C4 Perennial Grass Research Articles

Root Morphology and Anatomy of Field-Grown Erianthus arundinaceus

Erianthus species are perennial C4 grasses with such high biomass productivity and high tolerance to environmental stresses that they can be grown in marginal land to supply raw material for cellulosic bioethanol. Because high biomass production and strong tolerance to environmental stresses might be based on their large and deep-root system, we closely examined the morphology and anatomy of roots in first-year seedlings of field-grown Erianthus arundinaceus. The deep-root system of E. arundinaceus consists of many nodal roots growing with steep growth angles. Diameter of nodal roots with large variations (0.5 - 5 mm) correlates with the size and number of large xylem vessels. The microscopic observation shows that the nodal roots with dense root hairs developed soil sheath, hypodermis with lignified sclerenchyma in the outer cortex, and aerenchyma in the mid-cortex. In addition, starch grains were densely accumulated in the stele of nodal roots in winter. In the first year, E. arundinaceus developed less lateral roots than other reported grass species. The lateral roots formed a large xylem vessel in the center of the stele and no hypodermis in the outer cortex. Morphology and anatomy of E. arundinaceus root were discussed with reference to strong tolerance to environmental stresses.

Plant community response to the East Amarillo Complex wildfires in the Southern High Plains, USA

Severe wildfires are increasing in extent in the western US. We used a matched-pairs design with plots in burned and non-burned areas to study effects of the East Amarillo Complex (EAC) wildfires on mixed grass prairie mean community composition and variability in composition. Species composition and ground cover data were collected at 5 study sites each year for three years following the EAC. Fire effects on mean species composition were analyzed with permutational analyses of variance; temporal patterns were analyzed with permutational anova and nonmetric multidimensional scaling; and an index of multivariate dispersion was used to assess variability in plant community composition. We found weak immediate impacts (year 1) of wildfire on mean species composition, but strong impacts by year three. Two general patterns of changes in mean species composition emerged: at 3 study sites, there was a progressive divergence in similarity between burned and non-burned areas following wildfire whereas at 2 study sites, wildfire effects lessened over time. In contrast, a trend of increased homogeneity in burned vegetation relative to non-burned vegetation was apparent at all 5 study sites 2 to 3 years post-fire; burned areas also had higher species diversity, higher evenness but similar species richness 3 years post-fire. Ground cover composition, which was immediately impacted by wildfire through removal of residual dry matter, was fully recovered to non-burned conditions 3 years after wildfire. We observed little seedling recruitment either of native or exotic species following wildfire. Community composition changes in burned areas were likely the result of changing dominance relationships among plants that survived the wildfires rather than the result of recruitment of new individuals in gaps created by dead plants. Given the growth form of perennial C4 grasses and the fact that these grasslands evolved in the context of repeated fire, it is likely that changes in mean species composition and compositional variability are short-term responses.

Chloroplast DNA variation and genetic structure of Miscanthus sinensis in southwest China

Miscanthus sinensis is a dominant perennial C4 grass with the potential to being a feedstock crop in North America, Europe, and China. Variation in chloroplast DNA sequence was used to obtain information regarding the genetic diversity and structure of populations of M. sinensis in southwest China. Chloroplast DNA, trnL-F and rpl20-rps12 sequences from seventy-five individuals representing 14 populations of M. sinensis were used to study the sequence variation. Seven haplotypes and 16 polymorphic sites (2.7%) were identified. Five substitutions, 6 indels, and 5 existing substitutions and indels sites, were detected through splicing these two gene segments. The genetic diversity within the studied populations (diversity of haploids, h = 0.561, nucleotide diversity, π = 0.00504) was low, this may be affected by the relatively larger effect of genetic drift on the chloroplast DNA, reflecting smaller effective populations than nuclear DNA. Genetic variance within the populations was higher than that between the populations, suggesting that higher gene flow may exist within these populations. The results of parsimony network in seven haplotypes indicated that H1 and H2 may be ancient haplotypes, which may help guide future research on the origin of M. sinensis. Our results provide information on the genetic diversity and structure of M. sinensis and may assist future studies on the phylogeography of M. sinensis.

