The impact of vertic soil cracks on submerged macrophyte diaspore bank depth distribution in Mediterranean temporary wetlands

In temporary wetlands, a majority of diaspores are located in the upper layers of the soil. In general, it is assumed that the vertical movement of diaspores is caused by animal disturbance, and that reduction in this activity concentrates the diaspores in the topsoil. The occurrence of seasonal cracks and the dispersion of topsoil during dry periods are processes that have been underestimated in diaspore bank studies. Our results indicated that diaspore size can partially explain the density of diaspores of submerged macrophytes observed at four discrete soil depths in Doñana National Park temporary marshes (SW Spain). The top layers of the soil exhibit the largest number of species. Spore density of Riella spp., a liverwort, increases with depth, creating an important reserve of viable diaspores in deep soil. Downward movement of diaspores in soil cracks during dry periods, and the extended longevity of Riella diaspores together explain this inverse depth distribution.     

The impact of vertic soil cracks on submerged macrophyte diaspore bank depth distribution in Mediterranean temporary wetlands

In temporary wetlands, a majority of diaspores are located in the upper layers of the soil. In general, it is assumed that the vertical movement of diaspores is caused by animal disturbance, and that reduction in this activity concentrates the diaspores in the topsoil. The occurrence of seasonal cracks and the dispersion of topsoil during dry periods are processes that have been underestimated in diaspore bank studies. Our results indicated that diaspore size can partially explain the density of diaspores of submerged macrophytes observed at four discrete soil depths in Doñana National Park temporary marshes (SW Spain). The top layers of the soil exhibit the largest number of species. Spore density of Riella spp., a liverwort, increases with depth, creating an important reserve of viable diaspores in deep soil. Downward movement of diaspores in soil cracks during dry periods, and the extended longevity of Riella diaspores together explain this inverse depth distribution.     

Soil seed banks near rubbing trees indicate dispersal of plant species into forests by wild boar

Current knowledge about processes that generate long-distance dispersal of plants is still limited despite its importance for persistence of populations and colonization of new potential habitats. Today wild large mammals are presumed to be important vectors for long-distance transport of diaspores within and between European temperate forest patches, and in particular wild boars recently came into focus. Here we use a specific habit of wild boar, i.e. wallowing in mud and subsequent rubbing against trees, to evaluate epizoochorous dispersal of vascular plant diaspores. We present soil seed bank data from 27 rubbing trees versus 27 control trees from seven forest areas in Germany. The mean number of viable seeds and the plant species number were higher in soil samples near rubbing trees compared with control trees. Ten of the 20 most frequent species were more frequent, and many species exclusively appeared in the soil samples near rubbing trees. The large number of plant species and seeds – more than 1000 per tree – in the soils near rubbing trees is difficult to explain unless the majority were dispersed by wild boar. Hooked and bristly diaspores, i.e. those adapted to epizoochory, were more frequent; however, many species with unspecialized diaspores occurred exclusively near rubbing trees. As opposed to plant species closely tied to forests species which occur both in forest and open vegetation and non-forest species were more frequent near rubbing trees compared with controls. These findings are consistent with previous studies on diaspore loads in the coats and hooves of shot wild boars. However, our method allows to identify the transport of diaspores from the open landscape into forest stands, where they might especially emerge after disturbance, and a clustered distribution of epizoochorically dispersed seeds. Moreover, accumulation of seeds of wetness indicators near rubbing trees demonstrates directed dispersal of plant species inhabiting wet places among remote wallows.     

Exploitation of Fallen Diaspores by Ants: Are there Ant–Plant Partner Choices?

