Habitat and landscape effects on abundance of Missouri's grassland birds

Of 6 million ha of prairie that once covered northern and western Missouri, <36,500 ha remain, with planted, managed, and restored grasslands comprising most contemporary grasslands. Most grasslands are used as pasture or hayfields. Native grasses largely have been replaced by fescue (Festuca spp.) on most private lands (almost 7 million ha). Previously cropped fields set aside under the Conservation Reserve Program (CRP) varied from a mix of cool-season grasses and forbs, or mix of native warm-season grasses and forbs, to simple tall-grass monocultures. We used generalized linear mixed models and distance sampling to assess abundance of 8 species of breeding grassland birds on 6 grassland types commonly associated with farm practices in Missouri and located in landscapes managed for grassland-bird conservation. We selected Bird Conservation Areas (BCAs) for their high percentage of grasslands and grassland-bird species, and for <5% forest cover. We used an information-theoretic approach to assess the relationship between bird abundance and 6 grassland types, 3 measures of vegetative structure, and 2 landscape variables (% grassland and edge density within a 1-km radius). We found support for all 3 levels of model parameters, although there was less support for landscape than vegetation structure effects likely because we studied high-percentage-grassland landscapes (BCAs). Henslow's sparrow (Ammodramus henslowii) counts increased with greater percentage of grassland, vegetation height-density, litter depth, and shrub cover and lower edge density. Henslow's sparrow counts were greatest in hayed native prairie. Dickcissel (Spiza americana) counts increased with greater vegetation height-density and were greatest in planted CRP grasslands. Grasshopper sparrow (A. savannarum) counts increased with lower vegetation height, litter depth, and shrub cover. Based on distance modeling, breeding densities of Henslow's sparrow, dickcissel, and grasshopper sparrow in the 6 grassland types ranged 0.9–2.6, 1.4–3.2, and 0.1–1.5 birds/ha, respectively. We suggest different grassland types and structures (vegetation height, litter depth, shrub cover) are needed to support priority grassland-bird species in Missouri. © 2011 The Wildlife Society.     

Recent decline of the critically endangered Plains‐wanderer (Pedionomus torquatus), and the application of a simple method for assessing its cause: major changes in grassland structure

A monitored population of the critically endangered Plains‐wanderer (Pedionomus torquatus) on Victoria's Northern Plains declined by over 90% between 2010 and 2012 following an unusually wet year which led to flooding, excess grass growth and a major change in the structure of native grasslands. The Plains‐wanderer population remained very low on private land during 2013 and 2014 when dry conditions prevailed and domestic stock overgrazed most of its favoured grasslands on red soil. Numbers also remained very low on public reserves despite grassland structure gradually improving there by 2014. In 2015, the population partially recovered in some grasslands protected from overgrazing. Grassland structure is critically important for Plains‐wanderer conservation. The ‘golf ball technique’ proved to be a quick and effective method for measuring grassland structure; it offers a means of accelerating responses to habitat change because it can be easily used by land managers.     

Comparison of removal‐based methods for estimating abundance of five species of prairie songbirds

Estimating species abundance is important for land managers, especially for monitoring conservation efforts. The two main survey methods for estimating avian abundance are point counts and transects. Previous comparisons of these two methods have either been limited to a single species or have not included detection probability. During the 2012 breeding season, we compared and assessed the efficiency (precision for amount of effort) of point count time of detection (PCTD) and dependent double‐observer transect (TRMO) methods based on detection probabilities and abundance estimates of five species of songbirds that use a range of habitats in a prairie system in Montana dominated by sagebrush and grassland vegetation. Our focal species included Vesper Sparrows (Pooecetes gramineus), a generalist species found in both shrub and grassland habitat, shrub‐obligate Brewer's Sparrows (Spizella breweri), and McCown's Longspurs (Rhynchophanes mccownii), Horned Larks (Eremophila alpestris), and Western Meadowlarks (Sturnella neglecta), three species of grassland obligates that prefer different grass heights. Detection probabilities were significantly higher for TRMO surveys, with less variation for all five species and differences most pronounced for Brewer's Sparrows and Horned Larks. PCTD surveys required less field effort (~8–20 fewer people minutes per plot) than TRMO surveys because the TRMO surveys required two people. However, time spent on TRMO surveys provided between 0.38 and 87 times more precision per people minute than PCTD surveys. Our results suggest that TRMO surveys provide a more efficient (measured as time spent per unit of standard error) field‐based technique in sagebrush prairie systems for the species we investigated, resulting in more precise detection and abundance estimates.     

