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Home , Amphigerontia bifasciata, Caecilius (genus), Caecilius fuscopterus, Cuneopalpus, Elipsocidae, Elipsocus

JOURNAL OF FOREST SCIENCE, 62, 2016 (5): 211–222

doi: 10.17221/97/2015-JFS

The first contribution to the fauna of psocids (Insecta: ) in forests in the Drahanská vrchovina Hills (Czech Republic)

D. Mazáč

Department of Forest Protection and Wildlife Management, Faculty of Forestry and Wood Technology, Mendel University in Brno, Brno, Czech Republic

ABSTRACT: Taxocenosis of psocids (Psocoptera) was studied in the territory of the Drahanská vrchovina Hills in the Czech Republic. Representative research plots were selected in forest ecosystems with natural species composition and spatial structure (small-scale strictly protected areas) as well as in forest ecosystems with altered species composi- tion and spatial structure. Research was conducted in three altitudinal vegetation zones (AVZ): in 2nd communities of Fagi-querceta s. lat. (- forests), 3rd Querci-fageta s. lat. (oak-beech forests) and 4th Fageta abietis (beech forests with fir). Research plots are situated at altitudes ranging between 275 and 540 m a.s.l. In the 2013 growing season, totally 3,474 imagoes and 2,532 nymphs of 32 psocid species were collected. Of those, 748 imagoes of 25 psocid species were collected in Fagi-querceta. The occurrence of burmeisteri, Caecilius flavidus and cruciatus was eudominant. 2,194 imagoes of 23 psocid species were found in Querci-fageta, eudominant were there Caecilius flavidus and Caecilius burmeisteri. 532 imagoes of 18 psocid species were found in Fageta abietis, eudominant were there: Caecilius flavidus, subfasciatus and Caecilius burmeisteri. In respect to the species composition, 3rd AVZ and 4th AVZ are similar to each other while 2nd AVZ is less similar.

Keywords: taxocenoses; ecology; forest ecosystems; altitudinal vegetation zones

Abbreviations: AVZ – altitudinal vegetation zone, 1st AVZ – oak (Querceta s. lat.), 2nd AVZ – beech-oak (Fagi-querceta s. lat.), 3rd AVZ – oak-beech (Querci-fageta s. lat.), 4th AVZ – beech (Fageta abietis s. lat.), 5th AVZ – fir-beech (Abieto-fageta s. lat.), 6th AVZ – -beech (Piceti-fageta s. lat.), 7th AVZ – beech-spruce (Fageti-piceeta s. lat), 8th AVZ – spruce (Piceeta s. lat.), 9th AVZ – mountain (Pineta mugo s. lat.), COR – corticicolous, FOL – folicolous, FSC – forest site type complex, HOL – holarctic, PAL – palearctic, WPA – west palearctic

So far the component has been paid in- ral ecosystems, and animal components are sufficient attention in the studies of forest ecosys- dynamically balanced (Míchal 1994). tems, with just a few exceptions. As pointed out by In history, the animal component of ecosystems Holuša (2003a), studying and their com- was studied in various units of the differentiation munities faces numerous obstacles caused for exam- of natural conditions, from altitudinal vegetation ple by the hidden way of the life of certain species, zones (AVZs) to forest site type classification units complexity of obtaining the study material, com- of forest site type complexes (FSCs) or biogeocene plicated, time-consuming and technically demand- type groups of the geobiocenological system. In ing determination and often incomplete knowledge the context of altitudinal vegetation zones, the ex- of the autecology of some species. The purpose of istence of relations between numerous groups of studying animals is to find regularities in their spe- invertebrates and selected vertebrate groups was cies composition within ecosystems as well as direct investigated. Stolina (1959) studied gradations of and indirect relations between the animal and the Zeiraphera diniana and Choristoneura muriana, plant component of the studied ecosystem. In natu- or Ips typographus within AVZ. Králíček and Po-

J. FOR. SCI., 62, 2016 (5): 211–222 211 volný (1978) and Povolný (1979) studied Lepi- slezské Beskydy Mts. in Mazák National Nature Re- doptera within AVZ. Scolytidae were studied by serve (Holuša 2003b), V Podolánkách Nature Re- Zumr (1984). Horník (1976) compared Carabidae serve (Holuša 2007b) and in the Oravské Beskydy and Cicindelidae in AVZ with natural and altered Mountains in Slovakia (Holuša 2012b). The remain- tree composition. Šustek (1993, 2000) studied the ing biogeographical areas of the Czech Republic have occurrence of Carabidae in AVZ. In the context of been paid much less attention. The Drahanská vr- complex biogeocenological research (i.e. continual chovina Hills, with regard to biogeographical char- research on the plant and the animal component) on acteristics, is unique by its position at the boundary the Smrk mountain in the Moravskoslezské Besky- between Hercynicum and Pannonicum. This specific dy Mountains, Roháčová (2001) counted Cara- location preconditions the occurrence of character- bidae and Heteroptera, Boháč and Roháčová istic animal as well as plant species and communities. (2001) counted Staphylinidae, Kočárek and Roháčová (2001) counted Silphidae, Holecová and Roháčová (2001) counted Curculionidae and METHODS Holuša (2001) counted Psocoptera. Povolný (1983) characterized the structure of Calliphoridae This study uses vegetation zoning according to in altitudinal vegetation zones. Plíva (1971). Characteristics and definition of Studies of the occurrence of psocid communi- vegetation tiers are used according to Holuša and ties in altitudinal vegetation zones have recently Holuša (2008, 2010, 2011). focused on the Carpathian Mountains. The relation Altitudinal vegetation zones are units of the for- between the vegetation zone and the occurrence of est site classification system (Plíva 1991) as well psocids was studied by Holuša (2013) in 3rd AVZ, as of the biogeocenological system (Zlatník 1959, i.e. communities of Querci-fageta s. lat., in 4th AVZ, 1975, 1976). This work uses FSCs as basic units of i.e. communities of Fageta abietis s. lat. (Holuša the forest typology classification system. Forest 2012a), in 5th AVZ, i.e. communities of Abieti-fageta types are grouped in FSCs according to their envi- s. lat. (Holuša 2007a), in 6th AVZ, i.e. communities ronmental relations and phytocenological similari- of Piceti-fageta s. lat. (Holuša 2011), in 7th AVZ, i.e. ties of the undergrowth. communities of Fagi-piceeta s. lat. and in 8th AVZ, i.e. Study area (Fig. 1). From the viewpoint of the communities of Piceeta s. lat. (Holuša 2007a), and regional geo-relief structure the research plots are other localities in the Carpathians – in the Moravsko- situated in the subunits of the Adamovská vrcho-

