Beavers promote calicioid diversity in boreal forest landscapes

Beavers are ecosystem engineers that modify and maintain a range of special habitat types in boreal forests. They also produce large quantities of deadwood that provide substrate for many lignicolous organisms such as calicioid fungi (Ascomycota). We studied how calicioid diversity differed between boreal riparian forests with and without beaver activity. The results show that calicioid diversity were significantly higher at beaver sites compared to the other two forest site types studied. The large quantity and diverse forms of deadwood produced by beavers clearly promotes calicioid diversity in the boreal landscape. The specific lighting and humidity conditions within beaver wetlands could be the reason why they promote the success of certain calicioid species.     

Fine-scale structure and cross-taxon congruence of bird and beetle assemblages in an old-growth boreal forest mosaic

Aim Nestedness occurs when species present in depauperate sites are subsets of those found in species‐rich sites. The degree of congruence of site nestedness among different assemblages can inform commonalities of mechanisms structuring the assemblages. Well‐nested assemblages may still contain idiosyncratic species and sites that notably depart from the typical assemblage pattern. Idiosyncrasy can arise from multiple processes, including interspecific interactions and habitat preferences, which entail different consequences for species co‐occurrences. We investigate the influence of fine‐scale habitat variation on nestedness and idiosyncrasy patterns of beetle and bird assemblages. We examine community‐level and pairwise species co‐occurrence patterns, and highlight the potential influence of interspecific interactions for assemblage structure. Location Côte‐Nord region of Québec, Canada. Methods We sampled occurrences of ground‐dwelling beetles, flying beetles and birds at sites within old‐growth boreal forest. We examined the nestedness and idiosyncrasy of sites and sought relationships to habitat attributes. We analysed non‐random species co‐occurrence patterns at pairwise and community levels, using null model analysis and five ‘association’ indices. Results All three assemblages were significantly nested. There was limited congruence only between birds and flying beetles whose nestedness was related to canopy openness. For ground‐dwelling beetles, nestedness was related to high stand heterogeneity and sapling density, whereas site idiosyncrasy was inversely related to structural heterogeneity. For birds, site idiosyncrasy increased with canopy cover, and most idiosyncratic species were closed‐canopy specialists. In all assemblages, species idiosyncrasy was positively correlated with the frequency of negative pairwise associations. Species co‐occurrence patterns were non‐random, and for flying beetles and birds positive species pairwise associations dominated. Community‐level co‐occurrence summaries may not, however, always reflect these patterns. Main conclusions Nestedness patterns of different assemblages may not correlate, even when sampled at common locations, because of different responses to local habitat attributes. We found idiosyncrasy patterns indicating opposing habitat preferences, consistent with antagonistic interactions among species within assemblages. Analysis of such patterns can thus suggest the mechanisms generating assemblage structures, with implications for biodiversity conservation.     

Stand dynamics,regeneration patterns and long-term continuity in boreal old-growth Picea abies swamp-forests

Abstract. The occurrence of macrofossil charcoal (long axis > 0.5 mm) and Picea abies (Norway spruce) pollen in peat stratigraphies, in combination with size and age data from 2976 P. abies trees were used to analyse ecosystem continuity and stand-structure in ten old-growth swamp-forests in northern Sweden. All stands were dominated by P. abies, a species whose abundance increased westwards in Sweden between 3000 and 2000 yr B.P. In three stands no macrofossil charcoal was found and the maximum age of the peat, determined by ¹⁴C dating, varied from 1800 to 3600 yr B.P. In the other seven stands the number of levels containing charcoal varied from 1 to 23, but only between 1 and 7 levels were found after the appearance of spruce. Here the maximum age of the peat varied from 400 to 7900 yr B.P. The ten stands had an all-sized stand structure and a stand continuity of ca. 300 yr. The shape of the age structure was similar to an inverse J-curve. This indicates a continuous recruitment over time in a self-perpetuating ecosystem. In a short-term perspective (< 300 yr), the swamp-forests are characterized by individual trees continually emerging while others are dying. it is suggested that internal dynamics of continuous small-scale disturbances in combination with local site-specific factors determine the structure of these forests. in a long-term perspective, some of the present spruce swamp-forests within the northern boreal zone have functioned as true fire-free refugia since the establishment of P. abies populations while others have been affected by recurring fires, although not as frequently as forests on surrounding drier sites. The hypothesis that Scandinavian spruce swamp-forests in general have functioned as true longterm fire-free refugia is thus modified by the present results.     

