Microsite and elevational influences on early forest regeneration after catastrophic windthrow

Abstract. We compared vegetation establishment in 25 treefall pits and mounds along a hillside elevational gradient in a fourth-year catastrophic windthrow in eastern North America. Plant communities differed greatly between pits and mounds, with pit microsites having significantly greater species richness, total biomass, and total tree stem density. Species richness in pits and on mounds decreased with increasing elevation from the bottom of the hillside, although the effect of elevation on mound species richness was less than that of elevation on pit species richness. Biomass of Erechtites hieraciifolia decreased significantly, while that of Betula alleghaniensis increased significantly with elevation. However, total biomass of both pit and mound microsites was unrelated to elevation. Total stem density decreased with elevation in pits, but was unaffected by elevation on mounds. This study shows that both small-scale (microsite) effects and intermediate-scale effects influence the re-establishment of plant communities within this catastrophic windthrow. Consideration of both microsite and position along intermediate-scale gradients may allow more precise prediction of plant community composition and dynamics in recovery of disturbed areas.     

红松阔叶混交林不同大小林隙内丘坑复合体微气候动态变化

2012年5月,在小兴安岭凉水国家级自然保护区阔叶红松混交林2.55 hm²的固定样地内,调查了由掘根风倒形成的38对丘坑复合体所处的7个小林隙、5个中林隙和3个大林隙以及7个郁闭林分的基本状况.于2012年6—9月,每月选定6个典型晴天,测定处于大林隙、中林隙和小林隙以及郁闭林分内每个丘坑复合体不同微立地(坑底、坑壁、丘顶、丘面及完整立地)的土壤温度、土壤含水量和空气相对湿度.结果表明: 6—9月,丘顶的土壤温度平均值最大,坑底最小;坑底土壤含水量和空气相对湿度的平均值最大,丘顶最小.上述指标在大多数微立地之间差异显著.6—9月,位于不同大小林隙和郁闭林分的丘坑复合体土壤温度总平均值依次为:大林隙>中林隙>小林隙>郁闭林分;各月份土壤水分大小次序并不一致;6月、8月和9月丘坑复合体各个微立地月均空气相对湿度大小顺序均为郁闭林分>小林隙>中林隙>大林隙,7月的排列次序有所不同.上述指标在不同大小林隙及郁闭林分内丘坑复合体大多数微立地之间差异显著.不同大小林隙和郁闭林分内丘坑复合体各微立地月均土壤温度和空气相对湿度均为7月最大,9月最小;除完整立地6月月均土壤含水量最大以外,其余微立地均为7月最大,9月最小.丘坑复合体微气候的变化主要受林隙大小、微立地和时间等的影响.     

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

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

The effects of windthrow on plant species richness in a Central European beech forest

The effects of soil disturbance caused by the uprooting of a single or a few canopy trees on species richness and composition of vascular plant species and bryophytes were examined in a temperate beech forest (Fagus sylvatica) in northern Germany. We recorded the vegetation in 57 pairs of disturbed and adjacent undisturbed plots and established a chronosequence of mound ages to study the effect of time since microsite formation on plant species richness and composition. We found significant differences in plant species richness and composition between disturbed and adjacent undisturbed plots. Species richness of both vascular plants and bryophytes was higher in the disturbed than in the undisturbed plots, but these differences were more pronounced for bryophytes. We suggest that three main factors are responsible for this differential response. The availability of microsites on the forest floor that are suitable for the recruitment of bryophytes is lower than for vascular plants. Establishment of bryophytes in disturbed microsites is favoured by a greater abundance of propagules in the close vicinity and in the soil of the disturbed microsites, as well as by a greater variety of regeneration strategies in bryophytes than in vascular plants. Time since mound formation was a major factor determining plant species richness and composition. A significant decrease in the mean number of species was found from young mounds to intermediate and old mounds. However, differences were observed between vascular plants and bryophytes in the course of changes through time in species richness and composition. A large number of exclusive and infrequent vascular plant species was observed on young mounds, among them several disturbance specialists. We suggest that the establishment of many vascular plant species was infrequent and short-lived due to unfavourable light conditions and a low abundance of propagules. By contrast, the development of a litter layer was the main reason for the decreased mean number of bryophytes on old mounds. Our study supports the view that groups of species differing in important life history traits exhibit different responses to soil disturbance.     

