Importance of moss habitats for mesostigmatid mites (Acari

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http://journals.tubitak.gov.tr/zoology/

Turkish Journal of Zoology Turk J Zool(2018) 42: 673-683© TÜBİTAKdoi:10.3906/zoo-1712-6

Importance of moss habitats for mesostigmatid mites (Acari: Mesostigmata) in Romania

Minodora MANU1,*, Raluca Ioana BĂNCILĂ2,3, Marilena ONETE1

1Department of Ecology, Taxonomy and Nature Conservation, Institute of Biology Bucharest, Romanian Academy, Bucharest, Romania

2Faculty of Natural Sciences, University Ovidius Constanţa, Constanţa, Romania3Department of Biospeleology and Soil Edaphobiology, “Emil Racoviţă” Institute of Speleology, Romanian Academy, Bucharest, Romania

* Correspondence: [email protected]

1. IntroductionNatural forests are complex and mature terrestrial ecosystems. They are characterized by a wide variety of habitats (wood debris, litter fermentation layer, soil, moss layer, canopy, etc.) which offer proper environmental conditions for a high diversity of organisms (Cragg and Bardgett, 2001; Spiecker, 2003; Paquette and Messier, 2011; Garcia-Palacios et al., 2013). One of the most abundant invertebrate groups living in forest ecosystems are mites (Acari). The mite densities that have been reported from a square meter of surface and subsurface soil were between 50,000 and 250,000 individuals or even more (400,000 individuals) during the winter months (Wallwork, 1959; Peterson, 1982; Kethley, 1990; Krantz and Walter, 2009). Soil mites (Acari) play an important ecological role in forests, participating in soil formation processes (humification, mineralization, and nutrient flow), influencing fertility and productivity (Cragg and Bardgett, 2001; Zhang et al., 2001; Garcia-Palacios et al., 2013; Zhang et al., 2015). According to many studies, mites are useful indicators of the ecological stages of different habitats and their management measures, and are considered an appropriate taxon to use when we examine the hierarchical aspects of biodiversity (Ruf, 1998; Rutgers et al., 2009; Aspetti et al., 2010; Bolger et al., 2014). The majority of

Mesostigmata mites are predators, participating indirectly to the decomposition process, soil structure, and plant productivity, and directly to the population regulation of other edaphic invertebrate groups, such as springtails, enchytreids, and immature oribatids (Walter and Proctor, 1999). In forest ecosystems, soil mites from the order Mesostigmata are frequently collected from different forest microhabitats, including aphyllophorales fungi; black truffle; litter; soil; canopies; moss layer; rooting wood; bark beetle galleries; grass sod; excrement; dead wood; nests of birds, ants, or small mammals; rock cracks (Bajerlein et al., 2006; Gwiazdowicz, 2007; Salmane and Brumelis, 2008, 2010; Arroyo et al., 2010; Gwiazdowicz et al., 2011, 2012; Huhta et al., 2012; Kamczyc and Gwiazdowicz, 2013; Kamczyc and Skorupski, 2014; Queralt et al., 2014; Krawczyk et al., 2015; Dirilgen et al., 2016).

The literature shows that Mesostigmata fauna varies significantly between different microhabitats within forests (Madej et al., 2011). One of the most interesting forest microhabitats is moss. In Europe, the taxonomical or ecological studies on mesostigmatid mites have been focused mainly on moss from soil or moss from peatbogs (Mašán 2003a, 2003b; 2007; Kalȕz and Fenďa, 2005; Ujvári and Kontschán, 2007; Gwiazdowicz, 2007; Salmane and Brumelis, 2008; Skorupski et al., 2008; Mašán et al., 2008;

Abstract: This study aimed to characterize the composition of soil mite 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

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

Research Article

This work is licensed under a Creative Commons Attribution 4.0 International License.

https://orcid.org/0000-0002-4372-8208

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https://orcid.org/0000-0003-1439-9975

