Turkish Journal of Zoology Turk J Zool (2018) 42: 673-683 http://journals.tubitak.gov.tr/zoology/ © TÜBİTAK Research Article doi:10.3906/zoo-1712-6

Importance of moss habitats for mesostigmatid (: ) in Romania

1, 2,3 1 Minodora MANU *, Raluca Ioana BĂNCILĂ , Marilena ONETE  1 Department of Ecology, and Nature Conservation, Institute of Biology Bucharest, Romanian Academy, Bucharest, Romania 2 Faculty of Natural Sciences, University Ovidius Constanţa, Constanţa, Romania 3 Department of Biospeleology and Soil Edaphobiology, “Emil Racoviţă” Institute of Speleology, Romanian Academy, Bucharest, Romania

Received: 04.12.2017 Accepted/Published Online: 10.09.2018 Final Version: 12.11.2018

Abstract: This study aimed to characterize the composition of soil populations (Acari: Mesostigmata) from 3 moss habitats (rock moss, bark moss, and soil moss). In total, 15 natural forest ecosystems were analyzed (3 deciduous forests, 5 beech forests, 1 fir forest, 5 spruce forests, and 1 mixed forest), from 8 counties in Romania. A total of 240 soil samples, 97 species, and 3018 individuals were analyzed. The samples were taken from April 2012 until October 2013. The highest numerical abundance and species diversity was found in the soil moss, in comparison with bark moss, where the lowest values were recorded. Using statistical analysis, we found a significant effect of habitat type on abundance and species richness, with mite communities grouped into 3 distinct classes. If we take into consideration the high diversity values and the presence of characteristic species (53.59% from the total number of mites from Romania), we conclude that these moss habitats, situated in natural undisturbed forests, are very important from a conservation point of view.

Key words: Abundance, bark, mite, moss, richness, rock, soil

1. Introduction Mesostigmata mites are predators, participating indirectly Natural forests are complex and mature terrestrial to the decomposition process, soil structure, and plant ecosystems. They are characterized by a wide variety of productivity, and directly to the population regulation habitats (wood debris, litter fermentation layer, soil, moss of other edaphic invertebrate groups, such as springtails, layer, canopy, etc.) which offer proper environmental enchytreids, and immature oribatids (Walter and Proctor, conditions for a high diversity of organisms (Cragg and 1999). In forest ecosystems, soil mites from the order Bardgett, 2001; Spiecker, 2003; Paquette and Messier, 2011; Mesostigmata are frequently collected from different forest Garcia-Palacios et al., 2013). One of the most abundant microhabitats, including aphyllophorales fungi; black invertebrate groups living in forest ecosystems are mites truffle; litter; soil; canopies; moss layer; rooting wood; bark (Acari). The mite densities that have been reported from a beetle galleries; grass sod; excrement; dead wood; nests square meter of surface and subsurface soil were between of birds, ants, or small mammals; rock cracks (Bajerlein 50,000 and 250,000 individuals or even more (400,000 et al., 2006; Gwiazdowicz, 2007; Salmane and Brumelis, individuals) during the winter months (Wallwork, 2008, 2010; Arroyo et al., 2010; Gwiazdowicz et al., 2011, 1959; Peterson, 1982; Kethley, 1990; Krantz and Walter, 2012; Huhta et al., 2012; Kamczyc and Gwiazdowicz, 2009). Soil mites (Acari) play an important ecological 2013; Kamczyc and Skorupski, 2014; Queralt et al., 2014; role in forests, participating in soil formation processes Krawczyk et al., 2015; Dirilgen et al., 2016). (humification, mineralization, and nutrient flow), The literature shows that Mesostigmata fauna varies influencing fertility and productivity (Cragg and Bardgett, significantly between different microhabitats within 2001; Zhang et al., 2001; Garcia-Palacios et al., 2013; Zhang forests (Madej et al., 2011). One of the most interesting et al., 2015). According to many studies, mites are useful forest microhabitats is moss. In Europe, the taxonomical indicators of the ecological stages of different habitats or ecological studies on mesostigmatid mites have been and their management measures, and are considered an focused mainly on moss from soil or moss from peatbogs appropriate taxon to use when we examine the hierarchical (Mašán 2003a, 2003b; 2007; Kalȕz and Fenďa, 2005; Ujvári aspects of (Ruf, 1998; Rutgers et al., 2009; and Kontschán, 2007; Gwiazdowicz, 2007; Salmane and Aspetti et al., 2010; Bolger et al., 2014). The majority of Brumelis, 2008; Skorupski et al., 2008; Mašán et al., 2008; * Correspondence: [email protected] 673

