Mixed effects of a national protected area network on terrestrial and freshwater biodiversity.

Protected areas are considered fundamental to counter biodiversity loss. However, evidence for their effectiveness in averting local extinctions remains scarce and taxonomically biased. We employ a robust counterfactual multi-taxon approach to compare occupancy patterns of 638 species, including birds (150), mammals (23), plants (39) and phytoplankton (426) between protected and unprotected sites across four decades in Finland. We find mixed impacts of protected areas, with only a small proportion of species explicitly benefiting from protection-mainly through slower rates of decline inside protected areas. The benefits of protection are enhanced for larger protected areas and are traceable to when the sites were protected, but are mostly unrelated to species conservation status or traits (size, climatic niche and threat status). Our results suggest that the current protected area network can partly contribute to slow down declines in occupancy rates, but alone will not suffice to halt the biodiversity crisis. Efforts aimed at improving coverage, connectivity and management will be key to enhance the effectiveness of protected areas towards bending the curve of biodiversity loss.

Forest mosses sensitively indicate nitrogen deposition in boreal background areas.

Mosses take up nitrogen (N) mainly from precipitation through their surfaces, which makes them competent bioindicators of N deposition. We found positive relationships between the total N concentration (mossN%) of common terrestrial moss species (feather mosses Pleurozium schreberi and Hylocomium splendens, and a group of Dicranum species) and different forms of N deposition in 11-16 coniferous forests with low N deposition load in Finland. The mosses were collected either inside (Dicranum group) or both inside and outside (feather mosses) the forests. Deposition was monitored in situ as bulk deposition (BD) and stand throughfall (TF) and detected for ammonium (NH4+-N), nitrate (NO3--N), dissolved organic N (DON), and total N (Ntot, kg ha-1yr-1). Ntot deposition was lower in TF than BD indicating that tree canopies absorbed N from deposition in N limited boreal stands. However, mossN% was higher inside than outside the forests. In regression equations, inorganic N in BD predicted best the mossN% in openings, while DON in TF explained most variation of mossN% in forests. An asymptotic form of mossN% vs. TF Ntot curves in forests and free NH4+-N accumulation in tissues in the southern plots suggested mosses were near the N saturation state already at the Ntot deposition level of 3-5 kg ha-1yr-1. N leachate from ground litterfall apparently also contributed the N supply of mosses. Our study yielded new information on the sensitivity of boreal mosses to low N deposition and their response to different N forms in canopy TF entering moss layer. The equations predicting the Ntot deposition with mossN% showed a good fit both in forest sites and openings, especially in case of P. schreberi. However, the open site mossN% is a preferable predictor of N deposition in monitoring studies to minimize the effect of tree canopies and N leachate from litterfall on the estimates.

N2 fixation associated with the bryophyte layer is suppressed by low levels of nitrogen deposition in boreal forests.

Biological fixation of atmospheric nitrogen (N2) by bryophyte-associated cyanobacteria is an important source of plant-available N in the boreal biome. Information on the factors that drive biological N2 fixation (BNF) rates is needed in order to understand the N dynamics of forests under a changing climate. We assessed the potential of several cryptogam species (the feather mosses Hylocomium splendens and Pleurozium schreberi, a group of Dicranum bryophytes, two liverworts, and Cladina lichens) to serve as associates of cyanobacteria or other N2-fixing bacteria (diazotrophs) using acetylene reduction assay (ARA). We tested the hypotheses that the legacy of chronic atmospheric N deposition reduces BNF in the three bryophyte species, sampled from 12 coniferous forests located at latitudes 60-68° N in Finland. In addition, we tested the effect of moisture and temperature on BNF. All species studied showed a BNF signal in the north, with the highest rates in feather mosses. In moss samples taken along the north-south gradient with an increasing N bulk deposition from 0.8 to 4.4 kg ha-1 year-1, we found a clear decrease in BNF in both feather mosses and Dicranum group. BNF turned off at N deposition of 3-4 kg ha-1 year-1. Inorganic N (NH4-N + NO3-N) best predicted the BNF rate among regression models with different forms of N deposition as explanatory variables. However, in southern spruce stands, tree canopies modified the N in throughfall so that dissolved organic N (DON) leached from canopies compensated for inorganic N retained therein. Here, both DON and inorganic N negatively affected BNF in H. splendens. In laboratory experiments, BNF increased with increasing temperature and moisture. Our results suggest that even relatively low N deposition suppresses BNF in bryophyte-associated diazotrophs. Further, BNF could increase in northern low-deposition areas, especially if climate warming leads to moister conditions, as predicted.

Forest floor vegetation response to nitrogen deposition in Europe.

