JOURNAL OF FOREST SCIENCE, 53, 2007 (Special Issue): 3–10

Composition of psocid taxocenoses (Insecta: Psocoptera) in Fageti-Piceeta s. lat. and Piceeta s. lat. forests in the Western Carpathian Mts.

O. Holuša

Faculty of Forestry and Wood Technology, Mendel University of Agriculture and Forestry in Brno, Brno, Czech Republic

Abstract: Psocid taxocenoses (Psocoptera) were studied in forest ecosystems of the Western Carpathian Mts. during 1997–2001. As a study frame were used altitudinal vegetation zones (according to Plíva 1971, 1991). Lower units of forest typological system (forest type complexes) were used for a classification of ecological conditions as well. Within this work can be the term “mountain spruce forest” understood as following communities of altitudinal vegetation zones (AVZ): the 7th – Fageti-Piceeta s. lat. and the 8th – Piceeta s. lat. These AVZ occur in the study area in the Moravskoslezské Beskydy Mts. in the Czech Republic and the Oravské Beskydy Mts. in the Slovakia 2,461 adults comprising 16 species were found in total: 12 species (eudominant species Caecilius despaxi, Mesopsocus unipunc- tatus, dominant species Stenopsocus lachlani, Amphigerontia bifasciata and Caecilius burmeisteri) were found in the 7th AVZ and an equal number of species was found in the 8th AVZ (eudominant species Caecilius despaxi, Stenopsocus lachlani). Taxocenoses of psocids were evaluated by Detrended Correspondence Analysis (DCA) and Divisive Cluster Analysis (DvClA). Material was compared with other material gained from various altitudinal vegetation zones in the Outer Western Carpathians Mts. Characteristic species composition of psocids in the 7th and 8th altitudinal vegeta- tion zones were designated: the 7th AVZ – Caecilius despaxi – Amphigerontia bifasciata – Mesopsocus unipunctatus – Stenopsocus lachlani, the 8th AVZ is identical but with different species dominance.

Keywords: Psocoptera; taxocenoses; diversity; forest ecosystems; altitudinal vegetation zones; Fageti-Piceeta s. lat.; Piceeta s. lat.; Moravskoslezské Beskydy Mts.; Oravské Beskydy Mts.; Western Carpathian Mts.

In general, psocids are a rarely studied insect cenoses composition dependence on vegetation order. Thanks to their size, quiet coloration and tier in the Mazák Nature Reserve, located in the relatively difficult way of collecting and prepara- Moravskoslezské Beskydy Mts. (Holuša 2003b). tion, they are at the edge of entomologists’ inter- In the Podbeskydská pahorkatina Hills was further ests. The psocids were studied only in some areas of evaluated psocid occurrence within the frame of the Czech Republic, mostly in various mountains forest type complexes in the Nature Reserve Ka- of Moravia and Silesia – the Hrubý Jeseník Mts., menec (Holuša 2005). Moreover, Mückstein and Králický Sněžník Mt. (Obr 1949) and the Moravsko- Holuša (2003) studied the composition of psocid slezské Beskydy Mts. (Obr 1952, 1965). Only oc- taxocenoses in different ecosystem types and its casional captures are published from other areas. A dependence on naturalness level of forest ecosys- complex psocopterological research was initiated tems in the region of the Žďárské vrchy Hills. in a territory of the Czech Republic and Slovakia The aim of the systematic study of psocids, con- in year 1997. ducted in the Western Carpathian Mts. in years Only faunistic data are mostly known from our 1997–2001, was to define species diversity and country at present, however, Holuša (2001) also characteristic species composition of psocids in studied an ecological problem of psocid taxo- particular vegetation zones and to prove an ap-

