Growth of reindeer lichens and effects of reindeer grazing on ground cover vegetation in a Scots pine forest and a subarctic heathland in Finnish Lapland

Reindeer lichens are an important component of northern ecosystems. The aim of this study was to measure the growth rate of terricolous lichens as it is a key parameter involved in productivity of these ecosystems and an important part of lichen tolerance to reindeer grazing. Furthermore, the natural succession and the long-term effects of reindeer grazing on lichen community characteristics in two contrasting habitats were investigated as well as the interactions between lichen cover and mosses and vascular plants. Biomass and coverage measurements were conducted in a lichen woodland and in a subarctic heath with grazed and ungrazed areas in northern Finland. Measurements spanning over 13 yr of undisturbed development show that the growth rate of Cladina stellaris can be as high as >0.17 g g⁻¹ produced annually, although in average growth rates were much lower. During the succession of ground vegetation, C. stellaris , C. rangiferina , C. mitis and Cetraria nivalis increased in biomass in fenced areas and were reduced most in biomass by reindeer in unprotected areas. Reindeer grazing and trampling seem to change the vegetation towards a type that is dominated by small dwarf shrubs, bare soil and minute-cup lichens ( Cladonia spp.). Removing the lichen layer by reindeer may reduce natural regeneration of pine trees as implied by increasing numbers of pine seedlings with increasing lichen cover.     

Long- and short-term effects of reindeer grazing on tundra wetland vegetation

We studied long-term (50 years) and short-term (4 years) effects of summer grazing of reindeer on subarctic tundra wetland vegetation. The long-term effects of summer grazing were studied by comparing vegetation on Finnish and Norwegian sides of the fence line separating reindeer grazing regimes. The Finnish side was intensively grazed and trampled throughout the year, whereas the Norwegian side was grazed in winter. Experimental fences were erected to examine short-term effects of grazing exclusion. Both in the long- and short-term, summer grazing decreased the height of Salix lapponum whereas the short-term effects on willow cover were less clear than the long-term effects. In contrast, Carex spp. benefited from grazing. Long-term grazing had little effect on total bryophyte cover. Grazing had negligible effects on the nutrient content of leaves of S. lapponum and Eriophorum angustifolium. We conclude that tundra wetlands can withstand moderately high grazing pressure sustained over several decades.     

Ossification of the antler in the Lapland reindeer (Rangifer tarandus tarandus)

A histological and histochemical study of biopsy specimens from the Lapland reindeer antler indicated that the intercellular matrix of the cartilage that forms the partitions of longitudinal channels becomes calcified at virtually the initial stage of formation. The lacunae at the peripheries of the cartilaginous partitions are invaded by osteoblasts in a process comparable to endochondral ossification. The very centres of the partitions evidently become directly converted into bone without the presence of e.g. osteoclasts. The osseous partitions are remodelled by osteoclasts. The antler increases in diameter by periosteal apposition.     

Facilitation versus competition in grazing herbivore assemblages

The importance of facilitation versus competition in structuring herbivore species assemblages is a critical issue in theoretical ecology as well as for practical wildlife management. This paper examines the evidence for facilitation and clarifies our understanding in relation to the mechanisms and the spatial and temporal scales where they occur. Evidence for facilitation through stimulation of grass regrowth during the growing season appears stronger than that for increased resource access through removal of obstructing grass structures during the dormant season. Although facilitation may benefit the nutritional gains obtained by certain species in the short term, these benefits do not appear to be translated into the expected population consequences. We suggest this could be due to seasonal tradeoffs between facilitation and competition, as well as to restrictions on the spatial extent of trophic overlap.     