Ants frequently interact with fleshy fallen diaspores (fruits or seeds) not adapted for ant‐dispersal. Such interactions are usually considered as opportunistic, but recent evidence has indicated that these ants may differ in their effects on diaspore survival and plant recruitment. We investigated if partner choices are recognizable among genera of ants and plants, and if ant and plant traits may influence such preferences in cerrado (savanna‐like vegetation) from southeast Brazil. During a 2‐yr period, 521 ant–diaspore interactions were recorded through various methods, yielding 71 ant species and 38 plant species. Exploitation of fallen diaspores was common among several ant genera, and included carnivorous, omnivorous, and fungivorous ants. Contrary to others areas around the world, where true myrmecochory (seed dispersal by ants) is common among shrubs, ants also exploited diaspores from several cerrado trees. Plant life form, diaspore size, and ant body size did not seem to explain the pattern of interactions observed. Two subsets of preferential interactions, however, segregated fungivorous ants from another group composed of carnivorous and omnivorous ants, probably influenced by the chemical composition of the plant diaspores. Omnivorous ants usually remove the fleshy portion of diaspores on spot and probably provide limited benefits to plants. Carnivorous and fungivorous ants usually remove the whole diaspore to the nest. As each of these ant groups may influence the fitness of diaspores in different ways, there are possible subtle pathways for the evolution of partner choices between ants and these non‐myrmecochorous diaspores.     

植物锥形繁殖体结构及其适应

植物的锥形繁殖体是具有吸湿芒和锐利尖端的一种繁殖体类型。锥形繁殖体结构的功能几乎在植物生活史的各个阶段上都有体现,然而,其最基本的功能是通过繁殖体的打钻作用将自身埋入土壤。从植物繁殖体的形态结构出发,对锥形繁殖体的各项功能进行了综述,重点阐述植物繁殖体打钻作用及其机制方面的研究进展,面临的主要问题和未来的研究方向。     

Plant traits: Their role in the regeneration of alpine plant communities in sub‐arctic Finland

Abstract. Question: How do the relative frequencies of plant traits (clonality, growth form, seed weight, diaspore morphology) vary during the life cycle and how does this affect regeneration? Location: Alpine meadow and heath communities at Kilpisjärvi, sub‐Arctic Finland. Methods: Control plots and three treatments were used to measure relative species abundances for five life cycle stages: standing vegetation, seed rain, seed bank and seedlings emerging in gaps and in closed vegetation. Results: The relative frequencies of plant traits varied between the life cycle stages. The meadows were dominated by weakly clonal herbs, small or intermediate seeds and unappendaged diaspores, while the heaths were dominated by clonal dwarf shrubs, small seeds and fleshy fruits. In the meadows, species with small seeds dominated during the seed rain and in the seedling stage in gaps, while species with intermediate seeds dominated the seed bank and the seedling stage in closed vegetation. Species with unappendaged diaspores dominated throughout the life cycle. In the heaths, seed bank and seedling stage were practically absent. Conclusions: The observed differences in plant trait spectra between life cycle stages indicate that important environmental factors differ among the stages. Small seeds are advantageous for dispersal, whereas intermediate seeds have a greater probability of germinating and establishing in closed vegetation. Appendages facilitate dispersal, whereas unappendaged diaspores favour seed burial. Although the plant growth form spectrum largely reflects environmental constraints during the regeneration cycle, information on seed weight and diaspore morphology improves our knowledge of the relative importance of morphological adaptations of sexual structures in different stages during the life cycle.     

黄土丘陵沟壑区80种植物繁殖体形态特征及其物种分布

植物繁殖体的形态特征及其物种分布是植物长期适应环境的结果,对退化生态系统的更新与恢复具有重要的理论与实践指导意义。研究了80种植物的繁殖体形态特征及与其物种分布的关系,结果表明：1）繁殖体重量差别较大（0. 020—357.428 mg）,主要集中在1—9.999 mg（占52.5%）。2）繁殖体的形状指数FI变化范围较大（1.06—12.93）,61.3%的植物繁殖体的形状近圆球形。3）33种植物繁殖体具有明显的附属物,包括毛、翅、芒和刺等。4）87.5%的植物繁殖体表面具有各种纹饰（棱、被毛、纹和颗粒状等）;繁殖体颜色以褐色和黑色为主,少数繁殖体颜色鲜艳;6种植物繁殖体具有吸水分泌粘液的特性。5）黄土丘陵沟壑区繁殖体重量小或形状近圆球形的植物分布数量多、范围广。6）繁殖体具有毛、翅和芒附属物或吸湿分泌粘液的植物在黄土丘陵沟壑区较繁殖体无附属物的植物能够分布更多、更广。繁殖体这些有利于物种分布的形态特征对于植物适应该区干旱与土壤侵蚀干扰具有重要的生态学意义,而且可用于指导该区人工补播促进植被恢复的物种选择。     