Revegetation following experimental disturbance in a boreal old-growth Picea abies forest

Abstract. We studied revegetation patterns after experimental fine-scale disturbance (e.g. uprooting) in an old-growth Picea abies forest in southeastern Norway. An experimental severity gradient was established by manipulation of the depth of soil disturbance; two types of disturbed areas were used. Species recovery was recorded in the disturbed patches in three successive years after disturbance. The cover of vascular plants and, even more so the cover of bryophytes and lichens, recovered slowly after disturbance. The least severe treatments (removal of vegetation and removal of vegetation and the litter layer) was followed by the fastest recovery. The mean number of vascular plant species was usually higher three years after disturbance than before disturbance, while the opposite was true for bryophytes. Several vascular plant species that were abundant in intact forest floor vegetation (Vaccinium myrtillus, V. vitis-idaea and Deschampsia flexuosa) recovered during a three-year period primarily by resprouting from intact rhizomes and clonal in-growth. Other important recovery mechanisms included germination from a soil-buried propagule bank (e.g. Luzula pilosa, Plagiothecium laetum agg., Pohlia nutans and Polytrichum spp.) and dispersal of propagules into the disturbed patches (e.g. Betula pubescens and Picea abies). Microsite limitation seemed to occur in several species that were abundant in the soil propagule bank (e.g. the ferns Athyrium filix-femina, Gymnocarpium dryopteris and Phegopteris connectilis) but which did not appear in disturbed patches. Disturbance severity influenced revegetation patterns, recorded both as trajectories of vegetation composition in a DCA ordination space and as change in floristic dissimilarity. The length of the successional path (compositional change measured in β-diversity units) increased with increasing disturbance severity, and was also influenced by the area of the disturbed patch and the distance to intact vegetation. The rate of succession depended on the method by which it was measured; decreasing year by year in floristic space, while first decreasing and then increasing in ordination space. The reason for this difference is explained.     

50 years of change in a Swedish boreal old-growth Picea abies forest

Abstract. Changes in the tree layer (> 1.3m) and sapling layer (< 1.3m, including seedlings) of a Swedish boreal old-growth Picea abies (Norway spruce) forest from the 1930s to the 1980s were studied in permanent plots. The plots were established in 1938–1939 and re-analysed in 1983–1988. Regeneration, mortality, turnover rate in the tree layer and amount of decomposing logs as well as the time required for complete decomposition of logs were investigated using the detailed data from the 1930s. Ca. 25 % of the trees present during the first analysis were no longer alive. This mortality was balanced by recruitment from the sapling layer. The rate of mortality suggests a turnover time for the tree layer of ca. 200 yr. The number of spruces in the sapling layer has increased by ca. 85 %, hypothetically in response to an increase in amount of decomposing wood that can serve as nurse logs and stumps. The mean time for total decomposition was calculated as ca. 200 yr. Spruce regeneration on logs does not occur until the log is at least ca. 50 yr old. The survival pattern in the sapling layer suggests a high mortality rate at the seedling stage (≤ 1 yr) and a low mortality rate at the sapling stage. In conclusion, it is suggested that the amount of coarse woody debris available for regeneration, the occurrence of seedlings, and seedling mortality constitute concurrent factors through which climatic fluctuations, in a long-term perspective, direct stand recruitment and density. As a consequence, these boreal forests will be kept in a dynamical equilibrium.     