Chronic nitrogen (N) deposition is a threat to biodiversity that results from the eutrophication of ecosystems. We studied long-term monitoring data from 28 forest sites with a total of 1,335 permanent forest floor vegetation plots from northern Fennoscandia to southern Italy to analyse temporal trends in vascular plant species cover and diversity. We found that the cover of plant species which prefer nutrient-poor soils (oligotrophic species) decreased the more the measured N deposition exceeded the empirical critical load (CL) for eutrophication effects (P = 0.002). Although species preferring nutrient-rich sites (eutrophic species) did not experience a significantly increase in cover (P = 0.440), in comparison to oligotrophic species they had a marginally higher proportion among new occurring species (P = 0.091). The observed gradual replacement of oligotrophic species by eutrophic species as a response to N deposition seems to be a general pattern, as it was consistent on the European scale. Contrary to species cover changes, neither the decrease in species richness nor of homogeneity correlated with nitrogen CL exceedance (ExCLemp N). We assume that the lack of diversity changes resulted from the restricted time period of our observations. Although existing habitat-specific empirical CL still hold some uncertainty, we exemplify that they are useful indicators for the sensitivity of forest floor vegetation to N deposition.

Differences in the growth response of three bryophyte species to nitrogen.

The effect of nitrogen on biomass production, shoot elongation and relative density of the mosses Pleurozium schreberi, Hylocomium splendens and Dicranum polysetum was studied in a chamber experiment. Monocultures were exposed to 10 N levels ranging from 0.02 to 7.35 g N m(-2) during a 90-day period. All the growth responses were unimodal, but the species showed differences in the shape parameters of the curves. Hylocomium and Pleurozium achieved optimum biomass production at a lower N level than Dicranum. Pleurozium had the highest biomass production per tissue N concentration. Tolerance to N was the widest in Dicranum, whereas Hylocomium had the narrowest tolerance. Dicranum retained N less efficiently from precipitation than the other two species, which explained its deviating response. All species translocated some N from parent to new shoots. The results emphasize that the individual responses of bryophytes to N should be known when species are used as bioindicators.

Element accumulation in boreal bryophytes, lichens and vascular plants exposed to heavy metal and sulfur deposition in Finland.

Macronutrient (N, P, K, Mg, S, Ca), heavy metal (Fe, Zn, Mn, Cu, Ni, Cd, Pb) and Al concentrations in understorey bryophytes, lichens and vascular plant species growing in Scots pine forests at four distances from the Harjavalta Cu-Ni smelter (0.5, 2, 4 and 8 km) were compared to those at two background sites in Finland. The aim was to study the relationship between element accumulation and the distribution of the species along a pollution gradient. Elevated sulfur, nitrogen and heavy metal concentrations were found in all species groups near the pollution source. Macronutrient concentrations tended to decrease in the order: vascular plants>bryophytes>lichens, when all the species groups grew on the same plot. Heavy metal concentrations (except Mn) were the highest in bryophytes, followed by lichens, and were the lowest in vascular plants. In general, vascular plants, being capable of restricting the uptake of toxic elements, grew closer to the smelter than lichens, while bryophytes began to increase in the understorey vegetation at further distances from the smelter. A pioneer moss (Pohlia nutans) was an exception, because it accumulated considerably higher amounts of Cu and Ni than the other species and still survived close to the smelter. The abundance of most of the species decreased with increasing Cu and Ni concentrations in their tissues. Cetraria islandica, instead, showed a positive relationship between the abundance and Cu, Ni and S concentrations of the thallus. It is probable that, in addition to heavy metals, sporadically high SO(2) emissions have also affected the distribution of the plant species.

Copper resistance of the evergreen dwarf shrub Arctostaphylos uva-ursi: an experimental exposure.

The copper (Cu) resistance of Arctostaphylos uva-ursi was tested in a pot experiment (lasting 8 weeks) using rooted cuttings originating from an area near the Harjavalta Cu-Ni smelter, SW Finland. The fine roots were moderately infected by arbutoid mycorrhizae. The plants were exposed to five Cu levels (1, 10, 22, 46 and 100 mg l(-1)) given repeatedly together with a nutrient solution. The critical Cu concentration in the nutrient solution inhibiting the growth of A. uva-ursi was below 10 mg l(-1) Cu (EC(50) value for biomass production 3.3 mg l(-1) Cu). This concentration was clearly lower than the value we have found earlier for other dwarf shrubs under similar experimental conditions. Most of the Cu given accumulated in the roots and old stems. The results suggest that A. uva-ursi cuttings were relatively sensitive to Cu despite the ability of the adult clones to grow in Cu-contaminated soil. The adult clones extend their roots into the less toxic deeper soil layers, which may facilitate the avoidance of heavy metals.