Interaction Between “Safe Sites” and “Safe Sides” for Germination of Neurada procumbens (Neuradaceae) in the Middle East

The seed bank of Neurada procumbens, a prostrate annual common in sandy desert habitats, is vulnerable to surface disturbances. Moreover, its seeds are highly heteromorphic, having both spinose and non-spinose sides, which affects both the vertical distribution of seeds and diaspores, and the precise position in which they lie in the soil. Here we explore the ecological implications of two factors, “safe sites” within habitats and seed orientation or “safe sides”, on seedling establishment and the seed bank. In three natural populations, the vertical distribution of buried, germinable diaspores decreased with soil depth. Most diaspores occurred at the surface (0–1 cm below the surface), while none were observed below 10 cm. Seedling emergence decreased with depth and reached zero in diaspores below 5 cm. Seedling emergence also proved highly sensitive to the orientation of diaspores in soil. Diaspores lying on their smooth side with the spinose upper side directed upwards reached the highest seedling emergence of 61 %. Root length ranged from 10 to 16 cm, and the root to shoot ratio varied between 22.8 and 30.3 cm, depending on diaspore size. Diaspores near the surface may constitute a more transient component of the seed bank, as few of these remained viable for more than a year. Persistence of Neurada seed banks seems to depend not only on vertical movement of diaspores among habitat “safe sites” but also on different seed position alternatives.     

科尔沁沙地78种植物繁殖体重量和形状比较

研究了科尔沁沙地 78种植物的繁殖体 (2 3种为种子 ,5 5种为果实 )重量和形状 ,其重量差别很大 ,最小的单粒重不足 0 .1mg,最大的超过 130 mg;形状差异也很大 ,最小的方差不足 0 .0 3,最大的超过 0 .18。综合本研究和以前研究的全部 14 0种植物的研究结果 ,进行了分析。结果表明 :1计有 2 4种植物繁殖体单粒重小于 1mg并接近圆球形 (方差小于 0 .0 9) ,它们可能具有持久种子库 ;2 1年生植物 (平均方差为 0 .0 70 )与 2年生植物 (平均方差为 0 .12 9)间的形状差异显著 (P<0 .0 5 ) ,2年生植物的繁殖体更加扁、长 ;1年生植物也与多年生草本植物 (平均方差为 0 .10 9)之间形状差异显著 (P<0 .0 5 ) ,多年生草本植物的繁殖体更加扁、长 ;3有 5 5 %的豆科植物、70 %的藜科植物繁殖体接近圆球形 (方差 <0 .0 6 ) ,所有的菊科、萝摩科植物繁殖体都很扁、长 (方差 >0 .0 6 )。豆科植物显著比菊科、禾本科、藜科植物繁殖体的重量大 (P<0 .0 5 ) ;4繁殖体附属物在植物的传播和定居方面具有重要的作用 :6种萝摩科植物和 3种杨柳科植物具有绢毛 ,6 0 %的菊科植物具有冠毛 ,6种植物有翅 ,8种植物有宿存花柱或宿存花萼 ,这些植物可能易于被风传播 ;5 4 %的禾本科植物具芒 ,减小了传播可能 ;苍耳、雾冰藜、鹤虱、蒺藜、     

Experimental analysis of patch dynamics and community heterogeneity in tallgrass prairie