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Madej et al., 2011; Arroyo et al., 2012, 2013; Seniczak mountain areas close to the forests, in comparison with et al., 2014; Ács and Kontschan, 2014, 2015; Salmane those from hilly regions (Manu, 2011; Manu and Onete, and Spungis, 2015; Mitchell et al., 2016). These studies 2015). have demonstrated that moss represents ecological Taking into consideration these data, some questions corridors between isolated habitat patches, preventing have arisen. Are the moss habitats characterized by the or slowing down the process of disassembly of complex same composition of mesostigmatid fauna? Are these soil communities. Another positive role of this habitat is habitats important from the acarological conservation increased dispersal among habitat patches under harsh point of view? In this context, the main objectives of the climatic conditions, maintenance of population sizes of present study are to determine the species composition of vulnerable species, and favorable environment conditions. the mesostigmatid fauna from moss habitats, to study the On the other hand, soil microarthropod communities mesostigmatid communities from several moss habitats, from isolated habitats were found to be less resilient and to identify the distinct mite communities in the than those in more connected habitats (through moss), investigated samples. implying a role for dispersal in the recovery of impacted communities (Hoyle and Gilbert, 2004; Salmane and 2. Materials and methods Brumelis, 2008; Perdomo et al., 2012; Bolger et al., 2014). 2.1. Investigated areas A few studies were focused on moss from tree bark/trunk In order to investigate the mesostigmatid fauna from moss and the canopy, demonstrating that many of these species habitats (bark moss: BM; rock moss: RM; soil moss: SM), are essentially exclusively canopy dwellers (Arroyo et al., 15 forest ecosystems were analyzed (3 deciduous forests, 2010, 2012). The species composition of soil mites of the 5 beech forests, 1 fir forest, 5 spruce forests, and 1 mixed order Mesostigmata in the soil/litter collected from rock forest), from 8 Romanian counties (Figure 1). cracks and crevices in Szczeliniec Wielki and Błędne Skały The moss habitats were sampled randomly, taking rock labyrinths in the area of Stołowe Mountains National into consideration the presence of any type of them in the Park was reported (Kamczyc and Skorupski, 2014). investigated ecosystem. The samples were collected using a In Romania, most ecological studies from forest metal square (10 × 10 cm). The sample depth was 4 cm. The ecosystems were focused only on moss from soil, as a study was performed in 2012–2013. The elevation ranged component of the litter-fermentation layer (Solomon, between 378 and 1445 m a.s.l.. All investigated ecosystems 1980; Călugăr and Huţu, 2008; Manu, 2012; Manu et al., are mature (over 80 years) natural forests (Table 1). 2013). Only a few studies have been made on the moss 2.2. Mite samples from cliffs and rocky areas, revealing the affinity of mite The moss samples (Sphagnum sp. and Polytrichum sp.) populations for these types of ecosystems situated in were collected from soil, bark, and rocky areas, in the

Figure 1. Geographical description of the investigated ecosystems in Romania (https:// google-earth.en.softonic.com; accessed in 26.06.2017).

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Table 1. The geographical description of investigated forest ecosystems from Romania.

