@article{BergholzMayRistowetal.2017,
  author    = {Bergholz, Kolja and May, Felix and Ristow, Michael and Giladi, Itamar and Ziv, Yaron and Jeltsch, Florian},
  title     = {Two Mediterranean annuals feature high within-population trait variability and respond differently to a precipitation gradient},
  series = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie},
  volume    = {25},
  journal   = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie},
  publisher = {Elsevier},
  address   = {Jena},
  issn      = {1439-1791},
  doi       = {10.1016/j.baae.2017.11.001},
  pages     = {48 -- 58},
  year      = {2017},
  abstract  = {Intraspecific trait variability plays an important role in species adaptation to climate change. However, it still remains unclear how plants in semi-arid environments respond to increasing aridity. We investigated the intraspecific trait variability of two common Mediterranean annuals (Geropogon hybridus and Crupina crupinastrum) with similar habitat preferences. They were studied along a steep precipitation gradient in Israel similar to the maximum predicted precipitation changes in the eastern Mediterranean basin (i.e. -30\% until 2100). We expected a shift from competitive ability to stress tolerance with decreasing precipitation and tested this expectation by measuring key functional traits (canopy and seed release height, specific leaf area, N-and P-leaf content, seed mass). Further, we evaluated generative bet-hedging strategies by different seed traits. Both species showed different responses along the precipitation gradient. C. crupinastrum exhibited only decreased plant height toward saridity, while G. hybridus showed strong trends of generative adaptation to aridity. Different seed trait indices suggest increased bet-hedging of G. hybridus in arid environments. However, no clear trends along the precipitation gradient were observed in leaf traits (specific leaf area and leaf N-/P-content) in both species. Moreover, variance decomposition revealed that most of the observed trait variation (>> 50\%) is found within populations. The findings of our study suggest that responses to increased aridity are highly species-specific and local environmental factors may have a stronger effect on intraspecific trait variation than shifts in annual precipitation. We therefore argue that trait-based analyses should focus on precipitation gradients that are comparable to predicted precipitation changes and compare precipitation effects to effects of local environmental factors. (C) 2017 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.},
  language  = {en}
}
@phdthesis{Bergholz2018,
  author    = {Bergholz, Kolja},
  title     = {Trait-based understanding of plant species distributions along environmental gradients},
  doi       = {10.25932/publishup-42634},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus4-426341},
  school      = {Universit{\"a}t Potsdam},
  pages     = {128},
  year      = {2018},
  abstract  = {For more than two centuries, plant ecologists have aimed to understand how environmental gradients and biotic interactions shape the distribution and co-occurrence of plant species. In recent years, functional trait-based approaches have been increasingly used to predict patterns of species co-occurrence and species distributions along environmental gradients (trait-environment relationships). Functional traits are measurable properties at the individual level that correlate well with important processes. Thus, they allow us to identify general patterns by synthesizing studies across specific taxonomic compositions, thereby fostering our understanding of the underlying processes of species assembly. However, the importance of specific processes have been shown to be highly dependent on the spatial scale under consideration. In particular, it remains uncertain which mechanisms drive species assembly and allow for plant species coexistence at smaller, more local spatial scales. Furthermore, there is still no consensus on how particular environmental gradients affect the trait composition of plant communities. For example, increasing drought because of climate change is predicted to be a main threat to plant diversity, although it remains unclear which traits of species respond to increasing aridity. Similarly, there is conflicting evidence of how soil fertilization affects the traits related to establishment ability (e.g., seed mass). In this cumulative dissertation, I present three empirical trait-based studies that investigate specific research questions in order to improve our understanding of species distributions along environmental gradients. In the first case study, I analyze how annual species assemble at the local scale and how environmental heterogeneity affects different facets of biodiversity—i.e. taxonomic, functional, and phylogenetic diversity—at different spatial scales. The study was conducted in a semi-arid environment at the transition zone between desert and Mediterranean ecosystems that features a sharp precipitation gradient (Israel). Different null model analyses revealed strong support for environmentally driven species assembly at the local scale, since species with similar traits tended to co-occur and shared high abundances within microsites (trait convergence). A phylogenetic approach, which assumes that closely related species are functionally