Seasonal and spatial variance of seed bank species composition in an oligotrophic wet meadow

Seasonal and small-scale spatial variation of the seed bank in an oligotrophic wet meadow (Molinion) was assessed by two methods: (1) direct determination of seeds extracted from soil samples (6 cm in diameter, 10 cm depth) and (2) germination of seeds according to Ter Heerdt et al. [1996. An improved method for seed bank analysis: seedling emergence after removing the soil by sieving. Funct. Ecol. 10, 144–151]. Samples were taken three times during the year (July and November 2001, March 2002) from three distinct microhabitats (moss layer, gaps, Molinia caerulea tussocks). Using the extraction method, more seeds were found in the upper 0–3 cm soil layer than in the deeper 3–7 cm layer, significant differences in the number of seeds were found among microhabitats (with the lowest seed bank densities in Molinia tussocks). The highest number of seeds was found in the spring census, but the test was not significant (P<0.10). However, seasonal variation was found when damaged seeds were included in the analysis. Changes in damaged/undamaged and light/heavy seed ratios, and in numbers of grass/herbs/sedges seeds are also discussed. The spatial distribution of seeds is probably influenced to a great degree by abiotic factors, such as microtopography (round and heavy seeds of sedges prevail in gaps and in deeper soil layers). The species composition of the seed bank varied among microhabitats, depth, and throughout the year. No differences were found by the seed germination method.     

Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation

Understanding processes underlying spatial distribution of tree species is fundamental to studying species coexistence and diversity. This study modeled point patterns of tree distribution, expressed by Cartesian coordinates of individual trees within a mapped forest stand, for the purpose of identifying processes that may generate spatial patterns of tree communities. We used four primary point pattern processes (homogeneous Poisson process, inhomogeneous Poisson process, homogeneous Thomas process, and inhomogeneous Thomas process) to model tree distribution in two stem-mapped forests in Taiwan, Republic of China. These four models simulate spatial processes of habitat association and seed dispersal, allowing us to evaluate the potential contribution of habitat heterogeneity and dispersal limitation to the formation of spatial patterns of tree species. The results showed that the inhomogeneous Thomas process was the best fit model and described most of the species studied, suggesting that spatial patterns of tree species might be formed by the joint effects of habitat associations and dispersal limitation. The homogeneous Thomas process that models the effect of dispersal limitation was the second best model. We also found that the best fit models could be predicted by species attributes, including species abundance and dispersal mode. The significant traits, however, differed between the two study plots and demonstrated site-specific patterns. This study indicated that the interactive operation of niche-based (habitat heterogeneity) and neutral-based (dispersal limitation) may be important in generating spatial patterns of tree species in forest communities.     

The scale-dependent importance of habitat factors and dispersal limitation in structuring Great Lakes shoreline plant communities

Niche-based and neutral models of community structure posit distinct mechanisms underlying patterns in community structure; correlation between species’ distributions and habitat factors points to niche assembly while spatial pattern independent of habitat suggests neutral assembly via dispersal limitation. The challenge is to disentangle the relative contributions when both processes are operating, and to determine the scales at which each is important. We sampled shoreline plant communities on an island in Lake Michigan, varying the extent and the grain of sampling, and used both distance-based correlation methods and variance partitioning to quantify the proportion of the variation in plant species composition that was attributable to habitat factors and to spatial configuration independent of habitat. Our results were highly scale dependent. We found no distance decay of plant community similarity at the island scale (1−33 km). All of the explained variation (32%) in species composition among samples at this scale was attributed to habitat factors. However, at a site intensively sampled at a smaller scale (5−1,200 m), similarity of species composition did decay with distance. Using a coarse sampling grain (transects), habitat factors explained 40% of the variation, but the purely spatial component explained a comparable 22%. Analyzing plots within transects revealed variation in species composition that was still jointly determined by habitat and spatial factors (18 and 11% of the variance, respectively). For both grain sizes, most of the habitat component was spatially structured, reflecting an abrupt alongshore transition from sandy dunes to cobble beach. Space per se explained more variation in species composition at a second site where the habitat transition was more gradual; here, habitat acted as a less selective filter, allowing the signal of dispersal limitation to be detected more readily. We conclude that both adaptation to specific habitat factors and habitat-independent spatial position indicative of dispersal limitation determine plant species composition in this system. Our results support the prediction that dispersal limitation—a potentially, but not necessarily, neutral driver—is relatively more important at smaller scales.     

