Evidence for indirect effects of plant diversity and composition on net nitrification

Abiotic controls on net nitrification rates are well documented, but the potential effects of plants on this important ecosystem process are poorly understood. We evaluated four structural equation models to determine the relative importance of plant community composition, aboveground herbaceous production, and plant species richness on nitrifier abundance and net nitrification following restoration treatments in a ponderosa pine forest. Model selection criteria indicated that species richness was the best predictor of nitrifier abundance, but a model that included community composition effects also had some support in the data. Model results suggest that net nitrification was indirectly related to plant species richness via a positive relationship between species richness and nitrifier abundance. Community composition was indirectly related to nitrifier abundance through its relationship with species richness. Our model indicates that species-rich plant communities dominated by C₃ graminoids and legumes are associated with soils that have high abundances of nitrifiers. This study highlights the complexity of deciphering effects of ecological treatments on a system response when multiple interacting factors are simultaneously affected. Our results suggest that plant diversity and composition can both respond to forest thinning, prescribed fire and fuel manipulations, and can be factors that might indirectly influence an ecosystem process such as nitrification. Ecological restoration treatments designed to increase plant diversity and alter community composition may have cascading effects on below-ground processes.     

Short‐term bryoid and vascular vegetation response to reforestation alternatives following wildfire in conifer plantations

Question: How are dynamics of early‐seral post‐fire vascular plant and bryoid (terrestrial mosses, lichens, and fungi) vegetation impacted by reforestation activities, particularly manual vegetation removal and planting density? Does the relationship between vegetation dynamics and vegetation removal differ between harsh (west‐facing) and moderate (east‐facing) aspects? Location: Five high‐severity burn plantation forests of Pseudotsuga menziesii in southwestern Oregon, USA. Methods: Plantations severely burned in a recent wildfire were planted with conifer seedlings as a four‐species mixture or a monoculture, at two different densities, with and without manual vegetation removal. A subset of plots was also planted on a contrasting aspect within each plantation. The contrasting aspects differed in potential solar insolation and were indicative of moderate (eastern exposure) and harsh (western exposure) site conditions. Covers of shrub, herbaceous and bryoid vegetation layers were measured during reforestation activities 2–4 yr after the fire. Dynamics of structural layer cover and community composition were compared among treatments with analysis of variance and multivariate analyses (non‐metric multidimensional scaling and blocked multi‐response permutation procedure). Results: Structural layer cover and community composition differed between areas that received reforestation treatments and untreated areas. However, variability within treatments in a plantation was greater than variability within treatments across plantations. Effects of vegetation removal on composition and structure were more evident than effects of planting or altering planting density. Vegetation removal decreased cover of tall and low shrub and the bryoid layer, and increased herbaceous layer cover. Bryoid community and low shrub structural layer responses were more pronounced on moderate aspects than on harsh aspects. Vegetation removal shifted vascular plant community composition towards exotic and annual species. Conclusions: These reforestation treatments may be implemented without substantially altering early‐seral vegetation community composition dynamics, especially in areas with harsh site conditions. Site conditions, such as aspect, should be evaluated to determine need and potential effects of reforestation before implementation. Monitoring for exotic species establishment should follow reforestation activities.     

Restoring plant species diversity and community composition in a ponderosa pine-bunchgrass ecosystem

Monitoring of ecological restoration treatments often focuses on changes in community structure and function. We suggest that long-term changes in community composition also need to be explicitly considered when evaluating the success of restoration treatments. In 1992, we initiated an experiment in a ponderosa pine-bunchgrass ecosystem to evaluate responses to restoration treatments: (a) thinning the overstory vegetation (‘thinning’), (b) thinning plus forest floor manipulation with periodic prescribed burning (‘composite’), and (c) untreated ‘control.’ Treatments were further stratified by forest patch type: presettlement tree clumps (trees that established prior to the onset of fire exclusion in 1876), patches of retained postsettlement trees, patches where all postsettlement trees were removed, and remnant grass openings. Species richness did not differ among treatments for 10 years, but was highest in the composite treatment in 11th and 12th year after initial treatment. Community composition diverged among treatments 5 years after initial treatment, and compositional changes were greatest in the composite treatment. Species richness and composition differed among patch types prior to treatment. Remnant grass patches were the most diverse and presettlement patches were the least diverse. Following treatment, species richness in the postsettlement removed and retained patches, gradually approached levels found in remnant grass patches. Compositional differences among patch types changed a little by 2005. Species richness at the 2 m² scale increased only where the overstory was thinned and the understory was burned. However, these changes may not be detectable for many years, and can vary temporally in response to events such as severe droughts. Nonnative species establishment may be reduced by scheduling longer burn intervals or by refraining from burning where fuel loads are not hazardous, though these options may hinder goals of increasing diversity. Restoring species diversity and community composition continues to be more difficult than restoring ecosystem structure and function.     

