PROCEEDINGS OF THE LATVIAN ACADEMY OF SCIENCES. Section B, Vol. 69 (2015), No. 3 (696), pp. 120–131. DOI: 10.1515/prolas-2015-0017

ECOLOGY AND DIVERSITY OF URBAN PINE FOREST SOIL INVERTEBRATES IN RÎGA, LATVIA Dmitry Telnov and Ineta Salmane#

Institute of Biology, University of Latvia, 3 Miera Str., LV-2169 Salaspils, LATVIA # Corresponding author: [email protected]

Communicated by Viesturs Melecis

A study on ecology and diversity of soil invertebrates of urban pine and mixed pine forests was carried out in seven different sampling plots in Rîga during 2014. Ninety eight soil samples were processed and in total, 40 426 specimens were extracted (of them, 25 237 specimens were iden- tified to species level and 15 189 to order level). Indices (abundance, community similarity etc.) characterising faunal diversity and species communities of Rîga city soil fauna were estimated. The most numerous soil invertebrate groups were Collembola, and , ac- counting for 95% of all collected . There was rather high diversity of soil invertebrates in the disturbed urban forest habitats, but undisturbed soils harbour a greater species richness of fauna than disturbed soils. Key words: soil invertebrate fauna, urban forest, bioindication.

INTRODUCTION Urban forests are green islands in an anthropogenic land- scape and support biological diversity within surrounding Urbanisation is increasing worldwide. Today, 45% of the built-up areas and streets (McPherson et al, 1997). Urban human population lives in cities, and in industrialised coun- forests are typical but specific ecosystems for Latvia, where tries this proportion increases to about 80% (Magura et al., nearly 20% of urban areas are forested. Pine forests are the 2008). Urbanisation is associated with a variety of effects most typical among them. These forests are under long-term on the soil system, including pollution, conversion of indig- anthropogenic pressure and urban soil mesofauna are con- enous habitats to various forms of land use, habitat frag- sidered to be degraded from environmental stress and eco- mentation and loss, and soil community changes (Pickett et logical forest succession is progressing toward deciduous al., 2001; Santorufo et al., 2012). Habitat conditions in cit- forests caused by soil eutrophication. ies much differ from those in natural habitats (Niedbala et Understanding of urban ecosystems requires information al., 1990). Especially the soil is subject to major transfor- about the response of soil biotic communities to environ- mations, which are reflected by the species composition and mental changes within large cities (Smith et al., 2006), and abundance of soil fauna (Niedbala et al., 1990). As it is well particularly the reaction of soil mesofauna to forest type known, soil plays an irreplaceable role in the biosphere: it succession and environmental stress (Barbercheck et al., governs plant productivity, organic matter degradation and 2009). nutrient cycling (Santorufo et al., 2012). Soil living organ- isms, including soil invertebrate fauna affect ecosystem pro- Surprisingly few data are available on invertebrate fauna cesses, such as organic matter decomposition, nutrient min- and ecology of Rîga. Even faunistic data are limited to eralisation and cycling, microbial activity, and soil mixing half-a-dozen papers devoted to leafhoppers (Danka, 1973), (Murphy, 1955; Witt, 1997; Koehler, 1999; Liiri et al., blow flies (Danka, 1979), barklices (Danka and Spuris 2002; Huhta et al., 2005; Huhta, 2007; Santorufo et al., 1977), ground beetles (Stiprais, 1973a; Melecis et al., 2012). They are involved in creation of the soil system, 1999), and (Stiprais, 1973b). An ecological description which acts as an environment for organisms, and within of the territory of Rîga was published in 1973 (Danka and which they are adapted (Colemann and Crossley, 1996). Stiprais 1973) as an introductional paper in a series devoted The soil invertebrates in nature closely interact with abiotic to the entomofauna of Rîga. environmental factors, which affect their activities, and many of them respond sensitively to changes in the environ- Of epigeic and soil , ground beetles (Coleoptera: ment conditions, and thus may be used as indicators of soil Carabidae) and ants (: Formicidae) were cov- quality (Reddy 1986; Coleman, 2008). ered by sporadic studies in Rîga. Nearly one half of Latvian

120 Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. ground beetle fauna, or 128 species, were recorded from MATERIALS AND METHODS Rîga (Stiprais, 1973a), and supplemented by several species Study area. The study was carried out in Rîga, the capital found during later studies (Melecis et al., 1999). Ground 2 beetles are among the most successful epigeic arthropods in of Latvia. The present area of Rîga is 304 km ; of which urban environments, but such high species diversity is a re- ~56% comprises build up areas and roads, 28% green sult of high diversity of available ecological niches. Fifteen spaces and ~16% water bodies (www.riga.lv). The territory species have been recorded from the territory of Rîga of Rîga expanded rapidly in the post-Second World War pe- (Stiprais, 1973b), of them Formica species that were mostly riod, when large surrounding areas were incorporated into found in forested areas. the city. In Latvia, after the Second World War, the urban forests in the inner city were protected by legislation of the Some data are available from Lapiòa (Lapiòa, 1988) on soil Soviet period and most of present forested areas in Rîga are Mesostigmata of Rîga and Jûrmala. She mentioned 50 aged up to 100 years. These forests were mostly planted species from both urban forests and gardens. Some findings post-Second World War, or expanded around older forest of Mesostigmata mites in various habitats in Cçsis, Jûrmala, fragments. Ogre, Rîga, Salaspils, and Sigulda were published by I. Sal- In Latvia, nearly 20% of the urban area is covered by for- mane (Salmane, 2005a & b; Salmane and Meiere, 2005; ests (Donis, 2003). No less than one half of green spaces in Kontschán Salmane, 2006; 2007a & b; et al., 2008; Rîga are forested areas, which amounts to 4244 ha Salmane, 2009; Salmane and Telnov, 2009). Still, all these (www.riga.lv). Rîga forests consist of 15 forest tracts that records are fragmentary and no planned investigation or are connected with rural forests or are smaller, isolated for- monitoring of soil invertebrate fauna in Rîga or any other ests — remnants of ancient forest or planted forests city of Latvia has been made. (Straupe et al., 2012). Scots pine (Pinus sylvestris)isthe prevailing species in Rîga forests, except for stands on allu- Of non-epigeic species, leafhoppers (Homoptera: Auche- vial soils, which are mostly forested by black alder (Alnus norryncha), barklices (Psocoptera) and blow flies (Diptera: glutinosa). Pine forests in Rîga cover in total 46.9 km2 or Calliphoridae) have been mentioned in the Rîga fauna. 88% of total forest area in the city (Straupe et al., 2012). Eighty seven species of Auchenorrhyncha were recorded The pine forests in Rîga and its suburb area have mostly de- from the territory of Rîga, of them 74 species connected veloped on poor sandy soils. In total, Rîga city forests con- with grasslands and only few forest species (Danka, 1973). sist of 15 forest tracts (Jankovska et al., 2014) of various Only six species have been recorded from historical old grade of isolation. town of Rîga, where green spaces are very limited. Among twenty barklice species found in Rîga, 15 species were re- So far 154 vascular plant and 18 moss species have been re- corded from the inner city parks (species possible connected ported from Rîga urban pine forests (Straupe et al., 2012). with old-growth broad-leaved trees) (Danka and Spuris, 1977). Only nine species of blow flies (Diptera: Callipho- Study plots. Seven forest plots were selected for collection ridae) have been recorded from Rîga, of them three species of soil invertebrates. Of them, one plot was located on the from the central part of the city (Danka, 1979). left and six plots on the right side of River Daugava (Fig. 1). The characteristics of plots (numbered 1 to 7) are briefly No revisional account of molluscs exists for Rîga. described in Table 1.