不同林型树倒林隙内微立地类型的土壤微团聚体组成及其分形特征

采用野外调查和室内分析相结合的方法,分析小兴安岭地区阔叶红松林和云冷杉红松林两种林型下树倒林隙内丘坑微立地类型(丘顶和坑底)土壤微团聚体组成及其分形特征。结果表明: 两种林型的土壤微团聚体0.25～2 mm和0.05～0.25 mm粒级含量较高,分别为25.7%～50.7%和27.0%～42.8%,<0.002 mm粒级含量最小,为4.4%～8.9%。在两种林型的树倒林隙内,坑底和丘顶的土壤容重较大,且丘顶土壤养分含量均高于坑底。阔叶红松林树倒林隙内丘顶和坑底的土壤养分含量均高于云冷杉红松林。<0.002 mm土壤微团聚体与土壤物理和化学性质均无相关性,0.25～2 mm和0.002～0.02 mm土壤微团聚体分别与土壤非毛管孔隙度、总孔隙度、通气度、有机质、全磷、全氮和有机碳之间呈极显著正相关和极显著负相关。整体来看,阔叶红松林的土壤分形维数(D)和土壤特征微团聚体组成比例(PCM)大于云冷杉红松林。两种林型下,丘顶和坑底的土壤特征微团聚体粒级比值(RMD)均增高。土壤D和PCM与各土壤理化性质均无显著相关性,土壤RMD与土壤毛管孔隙度、总孔隙度、土壤容重和通气度呈显著负相关。丘顶和坑底微立地的形成会导致土壤较大粒级微团聚体减少,土壤微团聚体稳定性降低,土壤D和PCM增加,RMD显著增加;土壤RMD可作为定量化描述不同林型下丘坑微立地内土壤理化性质的指标。     

Mounding alters environmental filters that drive plant community development in a novel grassland

Earthen mounds are commonly used in ecological restoration to increase environmental heterogeneity, create favorable microclimates and retain soil resources that promote plant establishment. Although mounding is commonly employed in restoration, few microtopography studies focus on the long-term effects of mounding on restored plant community development. We assessed the vegetation and physical environment of earthen mounds installed at a novel grassland ten years after restoration to look for patterns in plant community development. We used permutational multiple analysis of variance (PERMANOVA) to identify differences in plant community composition and the associated mound-driven environmental variables, summer soil moisture and height above peak soil inundation, in relation to mound position. We used indicator species analysis (ISA) to classify the species that defined mound top and intermound space plant communities. We found that mound position drove plot height above flooding and soil moisture while plant community composition was driven by plot height above flooding, summer soil moisture, and mound position. ISA showed that species colonized mound microsites differently: most wetland species occurred between mounds and xeric stress tolerators largely occupied dry mound tops. We visualized these differences with non-metric multidimensional scaling (NMDS) ordination, finding that species sorted out in multivariate space based on mound position. We conclude that mounding can have relatively long-term effects on plant community development, even in highly disturbed, minimally maintained restoration projects.     

Gopher mounds decrease nutrient cycling rates and increase adjacent vegetation in volcanic primary succession