Fig. 1. The territory of the Drahanská vrchovina Hills

212 J. FOR. SCI., 62, 2016 (5): 211–222 vina Hills and the Moravian Karst, both forming geocenosis. During the collection, the greatest at- parts of the Drahanská vrchovina Hills unit, a part tention was focused on the synusia of woody of the Brněnská vrchovina Hills subsystem. The and undergrowth. The occurrence of psocids on Adamovská vrchovina Hills are characterized by the ground surface, on stones and rocks was also heterogeneous and varied topography. Its surface ascertained. The psocid material was obtained is built-up from a number of uplifted and sunken from accessible branches, low and branches small blocks. Typical features of the hilly land in- by knocking them off with a wooden bar 1.0 m clude a deep fault gap of the Svitava River with long into a sweep net. The sweep net was of circu- deeply incised meanders. The Moravian Karst dif- lar shape with a diameter of 0.4 m, and 0.8 m long fers considerably from the neighbouring areas. The handle. Psocids were knocked off the branches at geological bedrock consists of igneous rocks later heights up to 3.0 or 3.5 m, depending on accessi- covered with limestone layers (Bína, Demek 2012). bility. The branches were knocked off at a length The research plots are located in the Brno and of about 1.0 to 1.5 m from their end. Each tree or the Macocha bioregions of Czech Republic, which shrub type in the research plots was knocked off are part of the Hercynian subprovince. In the Brno separately. The undergrowth was swept with a bioregion the rich geological substrate and rug- sweep net of 0.4 m in diameter. Stubs, forest lit- ged terrain increase the overall biological diversity ter, ground surface, stones and other objects were (Culek 1996). The altitude of the research plots inspected visually. The knocked-off material was ranged from 275 to 540 m a.s.l. The plots are locat- removed from the sweep net and psocids were ed in quads 6665, 6666, 6765 and 6766 of the faunal sucked with an exhauster into a test tube and im- grid mapping (Pruner, Míka 1996). mediately fixed with 70% alcohol. Research plots were stabilized in forest ecosystems The material was collected from the plot centre in occurring in the Drahanská vrchovina Hills. The plots a spiral for each woody plant separately. One collec- were intended to represent a wide span of the degree tion included 50 strokes on more individuals of one of naturalness of forest stands (Fig. 2). The assess- woody plant species with 1 to 3 strokes per 1 m of ment of forest naturalness was based on classification the branch length. At sweeping, one collection rep- according to the survey of the natural species com- resents 50 sweeps, one sweep representing a sweep position of woody plants pursuant to the forest site net pull in the vegetation storey at a length of 2.5 m. type classification by Macků (2012), natural forests The collection was repeated twice at each plot for – naturalness degrees 4 (“high”) and 5 (“very high”), control, i.e. each woody plant (if possible) or vegeta- non-natural forests – naturalness degrees 3 (“medi- tion layer was subjected to 100 strokes. um”) and 2 (“low”). Forests stands with a low degree The collection was performed from May to Sep- of naturalness are characterized here by altered tree tember 2013. All research plots were visited twice species composition and spatial structure, and con- in that period. The following scale of dominance sist mostly of Abies alba monocultures in the 2nd AVZ was used to assess dominance: eudominant species – communities of Fagi-querceta s. lat. (partly also in more than 10%, dominant species 5–10%, subdomi- 3rd AVZ) and by Picea abies and Pseudotsuga men- nant species 2–5%, recedent species 1–2%, subrece- ziesii monocultures in 2nd, 3rd and 4th AVZ. All these dent species less than 1% (Losos et al. 1984). The forests are commercial forests without exception (§ 9 collected psocid material will be deposited in the of Act No. 289/1995 Coll.). Forest stands with a high collections of Otakar Holuša and partly stored in the degree of naturalness are represented by small-scale Beskyd Museum in Frýdek-Místek. The material was strictly protected areas, namely by nature reserves collected and determined by the author. and national nature reserves. Some research plots are situated in the nature reserve of the Moravian Karst, a large-scale strictly protected area. The range of for- RESULTS AND DISCUSSION ests of the varying naturalness is complemented by forest research plots with medium degree of natural- The total collected psocid material included 3,474 ness. Such a range from low to high was achieved in imagoes (Table 1) and 2,532 nymphs. The total num- all three AVZs. ber of found psocid species amounted to 32. Deter- The total number of plots amounted to 60, of mination and evaluation were performed only for which 20 are situated in the 2nd AVZ, 23 in the the imago material, while for the nymph material 3rd AVZ and 17 in the 4th AVZ. just counts were recorded. The following section The occurrence of psocids was studied within the shows the identified status of psocid taxocenoses research plots in individual layers of the forest bio- within the particular AVZs.

J. FOR. SCI., 62, 2016 (5): 211–222 213 Table 1. Abundance of psocids in altitudinal vegetation zones (AVZs), particularly in AVZs in the Drahanská vrchovina Hills

AVZ AVZ Psocid species Psocid species 2nd 3rd 4th total 2nd 3rd 4th total bifasciata 0 1 0 1 variegata 28 3 3 34 Amphigerontia contaminata 3 0 8 11 laticeps 0 4 0 4 Blaste quadrimaculata 4 0 0 4 Mesopsocus unipunctatus 1 0 0 1 Caecilius burmeisteri 416 574 75 1,065 2 11 14 27 Caecilius despaxi 0 13 0 13 8 5 0 13 Caecilius flavidus 86 1,097 188 1,371 1 2 0 3 0 4 3 7 Peripsocus parvulus 4 1 5 10 Caecilius piceus 52 129 25 206 4 6 0 10 cyanops 0 0 1 1 3 96 103 202 Ectopsocus meridionalis 3 8 3 14 Philotarsus parviceps 1 0 0 1 abdominalis 7 0 0 7 Philotarsus picicornis 2 6 13 21 Elipsocus annulatus 0 0 2 2 Psococerastis gibbosa 1 14 0 15 1 2 26 29 lachlani 0 8 0 8 Elipsocus moebiusi 23 59 21 103 Stenopsocus stigmaticus 1 0 4 5 85 133 26 244 Trichadenotecnum majus 10 14 12 36 pedicularia 1 0 0 1 Trichadenotecnum sexpunctatum 1 4 0 5 Total 748 2,194 532 3,474

2nd AVZ – beech-oak (Fagi-querceta s. lat.), 3nd AVZ – oak-beech (Querci-fageta s. lat.), 4th AVZ – beech-fir (Fageta abietis s. lat.)

