Relationships between land snail assemblage patterns and soil properties in temperate-humid forest ecosystems

Aim The objectives were to (1) analyse the combined effects of soil pH, Ca content and soil moisture on total density and species richness of land snails in forest ecosystems, (2) explore relationships between the quantitative composition of snail assemblages and habitat characteristics, (3) investigate the relationships between soil pH and density of some of the most frequent species, and (4) compare the data with those from studies conducted in other temperate‐humid regions of Europe. Location Study sites were selected from 15 landscape types including different lithologies within the area of Baden‐Württemberg (35,000 km²), SW Germany. Methods Snails were recorded quantitatively from 83 study sites, with four plots representing a total of 0.25 m² per site. Topsoil samples from each site were analysed for pH, exchangeable Ca, and Ca content of carbonates. Three categories of soil moisture (dry, intermediate and wet) were established and defined according to the (climatic) water balance. Numbers of individuals and species were brought in relation to soil moisture and soil pH. Cluster analyses were conducted to identify groups of quantitatively similar snail species assemblages. Results Topsoil pH (2.7–7.5) and soil Ca contents were closely correlated. On dry soils, total snail density and species richness are generally low and do not change with pH, but clearly increase with increasing pH on intermediate moisture soils and on wet soils. On the latter, numbers of individuals and species are generally much higher compared with intermediate moisture sites at the same value of soil pH. Changes of density in relation to soil pH vary between species. Depending on the species, density increases only in the lower or only in the higher range of pH, is not related to pH, or decreases with increasing pH. Furthermore, these patterns vary within the same species depending on the region. This became evident from comparisons with other studies, particularly between sites in SW Germany and southern Scandinavia. From cluster analyses, subgroups of snail assemblages of high quantitative similarity were identified. Group formation is explained by soil pH to some extent, and one subgroup showed a connection with coniferous woodland sites on acidic soils. No further environmental factors available from our data could explain the clustering of snail assemblages more detailed. Main conclusions Soil moisture is the strongest determinant of snail density and species richness at undisturbed woodland sites, but effects of soil moisture and soil pH on these patterns are closely interrelated on intermediate moisture soils and wet soils. However, the quantitative species composition of the land snail assemblages is related to soil properties to a lower degree than snail density and species richness, and other habitat characteristics such as vegetation or litter quality, can be important for species dominance in addition.     

Macrofauna assemblage composition and soil moisture interact to affect soil ecosystem functions

Changing climatic conditions and habitat fragmentation are predicted to alter the soil moisture conditions of temperate forests. It is not well understood how the soil macrofauna community will respond to changes in soil moisture, and how changes to species diversity and community composition may affect ecosystem functions, such as litter decomposition and soil fluxes. Moreover, few studies have considered the interactions between the abiotic and biotic factors that regulate soil processes. Here we attempt to disentangle the interactive effects of two of the main factors that regulate soil processes at small scales - moisture and macrofauna assemblage composition. The response of assemblages of three common temperate soil invertebrates (Glomeris marginata Villers, Porcellio scaber Latreille and Philoscia muscorum Scopoli) to two contrasting soil moisture levels was examined in a series of laboratory mesocosm experiments. The contribution of the invertebrates to the leaf litter mass loss of two common temperate tree species of contrasting litter quality (easily decomposing Fraxinus excelsior L. and recalcitrant Quercus robur L.) and to soil CO₂ fluxes were measured. Both moisture conditions and litter type influenced the functioning of the invertebrate assemblages, which was greater in high moisture conditions compared with low moisture conditions and on good quality vs. recalcitrant litter. In high moisture conditions, all macrofauna assemblages functioned at equal rates, whereas in low moisture conditions there were pronounced differences in litter mass loss among the assemblages. This indicates that species identity and assemblage composition are more important when moisture is limited. We suggest that complementarity between macrofauna species may mitigate the reduced functioning of some species, highlighting the importance of maintaining macrofauna species richness.     