MANU et al. / Turk J Zool

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Madej et al., 2011; Arroyo et al., 2012, 2013; Seniczak et al., 2014; Ács and Kontschan, 2014, 2015; Salmane and Spungis, 2015; Mitchell et al., 2016). These studies have demonstrated that moss represents ecological corridors between isolated habitat patches, preventing or slowing down the process of disassembly of complex soil communities. Another positive role of this habitat is increased dispersal among habitat patches under harsh climatic conditions, maintenance of population sizes of vulnerable species, and favorable environment conditions. On the other hand, soil microarthropod communities from isolated habitats were found to be less resilient than those in more connected habitats (through moss), implying a role for dispersal in the recovery of impacted communities (Hoyle and Gilbert, 2004; Salmane and Brumelis, 2008; Perdomo et al., 2012; Bolger et al., 2014). A few studies were focused on moss from tree bark/trunk and the canopy, demonstrating that many of these species are essentially exclusively canopy dwellers (Arroyo et al., 2010, 2012). The species composition of soil mites of the order Mesostigmata in the soil/litter collected from rock cracks and crevices in Szczeliniec Wielki and Błędne Skały rock labyrinths in the area of Stołowe Mountains National Park was reported (Kamczyc and Skorupski, 2014).

In Romania, most ecological studies from forest ecosystems were focused only on moss from soil, as a component of the litter-fermentation layer (Solomon, 1980; Călugăr and Huţu, 2008; Manu, 2012; Manu et al., 2013). Only a few studies have been made on the moss from cliffs and rocky areas, revealing the affinity of mite populations for these types of ecosystems situated in

mountain areas close to the forests, in comparison with those from hilly regions (Manu, 2011; Manu and Onete, 2015).

Taking into consideration these data, some questions have arisen. Are the moss habitats characterized by the same composition of mesostigmatid fauna? Are these habitats important from the acarological conservation point of view? In this context, the main objectives of the present study are to determine the species composition of the mesostigmatid fauna from moss habitats, to study the mesostigmatid communities from several moss habitats, and to identify the distinct mite communities in the investigated samples.

2. Materials and methods2.1. Investigated areasIn order to investigate the mesostigmatid fauna from moss habitats (bark moss: BM; rock moss: RM; soil moss: SM), 15 forest ecosystems were analyzed (3 deciduous forests, 5 beech forests, 1 fir forest, 5 spruce forests, and 1 mixed forest), from 8 Romanian counties (Figure 1).

The moss habitats were sampled randomly, taking into consideration the presence of any type of them in the investigated ecosystem. The samples were collected using a metal square (10 × 10 cm). The sample depth was 4 cm. The study was performed in 2012–2013. The elevation ranged between 378 and 1445 m a.s.l.. All investigated ecosystems are mature (over 80 years) natural forests (Table 1). 2.2. Mite samplesThe moss samples (Sphagnum sp. and Polytrichum sp.) 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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period April–October 2012–2013, using a square metal core. The surface of 1 moss sample was 10 × 10 cm. In total, 240 moss samples were analyzed (80 moss samples for each substratum). The samples were taken randomly. The extraction of the mites lasted from 10 to 14 days, using the Berlese–Tullgren method as modified by Balogh (1972). The samples were kept in a refrigerator until the next extraction. The mesostigmatid fauna were preserved in ethyl alcohol (90%). The mites’ numbering and identification were performed using a Zeiss stereomicroscope and an Axioscope A1 Zeiss microscope (Oberkochen, Germany). Some of the mites were mounted whole on glass slides in Hoyer’s medium (Krantz and Walter, 2009). Several mite specimens were dissected under a stereoscopic microscope after clearing in lactic acid. Each body part was mounted in Hoyer’s medium or polyvinyl alcohol–lactic acid mixture (PVA) medium.

The mites were identified to species level using published identification keys (Ghilyarov and Bregetova 1977; Hyatt, 1980; Karg, 1993; Mašán, 2003a, 2003b; Mašán and Fenďa, 2004; Kalȕz and Fenďa, 2005; Mašán, 2007; Mašán et al., 2008; Mašán and Halliday, 2010). Species were grouped in suborders: Gamasina (G), Antennophorina (A), and Uropodina (U). All identified species are in the mite collection of the Institute of Biology Ecological Station in Posada.2.3 Data processingStatistical analyses were conducted using R 3.2.1 (R 236 Development Core Team, 2006: http://www.r-project.org).