Spatio-temporal patterns of ground-dwelling ant assemblages in a lowland rainforest in southeast Cameroon

We studied the spatio-temporal patterns of ground-dwelling and -foraging ant assemblages in a continuous mixed secondary lowland rainforest on the northern periphery of the Dja Biosphere Reserve in southeast Cameroon. The effect of season and of the interaction of season with vegetation type on ant species density, activity and composition were investigated. We also checked for a possible impact of army ant activity on the ant assemblage. Ants were collected by pitfall trapping in seven vegetation types in each of three seasons during 12 months in 2003 and 2004–2005. Season significantly influenced the ground-dwelling ant assemblage. The late wet season had the lowest ant species density and a different species composition than the early wet and the dry season. Also inter-annual variation in ant species density and composition was detected and could partly be explained by an influence of Dorylus activity. In general, vegetation type and inter-annual variation had a larger influence on the ground-ant assemblage than season. Significant effects of interactions between season, year, vegetation type and army ants on the ant assemblage confirm that also in African rainforests ant assemblages are extremely dynamic and depend on a complex combination of availability of food resources, nest sites and predators. Received 4 October 2006; revised 4 June 2007; accepted 2 July 2007.     

Seasonal grazing effects by semi-domesticated reindeer on subarctic mountain birch forests

In northern Fennoscandia, the spatial and temporal grazing practices of semi-domesticated reindeer (Rangifer tarandus tarandus) vary, which implies different grazing effects dependent on natural conditions as well as management regime (i.e., timing and intensity of grazing). We compared density and biomass of main plant groups in semi-dry mountain birch forests exposed to either long-term summer or winter grazing in three reindeer herding districts in the northernmost Finland. Percent plant cover, height, and biomass of reindeer lichens (Cladonia spp.) and dwarf shrubs (Vaccinium uliginosum, Calluna vulgaris, and Betula nana) were lower on summer ranges compared with winter ranges. The biomass of other dwarf shrubs (Vaccinium myrtillus and V. vitis-idaea), and graminoids and herbs, and the % cover of non-vegetated bare soil and litter were, however, higher on summer ranges than on winter ranges. Young mountain birch shoots (Betula pubescens ssp. czerepanovii) were less frequent on summer ranges than on winter ranges. The total leaf biomass under the browsing height of reindeer (<1.5 m) was also lower on summer ranges compared with winter ranges. Especially in drier and nutrient poor mountain birch forests, intensive summer grazing reduces the quantity of lichens and total plant biomass which affects the ecological state and productivity of these forests and also reduces especially their winter grazing value for reindeer. Therefore, in addition to regulating the maximum sustained numbers of reindeer, pasture rotation systems that effectively protect dry and nutrient poor vegetation from summer grazing and trampling should be encouraged.     

Size-selective grazing on bacteria by natural assemblages of estuarine flagellates and ciliates

The small average cell size of in situ bacterioplankton, relative to cultured cells, has been suggested to be at least partly a result of selection of larger-sized cells by bacterivorous protozoa. In this study, we determined the relative rates of uptake of fluorescence-labeled bacteria (FLB), of various cell sizes and cell types, by natural assemblages of flagellates and ciliates in estuarine water. Calculated clearance rates of bacterivorous flagellates had a highly significant, positive relationship with size of FLB, over a range of average biovolume of FLB of 0.03 to 0.08 microns3. Bacterial cell type or cell shape per se did not appear to affect flagellate clearance rates. The dominant size classes of flagellates which ingested all types of FLB were 3- to 4-microns cells. Ciliates also showed a general preference for larger-sized bacteria. However, ciliates ingested a gram-positive enteric bacterium and a marine bacterial isolate at higher rates than they did a similarly sized, gram-negative enteric bacterium or natural bacterioplankton, respectively. From the results of an experiment designed to test whether the addition of a preferentially grazed bacterial strain stimulated clearance rates of natural bacterioplankton FLB by the ciliates, we hypothesized that measured differences in rates of FLB uptake were due instead to differences in effective retention of bacteria by the ciliates. In general, clearance rates for different FLB varied by a factor of 2 to 4. Selective grazing by protozoa of larger bacterioplankton cells, which are generally the cells actively growing or dividing, may in part explain the small average cell size, low frequency of dividing cells, and low growth rates generally observed for assemblages of suspended bacteria.     

