Changes in species diversity often result from species losses and gains. The dynamic nature of beta diversity (spatial variation in species composition) that derives from such temporal species ...turnover, however, has received relatively little attention. Here, we disentangled extinction and colonization components of beta diversity by using the sets of species that went locally extinct and that newly colonized the study sites. We applied this concept of extinction and colonization beta diversity to ground vegetation communities that have been repeatedly surveyed in forests where fire and harvesting were experimentally applied. We first found that fire and harvesting caused no effect on beta diversity 2 yr after the treatments. From this result, we might conclude that they did not alter the ways in which species assemble across space. However, when we analyzed the extinction and colonization beta diversity between pretreatment and 2 yr after the treatments, both measures were found to be significantly lower in burned sites compared to unburned sites (i.e., the groups of excluded and newly colonized species both showed low beta diversity in the burned sites). These results indicate that the fire excluded similar subsets of species across space, making communities become more heterogeneous, but at the same time induced spatially uniform colonization of new species, causing communities to homogenize. Consequently, the effects of these two processes canceled each other out. The relative importance of extinction and colonization components per se also changed temporally after the treatments. Fire and harvesting showed synergetic negative impacts on extinction beta diversity between pre-treatment and 10 yr after the treatments. Overall, analyses using extinction and colonization beta diversity allowed us to detect nonrandom disassembly and reassembly dynamics in ground vegetation communities. Our results suggest that common practices of analyzing beta diversity at one point in time can mask significant variation driven by disturbance. Acknowledging the extinction–colonization dynamics behind beta diversity is essential for understanding the spatiotemporal organization of biodiversity.
Background and aims
Forest management towards increased carbon (C) sequestration has repeatedly been suggested as a “natural climate solution”. We evaluated the potential of altered management to ...increase C sequestration in boreal
Pinus sylvestris
forest plantations.
Methods
At 29 forest sites, distributed along a 1300 km latitudinal gradient in Sweden, we studied interactive effects of fertilization and thinning on accumulation of C in standing biomass and the organic horizon over a 40 year period.
Results
Abstention from thinning increased the total C stock by 50% on average. The increase was significant (14% on average) even when C in the removed timber was included in the total ecosystem C pool. Fertilization of thinned stands increased stocks similarly regardless of including (11%) or excluding (12%) removed biomass, and fertilization combined with abstention from thinning had a synergistic effect on C stocks that generated an increase of 79% (35% when removed timber was included in the C stock). A positive effect of fertilization on C stocks was observed along the entire gradient but was greater in relative terms at high latitudes. Fertilization also reduced soil respiration rates.
Conclusion
Taken together, our results suggest that changed forest management practices have major potential to increase the C sink of boreal forests. Although promising, these benefits should be evaluated against the undesired effects that such management can have on economic revenue, timber quality, biodiversity and delivery of other ecosystem services.
In Sweden, the majority of forest area has been altered by industrial forestry over the decades. Almost 30 years ago, a shift towards biodiversity-oriented forest management practices occurred. Here ...we took advantage of long-term data collected by the Swedish National Forest Inventory to track developmental changes in forest structural components over this time. We assessed changes in structural components that play an important role in biodiversity (dead wood, large living trees, tree species composition, and understory vegetation) in four forest types with descending tiers of biodiversity protection: protected areas, woodland key habitats, low-productivity forests and production forests. Overall, we found a positive trend in the volumes of dead wood and large living trees, as well as in tree species diversity, while there was a general decline in understory vegetation coverage. Most observed changes were consistent with the intended outcomes of the current forest policy, adapted in the early 1990s. The implementation of retention forestry is likely driving some of the observed changes in forest structural components in the south. In contrast, we observed no changes in any of the focal structural components in the north, which could be attributed to the ongoing clear-cutting of forests previously managed less intensively. Dead wood and large living trees increased not only in managed, but also in unmanaged forests, likely reflecting historical management. The increased tree species diversity can be explained through current forest management practices that encourages maintenance of additional tree species. Decreasing understory vegetation coverage in both dense managed and unmanaged forests suggests that factors other than forestry contribute to the ongoing changes in understory vegetation in Swedish forests. Overall, the observed increase in structural components has not yet been reflected in documented improvements for red-listed forest species, which may be due to delays in species responses to small improvements, as well as a lack of detailed monitoring. Similarly, the increased availability of forest structural components might still be insufficient to meet the specific habitat requirements of red-listed species.
•Biodiversity important structural components were assessed in Swedish forests.•Dead wood and large living trees increased over time, but not in the north.•Tree species diversity increased due to more deciduous tree species.•Understory vegetation coverage declined, probably due to denser forests.•The observed trends are in line with the increased conservation concern.
•Airborne laser scanning data (ALS) helped assess forest conservation value.•ALS-detected deadwood can indicate forest conservation value.•Trees considered particularly important for conserving ...biodiversity can be detected.•Wall-to-wall maps of forest conservation value can be generated using ALS.
