Long-term patterns of shrub expansion in a C4-dominated grassland: fire frequency and the dynamics of shrub cover and abundance

Worldwide, grassland ecosystems have experienced a major shift in growth-form dominance as woody plant species have expanded and replaced native grasses. In the C(4)-dominated grasslands of central North America, a reduction in fire frequency is the most cited cause of this shift in growth forms as fire both enhances grass productivity and constrains the establishment and expansion of native woody vegetation. Using an 18-yr plant species composition data set, we quantified patterns of change in shrub cover, frequency, and species richness associated with three distinct fire regimes. During the study period (1983-2000), shrub cover increased most dramatically in sites in which the frequency of fire was once every 4 yr (intermediate frequency; 28.6%) followed by sites in which fire occurred only once during the 18-yr period (low frequency; 23.7%). Annual fire effectively prevented the recruitment of new woody species, but even with this high fire frequency, shrub cover increased slightly (3.7%). Comparatively, shrub species richness increased by three and six, respectively, in the intermediate- and low-frequency fire sites. These data indicate that within this grassland, periods without fire are necessary for recruitment of both new individuals and additional shrub species; however, once established, shrub cover will increase regardless of fire frequency and even annual fire will not reduce shrub abundance.     

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.     

Determinants of C3 forb growth and production in a C4 dominated grassland

Forbs are the most abundant species within the vascular flora of tallgrass prairie and they make the greatest contribution to biodiversity of all growth forms. However, little is known about the factors that determine their productivity and growth rates. The objective of this study was to assess the controls of forb growth (absolute and relative) and production in tallgrass prairie from a long-term burning experiment at the Konza Prairie in NE Kansas. Over the 14-year study, forbs comprised 9% of the total biomass production on sites with a high fire frequency vs. 29% on the low fire frequency site, with gramminoids accounting for the remainder. Although interannual variations in peak biomass of the grasses was strongly correlated with environmental variables related to water availability, there were no similar relationships for forbs, suggesting that production of forbs and grasses responded to interannual variations in climate in different ways. Multivariate analysis of climatic controls on growth rates of grasses and forbs yielded similar results. Although forbs had low biomass and absolute growth per unit ground area in frequently burned prairie, their relative growth rates were highest in such sites. Thus, it appears that reduced growth rates of individual forbs per se do not limit forb success in annually burned prairie. Instead, direct negative effects of fire on forbs (increased mortality) may be responsible. Determinants of forb growth and productivity in unburned prairie remain unresolved.     

Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Restoration of C4 grasses with seasonal fires in a C3/C4 grassland invaded by Prosopis glandulosa,a fire‐resistant shrub

Questions: Can prescribed fire restore C₄ perennial grasses in grassland ecosystems that have become dominated by fire‐resistant C₃ shrubs (Prosopis glandulosa) and C₃ grasses? Do fires in different seasons alter the direction of change in grass composition? Location: Texas, USA. Methods: We quantified short‐ and long‐term (12 yr post‐fire) herbaceous functional group cover and diversity responses to replicated seasonal fire treatments: (1) repeated‐winter fires (three in 5 yr), (2) repeated‐summer fires (two in 3 yr), and (3) alternate‐season fires (two winter and one summer in 4 yr), compared with a no‐fire control. Results: Summer fires were more intense than winter fires, but all fire treatments temporarily decreased Prosopis and C₃ annual grass cover. The alternate‐season fire treatment caused a long‐term increase in C₄ mid‐grass cover and functional group diversity. The repeated‐summer fire treatment increased C₄ short‐grass cover but also caused a long‐term increase in bare ground. The repeated winter fire treatment had no long‐term effects on perennial grass cover. Mesquite post‐fire regrowth had increasingly negative impacts on herbaceous cover in all fire treatments. Conclusions: Summer fire was necessary to shift herbaceous composition toward C₄ mid‐grasses. However, the repeated‐summer fire treatment may have been too extreme and caused post‐fire herbaceous composition to “over‐shift” toward less productive C₄ short‐grasses rather than C₄ mid‐grasses. This study provides some of the first long‐term data showing a possible benefit of mixing seasonal fires (i.e., the alternate‐season fire treatment) in a prescribed burning management plan to restore C₄ mid‐grass cover and enhance overall herbaceous diversity.     