Secondary metabolites in Cladina stellaris in relation to reindeer grazing and thallus nutrient content

We analysed concentrations of phenols and key macro-nutrients in a dominant mat-forming lichen, Cladina stellaris, in dry heath forests subjected to heavy reindeer grazing in Finnish Lapland. Lichen samples were collected in the beginning of two growth seasons from six to eight sites with old reindeer exclosures that served as control plots within sites. The concentration of perlatolic acid in lichen apices (top 10 mm) was higher in grazed plots than in control ones, whereas usnic acid did not seem to respond unequivocally to grazing. Moreover, there was a strong negative relationship between nitrogen and phenolic content of Cladina stellaris in intact plots but this relationship was absent in grazed ones. Changes in nitrogen and phosphorus contents caused by reindeer grazing were not correlated with changes in the level of phenolics in lichen thalli. The present result demonstrated that the carbon-nutrient balance (CNB) hypothesis may well explain some of the variation in lichen phenolic content but is not sufficient for explaining changes caused by reindeer grazing. We hypothesized that physical changes in the lichen microenvironment induced by reindeer have more profound impact on lichen phenolic content than alterations in thallus nutrient content.     

Effects of summer grazing by reindeer on composition of vegetation, productivity and nitrogen cycling

In this study, we investigated the effect of reindeer grazing on tundra heath vegetation in northern Norway. Fences, erected 30 yr ago, allowed us to compare winter grazed, lightly summer grazed and heavily summer grazed vegetation at four different sites. At two sites, graminoids dominated the heavily grazed zone completely, while ericoid dwarf shrubs had almost disappeared. In the other two areas, the increase of graminoids was almost significant. At one of the sites where graminoids dominated the heavily grazed area, we also measured plant biomass, primary production and nitrogen cycling. In this site, heavy grazing increased primary production and rate of nitrogen cycling, while moderate grazing decreased primary production. These results were inconsistent with the view that the highest productivity is found at intermediate grazing pressure. These results rather support the hypothesis that intensive grazing can promote a transition of moss-rich heath tundra into productive, graminoid-dominated steppe-like tundra vegetation. Moreover the results suggests that intermittent intensive reindeer grazing can enhance productivity of summer ranges.     

Rapid recovery of recently overexploited winter grazing pastures for reindeer in northern Norway

During the last 25 yr, Sami reindeer husbandry in parts of Finnmarksvidda in the Norwegian Arctic has been in a critical state because of overexploitation of lichen-dominated tundra, which serves as winter forage. To better understand the ecosystem’s capabilities for recovery we investigated vegetation cover changes over a 7-yr period, starting in 1998, at 52 sites dispersed over a large area at Finnmarksvidda. Two types of plots were established: one fenced from reindeer grazing and trampling and one open for reindeer. The investigations in 2005 showed that lichen cover had had a significant and rapid increase (up to 8.6-fold per year). The cover of vascular plants, mainly dwarf shrubs, also increased significantly, while barren areas and the cover of litter decreased significantly during the period. Mean relative growth rate of lichen biomass was 0.083 ± 0.011 per year in open plots, which is considered very rapid recovery compared to previous studies. Lichen recovery was significantly faster on leeward ridges than on exposed ridges, and fencing alone did not have any significant effects on lichen recovery, but in interaction with time, fencing contributed to increasing recovery rates. The lichen heath recovery was reciprocally correlated with reindeer density. In addition, lichen recovery was probably facilitated by recent climate changes, viz. shallower snow depths which made leeward tundra and forest floor vegetation accessible for reindeer, and increased summer precipitation rates which improved growth rates. The results from this study show that in a very short time there was a transition from an overexploited depauperate vegetation and barren ground state to a flourishing lichen-dominated vegetation state, suggesting that the injuries were repairable. The vegetation transitions which have taken place in the study area are considered to be reversible with fewer persistent effects.     

Soil dumping techniques and afforestation drive ground-dwelling beetle assemblages in a 25-year-old open-cast mining reclamation area