Fossorial mammals may affect nutrient dynamics and vegetation in recently initiated primary successional ecosystems differently than in more developed systems because of strong C and N limitation to primary productivity and microbial communities. We investigated northern pocket gopher (Thomomys talpoides) effects on soil nutrient dynamics, soil physical properties, and plant communities on surfaces created by Mount St. Helens’ 1980 eruption. For comparison to later successional systems, we summarized published studies on gopher effects on soil C and N and plant communities. In 2010, 18 years after gopher colonization, we found that gophers were active in ~2.5 % of the study area and formed ~328 mounds ha^?1. Mounds exhibited decreased species density compared to undisturbed areas, while plant abundance on mound margins increased 77 %. Plant burial increased total soil carbon (TC) by 13 % and nitrogen (TN) by 11 %, compared to undisturbed soils. Mound crusts decreased water infiltration, likely explaining the lack of detectable increases in rates of NO₃–N, NH₄–N or PO₄–P leaching out of the rooting zone or in CO₂ flux rates. We concluded that plant burial and reduced infiltration on gopher mounds may accelerate soil carbon accumulation, facilitate vegetation development at mound edges through resource concentration and competitive release, and increase small-scale heterogeneity of soils and communities across substantial sections of the primary successional landscape. Our review indicated that increases in TC, TN and plant density at mound margins contrasted with later successional systems, likely due to differences in physical effects and microbial resources between primary successional and older systems.     

Soil-foraging animals alter the composition and co-occurrence of microbial communities in a desert shrubland

Animals that modify their physical environment by foraging in the soil can have dramatic effects on ecosystem functions and processes. We compared bacterial and fungal communities in the foraging pits created by bilbies and burrowing bettongs with undisturbed surface soils dominated by biocrusts. Bacterial communities were characterized by Actinobacteria and Alphaproteobacteria, and fungal communities by Lecanoromycetes and Archaeosporomycetes. The composition of bacterial or fungal communities was not observed to vary between loamy or sandy soils. There were no differences in richness of either bacterial or fungal operational taxonomic units (OTUs) in the soil of young or old foraging pits, or undisturbed soils. Although the bacterial assemblage did not vary among the three microsites, the composition of fungi in undisturbed soils was significantly different from that in old or young foraging pits. Network analysis indicated that a greater number of correlations between bacterial OTUs occurred in undisturbed soils and old pits, whereas a greater number of correlations between fungal OTUs occurred in undisturbed soils. Our study suggests that digging by soil-disturbing animals is likely to create successional shifts in soil microbial and fungal communities, leading to functional shifts associated with the decomposition of organic matter and the fixation of nitrogen. Given the primacy of organic matter decomposition in arid and semi-arid environments, the loss of native soil-foraging animals is likely to impair the ability of these systems to maintain key ecosystem processes such as the mineralization of nitrogen and the breakdown of organic matter, and to recover from disturbance.     

The European ground squirrel increases diversity and structural complexity of grasslands in the Western Carpathians

Disturbances are important natural factors affecting biological diversity, community composition, and ecosystem structure. The European ground squirrel is a semi-fossorial organism, and through disturbances caused by burrowing activities, it can play an important role as an ecosystem engineer of grasslands in central and south-eastern Europe. The aim of this study was to assess the response of grassland vegetation to disturbances by the European ground squirrel. We conducted a pairwise survey within a 1-ha study site with homogenous environmental conditions. We compared the vegetation characteristics of 2?×?2-m plots placed on 30 mounds, with paired control plots situated at a distance of 10 m from each mound. The results showed that plots disturbed by the European ground squirrel achieved a higher species richness and diversity and a distinct species composition compared to the undisturbed control plots. Vertical structure of vegetation was also significantly different with a higher proportion of the high and medium vegetation layers on the mounds. Shifts in the composition of plant life forms and life strategies were reflected by the reduction of graminoids and plant competitors, and support of forbs on the mounds. These findings suggest that the European ground squirrel helps to maintain heterogeneity in grassland ecosystems and creates patches of higher diversity and higher structural complexity in the relatively homogenous grassland vegetation of the Western Carpathians.     

