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 PUBLICATIONS

I Vacht, P., Puusepp, L., Koff, T., & Reitalu, T. (2014). Variability of riparian soil diatom communities and their potential as indicators of anthropogenic disturbances. Estonian Journal of Ecology, 63(3), 168–284.          

             

               

                                                            

           

                                                                                                                                                                                                                                                                                          

         



                                     

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73     

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                      



74    

                                                                    

 

                            ′  ′                          ′  ′                                                                                                                                                                      

                



75     

                                              

             

      

                                                                                                                                                                                                                                    

                                                                 



76    

                                                                   

  

                                                                                               

              

    

                                                                                   ×                                                                                                            

   

                           

                           



77     

                                                                          

 

                       

  ′   ′                                                                                                                                   ρ                                                                                                                                                                                

  

                        



78    

       ±                                 

                         

  ±   ±   ±          ±   ±   ±          ±   ±   ±           ±   ±   ±           ±   ±   ±           ±   ±   ±            ±   ±   ±           ±   ±   ±           ±   ±   ±       

                                ±           ±                                    ±    ±      ≥ 

      ±    ±       ≥                                                                                                        ρ                                 ρ             ρ           

           ρ     

   

                  ±    

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79     

   ′    ±         ±                                                                                             ±      ±      ±              ± 

      ±                                             ± 

          ±                      

              

    ±   ±   ±      ±   ±   ±      ±   ±   ±      ±   ±   ±      ±   ±   ±      ±   ±   ±      ±   ±   ±      ±   ±   ± 

  ±   ±   ±                                

          ± 

       

  ±   ±   ±    ±   ±   ±    ±   ±   ± 

  ±   ±   ± 

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80    

                                   ±                   ±        ±          ±      ±                          ±                                                                                                                                                                                                         ±         ±                                                                                                                                                                                                               

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81     

                                           

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                       

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82    

                            

    

                                                       

                                                                                                                                                                                                                       

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83     

                                                                                                                                                    ρ     ρ                  

              

                                                                                                                       ±       ±                    ±    ±  

            

                                                                         



                                    

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84    

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                   č                                                         

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85     

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                       



86    

                                         



                                                                                                                                                      



                                               



                                                                                                                                           

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87     

                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                              

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88    

                                                       č                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                              

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89 90 II Vacht, P., Puusepp, L., & Koff, T. (2018). The use of oribatid mites and diatoms as combined indicators of contaminations from multiple origins in riparian zone forest soils in Estonia. Baltic Forestry, 21(1), 24–35. BALTIC FORESTRY

THE USE OF ORIBATID MITES AND DIATOMS AS COMBINED INDICATORS /.../ P. VACHT ET AL. The Use of Oribatid Mites and Diatoms as Com- bined Indicators of Contaminations from Multi- ple Origins in Riparian Zone Forest Soils in Es- tonia

PIRET VACHT A, LIISA PUUSEPP AB AND TIIU KOFF AB a School of Natural Sciences and Health, Tallinn University, Narva Rd 25, 10120, Tallinn, Estonia b Institute of Ecology, Tallinn University, Uus-Sadama 5, 10120, Tallinn, Estonia Corresponding author: Piret Vacht, Narva Rd 25, 10120, Tallinn, Estonia; +372 56 911 704, Email: [email protected]

Vacht, P., Puusepp, L. and Koff, T. 2018. The Use of Oribatid Mites and Diatoms as Combined Indicators of Contaminations from Multiple Origins in Riparian Zone Forest Soils in Estonia. Baltic Forestry 24(1): 24-35.

Abstract

This study investigates the combined use of oribatid mites and diatoms in pine forest riparian zone soils. We focus on these organism groups as bioindicators of long-term anthropogenic disturbances of various origins categorized as two contamination levels: 1) moderate level that integrates the effects of sulphate-rich mining water contamination and alkaline air pollution that affected the area in the 1980s; 2) mild level that has been affected only by the alkaline air pollution. Additionally, the oribatid mite and diatom communities of these two groups of study sites were compared to another study area, which has similar natural conditions but is uninfluenced by these anthropogenic pollutants. Changes in oribatid mite abundance and the presence or absence of specific diatom species were the most significant differences in comparison of the two contamination levels and the area with very low human impact. Based on the community dissimilarities, both bioindicators had difficulty differentiating between the contamination levels when viewed separately. However, when the community data of the two groups were pooled, their ability to indicate the studied contami- nation levels improved noticeably. The sulphate-rich mining water contamination showed no strong influence on the communities, therefore the full indicative potential of these two groups may not have been reached. Nevertheless, this study provided new information about the soil communities inhabiting riparian areas and through that helped us understand some of the dynamics in diatom and oribatid mite communities brought on by long-term but weak disturbances. Some promising species were proposed from both groups that might help predict the effects of sulphate-rich mining water contamination or alkaline air pollution. Also, the taxonomic list of oribatid mites provides a valuable insight into the Estonian riparian forest soil oribatid fauna.

Keywords: oribatid mites, diatoms, riparian zone, disturbances, anthropogenic influence.

Introduction dressing various disturbances such as trampling (e.g. Borcard and Matthey 1995) and draining (e.g. Markkula Oribatid mites (Acari: Oribatida) and diatoms 1986). However, a wide range of topics has still not been (Bacillariophyceae) are microscopic, highly species rich addressed, leaving vast knowledge gaps in their habitat and diverse components of the soil biota and are known preferences, ecology and sensitivity or tolerance towards to respond to environmental changes in their habitat. various disturbances (Lebrun and Van Straalen 1995). Oribatid mites, mostly known for being the dominant Diatoms and their bioindication abilities have been al- microarthropod group in the organic horizons of many most exclusively studied in marine and freshwater eco- soil ecosystems (Norton 1990), have previously been systems (e.g. Desrosiers et al. 2013, Hall and Smol 2010) studied in relation to, e.g. air quality (e.g. Kehl and but only on rare occasions from soil (e.g. Heger et al. Weigmann 1992, Seniczak et al. 2002), trace metal con- 2012, Kabirov and Gaisina 2009, Vacht et al. 2014, van tamination (e.g. Ivan and Vasiliu 2009, Skubaùa and Kafel Kerckvoorde et al. 2000) and wetland habitats (e.g. 2004), and forest management (e.g. Farská et al. 2014, Hargan et al. 2014, Poulíèková 2013). Lindo and Visser 2004). Also, the bioindication abilities Even though oribatid mites have been investigated of semi-aquatic oribatid mites have been studied, ad- more than diatoms living in the soil, the information about

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THE USE OF ORIBATID MITES AND DIATOMS AS COMBINED INDICATORS /.../ P. VACHT ET AL. both groups as bioindicators of anthropogenic distur- Materials and Methods bances in riparian habitats is scarce. In paleolimnological studies, two groups have been previously used together Site description as bioindicators (e.g Luoto et al. 2009), though in these The study areas are located in two kame fields: multi-proxy studies the full indicative potential of Kurtna, located in NE Estonia (59°16’ N, 79 27°34’ E), oribatid mites is often overlooked due to their low abun- representing the contaminated area (Figure 1); and the dance in lake sediments (Solh¸y and Solh¸y 2000). area with a very low human influence, Mustoja, in SE Riparian areas, which are known as biodiversity Estonia (57°53’ N 27°39’ E). Kurtna kame field was af- ‘hotspots’, are affected by changes occurring in catch- fected by alkaline air pollution with elevated levels of ment areas as well as in bodies of water through water- trace metals in the 1980s; the effects of this can be seen level fluctuations and water seepage (Grobbelaar 1983, in the moderately elevated soil pH values (Kont et al. Weilhoefer and Pan 2007). More specifically, riparian ar- 1994, 2007; Vacht et al. 2014). The sulphate content in a eas provide an ideal study system for observing distur- number of lakes (sites N1, N2, A, S and PK) in the Kurtna bances from various origins, e.g. water contamination study area has increased due to pumping of mining wa- and air pollution. Therefore, the advantages of both ters into the area (Terasmaa et al. 2013). Based on this bioindicator groups, diatoms, well-known aquatic indi- the study sites (Figure 1) can be divided into two groups cators, and oribatid mites, more known for terrestrial based on their contamination levels: (1) two of the Kurtna bioindication abilities, can be combined here. Under- study sites (J and M) that have been affected by alkaline standing the factors controlling the functioning of the air pollution, where the sulphate levels do not exceed 20 riparian soil biota is also considered fundamental for res- mg L-1 (Terasmaa et al. 2013) and can, therefore, be con- toration purposes (Fonseca and Joner 2007). sidered as mildly contaminated sites; (2) five of the Kurtna The present study investigates the effects of sul- study sites (PK, S, A, N2 and N1) that have been af- phate-rich mining waters pumped into a number of lakes fected by alkaline air pollution and the mining water con- in NE Estonia (Kurtna study area) and compares these tamination, where sulphate levels range from 160 to 270 results with other riparian zones in the vicinity, unaf- mg L-1 (Terasmaa et al. 2013) and can be considered as fected by the mining waters. These anthropogenic dis- moderately contaminated sites. In addition, three sites turbances differ from many commonly studied human in Mustoja kame field (M1, M2 and M3) which is an area influences because even though the contaminants have with no alkaline air pollution and no elevated sulphate been present for a long time (>50 years) (Rooma 1987) content. Omitting the possibility that the community dif- their levels can be considered mild. In addition, results ferences between two study areas could be induced by from the contaminated Kurtna study area are compared additional factors (e.g. water body type), the Mustoja with an area with a very low human influence located in study area was only used for comparison and not in- SE Estonia (Mustoja study area). cluded in gradient analysis. All of the study sites at Based on the knowledge about both bioindicator Kurtna and Mustoja had a similar hydric topsoil, rang- groups in question and numerous studies pointing out ing from gleyic podzols to histosols. The results of the the benefits of multi-proxy approaches in indicating con- analysis of the soil parameters measured (Vacht et al. tamination effects and other human influences on eco- 2014) are listed in Table 1. systems (Birks and Birks 2006, Lamentowicz 2015), the The vegetation on both kame fields belongs to the main hypothesis was proposed: Integrated use of dia- oligo-mesotrophic boreal forest type (Paal 1997) and tom and oribatid mite community patterns provides more Vaccinium myrtillus boreal forest site type alternating accurate bioindication than the two groups separately. with patches of drained wetlands characterized by Ledum In order to investigate the integrated use of dia- palustre L. and Pinus sylvestris L. The vegetation at the toms and oribatid mites as combined bioindicators of sampling sites reflected the variable moisture conditions sulphate-rich mining water contamination and alkaline of the riparian soils, containing Alnus incana L., air pollution, the following goals were set: (1) to analyse Equisetum fluviatile L. and Sphagnum mosses. Average the oribatid mite community structure and properties (e.g. yearly precipitation varies between 600 and 700 mm, ex- species richness, diversity and evenness) and the vari- ceeding the evaporation rate, and average yearly tem- ation of these properties depending on the level of an- perature ranges from 4.7 to 5.7 °C on both study areas thropogenic influence; (2) to compare these results to (Estonian Environmental Agency 2014). diatom community data that has been previously pub- lished (Vacht et al. 2014); (3) to assess the benefits of Sampling and chemical analyses using diatoms and oribatid mites as combined Soil microarthropods were sampled using a soil corer bioindicators. (196 cm3, 10 cm depth) in September of 2012 and 2013. Microarthropod sampling was done in five repetitions

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Figure 1. Locations of the study areas and study sites from each site, parallel to the lake or stream shore, ap- The effect of various disturbance levels (very low proximately 1–2 m from the water, where the effects of human influence, mildly contaminated and moderately water logging were clearly visible through soil moisture contaminated) on community parameters and species and vegetation characteristics. A total of 100 abundance was tested using ANOVA. Levene’s test for microarthropod samples were collected. Oribatid mites homogeneity of variances was used prior to ANOVA. were extracted by modified Berlese-Tullgren funnels un- The differences between the parameters and species til samples were dry (at least 48 hours). Adult oribatid abundance of the mildly and moderately disturbed com- mites were counted and identified to the species level, munities were determined by the Kruskal-Wallis test. The where damages made this impossible, to the genus level same test was used to compare the community param- (Weigmann 2006). Juvenile oribatid mites were excluded eters and species abundances of the two study areas. from the analysis. The preparation of 55 diatom samples Spearman correlation (H) was used to assess possible collected in 2012 has been previously described in detail associations between community parameters, abun- (Vacht et al. 2014). dances of specific species and environmental parameters. Samples for the analysis of soil properties were col- These exploratory statistical analyses were done includ- lected in September 2013. Standard methods were used ing all the identified species and measured environmen- to determine soil organic matter and carbonate content tal parameters. The relationship between the studied

(Heiri et al. 2001) and soil pH H2O. Total nitrogen, total communities and the environmental parameters in Kurtna carbon and exchangeable phosphorus contents were was investigated in more detail to understand the ef- measured using AL extraction; potassium, calcium and fects of sulphate rich mining water on riparian soil com- magnesium by AL extraction using the MP-AES. These munities. For this purpose, the covariability and signifi- analyses were conducted from a pooled sample from each cance of each variable was considered and a selection Kurtna study site and one pooled sample combining the was made amongst them: oribatid mite species compos- sites in Mustoja. ing more than 0.05% of the total abundance (18 species) and the potentially indicative diatom species (19 spe- Data analyses cies) (Vacht et al. 2014) were chosen. These data, using Microarthropod data from both 2012 and 2013 was the Hellinger transformation, along with six environmen- pooled for statistical analysis. Communities were com- tal parameters (organic matter, total nitrogen, exchange- pared by species richness, diversity (Shannon’s H’) and able phosphorous, carbonate and potassium content in evenness (Shannon’s J’). Due to high variability in the soil and sulphate content in the lake water) were included abundance of oribatid mites between sampling sites, rar- in the Canonical Correspondence Analysis (CCA) (Ter efied species richness was used to add a comparability Braak 1986). This enabled us to examine the relationship measure to the observed species richness (Heck et al. between community composition and the most signifi- 1975, Oksanen 2013a). cant environmental parameters. In order to arrange the

