Hans Henrik Bruun

Effects of altitude and topography on species richness of vascular plants, bryophytes and lichens in alpine communities

Abstract Question: What is the relationship between species richness of vascular plants, bryophytes and macrolichens, and two important gradients in the alpine environment, altitude and local topography? Location: Northernmost Fennoscandia, 250–1525 m a.s.l. corresponding to the range between timberline and mountain top. Methods: The vegetation was sampled in six mountain areas. For each 25 vertical metres, the local topographic gradient from wind-blown ridge to snowbed was sampled in quadrats of 0.8 m × 0.8 m. Patterns in species richness were explored using Poisson regression (Generalized Linear Models). Functional groups of species, i.e. evergreen and deciduous dwarf-shrubs, forbs, graminoids, mosses, hepatics and lichens were investigated separately. Results: Functional groups showed markedly different patterns with respect to both altitude and topography. Species richness of all vascular plants showed a unimodal relationship with altitude. The same was true for graminoids, forbs and lichens analysed separately, but forb richness peaked at much higher altitudes than total richness. The richness of dwarf-shrubs decreased monotonically with altitude, whereas richness of mosses and liverworts showed an increasing trend. Significant interactions between altitude and local topography were present for several groups. The unimodal pattern for total plant species richness was interpreted in terms of local productivity, physical disturbance, trophic interactions, and in terms of species pool effects. Conclusions: Patterns in local species richness result from the action of two opposing forces: declining species pool and decreasing intensity of competition with altitude. Nomenclature: Nilsson (1986) and Hallingbäck (1995, 1996) for vascular plants, lichens and bryophytes, respectively.

Local temperatures inferred from plant communities suggest strong spatial buffering of climate warming across Northern Europe

Recent studies from mountainous areas of small spatial extent (<2500 km(2) ) suggest that fine-grained thermal variability over tens or hundreds of metres exceeds much of the climate warming expected for the coming decades. Such variability in temperature provides buffering to mitigate climate-change impacts. Is this local spatial buffering restricted to topographically complex terrains? To answer this, we here study fine-grained thermal variability across a 2500-km wide latitudinal gradient in Northern Europe encompassing a large array of topographic complexities. We first combined plant community data, Ellenberg temperature indicator values, locally measured temperatures (LmT) and globally interpolated temperatures (GiT) in a modelling framework to infer biologically relevant temperature conditions from plant assemblages within <1000-m(2) units (community-inferred temperatures: CiT). We then assessed: (1) CiT range (thermal variability) within 1-km(2) units; (2) the relationship between CiT range and topographically and geographically derived predictors at 1-km resolution; and (3) whether spatial turnover in CiT is greater than spatial turnover in GiT within 100-km(2) units. Ellenberg temperature indicator values in combination with plant assemblages explained 46-72% of variation in LmT and 92-96% of variation in GiT during the growing season (June, July, August). Growing-season CiT range within 1-km(2) units peaked at 60-65°N and increased with terrain roughness, averaging 1.97 °C (SD = 0.84 °C) and 2.68 °C (SD = 1.26 °C) within the flattest and roughest units respectively. Complex interactions between topography-related variables and latitude explained 35% of variation in growing-season CiT range when accounting for sampling effort and residual spatial autocorrelation. Spatial turnover in growing-season CiT within 100-km(2) units was, on average, 1.8 times greater (0.32 °C km(-1) ) than spatial turnover in growing-season GiT (0.18 °C km(-1) ). We conclude that thermal variability within 1-km(2) units strongly increases local spatial buffering of future climate warming across Northern Europe, even in the flattest terrains.

COMMUNITY ASSEMBLY IN EXPERIMENTAL GRASSLANDS: SUITABLE ENVIRONMENT OR TIMELY ARRIVAL?

