Does Fungal Endophyte Infection Improve Tall Fescue’s Growth Response to Fire and Water Limitation?

Invasive species may owe some of their success in competing and co-existing with native species to microbial symbioses they are capable of forming. Tall fescue is a cool-season, non-native, invasive grass capable of co-existing with native warm-season grasses in North American grasslands that frequently experience fire, drought, and cold winters, conditions to which the native species should be better-adapted than tall fescue. We hypothesized that tall fescue’s ability to form a symbiosis with Neotyphodium coenophialum, an aboveground fungal endophyte, may enhance its environmental stress tolerance and persistence in these environments. We used a greenhouse experiment to examine the effects of endophyte infection (E+ vs. E−), prescribed fire (1 burn vs. 2 burn vs. unburned control), and watering regime (dry vs. wet) on tall fescue growth. We assessed treatment effects for growth rates and the following response variables: total tiller length, number of tillers recruited during the experiment, number of reproductive tillers, tiller biomass, root biomass, and total biomass. Water regime significantly affected all response variables, with less growth and lower growth rates observed under the dry water regime compared to the wet. The burn treatments significantly affected total tiller length, number of reproductive tillers, total tiller biomass, and total biomass, but treatment differences were not consistent across parameters. Overall, fire seemed to enhance growth. Endophyte status significantly affected total tiller length and tiller biomass, but the effect was opposite what we predicted (E−>E+). The results from our experiment indicated that tall fescue was relatively tolerant of fire, even when combined with dry conditions, and that the fungal endophyte symbiosis was not important in governing this ecological ability. The persistence of tall fescue in native grassland ecosystems may be linked to other endophyte-conferred abilities not measured here (e.g., herbivory release) or may not be related to this plant-microbial symbiosis.     

Low allelopathic potential of an invasive forage grass on native grassland plants: a cause for encouragement?

Tall fescue (Festuca arundinacea Schreb.), a highly competitive European grass that invades US grasslands, is reportedly allelopathic to many agronomic plants, but its ability to inhibit the germination or growth of native grassland plants is unknown. In three factorial glasshouse experiments, we tested the potential allelopathic effects of endophyte-infected (E+) and uninfected (E−) tall fescue on native grasses and forbs from Midwestern tallgrass prairies. Relative to a water control, at least one extract made from ground seed, or ground whole plant tissue of E+ or E− tall fescue reduced the germination of 10 of 11 species in petri dishes. In addition, the emergence of two native grasses in potting soil was lower when sown with E+ and E− tall fescue seedlings than when sown with seeds of conspecifics or tall fescue. However, when seeds of 13 prairie species were sown in sterilized, field-collected soil and given water or one of the four tall fescue extracts daily, seedling emergence was lower in one extract relative to water for only one species, and subsequent height growth did not differ among treatments for any species. We conclude that if tall fescue is allelopathic, its inhibitory effects on the germination and seedling growth of native prairie plants are limited, irrespective of endophyte infection. On the other hand, the apparent inability of these plants to detect tall fescue in field soil could hinder prairie restoration efforts if germination near this strong competitor confers fitness consequences. We propose that lack of chemical recognition may be common among resident and recently introduced non-indigenous plants because of temporally limited ecological interactions, and offer a view that challenges the existing allelopathy paradigm. Lastly, we suggest that tall fescue removal will have immediate benefits to the establishment of native grassland plants.     

Control of Exotic Annual Grasses to Restore Native Forbs in Abandoned Agricultural Land

Exotic annual grasses are a major challenge to successful restoration in temperate and Mediterranean climates. Experiments to restore abandoned agricultural fields from exotic grassland to coastal sage scrub habitat were conducted over two years in southern California, U.S.A. Grass control methods were tested in 5 m² plots using soil and vegetation treatments seeded with a mix of natives. The treatments compared grass‐specific herbicide, mowing, and black plastic winter solarization with disking and a control. In year two, herbicide and mowing treatments were repeated on the first‐year plots, plus new control and solarization plots were added. Treatments were evaluated using percent cover, richness and biomass of native and exotic plants. Disking alone reduced exotic grasses, but solarization was the most effective control in both years even without soil sterilization, and produced the highest cover of natives. Native richness was greatest in solarization and herbicide plots. Herbicide application reduced exotics and increased natives more than disking or mowing, but produced higher exotic forb biomass than solarization in the second year. Mowing reduced grass biomass and cover in both years, but did not improve native establishment more than disking. Solarization was the most effective restoration method, but grass‐specific herbicide may be a valuable addition or alternative. Solarization using black plastic could improve restoration in regions with cool, wet summers or winter growing seasons by managing exotic seedbanks prior to seeding. While solarization may be impractical at very large scales, it will be useful for rapid establishment of annual assemblages on small scales.     

