title: Responses of terrestrial arthropods to air pollution: a meta-analysis.
abstract: OBJECTIVE
  Arthropods, with over a million species described, are ubiquitous throughout different environments. Knowledge of their responses to human impact is crucial for understanding and predicting changes in ecosystem structure and functions. Our aim was to investigate the general patterns and to identify sources of variation in changes of the diversity, abundance and fitness of terrestrial arthropods (including Arachnida, Collembola and Insecta) in habitats affected by point polluters.
  
  
  METHODS
  We found 134 suitable studies which were published between 1965 and 2007. These data came from impact zones of 74 polluters in 20 countries with the largest representation from Russia (28 polluters), Poland (12 polluters) and the USA (six polluters). The database allowed calculation of 448 effect sizes (i.e. relative differences between measurements taken from polluted and control sites) on the effects of various point polluters like non-ferrous industries, aluminium plants, cement, magnezite, fertilising and chemical plants, power plants, iron- and steel-producing factories. We used meta-analysis to search for general effects and to compare between polluter types and arthropod groups, and linear regression to describe the latitudinal gradient and to quantify relationships between pollution and arthropod responses.
  
  
  RESULTS
  The overall effect of pollution on arthropod diversity did not differ from zero. However, species richness of soil arthropods (both living on the soil surface and within the soil) tended to decrease, and species richness of herbivores to increase, near point polluters. Abundance of terrestrial arthropods near point polluters decreased in general. This decrease resulted from strong adverse effects on soil arthropods, especially on decomposers and predators. Densities of herbivores increased, but a number of research biases that we discovered in published data may have led to overestimation of the latter effect. The dome-shaped density pattern along pollution gradients was discovered only in 5% of data sets. Among herbivores, only free-living defoliators and sap-feeders demonstrated higher densities in polluted sites; the effects of pollution on other guilds were not significant. Near the polluters, conifers suffered higher increase in damage from herbivores than deciduous woody plants and herbs. Overall effect of pollution on arthropod performance was negative; in particular, individuals from polluted sites were generally smaller than individuals from control sites. This negative effect weakened with increase in duration of the pollution impact, hinting evolution of pollution resistance in populations inhabiting polluted sites. Stepwise regression analysis demonstrated that pollution-induced changes in both the density and performance of arthropods depended on climate of the locality. Negative effects on soil fauna increased with increase in annual precipitation; positive effects on herbivore population density increased with increases in both mean July temperature and annual precipitation.
  
  
  CONCLUSIONS
  We detected effects of research methodology on the outcome of published studies. Many of them suffer from research bias-the tendency to collect data on organisms or under conditions in which one has an expectation of detecting significant effects. Pseudoreplicated studies (one polluted site contrasted to one control site) frequently reported larger effects than replicated studies (several polluted sites contrasted with several control sites). These methodological flaws especially influenced herbivory studies; we conclude that increase in herbivory in both heavily and moderately polluted habitats is not as frequent as it was earlier suggested. In contrast, the decrease in abundance of predators is likely to be a widespread phenomenon. Thus, our analysis supports the hypothesis that pollution may favour herbivore populations by creating an enemy-free space. Consistent declines in abundance of soil arthropods in impact zones of different polluters suggest that this group can potentially be used in bioindication of pollution-induced changes in terrestrial ecosystems.
  
  
  CONCLUSIONS
  Main effects of pollution on arthropod communities (decreased abundance of decomposers and predators and increased herbivory) may have negative consequences for structure and services of entire ecosystems. Responses of arthropods to pollution depend on both temperature and precipitation in such a way that ecosystem-wide adverse effects are likely to increase under predicted climate change.
  
  
  CONCLUSIONS
  Our analysis confirmed that local severe impacts of industrial enterprises on biota are well-suited to reveal the direction and magnitude of the biotic effects of aerial pollution, as well as to explore the sources of variation in responses of organisms and communities. Although we analysed the effects of point polluters, our conclusions can be applied to predict consequences of pollution impacts on regional and even global scales. We argue that possible interactions between pollution and climate should be accounted for in the analyses of global change impacts on biota.
authors.affiliation: Section of Ecology, Department of Biology, University of Turku, FI-20014, Turku, Finland. elezve@utu.fi
authors.forename: Elena L
authors.initials: EL
authors.lastname: Zvereva
authors.forename: Mikhail V
authors.initials: MV
authors.lastname: Kozlov