Impact of Formica exsecta Nyl. on seed bank and vegetation patterns in a subalpine grassland ecosystem

The mound building ant Formica exsecta Nyl. is widely distributed in grassland ecosystems of the Central European Alps. We studied the impact of these ants on seed bank and vegetation patterns in a 11 ha subalpine grassland, where we counted over 700 active ant mounds. The mounds showed a distinct spatial distribution with most of them being located in tall‐grass, which was rarely visited by ungulates (red deer; Cervus elaphus L.). Heavily grazed short‐grass, in contrast, seemed to be completely avoided by ants as only few mounds were found in this vegetation type. The species composition of the ant mound and grassland seed banks was quite similar, i.e. from 15 common plant species 12 were found in both seed bank types. We found the same proportions of myrmecochorous seeds in ant mound and grassland soil samples. In contrast, the number of seeds was 15 times higher in mound compared with the grassland soil samples. Also, the vegetation growing on ant mounds significantly differed from the vegetation outside the mounds: graminoids dominated on ant mounds, herbaceous and myrmecochorous species in the grassland vegetation. We found significant continuous changes in vegetation composition on gradients from the ant mound centre to 1 m away from the mound edge. Overall, F. exsecta was found to have a considerable impact on seed bank and vegetation patterns in the grassland ecosystem studied. These insects not only altered grassland characteristics in the close surrounding of their mounds, but also seem to affect the entire ecosystem including, for example, the spatial use of the grassland by red deer.     

Temporal vegetation dynamics and recolonization mechanisms on different-sized soil disturbances in tallgrass prairie

Assessing the various mechanisms by which plants revegetate disturbances is important for understanding the effects of disturbances on plant population dynamics, plant community structure, community assembly processes, and ecosystem function. We initiated a 2-yr experiment examining temporal vegetation dynamics and mechanisms of recolonization on different-sized soil disturbances created to simulate pocket gopher mounds in North American tallgrass prairie. Treatments were designed to assess potential contributions of the seed rain, soil seed bank, clonal propagation from the edges of a soil mound, and regrowth of buried plants. Small mounds were more rapidly recolonized than large mounds. Vegetative regrowth strategies were the dominant recolonization mechanisms, while the seed rain was considerably less important in maintaining the diversity of forbs and annuals than previously believed. All recolonization mechanisms influenced plant succession, but stem densities and plant mass on soil mounds remained significantly lower than undisturbed controls after two growing seasons. Because natural pocket gopher mounds are indistinguishable from undisturbed areas after two seasons, these results suggest that multiple modes of recruitment concurrently, albeit differentially, contribute to the recolonization of soil disturbances and influence tallgrass prairie plant community structure and successional dynamics.     

Microsite and litter cover effects on seed banks vary with seed size and dispersal mechanisms: implications for revegetation of degraded saline land

Seed movements and fates are important for restoration as these determine spatial patterns of recruitment and ultimately shape plant communities. This article examines litter cover and microsite effects on seed availability at a saline site revegetated with Eucalyptus sargentii tree rows interplanted with 5?C6 rows of saltbush (Atriplex spp.). As litter accumulation decreases with increasing distance from tree rows, soil seed banks were compared between paired bare and litter-covered zones within three microsites; tree row, saltbush row closest to tree row and saltbush mid-row (middle row of saltbush between tree rows). Germinable seed banks of the four most abundant species with contrasting seed sizes and dispersal mechanisms were assessed to test the hypotheses that: (i) microsites with litter cover contain higher seed densities than bare areas, but that (ii) microsite and litter effects will vary depending on seed size and dispersal mechanisms. Overall, litter cover increased seed densities, however, litter effects varied with seed size, with no effect on small-seeded species and litter increasing densities of large-seeded species. Seed bank composition also differed between tree and shrub microsites due to differences in seed morphology and dispersal mechanisms. Water-dispersed species were unaffected by microsite but densities of wind-dispersed species, including Atriplex spp., were higher in saltbush microsites. Densities of wind-dispersed species also differed between the two saltbush microsites despite similar litter cover. Future plantings should consider row spacing and orientation, as well as the dimensions of seeding mounds and associated neighbouring depressions, to maximize litter and seed-trapping by microsites.     

