Microbial colonization dynamics in temporary aquatic systems

Most studies of community development in insular systems have investigated the colonization dynamics of only a portion of the biotic community using islands that have the benefit of prior biotic modification. Two experiments assessed the predictive ability of equilibrium island theory with regard to the development of the protistan community in temporary aquatic islands (100-1 plastic swimming pools) void of any prior biotic modification. The first experiment ran for 170 d (March–September 1985) and manipulated the access of islands to animals that represent potentially important dispersal pathways for microbes. A second experiment (May–June 1986) investigated in greater detail the early stages of species accrual in the absence of dispersal pathways other than via the atmosphere. Polyurethane foam substrates were used as sampling devices to increase the accuracy and replicability of sampling and provide habitat for colonists. Sampling was determined to be asymptotic. Species accrual was asymptotic in both experiments, although it initially lagged behind that predicted by theory. Autotrophs approached equilibrium faster than heterotrophs, but at a lower species richness. The predicted number of autotroph species at equilibrium was lower in islands that were excluded from contact with birds compared to less exclosed islands. The colonization dynamics of the entire community was not significantly different among islands having different degrees of exclosure. In both experiments, rates of species immigration were nonmonotonic with respect to time and species richness. This relationship appeared to reflect the importance of species interactions during the initial accrual phase before equilibrium. Rates of extinction were positively correlated with both these parameters, although they tended to decrease with time during the equilibrium period in less exclosed islands. Turnover at equilibrium was significant and resulted in directional changes in species composition over time. Assortative processes appeared to be important since later colonists exhibited greater persistence. Colonizing species generally fall into three autecological categories: 1) those that were ubiquitous and had temporally predictable patterns of immigration (successional species); 2) those possessing temporally predictable distributions but not spatially ubiquitous distributions (dispersal limited species); 3) those that showed little temporal or spatial predictability in immigration (transient or allochthonous species). Individual islands exhibited various degrees of fluctuation in species number during the predicted equilibrium which were poorly correlated with exogenous environmental variables and physicochemical habitat parameters. The presence of predacious mosquito larvae(Culex spp.) invariably resulted in a sharp decreases in microbial species richness, while documented contact with rodents was followed by an increase in species number in an island so contaminated. Several aspects of microbial colonization of temporary waters that contradict equilibrium predictions appear to be strongly influenced by microbe-microbe as well as macrobe-microbe interactions.     

Associations between arbuscular mycorrhizal fungi and grasses in the successional context of a two-phase mosaic in the Chihuahuan Desert

The hypothesis that plant species are more responsive to mycorrhiza in late than in early successional stages was assessed in grasses from a successional process occurring in two-phase mosaics from the Mexican Chihuahuan Desert. We estimated the density of spores of arbuscular mycorrhizal (AM) fungi and the AM colonization of pioneer and late-successional grasses in the field. In growth chamber experiments, we tested the effect of the native AM fungal community on grasses growing in soils from different successional stages. Spore density was higher in late than in early successional stages. Late-successional species were more responsive to AM (positive AM responsiveness) whereas pioneer species were nondependent on mycorrhiza or if associated to AM fungi, the interaction showed a negative AM responsiveness for the seedling stage. Our findings showed that late successional species fitted the proposed models of mycorrhizal performance, but the two pioneer species differed in their AM condition and responsiveness. This further supports the idea that AM interactions are more complex along the successional processes than the predictions of the more widely cited hypotheses.     

植被恢复树种在不同实验光环境下叶片气体交换的生态适应特点

比较研究了华南丘陵地区植被恢复8种乔木树种在不同光条件下的光饱和点、日平均净光合速率、叶温、气孔导度日平均值、内在水分利用效率和光合辐射利用效率等生理生态特征。结果表明,火力楠和藜蒴具有演替后期种的特征,在荫蔽条件下生存和生长良好,它们不宜用作构建先锋群落的树种。大叶相思、马占相思和樟树是典型的阳生性树种,具有鲜明的演替后期种的特征;红椎和海南红豆显示除叶温以外的阳生性的特征。这些种类可被选作构建先锋群落的组成成分。桂西红荷则只能作为演替的过渡种类,它兼具有演替前期种和演替后期种的特征,尤其适用于对先锋群落进行林分改造。     

Effects of Small Rodent and Large Mammal Exclusion on Seedling Recruitment in Costa Rica1

