Long‐term Changes in Bird Communities after Wildfires in the Central Brazilian Amazon

We examined long‐term responses of an Amazonian bird assemblage to wildfire disturbance, investigating how understory birds reacted to forest regeneration 1, 3, and 10 years after a widespread fire event. The bird community was sampled along the Arapiuns and Maró river catchments in central Brazilian Amazonia. Sampling took place in 1998, 2000, and 2008 using mist‐nets in eight plots (four burned, four unburned sites). Species richness did not change significantly in unburned sites. In burned sites, however, we found significantly lower richness in 1998, higher richness in 2000, and similar richness in 2008. Multi‐dimensional scaling ordination showed consistent differences in bird communities both within burned sites sampled in different sampling years, and between burned and unburned sites in all years. Of the 30 most abundant species, 12 had not recovered 10 years after the fires, including habitat specialists such as mixed flocks specialists and ant‐followers. Fire‐disturbance favored three species (two hummingbirds and a manakin) in the short term only. All other species were either favored throughout the study (seven species of omnivores and small insectivores) or did not show a clear response (eight species). In burned sites, we also found significantly lower abundance of species sensitive to disturbances and habitat specialists over the entire study period. Although the bird community seems to be recovering in terms of richness, the overall community composition and abundance of some species in post‐burned and unburned sites remain very different, and have not recovered after 10 years of forest regeneration.     

Vegetation responses to burning in a rain forest in Borneo

During the 1997/98 ENSO (El Niño Southern Oscillation) event more than 5 million ha of East Kalimantan, Indonesia burned. Here we quantify the initial stages of regeneration (19982001), both in forest that burned and in unburned controls. Sapling and seedling density and species richness remained significantly lower in burned than in unburned forest and community composition remained substantially different between both forest types throughout the sampling period. The only pronounced edge effect was a significantly higher density of seedlings in the interior of unburned forest. Sapling density increased and seedling density declined in both unburned and burned forest during the four-year study period. In the unburned forest, sapling and seedling species richness remained stable, but sapling species richness declined significantly with time in the burned forest. The pioneer community in the burned forest was, furthermore, characterised by higher growth and recruitment than in the unburned forest but mortality did not differ between both forest types. Differences in environment (burned versus unburned: 2965% of variation explained) and the distance between sample sites (1323% of variation explained) explained substantial amounts of variation in sapling and seedling community similarity. Similarity was, however, only marginally (< 1% explained) related to the edge position and temporal variation (difference among sample events). Our results, four years after the initial burn, indicate that burned forest still differed greatly from unburned forest in terms of density, species richness and community composition. There was also no clear trend of a return to pre-disturbance conditions, which indicates that the burned forest may remain in a severely degraded state for a prolonged period of time.     

Butterfly,seedling, sapling and tree diversity and composition in a fire‐affected Bornean rainforest

Abstract: Fire‐affected forests are becoming an increasingly important component of tropical landscapes. The impact of wildfires on rainforest communities is, however, poorly understood. In this study the density, species richness and community composition of seedlings, saplings, trees and butterflies were assessed in unburned and burned forest following the 1997/98 El Niño Southern Oscillation burn event in East Kalimantan, Indonesia. More than half a year after the fires, sapling and tree densities in the burned forest were only 2.5% and 38.8%, respectively, of those in adjacent unburned forest. Rarefied species richness and Shannon's H’ were higher in unburned forest than burned forest for all groups but only significantly so for seedlings. There were no significant differences in evenness between unburned and burned forest. Matrix regression and Akaike's information criterion (AIC) revealed that the best explanatory models of similarity included both burning and the distance between sample plots indicating that both deterministic processes (related to burning) and dispersal driven stochastic processes structure post‐disturbance rainforest assemblages. Burning though explained substantially more variation in seedling assemblage structure whereas distance was a more important explanatory variable for trees and butterflies. The results indicate that butterfly assemblages in burned forest were primarily derived from adjacent unburned rainforest, exceptions being species of grass‐feeders such as Orsotriaena medus that are normally found in open, disturbed areas, whereas burned forest seedling assemblages were dominated by typical pioneer genera, such as various Macaranga species that were absent or rare in unburned forest. Tree assemblages in the burned forest were represented by a subset of fire‐resistant species, such as Eusideroxylon zwageri and remnant dominant species from the unburned forest.     

