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Formation of tussocks by sedges: effects of hydroperiod and nutrients.

Tussock formation is a global phenomenon that enhances microtopography and increases biodiversity by adding structure to ecological communities, but little is known about tussock development in relation to environmental factors. To further efforts to restore wetland microtopography and associated functions, we investigated Carex stricta tussock size in relation to elevation (a proxy for water depth) at a range of sites in southern Wisconsin, USA, and tested the effect of five hydroperiods and N+P addition (15 g N/m2 + 0.37 g P/m2) on tussock formation during a three-year mesocosm experiment. Wet meadows dominated by C. stricta averaged 4.9 tussocks/m2, with a mean volume of 1160 cm3 and height of 15 cm. Within sites, taller tussocks occurred at lower elevations, suggesting a structural adaptation to anoxic conditions. In our mesocosm experiment, C. stricta accelerated tussock formation when inundated, and it increased overall productivity with N + P addition. Within two growing seasons, continuous inundation (+18 cm) in the mesocosms led to tussocks that were nearly as tall as in our field survey (mean height in mesocosms, 10 +/- 1.3 cm; maximum, 17 cm). Plants grown with constant low water (-18 cm) only formed short mounds (mean height = 2 +/- 0.4 cm). After three growing seasons, the volume of the largest tussocks (3274 +/- 376 cm3, grown with +18 cm water depth and N + P addition) was 12 times that of the smallest (275 +/- 38 cm3, grown with -18 cm water depth and no N + P). Though tussock composition varied among hydroperiods, tussocks were predominantly organic (74-94% of dry mass) and composed of leaf bases (46-59%), fine roots (10-31%), and duff (5-13%). Only the plants subjected to high water levels produced the vertically oriented rhizomes and ascending shoot bases that were prevalent in field-collected tussocks. Under continuous or periodic inundation, tussocks achieved similar heights and accumulated similar levels of organic matter (range: 163-394 g C/m2), and we conclude that these hydroperiods can accelerate tussock formation. Thus, C. stricta has high utility for restoring wetland microtopography and associated functions, including carbon accumulation.