading to grass-dominated, low-diversity restorations not representative of native grasslands. These dominant grasses are strongly mycotrophic, while many subordinate forb species appear to be less dependent on mycorrhizal symbiosis. Therefore, manipulating arbuscular mycorrhizal fungi (AMF) may be useful in promoting establishment and growth of forb species in grass-dominated prairie restorations. To assess the potential role of mycorrhizae in affecting the productivity and community composition of restored tallgrass prairie, we conducted a 4-year field experiment on an 8-year-old grassland restoration at the Konza Prairie in northeastern Kansas, USA. At the initiation of our study, seeds of 12 forb species varying in degree of mycorrhizal dependence were added to established grass-dominated plots. Replicate plots were treated bi-weekly with a soil drench of fungicide (Topsin-M^®) over four growing seasons and compared to non-treated control plots to assess the role of AMF in affecting plant species composition, productivity, leaf tissue quality, and diversity in restored tallgrass prairie. Topsin applications successfully reduced mycorrhizal colonization of grass roots to approximately 60–80% relative to roots in control plots. Four years of mycorrhizal suppression reduced productivity of the dominant grasses and increased plant species richness and diversity. These results highlight the importance of mycorrhizae as mediators of plant productivity and community dynamics in restored tallgrass prairie and indicate that temporarily suppressing AMF decreases productivity of the dominant C4 grasses and allows for establishment of seeded forb species.     

Trait differences between grass species along a climatic gradient in South and North America

Question: Are trait differences between grasses along a gradient related to climatic variables and/or photosynthetic pathway? Location: Temperate grassland areas of South and North America. Methods: In a common garden experiment, we cultivated C₃ and C₄ grasses from grasslands under different climatic conditions, and we measured a set of 12 plant traits related to size and resource capture and utilization. We described (1) interspecific plant trait differences along a climatic gradient defined by the precipitation and temperature at the location where each species is dominant and (2) the association between those plant trait differences and the photosynthetic pathway of the species. Results: Trait differences between grasses were related to the precipitation at the area where each species is dominant, and to the photosynthetic pathway of the species. Leaf length, leaf width, plant height, leaf area per tiller, specific leaf area, leaf δ¹³C ratio, and nitrogen resorption efficiency increased while leaf dry matter content and nitrogen concentration in senesced leaves decreased as precipitation increased. A proportion of these changes along the gradient was related to the photosynthetic pathway because dominant grass species in cold areas with low precipitation are mainly C₃ and those from warm and wet areas are C₄. Conclusions: A previous worldwide analysis showed that traits of graminoid species measured in situ changed slightly along climatic gradients (< 10% variance explained). In contrast, under a common environment we observed that (1) grass traits changed strongly along a climatic gradient (30‐85% variance explained) and, (2) a proportion of those changes were related to the association between photosynthetic pathway of the species and precipitation.     

Consequences of shrub expansion in mesic grassland: Resource alterations and graminoid responses

Abstract. In the mesic grasslands of the central United States, the shrub Cornus drummondii has undergone widespread expansion in the absence of recurrent fire. We quantified alterations in light, water and N caused by C. drummondii expansion in tall‐grass prairie and assessed the hypothesis that these alterations are consistent with models of resource enrichment by woody plants. Responses in graminoid species, particularly the dominant C₄ grass Andropogon gerardii, were concurrently evaluated. We also removed established shrub islands to quantify their legacy effect on resource availability and assess the capability of this grassland to recover in sites formerly dominated by woody plants. The primary effect of shrub expansion on resource availability was an 87% reduction in light available to the herbaceous understorey. This reduced C uptake and N use efficiency in A. gerardii and lowered graminoid cover and ANPP at the grass‐shrub ecotone relative to undisturbed grassland. Shrub removal created a pulse in light and N availability, eliciting high C gain in A. gerardii in the first year after removal. By year two, light and N availability within shrub removal areas returned to levels typical of grassland, as had graminoid cover and ANPP were similar to those in open grassland. Recovery within central areas of shrub removal sites lagged behind that at the former grass‐shrub ecotone. These results indicate that the apparent alternative stable state of C. drummondii dominance in tall‐grass prairie is biotically maintained and driven by reductions in light, rather than resource enrichment. Within areas of shrub removal, the legacy effect of C. drummondii dominance is manifest primarily through the loss of rhizomes of the dominant grasses, rather than any long‐term changes in resource availability. C. drummondii removal facilitates grassland recovery, but the effort required to initiate this transition is a significant cost of woody plant expansion in mesic grasslands. Prevention of woody plant expansion in remnant tall‐grass prairies is, therefore, a preferred management option.     