The open-cast lignite mine “Hambach” (NW-Germany) and the accompanying land reclamation replace a large ancient forest. Reclamation of an 1000 ha large overburden dump started 25 years ago and is still in progress. Reclamation methods comprise combinations of different topsoil dumping techniques and reclamation measures such as seeding or afforestation. In accordance with the spatial heterogeneity hypothesis, we hypothesize that beetle assemblages from sites where the topsoil was left structured are more diverse than those from sites with levelled topsoil, and that richly structured mid-successional sites harbour more heterogeneous assemblages than afforested sites. We tested the influence of environmental variables on assemblage characteristics of pitfall-trapped beetles from 17 sites on the overburden dump. The results are based on 35,588 individuals from 339 species. Species assemblages differed significantly between almost all sites. Habitat age did not affect assemblage characteristics, while species richness was significantly influenced by surface structure. Sites that were not levelled and that additionally included pits that slope down for several meters exhibited the highest species richness. Abundance of beetles increased in parallel to the canopy cover, whereas assemblage heterogeneity (MVDISP) decreased. Successional and grassland sites usually exhibited higher diversity (Fisher's α) than afforested sites, but diversity was not significantly predicted by a single environmental factor. Instead, diversity increased along with increasing MVDISP-values. Afforestations speed up the successional process and some euryecious forest species are already well established, but stenecious forest specialists have not yet been found.     

The effect and extent of heavy grazing by reindeer in oligotrophic pine heaths in northeastern Fennoscandia

Thirty and fifty years old exclosures established in northeastern Fennoscandia in lichen-rich oligotrophic pine Pinus sylvestris forests on podzolised soil were used to study the effect of reindeer grazing on pine fine roots, microbial activity, and on bryophyte, dwarf shrub and lichen biomasses. There were significantly less lichens, especially Cladina stellaris , at grazed than at ungrazed sites. Coverage of other lichens like C. arbuscula and C. rangiferina and bryophytes, especially Dicranum spp., benefitted from grazing. The biomass of vascular plants, mainly Calluna vulgaris, Empetrum nigrum and Vaccinium vitis-idaea , was reduced at grazed sites, although their coverage was not influenced. Microbial activity was significantly lower at grazed sites. The influence of grazing is most obviously mediated by reduced soil moisture during dry periods at grazed sites. Fine root parameters (per soil and stem volumes) were lower at grazed sites (pPCA = 0.072), the first principal component consisting of a number of fine root tips, length and weight. Grazing decreased all exchangeable nutrients by 30–60% in organic layer. Based on PCA the decrease was significant for exchangeable nutrients, although of individual elements only P and S showed statistically significant difference. The extent of heavy grazing in northeastern Fennoscandia coniferous forest was revealed by remote sensing. It revealed extensive area in which reindeer lichens are reduced in northeastern Finland. The Finnish-Russian border can be clearly distinguished in the satellite image composite.     

Impact of grazing on vegetation dynamics in former ricefields

Abstract. In the Rhône delta, Juncus gerardi and Scirpus maritimus are often the dominant species in abandoned rice fields which are artificially flooded in early spring to improve forage production. Under these conditions they occur either in mixed communities, or form monospecific stands. Monitoring the vegetation dynamics in quadrats located in six abandoned rice fields artificially flooded from November to April confirmed the important role of grazing. In ungrazed plots, communities dominated by Scirpus maritimus mixed with Juncus gerardi developed fast. After 42 months of management Scirpus maritimus had established in nearly all quadrats and continued to expand, whereas Juncus gerardi had started to decline. In grazed plots Juncus gerardi alone dominated and continued to increase in cover up to the 42nd month. Scirpus maritimus established at low densities mainly in quadrats where Juncus gerardi was initially absent. Introduction of seeds of Scirpus maritimus in communities of Juncus gerardi under controlled conditions demonstrated the existence of the phenomenon of preemption. The increase in cover of Juncus gerardi suggests that the preemption of Juncus gerardi over Scirpus maritimus plays a more pronounced role in the field in the presence of grazing.     

The effect of reindeer grazing on decomposition, mineralization and soil biota in a dry oligotrophic Scots pine forest