We examined whether the experimental exclusion of large mammalian and small rodent seed predators had differing effects on seedling recruitment under natural seed rain conditions. In both primary and late‐successional secondary forested areas, exclosure experiments using natural seed densities were designed to assess seedling recruitment. To assess the differences in seedling recruitment, we monitored three exclosure treatments (1.2 m radius/1.5 m height) in two forest types (primary vs. late‐successional secondary forest): (1) fenced exclosures that excluded large mammals; (2) fenced exclosures that excluded both large and small mammals; and (3) open controls. Within each exclosure treatment, we marked and identified all seedlings at the beginning of the experiment (February 2001), followed the marked seedlings' fate for a year, and then marked and identified all new seedlings after a year. Two preliminary findings were generated from these data: for some tree species, small rodents and large mammals have differential effects on seedling recruitment, and the effect of excluding mammals did not differ with habitat type (primary vs. late‐successional secondary forest). These preliminary results highlight the need to examine further how the effects of small rodent and large mammal exclusion may affect species‐specific seed predation and seedling recruitment in a variety of habitat/land use types (e.g., primary forest, late‐successional forest, and early‐successional forest).     

Responses to nitrogen pulses and growth under low nitrogen availability in invasive and native tree species with differing successional status

Invasive species are frequently found in recently disturbed sites. To examine how these disturbance-dependent invasive species exploit resource pulses resulting from disturbance, twelve physiological and morphological traits, including age-dependent responsiveness in leaf traits to nitrogen pulse, were compared between Bischofia javanica, an invasive tree species in Ogasawara islands, and three native Ogasawara species, each having a different successional status. When exposed to a nitrogen pulse, invasive B. javanica showed higher increases in photosynthetic capacity, leaf area, epidermal cell number and cell size in leaves of broad age classes, and root nitrogen absorption ability than two native mid-/late or late-successional species, but showed no particular superiority to a native pioneer species in these responses. Under low nitrogen, however, it showed the largest relative growth rate among the four species, while the native pioneer showed the lowest growth. From these results, we concluded that the combination of moderately high responsiveness to resource pulses and the ability to maintain steady growth under resource limitations may give B. javanica a competitive advantage over a series of native species with different successional status from early to late-successional stages.     

Resolving the predator first paradox: Arthropod predator food webs in pioneer sites of glacier forelands

Primary succession on bare ground surrounded by intact ecosystems is, during its first stages, characterized by predator‐dominated arthropod communities. However, little is known on what prey sustains these predators at the start of succession and which factors drive the structure of these food webs. As prey availability can be extremely patchy and episodic in pioneer stages, trophic networks might be highly variable. Moreover, the importance of allochthonous versus autochthonous food sources for these pioneer predators is mostly unknown. To answer these questions, the gut content of 1,832 arthropod predators, including four species of carabid beetles, two lycosid and several linyphiid spider species caught in early and late pioneer stages of three glacier forelands, was screened molecularly to track intraguild and extraguild trophic interactions among all major prey groups occurring in these systems. Two‐thirds of the 2,310 identified food detections were collembolans and intraguild prey, while one‐third were allochthonous flying insects. Predator identity and not successional stage or valley had by far the strongest impact on the trophic interaction patterns. Still, the variability of prey spectra increased significantly from early to late pioneer stage, as did the niche width of the predators. As such the structure of pioneer arthropod food webs in recently deglaciated Alpine habitats seems to be driven foremost by predator identity while site and early successional effects contribute to a lesser extent to food web variability. Our findings also suggest that in these pioneer sites, predatory arthropods depend less on allochthonous aeolian prey but are mainly sustained by prey of local production.     

