Post‐fire understorey regeneration in boreal Pinus sylvestris forest sites with different fire histories

Abstract. We studied the characteristics of understorey regeneration on two sites with different fire history in a mature Pinus sylvestris forest in eastern Finland. The study area was a 4‐ha plot, which was divided into two parts based on fire history analysis. In one part the last fire event was a stand‐replacing fire in the early 19^th century, after which the whole stand regenerated, while the other part of the study plot was subsequently burnt by a surface fire in 1906. Understorey P. sylvestris individuals were much more abundant in the area of the 1906 burn compared to the old burn. In both areas the size frequency distribution of living trees was bimodal, with frequency peaks at the < 5 cm and 30–150 cm height classes. In the old burn small understorey trees were mainly associated with microsites created by treefall disturbances while in the 1906 burn most small understorey trees occurred on vegetation‐covered microsites. This indicates that with increasing time since last fire establishment of new understorey trees becomes more restricted by the availability of microsites created by treefall disturbances. In both areas the proportion of vigorous small understorey trees was highest on decayed wood. In the older burn uprooted pits and mounds also had a significant proportion of healthy small understorey trees, while the majority of trees classified as seriously weakened or dying were growing on microhabitats characterized by undisturbed vegetation. Ripley's K‐function analyses showed that spatial distribution of understorey trees was clustered in both areas in all microsite types and clustering at small scales was most pronounced in understorey trees growing in uprooted spots or in association with decayed wood. The bivariate analysis showed a significant repulsion effect between large trees and understorey trees at intermediate spatial scales, indicating that competition had an effect on understorey tree distribution and this effect was more pronounced in the younger burn. The analysis suggests that in Pinus sylvestris forests the abundance, quality and spatial pattern of understorey tree population may vary considerably as a function of disturbance history.     

Post‐fire shrub recruitment in a semi‐arid grassland: the role of microsites

Question: (1) Which factors regulate post‐fire recruitment and spread of the shrub Senecio bracteolatus in Patagonian grasslands? (2) What is the role of the grass Stipa speciosa on S. bracteolatus establishment in the post‐fire succession? Location: Northwest Patagonia, Argentina. Methods: We studied the effect of fire on S. bracteolatus recruitment and density by comparing these variables between burned and unburned grasslands. In burned areas, we compared abiotic characteristics and seedling establishment under the canopy of grasses (S. speciosa) and in gaps (inter‐tussock areas). Post‐fire interactions between S. bracteolatus seedlings and S. speciosa were studied using field and greenhouse experiments. Results: Density of S. bracteolatus was higher in burned than in unburned areas. In burned sites, seedlings were more abundant under tussock grasses, whereas juveniles were more abundant in gaps. Tussocks generated more attenuated micro‐environmental conditions than gaps during stressful summers. Gaps were more abundant in burned sites, while “under tussock” microsites were more frequent in unburned sites. In burned areas, tussocks allowed higher establishment of seedlings (facilitation), but gaps allowed more seedling growth and higher persistence of juveniles. Conclusions: Fire promoted S. bracteolatus recruitment in Patagonian grasslands by increasing the availability of favourable gap microsites. Grass protection for shrub seedlings became negative with time, probably due to competition with grasses. Gaps led to better performance and persistence of shrub plants. Six years after fire, higher shrub recruitment and adult density (observed as a trend) in burned grassland provides an opportunity for potential S. bracteolatus invasion.     

Persistence of resprouting species after fire in natural and post‐mine restored shrublands in southwestern Australia

