Habitat-based biodiversity responses to macroclimate and edaphic factors in European fen ecosystems.

Understanding large-scale drivers of biodiversity in palustrine wetlands is challenging due to the combined effects of macroclimate and local edaphic conditions. In boreal and temperate fen ecosystems, the influence of macroclimate on biodiversity is modulated by hydrological settings across habitats, making it difficult to assess their vulnerability to climate change. Here, we investigate the influence of macroclimate and edaphic factors on three Essential Biodiversity Variables across eight ecologically defined habitats that align with ecosystem classifications and red lists. We used 27,555 vegetation plot samples from European fens to assess the influence of macroclimate and groundwater pH predictors on the geographic distribution of each habitat type. Additionally, we modeled the relative influence of macroclimate, water pH, and water table depth on community species richness and composition, focusing on 309 plant specialists. Our models reveal strong effects of mean annual temperature, diurnal thermal range, and summer temperature on biodiversity variables, with contrasting differences among habitats. While macroclimatic factors primarily shape geographic distributions and species richness, edaphic factors emerge as the primary drivers of composition for vascular plants and bryophytes. Annual precipitation exhibits non-linear effects on fen biodiversity, with varying impact across habitats with different hydrological characteristics, suggesting a minimum requirement of 600 mm of annual precipitation for the occurrence of fen ecosystems. Our results anticipate potential impacts of climate warming on European fens, with predictable changes among habitat types and geographic regions. Moreover, we provide evidence that the drivers of biodiversity in boreal and temperate fens are closely tied to the ecological characteristics of each habitat type and the dispersal abilities of bryophytes and vascular plants. Given that the influence of macroclimate and edaphic factors on fen ecosystems is habitat specific, climate change research and conservation actions should consider ecological differentiation within functional IUCN ecosystems at continental and regional scales.

Conservation and restoration of Central European fens by mowing: A consensus from 20 years of experimental work.

While the importance of conservation mowing for mesic grasslands is generally accepted, its use for fens and fen grasslands interspersed within agricultural land is still controversial. Although fens may persist naturally, ongoing environmental changes increase productivity and accelerate succession. These processes can be mitigated through conservation management with appropriate settings. However, long-term management experiments are challenging and provide only locally valid results. Here, we analysed vegetation data (bryophytes and vascular plants) from seven management experiments (spanning 3-20 years) conducted in Central European poor, moderately-rich, and calcareous spring fens (Czech Republic, Slovakia). Two of these experiments examined the effects of restoration of abandoned fens, while five experiments examined changes in mowing regimes in managed fens (cessation, intensification, delay to autumn, and litter removal). Data were analysed using unidimensional and multidimensional methods separately for the initial, extended, and entire period. Mowing had a statistically significant effect on species composition except for the shortest (3-year) experiment. Litter removal did not compensate for mowing. Mowing twice or delayed mowing significantly affected the species composition of calcareous fens. In all cases, cessation of mowing significantly reduced the richness of species, especially those of conservation importance. In contrast, any mowing of abandoned fens increased species richness. The effects of mowing intensification or cessation on species richness and composition of a restored calcareous fen were evident in the first 2-3 years. Other effects were initially weak or nonsignificant but later became stronger, such as mowing delay and restoration removal of litter, which became significant only after nearly 20 years. We found that cessation or restoration of mowing usually triggers a rapid vegetation change, whereas it can take decades to detect the response caused by changes in mowing timing. Importantly, mowing can stabilise or even restore vegetation of fen ecosystems that have been weakened by their fragmentation in the temperate agricultural landscapes.

Rising temperature modulates pH niches of fen species.

Rising temperatures may endanger fragile ecosystems because their character and key species show different habitat affinities under different climates. This assumption has only been tested in limited geographical scales. In fens, one of the most endangered ecosystems in Europe, broader pH niches have been reported from cold areas and are expected for colder past periods. We used the largest European-scale vegetation database from fens to test the hypothesis that pH interacts with macroclimate temperature in forming realized niches of fen moss and vascular plant species. We calibrated the data set (29,885 plots after heterogeneity-constrained resampling) with temperature, using two macroclimate variables, and with the adjusted pH, a variable combining pH and calcium richness. We modelled temperature, pH and water level niches for one hundred species best characterizing European fens using generalized additive models and tested the interaction between pH and temperature. Fifty-five fen species showed a statistically significant interaction between pH and temperature (adj p Ë‚ .01). Forty-six of them (84%) showed a positive interaction manifested by a shift or restriction of their niche to higher pH in warmer locations. Nine vascular plants and no moss showed the opposite interaction. Mosses showed significantly greater interaction. We conclude that climate significantly modulates edaphic niches of fen plants, especially bryophytes. This result explains previously reported regional changes in realized pH niches, a current habitat-dependent decline of endangered taxa, and distribution changes in the past. A warmer climate makes growing seasons longer and warmer, increases productivity, and may lower the water level. These effects prolong the duration and intensity of interspecific competition, support highly competitive Sphagnum mosses, and, as such, force niches of specialized fen species towards narrower high-pH ranges. Recent anthropogenic landscape changes pose a severe threat to many fen species and call for mitigation measures to lower competition pressure in their refugia.

Land snail community patterns related to regional habitat conservation status of European spring fens.