Taxocenoses in the communities grees 3 and 2), the detected eudominant species in- of Fagi-querceta s. lat. (2nd AVZ) cluded Caecilius burmeisteri and the dominant spe- cies were Caecilius piceus and G. cruciatus (Fig. 2). In the Drahanská vrchovina Hills, the total num- P. abies yielded 40 imagoes. The eudominant species ber of collected psocid imagoes was 748 (25 species) included G. cruciatus, Elipsocus moebiusi, Trichade- (Table 1). In the forests with natural species com- notecnum majus, and Loen- position (naturalness degrees 4 and 5), eudominant sia variegata. The dominant species was Peripso- species included Caecilius flavidus and Graphopso- cus subfasciatus. P. menziesii yielded 38 imagoes. cus cruciatus. Quercus robur and Quercus petraea A rich species composition was found (13 species) yielded 40 imagoes (Table 2). The eudominant spe- with the eudominant occurrence of C. burmeisteri cies were C. flavidus and G. cruciatus. Carpinus and E. moebiusi. The dominant species included betulus yielded 36 imagoes with the eudominance G. cruciatus, L. variegata, Peripsocus phaeopterus of C. flavidus and G. cruciatus. In the forests of and Peripsocus alboguttatus. A. alba yielded a high non-natural species composition (naturalness de- number of 178 imagoes with eudominant C. bur-

450 30 Natural (degree 5 and 4) 400 25 Non-natural (degree 3 and 2) 350 300 20 250 15 (pcs) 200 150 10 100 5 50 0 0 Caecilius Grapho- Caecilius Caecilius flavidus psocus burmeisteri piceus cruciatus Elipsocus hyalinus Elipsocus moebiusi Peripsocus didymus Peripsocus parvulus Psococerastis gibbosa Philotarsus parviceps Philotarsus picicornis Elipsocus abdominalis Blaste quadrimaculata Peripsocus phaeopterus Peripsocus subfasciatus Peripsocus alboguttatus Stenopsocus stigmaticus Ectopsocus meridionalis Metylophorus nebulosus Trichadenotecnum majus Mesopsocus unipunctatus Amphigerontia contaminata Trichadenotecnum sexpunctatum

Fig. 2. Dominance of psocids in the Fagi-querceta s. lat (2nd AVZ) according to the degree of biotope naturalness (the diagram on the left with eudominant and dominant species occurrence, the remaining species on the right)

214 J. FOR. SCI., 62, 2016 (5): 211–222 1,200 140 Natural (degree 5 and 4) Non-natural (degree 3 and 2) 1,000 120 800 100 80

(pcs) 600 60 400 40 200 20 0 0 Caecilius Caecilius Caecilius flavidus piceus burmeisteri Caecilius despaxi Loensia variegata Elipsocus hyalinus Elipsocus moebiusi Peripsocus didymus Peripsocus parvulus Mesopsocus laticeps Stenopsocus lachlani Caecilius fuscopterus Psococerastis gibbosa Philotarsus picicornis Peripsocus phaeopterus Peripsocus subfasciatus Peripsocus alboguttatus Graphopsocus cruciatus Ectopsocus meridionalis Metylophorus nebulosus Trichadenotecnum majus Trichadenotecnum sexpunctatum

Fig. 3. Dominance of psocids in the communities of Querci-fageta s. lat (3rd AVZ) according to the degree of biotope natu- ralness (the diagram on the left with eudominant and dominant species occurrence, the remaining species on the right) meisteri and C. piceus. The undergrowth yielded sented by the species C. flavidus with a total num- 26 imagoes with the eudominant occurrence of ber of 1,097 specimens. In comparison with the C. flavidus, C. burmeisteri and G. cruciatus. The research into psocid taxocenoses in the Moravian species with the highest occurrence in the 2nd part of the Moravskoslezské Beskydy Mts. (Holuša AVZ was C. burmeisteri, in total 416 imagoes were 2001), the situation observed in the territory of the collected. Drahanská vrchovinaT Hills differed in the absence of Philotarsus parviceps as well as in lower counts of P. phaeopterus and dominant occurrence of Taxocenoses in the communities G. cruciatus. The same phenomenon was observed of Querci-fageta s. lat. (3rd AVZ) as in the Moravskoslezské Beskydy Mts., i.e. the colonization of the allochthonous P. abies by many In the 3rd AVZ, the total number of psocid imagoes species at considerable abundance. was 2,194 (23 species). Forests with natural species composition yielded the eudominant species C. fla- vidus (Fig. 3). Fagus sylvatica yielded 311 imagoes Taxocenoses in the communities with the eudominant (92%) occurrence of C. flavidus. of Fageta abietis s. lat. (4th AVZ) Q. robur and Q. petraea yielded 217 imagoes, again with the eudominance of C. flavidus (82%) together In the 4th AVZ, the total number of psocid ima- with G. cruciatus. C. betulus yielded 253 imagoes, goes was 532 in 18 species. Forests with the natural also with the eudominant (90%) occurrence of C. fla- species composition yielded the eudominant spe- vidus. Dominant species included G. cruciatus. cies C. flavidus and P. subfasciatus, and dominant cordata yielded 313 imagoes with the eudominant C. burmeisteri and G. cruciatus. F. sylvatica yielded (92%) occurrence of C. flavidus. Eudominant species 71 imagoes with the eudominant (89%) occurrence found in forests with altered tree species composition of C. flavidus. A. alba yielded 131 imagoes with the included C. burmeisteri, C. flavidus and C. piceus. eudominant (74%) P. subfasciatus. Dominant spe- Dominant species included G. cruciatus and P. subfas- cies included C. burmeisteri. Forests with altered ciatus. P. abies yielded 875 imagoes with the eudomi- tree species composition included the following nance of C. burmeisteri (63%) and C. piceus (Table 2). eudominant species: C. burmeisteri, C. piceus and Dominant species included P. subfasciatus and Philotarsus picicornis. Dominant species were Me- G. cruciatus. P. menziesii yielded 90 imagoes, eudo- tylophorus nebulosus, Elipsocus hyalinus, T. majus minant species being E. moebiusi, C. burmeisteri and and E. moebiusi. P. abies yielded 194 imagoes with P. subfasciatus. Dominant species included C. flavi- eudominant C. burmeisteri, E. hyalinus and C. pi- dus, T. majus and Psococerastis gibbosa. The under- ceus. Dominant species were E. moebiusi, G. cru- growth yielded in total 90 imagoes with the eudomi- ciatus, P. picicornis, T. majus and M. nebulosus. nant (98%) occurrence of C. flavidus. The highest total number of imagoes (188) in the The highest number of imagoes in the 3rd AVZ, 4th AVZ was found in the species C. flavidus (Fig. 4). i.e. communities of Querci-fageta s. lat., was repre- The occurrence of C. flavidus corresponds to the