Generation of Spatial Patterns in Boreal Forest Landscapes

Boreal forests are composed of a few plant species with contrasting traits with respect to ecosystem functioning and spatial patterning. Early successional deciduous species, such as birch and aspen, disperse seeds widely, do not tolerate low light and nitrogen availabilities, have rapidly decaying litter, and are highly preferred by herbivores. These later succeed to conifers, such as spruce and fir, which disperse seeds locally, tolerate low light levels and low nitrogen availability, have litter that decays slowly, and are unpalatable to most mammalian herbivores. Although there are also early successional conifers, such as jack pine and Scots pine, the aspen-birch-spruce-fir successional sequence is the most common over much of North America, and (without fir) in Fennoscandia and Siberia. The course of succession in these forests is controlled partly by seed dispersal and selective foraging by mammalian herbivores. Both of these processes are spatially dynamic, but little is known about how their spatial dynamics may affect ecosystem processes, such as nitrogen cycling or productivity. We present spatially explicit models that demonstrate the following: (a) Spatially explicit seed dispersal results in more clumped distribution of tree species and persistence of greater paper birch biomass than uniform seed rain across the landscape. Such results are consistent with current spatially explicit population models of dispersal and coexistence. (b) With localized seed dispersal, the concentrations of available soil nitrogen are distributed in larger patches with sharp transitions from low to high nitrogen availability near patch edges. In contrast, with a uniform seed rain, the distribution of soil nitrogen availability was more uniform and “hotspots” were more localized. Thus, the spatial pattern of an ecosystem process (nitrogen cycling) is determined by seed dispersal and competition for light among competing populations. (c) A dispersing herbivore, such as moose, that selectively forages on early successional deciduous species with high quality litter, such as aspen or birch, and discriminates against late successional conifers, such as spruce or fir, imposes higher-order repeated patterns of plant species and biomass distribution on the landscape. Thus, seed dispersal and herbivore foraging correlate properties in adjacent patches but in different ways, and different spatial patterns emerge. Other processes, such as insect outbreaks, fire, and water flow, also may correlate properties between adjacent patches and result in additional patterns. Received 8 February 1999; accepted 28 May 1999.     

Nest exclosures do not improve Streaked Horned Lark nest success

ABSTRACT Improving the conservation status of rare and declining species often requires multiple strategies targeted at several vital rates. We report on one of several ongoing management actions intended to benefit the declining population of Streaked Horned Larks (Eremophila alpestris strigata). To improve Streaked Horned Lark fecundity, we employed predator exclosures (wire cages) around nests (N= 33 exclosed and 32 not exclosed) in 2009 and 2010 at two sites in Oregon and two in Washington with the goal of excluding larger birds, the primary lark nest predators. We found no statistically significant effect of exclosures on nest success. For exclosed nests, lower rates of nest predation (exclosed = 12%, unexclosed = 48%) were offset by higher rates of nest abandonment (exclosed = 27%, unexclosed = 0%). Nest abandonment was likely caused by a variety of factors including American Kestrels (Falco sparverius) perching on exclosures, and predation of adults associated with exclosed nests. Our results suggest that the current exclosure design does not improve Streaked Horned Lark fecundity and may negatively affect adult survival. To improve exclosure effectiveness, we recommend modifications that prevent kestrels from perching on exclosures and deny their access to the nest. We also recommend that modifications be applied in an adaptive management framework that includes close monitoring to assess their effectiveness, and subsequent adaptation that might include continued structural modification of exclosures or discontinued use on some or all sites.     

Diversity and community biomass depend on dispersal and disturbance in microalgal communities

The evidence for species diversity effects on ecosystem functions is mainly based on studies not explicitly addressing local or regional processes regulating coexistence or the importance of community structure in terms of species evenness. In experimental communities of marine benthic microalgae, we altered the successional stages and thus the strength of local species interactions by manipulating rates of dispersal and disturbance. The treatments altered realized species richness, evenness and community biomass. For species richness, dispersal mattered only at high disturbance rates; when opening new space, dispersal led to maximized richness at intermediate dispersal rates. Evenness, in contrast, decreased with dispersal at low or no disturbance, i.e. at late successional stages. Community biomass showed a non-linear hump-shaped response to increasing dispersal at all disturbance levels. We found a positive correlation between richness and biomass at early succession, and a strong negative correlation between evenness and biomass at late succession. In early succession both community biomass and richness depend directly on dispersal from the regional pool, whereas the late successional pattern shows that if interactions allow the most productive species to become dominant, diverting resources from this species (i.e. higher evenness) reduces production. Our study emphasizes the difference in biodiversity–function relationships over time, as different mechanisms contribute to the regulation of richness and evenness in early and late successional stages.     

Why are leaf‐cutting ants more common in early secondary forests than in old‐growth tropical forests? An evaluation of the palatable forage hypothesis

