Post-burn succession on Brittany heathlands

Abstract. Vegetation succession after severe burning of a Brittany heathland was recorded in permanent plots for ten years. Mosses are the first colonizers, soon forming a dense layer which locally prevents establishment of new species. Ten years later, the moss layer is still an important component of the regenerating heathland. After ten years the pattern of species distribution is mosaic-like with grassy patches (Agrostis), heath patches (Erica, Flex), and wood (Betula). Processes which may be involved in this successional pattern are discussed. They are found to be pluriform: various strategy types, types of response to disturbance and various succession models operate simultaneously.     

Understanding ecological community succession: Causal models and theories,a review

Critical review of explanations for patterns of natural succession suggests a strong, common basis for theoretical understanding, but also suggests that several well known models are incomplete as explanations of succession. A universal, general cause for succession is unlikely, since numerous aspects of historical and environmental circumstances will impinge on the process in a unique manner. However, after disturbance, occupation of a site by any species causes changes in the conditions at the site. Sorting of species may result, since different species are adapted to different regions of environmental gradients. Such sorting can generate several patterns of species abundance in time, but commonly results in sequential replacements of species adapted to the varying conditions. This may be due to constraints on species' strategies, or life history traits, placed by the limited resources available to the organism. These constraints often result in inverse correlation between traits which confer success during early and late stages of succession. Facilitatory or inhibitory effects of species on each other are best understood in terms of these life history interactions, perhaps as restrictions on, or as moderation of, these processes.Strong support for the importance of correlations in life history traits stems from comparisons of simulated succession with and without these correlations. These simulations are reviewed in some detail, and followed by brief reviews of other prominent models for succession. Several aspects of the confusion and controversies in the successional literature are then discussed, with a view to a more optimistic synthesis and direction for successional ecology.     

Changes in microbial nutrient status during secondary succession and its modification by earthworms

Summary Microbial biomass, nutrient (N and P) status, and carbon and nutrient limitation of the microflora were investigated in soils from five different sites (field, 5-, 12-, and about 50-year-old fallow, beechwood), which represent different stages of a secondary succession from a wheat field to the climax ecosystem of a beechwood on limestone. In addition, the effect of faeces production by the substrate feeding earthworm species Octolasion lacteum (Örley) on the nutrient status of the soil microflora of these sites was studied. Humus had accumulated in the soil of the third fallow site, with an enhanced biomass of microflora. However, in the beechwood soil, which had the highest humus content, microbial biomass was lower than in the soil of the third fallow site and similar to that of the field and the two younger fallow sites. In general, soil microbial biomass was little affected by the passage of soil through the gut of O. lacteum. The soil microflora of the field, the 5-, 12-, and about 50-year-old fallow was limited by carbon, whereas in the beechwood soil phosphorus limited microbial growth. NItrogen availability to the soil microflora was low in the two younger fallow sites and high in the field and the third fallow. In the beechwood soil nitrogen supply did not affect microbial carbon utilization. Application of phosphorus stimulated glucose mineralization in the soil of the field, the third fallow, and the beechwood, but not in the two younger fallow sites. Therefor, the nutrient status of the soil microflora seems to have changed during secondary succession: presumably, during the first phase the availability of nitrogen decreased, whereas during the second phase microbial phosphorus supply became more important, which resulted in phosphorus limitation of the soil microflora in the climax ecosystem. The passage of soil through the gut of O. lacteum caused an alteration in the microbial nutrient status. Generally, microbial growth in earthworm casts was limited by carbon. The relative effect of the gut passage of the soils on microbial carbon utilization seems to increase during succession. Therefore, the effect of decomposer invertebrates on microbial nutrient supply seems to increase during secondary succession. In general, nitrogen did not limit microbial carbon utilization in earthworm casts. Phosphorus requirements of the soil microflora were lowered by the gut passage of the soil of the third fallow site and the beechwood, which indicates an increased phosphorus supply in earthworm casts. Howerver, this additional supply was not sufficient to enable optimal carbon utilization by the soil microflora. The results indicate that the effect of decomposer invertebrates on the soil microflora depends on the nutrient status of the ecosystem.     