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THE USE OF ORIBATID MITES AND DIATOMS AS COMBINED INDICATORS /.../ P. VACHT ET AL. study sites according to their community similarities and and Nanhermannia nana (Nicolet 1855). Kurtna and Mustoja through that observe the community response towards study area shared 71% of the oribatid mite species. Mildly the anthropogenic disturbance levels in question, the and moderately contaminated sites in Kurtna shared 61% of species data of both bioindicator groups were first used the oribatid mite species. to run an average linkage cluster analysis based on the The abundance of nine oribatid mite species (Table Bray-Curtis dissimilarity index separately, followed by 2) differed significantly in comparison of the mild and the same analysis using the combined species data. For moderate contamination levels. Three of those species the CCA, cluster analysis and community parameter cal- (Tectocepheus velatus (Michael 1880); Achipteria culations (e.g. rarefied species richness and species di- coleoptrata (Linné 1758) and Minunthozetes semirufus versity) R programming environment (R Core Team 2014) (C.L. Koch 1841)) were more abundant at the moderate with the ‘vegan’ (Oksanen 2013b) and ‘MASS’ (Venables contamination level, where the sulphate concentrations and Ripley 2002) package were used, other statistical were elevated. Oppiella (Rhinoppia) hygrophila analyses were conducted in IBM Statistic SPSS 20.0. (Mahunka 1987) had significantly higher abundance at the mildly contaminate sites. The abundances of all Results oribatid mite species found from the three contamina- tion levels are presented in the Appendix. Soil conditions The mean values for the measured soil characteris- Table 2. Mean abundance (per sample, ± SE) or absence (-) tics are presented in Table 1. There was no significant of the oribatid mite species that presented a significant dif- difference in the soil characteristics by contamination ference in their abundance in comparison of the mild and level; however, significant variation could be detected moderate contamination levels in Kurtna with the ?² values. between individual sampling sites (e.g. lower organic Significance levels are reported (*: P < 0.05; **: P < 0.01; ***: P < 0.001) matter content in site N1 compared to the other Kurtna sites). The alkaline air pollution at Kurtna could be ob- Difference served through mildly elevated soil pH (4.27) com- Species Mild Moderate between the two H2O contamination contamination contamination pared to Mustoja (3.94), but values of individual sam- levels ?(Dz)  ples ranged from acidic to near neutral at Kurtna (Vacht Euphthiracarus cribrarius et al. 2014). (Berlese 1904) 0.50 ± 0.40 - 4.945* Phthiracarus laevigatus (C.L. Koch 1841) 0.70 ± 0.47 - 4.945* Table 1. Measured soil parameters (Vacht et al. 2014) and Conchogneta traegardhi (Forsslund 1947) 14.80 ± 5.66 6.00 ± 3.15 5.69* their values on the two disturbance levels in Kurtna (mild Oppiella (Rhinoppia) hygrophila and moderate contamination (and the Mustoja (very low (Mahunka 1987) 7.00 ± 3.72 0.29 ± 0.19 12.567*** Oppiella nova human influence) study area (mean ± SE or result of multi- (Oudemans 1902) 29.60 ± 11.18 11.75 ± 4.34 5.064* ple measurements from one mixed sample, true average for Tectocepheus velatus (Michael 1880) 0.30 ± 0.30 2.08 ± 0.84 4.686* pH) together with the range of sulphate content measured Achipteria coleoptrata in lake water (Terasmaa et al. 2013) (Linne 1758) 0.80 ± 0.47 3.04 ± 0.70 5.644* Minunthozetes semirufus (C.L. Koch 1841) - 1.79 ± 1.37 4.173* Study area Very low human Mild Moderate Pilogalumna crassiclava influence contamination contamination (Berlese 1914) 0.40 ± 0.31 - 4.945* Study sites M1, M2 and M3 J and M PK, S, A, N1 and N2

Organic matter content (%) 44.6 ± 10.4 35.8 ± 8.6 42.6 ± 6.4 Carbonate content (%) 0.4 ± 0.1 0.9 ± 0.2 1.3 ± 0.3 Table 3 lists the oribatid mite abundance, observed C (%) 5,7 11.5 ± 0.5 20.5 ± 2.9 species richness, values of rarefied species richness, N (%) 0,2 0.5 ± 0.0 1.1 ± 0.1 P (mg/kg) 29,1 19.1 ± 1.2 34.6 ± 6.6 Shannon’s diversity and evenness for the three contami- K (mg/kg) 42,2 101.0 ± 9.5 121.2 ± 19.4 Ca (mg/kg) 224,4 1616.2 ±111.1 4731.1 ± 832.3 nation levels and offers a comparison to the diatom com- Mg (mg/kg) 53,2 166.9 ±18.7 375.4 ±55.3 munity parameters (species richness, diversity and even- pHH2O 3,9 3,8 4,8 SO (mg/L) - <20 160-270 ness) in Table 4. Comparison of the sites with mild and 4 moderate contamination levels at Kurtna revealed, there Oribatid mite communities and their relation to were a minor increase in oribatid mite evenness and a contamination levels small decrease in their diversity with increasing contami- In total the studied riparian soils held 76 oribatid mite nation level. Also, the abundance decreased with in- species (Appendix), belonging to 50 genera. The most nu- creasing disturbance level. Mustoja study area had a merous oribatid species that composed between 5% and significantly (P = 0.049) higher oribatid mite abundance 26% of the community of a specific site were Oppiella nova (182.1 ± 40.6 per samples) than the contaminated Kurtna (Oudemans 1902), Conchogneta traegardhi (Forsslund area on the whole (90.9 ± 15.0 per sample). Oribatid mite 1947), Steganacarus (Atropacarus) striculus (Koch 1835) species richness did not vary significantly.

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THE USE OF ORIBATID MITES AND DIATOMS AS COMBINED INDICATORS /.../ P. VACHT ET AL. Table 3. Oribatid mite community proper- Very low human Moderate Study area Mild contamination ties in the two disturbance levels in Kurtna influence contamination PK, S, A, N1 and (mild and moderate contamination) and as a Study sites M1, M2 and M3 J and M comparison in Mustoja (very low human N2 Oribatid mite abundance per sample 182.07 ± 40.56 136.80 ± 35.62 71.79 ± 14.02 influence) Oribatid mite abundance per m2 ×103 92.67 ± 20.65 69.63 ± 18.13 36.54 ± 7.13 Estimated oribatid mite species richness † 18 19 19 Observed oribatid mite species richness 39 46 27 Oribatid mite Shannon's diversity 2.59 2.75 2.54 Oribatid mite Shannon's J' evenness 0.71 0.72 0.77 † rarefied species richness (Heck et al. 1975, Oksanen 2013b)

Table 4. Diatom community parameters Very low human Mild Moderate Study area (Vacht et al. 2014) in the two disturbance influence contamination contamination levels in Kurtna (mild and moderate contam- PK, S, A, N1 and Study sites M1, M2 and M3 J and M ination) and as a comparison in Mustoja N2 (very low human influence). Observed spe- Estimated diatom species richness † 21 16 18 cies richness was not measured for diatoms Diatom Shannon's diversity 2.45 1.70 2.1 Diatom Shannon's J' evenness 0.82 0.68 0.76 † rarefied species richness (Heck et al. 1975, Oksanen 2013b)

Oribatid mite and diatom community similarities between study sites The site level cluster dendrograms (Figure 2) for diatoms and oribatid mites give an insight into the com- munity similarities between the studied sites. The oribatid mite dendrogram groups together two moderately con- taminated sites (N1 and N2) and also separates two other moderately contaminated sites (S and A) from the rest of the Kurtna sites. No clear separation was detected be- tween the Kurtna and Mustoja study areas. Diatom com- munities, however, form two main clusters, coinciding only on a few occasions with the contamination levels (e.g. M2 and M3 that belong to the Mustoja study area with very low human influence). Even though the Bray- Curtis dissimilarity matrix of the oribatid mite and diatom communities separately did not draw out a classification between the sampling sites in relation to the contamina- Figure 2. Results of the average linkage cluster analysis tion levels (Figure 2), the combined species data showed based on the Bray-Curtis dissimilarity matrix for diatom and a clear distinction between the study sites, and also oribatid mite communities as a community dendrogram bring- brought out some differences between the Kurtna study ing out the similarities and dissimilarities between the study sites (Figure 3). The cluster dendrogram formed three sites. See Figure 1 for the location of the study sites in the main clusters, separating some of the moderately dis- Kurtna and Mustoja kame fields turbed study sites in Kurtna (N1 and N2) from the uninfluenced Mustoja sites (M1, M2 an M3) and the rest of the Kurtna sites (J, A, M, S, PK).

Community relation to environmental parameters in the Kurtna study area Along with 18 oribatid mite and 19 diatom species, six environmental variables (sulphate content in lake water 2= 0.101, soil organic matter 2= 0.031, total nitro- gen2 = 0.009, phosphorous 2= 0.076, carbonate 2= 0.577 and potassium content 2= 0.005) were in- cluded in the CCA of the Kurtna study sites (Figure 4). Figure 3. Results of average linkage cluster analysis for com- These six environmental variables explained 30% of the bined groups based on the Bray-Curtis distance matrix in the variation in the combined species data while the first form of a community dendrogram. See Figure 1 for the loca- axis explained about 43.4% and the second 19.5% of the tion of the study sites in the Kurtna and Mustoja kame fields

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THE USE OF ORIBATID MITES AND DIATOMS AS COMBINED INDICATORS /.../ P. VACHT ET AL. constrained variability. This analysis indicated that the soil nutrient rather than organic matter content. The sig- communities were not strongly correlated with the meas- nificant (P < 0.05) correlations to various nutrient con- ured environmental parameters. Most of the moderate centrations were generally stronger (H > 0.6) for diatoms but statistically significant correlations were associated than the relationships between oribatid mite species and with organic matter (e.g. Chamobates borealis (Trägårdh the environmental parameters. 1902): H = 0.344, P = 0.043, Eupelops torulosus (Koch The oribatid species like Protoribotritia aberrans 1839): H= 0.354, P = 0.037, Steganacarus applicatus (Markel and Meyer 1959) and S. applicatus were less asso- (Sellnick 1920): H = 0.495, P = 0.003, S. striculus: ciated with the sulphate contamination. At the same time the H= 0.380, P = 0.024, S. carinatus (Koch 1841): H = 0.390, diatom species Eunotia exigua (Brébisson ex Kützing) P = 0.021). Some phthiracarid mites indicated moderate Rabenhorst, Tabellaria fenestrata (Lyngbye) Kützing, T. yet significant (p < 0.002) correlations towards changes flocculosa (Roth) Kützing and Achnanthidium kranzii in soil Ca and Mg content ( S. applicatus with Ca H = 0.4 (Lange-Bertalot) Round & Bukhtiyarova along with oribatid and with Mg H = 0.5; S. striculus with Ca H = 0.5 and with mite species O. nova (P = 0.024) and N. nana (P > 0.05) were Mg H = 0.5). The diatom species were correlated with abundant when the sulphate content in the lake water was below 20 mg L-1 (Terasmaa et al. 2013), while A. coleoptrata (Linne 1758) (P = 0.018) was more abundant at higher con- centrations.

Discussion and conclusions

Even though there was only a slight difference in

soil pHH O of the two study areas, the contamination gra- dient is still2 evident through elevated concentrations of trace metals and other geochemical changes that have been mentioned by previous studies comparing these soils (Kont et al. 1994, 2007, Rooma 1987). It is also pos- sible that today the alkaline air pollution may act as a neutralizing rather than alkalizing factor together with the moderately acidic sulphate rich mining water con- tamination, therefore, lessening the effects of these dis- turbances on the soil biota. The majority of the oribatid mite community con- sisted of wetland, coniferous forest litter- and soil-dwell- .ECKHA". Results of CCA as an ordination plot of combined ing species rather than strictly aquatic species (Luxton diatom and oribatid mite data from Kurtna study area with 1972, Schatz and Behan-Pelletier 2008, Weigmann 2006). six environmental parameters. Abbreviations: Sulph, sulphate This was surprising considering the fact that, for exam- content in lake water; OM, soil organic matter content; N, total nitrogen content in soil; P, content of exchangeable phos- ple, the edges of bog lakes are known to contain numer- phorus in soil; Carb, carbonate content; K, potassium con- ous aquatic species (Seniczak et al. 2013). Their absence tent; diatom species: AE, Achnanthes exigua; AH, A. hun- in the studied riparian zones could be linked to e.g. wa- garica; AK, Achnanthidium kranzii; AUG, Aulacoseira gran- ter-level fluctuations in the lakes (Terasmaa et al. 2013) ulata; EX, Eunotia exigua; EP, E. paludosa; FL, Fragilaria causing periodical drought periods that limit the suit- leptostauron; FP, F. pinnata; FZE, F. zeilleri var. elliptica; ability of these riparian soils for aquatic acarofauna. The HA, Hantzschia amphioxys; NM, Navicula mutica; NO, N. diatom community composed of numerous species that oblonga; NA, Nitzschia amphibia; PB, Pinnularia borealis; are known to live both in aquatic and terrestrial habitats, P P, P. perirrorata; RG, Rhopalodia gibba; SL, Staurosirella freshwater species, and also several species whose en- leptostauron; TF, Tabellaria fenestrata; TFL, T. flocculosa; oribatid mite species: AC, Achipteria coleoptrata; CB, vironmental optimum has not been well researched yet Chamobates borealis; CT, Conchogneta traegardhi; ET, (e.g. P. lata) (Vacht et al. 2014). Eupelops torulosus; HR, Hypochthonius rufulus; LB, Lioch- The percentage of shared species reflects the basic thonius brevis; MS, Minunthozetes semirufus; NN, Nanher- similarities in the habitat conditions of the two areas mannia nana; OA, Oribatula tibialis; ON, Oppiella nova; PA , and the two contamination levels in Kurtna. In both cases Protoribotritia aberrans; PG, Phthiracarus globosus; PLP, there were more shared oribatid mite species (71% and Platynothrus peltifer; SS, Steganacarus striculus; SA, Stega- 61% respectively) than diatoms (51% and 54% respec- nacarus applicatus; SC, S. carinatus; SCL, Scheloribates tively) (Vacht et al. 2014), which shows that oribatid mite laevigatus; T V, Tectocepheus velatus communities were more stable throughout the sampling