I evaluated the hypothesis that leaf‐cutting ants are more common in early successional forests than in old‐growth forests because pioneer species, which dominate in early successional habitats, appear more susceptible to leafcutters than shade‐tolerant species, which dominate primary forests (palatable forage hypothesis). The relative importance of pioneer and shade‐tolerant species as plant resources for leaf‐cutting ant colonies was evaluated (1) by literature review of leaf‐cutting ants’ diet, and (2) experimentally, using field assays to determine leafcutter's selectivity. Pioneer species were harvested three times more frequently than shade‐tolerant species and made up the largest component of the diet in all the studies reviewed. The amount harvested was not correlated with the plant species abundance. In addition, leaves from pioneer plants were selected eight times more than leaves from shade‐tolerant species in the field assays. These results support the palatable forage hypothesis. Leafcutters probably select pioneer leaves because of their low level of chemical defenses and high nutrient content. The high availability of pioneer species in early successional forest probably decreases the cost to locate palatable resources. Therefore, early successional habitats support more ant colonies than old‐growth forests. On the other hand, the effective defense mechanisms of mature plant species and the high dispersion of palatable plants could explain the low density of leaf‐cutting ant colonies in old‐growth forests. The palatable forage hypothesis is compared with other hypotheses that explain leaf‐cutting ant density. The preference of foundress queens for forest clearings, the dependence of small colonies on herbs, and the importance of pioneer plant species for mature colonies (palatable forage hypothesis) can be considered complementary, because they focus on different stages of the colony's life history. Consequently, the availability of pioneer plants appears to be one of the most influential factors determining mature leaf‐cutting ant nest densities in Neotropical forests.     

Comparative hydraulic architecture of tropical tree species representing a range of successional stages and wood density

Plant hydraulic architecture (PHA) has been linked to water transport sufficiency, photosynthetic rates, growth form and attendant carbon allocation. Despite its influence on traits central to conferring an overall competitive advantage in a given environment, few studies have examined whether key aspects of PHA are indicative of successional stage, especially within mature individuals. While it is well established that wood density (WD) tends to be lower in early versus late successional tree species, and that WD can influence other aspects of PHA, the interaction of WD, successional stage and the consequent implications for PHA have not been sufficiently explored. Here, we studied differences in PHA at the scales of wood anatomy to whole-tree hydraulic conductance in species in early versus late successional Panamanian tropical forests. Although the trunk WD was indistinguishable between the successional groups, the branch WD was lower in the early successional species. Across all species, WD correlated negatively with vessel diameter and positively with vessel packing density. The ratio of branch:trunk vessel diameter, branch sap flux and whole-tree leaf-specific conductance scaled negatively with branch WD across species. Pioneer species showed greater sap flux in branches than in trunks and a greater leaf-specific hydraulic conductance, suggesting that pioneer species can move greater quantities of water at a given tension gradient. In combination with the greater water storage capacitance associated with lower WD, these results suggest these pioneer species can save on the carbon expenditure needed to build safer xylem and instead allow more carbon to be allocated to rapid growth.     

How habitat heterogeneity shapes bacterial and protistan communities in temperate coastal areas near estuaries

In this study, we investigated microbial communities (bacteria and protist) in two coastal areas near the estuaries of the Liaohe (LH) River and Yalujiang (YLJ) River in the Northwestern Pacific Ocean. Due to the existence of Liaodong Peninsula and different levels of urbanization, geographical segregation and significant environmental heterogeneity were observed between these two areas. There were significantly different regional species pools and biogeographic patterns for both bacterial and protistan communities between LH and YLJ coastal areas. Species turnover was the main mechanism driving β-diversity patterns of both bacterial and protistan communities in each area. In addition, the contributed ratio of nestedness to the β-diversity patterns was significantly higher for protists compared to bacteria. Variation in regional species pools was found to be the dominant driver of differences of bacterial and protistan communities between the LH and YLJ coastal areas. For a single-studied area, local community assembly mechanisms, including heterogeneous selection and dispersal limitation, were found to shape the bacterial and protistan communities through calculation of the β-deviation index. Among them, the relative importance of heterogeneous selection and dispersal limitation on the community assembly varied according to microorganism type and habitat.     

Acknowledgments

In this study we investigated the temporal variability of N-source utilization of pioneer plant species in different early successional stages of dry acidic grasslands. Current theory states that plant species occupy distinct ecological niches and that there are species-specific, temporal N-uptake patterns. We hypothesized that small-scale dynamics in the natural habitat may affect niche differentiation among plant species. We investigated N-uptake patterns of two co-occurring plant species from different functional groups (Corynephorus canescens, Rumex acetosella) under natural conditions using ¹⁵N-labeled nitrate and ammonium in three different early successional stages during early and late summer. We found (1) marked seasonal dynamics with respect to N-uptake and N-source partitioning, and (2) different uptake rates across successional stages but a similar N-form utilization of both species. Nitrate was the main N-source in the early and later successional stages, but a shift towards enhanced ammonium uptake occurred at the cryptogam stage in June. Both species increased N-uptake in the later successional stage in June, which was associated with increasing plant biomass in C. canescens, whereas R. acetosella showed no significant differences in plant biomass and root/shoot-ratio between successional stages. Ammonium uptake decreased in both species across all stages with increasing drought. Nevertheless, the peak time of N-uptake differed between the successional stages: in the early successional site, with the lowest soil N, plants were able to extend N-uptake into the drier season when uptake rates in the other successional stages had already declined markedly. Hence, we found a pronounced adjustment in the realized niches of co-occurring plant species with respect to N-uptake. Our results indicate that ecological niches can be highly dynamic and that niche sharing between plant species may occur instead of niche partitioning.     