We used generalized linear mixed models (GLMM) (Dormann et al., 2007) to test whether the main community features (total abundance and species number) are related to habitat type. The models were fitted using the lme4 package (Bates and Maechler, 2010). In these models,

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

No. Type of forest County Location Toponym Elevation(meters) North East Moss

habitat

1 Deciduous forest Prahova Bucegi Mountains Stânca Sf. Ana 1113 45.214451 25.312906 soil, cliff

2 Beech forest Prahova Bucegi Mountains Poiana Stânii 1241 45.222134 25.313077 soil, bark

3 Fir forest Prahova Bucegi Mountains

Cascada Urlătoarea 946 45.233446 25.315363 soil, cliff,

bark

4 Spruce forest Prahova Bucegi Mountains

CuibulDorului 1546 45.192843 25.265589 soil, bark

5 Beech forest Alba Trascău Mountains

Zlatna-Valealui Paul 581 46.063086 23.152062 cliff

6 Beech forest Hunedoara Parâng Mountains Jieţului gorges 1126 45.242928 23.315408 cliff

7 Beech forest Hunedoara Parâng Mountains Parâng resort 1103 45.23303 23.260859 soil, bark

8 Spruce forest Suceava Călimani Mountains

Valea 12Apostoli 1159 633192.53 517401.19 soil, bark

9 Spruce forest Bistriţa-Năsăud

Călimani Mountains

BistriţaBârgăului 1615 624616.82 508011.28 cliff

10 Beech forest Argeş Făgăras Mountains Cumpăniţa 868 45.260598 24.36062 soil, bark,

cliff

11 Deciduous forest Gorj Cheile Sohodorului

Cheile Sohodorului 378 45.081253 23.082243 cliff

12 Spruce forest Argeş Leota Mountains

Rudăriţa-Valea Cheii 1247 435312.9 523719.54 soil

13 Deciduous forest fir and spruce Dâmboviţa Leota

Mountains Valea Raciu 1034 45.15668 25.18393 soil

14 Spruce forest Dâmboviţa Leota Mountains

ValeaFrumuşelu 1445 45.17464 25.1933 soil


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habitat type was introduced as a fixed factor and the sites were used as random factors. With the estimates from the models, we performed pairwise comparisons among habitat types using the R multcomp package (Hothorn et al., 2008).

To model the multivariate response of mite species assemblage to habitat type, we applied a constrained correspondence analysis (CCA). Before analysis, the mite abundance was ln (x + 1) transformed to maintain normal distribution and to avoid the ‘arch effect’ in CCA (Ter Braak, 1986). The permutation procedure (based on 9999 cycles) was used to test the significance of explanatory variables in CCA (Oksanen et al., 2006). For comparison of the 3 habitat types, we used a linear discriminant function (LDF). The CCA was performed using the vegan package (Oksanen et al., 2006); LDF, using the BiodiversityR package (Kindt, 2014).

At the same time, we determined the following parameters using PAST software: dominance (D), Shannon index of diversity (H), and equitability (J) (Hammer et al., 2001).

The numerical density per square meter was calculated using the formula (Σ no. of individuals/no. of samples) × 1 m2/surface area of the soil core (Botnariuc and Vădineanu, 1982). The surface area of the soil core was 100 cm2.

3. Results We collected a total of 3018 mites belonging to 97 species, grouped in 3 suborders: Anntenophorina (1 family, 1 genus, and 1 species); Gamasina (10 families, 27 genera, and 87 species); Uropodina (2 families, 5 genera, and 9 species). The following families were identified: Celaenopsidae (Anntenophorina), Epicriidae, Parasitidae, Veigaiidae, Rhodacaridae, Ascidae, Phytoseiidae, Macrochelidae, Pachylaelapidae, Laelapidae, Zerconidae, Eviphididae

(Gamasina), Trachytidae, and Uropodidae (Uropodina) (Table 2; Appendix 1). If we considered the numerical abundance, the total value in all moss habitats was 3018 individuals. The highest abundance was obtained in soil moss habitat (1622 individuals), in comparison with bark moss, where the lowest abundance was recorded (627 individuals). The same tendency was identified on the numerical density index. Total numerical density was 3764 ind./m2, but the highest value was described for mite communities from soil moss (2028 ind./m2), followed by those from rock moss (951 individuals/m2), and by those from bark moss (785 ind./m2) (Table 2).

From a total of 97 species, the highest number of species (71) were identified in soil moss (SM), in comparison with the other 2 moss habitats, where the values of this parameter were almost similar (46 species in RM and 41 species in BM). These results are confirmed by the values of the Shannon index of diversity, which recorded the highest value in SM, in comparison with the other 2 habitats (RM and BM) (Table 2; Appendix 1).

