Orchid pollination is traditionally considered to rely on intact pollinarium transfer by animal vectors. Species lacking a functional viscidium are generally classified as obligately autogamous. In this study, we investigated the reproductive biology of Epipactis bugacensis, a taxon long regarded as strictly self-pollinating. Floral visitor activity was assessed through repeated field observations, and pollinator dependence was tested using a pollinator-exclusion (net-covering) experiment at two Hungarian populations, combined with measurements of fruit set, capsule volume, seed number, and seed density. We documented a previously unreported pollen-transfer mechanism in E. bugacensis, whereby halictid bees fragment pollinia and transfer these fragments in their scopa to neighboring flowers enabling geitonogamous deposition and suggesting the potential for xenogamous pollen transfer. Other visitor taxa showed no evidence of effective pollen transport. Mesh coverage increased fruit set, capsule volume, and seed number, while seed density remained unchanged. Reproductive output declined from basal to apical positions along flowering shoots, revealing strong internal resource-allocation constraints. Overall, E. bugacensis is predominantly self-pollinating but not strictly obligate autogamous, and its reproductive success is governed primarily by microhabitat quality rather than pollinator availability.

Microrefugia play a key role in facilitating the persistence of biodiversity during climate change. Many occur in topographically complex landscapes shaped by various disturbances, but we know little about how the combined effects of topography and disturbance affect the capacity of refugia to support biodiversity. To better understand this, we inventoried taxonomic richness across four biological groups (soil microbiota, vascular plants, terrestrial snails, and ants), as well as climatic and soil conditions, in different microhabitats (south-facing slopes, north-facing slopes, and bottoms) of topographic depressions (dolines) and on the surrounding plateaus. Unique species assemblages and cooler, moister microclimatic conditions in dolines supported their importance as biodiversity hotspots and microrefugia. Relationships between indicators of disturbances (anthropogenic: historical logging; natural: canopy gaps) and species richness differed, depending on the biological group and microhabitat. While most biological groups seemingly recovered within 50 years following clear-felling, plants did not, highlighting the persistent impact of anthropogenic disturbances on refugial capacity. Plants were also the only group that displayed a significant response to the presence of small canopy gaps at doline bottoms, which promoted the occurrence of specific plant species. All biological groups displayed some response to microhabitats, although these responses differed among taxa. Therefore, high environmental heterogeneity appears to help facilitate the role of dolines as biodiversity hotspots and microrefugia. We conclude that the direction and magnitude of the effects of disturbances and topography are taxon-specific, due to species-specific responses to microenvironmental conditions. Disturbance history is an important consideration when identifying refugia for climate change management.