Population dynamics and dispersal of Lepthyphantes tenuis in an ephemeral habitat

The population dynamics of Lepthyphantes tenuis (Blackwall) were investigated in two fields of winter wheat in Sussex, England, with the aim of attributing the major factors controlling population fluctuations. Continuous measures of changes in density, reproductive rate and dispersal were obtained using a range of methodologies, including laboratory analysis, suction trapping, rotary sampling of the aerial fauna, caging, and detailed density estimations. The resulting data were then used to determine the major causes of the observed population fluctuations and to describe the processes of dispersal and reproduction in this population.Population densities were initially very low following ploughing operations, but increased throughout the season as a result of high reproductive rates. Investigation of the reproductive potential of the spiders showed that the number of eggs per egg-sac did not vary with season, and that egg development was temperature related. There was no evidence that the number of viable eggs per egg-sac varied during the season. There were three peaks of hatchling production per year suggesting two to three generations of spiders. Dispersal was not responsible for the major changes in field density throughout the season although dispersal activity was high for this species. Dispersal activity increased with age and was highest in females. Dispersal activity as a ratio of activity/field density for females, males, sub-adults, immatures and hatchlings was 4:2:3:1:1.Aerial dispersal is only possible under suitable weather conditions. It was found that female dispersal was correlated with weather conditions far more strongly than other population groups. Calculations showed that under suitable weather conditions (e.g., 25% of the time being suitable for dispersal), almost 4% of the adult females could be expected to leave the population daily. The overall effect is therefore to displace individuals from the population but to have little effect on densities. Dispersal in other stages was controlled by the suitability of weather conditions but also by other, undetermined factors. It is postulated that L. tenuis has a life history strategy whereby suitable habitats act as sources of spiders, mainly females, which are continually involved in dispersal. Dispersal in other population groups may be triggered by factors such as the avoidance of adverse conditions. This type of life-history strategy is typical of organisms, such as weeds, that are successful in ephemeral habitats and may indicate that this species originally evolved in unpredictable habitats such as dry river beds or coastal areas.     

Flooded meadow communities an analysis of their productivity in a wet year

The productivity of three plant communities differeing in moisture conditions was studied in the river basin of the Dyje near the village of Lan?hot (Southern Moravia). The communities were as follows:Serratuleto-Festucetum commutatae Balátová-Tulá?ková 1963,Gratiola officinalis—Carex praecox-suzae subass.Galium boreale Balátová-Tulá?ková 1963, andGratiola officinalis—Carex praecox-suzae subass.Rorippa silvestris Balátová-Tulá?ková 1963. The associationGratiola officinalis—Carex praecox-suzae subass.Galium boreale appeared as the most productive one, its biomass maximum W=400 g . m^?2 and the maximum R=0.042 g . g^?1 . day^?1 C=4.84 g . m^?2 . day^?1. Owing to extreme moisture conditions in the year 1966, the associationSerratuleto-Festucetum commutatae was also highly productive, as it reached the following maximum of dry matter production: 240 g . m^?2, R=0.0388g . g^?1 . day^?1 C=4.64 g . m^?2 . day^?1. The maximum value of dry matter in the associationGratiola officinalis—Carex praecox-suzae subass.Rorippa silvestris was 220 g.m^?2. Changes in dry matter production of shoots were evaluated statistically. The dry matter in the underground parts of plants in 1 square metre, collected from the 0–25 cm layer varied from 1,000 to 2,000 g.m^?2. Together with records of the increasing dry matter in the shoots the author kept records of the properties of dead material.     

Light limitation of phytoplankton development in an oligotrophic lake - Loch Ness, Scotland