Assignment and evaluation of ground beetle (Coleoptera: Carabidae) assemblages to sites on different scales in a grassland landscape

Carabid beetles and environmental parameters were investigated in 52 grassland sites with three replicate pitfall traps in each site and in the valley of the River Eider in Schleswig–Holstein (northern Germany) with 61 pitfall traps. Environmental parameters included water content of soil, sand content, organic matter content and pH. Ground beetle assemblages were derived by detrended correspondence analyses (DCA) and characterised by the specific environmental conditions as means for each assemblage. On the regional scale including all investigated sites of Schleswig–Holstein, five assemblages were differentiated. On the local scale including the investigated sites in the valley of the River Eider, three assemblages were found corresponding well with those found on the regional scale. Environmental conditions at the sites of the five assemblages were correlated with land use data, soil types, and water level stages provided by three maps of a geographic information system (GIS). The GIS maps were combined to develop smaller areas with land use, soil type and water level stage information. The characteristic environmental conditions were assigned to each area to derive the spatial distribution of the five ground beetle assemblages. Spatial prediction was correct for 65% of investigated sites. The potential area of each assemblage was estimated for the valley. The different grassland areas were evaluated as potential habitats for ground beetle species comparing total species richness with the regional species richness of each assemblage. The comparison shows that species richness in the evaluated assemblages is relatively low compared to the regional potential.     

Vegetation changes along gradients of long-term soil development in the Hawaiian montane rainforest zone

The development of the Hawaiian montane rainforest was investigated along a 4.1-million-year soil age gradient at 1200 m elevation under two levels of precipitation, the mesic (c. 2500 mm annual rainfall) vs. wet (>4000 mm) age gradient. Earlier analyses suggested that soil fertility and foliar nutrient concentrations of common canopy species changed unimodally on the same gradients, with peak values at the 20,000–150,000 yr old sites, and that foliar concentrations were consistently lower under the wet than under the mesic conditions. Our objectives were to assay the influences of soil aging and moisture on forest development using the patterns and rates of species displacements. The canopies at all sites were dominated by Metrosideros polymorpha. Mean height and dbh of upper canopy Metrosideros trees increased from the youngest site to peak values at the 2100–9000 yr sites, and successively declined to older sites. A detrended correspondence analysis applied to mean species cover values revealed that significant variation among sites occurred only on one axis (axis 1), for both soil-age gradients. Sample scores along axis 1 were perfectly correlated with soil age on the mesic gradient, and significantly correlated on the wet gradient. Higher rainfall appeared to be responsible for the higher rates of species turnover on the wet gradient probably through faster rock weathering and greater leaching of soil elements. We concluded that the changes in species cover values and size of the canopy species was a reflection of the changing pattern of nutrient availability associated with soil aging.     

Changes in ectomycorrhizal community structure on two containerized oak hosts across an experimental hydrologic gradient

Shifts in ectomycorrhizal (ECM) community structure were examined across an experimental hydrologic gradient on containerized seedlings of two oak species, Quercus montana and Quercus palustris, inoculated from a homogenate of roots from mature oak trees. At the end of one growing season, seedlings were harvested, roots were sorted by morphotype, and proportional colonization of each type was determined. DNA was subsequently extracted from individual root tips for polymerase chain reaction, restriction fragment length polymorphism, and rDNA sequencing of the ITS1/5.8S/ITS2 region to determine identities of fungal morphotypes. Twelve distinct molecular types were identified. Analysis of similarity showed that ECM fungal assemblages shifted significantly in composition across the soil moisture gradient. Taxa within the genus Tuber and the family Thelephoraceae were largely responsible for the changes in fungal assemblages. There were also significant differences in ECM community assemblages between the two oak host species. These results demonstrate that the structure of ECM fungal communities depends on both the abiotic and biotic environments and can shift with changes in soil moisture as well as host plant, even within the same genus.     

