Successional Models as Guides for Restoration of Riparian Forest Understory

We compare two successional models as guides for restoring native riparian understory species along a 160‐km stretch of the Sacramento River in California. In 2001 and 2007, we surveyed cover, frequency, and richness of native and exotic understory species in 15 sites planted (1989–1996) with overstory species to determine whether native understory species colonized naturally (passive relay floristics model). In 2007, we surveyed 20 additional sites (planted 1997–2003) in 14 of which understory species were planted (initial floristics model) to evaluate whether planting accelerated community recovery. We surveyed 10 remnant forests as references for successional trajectories. Mean cover and frequency of natives changed little over time in sites where they were not planted initially; increases in native cover in a few sites were primarily due to a single common species (Galium aparine). Species composition shifted from light‐demanding to shade‐adapted species, both exotic and native, in response to a doubling of overstory cover. Sites with high intensity understory plantings had greater cover and frequency of native understory species than unplanted sites, but were still low relative to reference forests. Light‐demanding natives (e.g., Artemisia douglasiana, Rubus ursinus, and grasses) established in sites where they were planted; however, a shade‐adapted species (Carex barbarae) did not survive well. Our research indicates that the passive relay floristics and the initial floristic composition approaches serve to restore a few common native understory species, but that planting species as site conditions become appropriate (active relay floristics model) will be needed to restore entire native understory communities.     

Stochastic losses of fire‐dependent endemic herbs revealed by a 65‐year chronosequence of dispersal‐limited woody plant encroachment

The factors responsible for maintaining diverse groundcover plant communities of high conservation value in frequently burned wet pine savannas are poorly understood. While most management involves manipulating extrinsic factors important in maintaining species diversity (e.g., fire regimes), most ecological theory (e.g., niche theory and neutral theory) examines how traits exhibited by the species promote species coexistence. Furthermore, although many ecologists focus on processes that maintain local species diversity, conservation biologists have argued that other indices (e.g., phylogenetic diversity) are better for evaluating assemblages in terms of their conservation value. I used a null model that employed beta‐diversity calculations based on Raup–Crick distances to test for deterministic herbaceous species losses associated with a 65‐year chronosequence of woody species encroachment within each of three localities. I quantified conservation value of assemblages by measuring taxonomic distinctness, endemism, and floristic quality of plots with and without woody encroachment. Reductions in herb species richness per plot attributable to woody encroachment were largely stochastic, as indicated by a lack of change in the mean or variance in beta‐diversity caused by woody encroachment in the savannas studied here. Taxonomic distinctness, endemism, and floristic quality (when summed across all species) were all greater in areas that had not experienced woody encroachment. However, when corrected for local species richness, only average endemism and floristic quality of assemblages inclusive of herbs and woody plants were greater in areas that had not experienced woody encroachment, due to the more restricted ranges and habitat requirements of herbs. Results suggest that frequent fires maintain diverse assemblages of fire‐dependent herb species endemic to the region. The stochastic loss of plant species, irrespective of their taxonomic distinctness, to woody encroachment suggests that the relevance of niche partitioning or phylogenetic diversity to the management of biodiversity in wet pine savannas is minimal.     

Contrasting changes in vegetation structure and diversity with time since fire in two Australian Mediterranean‐climate plant communities

Mallee‐heath and mallee communities occur in a mosaic across large areas of south‐western Australia, in topographically subdued and fire‐prone landscapes. Consequently, it could be expected that these communities would have historically experienced similar fire regimes, and would respond similarly to variation in aspects of the fire regime. We studied the response of mallee‐heath and mallee to time since the last fire, measuring species density, species–area relationships, diversity indices and vegetation structure. Floristic responses to time since fire accorded with the initial floristic composition model of plant succession, with declining species density and Shannon diversity with age in mallee‐heath. Mallee‐heath exhibited structural senescence when > approximately 45–55 years since fire, with increasing standing dead vegetation, bare ground and stagnating or declining size in sprouting Eucalyptus spp. Mallee showed no such evidence of senescence, and indeed continued to increase in stature beyond the mean fire interval but without the compositional change required to provide support for the relay floristic model of plant succession. These results indicate that mallee‐heath is a fire maintained community and as such is reliant upon periodic burning to maintain diversity and vigour. Mallee, in contrast, is modified but not maintained by fire (at least over the period of time since fire examined) and hence is less susceptible to fire interval effects. Indeed, structural attributes likely to be significant for fauna habitat and carbon sequestration continue to develop in mallee unburnt for 55 years or more. Different responses to time since fire will create challenges for management, particularly in fragmented landscapes where fire potentially interacts with other threatening processes.     

