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Home , Amphigerontia bifasciata, Caecilius fuscopterus, Elipsocus, Elipsocus abdominalis, Elipsocus hyalinus, Elipsocus pumilis

ACTA UNIVERSITATIS AGRICULTURAE ET SILVICULTURAE MENDELIANAE BRUNENSIS

Volume LX 6 Number 3, 2012

STRUCTURE AND DIVERSITY OF PSOCID TAXOCENOSES (INSECTA: ) IN THE FOREST ECOSYSTEMS OF THE FAGETA ABIETIS S. LAT. ZONE IN THE WESTERN CARPATHIAN MTS.

O. Holuša

Received: December 5, 2011

Abstract

HOLUŠA, O.: Structure and diversity of psocid taxocenoses (Insecta: Psocoptera) in the forest ecosystems of the Fageta abietis s. lat. zone in the Western Carpathian Mts. Acta univ. agric. et silvic. Mendel. Brun., 2012, LX, No. 3, pp. 51–60

Structure of psocid taxocenoses (Psocoptera) were studied in forest ecosystems of the Western Carpathian Mts. during 1997–2001. Vegetation tiers were used as a study frame. Lower units of forest site classifi cation system (forest type complexes) were used for a classifi cation of ecological conditions as well. Only a part of material, i.e. individuals that was found in the forest ecosystems of Fageta abietis s. lat. communities (= the 4th vegetation tier) was evaluated for purpose of this work. This vegetation tier is widespread in large part of Vsetínské vrchy hills, in the highest parts of Podbeskydská pahorkatina hills, and in the foothills of the Moravskoslezské Beskydy Mts. 1774 adults comprising 32 species were found in total in the 4th vegetation tier. As eudominant species, the following ones were found: fl avidus and C. burmeisteri, as dominant species, the following ones were found: Philotarsus picicornis, moebiusi and subfasciatus. In natural geobiocenoses with the level of naturalness of 1 or 2, the following species were found: as eudominant species: Caecilius fl avidus and Elipsocus moebiusi, as dominant species, the following one was found: unipunctatum. Taxocenoses of psocids were evaluated by Detrended Correspondence analysis (DCA) and Divisive Cluster analysis (DvClA). This material was compared to another material gained from various vegetation tiers in the Western Carpathians Mts. The characteristic species composition of psocids in the 4th vegetation tier was as follows: Caecilius fl avidus – Elipsocus moebiusi – Peripsocus parvulus.

Psocoptera, taxocenoses, diversity, forest ecosystems, vegetation tier – Fageta abietis s. lat., the Moravskoslezské Beskydy Mts., the Western Carpathian Mts.

Order Psocoptera has not been in focus of faunistic composition dependence on VTs in the or ecological studies. Complex psocopterological Moravskoslezské Beskydy Mts., in the Podbeskydská research was initiated recently by author in pahorkatina hills (HOLUŠA, 2005) and in the alluvium a territory of the Czech Republic and Slovakia in of the Odra river (the Protected landscape area of 1997. The aim of the systematic study of psocids Poodří) (HOLUŠA, 2007a). conducted in the Western Carpathian Mts. was to The aim of presented article is to analyse defi ne the species diversity and characteristic species composition and diversity of psocid taxocenoses of composition of psocids in particular vegetation the 4th VT (Fageta abietis s. lat.) i.e. and to discuss the tiers (= altitudinal vegetation zone, further only VT) diff erences with other known VTs i.e. the 5th (Abieti– and to prove an applicability of vegetation zones fageta s. lat.) (HOLUŠA, 2009), the 6th (Picei–fageta s. lat.) or lower units of geobiocenological or forest + site (HOLUŠA, 2011), the 7th (Fageti–piceeta s. lat.) and the 8th classifi cation systems in zoocenological studies. (Piceeta s. lat.) VTs (HOLUŠA, 2007c). HOLUŠA (2001, 2003a, b, 2007b, c) published The aim of this study is the using of frames and studies at ecological problem of psocid taxocenoses units of forest site classifi cation system (according