Ecological species groups of landform-level ecosystems dominated by jack pine in northern Lower Michigan,USA

A combination of field and tabular methods and multivariate analyses were used to develop groups of ground flora species (i.e., ecological species groups) that characterize and distinguish highly disturbed, landform-level ecosystems dominated by jack pine in northern Lower Michigan. The endangered Kirtland's warbler formerly or currently occupied the large glacial landforms for which species groups were developed. Eight such ecological species groups were created using 31 woody and herbaceous species sampled in 144 plots within a 20,000 km² geographic region of uniform climate and regional physiography. The groups were initially constructed using subjective, observation-based groupings of species with similar presence and abundance along soil moisture and fertility gradients. Species groups were corroborated using TWINSPAN and detrended correspondence analysis, and the environmental conditions indicated by each were described and contrasted based upon field observations and canonical correspondence analysis. Two of the eight species groups indicated very dry, infertile sites, and one was indicative of a very broad range of sites dominated by jack pine. The remaining five groups reflected a relatively gentle environmental gradient within the set of ecosystems we sampled, indicating small differences along a soil moisture gradient and less so along a soil fertility and light availability gradient. The groups were applied successfully for the majority of an area that had been repeatedly logged and/or burned for at least 120 years. In addition, the groups were successful when tested on sites with highly disturbed upper soil strata furrowed for the establishment of plantations between the 1960s and 1980s. Within the boundaries of the regional ecosystems for which they were developed, ecological species groups reflect the integrated effects of multiple site factors that control the height growth rates of jack pine trees that, in turn, determine the duration of Kirtland's warbler occupancy. While usually applied at the scale of ecosystem types, our results demonstrate that ecological species groups may also provide the ecological basis for distinguishing ecosystems at broader scales. When examined simultaneously in the field with physiography, microclimate, and soil factors, the groups are therefore useful in identifying and classifying ecosystem units at the scale of landforms, the appropriate scale of management for the Kirtland's warbler.     

Possible interactions between environmental factors in determining species optima

Questions: 1. Indicator values, such as those of Ellenberg, for different environmental factors are seen as independent. We tested for the presence of interactions between environmental factors (soil moisture and reaction) to see if this assumption is simplistic. 2. How close are Ellenberg indicator values (IV s) related to the observed optima of species response curves in an area peripheral to those where they have been previously employed and 3. Can the inclusion of bryophytes add to the utility of IVs? Location: South Uist, Outer Hebrides, Scotland, UK. Methods: Two grids (ca. 2000 m × 2000 m) were sampled at 50‐m intervals across the transition from machair to upland communities covering an orthogonal gradient of both soil pH (reaction) and soil moisture content. Percentage cover data for vascular plants, bryophytes and lichens were recorded, along with pH and moisture content of the underlying sand/soil/ peat. Reaction optima, derived from species response curves calculated using HOF models, were compared between wet and dry sites, and moisture optima between acidic and basic samples. Optima for the whole data set were compared to Ellenberg IVs to assess their performance in this area, with and without the inclusion of bryophytes. Results: A number of species showed substantially different pH optima at high and low soil moisture contents (18% of those tested) and different soil moisture optima at high and low pH (49%). For a number of species the IVs were poor predictors of their actual distribution across the sampled area. Bryophytes were poor at explaining local variation in the environmental factors and also their inclusion with vascular plants negatively affected the strength of relationships. Conclusions: A substantial number of species showed an interaction between soil moisture and reaction in determining their optima on the two respective gradients. It should be borne in mind that IVs such as Ellenberg's may not be independent of one another.     

Distinct edaphic habitats are occupied by discrete legume assemblages with unique indicator species in the Cape Peninsula of South Africa

Aims The Cape Peninsula is a small area (471 km 2) situated on the south-westernmost tip of the Core Cape Subregion (CCR) of South Africa. Within the Cape Peninsula, Fabaceae are the third most species-rich plant family (162 species) and they have the second highest number of endemic species after the Ericaceae. However, legumes are not the dominant taxa in the vegetation. They tend to show patchy distributions within the landscape and different species assemblages usually occupy particular niches at any given locality. The present study undertook to establish if edaphic factors influence legume species distribution in the Cape Peninsula and to determine the key indicator species for the different assemblages.Methods Soils from 27 legume sites, spanning all major geological substrates of the Cape Peninsula, were analysed for 31 chemical and physical properties. Legume species present at each site were recorded and a presence/absence matrix was generated. Cluster analysis and discriminant function analysis (DFA) were run to group the sites based on overall similarity in edaphic characteristics and to identify the soil parameters contributing towards discriminating the groups. Canonical correspondence analysis (CCA) was used to test for a correlation between legume species compositions and edaphic factors. The strength of the association between legume species and site groupings based on edaphic properties was assessed using indicator species analysis.Important findings Based on similarity in overall soil characteristics, the sites formed three clusters: one comprising sites of sandstone geology, one with dune sand sites and the third cluster comprising sites of both shale and granite geologies (hereafter referred to as soil types). The DFA confirmed the distinctness of these clusters and the CCA showed a significant correlation between legume species composition and edaphic factors. The key edaphic parameters were clay content, iron (Fe), potassium (K), sulphur (S) and zinc (Zn). These findings reveal that the Cape Peninsula is edaphically heterogeneous and edaphically distinct habitats contain discrete legume species assemblages that can be distinguished by unique indicator species. Furthermore, multiple soil parameters, rather than a single parameter, are involved. Therefore, edaphic factors play a significant role in driving the distribution of legume species in the Cape Peninsula and discrete legume species assemblages occupy distinct habitats.     