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THE USE OF ORIBATID MITES AND DIATOMS AS COMBINED INDICATORS /.../ P. VACHT ET AL. sites, thus pointing out their relative robustness as ties are affected as a whole was different. This means bioindicators (e.g. Gulvik 2007, Behan-Pelletier 1999). the two bioindicators show some complimentary quali- This difference could also be caused by the absence of ties. The low magnitude of change in community diver- strictly aquatic oribatid mite species compared to dia- sity was expected, explained by previous studies imply- tom communities that contained also species commonly ing changes rather in species composition than diver- found in freshwater habitats (Vacht et al. 2014). The dif- sity due to environmental changes in riparian areas ferent response of these two bioindicator groups can (Naiman and Déchamps 1997). Considering that the ripar- also be noticed in the percentage of shared species: ian zones are often biologically diverse (e.g. Naiman and There were more shared diatom species in the two con- Décamps 1997, Seniczak et al. 2006), and species rich tamination levels in Kurtna than in the two study areas (Malanson 1993, Naiman et al. 1993), the abundance and (Vacht et al. 2014), while for oribatid mites the situation relatively low species richness in the moderately contami- was reversed. While for diatoms the change in shared nated sites could be linked to the anthropogenic distur- species was relatively minor (Vacht et al. 2014), the per- bances (Naiman and Décamps 1997). However, the extent centage of shared species for oribatid mites changed and variability of these changes implies that the studied considerably. This indicates that not only was the change disturbances had a weak effect on the communities. in shared species of the two groups reversed regarding Nevertheless, many eurytopic oribatid mite species, the contamination gradient, but the response was also such as A. coleoptrata (Linnaeus 1758) and T. velatus different in its extent. (Michael 1880), had higher abundance at the highest Due to the nature of diatom extraction methodol- contamination level (Luxton 1979), which could be con- ogy, the abundance of diatoms was not evaluated and sidered as a form of bioindication. T. velatus, for exam- only the oribatid mites were investigated from this as- ple, is known for elevated abundance under the effects pect. The results showed a clear decrease in oribatid of various anthropogenic disturbances (e.g. Gulvik 2007, mite abundance with increasing contamination level, the Skubaùa and Zaleski 2012, Ivan and Vasiliu 2009). Many abundance at the moderate contamination level differed of the oribatid mite species found are known for their roughly two-fold compared to the mild contamination abilities to accumulate trace metals (e.g. T. velatus) level. On the whole, the range of oribatid mite abun- (Skubaùa and Zaleski 2012), which could also explain their dance was lower than some studies from Poland regis- abundance at Kurtna. A. coleoptrata is not as well known tered in fairly undisturbed forest floor (Skubaùa and for its indicativeness of strong contamination, it has Marzec 201, Lósková et al. 2013) but reaching notably rather been found from mildly or moderately affected higher abundances than e.g. industrial dumps (Skubaùa sites (Skubaùa and Zaleski 2012, Caruso and Migliorini 1995). Their abundance in the moderately contaminated 2006). The species that have previously been consid- sites can be compared to Slovakian sites affected by e.g. ered sensitive to air pollution (e.g. Seniczak and clear-cutting or wildfire (Lósková et al. 2013). Therefore, Dabrowski 1995, Kehl and Weigmann 1992) e.g. the abundance of oribatid mites can be considered nota- Carabodes labyrinthicus (Michael 1879), Trichoribates bly low only on the site with moderate contamination. trimculatus (Koch 1835) are represented in too low num- The oribatid mite species richness in the mildly con- bers in order to draw specific conclusions on their taminated Kurtna sites and the Mustoja study area was indicativeness to alkaline air pollution in the Kurtna similar to the number of species found, e.g. a pine for- study area. The species indicating most significantly the ests affected by air pollution in Poland (Seniczak et al. decrease in contamination levels with its increasing abun- 1998). The influence of contamination effects on oribatid dance was O. hygrophila which can for this reason be mites cannot be interpreted solely by the observed spe- considered as a potential indicator species of anthropo- cies richness because overall the number of oribatid mite genic disturbances. However, there are no specific link- species can be considered low compared to other stud- ages connecting the species to changes in sulphate con- ies (e.g. Horwood and Butt 2000, Luptáèik 2012). Diatom centrations or alkaline air pollution. In addition, there is communities had clearly higher estimated species rich- currently very little information on its environmental pref- ness at the fairly undisturbed Mustoja study area but erences in general making it impossible to draw further the two contamination levels in Kurtna showed no sig- conclusions. nificant difference (Vacht et al. 2014). Overall, the spe- The diatom communities at the highest contamina- cies richness of neither groups appeared to be strongly tion level, and the Kurtna study area as a whole, showed influenced by the contamination levels (Vacht et al. 2014). high levels of ubiquitous diatom flora (Vacht et al. 2014). The change in diversity and evenness with increasing Some species (e.g. P. lata) decreased in their abundance contamination level in Kurtna for the two bioindicator with increasing contamination level (Vacht et al. 2014) groups showed minor and sometimes contradictory re- but in most cases only a moderate shift was observable. sults, implying that the way these indicator communi- Overall, diatom communities were characterized by their

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high variability between study sites and single samples 1987) than due to the change in sulphate concentrations. (Vacht et al. 2014). This implies that using only the ripar- The two bioindicator groups have been used in numer- ian diatom flora, with very few species potentially suit- ous occasion as multi-proxy bioindicators in able for bioindication, as an indicator of sulphate-rich paleoecologically, especially paleolimnological, studies mining waters would give insufficient results. (e.g. Luoto 2009, Solh¸y and Solh¸y 2000). However, in Oribatid mite communities in riparian areas are gen- these studies, diatoms are usually the main indicating erally extensively studies; and the existing research (e.g. agents, while the use of species-specific bioindicator Seniczak et al. 2013, Seniczak 2011) does not mention qualities of oribatid mites are often overlooked because linkages between the communities nor the specific spe- of their low abundance (Solh¸y and Solh¸y 2000). In cies encountered in this study and sulphate concentra- soil ecosystem the situation is reversed, the abundance tions. Previous studies have reported that diatom com- of oribatid mites exceeding that of diatoms. This study munities respond to changes in the sulphate content confirmed that in riparian areas the two groups are pre- (e.g. De la Rey et al. 2004), but the species-specific sented in sufficient numbers creating a suitable envi- indicativeness of these observations is less studied (e.g. ronment for studying their combined bioindication abili- Luis et al. 2012). Our earlier detailed analysis of the dia- ties. Even though interface environments such as ripar- tom community structure and parameters in relation to ian zones are particularly sensitive to environmental lake water sulphate concentrations (Vacht et al. 2014) change (Malanson 1993, Naiman and Déchamps 1997), it noted R. gibba (Ehrenberg) Müller and F. zeilleri var. also true that the riparian zones are challenging ecosys- elliptica (Gasse) as species that react positively to el- tems for studying soil bioindicators in general due to evated sulphate concentration levels. Some research accelerated decomposition rates and the fragile balance suggests that sulphates may influence the diatom com- between nutrient buffering (Naiman and Déchamps 1997) munity through nutrient uptake (Saros and Fritz 2000). and rapid nutrient release (Edmonds 1980) adding to the The cluster analysis showed clearly that based on possible changes in environmental parameters. There- the community structure and through that the similari- fore, even though these ecosystems are suitable for the ties of specific sites; the differentiations between con- combined use of oribatid mites and diatoms, riparian zone tamination levels were less obvious when viewed solely soils also present a series of challenges in interpreting through diatom or oribatid mite communities than when the effects of anthropogenic disturbances on the com- observed through the combined use of the two indicator munity patterns. This could explain the reason why the groups. However, even then, the difference was only differentiation between the disturbances from various clear enough to separate the communities on two study origins remained weaker than originally anticipated. areas, and one is less clear between the two contamina- In conclusion, it can be indicated that oribatid com- tion levels in Kurtna. Therefore, it is likely that the sul- munities express significant differences between the mild phate concentrations are not the primary environmental and moderate contamination levels through changes in factors controlling the oribatid mite and diatom commu- their abundance and some species-specific variation. nities in these riparian soils. While oribatid communities contained mostly litter- and Neither of the studied bioindicators is flawless, in- soil-dwelling species rather than strictly aquatic spe- dicating immaculately the sulphate concentrations nor cies, diatom communities contained both aquatic and the three categorical contamination levels that were in- aerial species. This provided new insights into the di- vestigated. The high variability of these communities, versity of diatoms and oribatid mites in riparian zone presence of species whose species-specific indication forest soils in Estonia, proposing also some species from properties are not yet well understood and the unex- both groups as potential indicators of sulphate-rich min- pectedly weak influence of the measured environmental ing water contamination. The two bioindicator groups parameters on the community as a whole, present diffi- reacted differently to the studied disturbances meaning culties in interpreting the two indicator groups. While that the groups carry complementary information. Based some information exists on aquatic Oribatida indicating on the cluster analysis and comparative analyses of the changes in lake water (Solh¸y and Solh¸y 2000), there is changes in community parameters and structure in re- a knowledge gap in understanding the effects of water gards to contamination levels the integrated use of dia- contamination on mainly terrestrial oribatid communi- toms and oribatid mites provided more accurate ties. Based on previous knowledge and data gained from bioindication that the two groups separately. Their com- this study, it is currently possible to rather observe a bined use enabled the two study areas to be roughly shift in oribatid communities due to the difference be- separated, although based on the community structure tween the two study areas, that were differentiable of combined species data no clear separation could be through the long-term effects of alkaline air pollution made between the Kurtna sites with lower sulphate con- also rich in trace metals (Kont et al. 1994, 2007, Rooma centrations in lake water and the elevated ones. Even

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THE USE OF ORIBATID MITES AND DIATOMS AS COMBINED INDICATORS /.../ P. VACHT ET AL. though riparian zone soils are suitable study systems Farská, J., Prejzková, K. and Rusek, J. 2014. Management for the combined use of diatoms and oribatid mites, each intensity affects traits of soil microarthropod communi- ty in montane spruce forest. Applied Soil Ecology 75: carrying their own strengths as bioindicators, these ar- 71–79. eas offer challenges for interpreting the effects of long- Fonseca, C.R. and Joner, F. 2007. Two-sided edge ef fect term but mild anthropogenic disturbances on these studies and the restoration of endangered ecosystems. groups. More specifically, at the current magnitude the Restoration Ecology 15: 613–619. Grobbelaar, J.U. 1983. Availability to algae of N and P ad- disturbances may not have been sufficiently strong to sorbed on suspended solids in turbid waters of the Ama- bring out the full indicative potential of these indicator zon River. Archiv für Hydrobiologie 96: 302–306. groups. Gulvik, M. 2007. Mites (Acari) as indicators of soil biodi- versity and land use monitoring: a review. Polish Journal of Ecology 55: 415-440. Acknowledgments Hall, R.I. and Smol, J.P. 2010. Diatoms as indicators of lake eutrophication. In: J.P. Smol and E.F. Stoermer (Eds.): We are grateful for financial support from the En- The Diatoms: Applications for the Environmental and vironmental Conservation and Environmental Technol- Earth Sciences. Cambridge University Press, Cambridge, p. 122–151. ogy R&D Program Project “EDULOOD”, Tallinn Uni- Hargan, K.E., Rühland, K.M., Paterson, A.M., Finkelstein, versity’s Centre of Excellence initiative “Studies of natu- S.A., Holmquist, J.R., MacDonald, G.M., Keller, B. ral and man-made environments”, The Estonian Sci- and Smol, J.P. 2014. The influence of water table depth ence Foundation (grant ETF8189), the Japan Society and pH on the spatial distribution of diatom species in peatlands of the Boreal Shield and Hudson Plains, Cana- for the Promotion of Science (JSPS ID No. P13758), da. Botany 93: 57–74. Tartu Cultural Endowment, The Hoogstraal Memorial Heck, K.L., van Belle, G. and Simberloff, D. 1975. Ex- Acarolgy Student Fund and European Social Fund’s plicit calculation of the rarefaction diversity measurement Doctoral Studies and Internationalization Program and the determination of sufficient sample size. Ecology 56: 1459–1461. DoRa, carried out by Foundation Archimedes. We thank Heger, T.J., Straub, F. and Mitchell, E.A.D. 2012. 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Appendix. Mean abundances per sample (± SE, 196 cm 3, 10 cm depth, sampled in Septem- ber 2012 and 2013) or absences (-) of oribatid mites at the very low human influence level in Mustoja (sites M1, M2 and M3, n = 30), mildly contaminated sites in Kurtna (J and M, n = 20) and moderately contaminated sites in Kurtna (PK, S, A, N1 and N2, n = 50)