Seed rain,seedling establishment and clonal growth strategies on a glacier foreland

Abstract. Seed production, composition of the seed rain, germination, and seedling mortality, as well as vegetative growth characteristics of common pioneer plant species were studied on the foreland of the retreating Morteratsch glacier in the Swiss Alps. The frequency of diaspores trapped in different successional stages was related to their dispersal mode and was highly skewed towards a few species. Plenty of diaspores well adapted for dispersal by wind are a precondition for the most important pioneer species. Seeds from all pioneer species investigated had a good germination success, provided that the moisture content of the soil was high enough. However, requirement for seedling establishment differed among sites of increasing terrain age and among species. Only specialized pioneers such as Cerastium pedunulatum, Linaria alpina, Oxyria digyna and Saxifraga aizoides tolerate the cold and moist conditions near the glacier. However, these species are restricted to early successional stages. Seedlings of Epilobium fleischeri are affected not only by the cold and moist conditions near the glacier but also by moderately dry conditions on older sites. Availability of safe sites becomes crucial for most species with increasing age of sites and with drier conditions. Most species playing a dominant role during early succession and persisting during later successional stages have a distinctive ability to spread clonally and have a growth form with more or less widely spaced ramets: Achillea moschata. Cerastium pedunculatum. Epilobium fleischeri and Hieraium staticifolium. The growth strategy and demography of the clonal E. fleischeri is presented as an example. The life cycle of this species is characterized in succession by (1) the colonization of safe sites by small seeds adapted for wind dispersal, (2) horizontal spread by clonal growth, and (3) the persistence through phenotypic morphological plasticity in later successional stages. Seedling establishment and clonal growth are thus complementary mechanisms in plant succession on recently deglaciated terrain.     

Responses of Pioneer and Later‐Successional Plant Assemblages to Created Microtopographic Variation and Soil Treatments in Riparian Forest Restoration

Riparian forest restoration generally involves introduction of later‐successional tree species, but poor species suitability to severely altered or degraded site conditions results in high mortality and poor community development. Additionally, while microtopographic heterogeneity plays a crucial role in the development of natural riparian forests, little is known regarding effects of restored or created microtopography on the development of introduced plant communities. The objective of this study was to determine the influence of created microtopography and soil treatments on early development of introduced pioneer and later‐successional plant communities in riparian forest restoration. Ridges, flats, and a mound‐and‐pool complex were created, and pioneer and later‐successional tree assemblages were planted within plots in each of these three microtopographic positions. Straw‐based erosion control mats were placed on half the plots as a source of mulch. After two growing seasons, growth and survival of the pioneer assemblage were equal among microtopographic positions, but survival of the later‐successional assemblage was significantly higher on ridges (59%) than on mounds and pools (22%) and flats (26%). A suitability index indicated that performance of the later‐successional assemblage on ridges was higher than that of the pioneer assemblage for all microtopographic positions. Flood duration explained much of the variation in plant assemblage survival, and erosion control mats had little influence on seedling survival. Restoring microtopographic features has the potential to enhance species survival and promote community development. Microtopographic restoration may be as important in riparian forest restoration as proper species selection and hydrologic reestablishment, especially at severely disturbed sites.     