more similar to each other than distantly related ones, partly supported these results. However, I observed that species abundances within microsites were, surprisingly, more evenly distributed across the phylogenetic tree than expected (phylogenetic overdispersion). Furthermore, I showed that environmental heterogeneity has a positive effect on diversity, which was higher on functional than on taxonomic diversity and increased with spatial scale. The results of this case study indicate that environmental heterogeneity may act as a stabilizing factor to maintain species diversity at local scales, since it influenced species distribution according to their traits and positively influenced diversity. All results were constant along the precipitation gradient. In the second case study (same study system as case study one), I explore the trait responses of two Mediterranean annuals (Geropogon hybridus and Crupina crupinastrum) along a precipitation gradient that is comparable to the maximum changes in precipitation predicted to occur by the end of this century (i.e., -30\%). The heterocarpic G. hybridus showed strong trends in seed traits, suggesting that dispersal ability increased with aridity. By contrast, the homocarpic C. crupinastrum showed only a decrease in plant height as aridity increased, while leaf traits of both species showed no consistent pattern along the precipitation gradient. Furthermore, variance decomposition of traits revealed that most of the trait variation observed in the study system was actually found within populations. I conclude that trait responses towards aridity are highly species-specific and that the amount of precipitation is not the most striking environmental factor at this particular scale. In the third case study, I assess how soil fertilization mediates—directly by increased nutrient addition and indirectly by increased competition—the effect of seed mass on establishment ability. For this experiment, I used 22 species differing in seed mass from dry grasslands in northeastern Germany and analyzed the interacting effects of seed mass with nutrient availability and competition on four key components of seedling establishment: seedling emergence, time of seedling emergence, seedling survival, and seedling growth. (Time of) seedling emergence was not affected by seed mass. However, I observed that the positive effect of seed mass on seedling survival is lowered under conditions of high nutrient availability, whereas the positive effect of seed mass on seedling growth was only reduced by competition. Based on these findings, I developed a conceptual model of how seed mass should change along a soil fertility gradient in order to reconcile conflicting findings from the literature. In this model, seed mass shows a U-shaped pattern along the soil fertility gradient as a result of changing nutrient availability and competition. Overall, the three case studies highlight the role of environmental factors on species distribution and co-occurrence. Moreover, the findings of this thesis indicate that spatial heterogeneity at local scales may act as a stabilizing factor that allows species with different traits to coexist. In the concluding discussion, I critically debate intraspecific trait variability in plant community ecology, the use of phylogenetic relationships and easily measured key functional traits as a proxy for species' niches. Finally, I offer my outlook for the future of functional plant community research.},
  language  = {en}
}
@misc{BergholzKoberJeltschetal.2021,
  author    = {Bergholz, Kolja and Kober, Klarissa and Jeltsch, Florian and Schmidt, Kristina and Weiß, Lina},
  title     = {Trait means or variance},
  series = {Postprints der Universit{\"a}t Potsdam : Mathematisch-Naturwissenschaftliche Reihe},
  journal   = {Postprints der Universit{\"a}t Potsdam : Mathematisch-Naturwissenschaftliche Reihe},
  issn      = {1866-8372},
  doi       = {10.25932/publishup-51990},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus4-519905},
  pages     = {3357 -- 3365},
  year      = {2021},
  abstract  = {One of the few laws in ecology is that communities consist of few common and many rare taxa. Functional traits may help to identify the underlying mechanisms of this community pattern, since they correlate with different niche dimensions. However, comprehensive studies are missing that investigate the effects of species mean traits (niche position) and intraspecific trait variability (ITV, niche width) on species abundance. In this study, we investigated fragmented dry grasslands to reveal trait-occurrence relationships in plants at local and regional scales. We predicted that (a) at the local scale, species occurrence is highest for species with intermediate traits, (b) at the regional scale, habitat specialists have a lower species occurrence than generalists, and thus, traits associated with stress-tolerance have a negative effect on species occurrence, and (c) ITV increases species occurrence irrespective of the scale. We measured three plant functional traits (SLA = specific leaf area, LDMC = leaf dry matter content, plant height) at 21 local dry grassland communities (10 m × 10 m) and analyzed the effect of these traits and their variation on species occurrence. At the local scale, mean LDMC had a positive effect on species occurrence, indicating that stress-tolerant species are the most abundant rather than species with intermediate traits (hypothesis 1). We found limited support for lower specialist occurrence at the regional scale (hypothesis 2). Further, ITV of LDMC and plant height had a positive effect on local occurrence supporting hypothesis 3. In contrast, at the regional scale, plants with a higher ITV of plant height were less frequent. We found no evidence that the consideration of phylogenetic relationships in our analyses influenced our findings. In conclusion, both species mean traits (in particular LDMC) and ITV were differently related to species occurrence with respect to spatial scale. Therefore, our study underlines the strong scale-dependency of trait-abundance relationships.},