Data on Collembola are absent for Rîga almost completely. Consequently, no studies on ecology or urban pressure on communities for the territory of Rîga city have been previously published.

Our general objective was to determine if selected groups of soil arthropods and molluscs differed consistently according to relative level of pine and pine mixed forest succession and therefore could serve as potential biological indicators of ecosystem condition across forest ecosystems in the ur- ban environment. The aim was to discover differences or similarities in soil invertebrate communities among the ur- ban pine forest patches with variable vegetation and soil conditions. To test this concept, soil and epigeic arthropod and mollusc populations were examined in seven forest patches in relatively disturbed and undisturbed sites multi- ple times within one season. We selected Rîga as the largest and most populated city in NE Europe, as it is known as rich in forest ecosystems of various disturbance (Jankovska et al., 2014). Fig. 1. Sampling plots and their location in Rîga, 2014.

Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. 121 Table 1 BRIEF GEOBOTANICAL DESCRIPTION OF URBAN FOREST STUDY PLOTS IN RÎGA CITY (M. Laiviòð, unpublished data)

Study plots 1234567 Location in Rîga city Vakarbuïïi Bâbelîtis Berìi Dreiliòi Meþciems Gaiïezers Jugla Geographical co-ordinates 498073 / 513652 / 517165 / 514220 / 515085 / 515204 / 515558 / 6319074 6316596 6316626 6311228 6314333 6313904 6313346 Forest type vacciniosa vacciniosa/ vacciniosa/ myrtillosa mel myrtillosa hyloco-miosa hyloco-miosa myrtillosa myrtillosa Closeness of canopy layer 45% 55% 65% 70% 60% 90% 50% Pinus sylvestris 45% 55% 60% 50% 40% 30% 50% Acer platanoides 0 0 0 15% 5% 50% 0.5% Quercus robur 0 0 0 5% 10% 10% 0.5% Betula pendula 0 05%01%00 Closeness of shrub layer 3% 50% 2% 5% 80% 20% 15% Sorbus aucuparia 1% 35% 1% 4% 10% 0,5% 8% Acer platanoides 0 1% 1% 0.5% 12% 12% 4% Amelanchier spicata 0 10% 0.5% 0.5% 80% 4% 0.5% Quercus robur 1% 1% 0.5% 0 0 0 8% Closeness of herbaceous layer 50% 70% 65% 95% 30% 35% 85% Rubus idaeus 0 0 0 40% 8% 0.5% 0 Impatiens parviflora 0 0 0 30% 5% 12% 0 Vaccinium myrtillus 000015%4%40% Vaccinium vitis-idaea 0015%0000 Deschampsia flexuosa 15% 0 35% 0 0 0 18% Festuca ovina 8%010%0000 Fragaria vesca 030%00000 Dactylis glomerata 015%00000 Agrostis tenius 08%00000 Carex arenaria 6%000000 Bryophyte layer 65% 40% 70% 5% 0 0 40% Brachytecium rutabulum 0003%000 Plagiomnium affine 1% 1% 0 2% 0 0 0.5% Hylocomium splendens 30% 35% 25% 0 0 0 30% Pleurozium schreberi 0 5% 40% 0 0 0 10% Rhytidiadelphus triquetrus 30%000000 Grade of hypertrophica -tion * low medium medium high high high low

Null (0) means species was represented by less than a 0.5%; co-ordinates given using the Baltic Coordinate System (BCS). * based on unpublished data of soil analysis performed under same study.

The area of each plot was 706.5 m2. The selected plots rep- and plots 2 and 3 were considered transitional. Chemical resent various stages of pine forest succession. Plots 1, 3, characteristics of soil are given in Table 2. and 7 (Vakarbuïïi, Berìi and Jugla) represented typical open and light pine forests with very scarse understorey and Sampling methodology. Soil samples were taken at least shrub layer, while plots 2, 4, and 6 (Bâbelîtis, Dreiliòi, and 5 m from the plot border in a relatively homogenous forest Gaiïezers) had a rich understorey and shrub layer of decidu- patch. Fourteen samples were taken from each plot, once ous trees and shrubs (in Gaiïezers, Acer platanoides trees every two weeks from April to October of 2014. A total of contributed half of the canopy layer). Consequently, the 98 soil samples were processed during the present study. A herbaceous layer was poor in plots 1 and 6, and very dense soil corer (diameter 15 × 15 cm) was used for soil sampling. in 3 and 5. The bryophyte layer was well developed in pure Each sample was placed in a plastic container and processed Scots pine stands (plots 1, 3, and 7), and underdeveloped in in the laboratory on a modified Tullgren funnels. Inverte- plots 5 and 6. Plots 1 and 7 were considered background brates were extracted for 10 days for each sample. Molluscs plots (pine forest with the lowest level of eutrophication), were collected separately from the same area plot by sieving while plots 4, 5, and 6 were considered the most eutrophic, samples. Collected specimens were stored in 100% ethanol