Human activities have enormous impact on current biodiversity distribution across all spatial scales. Despite the numerous studies showing the difference between preserved and impaired sites, only little is known about the regional scale. Therefore, we selected four European regions differing in habitat conservation status (HCS) to explore if the variation in land snail communities reflects regional differences. We collected quantitative land snail samples at 169 isolated spring fen sites and measured environmental parameters. The species richness of habitat specialists expressed low variation and weak associations with local conditions in the two regions of adequate HCS, presumably because of their common occurrence throughout most sites. In contrast, the richness of matrix-derived species, i.e. predominantly habitat generalists, was highly variable in these two regions and also tightly associated with local conditions, especially moisture. In both the intermediate and the inadequate HCS region, these associations were much weaker as the fens are less extreme and allow for penetration of matrix-derived species. Population densities of Vertigo geyeri, an umbrella species internationally protected by the EU Habitats Directive, were highest in the two adequate HCS regions. Species composition was primarily controlled by moisture in the regions of adequate HCS, while in the remaining regions, those predictors that are less easily jeopardized by human impact, such as climate, water chemistry and terrain topography, gained importance. In the inadequate HCS region, none of the analysed predictors was associated with the main compositional gradient, suggesting a complete disruption of community-environment relationships. Our results suggest that the species richness and community responses to natural gradients might be substantially modified by human impact, although the effect of some other region-specific factors cannot be easily disentangled because of inevitably low number of studied regions.

Current Condition of Pannonic Salt Steppes at Their Distribution Limit: What Do Indicator Species Reveal about Habitat Quality?

Little is known about the suite of ecological conditions under which characteristic species may continue to develop under the pressure of recent habitat deterioration. We aimed to determine the niche of three indicator species of the priority habitat Pannonic salt steppes and to find out how their vegetation composition, land use, and soil chemistry mirror the current condition of their typical habitat. A plot-based vegetation survey was conducted in degraded and in pristine (reference) inland salt steppes in East-Central Europe. We confirmed decreased habitat quality at their northern geographical limit. Most of the sites there showed a strong prevalence of generalists (e.g., Elytrigia repens) and lack of specialists, both resulting from lowered habitat extremity and inappropriate land use (abandonment). A small proportion of plots (19%) were in the same good condition as the reference vegetation in the central area. Soil analyses revealed that the studied halophytes are able to persist on desalinized soils if the land use is suitable. The occurrence of the annual Camphorosma annua (Amaranthaceae) was driven largely by abiotic stress; grazing alone is insufficient for its long-term persistence, while the perennial Artemisia santonicum (Asteraceae) and Tripolium pannonicum (Asteraceae) have higher survival chances as they are able to coexist with generalists. Overall habitat quality can be reliably determined from the analyzed ecological conditions of indicator species. The outcomes of the presented work are relevant for conservation practice and can serve as a quick tool for assessing the current stage of other grassland habitats.

Habitat extremity and conservation management stabilise endangered calcareous fens in a changing world.

Calcareous fens represent an endangered type of peatlands, acting as refugia for stress-tolerant species in the currently changing landscapes. The resurveys across many regions have reported their recent disappearance or deterioration despite both the extreme habitat conditions (carbonate richness, presence of calcareous tufa, nutrient limitation, high water level) and conservation management. To test the stability of their biotic communities in different environmental and management configurations, we repeatedly sampled molluscs (terrestrial and aquatic), vascular plants, and bryophytes at 30 calcareous fens in the Inner Western Carpathians (Slovakia, Poland) after 13-17 years of warm summers and land-use changes. We found a small yet statistically significant effect of sampling period (old versus new survey) on the species composition of all three groups of organisms when the effect of various positions of sites along ecological gradients was controlled for. The compositional changes, interpreted with the help of Ellenberg Indicator Values, suggest an incipient succession towards grasslands and shrublands, driven by decreasing soil moisture and increasing nutrient availability. Although the number of habitat specialists did not change, the number of matrix-derived vascular plant and bryophyte species significantly increased, with six ubiquitous species of productive habitats being significantly more represented currently, while the richness of aquatic molluscs significantly decreased. Fens in which potentially strongly competitive plant species were less stressed because of less intense management and lower habitat extremity were more prone to such succession. There was no single factor that could predict the magnitude of composition changes; instead, tested factors were found to act synergistically. Conservation management was predominantly important for bryophytes, while extreme habitat conditions were predominantly important for terrestrial snails. We suggested a way how nature conservancy authorities can prioritise the management needs by applying an abiotic indicator system, with less environmentally extreme fens requiring more intense conservation management.

The age of island-like habitats impacts habitat specialist species richness.

While the effects of contemporaneous local environment on species richness have been repeatedly documented, much less is known about historical effects, especially over large temporal scales. Using fen sites in the Western Carpathian Mountains with known radiocarbon-dated ages spanning Late Glacial to modern times (16 975-270 cal years before 2008), we have compiled richness data from the same plots for three groups of taxa with contrasting dispersal modes: (1) vascular plants, which have macroscopic propagules possessing variable, but rather low, dispersal abilities; (2) bryophytes, which have microscopic propagules that are readily transported long distances by air; and (3) terrestrial and freshwater mollusks, which have macroscopic individuals with slow active migration rates, but which also often possess high passive dispersal abilities. Using path analysis we tested the relationships between species richness and habitat age, area, isolation, and altitude for these groups. When only matrix-derived taxa were considered, no significant positive relation was noted between species richness and habitat size or age. When only calcareous-fen specialists were considered, however, habitat age was found to significantly affect vascular plant richness and, marginally, also bryophyte richness, whereas mollusk richness was significantly affected by habitat area. These results suggest that in inland insular systems only habitat specialist (i.e., interpatch disperser and/or relict species) richness is influenced by habitat age and/or area, with habitat age becoming more important as species dispersal ability decreases.