Questions: Is post‐fire persistence of resprouting species lower in restored sites, and is survival related to lignotuber size? Location: Southwestern Australia, Eneabba, 300 km north of Perth. Methods: Post‐fire persistence of 10 lignotuberous shrub species was compared between three sites restored 8–24 years ago after mineral‐sand mining and three surrounding natural shrubland sites (8–24 years since previous fire). Results: Overall persistence of species was 11–93% in restored sites (mean 52%) and 79–100% in natural sites (mean 96%). Persistence increased with time since rehabilitation for five species with <25% of individuals in three species surviving in the youngest stand. For equivalent crown size, average lignotuber circumferences were 50% smaller at restored sites and this probably accounted for their higher post‐fire mortality. Apart from differences in the age of plants, restored sites had lower soil penetrability than natural sites, which may have restricted rootstock development. A tradeoff favoring a higher crown volume to lignotuber size ratio was apparent in nine of the ten species with greater crown volumes (by 37%) and smaller lignotubers (by 36%) in restored sites. Two resprouting species for which crown seed store was quantified had much higher fecundity in restored sites. Conclusions: Fires reduced resprouter persistence in restored sites owing to poor development/insufficient size of lignotubers. Further management after fires is required, including application of resprouter seeds/seedlings on restored topsoil, transplanting adult resprouters (where viable) from natural areas ahead of the mining front. Low intensity/patchy fires are recommended on long unburnt sites. Resprouter survival would have likely been much greater in the first place if a deeper sandy soil profile was rehabilitated, thereby providing a more suitable medium for lignotuber development.     

The effect of fire interval on post‐fire understorey communities in Yellowstone National Park

Questions: How does the time interval between subsequent stand‐replacing fire events affect post‐fire understorey cover and composition following the recent event? How important is fire interval relative to broad‐ or local‐scale environmental variability in structuring post‐fire understorey communities? Location: Subalpine plateaus of Yellowstone National Park (USA) that burned in 1988. Methods: In 2000, we sampled understorey cover and Pinus contorta density in pairs of 12–yr old stands at 25 locations. In each pair, the previous fire interval was either short (7–100 yr) or long (100–395 yr). We analysed variation in understorey species richness, total cover, and cover of functional groups both between site pairs (using paired t‐tests) and across sites that experienced the short fire intervals (using regression and ordination). We regressed three principal components to assess the relative importance of disturbance and broad or local environmental variability on post‐fire understorey cover and richness. Results: Between paired plots, annuals were less abundant and fire‐intolerant species (mostly slow‐growing shrubs) were more abundant following long intervals between prior fires. However, mean total cover and richness did not vary between paired interval classes. Across a gradient of fire intervals ranging from 7–100 yr, total cover, species richness, and the cover of annuals and nitrogen‐fixing species all declined while the abundance of shrubs and fire‐intolerant species increased. The few exotics showed no response to fire interval. Across all sites, broad‐scale variability related to elevation influenced total cover and richness more than fire interval. Conclusions: Significant variation in fire intervals had only minor effects on post‐fire understorey communities following the 1988 fires in Yellowstone National Park.     

Post‐fire succession and 20th century reduction in fire frequency on xeric southern Appalachian sites

Abstract. We document post‐fire succession on xeric sites in the southern Appalachian Mountains, USA and assess effects of 20th century reduction in fire frequency on vegetation structure and composition. Successional studies over 18 yr on permanent plots that had burned in 1976–1977 indicate that tree mortality and vegetation response varied with fuel load and fire season. In the first three years after fire, hardwood sprouts dominated tree regeneration. On sites where summer and autumn fires reduced litter depth to less than 1 cm, densities of shade‐intolerant Pinus seedlings increased steadily over this period. 4 to 8 yr after fire, large numbers of newly established seedlings and sprouts had grown to 1 – 10 cm DBH. By year 18 growth of these saplings led to canopy closure on most sites. Herbaceous cover and richness peaked in the first decade after fire, then declined. On similar sites that had not burned in more than 50 yr, regeneration of shade‐intolerant Pinus spp. and mean cover and richness of herbs were considerably lower than those observed on recently burned plots. Reconstructions of landscape conditions based on observed post‐fire succession and 20th century changes in fire regime suggest that reductions in fire frequency circa 1940 led to substantial changes in forest structure and decreases in cover and richness of herbaceous species.     