 Very low Mild Moderate Species human contamination contamination influence Brachychthonius berlesei (Willmann, 1928) 0.13 ± 0.13 - 0.08 ± 0.06 Liochthonius brevis (Michael, 1888) - 4.7 ± 3.89 0.38 ± 0.29 Liochthonius furcillatus (Willmann, 1942) - 0.50 ± 0.5 - Hypochthonius rufulus (C.L. Koch, 1835) 0.20 ± 0.10 9.10 ± 5.18 3.58 ± 1.17 Eniochthonius minutissimus (Berlese, 1903) - - 0.08 ± 0.06 Eulohmannia ribagai (Berlese, 1910) - 0.30 ± 0.30 0.04 ± 0.04 Phthiracarus ferrugineus (C.L. Koch, 1841) 0.47 ± 0.19 0.90 ± 0.61 1.00 ± 0.35 Phthiracarus globosus (C.L. Koch, 1841) 0.73 ± 0.28 3.20 ± 1.41 1.92 ± 0.44 Phthiracarus laevigatus (C.L. Koch, 1844) 0.33 ± 0.33 0.70 ± 0.47 - Steganacarus applicatus (Sellnick, 1920) - 0.80 ± 0.44 1.13 ± 0.40 Steganacarus (Tropacarus) carinatus (C.L. Koch, 7.20 ± 2.03 3.60 ± 2.63 6.29 ± 2.12 1841) Steganacarus magnus (Nicolet, 1855) 0.27 ± 0.15 0.70 ± 0.52 0.13 ± 0.07 Steganacarus (Atropacarus) striculus (C.L. Koch, 11.47 ± 7.87 12.70 ± 4.29 6.58 ± 1.90 1835) Euphthiracarus cribrarius (Berlese, 1904) 0.07 ± 0.07 0.50 ± 0.40 - Microtritia minima (Berlese, 1904) 5.87 ± 2.81 0.10 ± 0.10 - Rhysotritia ardua (C.L. Koch, 1841) - 0.20 ± 0.13 0.04 ± 0.04 Protoribotritia aberrans (Märkel & Meyer 1959) 1.40 ± 0.52 0.50 ± 0.17 2.13 ± 0.98 Platynothrus peltifer (C.L. Koch, 1839) 5.60 ± 2.76 4.40 ± 2.67 2.46 ± 0.53 Heminothrus longisetosus (Willmann, 1925) - - 0.08 ± 0.06 Nothrus borussicus (Sellnick, 1928) 1.87 ± 0.79 - 0.04 ± 0.04 Nothrus palustris (C.L. Koch, 1839) 1.73 ± 0.80 1.10 ± 0.64 0.13 ± 0.09 Nothrus pratensis (Sellnick, 1928) 0.20 ± 0.10 0.30 ± 0.21 0.46 ± 0.23 Nothrus silvestris (Nicolet, 1855) 10.67 ± 3.89 1.80 ± 1.04 0.38 ± 0.16 Malaconothrus monodactylus (Michael, 1888) 0.07 ± 0.07 0.40 ± 0.40 0.04 ± 0.04 Nanhermannia nana (Nicolet, 1855) 23.60 ± 14.12 20.60 ± 11.41 2.71 ± 1.05 Hermannia gibba (C.L. Koch, 1839) - - 0.04 ± 0.04 Hermanniella dolosa (Grandjean, 1931) 0.40 ± 0.19 0.30 ± 0.21 0.13 ± 0.09 Metabelba spp (Grandjean, 1936) 1.33 ± 0.33 2.00 ± 0.41 Belba spp (von Heyden, 1926) - 1.00 ± 0.00 - Damaeus grac ilipes (Kulczynski, 1902) 0.13 ± 0.13 - 0.04 ± 0.04 Spatiodamaeus verticillipes (Nicolet, 1855) 0.53 ± 0.32 0.20 ± 0.20 0.08 ± 0.06 Cepheus cepheiformis (Nicolet, 1855) - - 0.21 ± 0.17 Caleremaeus monilipes (Michael, 1882) 0.40 ± 0.21 - 0.13 ± 0.13 Liacarus spp (Michael, 1898) 1.00 ± 0.00 - - Adoristes ovatus (C.L. Koch, 1839) 0.20 ± 0.20 0.3 ± 0.15 0.25 ± 0.14

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Appendix continued) Very low Mild Moderate Species human contamination contamination influence Xenillus tegeocranus (Hermann, 1804) - 0.20 ± 0.13 - Astegistes pilosus (C.L. Koch, 1840) 1.07 ± 0.51 - 0.04 ± 0.04 Ceratoppia spp (Berlese, 1908) 1.00 ± 0.00 - - Gustavia microcephala (Nicolet, 1855) 0.07 ± 0.07 - 0.08 ± 0.06 Carabodes areolatus (Berlese, 1916) - - 0.25 ± 0.14 Carabodes labyrinthicus (Michael,1879) 0.47 ± 0.29 - 0.04 ± 0.04 Carabodes rugosior (Berlese, 1916) 12.27 ± 5.89 - 0.04 ± 0.04 Carabodes willmanni (Bernini, 1975) 1.07 ± 0.75 - 0.08 ± 0.08 Conchogneta traegardhi (Forsslund, 1947) 8.47 ± 8.33 14.80 ± 5.66 6.00 ± 3.15 Dissorhina ornata (Oudemans, 1900) - - 0.04 ± 0.04 Oppiella (Rhinoppia) hygrophila (Mahunka, 1987) 42.87 ± 12.83 7.00 ± 3.72 0.29 ± 0.19 Oppiella (Moritzoppia) translamellata (Willmann, 0.07 ± 0.07 2.50 ± 1.13 3.17 ± 1.27 1923) Oppiella nova (Oudemans, 1902) 3.00 ± 1.46 29.60 ± 11.18 11.75 ± 4.34 Tectocepheus minor (Berlese, 1903) 6.33 ± 2.84 0.20 ± 0.13 1.08 ± 0.51 Tectocepheus velatus velatus (Michael, 1880) 0.73 ± 0.37 0.30 ± 0.30 2.08 ± 0.84 Eupelops acromios (Hermann, 1804) - 0.10 ± 0.10 0.08 ± 0.06 Eupelops hirtus (Berlese, 1916) - - 0.04 ± 0.04 Eupelops tardus (C.L. Koch, 1835) 0.13 ± 0.09 0.10 ± 0.10 0.13 ± 0.09 Eupelops torulosus (C.L. Koch, 1839) 0.07 ± 0.07 0.50 ± 0.17 2.08 ± 1.49 Achipteria coleoptrata (Linné, 1758) 0.13 ± 0.13 0.80 ± 0.47 3.04 ± 0.70 Oribatula tibialis (Nicolet, 1855) 1.00 ± 0.50 0.10 ± 0.10 0.13 ± 0.13 Liebstadia pannonica (Willmann, 1951) 0.13 ± 0.09 0.40 ± 0.40 0.13 ± 0.13 Scheloribates laevigatus (C.L. Koch, 1835) 0.53 ± 0.24 0.90 ± 0.64 0.96 ± 0.49 Scheloribates latipes (C.L. Koch, 1844) 3.87 ± 1.15 0.10 ± 0.10 0.29 ± 0.15 Chamobates borealis (Trägårdh, 1902) 1.73 ± 0.63 7.50 ± 5.00 3.75 ± 1.80 Chamobates cuspidatus (Michael, 1884) 2.47 ± 1.15 - 0.04 ± 0.04 Chamobates pusillus (Berlese, 1895) - 0.10 ± 0.10 0.13 ± 0.07 Chamobates voigtsi (Oudemans, 1902) 17.93 ± 9.16 0.20 ± 0.20 0.79 ± 0.51 Ceratozetes mediocris (Berlese, 1908) 0.20 ± 0.20 0.20 ± 0.20 0.50 ± 0.27 Ceratozetes parvulus (Sellnick, 1922) 0.07 ± 0.07 0.10 ± 0.10 0.04 ± 0.04 Trichoribates trimaculatus (C.L. Koch, 1835) - - 0.04 ± 0.04 Euzetes globulus (Nicolet, 1855) 0.53 ± 0.17 0.60 ± 0.43 0.83 ± 0.50 Minunthozetes semirufus (C.L. Koch, 1841) 0.13 ± 0.13 - 1.79 ± 1.37 Mycobates bicornis (Strenzke, 1954) 0.40 ± 0.40 0.40 ± 0.22 0.54 ± 0.19 Punctoribates hexagonus (Berlese, 1908) 0.07 ± 0.07 1.10 ± 0.89 0.42 ± 0.18 Galumna flagellata (Willmann, 1925) 0.27 ± 0.12 - 0.04 ± 0.04 Galumna lanceata (Oudemans, 1900) 0.93 ± 0.45 0.40 ± 0.40 0.08 ± 0.06 Galumna obvia (Berlese, 1914) - 0.20 ± 0.20 0.25 ± 0.17 Pergalumna formicaria (Berlese, 1914) 0.27 ± 0.21 - 0.04 ± 0.04 Pergalumna nervosa (Berlese, 1914) 0.87 ± 0.27 0.10 ± 0.10 0.17 ± 0.10 Pilogalumna crassiclava (Berlese, 1914) 0.40 ± 0.29 0.40 ± 0.31 -

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104 III Vacht , P., Kuu, A., Puusepp, L., Koff, T., Kutti, S., Raamets, J., & Küttim, L. (2018). Diatoms and microarthropod communities of three airfields in Estonia – their differences and similarities and possible linkages to airfield properties. European Journal of Ecology, 4(2), 1–14. EUROPEAN JOURNAL OF ECOLOGY

EJE 2018, 4(2): 1-14, doi:10.2478/eje-2018-0008

Diatom and microarthropod communities of three airfields in Estonia – their differences and similarities and possible linkages to airfield properties

1 1 2 1,3 1,3 2 2 1,3 School of Natural Sci- Piret Vacht , Annely Kuu , Liisa Puusepp , Tiiu Koff , Sander Kutti , Jane Raamets , Liisa Küttim ences and Health, Tallinn University, Narva Rd 25, 10120, Tallinn, Estonia Corresponding author, ABSTRACT E-mail: [email protected] Even though airfields, which are often anthropologically modified natural areas, are continuously influenced by 2 School of Engineering, human activities, their soils are still dynamic ecosystems containing various habitats for microscopic groups of Tartu College of Tallinn organisms which are often ignored. In this exploratory study, the microarthropod fauna, Collembola (Hexapoda) University of Technology, and oribatid mites (Acari: Oribatida), and diatom (Bacillariophyta) flora were identified in three Estonian air- Puiestee 78, 51008, Tartu fields, both runway sides and snow-melting sites were investigated. The communities of these airfields shared 3 Institute of Ecol- approximately 10–60% of the species belonging to each studied bioindicator group. The shared species were ogy, Tallinn University, generally characteristic of a broad habitat spectrum. Communities were also characterized based on their species Uus-Sadama 5, 10120, richness and diversity and in relation to location and the purpose of different airfield areas (e.g. snow-melting Tallinn, Estonia sites vs. runway sides). Also, species indicative of a specific airfield or purpose of the area within the airfield were identified using Indicator Species Analysis. Some possible linkages between airfield properties and communities, e.g. airfield that had highest pollutant concentrations had also maintained high diversity and species richness, were noted. Despite the contamination levels the airfield soils had still maintained a functioning soil ecosystem.

KEYWORDS airfields, soil, Oribatida, Collembola, diatoms, bioindicators

© 2018 Piret Vacht et al. This is an open access article distributed under the Creative Commons Attribution-NonCommercial-NoDerivs license

INTRODUCTION et al. 2003). Research on urban microarthropod communities Civil aviation is a fast-growing industry driven by globalisa- are more common. The subjects range from roadside soils tion. Previously the environmental impacts of aircraft emis- (Eitminavičiūtė 2006a,b; Magro et al. 2013) to urban soil qual- sions (Moussiopoulos et al. 1997; Rissman et al. 2013), noise ity assessment (Santorufo et al. 2012). (Vanker et al. 2013), de-icing chemicals (Turnbull & Bevan Soil mites, including Oribatida (Acari), are considered 1995) and polycyclic aromatic hydrocarbons (Ray et al. 2008) fairly resistant towards anthropogenic activities (Skubała1997 ; have been studied. Also, airfield vegetation and wildlife studies Khalil et al. 2009; Santorufo et al. 2012), however their species are common (Servoss et al. 2000; Blackwell et al. 2008, 2009; specific response to various disturbances and environmental Belant et al. 2013; Fischer et al. 2013). However, in the con- changes make them valuable bioindicators (Paoletti & Bressan text of airport soils and anthropogenic soils in general, there 1996). Other soil microarthropods such as (Hexapo- is less information (Pytka et al. 2003; Baran et al. 2004; Ray da: Collembola) and other mite orders are considered less sen- et al. 2008; McNeill & Cancilla 2009; Werkenthin et al. 2014). sitive towards traffic related environmental changes, though Even more scarcely studies on airfield soil biota have been con- this does not lessen their importance as potential bioindicators ducted. Among the few exceptions is a study exploring the mi- (Paoletti & Bressan1996 ; Eitminavičiūtė 2006a), e.g. when and croarthropod communities in the area of Orio al Serio Airport where other groups are only present in low numbers. Many in Bergamo, Italy (Migliorini et al. 2003) and another studying microarthropod species are also known for tolerating exposure the development of earthworm communities on translocated to heavy metals due to their accumulative properties, which

grassland turf from Manchester airport, United Kingdom (Butt makes them suitable for studying aviation related disturbances European Journal of Ecology

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(Van Straalen & Van Wensem 1986; Heikens et al. 2001). Dia- 1. MATERIAL AND METHODS toms, mostly known from their bioindication abilities in water (Dickman 1998), are so far less used for in soil ecological re- 1.1. Study sites search, but can still be considered as potential indicators of soil Three Estonian airfields (Fig. 1) were selected based on their habitat properties (van Kerckvoorde et al. 2000; Van de Vijver accessibility, aviation history, availability of background infor- et al. 2008; Heger et al. 2012; Vacht et al. 2014). mation and natural conditions for studying soil communities. Based on previous studies from urban areas and traf- The main characteristics of these three airfields are shown in fic related disturbances (Santorufo et al. 2012) the following Table 1. Based on the soil maps provided by Estonian Land -hypotheses were set: Although, microarthropod and diatom Board (http://geoportaal.maaamet.ee/est/Kaardiserver/Mul communities vary between the airfields the communities at lakaart-p96.html), the soils of Tallinn airfield range from Rendzi the studied airfields also share a substantial percentage of spe- Leptosols to Histosols. In Tartu the soils have been classified as cies which can be considered typical to areas affected by inten- Luvisols influenced by excess moisture (gleying) and in Pärnu sive traffic related disturbances such as airfields; Diatom and airfield as Gleysols. Since their classification, several decades microarthropod (Collembola and oribatid mites) species indic- ago, they have been subject to draining in various extent. ative of specific airfields and the purpose of different airfield The three airfields have experienced aviation - relat areas can be identified. ed disturbances in various extents. For example, Tallinn air- This pilot study aims (1) to describe and analyze the field today acts as the primary airfield in Estonia with highest microarthropod fauna and diatom flora of Tallinn, Tartu and passenger and aircraft movement numbers and also receives Pärnu airfields and their community parameters; (2) to identify intensive maintenance activities. Tartu airfield has low usage the extent of the shared community of each bioindicator group intensity but has been most recently (< 15 years) reconstruct- in comparison of the three airfields and the components of the ed, including partial removal and filling of soil. Pärnu airfield, ,shared community; (3) to attempt to identify whether the stud- on the other hand, is today only used for a few regional flights ied airfield soils have maintained their functionality based on but has been under extensive military use during the Soviet their biological properties. These aims were achieved. era (1940-1991) which may have resulted in high exposure to aviation related pollutants. Therefore, it is impossible to place the airfields on a clearly definable anthropogenic disturbance gradient.

