Combining plant and animal traits to assess community functional responses to disturbance

There is a general consensus that functional traits are reliable indicators of adaptation of organisms to particular environmental characteristics. In this study we relate the combined distributions of species traits of plants and animals to disturbance regimes in chestnut forests of southern Switzerland affected by regular winter fires. We used co‐inertia analysis for combining the trait response of 471 invertebrate species (117 001 individuals) and 81 plant species at 23 sites with different fire and cutting histories. Trait response was assessed by calculating the variation in weighted mean traits averaged over the communities and by using mean traits in multivariate analyses. The analysis showed a strong association between plant and animal traits under fire constraints (Monte‐Carlo test, p=0.0045). Plants and animal distributions show parallel trends in responses to fire which selects traits relating to persistence (ability to survive), resilience (ability to recover) and mobility. Warmth‐demanding insects, herbivores, flying carnivores and pollinators were associated with recent fires, as were annual, ruderal and light‐demanding plant species with long flowering duration. Small arthropods feeding on dead wood and those with narrow habitat requirements were associated with low fire frequency and unburnt sites, as were competitive plants with large seeds favoring moist sites. The spatial association between plant and animal traits reflected adaptations that promote survival in the disturbance regime, while the disturbance acts as an environmental filter on the distribution and assemblage of the trait values within communities. This combined analysis of plant and invertebrate traits distributions illustrates how community and ecosystem responses can be monitored and the results generalized across localities and disturbance types. Analyses of traits that cross trophic levels provide powerful and promising tools for validating management procedures and controlling ecosystem functions.     

Annual plant community responses to density of small-scale soil disturbances in the Negev Desert of Israel

We investigated whether plant diversity and productivity in small-scale soil disturbances, which is known to be higher than in undisturbed soil, decreases as the density of the disturbances increases. We studied this in an experiment with soil diggings (15 cm diameter and 15 cm depth) dug at a range of densities, on a north- and a south-facing slope of a watershed in the central Negev Desert of Israel. The diggings were similar to the commonly occurring pits made by porcupines (Hystrix indica) as they forage for below-ground plant parts. We used four levels of digging density, within the naturally occurring range in the region, represented by a rectangular plot with rows of diggings dug at four distances between diggings. The plots were laid out in a blocked design with three replications on both slopes, with each block containing all four levels of digging density. In the spring of 1992, 1994 and 1995 we measured plant density, species richness and plant productivity in the diggings, and in adjacent equal-sized undisturbed control areas (“soil matrix”) and on the mounds made by the removed excess soil. Plant density, species richness and productivity of annual plants were higher in the diggings than in the undisturbed matrix, while these responses were very low on the mounds. Plant density, species richness and productivity in the diggings, but not in the matrix or mounds, decreased as digging density increased. This effect varied slightly with location within a watershed and with annual rainfall. The density of seeds captured in the diggings from outside the digging during the 1995 dispersal season decreased with increasing digging density, but only on one of the slopes. At the highest digging density, plant density and species number in the diggings did not decrease down the slope, as expected if interference between diggings in runoff water capture were the cause of the digging density effect. There was a weak decrease in biomass production in 1994–1995 down the slope. We used a simple mathematical model to estimate whether the distribution of rainfall intensities that occurred during the winter of 1994–1995 could result in differences between digging densities in the amount of water captured by the diggings, and whether this could explain the observed effect of digging density. The model showed that there were four events during which less water was captured by the diggings at high digging densities, except in the topmost row of diggings. Soil moisture measurements, however, showed very little difference between diggings at different digging densities. We explain our findings as the result of the interaction between the properties of the disturbance patch with its surroundings, as the diggings capture resources in the form of runoff water, and seeds moved primarily by wind. The additional resources and seeds captured in diggings increase plant density, species richness and productivity relative to the undisturbed matrix. However, the contrast in plant responses between the disturbed patches and undisturbed soil diminishes at higher digging densities. We explain this as interference among diggings at close proximity. As we did not detect a decrease in plant responses down the slopes, we conclude that interference is due to interception of the wind-driven, non-directional flow of seeds. Interception of the down-slope flow of runoff water by upslope diggings is insufficient to affect plant density, determined at the beginning of the season. Later in the season, runoff interception may become important for biomass production. Received: 16 August 1996 / Accepted: 26 October 1997     

Interactive community responses to disturbance in streams: disturbance history moderates the influence of disturbance types