Effects of fire and small-scale soil disturbances on species richness, community heterogeneity, and microsuccession were investigated in a central Oklahoma tallgrass prairie. In the fall of 1985, 0.2 m² soil disturbances were created on burned and unburned tallgrass prairie. Vegetation on and off disturbances was sampled at monthly intervals over two growing seasons. During the first growing season, the cover of forbs and annuals, and species richness were significantly greater on versus off disturbances, but these differences did not persist through the second year. The variation in species composition among disturbed plots (heterogeneity) was significantly greater compared to undisturbed areas throughout the study. Fire had no consistent effect on richness and heterogeneity of vegetation on soil disturbances but fire reduced heterogeneity on undisturbed vegetation. Rate of succession, based on an increase in cumulative cover of perennial grasses over time, did not differ among treatments during the first growing season. During the second year, rate of succession was significantly greater on burned soil disturbances compared to unburned soil disturbances. These results suggest that while small-scale soil disturbances have primarily short-lived effects on grassland community structure, disturbances do help to maintain spatial and temporal variation in tallgrass prairie communities. Unlike in undisturbed vegetation, however, species richness and heterogeneity on soil disturbances were little effected by fire, but the rate of colonization onto disturbances appeared to be enhanced by fire.     

Role of soil seed bank along a disturbance gradient in an alpine meadow on the Tibet plateau

We examined the role of the soil seed bank along a grazing disturbance gradient and its relationship with the vegetation of alpine meadows on the Tibet plateau, and discussed the implications for restoration. The seed bank had a high potential for restoration of species-rich vegetation; 62 species were identified in the vegetation and 87 in the seed bank, 39 species being common to both. Mean seed density was 3069–6105 viable seeds m⁻². The density of buried seeds increased significantly with increasing disturbance, indicating that restoration of disturbed areas is not seed limited. Seed density and species richness decreased with depth. The proportion of perennial species decreased with decrease in disturbance both in seed bank and in vegetation. A large portion of species with persistent seeds in the disturbed areas indicate that this seed type can be regarded a strategy of adaptation to current disturbances. Detrended correspondence analysis (DCA) showed significant differences of species composition between seed bank and vegetation, except for the seriously disturbed site. Our results suggest that the establishment of new species in severely disturbed areas is more dependent on the seed bank. By contrast, the restoration in less-disturbed and mature meadows does not rely on seed banks, and the establishment of the vegetation in these communities is more likely to rely on seed dispersal from the standing vegetation and on species with vegetative reproduction.     

Gap colonization in the Patagonian semidesert: seed bank and diaspore morphology

We report work on a cold, windy South American steppe dominated by tussock grasses and shrubs of small stature that, together, cover only half of the soil surface. Our objective was to find out why seedlings and juveniles of these dominant species are generally absent from the bare or poorly-populated spots (gaps) that exist between established individuals. We hypothesized that matrix-forming species fail to colonize gaps because of a lack of properly-placed seeds, contained in diaspores which because of their morphology are blown away from gaps that otherwise would constitute safe sites for recruitment. We evaluated diaspore size and wing loading (weight:area ratio) for all common species in the community, collected seed-bank samples in different occasions and microsites, and performed detailed field observations for one gap-dwelling species during several years. We found that: 1) Seeds of the dominant, matrix-forming species are uncommon in the soil bank of the center of the gaps between established vegetation. 2) Seeds of the dominant species are more abundant towards the edges of the gaps than at their center. 3) Diaspores of those species present in the seed bank of the gaps are smaller than diaspores of absent species; contrary to expectations, not all gap-dwelling species had larger diaspore wing loading than non-gap species. 4) Seeds and adult densities of the most common gap species (the annual Camissonia dentata ), were correlated between them and across subsequent years. We conclude that it is not an overall shortage of seeds that precludes the dominant species from becoming established in the gaps, but rather the seeds' uneven spatial distribution. We further argue that gaps would be suitable sites for recruitment, but large diaspore size makes seeds of the dominant species to be blown away by the strong westerly winds.     