51 52 O. Holuša

to PLÍVA, 1971, 1991; HOLUŠA & HOLUŠA, 2008, 2010, region of the Podbeskydský region (cf. CULEK, 1996), 2011) for zoocenological studies with a view to 4B represents forest type complexes 4B (i.e. Fagetum processing zoocoenoses characteristics for forest eutrophicum) and bk is an acronym for the species site units. Fagus sylvatica. Material was collected at all localities in the same METHODS way, therefore dominance is calculated as the sum of all the methods for individual habitats. Malaise trap The net of equally distributed geobiocenological was evaluated as a separate habitat due to diff erences research plots was situated in regions of eastern in obtaining material. Individual species are Moravia, eastern Silesia and northern Slovakia classifi ed into classes according to the dominance of in the territory of Polonic and Westcarpathian TISCHLER (1949). Dominance was fi rst calculated the biogeographical subprovinces (i.e. in the region of total for all geobiocenosis with varying degrees of the Western Carpathian Mts.). Plots were selected nature, and then only for geobiocenosis natural (i.e. in all VTs occurring in this region, i.e. from the the degree of naturalness 1–2 according to ELLENBERG rd th 3 (communities of Querci–fageta s. lat.) to the 9 1973, 1978)). (communities of Pineta mugi s. lat.) (cf. HOLUŠA, Diversity was evaluated by Shannon-Wiener (H ) 2003a). Plots were placed in such parts of forest S and Brillouin diversity index (HB). Both indexes, stands which represent a particular VT and in which Shannon-Wiener and Brillouin, were computed it was possible to collect a representative material according to KAESLER & MULVANY (1976a, b). Diversity of psocids. Approximately the same number of indexes of individual habitats were calculated from permanent plots was placed in all VTs. Permanent a total number of captured specimens (Tab. 1). Some plots were marked out in the best-preserved parts material was excluded from statistical processing of nature reserves (with the level of naturalness because of a small number of collected specimens 1 or 2 according to ELLENBERG (1973, 1978) and in some plots (i.e. species in a lower number than 5 additional plots were selected in modifi ed parts of specimens or 2 species even less than 3 specimens) nature reserves or in managed forests with the level to prevent a data distortion. of naturalness 3 or 4 according to ELLENBERG (1973, 1978). Detrended Correspondence Analysis – DCA The material was obtained from permanent Detrended Correspondence Analysis (DCA), sampling sites during the vegetation period (from according to GAUCH (1982), HILL (1974) and HILL & the beginning of May to the middle of September). GAUCH (1980), proceeds from the method of Principal The samples were collected by sweeping with Component Analysis (PCA), used for non-linear a sweep net of 50 cm mouth in diameter. Branches of data. Axes were adjusted in order to prevent criteria and bushes were beaten with the same sweep deformation by the axis ends in the DCA-analysis. net in the extent of about 1 m from the branch end The unit length of axes corresponds with the and up to approximately 2.5 m height. Individual average species dispersion. In diff erent parts of axis collection of adults also complemented this method. it remains unchanged. The DCA ordination method During sweeping and beating, 30 sweepings or has a quite heuristic character. Interpretation of beatings were carried out in each locality. At one axes and ordination positions of particular species locality Malaise trap was installed - type of Townes is based on their ecology with a view to habitat (TOWNES, 1972). The trap is used primarily for characteristics. Modifi ed SW Decorana was used to capturing Diptera (CHVÁLA, 1980). The material was process the DCA analysis, which was adapted for collected about 3 times a month. Malaise trap was zoocenological data processing (POVOLNÝ & ZNOJIL, installed by the