Diverse responses across soil parent materials during ecological restoration

A major challenge to advancing the science and practice of ecological restoration is working across large landscapes containing diverse sites that may respond differently to restoration. We conducted a 5‐year restoration experiment, replicated across 9 sites spanning 3 soil parent material types within a 9,000‐ha Pinus ponderosa forest landscape. We evaluated plant community response to restoration Pinus thinning, grazing, and aqueous smoke application. We measured vegetation before (2003) and 3 (2006) and 5 (2008) years after treatment. Plant community responses of species richness, cover, and composition were diverse, ranging from increases, decreases, or no change depending on soil parent material, tree thinning, and presence or exclusion of grazing. Restoration outcomes were under hierarchical control: soil parent material constrained response to Pinus thinning, which in turn influenced grazing effects. On limestone‐derived soil, responses included no change in species richness but increased plant cover with Pinus thinning. Both plant richness and cover increased on benmorite soil after thinning, and cover generally increased more without grazing. On rocky, basalt soil, plant richness increased but cover did not after any treatment. Diversity of responses to restoration has implications for: (1) setting goals or monitoring indicators tailored to inherent soil capability; (2) identifying where grazing most affects restoration outcomes; and (3) forecasting responses to restoration across landscapes. Diverse responses to restoration along physiographic gradients such as soil parent material warrant consideration when developing restoration across degraded landscapes.     

Chaparral shrub recovery after fuel reduction: a comparison of prescribed fire and mastication techniques

Fuel management techniques are commonly used in shrublands to reduce wildfire risk. However, more information about the ecological effects of these treatments is needed by managers and ecologists. In an effort to address this need, we performed a replicated (4 replicates per treatment) 48-ha experiment in northern California chaparral dominated by Adenostoma fasciculatum to determine the effects of two fuel reduction types (prescribed fire and mastication) and three different seasons of treatment (fall, winter, and spring) on shrub cover, height, and seedling density. Exclosures (2.5 m² each) were also used to assess herbivory effects. By the third post-treatment year, prescribed fire treatments had higher shrub cover (71 ± 2%) than mastication (43 ± 4%). There was no treatment effect on shrub height, species richness, or composition. Seedling density was initially higher in prescribed fire treatments (31 ± 4 seedlings m⁻²) than mastication (3 ± 0 seedlings m⁻²); however, prescribed fire treatments experienced substantial mortality, especially spring burning, resulting in lower densities 3 years after treatments (18 ± 0 seedlings m⁻² after fall and winter fire compared to 2 ± 0 seedlings m⁻² after spring fire). A. fasciculatum remained the dominant shrub species after the treatments, and Ceanothus cuneatus recruitment was higher in fall burning. Deer herbivory only affected shrub height, especially in masticated units, resulting in heights of 55 ± 2 cm in unexclosed areas compared to 66 ± 4 cm inside exclosures by the third post-treatment year. Overall, our findings suggest that fuel treatments play an important role in shrubland community dynamics, at least in the short-term, with implications for re-treatment frequency, community structure, and wildlife habitat.     

