Martin Kopecký

Non‐random extinctions dominate plant community changes in abandoned coppices

1. The plant community structure of European lowland forests has changed dramatically in the 20th century, leading to biodiversity decline at various spatial scales. However, due to methodological difficulties associated with simultaneous changes in species diversity and composition, ecological processes behind the changes are still poorly understood. 2. We analysed temporal changes in forest plant community after the mid-20th century abandonment of coppicing in a typical Central European forest, which had been managed as coppice for centuries. We used 122 semi-permanent plots first surveyed in the 1950s shortly after the last coppicing and again in the 2000s after half a century of natural succession. We used a novel Temporal Nestedness Analysis to disentangle the immigration and extinction processes underlying temporal changes in community structure and tested whether species gains and losses were ecologically random. 3. The studied vegetation has shifted from the species-rich assemblages of a relatively open and low-nutrient forest towards the impoverished flora of a closed-canopy forest dominated by a few shade-adapted species. The significant reduction of beta diversity, i.e. compositional heterogeneity among plots, indicated taxonomic homogenization of the forest understorey. Temporal species turnover was only a minor component of the community change and recent assemblages are nested subsets of the former ones. Ecologically non-random extinctions dominated these changes. Light-demanding species with a persistent seed-bank were the most prone to extinction, while species with high specific leaf area substantially increased in frequency. 4. Synthesis and applications. The dominant process after the abandonment of coppicing was the ecologically non-random extinction of light-demanding species leading to an impoverished, temporally nested plant community structure. This development is typical for many abandoned lowland coppice forests and poses a significant threat to forest biodiversity in Europe. If forestry and conservation policies continue to prefer closed-canopy stands, many endangered species are likely to pay their extinction debts. To restore declining or even locally extinct species, canopy opening in abandoned coppices is urgently needed.

Vegetation resurvey is robust to plot location uncertainty

AIM Resurveys of historical vegetation plots are increasingly used for the assessment of decadal changes in plant species diversity and composition. However, historical plots are usually relocated only approximately. This potentially inflates temporal changes and undermines results. LOCATION Temperate deciduous forests in Central Europe. METHODS To explore if robust conclusions can be drawn from resurvey studies despite location uncertainty, we compared temporal changes in species richness, frequency, composition and compositional heterogeneity between exactly and approximately relocated plots. We hypothesized that compositional changes should be lower and changes in species richness should be less variable on exactly relocated plots, because pseudo-turnover inflates temporal changes on approximately relocated plots. RESULTS Temporal changes in species richness were not more variable and temporal changes in species composition and compositional heterogeneity were not higher on approximately relocated plots. Moreover, the frequency of individual species changed similarly on both plot types. MAIN CONCLUSIONS The resurvey of historical vegetation plots is robust to uncertainty in original plot location and, when done properly, provides reliable evidence of decadal changes in plant communities. This provides important background for other resurvey studies and opens up the possibility for large-scale assessments of plant community change.

Testing the Stress-Gradient Hypothesis at the Roof of the World: Effects of the Cushion Plant Thylacospermum caespitosum on Species Assemblages

Many cushion plants ameliorate the harsh environment they inhabit in alpine ecosystems and act as nurse plants, with significantly more species growing within their canopy than outside. These facilitative interactions seem to increase with the abiotic stress, thus supporting the stress-gradient hypothesis. We tested this prediction by exploring the association pattern of vascular plants with the dominant cushion plant Thylacospermum caespitosum (Caryophyllaceae) in the arid Trans-Himalaya, where vascular plants occur at one of the highest worldwide elevational limits. We compared plant composition between 1112 pair-plots placed both inside cushions and in surrounding open areas, in communities from cold steppes to subnival zones along two elevational gradients (East Karakoram: 4850–5250 m and Little Tibet: 5350–5850 m). We used PERMANOVA to assess differences in species composition, Friedman-based permutation tests to determine individual species habitat preferences, species-area curves to assess whether interactions are size-dependent and competitive intensity and importance indices to evaluate plant-plant interactions. No indications for net facilitation were found along the elevation gradients. The open areas were not only richer in species, but not a single species preferred to grow exclusively inside cushions, while 39–60% of 56 species detected had a significant preference for the habitat outside cushions. Across the entire elevation range of T. caespitosum, the number and abundance of species were greater outside cushions, suggesting that competitive rather than facilitative interactions prevail. This was supported by lower soil nutrient contents inside cushions, indicating a resource preemption, and little thermal amelioration at the extreme end of the elevational gradient. We attribute the negative associations to competition for limited resources, a strong environmental filter in arid high-mountain environment selecting the stress-tolerant species that do not rely on help from other plants during their life cycle and to the fact the cushions do not provide a better microhabitat to grow in.