.

Figure 1. Location of study sites in north-eastern Europe (A) and in Estonia (B) and general sampling design within the airfields (C) where TL1 and TL2 mark the sampling sites on slow-melting sites, and TL3 through TL8 mark the runway side sampling sites.

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Table 1. Characteristics of Tallinn, Tartu and Pärnu airfields lot et al. 2011; Souffreau et al. 2013; Lowe et al., 2014). The nomenclature follows Krammer & Lange-Bertalot (1988; 1991) with additions from newer taxonomic literature (e.g Compère, 2001; Lange-Bertalot et al. 2011). Soil parameters were measured to characterize the airfields in general, not to link them directly to soil commu- nities, therefore, the soil samples from each airfields were pooled. These samples were analyzed at the Estonian Environ- mental Research Center with ICP-AES for Cu, Pb, Zn and Cd be- cause these metals are considered most indicative of traffic- re lated pollution (Davis et al. 2001). Samples were also analysed

for total-N, P and K, Ca, Mg and pHKCl at the Estonian Agricul- -tural Research Centre. Soil organic matter and carbonate con tent was measured using loss-on-ignition method (prepASH®, Precisa). Soil microbial respiration rates (basal respiration) and Substrate Induced Respiration (SIR) were determined using 1.2. Sampling procedure and laboratory analysis of samples manometric respirometers (Oxitop®, WTW). Sampling was conducted in September, 2013, and repeated in September 2014. In addition to six sampling points at runway 1.3. Data analysis sides also two snow-melting sites were sampled from each air- Species data from two years of sampling were pooled for nu- field. Fig. 1 shows the general sampling design at each airfield. merical analyses. Communities were described by observed Because sampling at the airfields was to some extent restricted Collembola and oribatid mite abundance per sample, pres- (e.g the allowed time for sampling and the volume of soil re- ence-absence data was noted for diatoms. Because all mi- moved from the site) eight samples were collected for this ex- croarthropod communities from the same volume of soil were plorative study from each airfield each year (in total 48 samples identified and counted, only their observed species richness of each bioindicator group). A ø 5 cm soil auger was used to were calculated. Diatom communities were described based collect samples from 0-10 cm of topsoil (196 cm3) for microar- on rarefied species richness (Heck et al. 1975; Oksanen 2012). thropod (oribatid mites and springtails) and diatom extraction, All bioindicator groups were characterized based on communi- microbial measurements and soil chemical analysis. ty diversity (Shannon’s H). Also the Oribatida Collembola ratio Microarthropods were extracted using Berlese-Tull- (O/C) was calculated for each airfield. The number of repeti- gren funnel until samples were fully dry (at least 72 hours) tions for soil chemical and microbial analysis was limited due and stored in 90% ethanol. For clearing purposes 80% lactic to fieldwork restrictions and therefore this data was only ana- acid was used for oribatid mites. Samples were hand sorted lyzed using descriptive statistical methods to give an overview and identified by species. Keys by Fjellberg (1998; 2007) and on general soil characteristics. Hopkin (2007) were used to identify adult Collembola. The no- Kruskal-Wallis test, ANOVA, and average linkage clus- menclature follows Bellinger et al (1996-2018). For the identi- ter analysis based on Bray-Curtis dissimilarity were used to fication of adult oribatid mites keys by Weigmann (2006) and compare the communities of Tallinn, Tartu and Pärnu airfields Niedbala (2008) were used. The nomenclature follows Weig- and the different land uses within airfields (runway sides and mann (2006). snow-melting fields). Indicator Species Analysis (ISA) (Dufrene For diatom analysis soil samples (N = 24) were mixed & Legendre 1997; De Cáceres & Legendre 2009) together with and 2 cm3 of soil was extracted, then treated with 10% HCl and Monte Carlo randomization technique for testing the statistical

30% H2O2 while heating the samples to remove carbonates and significance (P < 0.05) of Indicator Values (IV) was used to iden- -organic matter. Samples were decanted and centrifuged to re- tify potential indicator species. Community parameter calcu move other unnecessary components. The solution (0.1 ml) lations, descriptive and generalising analyzes were conducted was transferred to cover glass, dried and fixed with Naphrax. using R programming environment (R Development Core Team Samples were examined under 600–1000 × magnification (us- 2014) with the ‘indicspecies’ (De Cáceres & Legendre 2009), ing immersion oil nd = 1.516). From each sample 300–500 ‘MASS’ (Venables & Ripley 2002) and ‘vegan’ (Oksanen 2013) valves were identified and counted along random transects. package, and IBM SPSS Statistics 20.0. Damaged valves were counted as a separate individual when more than half of the valve was intact. Diatoms were identi- fied to species level and when this was impossible due to dam- 2. RESULTS ages, to genus level (Krammer & Lange-Bertalot 1988, 1991; Hamilton et al. 1992; Lange-Bertalot & Metzeltin 1996; Round 2.1. Soil properties & Bukhtiyarova1996 ; Lange-Bertalot & Metzeltin,1996 ; Lange- Although the soil maps provided by Estonian Land Board show Bertalot, 1997; Krammer, 2000; Compère, 2001; Lange-Berta- some differences between the three airfield soils and allof

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them have maintained some characteristics of the original soils community not shared by all studied airfields can be considered (e.g. higher carbonate content in Tallinn) they have all been af- site specific, possibly depending on the natural conditions (e.g. fected by aviation, airfield maintenance, and aviation related hydrological regime, soil development prior to aviation related construction activities e.g. draining, tilling, removal and filling. activities) and anthropogenic history (e.g. history and place- An overwhelming area of these soils can today be classified ment of the filling soil on site, type and quantity of de-icing as Technosols (IUSS Working Group WRB 2015), characterized chemicals) of each site. These site specific species included e.g. by a dense and tightly rooted mull type humus horizon that Bourletiella arvalis Fitch, 1863 (characteristic of Tartu airfield), contains various artifacts such as pieces of glass, aluminium, Micranurida pygmaea Börner, 1902 (characteristic of Pärnu air- plastic, paint and runway construction materials. Table 2 lists field) andFolsomia quadrioculata Tullberg, 1871 (characteristic the results from soil chemical analyses, including results from a of Tallinn airfield). previous study on Tallinn airfield soil (Keskküla 2011), and soil The list of all encountered species together with microbial parameter measurements. their abundance (per sample) at each airfield, separating be- tween the runway sides and snow-melting sites is given in Ap- 2.2. Community properties pendix (Table A.1.). No significant differences were detected The results from community properties are shown in Table 3 in the species abundance in comparison of the three airfields. separating between the runway sides and snow-melting sites. In three-way ISA (Tallinn, Tartu and Pärnu airfield), springtails On Pärnu airfield the oribatid mites outnumbered the Col- were not significantly strong indicators of any of the airfields lembola (O/C ratio 3.0). In Tallinn the O/C ratio was 0.8 and in singularly, but as a combination M. affinis (IV = 0.87, P = 0.02) Tartu 0.5. and T. vulgaris (IV = 0.81, P = 0.011) were indicative of Pärnu and Tallinn airfields. 2.3. Collembola community composition On Family level, Neanuridae were significantly more In total 31 species were identified, belonging to 22 abundant at snow-melting sites compared to runway sides (χ2 = genera. The most abundant species, forming 31% of the to- 5.715, df = 1, P = 0.017). The trend was opposite for Tomoceri- tal abundance, was Protaphorura armata Tullberg, 1869. The dae (χ2 = 6.611, df = 1, P = 0.01). Significant differences in spe- snow-melting sites of the three airfields shared five springtail cies abundance depending on the purpose of the airfield area species: Anurida granaria Nicolet, 1847, Metaphorura affinis (runway side or snow-melting sites) were rare, only a few spe- Börner, 1902, P. armata, Parisotoma notabilis Schaeffer, 1896 cies showed significant difference – e.g. T. vulgaris (χ2 = 6.611, and Ceratophysella sp. Runway sides shared 11 species, includ- df = 1, P = 0.01) and L. lanuginosus (χ2 = 7.689, df = 1, P = 0.006) ing almost all of those also found from snow-melting sites (ex- that were both only found on runway sides. Two-way ISA sup- cept Ceratophysella sp). In addition, the airfield runway sides ported the assumption that these two species can be consid- also shared Folsomia candida Willem 1902, Folsomia sexocula- ered significantly indicative of airfield runway sides (IV = 0.85,P ta Tullberg, 1871, Isotoma minor Schaeffer, 1896, Isotoma viri- = 0.011 and IV = 0.82, P = 0.017 respectively). The results from dis Bourlet, 1839, Isotomurus sp., Tomocerus vulgaris Tullberg, ISA, where the sites were divided into six groups (separating 1871 and Lepidocyrtus lanuginosus Gmelin, 1788. between airfields, their runway sides and snow-melting sites) An explorative average linkage cluster analysis re- showed that T. vulgaris (IV = 0.93, P = 0.01) and L. lanugino- vealed that based on the springtail communities the three air- sus (IV = 0.93, P = 0.003) are more specifically strongly and sig- fields cannot be distinguished from each-other. The part of the nificantly associated with the combination of Tallinn and Tartu runway sides. These species, however, can not be considered ideal indicators of these runway sides because they can also be Table 2. Results from soil chemical and microbial analysis. Heavy metal and oil found on other sites in addition to the two runway sides (A = product (C10-C40) concentration result ranges for Tallinn (*) are based on previ- 0.9459 and B = 0.9429 respectively) and they cannot be found ous research (Keskküla 2011), others are based on a single result from repreated measurements (N=10) or expressed as Mean ± SE as provided by the laboratory Table 3. Collembola, oribatid mites and diatom community parameters (observed species richness for Collembola and oribatid mites, rarefied species richness for diatoms and diversity expressed by Shannon’s H on the runway sides and snow- melting sites of Tallinn, Tartu and Pärnu airfields

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. though L. pannonica inhabited only runway side soils, no sig- nificant indicators were found that could identify different land use within airfields.

2.5. Diatom community composition In total 49 diatom species were identified belonging to 30 gen- era. The most abundant diatoms were Hantzschia amphioxys (Ehrenberg) Grunow, 1880 (26.1%), Aulacoseira granulata (Eh- renberg) Ehrenberg, 1843 (12.9%) and Luticola mutica (Kützing) Mann, 1990 (9.7%). The snow-melting sites shared five species, Figure 2. Average linkage cluster analysis of the oribatid mite samples collected from runways sides, abbreviation beginning with TL refer to Tallinn airfield, TR to which in addition to the three most abundant species were Tartu airfield and P to Pärnu airfield, numbers stand for sampling sites within the Fragilaria nitzschioides (Grunow) Lange-Bertalot, 2011 and airfield as shown on Fig.1. Navicula cari (Ehrenberg) Ehrenberg, 1836. The runway sides from all the sampled sites on Tallinn and Tartu runway sides (B of the three airfields shared eight species. These included the = 0.9167 for both). most abundant diatom species mentioned previously and also Pinnularia obscura (Krasske) Patrik & Reimer, 1966, N. cari, Hu- 2.4. Oribatida community composition midophila contenta (Grunow) Lowe et al., 2014 and F. nitzschi- In total 34 oribatid mite species were identified, belonging to oides. An explorative cluster analysis did not reveal notable sim- 22 genera. The majority of species found on airfield soils belong ilarities between the diatom communities of the three airfields, to suborder Brachypylina. The most abundant species were in comparison of the snow-melting sites and runway sides. Scheloribates laevigatus Koch, 1836 and Tectocepheus velatus There were 12 diatom species that were specific to velatus Michael, 1880. These two species made up 37% of the Pärnu airfield, including e.g. Fragilariforma virescens (Ralfs) total abundance, 19% and 18% respectively. While the snow- Williams & Round, 1988 and several species belonging to the melting sites of the three airfields shared only one oribatid mite Stauroneis genus. The list of all encountered species at each air- species, T. velatus velatus, the runway sides had five species field, separating between the runway sides and snow-melting incommon: Liebstadia pannonica Willmann, 1951, Platynothrus sites is given in Appendix (Table A.3.) peltifer Koch, 1839, S. laevigatus, Trichoribates incisellus Kram- The results from six-way ISA (separating between er, 1897 and T. velatus velatus. The list of all encountered spe- airfields, their runway sides and snow-melting sites) indicated cies together with their abundance (per sample) at each air- that Pinnularia lata (Brébisson) Rabehorst, 1853 (IV = 0.985 P field, separating between the runway sides and snow-melting = 0.016), Pinnularia borealis (Ehrenberg) Ehrenberg, 1843 (IV = sites at each airfield is given in Appendix (Table A.2.). 0.957 P = 0.041) and Diploneis finnica (Ehrenberg) Cleve, 1891 No significant differences in oribatid mite species (IV = 0.913 P = 0.027) are strongly and significantly associated abundance depending on the purpose of the airfield area (run- with the snow-melting sites of Pärnu airfield. These three spe- way side or snow-melting sites) or airfield were detected (P > cies can be used to indicate the snow-melting sites of Pärnu 0.05). The explorative average linkage cluster analysis showed airfield because they appear on all sampled sites (B = 1.0000) that at the snow-melting sites the oribatid mite communities and they are largely (but not completely) restricted to them were not differentiable based on the specific airfield. The com- (A=0.9706, A=0.9158 and A=0.8333 respectively). Also three munities belonging to the runway sides, however, formed a other diatom species – F. nitzschioides, L. mutica and D. finnica) distinct group together with the communities on the Tallinn air- were significantly (P > 0.05) linked to snow-melting sites of all field, and separated them from the communities of Tartu and the studied airfields. Pärnu (Fig. 2). No significant differences were observed in the abundance of oribatid mite species in comparison on runway sides and snow-melting sites nor in comparison of the three 3. DISCUSSION airfields. The concentrations of Pb and Zn were mildly elevated (Peter- Some site specific species such as Eupelops acromios sell et al. 1997) in Tallinn airfield. Pärnu airfield had elevated Hermann, 1804 and Nothrus silvestris Nicolet, 1855, character- Zn content compared to normal levels in the region. Levels of istic only of Tallinn airfield, andPunctoribates punctum Berlese, Cd were high on all airfields (Petersell et al. 1997). Neverthe- 1908, characteristic of Pärnu airfield, were encountered. When less, the concentration of measured heavy metals and oil prod- grouping the sites into six groups (separating between airfields, ucts (C10–C40) remained lower than the limits set by Estonian their runway sides and snow-melting sites) the ISA did not re- Environmental Ministry (Riigiteataja 2010; Pinedo et al. 2014). veal any oribatid mite species that were significantly indicative. While most soil parameters varied only moderately between In three-way ISA E. acromios was significantly indicative of Tal- the three airfields, elevated level of K was noted in Tallinn linn airfield (IV = 0.79, P = 0.01). S. laevigatus, T. velatus ve- and higher organic matter content in Tartu compared to the latus and Eupelops hygrophilus Knülle, 1954 were significantly other airfields. Overall, both K and organic matter content in (P < 0.04) linked to various combinations of two airfields. Even the humus horizon can be considered high (Kask & Niine 1997;