Effects of long‐term fire exclusion and frequent fire on plant community composition: A case study from semi‐arid shrublands

Time since last fire and fire frequency are strong determinants of plant community composition in fire‐prone landscapes. Our study aimed to establish the influence of time since last fire and fire frequency on plant community composition and diversity of a south‐west Australian semi‐arid shrubland. We employed a space‐for‐time approach using four fire age classes: ‘young’, 8–15 years since last fire; ‘medium’, 16–34; ‘old’, 35–50; and ‘very old’, 51–100; and three fire frequency classes: burnt once, twice and three times within the last 50 years. Species diversity was compared using one‐way ANOVA and species composition using PERMANOVA. Soil and climatic variables were included as covariables to partition underlying environmental drivers. We found that time since last fire influenced species richness, diversity and composition. Specifically, we recorded a late successional transition from woody seeders to long‐lived, arid‐zone, resprouting shrub species. Fire frequency did not influence species richness and diversity but did influence species composition via a reduction in cover of longer‐lived resprouter species – presumably because of a reduced ability to replenish epicormic buds and/or sufficient starch stores. The distinct floristic composition of old and very old habitat, and the vulnerability of these areas to wildfires, indicate that these areas are ecologically important and management should seek to preserve them.     

Soil carbon and nitrogen across a chronosequence of woody plant expansion in North Dakota

In annual crops, the partitioning of photosynthates to support root growth, respiration and rhizodeposition should be greater during early development than in later reproductive stages due to source/sink relationships in the plant. Therefore, seasonal fluctuations in carbon dioxide (CO₂) and nitrous oxide (N₂O) production from roots and root-associated soil may be related to resource partitioning by the crop. Greenhouse studies used ¹³C and ¹⁵N stable isotopes to evaluate the carbon (C) partitioning and nitrogen (N) uptake by corn and soybean. We also measured the CO₂ and N₂O production from planted pots as affected by crop phenology and N fertilization. Specific root-derived respiration was related to the ¹³C allocated to roots and was greatest during early vegetative growth. Root-derived respiration and rhizodeposition were greater for corn than soybean. The ¹⁵N uptake by corn increased between vegetative growth, tasseling and milk stages, but the ¹⁵N content in soybean was not affected by phenology. A peak in N₂O production was observed with corn at the milk stage, suggesting that the corn rhizosphere supported microbial communities that produced N₂O. Most of the ¹⁵N-NO₃ applied to soybean was not taken up by the plant and negative N₂O production during vegetative growth and floral initiation stages suggests that soybean roots supported the reduction of N₂O to dinitrogen (N₂). We conclude that crop phenology and soil N availability exert important controls on rhizosphere processes, leading to temporal variation in CO₂ and N₂O production.     

Species richness and cover along a 60-year chronosequence in old-fields of southeastern Spain

We analyse changes in plant cover and species richness along a 60-year chronosequence in semi-arid Mediterranean old-fields of southeastern Spain. The objectives were: (i) to study patterns of species richness along the abandonment gradient in semi-arid conditions (e.g., to test the “humped-back model” in our system); (ii) to test whether different broad life forms (annuals, forbs, grasses and woody species) showed different patterns along the abandonment gradient, and (iii) to examine to what extent plants with different dispersal strategies dominate at different stages of succession. The explained variance of the regression relating species richness to years since abandonment is improved when considering different life forms. The results suggest that cover and richness of different functional groups show a non-linear unimodal (often positive-skewed) pattern along the gradient (age since abandonment). Maximum total richness is found at young stages of abandonment (<20 years), when most life forms and dispersal strategies coexist. Annuals and perennial forbs reached their maximum richness during the first 10 years of abandonment. About 45% of total woody species richness is reached at this time as a consequence of early colonization of zoochorous shrubs. While the results showed a tendency towards a life-form replacement sequence, the pattern is not so clear when looking at the different dispersal strategies. The results complement previous results in Mediterranean conditions and emphasise the importance of considering different functional types when studying successional patterns. This revised version was published online in June 2006 with corrections to the Cover Date.     