Miscanthus agronomy and bioenergy feedstock potential on minesoils

The US government has mandated production of 79 million liters of biofuel from lignocellulose biomass and advanced fuels by 2022. To meet this requirement, the Department of Energy is encouraging research to develop herbaceous lignocellulose-based bioethanol for use as transport fuel. Miscanthus × giganteus was introduced in the US during 1930s, and is being widely studied for its potential to produce large biomass yield with minimum input in different soils. Miscanthus × giganteus is highly productive, sterile rhizomatous C4 perennial grass adapted to a wide range of climatic and soil conditions. Since the early 1980s, this crop has been studied under various climate and soil conditions in Europe and used to produce heat and electricity by combustion. This paper summarizes its agronomy and the characteristics which make it a potential dedicated bioenergy crop suitable for the reclaimed minesoils of the Appalachian region: Ohio, West Virginia, Virginia, Pennsylvania, Maryland, Kentucky, and Tennessee. The area which has been mined in the Appalachian region is estimated at 1.1 million hectares and only about 5% has been fully reclaimed. Using minesoils for miscanthus bioenergy feedstock production minimizes competition for arable land and sequesters soil organic carbon in these degraded lands. Reclaimed minesoils in the Appalachian region has potential to produce 9.22 × 106 Mg yr−1 dry biomass of Miscanthus × giganteus feedstock.

Testing for Thresholds in a Semiarid Grassland: The Influence of Prairie Dogs and Plague

State-and-transition models for semiarid grasslands in the North American Great Plains suggest that the presence of herbivorous black-tailed prairie dogs (Cynomys ludovicianus) on a site 1) creates a vegetation state characterized by increased dominance of annual forbs and unpalatable bunchgrasses and increased bare soil exposure and 2) requires long-term (> 40 yr) prairie dog removal to transition back to a vegetation state dominated by palatable perennial grasses. Here, we examine 1) how the recent history of prairie dog occupancy on a site (1–10 yr) influences the magnitude of prairie dog effects on vegetation composition and 2) how occupancy history affects vegetation dynamics following extirpation of prairie dogs. We used a natural experiment in the shortgrass steppe of northeastern Colorado, USA, where prairie dogs were extirpated from multiple sites during an outbreak of epizootic plague. On sites occupied by prairie dogs for 1–4 yr prior to extirpation, plant cover and composition recovered to conditions similar to unoccupied sites within a single growing season. Larger reductions in perennial C4 grasses occurred on sites occupied for the prior 7–10 yr compared to sites with shorter occupancy histories (< 6 yr). On sites occupied for the prior 7–10 yr, C4 perennial grasses recovered after 5 yr following prairie dog extirpation; in addition, C3 perennial graminoids and forbs remained more abundant (compared to sites with no history of prairie dogs) throughout the 5-yr period. Our findings showcase that prior site occupancy (up to 10 yr) by prairie dogs did not induce irreversible shifts in vegetation state in this semiarid grassland. Rather, vegetation changes induced by prairie dogs represent primarily a phase shift in landscapes where prairie dog populations are regulated by epizootic plague.

Management of Perennial Warm-Season Bioenergy Grasses. II. Seasonal Differences in Elephantgrass and Energycane Morphological Characteristics Affect Responses to Harvest Frequency and Timing

Elephantgrass (Pennisetum purpureum Schum.) and energycane (Saccharum spp. interspecific hybrid) are perennial C4 grasses with potential for use as bioenergy feedstocks. Their biomass production has been quantified, but differences in plant morphology and the relationship of morphology with biomass harvested and plant persistence are not well understood. The objective was to quantify monthly changes in morphological characteristics of elephantgrass (cv. Merkeron and breeding line UF1) and energycane (cv. L 79-1002) and relate these changes to biomass accumulation and plant responses to defoliation. All were evaluated monthly during full-season growth or when defoliated once in mid-season. Merkeron and UF1 elephantgrass generally showed similar morphological characteristics. Relative to energycane, elephantgrass had fewer tillers early in the growing season, less seasonal variation in tiller number, greater tiller mass and maximum leaf area index (LAI), and earlier spring development of LAI. Energycane showed slower leaf area development in spring, lower maximum LAI, and shorter period of increasing tiller mass and canopy height during the growing season relative to UF1. Elephantgrass had greater incidence of lodging than energycane when exposed to high wind, likely due to greater elephantgrass tiller mass. Morphological characteristics of tall-growing bioenergy grasses help to explain differences among them in biomass production and plant persistence responses to defoliation.