J. FOR. SCI., 53, 2007 (Special Issue): 3–10  plicability of vegetation zones or lower units of in a small test tube with 70% alcohol. All samples geobiocenological or forest typological systems in were collected and determined by author. The evi- zoocenological studies. dence material is deposited in 70% alcohol in the “Mountain spruce forest” is a commonly used author’s collection. Articles by Günther (1974) term, but its definition is usually not very clear and and Lienhard (1998) were used for determination; well understood. It is possible to use one of the nomenclature, zoogeographical distribution and vegetation classification systems – the geobioceno- ecological demands pursuant to Lienhard (1977, logical system (Zlatník 1959, 1976; Buček, Laci- 1998). na 1999) or the forest typological system (Plíva Samples were sorted into vectors, which repre- 1971, 1991) to specify it. “Mountain spruce forest” sent “habitats of psocids”. Following factors were is analogous to the 6th and 7th altitudinal vegetation taken into account for the purpose of material sort- zone according to the geobiocenological system ing: biogeographical region, ecological conditions and according to the forest typological system it (according to the forest type complexes) and tree or corresponds with the 7th and 8th AVZ (cf. Holuša shrub species, from which was material obtained 2003a). (samples were also distinguished according to the capture method; captured either in the herb layer or Methods by the Malaise trap). For example: BE5Ssm, where BE denotes the Beskydský biogeographical region A net of equally distributed geobiocenological (No. 3.10), 5S represent forest type complexes 5S research plots was situated in regions of eastern (i.e. Abieto-Fagetum mesotroficum) and sm is an ac- Moravia, eastern Silesia and northern Slovakia in ronym for the tree species Picea abies. the territory of Polonic and Westcarpathian bio- Diversity was evaluated by Shannon-Wiener (HS) geographical subprovinces (i.e. in the region of the and Brillouin diversity index (HB). Both indexes, Western Carpathians). Plots were selected in all al- Shannon-Wiener and Brillouin, were computed titudinal vegetation zones occurring in this region, according to Kaesler and Mulvany (1976a,b). i.e. from the 3rd (communities of Querci-Fageta s. Diversity indexes of individual habitats were calcu- lat.) to the 9th (communities of Pineta mugi s. lat.). lated from a total number of captured specimens, Plots were placed in such parts of forest stands, however, in case of a higher number of specimens which represent a particular altitudinal vegeta- these were reduced to a constant number (30, 60, tion zone and in which it was possible to collect 120 and 240) (Table 1). Some material was exclud- representative material of psocids. Approximately ed from statistical processing because of a small the same number of permanent plots was placed number of collected specimens in some plots (i.e. in all altitudinal vegetation zones. Permanent plots species in a lower number than 5 specimens or were marked out in the best-preserved parts of na- 2 species even less than 3 specimens) to prevent ture reserves and additional plots were selected in data distortion. modified parts of nature reserves or in managed forests. Detrended Correspondence Analysis – DCA Sampling was carried out in the same way in all AVZ during the research and material from the Detrended Correspondence Analysis (DCA), ac- 7th (i.e. Fageti-Piceeta s. lat.) and the 8th (i.e. Piceeta cording to Gauch (1982), Hill (1974) and Hill s. lat.) AVZ is presented in this study. The research and Gauch (1980), proceeds from the method was conducted in years 1997–2001. of Principal Component Analysis (PCA) used for Material was obtained from the permanent sam- non-linear data. In the DCA-analysis, axes were pling sites during the vegetation period (from the adjusted in order to prevent criteria deformation by beginning of May up to the middle of September). the axis ends. The unit length of axes corresponds Samples were collected by sweeping with a sweep with average species dispersion. This unit remains net of 50 cm mouth in diameter. Branches of trees without change in various parts of axes. The DCA and bushes were beaten with the same sweep net in ordination method has a quite heuristic character. the extent of about 1 m from the branch end and up Interpretation of axes and ordination positions of to approximately 2.5 m height. These methods were particular species is based on their ecology with a also complemented by an individual collecting of view to habitat characteristics. Modified SW Dec- adults. During sweeping and beating, 30 sweepings orana was used to process the DCA analysis, which or beatings were carried out in each locality. Caught was adapted for zoocenological data processing psocids were sucked into the exhauster and stored (Povolný, Znojil 1990).