Resource availability at three presumed stages of a heathland succession on the Lüneburger Heide,Germany

Abstract. The availability of maj or plant resources was investigated in three vegetation types that were assumed to represent different stages of a secondary succession on heathland on the Lüneburger Heide, northwestern Germany. Canopy transmission and absorption of photosynthetically active radiation (PAR), soil-water availability, and nutrient (Ca, K, Mg, N, P) availability were monitored with high spatial and temporal resolution in (1) a Calluna vulgaris heathland, (2) a pioneer birch-pine forest and (3) a late-successional oak-beech forest, situated close to each other on comparable geological substrate (diluvial). Mean fractional transmission of PAR during summer decreased from 0.48 in the heathland to 0.04 in the oak-beech forest while the fractional canopy absorption increased from 0.49 to 0.92. Soil-water availability as indicated by the soil-water potential, was significantly influenced by differential canopy interception loss and characteristic rooting patterns in the three vegetation types. Annual mean nutrient concentrations in the equilibrium soil solution were similar or, for some elements, increased from the heathland to the birch-pine and the oak-beech forest despite a growing demand. A marked increase was found for the total nutrient pools in the soil-organic layer between early and late successional vegetation types. On the poor glacial parent material, nutrient pools seem to be strongly dependent on stand productivity and litter production which increased with succession. Thus, for nutrients, facilitation seems to be important in this type of succession.     

Colonization of experimentally created gaps along an alpine successional gradient

The colonization of artificially created gaps was analyzed along an alpine successional gradient from pioneer to early, late, and old successional stages. The presence/absence of species and the abundances of seedlings and adults in the gaps were recorded and compared with those of the surrounding areas. We hypothesized that in the older successional stages, the gaps were likely to be colonized by clonal ingrowth of the surrounding species. In the younger stages, we expected to find a high presence of seedlings and adults recruited by seeds. Micro-succession in the gaps occurred at each successional stage, with all life forms among the colonizers. The abundance of seedlings was significantly higher in the gaps compared with the surrounding area. At the early and late successional stages, the surrounding areas provided safe sites for seedling establishment, with the abundance of adults recruited by seeds higher at the gap edges than in the gap centers. We can confirm the first hypothesis of a higher clonal ingrowth in the old successional stage. Clonal ingrowth also occurred in the younger successional stages. Despite the lower species richness in the gaps, a positive correlation was found between gap and surrounding species frequencies, which were the highest in the pioneer and the lowest in the old successional stage. We conclude that gaps are relevant for seedling recruitment along the entire primary succession gradient. New species invasions from greater distances were not observed in the gaps. The dominant species on each site were identified to be successful gap colonizers.     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

The role of successional stage and small‐scale disturbance for establishment of pioneer grass Corynephorus canescens

Question: Which mechanisms promote the maintenance of the protected pioneer grass Corynephorus canescens in a mosaic landscape? Which are the interactive effects of small‐scale disturbances, successional stage and year‐to‐year variation on early establishment probabilities of C. canescens? Location: Brandenburg, NE Germany. Methods: We measured emergence and survival rates over 3 yr in a sowing‐experiment conducted in three successional stages (C. canescens‐dominated site, ruderal forb site and pioneer forest) under two different regimes of mechanical ground disturbance (disturbed versus undisturbed control). Results: Overall, disturbance led to higher emergence in a humid year and to lower emergence in a very dry year. Apparently, when soil moisture was sufficient, the main factor limiting C. canescens' establishment was competition, while in the dry year, water became the limiting factor. Survival rates were not affected by disturbance. In humid years, C. canescens emerged in higher numbers in open successional stages while in the dry year, emergence rates were higher in late stages, suggesting an important role of late successional stages for the persistence of C. canescens. Conclusions: Our results suggest that small‐scale disturbances can promote germination of C. canescens. However, disturbances should be carefully planned. The optimal strategy for promoting C. canescens is to apply disturbances just before seed dispersal and not during dry years. At the landscape scale, a mosaic of different vegetation types is beneficial for the protected pioneer grass as facilitation by late‐successional species may be an important mechanism for the persistence of C. canescens, especially in dry years.     

How long can glacier foreland species live?

Age determinations provide information on longevity of the species and on their population structure. This knowledge is of particular interest for species involved in succession. On a glacier foreland of the Central Alps age determinations of six target species (Saxifraga oppositifolia, Oxyria digyna, Artemisia mutellina, Anthyllis vulneraria ssp. alpestris, Trifolium pratense ssp. nivale and Helianthemum grandiflorum) were carried out by means of herbchronology. The species were selected on three differently aged moraines in order to compare pioneer, early and late successional species. Pioneer species were expected to have young, invasive populations. This was suggested also for populations of late successional species growing towards the glacier front where they substitute the pioneer species. These hypotheses were only confirmed for the pioneer Saxifraga oppositifolia. The longest life span was found for Helianthemum grandiflorum on the oldest moraine. Plant size of all investigated individuals correlated with age. Significant differences where observed between the age of vegetative and generative individuals.