Patterns, determinants and models of woody plant diversity in China

What determines large-scale patterns of species richness remains one of the most controversial issues in ecology. Using the distribution maps of 11 405 woody species in China, we compared the effects of habitat heterogeneity, human activities and different aspects of climate, particularly environmental energy, water-energy dynamics and winter frost, and explored how biogeographic affinities (tropical versus temperate) influence richness-climate relationships. We found that the species richness of trees, shrubs, lianas and all woody plants strongly correlated with each other, and more strongly correlated with the species richness of tropical affinity than with that of temperate affinity. The mean temperature of the coldest quarter was the strongest predictor of species richness, and its explanatory power for species richness was significantly higher for tropical affinity than for temperate affinity. These results suggest that the patterns of woody species richness mainly result from the increasing intensity of frost filtering for tropical species from the equator/lowlands towards the poles/highlands, and hence support the freezing-tolerance hypothesis. A model based on these results was developed, which explained 76-85% of species richness variation in China, and reasonably predicted the species richness of woody plants in North America and the Northern Hemisphere.     

Influence of fire history on small mammal distributions: insights from a 100‐year post‐fire chronosequence

Aim Fire affects the structure and dynamics of ecosystems world‐wide, over long time periods (decades and centuries) and at large spatial scales (landscapes and regions). A pressing challenge for ecologists is to develop models that explain and predict faunal responses to fire at broad temporal and spatial scales. We used a 105‐year post‐fire chronosequence to investigate small mammal responses to fire across an extensive area of ‘tree mallee’ (i.e. vegetation characterized by small multi‐stemmed eucalypts). Location The Murray Mallee region (104,000 km²) of semi‐arid Australia. Methods First, we surveyed small mammals at 260 sites and explored the fire responses of four species using nonlinear regression models. Second, we assessed the predictive accuracy of models using cross‐validation and by testing with independent data. Third, we examined our results in relation to an influential model of animal succession, the habitat accommodation model. Results Two of four study species showed a clear response to fire history. The distribution of the Mallee Ningaui Ningaui yvonneae, a carnivorous marsupial, was strongly associated with mature vegetation characterized by its cover of hummock grass. The occurrence of breeding females was predicted to increase up to 40–105 years post‐fire, highlighting the extensive time periods over which small mammal populations may be affected by fire. Evaluation of models for N. yvonneae demonstrated that accurate predictions of species occurrence can be made from fire history and vegetation data, across large geographical areas. The introduced House Mouse Mus domesticus was the only species positively associated with recently burnt vegetation. Main conclusions Understanding the impact of fire over long time periods will benefit ecological and conservation management. In this example, tracts of long‐unburnt mallee vegetation were identified as important habitat for a fire‐sensitive native mammal. Small mammal responses to fire can be predicted accurately at broad spatial scales; however, a conceptual model of post‐fire change in community structure developed in temperate Australia is not, on its own, sufficient for small mammals in semi‐arid systems.     