Prescribed Fire and Herbicide Effects on Soil Processes During Barrens Restoration

Prescribed fire has become a common tool of natural area managers for removal of non‐indigenous invasive species and maintenance of barrens plant communities. Certain non‐native species, such as tall fescue (Festuca arundinacea), tolerate fire and may require additional removal treatments. We studied changes in soil N and C dynamics after prescribed fire and herbicide application in remnant barrens in west central Kentucky. The effects of a single spring burn post‐emergence herbicide, combined fire and herbicide treatments, and an unburned no‐herbicide control were compared on five replicate blocks. In fire‐plus‐herbicide plots, fescue averaged 8% at the end of the growing season compared with 46% fescue cover in control plots. The extent of bare soil increased from near 0 in control to 11% in burned plots and 25% in fire‐plus‐herbicide plots. Over the course of the growing season, fire had little effect on soil N pools or processes. Fire caused a decline in soil CO₂ flux in parallel to decreased soil moisture. When applied alone, herbicide increased plant‐available soil N slightly but had no effect on soil respiration, moisture, or temperature. Fire‐plus‐herbicide significantly increased plant‐available soil N and net N transformation rates; soil respiration declined by 33%. Removal of non‐native plants modified the chemical, physical, and biological soil conditions that control availability of plant nutrients and influence plant species performance and community composition.     

Observations of responses to re‐introducing fire in a Basalt Plains grassland after the removal of grazing: Implications for restoration

Natural grasslands in southern Australia commonly exist in altered states. One widespread altered state is grassland pasture dominated by cool‐season (C3) native grasses maintained by ongoing grazing. This study explores the consequences of removing grazing and introducing fire as a conservation management tool for such a site. We examined the abundance of two native and three exotic species, across a mosaic of fire regimes that occurred over a three‐year period: unburnt, summer wild‐fire (>2 years previous), autumn management fire (<1 year previously) and burnt in both fires. Given that one aim of conservation management is to increase native species at the expense of exotics, the impacts of the fires were largely positive. Native grasses were at higher cover levels in the fire‐managed vegetation than in the unburnt vegetation. Of the three exotic species, one was consistently at lower density in the burnt plots compared to the unburnt plots, while the others were lower only in those plots burnt in summer. The results show that the response of a species varies significantly between different fire events, and that the effects of one fire can persist through subsequent fires. Importantly, some of the effects were large, with changes in the density of plants of over 100‐fold. Fire is potentially a cost‐effective tool to assist the ecological restoration of retired grassland pastures at large scales.     

An Invasive Grass Increases Live Fuel Proportion and Reduces Fire Spread in a Simulated Grassland

Fire is a globally important ecosystem process, and invasive grass species generally increase fire spread by increasing the fuel load and continuity of native grassland fuelbeds. We suggest that invasive grasses that are photosynthetically active, while the native plant community is dormant reduce fire spread by introducing high-moisture, live vegetation gaps in the fuelbed. We describe the invasion pattern of a high-moisture, cool-season grass, tall fescue (Schedonorus phoenix (Scop.) Holub), in tallgrass prairie, and use spatially explicit fire behavior models to simulate fire spread under several combinations of fuel load, invasion, and fire weather scenarios. Reduced fuel load and increased extent of tall fescue invasion reduced fire spread, but high wind speed and low relative humidity can partially mitigate these effects. We attribute reduced fire spread to asynchrony in the growing seasons of the exotic, cool-season grass, tall fescue, and the native, warm-season tallgrass prairie community in this model system. Reduced fire spread under low fuel load scenarios indicate that fuel load is an important factor in fire spread, especially in invaded fuel beds. These results present a novel connection between fire behavior and asynchronous phenology between invasive grasses and native plant communities in pyrogenic ecosystems.     