 J. FOR. SCI., 53, 2007 (Special Issue): 3–10 B E 0.593 0.657 0.574 B H 1.116 1.303 1.200 240 S E 0.587 0.652 0.570 S H 1.069 1.250 1.148 s. lat. B – equatibility B E 0.744 0.613 0.608 0.240 0.663 0.642 Piceeta B H 1.574 1.132 1.159 0.370 1.258 1.191 120 S s. lat. and E 0.739 0.602 0.600 0.225 0.655 0.635 S H 1.469 1.053 1.080 0.331 1.174 1.114 Fageti-Piceeta B – Brillouin index of diversity, E H B 0.784 0.631 0.644 0.281 0.722 0.675 0.666 S E 1.523 1.109 1.002 0.312 1.346 1.155 1.245 60 S H 0.776 0.616 0.630 0.259 0.711 0.664 0.656 – equatibility, H S B E 1.365 0.988 0.902 0.266 1.204 1.038 1.113 B H 0.784 0.650 0.676 0.361 0.713 0.738 0.710 B E 1.481 1.009 1.005 0.311 1.206 1.101 1.198 30 S E 0.774 0.628 0.657 0.330 0.696 0.726 0.696 S H 1.237 0.845 0.847 0.255 1.010 0.941 1.009 – Shannon-Wiener index of diversity, E S B – E 0.687 0.761 0.323 0.811 1.000 0.575 0.476 0.516 0.896 0.627 0.207 0.865 0.627 0.916 0.818 0.549 0.928 0.548 0.625 B – H 1.582 0.836 0.224 0.562 1.099 1.118 0.660 1.133 1.242 1.010 0.372 0.950 1.303 1.270 0.898 1.206 1.494 0.602 1.217 c N S – E 0.684 0.743 0.281 0.774 1.000 0.570 0.433 0.512 0.922 0.596 0.200 0.881 0.624 0.913 0.810 0.545 0.952 0.512 0.618 S – H 1.504 0.700 0.167 0.347 0.597 1.083 0.489 1.095 0.798 0.788 0.349 0.599 1.272 1.062 0.711 1.168 1.015 0.464 1.143 7 4 3 6 5 8 N 21 17 17 20 21 13 16 199 369 357 310 500 393 143 1 3 2 2 3 7 4 9 4 5 6 3 8 4 3 9 5 3 7 10 Nsp Table 1. Values of indexes of diversity and equitability particular for of indexes psocid biotopes vegetation altitudinal zones 1. Values in the of Table Biotope BE7Fbk BE7Fsm BE7Sbk BE7Sjiv BE7Sjr BE7Spod BE7Ssm BE7Zbk BE7Zsm BE8Zbk BE8Zkos BE8Zma BE8Zpod BE8Zsm OR7Ssm OR8Sjr OR8Ssm OR8Zjan OR8Zkos OR8Zsm Nsp – number of species, N – number of specimens, H Indexesof diversity for individual habitats from total number of captured specimens (N), in case of greater number 60,of specimens 120 and 240 were reduced for constant number of specimens – 30,

J. FOR. SCI., 53, 2007 (Special Issue): 3–10  Divisive Cluster Analysis – DvClA column and on the right is one algorithm division of species spectrums in groups. The main field rep- Divisive Cluster Analysis (DvClA) represents resents the semiquantitative relative frequency of a method of hierarchic divisive classification particular species in groups corresponding with (Gower 1967; Orlóci 1976). The ordination of their biotopes. Explanations: – species does not groups is performed twice by “Reciprocal averag- occur, 1 – rare species, 2 – very scarce, 3 – scarce, ing” (RA). All vectors are projected into the main 4 – common, 5 – very common to subdominant, axis as a super-ellipsoid. In the second phase, par- 6 – dominant. Groups of psocid species and groups tial complexes of vectors are divided according to of habitats were organized to increase their clear- species ordinate in particular vectors and accord- ness so that there is an evident species transfer ing to abundance of particular species (indicators) within biotopes in the diagonal direction from the as well. These indicators are automatically selected left upper corner to the right lower corner. by the program in compliance with species spec- Acronyms of trees and shrubs (investigated tree trum of particular vectors (habitats) to end parts of species): sm – Picea abies, bk – Fagus sylvatica, kos ordination axis. Used modification – Twinspan al- – Pinus mugo, jan – Juniperus communis nana, jiv gorithm comes from a gradual division of habitats – Salix caprea, jr – Sorbus aucuparia; pod – copse, and species. Every processed file is ordinated by RA ma – Malaise trap. method, whereupon characteristic species (or bio- Next psocid communities were classified in the topes) are associated with axes ends. Central parts following study plots: 7F – Fageto-Piceetum aci- of axes are ordinated consequently. On the base of dophilum; 7S – Fageto-Piceetum mesotrophicum; acquired results, it is searched for species combi- 7Z – Fageto-Piceetum humile; 8S – Piceetum meso- nations, which are characteristic for parts of ordi- trophicum; 8Z – Sorbeto-Piceetum. nation axes and can be used as appropriate “tools for cuts” (Hill 1974). This method was modified results and discussion for the purpose of this study, because the first ver- sion is defined for phytocenological studies only. 2,461 adults comprising 16 species were found Column heads represent abbreviations of biotopes. in total: 12 species (eudominant species Caecilius Numbers in columns below indicate the division despaxi, Mesopsocus unipunctatus, dominant spe- of appropriate algorithm (every habitat is divided, cies Stenopsocus lachlani, Amphigerontia bifasciata marked 0 or 1). There are species names in the left a Caecilius burmeisteri) were found in the 7th AVZ