Patterns and mechanisms of plant succession after fire on Artemisia-grass sites in southeastern Idaho

Cover data for plant species on eight environmentally similar sites that were each burned in a different year (from 2 to 36 years ago) were used to construct a composite sequence of vegetational change after fire on Artemisia-grassland sites in southeastern Idaho. Some species were early successional such as Lithospermum ruderale, and some late successional: Artemisia tridentata, A. tripartita, and Gutierreza sarothrae. But many species: Purshia tridentata, Symphoricarpos oreophilus, Amelanchier alnifolia, Chrysothamnus viscidiflorus, Achillea millefolium, Agropyron dasystachyum, and A. spicatum were present in both early and later stages. Shannon and Simpson indices of diversity and species richness indicated little change in alpha diversity through time. This was attributed mainly to the limited change in species composition from early to later stages. The general pattern of succession is compatible with the tolerance model of Connell & Slatyer (1977) in most respects. Species traits relating to persistence through a disturbance or re-establishment on the site, and tolerance of competition shape the course of succession on a site. Perennial grasses and forbs which sprout from the base after fire are the first species to dominate the sites. Sprouting shrubs, which require some years to regrow to their pre-fire form, are prominent by the sixth year. Shrubs which rely on dispersal become co-dominants in later stages, at which time some herbaceous species are reduced oreliminated. The pattern of succession can differ due to presence or absence of species with particular traits.     

Colonization and early succession on anthropogenic soils

Abstract. This paper reports on vegetation development on permanent experimental plots during five years of succession. Nine (1 m²) plots were filled with three typical substrates from man-made habitats of urban and industrial areas in the region of Berlin. The three substrates (a commercial ‘topsoil’, a ruderal ‘landfill’ soil and a sandy soil), differ in organic matter and nutrient contents. Relevés of species composition and percent cover of each species present were made monthly during the growing season from the start of vegetation development. This paper describes the different successional pathways on topsoil and ruderal soil and the colonization process on sandy soil. On topsoil, ruderal annuals are dominant in the first year and are replaced by short-lived perennials from the second year. Those species were replaced by long-lived perennial herbs (Ballota nigra or Urtica dioica) from the third year of succession onwards. On the ruderal land-fill soil the early successional stages are less sharp and the perennial Solidago canadensis is able to dominate within one year after the succession was initiated. On sandy soil there is still an ongoing colonization process, where pioneer tree species like Betula pendula and Populus nigra play a main role. The importance of ‘initial floristic composition’, the role of substrate for community structure and the peculiarities of successional sequences on anthropogenic soils in the context of primary and secondary successions are discussed.     

Nutrient dynamics in milpa agriculture and the role of weeds in initial stages of secondary succession in Belize,C.A.

Nutrient levels, soil moisture and soil organic matter were monitored monthly during a three year period in a site cleared for shifting agriculture (milpa) and in adjacement undisturbed high bush forest soils in Belize. Throughout the study there was no evidence of a decline in available nutrient levels in the cleared site. Sharp changes in P and Mn were considered indicative of wetting and drying periods. Corn yields for the rainy season crop in the first two years were average for the region ca 2500 kg ha⁻¹ dry wt, but declined dramatically in the third year. Total weed biomass for the rainy season crop conversely increased every year and reduced grain yield probably through competition for nutrients (especially N), light and water. Substantial amounts of nutrients were also immobilized in the weed biomass, which served the agroecological purpose of preventing nutrient loss by leaching.     

华北盐渍化改造区蚯蚓种群次生演替与生产投入的关系——以河北省曲周试区为例

通过对河北省曲周县盐渍化改造区农田生态系统中蚯蚓物种多样性分析,发现蚯蚓种群的次生演替与农田肥料投稿有很好的相关性,合理的投稿特别是有机肥的投入可以加速农田生态系统中生态演替;该地区蝗蚓种群分布总的趋势是：第2代试区＞第1代试区＞第3代试区＞原貌区;不同土壤利用方式蚯蚓的分布和数量各不相同,菜地＞农田、边缘地和果园。     

Plant succession and rhizosphere microbial communities in a recently deglaciated alpine terrain