The number of species was strongly correlated with sampling effort (Figure 2).

In the SM habitat, the mite populations were represented by some dominant species (Leptogamasus parvulus, Neopodocinum mrciaki, Trachytes aegrota, Veigaia nemorensis), which had the highest number of individuals (for all mite communities, dominance [D] = 0.1). In the BM habitat, species such as Leptogamasus parvulus, Pergamasus mediocris, Leptogamasus tectegynellus, Veigaia nemorensis, Zercon berlesei, and Zercon carpathicus were dominant (for all mite communities, D = 0.11). In the RM habitat, the highest numbers of individuals were recorded for the following dominant species: Paragamasus similis, Veigaia nemorensis, Zercon schweizeri, Zercon berlesei, 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 97No. of genera 23 20 27 36No. of families 10 11 15 14No. of suborders 2 2 3 3No. of individuals 627 769 1622 3018Numerical density 785 951 2028 3764Dominance D 0.11 0.25 0.1Shannon Hʹ 2.63 2.99 2.31Equitability J 0.71 0.7 0.6


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those from SM habitat (for all mite communities, D = 0.25) (Table 2). Taking into consideration the equitability index, we observed that in BM and RM, the species were represented by the closed values of numerical abundance (equitability J = 0.71 and 0.7, respectively), while in SM this parameter demonstrates that the equitability between mite populations is lower (J = 0.6) (Table 2).

On the other hand, the species that recorded low values of numerical abundance in all moss habitats were Arctoseius brevicheles, Arctoseius cetratus, Celaenopsis badius, Discourella modesta, Epicrius canestrini, Hypoaspis

astronomica, Ololaelaps placentula, Paragamasus robustus, and Proctolealaps pygmaeus (Appendix 1).

We found a significant effect of habitat type on both abundance (F [1.237] = 18.538, P <0.0001) and species richness (F [1.237] = 33.1821, P <0.0001). The pairwise comparisons among habitat types indicated that both total abundance and species richness were significantly higher in SM than in BM and RM. There was no significant difference between RM and BM (Table 3).

The CCA of the association between abundance of 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

AbundanceRM–BM –1.775 2.193 –0.809 0.697SM–BM 10.650 2.193 4.855 <0.0001SM–RM 12.425 2.193 5.665 <0.0001Species richnessRM–BM 0.225 0.392 0.574 0.834SM–BM 28.875 0.392 7.369 <0.0001SM–RM 26.625 0.392 6.795 <0.0001


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the upper right quadrate are associated with SM (Figure 3). The first and second axes accounted for 68.03% and 31.97%, respectively. Species Zercon triangularis and Zercon peltadoides were strongly associated with BM. In the lower quadrate, 12 species were highlighted to be strongly associated with RM.

LDF showed that based on the community structure of the mite species, LDF1 explained 72.9% of the variance and separated the 3 habitats (Figure 4).

On the one hand, the majority of mite populations were classified into 3 distinct groups. These groups are defined by the characteristic species for each moss habitat. In soil moss, 30.92% were from species identified only in this habitat. 14.43% of the mites were identified only in rock moss, and 8.24% only in bark moss. On the other hand, this analysis revealed the presence of 17 common species, which represented 17.52% of all identified mites.

4. DiscussionAnalyzing the numerical densities of mite populations, we observed that the highest value was obtained in soil moss (2028 ind./m2), followed by the value from rock moss (951 ind./m2), and the lowest value was from bark moss (785

ind./m2). The same tendency was obtained for numerical density (Table 2). In terms of the species diversity, the best conditions for the mites’ development appear to be in soil moss (68.87% from the total number of species), in comparison with rock moss (44.62%) and bark moss (39.77%).