Ungulate herbivory reduces abundance and fluctuations of herbivorous insects in a boreal old-growth forest

Ecological theory predicts the strongest ecosystem effects of herbivory when dominant and ecologically important species are consumed. Bilberry, Vaccinium myrtillus, is such a key plant species, attractive to many other species in the boreal forests, for example ungulate and invertebrate herbivores. Large herbivores may remove substantial biomass and alter plant quality and therefore affect abundance and populations of invertebrate animals sharing the same food plant. We combined experimental exclusion of ungulates with a browsing intensity gradient to investigate the 15-year effect of ungulate (Cervus elaphus and Ovis aries) browsing on bilberry plant size and on bilberry-feeding herbivorous larvae (Lepidoptera and Symphyta), in a Norwegian old growth boreal forest ecosystem. Bilberry ramets in exclosure plots had nearly nine times higher dry mass and three times higher abundance of invertebrates feeding on them than in ungulate-access plots. Sweep-netting data verified these findings as larval numbers were twice as high in exclosure plots. The pattern in the large herbivore effects on bilberry size and abundance of herbivorous larvae were identical along the browsing gradient. Differences in larval abundance between treatments, as indicated by leaf-chewing, increased during the 15-year study period, and the community fluctuations were larger when ungulate herbivores were excluded. The browsing effect was moderated by plant quality as larval densities were lowest on both heavily-browsed and non-browsed plants, and highest on ramets that had 50–74% of annual shoots browsed. Our study supports previous findings in that bilberry is relatively disturbance tolerant and may recover quickly, but that ungulates may compete with herbivorous larvae for food biomass. Additionally, our results strongly indicates that population insect community peaks and fluctuations are dampened by ungulate consumption. Our findings add to the understanding on how ungulates may structure forest ecosystems directly and indirectly.     

Crustose lichens as indicators of forest continuity in boreal coniferous forests

The crustose lichen flora of fifteen areas in boreal coniferous forests from southern Sweden to Lapland was surveyed. Many of these areas are included in a national programme for monitoring long-term environmental change, the PMK programme. In all 190 species were recorded, 129 of them occurring on bark of conifers and lignum. Some species were found exclusively in forests with a long continuity. By using ordination and hierarchical classifications further species with similar habitat requirements were identified. An Indicator Species Index of Forest Continuity (ISIFC) was designed. Only two of the twenty index species occur on deciduous trees. The ISIFC was highly correlated with forest continuity and also with occurrence of species listed as threatened. Correlations with macroclimatic variables are considered sampling artifacts, and microclimatic factors are suggested to be of primary importance for species occurring in forests with a long continuity.     

Morphology influences water storage in hair lichens from boreal forest canopies

Hair lichens (Alectoria, Bryoria, Usnea) with high surface-area-to-mass ratios rapidly trap moisture. By photography and scanning we examined how internal water storage depended on morphological traits in five species. Specific thallus mass (STM, mg DM cm⁻²) and water holding capacity (WHC, mg H₂O cm⁻²) after shaking and blotting a fully hydrated thallus increased with thallus area. STM was ≈50% higher in Alectoria and Usnea thalli than in Bryoria. WHC was highest in Alectoria while percent water content of freshly blotted thalli was lowest in Usnea. Thallus area overlap ratio (TAO), assessing branch density of the thallus, was highest in the two thinnest Bryoria; lower in the thicker Usnea. Within species, hair lichens increased their water storage by increasing branch density rather than branch diameter. The taxonomically related genera Alectoria and Bryoria shared water storage characteristics, and differed from Usnea. Hair lichens in lower canopies have among the lowest water storage capacity reported in lichens.     

Gap regeneration of shade-intolerant Larix gmelini in old-growth boreal forests of northeastern China

Abstract. Stand mapping and vertical and oblique tree crown projections were used to study gap characteristics and gap effects on the regeneration and stand development of Larix gmelini. The hypothesis is that waves of advance Larix regeneration are recruited into the canopy layer following the creation of canopy openings. In old-growth Larix forests of the northern Da Xingan Ling Mts., at 52 °N, obliquely projected gaps (OPG) begin at a distance of 60 - 80% of the canopy tree height from the southernmost stems bordering the gap and thus the OPGs may extend beyond the northern boundaries of the vertically projected gaps (VPG). Changes in the environment and resource availability in the OPG result in increased Larix sapling survival. Due to a greater incidence of light, 10 - 30 yr old OPG saplings were more abundant than saplings in either a near-oblique projected gap (NOPG) or in the shadow of obliquely projected crowns (SOPC). The survival of saplings more than 30-yr old was highest when they were found in the OPG of one canopy opening and the VPG of another. This means that, following recruitment into an OPG, saplings then require the space found in a VPG to permit growth into the canopy. Thus, various-sized gaps contribute to the survival of different aged saplings by increasing the complexity of stand structure. Although individuals may regenerate in an OPG, successful recruitment into the canopy requires the available growing space of a VPG. This research suggests that shade intolerant Larix gmelini can maintain its canopy dominance without fire via gap regeneration.     