• 1 The underwater light climate in Loch Ness is described in terms of mixing depth (Z_m) and depth of the euphoric zone (Z_eu). During periods of complete mixing, Z_m equates with the mean depth of the loch (132 m), but even during summer stratification the morphometry of the loch and the strong prevailing winds produce a deep thermocline and an epilimnetic mixed layer of about 30 m or greater. Hence, throughout the year the quotient Z_m/Z_eu is exceptionally high and the underwater light climate particularly unfavourable for phytoplankton production and growth.
• 2 Phytoplankton biomass expressed as chlorophyll a is very low in Loch Ness, with a late summer maximum of less than 1.5 mg chlorophyll a m^-3 in the upper 30 m of the water column. This low biomass and the resulting very low photosynthetic carbon fixation within the water column are evidence that a severe restraint is imposed on the rate at which phytoplankton can grow in the loch.
• 3 The chlorophyll a content per unit of phytoplankton biovolume and the maximum, light-saturated specific rate of photosynthesis are both parameters which might be influenced by the light climate under which the phytoplankton have grown. However, values obtained from Loch Ness for both chlorophyll a content (mean 0.0045 mg mm^-3) and maximum photosynthetic rate (1–4 mg C mg Chla^-1 h^-1) are within the range reported from other lakes.
• 4 Laboratory bioassays with the natural phytoplankton community from Loch Ness on two occasions in late summer when the light climate in the loch is at its most favourable, suggest that even then limitation of phytoplankton growth is finely balanced between light and phosphorus limitation. Hence, for most of the year, when the light climate is less favourable, phytoplankton growth will be light limited.
• 5 Quotients relating mean annual algal biomass as chlorophyll a (c. 0.5 mg Chla m^-3) and the probable annual specific areal loading of total phosphorus (0.4–1.7 g TP m^-2 yr^-1) suggest that the efficiency with which phytoplankton is produced in Loch Ness per unit of TP loading is extremely low when compared with values from other Scottish lochs for which such an index has been calculated. This apparent inefficiency can be attributed to suppression of photosynthetic productivity in the water column due to the unfavourable underwater light climate.
• 6 These several independent sources of evidence lead to the conclusion that phytoplankton development in Loch Ness is constrained by light rather than by nutrients. Loch Ness thus appears to provide an exception to the generally accepted paradigm that phytoplankton development in lakes of an oligotrophic character is constrained by nutrient availability.

     

Seed,dispersal, microsite,habitat and recruitment limitation: identification of terms and concepts in studies of limitations

Recently, there is an increase in number of studies concerned with the effect of various types of limitations on species local population size and distribution pattern at the landscape scale. The terminology used to describe these limitations is, however, very inconsistent. Since the different terms often appear in conclusions of the papers, the inconsistency in their use obscures the message of these papers. In this study, we review the current uses of these terms, identify the basic concepts involved in the discussion of a limitation and link the concepts with the currently used terms. Finally, we discuss the experimental approaches used to assess the different types of limitations. We differentiated four basic concepts resulting from the combination of limitation by environment versus ability to grow and spread, and two spatial scales (local and regional scale). The two concepts at each spatial scale are expected to form a gradient of all possible combinations of the two respective types of limitations. In the considerations of various experimental approaches used to assess these limitations, we conclude that sowing experiments, i.e. seed addition into existing populations or seed introduction into unoccupied habitats, are the only reliable types of evidence for the different types of limitations.     

Spatial autocorrelation and dispersal limitation in freshwater organisms

Dispersal can limit the ranges of species and the diversity of communities. Despite its importance, little is known about its role in freshwater habitats and its relation to habitat type (lentic vs. lotic), especially for organisms with cryptic dispersal methods such as plankton. Poor dispersers are expected to show more clumped distributions or greater spatial autocorrelation (SA) in community composition than good dispersers. We examined patterns of SA across freshwater taxa with different dispersal modes (active vs. passive) and their association with habitat type (lake vs. stream) using 18 spatially explicit community composition data sets. We found significant relationships between SA and body size among taxa in lake habitats, but not in streams. However, the increase in SA with body size in lakes was driven entirely by fishes—organisms ranging in size from diatoms to macro-invertebrates showed equivalent levels of SA. These results support the idea that large organisms are less effective dispersers in aquatic environments, resulting in greater SA in community structure over broad scales. Streams may be effectively more connected than lakes as patterns of SA and body size were weaker in lotic habitats. Our data suggest that the critical threshold where greater body size increases dispersal limitation seems to come at the juncture between invertebrates and vertebrates rather than that between unicellular and multicellular organisms as has been previously suggested. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effects of nutrient limitation and stream discharge on the epilithic microbial community in an oligotrophic Arctic stream

Nutrient limitation of epilithic microbial activity is modified by stream discharge and drainage from the tundra surrounding the Kuparuk River, Alaska, USA. During 1984, after three weeks of whole stream enrichment with phosphorus, autotrophic activity per unit biomass had increased in the enriched section of the stream suggesting that phosphorus availability was limiting productivity. In contrast, after three weeks of phosphorus enrichment during 1985, heterotrophic and autotrophic activity was similar in the control and enriched sections of the stream. However, when ammonia or nitrate and phosphorus were added to an in situ bioassay chamber for two weeks, higher community biomass and heterotrophic activity resulted. Ten days later biomass significantly dropped in the unenriched section. Nitrate levels over this period increased four fold concomitantly with decreased stream discharge. Apparently during 1985, nitrogen was limiting epilithic microbial community in the phosphorus enriched section of the Kuparuk River. The significant negative relationship between nitrate concentration and stream discharge observed during 1984 supported the trends seen in 1985. These data suggest that nutrient concentrations which limit epilithic microbial activity and biomass are regulated by the stream discharge and drainage from the surrounding tundra.     