Elevation Moss No. Type of forest County Location Toponym North East (meters) habitat Bucegi 1 Deciduous forest Prahova Stânca Sf. Ana 1113 45.214451 25.312906 soil, cliff Mountains Bucegi 2 Beech forest Prahova Poiana Stânii 1241 45.222134 25.313077 soil, bark Mountains Bucegi Cascada soil, cliff, 3 Fir forest Prahova 946 45.233446 25.315363 Mountains Urlătoarea bark Bucegi Cuibul 4 Spruce forest Prahova 1546 45.192843 25.265589 soil, bark Mountains Dorului Trascău Zlatna-Valea 5 Beech forest Alba 581 46.063086 23.152062 cliff Mountains lui Paul Parâng 6 Beech forest Hunedoara Jieţului gorges 1126 45.242928 23.315408 cliff Mountains Parâng 7 Beech forest Hunedoara Parâng resort 1103 45.23303 23.260859 soil, bark Mountains Călimani Valea 12 8 Spruce forest Suceava 1159 633192.53 517401.19 soil, bark Mountains Apostoli Bistriţa- Călimani Bistriţa 9 Spruce forest 1615 624616.82 508011.28 cliff Năsăud Mountains Bârgăului Făgăras soil, bark, 10 Beech forest Argeş Cumpăniţa 868 45.260598 24.36062 Mountains cliff Cheile Cheile 11 Deciduous forest Gorj 378 45.081253 23.082243 cliff Sohodorului Sohodorului Leota Rudăriţa-Valea 12 Spruce forest Argeş 1247 435312.9 523719.54 soil Mountains Cheii Deciduous forest Leota 13 Dâmboviţa Valea Raciu 1034 45.15668 25.18393 soil fir and spruce Mountains Leota Valea 14 Spruce forest Dâmboviţa 1445 45.17464 25.1933 soil Mountains Frumuşelu period April–October 2012–2013, using a square metal The mites were identified to species level using core. The surface of 1 moss sample was 10 × 10 cm. In published identification keys (Ghilyarov and Bregetova total, 240 moss samples were analyzed (80 moss samples 1977; Hyatt, 1980; Karg, 1993; Mašán, 2003a, 2003b; for each substratum). The samples were taken randomly. Mašán and Fenďa, 2004; Kalȕz and Fenďa, 2005; Mašán, The extraction of the mites lasted from 10 to 14 days, 2007; Mašán et al., 2008; Mašán and Halliday, 2010). using the Berlese–Tullgren method as modified by Species were grouped in suborders: Gamasina (G), Balogh (1972). The samples were kept in a refrigerator Antennophorina (A), and Uropodina (U). All identified until the next extraction. The mesostigmatid fauna were species are in the mite collection of the Institute of Biology preserved in ethyl alcohol (90%). The mites’ numbering Ecological Station in Posada. and identification were performed using a Zeiss 2.3 Data processing stereomicroscope and an Axioscope A1 Zeiss microscope Statistical analyses were conducted using R 3.2.1 (R 236 (Oberkochen, Germany). Some of the mites were Development Core Team, 2006: http://www.r-project.org). mounted whole on glass slides in Hoyer’s medium (Krantz We used generalized linear mixed models (GLMM) and Walter, 2009). Several mite specimens were dissected (Dormann et al., 2007) to test whether the main community under a stereoscopic microscope after clearing in lactic features (total abundance and species number) are related acid. Each body part was mounted in Hoyer’s medium or to habitat type. The models were fitted using the lme4 polyvinyl alcohol–lactic acid mixture (PVA) medium. package (Bates and Maechler, 2010). In these models,

675 MANU et al. / Turk J Zool habitat type was introduced as a fixed factor and the sites (Gamasina), Trachytidae, and Uropodidae (Uropodina) were used as random factors. With the estimates from (Table 2; Appendix 1). If we considered the numerical the models, we performed pairwise comparisons among abundance, the total value in all moss habitats was 3018 habitat types using the R multcomp package (Hothorn et individuals. The highest abundance was obtained in soil al., 2008). moss habitat (1622 individuals), in comparison with bark To model the multivariate response of mite species moss, where the lowest abundance was recorded (627 assemblage to habitat type, we applied a constrained individuals). The same tendency was identified on the correspondence analysis (CCA). Before analysis, the mite numerical density index. Total numerical density was abundance was ln (x + 1) transformed to maintain normal 3764 ind./m2, but the highest value was described for mite distribution and to avoid the ‘arch effect’ in CCA (Ter communities from soil moss (2028 ind./m2), followed by Braak, 1986). The permutation procedure (based on 9999 those from rock moss (951 individuals/m2), and by those cycles) was used to test the significance of explanatory from bark moss (785 ind./m2) (Table 2). variables in CCA (Oksanen et al., 2006). For comparison of From a total of 97 species, the highest number of species the 3 habitat types, we used a linear discriminant function (71) were identified in soil moss (SM), in comparison (LDF). The CCA was performed using the vegan package with the other 2 moss habitats, where the values of this (Oksanen et al., 2006); LDF, using the BiodiversityR parameter were almost similar (46 species in RM and 41 package (Kindt, 2014). species in BM). These results are confirmed by the values of At the same time, we determined the following the Shannon index of diversity, which recorded the highest parameters using PAST software: dominance (D), Shannon value in SM, in comparison with the other 2 habitats (RM index of diversity (H), and equitability (J) (Hammer et al., and BM) (Table 2; Appendix 1). 2001). The number of species was strongly correlated with The numerical density per square meter was calculated sampling effort (Figure 2). using the formula (Σ no. of individuals/no. of samples) × 1 In the SM habitat, the mite populations were m2/surface area of the soil core (Botnariuc and Vădineanu, represented by some dominant species (Leptogamasus 1982). The surface area of the soil core was 100 cm2. parvulus, Neopodocinum mrciaki, Trachytes aegrota, nemorensis), which had the highest number of individuals 3. Results (for all mite communities, dominance [D] = 0.1). In the We collected a total of 3018 mites belonging to 97 species, BM habitat, species such as Leptogamasus parvulus, grouped in 3 suborders: Anntenophorina (1 family, 1 genus, Pergamasus mediocris, Leptogamasus tectegynellus, Veigaia and 1 species); Gamasina (10 families, 27 genera, and 87 nemorensis, Zercon berlesei, and Zercon carpathicus were species); Uropodina (2 families, 5 genera, and 9 species). dominant (for all mite communities, D = 0.11). In the RM The following families were identified: Celaenopsidae habitat, the highest numbers of individuals were recorded (Anntenophorina), Epicriidae, , , for the following dominant species: Paragamasus similis, , , , Macrochelidae, , Zercon schweizeri, Zercon berlesei, , Laelapidae, , Zercon triangularis, but the values are not as high as

Table 2: Population parameters of identified mite communities from inves- tigated moss habitats (BM – bark moss, RM – rock moss, and SM – soil moss).