The growth form of dominant grasses regulates the invasibility of Uruguayan grasslands

In this paper we report the results of a small‐scale experiment on patches of natural grasslands in Uruguay. We analyzed whether growth form of C4 grasses affects invasion success. The experimental plots represented three patch‐types arising from short‐term grazing effects and were dominated by prostrate C4 grasses, erect C4 grasses and codominated by prostrate C4 grasses and forbs. We seeded the plots with four exotic species: one perennial C4 grass, one annual C3 grass, one perennial forb and one annual legume. Contrary to many studies conducted at small spatial scales, our results suggest that in our system, the processes that control invasibility are not directly linked with resident species richness or biomass, but rather are driven by growth form. Emergence of invaders did not differ among patch‐types, but seedlings survived less in plots dominated by erect grasses. We suggest that the erect growth form of C4 grass species is more successful in preventing invasion by mediating seedling competition for light under the canopy for a longer period of time. Grazing management, through its ability to regulate canopy architecture, is a potentially important tool in controlling invasion.

Effects of a 20-year old Miscanthus × giganteus stand and its removal on soil characteristics and greenhouse gas emissions

Miscanthus is a perennial rhizomatous C4 grass with high yield potentials and low nutrient needs, thus a promising candidate for the production of cellulosic biomass. While optimal management options and yields attainable on a commercial scale are still debated, no study has yet addressed its removal and potential effects on following crops. Here, we present results from a trial involving a 20-year old Miscanthus stand on i) soil C, N, P and K stocks, compared with an adjacent field cultivated with a rotation of annual arable food crops, ii) the greenhouse gases (GHG) emissions following the removal of Miscanthus and iii) the immediate short-term effects on the following land use (re-cultivation to wheat or set-aside).Compared to the adjacent field under annual crops, the Miscanthus plot had a larger soil organic C stock (by 13 t C ha−1) but a similar N stock, and lower P and K stocks (with differences of 100 kg P ha−1 and 1170 kg K ha−1, respectively). These losses imply that some degree of fertilization may be necessary as compensation. The effects of Miscanthus removal for the following wheat were significant on crop N content but negligible on grain yield.1.5 t CO2 ha−1 of CO2 were released after the Miscanthus removal and the N2O emissions increased from 150 g N2O-N ha−1 to 493 g N2O-N ha−1 during the following year. These results highlight the importance of investigation of the end-of-life stage of perennial crops for an accurate assessment of their environmental impacts.

Soil moisture--a regulator of arbuscular mycorrhizal fungal community assembly and symbiotic phosphorus uptake.

Multiple species of arbuscular mycorrhizal fungi (AMF) can colonize roots of an individual plant species but factors which determine the selection of a particular AMF species in a plant root are largely unknown. The present work analysed the effects of drought, flooding and optimal soil moisture (15-20 %) on AMF community composition and structure in Sorghum vulgare roots, using PCR-RFLP. Rhizophagus irregularis (isolate BEG 21), and rhizosphere soil (mixed inoculum) of Heteropogon contortus, a perennial C4 grass, collected from the semi-arid Delhi ridge, were used as AMF inocula. Soil moisture functioned as an abiotic filter and affected AMF community assembly inside plant roots by regulating AMF colonization and phylotype diversity. Roots of plants in flooded soils had lowest AMF diversity whilst root AMF diversity was highest under the soil moisture regime of 15-20 %. Although plant biomass was not affected, root P uptake was significantly influenced by soil moisture. Plants colonized with R. irregularis or mixed AMF inoculum showed higher root P uptake than non-mycorrhizal plants in drought and control treatments. No differences in root P levels were found in the flooded treatment between plants colonized with R. irregularis and non-mycorrhizal plants, whilst under the same treatment, root P uptake was lower in plants colonized with mixed AMF inoculum than in non-mycorrhizal plants.