It is hard to defend the view that biotic communities represent a simple and predictable response to the abiotic environment. Biota and the abiotic environment interact, and the environment of an individual certainly includes its neighbors and visitors in the community. The complexity of community assembly calls forth a quest for general principles, yet current results and theories on assembly rules differ widely. Using a grassland microcosm as a model system, we manipulated fertility, disturbance by defoliation, soil/microclimate, and arrival order of species belonging to two groups differing in functional attributes. We analyzed the outcome of community assembly dynamics in terms of species richness, invasibility, and species composition. The analyses revealed strong environmental control over species richness and invasibility. Species composition was mainly determined by the arrival order of species, indicating that historical contingency may change the outcome of community assembly. The probability for multiple equilibria appeared to increase with productivity and environmental stability. The importance of arrival order offers an explanation of the difficulties in predicting local occurrences of species in the field. In our experiment, variation in fertility and disturbance was controlling colonization with predictable effects on emergent community properties such as species richness. The key mechanism is suggested to be asymmetric competition, and our results show that this mechanism is relatively insensitive to the species through which it works. While our analyses indicate a positive and significant correlation between richness and invasibility, the significance disappears after accounting for the effect of the environment. The importance of arrival order (historical contingency) and environmental control supports the assumption of the unified neutral theory that different species within a trophic level can be considered functionally equivalent when it comes to community assembly. However, our results indicate that variation in asymmetric competition is the key factor determining the richness of the resulting communities, and this is far from neutral.

The Past Impact of Livestock Husbandry on Dispersal of Plant Seeds in the Landscape of Denmark

Abstract The recent decline in species richness in (semi)-natural habitats in northern Europe has largely been attributable to habitat destruction, and to subsequent limitation in seed dispersal among fragments. However, some habitat types were probably split up already in the historical landscape, but the segregated parts were probably not isolated to the present degree. This paper seeks evidence for livestock as vectors for propagules at 3 spatial scales in the past cultural landscape. Three main scales at which livestock acted as seed dispersers are important: free movement in the landscape (1–10 km), driving animals to mast feeding or to manors (10–50 km), and the export of living animals (hundreds of km). The emerging picture is for most plant species a dramatically decreased chance of dispersal in the modern landscape. The consequence is probably decreasing species richness in (semi)-natural plant communities, such as pasture, meadow, and heathland.

Strong microsite control of seedling recruitment in tundra.

The inclusion of environmental variation in studies of recruitment is a prerequisite for realistic predictions of the responses of vegetation to a changing environment. We investigated how seedling recruitment is affected by seed availability and microsite quality along a steep environmental gradient in dry tundra. A survey of natural seed rain and seedling density in vegetation was combined with observations of the establishment of 14 species after sowing into intact or disturbed vegetation. Although seed rain density was closely correlated with natural seedling establishment, the experimental seed addition showed that the microsite environment was even more important. For all species, seedling emergence peaked at the productive end of the gradient, irrespective of the adult niches realized. Disturbance promoted recruitment at all positions along the environmental gradient, not just at high productivity. Early seedling emergence constituted the main temporal bottleneck in recruitment for all species. Surprisingly, winter mortality was highest at what appeared to be the most benign end of the gradient. The results highlight that seedling recruitment patterns are largely determined by the earliest stages in seedling emergence, which again are closely linked to microsite quality. A fuller understanding of microsite effects on recruitment with implications for plant community assembly and vegetation change is provided.

Gradient analysis of dry grassland vegetation in Denmark

Abstract. We present a gradient analysis of 620 vegetation samples covering most of the floristic and environmental variation in semi-natural grassland vegetation on well-drained soils in Denmark. Vegetation was sampled using frequency in subplots. Explanatory variables were surface inclination, aspect, pH, geographical co-ordinates together with indications of soil type. Detrended Correspondence Analysis revealed four floristic gradients that could be interpreted in ecological terms by measured variables supplemented with site calibrations based on weighted averaging of Ellenbergs indicator values. All four axes were interpreted using rank correlation statistics, and linear and non-linear multiple regression of sample scores on explanatory variables. The first gradient was from dry calcareous to humid acidic grasslands; the second reflected an underlying gradient in fertility; the third reflected regional differentiation and the fourth was associated with variation in intensity of competition as indicated by association with calibrated Grime-CSR values for the plots. We applied subset ordination to the data as a supplement to traditional permutation and correlation statistics to assess the consistency of ordination results. DCA axes 1 and 2 were consistent in space and time. This gradient analysis is discussed in a context of plant strategy theory and species diversity models. Ecocline patterns lend support to the view that grazing not only favours the ruderal strategy but also the stress-tolerant strategy. The low rank of competition as an explanatory variable for the floristical gradients supports the notion that competitive effects play a subordinate role for species composition compared to microclimate and soil conditions in infertile semi-natural grasslands.