Defensive compound concentration in boreal lichens in response to simulated nitrogen deposition

Nitrogen (N) deposition is expected to increase in northwestern Europe the next 50–100 years. The effects of higher N availability on lichens will presumably depend on their capacity to acquire carbon (C), that is, of the timing and duration of the wet and active state. If lichens respond like plants, their C and N status may affect their concentration of carbon-based secondary compounds (CBSCs), and thus their defence against herbivores, detrimental radiation, pathogens and parasites. In the present study we have manipulated N availability and timing and duration of the metabolically active state by spraying lichen transplants in an old spruce forest with rainwater or rainwater with added N corresponding to 50 kg N ha⁻¹ yr⁻¹. The spraying was applied either at night, in the morning or at noon to also investigate the effect of timing and duration of the active state. Concentrations of N, chlorophyll a (Chl a ) and CBSCs were measured before and after one summer's spraying of 10 thalli in each of four different lichen species; Alectoria sarmentosa, Lobaria scrobiculata, Platismatia glauca , and Xanthoria aureola . The added N was readily taken up by all the lichen species. A. sarmentosa, P. glauca , and X. aureola increased their Chl a concentration in response to increased N, while L. scrobiculata increased Chl a in response to increased active time. None of the studied species reduced their concentration of secondary compounds during the experimental period, but in P. glauca the concentration of all compounds were significantly lower in N-treated thalli compared with those that got only rainwater. The results are consistent with a high degree of constitutive defence in three of four species, and we conclude that all the investigated lichens seem to have rather robust chemical defence systems despite considerable manipulation of the environmental conditions.     

Beetle diversity in a matrix of old-growth boreal forest: influence of habitat heterogeneity at multiple scales

The relative contribution of compositional and structural heterogeneity on biodiversity is currently ambiguous because field studies generally integrate these two sources of habitat heterogeneity into a single index. We established the relationship between species richness of ground-dwelling and flying beetles and compositional and structural attributes of forest heterogeneity. The relationship was evaluated at two spatial scales: the scale of forest stand, corresponding to an 11.3  m radius, and the scale of landscape, corresponding to either a 400 or 800  m radius. Seventy stands were sampled in the matrix of old-growth boreal forest of the North Shore region of Québec, Canada, during the summers of 2004 and 2005. A total of 133 ground-dwelling beetle species (range: 4–42 species per site) were captured in the pitfall traps and 251 flying species (range 16–58 species per site) in flight-interception traps. We found that the most relevant type of heterogeneity to explain variations in species richness and the significance of landscape scale information varied between groups of beetles. Compositional heterogeneity (i.e. the number of species of forest trees and shrubs) at the stand scale best predicted species richness in ground-dwelling beetles. On the other hand, it was the combined influence of structural and compositional habitat heterogeneity at stand and landscape scales that best explained richness patterns in flying beetles. Our study outlines the significance of considering multiple types and spatial scales of habitat heterogeneity when describing patterns of species richness.     

High cyanobiont selectivity of epiphytic lichens in old growth boreal forest of Finland

Here, cyanobiont selectivity of epiphytic lichen species was examined in an old growth forest area in Finland. Samples of the eight lichen species were collected from the same aspen (Populus tremula) and adjacent aspens in the same stand. The cyanobionts of these samples were compared with free and symbiotic Nostoc obtained from other habitats and geographic regions. Our results, based on the phylogenetic analysis of a partial small subunit of the ribosomal DNA (16S rDNA) and the rbcLX gene complex did not show any correlation with the geographic origin of the samples at any spatial scale. Instead, there was a correlation between the cyanobionts and the alleged taxonomy of their mycobionts. The results indicate that the lichen species examined are highly selective towards their cyanobiont partners. Only Lobaria pulmonaria proved to be more flexible, being able to associate with a wide range of Nostoc. A same Nostoc strain was found to form associations with taxonomically unrelated lichens indicating that the cyanobiont-mycobiont associations as a whole were not highly specific in the examined species.     