Bird Community Changes in Response to Single and Repeated Fires in a Lowland Tropical Rainforest of Eastern Borneo

Our current understanding of bird community responses to tropical forest fires is limited and strongly geographically biased towards South America. Here we used the circular plot method to carry out complete bird inventories in undisturbed, once burned (1998) and twice burned forests (1983 and 1998) in East Kalimantan (Indonesia). Additionally, environmental variables were measured within a 25 m radius of each plot. Three years after fire the number of birds and bird species were similar for undisturbed and burned forests, but species diversity and turnover were significantly lower in the burned forests. The bird species composition also differed significantly between undisturbed and burned forests, with a strong decline of closed forest preferring bird species accompanied by a strong increase in degraded forest preferring species in burned forests. These differences were strongly related to differences in environmental conditions such as shifts in vegetation cover and layering and differences in ground and understorey vegetation structure. We also found significant shifts in body mass distribution, foraging height and feeding guilds between the bird communities in unburned and burned forests. Surprisingly, repeated burning did not lead to increasing impoverishment of the avifauna, and both once and twice burned forests still contained most of the bird species that were also present in undisturbed forest, even though their densities were considerably lowered.     

Diversity and community composition of butterflies and odonates in an ENSO-induced fire affected habitat mosaic: a case study from East Kalimantan, Indonesia

Little is known about the diversity of tropical animal communities in recently fire‐affected environments. Here we assessed species richness, evenness, and community similarity of butterflies and odonates in landscapes located in unburned isolates and burned areas in a habitat mosaic that was severely affected by the 1997/98 ENSO (El Niño Southern Oscillation) event in east Kalimantan, Indonesian Borneo. In addition related community similarity to variation in geographic distance between sampling sites and the habitat/vegetation structure Species richness and evenness differed significantly among landscapes but there was no congruence between both taxa. The species richness of butterflies was, for example, highest in sites located in a very large unburned isolate whereas odonate species richness was highest in sites located in a small unburned isolate and once‐burned forest. We also found substantial variation in the habitat/vegetation structure among landscapes but this was mainly due to variation between unburned and burned landscapes and variation among burned landscapes. Both distance and environment (habitat/vegetation) contributed substantially to explaining variation in the community similarity (beta diversity) of both taxa. The contribution of the environment was, however, mainly due to variation between unburned and burned landscapes, which contained very different assemblages of both taxa. Sites located in the burned forest contained assemblages that were intermediate between assemblages from sites in unburned forest and sites from a highly degraded slash‐and‐burn area indicating that the burned forest was probably recolonised by species from these disparate environments. We, furthermore, note that in contrast to species richness (alpha diversity) the patterns of community similarity (beta diversity) were highly congruent between both taxa. These results indicate that community‐wide multivariate measures of beta diversity are more consistent among taxa and more reliable indicators of disturbance, such as ENSO‐induced burning, than univariate measures.     

不同林型兴安落叶松林火烧迹地物种组成及多样性研究

以内蒙古大兴安岭兴安落叶松林火烧迹地为研究对象,采用分层取样的方法对群落中植被进行了调查,分析比较了3种不同林型兴安落叶松林群落恢复过程中物种组成及多样性的差异。结果表明:不同林型兴安落叶松林群落各层α多样性指数总体表现为草本层灌木层乔木层。不同林型兴安落叶松林火烧迹地群落恢复12年后,草本层的Simpson和Shannon-Wiener指数均小于对照样地并呈现出草类-兴安落叶松林杜香-兴安落叶松林杜鹃-兴安落叶松林的趋势,而在灌木层则各指数均大于对照样地并呈现出草类-兴安落叶松林杜鹃-兴安落叶松林杜香-兴安落叶松林的趋势。     

Fire effects on the composition of a bird community in an Amazonian savanna (Brazil).