350 3VS 2VS 300 4VS 5VS 250 6VSȱBE 7VSȱBE 200 8VSȱBE 9VSȱOR 150 8VSȱOR

100 7VSȱOR 6VSȱOR

50

0 0 50 100 150 200 250 300 350 400 450 500

Fig. 1. DCA analysis of psocid biotopes (axis x – gradient of altitudinal vegetation zones, q – gradient of hydricity)

 J. FOR. SCI., 53, 2007 (Special Issue): 3–10 and an equal number of species was found in the A-I-b (not illustrated in Fig. 2) and habitats with 8th AVZ (eudominant species Caecilius despaxi, Picea abies and Salix caprea occur in group B-II-b-1, Stenopsocus lachlani). Species spectrum and domi- i.e. the 5th–9th AVZ group. nancy found in the 7th and 8th AVZ in the Moravsko- In the DCA-analysis, habitats of the 7th AVZ cre- slezské Beskydy Mts. differ from those in the ate a field, which is located on the left side of the Oravské Beskydy Mts. mainly by representation of whole dotted field (along x-axis). It forms the high- Mesopsocus unipunctatus. est AVZ together with fields of the 8th and 9th AVZ. Resulting from the comparison of tree coloniza- Only single habitats of the 7th AVZ occur in the tion, Picea abies was the most colonized tree species field of the 4th and 5th AVZ. in community 7F and 8S. There were found higher From the view of hydricity (q-axis), habitats of values of diversity indexes in the communities 7F the 7th AVZ are on the same level as those of the and 8Z (Table 1) and the highest value was calcu- 4th–6th AVZ. lated for Picea abies in forest type complex 7F. Diversity indexes HS reach values from 0.17 to The DCA-analysis might be interpreted as fol- 1.50, HB 0.22–1.59. The highest values were cal- lows, the x-axis denotes an influence of altitudinal culated for habitat BE7Fsm with reduced number vegetation zones and q-axis refers to an influence of N30 HS 1.24 and HB 1.48, higher values also showed hydricity. These factors might raise a presumption habitat BE7Ssm with reduced number N30 HS 0.85 of mutual correlation, but all AVZ included habi- and HB 1.01. tats with high hydricity – flooded habitats, water Characteristic species composition of the 7th AVZ logging and peaty habitats as well as dry or desic- was defined:Caecilius despaxi – Amphigerontia bi- cate habitats. Because every AVZ comprehends a fasciata – Mesopsocus unipunctatus – Stenopsocus large scale of habitats – from dry to peaty habitats, lachlani. These species, occurring in the 7th AVZ, hydricity of habitat does not correlate with altitude are missing in the lower and middle altitudinal veg- within collected material. Habitats of the 7th AVZ etation zones. are situated “higher” than habitats of the 8th AVZ in the graph of x-q axis (Fig. 1) and thus it is pos- Taxocenosis of the 8th (Piceeta s. lat.) altitudinal sible to state that biotopes of the 8th AVZ are more vegetation zone “moist”. A field of habitats of the 7th AVZ is situ- ated along the x-axis, i.e. along altitudinal vegeta- Eudominant species Caecilius despaxi, Stenopso- tion zones. The difference is then in the hydricity cus lachlani were found on the base of total domi- of habitats of the Oravské and Moravskoslezské nancy in the 8th AVZ. In the natural communities Beskydy Mts. habitats of the 7th and 8th AVZ in the were identified Stenopsocus lachlani and Caecilius Moravskoslezské Beskydy Mts. create a homog- despaxi as eudominant and Caecilius burmeisteri enous dotted field situated “higher” than a habitat as dominant species. The most diverse species field of the Oravské Beskydy Mts. spectrum with the highest abundance was on Picea abies. Other tree species are colonized by a higher Taxocenosis of the 7th (Fageti-Piceeta s. lat.) number of psocid species as well, however, in lower altitudinal vegetation zone abundances. In the DvClA-analysis, habitats of the 8th AVZ Eudominant species Caecilius despaxi, Mesopso- create group B-II-b-1, only individually they oc- cus unipunctatus and dominant species Stenopso- cur in group B-II-a. Group B-II-b covers biotopes cus lachlani, Amphigerontia bifasciata, Caecilius of the 5th–9th AVZ and group B-II-a biotopes of burmeisteri were found on the base of total domi- the 4th–8th AVZ where only several habitats (Pinus nancy in the 7th AVZ. In the natural communities, mugo, Juniperus communis nana) come under. In Caecilius despaxi, Mesopsocus unipunctatus were the DCA-analysis, habitats of the 8th AVZ lie along eudominant and as dominant species were identi- the x-axis on the left side. This dotted field is not fied Caecilius burmeisteri, Amphigerontia bifascia- situated along the x-axis in the same way as the ta and Stenopsocus lachlani. Picea abies was the field of the 7th AVZ because the field of the 8th AVZ most abundantly colonized tree species, whereas shows higher moisture according to the gradient of Fagus sylvatica was colonized by a poorer species the q-axis. spectrum (max. 4). Diversity indexes HS reach values 0.35–1.27, HB In the DvClA-analysis, habitats of the 7th AVZ 0.37–1.49. The highest values were found within occur in two groups. Habitats of broad-leaf trees habitats BE8Zsm with reduced number N30 HS 1.01 (Fagus sylvatica, Sorbus aucuparia) form groups and HB 1.21, similarly high values of indexes were

