Environmental filtering predicts plant-community trait distribution and diversity: Kettle holes as models of meta-community systems.

Meta-communities of habitat islands may be essential to maintain biodiversity in anthropogenic landscapes allowing rescue effects in local habitat patches. To understand the species-assembly mechanisms and dynamics of such ecosystems, it is important to test how local plant-community diversity and composition is affected by spatial isolation and hence by dispersal limitation and local environmental conditions acting as filters for local species sorting.We used a system of 46 small wetlands (kettle holes)-natural small-scale freshwater habitats rarely considered in nature conservation policies-embedded in an intensively managed agricultural matrix in northern Germany. We compared two types of kettle holes with distinct topographies (flat-sloped, ephemeral, frequently plowed kettle holes vs. steep-sloped, more permanent ones) and determined 254 vascular plant species within these ecosystems, as well as plant functional traits and nearest neighbor distances to other kettle holes.Differences in alpha and beta diversity between steep permanent compared with ephemeral flat kettle holes were mainly explained by species sorting and niche processes and mass effect processes in ephemeral flat kettle holes. The plant-community composition as well as the community trait distribution in terms of life span, breeding system, dispersal ability, and longevity of seed banks significantly differed between the two habitat types. Flat ephemeral kettle holes held a higher percentage of non-perennial plants with a more persistent seed bank, less obligate outbreeders and more species with seed dispersal abilities via animal vectors compared with steep-sloped, more permanent kettle holes that had a higher percentage of wind-dispersed species. In the flat kettle holes, plant-species richness was negatively correlated with the degree of isolation, whereas no such pattern was found for the permanent kettle holes.Synthesis: Environment acts as filter shaping plant diversity (alpha and beta) and plant-community trait distribution between steep permanent compared with ephemeral flat kettle holes supporting species sorting and niche mechanisms as expected, but we identified a mass effect in ephemeral kettle holes only. Flat ephemeral kettle holes can be regarded as meta-ecosystems that strongly depend on seed dispersal and recruitment from a seed bank, whereas neighboring permanent kettle holes have a more stable local species diversity.

Biogeochemistry of natural ponds in agricultural landscape: Lessons learned from modeling a kettle hole in Northeast Germany.

Kettle holes, small shallow ponds of glacial origin, represent hotspots for biodiversity and biogeochemical cycling. They abound in the young moraine landscape of Northeast Germany, potentially modulating element fluxes in a region where intensive agriculture prevails. The Rittgarten kettle hole, with semi-permanent hydroperiod and a surrounding reed belt, can be considered as a representative case study for such systems. Aiming to provide insights into the biogeochemical processes driving nutrient and primary producer dynamics in the Rittgarten kettle hole, we developed a mechanistic model that simulates the carbon, nitrogen, phosphorus and oxygen, phytoplankton, and free-floating macrophyte biomass dynamics. After model calibration and sensitivity analysis, our modeling exercise quantified the simulated nutrient fluxes associated with all the major biogeochemical processes considered by the model. Seasonality of nutrient concentrations, magnitude of primary productivity rates, and biogeochemical process characterization in the pond were reasonably reproduced by the model from July 2013 to July 2014. Our results suggest that the establishment of a phytoplankton community well-adapted to low light availability, together with the differential use of N and P from free-floating macrophytes and phytoplankton can explain their coexistence in kettle holes. Sediment nutrient release along with decomposition of decaying submerged macrophyte are essential drivers of internal nutrient cycling in kettle holes. Our results also suggest that the Rittgarten kettle hole act as a net source of CO2 to the atmosphere on an annual scale, which offers a testable hypothesis for kettle holes with structural and functional similarities. We conclude by discussing the need to shed light on the effects of water level fluctuations on nutrient dynamics and biological succession patterns, as well as the relative importance of external sources and internal nutrient recycling mechanisms.

Land-use and hydroperiod affect kettle hole sediment carbon and nitrogen biogeochemistry.