Using consecutive prescribed fires to reduce shrub encroachment in grassland by increasing shrub mortality

Woody plant encroachment into open grasslands occurs worldwide and causes multiple ecological and management impacts. Prescribed fire could be used to conserve grassland habitat but often has limited efficacy because many woody plants resprout after fire and rapidly reestablish abundance. If fire‐induced mortality could be increased, prescribed fire would be a more effective management tool. In California's central coast, shrub encroachment, especially of Baccharis pilularis (coyote brush), is converting coastal prairie into shrub‐dominated communities, with a consequent loss of native herbaceous species and open grassland habitat. B. pilularis has not been successfully controlled with single prescribed fire events because the shrub resprouts and reestablishes cover within a few years. We investigated whether two consecutive annual burns would control B. pilularis by killing resprouting shrubs, without reducing native herbaceous species or encouraging invasive plants. As expected, resprouting did occur; however, 2 years after the second burn, B. pilularis cover on burned plots was only 41% of the cover on unburned plots. Mortality of B. pilularis more than doubled following the second burn, likely maintaining a reduction in B. pilularis cover for longer than a single burn would have. Three native coastal prairie perennial grasses did not appear to be adversely affected by the two burns, nor did the burns result in increased cover of invasive species. Managers wanting to restore coastal prairie following B. pilularis encroachment should consider two consecutive annual burns, especially if moderate fire intensity is achievable.     

Fungal endophyte infection increases tall fescue's survival,growth, and flowering in a reconstructed prairie

Grasslands in North America are increasingly threatened by land conversion and ecological degradation, prompting restoration efforts to increase native plant species diversity and improve wildlife habitat. A major challenge is the removal and management of nonnative invasive species such as tall fescue (Schedonorus arundinaceus), which has a symbiotic association with a fungal endophyte (Epichloë coenophiala) that modifies its ecological interactions. Using transplanted clumps of the cultivar Kentucky‐31, we tested the effects of endophyte infection on tall fescue's survival and performance (tiller production, flowering, and basal area) for 5 years in a central Kentucky reconstructed prairie. We predicted that endophyte infected (E+) clumps would have increased performance compared to endophyte‐free (E?) clumps. Overall, E+ clumps had greater survival, tiller production, flowering tiller production, and basal area, but not reproductive effort (proportion of tillers flowering) as compared to E? clumps. However, survival and trends in tiller number and basal area over the 5‐year period suggested experimental tall fescue populations were in decline in the reconstructed prairie, although the E? population declined more rapidly. Our study provides evidence that endophyte infection improved tall fescue's growth and survival in a postreconstruction plant community, at least in the early years following reconstruction, and may increase the invasive potential of this nonnative species in prairie restorations.     

Endophytes inconsistently affect plant communities across Schedonorus arundinaceus hosts

Fungal endophytes in cool-season grasses may affect communities at multiple trophic levels. However, it is unclear whether community-scale endophyte effects arise due to the endophyte itself or as a result of unique, endophyte–host interactions. We used a long-term field experiment to test whether common-toxic (CT) and non-ergot alkaloid-producing (novel) endophytes in Schedonorus arundinaceus (tall fescue) forage cultivars consistently affect communities across tall fescue hosts. Tilled plots (2 × 2 m; Guelph, ON) were seeded with Georgia 5 and Jesup cultivars containing either the CT or AR542 (novel) endophyte and allowed to be re-colonized by plant species from the local propagule pool. Non-seeded control plots were included to assess effects of seeding the non-native grass. We assessed plant, invertebrate, soil moisture, and soil nutrient responses to the endophyte–cultivar treatments after four growing seasons. Seeding tall fescue affected plant species abundances, but not richness, and did not consistently alter soil moisture and nutrient pools. Endophyte identity in the tall fescue cultivars affected the communities, but effects were not consistent between cultivars. Within Georgia 5, the AR542 endophyte reduced tall fescue abundance and altered the invertebrate community relative to CT plots. Within Jesup, the AR542 endophyte reduced species evenness and decreased soil moisture during dry periods relative to CT plots. Endophyte effects were not consistent between cultivars, and it is probable that the community-scale effects of endophyte infection in tall fescue cultivars arise due to unique interactions between cultivar and endophyte.     