J. FOR. SCI., 62, 2016 (5): 211–222 215

, 1 2 7 1 4 7 1 1 4 3 8 5 1 5 34 29 14 11 13 10 13 21 15 36 27 10 244 103 206 202 Total 1,371 1,065 3,474 3 2 1 1 1 8 2 3 13 26 15 63 22 12 11 10 sm 193 torminalis 3 1 2 2 92 po 100 6 1 4 2 3 1 1 4 jd 12 97 , powth – undergro 131 th 1 1 1 28 31 hb 4 532 , jrb – Sorbus 1 4 5 db alba 1 1 bzh 2 1 1 1 3 63 71 bk 2 4 4 5 1 5 1 8 3 2 2 1 60 10 12 79 sm 555 121 875 2 6 3 7 1 5 6 5 3 27 13 90 12 dgl 1 2 2 2 1 3 1 1 2 4 19 bo 1 1 88 90 po 2 2 1 3 1 1 1 3 lp 10 289 313 rd 3 2,194 3 1 jl 17 21 1 1 2 1 5 jd 6 4 1 1 14 hb 227 253 2 2 27 10 db 176 217 1 1 1 1 1 4 1 1 1 14 bk 285 311 AVZ 4 4 1 2 4 1 19 22 57 smp 4 1 7 5 2 2 2 1 6 1 1 2 12 18 sm 232 296 AVZ – beech-firabietis s., bo lat.), sylvatica ( Fageta – PinusFagus bk – sylvestris AVZ , brbr – Euonymus th 6 1 2 1 1 1 jd 32 134 178 3 3 5 1 1 1 1 1 1 2 1 2 16 38 dgl s. lat.), 4 1 6 9 1 1 bo 3 1 5 1 1 1 2 1 11 26 po

2 2 tr 2 3 3 8 sv 0 ptz nd 2 748 3 3 lp 1 4 1 8 14 jvb AVZ – oak-beech ( Querci-fageta AVZ 5 6 1 js nd 1 1 2 4 jrb 8 1 1 hl 10 2 9 1 1 1 1 21 36 hb 2 1 dr 10 13 23 17 40 db 5 3 8 brbr

AVZ – beech-oak s. ( Fagi-querceta lat.), 3 AVZ nd Loensia variegata Lachesilla pedicularia Elipsocus hyalinus Elipsocus moebiusi Elipsocus abdominalis Elipsocus annulatus Ectopsocus meridionalis Graphopsocus Graphopsocus cruciatus Cuneopalpus Cuneopalpus cyanops Caecilius burmeisteri Caecilius piceus Caecilius fuscopterus Blaste quadrimaculata Caecilius despaxi Caecilius flavidus Amphigerontia Amphigerontia contaminata Total/AVZ verticalthe dash-and-d ot line divides and altered tree species natural composition within each AVZ 2 Amphigerontia Amphigerontia bifasciata verrucosus , db – Quercus and Quercus robur petraea , dglPseudotsuga – menziesii , dr – Cornus mas , hb – Carpinus betulus, hl – , jdAbies – Tilia cordata excelsior, ptz – Ligustrum, sm – Picea vulgare – js campestre , lp – Tilia – Picea abies, jvb, smp Acer – , sv pungens – Cornus, tr – Prunus sanquinea avium Table 2. Abundance of psocids 2. Abundance on trees vegetation and undergrowth in altitudinal zones (AVZs) Table Metylophorus Metylophorus nebulosus Mesopsocus Mesopsocus unipunctatus Mesopsocus Mesopsocus laticeps Psocid species/ tree species Peripsocus parvulus Peripsocus phaeopterus Stenopsocus stigmaticus - Trichadenotec majus num Peripsocus didymus Philotarsus picicornis Psococerastis gibbosa Stenopsocus lachlani ctatum - Trichadenotec sexpun-num Total Peripsocus alboguttatus Peripsocus subfasciatus Philotarsus parviceps