Revegetation following soil disturbance in a California meadow: the role of propagule supply

Revegetation following a disturbance event initially should be constrained by the abundance and types of propagules available at the disturbed site. I tested this idea by conducting two experiments in which I created artificial soil disturbances by excavating or burying pre-existing grassland vegetation. In the first experiment, I varied disturbance intensity (depth), to investigate the consequences for revegetation when numbers of surviving propagules (dormant seeds and bulbs) were altered. In the second experiment, I varied the timing of disturbance, to investigate the consequences when disturbed sites experienced differing exposures to seasonal patterns of clonal growth and seed dispersal. I sampled these experiments from 1991 to 1993, and have interpreted their results using measurements of the seed bank, the bulb bank, and the seed rain. In the first (depth) experiment, bulbs declined in abundance with burial depth and were scarcer in deeper excavations. In contrast, numbers of annual graminoids initially showed no trends with respect to disturbance depth. These results reflect the depth distributions of the seed and bulb banks. Since bulbs occur deeply in the soil, progressively deeper disturbances left fewer survivors. Similarly, perennial graminoids could grow through the shallowest burials. In contrast, since the annual-graminoid-dominated seed bank is concentrated near the soil surface, disturbance depth mattered less to these species: any disturbance removing the surface layer was equally destructive. In the second (timing) experiment, more annual graminoids initially occurred in older plots. This result reflects seasonal patterns of seed production: plots exposed to more of the annual-graminoid-dominated seed rain supported higher densities of annual graminoids as a result. In subsequent years, the vegetation of most plots in both experiments was increasingly dominated by annual graminoids, again as a consequence of their great abundance in the seed rain. These results indicate that interactions between soil disturbances and sources of propagules play an important role in controlling early stages of succession in newly created gaps. They also suggest that disturbance may play different roles in communities characterized by species with different reproductive strategies. Understanding sources of colonists will improve our ability to predict the effects of disturbance.     

Ants as diaspore removers of non-myrmecochorous plants: a meta-analysis

Diaspore (e.g. seeds, fruits) dispersal is pivotal for plant communities and often involves several steps and different dispersing agents. Most studies focusing on diaspore dispersal by animals have highlighted the role of vertebrates, neglecting the role of ants in the diaspore dispersal of non-myrmecochorous plants. Diaspore dispersal by ants is especially relevant in the current scenario of declining of vertebrate populations and, consequently, collapse of the dispersal system of large-seeded plants. Although ants can never compensate for the dispersal service provided by vertebrates, they can mitigate the impact of vertebrate decline via removal of diaspores deposited on the ground. We have used a meta-analytical approach to investigate the contribution of ants in the removal of non-myrmecochorous diaspores (through vertebrate exclusion experiments). We considered the number of diaspore removal as effect size and factors such as plant growth forms, diaspore and ant size, habitat type as moderators. In addition, we investigated the role of such factors on the diaspore removal distance by ants. Ants played complementary role to non-myrmecochorous diaspore removal services provided by vertebrates (mean Hedges’ g of −0.30). The ant diaspore removal was 69% higher for diaspores from shrubs than that of tree diaspores and removal of small-sized diaspores were 69% and 70% higher in comparison to medium- and large-sized diaspores, respectively. Regarding the diaspore removal distance by ants, those of tree species were removed 32% farther than those of shrub species, and diaspores were removed three- times farther in the savanna than in rainforest ecosystems. Our results highlight the shrubs and small-sized diaspores. Regarding the diaspore removal distance, the ants can be crucial for the dispersal of tree diaspores and in the savanna ecosystems. Finally, considering the biodiversity crisis, the ants may play an even more important role than appreciated in diaspores dispersal.     