Burrowing activities of kangaroo rats and patterns in plant species dominance at a shortgrass steppe-desert grassland ecotone

Abstract. Our objective was to evaluate the effects of burrowing activities by banner-tail kangaroo rats (Dipodomys spectabilis Merriam) on plant community structure and species dominance for two patch types at the ecotone between shortgrass steppe and desert grassland in New Mexico, USA. 10 mounds produced by kangaroo rats were selected in patches dominated by Bouteloua gracilis (the dominant in shortgrass steppe communities) and 10 mounds were selected in patches dominated by B. eriopoda (the dominant in Chihuahuan desert grasslands). Plant cover and density by species were sampled from three locations associated with each mound: the mound proper, the edge of the mound in the transition area, and the off-mound vegetation. Similar cover of B. eriopoda for the edges of mounds in both patch types indicates the ability of this species to respond to animal disturbances regardless of the amount of cover in the surrounding undisturbed vegetation. By contrast, cover of B. gracilis was low for all mounds and mound edges in patches dominated by this species. Much higher cover of B. eriopoda on mound edges compared to the undisturbed vegetation in B. gracilis-dominated patches indicates that kangaroo rats have important positive effects on this species. Lower cover of perennial grasses and higher cover of forbs, shrubs, and succulents on the edges of mounds in B. eriopoda-dominated patches compared to patches dominated by B. gracilis indicate the importance of surrounding vegetation to plant responses on disturbed areas. Our results show that kangaroo rats have important effects on both species dominance and composition for different patch types, and may provide a mechanism for small-scale dominance patterns at an ecotone; thus providing further support for their role as keystone species in desert grasslands.     

Origin of mound-field landscapes: a multi-proxy approach combining contemporary vegetation, carbon stable isotopes and phytoliths

Background and aims

Seasonally flooded South American savannas harbor different kinds of mound-field landscapes of largely unknown origin. A recent study used soil carbon-isotope depth profiles and other proxies to infer vegetation history in murundu landscapes in Brazil. Results suggested that differential erosion, not building-up processes (e.g., termite mounds), produced mounds. We tested this approach to inferring mound origin in a mound-field landscape in French Guiana.

Methods

We examined carbon-isotope depth profiles of soil organic matter, phytolith profiles and contemporary vegetation composition in mounds and inter-mounds.

Results

Relative abundance of C3 and C4 plants across habitats was very different from that in murundu landscapes; C3 plants were better represented in inter-mounds than on mounds. Habitat differences in C3/C4 distribution were subtler than in murundu landscapes, limiting inference of vegetation history based on carbon isotopes. Still, carbon-isotope and phytolith depth profiles gave similar pictures of vegetation history, both favoring a building-up hypothesis, corroborating other evidence that these mounds are vestiges of ancient agricultural raised fields.

Conclusions

Carbon-isotope depth profiles are unlikely to be adequate for deciphering origin of mound-field landscapes from vegetation history in seasonally flooded savannas. Including data on current vegetation and phytoliths makes inferences more robust.     

Seedling and Sapling Dynamics of Treefall Pits in Puerto Rico1

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N–fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N–fixing sym–bionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer–term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.     

Spatial distribution of ant mounds and effects on soil physical properties in wetlands of the Sanjiang plain, China