If we make a comparison with other types of habitats from Europe, we discovered that the obtained values of the number of species are similar to those obtained for mesostigmatids from tree hollows (96 species), wood debris (91 species), and Aphyllophorales fungi (100 species), but are higher in comparison with those obtained from nests of small vertebrates (44–56 species), ant nests (26 species), bird nests (37 species), canopies (22 species), bark beetles (16 species), rock cracks and labyrinths (27 species), rotting wood (46 species), sod (23 species), leaf litter (35 species), black truffle (58 species), and forest soil (52–60 species). From a conservation point of view, we considered that these moss habitats are very important in comparison to others, due to the relatively high species diversity. If we take into consideration the numerical abundance of mites, the obtained value from moss habitats was close to 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 = Pachyseius 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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those from ant nests (136 individuals), canopies (176 individuals), bark beetles (1804 individuals), black truffle (305 individuals), sod (192 individuals), or rock cracks and labyrinths (251 individuals); but it was much lower than those of this parameter recorded in tree hollows (9006 individuals), nests of micromammals (27,097 individuals), bird nests (13,355 individuals), or forest leaf litter (6964 individuals) (Gwiazdowicz and Klemt, 2004; Mašán and Stanko, 2005; Bajerlein et al., 2006; Gwiazdowicz, 2007; Arroyo et al., 2010; Salmane and Brumelis, 2010; Gwiazdowicz et al., 2011, 2012; Kaczmarek et al., 2011, 2015; Kamczyc and Gwiazdowicz, 2013; Kamczyc and Skorupski, 2014; Queralt et al., 2014; Manu and Ion, 2014).

In general, habitats rich in organic matter are favorable for soil mites (Huhta et al., 2004; Gwiazdowicz et al., 2011; Kaczmarek et al., 2011; Manu, 2012; Garcia-Palacios et al., 2013; Bolger at al., 2014; Zhang et al., 2011, 2015). Moss habitats in particular have been associated with higher predatory mite diversity (Hoyle and Glibert, 2004; Perdomo et al., 2012). Mosses retain moisture by preventing evaporation in drought periods, thereby improving food resources and increasing habitat diversity (Salmane and Brumelis, 2008; Salmane and Spungis, 2015).

As for species diversity in soil moss, we observed that the values obtained in our study were much higher in comparison with those of similar habitats in Norway, Latvia, Ireland, Poland, and United Kingdom, which varied from 5 to 43 species (Gwiazdowicz and Kmita, 2004; Arroyo et al., 2010; Salmane and Brumelis, 2010; Madej et al., 2011; Seniczak et al., 2014; Mitchell et al., 2016). The same tendency was observed with the number of species from bark moss; in Ireland, e.g., this parameter had lower recorded values (3–11 species) (Arroyo et al., 2010). For rock moss, the single terms of comparison for the number of mesostigmatid species and abundance were obtained from habitats such as rocky cracks and labyrinths or from cliffs in Poland and Romania. The obtained parameters were lower, varying between 17 and 27 mite species, and from 134 to 251 individuals (Kamczyc and Skorupski, 2014; Manu and Onete, 2015).

Analyzing the mite community composition across a European transect, we observed that the species diversity is higher in continental areas than in alpine, Atlantic, or Mediterranean bioregions (Seniczak et al., 2014; Dirilgen et al., 2015). At the same time, the forests in the continental region provide the most favorable habitats for mites, in

Figure 4. Linear discriminant function scores of mite species from BM – bark moss, RM – rock moss, and SM – soil moss.


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comparison with meadows, shrubs, or other types of ecosystems (Gwiazdowicz, 2007; Călugăr and Huţu, 2008; Skorupski et al., 2008; Manu, 2011; Manu et al., 2013; Seniczak et al., 2015).

It is known that natural, undisturbed, mature forests, such as those of Romania, are complex and stable ecosystems. They are an inexhaustible source of ecological information about biodiversity, structure, natural processes, and overall functioning (Schnitzlera and Borleab, 1998; Parviainen, 2005; Pătru-Stupariu et al., 2013). These characteristics are due to factors that provide the ecosystem’s stability, such as species diversity (interactions, life strategies), trophic complexity (food web structure), and nutrient or energy flux. In natural forests of Romania, the biodiversity of these habitats (such as moss) is higher in comparison with artificial (planted) or disturbed ones (Spiecker, 2003; Moraza, 2009; Gwiazdowicz et al., 2011, Manu et al., 2013; Manu and Ion, 2014). If we extrapolate this affirmation to the moss habitats, it could explain the high mite diversity in comparison with other countries with temperate climates (e.g., Poland, Latvia, or United Kingdom).