Both disturbance history and disturbance type act to structure communities through selecting for particular species traits but they may also interact. For example, flooding selects for species with flood‐resistant traits in streams, but those traits could make communities susceptible to other disturbances and so could cause shifts in community composition due to anthropogenic climate change. To better understand the interactive influences of disturbance history and type on community composition, we investigated the response of macroinvertebrate communities to disturbance using in‐stream channels. Using a split‐plot design, individual channels in five ‘stable’ streams and five ‘frequently disturbed’ streams (disturbance history) were subject to different disturbance type treatments (flooding, drying and a control). Disturbance type independently drove effects on species diversity, but all other effects of disturbance type depended on disturbance history. In particular, the interaction of disturbance type and history determined overall community response. Both disturbance types tested produced similar community responses in frequently disturbed streams, including changes in community composition and alterations to the abundance of less mobile taxa, but low‐flow had a significantly greater effect in stable streams. Macroinvertebrate drift was greatest in the rock‐rolling treatments and significantly less in the low‐flow treatment for both disturbance histories. Therefore, disturbance history moderated the effects of disturbance type and determined the mechanism of community response by determining how well species were adapted to disturbance. This outcome suggests that previous disturbances strongly influence how vulnerable communities are to changes in disturbance, and so should be considered when predicting how changes in disturbance regimes will affect future community composition.     

Community and population-level responses to disturbance in a sandflat community

The responses of soft sediment infauna were investigated in an intertidal sandflat to determine patterns of recolonization and succession at the community and population level. Experimental disturbance plots, 1 m², were initiated in August and sampled for 4.5 months along with ambient sediments. Sediment grain-size was used as a general indicator of the physical state of the disturbance patches, and grain-size distributions among disturbance and ambient patches became similar after ∼2.5 months. Recolonization varied among the dominant infaunal taxa. Densities of infauna that were most abundant in the habitat, primarily syllid polychaetes, did not recover to ambient levels until 3-4 months after disturbance, when ambient densities were falling to winter lows. Multivariate analysis indicated that community recovery occurred by the end of the study period after 4.5 months. Although community structure recovered by the end of the study, the population structure of the dominant species Parapionosyllis longicirrata remained significantly different among ambient and disturbed patches. On all sampling dates except one, disturbance patches had a higher number of larger individuals than ambient sediments. Previous studies have shown that bedload transport of juveniles and adults, and other processes, can cause recolonization to be relatively rapid on intertidal sandflats. However, our results indicate that recovery times may be on the order of months at large disturbance sizes. Therefore, rapid responses may occur primarily in the case of small-scale (<1 m²) disturbance patches. Secondly, recovery at the community level does not necessarily mean that population-level characteristics of species comprising the community have recovered. Population-level differences may be longer lasting than indicated by community level indicators of recovery.     

Measuring community responses to large-scale disturbance in conservation biogeography

Aim   Which community metrics should be used to reflect community response to large-scale habitat alterations is unclear. Here, we assess what and how community changes should be measured to accurately track community responses to large-scale disturbance in space and/or time.
Location   France.
Method   We first developed a simulation model to examine temporal changes in the species composition of large-scale metacommunities. Using this model, we assessed how species richness, Shannon index, trends of particular subset of species or community indices of habitat specialization were influenced by different disturbance scenarios, and whether these indices were biased by imperfect detectability. We further used more than 1000 empirical bird communities from the French Breeding Bird Survey recently exposed to disturbances of various intensities as a case study.
Results   Our simulation and empirical results both demonstrate that species richness and diversity measures can show confusing trends and even provide misleading messages of communities' fate. In contrast, reflecting the composition of the community in terms of habitat specialist and generalist species was more robust and powerful to reflect disturbance effects.
Main conclusions   We highlight the weakness of using community metrics that fail to incorporate ecological difference among species when summarizing community-level trends in disturbed landscapes.     

Historical forest disturbance mediates soil microbial community responses to drought

Despite the abundance of studies demonstrating the effects of drought on soil microbial communities, the role of land use legacies in mediating these drought effects is unclear. To assess historical land use influences on microbial drought responses, we conducted a drought-rewetting experiment in soils from two adjacent and currently forested watersheds with distinct land use histories: an undisturbed ‘reference’ site and a ‘disturbed’ site that was clear-cut and converted to agriculture ~60 years prior. We incubated intact soil cores at either constant moisture or under a drought-rewet treatment and characterized bacterial and fungal communities using amplicon sequencing throughout the experiment. Bacterial alpha diversity decreased following drought-rewetting while fungal diversity increased. Bacterial beta diversity also changed markedly following drought-rewetting, especially in historically disturbed soils, while fungal beta diversity exhibited little response. Additionally, bacterial beta diversity in disturbed soils recovered less from drought-rewetting compared with reference soils. Disturbed soil communities also exhibited notable reductions in nitrifying taxa, increases in putative r-selected bacteria, and reductions in network connectivity following drought-rewetting. Overall, our study reveals historical land use to be important in mediating responses of soil bacterial communities to drought, which will influence the ecosystem-scale trajectories of these environments under ongoing and future climate change.     