Sporostigma,a new calicioid genus in Arthoniales

Abstract:The new genus Sporostigma is introduced for the remarkable species Arthonia melaspora. The genus is characterized by mazaediate ascomata and dark brown ascospores. Relationships with other genera are discussed.     

Nonlinear responses of ecosystem carbon fluxes to nitrogen deposition in an old-growth boreal forest

Nitrogen (N) deposition is known to increase carbon (C) sequestration in N-limited boreal forests. However, the long-term effects of N deposition on ecosystem carbon fluxes have been rarely investigated in old-growth boreal forests. Here we show that decade-long experimental N additions significantly stimulated net primary production (NPP) but the effect decreased with increasing N loads. The effect on soil heterotrophic respiration (Rh) shifted from a stimulation at low-level N additions to an inhibition at higher levels of N additions. Consequently, low-level N additions resulted in a neutral effect on net ecosystem productivity (NEP), due to a comparable stimulating effect on NPP and Rh, while NEP was increased by high-level N additions. Moreover, we found nonlinear temporal responses of NPP, Rh and NEP to low-level N additions. Our findings imply that actual N deposition in boreal forests likely exerts a minor contribution to their soil C storage.     

High occurrence of red-listed bryophytes and lichens in mature managed forests in boreal Sweden

Successful biodiversity conservation needs to be based on reliable data regarding the distribution and abundance of species. Red Lists summarize the status of rare and declining species. Here we present the first detailed survey (10 m × 10 m resolution) of nationally red-listed bryophytes and lichens at stand level in boreal forests. 30 managed forest production stands in boreal Sweden, soon to be harvested, were surveyed in detail regarding red-listed bryophytes and lichens. On the 180 ha totally investigated, 33 red-listed species were found, representing 35% of all forest red-listed bryophytes and lichens so far recorded from this part of Sweden. The mean number of red-listed species per stand was 10 and varied between 5 and 16. Excluding three species which were so common that, given the time-limit, it was not possible to record all of their occurrences, there were on average 6 number of observations ha⁻¹. The presence and density exceed those earlier found in ‘key habitats’, defined as hot-spots for red-listed species. Almost all observations were made on trees, with 48% on living trees and 52% on dead. Altitude, area and stem volume explained 67% of the red-list species density. The results highlight the importance of unprotected areas for rare and declining species and point to a hitherto unacknowledged high biodiversity value of mature managed boreal forest stands, shaped by a less intense forestry than today's. It also raises questions regarding the use of red-listed species to identify areas of importance to biodiversity. Objective and systematic surveys of red-listed species in the forest landscape, including younger stands, are urgent.Der erfolgreiche Schutz der Biodiversität muss auf verlässlichen Daten über die Verbreitung und Abundanz der Arten basieren. Rote Listen fassen den Status seltener und abnehmender Arten zusammen. Hier präsentieren wir die erste detaillierte Erfassung (10 m × 10 m Auflösung) von Bryophyten und Flechten der nationalen Rote Liste auf Bestandsniveau in borealen Wäldern. Im borealen Schweden wurden in 30 bewirtschafteten Produktionswälder, die bald geerntet werden sollen, die Bryophyten und Flechten der Roten Liste im Detail erfasst. Auf den insgesamt untersuchten 180 ha wurden 33 Arten der Roten Liste gefunden, die 35% aller Wald-Bryophyten und-Flechten der Roten Liste repräsentieren, die bisher in diesem Teil Schwedens erfasst wurden. Die mittlere Anzahl derRote-Liste-Arten pro Bestand war 10 und variierte zwischen 5 und 16. Unter Ausschluss von drei Arten, die so gewöhnlich waren, dass es bei dem gegebenen Zeitlimit nicht möglich war alle ihre Vorkommen zu erfassen, gab es im Durchschnitt 6 Beobachtungen ha⁻¹. Vorkommen und Dichten übertrafen die früher in “Schlüsselhabitaten” gefundenen, die als ‘hot spots’ bezeichnet werden. Fast alle Beobachtungen wurden auf Bäumen gemacht, mit 48% auf lebenden und 52% auf toten Bäumen. Die Höhe, das Gebiet und das Stammvolumen erklärten 67% der Dichte der Rote-Liste-Arten. Die Ergebnisse heben die Wichtigkeit nicht-geschützter Gebiete für seltene und abnehmende Arten hervor und weisen auf einen bisher nicht wahrgenommenen hohen Biodiversitätswert von alten bewirtschafteten borealen Waldbeständen hin, die von einer weniger intensiven Forstwirtschaft als der heutigen geformt wurden. Es stellen sich ebenso Fragen, die die Verwendung der Rote-Liste-Arten betreffen, um für die Biodiversität wichtige Gebiete zu identifizieren. Objektive und systematische Erfassungen der Rote-Liste-Arten in Waldlandschaften inklusive der jüngeren Bestände sind dringend.     