Competition by herbs as a limiting factor in shrub invasion in grassland: a test with different growth forms

Abstract. We tested the hypothesis that seedling establishment, the critical stage in the invasion of grassland by shrubs, is limited by competition with perennial grasses in seasonally wet/dry savannas. We placed seeds of two invasive exotic shrubs – Cryptostegia grandiflora, a woody vine, and Acacia nilotica, an arborescent legume – into pots with a wide range of existing above- and below-ground herbaceous biomass provided by either a tussock or a stoloniferous perennial grass. We also imposed different levels of watering frequency (5, 10 and 21 d), nutrient addition (+ and -) and grass clipping intensity (no clipping, clipped to 5 cm and clipped to 25 cm). There was no effect of any treatment on shrub seedling emergence or survival and all of the seedlings that emerged survived the 90-d growing period. Herbaceous competition also failed to have an effect on biomass accumulation in shrub seedlings. More frequent watering significantly increased above- and below-ground biomass accumulation for both shrub species and nutrient addition significantly increased Cryptostegia biomass accumulation. Based on these results, we question the proposition that reduction in competition by herbs via livestock grazing has been a significant factor in determining the rate or pattern of exotic shrub increase in the seasonally wet/dry tropics. We also question the suitability of the two-layer soil moisture hypothesis as a basis for management practices to control the ingress of woody species into grasslands and open savannas.     

Phosphorus responses of C3 and C4 species

An hypothesis was formulated that phosphorus (P) partitioningin tissues of C₄ leaves would permit C₄ plants to resist P deficiencybetter than C₃ plants. To test this hypothesis, 12 C₃, C₄, andC₃–C₄ intermediate species were grown at adequate, deficient,and severely deficient P supply in a solid-phase-buffered sandculture system to characterize photosynthetic and growth responses.Species differed considerably in response to P stress. The growthof C₃ species was more sensitive to P supply than C₄ species,but C₃ and C₄ species had similar photosynthetic P use efficiency,and C₄ species did not have low leaf P content, contrary toour hypothesis. In fact, leaf photosynthetic rates were notcorrelated with growth responses. Moncots had lower leaf P contentand better maintenance of leaf production under P stress thandicots, because of greater inhibition of branching (dicots)than of tillering (monocots). The most P efficient species inthis survey was Brachiaria, a C₄ monocot that increased rootbiomass allocation under stress while maintaining P allocationto the shoot. It is concluded that C₄ species are not inherentlymore P efficient than C₃ species, but that monocots are moreP efficient than dicots, because of contrasting P and biomassallocation under stress. Key words: Phosphorus deficiency, C₃ plants, C₄ plants, growth response     

Contribution of root respiration to soil respiration in a C3/C4 mixed grassland

The spatial and temporal variations of soil respiration were studied from May 2004 to June 2005 in a C₃/C₄ mixed grassland of Japan. The linear regression relationship between soil respiration and root biomass was used to determine the contribution of root respiration to soil respiration. The highest soil respiration rate of 11-54 Μmol m^-2 s^-1 was found in August 2004 and the lowest soil respiration rate of 4.99 Μmol m^-2 s^-1 was found in April 2005. Within-site variation was smaller than seasonal change in soil respiration. Root biomass varied from 0.71 kg m^-2 in August 2004 to 102 in May 2005. Within-site variation in root biomass was larger than seasonal variation. Root respiration rate was highest in August 2004 (5.7 Μmol m^-2 s^-1) and lowest in October 2004 (1.7 Μmol m^-2 s^-1). Microbial respiration rate was highest in August 2004 (5.8 Μmol m^-2 s^-1) and lowest in April 2005 (2.59 Μmol m^-2 s^-1). We estimated that the contribution of root respiration to soil respiration ranged from 31% in October to 51% in August of 2004, and from 45% to 49% from April to June 2005.     