In sustainable forest resource management, establishing forest conservation areas is important to maintain forest biodiversity. However, assessing the conservation value of forests is challenging because the target areas are often both large and remote. We explored using dense airborne laser scanning (ALS) data to estimate conservation values. Field data were collected at sites in Sweden for standing deadwood (S trees), laying deadwood (L trees), and trees considered particularly important for conserving biodiversity (N trees), and forest conservation values were determined using a commonly employed method in the field. A template matching method was then used to detect L, S and N trees from ALS data. L trees were identified from linear features in the point cloud above the ground, with a 52 % detection error rate. S and N trees were identified from unusually small or large crown diameters, with 71 % and 83 % error rates, respectively. We also tested the relationships between the three types of indicator tree, their summed values and the field inventory-assessed conservation values. Regressions between the assessed conservation values and ALS indicators were most robust comparing the three test sites when using the summed number of L, S, and N trees. A wall-to-wall map covering a 3 km × 4 km area was generated using Kernel density estimation of the summed number of ALS-derived indicators, to represent relative conservation values. The map was validated using 10 1-ha plots, and yielded an R2 value of 0.6 for predicted conservation values at the plot level. We conclude that ALS data can be used to map forest conservation values and inform decisions about which forests should be used for timber production and which should be set aside as conservation areas. The maps could also be used as a data source for habitat analysis.
Stratigraphic records from peatlands suggest that the shift from a rich fen (calcareous fen) to an ombrotrophic bog can occur rapidly. This shift constitutes a switch from a species-rich ecosystem to ...a species-poor one with greater carbon storage. In this process, the invasion and expansion of acidifying bog species of
Sphagnum
(peat mosses) play a key role. To test under what conditions an acidifying bog species could invade a rich fen, we conducted three experiments, contrasting the bog species
S. fucsum
with the rich-fen species
S. warnstorfii
and
S. teres
. We first tested the effect of calcareous water by growing the three species at different constant height above the water table (HWT; 2, 7, and 14 cm) in a rich-fen pool and measured maximum photosynthetic rate and production and difference in length growth as an indicator of competition. In none of the species was the photosynthetic capacity negatively affected when placed at low HWT, but
S. fuscum
was a weaker competitor at low HWT. In our second experiment we transplanted the three species into microhabitats with different and naturally varying HWT in a rich fen. Here,
S. fuscum
nearly ceased to photosynthesize when transplanted to low HWT (brown moss carpet), while it performed similarly to the two rich-fen species at the intermediate level (
S. warnstorfii
hummock level). In contrast to
S. fuscum
, the rich-fen sphagna performed equally well in both habitats. The brown moss carpet was seasonally flooded, and in our third experiment we found that
S. fuscum
, but not
S. teres
, was severely damaged when submerged in rich-fen water. Our results suggest two thresholds in HWT affecting the ecosystem switch: one level that reduces the risk of submergence and a higher one that makes bog sphagna competitive against the rich-fen species.
Herbivorous insects can influence grassland ecosystem functions in several ways, notably by altering primary production and nutrient turnover. Interactions between above- and belowground herbivory ...could affect these functions; an effect that might be modified by nitrogen (N) addition, an important global change driver. To explore this, we added above- (grasshoppers) and belowground (wireworms) insect herbivores and N into enclosed, equally composed, grassland plant communities in a fully factorial field experiment. N addition substantially altered the impact of above- and belowground herbivory on ecosystem functioning. Herbivory and N interacted such that biomass was reduced under above ground herbivory and high N input, while plant biomass remained stable under simultaneous above- and belowground herbivory. Aboveground herbivory lowered nutrient turnover rate in the soil, while belowground herbivory mitigated the effect of aboveground herbivory. Soil decomposition potential and N mineralization rate were faster under belowground herbivory at ambient N, but at elevated N this effect was only observed when aboveground herbivores were also present. We found that N addition does not only influence productivity directly (repeatedly shown by others), but also appears to influence productivity by herbivory mediated effects on nutrient dynamics, which highlights the importance of a better understanding of complex biotic interactions.
Insect herbivores are important drivers of ecosystem processes in grasslands, and can mediate the grassland's response to environmental change. For example, recent evidence shows that above‐ and ...belowground herbivory, individually and in combination, can modify how a plant community responds to nitrogen (N) eutrophication, an important driver of global change. However, knowledge about how such effects extend to the associated soil food web is lacking. In a mesocosm experiment, we investigated how communities of soil nematodes – an abundant and functionally important group of soil organisms – responded to above‐ and belowground insect herbivory at contrasting N levels. We found that the strongest influence of above‐ and belowground herbivory on the nematode community appeared at elevated N. The abundance of root‐feeding nematodes increased when either above‐ or belowground insect herbivores were present at elevated N, but when applied together the two herbivore types cancelled out one another's effect. Additionally, at elevated N aboveground herbivory increased the abundance of fungal‐feeders relative to bacterial‐feeders, which indicates changes in decomposition pathways induced by N and herbivory. Belowground herbivory increased the abundance of omnivorous nematodes. The shifts in both the herbivorous and detrital parts of the soil food web demonstrate that above‐ and belowground herbivory does not only mediate the response of the plant community to N eutrophication, but in extension also the soil food web sustained by the plant community. We conclude that feedbacks between effects of above‐ and belowground herbivory mediate the response of the grassland ecosystem to N eutrophication.