216 J. FOR. SCI., 62, 2016 (5): 211–222

, 1 2 7 1 4 7 1 1 4 3 8 5 1 5 34 29 14 11 13 10 13 21 15 36 27 10 244 103 206 202 Total 1,371 1,065 3,474 3 2 1 1 1 8 2 3 13 26 15 63 22 12 11 10 sm 193 torminalis 3 1 2 2 92 po 100 6 1 4 2 3 1 1 4 jd 12 97 , powth – undergro 131 th 1 1 1 28 31 hb 4 532 , jrb – Sorbus 1 4 5 db alba 1 1 bzh 2 1 1 1 3 63 71 bk 2 4 4 5 1 5 1 8 3 2 2 1 60 10 12 79 sm 555 121 875 2 6 3 7 1 5 6 5 3 27 13 90 12 dgl 1 2 2 2 1 3 1 1 2 4 19 bo 1 1 88 90 po 2 2 1 3 1 1 1 3 lp 10 289 313 rd 3 2,194 3 1 jl 17 21 1 1 2 1 5 jd 6 4 1 1 14 hb 227 253 2 2 27 10 db 176 217 1 1 1 1 1 4 1 1 1 14 bk 285 311 AVZ 4 4 1 2 4 1 19 22 57 smp 4 1 7 5 2 2 2 1 6 1 1 2 12 18 sm 232 296 AVZ – beech-firabietis s., bo lat.), sylvatica ( Fageta – Pinus bk – Fagus sylvestris AVZ , brbr – Euonymus th 6 1 2 1 1 1 jd 32 134 178 3 3 5 1 1 1 1 1 1 2 1 2 16 38 dgl s. lat.), 4 1 6 9 1 1 bo 3 1 5 1 1 1 2 1 11 26 po

2 2 tr 2 3 3 8 sv 0 ptz nd 2 748 3 3 lp 1 4 1 8 14 jvb AVZ – oak-beech ( Querci-fageta AVZ 5 6 1 js nd 1 1 2 4 jrb 8 1 1 hl 10 2 9 1 1 1 1 21 36 hb 2 1 dr 10 13 23 17 40 db 5 3 8 brbr

AVZ – beech-oak s. ( Fagi-querceta lat.), 3 AVZ nd Loensia variegata Lachesilla pedicularia Elipsocus hyalinus Elipsocus moebiusi Elipsocus abdominalis Elipsocus annulatus Ectopsocus meridionalis Graphopsocus Graphopsocus cruciatus Cuneopalpus Cuneopalpus cyanops Caecilius burmeisteri Caecilius piceus Caecilius fuscopterus Blaste quadrimaculata Caecilius despaxi Caecilius flavidus Amphigerontia Amphigerontia contaminata Total/AVZ verticalthe dash-and-d ot line divides and altered tree species natural composition within each AVZ 2 Amphigerontia Amphigerontia bifasciata verrucosus , db – Quercus and Quercus robur petraea , dgl – Pseudotsuga menziesii , dr – Cornus mas, hb – Carpinus betulus , hl – Crataegus monogyna , jd – Abies Tilia cordata excelsior, ptz – Ligustrum, sm – Picea vulgare – Fraxinus js campestre , lp – Tilia – Picea abies, jvb, smp – Acer , sv pungens – Cornus, tr – Prunus sanquinea avium Table 2. Abundance of psocids 2. Abundance on trees vegetation and undergrowth in altitudinal zones (AVZs) Table Metylophorus Metylophorus nebulosus Mesopsocus Mesopsocus unipunctatus Mesopsocus Mesopsocus laticeps Psocid species/ tree species Peripsocus parvulus Peripsocus phaeopterus Stenopsocus stigmaticus - Trichadenotec majus num Peripsocus didymus Philotarsus picicornis Psococerastis gibbosa Stenopsocus lachlani ctatum - Trichadenotec sexpun-num Total Peripsocus alboguttatus Peripsocus subfasciatus Philotarsus parviceps

J. FOR. SCI., 62, 2016 (5): 211–222 217 200 25 Natural (degree 5 and 4) 180 Non-natural (degree 3 and 2) 160 20 140 120 15 100 (pcs) 80 10 60 40 5 20 s s i s i s s s s 0 s s

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Fig. 4. Graphical representation of the occurrence of psocids in the communities of Fageta abietis s. lat. (4th AVZ) ac- cording to the degree of biotope naturalness (the diagram on the left with eudominant and dominant species occurrence, the remaining species on the right) situation observed in the Moravian part of the such as Dictamnus albus, Brachypodium pinnatum Moravskoslezské Beskydy Mts. (Holuša 2001). Un- etc. Research plots in the 2nd AVZ also include forest like the Moravskoslezské Beskydy Mts., the Drahan- stands on azonal xerothermophilous sites. The sec- ská vrchovina Hills showed lower counts of E. moe- ond important reason for the impaired similarity is biusi and Mesopsocus unipunctatus in the natural the absence of beech communities. biogeocenosis. E. moebiusi dominated on allochtho- nous woody plants. The similarity of altitudinal vegetation zones was Comments on the significant psocid species examined with the help of the principal compo- nent analysis (Fig. 5). It follows from the analysis that 3rd AVZ and 4th AVZ are similar with regard to Caecilius flavidus (Stephens, 1836) psocid taxocenoses. Considerable similarity is given HOL; FOL – on trees, fallen leaves, sometimes also by beech communities naturally occurring in both on the vegetation and forest litter AVZs. Dissimilarity of the 2nd AVZ has two reasons. The species is eudominant in the Drahanská vr- The first of them is the extreme character of the sites chovina Hills in 2nd, 3rd and 4th AVZ (Table 1), and is predetermined by the occurrence of FSC 2X (Cor- represented on a majority of tree species, F. sylvatica neto-Fagi Quercetum xerothermicum) (Table 3). The yielded the highest numbers. In 2nd AVZ, the number main edificators are thermophilic shrubs and plants was lower due to the absence of beech trees but the species was also found on a majority of tree species. 1.0 Considerable occurrence was also found in the un- dergrowth. This was the most frequent species occur- ring in the area. On Králický Sněžník Mt. eudominant th th 0.5 occurrence was observed in 4 and 6 AVZ (Gol, Holuša 2011). Research in the Moravskoslezské Beskydy Mts. (Holuša 2001, 2003a) revealed the 0.0 abundant occurrence of C. flavidus in 5th and 6th AVZ, mainly on F. sylvatica. Factor 2: 20.06% Factor