Understorey plant and soil responses to disturbance and increased nitrogen in boreal forests

Question: How do N fertilization and disturbance affect the understorey vegetation, microbial properties and soil nutrient concentration in boreal forests? Location: Kuusamo (66°22′N; 29°18′E) and Oulu (65°02′N; 25°47′E) in northern Finland. Methods: We conducted a fully factorial experiment with three factors: site (two levels), N fertilization (four levels) and disturbance (two levels). We measured treatment effects on understorey biomass, vegetation structure, and plant, soil and microbial N and C concentrations. Results: The understorey biomass was not affected by fertilization either in the control or in the disturbance treatment. Fertilization reduced the biomass of deciduous Vaccinium myrtillus. Disturbance had a negative effect on the biomass of V. myrtillus and evergreen Vaccinium vitis‐idaea and decreased the relative proportion of evergreen species. Fertilization and disturbance increased the biomass of grass Deschampsia flexuosa and the relative proportion of graminoids. The amount of NH₄⁺ increased in soil after fertilization, and microbial C decreased after disturbance. Conclusions: Our results suggest that the growth of slow‐growing Vaccinium species and soil microbes in boreal forests are not limited by N availability. However, significant changes in the proportion of dwarf shrubs to graminoids and a decrease in the biomass of V. myrtillus demonstrate the susceptibility of understorey vegetation to N enrichment. N enrichment and disturbance seem to have similar effects on understorey vegetation. Consequently, increasing N does not affect the rate or the direction of recovery after disturbance. Moreover, our study demonstrates the importance of understorey vegetation as a C source for soil microbes in boreal forests.     

Bird community responses to disturbance in a forestry concession in lowland Bolivia

Bird community characteristics of three sites with different levels of disturbance were studied using transect surveys during the dry season in a subtropical humid forest in Bolivia. One area had been unintentionally burned 4 years prior and selectively harvested (DIS) 1 year prior to sampling. A second area had been selectively harvested 1 year prior to sampling and had no recent history of fire (HAR). Species richness, as assessed by species–time curves and rarefaction, was higher in both altered areas than in undisturbed forest (INT). In general, frugivores and omnivores were more abundant in both altered areas compared to intact forest. Canopy frugivores, understory omnivores and multiple-strata omnivores were most abundant in HAR. Canopy frugivores, near-ground insectivores, understory and multiple-strata omnivores were least abundant in INT, although INT had the highest abundances of canopy insectivores and near-ground omnivores. Richness and abundance of widespread species with low habitat specificity was higher in both areas that experienced disturbance compared to intact forest. Differences in bird community structure between disturbed and intact forest at this site are attributed primarily to the addition of widespread species with less narrow habitat requirements, and possibly to changes in the distribution of different food types.     

Ecology of plant resprouting: populations to community responses in fire-prone ecosystems

Resprouting has emerged as a key functional trait in plant ecology over the past decade with more than 400 papers published since 2000 (Web of Science). This special issue of Plant Ecology brings together a set of papers that advance our understanding of this functional trait, in the quest for developing a better conceptual framework for predicting community response to disturbance. This special issue highlights current research on all aspects of the effects of fire and other disturbances on plant resprouting behaviour and the importance of the persistence niche in structuring plant communities.     

Cardiorespiratory responses of the woodchuck and porcupine to CO2 and hypoxia

Summary The burrow-dwelling woodchuck (Marmota monax) (mean body wt.=4.45±1 kg) was compared to a similar-sized (5.87±1.5 kg) but arboreal rodent, the porcupine (Erithrizon dorsatum), in terms of its ventilatory and heart rate responses to hypoxia and hypercapnia, and its blood characteristics.V _T,f,T _I andT _E were measured by whole-body plethysmography in four awake individuals of each species. The woodchuck has a longerT _E/T _TOT (0.76±0.03) than the porcupine (0.61±0.03). The woodchuck had a higher threshold and significantly smaller slope to its CO₂ ventilatory response compared to the porcupine, but showed no difference in its hypoxic ventilatory response. The woodchuck P₅₀ of 27.8 was hardly different from the porcupine value of 30.7, but the Bohr factor, –0.72, was greater than the porcupine's, –0.413. The woodchuck breathing air has PaCO₂=48 (±2) torr, PaO₂=72 (±6), pHa=7.357 (±0.01); the porcupine blood gases are PaCO₂=34.6 (±2.8), PaO₂=94.9 (±5), pHa=7.419 (±0.03), suggesting a difference in PaCO₂/pH set points. The woodchuck exhibited no reduction in heart rate with hypoxia, nor did it have the low normoxic heart rate observed in other burrowing mammals.     