The effects of fire on the composition of a bird community were investigated in an Amazonian savanna near Alter-do-Ch?o, Pará (Brazil). Mist-net captures and visual counts were used to assess species richness and bird abundance pre- and post-fire in an approximately 20 ha area. Visual counts along transects were used to survey birds in an approximately 2000 ha area in a nearby area. Results using the same method of ordination analysis (multidimensional scaling) showed significant effects of fire in the 20 ha and 2000 ha areas and strongly suggest direct effects on bird community composition. However, the effects were different at different spatial scales and/or in different years, indicating that the effects of fire vary spatially and/or temporally. Bird community composition pre-fire was significantly different from that found post-fire. Using multiple regression analysis it was found that the numbers of burned and unburned trees were not significantly related to either bird species richness or bird abundance. Two months after the fire, neither bird species richness nor bird abundance was significantly related to the number of flowering trees (Lafoensia pacari) or fruiting trees (Byrsonima crassifolia). Since fire is an annual event in Alter-do-Ch?o and is becoming frequent in the entire Amazon, bird community composition in affected areas could be constantly changing in time and space.     

Impact of Past Forest Fires on Bird Populations in Flooded Forests of the Cuini River in the Lowland Amazon

Blackwater floodplain forests of the Rio Negro are susceptible to understory fires. Bird composition was distinct between burned and unburned floodplain forest but not between young (12–18 yr) and old burns (>25 yr), indicating low resilience after fire. Forest regeneration is slow, with open grassy areas persisting >80 yr.     

Post-fire recovery of ant communities in Submediterranean Pinus nigra forests

This study analyzes the variations in the structure and composition of ant communities in burned Pinus nigra forests in central Catalonia (NE Spain). Pinus nigra forests do not recover after fire, changing to shrublands and oak coppices. For this reason, we suggest that ant communities of burned P. nigra forests will change after fire, because the post‐fire scenario, in particular with the increase of open areas, is different to the unburned one, and more favourable for some species than for others. In four locations previously occupied by P. nigra forests where different fires occurred 1, 5, 13 and 19 yr before the sampling, we sampled the structure and composition of ant communities with pitfall traps, tree traps and net sweeping in unburned plots and in plots affected by canopy and understory fire. The results obtained suggest that canopy and understory fire had little effect on the structure of ant communities. Thus, many variables concerning ant communities were not modified either by fire type (understory or canopy fire) or by time since fire. However, a number of particular species were affected, either positively or negatively, by canopy fire: three species characteristic of forest habitats decreased after fire, while eight species characteristic of open habitats increased in areas affected by canopy fire, especially in the first few years after fire. These differences in ant community composition between burned and unburned plots imply that the maximum richness is achieved when there is a mixture of unburned forests and areas burned with canopy fire. Moreover, as canopy cover in P. nigra forests burned with canopy fire is not completed in the period of time studied, the presence of the species that are characteristic of burned areas remains along the chronosequence studied, while the species that disappear after fire do not recover in the period of time considered. Overall, the results obtained indicate that there is a persistent replacement of ant species in burned P. nigra forests, as is also the case with vegetation.     

Diversity Patterns of Bornean Butterfly Assemblages

Borneo contains a diverse rainforest butterfly community, but its forests are under threat from logging and ENSO- (El Niño Southern Oscillation) induced fires. Contrasts in butterfly assemblage structure were examined in nine 450 ha landscapes in logged forest, primary unburned continuous and isolated forest, and forest affected by surface fires during the 1997/98 ENSO event. Temporally the effect of the 1997/98 ENSO event was followed in a single burned landscape from 1997 to 2004. In total, 517 species were present in 190 sampling sites. There was a five-fold difference in species richness among landscapes, with highest richness in continuous landscapes and lowest richness in burned landscapes. Richness was also higher in logged forest than proximate unlogged forest. Temporally, species richness dropped dramatically from 1997 to 1998, but afterwards increased remaining, however, substantially lower than pre-ENSO (1997) sampling. Sites in burned landscapes were distinct from other sites in terms of vegetation structure with the slash-and-burn area the most dissimilar to other landscapes. There was much less structure among unburned landscapes. The pattern of butterfly community composition was similar to that of vegetation structure with the community from the slash-and-burn area the most distinct. However, there was much less overlap among sites from different landscapes. Temporally, 1998 possessed the most distinct assemblage when compared to assemblages from other years. The community composition was, however, slowly returning to a pre-disturbance composition. Variance in community composition explained by environmental and spatial factors differed substantially among landscapes. The spatial fraction was the only explanatory component in recently burned landscapes and a proximate small unburned isolate, but explained no variation in logged landscapes. The environmental fraction explained substantial amounts of variation in logged landscapes and the slash-and-burn area. When all landscapes were pooled high proportions of variation in butterfly community composition were explained by both geographic distance between sites and environmental variables. In contrast when only unburned landscapes were considered, most variation was explained by the geographic distance among them. Despite differences among landscapes there was a general pattern of relatively sharp decline in similarity at short distances that levels out over greater distances, a result that agrees with previous studies on other tropical species assemblages.     