Insect oviposition preference between Epichloë‐symbiotic and Epichloë‐free grasses does not necessarily reflect larval performance

Variation in plant communities is likely to modulate the feeding and oviposition behavior of herbivorous insects, and plant‐associated microbes are largely ignored in this context. Here, we take into account that insects feeding on grasses commonly encounter systemic and vertically transmitted (via seeds) fungal Epichloë endophytes, which are regarded as defensive grass mutualists. Defensive mutualism is primarily attributable to alkaloids of fungal origin. To study the effects of Epichloë on insect behavior and performance, we selected wild tall fescue (Festuca arundinacea) and red fescue (Festuca rubra) as grass–endophyte models. The plants used either harbored the systemic endophyte (E+) or were endophyte‐free (E?). As a model herbivore, we selected the Coenonympha hero butterfly feeding on grasses as larvae. We examined both oviposition and feeding preferences of the herbivore as well as larval performance in relation to the presence of Epichloë endophytes in the plants. Our findings did not clearly support the female's oviposition preference to reflect the performance of her offspring. First, the preference responses depended greatly on the grass–endophyte symbiotum. In F. arundinacea, C. hero females preferred E+ individuals in oviposition‐choice tests, whereas in F. rubra, the endophytes may decrease exploitation, as both C. hero adults and larvae preferred E? grasses. Second, the endophytes had no effect on larval performance. Overall, F. arundinacea was an inferior host for C. hero larvae. However, the attraction of C. hero females to E+ may not be maladaptive if these plants constitute a favorable oviposition substrate for reasons other than the plants' nutritional quality. For example, rougher surface of E+ plant may physically facilitate the attachment of eggs, or the plants offer greater protection from natural enemies. Our results highlight the importance of considering the preference of herbivorous insects in studies involving the endophyte‐symbiotic grasses as host plants.     

Epichloae infection in a native South African grass,Festuca costata Nees

Fungal endophytes have been documented in almost all terrestrial plant groups. Although the endophyte infection syndrome in agronomic cultivars is well studied, relatively little work addresses questions of spatial ecology and fire effects on epichloae endophyte infection in native grasses, and none, to our knowledge, in sub‐Saharan Africa. We sampled seven populations of the native Festuca costata Nees along the spline of the Drakensberg range in South Africa at several spatial scales, including both recently burned and unburned stands. We tested epichloae presence and prevalence with immunoblot assays, PCR and genetic sequencing. We found epichloae endophytes were present and prevalent (38–98% infection rates depending on location). Variation in infection rates occurred primarily among locations, but also among bunches. There was little evidence that endophyte infection rates varied with fire. Novel evidence of epichloae infection of a native Festuca in South Africa opens the door to several new research questions, from the phylogenetic relationship between epichloae of sub‐Saharan Africa and other continents to the ecological advantages or disadvantages that endophytes confer upon their hosts, especially in a fire‐prone ecosystem vulnerable to global environmental change.     

Interannual climate variability mediates changes in carbon and nitrogen pools caused by annual grass invasion in a semiarid shrubland

Exotic plant invasions alter ecosystem properties and threaten ecosystem functions globally. Interannual climate variability (ICV) influences both plant community composition (PCC) and soil properties, and interactions between ICV and PCC may influence nitrogen (N) and carbon (C) pools. We asked how ICV and non-native annual grass invasion covary to influence soil and plant N and C in a semiarid shrubland undergoing widespread ecosystem transformation due to invasions and altered fire regimes. We sampled four progressive stages of annual grass invasion at 20 sites across a large (25,000 km²) landscape for plant community composition, plant tissue N and C, and soil total N and C in 2013 and 2016, which followed 2 years of dry and wet conditions, respectively. Multivariate analyses and ANOVAs showed that in invasion stages where native shrub and perennial grass and forb communities were replaced by annual grass-dominated communities, the ecosystem lost more soil N and C in wet years. Path analysis showed that high water availability led to higher herbaceous cover in all invasion stages. In stages with native shrubs and perennial grasses, higher perennial grass cover was associated with increased soil C and N, while in annual-dominated stages, higher annual grass cover was associated with losses of soil C and N. Also, soil total C and C:N ratios were more homogeneous in annual-dominated invasion stages as indicated by within-site standard deviations. Loss of native shrubs and perennial grasses and forbs coupled with annual grass invasion may lead to long-term declines in soil N and C and hamper restoration efforts. Restoration strategies that use innovative techniques and novel species to address increasing temperatures and ICV and emphasize maintaining plant community structure—shrubs, grasses, and forbs—will allow sagebrush ecosystems to maintain C sequestration, soil fertility, and soil heterogeneity.     