122 Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. S Table 2 =- Hppå iiln = SOIL CHARACTERISTICS BY PLOTS IN RÎGA FORESTS (G. Tabors, i 1 unpublished data) where p is the proportion (n/N) of individuals of one species counted (n) divided by the total number of individuals Plots 1234576 counted (N), and s is the number of species ph 3 4.6 3.2 5.2 4.7 3.8 5.1 Type of humus moder moder mor moder moder moder mull Evenness. Species evenness refers to how close in numbers O layer 5 cm 6 cm 8 cm 2 cm 2 cm 5 cm 2 cm each species in an environment are. Mathematically it is de- fined as a measure of biodiversity which quantifies how Ah, AhE layer 6 cm 10 cm 10 cm 26 cm 6 cm 9 cm 21 cm equal the community is numerically. J' is constrained be- Cu 1.6 1.2 0.7 3.3 1.65 1.35 1.3 tween 0 and 1. Lower variation in a community between Fe 130 175 105 200 195 85 130 species gives a higher evenness index: H¢ (for , Isopoda, Pseudoscorpionida, Opiliones, J¢ = ¢ Chilopoda and Insecta) or in ethanol and glycerine mix (for H max ). In total, 40 426 specimens were extracted and pro- where Hi is the number derived from the Shannon diversity cessed. i i index and H max is the maximum value of H , equal to: Determination of specimens was based on various scientific S sources. Due to the time limitation, it was not possible to ¢ =-11 = H max å ln lnS . finish identification of all Acari and Collembola to species i=1 SS level. where S is the total number of species. Data analysis. Community similarity index. Sorenson’s co- efficient of community similarity gives an estimate of RESULTS shared taxa by a value between 0 and 1 (complete commu- nity overlap is equal to 1; complete community dissimilarity In total 40 426 specimens were extracted and processed is 0). from 98 samples of seven sampling plots in Rîga city for- ests. Of them, 25 237 specimens identified to species level, Sorenson’s Coefficient (CC) and another 15 189 specimens to order level. 2C CC = Number of specimens per plot. The highest number of SS12+ specimens was recorded for plot 1 (Vakarbuïïi, less where C is the number of species the two communities have eutrophic forest patch) and plot 4 (Dreiliòi, eutrophic forest in common, S1 is the total number of species found in the patch), and the lowest for plot 5 (Meþciems, eutrophic for- community (plot) 1, and S2 is the total number of species est patch) (Fig. 2). found in community (plot) 2. Number of specimens per month of study. The number of Diversity and similarity. Diversity of mesofauna was esti- invertebrate specimens increased almost continuously from mated using the Shannon diversity index and the Simpson April and reached a maximum in October (Fig. 3). Mini- diversity index, which area affected by both richness and mum number of extracted specimens was in April for all evenness. seven plots, but maximum occurred in different months for the plots (Table 3). The Simpson Index is a dominance index that gives more weight to common or dominant species. In this case, a few rare species with only a few representatives will not affect the diversity.

Simpson Index (D)

= 1 D S 2 å pi i=1 where p is the proportion (n/N) of individuals of one species counted (n) divided by the total number of individuals counted (N), and s is the total number of species.

Shannon Index (H) The Shannon Index is an information statistic index, which Fig. 2. Number of invertebrate specimens per plot in Rîga, 2014. 1–7 – plot assumes all species are represented in a sample and that numbers:1–Vakarbuïïi,2–Bâbelîtis,3–Berìi, 4 – Dreiliòi, 5 – they are sampled randomly. Meþciems, 6 – Gaiïezers, 7 – Jugla.

Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. 123 Fig. 3. Number of invertebrate specimens per month from April to Octo- Fig. 4. Species diversity of sampled invertebrates by taxonomical group, ber, Rîga, 2014. Rîga, 2014.

Table 3 NUMBER OF EXTRACTED INVERTEBRATE SPECIMENS BY PLOT Biological diversity of extracted specimens. Extracted AND MONTH, RÎGA, 2014 specimens were from the phylum Mollusca (304 speci- mens), Crustacea (66 specimens), and Arthropoda (40 056 Plot April May June July August September October specimens). Three arthropod orders (Collembola, Mesostig- No. mata, and Oribatida) were represented by 38 376 specimens 1 594 732 1038 1037 1412 1189 1269 or 95% of all extracted specimens. The remaining higher 2 339 768 613 659 643 790 922 ranking taxonomical groups were represented by 2 050 3 376 1402 617 904 936 1004 1151 specimens in total (Fig. 4, Tables 4 and 5). 4 744 879 954 1090 982 1107 1094 5 308 503 519 633 708 677 758 In total, 148 species were identified, and 15 189 specimens 6 719 769 748 891 1076 1053 978 were identified to the order or family rank. Order 7 321 502 560 880 827 783 968 Coleoptera was the richest group with 38 species, followed

Table 4 MAIN TAXONOMICAL INVERTEBRATE GROUPS BY PLOT IN URBAN FORESTS OF RÎGA, 2014 Plot No. Mollusca Mesostigmata Oribatida Collembola Chilopoda Coleoptera Hymenoptera Totally (Formicidae) 1 0 2252 1092 3588 16 54 263 7265 2 9 1924 684 1892 11 65 120 4705 3 0 3092 873 2170 14 37 202 6388 4 83 1109 1484 3882 9 200 25 6792 5 48 1245 923 1759 17 73 29 4094 6 158 851 1767 3082 4 227 84 6173 7 6 2005 795 1907 14 53 52 4832 Totally 304 12478 7618 18280 85 709 775

Only significantly represented taxonomical groups are represented in this table.

Table 5 MAIN TAXONOMICAL INVERTEBRATE GROUPS BY MONTH IN RÎGA FORESTS, 2014 Taxonomical group April May June July August September October Mesostigmata 982 2085 1473 1684 1796 2028 2430 Oribatida 904 966 930 1047 1078 1164 1529 Collembola 1443 2253 2204 2977 3277 3093 3033 Coleoptera 31 110 159 127 121 109 52 Hymenoptera (Formicidae) 0 62 197 175 198 104 39 Chilopoda 7 13 16 17 14 14 4 Isopoda 13 11 10 11 11 8 2 Mollusca 21 43 36 45 58 53 48

Maximum quantity per study period is marked with bold.

124 Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. Fig. 5. Number of sampled invertebrate specimens by taxonomical group, Rîga, 2014. Fig. 6. Species abundance (total number of individuals per species), Rîga, 2014. Table 6 NUMBER OF IDENTIFIED INVERTEBRATE SPECIES PER PLOT AND MONTH, RÎGA FORESTS, 2014

Plot April May June July August September October No. 1 59 64 59 65 68 61 62 2 41 64 63 64 61 65 61 3 50 53 61 67 67 58 54 4 52 62 70 63 71 77 62 55557 54 61 59 62 59 6 48 65 67 66 75 72 63 7 45 53 71 65 56 65 67