Tree-rings mirror management legacy: dramatic response of standard oaks to past coppicing in Central Europe.

Background Coppicing was one of the most important forest management systems in Europe documented in prehistory as well as in the Middle Ages. However, coppicing was gradually abandoned by the mid-20th century, which has altered the ecosystem structure, diversity and function of coppice woods. Methodology/Principal Findings Our aim was to disentangle factors shaping the historical growth dynamics of oak standards (i.e. mature trees growing through several coppice cycles) in a former coppice-with-standards in Central Europe. Specifically, we tried to detect historical coppicing events from tree-rings of oak standards, to link coppicing events with the recruitment of mature oaks, and to determine the effects of neighbouring trees on the stem increment of oak standards. Large peaks in radial growth found for the periods 1895–1899 and 1935–1939 matched with historical records of coppice harvests. After coppicing, the number of newly recruited oak standards markedly grew in comparison with the preceding or following periods. The last significant recruitment of oak standards was after the 1930s following the last regular coppicing event. The diameter increment of oak standards from 1953 to 2003 was negatively correlated with competition indices, suggesting that neighbouring trees (mainly resprouting coppiced Tilia platyphyllos) partly suppressed the growth of oak standards. Our results showed that improved light conditions following historical coppicing events caused significant increase in pulses of radial growth and most probably maintained oak recruitment. Conclusions/Significance Our historical perspective carries important implications for oak management in Central Europe and elsewhere. Relatively intense cutting creating open canopy woodlands, either as in the coppicing system or in the form of selective cutting, is needed to achieve significant radial growth in mature oaks. It is also critical for the successful regeneration and long-term maintenance of oak populations.

Experimental restoration of coppice-with-standards: Response of understorey vegetation from the conservation perspective

A substantial part of European lowland woodlands was managed as coppices or wood pastures for millennia. However, traditional management forms were almost completely abandoned in Central Europe by the middle of the 20th century. Combined with the effects of nitrogen deposition and herbivore pressure, shifts in management resulted in biodiversity loss affecting particularly light-demanding oligotrophic plant species. Experimental thinning was applied in a former oak coppice-with-standards in an attempt to restore vanishing understorey plant communities. Two levels of thinning intensity and zero management as control were used on 90 plots. Ten years after the treatment, significant changes in species composition and diversity were observed in heavily thinned plots, while moderate thinning had mostly insignificant effects. Light-demanding oligotrophic species significantly increased, indicating positive consequences of restoration. However, heavy thinning also brought about the expansion of native ruderal species. Alien species remained unchanged. We conclude that the restoration of coppice-with-standards can be an efficient tool to support vanishing light-demanding woodland species. Combined with biodiversity benefits, the increasing demand for biofuel may contribute to the renaissance of traditional management forms in forestry.

Resurveying historical vegetation data – opportunities and challenges

Background Resurveying historical vegetation plots has become more and more popular in recent years as it provides a unique opportunity to estimate vegetation and environmental changes over the past decades. Most historical plots, however, are not permanently marked and uncertainty in plot location, in addition to observer bias and seasonal bias, may add significant error to temporal change. These errors may have major implications for the reliability of studies on long-term environmental change and deserve closer attention of vegetation ecologists. Material & Methods Vegetation data obtained from the resurveying of non-permanently marked plots are assessed for their potential to study environmental-change effects on plant communities and the challenges the use of such data have to meet. We describe the properties of vegetation resurveys distinguishing basic types of plots according to relocation error, and we highlight the potential of such data types for studying vegetation dynamics and their drivers. Finally, we summarise the challenges and limitations of resurveying non-permanently marked vegetation plots for different purposes in environmental change research. Results and Conclusions Resampling error is caused by three main independent sources of error: error caused by plot relocation, observer bias, and seasonality bias. For relocation error, vegetation plots can be divided into permanent and non-permanent plots, while the latter are further divided into quasi-permanent (with approximate relocation) and non-traceable (with random relocation within a sampled area) plots. To reduce the inherent sources of error in resurvey data, the following precautions should be followed: (i) resurvey historical vegetation plots whose approximate plot location within a study area is known; (ii) consider all information available from historical studies in order to keep plot relocation errors low; (iii) resurvey at times of the year when vegetation development is comparable to the historical survey to control for seasonal variability in vegetation; (iv) keep a high level of experience of the observers to keep observer bias low; and (v) edit and standardise datasets before analyses.