Host Status of Endophyte-Infected and Noninfected Tall Fescue Grass to Meloidogyne spp

Tall fescue grass cultivars with or without endophytes were evaluated for their susceptibility to Meloidogyne incognita in the greenhouse. Tall fescue cultivars evaluated included, i) wild-type Jesup (E+, ergot-producing endophyte present), ii) endophyte-free Jesup (E-, no endophyte present), iii) Jesup (Max-Q, non-ergot producing endophyte) and iv) Georgia 5 (E+). Peach was included as the control. Peach supported greater (P ≤ 0.05) reproduction of M. incognita than all tall fescue cultivars. Differences in reproduction were not detected among the tall fescue cultivars and all cultivars were rated as either poor or nonhosts for M. incognita. Suppression of M. incognita reproduction was not influenced by endophyte status. In two other greenhouse experiments, host susceptibility of tall fescue grasses to two M. incognita isolates (BY-peach isolate and GA-peach isolate) did not appear to be related to fungal endophyte strain [i.e., Jesup (Max-Q; nontoxic endophyte strain) vs. Bulldog 51 (toxic endophyte strain)]. Host status of tall fescue varied with species of root-knot nematode. Jesup (Max-Q) was rated as a nonhost for M. incognita (BY-peach isolate and GA-peach isolate) and M. hapla, a poor host for M. javanica and a good host for M. arenaria. Bulldog 51 tall fescue was also a good host for M. arenaria and M. javanica, but not M. incognita. Jesup (Max-Q) tall fescue may have potential as a preplant control strategy for M. incognita and M. hapla in southeastern and northeastern United States, respectively.     

Fungal endophyte presence and genotype affect plant diversity and soil-to-atmosphere trace gas fluxes

Aims

Novel fungal endophyte (Neotyphodium coenophialum; Latch, Christensen and Samuels; Glenn, Bacon, and Hanlin) genotypes in symbiosis with tall fescue (Lolium arundinaceum; Schreb. Darbysh.) have been recently introduced to agricultural seed markets. These novel endophytes do not produce the full suite of toxins that the ‘common toxic’ form does, and therefore, may not have the same consequences on plant and soil processes. Here, we evaluated the effects of endophyte presence and genotype on ecosystem processes of tall fescue stands.

Methods

We quantified the effects of the presence of the common toxic endophyte (CT), two novel endophyte genotypes (AR-542, AR-584), no endophyte (endophyte free, E-), and a mixture of all endophyte statuses (mix) within a single genotype of tall fescue (PDF) on various soil and plant parameters.

Results

Endophyte presence and genotype affected tall fescue cover and plant species diversity: cover—CT, AR-542, AR -584, mix > E- and species diversity—E- > AR-542, AR -584 > CT, mix. Most measured soil parameters had significant endophyte effects. For example, higher fluxes of soil CO₂ and N₂O were measured from stands of AR-542 than from the other endophyte treatments.

Conclusions

These results indicate that endophyte presence and genetic identity are important in understanding the ecosystem-scale effects of this agronomically important grass-fungal symbiosis.     

Experimental Restoration of Native Vegetation in Indiana Dunes National Lakeshore