Fig. 7. Soil invertebrate species diversity by plot, Rîga, 2014. 1–7 – plot by Oribatida (29 species), Mesostigmata (25 species), numbers: 1 – Vakarbuïïi, 2 – Bâbelîtis, 3 – Berìi, 4 – Dreiliòi, 5 – Collembola (16 species), and Mollusca (15 species). All of Meþciems, 6 – Gaiïezers, 7 – Jugla. the other orders were represented by less than five species (Fig. 5). The lowest diversity was recorded in April in 6 Faunal composition or species overlap. Faunal composi- plots, while the highest diversity occurred in different tion of the studied pine forests in Rîga appeared quite simi- months in the plots (Table 6). lar. Among 137 identified species, 50 species (36%) were found in all 7 plots and only six species were “restricted” to Abundance. To determine possible indicators of distur- a single plot (Fig. 8). bance, 148 species were used. Abundance was pooled across sample dates (“cumulative abundance”) and fre- Community similarity index. Sorenson’s coefficient of quency of detection, expressed as the percentage of samples similarity among the studied invertebrate communities was of the total number of samples was calculated. 0.6–0.91, which indicates strong to very strong similarity for the soil fauna (Table 7). The total combined similarity Species abundance. The most numerous species (for all index for all plots was 0.5. plots in general) were Mesostigmata mites Parazercon Diversity and similarity. Epigeic invertebrate fauna diver- sarekensis (1456 specimens), Ololaelaps placentula (1275 sity of the urban pine forests in Rîga was similar (Table 8) specimens), ostrinus (1021 specimens), Leioseius and no significant differences were found. Highest diversity bicolor (994 specimens) and collembola Arrhopalites occurred in plot 1 (Vakarbuïïi, low eutrophication), and the principalis (1193 specimens) (Fig. 6). Twenty species were lowest in plot 3 (Berìi, medium eutrophication), according represented by five or less specimens. to the data identified to species level (Table 8). Diversity of identified species per plot. The highest num- ber of species was recorded from plot 5 (106 species), 7 DISCUSSION (106 species), and 6 (103 species), and the lowest number (91 species) from plot 3 (Fig. 7). These values are only rela- Our results show rather high diversity of soil invertebrates tive, because of the high number of unidentified specimens. and similar faunal composition in the disturbed urban forest

Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. 125 The highest total number of invertebrates was collected in pine forest with sparse understorey, dense moss layer and a moder type of humus (plot 1), and the least in pine-mixed forest with a dense herbaceous plant layer and moder type of humus (plot 5) (Fig. 2, Table 2).

The highest total number of determined species was re- corded in pine-mixed forest with dense herbaceous plant layer and moder type of humus (plot 5) and in pine forest without understorey, with a dense moss layer and moder type of humus (plot 7). The lowest total number of species was recorded in pine forest without understorey and dense Fig. 8. Proportion (number of species) of species overlaps by plots, Rîga, herbaceous plant and a moss layer and mor type of humus 2014. 1–7 – plot numbers: 1 – species only recorded in single plot (no plot (plot 3) (Fig. 7). Plots with a higher range of variable overlapping), 2–7 – species recorded from 2 to 7 plots (overlapping from 2 to 7 plots). microhabitats are inhabited by a higher number of inverte- brates. Some authors have proposed the number of speci- mens better reflect soil properties than the species diversity Table 7 (Santorufo et al., 2012). SORENSON’S COEFFICIENT OF COMMUNITY SIMILARITY AMONG 7 INVERTEBRATE COMMUNITIES IN RÎGA FORESTS A clear difference for the forest types was observed among STUDIED IN APRIL–OCTOBER, 2014 the investigated groups. Saprophagous invertebrates, like Collembola, Oribatida mites and Mollusca had the highest Plot No 4 123456 6 numbers in moder or mull humus type with relatively high 1 ------pH levels and a thick well decomposed humus layer (plots 4, 6) (Tables 2, 4). The main litter decomposers are soil in- 2 0.85 ----- vertebrates, e.g. saprophagous mites and Collembola, and 3 0.90 0.85 ---- the most important besides lumbricids are moss mites 4 0.66 0.73 0.63 - - - (Petersen and Luxton, 1982; Evans, 1992). Oribatida mites 5 0.85 0.85 0.80 0.81 - - were the most numerous in pine-mixed forest plots in our 6 0.64 0.68 0.63 0.60 0.77 - investigation. Litter-feeding mites (e.g. Oribatida) prefer 7 0.91 0.86 0.88 0.63 0.75 0.64 leaf as opposed to conifer litter (Murphy, 1955). In the cur- Total communities similarity for plots 1–7: 0.50 rent investigation, 29 moss mite species were recorded, and dominant species of them were Carabodes subarcticus habitats. Among 137 identified species, 50 species (36%) Trägårdh, 1902, Carabodes labyrinthicus (Michael, 1879), were found in all seven plots and only six species were “re- and Oribatula tibialis (Nicolet, 1855). In total, 80 moss stricted” to a single plot with no overlap (Fig. 8). mite species were recorded during a long-term investigation in natural pine forests of Latvia (Kagainis et al., 2014). Of The most numerous soil invertebrate groups in our investi- them, Oppiella (Oppiella) nova (Oudemans, 1902), gation were Collembola, Oribatida, and Mesostigmata, Tectocepheus velatus velatus (Michael, 1880), and which accounted for 95% of all collected animals. Other au- Suctobelbella falcata (Forsslund, 1941) were dominant spe- thors have found extremely high number of springtails and cies. In our investigation, these species were found in low mites in urban soils (Kuznetzova, 1994; Krestyaninova and numbers, which may be due to the high number of undeter- Kuznetzova, 1996; McIntyre et al., 2001). This high abun- mined species or to the differences in natural and urban for- dance is due to their ability to survive and reproduce faster est habitats. In total, 141 Oribatida species were recorded in in the urban environment, i.e. tolerance of urbanisation in pine forests of Lithuania (Eitminavichute, 2003) and two of their high variety. Kuznetsova (1995) confirms that despite those T. velatus velatus and O. (Oppiella) nova were domi- of negative antropogenic factors like pollution, human tram- nant (Eitminavichute et al., 2008). Manu and Honciuc pling, organic matter and moisture deficit etc., springtails (2010) recorded 54 Oribatida species in parks of Bucharest can form diverse and abundant communities in urban soils. city in Romania.

Table 8 DIVERSITY AND SIMILARITY OF SOIL INVERTEBRATE FAUNA BY PLOT IN RÎGA FORESTS, STUDIED IN APRIL–OCTOBER, 2014

Plot No 4 1234567 Hypertophication 4 low medium medium high high high low Index 6 Simpson 2.451 2.697 2.744 2.665 2.830 2.623 2.730 Shannon 109.415 81.840 29.558 97.797 85.150 97.912 69.067 Evenness 0.042 0.056 0.153 0.047 0.055 0.047 0.068