Parameters BM RM SM Total No. of species 41 46 71 97 No. of genera 23 20 27 36 No. of families 10 11 15 14 No. of suborders 2 2 3 3 No. of individuals 627 769 1622 3018 Numerical density 785 951 2028 3764 Dominance D 0.11 0.25 0.1 Shannon Hʹ 2.63 2.99 2.31 Equitability J 0.71 0.7 0.6

676 MANU et al. / Turk J Zool those from SM habitat (for all mite communities, D = astronomica, Ololaelaps placentula, Paragamasus robustus, 0.25) (Table 2). Taking into consideration the equitability and Proctolealaps pygmaeus (Appendix 1). index, we observed that in BM and RM, the species were We found a significant effect of habitat type on both represented by the closed values of numerical abundance abundance (F [1.237] = 18.538, P <0.0001) and species (equitability J = 0.71 and 0.7, respectively), while in SM richness (F [1.237] = 33.1821, P <0.0001). The pairwise this parameter demonstrates that the equitability between comparisons among habitat types indicated that both total mite populations is lower (J = 0.6) (Table 2). abundance and species richness were significantly higher On the other hand, the species that recorded low in SM than in BM and RM. There was no significant values of numerical abundance in all moss habitats were difference between RM and BM (Table 3). Arctoseius brevicheles, , Celaenopsis The CCA of the association between abundance of badius, Discourella modesta, Epicrius canestrini, Hypoaspis mite species and the habitat showed that mite species in

Figure 2. Site-based accumulation curve for species richness of mite community for each habitat type: BM – bark moss (open circle), RM – rock moss (open triangle), and SM – soil moss (cross). The bars represent the 95% confidence intervals.

Table 3. The pairwise comparisons (multiple comparisons of means: Tukey contrasts) showing the effect of habitat type (BM – bark moss, RM – rock moss, and SM – soil moss) on the total abundance and species richness for the mite species.

Estimate SE Z P Abundance RM–BM –1.775 2.193 –0.809 0.697 SM–BM 10.650 2.193 4.855 <0.0001 SM–RM 12.425 2.193 5.665 <0.0001 Species richness RM–BM 0.225 0.392 0.574 0.834 SM–BM 28.875 0.392 7.369 <0.0001 SM–RM 26.625 0.392 6.795 <0.0001

677 MANU et al. / Turk J Zool the upper right quadrate are associated with SM (Figure ind./m2). The same tendency was obtained for numerical 3). The first and second axes accounted for 68.03% and density (Table 2). In terms of the species diversity, the 31.97%, respectively. Species Zercon triangularis and best conditions for the mites’ development appear to be Zercon peltadoides were strongly associated with BM. in soil moss (68.87% from the total number of species), In the lower quadrate, 12 species were highlighted to be in comparison with rock moss (44.62%) and bark moss strongly associated with RM. (39.77%). LDF showed that based on the community structure If we make a comparison with other types of habitats of the mite species, LDF1 explained 72.9% of the variance from Europe, we discovered that the obtained values of and separated the 3 habitats (Figure 4). the number of species are similar to those obtained for On the one hand, the majority of mite populations mesostigmatids from tree hollows (96 species), wood debris were classified into 3 distinct groups. These groups are (91 species), and Aphyllophorales fungi (100 species), but defined by the characteristic species for each moss habitat. are higher in comparison with those obtained from nests In soil moss, 30.92% were from species identified only in of small vertebrates (44–56 species), ant nests (26 species), this habitat. 14.43% of the mites were identified only in bird nests (37 species), canopies (22 species), bark beetles rock moss, and 8.24% only in bark moss. On the other (16 species), rock cracks and labyrinths (27 species), hand, this analysis revealed the presence of 17 common rotting wood (46 species), sod (23 species), leaf litter (35 species, which represented 17.52% of all identified mites. species), black truffle (58 species), and forest soil (52–60 species). From a conservation point of view, we considered 4. Discussion that these moss habitats are very important in comparison Analyzing the numerical densities of mite populations, we to others, due to the relatively high species diversity. If observed that the highest value was obtained in soil moss we take into consideration the numerical abundance of (2028 ind./m2), followed by the value from rock moss (951 mites, the obtained value from moss habitats was close to ind./m2), and the lowest value was from bark moss (785 that of wood debris (3621 individuals), and higher than