Ants constitute a dominant element of soil mesofauna due to their biomass, abundance, richness of species and distribution within terrestrial ecosystems. They are important regulators of soil aggregate structure as they translocate large amounts of soil from the bottom to the soil surface. In doing so, they form biogenic structures (BS) made up of aggregates of different sizes and characteristics, i.e. ant mounds. These BS have varying characteristics according to the ant species and the soil where they carry their activities. Ants are considered soil engineers because of their effects on soil properties, availability of resource and flow of energy and nutrients in soil. Thus, it is important to gain information on their distribution and abundance. Relatively little is known about the spatial distribution of mounds and their role in the soil physical properties in wetlands of the Sanjiang plain, China. We conducted a survey of ant mounds and measured the density, height, and diameter and material composition of different ant mounds. The ecological characteristics of wetlands that ant mounds wide occurrence were also investigated, including soil type, hydrology characters and plant composition. Differences in soil particle composition, bulk density and soil moisture between ant mound and natural meadow were measured to assess the influences of ant mounds on soil physical properties. We also studied the effects of ant mounds on the microtopography of meadows. Ant mounds were found mainly in the transition zone between terrestrial and aquatic habitats, with wetland type, including Calamagrostis augustifolia wet meadow, C.augustifolia marsh meadow, shrubs marsh meadow and Carexmeyeriana–Carexappendiculata wetland, being a significant factor. Most of the mounds detected were inhabited by Lasius flavus Fabricius, Lasius niger Linnaeus and Formica sanguinea Latreille, which occupied 52.9%, 26.5% and 20.6% of the mounds surveyed, respectively. The density, height, diameter and mound composition were significantly different among the mounds of F. sanguinea Latreille, L. flavus Fabricius and L. niger Linnaeus. The average density and diameter of L. flavus mounds was significantly higher than those of other ant mounds. The average height of F. sanguinea mounds was highest among the mounds detected. Mound building activities changed soil particle size distribution, with the silt and clay content of mounds higher than for non-mound soil. Compared with adjacent, non-mound soil, the bulk density (0–30 cm) and water content (0–25 cm) of mound soil were significantly lower, but there were no significant differences between the mound soil of F. sanguinea Latreille and L. flavus Fabricius. The spatial distribution of ant mounds with different height and diameter also changed the micro-geomorphology of the soil surface, increasing the degree of fluctuation of the microtopography. The ant distribution characteristics and their ecological roles respond to a wide range of environmental alterations. The biogenic structures of ant and the specific environment associated with them have been defined as the “functional domain”, a sphere of influence that may significantly affect soil processes at certain spatial and temporal scales. Our results suggest that the distribution and structure of ant mounds can indicate wetland environmental changes, with mounds influencing ecosystem functions and enhancing wetland degradation.     

Forest microsite effects on community composition of ectomycorrhizal fungi on seedlings of Picea abies and Betula pendula

Niche differentiation in soil horizons, host species and natural nutrient gradients contribute to the high diversity of ectomycorrhizal fungi in boreal forests. This study aims at documenting the diversity and community composition of ectomycorrhizal fungi of Norway spruce ( Picea abies ) and silver birch ( Betula pendula ) seedlings in five most abundant microsites in three Estonian old-growth forests. Undisturbed forest floor, windthrow mounds and pits harboured more species than brown- and white-rotted wood. Several species of ectomycorrhizal fungi were differentially represented on either hosts, microsites and sites. Generally, the most frequent species in dead wood were also common in forest floor soil. Ordination analyses suggested that decay type determined the composition of EcM fungal community in dead wood. Root connections with in-growing mature tree roots from below affected the occurrence of certain fungal species on seedling roots systems in dead wood. This study demonstrates that ectomycorrhizal fungi differentially establish in certain forest microsites that is attributable to their dispersal and competitive abilities. Elevated microsites, especially decayed wood, act as seed beds for both ectomycorrhizal forest trees and fungi, thus affecting the succession of boreal forest ecosystems.     

Vegetation of ant-hills in a mountain grassland:effects of mound history and of dominant ant species

