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For 29 consecutive years, the populations of three species of brown frogs, Rana arvalis, R. dalmatina and R. temporaria, were monitored in an open area in South Zealand, Denmark, with no direct influence of agriculture. Population sizes were recorded by counting egg clumps and showed large variations from year to year. The total population of R. arvalis differed by a factor of 100 between the years with the lowest and the highest numbers. A total of 19 initially unoccupied suitable waterbodies could potentially be colonised by the frogs. Rana dalmatina colonised all of the ponds, mostly in the very first year of existence. Rana arvalis colonised 17 ponds after an average of 10.5 years. Rana temporaria colonised eight ponds after an average of 13.4 years. Colonisation by R. dalmatina was independent of changes in total population size, whereas colonisation by R. arvalis predominantly occurred in years with considerable population increases. The results are discussed in relation to the movement patterns and philopatry of juvenile frogs of the three species. Juvenile R. dalmatina disperse far from the breeding site, but most individuals return to their natal site. This allows the species to be an efficient coloniser of new waterbodies and, at the same time, to have stable occurrence at the original site. The two other species show a more erratic type of dispersal and especially R. temporaria often shifts breeding site from year to year.

Abstract Background In light of the biodiversity crisis and our limited ability to explain variation in biodiversity, tools to quantify spatial and temporal variation in biodiversity and its underlying drivers are critically needed. Inspired by the recently published ecospace framework, we developed and tested a sampling design for environmental and biotic mapping. We selected 130 study sites (40 × 40 m) across Denmark using stratified random sampling along the major environmental gradients underlying biotic variation. Using standardized methods, we collected site species data on vascular plants, bryophytes, macrofungi, lichens, gastropods and arthropods. To evaluate sampling efficiency, we calculated regional coverage (relative to the known species number per taxonomic group), and site scale coverage (i.e., sample completeness per taxonomic group at each site). To extend taxonomic coverage to organisms that are difficult to sample by classical inventories (e.g., nematodes and non-fruiting fungi), we collected soil for metabarcoding. Finally, to assess site conditions, we mapped abiotic conditions, biotic resources and habitat continuity. Results Despite the 130 study sites only covering a minute fraction (0.0005%) of the total Danish terrestrial area, we found 1774 species of macrofungi (54% of the Danish fungal species pool), 663 vascular plant species (42%), 254 bryophyte species (41%) and 200 lichen species (19%). For arthropods, we observed 330 spider species (58%), 123 carabid beetle species (37%) and 99 hoverfly species (33%). Overall, sample coverage was remarkably high across taxonomic groups and sufficient to capture substantial spatial variation in biodiversity across Denmark. This inventory is nationally unprecedented in detail and resulted in the discovery of 143 species with no previous record for Denmark. Comparison between plant OTUs detected in soil DNA and observed plant species confirmed the usefulness of carefully curated environmental DNA-data. Correlations among species richness for taxonomic groups were predominantly positive, but did not correlate well among all taxa suggesting differential and complex biotic responses to environmental variation. Conclusions We successfully and adequately sampled a wide range of diverse taxa along key environmental gradients across Denmark using an approach that includes multi-taxon biodiversity assessment and ecospace mapping. Our approach is applicable to assessments of biodiversity in other regions and biomes where species are structured along environmental gradient.