Figure 3. Biplots of the CCA model of the mite species abundance in relation to habitat type (Ht): BM – bark moss, RM – rock moss, and SM – soil moss. The abbreviations of species names: Amblimeri = Amblygamasus meridionalis; Holocaes = Holoparasitus caesus; Holoexci = Holoparasitus excisus; Leptparv = Leptogamasus parvulus; Lepttect = Leptogamasus tectegynellus; Lept1 = Leptogamasus sp. 1; Olopvyso = Olopachys vysotskajae; Pachhume = humeralis; Pergcras = Pergamasus crassipes; Pergmedi = Pergamasus medi- ocris; Prozkoch = Prozercon kochi; Tracaegr = Trachytes aegrota; Uropsp = Uropoda sp.; Zercarcu = Zercon arcuatus; Zercberl = Zercon berlesei; Zercpelt = Zercon peltatus; Zercpelt1 = Zercon peltadoides; Zerctria = Zercon triangularis; Veigcerv = Veigaia cerva; Veignemo = Veigaia nemorensis.

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Figure 4. Linear discriminant function scores of mite species from BM – bark moss, RM – rock moss, and SM – soil moss. those from ant nests (136 individuals), canopies (176 As for species diversity in soil moss, we observed individuals), bark beetles (1804 individuals), black truffle that the values obtained in our study were much higher (305 individuals), sod (192 individuals), or rock cracks and in comparison with those of similar habitats in Norway, labyrinths (251 individuals); but it was much lower than Latvia, Ireland, Poland, and United Kingdom, which those of this parameter recorded in tree hollows (9006 varied from 5 to 43 species (Gwiazdowicz and Kmita, 2004; individuals), nests of micromammals (27,097 individuals), Arroyo et al., 2010; Salmane and Brumelis, 2010; Madej et bird nests (13,355 individuals), or forest leaf litter (6964 al., 2011; Seniczak et al., 2014; Mitchell et al., 2016). The individuals) (Gwiazdowicz and Klemt, 2004; Mašán same tendency was observed with the number of species and Stanko, 2005; Bajerlein et al., 2006; Gwiazdowicz, from bark moss; in Ireland, e.g., this parameter had lower 2007; Arroyo et al., 2010; Salmane and Brumelis, 2010; recorded values (3–11 species) (Arroyo et al., 2010). For Gwiazdowicz et al., 2011, 2012; Kaczmarek et al., 2011, rock moss, the single terms of comparison for the number 2015; Kamczyc and Gwiazdowicz, 2013; Kamczyc and of mesostigmatid species and abundance were obtained Skorupski, 2014; Queralt et al., 2014; Manu and Ion, 2014). from habitats such as rocky cracks and labyrinths or from In general, habitats rich in organic matter are favorable cliffs in Poland and Romania. The obtained parameters for soil mites (Huhta et al., 2004; Gwiazdowicz et al., 2011; were lower, varying between 17 and 27 mite species, and Kaczmarek et al., 2011; Manu, 2012; Garcia-Palacios et from 134 to 251 individuals (Kamczyc and Skorupski, al., 2013; Bolger at al., 2014; Zhang et al., 2011, 2015). 2014; Manu and Onete, 2015). Moss habitats in particular have been associated with Analyzing the mite community composition across a higher predatory mite diversity (Hoyle and Glibert, 2004; European transect, we observed that the species diversity Perdomo et al., 2012). Mosses retain moisture by preventing is higher in continental areas than in alpine, Atlantic, or evaporation in drought periods, thereby improving food Mediterranean bioregions (Seniczak et al., 2014; Dirilgen resources and increasing habitat diversity (Salmane and et al., 2015). At the same time, the forests in the continental Brumelis, 2008; Salmane and Spungis, 2015). region provide the most favorable habitats for mites, in