This study describes how early and late successional plant species affect soil microorganisms in alpine ecosystems. We quantify the relative importance of plant species and soil properties as determinants of belowground microbial communities. Sixteen plant species were selected from six successional stages (4–14–20–43–75–135 years) within the foreland of the Rotmoosferner glacier, Austria, and at one (reference) site outside the foreland. The size, composition and function of the communities of microorganism in the bulk soil and the rhizosphere were characterized by ninhydrin-reactive nitrogen, phospholipid fatty acids and enzyme activities (β-glucosidase, β-xylosidase, N-acetyl-β-glucosaminidase, leucine aminopeptidase, acid phosphatase, sulphatase). The results show that the microbial data could be grouped according to early (up to 43 years) and late-colonizing plant species (75 or more years). In early succession, no plant species or soil age effect was detected on the microbial biomass, phospholipid fatty acids, or enzyme activity. The rhizosphere microbial community was similar to that in the bulk soil, which in turn was determined by the abiotic environmental conditions. In late succession, improved soil conditions probably mediated plant species effects on the belowground microbial community. The most pronounced rhizosphere effects were attributed to plant species of the 75- and 135-year-old sites. The microbial colonization (size, composition, activity) of the bulk soil predominantly followed changes in vegetation cover, plant life forms and soil organic matter. In summary, the observed successional pattern of the above- and belowground communities provides an example of the facilitation models of primary succession.     

Bacterial diversity in relation to secondary production and succession on surfaces of the kelp Laminaria hyperborea

Kelp forests worldwide are known as hotspots for macroscopic biodiversity and primary production, yet very little is known about the biodiversity and roles of microorganisms in these ecosystems. Secondary production by heterotrophic bacteria associated to kelp is important in the food web as a link between kelp primary production and kelp forest consumers. The aim of this study was to investigate the relationship between bacterial diversity and two important processes in this ecosystem; bacterial secondary production and primary succession on kelp surfaces. To address this, kelp, Laminaria hyperborea, from southwestern Norway was sampled at different geographical locations and during an annual cycle. Pyrosequencing (454-sequencing) of amplicons of the 16S rRNA gene of bacteria was used to study bacterial diversity. Incorporation of tritiated thymidine was used as a measure of bacterial production. Our data show that bacterial diversity (richness and evenness) increases with the age of the kelp surface, which corresponds to the primary succession of its bacterial communities. Higher evenness of bacterial operational taxonomical units (OTUs) is linked to higher bacterial production. Owing to the dominance of a few abundant OTUs, kelp surface biofilm communities may be characterized as low-diversity habitats. This is the first detailed study of kelp-associated bacterial communities using high-throughput sequencing and it extends current knowledge on microbial community assembly and dynamics on living surfaces.     

Nitrogen-fixers <Emphasis Type="Italic">Alnus</Emphasis> and <Emphasis Type="Italic">Lupinus</Emphasis> influence soil characteristics but not colonization by later successional species in primary succession on Mount St. Helens

Changes to the primary successional environment caused by colonizing plants that present symbiotic associations with nitrogen-fixing bacteria were investigated at two areas on Mount St. Helens. One area was occupied by alder (Alnus viridis) thickets and old lupine (Lupinus lepidus) patches and the other area by young lupine patches and pumice barrens. Alder thicket soils had higher levels for a few soil nutrients and had greater cover by other pioneer species as compared to old lupine patches. Many soil nutrients, including nitrogen and soil organic matter, were below detection limits in old lupine patches but not in alder thicket soils. Young lupine patch soils were generally not different from barren site soils but had greater cover by other pioneer species. Below detection nitrogen and soil organic matter levels also occurred in many barren soil samples but not in young lupine patch soils. Barren soils were moister than were the other sites. The apparent increase in soil fertility has not led to invasion by later successional species, perhaps due to dry conditions or to other inhibitory factors. Seedbanks, composed of early successional species, appear to be developing in these areas.     

Life history and secondary production of Glossosoma nigrior Banks (Trichoptera: Glossosomatidae) in two Alabama streams with different geology