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.     

Abundance,territory sizes,and pairing success of male Henslow's Sparrows in restored warm‐ and cool‐season grasslands

Henslow's Sparrows (Ammodramus henslowii) are known to breed in restored grasslands consisting of either warm‐ or cool‐season grasses, but additional information is needed concerning their breeding biology in these two types of grasslands. We compared the abundance, territory sizes, and pairing success of male Henslow's Sparrows in grasslands in west‐central Missouri in 2010 and 2011 using a paired‐treatment design, where warm‐ and cool‐season grasslands were located in close proximity. Selection indices indicated no apparent preference by male Henslow's Sparrows for either type of grassland, and the territory sizes and pairing success of males in cool‐ and warm‐season grasslands did not differ. In addition, we found no significant differences in habitat structure between these grassland types. Thus, our results suggest that the warm‐ and cool‐season grasslands in our study provided suitable habitat for Henslow's Sparrows. Other investigators have drawn similar conclusions for Henslow's Sparrows and for grassland birds in general, with the structural characteristics and, for some species, the size of grasslands being more important for grassland birds than plant species composition.     

Multi-scale impacts of crested wheatgrass invasion in mixed-grass prairie

Most of North America’s northern Great Plains have been cultivated for crop production, leaving remnants of natural mixed-grass prairie fragmented and threatened by alien plant invasions. The region’s most widespread alien perennial forage crop, crested wheatgrass (Agropyron cristatum sensu amplo), has invaded native grassland and raised concerns regarding its ecological impact. To evaluate impacts at multiple scales of organization, adjacent invaded and uninvaded mixed-grass prairie were sampled at eight widely separated locations. At the population level, native C₃ mid-grasses and forbs were less abundant in invaded grasslands, while native C₃ and C₄ short-grass abundance was not different. At community and landscape levels, diversity was lower in invaded grasslands largely because of lower forb species richness and cover, and crested wheatgrass dominance of both cover (14% basal cover) and seedbank (404 seeds m⁻²). At the ecosystem level, both vegetation and litter biomass were greater in invaded grasslands, however, below ground organic matter (roots and litter), soil organic carbon, total nitrogen and phosphorus were not different. Crested wheatgrass invasion of mixed-grass prairie was associated with lower diversity within and among plant communities, and appears to simplify the composition of mixed-grass prairie landscapes. Hypotheses for crested wheatgrass dominance and persistence following invasion are suggested.     

Burning effects on detritivory and litter decay in Campos grasslands

Disturbances are primary forces creating spatial heterogeneity in ecosystems, and inducing changes on biological communities, abiotic characteristics and ecological processes. Here we focus on the effects of fire disturbance in the decomposition process at subtropical Campos grasslands in South Brazil, where burns are traditionally used to reduce shrub encroachment, and improve forage quality. We experimentally investigated how burns and the changes they produce in grassland habitat conditions affect soil fauna detritivory and surface leaf‐litter decaying patterns over one year. Previously to fire, we found significant correlation of litter decay with plant evenness and detritivory rates in non‐disturbed grasslands. One month after fire grassland patches presented reduced soil fauna densities and surface feeding activity possible because of the mortality caused directly by heating, and/or due to harsh microenvironmental filters to fauna colonization and permanency (e.g. decreased humidity). At 6–7 months after fire however these features did not differ any more from the paired unburned plots. On the other hand, canopy openness accelerated the decaying of leaf‐litter in burned patches by allowing increased action of abiotic factors as solar radiation potentially triggering photodegradation. These effects seemed to last less than one year. Overall, our results bring insights regarding drivers of soil ecological processes at local scales in subtropical grasslands, and suggest that detritivory and litter decay processes are sensitive to fire, but resilient following grassland recovering.     