679 MANU et al. / Turk J Zool comparison with meadows, shrubs, or other types of wide ecological potency. These species are common for all ecosystems (Gwiazdowicz, 2007; Călugăr and Huţu, 2008; 3 moss habitats. In general, they prefer spruce and mixed Skorupski et al., 2008; Manu, 2011; Manu et al., 2013; forests with soil layers rich in organic matter (detritus, Seniczak et al., 2015). moss, litter, mouldering wood substrates of various degree It is known that natural, undisturbed, mature of decomposition) (Masan and Fend’a, 2004; Skorupski et forests, such as those of Romania, are complex and al., 2008; Salmane and Brumelis, 2010; Arroyo et al., 2012; stable ecosystems. They are an inexhaustible source of Manu and Ion, 2014). ecological information about biodiversity, structure, According to the linear discriminant function, mite natural processes, and overall functioning (Schnitzlera populations formed 3 distinct groups. On the one hand, and Borleab, 1998; Parviainen, 2005; Pătru-Stupariu et soil, bark, and rock mosses provide specific environmental al., 2013). These characteristics are due to factors that conditions. Some studies revealed that microarthropod provide the ecosystem’s stability, such as species diversity communities depend on habitat connectivity, temperature, (interactions, life strategies), trophic complexity (food web and trophic sources (Perdomo et al., 2012; Garcia-Palacios structure), and nutrient or energy flux. In natural forests of et al., 2013; Mitchell et al., 2016). Soil moss habitat has a Romania, the biodiversity of these habitats (such as moss) is direct connectivity with litter-fermentation and humus higher in comparison with artificial (planted) or disturbed layers, providing favorable environmental conditions for ones (Spiecker, 2003; Moraza, 2009; Gwiazdowicz et mites (Hoyle and Gilbert, 2004; Salmane and Brumelis, al., 2011, Manu et al., 2013; Manu and Ion, 2014). If we 2008, 2010; Călugăr and Huţu, 2008; Arroyo et al., 2010; extrapolate this affirmation to the moss habitats, it could Manu, 2012). On the other hand, moss from the rock and explain the high mite diversity in comparison with other bark provides isolated habitats. There are not significant countries with temperate climates (e.g., Poland, Latvia, or exchanges of matter and energy between the main United Kingdom). habitat (moss on rock or bark) and its substrate, and the Other studies concerning Mesostigmata mites from abiotic conditions are more hostile (the lack of organic different types of ecosystems, which integrated soil moss matter, increased temperature, decreased humidity, lower habitat (such as peat bogs, fens, or different types of vegetation cover) (Manu et al., 2011, 2013; Kamczyc and forest), mainly dominated by Sphagnum sp. or Polytrichum Skorupski, 2014; Manu and Onete, 2015). sp., offered varied information. In bogs in Ireland, 4 to At the same time, according to the LDF analysis, 14 mesostigmatid species were described; in Poland, some species are common for all types of moss habitats, 35 species; in Latvia, 45 species (Skorupski et al., 2008; such as meridionalis, Holoparasitus caesus, Salmane and Brumelis, 2008; Wisdom et al., 2011; Arroyo Leptogamasus parvulus, furcifer, Pergamasus et al., 2013). In fens in Latvia, Salmane and Spungis (2015) crassipes, Trachytes aegrota, T. pauperior, Veigaia described 28 mite species; in spruce forests (provided nemorensis, V. transisalae, and Zercon berlesei. All of these with a rich moss layer), 25 species with 1560 individuals. species are very mobile predatory mites, continuously Studies from Romania made in spruce forests have shown searching for food. They have wide ecological plasticity. high diversity (68 species), these values being close to that Veigaia nemoresis is an edaphic–detriticole species with obtained in the soil moss habitats studied in the present the widest distribution in Romania, as well in Europe, from research (Manu, 2012). lowlands up to the alpine zone. It occurs in various soil According to constrained correspondence analyses, microhabitats (roots, rock cracks, etc.) (Manu et al., 2017). the most favorable habitat for Mesostigmata mites was Veigaia transisalae has a narrow distribution in Romania, soil moss, in comparison with bark moss and rock moss. from lowlands up to montane areas. It is frequent in soil Different studies revealed that mosses have a buffering microhabitats in coniferous forests (Manu et al., 2017). effect on soil temperature, and they may be very efficient in Species Zercon berlesei is well adapted to xerothermophilous capturing N and P from precipitation. Soil mosses prevent phytocoenosis and to the chasmophytic vegetation of scree humus moisture from evaporation, thereby improving slopes and rocky areas, but is also found in coniferous food resources and offering favorable conditions for forest, where grass rhizosphere, moss, soil detritus, and other invertebrates which represent a trophic source for needle litter are its natural microhabitats (Mašàn and predatory mites, such as those from the Mesostigmata Fenďa, 2004). Pachylaelaps furcifer is distributed from order (Salmane and Brumelis, 2008; Madej et al., 2011; the lowlands up to 1500 m a.s.l. It is found in deciduous Perdona et al., 2012; Garcia-Palacios, 2013; Salmane and and acid coniferous forests, but strongly prefers moist Spungis, 2015). and humid habitats (Mašán, 2007). Trachytes aegrota and Dominant species such as Leptogamasus parvulus, L. T. pauperior have wide ecological tolerance and inhabit tectegynellus, Neopodocinum mrciaki, Trachytes aegrota, various habitats (Mašán, 2003b, Bloszyk et al., 2006). All Zercon berlesei, and Z. triangularis are predators, with a of these species have been identified in Europe not only