Epiphytes from a deep-water characean meadow in an oligotrophic New Zealand lake: species composition,biomass and photosynthesis

1. The epiphytic flora of a characean meadow in Lake Coleridge, a deep, oligotrophic lake on the South Island of New Zealand, was dominated by diatoms, particularly Eunotia pectinalis and Achnanthes minutissima. The meadows occupied a depth range from 5 to 30 m. Adnate taxa predominated at all depths below 5 m, while increased taxonomic diversity at 5 m resulted from an increased abundance of erect taxa, including chlorophytes and stalked diatoms. 2. Seasonal changes in epiphyte biomass were followed using artificial substrata and by estimating epiphyte chlorophyll a concentration on host plants. The latter required development of a novel technique utilizing the consistent relationship between fucoxanthin and chlorophyll a concentrations in the epiphyton. Epiphyte chlorophyll a on host plants varied with depth and host species between 0.1 and 0.3 mg g^–1 dry weight. Maximum epiphyte biomass was at 10–15 m depth. At depths of 15 m and less, epiphyte chlorophyll a reached a maximum of ≈ 200–300 mg m^–2 in mid-summer, while at greater depths maximum biomass was less and coincided with a period of clear water in spring. 3. Photosynthetic carbon fixation was estimated from photosynthesis–radiation curves and estimates of radiation flux at sampling depths. At depths greater than 10 m, variability of the vertical extinction coefficient of lake water rather than seasonal fluctuations in incident radiation were responsible for determining the temporal pattern of production. Chlorophyll a-specific photosynthesis was estimated to peak in summer at 5 m (8 mg mg^–1 day^–1), and in spring at all other depths. 4. Annual epiphyte production was estimated as 27 g C m^–2 year^–1 at 5 m depth, falling to 15 g C m^–2 year^–1 at 15 m and 1 g C m^–2 year^–1 at 30 m. Areal biomass changes tended to be temporally but not quantitatively coupled to estimated in situ photosynthesis, and we hypothesize that epiphyte biomass may have been controlled by grazing gastropod snails.     

Diversity and persistence of arbuscular mycorrhizas in a low-Arctic meadow habitat

Little is known about the ecology and diversity of arbuscular mycorrhizal (AM) fungi in Arctic ecosystems. Here, the diversity and composition of the AM fungal community and its response to host plant community composition were studied in a low-Arctic meadow habitat. The natural vegetation in two low-Arctic meadow sites was manipulated. Plots with natural vegetation, monoculture and no vegetation were established. Seeds of Solidago virgaurea were sown into the plots and the AM fungal community in the seedling roots was analysed using the terminal restriction fragment length polymorphism (T-RFLP) method. The vegetation manipulation treatments affected the community composition but not the diversity of AM fungi found in S. virgaurea roots. The diversity of AM fungi was higher in S. virgaurea roots in the site with naturally higher plant species diversity. These results show that AM fungi in low-Arctic meadows are able to survive for a period of 2 yr without a host plant. This ability buffers the AM fungal community against short-term changes in host plant community composition and diversity.     

Spatial models of pollen dispersal in the forage grass meadow fescue

Several bivariate probability distributions, generated by different underlying dispersal mechanisms, are fitted to the observed frequencies of an isozyme marker gene using a maximum likelihood approach. The pollen dispersal data were generated using two experimental populations of meadow fescue (Festuca pratensis Huds.), homozygous for different allozymes at the (Pgi-2) locus, arranged in a circular donor–acceptor field design. The contribution of a plant depends on plant position, fecundity and flowering time, factors which are taken into account when fitting the different models. Several approximate likelihood-ratio tests are done between alternative nested models, and a wind threshold model with bimodality in the wind direction is selected. The evolutionarily important variances and expectations of gene displacement under the selected model are calculated. It is also shown that the underlying probability distribution is significantly more than exponentially leptokurtic. By fitting a distribution of deposition in all three dimensions to the data, taking into account differences in plant height, separate estimates of additional physical parameters are obtained, showing that gravity and vertical random movements are more important than intervening vegetation in limiting pollen dispersal in meadow fescue. According to the model, plants with a high seed yield contribute pollen over-proportionally to neighbouring plants. This revised version was published online in July 2006 with corrections to the Cover Date.     