Seed Stores for Restoration of Species-Rich Shrubland Vegetation Following Mining in Western Australia

The extreme species richness of native shrubland vegetation (kwongan) near Eneabba, Western Australia, presents a major problem in the restoration of sites following mineral sand mining. Seed sources available for post-mining restoration and those present in the native kwongan vegetation were quantified and compared. Canopy-borne seeds held in persistent woody fruits were the largest seed source of perennial species in the undisturbed native vegetation and also provided the most seeds for restoration. In undisturbed vegetation, the germinable soil seed store (140–174 seeds · m^?2) was only slightly less than the canopy-borne seed store (234–494 seeds · m^?2), but stockpiled topsoil provided only 9% of the germinable seeds applied to the post-mining habitat. The age of stockpiled soil was also important. In the three-year-old stockpiled topsoil, the seed bank was only 10.5 seeds · m^?2 in the surface 2.5 cm, compared to 56.1 to 127.6 seeds · m^?2 in fresh topsoil from undisturbed vegetation sites. In the stockpiled topsoil, most seeds were of annual species and 15–40% of the seeds were of non-native species. In the topsoil from undisturbed vegetation, over 80% of the seeds were of perennial species, and non-native species comprised only 2.7% of the seed bank. Additional seeds of native species were broadcast on restoration areas, and although this represented only 1% of the seed resources applied, the broadcast seed mix was an important resource for increasing post-mining species richness. Knowledge of the life-history characteristics of plant species may relate to seed germination patterns and assist in more accurate restoration where information on germination percentages of all species is not available.     

Establishment and reproduction of Aeschynomene virginica (L.) (Fabaceae) a rare,annual, wetland species in relation to vegetation removal and water level

Aeschynomene virginica is a rare annual plant found in freshwater tidal wetlands of the eastern United States. We hypothesized that standing vegetation and water inundation were two important environmental factors in its population dynamics. To test these hypotheses, we sowed seeds into plots with undisturbed vegetation or plots with all aboveground vegetation removed in 1998 and 1999. Presence/absence of seedlings was noted and seedling survival to reproduction, final size, and seed set were measured throughout both growing seasons. Seedling establishment from germination to the first true leaf stage increased with decreasing water depth. Vegetation removal plots had greater seedling establishment, higher seedling survival, and higher seed set per plant than non-removal plots. In a greenhouse study designed to test the effects of water level on seed germination and seedling establishment, no seedlings established in submerged soils, and seed germination and seedling establishment were lower in waterlogged soil than in wet soil. Physical stress associated with deeper water likely limits the distribution of A. virginica to higher elevations, where seeds that colonize patches with low vegetative cover are more likely to produce reproductive adults that produce more seeds relative to patches with established vegetation. A. virginica appears to be a fugitive species specializing on open habitat patches in tidal wetlands. This species may be dependent on disturbances for population establishment and maintenance.     

Species,Diaspore Volume and Body Mass Matter in Gastropod Seed Feeding Behavior

Background

Seed dispersal of ant-dispersed plants (myrmecochores) is a well studied ecosystem function. Recently, slugs have been found to act as seed dispersers of myrmecochores. The aim of our study was to (1) further generalize the finding that gastropods feed on seeds of myrmecochores and hence may act as seed dispersers, (2) to test whether gastropod body mass and the volume of diaspores have an influence on the seed dispersal potential.

Methodology and Principal Findings

We assessed the seed dispersal potential of four slug and snail species with a set of seven myrmecochorous plant species from seven different plant families common to Central European beech forests. Diaspores differed in shape and size. Gastropods differed in their readiness to feed on diaspores and in the proportion of seeds that were swallowed as a whole, and this readiness generally decreased with increasing diaspore size. Smaller Arionid slugs (58 mm body length; mean) mostly fed on the elaiosome but also swallowed small diaspores and therefore not only act as elaiosome consumers, a nutrient rich appendage on myrmecochorous diaspores, but may also disperse seeds. Large Arionid slugs (>100 mm body length) swallowed diaspores of all sizes. Diaspores swallowed by gastropods were defecated without damage. Within-species variability in body size also affect seed dispersal potential, as larger individuals of the red slug (Arion rufus) swallowed more diaspores of wood anemone (Anemone nemorosa) than smaller ones.

Conclusions and Significance

Our results help to generalize the finding that gastropods consume and potentially disperse seeds of myrmecochores. The dispersal potential of gastropods is strongly influenced by diaspore size in relation to gastropod size.     