Reindeer grazing in the Fennoscandian area has a considerable influence on the ground vegetation, and this is likely in turn to have important consequences for the soil biota and decomposition processes. The effects of reindeer grazing on soil biota, decomposition and mineralization processes, and ecosystem properties in a lichen‐dominated forest in Finnish Lapland were studied inside and outside a large long term fenced reindeer exclosure area. Decomposition rates of Vaccinium myrtillus leaves in litter bags were retarded in the grazed area relative to the ungrazed area, as well as in subplots from which lichens had been artificially removed to simulate grazing. The effect of reindeer grazing on soil respiration and microbial C was positive in the lichen and litter layers of the soil profile, but retarded in the humus layer. There was no effect of grazing on gross N mineralization and microbial biomass N in the humus and upper mineral soil layer, but net N mineralization was increased by grazing. In these layers soil respiration was reduced by grazing, indicating that reindeer effects reduce the ratio of C to N mineralized by soil microorganisms. Grazing stimulated populations of all trophic groupings of nematodes in the lichen layer and microbe feeding nematodes in the litter layer, indicating that grazing by reindeer has multitrophic effects on the decomposer food‐web. Grazing decreased lichen and dwarf shrub biomasses and increased the mass of litter present in the litter layer on an areal basis, but did not significantly alter total C storage per unit area in the humus and mineral soil layers. The N concentration of lichens was increased by grazing, but the N concentrations of both living and dead Pinus sylvestris needles and Empetrum hermaphroditum leaves were not affected.
There was some evidence for each of three mechanisms which could account for the grazing effects that we observed in our study. Firstly, reindeer may have changed the composition and quality of litter input by affecting plant species composition and through addition of N from urine and faeces, resulting in a lack of available C relative to N for decomposer organisms. Secondly, the organic matter in the soil may be older in the grazed area, because of reduction of recent production of lichen litter relative to the ungrazed area. The organic matter in the grazed area may have been in a different phase of decomposition from that in the exclosure. Thirdly, the soil microclimate is likely to be affected by reindeer grazing through physical removal of lichen cover on the ground, and this can have a significant influence on soil microbial processes. This is supported by the strong observed effects of experimental removal of lichens on decomposer processes. The impact of reindeer grazing on soil processes may be a result of complex interactions between different mechanisms, and this could help to explain why the below‐ground effects of reindeer grazing have different consequences to those which have been observed in recent investigations on other grazing systems.     

Alterations in the biomass-specific productivity of periphyton assemblages mediated by fish grazing

1. In an experimental flume, we examined the effects of a biomass reduction and alteration of taxonomic composition, because of grazing by the fish Plecoglossus altivelis, on the net biomass accumulation of periphyton. 2. Grazed and ungrazed assemblages with different biomass and taxonomic composition were first prepared in fish enclosures and exclosures, respectively. These assemblages were then set out in the flume and incubated for 2 days under grazing‐free conditions to examine (i) the relationship between biomass and biomass accumulation rate and (ii) the effect of taxonomic composition on the relationship between these two. 3. The grazed and ungrazed assemblages were dominated by upright filamentous cyanobacteria and diatoms, respectively. The rate of biomass accumulation decreased with increasing periphyton biomass in both the grazed and ungrazed assemblages, and was lower in the grazed than the ungrazed assemblages at any biomass level. 4. The results showed that the reduction in biomass and the alteration of taxonomic composition due to fish grazing have opposite effects on biomass‐specific productivity. Biomass accumulation rate increased in response to biomass reduction, although a shift in dominance from diatoms to upright filamentous cyanobacteria decreased the overall productivity of the periphyton.     

Ground-beetle (Coleoptera,Carabidae) assemblages in the forest-steppe of the southern Vitim Plateau

A list of species and the structure of ground beetle assemblages in the forest-steppe of Eravninskaya (= Sosnovoozerskaya) Depression situated in the permafrost area are presented. Ground beetle assemblages dwelling in larch, mixed, and small-leaved forests (intact and after fires), and also in the meadow steppe and waterlogged meadows with varying degrees of humidity were studied. The data obtained demonstrate that the communities examined are intermediate between those revealed in adjacent territories of the Selenga River basin, mid-altitude mountains, and of the northern Transbaikalia. The gentle sauce-shaped relief of Eravninskaya Depression is responsible for the concentric circular distribution of carabid assemblages, occasionally interrupted by forest-steppe fragments in isolated forests and meadow steppes. All the hill slopes surrounding the depression are populated by taiga carabid assemblages, whereas the bottom of the depression is the habitat for meadow-steppe and forest-steppe assemblages.     