Different growth responses of C3 and C4 grasses to seasonal water and nitrogen regimes and competition in a pot experiment

Understanding temporal niche separation between C₃ and C₄ species(e.g. C₃ species flourishing in a cool spring and autumn whileC₄ species being more active in a hot summer) is essential forexploring the mechanism for their co-existence. Two parallelpot experiments were conducted, with one focusing on water andthe other on nitrogen (N), to examine growth responses to wateror nitrogen (N) seasonality and competition of two co-existingspecies Leymus chinensis (C₃ grass) and Chloris virgata (C₄grass) in a grassland. The two species were planted in eithermonoculture (two individuals of one species per pot) or a mixture(two individuals including one L. chinensis and one C. virgataper pot) under three different water or N seasonality regimes,i.e. the average model (AM) with water or N evenly distributedover the growing season, the one-peak model (OPM) with morewater or N in the summer than in the spring and autumn, andthe two-peak model (TPM) with more water or N in the springand autumn than in the summer. Seasonal water regimes significantlyaffected biomass in L. chinensis but not in C. virgata, whileN seasonality impacted biomass and relative growth rate of bothspecies over the growing season. L. chinensis accumulated morebiomass under the AM and TPM than OPM water or N treatments.Final biomass of C. virgata was less impacted by water and Nseasonality than that of L. chinensis. Interspecific competitionsignificantly decreased final biomass in L. chinensis but notin C. virgata, suggesting an asymmetric competition betweenthe two species. The magnitude of interspecific competitionvaried with water and N seasonality. Changes in productivityand competition balance of L. chinensis and C. virgata undershifting seasonal water and N availabilities suggest a contributionof seasonal variability in precipitation and N to the temporalniche separation between C₃ and C₄ species. Key words: Chloris virgata, competition, growth, Leymus chinensis, nitrogen seasonality, water seasonality Received 19 November 2007; Revised 29 January 2008 Accepted 4 February 2008     

Variation in growth rate in shoot populations of the clonal dwarf shrub Linnaea borealis

This study was based on two years of field observations of a Linnaea borealis population in a coniferous forest in Sweden. Detailed information on performance and survivorship of shoots were gathered from consecutively made drawings of 90 individually marked shoot systems of Linnaea distributed among 15 plots in the forest. Branching of the shoot systems was correlated to growth of the main shoot. Increasing light flux negatively affected growth of Linnaea . A tendency for a competitive effect on Linnaea of Vaccinium spp. was also found. In contrast, the surrounding mosses, and intraspecific density of Linnaea had no significant effect on the shoot systems. The growth rate of the shoot population was analysed by matrix models. Population growth rate was most dependent on survivorship and growth of main shoots in Linnaea , while the survivorship of lateral shoots was of less importance. In spite of an average decrease in the population size as revealed by the matrix model simulation, the incorporation of the observed spatial variation in shoot growth increment, branching and shoot survivorship, yielded varying population growth rates. Parts of the population increased in size. The importance of choice of spatial scale in population analyses of forest understory plants is emphasized.     

Influence of summer rainfall on root and shoot growth of a cold-winter desert shrub,Atriplex confertifolia

Summary The influence of irrigation and nitrogen fertilization in early summer on root and shoot growth of Atriplex confertifolia, a C₄ shrub species, was examined in a cold-winter desert community in northern Utah. Soil water and xylem pressure potentials were monitored during the summer period.At the time of watering the surface soil (0–30 cm) was dry but there were turgid fine roots in this horizon. Watering of the soil reduced plant water stress from-30 to-15 bars (dawn values) indicating that roots near the surface were capable of absorbing water, and induced root growth in the 0–30 cm zone. The addition of N to the water treatment did not further increase root production. However, watering and watering +N fertilizer failed to stimulate shoot elongation or any dry weight increase of shoots. This shoot dormancy during summer is not typical of C₄ plants and is probably associated with adaptation to the cool arid environment.This work was carried out while the senior author was on study leave from CSIRO     

