Spatial scaling of pollen-plant diversity relationship in landscapes with contrasting diversity patterns
We found a significant positive relationship between pollen- and plant richness regardless of differences in plant diversity, landscape structure and environmental conditions between the two study regions. This finding represents a major step stone towards more accurate paleoecological reconstructions of plant diversity in temperate Central Europe, as previous studies on this topic have mostly been conducted in boreal and boreal-nemoral zones8,11, in high mountain habitats10 or in southern Europe9,12.Methodological differences e.g., in diversity indices, data transformations or sample sizes used make comparison between studies difficult. Nevertheless, the strongest relationships seem to be found when habitats with contrasting patterns of plant diversity are compared, such as forests and alpine vegetation7 or forests, peatlands and grasslands11. Also in our study, we found the strongest correlations when complete datasets combining forested and open habitats were analysed together for both study regions. As it is well known that plant richness is generally lower in forests than in open landscapes across temperate and boreal regions28, this finding may seem rather trivial. However, it is important for paleoecological reconstruction because Holocene changes in diversity in temperate regions were largely driven by changes in the relative abundance of major habitat types (such as forests, grasslands, wetlands and man-made habitats), and not just by changes in species richness within these habitats5,6.Regarding individual habitats, the pollen-plant diversity relationship is often rather strong and significant in grasslands and other open habitats8,11; for example the WCM open-habitat subset in this study. Open habitats are generally richer in species, thus providing a longer gradient of species richness compensating for the taxonomical imprecision of the pollen analysis. In forested sites with less species, we found mostly non-significant relationships. Moreover, two other factors may play a role.First, high pollen productivity of trees biases the diversity relationship according to the studies from northern Europe16. However, a study from an elevational transect in southern Norway showed that the strongest bias in representation occurs only in the boreal forest biome, which is dominated by high pollen producers10. Our dominant vegetation component, Picea and Quercus, have intermediate to high pollen productivity (2–2.5), whereas true high pollen producers such as Alnus and Betula ( > 3) are less abundant in our study area (Supplementary Fig. S2). Adjustment of pollen counts by PPEs led to stronger relationship between pollen and floristic richness only in the WCM open-habitat subset (Supplementary Fig. S4).Second, interception of pollen by the tree canopies29 and subsequent washout to the forest floor affects the diversity relationship of forest sites more than pollen productivity. This noise described also as a vegetation filtering30 can be illustrated in our dataset by pollen of long-distance transport from Ambrosia artemisiifolia-type, which has the closest source populations ca. 50 km south-eastwards from WCM region31; or pollen of Artemisia, growing in open habitats. Both pollen taxa are more abundant in the forest than in open sites (Supplementary Fig. S3).Regarding the application of these results for the interpretation of fossil record, we suggest to consider only marked changes of pollen richness in the past and to avoid overinterpretation of small differences, as the non-significant relationships obtained in both forest datasets suggest some limitations of the method.We showed that the pollen-plant diversity relationship may be at least partly disentangled by knowing the exact spatial position of plant species in broader surroundings of the pollen sampling sites. Changes in the relationship with changing spatial scale are largely driven by the numbers of species newly appearing as the radius of surveyed area increases, especially as new habitats are added (Fig. 5, Supplementary Fig. S5). Remarkably, in the BMH region it increases with distance, whereas the opposite trend was observed in the WCM region. This discrepancy may be explained by non-uniform richness patterns in different habitats and by different landscape structure (i.e. spatial arrangement of different habitats) in the two study regions.At open-habitat sites in the WCM area, most species generally appeared within the first 40 m. This observation is consistent with the knowledge of extremely high fine-scale plant diversity in the local steppic meadows, where a substantial portion of the species pool occurs on a scale of tens of square meters32. Moreover, the grain size of the habitat mosaic in the WCM region is finer than in the BMH region. Therefore, the closest pollen-plant diversity relationship across habitats in the WCM region is achieved over shorter distances. Although habitats such as built-up areas and roads occurring at distances greater than 40 m may be species-rich and compositionally different from the grasslands and forests, it appears