Desert grassland dynamics estimated from carbon isotopes in grass phytoliths and soil organic matter

Abstract. We document the potential for using carbon isotopes in both soil organic matter (SOM) and grass phytoliths in soil to increase the temporal and taxonomic resolutions of long term vegetation dynamics. Carbon isotope values from both SOM and phytoliths are expected to describe both the age of material through ¹⁴C dating, and the photosynthetic pathway of the source plant material through ratios of ¹²C/¹³C. Taxonomic resolution is increased because the phytoliths examined are specific to grasses, whereas the SOM reflects the contribution of all the vegetation. Temporal resolution is increased because phytoliths are less mobile in the soil profile than SOM, and can therefore provide older dates from the same soil depth. Our results, from a desert grassland site in southwestern North America, largely confirm these expectations, and show that C₄ species have dominated the grass composition for the last 8000 yr, C₃ non‐grass vegetation increased about 100–350 yrBP, and no significant C₃ grass or non‐grass vegetation existed between 350–2000 yr BP.     

气候条件对典型草原牧草生长发育影响浅析

采用兴安盟扎赉特旗2004年～2011年气候生态环境监测站监测的典型草原牧草生长发育及地上生物量等基本资料。典型草原主要牧草种类有贝加尔针茅、野古草、羊草等禾草,伴生中旱生杂草、灌木及半灌木组成,草丛一般高30～50 cm。本文详细分析了当地气候条件对典型草原牧草生长发育及产量的影响。结果表明,典型草原牧草的高度、盖度及地上生物量受温度和降水量影响最明显,其中降水量的影响超过温度的影响。降水量多的年份地上生物量明显增加,而干旱年份地上生物量减少,其最大值一般出现在7月份,并滞后于降水量最大值。     

Dominant Grasses Suppress Local Diversity in Restored Tallgrass Prairie

Warm‐season (C₄) grasses commonly dominate tallgrass prairie restorations, often at the expense of subordinate grasses and forbs that contribute most to diversity in this ecosystem. To assess whether the cover and abundance of dominant grass species constrain plant diversity, we removed 0, 50, or 100% of tillers of two dominant species (Andropogon gerardii or Panicum virgatum) in a 7‐year‐old prairie restoration. Removing 100% of the most abundant species, A. gerardii, significantly increased light availability, forb productivity, forb cover, species richness, species evenness, and species diversity. Removal of a less abundant but very common species, P. virgatum, did not significantly affect resource availability or the local plant community. We observed no effect of removal treatments on critical belowground resources, including inorganic soil N or soil moisture. Species richness was inversely correlated with total grass productivity and percent grass cover and positively correlated with light availability at the soil surface. These relationships suggest that differential species richness among removal treatments resulted from treatment induced differences in aboveground resources rather than the belowground resources. Selective removal of the dominant species A. gerardii provided an opportunity for seeded forb species to become established leading to an increase in species richness and diversity. Therefore, management practices that target reductions in cover or biomass of the dominant species may enhance diversity in established and grass‐dominated mesic grassland restorations.     