J. FOR. SCI., 53, 2007 (Special Issue): 3–10 

0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 1 0 1 0 00 0 1 0 00 0 1 0 00 1 1 0 10 0 1 0 10 0 1 0 10 1 1 0 10 1 1 1 00 0 1 1 00 1 1 1 00 1 1 1 1 1 1 0 01 0 1 0 01 0 1 0 01 1 1 0 0 01 1 1 0 0 01 1 1 1 0 01 1 1 1 0 01 1 1 1 0 01 1 1 1 0 1 1 1 0 1 1 1 1 0 1 1 1 11 0 1 0 1 11 0 1 1 1 11 0 1 1 1 11 1 1 1 11 1 0 1 01 0 0 1 01 0 0 1 01 1 0 1 11 0 0 1 11 0 0 1 11 1 0 0 0 1 0 0 11 0 0 0 0 11 0 0 1 0 11 0 0 1 0 1 1

OR9Zsm

OR9Zkos OR9Zjan

B-II-b-2

OR9Ksm

OR9Kkos

OR8Zsm

OR8Ssm

OR8Sjr

OR7Ssm

BE8Zsm

BE8Zma

BE8Zkos

BE8Zbk

BE7Zsm

BE7Ssm

BE7Sjiv

BE7Fsm BE6Zsm

B-II-b-1 BE6Ssm

BE6Sjd

BE6Fsm

BE5Sjd

VS4Sjd

VS4Djd BE7Fbk

231234515145--151-12-3 -- 1 BE6Gsm

----322513-43-114--45-6----3226253661366226214 12 5 -- 2

BE5Fsm

BE5Fjd

OR6Bsm

OR8Zkos

BE8Zpod

OR5Sjd

BE7Spod

BE6Spod B-II-a-2

VS5Bjd

VS5Asm

VS3Ltrn

OR8Zjan

BE4Ssm PB4Dsm

------11-3 ---1-11111513-451145226525 13 3 BE6Sbk

B-II-a-1 BE5Sbk

11100000 001 01111110000000000000000000 VS4Emd

------1---11-----1--1------1 VS4Dsm

------11------1------1------111------11--- 2233----1-3-3-22---42-11-1 1- - -- - 1- 2 PB3Hsm

AVZ areAVZ marked with red colour (with regard to the table extent, right third of the whole graph, i.e.