Invasion by <Emphasis Type="Italic">Aegilops triuncialis</Emphasis> (Barb Goatgrass) Slows Carbon and Nutrient Cycling in a Serpentine Grassland

Invasive plant species alter plant community composition and ecosystem function. In the United States, California native grasslands have been displaced almost completely by invasive annual grasses, with serpentine grasslands being one of the few remaining refugia for California grasslands. This study examined how the invasive annual grass, Aegilops triuncialis, has altered decomposition processes in a serpentine annual grassland. Our objectives were to (1) assess howA. triuncialis alters primary productivity and litter tissue chemistry, (2) determine whether A. triuncialis litter is more recalcitrant to decomposition than native litter, and (3) evaluate whether differences in the soil microbial community in A. triuncialis-invaded and native-dominated areas result in different decomposition rates of invasive and/or native plant litter. In invaded plant patches, A. triuncialis was approximately 50% of the total plant cover, in contrast to native plant patches in which A. triuncialis was not detected and native plants comprised over 90% of the total plant cover. End-of-season aboveground biomass was 2-fold higher in A. triuncialis dominated plots compared to native plots; however, there was no significant difference in belowground biomass. Both above- and below-ground plant litter from A. triuncialis plots had significantly higher lignin:N and C:N ratios and lower total N, P, and K than litter from native plant plots. Aboveground litter from native plots decomposed more rapidly than litter from A. triuncialis plots, although there was no difference in decomposition of belowground tissues. Soil microbial community composition associated with different soil patch types had no effect on decomposition rates. These data suggest that plant invasion impacts decomposition and nutrient cycling through changes in plant community tissue chemistry and biomass production.     

Dominance of an invasive earthworm in native and non-native grassland ecosystems

More attention is currently being focused on earthworm invasions; however, in many ecosystems the relative abundance of native and invasive earthworm species is unknown. We characterized earthworm populations of two grassland types within the Palouse region: native prairie remnants and Conservation Reserve Program (CRP) set asides planted with exotic grasses. The earthworm community in both grassland types was completely dominated by the exotic-invasive Aporrectodea trapezoides. Only one individual of a native species, Driloleirus americanus (the giant Palouse earthworm), was found in a prairie remnant. No differences were found between prairie remnants and CRP sites for mean earthworm density (24–106 individuals m⁻²) or fresh weight (12–45 g m⁻²). Our results suggest that the combined effects of land-use change, habitat fragmentation and competitive interactions have resulted in the decimation of native earthworm populations and dominance of invasive earthworms in native and non-native grasslands of the Palouse region.     

Reducing Competitive Suppression of a Rare Annual Forb by Restoring Native California Perennial Grasslands

Populations of the rare annual forb Amsinckia grandiflora may be declining because of competitive suppression by exotic annual grasses, and may perform better in a matrix of native perennial bunchgrasses. We conducted a field competition experiment in which Amsinckia seedlings were transplanted into forty 0.64‐m² experimental plots of exotic annual grassland or restored perennial grassland. The perennial grassland plots were restored using mature 3 cm‐diameter plants of the native perennial bunchgrass Poa secunda planted in three densities. The exotic annual grassland plots were established in four densities through manual removal of existing plants. Both grass types reduced soil water potential with increasing biomass, but this reduction was not significantly different between grass types. Both grass types significantly reduced the production of Amsinckia inflorescences. At low and intermediate densities (dry biomass per unit area of 20–80 g/m²), the exotic annual grasses reduced Amsinckia inflorescence number to a greater extent than did Poa, although at high densities (>90 g/m²) both grass types reduced the number of Amsinckia inflorescences to the same extent. The response of Amsinckia inflorescence number to Poa biomass was linear, whereas the same response to the annual grass biomass is logarithmic, and appeared to be related to graminoid cover. This may be because of the different growth forms exhibited by the two grass types. Results of this research suggest that restored native perennial grasslands at intermediate densities have a high habitat value for the potential establishment of the native annual A. grandiflora.     