The role of plant resources in forest succession: changes in radiation, water and nutrient fluxes, and plant productivity over a 300-yr-long chronosequence in NW-Germany

In the past insufficient attention has been paid to quantitative measurements of resource fluxes in ecosystems that undergo successional change. In this study, simultaneous changes in seven plant resources (photosynthetically active radiation (PAR), water, nitrogen, phosphorus, calcium, magnesium and potassium) are quantified by a chronosequence approach for a 300-yr-long secondary succession on poor soil from Calluna vulgaris heathland to Fagus sylvatica-Quercus petraea late-successional forest (heathland-to-forest succession).Above-ground net primary production increases sevenfold, and total above-ground phytomass about fortyfold during heathland-to-forest succession. Plant organs that capture resources increase much more slowly (leaf area index: threefold; fine root biomass: 1.3-fold). The increase in productivity is based both on higher absorptivity and conversion efficiency of PAR by the canopies of the successional plants.Accumulation of organic material on the forest floor significantly improves soil water availability. Evapotranspiration losses increase early in succession as the growing vegetation increases in both height and leaf area but tend to decrease again in the late-successional community. Drainage losses are at their minimum at the conifer-dominated pioneer forest stage.Accumulation of available nutrients in the soil is a key process in heathland-to-forest succession that significantly improves plant nutrient availability but leads to only minor changes in carbon/nutrient ratios and humus quality. Litter decomposition rates increase and result in a more rapid nutrient turnover in late successional stages. External nutrient inputs (from the atmosphere and soil weathering) significantly contribute to plant nutrient supply early in succession, whereas the internal cycling of nutrients through litter fall and nutrient mineralisation by far exceeds external inputs at the late stages.Vitousek & Reiners' (1975) ecosystem nutrient loss hypothesis is supported by the heathland-to-forest succession data. Odum's (1969) hypotheses on how nutrient cycles change during the course of succession is, in one part, rejected, in part supported. Tilman's (1988) hypothesis on nutrient limitation early, and light limitation late in primary succession is rejected.     

Direct effects of warming increase woody plant abundance in a subarctic wetland

Both the direct effects of warming on a species’ vital rates and indirect effects of warming caused by interactions with neighboring species can influence plant populations. Furthermore, herbivory mediates the effects of warming on plant community composition in many systems. Thus, determining the importance of direct and indirect effects of warming, while considering the role of herbivory, can help predict long‐term plant community dynamics. We conducted a field experiment in the coastal wetlands of western Alaska to investigate how warming and herbivory influence the interactions and abundances of two common plant species, a sedge, Carex ramenskii, and a dwarf shrub, Salix ovalifolia. We used results from the experiment to model the equilibrium abundances of the species under different warming and grazing scenarios and to determine the contribution of direct and indirect effects to predict population changes. Consistent with the current composition of the landscape, model predictions suggest that Carex is more abundant than Salix under ambient temperatures with grazing (53% and 27% cover, respectively). However, with warming and grazing, Salix becomes more abundant than Carex (57% and 41% cover, respectively), reflecting both a negative response of Carex and a positive response of Salix to warming. While grazing reduced the cover of both species, herbivory did not prevent a shift in dominance from sedges to the dwarf shrub. Direct effects of climate change explained about 97% of the total predicted change in species cover, whereas indirect effects explained only 3% of the predicted change. Thus, indirect effects, mediated by interactions between Carex and Salix, were negligible, likely due to use of different niches and weak interspecific interactions. Results suggest that a 2°C increase could cause a shift in dominance from sedges to woody plants on the coast of western Alaska over decadal timescales, and this shift was largely a result of the direct effects of warming. Models predict this shift with or without goose herbivory. Our results are consistent with other studies showing an increase in woody plant abundance in the Arctic and suggest that shifts in plant–plant interactions are not driving this change.     

Unpredictability of fish recruitment: interannual variation in young–of–the–year abundance

In 1989–1998, vendace larvae Coregonus albula were sampled in Finnish lakes following a stratified random sampling design. The abundance of young–of–the–year vendace after the first growing season was estimated using catch–per–unit–effort statistics. The number or total area of nursery places hardly limited the recruitment of vendace in the study lakes. The major proportion of prerecruit mortality of vendace occurred in the larval phase and larval sampling produced significant information on young–of–the–year survival. Although larval abundances and recruitment were clearly associated only in Lake Onkamo, generally high larval abundance was needed to produce high number of recruits.     