PB3Hma th

PB4Bsm

BE6Pbk BE5Nsm

and 8 BE5Bsm

th ------11--1------1-51--11-141-1 -1 1 BE5Bjd

----1-1 ------1----1------1------

BE6Osm

BE5Ssm

BE5Lsm B-I-b

BE6Rsm BE6Psm

3 3 4 4 2 2 3 1 2 - 1 364431-1-1-- --11--1------11 134 VS4Bsm

VS5Bsm

PB2Lsm BE5Hsm

11154632411-1213 1212453511--1164 1--1

OD1Lsm

OR6Bjd

BK4Bsm

BK3Bsm

BE6Ppod

BE6Pjd

PB4Bma BE6Ojiv

B-I-a

VS4Bjd

VS3Lsm

OP3Hsm BE4Bjd

111111000000000011000001 0000111110000011 00010011101111 0011111111 00011 00001111111111111111111111 10 0 VS4Bbo

--1-1------1------112------1------1------2------11-1----1------2--6 -112------1131--42111---1-1 ----1------1-111-----1 -----1------1 -1------1------1----1------1- -1-4---2------11 ---1------31-2141-1------5355361123-1--21 ------1-11------431----1------22------11-2-312111-211------3--1111--11 ------11--1------6 ------1------1-----1 ------2------1-----1 ------1------2------1 ------1------14-1------2 ------1------111---11 331555121-1-1262 ------2------1-----11---1--11111------1------12-211111323 -1-4 ------1---24--1-----2------2------12---1------1------1------1--1------2 ------1------211------3 ------2------1------1------2246662613543443 ------1------1------11----1---1------251--1------1 ------1-1------11-1---1------1-141--2------1 ------1123321222 ------21-1 ------2---1------11111 -- 1------11 ------1------2-11--1--1 334 ------1------1------1-3-1111-3---1-2 1------1 - 54 1 5 -- - 1------1 ---1------1------1------1 ------1-3----1--1------1-1-1--21221------1------311422-4-2-111111---1--1---11 ------2------11111111111110000000000000 534646352-211---0000000000000 -----0111111111111 ------1------1111111111111111 - 0000000000000000 1111111111111111 1111111110000000 ----- 1111111111 1111111111 0000000000 ------0000000000 -- - 11111 11111 00000 -- 11111 11111111111111111111111111 - 11111111111111111111111111 11111111111111111111111111 00000000000000000000000000 -- - 11 11 1 11 1 11 1 1 Biotopes Groups of biotopes Elipsocus moebiusi Mesopsocus laticeps Stenopsocus immaculatus Stenopsocus stigmaticus Caecilius flavidus Elipsocus pumilis Amphigerontia contaminata Caecilius gynapterus Lachesilla quercus Ectopsocus meridionalis Graphopsocus cruciatus Psococerastis gibbosa Peripsocus phaeopterus Philotarsus parviceps Peripsocus subfasciatus Caecilius atricornis Peripsocus parvulus Trichadenotecnum majus Blaste quadrimaculata Kolbia quisquiliarum Epipsocus lucifugus Liposcelis corrodens Elipsocus abdominalis Enderleinella obsoleta Hemineura dispar Reuterella helvimaculata Caecilius piceus Peripsocus didymus Peripsocus alboguttatus Metylophorus nebulosus Loensia fasciata Loensia variegata Caecilius fuscopterus Elipsocus hyalinus Stenopsocus lachlani Trogium pulsatorium Caecilius burmeisteri Loensia pearmani Philotarsus picicornis Cuneopalpus cyanops Amphigerontia bifasciata Trichadenotecnum sexpunctatum Lachesilla pedicularia Mesopsocus unipunctatus Caecilius despaxi Fig. 2. Results of DvClA-analysis – Twinspan algorithm; biotopes of the 7 subgroup B, is illustrated only)

 J. FOR. SCI., 53, 2007 (Special Issue): 3–10 found within habitat OR8Zsm with reduced num- frame for animal studies. According to results, pso- ber N30 HS 1.01 and HB 1.20. cid taxocenoses are dependent on the main eco- Characteristic species composition of the 8th AVZ logical factors of environment, therefore AVZ are is identical with the 7th AVZ: Caecilius despaxi the most appropriate units considering changes of – Amphigerontia bifasciata – Mesopsocus unipunc- the main ecological factors in landscape segments. tatus – Stenopsocus lachlani. However, it differs in This study also confirmed that AVZ are the main dominancy of Caecilius despaxi (lower) and Lache- factor with the greatest influence on variability of silla pedicularia is more abundant. psocid taxocenoses. Finally, the order of psocids can serve as a suitable tool for the geobiocenologi- Conclusion cal classification of ecosystems.