126 Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. Beside the Oribatida mites, Collembola also are significant on their distribution in urban forests. Still, they were found decomposers. In the current investigation the highest num- in higher numbers in plots 4 and 6, pine-mixed forest habi- ber of Collembola was recorded in plot 4. Jucevica and tats with thick decomposed humus layer and higher pH val- Melecis (2005) recorded 66 Collembola species in natural ues. It is well known that Isopoda generally prefer calcare- pine forests of Latvia. In our investigation, of the deter- ous habitats with high pH, which is not the case in pine mined springtail species, the dominant species were forests (Huhta et al., 2005). Arrhopalites principalis Stach, 1945 and Isotomiella minor (Schäffer, 1896). Still, we have to take in consideration the The total number of extracted invertebrate specimens in- very high number of unidentified Collembola (about 11 000 creased almost continuously from April and reached a max- specimens). Jucevica and Melecis (2006) mentioned imum in October (Fig. 3). Of the most numerous groups, Parisotoma notabilis (Schäffer, 1896) and Isotomiella mi- Collembola reached a maximum in august, Mesostigmata nor to be dominant species in natural pine forests of Latvia. mites had two maximums (May and October), and Oribatida mites in October. This is in contrast with the results of study To a lesser extent, also terrestrial molluscs are important on soil arthropods in natural pine forests of Latvia (Sal- decomposers of plant residues. The most numerous mane, 2000), where the highest number of soil Oribatida molluscs were found in plots 6 and 4, which were relatively mites was found in April. Collembola and Mesostigmata wet and rich in leaf litter pine-mixed forest habitats. This mites had two maximums — in April and September and in type of habitat is favourable for them (Kappes, 2005; April and August, respectively. Increase in the number of Niemelä, 1997). Molluscs were consequently absent in plots invertebrates during autumn 2014 was possibly due to the 1 and 3, which were dry pure pine forests. heat island effect characteristic for large cities (Pickett et al., 2011). Predators, like Mesostigmata mites, Chilopoda and Formicidae, had the highest numbers in plots 1, 3 or 5 with In general, undisturbed soils harbour a greater species rich- mor or moder type of humus and a thin decomposed humus ness of mite fauna than disturbed soils. The number of layer (Tables 2, 4). Mesostigmata mites are predators on mites in litter and soil in mature forests of U.S. Pacific various small invertetebrates like Collembola, Enchy- Northwest was greater than in clearcut plots (Barbercheck traeidae, various mites and other invertebrates (Koehler, et al., 2009). 1999). We found a surprisingly high total number of Meso- stigmata mites, even higher than for moss mites (Table 4). Several authors have investigated impact of habitat frag- During previous investigations in natural pine forests of mentation on soil fauna and found that this is an important Latvia, Oribatida mites had a significantly higher number of factor for species diversity (Bolger et al., 2000). However, specimens than for Mesostigmata (Salmane 2000). Lapiòa we lack sufficient data from one year of investigations to (Lapina, 1988) recorded 22 species in Rîga city forests, of make global conclusions on soil invertebrate fauna in Rîga which 17 were found in our investigation. In Rîga, 25 for- forest habitats. ests Mesostigmata species were recorded, compared to 41 in natural pine forests (Salmane, unpublished data). In natural One season of study is definitively not sufficient for making pine forests, dominants were nemorensis (C. L. global conclusions on faunal differences in soil invertebrate Koch, 1839), Pergamasus vagabundus Karg, 1968, communities in urban forests affected by environmental dis- Parazercon sarekensis Willmann 1939 and Prozercon kochi turbance. Sellnick 1943, while in Rîga city dominants were Olo- laelaps placentula (Berlese, 1887), Parazercon sarekensis, (C. L. Koch, 1836) and Leioseius bicolor ACKNOWLEDGEMENTS (Berlese, 1918). In pine forests of Lithuania, 77 Meso- We are indebted to our colleagues Mâris Laiviòð and Gunta stigmata species were recorded (Eitminavichute, 2003). Èekstere (both from Institute of Biology, University of Lat- Heldt (1995) recorded 37 Mesostigmata species in broad- via, Salaspils, Latvia) for providing botanical and chemical leaved forest habitats in Bremen, northern Germany, of data on the sampling plots, and to Guntis Tabors (Univer- which the dominant were Hypoaspis claviger (Berlese, sity of Latvia, Department of Biology, Rîga) for soil data. 1883), Rhodacarus elbius Karg, 1971, and Zercon peltatus C. L. Koch, 1836. Manu and Honciuc (2010) recorded 18 Helpful colleagues from various scientific institutions Gamasina species in parks of Bucharest in Romania. across Europe are thanked for assistance with identification of specimens. Most of the identified Coleoptera are predators on larger in- vertebrates. Among Coleoptera, predators like Carabidae The study was financially supported by the European Social and Staphylinidae were the most abundant groups. Occa- Fund, the project No. 2013/0060/1DP/1.1.1.2.0/13/APIA/ sional findings of phytophagous (like Curculionidae) spe- VIAA/041. cies signalised increasing presence of deciduous trees and shrubs (pine forest succession due to eutrophication). REFERENCES In our investigation, Isopoda were recorded in small num- Anonymous. Riga City Council Portal. http://www.riga.lv (accessed 12 Jan- bers, and therefore it is impossible to make any conclusions uary 2015).

Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. 127 Barbercheck, M. E., Neher, D. A., Anas, O., El-Allaf, S. M., Weicht, T. R. Kuznetzova, N. A. (1994). Collembolan guild structure as an indicator of tree (2009). Response of soil invertebrates to disturbance across three resource plantation conditions in urban areas. Memor. Zool., 49, 197–205. regions in North Carolina. Environ. Monitoring Assess., 152, 283–298. Kuznetzova, N. A. (1995). Structure of collembolan microcommunities of Bolger, D. T., Suarez, A. V., Crooks, K. R., Morrison, S. A., Case, T. J. small isolated areas of urban plantations. Bulletin Entomol. Pologne, (2000). Arthropods in urban habitat fragments in southern California: 64,149–158. Area, age and edge effects. Ecol. Appl., 10, 1230–1248. Lapina, I. M. (1988). Gamasina mites of Latvia [Ëàïèíÿ, È. Ì. Ãàìàçîâûå Coleman, D. C. (2008). From peds to paradoxes: Linkages between soil biota êëåùè Ëàòâèè]. Zinatne, Riga. 200 pp [in Russian]. and their influences on ecological processes. Soil Biol. Biochem., 40, Liiri, M., Setälä, H., Haimi, J., Pennanen, T., Fritze, H. (2002). Relationship 271–289. between soil microarthropod species diversity and plant growth does not Danka, L. (1973). Materiâli par Rîgas kukaiòu faunu, II. Cikâdes – change when the system is disturbed. Oikos, 96, 137–149. Auchenorrhyncha [Materials on the Entomofauna of Rîga, II. Auchen- Magura, T., Tóthmérész, B., Hornung, E., Horváth, R. (2008). Urbanisation orrhyncha]. Latvijas Entomologs, 15, 8–17 (in Latvian, summary in Ger- and ground-dwelling invertebrates. In: Wagner, N. (Ed.). Urbanization: man). 21st Century Issues and Challenges. Nova Science Publishers Inc., New Danka, L. (1979). Beiträge zur Insektenfauna von Riga. VII. Schmeiss- York, pp. 213–255. fliegen - Calliphoridae. Latvijas Entomologs, 22, 28–34 (in Latvian, sum- Manu, M., Honciuc, V. (2010). Rank correlations at the level of soil mites mary in German). (Acari: Gamasida, Oribatida) from central parks of Bucharest city, Roma- Danka, L., Spuris, Z. (1977). Beiträge zur Insektenfauna von Riga, VI. nia. Acta Entomol. Serb., 15 (1), 129–140. Staubläuse – Psocoptera. Latvijas Entomologs, 20, 5–13 (in Latvian). McDonnell, M. J., Pickett, S. T. A., Groffman, P., Bohlen, P., Pouyat, R. V., Danka, L., Stiprais, M. (1973). Materiâli par Rîgas kukaiòu faunu, I. Rîgas Zipperer, W. C., Parmelee, R. W., Carreiro, M. M., Medley, K. (1997). teritorijas ekoloìisks raksturojums [Materials on entomofauna of Rîga, I. Ecosystem processes along an urban-to-rural gradient. Urban Ecosyst., 1, The ecological description of the territory of Rîga]. Latvijas Entomologs, 21–36. 15, 3–7 (in Latvian). McPherson, E. G., Nowak, D., Heisler, G., Grimmond, S., Souch, C., Grant, Donis, J. (2003). Designating a greenbelt around the city of Riga, Latvia. Ur- R., Rowntree, R. (1997). Quantifying urban forest structure, function, and ban Forestry & Urban Greening, 2, 31–39. value: The Chicago Urban Forest Climate Project. Urban Ecosyst., 1, Eitminavichute, I. (2003). Soil Mites of Lithuania (Acari) Catalogue. 49–61. Vilnius, Institute of Ecology of Vilnius University. 168 pp. McIntyre, N. E., Rango, J., Fagan, W. F., Faethm, S. H. (2001). Ground ar- Eitminavichute, I., Matusevichute, A., Augustaitis, A. (2008). Dynamic and thropod community structure in a heterogeneous urban environment. seasonal fluctuations of microarthropods complex in coniferous forest soil. Landscape Urban Planning, 52, 257–274. Ekologija, 54, 201–215. Melecis, V., Saviès, F., Puíîte, I. (1999). Rîgas centra skrejvaboles Evans, G. O. (1992). Principles of Acarology. CAB International (Coleoptera, Carabidae) [The Carabidae of the Rîga City centre]. In: Wallingford. 576 pp. Daugavpils Pedagoìiskâs universitâtes 8. Zinâtniskâs konferences rakstu Heldt, S. (1995). Zur Kenntnis der Raubmilbenfauna (Acari: Gamasina) krâjums A11 (Dabaszinâtnes, dabaszinâtòu didaktika, matemâtika, Bremens: II. Die Besiedlung ausgewählter Grünland- und Waldstandorte datorzinâtne). Daugavpils, pp. 26–28 [in Latvian]. im Bürgerpark. Abhandlungen Naturwissenschaftlicher Verein Bremen, 43 Murphy, P. W. (1955). Ecology of the fauna of forest soils. In: Kevan, D. K., (1), 91–115. McE. (Ed.). Soil Zoology. Proceedings of the University of Nottingham, Huhta, V. (2007). The role of soil fauna in ecosystems: A historical review. Second Easter School in Agricultural Science. London, pp 99–124. Pedobiologia, 50, 489–495. Niedbala, W., Bloszyk, J., Kaliszewski, M., Kazmierski, A., Olszanowski, Z. Huhta, V., Räty, M., Ahlroth, P., Hänninen, S.-M., Mattila, J., Penttinen, R., (1990). Structure of soil mite (Acari) communities in urban green of War- Rintala, T. (2005). Soil fauna of deciduous forests as compared with spruce saw. Fragmenta Faunistica, 33 (3), 21–44. forests in central Finland. Memoranda Societatis pro Fauna et Flora Niemelä, J. (1997). Invertebrates and boreal forest management. Conserv. Fennica, 81, 52–70. Biol., 11, 601–610. Jankovska, I., Straupe, I., Brumelis, G., Pavlyuk, D. (2014). Urban forests of Petersen, N. H., Luxton, M. (1982). A comparative analysis of soil fauna Riga, Latvia – pressures, naturalness, attitudes and management. Baltic populations and their role in decomposition processes. Quantitative ecol- Forestry, 20 (2), 342–351. ogy of microfungi and animals in soil and litter. Oikos, 39 (3), 287–376. Jucevica, E., Melecis, V. (2005). Long-term effects of climate warming on Pickett, S. T. A., Cadenasso, M. L., Grove, J. M., Nilon, C. H., Pouyat, R. V., forest soil Collembola. Acta Zoologica Lituanica, 15 (2), 124–126. Zipperer, W. C., Costanza, R. (2001). Urban ecological systems: Linking Jucevica, E., Melecis, V. (2006). Global warming affect Collembola commu- terrestrial ecological, physical, and socioeconomic components of metro- nity: A long-term study. Pedobiologia, 50, 177–184. politan areas. Annu. Rev. Ecol. System., 32, 127–57. Kagainis, U., Spuòìis, V., Melecis, V. (2014). The armoured mite fauna Pickett, S. T. A., Cadenasso, M. L., Grove, J. M., Boone, C. G. D., Groffman, (Acari: Oribatida) from a long-term study in the Scots pine forest of the P. M., Irwin, E., Kaushal, S. S., Marshall, V., McGrath, B. P., Nilon, C. H., Northern Vidzeme Biosphere Reserve, Latvia. Fragmenta Faunistica, 57 Pouyat, R. V., Szlavecz, K., Troy, A., Warrenm, P. (2011). Urban ecologi- (2), 141–149. cal systems: Scientific foundations and a decade of progress. J. Environ. Manag., 92, 331–362. Kappes, H. (2005). Influence of coarse woody debris on the gastropod com- munity of a managed calcareous beech forest in Western Europe. J. Moll. Reddy, M. V. (1986). Soil-inhabiting arthropods as indicators of environ- Stud., 71 (2), 85–91. mental quality. Acta Biol. Hung., 37 (1), 79–84. Koehler, H. H. (1999). Predatory mites (Gamasina, Mesostigmata). Agricult. Salmane, I. (2000). Investigation on the seasonal dynamics of soil Gamasina Ecosyst. Environ., 74, 395–410. mites (Acari: Mesostigmata) in Pinaceum myrtillosum, Latvia. Ekológia (Bratislava), 19, supplementum 3, 245–252. Kontschán, J., Salmane I. (2008). New records of the Uropodina mites of Latvia and description of two new species (Acari: Mesostigmata). , Salmane, I. (2005a). List of Mesostigmata mites (Acari, Parasitiformes) as- 19 (2), 335–341. sociated with Aphyllophorales fungi (Basidiomycetes) in Latvia. Latvijas Krestyaninova, A. I., Kuznetzova, N. A. (1996). Dynamics of collembolan Entomologs, 42, 57–71. (, Collembola) association in the soil of an urban boulevard. Salmane, I. (2005b). Addition to the Latvian Mesostigmata (Acari, Parasiti- Entomol. Rev., 76 (9), 1220–1230. formes) check-list. Latvijas Entomologs, 42, 58–62.