Reliability of Ellenberg indicator values for moisture,nitrogen and soil reaction: a comparison with field measurements

Abstract. Ellenberg indicator values for moisture, nitrogen and soil reaction were correlated with measured soil and vegetation parameters. Relationships were studied through between‐species and between‐site comparisons, using data from 74 roadside plots in 14 different plant communities in The Netherlands forming a wide range. Ellenberg moisture values correlated best with the average lowest moisture contents in summer. Correlations with the annual average groundwater level and the average spring level were also good. Ellenberg N‐values appeared to be only weakly correlated with soil parameters, including N‐mineralization and available mineral N. Instead, there was a strong relation with biomass production. We therefore endorse Hill & Carey's (1997) suggestion that the term N‐values be replaced by ‘productivity values'. For soil reaction, many species values appeared to need regional adjustment. The relationship with soil pH was unsatisfactory; mean indicator values were similar for all sites at pH > 4.75 because of wide species tolerances for intermediate pH levels. Site mean reaction values correlated best (r up to 0.92) with the total amount of calcium (exchangeable Ca²⁺ plus Ca from carbonates). It is therefore suggested that reaction values are better referred to as ‘calcium values'. Using abundance values as weights when calculating mean indicator values generally improved the results, but, over the wide range of conditions studied, differences were small. Indicator values for bryophytes appeared well in line with those for vascular plants. It was noted that the frequency distributions of indicator values are quite uneven. This creates a tendency for site mean values to converge to the value most common in the regional species pool. Although the effect on overall correlations is small, relationships tended to be less linear. Uneven distributions also cause the site mean indicator values at which species have their optimum to deviate from the actual Ellenberg values of these species. Suggestions for improvements are made. It is concluded that the Ellenberg indicator system provides a very valuable tool for habitat calibration, provided the appropriate parameters are considered.     

Chironomidae (Diptera) larvae assemblages differ along an altitudinal gradient and temporal periods in a subtropical montane stream in Northwest Argentina

The objective of this study was to analyze the spatial and temporal dynamics of Chironomidae larvae assemblages in a subtropical mountain river basin of northwestern Argentina. We hypothesized that they would respond to (1) environmental changes along the altitudinal gradient in the Yungas forests; (2) environmental changes caused by spates (e.g., increased discharge or sediment transportation). We sampled five sites along an altitudinal gradient of ca. 1,500 m (from 680 to 2,170 m) during low-water and high-water period for 2 years. The Chironomidae larvae assemblages varied along the altitudinal gradient and between sampling periods based on an indicator species analyses and an ordination analyses (NMDS). The best indicator for high altitude sites was “Genus X” followed by Onconeura sp.2, “Genus 9”, “Genus 10”, and Rheotanytarsus sp.2; while Rheotanytarsus lamellatus, Oliveiriella sanjavieri, and Thienemanniella sp. were good indicators for low altitude sites. O. sanjavieri and R. lamellatus were the best indicators for high-water period. The most relevant environmental variables influencing their spatial and temporal distribution were altitude, water temperature, conductivity, and pH. This study is the first to analyze and describe in detail the spatial and temporal distribution of Chironomidae larvae assemblages in northwestern Argentina, and most of the taxa reported here are new records for the region.