Carbon and water relations of juvenile Quercus species in tall‐grass prairie

Abstract. In ecosystems where environments are extreme, such as deserts, adult plant species may facilitate the establishment and growth of seedlings and juveniles. Because high temperatures and evaporative demand characterize tall‐grass prairies of the central United States (relative to forests), we predicted that the grassland‐forest ecotone, by minimizing temperature extremes and moderating water stress, may function to facilitate the expansion of Quercus species into undisturbed tall‐grass prairie. We assessed the carbon and water relations of juvenile Quercus macrocarpa and Q. muhlenbergii, the dominant tree species in gallery forests of northeast Kansas, in ecotone and prairie sites. To evaluate the potentially competitive effects of neighboring herbaceous biomass on these oaks, juveniles (< 0.5 m tall) of both species also were subjected to either: (1) removal of surrounding above‐ground herbaceous biomass, or (2) control (prairie community intact) treatments. Herbaceous biomass removal had no significant effect on gas exchange or water relations in these oak species in either the prairie or the ecotone environment. Although the ecotone did alleviate some environmental extremes, photosynthetic rates and stomatal conductance were ca. 20 % higher (p < 0.05) in both oaks in prairie sites vs. the ecotone. Moreover, although leaf temperatures on average were higher in oaks in the prairie, high leaf temperatures in the ecotone had a greater negative effect on photosynthesis. These data suggest that the grassland‐forest ecotone did not facilitate the growth of Quercus juveniles expanding into this grassland. Moreover, the carbon and water relations of juvenile oaks in the prairie appeared to be unaffected by the presence of the dominant C₄ grasses.     

Belowground impacts of perennial grass cultivation for sustainable biofuel feedstock production in the tropics

Perennial grasses can sequester soil organic carbon (SOC) in sustainably managed biofuel systems, directly mitigating atmospheric CO₂ concentrations while simultaneously generating biomass for renewable energy. The objective of this study was to quantify SOC accumulation and identify the primary drivers of belowground C dynamics in a zero‐tillage production system of tropical perennial C4 grasses grown for biofuel feedstock in Hawaii. Specifically, the quantity, quality, and fate of soil C inputs were determined for eight grass accessions – four varieties each of napier grass and guinea grass. Carbon fluxes (soil CO₂ efflux, aboveground net primary productivity, litterfall, total belowground carbon flux, root decay constant), C pools (SOC pool and root biomass), and C quality (root chemistry, C and nitrogen concentrations, and ratios) were measured through three harvest cycles following conversion of a fallow field to cultivated perennial grasses. A wide range of SOC accumulation occurred, with both significant species and accession effects. Aboveground biomass yield was greater, and root lignin concentration was lower for napier grass than guinea grass. Structural equation modeling revealed that root lignin concentration was the most important driver of SOC pool: varieties with low root lignin concentration, which was significantly related to rapid root decomposition, accumulated the greatest amount of SOC. Roots with low lignin concentration decomposed rapidly, but the residue and associated microbial biomass/by‐products accumulated as SOC. In general, napier grass was better suited for promoting soil C sequestration in this system. Further, high‐yielding varieties with low root lignin concentration provided the greatest climate change mitigation potential in a ratoon system. Understanding the factors affecting SOC accumulation and the net greenhouse gas trade‐offs within a biofuel production system will aid in crop selection to meet multiple goals toward environmental and economic sustainability.     

Grazing-induced changes in plant composition affect litter quality and nutrient cycling in flooding Pampa grasslands

Changes in plant community composition induced by vertebrate grazers have been found to either accelerate or slow C and nutrient cycling in soil. This variation may reflect the differential effects of grazing-promoted (G+) plant species on overall litter quality and decomposition processes. Further, site conditions associated with prior grazing history are expected to influence litter decay and nutrient turnover. We studied how grazing-induced changes in plant life forms and species identity modified the quality of litter inputs to soil, decomposition rate and nutrient release in a flooding Pampa grassland, Argentina. Litter from G+ forbs and grasses (two species each) and grazing-reduced (G−) grasses (two species) was incubated in long-term grazed and ungrazed sites. G+ species, overall, showed higher rates of decomposition and N and P release from litter. However, this pattern was primarily driven by the low-growing, high litter-quality forbs included among G+ species. Forbs decomposed and released nutrients faster than either G+ or G− grasses. While no consistent differences between G+ and G− grasses were observed, patterns of grass litter decay and nutrient release corresponded with interspecific differences in phenology and photosynthetic pathway. Litter decomposition, N release and soil N availability were higher in the grazed site, irrespective of species litter type. Our results contradict the notion that grazing, by reducing more palatable species and promoting less palatable ones, should decrease nutrient cycling from litter. Plant tissue quality and palatability may not unequivocally link patterns of grazing resistance and litter decomposability within a community, especially where grazing causes major shifts in life form composition. Thus, plant functional groups defined by species’ “responses” to grazing may only partially overlap with functional groups based on species “effects” on C and nutrient cycling.     

