Differences in colonization strategies of three common pioneer woody species in post mining heaps.

Willow (Salix caprea), birch (Betula pendula) and aspen (Populus tremula) are common pioneer woody species, however little is known about colonization strategies in large-scale disturbances. Here we have compared the strategies of establishment of these pioneer woody species in unreclaimed sites on a large (1957 ha) spoil heap in Czechia. For all species, seedlings numbers peaked in the 17 year old (successional age - time since overburden heaping) plot, suggesting that initial soil development promotes seedling establishment while covering of the surface by litter and organic layers reduces the establishment of pioneer species. The proportion of willow decreased from the edge of the heap and analysis of the age structure suggests that willow establishment was correlated with the presence of older willows in the vicinity of willows of certain ages (13 and 23 years being particularly important). The proportion of birch increased with its distance from the heap edge, and it is correlated with suitable weather conditions in the year of establishment, mainly July rainfall. Aspen proportion does not change significantly with its distance from the heap edge and year of establishment. It correlates with the number of trees in both surrounding and climatic conditions. Detailed analysis of young trees shows that vegetative propagation by root suckers (offspring) is rare in birch. In willow they represent about half of the trees while in aspen all of the young trees were root suckers derived from older aspen trees. This indicates a different colonization strategy of individual species. Birch is capable of long-distance seed transfer, which establishes most of the population, and its establishment is highly influenced by climatic conditions. Willow spreads massively over a short distance, and its establishment is highly influenced by the presence of 21-23 year old willow individuals in the vicinity. Only few scattered trees are able to establish at longer distances, where they can later spread locally by seeds. Aspen spreads over a long distance in low numbers but when some trees are established it spreads massively locally by clones.

Carbohydrate storage in herbs: the forgotten functional dimension of the plant economic spectrum.

BACKGROUND AND AIMS: Although the plant economic spectrum seeks to explain resource allocation strategies, carbohydrate storage is often omitted. Belowground storage organs are the centre of herb perennation, yet little is known about the role of their turnover, anatomy and carbohydrate storage in relation to the aboveground economic spectrum. METHODS: We collected aboveground traits associated with the economic spectrum, storage organ turnover traits, storage organ inner structure traits and storage carbohydrate concentrations for ~80 temperate meadow species. KEY RESULTS: The suites of belowground traits were largely independent of one another, but there was significant correlation of the aboveground traits with both inner structure and storage carbohydrates. Anatomical traits diverged according to leaf nitrogen concentration on the one hand and vessel area and dry matter content on the other; carbohydrates separated along gradients of leaf nitrogen concentration and plant height. CONCLUSIONS: Contrary to our expectations, aboveground traits and not storage organ turnover were correlated with anatomy and storage carbohydrates. Belowground traits associated with the aboveground economic spectrum also did not fall clearly within the fast-slow economic continuum, thus indicating the presence of a more complicated economic space. Our study implies that the generally overlooked role of storage within the plant economic spectrum represents an important dimension of plant strategy.

Strong impact of management regimes on rhizome biomass across Central European temperate grasslands.

Grassland ecosystems account for approximately 40% of terrestrial biomes globally. These communities are characterized by a large allocation to belowground biomass, often exceeding its aboveground counterpart. However, this biomass investment cannot be entirely attributed to the acquisitive function of roots. Grassland plants also allocate to non-acquisitive, stem-derived, belowground organs, such as rhizomes. These organs are responsible for the key plant functions of space occupancy, resprouting after damage, and seasonal rest. However, biomass investment to rhizomes has rarely been studied. Here we gathered community-level aboveground and rhizome biomass data for 52 temperate grasslands in Czech Republic (Central Europe), differing in management intensity. We found that rhizome biomass scaled linearly with aboveground biomass, and more intensive management disproportionally (negatively) affected rhizome biomass. This finding may have important implications for the persistence of temperate grassland plants and their provision of ecosystem services (e.g., soil carbon sequestration, soil stabilization) in relation to changing environments.

Functionally distinct assembly of vascular plants colonizing alpine cushions suggests their vulnerability to climate change.