680 MANU et al. / Turk J Zool in soil moss, but also in bark and rock moss (Mašàn and diversity were highest in comparison with the other 2 Fenďa, 2004; Salmane and Kontschan, 2005; Salmane and habitats. Brumelis, 2008, 2010; Mašán et al., 2008; Skorupski et al., If we take into consideration the high values of diversity 2008; Madej et al., 2011; Arroyo et al., 2010, 2012, 2013; and the presence of characteristic species, we conclude that Kamczyc and Skorupski, 2014; Ács and Kontschan, 2014; these moss habitats, situated in natural undisturbed forests, Salmane and Spungis, 2015). are very important from the acarological conservation In conclusion, according to the statistical analysis, point of view. This study could be a reference point for distinct mite populations were identified in soil moss, bark other acarological research in disturbed or anthropized moss, and rock moss. At the same time, all investigated forest ecosystems. moss habitats were characterized by dominant species (Leptogamasus parvulus, Pachylaelaps furcifer, Pergamasus Acknowledgments crassipes, Trachytes aegrota, T. pauperior, Veigaia This study was carried out in the framework of the project nemorensis, V. transisalae, and Zercon berlesei) which are RO1567-IBB01/2018 from Institute of Biology-Bucharest, common for temperate areas of Europe, and which have a Romanian Academy. We thank Simona Plumb, Rodica wide ecological plasticity. The most favorable habitat was Iosif, and Lazăr Dumitru for their assistance in the lab and the soil moss, where numerical abundance and species the field.

References

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Appendix 1. Numerical abundance of the mesostigmatid mites identified from investigated moss habitats (BM – bark moss, RM – rock moss, and SM – soil moss).

No. Species Code Suborder BM RM SM Total Family Celaenopsidae 1 Celaenopsis badius Cela.badi A 0 0 1 1 Family Epicriidae 2 Epicrius bureschi Epic.bure G 0 0 4 4 3 Epicrius canestrini Epic.cane G 0 0 1 1 4 Epirius tauricus Epic.taur G 0 0 6 6 Family Parasitidae 5 Holoparasitus caesus Holo.caes G 2 5 9 16 6 Holoparasitus calcaratus Holo.calc G 0 1 1 2 7 Holoparasitus excisus Holo.exci G 1 3 0 4 8 Holoparasitus fortunatus Holo.fort G 2 6 1 9 9 Holoparasitus sp. Holo.sp G 0 0 7 7 10 Leptogamasus doinae Lept.doin G 0 0 7 7 11 Leptogamasus obesus Lept.obes G 0 1 0 1 12 Leptogamasus parvulus Lept.parv G 103 7 134 244 13 Leptogamasus sp. 2 Lept.sp2 G 10 11 10 31 14 Leptogamasus sp. 1 Lept.sp1 G 0 18 0 18 15 Leptogamasus sp. 3 Lept.sp3 G 0 9 5 14 16 Leptogamasus tectegynellus Lept.tect G 87 15 44 146 17 Lysigamasus conus Lysi.conu G 0 0 7 7 18 Lysigamasus lapponicus Lysi.lapp G 1 0 5 6 19 Lysigamasus neoruncatellus Lysi.neor G 1 0 12 13 20 Lysigamasus runcatelus Lysi.runc G 0 2 0 2 21 Lysigamasus sp. Lysi.sp G 0 0 6 6 22 Paragamasus robustus Para.robu G 1 0 0 1 23 Paragamasus similis Para.simi G 0 20 5 25 24 Paragamasus sp. Par.sp G 0 0 16 16 25 Parasitus lunulatus Para.lunu G 0 0 2 2 26 Parasitus sp. Pars.sp G 2 0 2 4 27 Pergamasus brevicornis Perg.brev G 0 0 3 3 28 Pergamasus crassipes Perg.cras G 14 10 33 57 29 Pergamasus laetus Perg.laet G 0 0 23 23 30 Pergamasus mediocris Perg.medi G 24 0 3 27 31 Pergamasus sp. Perg.sp G 2 0 3 5 32 Vulgarogamasus trouessarti Vulg.trou G 2 0 0 2 Family Veigaiidae 33 Veigaia cerva Veig.cerv G 8 1 13 22 34 Veigaia exigua Veig.exig G 0 3 3 6 35 Veigaia kochi Veig.koch G 0 3 12 15 36 Veigaia nemorensis Veig.nemo G 96 43 394 533 37 Veigaia planicola Veig.plan G 3 0 0 3

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Appendix 1. (Continued).