Prairie plant guilds: a multivariate analysis of prairie species based on ecological and morphological traits

An ecomorphological analysis of the tallgrass prairie of central North America divided representative species of the native grassland flora into eight guilds or groups of species with similar life-form, phenology, and ecology. The guilds, segregated by multivariate analysis, are: (1) warm-season graminoids with Kranz anatomy and the Hatch-Slack photosynthetic pathway (C4 grasses); (2) cool-season graminoids without Kranz anatomy, but with the common Calvin or C3 photosynthetic pathway (C3 grasses and sedges); (3) annuals and biennial forbs; (4) ephemeral spring forbs; (5) spring forbs; (6) summer/fall forbs; (7) legumes; and (8) woody shrubs. The study was based on 158 plant species indigenous to three upland prairie sites in northeastern Kansas. Each species was scored for 32 traits which fall into five broad categories: plant habit, leaf characteristics, stem structures, root structures, and reproductive traits, including phenology. A multivariate, detrended correspondence analysis sorted the 158 species into the eight principal groups or guilds. These groups were further supported by a cluster analysis and discriminant function analysis of the same data set. The discriminant function analysis determined that 94.3% of the species were correctly classified in their respective guilds, and that the guilds were statistically different. Results indicate that guild analysis offers a basis for detailed classification of grassland vegetation that is more ecologically focused than species composition, as the myriad of species (about 1,000 prairie species on the central plains of North America) vary in presence, cover, and importance with their individualistic distribution.Abbreviations C3= C3 photosynthesis - C4= C4 photosynthesis - LSD= least significant difference     

Maintenance of leaf N controls the photosynthetic CO2 response of grassland species exposed to 9 years of free‐air CO2 enrichment

Determining underlying physiological patterns governing plant productivity and diversity in grasslands are critical to evaluate species responses to future environmental conditions of elevated CO₂ and nitrogen (N) deposition. In a 9‐year experiment, N was added to monocultures of seven C₃ grassland species exposed to elevated atmospheric CO₂ (560 μmol CO₂ mol^?1) to evaluate how N addition affects CO₂ responsiveness in species of contrasting functional groups. Functional groups differed in their responses to elevated CO₂ and N treatments. Forb species exhibited strong down‐regulation of leaf N_mass concentrations (?26%) and photosynthetic capacity (?28%) in response to elevated CO₂, especially at high N supply, whereas C₃ grasses did not. Hence, achieved photosynthetic performance was markedly enhanced for C₃ grasses (+68%) in elevated CO₂, but not significantly for forbs. Differences in access to soil resources between forbs and grasses may distinguish their responses to elevated CO₂ and N addition. Forbs had lesser root biomass, a lower distribution of biomass to roots, and lower specific root length than grasses. Maintenance of leaf N, possibly through increased root foraging in this nutrient‐poor grassland, was necessary to sustain stimulation of photosynthesis under long‐term elevated CO₂. Dilution of leaf N and associated photosynthetic down‐regulation in forbs under elevated [CO₂], relative to the C₃ grasses, illustrates the potential for shifts in species composition and diversity in grassland ecosystems that have significant forb and grass components.     