that high fine-scale plant diversity (in our case in WCM open-habitat subset) limits the influence of the surrounding landscape on pollen richness and reduces the source area of pollen richness. Several studies of the relevant source area of pollen report analogous results33,34,35. A weakening relationship between pollen diversity and plant diversity with distance has also been observed in the Mediterranean region9, although their interpretations are limited by field survey methodology.The appearance of open habitats within forests led to the increase of species numbers and the local maxima of adjusted R2 in both regions. While in the BMH forest the appearance of forest roads at about 70 m was crucial, meadows and orchards at about 250 m played a similar role in the WCM forest subset. In the WCM open-habitat subset diversity patterns in the first tens of metres were crucial, while in the BMH open-habitat subset increased correlation of floristic and pollen richness appeared only at 400 and 550 m; at this distance many species appeared due to the frequent transition of meadow complexes to shrubby habitats and built-up areas. Also other studies from semi-open landscapes found a high correlation between pollen richness and landscape openness17,26,27.Estimating the source area of pollen variance as a regression of pollen and floristic variance implies that the resulting distance of 100–250 m represents all datasets. Although they differ in species richness, openness and habitats, the relationship between variances is fairly linear. The exception is the WCM open-habitat subset suggesting that the spatial scale at which the pollen variance corresponds to the floristic variance cannot be generalized.The strong effect of high pollen richness in the WCM open-habitat subset is also visible in the comparison of pollen and floristic variance. At 150 m, the WCM open-habitat subset had much lower floristic variance than the other subsets. Floristic variance in this subset corresponding to the pollen variance and the pattern of the other datasets lay at 6 m (Fig. 6b). Again, this may be caused by the high fine-scale diversity of the meadows, which include most pollen types present in the surrounding landscape. Only a few new species appeared in broader surroundings and at 150 m, WCM open habitats are more similar than other analysed habitats. The fact that extremely high alpha diversity is compensated by low beta diversity has already been reported from the open habitats of the White Carpathians36. The linearity and the significance of the variance relationship within the rest of the datasets indicate robustness and possible applicability to a variety of fossil records.The mechanism of establishing the source area of pollen variance was similar to that mentioned for the source area of pollen richness. The appearance of new habitats with new species (Fig. 5) like open habitat for forest sites (WCM forest subset) or built-up areas for open sites (BMH open-habitat subset), caused small to negligible increases of floristic variance. Moreover, the high yet insignificant relationship of the variances at the distance between 250 and 600 m (Fig. 6a) corresponds to the distance of the second range of fit between floristic and pollen richness (Fig. 4a).Beta diversity, understood as directional turnover (temporal or spatial), is becoming more frequently used in pollen analysis22,24 than beta diversity as a non-directional variation. According to Nieto-Lugilde et al.25 pollen-based turnover correlates with forest-inventory-based turnover. We extend this finding from woody taxa to all species and from directional turnover to non-directional variance. Moreover, forest sites with high contributions to pollen beta diversity also show an increased contribution to floristic beta diversity (Fig. 4b).The reference data on plant diversity report 1477 species in 15 mapping squares covered by our survey for the BMH region and 2045 species in 14 squares for the WCM region37. It means that we recorded 54.1 and 53.7%, respectively, of the known regional species pool in the two regions. We consider this as a rather good result and the close agreement in representativeness between the two regions speaks for consistency in data quality between the datasets. We advise that future studies covering wider areas and various biomes should preferentially use high-quality floristic data collected in targeted field surveys rather than database data or data from simplified field surveys. Only then we will be able to understand the pollen-plant diversity relationships more realistically and in a spatially explicit manner.In order to interpret fossil pollen richness in the light of our present results, we need to consider landscape openness, which can be roughly inferred from the ratio of arboreal and non-arboreal pollen. Variation of pollen richness during the forest phases of the records should be interpreted more carefully, especially in cases of low variation. In all other cases, the pollen richness is significantly linked to the plant richness within a distance of ten to several hundreds of meters, depending on the distance of the expected species-rich patches. More
