Abstract
The Balkan Peninsula is considered one of the most important centres of orchid diversity in Europe. However, the patterns of orchid species richness in the Central Balkans have not been sufficiently studied so far. The aim of this study was, therefore, to identify the centres of orchid diversity and the factors that influence the spatial variation in orchid species richness in the Central Balkans. For the analyses, the area of the Central Balkans was divided into 10 × 10 km grid cells. The environmental variables determined for each grid cell and used in the analyses were altitude, bioclimatic variables, geological substrates and habitat types. A random forest (RF) analysis was used to identify the environmental predictors most strongly associated with species richness. In addition to the total number of taxa, orchids with three belowground organ types were analysed separately: (a) rhizomatous orchids, (b) orchids with palmately lobed and fusiform tubers ("palmate tuberous orchids") and (c) orchids with spherical or ovoid tubers ("ovoid tuberous orchids"). In the Central Balkans, 54 orchid species and subspecies have been recorded, and the most important centres of diversity are the Tara, Zvijezda, Jadovnik and Zlatar Mountains and the Ovčar-Kablar Gorge. In general, two groups of grid cells with the largest number of orchid taxa, i.e., hotspots, stood out: (1) grid cells with a large altitudinal range and (2) grid cells occupied by gorges and ravines. The most important gradients influencing orchid species richness are specific habitat types and altitudinal ranges, while climatic factors and geological substrates are less important. The most important factors affecting the richness of total and rhizomatous orchids are altitudinal range and habitat types (Abieti-Fagenion, Ostryo-Carpinion orientalis and Pinion nigrae forests), highlighting the important role of habitat heterogeneity. The maximum altitude, percentage of Abieti-Fagenion and Vaccinio-Picetea forests and the minimum value of the mean temperature of the driest quarter are the most important factors for determining the richness of palmate tuberous orchids, whereas the percentage of xero-thermophilous habitat types (Ostryo-Carpinion orientalis, Asplenietea trichomanis and Pinion nigrae) has the greatest influence on the richness of ovoid tuberous orchids. These results confirm the hypothesis concerning the origin and development of underground organs in orchids, emphasising that palmate tuberous orchids are best adapted to cold and humid habitat conditions, whereas ovoid tuberous orchids have the ability to grow in habitats with very warm and dry conditions. This study provides a good basis for better orchid conservation planning and underlines the importance of belowground strategies as a feature of orchid life history that should be considered when studying patterns of orchid diversity.