Contrasting needs of grassland dwellers: habitat preferences of endangered steppe beetles (Coleoptera)

Temperate grasslands are local biodiversity hotspots. In Europe, their extent was mostly reduced to isolated habitat patches, whose biota is subject to extinction debt. Knowledge on requirements of dry-grassland inhabitants is thus vital to slow down decline of grassland biodiversity. We studied habitat requirements of eight flightless steppe beetles, including some of the most endangered dry-grassland specialists of the continent. The beetles were sampled using 167 pitfall traps at a pannonian dry-grassland fragment, the Pouzdrany steppe, SE Czech Republic, from March to November 2006. The number of each species captures in each trap was related to vegetation and abiotic habitat characteristics; captures of all beetles were related to each other. Two of the studied species required relatively humid microhabitats, including tall-grass steppe with litter (Carabus hungaricus, Carabidae) and grassland of high herb cover (Meloe proscarabaeus, Meloidae). Others were associated with xeric habitats (e.g. Meloe scabriusculus) and their early-successional stages, including short-turf vegetation (Dorcadion fulvum, D. pedestre, Cerambycidae) and/or bare-ground patches (Blaps lethifera, Tenebrionidae; Meloe decorus, M. uralensis). Our findings point to key importance of early-successional vegetation for grassland biodiversity, and to the fact that locally co-occurring and closely related grassland specialists may exhibit contrasting habitat needs. Spatially and temporarily highly diversified patch management creating a fine scale mosaic of various seral stages from bare soil to tall-grass steppe is therefore the most appropriate approach for managing isolated grasslands. Prescribed burning and support of burrowing herbivores are recommended and discussed together with other measures for restoration of habitat diversity in dry-grassland fragments.     

Fine-scale habitat associations of Oklahoma's longspurs

Overwintering is a key demographic stage for migratory birds but remains poorly understood, especially among multiple declining grassland bird species. The non-breeding ranges all 4 species of longspur (i.e., chestnut-collared [Calcarius ornatus], Smith's [C. pictus], Lapland [C. lapponicus], thick-billed [Rhynchophanes mccownii]) overlap in Oklahoma and the Texas Panhandle, USA, making this region ideal to study their wintering ecology. We evaluated the relationship between wintering longspur occurrence and fine-scale habitat characteristics using a combination of standardized bird surveys and vegetation plot sampling. Our study encompassed large, representative tracts of 3 prairie ecosystems (i.e., shortgrass, mixed-grass, and tallgrass prairies) that intersect within the Southern Great Plains, during winters of 2018–2019 and 2019–2020. Using randomization tests and classification trees, we characterized longspur habitats and compared these associations across the 3 prairie ecosystems. Fine-scale winter habitats (horizontal structure, vertical structure, and species compositions) varied among all 4 longspur species, varied at very fine scales, and differed between grassland types. Our findings can be applied to the management of grasslands such as decreasing vegetation height in mixed-grass prairies for chestnut-collared longspurs or removing woody vegetation in shortgrass prairies for thick-billed longspurs to help develop full-life cycle conservation for longspurs, which have experienced population declines.     

Regional analysis of litter quality in the central grassland region of North America

Abstract. The central grassland region of North America is characterized by large gradients of temperature and precipitation. These climatic variables are important determinants of the distribution of plant species, and strongly influence plant morphology and tissue chemistry. We analysed regional patterns of plant litter quality as they vary with climate in grassland ecosystems throughout central North America including tall‐grass prairie, mixed grass prairie, shortgrass steppe, and hot desert grasslands. An extensive database from the International Biological Program and the Long‐Term Ecological Research Program allowed us to isolate the effects of climate from those of plant functional types on litter quality. Our analysis of grass species confirms a previously recognized positive correlation between C/N ratios and precipitation. Precipitation exhibited a similar positive relationship with lignin/N and percent lignin. Although there was no significant correlation between temperature and C/N, there was a significant positive relationship between temperature and both percent lignin and lignin/N. Among functional types, C₃ grasses had a slightly lower C/N ratio than C₄ grasses. Tall grass species exhibited higher C/N, lignin/N, and percent lignin than short grass species. This understanding of the regional patterns of litter quality and the factors controlling them provides us with a greater knowledge of the effect that global change and the accompanying feedbacks may have on ecosystem processes.     