Plant community responses to thinning in densely regenerating Acacia harpophylla forest

Thinning is thought to be particularly effective at accelerating plant community recovery in some dense regenerating forests where one species dominates. However, longer‐term experimental data are required to confirm the efficacy of thinning for this purpose. We investigated plant community responses 8 years after thinning in densely resprouting brigalow (Acacia harpophylla) forests in Australia. We examined the abundance, diversity, composition, and size of all woody recruits, as well as the functional composition of woody plant assemblages using functional traits. The average prethinning density of A. harpophylla was 15,600 stems/ha. Four treatments were applied to 25 × 25 m plots in a randomized block design: control (no thinning) and random thinning (ringbarking) down to 4,000, 2,000, and 1,000 stems/ha. Reference plots were also established in a nearby mature forest. Thinning accelerated the recruitment of certain shrub species and increased some measures of diversity. However severe thinning led to an overabundance of exploitative shrub species (high‐specific leaf area and low stem‐specific density), akin to pioneer dominance during early stages of rainforest succession. Species with more conservative growth strategies have so far failed to recruit, potentially due to dispersal limitation. Longer‐term monitoring is required to determine if thinning does in fact accelerate compositional recovery in this endangered Australian ecosystem.     

Vegetation recovery after fuel management in Mediterranean shrublands

Question: What is the effect of fuel management practices in the recovery capacity of seeder‐dominated shrublands? Location: Ulex parviflorus shrubland localities in Mediterranean regions of eastern Spain. Methods: We applied prescribed burning and brush‐chipping as fuel management techniques in three young and three mature shrublands, and evaluated the effects in the following four years. Results: Canopy opening by the treatments allowed increasing species richness through the four years of secondary succession. The treatments produced a change in community structure and dominant species, from the woody seeder Ulex parviflorus to the resprouter grass Brachypodium retusum. Vegetation response was conditioned by both shrubland developmental stage and treatment applied. Burning resulted in more severe modification of the ecosystem, increasing bare soil cover. Four years after fuel management in different aged Mediterranean gorse shrublands, vegetation response followed a similar pattern with the exception of the young, brush‐chipped shrublands. The treatments applied for controlling Ulex parviflorus were seen to be very effective, with the exception of brush‐chipping in young shrublands. Conclusions: Selective brush‐chipping in middle‐aged or mature gorse shrublands would combine a drastic reduction in fire hazard with ecosystem conservation and regeneration.     

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

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

Understory vegetation response to mechanical mastication and other fuels treatments in a ponderosa pine forest

Questions: What influence does mechanical mastication and other fuel treatments have on: (1) canopy and forest floor response variables that influence understory plant development; (2) initial understory vegetation cover, diversity, and composition; and (3) shrub and non‐native species density in a second‐growth ponderosa pine forest. Location: Challenge Experimental Forest, northern Sierra Nevada, California, USA. Methods: We compared the effects of mastication only, mastication with supplemental treatments (tilling and prescribed fire), hand removal, and a control on initial understory vegetation response using a randomized complete block experimental design. Each block (n=4) contained all five treatments and understory vegetation was surveyed within 0.04‐ha plots for each treatment. Results: While mastication alone and hand removal dramatically reduced the midstory vegetation, these treatments had little effect on understory richness compared with control. Prescribed fire after mastication increased native species richness by 150% (+6.0 species m²) compared with control. However, this also increased non‐native species richness (+0.8 species m²) and shrub seedling density (+24.7 stems m²). Mastication followed by tilling resulted in increased non‐native forb density (+0.7 stems m²). Conclusions: Mechanical mastication and hand removal treatments aided in reducing midstory fuels but did not increase understory plant diversity. The subsequent treatment of prescribed burning not only further reduced fire hazard, but also exposed mineral soil, which likely promoted native plant diversity. Some potential drawbacks to this treatment include an increase of non‐native species and stimulation of shrub seed germination, which could alter ecosystem functions and compromise fire hazard reduction in the long‐term.     

Selective and Non-Selective Control of Invasive Plants: The Short-Term Effects of Growing-Season Prescribed Fire, Herbicide, and Mowing in Two Texas Prairies