Other studies concerning Mesostigmata mites from different types of ecosystems, which integrated soil moss habitat (such as peat bogs, fens, or different types of forest), mainly dominated by Sphagnum sp. or Polytrichum sp., offered varied information. In bogs in Ireland, 4 to 14 mesostigmatid species were described; in Poland, 35 species; in Latvia, 45 species (Skorupski et al., 2008; Salmane and Brumelis, 2008; Wisdom et al., 2011; Arroyo et al., 2013). In fens in Latvia, Salmane and Spungis (2015) described 28 mite species; in spruce forests (provided with a rich moss layer), 25 species with 1560 individuals. Studies from Romania made in spruce forests have shown high diversity (68 species), these values being close to that obtained in the soil moss habitats studied in the present research (Manu, 2012).

According to constrained correspondence analyses, the most favorable habitat for Mesostigmata mites was soil moss, in comparison with bark moss and rock moss. Different studies revealed that mosses have a buffering effect on soil temperature, and they may be very efficient in capturing N and P from precipitation. Soil mosses prevent humus moisture from evaporation, thereby improving food resources and offering favorable conditions for other invertebrates which represent a trophic source for predatory mites, such as those from the Mesostigmata order (Salmane and Brumelis, 2008; Madej et al., 2011; Perdona et al., 2012; Garcia-Palacios, 2013; Salmane and Spungis, 2015).

Dominant species such as Leptogamasus parvulus, L. tectegynellus, Neopodocinum mrciaki, Trachytes aegrota, Zercon berlesei, and Z. triangularis are predators, with a

wide ecological potency. These species are common for all 3 moss habitats. In general, they prefer spruce and mixed forests with soil layers rich in organic matter (detritus, moss, litter, mouldering wood substrates of various degree of decomposition) (Masan and Fend’a, 2004; Skorupski et al., 2008; Salmane and Brumelis, 2010; Arroyo et al., 2012; Manu and Ion, 2014).

According to the linear discriminant function, mite populations formed 3 distinct groups. On the one hand, soil, bark, and rock mosses provide specific environmental conditions. Some studies revealed that microarthropod communities depend on habitat connectivity, temperature, and trophic sources (Perdomo et al., 2012; Garcia-Palacios et al., 2013; Mitchell et al., 2016). Soil moss habitat has a direct connectivity with litter-fermentation and humus layers, providing favorable environmental conditions for mites (Hoyle and Gilbert, 2004; Salmane and Brumelis, 2008, 2010; Călugăr and Huţu, 2008; Arroyo et al., 2010; Manu, 2012). On the other hand, moss from the rock and bark provides isolated habitats. There are not significant exchanges of matter and energy between the main habitat (moss on rock or bark) and its substrate, and the abiotic conditions are more hostile (the lack of organic matter, increased temperature, decreased humidity, lower vegetation cover) (Manu et al., 2011, 2013; Kamczyc and Skorupski, 2014; Manu and Onete, 2015).

At the same time, according to the LDF analysis, some species are common for all types of moss habitats, such as Amblyseius meridionalis, Holoparasitus caesus, Leptogamasus parvulus, Pachylaelaps furcifer, Pergamasus crassipes, Trachytes aegrota, T. pauperior, Veigaia nemorensis, V. transisalae, and Zercon berlesei. All of these species are very mobile predatory mites, continuously searching for food. They have wide ecological plasticity. Veigaia nemoresis is an edaphic–detriticole species with the widest distribution in Romania, as well in Europe, from lowlands up to the alpine zone. It occurs in various soil microhabitats (roots, rock cracks, etc.) (Manu et al., 2017). Veigaia transisalae has a narrow distribution in Romania, from lowlands up to montane areas. It is frequent in soil microhabitats in coniferous forests (Manu et al., 2017). Species Zercon berlesei is well adapted to xerothermophilous phytocoenosis and to the chasmophytic vegetation of scree slopes and rocky areas, but is also found in coniferous forest, where grass rhizosphere, moss, soil detritus, and needle litter are its natural microhabitats (Mašàn and Fenďa, 2004). Pachylaelaps furcifer is distributed from the lowlands up to 1500 m a.s.l. It is found in deciduous and acid coniferous forests, but strongly prefers moist and humid habitats (Mašán, 2007). Trachytes aegrota and T. pauperior have wide ecological tolerance and inhabit various habitats (Mašán, 2003b, Bloszyk et al., 2006). All of these species have been identified in Europe not only


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in soil moss, but also in bark and rock moss (Mašàn and Fenďa, 2004; Salmane and Kontschan, 2005; Salmane and Brumelis, 2008, 2010; Mašán et al., 2008; Skorupski et al., 2008; Madej et al., 2011; Arroyo et al., 2010, 2012, 2013; Kamczyc and Skorupski, 2014; Ács and Kontschan, 2014; Salmane and Spungis, 2015).