Conservation planning in an agricultural landscape: the case of Sharpe's Longclaw

Lens, L., Muchai, M., Bennun, L.A. & Duchateau, L. 2000. Conservation planning in an agricultural landscape: the case of Sharpe's Longclaw. Ostrich 71 (1 & 2): 300–303.

We studied the distribution and habitat use of Sharpe's Longclaw, Macmnyx sharpei, a Kenyan montane grassland endemic, in a fragmented agricultural landscape at two spatial scales. Monthly counts in a fixed number of plots selected in contrasting habitat types provided insight in the species' distribution in relation to the available habitat. Simultaneously, focused ecological work in a smaller area provided information on its behaviour, movements and habitat choice, and allowed us to interpret the wider distribution patterns in a biologically meaningful way. Sharpe's Longclaw occurs only in grasslands, not in cultivated fields or plantations of exotic trees. It prefers short grassland with a well-developed tussock structure, apparently because tussocks provide good cover while foraging and nesting. However, as tussock grass is unpalatable to livestock, this grassland type is being ploughed up, re-seeded and converted to cultivation at an alarming rate, and the remaining patches are becoming ever more isolated from each other. Although only limited information on the species' population dynamics and dispersal ability is available, there is a clear need to conserve and manage the remaining natural grassland habitat. Since most of the land within the species' distribution range is privately owned, keeping viable populations of Sharpe's Longclaw may be extremely difficult.     

Reintroducing a keystone burrowing rodent to restore an arid North American grassland: challenges and successes

Prairie dogs (Cynomys spp.) are important ecosystem engineers in North America's central grasslands, and are a key prey base for numerous predators. Prairie dogs have declined dramatically across their former range, prompting reintroduction efforts to restore their populations and ecosystem functions, but the success of these reintroductions is rarely monitored rigorously. Here, we reintroduced 2,400 Gunnison's prairie dogs (C. gunnisoni) over a period of 6 years to the Sevilleta National Wildlife Refuge, in central New Mexico, U.S.A., a semi‐arid grassland ecosystem at the southern edge of their range. We evaluated the population dynamics of prairie dogs following their reintroduction, and their consequent effects on grassland vertebrates. We found postrelease survival of prairie dogs stabilized at levels typical for the species (ca. 50%) after approximately 1 month, while average annual recruitment was ca. 0.35 juveniles per female, well below what was required for a self‐sustaining, stable population. Extreme drought conditions during much of the study period may have contributed to low recruitment. However, recruitment increased steadily over time, indicating that the reintroduced colony may simply need more time to establish in this arid system. We also found well‐known associates of prairie dog colonies, such as American badgers (Taxidea taxus) and burrowing owls (Athene cunicularia), were significantly more common on the colonies than off. After 7 years, we have yet to meet our goal of establishing a self‐sustaining population of Gunnison's prairie dogs in this semi‐arid grassland. But despite the uncertainty and challenges, our work shows that reestablishing keystone species can promote ecosystem restoration.     

Sprague's pipit breeding biology and reproductive success in planted and native grasslands