We investigated the effects of prescribed fire, herbicide treatment, and sod removal on the eradication of exotic grasses and the establishment of native plant species in 24 experimental restoration plots in three razed residential sites within the boundary of Indiana Dunes National Lakeshore. During 1992–1995, herbicide treatment and sod removal decreased the combined cover of Poa pratensis (Kentucky blue grass) and Agropyron repens (quackgrass) significantly (from 82% to 13%, and 85% to 8%, respectively), whereas fire did not suppress such exotic lawn grasses. In 1993, several opportunistic species, represented by Cyperus spp. (umbrella sedges), Digitaria sanguinalis (crab grass), and Ambrosia artemisiifolia (common ragweed), filled the gaps left by the removal of lawn grasses. For the same period, Detrended Correspondence Analysis revealed a clear vegetation divergence between the control-fire plots and the herbicide-sod removal plots. While Poa pratensis and Agropyron repens continued to dominate the control and fire plots, the planted native species, represented by Schizachyrium scoparium (little blue-stem), Sorghastrum nutans (Indian grass), Rudbeckia hirta (black-eyed Susan), and Monarda punctata (horsemint), began to dominate in the herbicide and sod removal plots from 1994. In both herbicide and sod removal plots, the ground cover of grasses (68%) was much higher than the forbs (10%). The herbicide plots, where exotic species were removed but nitrogen-rich top soils were not removed, showed a higher diversity of planted native species than the sod removal plots (where both exotic species and top soils were removed) and the control-fire plots (where neither was removed). This finding suggests that an optimum but not excessive concentration of soil nitrogen is needed to support a maximum species diversity in such infertile substrate as sandy soil. In addition, the decrease in potassium in all plots, regardless of treatment, suggests that potassium may become a limiting factor for our restored native vegetation.     

Relationships between Methylobacteria and Glyphosate with Native and Invasive Plant Species: Implications for Restoration

After removing invasive plants, whether by herbicides or other means, typical restoration design focuses on rebuilding native plant communities while disregarding soil microbial communities. However, microbial–plant interactions are known to influence the relative success of native versus invasive plants. Therefore, the abundance and composition of soil microorganisms may affect restoration efforts. We assessed the effect of herbicide treatment on phytosymbiotic pink‐pigmented facultative methylotrophic (PPFM) bacteria and the potential consequences of native and invasive species establishment post‐herbicide treatment in the lab and in a coastal sage scrub (CSS)/grassland restoration site. Lab tests showed that 4% glyphosate reduced PPFM abundance. PPFM addition to seeds increased seedling length of a native plant (Artemisia californica) but not an invasive plant (Hirschfeldia incana). At the restoration site, methanol addition (a PPFM substrate) improved native bunchgrass (Nassella pulchra) germination and size by 35% over controls. In a separate multispecies field experiment, PPFM addition stimulated the germination of N. pulchra, but not that of three invasive species. Neither PPFM nor methanol addition strongly affected the growth of any plant species. Overall, these results are consistent with the hypothesis that PPFMs have a greater benefit to native than invasive species. Together, these experiments suggest that methanol or PPFM addition could be useful in improving CSS/grassland restorations. Future work should test PPFM effects on additional species and determine how these results vary under different environmental conditions.     

Influence of temperature on alkaloid levels and fall armyworm performance in endophytic tall fescue and perennial ryegrass

The symbiotic relationships between Neotyphodium endophytes (Clavicipitacea) and certain cool‐season (C3) grasses result in the synthesis of several alkaloids that defend the plant against herbivory. Over a 3 month period we evaluated the effects of temperature on the expression of these alkaloids in tall fescue, Festuca arundinacea Schreb, and perennial ryegrass, Lolium perenne L. (Poaceae). Response surface regression analysis indicated that month, temperature, and their interaction had an impact on the alkaloid levels in both grasses. We aimed to identify the alkaloids most closely associated with enhanced resistance to the fall armyworm, Spodoptera frugiperda JE Smith (Lepidoptera: Noctuidae), and clarify the role of temperature in governing the expression of these alkaloids. The dry weights and survival of fall armyworms feeding on endophyte‐infected tall fescue or perennial ryegrass were significantly lower than for those feeding on uninfected grass, whereas endophyte infection had no significant influence on survival. For tall fescue, a four‐alkaloid model consisting of a plant alkaloid, perloline, and the fungal alkaloids ergonovine chanoclavine, and ergocryptine, explained 47% of the variation in fall armyworm dry weight, whereas a three‐alkaloid model consisting of the plant alkaloid perloline methyl ether and the fungal alkaloids ergonovine and ergocryptine explained 70% of the variation in fall armyworm dry weight on perennial ryegrass. Although temperature had a significant influence on overall alkaloid expression in both grasses, the influence of temperature on individual alkaloids varied over time. The levels of those alkaloids most closely linked to armyworm performance increased linearly or curvilinearly with increasing temperature during the last 2 months of the study. We conclude that the growth temperature of grasses can influence the performance of fall armyworm, and that this effect may be mediated through a set of plant‐ and endophyte‐related alkaloids.     

