River Restoration, Habitat Heterogeneity and Biodiversity: A Failure of Theory or Practice?

1. Stream ecosystems are increasingly impacted by multiple stressors that lead to a loss of sensitive species and an overall reduction in diversity. A dominant paradigm in ecological restoration is that increasing habitat heterogeneity (HH) promotes restoration of biodiversity. This paradigm is reflected in stream restoration projects through the common practice of re-configuring channels to add meanders and adding physical structures such as boulders and artificial riffles to restore biodiversity by enhancing structural heterogeneity. 2. To evaluate the validity of this paradigm, we completed an extensive evaluation of published studies that have quantitatively examined the reach-scale response of invertebrate species richness to restoration actions that increased channel complexity ⁄HH. We also evaluated studies that used manipulative or correlative approaches to test for a relationship between physical heterogeneity and invertebrate diversity in streams that were not in need of restoration. 3. We found habitat and macroinvertebrate data for 78 independent stream or river restoration projects described by 18 different author groups in which invertebrate taxa richness data in response to the restoration treatment were available. Most projects were successful in enhancing physical HH; however, only two showed statistically significant increases in biodiversity rendering them more similar to reference reaches or sites. 4. Studies manipulating structural complexity in otherwise healthy streams were generally small in scale and less than half showed a significant positive relationship with invertebrate diversity. Only one-third of the studies that attempted to correlate biodiversity to existing levels of in-stream heterogeneity found a positive relationship. 5. Across all the studies we evaluated, there is no evidence that HH was the primary factor controlling stream invertebrate diversity, particularly in a restoration context. The findings indicate that physical heterogeneity should not be the driving force in selecting restoration approaches for most degraded waterways. Evidence suggests that much more must be done to restore streams impacted by multiple stressors than simply re-configuring channels and enhancing structural complexity with meanders, boulders, wood, or other structures. 6. Thematic implications: as integrators of all activities on the land, streams are sensitive to a host of stressors including impacts from urbanisation, agriculture, deforestation, invasive species, flow regulation, water extractions and mining. The impacts of these individually or in combination typically lead to a decrease in biodiversity because of reduced water