Abstract (eng)
Insect decline has become a widely discussed topic in science and mainstream media in recent years. But besides the general conclusion that insects are disappearing, we often still lack in information what drivers are causing this phenomenon. Multiple predictors are in discussion to negatively affect insect biodiversity. But are some factors more important than others? And how do these multiple predictors influence the different aspects of biodiversity? These are the questions I want to address in my thesis.
I chose 60 study sites situated within two Mediterranean coastal forest reserves in North-eastern Italy – Pineta san Vitale (PsV) and Pineta di Classe (PdC) – that once formed one contiguous coastal forest but are today isolated from each other and surrounded by anthropogenically modified landscapes. By analyzing multiple possible drivers, my aim was to achieve a deeper understanding of how biotic communities in isolated conservation areas are influenced by multiple environmental gradients and how biodiversity in such reserves could be better preserved in future.
First, I analyzed how vegetation is influenced by environmental predictors. Plants, building the first trophic level, are the basis of every terrestrial food-web and therefore are essentially determining, how a habitat looks like and which species of other groups of organisms may colonize it. This is even more important for phytophagous organisms that have direct trophic links with often rather specific host plants. The composition and species diversity of plants can be driven by local circumstances, such as abiotic soil conditions, and landscape-scale attributes, such as land use in the surroundings of a reserve. In the case of my 60 study sites, which all were located within mixed oak and pine forest, plant species richness and functional dispersion at the local site level were positively affected by the proximity to water canals. However, near these canals, there were also more salt tolerating plants and the vegetation was characterized by higher mean nutrient indicator values. Furthermore, mean nutrient indicator values inside the reserves were increasing with higher proportions of agricultural land around the conservation areas. These findings indicate nutrient spill-over as well as salt water intrusion, which both are mainly caused by human actions outside the proper reserves. The amount of anthropogenically modified areas in the surroundings also led to the disappearance of plants with more restricted distributional ranges, being replaced by widely distributed ones. So, landscape-scale human actions might indirectly drive biotic homogenization of plant communities inside conservation areas.
Based on these results, the next question was: Do these findings translate to the next trophic level? How are moths – as an example for a species-rich, mobile, predominately phytophagous insect group – influenced by local and landscape-scale predictors? From 2015 to 2017, I sampled moths at all the 60 locations, using automated light traps. Every site was sampled twice, one time in early summer (June) and one time in late summer (August). In total, 23870 individuals of 392 species (comprising 32 families) were available for analysis.
The first aspect of moth diversity I examined was small-scaled α-diversity and the number of caught individuals per site. To describe the environmental conditions that possibly might affect moth diversity I used 14 local site descriptors, like plant diversity and forest structure, and another 14 landscape-scale factors, like landscape diversity and the proportion of modified areas around light-trapping sites. These 28 factors were condensed through a Principal Component Analysis, resulting in five local and five landscape-scale PC-axes being used as predictors in linear mixed effects models. Small-scaled α-diversity patterns of moths were mainly influenced by local site characteristics. Especially the number of plant species and a near-natural forest structure turned out to positively affect moth diversity. The diversity of nearby natural habitat structures also promoted higher local moth diversity, while landscape characteristics were less relevant. However, the diversity of near natural landscape elements within a 500-1000m radius positively affected the number of moth individuals. So, while favorable local conditions might be able to preserve the local α-diversity of moths, the landscape context plays a major role for individual numbers and regional γ-diversity. Integrating landscape-scale actions in conservation management therefore might be crucial not only for preserving the regional diversity, but also for maintaining a high number of insects, being important as food resource for higher trophic levels like birds and bats, but also likely relevant as pollinators of many flowering plants.
By zooming into moth species composition (differentiation diversity), I found local and landscape-scale predictors to play equally important roles in shaping moth assemblages. Proportional β-diversity addi-tionally was used to get an insight in how the different predictors affect moth communities. On the local scale, differentiation diversity was correlated to forest age and density. Additionally, mean soil humidity and nutrient content (described via plant indicator values) affected species composition. With the lack of typical forest species at dry, dense and young forest sites, it seems that subtractive heterogenization was the process underlying the observed β-diversity patterns. Looking at the landscape context, moth community composition changes were idiosyncratic relative to the landscape context. For PdC, the proportion of human-modified areas within 500m radius played an important role in shaping moth communities. In PsV, where less agricultural fields occur in the surroundings, the proximity to the industrial harbor of Ravenna significantly affected moth community composition. Here the loss of rare species near urban areas is believed to cause the observed patterns in β-diversity. In conclusion, anthropogenic actions seem to act very specifically on local communities. These findings emphasize that instead of perpetuating generalizations based on large-scale meta-analyses, it is always important to investigate focal areas of interest individually.
In an attempt to link moth community composition with ecosystem functions, I finally analyzed func-tional richness, redundancy and niche occupation of these insects. Biodiversity erosion can directly affect ecosystem resilience, because species richness itself can be described as the natural insurance capital of an ecosystem. With more species around, there is a higher chance to compensate disturbance, because even if some species disappear, others might take their place and guarantee that the functional niche keeps being occupied. To investigate the functional diversity of moths in PsV and PdC, I collated information on 387 species and 14 ecological traits. The analysis of the functional dendrogram led me to define 25 different functional groups, which were used to score niche occupation and functional redundancy. Similar to my findings on species composition both, local and regional environmental predictors played a significant role for shaping functional diversity. The humidity-nutrient gradient again was the strongest local predictor, but also plant diversity positively affected functional moth richness, redundancy and niche occupation at the site scale. On the landscape scale, landscape diversity around the light-trapping sites was most important. With diverse near-natural landscapes around, functional richness and redundancy was increasing, as well as niche occupancy. Landscape fragmentation, in contrast, had negative effects on the investigated functional components.
Collectively, these findings underline 1) the importance of a near-natural, old grown open forest struc-ture with a well developed understory and a diverse, species rich vegetation on the local scale. Such forests cannot only preserve the local α-diversity, but also might show better ecosystem function and resilience, as they maintain higher functional richness and redundancy. 2) The indispensability of land-scape-scale actions for successful conservation management can be concluded throughout all studied aspects of diversity. Landscape diversity does not only play a major role for preserving insect numbers, but also for γ-diversity, functional richness and redundancy. Finally, the analysis of species composition confirmed that landscape-scale anthropogenic impacts do not end at the reserve borders – they also affect biota inside conservation areas. If we want to counteract insect decline, we cannot only rely on isolated conservation area patches. We also have to think about ameliorating management methods towards a more sustainable land use in order to preserve the multifunctionality of ecosystems.