澳大利亚森林火灾的管理与火生态的研究

澳大利亚是火灾频发的地区．每年因森林火灾的危害都要造成相当的社会、经济损失及生态环境的破坏,故火生态的研究及火的管理在澳大利亚的生态学研究中一直占有重要地位．本文主要讨论了澳洲森林大火起燃的物理过程和机制、可燃物的特征、林火的特点、习性及对生态环境的影响和如何控制和减少火灾的危害性,达到对火进行利用、控制和管理的目的．     

Land‐use and fire history effects on post‐fire vegetation dynamics in eastern Spain

Question: Is post‐fire, medium‐term vegetation dynamics determined by land‐use or fire history prior to fire? Location: South‐facing slope in the Gallinera valley, Alicante province, eastern Spain. Methods: After mapping the land‐use and fire history of the study site using photo‐interpretation, we sampled vegetation structure on a set of plots representing the most frequent land‐use and fire history combinations on an area burned six years before sampling. We studied the effects of land‐use history, comparing the one‐fire land‐use trajectories. We analysed the effects of fire history; comparing one‐ and two‐fire plots for both previously cropped and uncropped areas. Results: Most variables were not significantly different between the earliest abandoned plots (abandoned at least 38 years before the fire) and the uncropped plots. On the most recently abandoned plots (abandoned between one and four years before the fire), the therophyte richness and the ratio of seeder: resprouter richness were significantly greatest. Different fire recurrences did not determine different post‐fire vegetation on either the uncropped or the early abandoned plots (all dominated by fire‐recruited seeder shrubs). The most recently abandoned plots had a lower resilience to fire. Conclusions: Land‐use history and recent pre‐fire land use, in particular, determined the post‐fire vegetation in the medium term. The vegetation composition converged during secondary succession among land‐use histories. Increasing fire recurrence had a small effect on mature plant communities, due to the combination of life‐history traits determining the response to fire of the dominant species.     

Life‐history traits predict perennial species response to fire in a desert ecosystem

The Mojave Desert of North America has become fire‐prone in recent decades due to invasive annual grasses that fuel wildfires following years of high rainfall. Perennial species are poorly adapted to fire in this system, and post‐fire shifts in species composition have been substantial but variable across community types. To generalize across a range of conditions, we investigated whether simple life‐history traits could predict how species responded to fire. Further, we classified species into plant functional types (PFTs) based on combinations of life‐history traits and evaluated whether these groups exhibited a consistent fire‐response. Six life‐history traits varied significantly between burned and unburned areas in short (up to 4 years) or long‐term (up to 52 years) post‐fire datasets, including growth form, lifespan, seed size, seed dispersal, height, and leaf longevity. Forbs and grasses consistently increased in abundance after fire, while cacti were reduced and woody species exhibited a variable response. Woody species were classified into three PFTs based on combinations of life‐history traits. Species in Group 1 increased in abundance after fire and were characterized by short lifespans, small, wind‐dispersed seeds, low height, and deciduous leaves. Species in Group 2 were reduced by fire and distinguished from Group 1 by longer lifespans and evergreen leaves. Group 3 species, which also decreased after fire, were characterized by long lifespans, large non‐wind dispersed seeds, and taller heights. Our results show that PFTs based on life‐history traits can reliably predict the responses of most species to fire in the Mojave Desert. Dominant, long‐lived species of this region possess a combination of traits limiting their ability to recover, presenting a clear example of how a novel disturbance regime may shift selective environmental pressures to favor alternative life‐history strategies.     

High‐severity fire corroborated in historical dry forests of the western United States: response to Fulé et al.

Accurate assessment of changing fire regimes is important, since climatic change and people may be promoting more wildfires. Government wildland fire policies and restoration programmes in dry western US forests are based on the hypothesis that high‐severity fire was rare in historical fire regimes, modern fire severity is unnaturally high and restoration efforts should focus primarily on thinning forests to eliminate high‐severity fire. Using General Land Office (GLO) survey data over large dry‐forest landscapes, we showed that the proportion of historical forest affected by high‐severity fire was not insignificant, fire severity has not increased as a proportion of total fire area and large areas of dense forest were present historically (Williams & Baker, Global Ecology and Biogeography, 21 , 1042–1052, 2012; W&B). In response, Fulé et al. (Global Ecology and Biogeography, 2013, doi: 10.1111/geb.12136; FE) suggest that our inferences are unsupported and land management based on our research could be damaging to native ecosystems. Here, we show that the concerns of FE are unfounded. Their criticism comes from misquoting W&B, mistaking W&B's methods, misusing evidence (e.g. from Aldo Leopold) and missing substantial available evidence. We also update corroboration for the extensive historical high‐severity fire shown by W&B. We suggest that restoration programmes are misdirected in seeking to reduce all high‐severity fire in dry forests, given findings from spatially extensive GLO data and other sources.     