BACKGROUND AND AIMS: Alpine cushion plants can initially facilitate other species during ecological succession, but later on can be negatively affected by their development, especially when beneficiaries possess traits allowing them to overrun their host. This can be reinforced by accelerated warming favouring competitively strong species over cold-adapted cushion specialists. However, little empirical research has addressed the trait-based mechanisms of these interactions. The ecological strategies of plants colonizing the cushion plant Thylacospermum caespitosum (Caryophyllaceae), a dominant pioneer of subnival zones, were studied in the Western Himalayas. METHODS: To assess whether the cushion colonizers are phylogenetically and functionally distinct, 1668 vegetation samples were collected, both in open ground outside the cushions and inside their live and dead canopies, in two mountain ranges, Karakoram and Little Tibet. More than 50 plant traits related to growth, biomass allocation and resource acquisition were measured for target species, and the phylogenetic relationships of these species were studied [or determined]. KEY RESULTS: Species-based trait-environment analysis with phylogenetic correction showed that in both mountain ranges Thylacospermum colonizers are phylogenetically diverse but functionally similar and are functionally different from species preferring bare soil outside cushions. Successful colonizers are fast-growing, clonal graminoids and forbs, penetrating the cushion by rhizomes and stolons. They have higher root-to-shoot ratios, leaf nitrogen and phosphorus concentrations, and soil moisture and nutrient demands, sharing the syndrome of competitive species with broad elevation ranges typical of the late stages of primary succession. In contrast, the species from open ground have traits typical of stress-tolerant specialists from high and dry environments. CONCLUSION: Species colonizing tight cushions of T. caespitosum are competitively strong graminoids and herbaceous perennials from alpine grasslands. Since climate change in the Himalayas favours these species, highly specialized subnival cushion plants may face intense competition and a greater risk of decline in the future.

Rough wave-like heaped overburden promotes establishment of woody vegetation while leveling promotes grasses during unassisted post mining site development.

Geodiversity plays an important role in species establishment during spontaneous succession. At post-mining sites in the Czech Republic in 2003, we established plots in which the surface of the heaped overburden was either kept wave-like or leveled. Based on surveys conducted from 2006 to 2015, leveled plots were increasingly dominated by grasses and herbs (and especially by the grass Calamagrostis epigejos) while the wave-like plots were increasingly dominated by the trees Salix caprea and Betula pendula. In 2015, a detailed survey was conducted of the dominant species. Both S. caprea and B. pendula occurred more often in wave-like plots than in leveled plots; this was particularly true for trees taller than 1 m, which were absent in leveled plots. In wave-like plots, leaf and root biomasses of both woody species were higher on the wave slopes than on the wave depressions. Nitrogen content was higher but content stress indicating proline in leaves of S. caprea was lower in wave-like plots than in leveled plots. In wave-like plots, both woody species occurred mainly on wave slopes but C. epigejos occurred mainly in the depressions. We speculate that trees were more abundant in wave-like plots than in leveled plots because the waves trapped tree seeds and snow and because the soil porosity was greater in wave-like than in leveled plots. Grasses may have preferred the leveled plots because soil porosity was lower and clay content was higher in leveled than in wave-like plots.

Applying the dark diversity concept to nature conservation.

Linking diversity to biological processes is central for developing informed and effective conservation decisions. Unfortunately, observable patterns provide only a proportion of the information necessary for fully understanding the mechanisms and processes acting on a particular population or community. We suggest conservation managers use the often overlooked information relative to species absences and pay particular attention to dark diversity (i.e., a set of species that are absent from a site but that could disperse to and establish there, in other words, the absent portion of a habitat-specific species pool). Together with existing ecological metrics, concepts, and conservation tools, dark diversity can be used to complement and further develop conservation prioritization and management decisions through an understanding of biodiversity relativized by its potential (i.e., its species pool). Furthermore, through a detailed understanding of the population, community, and functional dark diversity, the restoration potential of degraded habitats can be more rigorously assessed and so to the likelihood of successful species invasions. We suggest the application of the dark diversity concept is currently an underappreciated source of information that is valuable for conservation applications ranging from macroscale conservation prioritization to more locally scaled restoration ecology and the management of invasive species.

The effect of lignin photodegradation on decomposability of Calamagrostis epigeios grass litter.

The common grass Calamagrostis epigeions produces a large amount of dead biomass, which remain above the soil surface for many months. In this study, we determined how exposure of dead biomass above the soil affects its subsequent decomposition in soil. Collected dead standing biomass was divided in two parts, the first one (initial litter) was stored in a dark, dry place. The other part was placed in litterbags in the field. The litterbags were located in soil, on the soil surface, or hanging in the air without contact with soil but exposed to the sun and rain. After 1 year of field exposure, litter mass loss and C and N content were measured, and changes in litter chemistry were explored using NMR and thermochemolysis-GC-MS. The potential decomposability of the litter was quantified by burying the litter from the litterbags and the initial litter in soil microcosms and measuring soil respiration. Soil respiration was greater with litter that had been hanging in air than with all other kinds of litter. These finding could not be explained by changes in litter mass or C:N ratio. NMR indicated a decrease in polysaccharides relative to lignin in litter that was buried in soil but not in litter that was placed on soil surface or that was hanging in the air. Thermochemolysis indicated that the syringyl units of the litter lignin were decomposed when the litter was exposed to light. We postulate that photochemical decay of lignin increase decomposability of dead standing biomass.