Kettle holes are glaciofluvially created depressional wetlands that collect organic matter (OM) and nutrients from their surrounding catchment. Kettle holes mostly undergo pronounced wet-dry cycles. Fluctuations in water table, land-use, and management can affect sediment biogeochemical transformations and perhaps threaten the carbon stocks of these unique ecosystems. We investigated sediment and water of 51 kettle holes in NE Germany that differ in hydroperiod (i.e. the duration of the wet period of a kettle hole) and land-use. Our objectives were 1) to test if hydroperiod and land management were imprinted on the isotopic values (δ13C, δ15N) and C:N ratios of the sediment OM, and 2) to characterize water loss dynamics and kettle hole-groundwater connectivity by measuring the stable δ18O and δD isotope values of kettle hole water over several years. We found the uppermost sediment layer reflected recent OM inputs and short-term processes in the catchment, including land-use and management effects. Deeper sediments recorded the degree to which OM is processed within the kettle hole related to the hydroperiod. We see clear indications for the effects of wet-dry cycles for all kettle holes, which can lead to the encroachment of terrestrial plants. We found that the magnitude of evaporation depended on the year, season, and land-use type, that kettle holes are temporarily coupled to shallow ground water, and, as such, kettle holes are described best as partially-closed to open systems.

Sediment cores from kettle holes in NE Germany reveal recent impacts of agriculture.

Glacial kettle holes in young moraine regions receive abundant terrigenous material from their closed catchments. Core chronology and sediment accumulation were determined for two semi-permanent kettle holes, designated RG and KR, on arable land close to the villages of Rittgarten and Kraatz, respectively, in Uckermark, NE Germany. Core dating ((210)Pb, (137)Cs) revealed variable sediment accretion rates through time (RG 0.4-23.1 mm a(-1); KR 0.2-35.5 mm a(-1)), with periods of high accumulation corresponding to periods of intensive agricultural activity and consequent erosional inputs from catchments. Sediment composition (C, N, P, S, K, Ca, Fe, Mn, Zn, Cu, Mo, Pb, Cd, Zr) was used to determine sediment source and input processes. At RG, annual P input increased from 0.65 kg ha(-1) in the early nineteenth century to 1.67 kg ha(-1) by 2013. At KR, P input increased from 0.6 to 4.1 kg ha(-1) over the last century. There was a concurrent increase in Fe input in both water bodies. Thus, Fe/P ratios showed no temporal trend and did not differ between RG (18.5) and KR (18.4), indicating similar P mobility. At RG, the S/Fe ratio increased from 0.4 to 2.3, indicating more iron sulphides and thus higher P availability, coinciding with high coverage of duckweed (Spirodela polyrhiza (L.)) and soft hornwort (Ceratophyllum submersum L.). At KR, however, this ratio remained low and relatively unchanged (0.3 ± 0.4), indicating more efficient Fe-P binding and lower hydrophyte productivity. Trends in sediment composition indicate a shift towards eutrophication in both kettle holes, but with differences in timing and magnitude. Other morphologically similar kettle holes in NE Germany that are prone to erosion could have been similarly impacted but may differ in the extent of sediment infilling and degradation of their ecological functions.

Screening of prevailing processes that drive surface water quality of running waters in a cultivated wetland region of Germany - a multivariate approach.

The Spreewald wetland is a large, peaty, inland delta wetland in which the water level is managed by weirs in cascade belts across an extensive, partly channelized running water system. To model the quality of the water, 946 surface water samples from 43 sites were analyzed for 29 water quality parameters in two monitoring programs spanning a period of six years. In this study, we pursued a multivariate approach using nonlinear principal component analysis (Isomap) to identify the prevailing processes that control the water quality of the complex surface water system. The first four principal components explained 79% of the variance in the dataset. These components were interpreted as anthropogenic impact factors, such as groundwater exfiltration from degraded peat areas and the influence of coal mining drainage with respect to SO(4), as well as groundwater exfiltration from mineral aquifers, and phytoplankton growth and competition. A sub-area of the Spreewald wetland, characterized by a sandy aquifer overlain by degraded peat, had the greatest impact on downstream surface water quality for most of the investigated parameters. In order to achieve better water quality in the Spreewald wetland, pollutant input - particularly SO(4) input from the tributaries - must be controlled by enhancing the wetland's buffer capacity in the catchment, and peat mineralization and groundwater exfiltration must be minimized by raising the water table of the peatland area and receiving waters. The results show that Isomap is a very powerful tool for gaining a better insight into the dominating processes defining the surface water quality of complex wetland systems. Nevertheless, to be able to draw the right conclusions from multivariate statistical approaches such as Isomap it is necessary to possess basic knowledge of the structure of the system and of the processes that may occur.