Dynamics of soil organic carbon pools after agricultural abandonment

Abandonment of agricultural land and the subsequent recolonization by natural vegetation is known to cause increases in C contents, contributing to reduction in atmospheric CO2 concentrations. Assessment of the possible mitigation of CO2 excess requires understanding the SOC dynamics, the origin of C pools and the pathways of their transformation. The aims of this work were to assess, by using the δ13C signature, the changes of old and new organic C in total (soil organic carbon, SOC) and labile (microbial biomass C, MBC, dissolved organic C, DOC, CO2 efflux from soil) pools after vegetation change from vineyard (C3) to grassland (C4) under semiarid Mediterranean climate. Colonization of abandoned vineyard by the perennial C4-grass Hyparrhenia hirta after 15 or 35years increased topsoil C stocks by 13% and 16%, respectively. Such an increase was attributed to new above- and below-ground biomass C input from H. hirta. The maximal incorporation of new C was observed in MBC, whereas the DOC derived mainly from old SOC. Based on δ13C values of SOC, MBC, DOC and CO2 in C3 soil and in soils after 15 and 35years of C4 grass colonization, 13C fractionation per se from changes in isotopic composition by preferential utilization of substrates with different availability was separated. MBC in C3–C4 soil used more recent (13C-enriched) versus old C (relatively 13C-depleted) sources. The Δ13C by decomposition of SOC to CO2 (δ13C of CO2 minus δ13C of SOC) was higher than Δ13C by microbial respiration (CO2 minus MBC), demonstrating that under semiarid climate, soil microorganisms do not always preferentially decompose the most available SOC pools. The use of δ13C signature of SOC after C3–C4 vegetation change combined with soil incubation is a powerful tool to assess the exchange between old and new C in pools of various availability.

Diversity of nitrogen-fixing bacteria associated with switchgrass in the native tallgrass prairie of northern Oklahoma.

Switchgrass (Panicum virgatum L.) is a perennial C4 grass native to North America that is being developed as a feedstock for cellulosic ethanol production. Industrial nitrogen fertilizers enhance switchgrass biomass production but add to production and environmental costs. A potential sustainable alternative source of nitrogen is biological nitrogen fixation. As a step in this direction, we studied the diversity of nitrogen-fixing bacteria (NFB) associated with native switchgrass plants from the tallgrass prairie of northern Oklahoma (United States), using a culture-independent approach. DNA sequences from the nitrogenase structural gene, nifH, revealed over 20 putative diazotrophs from the alpha-, beta-, delta-, and gammaproteobacteria and the firmicutes associated with roots and shoots of switchgrass. Alphaproteobacteria, especially rhizobia, predominated. Sequences derived from nifH RNA indicated expression of this gene in several bacteria of the alpha-, beta-, delta-, and gammaproteobacterial groups associated with roots. Prominent among these were Rhizobium and Methylobacterium species of the alphaproteobacteria, Burkholderia and Azoarcus species of the betaproteobacteria, and Desulfuromonas and Geobacter species of the deltaproteobacteria.

Excreting and non-excreting grasses exhibit different salt resistance strategies.

The combination of traits that makes a plant successful under saline conditions varies with the type of plant and its interaction with the environmental conditions. Knowledge about the contribution of these traits towards salt resistance in grasses has great potential for improving the salt resistance of conventional crops. We attempted to identify differential adaptive response patterns of salt-excreting versus non-excreting grasses. More specifically, we studied the growth, osmotic, ionic and nutrient (carbon/nitrogen) relations of two salt-excreting (Aeluropus lagopoides and Sporobolus tremulus) and two non-excreting (Paspalum paspalodes and Paspalidium geminatum) perennial C4 grasses under non-saline and saline (0, 200 and 400 mM NaCl) conditions. Growth and relative growth rate decreased under saline conditions in the order P. geminatum > S. tremulus = A. lagopoides > P. paspalodes. The root-to-shoot biomass allocation was unaffected in salt-excreting grasses, increased in P. paspalodes but decreased in P. geminatum. Salt-excreting grasses had a higher shoot/root Na(+) ratio than non-excreting grasses. K(+), Ca(2+) and Mg(2+) homoeostasis remained undisturbed among test grasses possibly through improved ion selectivity with rising substrate salinity. Salt-excreting grasses increased leaf succulence, decreased ψs and xylem pressure potential, and accumulated proline and glycinebetaine with increasing salinity. Higher salt resistance of P. paspalodes could be attributed to lower Na(+) uptake, higher nitrogen-use efficiency and higher water-use efficiency among the test species. However, P. geminatum was unable to cope with salt-induced physiological drought. More information is required to adequately document the differential strategies of salt resistance in salt-excreting and non-excreting grasses.