Conservation of North American grasslands is hampered by the impact of invasive herbaceous species. Selective control of these plants, although desirable, is complicated by the shared physiology and phenology of the invader and the native components of the invaded plant community. Fortunately, there is evidence that some management practices, such as prescribed fire, herbicide, and mowing, can cause differential responses in native and invasive grassland species. However, timing of treatment is critical, and fire has been shown to increase rates of invasion when implemented during the dormant season. Bothriochloa ischaemum, an introduced C4 Eurasian grass is an increasing problem in grasslands, particularly in southern and central regions of North America. To date, there has been little success in effective selective control. Two invaded grassland sites representative of Blackland Prairie and Edwards Plateau ecoregions were subjected to two growing‐season prescribed fire treatments, single and double herbicide applications, and single and double mowing treatments. Mowing had no effect on either B. ischaemum or other dominant species at either site one‐year posttreatment. However, growing‐season fire and herbicide were both effective at reducing the abundance of B. ischaemum, with other codominant species responding either negatively to herbicide or neutrally or positively to fire. The vulnerability of B. ischaemum to growing‐season fire may be associated with the ecology of its native range. The negative growth response to growing‐season fire, combined with its lower implementation costs, indicates that this method warrants further investigation as a selective management tool for other problematic species in invaded grasslands.     

The effect of long‐term repeated burning and fire exclusion on above‐ and below‐ground Blackbutt (Eucalyptus pilularis) forest vegetation assemblages

We used a long‐term fire experiment in south‐east Queensland, Australia, to determine the effects of frequent prescribed burning and fire exclusion on understorey vegetation (<7.5 m) richness and density in Eucalyptus pilularis forest. Our study provided a point in time assessment of the standing vegetation and soil‐stored vegetation at two experimental sites with treatments of biennial burning, quadrennial burning since 1971–1972 and no burning since 1969. Vegetation composition, density and richness of certain plant groups in the standing and soil‐stored vegetation were influenced by fire treatments. The density of resprouting plants <3 m in height was higher in the biennially burnt treatment than in the unburnt treatment, but resprouters 3–7.5 m in height were absent from the biennial burning treatment. Obligate seeder richness and density in the standing vegetation was not significantly influenced by the fire treatments, but richness of this plant group in the seed bank was higher in the quadrennial treatment at one site and in the long unburnt treatment at the other site. Long unburnt treatments had an understorey of rainforest species, while biennial burning at one site and quadrennial burning at the other site were associated with greater standing grass density relative to the unburnt treatment. This difference in vegetation composition due to fire regime potentially influences the flammability of the standing understorey vegetation. Significant interactions between fire regime and site, apparent in the standing and soil‐stored vegetation, demonstrate the high degree of natural variability in vegetation community responses to fire regimes.     

Climate change effects on plant biomass alter dominance patterns and community evenness in an experimental old‐field ecosystem

Atmospheric and climatic change can alter plant biomass production and plant community composition. However, we know little about how climate change‐induced alterations in biomass production affect plant species composition. To better understand how climate change will alter both individual plant species and community biomass, we manipulated atmospheric [CO₂], air temperature, and precipitation in a constructed old‐field ecosystem. Specifically, we compared the responses of dominant and subdominant species to our climatic treatments, and explored how changes in plant dominance patterns alter community evenness over 2 years. Our study resulted in four major findings: (1) all treatments, elevated [CO₂], warming, and increased precipitation increased plant community biomass and the effects were additive rather than interactive, (2) plant species differed in their response to the treatments, resulting in shifts in the proportional biomass of individual species, which altered the plant community composition; however, the plant community response was largely driven by the positive precipitation response of Lespedeza, the most dominant species in the community, (3) precipitation explained most of the variation in plant community composition among treatments, and (4) changes in precipitation caused a shift in the dominant species proportional biomass that resulted in lower community evenness in the wet relative to dry treatments. Interestingly, compositional and evenness responses of the subdominant community to the treatments did not always follow the responses of the whole plant community. Our data suggest that changes in plant dominance patterns and community evenness are an important part of community responses to climatic change, and generally, that such compositional shifts can alter ecosystem biomass production and nutrient inputs.     