128 Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. Salmane, I. (2006). New Mesostigmata (Acari, Parasitiformes) species in Santorufo, L., van Gestel, C. A. M., Rocco, A., Maisto, G. (2012). Soil inver- fauna of Latvia. Latvijas Entomologs, 43, 52–56. tebrates as bioindicators of urban soil quality. Environ. Poll., 161, 57–63.

Salmane, I. (2007a). New and rare Mesostigmata mites (Acari, Parasiti- Smith, J., Chapman, A., Eggleton, P. (2006). Baseline biodiversity surveys formes) in Latvia. Latvijas Entomologs, 44, 125–126. of the soil macrofauna of London’s green spaces. Urban Ecosyst., 9, 337–349. Salmane, I. (2007b). Mesostigmata Mite (Acari, Parasitiformes) fauna of Stiprais, M. (1973a). Materiâli par Rîgas kukaiòu faunu, III. Skrejvaboles – wood-related microhabitats in Latvia. Latvijas Entomologs, 44, 75–92. Carabidae [Materials on the entomofauna of Rîga, III. Ground Beetles – Salmane, I. (2009). Some new and rare Mesostigmata (Acari, Parasitiformes) Carabidae]. Latvijas Entomologs, 15, 18–28 [in Latvian]. in the fauna of Latvia. Latvijas Entomologs, 47, 71–75. Stiprais, M. (1973b). Materiâli par Rîgas kukaiòu faunu, IV. Skudras – Formicidae [Materials on the entomofauna of Rîga, IV. Ants – Salmane, I., Meiere, D. (2005). Mesostigmata mites (Acari, Parasitiformes) Formicidae]. Latvijas Entomologs, 15, 30–32 [in Latvian]. associated with Aphyllophorales (Fungi, Basidiomycetes) in Latvia. Phytophaga, 14, 243–246. Straupe, I., Jankovska, I., Ozoliòa, I., Donis, J. (2012). The evaluation of pine forest vegetation in Riga city, Latvia. In: Recent Researches in Environ- Salmane, I., Teïnov, D. (2009). Introduction to the Mesostigmata mite mental Science and Landscaping. 5th WSEAS International Conference on (Acari, Parasitiformes) fauna associated with beetles (Insecta, Coleoptera) Landscape Architecture (LA ’12) University of Algarve, Faro, Portugal, in Latvia. Latvijas Entomologs, 47, 58–70. May 2–4. Portugal, pp. 20–25.

Received 16 July 2015

AUGSNES BEZMUGURKAULNIEKU BIOLOÌISKÂ DAUDZVEIDÎBA URBÂNAJOS PRIEÞU MEÞOS RÎGÂ, LATVIJÂ Rîgas pilsçtâ septiòos prieþu vai prieþu–jaukto meþu parauglaukumos veikts augsnes bezmugurkaulnieku bioloìiskâs daudzveidîbas pçtîjums. Kopumâ ievâkti 98 augsnes paraugi, kuros konstatçti 40 426 îpatòi, no kuriem 25 237 noteikti lîdz sugai, bet pârçjie — lîdz kârtas vai dzimtas lîmenim. Pçtîjumâ apskatîta daþâdu augsnes bezmugurkaulnieku sabiedrîbas raksturojoðo lielumu, kâ piemçram, îpatòu skaita, sugu skaita, sabiedrîbu lîdzîbu u.c. râdîtâju saistîba ar Rîgas pilsçtas urbâno meþu biotopiem.

Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. 129 Appendix 1 LIST OF TAXA IDENTIFIED TO SPECIES LEVEL IN URBAN FORESTS OF RÎGA CITY

Mollusca Parasitidae 27. Holoparasitus excipuliger (Berlese, 1905) Agriolimacidae 28. Parasitus kraepelini Berlese, 1903 1. Krynickillus melanocephalus Kaleniczenko, 1851 29. Pergamasus crassipes (Linnaeus, 1758) Arionidae 30. Pergamasus holzmannae Micherdzinsky, 1969 2. Arion subfuscus (O. F. Müller, 1774) 31. Pergamasus lapponicus Trägardh, 1910 Bradybaenidae 32. Pergamasus vagabundus Karg, 1968 3. Bradybaena fruticum (O. F. Müller, 1774) 33. Pergamasus wasmanni (Oudemans, 1902) Discidae Rhodacaridae 4. Discus ruderatus (Férussac, 1821) 34. Asca aphidioides (Linnaeus, 1758) Euconulidae 35. Asca bicornis (Canestrini, Fanzago, 1877) 5. Euconulus fulvus (O. F. Müller, 1774) 36. Dendrolaelaps rotundus Hirschmann, 1960 Helicidae 6. Arianta arbustorum (Linnaeus, 1758) 37. Veigaia cervus (Krämer, 1876) 7. Cepaea hortensis (O. F.Müller, 1774) 38. Veigaia exiqua (Berlese, 1917) Limacidae 39. Veigaia kochi (Trägardh, 1901) 8. Limax maximus Linnaeus, 1758 40. Veigaia nemorensis (C. L. Koch, 1839) 9. Malacolimax tenellus O. F. Müller, 1774 41. Parazercon sarekensis Willmann, 1939 10. Perpolita hammonis (Str¸m, 1765) 42. Prozercon kochi Sellnick, 1943 Punctidae 43. Zercon zelawaiensis Sellnick, 1944 11. Punctum pygmaeum (Draparnaud, 1801) 44. Zercon spatulatus C. L. Koch, 1839 Succineidae 45. Zercon forsslundi Sellnick, 1958 12. Succinea putris (Linnaeus, 1758) Oribatida Trichiidae Achipteriidae 13. Trichia hispida (Linnaeus, 1758) 46. Parachipteria punctata (Nicolet, 1855) Valloniidae Astegistidae 14. Vallonia pulchella (O. F. Müller, 1774) 47. Furcoribula furcillata (Nordenskiöld, 1901) Vertiginidae Camisiidae 15. Vertigo substriata (Jeffreys, 1833) 48. Heminothrus longisetosus Willmann, 1925 Arthropoda Malacostraca 49. Carabodes labyrinthicus (Michael, 1879) Isopoda 50. Carabodes ornatus Storkan, 1925 Oniscidae 51. Carabodes subarcticus Trägardh, 1902 16. Oniscus asellus Linnaeus, 1758 Chamobatidae Porcellionidae 52. Chamobates cuspidatus (Michael, 1884) 17. Porcellio scaber Latreille, 1804 Ceratozetidae 53. Diapterobates humeralis (Hermann, 1804) Pseudoscorpionida 54. Trichoribates novus (Sellnick, 1928) Chernetidae Dameidae 18. Chernes hahnii (C. L. Koch, 1839) 55. Spatiodamaeus verticillipes (Nicolet, 1855) Opiliones Eremaeidae Nemastomatidae 56. Eremaeus hepaticus C. L. Koch, 1835 19. Nemastoma lugubre (O. F. Müller, 1776) 57. Eueremaeus silvestris (Forsslund, 1956) Phalangiidae Euphthiracaridae 20. Phalangium opilio Linnaeus, 1758 58. Rhysotritia ardua (C. L. Koch, 1840) Galumnidae Acari 59. Galumna lanceata (Oudemans, 1900) Mesostigmata 60. Pergalumna nervosa (Berlese, 1914) Aceosejidae Liacaridae 21. Leioseius bicolor (Berlese, 1918) 61. Adoristes ovatus (C. L. Koch, 1839) 22. Leioseius insignis Hirschmann, 1963 Nothridae Eviphidae 62. Nothrus silvestris Nicolet, 1855 23. Eviphis ostrinus (C. L. Koch, 1836) Oppiidae Laelapidae 63. Microppia minus (Paoli, 1908) 24. Hypoaspis aculeifer (Canestrini, 1883) 64. Oppiella nova (Oudemans, 1902) 25. Hypoaspis vacua (Michael, 1891) Oribatulidae 26. Ololaelaps placentula (Berlese, 1887) 65. Oribatula tibialis (Nicolet, 1855)