In conclusion, according to the statistical analysis, distinct mite populations were identified in soil moss, bark moss, and rock moss. At the same time, all investigated moss habitats were characterized by dominant species (Leptogamasus parvulus, Pachylaelaps furcifer, Pergamasus crassipes, Trachytes aegrota, T. pauperior, Veigaia nemorensis, V. transisalae, and Zercon berlesei) which are common for temperate areas of Europe, and which have a wide ecological plasticity. The most favorable habitat was the soil moss, where numerical abundance and species

diversity were highest in comparison with the other 2 habitats.

If we take into consideration the high values of diversity and the presence of characteristic species, we conclude that these moss habitats, situated in natural undisturbed forests, are very important from the acarological conservation point of view. This study could be a reference point for other acarological research in disturbed or anthropized forest ecosystems.

AcknowledgmentsThis study was carried out in the framework of the project RO1567-IBB01/2018 from Institute of Biology-Bucharest, Romanian Academy. We thank Simona Plumb, Rodica Iosif, and Lazăr Dumitru for their assistance in the lab and the field.

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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 Celaenopsidae1 Celaenopsis badius Cela.badi A 0 0 1 1

Family Epicriidae2 Epicrius bureschi Epic.bure G 0 0 4 43 Epicrius canestrini Epic.cane G 0 0 1 14 Epirius tauricus Epic.taur G 0 0 6 6

Family Parasitidae5 Holoparasitus caesus Holo.caes G 2 5 9 166 Holoparasitus calcaratus Holo.calc G 0 1 1 27 Holoparasitus excisus Holo.exci G 1 3 0 48 Holoparasitus fortunatus Holo.fort G 2 6 1 99 Holoparasitus sp. Holo.sp G 0 0 7 710 Leptogamasus doinae Lept.doin G 0 0 7 711 Leptogamasus obesus Lept.obes G 0 1 0 112 Leptogamasus parvulus Lept.parv G 103 7 134 24413 Leptogamasus sp. 2 Lept.sp2 G 10 11 10 3114 Leptogamasus sp. 1 Lept.sp1 G 0 18 0 1815 Leptogamasus sp. 3 Lept.sp3 G 0 9 5 1416 Leptogamasus tectegynellus Lept.tect G 87 15 44 14617 Lysigamasus conus Lysi.conu G 0 0 7 718 Lysigamasus lapponicus Lysi.lapp G 1 0 5 619 Lysigamasus neoruncatellus Lysi.neor G 1 0 12 1320 Lysigamasus runcatelus Lysi.runc G 0 2 0 221 Lysigamasus sp. Lysi.sp G 0 0 6 622 Paragamasus robustus Para.robu G 1 0 0 123 Paragamasus similis Para.simi G 0 20 5 2524 Paragamasus sp. Par.sp G 0 0 16 1625 Parasitus lunulatus Para.lunu G 0 0 2 226 Parasitus sp. Pars.sp G 2 0 2 427 Pergamasus brevicornis Perg.brev G 0 0 3 328 Pergamasus crassipes Perg.cras G 14 10 33 5729 Pergamasus laetus Perg.laet G 0 0 23 2330 Pergamasus mediocris Perg.medi G 24 0 3 2731 Pergamasus sp. Perg.sp G 2 0 3 532 Vulgarogamasus trouessarti Vulg.trou G 2 0 0 2

Family Veigaiidae33 Veigaia cerva Veig.cerv G 8 1 13 2234 Veigaia exigua Veig.exig G 0 3 3 635 Veigaia kochi Veig.koch G 0 3 12 1536 Veigaia nemorensis Veig.nemo G 96 43 394 53337 Veigaia planicola Veig.plan G 3 0 0 3


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38 Veigaia transisalae Veig.tran G 5 2 3 10Family Rhodacaridae