Assessing Ecosystem Restoration Alternatives in Eastern Deciduous Forests: The View from Belowground

Both structural and functional approaches to restoration of eastern deciduous forests are becoming more common as recognition of the altered state of these ecosystems grows. In our study, structural restoration involves mechanically modifying the woody plant assemblage to a species composition, density, and community structure specified by the restoration goals. Functional restoration involves reintroducing dormant‐season, low‐severity fire at intervals consistent with the historical condition. Our approach was to quantify the effects of such restoration treatments on soil organic carbon and soil microbial activity, as these are both conservative ecosystem attributes and not ones explicitly targeted by the restoration treatments, themselves. Fire, mechanical thinning, and their combination all initially resulted in reduced soil organic C content, C:N ratio, and overall microbial activity (measured as acid phosphatase activity) in a study site in the southern Appalachian Mountains of North Carolina, but only the effect on microbial activity persisted into the fourth post‐treatment growing season. In contrast, in a similar forest in the central Appalachian Plateau of Ohio, mechanical thinning resulted in increased soil organic C, decreased C:N ratio, and decreased microbial activity, whereas fire and the combination of fire and thinning did not have such effects. In addition, the effects in Ohio had dissipated prior to the fourth post‐treatment growing season. Mechanical treatments are attractive in that they require only single entries; however, we see no indication that mechanical–structural restoration actually produced desired belowground changes. A single fire‐based/functional treatment also offered little restoration progress, but comparisons with long‐term experimental fire studies suggest that repeated entries with prescribed fire at intervals of 3–8 years offer potential for sustainable restoration.     

Are long‐unburnt eucalypt forest patches important for the conservation of plant species diversity?

Question: Are long‐unburnt patches of eucalypt forest important for maintaining floristic diversity? Location: Eucalyptus forests of southeastern New South Wales, Australia. Methods: Data from 976 sites representing a range of fire history from three major vegetation formations – shrubby dry sclerophyll forest (SF), grassy dry SF and wet SF – were analysed. Generalized linear models were used to examine changes in species richness with increasing time since wildfire and analysis of similarities to examine changes in community composition. Chi‐squared tests were conducted to examine the distribution of individual species across four time since fire categories. Results: Plant species relationships to fire varied between the three formations. Shrubby dry SF supported lower plant species richness with increasing time since wildfire and this was associated with shifts in community composition. Grassy dry SF showed significant shifts in community composition and species richness in relation to time, with a peak in plant species richness 20–30 yr post fire (either prescribed fire or wildfire). Wet SF increased in species richness until 10–20 yr post wildfire then displayed a general declining trend. Species richness in each vegetation type was not related to the fire frequencies and fire intervals observed in this study. Conclusions: Long‐unburnt (30–50 yr post wildfire) forests appeared to play a minor role in the maintenance of plant species diversity in dry forest systems, although this was more significant in wet forests. Maintenance of a range of fire ages within each vegetation formation will assist in maintaining floristic diversity within regions.     

Long-Term Effects of Prescribed Fire and Thinning on Residual Tree Growth in Mixed-Oak Forests of Southern Ohio

Long-term (10 years) growth responses of residual trees to prescribed fire and thinning were evaluated using standard dendrochronological protocols to understand the broader effects of the treatments on mixed-oak forest ecosystems in southern Ohio. Analysis of 696 increment cores (348 trees ≥ 25 cm DBH; five species) from 80 0.1 ha permanent plots distributed evenly across four treatments (control, thin, thin + burn, burn) indicated substantial increase in tree basal area increment (BAI) following the treatment. Post-treatment mean BAI of trees from the three active treatments ranged from 20.52 to 23.55 cm² y^?1 compared with pre-treatment values of 16.86–17.07 cm² y^?1. BAI rates (averaging 15.13 and 16.33 cm² y^?1, respectively, for pre- and post-treatments) in the control plots did not change much over time. Mechanical treatments were more effective than prescribed fire at enhancing BAI of trees. However, basal area growth depended to some degree on the severity of prescribed fire. Analysis of percent BAI change revealed an interesting temporal trend with moderate to major growth releases during the first 5-year post-treatment period, and a slight attenuation thereafter, suggesting the need for periodic application of treatments to sustain growth over a longer timescale. Growth responses varied greatly among species, with yellow-poplar and hickories exhibiting the highest and lowest post-treatment BAI rates of 31.11 and 15.71 cm² y^?1, respectively. Given their variable growth responses, integrating residual trees into current monitoring programs may help in elucidating the consequences of prescribed fire and thinning on forest dynamics and development.     