–0.5 Caecilius burmeisteri (Brauer, 1876) HOL; FOL – this species prefers coniferous trees, occasionally also inhabiting broadleaves –1.0 The species is eudominant in the Drahanská vr- –1.0 –0.5 0.0 0.5 1.0 chovina Hills in 2nd, 3rd and 4th AVZ (Table 1), and Factor 1: 77.41% was found exclusively on conifers, especially on Fig. 5. Similarity of psocid taxocenoses according to the P. abies and A. alba (Table 2). Research in the Mo- altitudinal vegetation zones by using the PCA analysis in ravskoslezské Beskydy Mts. (Holuša 2001, 2003a) Statistica (Version 12, 2013) and on Králický Sněžník Mt. (Gol, Holuša 2011)

218 J. FOR. SCI., 62, 2016 (5): 211–222 Querceto - Longitude 16°40'24.44'' 16°40'59.66'' 16°41'05.67'' 16°41'19.37'' 16°41'22.22'' 16°41'32.92'' 16°40'57.60'' 16°41'15.44'' 16°41'39.84'' 16°40'53.00'' 16°41'35.84'' 16°41'39.09'' 16°41'20.95'' 16°41'24.26'' 16°41'42.02'' 16°41'05.46'' 16°41'24.05'' 16°41'21.98'' 16°41'25.90'' 16°41'17.77'' 16°41'12.65'' 16°39'37.85'' 16°39'49.14'' 16°39'37.42'' 16°39'52.42'' 16°39'50.68'' 16°39'44.48'' 16°39'43.77'' 16°41'12.37'' 16°41'12.11'' Latitude xerothermicum , 3B – 49°13'15.72'' 49°13'58.28'' 49°14'03.23'' 49°13'47.78'' 49°13'42.93'' 49°13'37.42'' 49°13'14.02'' 49°19'11.39'' 49°18'14.04'' 49°13'15.81'' 49°19'10.16'' 49°19'07.13'' 49°19'16.15'' 49°19'10.10'' 49°18'16.69'' 49°13'57.13'' 49°19'16.78'' 49°13'43.31'' 49°13'38.28'' 49°19'18.99'' 49°19'13.39'' 49°16'32.53'' 49°16'35.13'' 49°16'26.63'' 49°16'18.18'' 49°16'14.27'' 49°16'26.89'' 49°16'16.33'' 49°19'19.08'' 49°19'15.87'' 5 4 2 2 3 2 3 5 5 2 5 5 5 5 5 3 5 3 3 5 4 5 5 5 5 5 5 5 3 3 Nat. Fagetum lapidosum lapidosum , 4A – Fagetum mesotrophicum S S E E – – – – W W SE SE SE SE SE NE NE NE NE SW SW SW SW NW NW NW NW NW NW NW Exp. Fagi Quercetum Fagi Fagetum 3J 2S 4S 3S 3S 3S 2B 2B 4B 4B 2B 4B 4B 2X 2X 2X 4A 3A 3D 3D 2H 2H 2H 2H 3H 3H 3H 4W 4W 4A9 FSC 400 370 370 380 400 370 400 400 445 440 475 390 370 455 470 445 425 380 400 455 385 275 310 280 310 360 350 360 385 380 m a.s.l. Olomučany NR Malužín NR Býčí skála Kanice, Šumbera Kanice, NNR Habrůvecká bučina Cadastral area, local name , 2X – CornetoQuercetum deluvium acerosum - , 3S – Querceto trophicum - illimerosum Fagetum Longitude 16°38'36.45'' 16°38'40.02'' 16°38'19.22'' 16°38'29.58'' 16°39'01.49'' 16°39'02.73'' 16°38'54.90'' 16°37'56.17'' 16°36'59.52'' 16°36'48.43'' 16°36'51.14'' 16°37'04.47'' 16°38'57.15'' 16°38'30.87'' 16°38'35.34'' 16°37'18.86'' 16°37'57.89'' 16°36'58.15'' 16°37'26.74'' 16°38'27.94'' 16°38'21.94'' 16°40'40.62'' 16°40'58.95'' 16°40'57.29'' 16°41'01.59'' 16°40'38.04'' 16°40'48.91'' 16°40'25.07'' 16°40'45.92'' 16°40'29.31'' , 3H – Querceto - Latitude 49°17'36.30'' 49°17'35.76'' 49°17'31.08'' 49°17'27.95'' 49°17'45.68'' 49°17'45.50'' 49°17'35.63'' 49°17'57.09'' 49°21'37.55'' 49°21'42.31'' 49°21'41.19'' 49°21'31.12'' 49°17'35.74'' 49°17'31.36'' 49°17'28.32'' 49°21'47.04'' 49°21'34.49'' 49°21'41.86'' 49°21'50.15'' 49°17'37.84'' 49°17'37.11'' 49°13'24.08'' 49°13'43.30'' 49°13'40.36'' 49°13'47.94'' 49°13'42.15'' 49°13'45.47'' 49°13'26.16'' 49°13'32.85'' 49°13'21.44'' eluvium 2 3 3 3 2 2 2 2 3 3 4 4 2 4 5 4 3 3 4 4 4 5 5 5 5 5 5 5 5 5 Nat. S S S S – – – – – – – – – – – – – W W W W SE SW SW NW NW NW NW NW NW Exp. 3J 3S 4S 4S 4S 4S 4S 3B 3B 3B 3B 3B 3B 3B 3B 4B 4B 3B 2X 2X 2X 2X 2X 4A 2X 2X 2X 3D 2D 3H acerosum Fagetum FSC - 360 465 425 290 290 310 400 500 500 315 420 370 510 500 500 480 425 540 510 425 400 415 415 415 410 410 400 420 400 370 m a.s.l. Table 3. Description of research plots Drahanská vrchovina in the Hills Table Cadastral area, local name Vranov u Brna, Coufavá Vranov , Exp. mesotrophicum – geographic degree Nat. exposure, – naturalness , 4S – Fagetum trophicum illimerosum , 4H – Fagetum eutrophicum , 4B – Fagetum acidophilum NR Coufavá Hořice u Blanska,Hořice Hořice Nad Hořicemi u Blanska Hořicemi Nad NNR Hádecká planinka NR – Nature Reserve,- Reserve,NR – Nature Nature NNR – National complex: 2D – Fageto FSC site – forest , 3D – Querceto eutrophicum Fagetum

J. FOR. SCI., 62, 2016 (5): 211–222 219 revealed the occurrence of C. burmeisteri on P. abies, In the Drahanská vrchovina Hills subdominant oc- and only sporadic occurrence in higher AVZs and on currence was observed in 2nd AVZ, more on conifers. F. sylvatica. Individually, the species occurred on oth- Higher numbers were found on solitary coniferous er tree species and in the undergrowth. The second trees. Individual occurrence on conifers was recorded most frequent species occurring in the area. in 3rd and 4th AVZ.