end of IV. the beginning of X. Caught 1990). psocids were sucked into an exhauster and stored in a small test tube with 70% alcohol. All samples were Divisive Cluster Analysis – DvClA collected and identifi ed by the author. The evidence Divisive Cluster Analysis (DvClA) represents material is deposited in 70% alcohol in the author’s a method of hierarchic divisive classifi cation collection. Papers of GÜNTHER (1974) and LIENHARD (GOWER, 1967; ORLÓCI, 1975). The ordination of (1998) were used for identifi cation; nomenclature, groups is performed twice by “Reciprocal averaging” zoogeographical distribution and ecological (RA). All vectors are projected into the main axis demands were used according to LIENHARD (1977, as a super-ellipsoid. In the second phase, partial 1998). complexes of vectors are divided according to Samples were sorted into vectors “habitats of species ordinate in particular vectors and according psocids”, where the following factors were taken to abundance of particular species (indicators) as into account: biogeographical region, ecological well. These indicators are automatically selected conditions (according to the forest type complexes) by the program in compliance with the species and tree or shrub species, from which the material spectrum of particular vectors (habitats) for the was obtained (samples were also distinguished end parts of ordination axis. Used modifi cation according to the capture method; captured either ‘Twinspan algorithm’ comes from a gradual division in a herb layer or by the Malaise trap). For example: of habitats and species. Every processed fi le is PB4Bbk, where PB denotes the biogeographical ordinated by RA method, whereupon characteristic Structure and diversity of Psocid taxocenoses (Insecta: Psocoptera) in the forest ecosystems of the Fageta abietis s. 53 B E B H N S E S H VS4Bos 4 8 0.727 0.743 1.074 0.774 biotope Nsp N VS4Ddbz 2 6 0.299 0.598 0.451 0.650 B E B H N . lat. S E S ------H biotope Nsp N B E B H N S E S H – Brillouin index of diversity – Brillouin index Values of Indexes of diversity and equitability for particular „habitats of psocids“ in the vegetation tier of Fageta abietis s Fageta tier of psocids“ in the vegetation of particular „habitats for and equitability diversity of Indexes of Values – Shannon-Wiener index of diversity index – Shannon-Wiener – Equitability – Equitability S B S B biotope Nsp N BE4BbkBE4Bdbl 4BE4Bhb 4 13 3 52 0.835 36 0.837 0.775 0.658 1.091 0.628 0.925 0.787 0.632 PB4Bsm 0.667 0.709 PB4Dbk 0.645 9 PB4Ddbl 3 63 1 15 1.216 4 0.356 0.616 0.395 1.388 0.485 0.632 0.442 VS4Bjd VS4Blpm 14 2 36 15 1.802 0.181 0.840 0.309 2.215 0.245 0.839 0.353 BE4BjdBE4Blpm 2BE4Bdbl 4BE4Bsm 2 4 18 1 0.347 0.735 3 0.774 0.646 9 0.366 0.562 0.926 1.000 0.811 0.668 0.637 - PB4Dsm PB4Djd 0.918 3 4 PB4Hsm - 2 16 4 0.464 - 13 0.795 0.512 1.000 0.197 0.602 - 1.386 0.344 0.548 1.000 0.271 VS4Bsm PB4Wsvk VS4Bpod 0.391 22 1 2 VS4Btrn 864 5 22 13 2.038 0.673 0.506 43 2.089 0.882 0.896 0.676 0.617 0.622 0.890 1.025 0.637 PB4AjlhPB4Ajs 2PB4Ajv 1PB4Bbk 4 2PB4Bhb 32 6 0.448PB4Bjd 3 1.000 4PB4Blpm 41 - 0.693 3PB4Bma 0.347 4 24 1.000 0.765 0.774PB4Bpod VS4Bbb 0.484 8 0.444 - 5 0.562 22 2 0.914 0.464 2 0.811 0.680 0.517 0.510 16 0.544 VS4Bbo - VS4Bdbl 1.000 16 0.442 0.495 6 1.350 VS4Bdbz 6 1.055 0.663 3 0.173 0.860 0.960 0.478 0.299 - 3 0.293 1.808 VS4Bhb VS4Bhh 0.598 8 12 0.234 0.869 VS4Bbk 0.451 3 0.337 7 3 VS4Bjb 0.541 1.152 0.650 4 0.621 1.000 0.764 VS4Djd 104 6 7 0.721 1.733 1.000 44 0.192 4 0.657 0.967 1.079 19 0.534 0.182 VS4Emd 0.384 VS4Ejb 0.982 0.699 0.217 0.924 18 0.305 VS4Esvk 3 0.796 0.197 2 0.646 0.462 0.774 0.725 2 VS4Fbk 1.202 0.333 0.680 5 VS4Sjd VS4Dsm 9 0.671 2 0.974 9 VS4Ssm 0.599 3 8 0.702 0.398 0.881 7 0.244 6 0.741 0.950 15 0.454 6 0.530 0.299 0.865 31 0.349 0.764 1.329 0.598 0.682 0.503 0.865 1.304 0.451 0.910 1.807 0.793 0.650 1.011 0.869 1.553 0.921 0.798 E H Nsp – number of species N – number of specimens H I: E 54 O. Holuša

