Meso-scale environmental heterogeneity drives plant trait distributions in fragmented dry grasslands.

Environmental heterogeneity shapes the patterns of resources and limiting factors and therefore can be an important driver of plant community composition through the selection of the most adaptive functional traits. In this study, we explored plant trait-environment relationships in environmentally heterogeneous microsite complexes at the meso-scale (few meters), and used ancient Bulgarian and Hungarian burial mounds covered by dry grasslands as a model habitat. We assessed within-site trait variability typical of certain microsites with different combinations of environmental parameters (mound slopes with different aspects, mound tops, and surrounding plain grasslands) using a dataset of 480 vegetation plots. For this we calculated community-weighted means (CWMs) and abundance models. We found that despite their small size, the vegetation on mounds was characterized by different sets of functional traits (higher canopy, higher level of clonality, and heavier seeds) compared to the plain grasslands. North-facing slopes with mild environmental conditions were characterized by perennial species with light seeds, short flowering period, and a high proportion of dwarf shrubs sharply contrasted from the plain grasslands and from the south-facing slopes and mound tops with harsh environmental conditions. Patterns predicted by CWMs and abundance models differed in the case of certain traits (perenniality, canopy height, and leaf dry matter content), suggesting that environmental factors do not necessarily affect trait optima directly, but influence them indirectly through correlating traits. Due to the large relative differences in environmental parameters, contrasts in trait composition among microsites were mostly consistent and independent from the macroclimate. Mounds with high environmental heterogeneity can considerably increase variability in plant functional traits and ecological strategies at the site and landscape levels. The large trait variation on topographically heterogeneous landscape features can increase community resilience against climate change or stochastic disturbances, which underlines their conservation importance.

First Survey of the Vascular and Cryptogam Flora on Bulgaria's Ancient Mounds.

This work represents the first study of the floristic diversity on Bulgaria's ancient mounds. The objective of this research was to assess the importance of the mounds for the preservation of the native vascular and cryptogam flora. Our sampling design included 111 ancient mounds distributed throughout the country. We recorded a total of 1059 vascular plants, 58 bryophytes and 61 lichen taxa. Despite their small area, the mounds were shown to preserve nearly a quarter of the Bulgarian flora. The vegetation cover on the mounds included 61% perennials indicating a long-term persistence and stability. The majority (98%) of the established vascular plants were native species. Although the conservation significance of the vascular plant species were not common, we recorded 2 critically endangered, 9 endangered and 14 Balkan endemics during the present study. The lichen Arthopyrenia salicis was recorded for the first time in Bulgaria and a new locality of the rare bryophyte Ceratodon conicus was discovered. The established compositional difference between plots from the northern and southern slopes of the mounds (88.95%) is a testament to the high local habitat diversity. The prevalence of species characteristic for Festuco-Brometea suggests that the mounds preserve fragments of native grasslands and steppes. The variation in cover of agricultural and other human modified areas in the mounds' immediate surroundings did not substantially affect their species richness. We argue that the ancient mounds should be taken into consideration in future green space planning.

Linking environmental heterogeneity and plant diversity: The ecological role of small natural features in homogeneous landscapes.

Small natural features (SNFs), such as road verges, midfield islets, rocky outcrops and ancient burial mounds, provide safe havens for species of natural habitats in human-modified landscapes; therefore, their great ecological importance is in contrast to their small size. SNFs often have a high topographical heterogeneity and abiotic conditions, which differ from their surroundings; therefore, they provide a unique opportunity for establishing links between environmental heterogeneity (EH) and biodiversity. However, no study has so far investigated the EH components of topographically heterogeneous SNFs in a comprehensive framework, by linking environmental and biotic parameters. To fill this knowledge gap, we evaluated the EH components and their effect on biodiversity on ancient mounds covered by semi-natural grasslands in the Pannonian (Hungary) and Continental (Bulgaria) biogeographical regions. We designated 16 study sites, each containing a few-metre-high mounds with five microsites (top, north-, east-, south- and west-facing slopes) and a nearby plain grassland. At each microsite, we measured soil moisture, soil chemical properties, solar radiation and microclimate; and recorded the cover of vascular plants in a total of 480 plots. On the mounds, topographical heterogeneity was associated with sharp differences in microclimate and soil properties. Besides the contrast between mild north-facing and harsh south-facing slopes, east- and west-facing slopes also sustained unique microsites characterised by dynamic diurnal changes in air temperature and vapour pressure deficit. Various combinations of the EH components resulted in unique plant species compositions within the microsites, and supported the co-occurrence of species typical of contrasting habitat types, even within a couple of metres. By combining high-resolution measurements of abiotic factors with fine-scale vegetation sampling, our study provides evidence that widespread SNFs with complex topography harbour several grassland-specialist plant species and introduce a high level of EH to otherwise homogeneous plain landscapes, which cover one third of the global land area.

Invader presence disrupts the stabilizing effect of species richness in plant community recovery after drought.

Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity-stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity-stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non-native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader × biodiversity × drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi-natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning.

Species richness effects on grassland recovery from drought depend on community productivity in a multisite experiment.

Biodiversity can buffer ecosystem functioning against extreme climatic events, but few experiments have explicitly tested this. Here, we present the first multisite biodiversity × drought manipulation experiment to examine drought resistance and recovery at five temperate and Mediterranean grassland sites. Aboveground biomass production declined by 30% due to experimental drought (standardised local extremity by rainfall exclusion for 72-98 consecutive days). Species richness did not affect resistance but promoted recovery. Recovery was only positively affected by species richness in low-productive communities, with most diverse communities even showing overcompensation. This positive diversity effect could be linked to asynchrony of species responses. Our results suggest that a more context-dependent view considering the nature of the climatic disturbance as well as the productivity of the studied system will help identify under which circumstances biodiversity promotes drought resistance or recovery. Stability of biomass production can generally be expected to decrease with biodiversity loss and climate change.