We studied life history and secondary production of a caddisfly scraper, Glossosoma nigrior, in two Alabama streams. Collier Creek, located within the Appalachian Plateau physiographic province, is underlain by sandstone bedrock, while Hendrick Mill Branch is located in the Valley and Ridge physiographic province with limestone bedrock. G. nigrior populations in both streams exhibited trivoltine life histories, which were attributed to the higher water temperature regimes than those found in more northern streams. Mean larval density (556 m⁻²) and biomass (B) (49.2 mg AFDM m⁻²) were much higher in Hendrick Mill Branch than Collier Creek (78 m⁻² and 6.7 mg AFDM m⁻²). G. nigrior in Hendrick Mill Branch maintained continuous larval growth and higher larval density than Collier Creek throughout the year mainly due to a greater availability of optimal habitat, a more stable hydrology, and warmer winter water temperature. These factors also resulted in the much higher annual secondary production (P) in Hendrick Mill Branch (965 mg AFDM m⁻²; P/B = 18.3) than Collier Creek (115 mg AFDM m⁻²; P/B = 17.9). Gut content analysis revealed that algae (>50%) and detritus (>40%) were the major diet items for G. nigrior, and the majority of secondary production (>80%) was contributed by the consumption of algae. Glossosoma populations play an important role in trophic linkage in these streams with their high production and grazing activities. Handling editor: D. Dudgeon     

Experimental increase of flying costs in a pelagic seabird: effects on foraging strategies,nutritional state and chick condition

A central point in life history theory is that parental investment in current reproduction should be balanced by the costs in terms of residual reproductive value. Long-lived seabirds are considered fixed investors, that is, parents fix a specific level of investment in their current reproduction independent to the breeding requirements. We tested this hypothesis analysing the consequences of an experimental increase in flying costs on the foraging ecology, body condition and chick condition in Cory’s shearwaters Calonectris diomedea. We treated 28 pairs by reducing the wing surface in one partner and compared them with 14 control pairs. We monitored mass changes and incubation shifts and tracked 19 foraging trips per group using geolocators. Furthermore, we took blood samples at laying, hatching and chick-rearing to analyse the nutritional condition, haematology, muscle damage and stable isotopes. Eighty-day-old chicks were measured, blood sampled and challenged with PHA immune assay. In addition, we analysed the effects of handicap on the adults at the subsequent breeding season. During incubation, handicapped birds showed a greater foraging effort than control birds, as indicated by greater foraging distances and longer periods of foraging, covering larger areas. Eighty-day-old chicks reared by treated pairs were smaller and lighter and showed a lower immunity than those reared by control pairs. However, oxygen demands, nutritional condition and stable isotopes did not differ between control and handicapped birds. Although handicapped birds had to increase their foraging effort, they maintained physical condition by reducing parental investment and transferred the experimentally increased costs to their partners and the chick. This result supports the fixed investment hypothesis and is consistent with life history theory.     

The challenge of plant regeneration after fire in the Mediterranean Basin: scientific gaps in our knowledge on plant strategies and evolution of traits

Though observations on re-colonisation of post-fire sites in the Mediterranean Basin are plentiful, there still is an ongoing debate on the interrelation of fire regimes and species traits related to fire adaptation. Most of the studies found are restricted to particular species or claim to present community attributes. Therefore they often lack information for the evaluation of evolutionary evidence and historical contingency of the local fire regime and other abiotic conditions, which may act as selective pressure for plant regeneration strategies. Indeed, knowledge about the success of regeneration mechanisms and their interrelation with ecological factors is essential for the interpretation of the high spatio-temporal variability found in post-fire species performance. Such knowledge would be necessary to assess the potential of different regeneration mechanisms to cope with ongoing land-use and climate change—a crucial scientific challenge. A summary is given of the knowledge about the limits and potential of plant regeneration mechanisms after fire in the Mediterranean Basin, along with corresponding studies conducted in other parts of the world with similar climatic conditions in order to present the fullest possible picture. Moreover, the positive or negative impacts of particular parameters of a fire regime on different regeneration strategies (post-fire seeders, resprouters, and facultative resprouters) are explained and discussed in the light of published literature. To conclude, reference is made to scientific gaps that need to be filled in order to analyse species resistance and community resilience absorbing possible climate or land use changes.     