A footprint of past climate change on the diversity and population structure of Miscanthus sinensis

Miscanthus is a perennial C4 grass that is a leading potential feedstock crop for the emerging bioenergy industry in North America, Europe and China. However, only a single, sterile genotype of M. × giganteus (M×g), a nothospecies derived from diploid M. sinensis (Msi) and tetraploid M. sacchariflorus (Msa), is currently available to farmers for biomass production. To facilitate breeding of Miscanthus, this study characterized genetic diversity and population structure of Msi in its native range of East Asia. A total of 767 accessions were studied, including 617 Msi from most of its native range in China, Japan and South Korea, and 77 ornamental cultivars and 43 naturalized individuals from the USA. Accessions were evaluated with 21 207 restriction site-associated DNA sequencing single nucleotide polymorphism (SNP) markers, 424 GoldenGate SNPs and ten plastid microsatellite markers. Six genetic clusters of Msi from geographically distinct regions in Asia were identified. Genetic data indicated that (1) south-eastern China was the origin of Msi populations found in temperate eastern Asia, which is consistent with this area probably having been a refugium during the last glacial maximum (LGM); (2) Msi migrated directly from south-eastern China to Japan before migrating to the same latitudes in China and Korea, which is consistent with the known sequence of warming post-LGM; (3) ornamental Msi cultivars were derived from the southern Japan population, and US naturalized populations were derived from a sub-set of the ornamental cultivars; and (4) many ornamental cultivars previously described as Msi have hybrid ancestry from Msa and Msi, whereas US naturalized populations of Msi do not. Population structure of Msi was driven by patterns of warming since the LGM, and secondarily by geographical barriers. This study will facilitate germplasm conservation, association analyses and identification of potential heterotic groups for the improvement of Miscanthus as a bioenergy crop.

Patrones de invasión de los pastizales de Uruguay a escala regional

Ecología Austral, 24:83-93 (2014)Studies on the relationship between the richness of native and exotic plants have generated conflicting evidence depending, among others, on the spatial scale of analysis. This study assessed the invasion by exotic species in natural grasslands of Uruguay at a regional scale. We analyzed 313 vegetation census from four geomorphological regions: Cuesta Basaltica, Cuenca Sedimentaria, Centro Sur and Sierras del Este. The results showed that the relationship between native and exotic richness was negative for Centro Sur while the effect of native species richness on exotic plant cover was negative for Centro Sur and Cuenca Sedimentaria. In most analyses, the functional type C4 grasses was negatively associated with the exotic richness and cover, and strongly inhibits Cynodon dactylon, the only C4 exotic grass species present in the sites. Conversely, C3 species showed irregular patterns of association with the exotics. Also, exotic richness was positively related with the percentage of agricultural use and vegetation cover. Our results suggest that propagule pressure and productivity favour the introduction and establishment of exotic species, but they are largely controlled by the composition of the resident community, in particular by the presence of C4 perennial grasses, which are the group with the highest cover in these communities. This could explain the very low incidence of exotic species in Uruguayan grasslands.

Elevated CO2-mitigation of high temperature stress associated with maintenance of positive carbon balance and carbohydrate accumulation in Kentucky bluegrass.