Nitrogen deposition and plant species interact to influence soil carbon stabilization

Anthropogenic nitrogen (N) deposition effects on soil organic carbon (C) decomposition remain controversial, while the role of plant species composition in mediating effects of N deposition on soil organic C decomposition and long‐term soil C sequestration is virtually unknown. Here we provide evidence from a 5‐year grassland field experiment in Minnesota that under elevated atmospheric CO₂ concentration (560 ppm), plant species determine whether N deposition inhibits the decomposition of soil organic matter via inter‐specific variation in root lignin concentration. Plant species producing lignin‐rich litter increased stabilization of soil C older than 5 years, but only in combination with elevated N inputs (4 g m^?2 year^?1). Our results suggest that N deposition will increase soil C sequestration in those ecosystems where vegetation composition and/or elevated atmospheric CO₂ cause high litter lignin inputs to soils.     

Plant community and tissue chemistry responses to fertilizer and litter nutrient manipulations in a temperate grassland

Human-mediated nutrient amendments have widespread effects on plant communities. One of the major consequences has been the loss of species diversity under increased nutrient inputs. The loss of species can be functional group dependent with certain functional groups being more prone to decline than others. We present results from the sixth year of a long-term fertilization and litter manipulation study in an old-field grassland. We measured plant tissue chemistry (C:N ratio) to understand the role of plant physiological responses in the increase or decline of functional groups under nutrient manipulations. Fertilized plots had significantly more total aboveground biomass and live biomass than unfertilized plots, which was largely due to greater productivity by exotic C₃ grasses. We found that both fertilization and litter treatments affected plant species richness. Species richness was lower on plots that were fertilized or had litter intact; species losses were primarily from forbs and non-Poaceae graminoids. C₃ grasses and forbs had lower C:N ratios under fertilization with forbs having marginally greater %N responses to fertilization than grasses. Tissue chemistry in the C₃ grasses also varied depending on tissue type with reproductive tillers having higher C:N ratios than vegetative tillers. Although forbs had greater tissue chemistry responses to fertilization, they did not have a similar positive response in productivity and the number of forb species is decreasing on our experimental plots. Overall, differential nutrient uptake and use among functional groups influenced biomass production and species interactions, favoring exotic C₃ grasses and leading to their dominance. These data suggest functional groups may differ in their responses to anthropogenic nutrient amendments, ultimately influencing plant community composition.     

Plant nitrogen concentration, use efficiency, and contents in a tallgrass prairie ecosystem under experimental warming

Plant nitrogen (N) relationship has the potential to regulate plant and ecosystem responses strongly to global warming but has not been carefully examined under warmed environments. This study was conducted to examine responses of plant N relationship (i.e. leaf N concentration, N use efficiency, and plant N content in this study) to a 4‐year experimental warming in a tallgrass prairie in the central Great Plains in USA. We measured mass‐based N and carbon (C) concentrations of stem, green, and senescent leaves, and calculated N resorption efficiency, N use efficiency, plant N content, and C : N ratios of five dominant species (two C₄ grasses, one C₃ grass, and two C₃ forbs). The results showed that warming decreased N concentration of both green and senescent leaves, and N resorption efficiency for all species. N use efficiencies and C : N ratios were accordingly higher under warming than control. Total plant N content increased under warming because of warming‐induced increases in biomass production that are larger than the warming‐induced decreases in tissue N concentration. The increases in N contents in both green and senescent plant tissues suggest that warming enhanced both plant N uptake and return through litterfall in the tallgrass ecosystem. Our results also suggest that the increased N use efficiency in C₄ grasses is a primary mechanism leading to increased biomass production under warming in the grassland ecosystem.     