Effects of urbanization on ground beetle assemblages (Coleoptera, Carabidae) of grassland habitats in western Canada

We pitfall-trapped carabid beetles across an urban-rural gradient in central Alberta, Canada to assess the relative contributions of landscape structure and habitat quality in explaining variation in composition of assemblages. The gradient ranged from the "urban" zone in the city of Edmonton, through a "suburban" zone just outside the city, into a "rural" zone 25–60 km east of the city. We trapped beetles at three replicated sites for each of two habitats (unmanaged grasslands and highly managed graveyards) in each of the three zones. Most of the 24 species (3162 individuals) captured were commonly associated with open habitats. Pterostichus melanarius , an introduced species, made up 80% of the total catch. Carabid assemblages of grasslands and graveyards were distinct, with lower species richness and lower catch rates of native, introduced and brachypterous carabids in graveyards. Graveyard assemblages also had proportionally fewer brachypterous carabid individuals (32 vs 52%) and species (20 vs 36%), suggesting that populations recruit locally from elsewhere after disturbances. Native carabids were least abundant in the urban areas. Brachypterous carabids were similarly abundant across the gradient, but more occurred in the urban zone than elsewhere (32 vs 5–10%). Samples from graveyards were characterized by lower beta diversity (variation among samples) than those from grasslands, but beta diversity was more-or-less equally variable across the gradient within habitats. The amount of unmanaged grassland habitat and the degree of urbanization significantly influence carabid assemblages, and therefore are relevant to management and conservation of urban grassland areas.     

Non-parallel changes in soil microbial carbon and nitrogen dynamics due to reindeer grazing in northern boreal forests

Reindeer grazing has a considerable influence on mineralization processes in northern Fennoscandian boreal forests, but the mechanisms underlying the observed differences between grazed and ungrazed areas are not well understood. We studied the below-ground impacts of reindeer grazing by comparing the carbon and nitrogen mineralization rates inside and outside long-term fenced reindeer exclosure areas in five oligotrophic, lichen-dominated and five mesotrophic, dwarf-shrub dominated forests. The soil C mineralization rates and microbial metabolic activity (qCO₂) were significantly lower in the grazed than the ungrazed areas in both oligotrophic and mesotrophic forests. The reductions occurred irrespective of the impact on soil moisture. We conclude that reindeer grazing causes a reduction in the supply of labile C substrates to microbes, resulting in reduced organic matter decomposition rates through changes in the activity of the microbial biomass. Simultaneously, grazing had no consistent effect on the microbial N dynamics, but the impact ranged from no change to increased or decreased in N mineralization rates at the different study sites. The impact of grazing on the N mineralization potential thus seems to be site-specific and uncoupled from the impact of grazing on soil C mineralization. Reciprocal transplant incubations showed no interactions between N mineralization rates and the reindeer-mediated impact on the soil microclimate. We suggest that plant root damage due to trampling by reindeer may be an important mechanism for the deceleration of soil C cycling. In some cases, however, the impact of grazing on the soil active N pool may be strong enough to outweigh the reduction in soil organic matter decomposition, and by these means uncouple soil N dynamics from soil C quality.     

Distribution of ground-dwelling beetle assemblages (Coleoptera) across ecotones between natural oak forests and mature pine plantations in North China

This paper studied edge effects resulting from logging to reforestation on the distribution of ground-dwelling beetles (Coleoptera) across ecotones between natural oak forests and mature pine plantations established after harvesting of natural forests. Using pitfall traps, ground-dwelling beetles were investigated at three replicated plots (ecotones) with three sampling positions of slope (lower, middle and upper) for each plot. Rarefaction estimates of species richness indicated that traps on natural forests and transition zones had more species than mature plantations did, and traps on the middle slope had more species than on the lower and upper slopes did. Results of an ANOVA analysis, which used forest type and slope position as factors and number of species and individuals as the response variables, showed a significant effect of forest type and slope position, and a significant interaction between forest type and slope position. Multivariate analyses (DCA and CCA) showed that beetles of transition zones were more similar to those of natural forests than to those of mature plantations, and that some environmental characteristics, i.e., proportion of broad-leaved trees, canopy cover and elevation (slope position), significantly affected species abundances. We conclude that the logging of natural oak forests and the reforestation of pine plantations can result in subtle variation in the composition and distribution of beetle assemblages at a local scale and such variation should be taken into account when conservation issues are involved.