Fire mediated patterns of population densities in mountain big sagebrush bird communities

We employed a chronosequence approach to evaluate patterns of bird abundance in relation to post-fire vegetation recovery in mountain big sagebrush (Artemisia tridentata vaseyana). We estimated population density for 12 species of birds within the perimeters of 4 fires that had undergone 8–20 years of vegetation recovery and on adjacent unburned areas in the northwestern Great Basin, USA. Six species showed negative responses to fire persisting up to 20 years. Two species showed positive responses with effects persisting for <20 years. Understory vegetation was similar between burned and unburned areas irrespective of recovery time, and shrub canopy cover was similar between burned and unburned sites after 20 years of recovery. Persistent reductions in bird densities lead us to conclude that shrub canopy cover alone is not a sufficient metric for predicting recovery of songbird abundances following disturbance in mountain big sagebrush. © 2013 The Wildlife Society.     

大兴安岭火烧迹地恢复初期土壤微生物群落特征

对大兴安岭兴安落叶松2003年重度和中度火烧迹地以及未过火样地的土壤微生物群落进行了考察,旨在揭示火烧迹地恢复初期土壤微生物群落变化特征。研究结果表明火烧迹地土壤养分(全氮、全碳、土壤有机质、有效氮)和土壤水分与未过火对照样地存在显著差异;火烧迹地土壤微生物量碳氮、微生物代谢活性以及碳源利用能力均显著高于对照样地;但火烧迹地与对照样地土壤微生物群落结构指标土壤微生物量碳氮比(MBC/MBN)以及多样性指数没有显著差异。相关分析结果表明:土壤微生物量、代谢活性和碳源利用能力与土壤养分指标(全碳、全氮、速效氮、有机质)和土壤水分含量有显著相关性。主成分分析的结果表明火烧与否是火烧样地与对照样地土壤微生物对碳源利用能力差异的原因。所有样地土壤微生物群落真菌比例较高,可能与该地区土壤酸碱度有关(pH=4.12—4.68)。经过6a的恢复,重度和中度火烧迹地的土壤养分和水分、土壤微生物群落的生长、代谢、以及群落多样性仍存在差异,但均不显著,表明此时火烧程度对土壤微生物群落的影响已很微弱。     

Differences in bird communities in postfire silvicultural practices stands within pine forest of South Korea

We examined differences in bird communities in relation to characteristics of habitat structure in a pine forest, Samcheok, South Korea. An unburned stand, a stand burned 7 years earlier and then naturally restored, and a stand where Japanese red pine Pinus densiflora seedlings were planted after the fire were used for the survey. Habitat structure was dramatically changed by postfire silvicultural practices. Number of stand trees, shrubs, seedlings, snags, and vegetation coverage were significantly different among study stands. We made 1,421 detections of 46 bird species during 23 separate line transect surveys per stand between February 2007 and December 2008. The mean number of observed bird species and individuals, bird species diversity index (H′), and Simpson’s diversity index (D _s) were highest in the unburned stand and lowest in the pine seedling stand. There were more species and individuals of forest-dwelling birds in the unburned stand than both burned stands. Canopy and cavity nesters, foliage searchers, bark gleaners, and timber drillers were significantly higher in the unburned stand. In the pine seedling stand, densities of birds that prefer open field and shrub cover were higher. Stand structure was simplified in the pine seedling stand by postfire practices. Because of differences in habitat structure and bird communities, postfire practices in the burned stand should be re-evaluated. Also, management strategies for pine forest after forest fires are needed based on results of long-term experiments.     