Spatial variation in insect community and species responses to habitat loss and plant community composition

Several experimental studies have examined species responses to manipulations of habitat area and spatial arrangement, but plant composition and spatial variation in species distributions also affect animal responses to habitat alteration. We used an experimental approach to study the combined effects of habitat area, edge, and plant community composition on the spatial structure of insect species richness and composition. The abundance of three guilds (herbivores, predators and parasitoids) and individual species were also analyzed. Habitat patches were created that differed in area and edge by selectively mowing portions of 15 m×15 m plots in a 1.7-ha old field. Spatial and environmental variables were used to predict insect responses in separate multiple regression and ordination models. The variation in species responses due to spatial and environmental variables was then partitioned by combining these variables into an overall regression or ordination. Spatial and environmental variables contributed similar percentages to the total variance in insect species richness, abundance or composition. No significant effects of habitat area were observed in any response variable. Herbivore abundance showed positive responses to legume or grass cover, as well as spatial variation that was unrelated to environmental variables. Predators and parasitoids had greater effects of plant species richness and habitat edge, and less unexplained spatial variation. Individual species differed in their responses to plant variables, depending on host specialization or intraspecific aggregation. Our study highlights the importance of plant community composition and spatial variation apart from environmental variables. Spatial variation stems both from species responses to environmental features as well as species differences in habitat specialization and intraspecific aggregation.     

Riparian disturbance and community structure of a Quercus-Ulmus forest in central Japan

Population structure and ecological characteristics of component species of a riparian Ulmus-Quercus forest in central Japan were analyzed with special reference to riparian disturbance regime. Though the dbh distribution of the whole community was L-shaped, those of several component tree populations had several modes, suggesting intermittent regeneration periods in the past. Correlation of spatial distributions among tree populations and subpopulations showed 6 major groups reflecting riparian disturbances in the past and different establishment patterns among species. A cluster of small-sized tree populations (Salix sachalinensis, Alnus hirsuta and Populus maximowiczii <30 cm dbh) were distributed on the lower terrace along the active river channel, while large-sized subpopulations (dbh 60 cm) of Quercus crispula and Ulumus davidiana var. japonica occurred on the higher terrace. The Phellodendron amurense population also occurred on the higher terrace in small clumps though the trees were small (less than 55 cm dbh). Subpopulations of intermediate-sized individuals (30 dbh < 60 cm) of Q. crispula and U. davidiana var. japonica, together with Betula and Acer spp. populations occurred on the intermediate terrace. Dendrochronological analyses indicated that the large and intermediate-sized tree groups were established about 330 and 90 years ago, respectively, while the small-sized tree group established about 35 years ago. A survey of historical disturbances showed that these periods of establishment of the former two groups almost coincided with the historically major floods occurring in 1662 and 1902. However, the disturbance that resulted in the establishment of the youngest group could not be precisely identified. Thus, the forest is a mosaic of three differently-aged patches, which is closely related to the frequency and scale of riparian disturbances. Longevity of trees and the preferred conditions for seed germination and/or seedling establishment were particularly important for the guild structure in this forest community.     

The responses of a community to disturbance: The importance of successional age and species' life histories

