Journal of Ecology 2009, 97, 603–608 doi: 10.1111/j.1365-2745.2009.01503.x
© 2009 The Authors. Journal compilation © 2009 British Ecological Society
Blackwell Publishing Ltd
FORUM
Species diversity and productivity: why do results of
diversity-manipulation experiments differ from natural
patterns?
Lin Jiang
1
*, Shiqiang Wan
2
and Linghao Li
2
1
School of Biology, Georgia Institute of Technology, 310 Ferst Drive, Atlanta, GA 30332, USA; and
2
Key Laboratory
of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
Summary
1. Experiments that directly manipulate species diversity often report a positive diversity effect on
productivity, whereas observations of natural communities reveal various productivity–diversity
relationships and nutrient addition to natural plant communities generally results in negative
productivity–diversity relationships.
2. We hypothesize that this apparent paradox may be potentially explained by the reduced roles
of complementarity and positive selection effects, and the increased importance of competitive
exclusion in natural communities compared to diversity-manipulation experiments. This hypothesis
arises from the difference in species distribution and abundance patterns between immature
synthetically assembled communities in diversity-manipulation experiments and more mature
natural communities.
3. Our hypothesis applies best to small-scale studies within homogenous habitats and comple-
ments the environmental heterogeneity hypothesis that explains diversity–productivity patterns
across heterogeneous habitats.
4. Synthesis. Our analysis highlights important differences between synthetic communities in
diversity-manipulation experiments and natural communities that may translate into different
diversity–productivity patterns, and cautions against indiscriminate extrapolations of results
of diversity-manipulation experiments to natural communities.
Key-words: biodiversity and ecosystem functioning, competitive exclusion, complementarity,
environmental heterogeneity, productivity, selection effects, species diversity
Introduction
During the past two decades ecologists have devoted con-
siderable effort to understanding functional significances of
biodiversity (Kinzig et al. 2002; Loreau et al. 2002; Hooper
et al. 2005; Srivastava & Vellend 2005). Much of the experimental
work in this area has been conducted in synthetically assembled
plant communities involving direct manipulations of plant
diversity. This research has thus far revealed generally positive
effects of biodiversity on ecosystem functioning and in
particular positive effects of plant species diversity on primary
productivity (Balvanera et al. 2006; Cardinale et al. 2006, 2007).
This positive relationship between plant species diversity and
productivity, however, is in striking contrast with emerging
patterns in natural communities that are revealed by studies
exploring productivity as a driver of species diversity – a
perspective that has historically received more attention (e.g.
Hutchinson 1959; Connell & Orias 1964). Such studies include
observations of diverse forms of productivity–diversity
relationships (Waide et al. 1999; Mittelbach et al. 2001; Gillman
& Wright 2006), and nitrogen fertilization experiments that
reported nearly ubiquitous negative productivity–diversity
relationships (Gough et al. 2000; Crawley et al. 2005; Suding
et al. 2005). Understandably, these patterns of different
diversity–productivity relationships have led to debates on
whether results of diversity-manipulation experiments are
relevant for natural communities (Thompson et al. 2005;
Hector et al. 2007) and some conceptual models have been
proposed to reconcile these different patterns (Loreau et al.
2001; Schmid 2002; Hector et al. 2007). The central message *Correspondence author. E-mail: lin.jiang@biology.gatech.edu