Caecilius piceus (Kolbe, 1882) Trichadenotecnum majus (Kolbe, 1880) WPA; FOL – this species exclusively inhabits co- HOL; COR – on both broadleaves and conifers, al- niferous trees; occasionally it appears also on broad- ways preferring shaded biotopes with higher relative leaves, vegetation and coniferous litter humidity In the Drahanská vrchovina Hills, it is dominant in Recedent occurrence in the Drahanská vrcho- 2nd, 3rd and 4th AVZ (Table 1), occurring mostly on co- vina Hills was recorded in 2nd, 3rd and 4th AVZ. niferous trees, namely on P. abies and A. alba. Indi- In 2nd and 3rd AVZ, the highest occurrence was vidual occurrence was recorded also on the other tree on conifers, and only individual occurrence was species and in the undergrowth (Table 2). observed on broadleaves. In 4th AVZ, the spe- cies occurred only on coniferous trees (Table 2). The observed occurrence of psocid taxocenoses Peripsocus subfasciatus (Rambur, 1842) in the research plots of the Drahanská vrchovina HOL – probably missing in the Mediterranean; Hills can be compared with the results of research KOR – both on live and dry branches of conifers, less done mainly in the Moravskoslezské Beskydy on broadleaves Mts. and Podbeskydská pahorkatina Hills (com- Eudominant occurrence in the Drahanská vrchovi- pare Holuša 2003a). The only exception is the nd na Hills was observed in 4th AVZ on P. abies, subdom- 2 AVZ, which does not occur in these regions. inant occurrence in 3rd AVZ also on P. abies (Table 2). As shown in Figs 2–4, psocid species occurring in Individual occurrences on the other tree species were the individual AVZs depend on particular woody recorded in 2nd AVZ. The species occurred mainly in plants – on broadleaved tree species (Quercus sp., younger spruce stands (growth stages of small pole F. sylvatica, C. betulus etc.) in natural forests, and and pole timber). usually on conifers (mainly P. abies, A. alba and P. menziesii) in non-natural forests. In the central part of the diagram, there are species occurring Graphopsocus cruciatus (Linnaeus, 1768) in both natural forests and in forests with altered PAL; FOL – on deciduous trees and shrubs, partly tree species composition. Exactly these species can also on coniferous trees and in the undergrowth storey be used for the bioindication of forest sites. These Dominant occurrence in the Drahanská vrchovina species are not bound to a particular woody plant, Hills in 2nd and 3rd AVZ; the species occurs on both group of woody plants or other components of the phytocenosis but rather to macroclimate. broadleaved and coniferous trees. Subdominant oc- nd currence was recorded in 4th AVZ on both broad- 2 AVZ exhibited important species occurring leaves and conifers. in both natural and non-natural forests with al- tered tree species composition – C. flavidus, G. cru- ciatus, E. moebiusi, Peripsocus parvulus, T. majus, E. abdominalis. Elipsocus moebiusi (Tetens, 1891) The 3rd AVZ exhibited important species occur- PAL; COR – tree and shrub branches, shed leaves ring in both natural and non-natural forests with Subdominant occurrence in the Drahanská vrcho- altered tree species composition, mainly C. flavi- nd rd th vina Hills in 2 , 3 and 4 AVZ, mainly on the repre- dus, Ectopsocus meridionalis, P. gibbosa, Peripso- sented coniferous tree species, especially in the lower, cus didymus and Trichadenotecnum sexpunctatum. dry parts of crowns of younger coniferous stands at Another interesting species occurring in the psocid the growth stage of small-pole and pole timber. taxocenoses was Caecilius despaxi. This species is found in the communities of psocids occurring in the Moravskoslezské Beskydy Mts. only from the Loensia variegata (Latreille, 1799) 5th AVZ, in which the altitudinal vegetation zone WPA; COR – living on live and dry branches of considerably differs from the th4 AVZ. C. despaxi is both broadleaves and conifers, also on shrub branches described as a “companion” to forests with P. abies and occasionally on stones and occurs up to the 9th AVZ – communities of