Summary The responses of different successional stages of a temperate intertidal algal community to disturbance were investigated with a field experiment. The experiment was conducted in a low intertidal boulder field in southern California. In this habitat, the top surfaces of boulders are covered with algae. The composition of the assemblage on any particular boulder depends on the length of time since it was last overturned by wave action. When a boulder is overturned, the algae on what was formerly the top surface, are killed in whole or part by a combination of sea urchin grazing, anoxia, light levels below compensation intensity, and mechanical damage caused by crushing or abrasion. The length of time that a boulder remains overturned and the local abundance of sea urchins determines the intensity of the disturbance. When the boulder is righted, recolonization begins either by vegetative regrowth of survivors and/or by spores from outside.Using a three-factorial design, this natural form of disturbance was experimentally mimicked and the responses of three different successional stages of the algal community monitored. Boulders in each successional category were overturned for periods of 17, 27 and 54 days in areas with and without sea urchins, then righted. Two aspects of community response to perturbation were evaluated. These were (1) the assemblage's ability to resist change and (2) its ability, if altered, to adjust to some semblance of its original state. The resistance of each assemblage and of its component species to change was measured by the percent decrease in algal cover and by the decline in percent similarity of the community to its original composition. The recovery rate of each assemblage and of the cover lost by each species during the first 35 days following a disturbance was measured by the rate of increase in percent similarity to the original composition and the percent reestablishment of lost cover.The experimental evidence demonstrates that the successional stages of the producer level of an intertidal algal community differ significantly in their responses to disturbance. Early successional communities suffer more damage from a given level of perturbation but recover more quickly than either middle or late successional communities. Damage to any particular assemblage of algae, irrespective of successional age, is more extensive and recovery slower, the longer the boulder is overturned and/or sea urchins are present. Several thresholds in these responses were also identified.Differences in community responses and non-linearities in these responses were attributable to the life history characteristics of the component species rather than emergent properties of the assemblage. These characteristics have evolved in response to a variety of recurrent natural disturbances. This interpretation is in agreement with recent critical reevaluations of the trends and mechanisms of successional change in natural communities.     

Riparian plant community responses to increased flooding: a meta‐analysis

A future higher risk of severe flooding of streams and rivers has been projected to change riparian plant community composition and species richness, but the extent and direction of the expected change remain uncertain. We conducted a meta‐analysis to synthesize globally available experimental evidence and assess the effects of increased flooding on (1) riparian adult plant and seedling survival, (2) riparian plant biomass and (3) riparian plant species composition and richness. We evaluated which plant traits are of key importance for the response of riparian plant species to flooding. We identified and analysed 53 papers from ISI Web of Knowledge which presented quantitative experimental results on flooding treatments and corresponding control situations. Our meta‐analysis demonstrated how longer duration of flooding, greater depth of flooding and, particularly, their combination reduce seedling survival of most riparian species. Plant height above water level, ability to elongate shoots and plasticity in root porosity were decisive for adult plant survival and growth during longer periods of flooding. Both ‘quiescence’ and ‘escape’ proved to be successful strategies promoting riparian plant survival, which was reflected in the wide variation in survival (full range between 0 and 100%) under fully submerged conditions, while plants that protrude above the water level (>20 cm) almost all survive. Our survey confirmed that the projected increase in the duration and depth of flooding periods is sufficient to result in species shifts. These shifts may lead to increased or decreased riparian species richness depending on the nutrient, climatic and hydrological status of the catchment. Species richness was generally reduced at flooded sites in nutrient‐rich catchments and sites that previously experienced relatively stable hydrographs (e.g. rain‐fed lowland streams). Species richness usually increased at sites in desert and semi‐arid climate regions (e.g. intermittent streams).     

Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change – fertilisation and herbivore loss – are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics.     

Interactive effects of fertilization and disturbance on community structure and resource availability in an old-field plant community

Summary The interactive effects of fertilization and disturbance on plant community structure and resource availability were studied by supplying four levels of nitrogen and applying four intensities of tilling to a 30 year old field in a factorial design for 2 year. Live above-ground biomass, root biomass, and litter generally increased with nitrogen supply and decreased with disturbance. Species composition varied significantly, with annuals increasing with both nitrogen and disturbance, but with perennials unaffected by nitrogen and decreased by disturbance. Species diversity decreased with disturbance, but decreased with nitrogen only in undisturbed vegetation. Root: shoot ratios decreased with added nitrogen, leaf allocation decreased with disturbance, and flowering allocation increased. Surprisingly, stem allocation was unaffected by disturbance. This result reflected a shift from vertical stems to horizontal stems as disturbance increased. Resource measurements suggested that the vegetation responded to interactions between the treatments as well as to direct treatment effects. Variation in light penetration was reduced by fertilization in undisturbed vegetation but not in tilled plots; variability was not directly affected by disturbance. The availability of nitrogen, the limiting soil nutrient, increased with fertilization but was not significantly affected by disturbance. In contrast, the ratio of ammonium to nitrate was significantly reduced by disturbance but unaffected by supply rates, suggesting that nitrogen may have had different effects under different disturbance regimes, even though its total availability was constant. While many community responses to fertilization and disturbance conformed to those reported earlier, resource and allocation measurements indicated that their interactions are not always predictable from their separate effects.