Compositions of psocid taxocenoses are influ- References enced by tree species composition in “mountain spruce forests” that correspond with the 7th and Buček A., Lacina J., 1999. Geobiocenologie II. [Skripta.] 8th AVZ. It is mainly valid for the 7th AVZ, where Brno, MZLU, LDF: 240. Fagus sylvatica is still edificator (it means subdom- Gauch H.G. Jr., 1982. Nose reduction by eigenvector ordina- inat tree). This influence is not important in the tions. Ecology, 63: 1643–1649. 8th AVZ because Fagus sylvatica occurs only indi- Gower J.C., 1967. A comparsion of some methods of cluster vidually here and in the stage of low tree or shrub. analysis. Biometrics, 23: 623–637. There are no significant differences in taxoce- Günther K.K., 1974. Die Tierwelt Deutschlands. 61. Teil. noses of the 7th and 8th AVZ, although the species Staubläse, Psocoptera. Jena, VEB Gustav Fischer: 314. spectrums are not identical. The taxocenoses differ Hill M.O., 1974. Correspondence analysis – a neglected in dominances, but characteristic species combina- multivariate method. Applied Statistics, 23: 340–354. tions of psocids are the same. This result supports Hill M.O., Gauch H.G. Jr., 1980. Detrended correspond- a correct classification of the 7th AVZ as “spruce ence analysis: an improved ordination technique. Vegetatio, forests”. 42: 47–58. It is possible to say that altitudinal vegetation Holuša O., 2001. Příspěvek k poznání fauny pisivek (In- zones proved to be a suitable frame for the defi- secta: Psocoptera) Přírodní rezervace Smrk (Beskydský nition of “mountain spruce forest” as well as for bioregion, Česká republika). Práce a studie Muzea Beskyd, zoocenological studies. AVZ and lower units of 11: 83–97. geobiocenological, respectively forest typological Holuša O., 2003a. Vegetační stupňovitost a její bioindikace system, together with description of tree species pomocí řádu pisivek (Insecta: Psocoptera). [Dizertační composition and naturalness level form a perfect práce.] Brno, MZLU, LDF: 258. base for studies focused on the animal taxoceno- Holuša O., 2003b. Fauna pisivek (Insecta: Psocoptera) ses structure. Furthermore, they might be a perfect Národní přírodní rezervace Mazák (Beskydský bioregion, tool for evaluation of changes in forest ecosystems Česká republika). Práce a studie Muzea Beskyd (Přírodní in the future. We confirmed the hypothesis that vědy), 13: 83–98. psocids, as a part of forest ecosystem, fully com- Holuša O., 2005. Fauna pisivek (Insecta: Psocoptera) ply with the theorem of geobiocenoses (Zlatník Přírodní památky Kamenec v Podbeskydské pahorkatině 1976). Geobiocenoses are composed of specific bio- (Podbeskydský bioregion, Česká republika). Práce a studie cenoses in conjunction with abiotic environment; Muzea Beskyd (Přírodní vědy), 15: 75–89. the biocenose is formed not only by plants or trees Kaesler R.L., Mulvany P.S., 1976a. Fortran IV program as the main community determinants, but an im- to compute diversity indices from information theory. portant part constitutes the zoocenose as well. On Computer & Geosciences, 2: 509–514. the basis of long-standing studies of “forest pests”, Kaesler R.L., Mulvany P.S., 1976b. Fortran IV program Stolina (1975) considers the geobiocenological to compute replicated diversity indices for random samples units, AVZ and groups of forest types, as suitable of specified size. Computer & Geosciences, 2: 515–519. frames for autecological studies of species. These Lienhard C., 1977. Die Psocopteren des Schweizerischen studies can consequently serve as determinants of Nationalparks und seiner Umgebung (Insecta: Psocoptera). habitat specifications (i.e. occurrence, localities of Ergebnisse der wissenschaftlichen Untersuchungen im occurrence, survival ability). Schweizerischen Nationalpark, Band 14, Nr. 75: 417–551. Altitudinal vegetation zones are units, which Lienhard C., 1998. Psocoptères Euro-méditerranéens. complexly conjugate ecological factors of ecosys- Faune de France, Vol. 83. Paris, Fédération Française des tems in landscape segments and they are a perfect Sociétés de Sciences Naturelles: 517.