Fire Season Effects on Flowering Characteristics and Germination of Longleaf Pine (Pinus palustris) Savanna Grasses

Pinus palustris (longleaf pine) savannas depend on contiguous grass cover to facilitate frequent surface fires. Enhanced flowering, seed production, and germination may be linked to season of fires. We assessed the effect of month of prescribed fire (February, April, May, and July) on percentage of plants with flowering culms (FCs%), FC/area, FC/plant, seed production, and germination for five warm seasons, fall‐flowering grasses. Multivariate analysis indicated the response of flowering and fruiting to burn month varied among the grass species. The dominant species, Sporobolus junceus and Schizachyrium scoparium var. stoloniferum, had greater numbers of most flowering characteristics when burnt during April, May, and July. Aristida purpurascens had increased FC/plant after May and July burns. In contrast, Ar. mohrii had the fewest FC/plant and seeds/FC when burnt in July. Germination was greatest (26–60%) for Ar. purpurascens. Seeds collected following July burns for Ar. purpurascens and Ar. ternarius and after May burns for S. junceus were within the highest germination values recorded. Germination of Sc. scoparium var. stoloniferum was very low after February and July burns (≤5%). With Ar. mohrii, only seed collected following February (2%) and April burns (3%) germinated. April, May, and July fires increased seed production of dominant matrix grasses, thus facilitating the potential for recruitment of these species and facilitating seed collection from potential donor sites for ground‐layer restoration projects. Varying prescribed fire burn month captured variation in flowering characteristics among these grasses.     

Savannas after afforestation: Assessment of herbaceous community responses to wildfire versus native tree planting

Afforestation and fire exclusion are pervasive threats to tropical savannas. In Brazil, laws limiting prescribed burning hinder the study of fire in the restoration of Cerrado plant communities. We took advantage of a 2017 wildfire to evaluate the potential for tree cutting and fire to promote the passive restoration of savanna herbaceous plant communities after destruction by exotic tree plantations. We sampled a burned pine plantation (Burned Plantation); a former plantation that was harvested and burned (Harvested & Burned); an unburned former plantation that was harvested, planted with native trees, and treated with herbicide to control invasive grasses (Native Tree Planting); and two old-growth savannas which served as reference communities. Our results confirm that herbaceous plant communities on post-afforestation sites are very different from old-growth savannas. Among post-afforestation sites, Harvested & Burned herbaceous communities were modestly more similar in composition to old-growth savannas, had slightly higher richness of savanna plants (3.8 species per 50-m²), and supported the greatest cover of native herbaceous plants (56%). These positive trends in herbaceous community recovery would be missed in assessments of tree cover: whereas canopy cover in the Harvested & Burned site was 6% (less than typical of savannas of the Cerrado), the Burned Plantation and Native Tree Planting supported 34% and 19% cover, respectively. By focusing on savanna herbaceous plants, these results highlight that tree cutting and fire, not simply tree planting and fire exclusion, should receive greater attention in efforts to restore savannas of the Cerrado.     

Effects of Prescribed Fires on Young Valley Oak Trees at a Research Restoration Site in the Central Valley of California

Woodland restoration sites planted with Quercus lobata (valley oak) often have serious invasions of nonnative annual grasses and thistles. Although prescribed fire can effectively control these exotics, restoration managers may be reluctant to use fire if it causes substantial mortality of recently planted saplings. We studied the effects of prescribed fires on the survival and subsequent growth of 5‐ and 6‐year‐old valley oak saplings at a research field near Davis, California. One set of blocks was burned in summer 2003 at a time that would control yellow star thistle, a second set of blocks was burned in spring 2004 at a time that would control annual grasses, and a third set was left unburned. Very few oaks died as a result of either fire (3–4%). Although a large proportion was top‐killed (66–72%), virtually all these were coppiced and most saplings over 300 cm tall escaped top‐kill. Tree height, fire temperature, and understory biomass were all predictive of the severity of sapling response to fire. Although the mean sapling height was initially reduced by the fires, the growth rates of burned saplings significantly exceeded the growth rates of unburned control trees for 2 years following the fires. By 2–3 years after the fires, the mean height of spring‐ and summer‐burned saplings was similar to that of the unburned control saplings. The presence of valley oak saplings does not appear to preclude the use of a single prescribed burn to control understory invasives, particularly if saplings are over 300 cm tall.