Landscape genomics and a common garden trial reveal adaptive differentiation to temperature across Europe in the tree species Alnus glutinosa

The adaptive potential of tree species to cope with climate change has important ecological and economic implications. Many temperate tree species experience a wide range of environmental conditions, suggesting high adaptability to new environmental conditions. We investigated adaptation to regional climate in the drought‐sensitive tree species Alnus glutinosa (Black alder), using a complementary approach that integrates genomic, phenotypic and landscape data. A total of 24 European populations were studied in a common garden and through landscape genomic approaches. Genotyping‐by‐sequencing was used to identify SNPs across the genome, resulting in 1990 SNPs. Although a relatively low percentage of putative adaptive SNPs was detected (2.86% outlier SNPs), we observed clear associations among outlier allele frequencies, temperature and plant traits. In line with the typical drought avoiding nature of A. glutinosa, leaf size varied according to a temperature gradient and significant associations with multiple outlier loci were observed, corroborating the ecological relevance of the observed outlier SNPs. Moreover, the lack of isolation by distance, the very low genetic differentiation among populations and the high intrapopulation genetic variation all support the notion that high gene exchange combined with strong environmental selection promotes adaptation to environmental cues.

A multivariate approach to plant community distribution in the coastal dune zonation of NW Denmark

A near-natural coastal dune system in NW Jutland, Denmark was investigated. The area contains a gradient in natural soil mobility with the highest values in dynamic white dunes dominated by Ammophila arenaria, and low mobility in fixed decalcified dune heath with Empetrum nigrum and Calluna vulgaris. A total of 267 plots was investigated and presence of mosses and rooted vascular plants recorded. Cluster analysis identified five communities: white dune, grey dune, outer dune heath, inner dune heath and humid dune heath. These communities were ordered along two main axes in a Detrended Correspondence Analysis (DCA). The relationship between DCA axes and several environmental variables, including soil characteristics and Ellenberg indicator values, were investigated by correlation. Two major complex gradients underlying compositional variation were identified; one short coenocline with an underlying steep gradient in habitat maturity, soil stability and nutrient-limited productivity, and one longer gradient in water-limited productivity and peat accumulation only applying to the older successional stages. Distance from the coast was an important correlate of several of the measured variables. Microclimatic indices, i.e. heat index and wind index, had limited value as descriptors of community patterns as they were not correlated with the DCA axes or with other environmental variables. However, within communities the microclimatic indices correlated with several explanatory variables. (Less)

Prediction of grassland quality for environmental management

Fifty plots on grassland on well-drained soil were investigated for land use history and analysed for the species composition of vascular plants, macrofungi, bryophytes, lichens, hepatics and for 20 environmental variables. A DCA ordination showed an overall influence of land use history on species composition. DCA axis 1 was strongly correlated with species diversity of vascular plants. High species diversity is proposed to result from the combined action of high pH, high CaCO 3 content, high age of turf, grazing and absence of fertilization. A DCCA ordination lead to the proposal of 22 species as indicators of old, unameliorated grasslands in Denmark.

Developing a classifier for the Habitats Directive grassland types in Denmark using species lists for prediction

Abstract This paper describes the use of supervised methods for the classification of vegetation. The difference between supervised classification and clustering is outlined, with reference to their current use in vegetation science. In the paper we describe the classification of Danish grasslands according to the Habitats Directive of the European Union, and demonstrate how a supervised classification can be used to achieve a standardized and statistical interpretation within a local flora. We thereby offer a statistical solution to the legal problem of protection of certain selected habitat types. The Habitats Directive protects three types of Danish grassland habitats, whereas two remaining types fall outside protection. A classification model is developed, using available Danish grassland data, for the discrimination of these five types based on their species composition. This new Habitats Directive classification is compared to a previously published unsupervised classification of Danish grassland vegetation. An indicator species analysis is used to find significant indicator species for the three protected habitat types in Denmark, and these are compared to the characteristic species mentioned in the interpretation manual of the Habitats Directive. Eventually, we discuss the pros and cons of supervised and unsupervised classification and conclude that supervised methods deserve more attention in vegetation science. Abbreviations: DCA = Detrended Correspondence Analysis; CCA = Canonical Correspondence Analysis; CVA = Canonical Variates Analysis; HD = Habitats Directive; TWINSPAN = Two-way indicator species analysis. Nomenclature: Tutin et al. (1964–1993).