Litter Decomposition in a California Annual Grassland: Interactions Between Photodegradation and Litter Layer Thickness

In annual grasslands that experience a mediterranean-type climate, the synchrony between plant senescence and peak solar radiation over summer results in high litter sun exposure. We examined the decomposition of both shaded and sun-exposed litter over summer and inferred the effects of photodegradation from changes in mass loss and litter chemistry. The carry-over effects of summer litter exposure on wet season decomposition were also assessed, and the attenuation of photodegradation with litter layer thickness was used to estimate the proportion of grass litter lignin susceptible to photodegradation under different treatments of a factorial global change experiment. Over summer, mass loss from grass and forb litter exposed to ambient sunlight ranged from 8% to 10%, whereas lignin decreased in grass litter by approximately 20%. After one year of decomposition, mass losses from grass leaves exposed to sunlight over summer were more than double the mass losses from summer-shaded leaves. When shade litter layer thickness was varied, mass losses over summer for all treatments were also approximately 8%; however, lignin decreased significantly only in the low shade treatments (0–64 g m⁻² of shade litter). Aboveground production of annual grasses nearly quadrupled in response to the combined effects of N addition, elevated atmospheric CO₂, increased precipitation and warming. The estimated proportion of grass litter lignin experiencing full photodegradation ranged from 100% under ambient conditions to 31–62% in plots receiving the combined global change treatments. These results reveal an important role of sun exposure over summer in accelerating litter decomposition in these grasslands and provide evidence that future changes in the quantity of litter deposition may modulate the influence of photodegradation integrated across the litter layer.     

The role of hemiparasitic plants: influencing tallgrass prairie quality,diversity, and structure

Wood betony, Orobanchaceae (Pedicularis canadensis) and bastard toadflax, Santalaceae (Comandra umbellata) are two root‐hemiparasitic plant species found in tallgrass prairie communities. Natural resource managers are interested in utilizing these species as “pseudograzers” in grasslands to reduce competitively dominant grasses and thereby increase ecological diversity and quality in prairie restorations and urban plantings. We performed an observational field study at 5 tallgrass prairie sites to investigate the association of hemiparasite abundance with metrics of phylogenetic and ecological diversity, as well as floristic quality. Although no reduction in C₄ grasses was detected, there was a significant association between hemiparasite abundance and increased floristic quality at all 5 sites. Hemiparasite abundance and species richness were positively correlated at one restoration site. In a greenhouse mesocosm experiment, we investigated response to parasitism by P. canadensis in 6 species representing different plant functional groups of the tallgrass prairie. The annual legume partridge pea, Fabaceae (Chamaecrista fasciculata) had the greatest significant dry biomass reduction among 6 host species, but the C₄ grass big bluestem, Poaceae (Andropogon gerardii) had significantly greater aboveground biomass when grown with the hemiparasite. Overall, host species biomass as a total community was significantly reduced in mesocosms, consistent with other investigations that demonstrate influence on community structure by hemiparasitic plant species. Although hemiparasites were not acting as pseudograzers, they have the potential to influence community structure in grassland restorations and remnants.     

Interspecific variation in plant responses to mycorrhizal colonization in tallgrass prairie

Symbiotic associations between plants and arbuscular mycorrhizal fungi are ubiquitous and ecologically important in many grasslands. Differences in species responses to mycorrhizal colonization can have a significant influence on plant community structure. The growth responses of 36 species of warm- and cool-season tallgrass prairie grasses and 59 tallgrass prairie forbs to arbuscular mycorrhizal (AM) fungal colonization were assessed in greenhouse studies to examine the extent of interspecific variation in host-plant benefit from the symbiosis and patterns of mycorrhizal dependence among host plant life history (e.g., annual, perennial) and taxonomic (e.g., grass, forb, legume, nonlegume) groups and phenological guilds. There was a strong and significant relationship between phenology of prairie grasses and mycorrhizal responsiveness, however this relationship was less apparent in forbs. Perennial warm-season C(4) grasses and forbs generally benefited significantly from the mycorrhizal symbiosis, whereas biomass production of the cool-season C(3) grasses was not affected. The root systems of the cool-season grasses were also less highly colonized by the AM fungi, as compared to the warm-season grasses or forbs. Unlike the native perennials, annuals were generally not responsive to mycorrhizal colonization and were lower in percentage root colonization than the perennial species. Plant growth responsiveness and AM root colonization were positively correlated for the nonleguminous species, with this relationship being strongest for the cool-season grasses. In contrast, root colonization of prairie legumes showed a significant, but negative, relationship to mycorrhizal growth responsiveness.