Forage availability for white-tailed deer following silvicultural treatments in hardwood forests

Closed-canopy upland hardwood stands often lack diverse understory structure and composition, limiting available nutrition for white-tailed deer (Odocoileus virginianus) as well as nesting and foraging structure for other wildlife. Various regeneration methods can positively influence understory development; however, non-commercial strategies are needed to improve available nutrition in many stands, as some contain timber that is not ready to harvest and others are owned by landowners who are not interested in harvesting timber. Applications of herbicide and prescribed fire have improved availability of food and cover for deer and other wildlife in pine (Pinus spp.) systems. However, this strategy has not been evaluated in hardwood systems. To evaluate the influence of fire and herbicide treatments on available deer forage in upland hardwood systems, we measured forage availability and calculated nutritional carrying capacity (NCC) at 14% crude protein mixed diet, following 7 silvicultural treatments, including controls, in 4 mixed upland hardwood stands July–September 2007 and 2008. We compared NCC among forest treatments and within 4 paired warm-season forage food plots to evaluate the usefulness of food plots in areas where forests are managed. Nutritional carrying capacity estimates (deer days/ha) were greatest following canopy reduction with prescribed fire treatments in both years. Understory herbicide application did not affect species composition or NCC 1 year or 2 years post-treatment. Production of forage plantings exceeded that of forest treatments both years with the exception of early-maturing soybeans and retention cut with fire 2 years post-treatment. We encourage land managers to use canopy reducing treatments and low-intensity prescribed fire to increase available nutrition and improve available cover where needed in upland hardwood systems. In areas where deer density may limit understory development, high-quality forage food plots may be used to buffer browsing while strategies to reduce deer density and stimulate the forest understory are implemented. © 2011 The Wildlife Society.     

Effects of fire alone or combined with thinning on tissue nutrient concentrations and nutrient resorption in <Emphasis Type="Italic">Desmodium nudiflorum</Emphasis>

This study examined tissue nutrient responses of Desmodium nudiflorum to changes in soil total inorganic nitrogen (TIN) and available phosphorus (P) that occurred as the result of the application of alternative forest management strategies, namely (1) prescribed low-intensity fire (B), (2) overstory thinning followed by prescribed fire (T + B), and (3) untreated control C), in two Quercus-dominated forests in the State of Ohio, USA. In the fourth growing season after a first fire, TIN was significantly greater in the control plots (9.8 mg/kg) than in the B (5.5 mg/kg) and T + B (6.4 mg/kg) plots. Similarly, available P was greater in the control sites (101 μg/g) than in the B (45 μg/kg) and T + B (65 μg/kg) sites. Leaf phosphorus ([P]) was higher in the plants from control site (1.86 mg/g) than in either the B (1.77 mg/g) or T + B plants (1.73 mg/g). Leaf nitrogen ([N]) and root [N] showed significant site–treatment interactive effects, while stem [N], stem [P], and root [P] did not differ significantly among treatments. During the first growing season after a second fire, leaf [N], stem [N], litter [P] and available soil [P] were consistently lower in plots of the manipulated treatments than in the unmanaged control plot, whereas the B and T + B plots did not differ significantly from each other. N resorption efficiency was positively correlated with the initial foliar [N] in the manipulated (B and T + B) sites, but there was no such relation in the unmanaged control plots. P resorption efficiency was positively correlated with the initial leaf [P] in both the control and manipulated plots. Leaf nutrient status was strongly influenced by soil nutrient availability shortly after fire, but became more influenced by topographic position in the fourth year after fire. Nutrient resorption efficiency was independent of soil nutrient availability. These findings enrich our understanding of the effects of ecosystem restoration treatments on soil nutrient availability, plant nutrient relations, and plant–soil interactions at different temporal scales.     

Prescribed burning of northern heathlands: <Emphasis Type="Italic">Calluna</Emphasis> <Emphasis Type="Italic">vulgaris</Emphasis> germination cues and seed-bank dynamics

The European coastal heathlands are important habitats for international conservation. Today, these low-intensity farming systems are threatened by the cessation of traditional management regimes, such as grazing and prescribed burning. In natural systems, the effects of fire on germination responses are often explained by adaptation to fire over extended periods of time. However, Northern heathlands are semi-natural systems with only a limited fire history. We investigated whether and how the keystone species in this system, Calluna vulgaris, responded to prescribed burning, based on previous findings where Calluna germinable seed-bank densities showed a pronounced peak right after fire. Our main findings were (i) an ecophysiological response to smoke; (ii) a potential explanation for this pattern, revealed by a seed-bank experiment where we managed to re-create the germination pattern experimentally by using an aqueous plant-derived smoke solution; and (iii) a history of anthropogenic use of fire and the development of heathlands in the region documented through palaeoecological investigations.     