广西马山岩溶植被年龄序列的群落特征

石漠化地区是我国西南植被恢复和生态重建的重点和难点地区。通过对马山县岩溶植被年龄序列(石漠、草丛、灌丛、小乔林和成熟林)5个演替阶段15个样地(20 m×50 m)的系统取样调查,研究了停止人为干扰后岩溶植被的更新、演替及群落特征的变化。结果表明:沿石漠、草丛、灌丛、小乔林、成熟林的顺向演替发展,群落各层次的覆盖度存在显著差异(P<0.05),乔木层覆盖度以成熟林最大,灌木层覆盖度以小乔林最高,而草本层覆盖度以灌丛最高;重要值≥10.00的科、属、种最大值出现在小乔林;不同演替阶段群落不同层次的结构明显不同,乔木层的植物密度存在显著差异(P<0.05),以小乔林最高;成熟林灌木层的植物密度显著低于小乔林和草丛(P<0.05),与石漠和灌丛差异不显著(P>0.05);森林阶段草本层的植物密度显著低于灌丛、草丛和石漠(P<0.05);群落的物种丰富度随着顺向演替发展而增加,但不同演替阶段不同层次的丰富度变化不同,草本层的物种丰富度以草丛阶段最大,成熟林最小;灌木层的丰富度以小乔林阶段最大,石漠阶段最小;乔木层的丰富度以小乔林最大,成熟林有所下降;不同演替阶段群落草本层的生态优势度无显著差异(P>0.05),而灌木层和乔木层均存在显著差异(P<0.05)。岩溶植被恢复可分为恶劣物理环境阈值和顶极种缺乏的阈值两个阶段,每一个阶段内由不同功能特征的驱动种和关键种决定着群落的演替方向和速度,越接近演替后期,顶极种越丰富,群落驱动种和关键种向高级、大型和长寿植物发展的趋势越明显。     

Salt stress limitation of seedling recruitment in a salt marsh plant community

Summary Seedling recruitment in salt marsh plant communities is generally precluded in dense vegetation by competition from adults, but is also relatively rare in disturbance-generated bare space. We examined the constraints on seedling recruitment in New England salt marsh bare patches. Under typical bare patch conditions seed germination is severely limited by high substrate salinities. We examined the germination requirements of common high marsh plants and found that except for one notably patch-dependent fugitive species, the germination of high marsh plants is strongly inhibited by the high soil salinities routinely encountered in natural bare patches. Watering high marsh soil in the greenhouse to alleviate salt stress resulted in the emergence of up to 600 seedlings/225 cm². The vast majority of this seed bank consisted of Juncus gerardi, the only common high marsh plant with high seed set. We tested the hypothesis that salt stress limits seedling contributions to marsh patch secondary succession in the field. Watering bare patches with fresh water partially alleviated patch soil salinities and dramatically increased both the emergence and survival of seedlings. Our results show that seedling recruitment by high marsh perennial turfs is limited by high soil salinities and that consequently their population dynamics are determined primarily by clonal growth processes. In contrast, populations of patch-dependent fugitive marsh plants which cannot colonize vegetatively are likely governed by spatially and temporally unpredictable windows of low salinities in bare patches.     

Effects of seed burial and fire on seedling and sapling recruitment, survival and growth of African savanna woody plant species

Seed predation and seedling mortality can act as strong demographic “bottlenecks” to sapling recruitment in African savanna woodlands. Fire also limits tree recruitment from saplings by suppressing their growth. I conducted field experiments with 13 woody plant species to assess the effects of seed burial on seedling emergence rates and effects of fire on seedling and sapling survival and growth rates over a period of 8 years at a savanna plot in central Zambia, southern Africa. Seed removal rates by small rodents varied among years and buried seeds had significantly higher emergence rates than seeds exposed to predators in most but not all the species. Annual burning reduced sapling growth in some species but in other species saplings experienced successive shoot die back even in the absence of fire. The findings show that for some woody species, seed predation is an important constraint to seedling recruitment but not for others and annual fires are important hindrances to demography and growth for some species but not others. Thus, demographic “bottlenecks” occur at different life history stages in different savanna woody species and these have the potential to alter woody tree competitive relationships and ultimately savanna structure.     

Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.