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Petersell et al. 1997). While in Tallinn the elevated K content is of the three airfields did not reveal any significant differences in ,likely linked to historic soil conditions, the high organic matter abundances, indicating that solely based on these parameters content in Tartu airfield soil is more of a mystery, possibly tied the airfields cannot be ranked according to their disturbance to characteristics of the filling soil used in some parts of the level. airfield. Soil pH values, varying from slightly acidic to neutral, The community similarity of different airfields and reflected the historic soil conditions, indicating also that the ad- airfield area purposes together with the number and percent- dition of construction and other aviation related substances has age of shared species suggest fairly homogeneous Collembolan not raised the soil pH as often occurs in soils under anthropo- communities. The typical airfield springtail community consists genic influence (Maechling et al. 1974). of A.granaria, M. affinis, P. armata, P. notabilis, and likely also The SIR results showed similar results as measured species belonging to Folsomia and Isotoma genera. from Estonian road-side grasslands (Vacht 2012), but exceeded Most of the oribatid mites found in this study (e.g. P. the levels measured from e.g. semi-coke heap habitats (Kalda punctum, S. laevigatus) are widespread eurytopic species, usu- et al. 2015). Basal respiration rate was higher than measured ally forming the main body of edaphic communities in most dis- from road-side soils (Vacht 2012). Soil basal respiration has turbed landscapes in Europe (Caruso et al. 2009). The species been shown to indicate toxicity effects (e.g. Pb, Zn, Cu) in soils composition of Oribatid mites resembled those of Lithuanian (Romero-Freire et al. 2016), but not in soils with high organic urban soils (Eitminavičiūtė 2006a,b) containing dominant T. ve- matter and carbonate content such as the soils encountered in latus velatus and S. laevigatus. The latter has also been found this study. Also the indicativeness of substrate induced respira- from the area of Orio al Serio Airport in Northern Italy (Migli- tion has been known to depend greatly on soil pH (Cheng & orini et al. 2003). The low abundance of Oppiella nova Oude- Coleman 1989). The elevated respiration rate in Pärnu may be mans, 1902 could be due to high soil pH as noted by previous related to the increased levels of oil products in the airfield soil studies (Eitminavičiūtė 2006a,b). Also typically moss-dwelling (Hund & Schenk 1994). (Smrž 1994) Scutovertex minutus Koch, 1835 was found from all Diatom and oribatid mite species richness (rarefied sites, however, only abundantly from one of the snow-melting species richness for diatoms) and diversity showed similar sites in Tartu (site TR1). Previously this species has been found trends in comparison of the three airfields. Collembolan diver- from the immediate proximity to the roads (Eitminavičiūtė sity varied greatly, showing no strong signs on indicativeness to 2006a,b) possibly showing its high tolerance towards de-icing airfield nor airfield area purpose. The oribatid mite and diatom agents and preference for hydric conditions. Some research community diversity were similar showing high diversity in the suggest the species has a high tolerance for fluctuating abiotic snow-melting sites of Tallinn airfield and lowest diversity in the soil parameters (e.g. moisture) (Smrž 1994). Also T. incisellus, runways sides of Tartu airfield. In Tallinn this difference could L. pannonica and Peloptulus phaenotus Koch, 1844 have been be caused by the nature of de-icing agents used, and in Tartu previously encountered in heavy metal contaminated soils (Ca- the decreased diversity could potentially be linked to recent ruso et al. 2009; Skubała & Zaleski 2012). Therefore, the com- runway construction work. munity that can be considered typical to airfields contains the Springtail communities were dominated by P. armata following species: T. velatus velatus, L. pannonica, P. peltifer, S. and P. notabilisi, which are known to inhabit highly disturbed laevigatus and T. incisellus. Compared to springtails the oribatid areas (Migliorini et al. 2003; Eitminavičiūtė 2006a,b). Collem- mite communities show more variability, especially in compari- bolans are considered as a relatively resistant group to anthro- son of different airfield area purposes. This is why in the future, pogenic disturbances (Bengtsson & Rundgren 1988; Haimi & especially the effects of airfield de-icing agents on oribatid mite Siira-Pietikäinen1996 ), however, their community composition communities should be studied in more detail. is known to respond to human-induced disturbances, especial- The dominant diatom species were the cosmopolitan ly by decreasing numbers of Isotomidae and Onychiuridae and H. amphioxys, A. granulata and L. mutica. All the studied air- increasing abundance of less demanding and fields contained the same base soil community, that has also Neanuridae (Sousa et al. 2003; Magro et al. 2013). This study, previously been found from many soil habitats both in Estonia however, did not encounter a strong shift in most of these (Vacht et al. 2014) and elsewhere (Soare & Dobrescu 2010; groups in comparison of the three airfields nor in comparison Heger et al. 2012) that represent a tolerant algal community .of the snow-melting sites and runway sides. Only Neanuridae which has little bioindication value that is particular to airfields (e.g. A. granaria and M. pygmaea) were more abundant at Nevertheless, the shared species of the three airfields (e.g. F. the snow-melting sites, which may indicate that these areas nitzschioides, H. amphioxys, N. cari, L. mutica) can be expected present a suitable niche for these species. For Entomobryidae to be found on also other similar airfields. Also it is possible that (e.g. L. lanuginosus and L. cyaneus) the change in abundance some species (e.g. D. finnica) may indicate snow-melting sites was opposite, making it difficult to interpret either the runway mostly due to increased moisture, which are less suitable habi- sides nor snow-melting sites as the more demanding habitat. tats for microarthropod communities. This is one of the reasons Some studies also point out that epedaphic (Pernin et al. 2006) why microarthropods and diatoms should be considered as L. lanuginosus may be indifferent to soil contamination levels complementary bioindicator groups in soil ecological research. (Austruy et al. 2016). The comparison the Collembolan families The ISA revealed that in comparison of the three bioindicator

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112 EUROPEAN JOURNAL OF ECOLOGY groups, diatoms and springtails were the most indicative on pollutants have maintained fully functional soil biological com- specific airfield or land use. Based on this, oribatid mites can be munities. Microarthropod and diatom community parame- considered fairly robust towards airfield related disturbances. ters show more similar variation than springtail communities Soil functionality is closely related to the concept of depending on the airfield and the purpose of the particular soil quality which in turn has the capability to sustain among airfield area. All three bioindicator groups had a substantial other things biotic communities (Gardi et al. 2002). All the air- percentage of species that can be considered typical to areas field soils are functional soils, containing a well adapted and affected by disturbances, in this case airfields. These species comparatively diverse community of diatoms, springtails and include springtails e.g. M. affinisand P. notabilis, oribatid mites oribatid mites. It has been noted on several occasions that us- T. velatus velatus, L. pannonica and T. incisellus, and diatoms F. ing bioindicators should be incorporated into evaluation of soil nitzschioides, H. amphioxys and L. mutica. functionality and quality of anthropogenic soils (Hartley et al. Collembola species L. lanuginosus and T. vulgaris 2008). The community composition found on these airfields were significantly indicative of airfield runway sides but no indicates that all main functional groups of microarthropods strong indicators of specific airfield were discovered. Oribatid are present, though fungal feeders were prominent (e.g. T. ve- mites were not indicative of different airfield purposes, only E. latus velatus, S. laevigatus). This means that the habitat must acromios was noted as indicative of Tallinn airfield. contain also sufficient food sources for all groups present.. Also Three diatom species, F. nitzschioides, Luticola mutica high ratio in Acarina/Collembola may suggest high soil quality and D. finnica, were significantly linked to airfield snow-melting because there are known to be less mites than springtails in sites. Also, many diatom species were specific to Pärnu airfield. degraded soils (Parisi et al. 2005). The results of the O/C ra- In conclusion, it is clear that despite aviation related tio indicates well that in an anthropogenic grassland such as disturbances these three airfields have maintained a functional airfields it is vital to include both oribatid mite and springtail soil biological community that has adapted well to these condi- identification due to the variation of the dominant group, to tions. Also, the shifts in O/C ratio and variation in the commu- receive a full overview of the microarthropod communities nity parameters of the three groups indicates the potential of before combining the data with diatom community data. The these three groups as complementary bioindicators of anthro- ratio reflects also the dominant soil process – humification over pogenic disturbances. Further studies are needed to confirm mineralization in Pärnu and mineralization over humification in the connections between these bioindicators and specific soil Tallinn and Tartu. The elevated O/C ratio ( >1 ) can be also con- parameters. sidered a measure of environmental stability (Bachelier 1986) which means Pärnu airfield can be considered as the most envi- Acknowledgements: We are grateful for financial support from ronmentally stable out of the three airfields studied. Tallinn University’s Centre of Excellence initiative “Studies of Natural and Man-made Environments” and “Natural Sciences and Sustainable Development”, and The Estonian Science 4. CONCLUSION Foundation. We are also grateful for The Hoogstraal Memorial While an overwhelming area of the three airfields show char- Acarology Student Fund, Tartu Cultural Endowment, and Eu- acteristics of Technosols, they also show properties of the ropean Social Fund’s Doctoral Studies and Internationalization natural soils which despite the moderately elevated levels of Program DoRa, carried out by Foundation Archimedes.

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Retrieved from: http://cc.oulu.fi/~jarioksa/opetus/metodi/ Skubała, P. & Zaleski, T. (2012) Heavy metal sensitivity and bioconcen sessio1.pdf. tration in oribatid mites (Acari, Oribatida) Gradient studyin Paole�, M.G. & Bressan, M. (1996) Soil Invertebrates as Bioindicators meadow ecosystems. Science of the Total Environment, 414, of Human Disturbance. Critical Review in Plant Sciences, 15, 21- 364–372. 62. Smrž, J. (1994) Survival of Scutovertex minutus (Koch) (Acari: Oribatida) Parisi, V., Menta, C., Gardi, C., Jacomini, C. & Mozzanica, E. (2005) Mi- under differing humidity conditions. Pedobiologia, 38, 448–454. croarthropod communities as a tool to assess soil quality and Soare, L.C. & Dobrescu, C.M. (2010) Preliminary data on edaphic algae biodiversity: A new approach in Italy. Agriculture, Ecosystems & in the city of Piteşti (Romania). Analele Universitatii din Oradea, Environment, 105, 323–333. Fascicula Biologie, 17, 186–189. 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Vacht, P. (2012) Soolatamise mõju maanteeäärsete muldade faunale ja Van Straalen, N.M. & Van Wensem, J. (1986) Heavy Metal Content of diatomeefloorale. Master’s thesis, Tallinn University, Tallinn, (in Forest Litter as Related to Body-Size and Trophic Estonian). Level. Environmental Pollution, 42, 209–221. Vacht, P., Puusepp, L., Koff, T. & Reitalu, T. (2014). Variability of ripar- Vanker, S., Enneveer, M. & Mäsak, M. (2013) Implementation of mea- ian soil diatom communities and their potential as indicators sures to reduce aviation noise at Tallinn airport. In: M.J. Crocker of anthropogenic disturbances. Estonian Journal of Ecology, 63, (Ed.), Book of Abstracts of 20th International Congress on Sound 168–184. & Vibration (7–11 July 2013, Bangkok, Thailand), International Van de Vijver, B., Gremmen, N. & Smith, V. (2008) Diatom communities Institute of Acoustics and Vibration, Bangkok, 400-404. from the sub-Antarctic Prince Edward Islands: diversity and dis- Venables, W.N. & Ripley, B.D. (2002). Modern Applied Statistics with S. .tribution patterns. Polar Biology, 31, 795–808. New York: Springer van Kerckvoorde, A., Trappeniers, K., Nijs, I. & Beyens, L. (2000) Terres- Weigmann, G. (2006). Hornmilben (Oribatida). Die Tierwelt Deutsch- trial soil diatom assemblages from different vegetation types in lands 76. Teil. Keltern: Goecke & Evers. Zackenberg (Northeast Greenland). Polar Biology, 23, 392–400. Werkenthin, M., Kluge, B. & Wessolek, G. (2014). Metals in European roadside soils and soil solution - A review. Environmental Pollu- tion, 189, 98–110.