J. FOR. SCI., 53, 2007 (Special Issue): 3–10  Mückstein P., Holuša O., 2003. Composition of psocid Plíva K., 1991. Funkčně integrované lesní hospodářství. taxocenoses (Insecta: Psocoptera) in dependence of level of 1 – Přírodní podmínky v lesním plánování. Brandýs nad naturalness of forest ecosystems in the Žďárské vrchy hills. Labem, ÚHÚL: 263. Journal of Forest Science, 49: 208–219. Povolný D., Znojil V., 1990. Vergleich zwischen Sar- Obr S., 1949. Pisivky ze Slezských Jeseníků a Králického cophagini-Taxozönosen (Insecta, Diptera) Thürigens und Sněžníku. Přírodovědný sborník Ostravského kraje, 10: der Tschechoslowakei. Rudolstädter Naturhistorische 219–234. Schriften, 3: 43–61. Obr S., 1952. Pisivky ze Slezských Beskyd. Přírodovědný STOLINA M., 1975. Geobiocenologické jednotky v štúdiu les- sborník Ostravského kraje, 13: 216–231. ného fytofágneho hmyzu. Lesnícky časopis, 74: 307–322. Obr S., 1965. Pisivky Moravských Beskyd. Spisy Přírodově- Zlatník A., 1959. Přehled slovenských lesů podle skupin decké fakulty UJEP Brno, Serie M 21, L 24, 1965/2 (č. 460): lesních typů. Spisy Vědecké laboratoře biocenologie a ty- 51–80. pologie lesa Lesnické fakulty VŠZ v Brně, č. 3: 1–159. Orlóci R., 1976. Multivariate Analysis in Vegetation Re- Zlatník A., 1976. Přehled skupin typů geobiocénů původně search. The Hague, Boston, Dr. W. Junk Publishing: 451. lesních a křovitých v ČSSR. Brno, Zprávy Geografického Plíva K., 1971. Typologický systém ÚHÚL. Brandýs nad ústavu ČSAV, 13: 55–65. Labem, ÚHÚL: 90.

Složení taxocenóz pisivek (Insecta: Psocoptera) v lesních ekosystémech bukových smrčin (Fageti-Piceeta s. lat.) a smrčin (Piceeta s. lat.) v západních Karpatech

AbstraKt: Během období 1997–2001 byly v lesních ekosystémech v oblasti západních Karpat studovány taxocenó- zy pisivek (Psocoptera). Jako rámce pro studium byly použity vegetační stupně podle systému geobiocenologie, resp. lesnické typologie (Plíva 1971, 1991). Pro klasifikaci stanoviště (tj. ekologických podmínek) byly použity jednotky – soubory lesních typů lesnicko-typologického systému. V práci pod pojmem “horské smrkové lesy – přirozené smrčiny” jsou chápány vegetační stupně (VS): 7. (tj. buko-smrkový) – společenstva Fageti-Piceeta s. lat. a 8. (tj. smrkový) – společenstva Piceeta s. lat. Tyto vegetační stupně se vyskytují v oblasti Moravskoslezských Beskyd na území České republiky a v oblasti Oravských Beskyd na Slovensku. Celkově byly zjištěno 2 461 imag v 16 druzích: v 7. VS bylo zjištěno 12 species (eudominantní druhy Caecilius despaxi, Mesopsocus unipunctatus, dominantní dru- hy Stenopsocus lachlani, Amphigerontia bifasciata a Caecilius burmeisteri), stejný počet druhů byl zjištěn v 8. VS (eudominantní druhy Caecilius despaxi, Stenopsocus lachlani). Taxocenózy pisivek byly vyhodnoceny statistickými metodami – detrendovanou korespondenční analýzou (DCA) a shlukovou divizní analýzou (DvClA). Materiál byl vyhodnocen v rámci širšího srovnání materiálu pocházejícího i z ostatních VS v rámci vnějších západních Karpat. Charakteristické druhové kombinace pisivek pro jednotlivé VS byly zjištěny: pro 7. VS – Caecilius despaxi – Amphi- gerontia bifasciata – Mesopsocus unipunctatus – Stenopsocus lachlani, pro 8. VS je charakteristická druhová kom- binace identická s rozdílem v druhové dominanci.

Klíčová slova: Psocoptera; taxocenózy; diverzita; lesní geobiocenózy; vegetační stupně; Fageti-Piceeta s. lat.; Piceeta s. lat.; Moravskoslezské Beskydy; Oravské Beskydy; západní Karpaty

Corresponding author:

Ing. Otakar Holuša, Ph.D., Mendelova zemědělská a lesnická univerzita v Brně, Lesnická a dřevařská fakulta, Lesnická 37, 613 00 Brno, Česká republika tel.: + 420 606 960 769, fax: + 420 555 559 865, e-mail: [email protected]

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