  language  = {en}
}
@article{BergholzKoberJeltschetal.2021,
  author    = {Bergholz, Kolja and Kober, Klarissa and Jeltsch, Florian and Schmidt, Kristina and Weiß, Lina},
  title     = {Trait means or variance},
  series = {Ecology and evolution},
  volume    = {11},
  journal   = {Ecology and evolution},
  number    = {7},
  publisher = {John Wiley \& Sons, Inc.},
  issn      = {2045-7758},
  doi       = {10.1002/ece3.7287},
  pages     = {3357 -- 3365},
  year      = {2021},
  abstract  = {One of the few laws in ecology is that communities consist of few common and many rare taxa. Functional traits may help to identify the underlying mechanisms of this community pattern, since they correlate with different niche dimensions. However, comprehensive studies are missing that investigate the effects of species mean traits (niche position) and intraspecific trait variability (ITV, niche width) on species abundance. In this study, we investigated fragmented dry grasslands to reveal trait-occurrence relationships in plants at local and regional scales. We predicted that (a) at the local scale, species occurrence is highest for species with intermediate traits, (b) at the regional scale, habitat specialists have a lower species occurrence than generalists, and thus, traits associated with stress-tolerance have a negative effect on species occurrence, and (c) ITV increases species occurrence irrespective of the scale. We measured three plant functional traits (SLA = specific leaf area, LDMC = leaf dry matter content, plant height) at 21 local dry grassland communities (10 m × 10 m) and analyzed the effect of these traits and their variation on species occurrence. At the local scale, mean LDMC had a positive effect on species occurrence, indicating that stress-tolerant species are the most abundant rather than species with intermediate traits (hypothesis 1). We found limited support for lower specialist occurrence at the regional scale (hypothesis 2). Further, ITV of LDMC and plant height had a positive effect on local occurrence supporting hypothesis 3. In contrast, at the regional scale, plants with a higher ITV of plant height were less frequent. We found no evidence that the consideration of phylogenetic relationships in our analyses influenced our findings. In conclusion, both species mean traits (in particular LDMC) and ITV were differently related to species occurrence with respect to spatial scale. Therefore, our study underlines the strong scale-dependency of trait-abundance relationships.},
  language  = {en}
}
@article{BergholzSittelRistowetal.2022,
  author    = {Bergholz, Kolja and Sittel, Lara-Pauline and Ristow, Michael and Jeltsch, Florian and Weiss, Lina},
  title     = {Pollinator guilds respond contrastingly at different scales to landscape parameters of land-use intensity},
  series = {Ecology and evolution},
  volume    = {12},
  journal   = {Ecology and evolution},
  number    = {3},
  publisher = {Wiley},
  address   = {Hoboken},
  issn      = {2045-7758},
  doi       = {10.1002/ece3.8708},
  pages     = {11},
  year      = {2022},
  abstract  = {Land-use intensification is the main factor for the catastrophic decline of insect pollinators. However, land-use intensification includes multiple processes that act across various scales and should affect pollinator guilds differently depending on their ecology. We aimed to reveal how two main pollinator guilds, wild bees and hoverflies, respond to different land-use intensification measures, that is, arable field cover (AFC), landscape heterogeneity (LH), and functional flower composition of local plant communities as a measure of habitat quality. We sampled wild bees and hoverflies on 22 dry grassland sites within a highly intensified landscape (NE Germany) within three campaigns using pan traps. We estimated AFC and LH on consecutive radii (60-3000 m) around the dry grassland sites and estimated the local functional flower composition. Wild bee species richness and abundance was positively affected by LH and negatively by AFC at small scales (140-400 m). In contrast, hoverflies were positively affected by AFC and negatively by LH at larger scales (500-3000 m), where both landscape parameters were negatively correlated to each other. At small spatial scales, though, LH had a positive effect on hoverfly abundance. Functional flower diversity had no positive effect on pollinators, but conspicuous flowers seem to attract abundance of hoverflies. In conclusion, landscape parameters contrarily affect two pollinator guilds at different scales. The correlation of landscape parameters may influence the observed relationships between landscape parameters and pollinators. Hence, effects of land-use intensification seem to be highly landscape-specific.},