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APPENDIX

Table A1 Collembola

Tallinn Tartu Pärnu

Runway Snow-mel- Runway Snow-mel- Runway Snow-mel- sides ting sites sides ting sites sides ting sites Collembola Anurida granaria (Nicolet, 1847) 0.3 ± 0.2 1.0 ± 1.0 0.2 ± 0.2 3.0 ± 3.0 0.2 ± 0.2 0.5 ± 0.5 Bourletiella arvalis (Fitch, 1863) - - 0.2 ± 0.2 0.5 ± 0.5 - - Ceratophysella bengtssoni (Ågren, 1904) 4.7 ± 2.1 - - 3.0 ± 3.0 0.2 ± 0.2 1.5 ± 0.5 Ceratophysella sp. 2.3 ± 2.1 0.5 ± 0.5 3.0 ± 2.2 21.0 ± 20.0 0.3 ± 0.3 2.5 ± 2.5 Dicyrtomina sp. - 0.5 ± 0.5 - 0.5 ± 0.5 1.5 ± 1.5 - Folsomia candida (Willem, 1902) 3.5 ± 1.5 - 2.2 ± 1.8 8.5 ± 8.5 0.2 ± 0.2 0.5 ± 0.5 Folsomia fimetaria (Linnaeus, 1758) - - 0.5 ± 0.2 - 0.2 ± 0.2 0.5 ± 0.5 Folsomia quadrioculata (Tullberg, 1871) 0.2 ± 0.2 0.5 ± 0.5 - - - - Folsomia sexoculata (Tullberg, 1871) 3.8 ± 2.3 7.5 ± 5.5 0.7 ± 0.3 0.5 ± 0.5 3.3 ± 2.6 - Friesea sp. - - - 1.5 ± 1.5 - - Heteromurus nitidus (Templeton, 1835) 0.3 ± 0.2 - - - - - Hypogastrura sp. 2.7 ± 1.8 0.5 ± 0.5 0.5 ± 0.3 11.0 ± 11.0 - - Isotoma minor (Schaeffer, 1896) 5.2 ± 1.7 - 1.0 ± 0.7 31.5 ± 31.5 1.2 ± 0.5 2.0 ± 0.0 Isotoma viridis (Bourlet, 1839) 1.5 ± 0.9 - 1.2 ± 0.3 0.5 ± 0.5 0.7 ± 0.4 1.0 ± 1.0 Isotomurus palustris (Müller, 1776) 1.7 ± 0.6 - - - 0.2 ± 0.2 1.0 ± 0.0 Isotomurus sp. 1.8 ± 1.1 - 0.8 ± 0.5 - 0.3 ± 0.2 - Tomocerus vulgaris (Tullberg, 1871) 0.3 ± 0.2 - 1.5 ± 0.8 - - - Lepidocyrtus lanuginosus (Gmelin, 1788) 3.0 ± 1.5 - 2.5 ± 1.1 - 0.3 ± 0.2 - Metaphorura affinis (Börner, 1902) 2.3 ± 1.0 2.5 ± 2.5 7.5 ± 2.9 2.5 ± 2.5 0.5 ± 0.5 1.5 ± 1.5 Micranurida pygmaea (Börner, 1901) - - - - 0.2 ± 0.2 1.0 ± 1.0 Parisotoma notabilis (Schaeffer, 1896) 17.2 ± 7.5 2,5 ± 0.5 3.0 ± 1.9 1.5 ± 0.5 3.3 ± 1.7 1.0 ± 1.0 Proisotoma minuta (Tullberg, 1871) 0.3 ± 0.2 - 0.7 ± 0.5 35.0 ± 35.0 - 0.5 ± 0.5 Protaphorura armata (Tullberg, 1869) 1.7 ± 1.1 1.5 ± 1.5 4.7 ± 1.9 70.0 ± 70.0 0.2 ± 0.2 0.5 ± 0.5 Pseudisotoma sensibilis (Tullberg, 1876) - - 1.2 ± 1.2 - - 2.5 ± 2.5 - - Pseudosinella sexoculata (Schött, 1902) - - - 6.0 ± 6.0 Sminthurides schoetti (Axelson, 1903) - - - - 0.3 ± 0.3 - Sminthurides sp. - - - - 0.7 ± 0.4 - Sminthurinus aureus (Lubbock, 1862) 0.2 ± 0.2 - - 0.5 ± 0.5 - - Tomocerus vulgaris (Tullberg, 1871) 2.7 ± 0.9 - 3.2 ± 0.7 - 0.3 ± 0.3 - buski (Lubbock, 1870) 1.2 ± 0.8 - 1.2 ± 1.0 - - - Xenylla sp. 2.2 ± 1.3 - 0.5 ± 0.3 1.0 ± 1.0 - -

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Table A2 Oribatid

Tallinn Tartu Pärnu Snow-melting Snow-melting Snow-melting Runway sides Runway sides Runway sides sites sites sites Oribatida Liochthonius brevis (Michael, 1888) - - - 4.0 ± 4.0 0.3 ± 0.3 1.0 ± 1.0 Liochthonius sellnicki (Thor, 1930) - - - - 0.5 ± 0.5 - Steganacarus (Atropacarus) sticulus (Koch, - - - - 0.5 ± 0.5 - 1836) Steganacarus (Tropacarus) carinatus 0.2 ± 0.2 - - - 0.2 ± 0.2 - (Koch, 1841) Rhysotrita ardua (Koch, 1841) - - - - 0.2 ± 0.2 - Platynothrus peltifer (Koch, 1839) 0.2 ± 0.2 - 0.7 ± 0.7 - 4.2 ± 2.1 1.0 ± 1.0 Nothrus silvestris (Nicolet, 1855) 0.2 ± 0.2 0.5 ± 0.5 - - - - Conchogneta traegardhi (Forsslund, 1947) 0.2 ± 0.2 - - - - - Oppiella nova (Oudemans, 1902) - - 0.8 ± 0.8 - 0.3 ± 0.2 - Oppiella translamellata (Willmann, 1923) 0.2 ± 0.2 - 0.8 ± 0.5 - - - Tectocepheus velatus velatus (Michael, 3.2 ± 1.5 2.0 ± 2.0 10.3 ± 5.3 18.0 ± 13.0 0.2 ± 0.2 - 1880) Scutovertex minutus (Koch, 1835) - 1.0 ± 1.0 - 23.5 ± 23.5 0.2 ± 0.2 - Eupelops tardus (Koch, 1835) 0.8 ± 0.5 - - - - - Eupelops acromios (Hermann, 1804) 1.7 ± 0.8 0.5 ± 0.5 - - - - Eupelops hygrophilus (Knülle, 1954) 3.3 ± 2.2 - - - 1.2 ± 0.6 4.0 ± 4.0 Eupelops occultus (Koch, 1835) - - 0.3 ± 0.3 - - - Peloptulus phaenotus (Koch, 1844) 0.3 ± 0.3 - - - 3.7 ± 2.1 1.0 ± 1.0 Liebstadia pannonica (Willmann, 1951) 0.2 ± 0.2 - 0.3 ± 0.3 - 0.3 ± 0.2 - Scheloribates laevigatus (Koch, 1836) 1.7 ± 1.7 - 14.8 ± 9.9 0.5 ± 0.5 4.7 ± 1.8 4.5 ± 3.5 Scheloribates latipes (Koch, 1844) 0.7 ± 0.7 - - - 0.5 ± 0.3 0.5 ± 0.5 Chamobates voigtsi (Oudemans, 1902) 0.2 ± 0.2 - - - - - Ceratozetes mediocris (Berlese, 1908) 0.2 ± 0.2 - - - - - Ceratozetes parvulus (Sellnick, 1922) 0.7 ± 0.5 - - - - - Ceratozetes gracilis (Michael, 1884) 0.2 ± 0.2 - - - - - Trichoribates incisellus (Kramer, 1897) 0.8 ± 0.5 - 0.7 ± 0.5 1.0 ± 1.0 0.2 ± 0.2 - Mycobates sarekensis (Trägårdh, 1910) - - 0.7 ± 0.7 0.5 ± 0.5 0.2 ± 0.2 - Mycobates tridactylus (Willmann, 1929) 0.2 ± 0.2 - - - - - Punctoribates punctum (Berlese, 1908) - - - - 0.2 ± 0.2 1.5 ± 1.5 Haplozetes vindobonesis (Willmann, 1935) 0.7 ± 0.7 - - - - - Peloribates longipilosus (Csiszár, 1962) 1.7 ± 1.7 - - - - - Galumna sp. 0.8 ± 0.3 - - - - - Galumna lanceata (Oudemans, 1900) 0.5 ± 0.5 - - - - - Galumna obvia (Berlese, 1914) 0.3 ± 0.3 - - - - - Pergalumna nervosa (Berlese, 1914) 0.7 ± 0.7 - - - - -

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Table A3 Diatom

Tallinn Tartu Pärnu Runway Snow-mel- Runway Snow-mel- Runway Snow-mel- sides ting sites sides ting sites sides ting sites Diatoms Achnanthes delicatula (Kützing) Grunow, 1880 - - - - - + Achnanthes hungarica (Grunow) Grunow, 1880 - - - - + - Actinella sp. (Lewis) Lewis, 1864 + - - - - - Amphipleura sp. (Kützing) Kützing, 1844 - - - - - + Aulacoseira granulata (Ehrenberg) Ehrenberg, 1843 + + + - + + Aulacoseira karelica (Mölder) Simonsen, 1979 + - - - - - Aulacoseira sp. (Ehrenberg) Thwaites, 1848 + - - + - - Caloneis sp. (Cleve) Cleve, 1894 - - - + - - Cocconeis fluviatilis (Wallace) Wallace, 1960 - - - - + - Cymbella affinis (Kützing) Kützing, 1844 - - + + - + Cymbella radiosa (Héribaud-Joseph) Héribaud-Joseph, + - - - - - 1903 Diploneis finnica(Ehrenberg) Cleve, 1891 - + - - - + Epithemia sp. Kützing, 1844 + - - - - + Epithemia turgida (Ehrenberg) Kützing, 1844 - - - + - + Eunotia paludosa Grunow 1862 - - + - - - Fragilaria nitzschioides (Grunow) Van Heurck 1881 + + + + + + Fragilaria sp. (Müller) Lyngbye, 1819 + - - - + - Fragilariforma virescens (Ralfs) Williams and Round, - - - - + - 1988 Gomphonema parvulus (Kützing) Kützing, 1849 + - - + - + Hantzschia amphioxys (Ehrenberg) Grunow, 1880 + + + + + + Humidophila contenta (Grunow) Lowe et al., 2014 + - + - + + Luticola mutica (Kützing) Mann, 1990 + + + + + + Luticola nivalis (Ehrenberg) Mann, 1990 - + - - - - Luticola venticosa (Kützing) Mann, 1990 + - - + - + Mastogloia sp. (Thwaites) Smith, 1856 - - - - - + Mayamaea sp. (Kützing) Lange-Bertalot, 1997 + + - - + + Navicula cari (Ehrenberg) Ehrenberg, 1836 + + + + + + Navicula cincta (Ehrenberg) Ralfs, 1861 - - - + + - Navicula subhamulata (Grunow) Grunow, 1880 + - - - + + Nitzschia alpina (Hustedt) Hustedt, 1943 + - - - + + Nitzschia amphibia (Grunow) Grunow, 1862 - - - + + + Nitzschia sp. Hassall, 1845 - - - + - - Pinnularia borealis (Ehrenberg) Ehrenberg, 1843 + + - - + + Pinnularia lata (Brébisson) Rabehorst, 1853 + - - - - + Pinnularia obscura (Krasske) Patrick and Reimer, 1966 + - - + + + Pinnularia rabenhorsti(Grunow) Krammer, 2000 - - - - + + Pinnularia sp. Ehrenberg, 1843 + - + - + - Pinnularia subcapitata Gregory, 1856 - - - - + - Pseudostaurosira sp. Williams and Round, 1988 - - - - - +

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continued Table A3 Diatom

Sellaphora alastos (Hohn and Hellerman) Lange-Berta------+ lot and Metzeltin, 1996 Stauroforma exiguformis Flower, Jones and Round, - - - - - + 1996 Stauroneis kriegeri (Patrick) Patrick, 1945 - - - - + - Stauroneis sp. Ehrenberg, 1842 - - - - - + Staurosira construens var. venter (Ehrenberg) Hamil------+ ton, 1992 Staurosirella leptostauron (Ehrenberg) Williams and + + - - - + Round, 1987 Stephanodiscus sp. (Ehrenberg) Eherenberg, 1845 + - - - - + Suriella ovata Kützing, 1844 - - - - + - Tabellaria fenestrata (Lyngbye) Kützing, 1844 + - + - - + Ulnaria ulna (Kützing) Compère, 2001 - - - - - +

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120 IV Vacht, P., Niglas, H., Kuu, A., Koff, T., Kutti, S., & Raamets, J. (2019). Oribatid mite (Acari: Oribatida) communities of urban brownfields in Tallinn, Estonia, and their potential as bioindicators of wasteland successional stage. Acarologia, 59(1), 26−32. Oribatid mite (Acari: Oribatida) communities of urban brownfields in Tallinn, Estonia, and their potential as bioindicators of wasteland successional stage

Piret Vachta , Helin Niglasa , Annely Kuub , Tiiu Koffa,c , Sander Kuttib , Jane Raametsb

a School of Natural Sciences and Health, Tallinn University, Narva Rd 25, 10120, Tallinn, Estonia. b School of Engineering, Tartu College of Tallinn University of Technology, Puiestee 78, 51008, Tartu, Estonia. c Institute of Ecology, Tallinn University, Uus-Sadama 5, 10120, Tallinn, Estonia.

Original article

ABSTRACT Urban brownfields are ecologically valuable ecosystems that have been and are under various anthropogenic influences. Because brownfields are subject to rapid development in urban areas, their biological communities, including soil mesofauna, are overlooked and seldom researched, even though they could provide valuable insight into the ecological functioning of these areas. This exploratory study describes the community characteristics of oribatid mite fauna of 12 brownfields in Tallinn, Estonia, and analyzes the potential of oribatid mites as bioindicators of brownfield successional stage. This study provides the first faunistic list of oribatid mites from the brownfields of Tallinn. No significant changes in species abundance or richness depending on the successional stage were discovered. While some changes in community structure were noted, due to low overall abundance, no clear indicators of brownfield successional stage were identified. For future research on urban brownfield oribatid fauna with the intention of finding potential bioindicators of brownfield properties, increasing the sampling size and inclusion of other groups of soil mesofauna is recommended.