39 Rhodacarellus silesiacus Rhod.sile G 0 0 2 2Family Ascidae

40 Arctoseius brevichelis Arct.brev G 0 0 1 141 Arctoseius cetratus Arct.cetr G 1 0 0 142 Arctoseius resinae Arct.resi G 0 0 7 743 Arctoseius semiscissus Arct.semi G 0 0 1 144 Asca aphidoides Asca.aphi G 0 7 0 745 Cheroseius sp. Chro.sp G 1 0 0 146 Gamasellodes bicolor Gama.bico G 6 1 0 747 Protogamasellus singularis Prot.sing G 2 0 0 248 Proctolaelaps pygmaeus Proc.pygm G 1 0 0 1

Family Phytoseiidae49 Amblyseius meridionalis Ambl.meri G 6 1 5 1250 Amblyseius obtusus Ambl.obtu G 1 0 6 7

Family Macrochelidae51 Geholaspis berlesei Geho.berl G 0 0 2 252 Geholaspis longisetosus Geho.long G 0 1 2 353 Macrocheles montanus Macr.mont G 0 2 1 354 Macrocheles recki Macr.reck G 0 12 0 1255 Neopodocinum mrciaki Neop.mrci G 3 2 244 249

Family Pachylaelapidae56 Olopachys vysotskajae Olol.vyso G 6 0 2 857 Pachydellus furcifer Pach.fu G 3 2 28 3358 Pachydellus vexillifer Pach.vexi G 0 0 2 259 Pachylaelaps pectinifer Pach.pect G 0 0 2 260 Pachyseius humeralis Pach.hume G 2 1 0 3

Family Laelapidae61 Hypoaspis aculeifer Hypo.acul G 2 0 20 2262 Hypoaspis astronomica Hypo.astr G 0 1 0 163 Hypoaspsis oblonga Hypo.oblo G 9 0 1 1064 Ololaelaps placentula Olol.plac G 0 0 1 1

Family Zerconidae65 Parazercon radiatus Para.radi G 7 0 21 2866 Prozercon carpathicus Proz.carp G 0 0 1 167 Prozercon carsticus Proz.cars. G 0 3 0 368 Prozercon fimbriatus Proz.fimb G 0 1 0 169 Prozercon kochi Proz.koch G 3 0 21 2470 Prozercon sellnicki Proz.sell G 1 0 4 571 Prozercon sp. Proz.sp G 0 4 0 472 Prozercon traegardhi Proz.trae G 0 0 32 32

73 Zercon arcuatus Zerc.arcu G 0 0 30 30

Appendix 1. (Continued).


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74 Zercon baloghi Zerc.balo G 0 8 0 875 Zercon berlesei Zerc.berl G 109 365 27 50176 Zercon carpathicus Zerc.carp G 43 0 26 6977 Zercon fageticola Zerc.fage G 0 7 0 778 Zercon foveolatus Zerc.fove G 1 3 13 1779 Zercon hungaricus Zerc.hung G 0 4 0 480 Zercon peltadoides Zerc.pelt1 G 1 11 38 5081 Zercon peltatus Zerc.pelt G 0 15 8 2382 Zercon romagniolus Zerc.roma G 0 0 10 1083 Zercon schweizeri Zerc.schw G 0 20 0 2084 Zercon sp. 1 Zerc.sp1 G 0 8 0 885 Zercon sp. 2 Zerc.sp2 G 0 20 6 2686 Zercon sp. 3 Zerc.sp3 G 0 0 11 1187 Zercon triangularis Zerc.tria G 0 92 12 104

Family Eviphididae88 Eviphis ostrinus Evip.ostr G 0 1 2 3

Family Trachytidae89 Trachytes aegrota Trac.aegr U 14 12 166 19290 Trachytes irenae Trac.iren U 0 0 16 1691 Trachytes lambda Trac.lamb U 0 0 4 492 Trachytes pauperior Trac.paup U 13 4 45 6293 Trachytes sp. Trac.sp U 0 0 22 22

Family Uropodidae94 Discourella modesta Disc.mode U 0 1 0 195 Polyaspinus sp. Poly.so U 0 0 4 496 Uroobovella sp. Uroo.sp U 0 0 1 197 Uropoda sp. Urop.sp U 28 0 0 28

Total individuals 627 769 1622 3018

Appendix 1. (Continued).

Documents

Importance of moss habitats for mesostigmatid mites (Acari