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REPORT
Top predator control of plant biodiversity
and productivity in an old-field ecosystem
Abstract
Oswald J. Schmitz Predators can have strong indirect effects on plants by altering the way herbivores
School of Forestry and impact plants. Yet, many current evaluations of plant species diversity and ecosystem
Environmental Studies and function ignore the effects of predators and focus directly on the plant trophic level.
Department of Ecology and This report presents results of a 3-year field experiment in a temperate old-field
Evolutionary Biology, Yale
ecosystem that excluded either predators, or predators and herbivores and evaluated
University, 370 Prospect Street,
the consequence of those manipulations on plant species diversity (richness and
New Haven, CT 06511, USA
evenness) and plant productivity. Sustained predator and predator and herbivore
E-mail:
oswald.schmitz@yale.edu
exclusion resulted in lower plant species evenness and higher plant biomass production
than control field plots representing the intact natural ecosystem. Predators had this
diversity-enhancing effect on plants by causing herbivores to suppress the abundance
of a competitively dominant plant species that offered herbivores a refuge from
predation risk.
Keywords
Diversity and productivity, grasshoppers, indirect effects, old-field ecosystem, plant
diversity, predators and ecosystem function, spiders, top-down control, trait-mediated
effects, trophic cascade.
RESULTS
In the absence of herbivores and predators, S. rugosa
Figure 2 Comparison of microclimatic conditions important to
percentage cover was negatively related to plant species
plant growth in an open plot (open squares) and a paired plot
richness and evenness (Fig. 3). Because the strength of the
surrounded by aluminium sheet metal (solid squares) to evaluate the
potential for a systematic bias due to a sheet-metal effect. The upper correlations could be determined by two outlying values of
frame presents values of photosynthetically active radiation (PAR) at S. rugosa percentage cover (51 and 61%), I reanalysed the
-h sampling intervals throughout the day at 0 cm (lower curves) data after removing those values. The correlation between
and 60 cm (upper curves) above the ground (the trends for 20 and species richness and S. rugosa cover and evenness and
40 cm are similar). Lower frames present daily mean 1 SE cover declined to 0.42 (P < 0.03) and 0.65 (P < 0.02),
(n 19) wind speeds and ambient temperatures. respectively. This suggests that species evenness is a
comparatively more robust descriptor of the relationship
between plant diversity and S. rugosa abundance.
evenness is one measure that combines the two mechanistic Randomized block ANOVA on July 2001 data (chosen
factors dominance and plant species abundance that are because it represents the height of the growing season and
important in this and other old-field systems (Wilsey & thus avoids the confounding effects of plant senescence
Potvin 2000). More generally, variation in plant diversity as a beginning in August) revealed that there were significant,
result of biotic interactions among plant species often cumulative treatment effects on percentage P. pratensis
results from evenness effects (Chapin et al. 2000; Stirling & cover (P < 0.05, d.f. 2, 18) and percentage S. rugosa cover
Wilsey 2001). I quantified evenness using the standard (P < 0.02, d.f. 2, 18). Block effects were not significant
Shannon index, J (Rpi log pi) log S, where pi is the (P > 0.15). A Tukey test revealed that 3 years of predator
abundance (percentage cover) of species i in a plot. exclusion resulted in significantly lower P. pratensis and
Evenness values close to 0 indicate one or a few species S. rugosa cover relative to the one-trophic level, plant-only
dominate and many other species are rare; values close to 1 treatment (Fig. 4). Sustained predator presence (three
indicate that species are equally abundant. trophic level treatment) resulted in increased P. pratensis
As an index of productivity, I estimated dry plant cover and decreased S. rugosa cover relative to the predator
biomass production (g plot)1 d)1) by calculating the exclusion treatment. This indirect positive effect of pred-
differences in plant biomass between 31 May 2001 and ators on P. pratensis and negative indirect effect on S. rugosa
19 June 2001; and 19 June 2001 and 19 July 2001, and (Fig. 4) is consistent with previous studies in this system
dividing each difference by the intervening number of days. (Beckerman et al. 1997; Schmitz 1998), but observed at a 4
These two periods represent independent growth condi- larger scale and over a 3 longer period.
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