Keywords brownfields; wastelands; Oribatida; bioindication; technosol

Introduction Urban brownfields, also known as wastelands (Atkinson et al. 2014; Mathey et al. 2018) Received 07 July 2018 are areas that have been under development in past, but have not been in formal use since Accepted 06 January 2019 Published 15 January 2019 (CABERNET 2006; Siebielec et al. 2012). The term often implies that the area is affected by contamination (CABERNET 2006; Mathey et al. 2018), however, this is not always the Corresponding author Piret Vacht: [email protected] situation depending on the characteristics of previous use. Detailed information about the history of these areas is, however, not always available. This makes decision making about Academic editor future developments of the urban brownfields challenging. Ekaterina Sidorchuk In Tallinn, Estonia, wastelands make up about 7% of the city area (Karro-Kalberg 2011). These areas are often considered as vacant space in cities and are therefore under rapid changes, DOI including consturction and road development, due to the current urban processes. Wastelands 10.24349/acarologia/20194310 are ecologically important urban green spaces that offer various ecosystem services, including Copyright habitats for miscellaneous fauna and flora (Herbst and Herbst 2006; Strauss and Biedermann Vacht et al. 2006; Pueffel et al. 2018). Distributed under Creative Commons CC-BY 4.0

How to cite this article Vacht et al. (2019), Oribatid mite (Acari: Oribatida) communities of urban brownfields in Tallinn, Estonia, and their potential as bioindicators of wasteland successional stage. Acarologia 59(1): 26-32; DOI 10.24349/acarologia/20194310

123 Previous studies have shown that microarthropods are one of the first groups of soil biota to inhabit novel areas (Hågvar et al. 2009; Hågvar 2012). However the communities tend to be more species-rich in older soils, influenced positively by the presence of vegetation (Madej et al. 2011; Wissuwa et al. 2012). Oribatid mites (Acari: Oribatida) are known as indicators of soil properties (Steiner 1995; Behan-Pelletier 1999; Magro et al. 2013), but they are not often researched in harsh environments such as urban wasteland Technosols (Madej et al. 2011; Rumble et al. 2018). The former use of wastelands may also affect the soil conditions and therefore influence the soil biota, including oribatid mite communities (Zaitsev and Straalen 2001). There are few studies addressing the soil acarofauna of urban areas (Weigmann and Jung 1992; Eitminavičiūtė 2006a, b; Minor and Cianciolo 2007). In Estonia, soil oribatid mites communities have so far not been studied in urban areas, only data from some coastal areas (Eitminavičiūtė 1976), coniferous forest riparian zones (Vacht et al. 2014) and Estonian airfields (Vacht et al. In Press) have been published. Because soil biota, including microarthropods, play an important part in major soil processes and functions, their presence or absence and diversity should be monitored in order to gain knowledge about the stability and functioning of these areas. Therefore, it is important to map what kind of soil biological communities can be found in urban brownfields of Tallinn and whether these can be used as bioindicators helping to evaluate the ecological importance of these areas. The aim of this explorative study was (1) to characterize the oribatid mite communities (community structure, species richness, evenness and diversity) of various urban wastelands in Tallinn that differ in their properties, past usage and successional stage; (2) to find out whether oribatid mites are suitable bioindicators of wasteland successonal stage.

Methods Sampling from 12 urban brownfields in Tallinn, Estonia, was conducted in September 2014 and September 2015. Each year four samples were collected from each study area. Table 1 shows the locations and characteristics of the sampling sites. The study areas included in this study have been out of formal use for up to 50 years, classifying into the three youngest (I, II and III) successional stages distinguished by Mathey and Rink (2010), that do not yet have high and dense vegetation cover. Samples were collected using ø 5 cm soil auger (0–10 cm of topsoil, in total 196 cm3) from sampling sites most characteristic to the wasteland. Because this was an exploratory study, samples for soil chemical analyses were collected from each site and pooled, for general background information without aim to link them directly to the soil biological communities. Soil organic matter and carbonate content was analyzed using prepASH® 340 at Tallinn University Institute of Ecology, soil P, K, Ca and Mg content were analyzed, and

pHKCl measured using standard methods at the Estonian Agricultural Research Center. Microarthropods were extracted using Berlese-Tullgren funnel until samples were fully dry (minimum of 72 hours) and stored in 90% ethanol. For clearing purposes 80% lactic acid was used on oribatid mites. Samples (N = 96) were hand sorted, oribatid mites identified (Weigmann 2006; Niedbała 2008)and counted using cavity slides on light microscope Olympus BX51 at 200–600 × magnification. The nomenclature follows Weigmann (2006). Samples have been deposited at Tallinn University Institute of Ecology. The oribatid mite species data from two years were pooled for numerical analysis. Community parameter calculations were conducted using R programming environment (R Development Core Team 2014) using ’vegan’ package (Oksanen 2018). Due to very low abundance of oribatid mites in the samples multivariate community analysis could not be applied.

Vacht et al. (2019), Acarologia 59(1): 26-32; DOI 10.24349/acarologia/20194310 27

124 Table 1 Study area characteristics, including coordinates, site successional stage (Mathey and Rink 2010) classification (I refering sites less than 3 year old, II to sites 3 to 10 years old, III to sites 10 to 50 years old) and short description of site purpose or previous use and vegetation characteristics.

Characteristics Sampling Coordinates Successional Site description Characteristic vascular plants site stage symbol L N 59°24′55.33″ I Former urban Achillea millefolium , Arctium tomentosum , E 24°52′8.79″ agriculture site, Artemisia vulgaris, Atriplex patula , Cirsium near airfield arvense , Echium vulgare , Scorzonera humilis , Trifolium medium, Tussilago farfara, Urtica dioica, Verbascum nigrum F N 59°25′8.81″ I Illegal garbage Arctium tomentosum, Artemisia vulgaris, Atriplex E 24°45′28.3″ dump, construction patula, Betula pendula, Cirsium vulgare, Epilobium waste angustifolium, Melilotus officinalis , Salix spp, Tanacetum vulgare P N 59°27′3.49″ II Construction site, Acer platanoides, Achillea millefolium , Alchemilla E 24°49′11.27″ construction waste vulgaris, Poaceae, Polygala vulgaris, Rosa spp, Scorzonera humilis, Taraxacum officinale , Tussilago farfara , Sorbus aucuparia V N 59°25′18.32″ II Former timber mill Acer platanoides, Anchusa officinalis, Betula E 24°44′58.84″ factory site pendula, Fraxinus excelsior, Poaceae, Populus tremula , Salix spp M N 59°25′51.42″ III Snow-melting site Acer platanoides, Achillea millefolium , Melilotus E 24°40′58.3″ albus , Phragmites australis , Plantago major, Salix spp, Solidago virgaurea , Taraxacum officinale , Tussilago farfara S N 59°25′11.98″ III Semi-natural area Achillea millefolium , Artemisia vulgaris, Alnus E 24°37′23.31″ with illegal garbage incana, Poaceae, Populus tremula, Ribes nigrum, dump Solidago virgaurea , Taraxacum officinale R N 59°24′50.61″ III Construction waste Aegopodium podagraria, Angelica sylvestris, E 24°39′41.89″ Artemisia vulgaris, Rubus idaeus, Rumex obtusifolius , Urtica dioica A N 59°27′19.43″ III Railway side, no Aegopodium podagraria , Convolvulus arvensis , E 24°41′45.21″ longer in use Equisetum pratense, Phragmites australis , Salix spp, Urtica dioica T N 59°27′0.06″ III Railway side, in Achillea millefolium, Aegopodium podagraria , E 24°43′12.38″ infrequent use Fraxinus excelsior, Medicago falcata, Poaceae , Rumex obtusifolius , Salix spp, Sedum acre K N 59°26′56.23″ III Construction site, Acer platanoides, Arctium tomentosum, Artemisia E 24°44′38.91″ contruction waste absinthium , Bunias orientalis , Salix spp, Saponaria officinalis U N 59°26′26.35″ III Construction site, Alnus incana, Artemisia absinthium , Chelidonium E 24°46′7.02″ former harbour majus, Impatiens parviflora, Populus tremula, Salix warehouse site spp, Taraxacum officinale X N 59°26′48.93″ III Former military Artemisia absinthium , Chrysosplenium E 24°50′15.17″ airfield, alternifolium , Lamium album , Salix spp, construction waste Symphoricarpos albus , Ulmus glabra

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125 Results and discussion The studied wasteland soils can be classified as Technosols due to the high volume of anthro- pogenic artifacts, mostly construction waste, found in them. The amount of anthropogenic deposits in the studied soils together their low organic matter content (Table 2) refers to their low water holding capacity, which can be considered unfavorable habitat characteristics for most soil mesofauna. Urban soils in Tallinn have generally neutral to mildly alkaline pH, which was confirmed by the results. Some study areas stood out by notably higher phosphorus or potassium content. Table 2 shows the results of the soil chemical analysis. From 96 samples 300 oribatid mites were identified. The abundance of oribatid mites differed greatly between the study sites. Even after pooling the data from two years the abundance per brownfield (N=8) varied from just one individual found in K and A sites to 34 – 42 individuals at P, V and X sites. Even though the youngest successional stage had lower oribatid mite abundance than stages II and III, these older stages (II and III) also had sites with only one individual. In total 17 species were encountered. The species list with abundance data (N=8) is shown in Table 3. Highest diversity (Shannon´s H) was encountered in the study area F (H=1.8), followed by the site P (H=1.7) and site V (H=1.4). Community evenness was highest in the study areas R, T and F (1.0, 0.9 and 0.9 respectively). Species richness did not increase with increasing successional stage. Some changes in species composition with increasing successional stage were noted. For example, while Platynothrus peltifer C. L. Koch, 1839 was the species encountered only on a site belonging to the youngest successional stage (only represented by two individuals), most species that were present at stage I were also encountered at the older successional stages. The slowly reproducing P. peltifer is known to be more abundant in near-neutral to alkaline soil conditions (van Straalen and Verhoef 1997) and sensitive to heavy metals (Khalil et al. 2009). The presence of this species in urban wasteland ecosystem together with the species’ known bioindicator properties is a promising combination for further studies. Eupelops torulosus C. L. Koch, 1939 and Trichoribates novus Sellnick, 1928 were only present at stages II and III. Both of these species are known to inhabit soils that already have vegetation – E. torulosus is often found in forest ecosystems (Siira-Pietikäinen et al. 2008; Vacht et al. 2018), T. novus in urban lawns (Weigmann and Kratz, 1987). Several species were indifferent to the successional stage of the wasteland (e.g Galumna lanceata Oudemans, 1900, Rhysotritia ardua C.L. Koch, 1841, Tectocepheus velatus velatus Knülle, 1954). All of these species are frequently encountered in urban and other human influenced ecosystems (Zaitsev and Straalen 2001; Caruso and Migliorini 2006; Eitminavičiūtė 2006b; Minor and Cianciolo 2007; Ivan and Vasiliu 2009;

Table 2 Results of soil chemical analysis to give an overview of wasteland soil conditions.

Soil parameters Sampling Organic Carbonates pH P (mg/kg) K (mg/kg) Ca (mg/kg) Mg (mg/kg) site symbol KCl matter (%) (%) L 7.3 48 126 7837 202 2.7 8.7 F 7.6 41 243 12800 286 4.8 16.4 P 7.6 51 143 6555 175 11.0 38.8 V 7.6 74 220 6131 219 9.7 8.6 M 7.6 31 82 8968 128 4.9 11.4 S 7.0 109 114 2761 99 4.5 0.5 R 7.4 95 140 4040 138 6.2 2.3 A 7.3 17 729 4010 129 8.0 3.1 T 7.3 78 172 5714 211 8.3 3.0 K 7.1 302 267 6312 232 10.9 16.0 U 7.2 206 213 5723 228 5.1 9.8 X 7.3 46 286 11320 197 14.8 23.2

Vacht et al. (2019), Acarologia 59(1): 26-32; DOI 10.24349/acarologia/20194310 29

126 Sheet1 Table 3 Species list and number of specimens collected from the study area (total N=96, each count is based on data from 8 samples of 196 cm3 of soil).

Sampling sites and successional stage I II III Species L F P V M S R A T K U X Rysotritia ardua (Koch, 1841) - 2 - 1 ------1 Trhypochthonius spp (Berlese, 1904) - - - 1 ------Platynothrus peltifer (Koch, 1839) - 2 ------Hermannia convexa (Koch, 1839) - - - 1 ------Tectocepheus velatus velatus (Michael, 1880) - 1 - 2 - - 2 - - - - - Scutovertex minutus (Koch, 1835) - - 1 ------Eupelops torulosus (Koch, 1839) - - 2 1 - 1 1 - - - 1 4 Eupelops hygrophilus (Knülle, 1954) - 3 4 - - - 1 - 2 - - 1 Achipteria coleoptrata (Linnaeus, 1758) - 2 3 21 4 - - 1 3 - - - Achipteria nitens (Nicolet, 1855) - - 1 7 ------Achipteria sellnicki (van der Hammen, 1952) - 1 ------1 1 - Scheloribates laevigatus (Koch, 1836) 1 5 16 ------31 Chamobates voigtsi (Oudemans, 1902) - - 3 2 ------Ceratozetes minutissimus (Willmann, 1951) ------1 Trichoribates novus (Sellnick, 1928) - - - 1 - - 1 - - - - 2 Mycobates tridactylus (Willmann, 1929) - - 1 ------Galumna lanceata (Oudemans, 1900) 1 - 3 - - - - - 1 - - 2

Vacht et al. 2018), for example, T. velatus velatus is often one of the most abundant species in urban ecosystems (Eitminavičiūtė 2006b; Minor and Cianciolo 2007). The number of individuals encountered in this study was very low compared to our previous research from areas under anthropogenic influences (Vacht et al. 2018; Vacht et al. In Press) and not sufficient for determining the full potential of oribatid mites as bioindicators of brownfield properties, including changes in community patterns in relation to successional stage, therefore this study is unable to recommend using oribatid mites alone as indicators of wasteland properties, including successonal stage. Increasing the sampling size and combining the results with other organism groups (e.g. Collembola, Mesostigmata) should be considered.

Conclusion Due to high concentration of anthropogenic imputs and low organic matter content, urban wasteland Technosols encountered in this study, are very poor habitats for oribatid mites, resulting in low abundance and diversity. This study provided novel species data on brownfield oribatid mite communities in Tallinn, Estonia. In comparison of three successional stages no significant changes were detected in abundance or diversity of the communities, but some changes in community structure were noted. Due to the low total abundance of oribatid mites, further studies are required to test, whether oribatid mites can be used as indicators of successional stage in urban brownfields. In future studies in urban brownfields we recommend increasing the sampling size. In order to assess the ecological functioning of the areas, the bioindicative data from their oribatid mite communities should be combined with that from the other groups of soil mesofauna.

Acknowledgements We are grateful for financial support from Tallinn University’s Centre of Excellence initiatives ”Studies of natural and man-made environments” and ”Natural Sciences and Sustainable Development”, and also Tallinn University´s StudentPage Research1 Project Grant.

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