Clonal plants in a spatially heterogeneous environment: effects of integration on Serengeti grassland response to defoliation and urine-hits from grazing mammals

It has been hypothesized that clonal integration between ramets inenvironments with spatially variable rates of herbivory and nutrientavailabilities leads to increased growth and fitness in the genet. An increasein genet growth could potentially influence ecosystem processes such as primaryproductivity and nutrient cycling. I tested the idea that clonal integrationwould lead to greater aboveground productivity, compensatory response todefoliation, and N re-distribution in a factorial experiment in fieldplots in the Serengeti Ecosystem, Tanzania. Each plot had either all stolonssevered or left intact, had repeated defoliation (to simulate grazing bythe African buffalo) or none, and was either located next to a plot thatreceived urea (to simulate a urine-hit) or next to a plot thatremained untreated. Plots that received stolon severing treatments had32% less peak biomass than did connected control plots, and this suggeststhat plants grew better when ramets remained connected. However, compensatoryresponse by plots to repeated defoliation was inconsistent with the hypothesizedbenefits of ramet integration at the ecosystem level: productivityresponse to defoliation was similar between plots with connected vs. severedramets (i.e. no stolon severing × defoliation interaction wasfound). When averaged across other treatments, defoliated plots hadincreased productivity compared to nondefoliated plots during the growingseason. Thus, ramet connection and defoliation did increase productivity whenthey were considered alone, but productivity response to defoliation wasunaffected by ramet connections. Urea additions, which led to a 78%increase in productivity in adjacent urine-hits, had noconsistent effect on productivity but did increase leaf percent N in adjoiningstudy plots. Thus, in the Serengeti, urine hits probably have very localizedeffects on productivity during the initial growing season.