Comparative gas exchange and growth responses of C3 and C4 beach species grown at different salinities

Summary Comparative laboratory gas exchange and relative growth rate experiments were conducted on three native California coastal strand species at four salinity treatment levels. Relative mesophyll conductance sensitivities to salinity of Atriplex leucophylla (Moq.) D. Dietr. (C₄) and Atriplex californica Moq. in D.C. (C₃) were nearly identical. Mesophyll conductances of both species were stimulated by moderate levels of salinity. Mesophyll conductances of Abronia maritima Nutt. ex Wats. (C₃) were highest in the absence of salinity and depressed by increasing levels of salinity. Increasing levels of salinity generally decreased net photosynthesis and leaf conductances but increased water use efficiencies. The C₄ species, Atriplex leucophylla, had higher mesophyll conductances and water use efficiencies at all salinity levels than the C₃ species. The effects of salinity on relative growth and net assimilation rates of greenhouse grown plants were not directly correlated with the effects on net photosynthesis measured in the laboratory. Growth of Abronia maritima was greatly stimulated by low levels of salinity whereas photosynthesis was substantially inhibited. The possible significance of C₄ photosynthesis in relation to salinity is discussed.     

Density-dependent growth responses in two clonal herbs: regulation of shoot density

Summary It has been shown in clonal perennial herbs that shoot natality decreases, and shoot mortality increases, in stands of increasing density. In a two-year garden experiment, we have tested Hutchings' (1979) hypothesis that these responses are the result of physiological integration, i.e. the exchange of resources and growth substances between shoots of a single clone. Dense monocultures of two rhizomatous graminoids, Brachypodium pinnatum and Carex flacca, were created that differed more than 10-fold in the density of clones (genets), but that had similar densities of shoots. A more effective shoot density control was expected in stands with the smaller clone densities (larger clones) due to more extensive clonal connections. Shoot turnover was evaluated by counting living and dead shoots at different times. In the summer of the second year, when shoot densities and stand structure were similar between treatments, shoot natality (the number of shoots born per plot) and shoot mortality (the number of shoots that died per plot) were usually unrelated to clone density in either species. If there was a significant treatment effect, it could be attributed to (small) differences in shoot density. Over the whole range of shoot densities, natality was negatively density-dependent. The number of shoots that died in a given growth period was proportional to the number of shoots present, suggesting that mortality rates were density independent. In Carex, however, there were some indications that mortality rate increased with increasing density. Our study confirms that clonal herbaceous species can effectively prevent an overproduction of shoots, but in contrast to Hutchings' (1979) propositions, we found no evidence that physiological integration may be the responsible mechanism. An alternative explanation for the observed patterns is proposed.     

The small scale spatial pattern of C3 and C4 grasses depends on shrub distribution

At micro‐site scale, the spatial pattern of a plant species depends on several factors including interactions with neighbours. It has been seen that unfavourable effects generate a negative association between plants, while beneficial effects generate a positive association. In grasslands, the presence of shrubby species promotes a particular microenvironment beneath their canopy that could affect differently the spatial distribution of plants with different tolerance to abiotic conditions. We measured photosynthetic active radiation, air temperature and wind speed under shrub canopies and in adjacent open sites and analysed the spatial distribution of four grass species (two C₃ and two C₄) in relation to shrub canopy in a grazed sub‐humid natural grassland in southern Uruguay. Radiation, air temperature and wind speed were lower under shrubs than in adjacent open sites. The spatial distribution of grasses relative to the shrub canopy varied depending on the photosynthetic metabolism of grasses. C₄ grasses showed a negative association or no correlation with the shrubs, whereas C₃ grasses showed a positive association. Our results highlight the importance of the photosynthetic metabolism of the grasses in the final outcome of interactions between grasses and shrubs. Micro‐environmental conditions generated underneath shrubs create a more suitable site for the establishment of C₃ than for C₄ grasses. These results show that facilitation could be more important than previously thought in sub‐humid grasslands.     