Display omitted
•Habitat heterogeneity and resource amount and quality were tested as indicators.•Habitat heterogeneity predicts the richness of species of conservation concern.•Dead wood and tree ...age predict the richness of red-listed species.•Thresholds for habitat heterogeneity score and deadwood were calculated.•Habitat heterogeneity score is useful tool in practical forestry and conservation.
Reliable assessment measures are crucial for tracking changes in biodiversity and for evaluating the state of biodiversity. Two of the main drivers of biodiversity are habitat heterogeneity and resource amount. These drivers are used as proxies of biodiversity but assessing both is costly, limiting their practical use. To test which of the drivers best predicts the number and abundance of sessile species of conservation concern (including macrofungi, lichens, bryophytes, and vascular plants), we assessed forest stand heterogeneity using a method developed in Sweden (‘Habitat Heterogeneity Score HHS’), and quantified the resource amount and quality of ecologically important structural variables (deadwood volume, basal area of living trees, proportion of broadleaved trees, and the age of the oldest tree in the stand). We conducted the assessments in 77 boreal conifer-dominated forest stands in two regions of Sweden. Despite some group-specific organism differences, HHS was the best predictor of both number and abundance of all species of conservation concern, regardless of the region. Further, HHS was the best predictor of red-listed species number and abundance in the southern region, while a model including the volume of deadwood and the age of the oldest tree performed best in the northern region. Deadwood (CWD) volume was the single best resource amount predictor of the number and abundance of species of conservation concern, emphasizing the critical role that dead trees have for biodiversity. In addition, we calculated threshold values for deadwood volume and HHS depicting the level above which the number of red-listed species is significantly higher, and found this value to be higher in the southern region (22.4 m3 ha−1 deadwood and a HSS value of 17) than in the north (20.0 m3 ha−1 and 16). These values can be used as guidance when identifying coniferous forests with high enough qualities to support red-listed species. To conclude, the method of assessing habitat heterogeneity presented in this study is a practical and reliable way to identify forests of high biological diversity, and can therefore be part of the toolbox for sustainable forestry in boreal forests.
Context
Biodiversity is highly affected by industrial forestry, which leads to the loss and fragmentation of natural habitats. To date, most conservation studies have evaluated associations among a ...single species group, forest type, or spatial scale.
Objective
The objective was to evaluate the richness of multiple species groups across various forest types and characteristics at multiple scales.
Methods
We used the occurrence data for 277 species of conservation interest from 455 stands of high conservation value, including four species groups and four forest types.
Results
Local, landscape, and regional forest characteristics influenced biodiversity in a non-uniform pattern among species groups and forest types. For example, an increased local spruce basal area in spruce forests was associated with higher vascular plant and bryophyte richness values, whereas macrofungi and lichen richness were positively correlated with deadwood availability, but negatively correlated with the spruce volume in the landscape. Furthermore, landscapes with twice as much mature forest as the average, had more than 50% higher richness values for vascular plants, macrofungi, and lichens.
Conclusion
Among sessile species groups in northern forests, a uniform conservation strategy across forest types and scales is suboptimal. A multi-faceted strategy that acknowledges differences among species groups and forest types with tailored measures to promote richness is likely to be more successful. Nevertheless, the single most common measure associated with high richness across the species groups and forest types was mature forest in the landscape, which suggests that increasing old forests in the landscape is a beneficial conservation strategy.
With an increasing demand for forest-based products, there is a growing interest in introducing fast-growing non-native tree species in forest management. Such introductions often have unknown ...consequences for native forest biodiversity. In this study, we examine epiphytic lichen species richness and species composition on the trunks of non-native Pinus contorta and compare these to the native Pinus sylvestris and Picea abies in managed boreal forests in northern Sweden across a chronosequence of age classes. Overall, we recorded a total of 66,209 lichen occurrences belonging to 57 species in the 96 studied forest stands. We found no difference in species richness of lichens between stands of P. contorta and P. sylvestris, but stands of P. abies had higher total species richness. However, species richness of lichens in stands of P. abies decreased with increasing stand age, while no such age effect was detected for P. contorta and P. sylvestris. Lichen species composition progressively diverged with increasing stand age, and in 30-year-old stands all three tree species showed species-specific assemblages. Epiphytic lichen assemblages in stands of 30-year-old P. contorta were influenced by greater basal area, canopy closure, and average diameter at breast height, P. abies stands by higher branch density and canopy closure, and stands of P. sylvestris by greater bark crevice depth. Differences in lichen species richness and composition were mainly explained by canopy closure and habitat availability, and the greater canopy closure in mature P. abies stands promoted the colonization and growth of calicioid lichen species. Our results indicate that the non-native P. contorta have similar species richness as the native P. sylvestris. The main difference in lichen species richness and composition is between P. abies and Pinus spp. in managed forests of boreal Sweden.