DISPERSAL ECOLOGY OF CAREX PEDUNCULATA (CYPERACEAE), A NEW NORTH AMERICAN MYRMECOCHORE

Carex pedunculata is the first North American species of the Cyperaceae that is identified as a myrmecochore. Many morphological and phenological features of this species and its breeding system are interpreted as adaptive for seed dispersal by ants. In laboratory tests, workers of the ant species Aphaenogaster rudis carry the diaspores to the nest, eat the elaiosomes, carry larvae to the elaiosomes to feed, and deposit diaspores whose elaiosomes have been eaten with other nest debris. The achenes then germinate. Achenes will also germinate without any handling by ants. Workers will also transport diaspores with uneaten elaiosomes when the nest is disturbed. Greenhouse tests show that seedling growth is greatly inhibited if a diaspore remains near the parent plant and cohort seedlings. Field studies of natural populations identify rotting logs (the location of ant nests) as forest floor microsites for colonization of C. pedunculata and other myrmecochores. Ant nesting behavior may pattern much of the herb stratum. This species is self-compatible, and single seeds may start successful new populations. Three processes contribute to population growth: vegetative growth, germination of untransported diaspores, and germination of ant-transported diaspores.     

Community and population dynamics of serpentine grassland annuals in relation to gopher disturbance

Summary This study examines the effects of soil disturbance by gophers on patterns of species abundance in an annual grassland community on serpentine soil. We assessed production, dispersal and storage of seed, germination, survivorship and growth of the most abundant species in undisturbed vegetation and on gopher mounds. Fewer seeds of the dominant species were dispersed onto gopher mounds due to the limited movement of seeds from within the closed vegetation. Species with taller flowering stalks were more likely to colonise gopher mounds. The timing of gopher disturbance in relation to the timing of seed fall determined which species could colonise mounds. Lower numbers of seeds falling onto gopher mounds resulted in lower seedling densities of several species compared with undisturbed areas. Survivorship of the commonest species differed between undisturbed areas and gopher mounds formed at different times of year. This resulted in characteristic spectra of species abundance on the different microhabitats, giving rise to distinct spatial patterning in the community. Plants growing on gopher mounds were generally larger and produced more seed than plants in undisturbed vegetation. We suggest that continued gopher disturbance is a factor allowing several species, including perennial grasses, to persist in this community.     

Recovery of pristine boreal forest floor community after selective removal of understorey, ground and humus layers

In boreal spruce forests that rarely experience extensive disturbances, fine-scale vegetation gaps are important for succession dynamics and species diversity. We examined the community implications of fine-scale gap disturbances by selective removal of vegetation layers in a pristine boreal spruce forest in Northern Finland. The aim was to investigate how the speed of recovery depends on the type of disturbance and the species growth form. We also wanted to know if there appeared changes in species composition after disturbance. Five different treatments were applied in the study: Control, removal of the ground layer (bryophytes and lichens), removal of the understorey layer (dwarf shrubs, herbs and graminoids), removal of both the ground and understorey layers, and complete removal of the vegetation and humus layers above the mineral soil. The vegetation recovery was monitored in terms of cover and species numbers over a 5-year period. Understorey layer cover, composed mainly of clonal dwarf shrubs, recovered completely in 4 years in treatments where the humus layer remained intact, whereas ground layer cover did not reach the control level in plots from where bryophytes and lichens were removed. Recovery was faster in terms of species number than species cover. Bryophytes, graminoids and dominant dwarf shrubs appeared in all disturbed plots quickly after disturbance. Seedlings of trees appeared exclusively in disturbed plots. Graminoids dominated after the removal of humus layer. The results indicate that the regeneration of forest floor after small gap disturbance occurs mainly by re-establishment of the dominant species. Although destruction of the humus layer leaves a long-lasting scar to the forest floor, exposing of mineral soil may enhance the sexual reproduction of dominant species and the colonization of weaker competitors.