Elevated CO2 concentration may promote plant growth while high temperature is inhibitory for C3 plant species. The interactive effects of elevated CO2 and high temperatures on C3 perennial grass growth and carbon metabolism are not well documented. Kentucky bluegrass (Poa pratensis) plants were exposed to two CO2 levels (400 and 800 μmol mol−1) and five temperatures (15/12, 20/17, 25/22, 30/27, 35/32°C, day/night) in growth chambers. Increasing temperatures to 25°C and above inhibited leaf photosynthetic rate (Pn) and shoot and root growth, but increased leaf respiration rate (R), leading to a negative carbon balance and a decline in soluble sugar content under ambient CO2. Elevated CO2 did not cause shift of optimal temperatures in Kentucky bluegrass, but promoted Pn, shoot and root growth under all levels of temperature (15, 20, 25, 30, and 35°C) and mitigated the adverse effects of severe high temperatures (30 and 35°C). Elevated CO2-mitigation of adverse effects of high temperatures on Kentucky bluegrass growth could be associated with the maintenance of a positive carbon balance and the accumulation of soluble sugars and total nonstructural carbohydrates through stimulation of Pn and suppression of R and respiratory organic acid metabolism.

Biomass Production by Warm‐Season Grasses as Affected by Nitrogen Application in Ontario

There is little information on the production levels of prairie C4 perennial grasses in the biomass context in eastern Canada. The objective of this study is to evaluate the effect of species/cultivar entries and N fertilizer application on biomass production of prairie perennial biomass grasses. Two switchgrass (Panicum virgatum L.) cultivars, two big bluestem (Andropogon gerardii Vit.) cultivars and one indiangrass (Sorghastrum nutans Nash.) cultivar were seeded in 2009 in eastern Ontario to evaluate their biomass production potential. The crops were seeded in a silt loam soil at 1000 pure live seeds m−2. No fertilizer was added in 2009 or 2010. In 2011 and 2012 treatment N50 received 50 kg N ha−1 while treatment N0 was kept as an unfertilized control. The crop biomass production, canopy height, leaf SPAD absorbance, leaf area index, and harvest moisture content were determined. Switchgrass production exceeded big bluestem and indiangrass, especially in the first 2 yr after establishment. Despite the drought, dry matter production for the different entries in 2012 ranged between 5.07 and 8.13 Mg ha−1. Biomass production was improved by N application only for big bluestem and indiangrass but not for switchgrass. However, N application increased leaf SPAD absorbance and leaf area index for all entries. Moisture content at harvest was species dependent. Our results suggest that switchgrass might need lower N levels than big bluestem and indiangrass. With the stands reaching their peak production, this study shows the high production potential for some of these grasses under Ontario conditions.

Granivory reduces biomass and lignin concentrations of plant tissue during grassland assembly

Small mammals can influence grassland assembly by selecting against palatable plants – the community can become dominated by the plants they avoid. This predation-based selection could have indirect effects on community biomass and tissue quality, especially given how untasty plants may have higher concentrations of recalcitrant carbon compounds including lignin. We tested small mammal effects on biomass and tissue quality of roots and shoots in a two-year-old 18ha restored tallgrass prairie with established zones of high and low plant predation. We focused on the three dominant herbaceous functional groups of tallgrass prairie (perennial forbs, C3 and C4 grasses), and targeted the early stages of assembly given that plant predation by small animals can unfold quickly and is difficult to subsequently quantify. We predicted rodent predation to create communities with reduced biomass but an increased abundance of lignin-rich plants; we only observed the former. Rodents reduced aboveground biomass by 46% but preferentially targeted lignin-rich plants, with the latter result explained by the predominance of granivory over herbivory – there was no opportunity for selection based on tissue palatability. Based strictly on aboveground biomass, we estimated small mammals reduced standing stocks of recalcitrant carbon by 65kgha−1, with reductions in belowground stocks almost certainly higher given that root:shoot ratios averaged 21:1. Given that the quantity and quality of plant production can affect ecosystem functions including decomposition and the regulation of soil carbon stocks, our work suggests that non-random plant predation may substantially affect rates of soil carbon accumulation in the early stages of grassland development.