Vegetation in grasslands with well-developed long-lastingant-hills in the Slovenské Rudohorie Mts., Slovakia, was studiedin relation to (i) position on the mound, (ii) ant speciesforming the mound, and (iii) history of the mound. Permanent plotrecordings of mound size and dominant ant species started fifteen years priorthe study began provided information on the history of individual mounds.The mound vegetation bears a striking similarity to vegetation insimilar habitats across a large part of Europe due to presence of species suchas Agrostis capillaris, Dianthusdeltoides, Polytrichum commune agg.,Thymus pulegioides, and Veronicaofficinalis. Out of the three major ant species-groups presentat the site (Lasius flavus, Tetramoriumcaespitum and Formica spp.), L.flavus had the most pronounced and the most lasting effect on themound vegetation. The dominance of the plant species listed above increased withthe time span over which the mound was inhabited by L.flavus. The effects of other species on vegetation composition,though discernible from short-term observation, disappeared over severalyears. The mounds proper did not differ from the undisturbed grassland in theproportion of myrmecochorous plants or plants with specific seed size ordormancy type. However, there was a highly significant concentration ofmyrmecochorous plants in the grassland patches immediately neighbouring themounds; this is likely to be due to seeds deposited there by the workers fromthe nest after the elaiosomes had been consumed. The mound vegetation wascomposed mainly of species with long stolons or rhizomes; however, there was nosignificant difference in formation type or length of stolons/rhizomesbetween mounds and the rest of the grassland or among mounds formed by differentant species.     

The role of a mound-building ecosystem engineer for a grassland butterfly

Both land use intensification and abandonment within grasslands lead to a homogenisation of vegetation structure. Therefore, specially structured microsites such as vegetation gaps with bare ground play an important role for species conservation within grasslands. Vegetation gaps are crucial for the establishment of low-competitive plant species and offer special microclimatic conditions essential for the development of the immature stages of many invertebrate species. The influence of small-scale soil disturbance in the form of mounds created by ecosystem engineers such as ants or moles on biodiversity is therefore of special scientific concern. The effects of mound-building species on plant species diversity have been extensively studied. However, knowledge on the significance of these species for the conservation of other animals is rare. In this study we analyse the importance of mounds created by the European mole (Talpa europaea) as an oviposition habitat for the small copper (Lycaena phlaeas) within Central European mesotrophic grasslands. Our study showed that host plants occurring at molehills were preferred for oviposition. Oviposition sites were characterised by an open vegetation structure with a high proportion of bare ground (with a mean coverage of about 50 %), a low cover of herbs and low-growing vegetation (mean height: 4.5 cm). Our study clearly illustrates the importance of small-scale soil disturbance for immature stages of L. phlaeas and the conservation of this species within mesotrophic grasslands. Mound-building ecosystem engineers, such as T. europaea, act as important substitutes for missing dynamics within mesotrophic grasslands by diversifying vegetation structure and creating small patches of bare soil.     

Vole mound effects and disturbance rate in a mediterranean plant community under different grazing and irrigation regimes

A factorial field experiment was used to assess the influence of soil-disturber mammals in the structure of a 9-year-old Mediterranean annual plant community subjected to different sheep grazing and irrigation regimes. We estimated the disturbance rate (mound building activity) by Mediterranean voles, their effects on vegetation and the mechanisms of these effects during a period of vole outbreak. The effects on vegetation were analysed at the levels of species, functional groups and plant community. Disturbance rate was high and voles can disturb the entire soil surface once every four or five years. The availability of certain trophic resources (perennial plants) appeared to drive vole expansion in the experimental plots and it was independent of the irrigation and grazing treatments. Mound building activities largely affected vegetation but conserved plot differences. Total vegetation cover, absolute cover of all functional groups, mean vegetation height and species richness were less on mounds than on undisturbed ground. These effects did not change the relative abundance of annuals, perennials, grasses and forbs. Only the relative abundance of small-seeded species decreased on mounds. As the proportion of these seeds was similar in both types of patches, we suggest that small-seeded species had more difficulties for germinating or emerging when they are buried during mound formation. Irrigation and sheep grazing promoted large changes in the vegetation parameters but these effects were, in general, similar on mounds and undisturbed ground. Our results show that the availability of germinable seeds may be the major limitation for mound revegetation, probably due to the scarcity of seeds existing at the depths from which soils are excavated. Our results also suggested a resource limitation on mounds. The results provide additional evidence that soil disturbances by small herbivore mammals exert relevant ecological effects on abandoned Mediterranean croplands. We discuss the ecological implications of vole mound-building activities for plant succession, plant species conservation and forage resource availability for livestock. This revised version was published online in June 2006 with corrections to the Cover Date.