Thermal shock and germination in North‐West European Genisteae: implications for heathland management and invasive weed control using fire

Question: Is the stimulation of germination by thermal shock (resulting from the passage of fire) commonly observed for Mediterranean‐climate Fabaceae also apparent for NW European Genisteae (Fabaceae) species? Location: Southern England and NW France. Methods: The germination of Cytisus scopiarius, Genista anglica, Ulex europaeus, Ulex gallii and Ulex minor was examined following exposure to a range of temperatures (50°C, 65°C, 80°C, 95°C and 110°C) applied to seeds for 5 min. A sixth Mediterranean‐origin species (Spartium junceum) was also included since it is a common invasive in NW Europe and North America. Results: All five native NW European species displayed increased germination following thermal shock, even when seeds were heated to 110°C. However, there was some variation depending on provenance: in contrast to seeds collected from southern England, germination of French C. scopiarius seeds was unaffected by temperature. Spartium junceum germinated most at 95°C, but was the only species to show reduced germination when seeds were heated to 110°C. Conclusions: The NW European Genisteae appear to be pre‐adapted to the high temperatures associated with fire; a response attributable to their evolutionary origins in the fire‐prone ecosystems of the Mediterranean Basin. Consequently, projected increases in fire frequency linked to climate change may stimulate their regeneration in NW European heathlands, potentially altering the species composition of these ecosystems. Additionally, a clearer understanding of the interaction between thermal shock and germination may explain why fire has so frequently been ineffective in controlling invasive Genisteae populations worldwide.     

Early post‐fire regeneration in subalpine heathland and grassland in the Victorian Alpine National Park,south‐eastern Australia

Landscape fire (at the scale of square kilometres or more) is relatively rare in the alpine and subalpine environments of Australia. In early 1998, a major fire (the ‘Caledonia Fire’), burnt approximately 35 000 ha, of which approximately 3000 hectares was subalpine heathland, grassland and wetland within the Victorian Alpine National Park. This fire was one of only three landscape‐scale fires that have occurred anywhere in the treeless vegetation of the Victorian Alps in the past 100 years, the others being in 1939 and 1985. Monitoring of regeneration in subalpine vegetation commenced 3 weeks postfire. Sites were established in burnt grassland at Holmes Plain (1400 m a.s.l.) and burnt grassland and heathland at Wellington Plain (1480 m a.s.l.), and in unburnt grassland at both sites. In burnt grassland and heathland, the fire consumed much of the vegetation, leaving extensive areas of bare ground. The cover of dense vegetation declined from > 70% prefire, to approximately 15% immediately postfire. Bare ground at the Holmes and Wellington Plains sites ranged from 70% to 85% immediately postfire. By May 2000, approximately 2.5 years postfire, dense vegetation cover in grassland had increased to approximately 20%, and bare ground had decreased to an average of approximately 30%. In unburnt grassland, dense vegetation cover was generally > 95%, and the amount of bare ground less than 5%. The tussock‐forming snow grasses resprouted vigorously following fire, and had flowered prolifically after 1 year. In heathland, most of the shrubs were incinerated, leaving close to 100% bare soil. Since then, a number of grasses and some dominant shrubs have resprouted vigorously, with some seedling regeneration. By May 2000, in heathland, bare soil was still > 50% and dense vegetation < 20%. Such ground cover conditions during this early postfire period were well below prefire levels, and well below the levels necessary to protect alpine soils from erosion. The Caledonia Fire has provided a rare opportunity to study ecological processes associated with postfire regeneration in treeless subalpine landscapes.