species (or biotopes) are associated with axes ends. tree level. cordata, Tilia platyphyllos, and Central parts of axes are ordinated consequently. On excelsior occur in the co-dominant tree level. Also the base of gained results, it is searched for species Acer platanoides, , Ulmus glabra, combinations, which are characteristic of parts of and Padus avium are co-dominant and overtopped ordination axes and can be used as appropriate tree levels. Tilia cordata, Tilia platyphyllos, and Acer “tools for cuts“ (HILL, 1974). This method was platanoides have thein ecological optimum. The modifi ed for the purpose of this study, because 4th VT occurs in the altitude from 300–640 m a.s.l. the fi rst version was defi ned for phytocenological with the centre of occurrence in the interval of studies only. Column heads represent abbreviations 380–440 m a.s.l., locally up to 680 m a.s.l. Soils of biotopes. Numbers in columns below indicate arose on the fl ysh series of rocks of sandstones and the division of appropriate algorithm (every habitat shales, insular on the deluvial loams. There are is divided, marked 0 or 1). There are species names represented the following soil types: Cambisols in the le column and on the right there is one (typical, eutrophic, sporadically also dystric and algorithm division of species spectrums in groups. pseudogley), are less frequent then podzols (cambic, The main fi eld represents the semiquantitative typical). 4th VT occupies large areas in the Natural relative frequency of particular species in groups forest area (further only NFA) 41, coherent areas in corresponding with their biotopes. Explanations: the highest parts of NFA 39, and some proportions – species does not occur, 1 – rare species, 2 – very of the lowest parts of NFA 40. The most widespread scarce, 3 – scarce, 4 – common, 5 – very common communities in the study area are the following to subdominant, 6 – dominant. Groups of psocid ones: Forest site type complex (further only FTC) 4B species and groups of habitats were organized to (Fagetum eutrophicum), FTC 4S (Fagetum mesotrophicum), increase their clearness so that there is an evident and FTC 4D (Fagetum acerosum deluvium). 4th VT occurs species transfer within biotopes in the diagonal on the 35.2% of the study area (including also non- direction from the le upper corner to the right forest land). lower corner. Acronyms of biogeographical regions: BE – RESULTS AND DISCUSSION biogeographical region of the Beskydský region, PB – biogeographical region of the Podbeskydský 1774 adults comprising 32 species were found th region, VS – biogeographical region of the Vsetínský in total in the 4 VT. The following eudominant region; codes of forest type complexes (according species were found: Caecilius fl avidus, C. burmeisteri, to Plíva 1971, 1991): 4S – Fagetum mesotrophicum, and as dominant species, the following ones were 4B – Fagetum eutrophicum, 4F – Fagetum lapidosum found: Philotarsus picicornis, Elipsocus moebiusi and mesotrophicum, 4H – Fagetum illimerosum trophicum, 4W . In natural geobiocenoses – Fagetum calcarium, 4D – Fagetum acerosum deluvium, 4A with the level of naturalness of 1 or 2, the following – Tilieto-fagetum acerosum lapidosum, for solitaire trees eudominant species were found: Caecilius fl avidus, was used code of “4E” i.e. without classifi cation of Elipsocus moebiusi. Dominant species was Mesopsocus natural community; acronyms of trees and shrubs unipunctatum. Resulting from the comparison (investigated tree species): bb – , bk of the tree colonization, the most numerous – Fagus sylvatica, bo – Pinus sylvestris, dbl – Quercus taxocenosis was found at Picea abies, which resulted robur, dbz – Quercus petraea, hb – Carpinus betulus, hh in a signifi cant change in dominance (see above). – Crataegus laevigata, jb – Malus sylvestris, jd – Abies Abies alba was colonized by numerous taxocenoses, alba, jlh – Ulmus glabra, js – , jv – Acer while the Fagus sylvatica inhabits the very poor range platanoides, lpm – Tilia cordata, md – Larix decidua, os – of psocids species in low abundances (see Tab. 1). th th Populus tremula, sm – Picea abies, sv – Prunus domestica, In comparison with higher VTs (i.e. 5 and 6 ), svk – Cornus sanguinea, trn – , ma – there was a change between species dominance. th malaise trap, pod – undergrowth. Eudominant species of 4 VT – Caecilius fl avidus, occurs in the higher VTs only as dominant species. The characteristic of the 4th vegetation tier This representation is related to the change in tree (Fageta abietis s. lat.) species composition – reducing the representation of broadleave tree species in 5th and then more in 6th According to HOLUŠA & HOLUŠA (2008) 4th VT VT. Contrary species Caecilius despaxi, which in 4th VT (Fageta abietis s. lat.) represents geobiocenoses where occurs only sporadically, in 5th VT is scarce species, Fagus sylvatica is a dominant species in natural in the 6th VT becomes a eudominant species. conditions, it creates the main level of stands Similarly, species Mesopsocus unipunctatus and (the maximum the height over 50 m). Abies alba Stenopsocus lachlani that occur rarely in the 4th, occur occurs in the co-dominant level, sporadically in in 6th VT as a eudominant species. Resulting from the level exceeding the main level. It can reach the the comparison of tree colonization, Picea abies was representation of ± 20% and the height of up to 50 m. the most colonized tree species in all VT: 4th, 5th and Quercus sp. – Quercus petraea and Quercus robur – occur 6th. In contrast to the tree species, Abies alba and Fagus only as interspersed species with the representation sylvatica, which were colonised by lower number of to 10%, they do not reach the co-dominant tree species, u Fagus sylvatica also in lower abundances. level. Carpinus betulus is represented regularly, but in the forest stand structure only in the overtopped Structure and diversity of Psocid taxocenoses (Insecta: Psocoptera) in the forest ecosystems of the Fageta abietis s. 55

