Historical contingency is the impact of past events, like the timing and order of species arrival, on community assembly, and can sometimes result in alternative stable states of ecological communities. Large herbivores, wild and domestic, can cause profound changes in the structure and functioning of plant communities and therefore probably influence historical contingency; however, little empirical data on the stability of such shifts or subsequent drivers of stability are available. We studied the centennial legacy of reindeer (Rangifer tarandus) pressure on arctic tundra vegetation by considering historical milking grounds (HMGs): graminoid-and forb-dominated patches amid shrub-dominated tundra, formed by historical Sami reindeer herding practices that ended approximately 100 years ago. Our results show that the core areas of all studied HMGs remained strikingly stable, being hardly invaded by surrounding shrubs. Soil nitrogen concentrations were comparable to heavily grazed areas. However, the HMGs are slowly being reinvaded by vegetative growth of shrubs at the edges, and the rate of ingrowth increased with higher mineral N availability. Furthermore, our data indicate that several biotic feedbacks contribute to the stability of the HMGs: increased nutrient turnover supporting herbaceous vegetation, strong interspecific competition preventing invasion and herbivore damage to invading shrubs. In particular, voles and lemmings appear to be important, selectively damaging shrubs in the HMGs. We concluded that HMGs provide clear evidence for historical contingency of herbivore effects in arctic ecosystems. We showed that several biotic feedbacks can contribute to subsequent vegetation stability, but their relative importance will vary in time and space.
The present review aims to summarize current knowledge in the topic of wood ash application to boreal forest and aquatic ecosystems, and the different effects derived from these actions. Much research has been conducted regarding the effects of wood ash application on forest growth. Present studies show that, generally speaking, forest growth can be increased on wood ash-ameliorated peatland rich in nitrogen. On mineral soils, however, no change or even decreased growth have been reported. The effects on ground vegetation are not very clear, as well as the effects on fungi, soil microbes, and soil-decomposing animals. The discrepancies between different studies are for the most part explained by abiotic factors such as variation in fertility among sites, different degrees of stabilization, and wood ash dosage used, and different time scales among different studies. The lack of knowledge in the field of aquatic ecosystems and their response to ash application is an important issue for future research. The few studies conducted have mainly considered changes in water chemistry. The biotoxic effects of ash application can roughly be divided into two categories: primary and secondary. Among the primary effects is toxicity deriving from compounds in the wood ash and cadmium is probably the worst among these. The secondary effects of wood ash are generally due to its alkaline capacity and a release of ions into the soil and soil water, and finally, watercourses and lakes. Given current knowledge, we would recommend site- and wood ash-specific application practices, rather than broad and general guidelines for wood ash application to forests.
The limited availability of historical and archaeological evidence means that much is still unknown about the development of Sami reindeer herding in Fennoscandia in both the recent and more distant past. To address this problem, high-resolution palynological analyses, 14 C and 210 Pb dating were undertaken on two adjacent (\25 m apart) peat profiles collected at a recently abandoned reindeer gathering pen (renvall) near Jokkmokk (*66.6°N, 19.8°E) in the boreal forest of northern Sweden. The aim was to assess the impact of Sami reindeer herding on the local environment through a study of pollen, coprophilous fungal spores, microscopic charcoal and sedimentology. The samples collected from within an annex to the renvall indicate cycles of use and abandonment of the pen on a multi-decadal timescale between *AD 1800-2008, most obviously in the coprophilous fungal spore archive. The pattern and timing of these cycles confirm events previously known only from oral histories. Although the local pollen assemblage zones associated with the phasing of activity were reproducible in a second peat core beyond the boundary of the renvall, the coprophilous fungal spore signal in this paired profile was much less distinctive, possibly due to the typically shorter dispersal distances for these microfossils in comparison to pollen grains.
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