@article{WulfKolk2014,
  author    = {Wulf, Monika and Kolk, Jens},
  title     = {Plant species richness of very small forests related to patch configuration, quality, heterogeneity and history},
  series = {Journal of vegetation science},
  volume    = {25},
  journal   = {Journal of vegetation science},
  number    = {5},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {1100-9233},
  doi       = {10.1111/jvs.12172},
  pages     = {1267 -- 1277},
  year      = {2014},
  abstract  = {Questions: (i) Is plant species richness related to patch configuration, quality, heterogeneity and history in very small forest patches? (ii) Is there a similar effect of patch configuration, quality, heterogeneity and history on all plant species or does it vary for different plant groups? (iii) Does the importance of patch configuration, quality, heterogeneity and history vary for plant species richness between patch size classes? Location: Prignitz region, western part of Brandenburg, Germany Methods: Plant species lists for 183 forest patches (0.08-12.7 ha) were compiled. We conducted a hierarchical partitioning analysis to test which of the four patch variables - configuration, quality, heterogeneity and history - explained most of the variance in plant species richness and richness of various plant groups. Results: Irrespective of patch size class, species richness is mainly predicted by patch quality and heterogeneity. In particular, generalists are primarily affected by the soil moisture and nutrient level, and specialists additionally by patch heterogeneity and history. A higher historical habitat quality, together with long habitat continuity, led to an increase in forest specialists and geophytes in all patches. In the >5-ha patches only historical habitat quality accounted for a significant increase in the number of species with short-distance dispersal capacity in all species. Regardless of patch size, the number of locally infrequent species increased significantly with historical habitat quality. The effect of patch configuration was negligible, except for space-filling trees and shrubs, when considering all patches. Conclusions: Our more comprehensive statistical approach shows, in general, that all complex variables have an effect on species richness and should be considered in future studies. The fact that patch heterogeneity and quality are the main predictors of species richness indicates that very small forest patches covered a range of highly structured patches and sites having various growth conditions. We assume that the century-old tradition of private ownership with smallholder social structure is the main reason for variability in stand structure and species assemblage, resulting in the uniqueness of each patch. Since the number of locally infrequent plant species increases significantly with historical habitat quality, we suggest that it is particularly important to consider this variable in future studies on the decline of infrequent plant species.},
  language  = {en}
}
@article{KolkNaafWulf2017,
  author    = {Kolk, Jens and Naaf, Tobias and Wulf, Monika},
  title     = {Paying the colonization credit},
  series = {Biodiversity and conservation},
  volume    = {26},
  journal   = {Biodiversity and conservation},
  publisher = {Springer},
  address   = {Dordrecht},
  issn      = {0960-3115},
  doi       = {10.1007/s10531-016-1271-y},
  pages     = {735 -- 755},
  year      = {2017},
  abstract  = {Massive historical land cover changes in the Central European lowlands have resulted in a forest distribution that now comprises small remnants of ancient forests and more recently established post-agricultural forests. Here, land-use history is considered a key driver of recent herb-layer community changes, where an extinction debt in ancient forest remnants and/or a colonization credit in post-agricultural forests are being paid over time. On a regional scale, these payments should in theory lead toward a convergence in species richness between ancient and post-agricultural forests over time. In this study, we tested this assumption with a resurvey of 117 semi-permanent plots in the well-studied deciduous forests of the Prignitz region (Brandenburg, NE Germany), where we knew that the plant communities of post-agricultural stands exhibit a colonization credit while the extinction debt in ancient stands has largely been paid. We compared changes in the species richness of all herb layer species, forest specialists and ancient forest indicator species between ancient and post-agricultural stands with linear mixed effect models and determined the influence of patch connectivity on the magnitude of species richness changes. Species richness increased overall, but the richness of forest specialists increased significantly more in post-agricultural stands and was positively influenced by higher patch connectivity, indicating a convergence in species richness between the ancient and postagricultural stands. Furthermore, the richness of ancient forest indicator species only increased significantly in post-agricultural stands. For the first time, we were able to verify a gradual payment of the colonization credit in post-agricultural forest stands using a comparison of actual changes in temporal species richness.},
  language  = {en}
}
@phdthesis{Kolk2019,
  author    = {Kolk, Jens},
  title     = {The long-term legacy of historical land cover changes},
  doi       = {10.25932/publishup-43939},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus4-439398},
  school      = {Universit{\"a}t Potsdam},
  pages     = {196},
  year      = {2019},
  abstract  = {Over the last years there is an increasing awareness that historical land cover changes and associated land use legacies may be important drivers for present-day species richness and biodiversity due to time-delayed extinctions or colonizations in response to historical environmental changes. Historically altered habitat patches may therefore exhibit an extinction debt or colonization credit and can be expected to lose or gain species in the future. However, extinction debts and colonization credits are difficult to detect and their actual magnitudes or payments have rarely been quantified because species richness patterns and dynamics are also shaped by recent environmental conditions and recent environmental changes. In this thesis we aimed to determine patterns of herb-layer species richness and recent species richness dynamics of forest herb layer plants and link those patterns and dynamics to historical land cover changes and associated land use legacies. The study was conducted in the Prignitz, NE-Germany, where the forest distribution remained stable for the last ca. 100 years but where a) the deciduous forest area had declined by more than 90 per cent (leaving only remnants of "ancient forests"), b) small new forests had been established on former agricultural land ("post-agricultural forests"). Here, we analyzed the relative importance of land use history and associated historical land cover changes for herb layer species richness compared to recent environmental factors and determined magnitudes of extinction debt and colonization credit and their payment in ancient and post-agricultural forests, respectively. We showed that present-day species richness patterns were still shaped by historical land cover changes that ranged back to more than a century. Although recent environmental conditions were largely comparable we found significantly more forest specialists, species with short-distance dispersal capabilities and clonals in ancient forests than in post-agricultural forests. Those species richness differences were largely contingent to a colonization credit in post-agricultural forests that ranged up to 9 species (average 4.7), while the extinction debt in ancient forests had almost completely been paid. Environmental legacies from historical agricultural land use played a minor role for species richness differences. Instead, patch connectivity was most important. Species richness in ancient forests was still dependent on historical connectivity, indicating a last glimpse of an extinction debt, and the colonization credit was highest in isolated post-agricultural forests. In post-agricultural forests that were better connected or directly adjacent to ancient forest patches the colonization credit was way smaller and we were able to verify a gradual payment of the colonization credit from 2.7 species to 1.5 species over the last six decades.},
  language  = {en}
}