Data on different seed harvesting methods used in grassland restoration on ex-arable land.

We present data of the grassland restoration experiment performed in the Bílé Karpaty Mts. (White Carpathians, Czech Republic) in dry species-rich meadows. First we harvested seed material in a preserved source meadow (donor site hereafter) by brush harvesting the vegetation once (B1 hereafter), brush harvesting three times during the season (B3 hereafter), and by cutting green hay (GH hereafter). Then we determined the species composition and seed quantity of the harvested material. Furthermore, we transferred the seeds to an experimental site on ex-arable land (receptor site hereafter), and monitored the development of the meadow communities in the following five years. Data are interpreted in: Á-J. Albert, O. Mudrák, I. Jongepierová, K. Fajmon, I. Frei, M. Sevcíková, J. Klimesová, J. Dolezal, Grassland restoration on ex-arable land by transfer of brush-harvested propagules and green hay. Agriculture, Ecosystems & Environment 272 (2019), 74-82.

Measuring size and composition of species pools: a comparison of dark diversity estimates.

Ecological theory and biodiversity conservation have traditionally relied on the number of species recorded at a site, but it is agreed that site richness represents only a portion of the species that can inhabit particular ecological conditions, that is, the habitat-specific species pool. Knowledge of the species pool at different sites enables meaningful comparisons of biodiversity and provides insights into processes of biodiversity formation. Empirical studies, however, are limited due to conceptual and methodological difficulties in determining both the size and composition of the absent part of species pools, the so-called dark diversity. We used >50,000 vegetation plots from 18 types of habitats throughout the Czech Republic, most of which served as a training dataset and 1083 as a subset of test sites. These data were used to compare predicted results from three quantitative methods with those of previously published expert estimates based on species habitat preferences: (1) species co-occurrence based on Beals' smoothing approach; (2) species ecological requirements, with envelopes around community mean Ellenberg values; and (3) species distribution models, using species environmental niches modeled by Biomod software. Dark diversity estimates were compared at both plot and habitat levels, and each method was applied in different configurations. While there were some differences in the results obtained by different methods, particularly at the plot level, there was a clear convergence, especially at the habitat level. The better convergence at the habitat level reflects less variation in local environmental conditions, whereas variation at the plot level is an effect of each particular method. The co-occurrence agreed closest the expert estimate, followed by the method based on species ecological requirements. We conclude that several analytical methods can estimate species pools of given habitats. However, the strengths and weaknesses of different methods need attention, especially when dark diversity is estimated at the plot level.

Vegetation dynamics at the upper elevational limit of vascular plants in Himalaya.

A rapid warming in Himalayas is predicted to increase plant upper distributional limits, vegetation cover and abundance of species adapted to warmer climate. We explored these predictions in NW Himalayas, by revisiting uppermost plant populations after ten years (2003-2013), detailed monitoring of vegetation changes in permanent plots (2009-2012), and age analysis of plants growing from 5500 to 6150 m. Plant traits and microclimate variables were recorded to explain observed vegetation changes. The elevation limits of several species shifted up to 6150 m, about 150 vertical meters above the limit of continuous plant distribution. The plant age analysis corroborated the hypothesis of warming-driven uphill migration. However, the impact of warming interacts with increasing precipitation and physical disturbance. The extreme summer snowfall event in 2010 is likely responsible for substantial decrease in plant cover in both alpine and subnival vegetation and compositional shift towards species preferring wetter habitats. Simultaneous increase in summer temperature and precipitation caused rapid snow melt and, coupled with frequent night frosts, generated multiple freeze-thaw cycles detrimental to subnival plants. Our results suggest that plant species responses to ongoing climate change will not be unidirectional upward range shifts but rather multi-dimensional, species-specific and spatially variable.

Spontaneous vegetation succession at different central European mining sites: a comparison across seres.

We performed detrended correspondence analysis (DCA) ordination to compare seven successional seres running in stone quarries, coal mining spoil heaps, sand and gravel pits, and extracted peatlands in the Czech Republic in central Europe. In total, we obtained 1,187 vegetation samples containing 705 species. These represent various successional stages aged from 1 to 100 years. The successional seres studied were more similar in their species composition in the initial stages, in which synathropic species prevailed, than in later successional stages. This vegetation differentiation was determined especially by local moisture conditions. In most cases, succession led to a woodland, which usually established after approximately 20 years. In very dry or wet places, by contrast, where woody species were limited, often highly valuable, open vegetation developed. Except in the peatlands, the total number of species and the number of target species increased during succession. Participation of invasive aliens was mostly unimportant. Spontaneous vegetation succession generally appears to be an ecologically suitable and cheap way of ecosystem restoration of heavily disturbed sites. It should, therefore, be preferred over technical reclamation.