Plant succession and interaction between soil and plants after land reclamation on the west coast of Korea

Plant succession and the interaction between soil and plants after reclamation were investigated on the west coast of Korea. Our study included one natural tidal flat site (Namdong, 3 km²), and five sites that differed in the number of years since being reclaimed: Hyundai A, 6 km² (1 yr); Hyundai B, 5 km² (2 yr); Jangdeog, 5 km² (8 yr); Mado, 2.5 km² (12 yr); and Baegseog, 1 km² (30 yr). The number of plant species occurring at each site was 10, 9, 15, 30, 28, and 38, respectively. Based on distribution ranges of the plants along gradients of salt and moisture contents in the soil, and species associations and life forms, plant succession was divided into two sere groups, hydrarch and xerarch. The major species of the former werePhragmites communis andTypha angustata, but the sequence of the sere could not be identified. Species associated with the latter were [Suaeda japonica] → [Salicornia herbacea, S. japonica, Atriplex subcordata, Suaeda asparagoides] → [Aster tripolium, Carex scabrifolia, Zoysia sinica, Limonium tetragonum] → [Artemisia scoparia, Calamagrostis epigeios, andSetaria viridis] → [Imperata cylindrica var.koenigii, Sonchus brachyotus, S. viridis] → [Aeschynomene indica, Lotus corniculatus var.japonicus, Trifolium repens, and other non-halo-phytes]. In certain circumstances, the first and second stages replacedS. japonica withC. scabrifolia andZ sinica. The progression of interactions between soil and plants through succession was salt leaching → increase in species richness and biomass → increase in soil organic matter → increase in total nitrogen and decrease in bulk density of soil, and/or salt leaching → increase in phytomass → decrease in soil-available phosphorus.     

Subalpine vegetation pattern three decades after stand‐replacing fire: effects of landscape context and topography on plant community composition,tree regeneration,and diversity

Objective: Our purpose was to characterize vegetation compositional patterns, tree regeneration, and plant diversity, and their relationships to landscape context, topography, and light availability across the margins of four stand‐replacing subalpine burns. Location: Four 1977 to 1978 burns east of the Continental Divide in Colorado: the Ouzel burn, a burn near Kenosha Pass, the Badger Mountain burn, and the Maes Creek burn. Methods: Vegetation and environmental factors were sampled in 200 0.01‐ha plots on transects crossing burn edges, and stratified by elevation. We utilized dissimilarity indices, mixed‐effects models, and randomization tests to assess relationships between vegetation and environment. Results: Three decades after wildfire, plant communities exhibited pronounced compositional shifts across burn edges. Tree regeneration decreased with increasing elevation and distance into burn interiors; concomitant increases in forbs and graminoids were linked to greater light availability. Richness was roughly doubled in high‐severity burn interiors due to the persistence of a suite of native species occurring primarily in this habitat. Richness rose with distance into burns, but declined with increasing elevation. Only three of 188 plant species were non‐native; these were widespread, naturalized species that comprised <1% total cover. Conclusions: These subalpine wildfires generated considerable, persistent increases in plant species richness at local and landscape scales, and a diversity of plant communities. The findings suggest that fire suppression in such systems must lead to reduced diversity. Concerns about post‐fire invasion by exotic plants appear unwarranted in high‐elevation wilderness settings.     

A model analysis of N and P limitation on carbon accumulation in Amazonian secondary forest after alternate land-use abandonment

Productivity and carbon (C) storage in many mature tropical forests are considered phosphorus (P) limited because of advanced soil weathering. However, disturbance can shift limitation away from P and toward nitrogen (N) because of disproportionately large N losses associated with its mobility relative to P in ecosystems. This shift was illustrated by model analyses in which large disturbances including timber extraction and slash-burn were simulated in a P-limited tropical forest. Re-accumulation of ecosystem C during secondary forest growth was initially N-limited, but long term limitation reverted to P. Mechanisms controlling shifts between N and P limitation included: (1) N volatility during slash combustion produced ash that increased soil solution P more than N, (2) a wide N:P ratio in residual fuel and belowground necromass relative to soil organic matter (SOM) N:P produced a simultaneous P sink and N source during decomposition, (3) a supplemental (to aerosol deposition) external N source via biological N fixation. Redistribution of N and P from low C:nutrient SOM to high C:nutrient vegetation was the most important factor contributing to the resilience of ecosystem C accumulation during secondary growth. Resilience was diminished when multiple harvest and re-growth cycles depleted SOM. Phosphorus losses in particular resulted in long-term reductions of C storage capacity because of slow re-supply rates via deposition and the absence of other external sources. Sensitivity analyses limiting the depth of microbially active SOM in soil profiles further illustrated the importance of elements stored in SOM to ecosystem resilience, pointing to a need for better knowledge on the functioning of deeply buried SOM.