350

300 3rd VT

4th VT 250 5th VT

200 6th VT

7th VT 150 8th VT

9th VT 100

50

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

1: DCA analysis of psocid biotopes with marked biotopes of Fageta abietis (4th VT) tier (axis x – gradient of vegetation tiers, axis q – gradient of hydricity)

The DCA-analysis might be interpreted as follows: The characteristic species combination of psocids the x-axis denotes an infl uence of VTs i.e. vertical for the 4th VT was established as follows: Caecilius zonality and the q-axis refers to an infl uence of fl avidus – Elipsocus moebiusi – Peripsocus parvulus. Unlike hydricity (cf. HOLUŠA, 2007c). Habitats of 4th VT create 3rd VT species Ectopsocus meridionalis is missing in 4th a large dotted elongated as in the gravity gradient VT. Also species Enderleinella obsoleta, Cuneopalpus vegetation (x-axis) as well as hydricity gradient cyanops and Caecilius piceus are characteristic for the (q-axis) (see Fig. 1). Dotted fi eld is quite similarly to 4th VT (like for the 5th VT). The “” corticolous the fi eld of the 5th VT, but more elongated along the species are starting to dominate in the changed q-axis. Extensiveness of dotted fi eld corresponds to vegetation (i.e. spruce stands) such as Caecilius the large number of diff erent tree species and the burmeisteri, Philotarsus picicornis, Stenopsocus lachlani, nature of the studied sites in 4th VT. Dotted fi eld in just that at this tree species “descend” to lower VTs. the x – q axis is shi ed higher than fi eld of the 3rd VT. Compared with species combinations of 5th VT, From the viewpoint of hydricity, the fi eld of 4th VT is which was identifi ed as follows: Caecilius fl avidus larger than the fi eld of 5th VT. Generally, habitats of – C.burmeisteri – C.despaxi – nebulosus 4th VT are “drier” than habitats of 3rd and 5th VTs. – Philotarsus picicornis. Elipsocus moebiusi is missing Habitats of 4th VT are represented in more groups in 5th VT, in contrast to 4th VT. The occurrence of in the DvClA-analysis, according to species, where Caecilius despaxi was found as the diff erential feature it was collected. Habitats of broad-leaf trees are between 4th and 5th VTs. 6th VT has totally diff erent sorted into following groups: A-I-b, individually in species combination, which was as follows: Caecilius A-II-a-2 (see Fig. 2.). In the A-I-b group are habitats despaxi – Stenopsocus lachlani – Mesopsocus unipunctatus of 4th VT grouped into larger complexes. Habitats of – Reuterella helvimacula. Caecilius despaxi which is coniferous are sorted into following groups: B-I-a, clearly more dominant, in contrast to the 5th VT. And B-I-b and-II-a. Habitats of coniferous (especially also Reuterella helvimacula is more dominant. There Picea abies) are sorted into higher VTs. Habitats is a conspicuous change of characteristic psocid of 4th VT are represented in the widest possible species, however also in the dominance of Caecilius scale, compared to other VTs, the 4th VT is the most despaxi diagnostic species which occurs in the 6th VT diversifi ed. as a eudominant species. There were found higher values of diversity In conclusion, it is the fi rst step towards to the indexes in the 4B community for the VS4Bsm knowledge of taxocenoses of model group of psocids biotope, i.e. HS 1.52 and HB 1.87, and high psocids in the forest site classifi cation frames – VTs, value was also found for the VS4Bjd psocid biotope, in this article the composition in the zone of Fageta i.e. HS 1.80 and HB 2.21 (see Table 1). Generally abietis s. lat. The data is based on relatively small speaking, the indexes of diversity HS are in the sample of the material, and should take in account interval of 0.18–2.04 and the HB indexes are in the the short-term of study (two growing seasons). It interval of 0.22–2.21. is therefore necessary to continue research and to monitor changes of psocid taxocenoses.