Differences in tree and shrub growth responses to climate change in a boreal forest in China

Global climate change has led to rising temperatures and drought in boreal forests in Northeast China. In some areas, shrubs and trees coexist in high altitude and high latitude areas, and their differences with global warming may lead to significant changes in vegetation composition and distribution. Therefore, we compared the relationships between climate and growth for the most widely distributed dwarf shrub (Pinus pumila) and the two dominant tree species (Larix gmelinii and Pinus sylvestris var. mongolica) in boreal forests in the Daxing’an Mountains, China. A total of 340 tree-ring cores from 172 trees and 64 discs from shrubs were collected from four sites and compared the responses of shrub and tree growth to climate patterns using dendrochronological methods. The shrub and two tree species responded differently to interannual climate variance. The negative effect of growing season temperature was greater on growth of L.gmelinii and P. sylvestrisvar.mongolica than on P. pumila, and the promoting effect of winter and spring precipitation was greatest on P. pumila. Compared with the two tree species, P. pumila had a higher temperature threshold and grew over a shorter growing season. Our findings suggested that L. gmelinii and P. sylvestrisvar.mongolica are more susceptible to global warming than the shrubs that coexist with them. However, P.pumila should be studied from an individual perspective in the future due to the dwarf morphology of shrubs and their complex microenvironment.     

Shoot growth dynamics and size-dependent shoot fate of a clonal plant,Festuca rubra, in a mountain grassland

The relation of the within-season and between-season patterns of shoot growth were compared in a clonal grass with long-lived shoots,Festuca rubra, in a mown mountain grassland. The growth rate of shoot length from spring to summer in a year was almost constant for each shoot irrespective of spring shoot length each year. The annual shoot growth rate from spring to spring was negatively correlated with the shoot length in the first spring. Shoots of different length and age therefore tended to converge over time to a population of identical shoot size, suggesting an equalizing effect of growth pattern on size structure. Shoot size (shoot length and number of leaves) influenced the fates of shoots. Larger shoots showed an increased incidence of both flowering and formation of intravaginal daughter shoots and a decreased incidence of death in the subsequent time period. The fates of shoots were independent of their age. Although the negatively size-dependent springto-spring annual shoot growth rate acted to decrease shoot size variation, the remaining variation within the shoot population was still sufficient to generate different fates of shoots. These fates were not related to the previous life history of individual shoots. There was a significantly positive effect of the shoot size at initiation on its life expectancy. This was mainly attributable to the positively size-dependent survival rate of shoots in the early stage (<1 year old) of shoot life history. Later on (> 1 year old), shoot size had little effect on the survival rate of shoots. Once small young shoots have survived this early stage (< 1 year old) in life history, they can grow vigorously, little affected by competition regardless of shoot size, and converge to a stable size structure of shoots of similar size. Only shoot size in the early stage ( < 1 year old) of life history is important for the persistence of a shoot population.     

Plant–community responses to shrub cover in a páramo grassland released from grazing and burning

Shrub encroachment can follow grazing or burning release in páramo grasslands. While encroachment decreases herbaceous species richness in some grassland systems, the effects of this process on the herbaceous community in páramo grasslands are currently unknown. We collected data on shrub cover, herbaceous‐species cover and species composition in a páramo grassland 12 years after release from burning and cattle grazing near Zuleta, Ecuador. Topographic and soil measures were also included as predictor variables of differences in community composition. Contrary to studies in other systems, shrub cover did not have a significant effect on herbaceous‐species richness, whereas shrub‐species richness significantly increased with shrub cover. However, shrub cover was associated with significant shifts in herbaceous–community composition. Most notably, there was an increase in some shade‐tolerant forbs and tall‐statured wetland grasses with increasing shrub cover, and a corresponding decrease in some short‐statured grasses and early successional forbs. These results could indicate that the ameliorative effects of shrubs (e.g. frost and wind protection) in harsh alpine environments may partially compensate for the expected competitive effect of shrubs due to shading.