Effects of late growing-season and late dormant-season prescribed fire on herbaceous vegetation in restored pine-grassland communities

Abstract. We compared the effects of late dormant-season and late growing-season prescribed fires on herbaceous species in restored shortleaf pine- (Pinus echinata) grassland communities in the Ouachita Highlands of western Arkansas. Herbaceous species richness, diversity, and total forb and legume abundance increased following fire. Late growing-season burns reduced distribution and abundance of panicums (primarily Panicum boscii, P. dichotomum, and P. linearifolium) while late dormant-season burns increased Panicum distribution and abundance. Density of legumes (such as Stylosanthes biflora) increased following frequent or annual dormant-season fires. However, season of fire influenced the distribution and abundance of fewer than 10 % of the species. Fire plays an essential role in pine-grassland communities by creating and maintaining open canopy conditions that perpetuate understory herbaceous plant communities.     

Vegetative and seedling regeneration after fire in planted Sardinian pinewood compared with that in other areas of Mediterranean-type climate

Aim Fire is a key disturbance in Mediterranean‐type climates. It has effects on plant community structure and composition and on the evolution of different groups of the flora. This study aimed to quantify changes in demography and vegetative regeneration caused by fire in key species in Mediterranean vegetation and to examine the hypothesis of convergence in characteristics of species in relation to fire by comparing the occurrence of bioecological groups of plants in the Mediterranean basin with groups of plants in other areas of Mediterranean‐type climate. Methods Changes in the demography and species composition after fire were studied in a plant community developed from a pinewood (Pinus halapensis Miller plantation) in north‐western Sardinia. The demography of the three main species dominating the community after fire, Cistus monspeliensis L., Pistacia lentiscus L. and Chamaerops humilis L., was quantified by quadrat measurements in areas of the plantation which had never been burnt and in different parts 1 and 8 years after fire. Vegetative regeneration of resprouting species was also quantified. From the results of the survey and examination of published material, comparison was made between bioecological groups of species in different regions with Mediterranean‐type climate to assess the similarities and differences in the biogeographic distribution of what can be broadly considered fire‐related functional groups of plants. Results Seeder and sprouter species showed contrasting population responses to fire in an area planted to pines over 50 years ago but subsequently in part subjected to fire. Mean species richness at a 25 m² scale decreased by c. 28% over 8 years after fire. C. monspeliensis in the unburnt pinewood was sparse and sporadic but recruitment after fire was prolific and produced plant densities similar to those of similar seeder species in other Mediterranean Basin areas and in Californian chaparral. However density was very variable between burnt areas of different ages. In particular, seedling recruitment was much lower under the canopies of P. lentiscus than away from the canopy. Sprouter species (P. lentiscus and C. humilis) had recruited into the pinewood over time since planting of the pines but did not recruit after fire. Above‐ground parts of plants were killed by fire and resprout stem density per plant increased initially but later decreased slightly due to self thinning of stems on each plant. Main conclusions Plant demographic and floristic changes after fire in the Sardinian pinewood were similar to those reported for other Mediterranean plant communities. Resprouting species only recovered by vegetative means and, in contrast to seeder species, underwent thinning of stems within, rather than between, plants, so plant density did not change after fire in resprouting species. The Tertiary relic, strongly resprouting, animal‐dispersed group of species in the Mediterranean Basin has a homologous group in the North American mediterraneoid flora but not in the Southern Hemisphere. In contrast, the general group of resprouting species which recruit after and/or between fires in the Southern Hemisphere mediterraneoid areas corresponds to only a small group of taxa in the Mediterranean basin. The bioecological group of seeder species, which are killed by and establish prolifically after fire, are represented in both Northern and Southern Hemisphere Mediterranean‐type heathlands but are much more species rich in the south.