220 J. FOR. SCI., 62, 2016 (5): 211–222 Pineta mugo (Holuša 2003a). The species was found AVZs was described (Holuša 2007a, 2011, 2012a, in 13 imago specimens in 3rd AVZ while no occur- 2013). rence was observed in the other AVZs. Similarly like In the 2013 growing season, the total number of in 4th AVZ of the Moravskoslezské Beskydy Mts., the psocid imagoes found in the area of interest was allochthonous character of P. abies was confirmed 3,474. The number covered 33 species, which is also in 3rd AVZ of the Drahanská vrchovina Hills due more than 25% of the total number of species to very low counts of C. despaxi in stands with altered known in the Czech Republic. Species dominance tree species composition as compared with the den- was evaluated for the respective study areas, veg- sity in stands with the natural occurrence of P. abies. etation zones and tree species. The most frequent The occurrence of Stenopsocus lachlani, which was species was C. flavidus, dominating on F. sylvatica, found in eight individuals in stands with altered tree Q. robur, C. betulus and T. cordata, i.e. in forest species composition is worth noticing. The species is ecosystems with the natural species composition characterized in the Moravskoslezské Beskydy Mts. corresponding to the 2nd, 3rd and 4th AVZ. The spe- as accompanying P. abies with eudominant and dom- cies is often denoted a “companion of beech” in inant occurrence from 3rd AVZ, to which it descends forest ecosystems. A species dominant in altered from higher AVZs (Holuša 2003a). Similarly like in biogeocenoses mainly composed of P. abies mono- C. despaxi, its low abundance may be explained by cultures was C. burmeisteri. This species is usu- the occurrence of allochthonous P. abies. ally bound to coniferous stands. Another species The species found in th4 AVZ and occurring both bound to conifers and occurring in abundance in in forest stands with the natural species composi- these stands was P. subfasciatus. As to the occur- tion and in altered forest stands included G. cru- rence of the psocid species, the most important are ciatus, E. hyalinus, E. meridionalis, E. moebiusi and those found in forest ecosystems with a high degree P. parvulus. Interesting was the occurrence of Stenop- of naturalness and at the same time in forest eco- socus stigmaticus because the research in the Mo- systems with a low degree of naturalness. ravskoslezské Beskydy Mts. indicated its attendance mainly in floodplain biogeocenoses, in water-logged sites. No occurrence was reported in other AVZs References (Holuša 2003a). In the Drahanská vrchovina Hills, the species occurred in 4 specimens in 4th AVZ (plus Bína J., Demek J. (2012): Z nížin do hor: geomorfologické 1 specimen in 2nd AVZ). The species was found in for- jednotky České republiky. Prague, Academia: 343. est stands with unaltered tree species composition. Culek M. (1996): Biogeografické členění České republiky. Prague, Enigma: 347. Boháč J., Roháčová M. (2001): Společenstva drabčíkovitých CONCLUSIONS (Coleoptera: Staphylinidae) Přírodní rezervace Smrk v Moravskoslezských Beskydech. Práce a studie Muzea The purpose of the study was to establish the struc- Beskyd, 11: 53–65. ture of the taxocenoses of a model group of psocids in Gol J., Holuša O. (2011): Struktura taxocenóz pisivek (Insecta: forest ecosystems of the Drahanská vrchovina Hills. Psocoptera) v lesních geobiocenózách v oblasti Králického The main intention was to determine the abundance Sněžníku. Acta Musei Beskidensis, 3: 71–84. of populations and the psocid species composition in Holecová M., Roháčová M. (2001): Druhové zastúpenie the individual AVZs and on individual tree species. nosáčikovitých (Coleoptera, Curculionidae) v epigeóne The objectives included a comparison of the species horského lesa (Moravskoslezské Beskydy, Česká republika). composition of psocid taxocenoses in forest stands Práce a studie Muzea Beskyd, 11: 75–82. with the natural species composition and in altered Holuša O. (2001): Příspěvek k poznání fauny pisivek (Insecta: forest stands and in the Drahanská vrchovina Hills Psocoptera) přírodní rezervace Smrk (Beskydský bioregion, with the situation recorded in the Moravskoslez- Česká republika). Práce a studie Muzea Beskyd, 11: 83–97. ské Beskydy Mts. Results of one-year research were Holuša O. (2003a): Vegetační stupňovitost a její bioindikace processed into the occurrence of psocids in natural, pomocí řádu pisivek (Insecta: Psocoptera). [Ph.D. Thesis.] close to natural and non-natural forest biogeoceno- Brno, Mendel University in Brno: 258. ses. Bioindicative properties of psocids were already Holuša O. (2003b): Fauna pisivek (Insecta: Psocoptera) proved by the core study from the area of the Moravs- Národní přírodní rezervace Mazák (Beskydský bioregion, koslezské Beskydy Mts. (Holuša 2003a). Subsequent Česká republika). Práce a studie Muzea Beskyd, 13: 83–98. studies in the same territory confirmed these bio- Holuša O. (2007a): Composition of psocid taxocenoses (In- indicative properties and the relation of psocids to secta: Psocoptera) in Fageti-Piceeta s. lat. and Piceeta s. lat.

J. FOR. SCI., 62, 2016 (5): 211–222 221 forests in the Western Carpathian Mts. Journal of Forest Macků J. (2012): Methodology for establishing of forest stands. Science, 53 (Special Issue): 3–10. Acta Universitatis Agriculturae et Silviculturae Mendelianae Holuša O. (2007b): Příspěvek k poznání fauny pisivek (Insecta: Brunensis, 60: 161–165. Psocoptera) Přírodní rezervace V Podolánkách v Moravsko- Míchal I. (1994): Ekologická stabilita. Brno, Veronica: 275. slezských Beskydech (Beskydský bioregion, Česká republika). Plíva K. (1971): Typologický systém ÚHÚL. Brandýs nad Labem, Práce a studie Muzea Beskyd, 19: 127–141. Institute for Forest Economy in Brandýs nad Labem: 90. Holuša O. (2011): Psocid taxocenoses structure and diversity Plíva K. (1991): Funkčně integrované lesní hospodářství. 1 – (Insecta: Psocoptera) in the forest ecosystems of the Piceeti- Přírodní podmínky v lesním plánování. Brandýs nad Labem, fageta s. lat. zone in the Western Carpathian Mts. 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International Symposium über Entomofaunistik in in the Oravské Beskydy Mts. (Slovakia). Acta Universitatis Agri- Mitteleuropa (SEEC), Budapest, Aug 15–20, 1983: 22–25. culturae et Silviculturae Mendelianae Brunensis, 60: 117–128. Pruner L., Míka P. (1996): Seznam obcí a jejich částí v České Holuša O. (2013): Psocid taxocenoses (Insecta: Psocoptera) in republice s čísly mapových polí pro síťové mapování fauny. the forest ecosystems of the Querci-fageta s. lat. zone in the Klapalekiana, 32 (Suppl.): 1–115. Western Carpathian Mts. and Polonic subprovincy. Acta Roháčová M. (2001): Entomocenózy Přírodní rezervace Smrk Universitatis Agriculturae et Silviculturae Mendelianae (Moravskoslezské Beskydy) na příkladě střevlíkovitých (Co- Brunensis, 61: 637–646. leoptera: Carabidae) a ploštic (Heteroptera). Práce a studie Holuša O., Holuša J. (2008): Characteristics of 3rd (Querci- Muzea Beskyd, 11: 23–51. fageta s. lat.) and 4th (Fageta (abietis) s. lat.) vegetation tiers of Stolina M. 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Corresponding author: Ing. David Mazáč, Mendel University in Brno, Faculty of Forestry and Wood Technology, Department of Forest Protection and Wildlife Management, Zemědělská 3, 613 00 Brno, Czech Republic; e-mail: [email protected]

222 J. FOR. SCI., 62, 2016 (5): 211–222