Spatial patterns of photobiont diversity in some Nostoc-containing lichens

Patterns of photobiont diversity were examined in some Nostoc -containing lichens using the nucleotide sequence of the cyanobacterial tRNA^Leu (UAA) intron. Lichen specimens collected in northwestern USA were analysed and the sequence data were compared with tRNA^Leu(UAA) intron sequences previously obtained from lichens in northern Europe. Generally, it is the species identity of a lichen rather than the geographical origin of the specimen that determines the identity of the cyanobiont. Identical intron sequences were found in Peltigera membranacea specimens collected in Oregon (USA) and in Sweden, and very similar sequences were also found in Nephroma resupinatum thalli collected in Oregon and Finland. Furthermore, in mixed assemblages where two Peltigera species grew in physical contact with each other, the different lichen species housed different photobiont strains. There is however not a one-to-one relation between mycobiont and photobiont as some intron sequences were found in more than one lichen species, and different intron sequences were found in different samples of some lichen taxa. Peltigera venosa exhibited a higher level of photobiont diversity than any other lichen species studied, and several intron sequences could for the first time be obtained from a single thallus. It is not clear whether this is evidence of lower cyanobiont specificity, or reflects an ability to exhibit different degrees of lichenization with different Nostoc strains. In one specimen of P. venosa , which contained bipartite cyanosymbiodemes and tripartite, cephalodiate thalli, both thallus types contained the same intron sequence.     

Chaenothecopsis quintralis, a new species of calicioid fungus

Chaenothecopsis quintralis from southwestern Argentina is described and illustrated as a new species in the family Mycocaliciaceae. It has been found in three localities in the Andean Patagonian temperate forests, growing strictly on dung of an endemic marsupial Dromiciops gliroides. The new species is distinguished by the hemispherical, black capitulum of ascoma, the presence of asci with croziers, one-celled brown ascospores, and its fimicolous habitat. Analysis of partial nuclear large subunit rDNA (LSU) sequences showed that this taxon is within Mycocaliciales.     

Above- and belowground patterns in a subalpine grassland-shrub mosaic

Many mountain pastures consist of a mosaic of grassland and shrub communities. Ongoing changes in mountain agriculture have affected the balance between the two elements of the mosaic. In order to understand the consequences of these changes for ecosystem functioning, we studied patterns in vegetation, root structure and soil properties along transects of varying grassland-to-shrub proportions. Our hypothesis was that differences in the vegetation aboveground are accompanied by differences belowground, related to soil properties and depth. The research was conducted at a subalpine site in the Trentino region (South-eastern Alps), consisting of Nardus stricta grasslands alternating with shrub patches of Rhododendron ferrugineum. Our investigation showed that the composition of vegetation was mainly governed by R. ferrugineum cover and less by soil properties. Plant species richness peaked at low to intermediate degrees of shrub cover and composition between transects became more similar with increasing shrub cover. Where R. ferrugineum cover was higher, Hemicryptophytes caespitosae were replaced by Nano-phanerophytes with consequences for belowground structures. At increasing shrub cover, root length density decreased, especially in the top soil, while root weight density remained stable and C content increased insignificantly. We discuss that theses structural changes along the gradient of R. ferrugineum cover affect a number of ecosystem services. The presented evidence suggests that maintaining grasslands with a low cover of R. ferrugineum balances a number of services, namely plant species diversity, carbon stabilization in soil and the prevention of soil erosion.