Community structure of soil phototrophs along environmental gradients in arid Himalaya

The well-developed biological soil crusts cover up to 40% of the soil surface in the alpine and subnival zones of the Tibetan Plateau, accounting for a vast area of Asia. We investigated the diversity and biomass of the phototrophic part (Cyanobacteria) of the microbial community inhabiting biological soil crusts and uncrusted soils in their surroundings on the elevation gradient of 5200-5900 m a.s.l. The influence of soil physico-chemical properties on phototrophs was studied. The ability of high-altitude phototrophs to fix molecular nitrogen was also determined under laboratory conditions. The biological soil crust phototroph community did not differ from that living in uncrusted soil in terms of the species composition, but the biomass is three-to-five times higher. An increasing trend in the cyanobacterial biomass from the biological soil crusts with elevation was observed, with the genera Nostoc spp., Microcoleus vaginatus and Phormidium spp. contributing to this increase. Based on the laboratory experiments, the highest nitrogenase activity was recorded in the middle elevations, and the rate of nitrogen fixation was not correlated with the cyanobacterial biomass.

Combining Biodiversity Resurveys across Regions to Advance Global Change Research

More and more ecologists have started to resurvey communities sampled in earlier decades to determine long-term shifts in community composition and infer the likely drivers of the ecological changes observed. However, to assess the relative importance of and interactions among multiple drivers, joint analyses of resurvey data from many regions spanning large environmental gradients are needed. In this article, we illustrate how combining resurvey data from multiple regions can increase the likelihood of driver orthogonality within the design and show that repeatedly surveying across multiple regions provides higher representativeness and comprehensiveness, allowing us to answer more completely a broader range of questions. We provide general guidelines to aid the implementation of multiregion resurvey databases. In so doing, we aim to encourage resurvey database development across other community types and biomes to advance global environmental change research.

Vascular plants at extreme elevations in eastern Ladakh, northwest Himalayas

Background: Due to the dry continental climate, the mountains of eastern Ladakh are unglaciated up to 6200–6400 m, with relatively large areas of developed soils between 5600 and 6000 m covered by sparse subnival vegetation. However, there are no studies on the composition of plant assemblages from such extreme elevations, their microclimates, vertical distributions and adaptive strategies. Aims: The subnival vegetation was described and the relationship between microclimate, species distribution and species functional traits was analysed. Methods: In total, 481 vegetation samples from 91 permanent plots, a floristic database of Ladakh and extensive microclimate measurements were used. Measurements of 15 functional traits were made and their relationship with species distribution between 4600 and 6150 m was tested. Results: The subnival zone was characterised by extreme diurnal temperature fluctuations, a short growing season (between 88 and 153 days) and low soil temperature during the growing season (between 2.9 °C and 5.9 °C). It hosted 67 species, mainly hemicryptophytes, and ranged from ca. 5600 m to the highest known occurrence of vascular plants in the region (6150 m). The most common plant families were Brassicaceae, Asteraceae, Poaceae, Fabaceae and Cyperaceae. Subnival specialists with narrow elevational ranges represented 42% of the flora; these species were shorter, had relatively higher water content and water-use efficiency and contained more nutrients and soluble carbohydrates than species with a wider elevational range. Conclusions: The subnival vegetation of eastern Ladakh is dominated by generalist species with wide vertical ranges and not by high-elevation specialists. These findings, in view of the vast unglaciated areas available for range extension, suggest a relatively high resilience of the subnival flora to climate change in this region.

Gardening in the zone of death: an experimental assessment of the absolute elevation limit of vascular plants.

Vascular plants in the western Tibetan Plateau reach 6000 m-the highest elevation on Earth. Due to the significant warming of the region, plant ranges are expected to shift upwards. However, factors governing maximum elevational limits of plant are unclear. To experimentally assess these factors, we transplanted 12 species from 5750 m to 5900 m (upper edge of vegetation) and 6100 m (beyond range) and monitored their survival for six years. In the first three years (2009–2012), there were plants surviving beyond the regional upper limit of vegetation. This supports the hypothesis of dispersal and/or recruitment limitation. Substantial warming, recorded in-situ during this period, very likely facilitated the survival. The survival was ecologically a non-random process, species better adapted to repeated soil freezing and thawing survived significantly better. No species have survived at 6100 m since 2013, probably due to the extreme snowfall in 2013. In conclusion, apart from the minimum heat requirements, our results show that episodic climatic events are decisive determinants of upper elevational limits of vascular plants.