An examination of scale of assessment,logging and ENSO-induced fires on butterfly diversity in Borneo

The impact of disturbance on species diversity may be related to the spatial scales over which it occurs. Here I assess the impact of logging and ENSO (El Niño Southern Oscillation) -induced burning and forest isolation on the species richness (477 species out of more than 28,000 individuals) and community composition of butterflies and butterfly guilds using small (0.9 ha) plots nested within large (450 ha) landscapes. The landscapes were located in three habitat classes: (1) continuous, unburned forest; (2) unburned isolates surrounded by burned forest; and (3) burned forest. Plots with different logging histories were sampled within the two unburned habitat classes, allowing for independent assessment of the two disturbance factors (logging and burning). Disturbance within habitat classes (logging) had a very different impact on butterfly diversity than disturbance among habitat classes (due to ENSO-induced burning and isolation). Logging increased species richness, increased evenness, and lowered dominance. Among guilds based on larval food plants, the species richness of tree and herb specialists was higher in logged areas but their abundance was lower. Both generalist species richness and abundance was higher in logged areas. Among habitat classes, species richness was lower in burned forest and isolates than continuous forest but there was no overall difference in evenness or dominance. Among guilds, generalist species richness was significantly lower in burned forest and isolates than continuous forest. Generalist abundance was also very low in the isolates. There was no difference among disturbance classes in herb specialist species richness but abundance was significantly higher in the isolates and burned forest than in continuous forest. Tree specialist species richness was lower in burned forest than continuous forest but did not differ between continuous forest and isolates.The scale of assessment proved important in estimating the impact of disturbance on species richness. Within disturbance classes, the difference in species richness between primary and logged forest was more pronounced at the smaller spatial scale. Among disturbance classes, the difference in species richness between continuous forest and isolates or burned forest was more pronounced at the larger spatial scale. The lower levels of species richness in ENSO-affected areas and at the larger (landscape) spatial scale indicate that future severe ENSO events may prove one of the most serious threats to extant biodiversity.     

The responses of leaf litter ant communities to wildfires in the Brazilian Amazon: a multi-region assessment

Fire is frequently used as tool for land management in the Amazon, but often escapes into surrounding forests, with potentially severe impacts for forest biodiversity. We investigated the effects of single wildfires on ant communities in four geographically distinct regions of the Brazilian Amazon (Roraima, Pará, Acre and Mato Grosso) where forests had burned between 8 months and 10 years before our sampling. We established 7–12 transects, 500 m each, in burned and unburned forests in each region to investigate the effects of fire on forest structure and leaf litter ant communities, which were sampled using Winkler sacks. Fire effects on forest structure were more drastic in the most recently burned forests in Acre and Mato Grosso, while the impacts of older burns in Roraima and Pará were more subtle. Ant species richness was not different between burnt and unburned areas, but community composition differed between burned and control forests in all regions except Mato Grosso. At the species level, indicator species analysis showed that a limited number of species were significant indicators of unburned control forests in all regions, except Acre. Forests structure variables and leaf litter volume were all important in shaping ant communities, but their relative importance varied between regions. Our results indicate that burned forest have different ant species communities from unburned forests, and those differences are still apparent 10 years after the disturbance, highlighting the importance of effective policies for fire management in Amazon.     

Limited Edge Effects Along a Burned‐Unburned Bornean Forest Boundary Seven Years after Disturbance

Large parts of the everwet tropics have been burned, leaving many unburned–burned forest edges. Here we studied a Bornean forest edge to determine: (1) how unburned and burned forest differ in vegetation structure, diversity, composition and plant functional traits 7 yr after fire, and (2) if these variables showed significant edge effects. Environmental and inventory data from 120 plots (0.01 ha each), covering both sides of a ～1.3 km forest boundary were sampled. Differences in vegetation structure, diversity, composition and plant functional traits were analyzed in relation to disturbance type (Mann–Whitney tests) and edge distance (partial correlation analysis that controlled for confounding effects of elevation, slope and fire intensity). Seven years after fire, burned forest differed significantly from unburned forest in most measured variables while few significant edge effects were detected, i.e., there existed a sharp delimitation between the two forest types. The regeneration of the burned forest depended almost entirely on in situ recruitment with little input of late successional species from the neighboring old growth forest. On the other hand, old growth forest showed few signs of edge degradation. A possible explanation for these results might be related to the absence of a mast fruiting event during these first 7 yr of forest recovery, resulting in low levels of late successional species seed input into the burned forest, combined with the quick development of a closed canopy in the burned forest by early successional species that shielded the unburned forest from adverse edge effects.     

