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Abstract Nutrient availability and grazing are known as main drivers of grassland plant diversity, and increased nutrient availability and long‐term cessation of grazing often decrease local‐scale plant diversity. Experimental tests of mechanisms determining plant diversity focus mainly on vascular plants (VP), whereas non‐vascular plants (NVP, here bryophytes) have been ignored. It is therefore not known how the current models based on VPs predict the rates of total (NVP + VP) losses in plant diversity.Here we used plant community data, including VPs and NVPs, from nine sites in Europe and North America and belonging to the Nutrient Network experiment, to test whether neglecting NVPs leads to biased estimates of plant diversity loss rates. The plant communities were subjected to factorial addition of nitrogen (N), phosphorus (P), potassium with micronutrients (K+μ), as well as a grazing exclusion combined with multi‐nutrient fertilization (NPK+μ) treatment.We found that nutrient additions reduced both NVP and VP species richness, but the effects on NVP species richness were on average stronger than on VPs: NVP species richness decreased 67%, while VP species richness decreased 28%, causing their combined richness to decrease 38% in response to multi‐nutrient (NPK+μ) fertilization. Thus, VP diversity alone underestimated total plant diversity loss by 10 percentage points.Although NVP and VP species diversities similarly declined in response to N and NPK+μfertilizations, the evenness of NVPs increased and that of VPs remained unchanged. NP, NPK+μfertilization and NPK+μfertilization combined with grazing exclusion, associated with decreasing light availability at ground level, led to the strongest loss of NVP species or probability of NVP presence. However, grazing did not generally mitigate the fertilization effects.Synthesis. In nine grassland sites in Europe and North America, nutrient addition caused a larger relative decline in non‐vascular plant (NVP) than vascular plant species richness. Hence, not accounting for NVPs can lead to underestimation of losses in plant diversity in response to continued nutrient pollution of grasslands.
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Nutrient addition drives declines in grassland species richness primarily via enhanced species loss
Abstract Declines in grassland diversity in response to nutrient addition are a general consequence of global change. This decline in species richness may be driven by multiple underlying processes operating at different time‐scales. Nutrient addition can reduce diversity by enhancing the rate of local extinction via competitive exclusion, or by reducing the rate of colonization by constraining the pool of species able to colonize under new conditions. Partitioning net change into extinction and colonization rates will better delineate the long‐term effect of global change in grasslands.We synthesized changes in richness in response to experimental fertilization with nitrogen, phosphorus and potassium with micronutrients across 30 grasslands. We quantified changes in local richness, colonization, and extinction over 8–10 years of nutrient addition, and compared these rates against control conditions to isolate the effect of nutrient addition from background dynamics.Total richness at steady state in the control plots was the sum of equal, relatively high rates of local colonization and extinction. On aggregate, 30%–35% of initial species were lost and the same proportion of new species were gained at least once over a decade. Absolute turnover increased with site‐level richness but was proportionately greater at lower‐richness sites relative to starting richness. Loss of total richness with nutrient addition, especially N in combination with P or K, was driven by enhanced rates of extinction with a smaller contribution from reduced colonization. Enhanced extinction and reduced colonization were disproportionately among native species, perennials, and forbs. Reduced colonization plateaued after the first few (<5) years after nutrient addition, while enhanced extinction continued throughout the first decade.Synthesis. Our results indicate a high rate of colonizations and extinctions underlying the richness of ambient communities and that nutrient enhancement drives overall declines in diversity primarily by exclusion of previously established species. Moreover, enhanced extinction continues over long time‐scales, suggesting continuous, long‐term community responses and a need for long‐term study to fully realize the extinction impact of increased nutrients on grassland composition.
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- PAR ID:
- 10383013
- Author(s) / Creator(s):
- ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; more »
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Journal of Ecology
- Volume:
- 111
- Issue:
- 3
- ISSN:
- 0022-0477
- Page Range / eLocation ID:
- p. 552-563
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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