Differential performance and behavior of the corn leaf aphid, Rhopalosiphum maidis, on three species of the biomass crop miscanthus

Miscanthus, a perennial C4-grass, is identified as a promising candidate for biomass production owing to its high yield potential under low nitrogen input and its low susceptibility to diseases and pests. However, some significant insect pests of conventional cereal grain crops such as the corn leaf aphid (Rhopalosiphum maidis) are able to develop on miscanthus. In the prospect of extensive planting of miscanthus for bioenergy, this may lead to the worsening of pest problems in the surrounding conventional crops. The aim of this study was to assess, through laboratory experiments, the ability of R. maidis to colonize three main miscanthus species studied in Europe for biomass production, i.e., Miscanthus×giganteus, M. sacchariflorus, and M. sinensis. In this purpose, comparative analyses of aphid's performance and plant penetration behavior were performed. Despite an altered plant penetration behavior on miscanthus, R. maidis was able to colonize (i.e., to develop and reproduce) the three miscanthus species tested and exhibited a behavior adapted to the transmission of both persistent and non-persistent phytoviruses. M. sacchariflorus and, to a lesser extent, M. sinensis were found less suitable and acceptable host plant for R. maidis than M.×giganteus. Results are discussed in terms of agronomic risks and implications for miscanthus improvement programs.

Seasonality and facilitation drive tree establishment in a semi-arid floodplain savanna

A popular hypothesis for tree and grass coexistence in savannas is that tree seedlings are limited by competition from grasses. However, competition may be important in favourable climatic conditions when abiotic stress is low, whereas facilitation may be more important under stressful conditions. Seasonal and inter-annual fluctuations in abiotic conditions may alter the outcome of tree-grass interactions in savanna systems and contribute to coexistence. We investigated interactions between coolibah (Eucalyptus coolabah) tree seedlings and perennial C4 grasses in semi-arid savannas in eastern Australia in contrasting seasonal conditions. In glasshouse and field experiments, we measured survival and growth of tree seedlings with different densities of C4 grasses across seasons. In warm glasshouse conditions, where water was not limiting, competition from grasses reduced tree seedling growth but did not affect tree survival. In the field, all tree seedlings died in hot dry summer conditions irrespective of grass or shade cover, whereas in winter, facilitation from grasses significantly increased tree seedling survival by ameliorating heat stress and protecting seedlings from herbivory. We demonstrated that interactions between tree seedlings and perennial grasses vary seasonally, and timing of tree germination may determine the importance of facilitation or competition in structuring savanna vegetation because of fluctuations in abiotic stress. Our finding that trees can grow and survive in a dense C4 grass sward contrasts with the common perception that grass competition limits woody plant recruitment in savannas.

Drought resistance and soil water extraction of a perennial C4 grass: contributions of root and rhizome traits.

Previously, we showed that genotypic differences in soil water extraction were associated with drought response, but we did not study underground root and rhizome characteristics. In this study, we demonstrate a similar relationship between drought resistance and soil water extraction but investigate the role of underground organs. Eighteen bermudagrass genotypes (Cynodon spp.) from four climatic zones were assessed under continuous drought at two locations with contrasting soils and climates. The criterion for drought resistance was the duration required to reach 50% green cover (GC50) after water was withheld. GC50, physiological traits, rhizome dry matter (RhDM), root length density (RLD) and average root diameter (ARD) were determined in both locations; water extraction was measured in one location. Large genotypic variation for drought resistance was observed in both locations, with GC50 being 187-277 days in a clay soil and 15-27 days in a sandy soil. Drought-resistant genotypes had greater soil water extraction and a higher water uptake rate. GC50 was correlated with relative water content (r=0.76), canopy temperature differential (r=-0.94) and photosynthetic rate (r=0.87) measured during drought; RhDM (r=0.78 to ~0.93) before and after drought; and ARD after drought (r=0.82 to ~0.94); GC50 was not correlated with RLD. Ecotypes collected from the Australian Mediterranean zone had superior drought resistance and were characterised by a large rhizome network. This is the first comprehensive study with perennial C4 grasses describing the association between water extraction, root distribution, rhizomes and drought resistance.