Effects of forest wildfire on soil microbial-community activity and chemical components on a temporal-seasonal scale

Background

Soil response and rehabilitation after wildfires are affected by natural environmental factors such as seasonality, and other time-dependent changes, such as vegetation recovery (e.g., % soil cover). These changes affect soil microbial-community activity. During summer 2006, almost 1,200 hectares (ha) of coniferous forest in northern Israel, including Byria Forest, burned.

Methods

Soil samples were collected seasonally from severely burned and unburned areas, on a time scale of 7?days to 4?years after wildfire. Chemical and microbial parameters of the forest soil system were examined.

Results

Results obtained show that increase in total soluble nitrogen (TSN) in burned areas may limit microbial activity during the first year after wildfire. Two years after wildfire, soil TSN levels in burned areas decreased to unburned levels after plant growth, allowing the microbial community to proliferate.

Conclusions

Wildfire had a significant impact on TSN, soil moisture (SM), and microbial nitrogen (MBN) compared to seasonality. These parameters are recommended for monitoring post-fire soil state. The direct effect of wildfire on soil constituents at the study site was stronger during the first 2–4?years. Indirect changes due to vegetation cover could have a longer effect on burned soil systems and should be further examined.     

Dependence on Fire in Wood-living Insects: An Experiment with Burned and Unburned Spruce and Birch Logs

Several boreal wood-living insect species breed exclusively in recently burned forest. However, the reason for this dependence on fire is largely unknown. Here wood-living insects and other arthropods were sampled from burned and unburned logs of birch and spruce in a burned forest, together with unburned logs at a clearing and in an uncut forest, during two years of succession after tree death. Burned spruce logs hosted fewer beetles than unburned logs. Notably, bark-beetles and their associated fauna, responded negatively to fire-scorching of the logs while arthropods that feed on ascomycete fungi responded positively. Fire-scorched logs more often had visible ascomycete fungi, and lost their bark faster than unburned logs. However, despite this obvious effect of fire-scorching of the logs, the species composition in burned and unburned logs at the burned site was more similar than in unburned logs at the three different sites. A larger diversity of beetles, when measured with rarefaction, was found for fire-scorched logs. When sites were compared, birch logs had the most diverse fauna at the burned site and spruce logs in the uncut forest. Pyrophilous insect species were almost exclusively confined to the burned forest, but occurred in both burned and unburned logs. These species may be divided into two groups: (1) mycophagous species that need burned substrate per se because ascomycete fungi are favoured by burning, and (2) phloem-feeders and predators that are favoured by some habitat characteristic of recently burned forest rather than of burned wood.     

Interaction of multiple disturbances: importance of disturbance interval in the effects of fire on rehabilitating mined areas

Abstract Multiple disturbance regimes are increasingly common as novel anthropogenic disturbances are added to existing natural disturbances. However, it is generally unknown whether simultaneous or sequential effects of different forms of disturbance are predictable from the independent effects of each disturbance. This study examines the short‐term effects of sequential disturbance by mineral sand‐mining followed by fire in a forest community in south‐eastern Australia. Four combinations of disturbance were sampled: unburned mined, burned mined, unburned forest (unmined) and burned forest (unmined, with between‐fire interval matching the disturbance interval between mining and fire of the burned mined treatment). All combinations were sampled approximately 12 months following fire on the burned sites. The impact of fire after mining depended on disturbance interval. Sites burned 0.5–2.4 years since mining had fewer native vascular plant species than unburned mined sites of the same mined age, whereas sites with 10–16 years or 20–26 years between mining and fire had greater native species richness than unburned mined sites of the same age. Burning 20–26 years after mining brought native species richness within the range of burned forest. For both unmined and mined sites native seedling densities increased with burning, and with longer disturbance intervals. Weed species richness and weed seedling densities were greater on mined sites than in forest, and burning mined sites elevated weed seedling densities further, particularly for short intervals. Both disturbance interval and fire intensity are likely to have contributed to these results, as intensity on mined areas increased with interval, and at 20–26 years post‐mining was equivalent to unmined forest. These results suggest that fire could be used to promote rehabilitation of these mined areas after at least 10 years, but should be excluded from earlier stages of post‐mining regeneration. However, other sources of spatial and temporal variability should be considered in addition to interval and intensity, as variation among mined areas was correlated with post‐fire weather conditions and available weed sources. Finally, the combined effects of mining and fire could not be predicted from knowledge of the disturbances operating separately, indicating that effects of multiple disturbance may be synergistic rather than additive.