  language  = {en}
}
@article{BergholzSittelRistowetal.2022,
  author    = {Bergholz, Kolja and Sittel, Lara-Pauline and Ristow, Michael and Jeltsch, Florian and Weiß, Lina},
  title     = {Pollinator guilds respond contrastingly at different scales to landscape parameters of land-use intensity},
  series = {Ecology and Evolution},
  volume    = {12},
  journal   = {Ecology and Evolution},
  number    = {3},
  publisher = {John Wiley \& Sons, Inc.},
  address   = {Hoboken (New Jersey)},
  issn      = {2045-7758},
  doi       = {10.1002/ece3.8708},
  pages     = {11},
  year      = {2022},
  abstract  = {Land-use intensification is the main factor for the catastrophic decline of insect pollinators. However, land-use intensification includes multiple processes that act across various scales and should affect pollinator guilds differently depending on their ecology. We aimed to reveal how two main pollinator guilds, wild bees and hoverflies, respond to different land-use intensification measures, that is, arable field cover (AFC), landscape heterogeneity (LH), and functional flower composition of local plant communities as a measure of habitat quality. We sampled wild bees and hoverflies on 22 dry grassland sites within a highly intensified landscape (NE Germany) within three campaigns using pan traps. We estimated AFC and LH on consecutive radii (60-3000 m) around the dry grassland sites and estimated the local functional flower composition. Wild bee species richness and abundance was positively affected by LH and negatively by AFC at small scales (140-400 m). In contrast, hoverflies were positively affected by AFC and negatively by LH at larger scales (500-3000 m), where both landscape parameters were negatively correlated to each other. At small spatial scales, though, LH had a positive effect on hoverfly abundance. Functional flower diversity had no positive effect on pollinators, but conspicuous flowers seem to attract abundance of hoverflies. In conclusion, landscape parameters contrarily affect two pollinator guilds at different scales. The correlation of landscape parameters may influence the observed relationships between landscape parameters and pollinators. Hence, effects of land-use intensification seem to be highly landscape-specific.},
  language  = {en}
}
@misc{BergholzSittelRistowetal.2022,
  author    = {Bergholz, Kolja and Sittel, Lara-Pauline and Ristow, Michael and Jeltsch, Florian and Weiß, Lina},
  title     = {Pollinator guilds respond contrastingly at different scales to landscape parameters of land-use intensity},
  series = {Zweitver{\"o}ffentlichungen der Universit{\"a}t Potsdam : Mathematisch-Naturwissenschaftliche Reihe},
  journal   = {Zweitver{\"o}ffentlichungen der Universit{\"a}t Potsdam : Mathematisch-Naturwissenschaftliche Reihe},
  number    = {1298},
  issn      = {1866-8372},
  doi       = {10.25932/publishup-57730},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus4-577307},
  pages     = {11},
  year      = {2022},
  abstract  = {Land-use intensification is the main factor for the catastrophic decline of insect pollinators. However, land-use intensification includes multiple processes that act across various scales and should affect pollinator guilds differently depending on their ecology. We aimed to reveal how two main pollinator guilds, wild bees and hoverflies, respond to different land-use intensification measures, that is, arable field cover (AFC), landscape heterogeneity (LH), and functional flower composition of local plant communities as a measure of habitat quality. We sampled wild bees and hoverflies on 22 dry grassland sites within a highly intensified landscape (NE Germany) within three campaigns using pan traps. We estimated AFC and LH on consecutive radii (60-3000 m) around the dry grassland sites and estimated the local functional flower composition. Wild bee species richness and abundance was positively affected by LH and negatively by AFC at small scales (140-400 m). In contrast, hoverflies were positively affected by AFC and negatively by LH at larger scales (500-3000 m), where both landscape parameters were negatively correlated to each other. At small spatial scales, though, LH had a positive effect on hoverfly abundance. Functional flower diversity had no positive effect on pollinators, but conspicuous flowers seem to attract abundance of hoverflies. In conclusion, landscape parameters contrarily affect two pollinator guilds at different scales. The correlation of landscape parameters may influence the observed relationships between landscape parameters and pollinators. Hence, effects of land-use intensification seem to be highly landscape-specific.},
  language  = {en}
}
@article{BergholzJeltschWeissetal.2015,
  author    = {Bergholz, Kolja and Jeltsch, Florian and Weiß, Lina and Pottek, Janine and Geißler, Katja and Ristow, Michael},
  title     = {Fertilization affects the establishment ability of species differing in seed mass via direct nutrient addition and indirect competition effects},
  series = {Oikos},
  volume    = {124},
  journal   = {Oikos},
  number    = {11},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {0030-1299},
  doi       = {10.1111/oik.02193},
  pages     = {1547 -- 1554},
  year      = {2015},