Distinguishing the roles of dispersal in diversity maintenance and in diversity limitation

Considerable recent research effort has gone into studying how dispersal might affect the diversity of local communities. While this general topic has received attention from theoretical and empirical ecologists alike, the research focus has differed between the two groups; theoretical ecologists have explored the role of dispersal in the maintenance of diversity within local communities, whereas empirical ecologists have sought to quantify the role of dispersal in limiting local diversity. We argue that there is no necessary relationship between these two components of diversity and we therefore need to develop empirical approaches to quantify the dispersal-maintained component of diversity, as well as the dispersal-limited component. We develop one such approach in this paper, based on a quantitative partitioning of the natural regeneration within intact communities onto different sources of recruits (local communityvs. dispersal across different spatial or temporal scales).     

Nutrient limitation along a productivity gradient in wet meadows

Conservation management in meadows often focuses on reducing soil fertility and consequently community productivity as to promote and sustain species-rich vegetations. The productivity level to which nutrients are limiting growth is, however, unclear, as well as the relationship between productivity and the type of nutrient limitation. We carried out a fertilization experiment with N, P and K in six annually mown meadows along an aerial phytomass gradient (200–650 g m^–2). All meadows were found to be growth-limited by nutrients. Low-productive meadows were N-limited, or N+P co-limited, whereas our higher productive meadows were co-limited by a combination of N, P and/or K. The results from our experiments were compared with the results from 45 other fertilization experiments with N, P and K in grasslands and wetlands (aerial phytomass range 50–1500 g m^–2). Our results were consistent in nitrogen being the most frequent (co)-limiting nutrient, and regarding the equal frequence of occurrence of P (co)-limitation and K (co)-limitation (both in ca. 25–30% of all sites). Co-limitation occurred more often in our sites than in the other experiments. There was no clear relationship between aerial phytomass and type of nutrient limitation, except that K (co)-limitation only occurred at sites with phytomass above 200 g m^–2, and P (co)-limitation below 600 g m^–2. A comparison of productivity and nutrient concentrations in aerial phytomass among two years indicated that the type of nutrient limitation is not a static site characteristic but may vary with dynamic environmental conditions such as soil wetness; drought seems to enhance N-availability which may induce P- and K-limitation.     

Patterns of dispersal and dynamics among habitat patches varying in quality

Both source-sink theory and extensions of optimal foraging theory ("balanced dispersal" theory) address dispersal and population dynamics in landscapes where habitat patches vary in quality. However, studying dispersal mechanisms empirically has proven difficult, and dispersal is rarely tied back to long-term spatial dynamics. We used a manipulable laboratory system consisting of bacteria and protozoa to investigate the ability of source-sink and optimal foraging theories to explain both dispersal and emergent spatial dynamics. Consistent with source-sink models and contrary to balanced dispersal models, there was a consistent net flux of protist individuals from high to low resource patches. However, unlike the simplest source-sink models, intermediate rates of dispersal led to highest abundances in low resource patches. Side experiments found strong density dependence in local population dynamics and differences in average protist body size in high and low resource patches. Parameterization and analysis of a two-patch model showed that high migration from high to low resource patches could have depressed population density in low resource patches, creating pseudosinks. The movement of individuals and biomass from sources to sinks (a form of ecosystem subsidy) resulted in the convergence of body size and population densities in sources and sinks. Our results indicate a need to carefully consider movement patterns and interaction with local dynamics in potential source-sink systems.     

How dispersal limitation shapes species-body size distributions in local communities

A critical but poorly understood pattern in macroecology is the often unimodal species-body size distribution (also known as body size-diversity relationship) in a local community (embedded in a much larger regional species pool). Purely neutral community models that assume functional equivalence among species are incapable of explaining this pattern because body size is the key determinant of functional differences between species. Several niche-based explanations have been offered, but none of them is completely satisfactory. Here we develop a simple model that unites a neutral community model with niche-based theory to explain the relationship. In the model, species of similar size are assumed to belong to the same size guild. Within a size guild, all individuals are equivalent in their competition for resources, sensu Hubbell's neutral community model; they have the same speciation rate and dispersal capacities. Between size guilds, however, the total number of individuals, the speciation rate, and the dispersal capacities differ, but using known allometric scaling laws for these properties, we can describe the differences between size guilds. Our model predicts that species richness reaches an optimum at an intermediate body size, in agreement with observations. The optimum at intermediate body size is basically the result of a trade-off between, on the one hand, allometric scaling laws for the number of individuals and the speciation rate that decrease with body size and, on the other hand, the scaling law for active dispersal that increases with body size.