1. Two predominant anthropogenic impacts on ecosystems, nutrient enrichment and the removal of consumers, are predicted to interact in their effects on producer diversity. Yet, measures of diversity alone may not capture changes occurring in the underlying mechanisms structuring communities. Furthermore, evidence for these interactions in rocky intertidal systems is mixed and may be confounded by variable baseline productivity or short experimental durations that do not capture seasonality, environmental heterogeneity or successional processes. 

2. We conducted a 2-year experiment examining the main and interactive effects of nutrients and herbivores in a low-productivity environment on the northern shore of Cook Strait near Wellington, New Zealand. We explicitly accounted for the small-scale spatial environmental heterogeneity characteristic of rocky shores. 

3. Rapid and dramatic shifts in diversity metrics and cover of algal communities in the first year were driven by removal of herbivores. Within two months of herbivore exclusion, open space decreased from > 90% to < 15%, regardless of nutrient addition, and both algal diversity and evenness plummeted while species richness rose. 

4. In contrast to theoretical predictions, strong interactions between top-down and bottom-up forces on diversity were rare; rather, over time, the addition of nutrients negatively impacted diversity. Further, where herbivores were removed, nutrients inhibited succession beyond ephemeral forms, contrasting with the tolerance model of succession observed under ambient nutrients. 

5. The effects of nutrients were only elucidated by accounting for variation in habitat complexity and seasonality, demonstrating that environmental heterogeneity can mask bottom-up processes. Moreover, shifts in relative species abundances revealed that after 10 months, nutrient addition caused increased community turnover regardless of herbivore presence or absence, indicating reduced stability not apparent in simple measures of diversity. 

6. Synthesis. Our results suggest that nutrient addition and herbivore reduction exert strong but mostly independent control on community biodiversity, where context dependency in habitat complexity, seasonality, and successional stage played key roles in determining community outcomes. These findings demonstrate the importance of longer-term field studies incorporating environmental context and multiple metrics of diversity to reveal the mechanisms underlying effects of nutrient enrichment and herbivore loss on producer community structure.

Clausing_readme

Read me file with abbreviations and explanations for all data files.

Clausing_metadata

This file provides treatment information and measures of environmental parameters for all experimental plots.

Clausing_algal_count_data

This file represents the raw algal community data from experimental plots using the point-intercept surveys. Experimental treatments of nutrient enrichment and herbivore removal were established at the beginning of March 2010 at which time algal surveys were conducted (t0). Surveys were conducted monthly for the first 4 months and quarterly thereafter for the two-year experimental period. 

Clausing_herbivore_data

This file contains the raw herbivore survey data from experimental plots. Experimental treatments of nutrient enrichment and herbivore removal were established at the beginning of March 2010 at which time herbivores were surveyed (time 0) and then  densities were manipulated. Subsequent surveys were conducted monthly for the first 4 months and quarterly thereafter for the two-year experimental period.