38 Veigaia transisalae Veig.tran G 5 2 3 10 Family Rhodacaridae 39 silesiacus Rhod.sile G 0 0 2 2 Family Ascidae 40 Arctoseius brevichelis Arct.brev G 0 0 1 1 41 Arctoseius cetratus Arct.cetr G 1 0 0 1 42 Arctoseius resinae Arct.resi G 0 0 7 7 43 Arctoseius semiscissus Arct.semi G 0 0 1 1 44 Asca aphidoides Asca.aphi G 0 7 0 7 45 Cheroseius sp. Chro.sp G 1 0 0 1 46 Gamasellodes bicolor Gama.bico G 6 1 0 7 47 Protogamasellus singularis Prot.sing G 2 0 0 2 48 Proctolaelaps pygmaeus Proc.pygm G 1 0 0 1 Family Phytoseiidae 49 Amblyseius meridionalis Ambl.meri G 6 1 5 12 50 Amblyseius obtusus Ambl.obtu G 1 0 6 7 Family Macrochelidae 51 Geholaspis berlesei Geho.berl G 0 0 2 2 52 Geholaspis longisetosus Geho.long G 0 1 2 3 53 Macrocheles montanus Macr.mont G 0 2 1 3 54 Macrocheles recki Macr.reck G 0 12 0 12 55 Neopodocinum mrciaki Neop.mrci G 3 2 244 249 Family Pachylaelapidae 56 Olopachys vysotskajae Olol.vyso G 6 0 2 8 57 Pachydellus furcifer Pach.fu G 3 2 28 33 58 Pachydellus vexillifer Pach.vexi G 0 0 2 2 59 Pachylaelaps pectinifer Pach.pect G 0 0 2 2 60 Pachyseius humeralis Pach.hume G 2 1 0 3 Family Laelapidae 61 Hypoaspis aculeifer Hypo.acul G 2 0 20 22 62 Hypoaspis astronomica Hypo.astr G 0 1 0 1 63 Hypoaspsis oblonga Hypo.oblo G 9 0 1 10 64 Ololaelaps placentula Olol.plac G 0 0 1 1 Family Zerconidae 65 Parazercon radiatus Para.radi G 7 0 21 28 66 Prozercon carpathicus Proz.carp G 0 0 1 1 67 Prozercon carsticus Proz.cars. G 0 3 0 3 68 Prozercon fimbriatus Proz.fimb G 0 1 0 1 69 Prozercon kochi Proz.koch G 3 0 21 24 70 Prozercon sellnicki Proz.sell G 1 0 4 5 71 Prozercon sp. Proz.sp G 0 4 0 4 72 Prozercon traegardhi Proz.trae G 0 0 32 32 73 Zercon arcuatus Zerc.arcu G 0 0 30 30

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Appendix 1. (Continued).

74 Zercon baloghi Zerc.balo G 0 8 0 8 75 Zercon berlesei Zerc.berl G 109 365 27 501 76 Zercon carpathicus Zerc.carp G 43 0 26 69 77 Zercon fageticola Zerc.fage G 0 7 0 7 78 Zercon foveolatus Zerc.fove G 1 3 13 17 79 Zercon hungaricus Zerc.hung G 0 4 0 4 80 Zercon peltadoides Zerc.pelt1 G 1 11 38 50 81 Zercon peltatus Zerc.pelt G 0 15 8 23 82 Zercon romagniolus Zerc.roma G 0 0 10 10 83 Zercon schweizeri Zerc.schw G 0 20 0 20 84 Zercon sp. 1 Zerc.sp1 G 0 8 0 8 85 Zercon sp. 2 Zerc.sp2 G 0 20 6 26 86 Zercon sp. 3 Zerc.sp3 G 0 0 11 11 87 Zercon triangularis Zerc.tria G 0 92 12 104 Family Eviphididae 88 ostrinus Evip.ostr G 0 1 2 3 Family Trachytidae 89 Trachytes aegrota Trac.aegr U 14 12 166 192 90 Trachytes irenae Trac.iren U 0 0 16 16 91 Trachytes lambda Trac.lamb U 0 0 4 4 92 Trachytes pauperior Trac.paup U 13 4 45 62 93 Trachytes sp. Trac.sp U 0 0 22 22 Family Uropodidae 94 Discourella modesta Disc.mode U 0 1 0 1 95 Polyaspinus sp. Poly.so U 0 0 4 4 96 Uroobovella sp. Uroo.sp U 0 0 1 1 97 Uropoda sp. Urop.sp U 28 0 0 28 Total individuals 627 769 1622 3018

3