Much of the native grasslands in agricultural regions have been converted to cropland or tilled and seeded with non‐native grasses for livestock production. Several grassland songbird species occupy planted grasslands, but occupancy or density may not be a reliable indicator of habitat quality. I studied the breeding biology of Sprague's pipit Anthus spragueii from 2004 to 2008 in Saskatchewan, Canada. My objective was to determine the extent to which the breeding biology, density and reproductive success of pipits varied in planted and native grasslands. Peak clutch initiation occurred in mid‐ to late‐May in planted and native grassland. Peak pipit density also occurred in May, but density drastically declined over the breeding season in planted grassland. Clutch size varied among years and declined over the breeding season, but was similar in planted (4.7  0.1 SE) and native grasslands (4.5  0.1 SE). Daily nest survival rates varied with age of the nest and date, but the relationships differed in the two habitats and was likely a result of lower nestling survival in planted grassland compared to native grassland. The number of young fledged per nest increased as the season progressed and tended to be greater in native (1.2  0.1 SE) than planted (0.9  0.2 SE) grasslands. Seasonal productivity was much greater in native grassland. Only three nests were initiated after May in planted grassland and all were unsuccessful, whereas pipit young fledged at higher rates from nests initiated in native grassland in June and July than planted grassland nests initiated in May. The number of fledged young from successful nests did not vary strongly with habitat, date or year. This research indicates that planted grasslands attract pipits at the beginning of the breeding season, but habitat suitability and reproductive success substantially declines as the breeding season progresses compared to that found in native grassland.     

Microhabitat use, giving-up densities and temporal activity as short- and long-term anti-predator behaviors in common voles

We used depletable food patches to determine the effect of microhabitat (mowed versus unmowed adjacent grasslands) and time (day versus night) on the foraging behavior of common voles (Microtus arvalis). The food remaining after 12‐h periods (giving‐up density, GUD) measured the vole's habitat selection under predation risk. In accord with several other rodent species and the effects of avian predators, voles had significantly lower GUDs in the unmowed than mowed portion of the grassland. GUDs in patches along the border between adjacent habitats were more similar to the risky mowed grassland than the safe unmowed grass. Time interacted strongly with microhabitat. In the mowed grass, voles had significantly higher GUDs at night than day. Whereas in the unmowed grass, GUDs were significantly higher during the day than night. Vole GUDs did not vary with time along the boundary. This suggests that predators are more abundant or effective in the mowed grass at night (owls?), and in the unmowed grass during the day (weasels?). In terms of predation risk, the voles perceived the mowed grass at night as the riskiest and the unmowed grass at night as the safest. Voles may have difficulties assessing resources under high predation risk: GUDs among patches were well equalized in the unmowed microhabitat whereas in the mowed grass only day GUDs did not vary significantly among patches. We linked these results to the vole's day‐night‐activity and life span. For the 533 voles live‐trapped at the study area, the ratio of day versus night captures for each individual served as an activity index and the span between first and last capture measured minimum life span. In accord with higher GUDs at night, very few individuals behaved selectively towards the night, but individual life expectancy increased with temporal opportunism. Microhabitat differences in GUDs reflect short‐term strategies of predator avoidance and the trapping data reflect long‐term patterns of anti‐predator behavior.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

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.     

The heat is on: frequent high intensity fire in bracken (<Emphasis Type="Italic">Pteridium aquilinum</Emphasis>) drives mortality of the sprouting tree <Emphasis Type="Italic">Protea</Emphasis><Emphasis Type="Italic">caffra</Emphasis> in temperate grasslands

We examined the effect of fire frequency and intensity on a Protea caffra tree population in the temperate montane grasslands of north-western KwaZulu-Natal, South Africa. We assessed the effect of fire by comparing the population structure of the resprouter P. caffra in discrete bracken (Pteridium aquilinum) patches with that in the surrounding grassland matrix. Fuel biomass did not differ between grassland and bracken, but bracken fuel was significantly drier than grass. Above-ground fire temperatures and fireline intensity, measured by P. caffra char height, were significantly higher in the bracken habitat. Forty-two percent of the P. caffra population in grassland and in bracken persisted by coppice resprouts, having lost their original stem to fire damage. Exposure to higher intensity bracken fire suppressed P. caffra regeneration and caused greater adult mortality compared with trees in grassland. Consequently, the P. caffra population in bracken was skewed towards old age with most trees severely fire damaged. The high incidence of small trees in grassland indicates that a regular fire interval of 2–3 years does not negatively affect regeneration of P. caffra. However, in bracken patches regular high intensity fires cause high mortality among all P. caffra size classes and will ultimately result in local extinction. Bracken thus has the potential to significantly alter tree–grass interactions in these montane grasslands.