  abstract  = {Fertilization causes species loss and species dominance changes in plant communities worldwide. However, it still remains unclear how fertilization acts upon species functional traits, e.g. seed mass. Seed mass is a key trait of the regeneration strategy of plants, which influences a range of processes during the seedling establishment phase. Fertilization may select upon seed mass, either directly by increased nutrient availability or indirectly by increased competition. Since previous research has mainly analyzed the indirect effects of fertilization, we disentangled the direct and indirect effects to examine how nutrient availability and competition influence the seed mass relationships on four key components during seedling establishment: seedling emergence, time of seedling emergence, seedling survival and seedling growth. We conducted a common garden experiment with 22 dry grassland species with a two-way full factorial design that simulated additional nutrient supply and increased competition. While we found no evidence that fertilization either directly by additional nutrient supply or indirectly by increased competition alters the relationship between seed mass and (time of) seedling emergence, we revealed that large seed mass is beneficial under nutrient-poor conditions (seedlings have greater chances of survival, particularly in nutrient-poor soils) as well as under competition (large-seeded species produced larger seedlings, which suffered less from competition than small-seeded species). Based on these findings, we argue that both factors, i.e. nutrient availability and competition intensity, ought to be considered to understand how fertilization influences seedling establishment and species composition with respect to seed mass in natural communities. We propose a simple conceptual model, in which seed mass in natural communities is determined by competition intensity and nutrient availability. Here, we hypothesize that seed mass shows a U-shaped pattern along gradients of soil fertility, which may explain the contrasting soil fertility-seed mass relationships found in the recent literature.},
  language  = {en}
}
@article{BergholzMayGiladietal.2017,
  author    = {Bergholz, Kolja and May, Felix and Giladi, Itamar and Ristow, Michael and Ziv, Yaron and Jeltsch, Florian},
  title     = {Environmental heterogeneity drives fine-scale species assembly and functional diversity of annual plants in a semi-arid environment},
  series = {Perspectives in plant ecology, evolution and systematics},
  volume    = {24},
  journal   = {Perspectives in plant ecology, evolution and systematics},
  publisher = {Elsevier},
  address   = {Jena},
  issn      = {1433-8319},
  doi       = {10.1016/j.ppees.2017.01.001},
  pages     = {138 -- 146},
  year      = {2017},
  abstract  = {Spatial environmental heterogeneity is considered a fundamental factor for the maintenance of plant species richness. However, it still remains unclear whether heterogeneity may also facilitate coexistence at fine grain sizes or whether other processes, like mass effects and source sink dynamics due to dispersal, control species composition and diversity at these scales. In this study, we used two complimentary analyses to identify the role of heterogeneity within 15 m x 15 m plots for the coexistence of species-rich annual communities in a semi-arid environment along a steep precipitation gradient. Specifically, we: (a) analyzed the effect of environmental heterogeneity on species, functional and phylogenetic diversity within microsites (alpha diversity, 0.06 m(2) and 1 m(2)), across microsites (beta diversity), and diversity at the entire plot (gamma diversity); (b) further we used two null models to detect non-random trait and phylogenetic patterns in order to infer assembly processes, i.e. whether co-occurring species tend to share similar traits (trait convergence) or dissimilar traits (trait divergence). In general, our results showed that heterogeneity had a positive effect on community diversity. Specifically, for alpha diversity, the effect was significant for functional diversity, and not significant for either species or phylogenetic diversities. For beta diversity, all three measures of community diversity (species, functional, and phylogenetic) increased significantly, as they also did for gamma diversity, where functional measures were again stronger than for species or phylogenetic measures. In addition, the null model approach consistently detected trait convergence, indicating that species with similar traits tended to co-occur and had high abundances in a given microsite. While null model analysis across the phylogeny partly supported these trait findings, showing phylogenetic underdispersion at the 1m(2) grain size, surprisingly when species abundances in microsites were analyzed they were more evenly distributed across the phylogenetic tress than expected (phylogenetic overdispersion). In conclusion, our results provide compelling support that environmental heterogeneity at a relatively fine scale is an important factor for species co-existence as it positively affects diversity as well as influences species assembly. Our study underlines the need for trait-based approaches conducted at fine grain sizes in order to better understand species coexistence and community assembly. (C) 2017 Elsevier GmbH. All rights reserved.},
  language  = {en}
}