130 Proc. Latvian Acad. Sci., Section B, Vol. 69 (2015), No. 3. Phenopelopidae 100. Rhyparochromus pini (Linaneus, 1758) 66. Eupelops torulosus (C. L. Koch, 1840) 101. Scolopostethus affinis (Schilling, 1829) Phthiracaridae Pentatomidae 67. Phthiracarus laevigatus (C. L. Koch, , 1844) 102. Palomena prasina (Linnaeus, 1761) 68. Steganacarus carinatus (C. L. Koch, 1841) Coleoptera 69. Steganacarus striculus (C. L. Koch, 1835) Carabidae Scheloribatidae 103. Calathus micropterus (Duftschmid, 1812) 70. Hemileius initialis (Berlese, 1908) 104. Carabus granulatus granulatus Linnaeus, 1758 71. Liebstadia similis (Michael, 1888) 105. Carabus hortensis hortensis Linnaeus, 1758 72. Scheloribates laevigatus (C. L. Koch, 1836) 106. Cychrus caraboides (Linnaeus, 1758) 73. Scheloribates latipes (C. L. Koch, 1844) 107. Notiophilus aquaticus (Linnaeus, 1758) Suctobelbidae 108. Notiophilus germinyi Fauvel, 1863 74. Suctobelbella subtrigona (Oudemans, 1916) 109. Platynus assimilis (Paykull, 1790) Tectocepheidae 110. Pterostichus niger niger (Schaller, 1783) 75. Tectocepheus velatus velatus (Michael, 1880) 111. Pterostichus nigrita (Paykull, 1790) Chilopoda 112. Pterostichus oblongopunctatus oblongopunctatus (Fabricius, 1787) Lithobiomorpha Coccinellidae Geophilidae 113. Coccinella septempunctata Linnaeus, 1758 76. Geophilus carpophagus Leach, 1814 Curculionidae 77. Pachymerium ferrugineum (C. L. Koch, 1835) 114. Brachyderes incanus (Linnaeus, 1758) Julidae 115. Hylobius abietis (Linnaeus, 1758) 78. Ommatoiulus sabulosus (Linnaeus, 1758) 116. Magdalis memnonia (Gyllenhal, 1837) Lithobiidae 117. Pissodes pini (Linnaeus, 1758) 79. Lithobius forficatus (Linnaeus, 1758) 118. Strophosoma capitatum (DeGeer, 1775) Insecta Geotrupidae Collembola 119. Geotrupes stercorarius (Linnaeus, 1758) Arrhopalitidae Lucanidae 80. Arrhopalites principalis Stach, 1945 Platycerus caraboides caraboides (Linnaeus, 1758) Dicyrtomidae Pselaphidae 81. Dicyrtoma fusca (Lucas, 1842) 120. Bryaxis puncticollis (Denny, 1825) Entomobryidae 121. Trimium brevicorne (Reichenbach, 1816) 82. Orchesella flavescens (Bourlet, 1839) Ptiliidae Isotomidae 122. Acrotrichis intermedia (Gillmeister, 1845) 83. Isotoma viridis Bourlet, 1839 Scaphidiidae 84. Isotomiella minor (Schäffer, 1896) 123. Scaphisoma agaricinum (Linnaeus, 1758) 85. Isotomurus maculatus (Schäffer, 1896) Scydmaenidae 86. Parisotoma notabilis (Schäffer, 1896) 124. Stenichnus collaris (Müller, Kunze, 1822) 87. Pseudanurophorus binoculatus Kseneman, 1934 Silphidae Katiannidae 125. Phosphuga atrata (Linnaeus, 1758) 88. Sminthurinus aureus (Lubbock, 1862) Staphylinidae Neanuridae 126. Acrulia inflata (Gyllenhal, 1813) 89. Friesea mirabilis (Tullberg, 1871) 127. Amischa analis (Gravenhorst, 1802) 90. Friesea truncata Cassagnau, 1958 128. Anotylus rugosus (Fabricius, 1775) Onychiuridae 120. Atheta crassicornis (Fabricius, 1792) 91. Mesaphorura tenuisensillata Rusek, 1974 130. Bisnius fimetarius (Gravenhorst, 1802) 92. Protaphorura armata (Tullberg, 1869) 131. Carpelimus corticinus (Gravenhorst, 1806) 93. Protaphorura subuliginata Gisin, 1956 132. Dinaraea aequata (Erichson, 1837) Sminthuridae 133. Drusilla canaliculata (Fabricius, 1787) 94. Allacma fusca (Linnaeus, 1758) 134. Gabrius splendidulus (Gravenhorst, 1802) Sminthurididae 135. Geostiba circellaris (Gravenhorst, 1806) 95. Sphaeridia pumilis (Krausbauer, 1898) 136. Mocyta fungi (Gravenhorst, 1806) Tomoceridae 137. Nudobius lentus (Gravenhorst, 1806) 96. Pogonognathellus flavescens (Tullberg, 1871) 138. Placusa tachyporoides (Waltl, 1838) Diplura 139. Sepedophilus littoreus (Linnaeus, 1758) Campodeidae 140. Sepedophilus testaceus (Fabricius, 1793) 97. Campodea staphylinus Westwood, 1842 141. Stenus clavicornis (Scopoli, 1763) Dermaptera Hymenoptera Forficulidae Formicidae 98. Forficula auricularia Linnaeus, 1758 142 Formica rufa Linnaeus, 1761 Heteroptera 143. Lasius flavus (Fabricius, 1781) Lygaeidae 144. Lasius niger (Linnaeus, 1758) 99. Gastrodes grossipes (DeGeer, 1773) 145. laevinodis Nylander, 1846

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