56 O. Holuša

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OD1Lolx PB4Bjd ------1------1------1-1--1------1------1121--1------1---1------1-- -212111111113243434233444334245322422 - ----111---11111121-212-3-311-1 2213------1--1------211111111211------1------4------42-216442124243423342223-31511 ------1------1------1------1------1-1------1-1----1------1------1 ----11--11----1------1--1------1------21-1--2----11--1---11------1------1------11-1--1-----111--1-1------1--1------1---1------1---134---12231212-31341131211 311-11---1----1---11-111--111------12------3------1------1------1------1------311111332-111-11111111112--1211 ------1------412324222-11111-1111---11111212 ------1------1------3111111----1------1------1------1------1------1------1------111------1-1------1------121------1------1------1------1------1------1---1------1---1------1------1 -----1------1------1------1------2 ------1-1------3-- -1------1------1 ------1------1 1111 ----1--1------11------1----1------1 ------1111111111111111111111111111111111111 11111111111111111111111111111111111111111 11110000000000000111111111111111111111111 111111111111 11111111111111000000000000000000000000000 111111111111 00000111111111001000000000000000000000000 111111111111 1111 001111111111 110000000001 0000000000000000000000000000000000000 0000 000000000000 1112111-123341311-1-11-111---1--1------411-1211-1-11-1-1--111--11113-11------53424234-1123311221111211-22111 1--1---1----1-1--11----1-11---1-----1 ------11--1------1------i i i r i Results of DvClA-analysis ‘Twinspan algorithm’; biotopes of the 4 of biotopes algorithm’; ‘Twinspan DvClA-analysis of Results esopsocus laticeps etylophorus nebulosus esopsocus unipunctatus Groups of biotopesGroups A-I-b emineura dispa lipsocus pumilis ctopsocus meridionalis pipsocus lucifugus lipsocus abdominalis nderleinella obsoleta euterella helvimaculata lipsocus hyalinus achesilla quercus iposcelis corrodens oensia fasciata oensia variegata oensia pearman achesilla pedicularia tenopsocus immaculatus tenopsocus stigmaticus tenopsocus lachlan 2: Elipsocus moebiusi Elipsocus M S S Caecilius flavidus E contaminata Caecilius gynapterus L E Graphopsocus cruciatus Psococerastis gibbosa Philotarsus parviceps Peripsocus subfasciatus Caecilius atricornis Peripsocus parvulus Trichadenotecnum majus Blaste quadrimaculata Kolbia quisquiliarum E L E E H R Caecilius piceus M L L E S Trogium pulsatorium Caecilius burmeister L Philotarsus picicornis Cuneopalpus cyanops Amphigerontia bifasciata Trichadenotecnum sexpunctatum L M Caecilius despax

Structure and diversity of Psocid taxocenoses (Insecta: Psocoptera) in the forest ecosystems of the Fageta abietis s. 57

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VS4Bsm

VS5Bsm

PB2Lsm

BE5Hsm

PB3Loll

PB3Hlpm

PB2Ltrn

PB1Tvrp

PB1Toll

PB1Tkro

PB4Djd BE5Ejb

3121111- - - --

VS3Lolx

VS3Lols

PB3Hdbl

PB3Hbk

OP3Hkro

OP3Hbk

OD1Lkl

BK3Bdbz

BK2Cdbp

BE6Obk

VS4Bos

PB3Lkl

OP1Lvrk BK2Ctrn

00000000000000000000000000000000000000000000 1111111111111111111111111 0000000000000000000000000 111111111111111 1111111111111111100000000 000000000000000 111 1111111111111111 0000111111111100111111 110000000000000 000 0000000000000000 110111111111111 000 1111111111111111 11 1111111110000000 00 111111000000 000011000001 0000111110000011 00010011101111

PB3Lstr

PB3Lsm

PB3Lkro

PB3Gbc

PB2Loll

PB2Llis

PB2Ljiv

PB2Lbc

PB1Ljiv

OP3Hkl

OP1Loll

OD1Lols

OD1Ljlh

OD1Lbc VS4Ssm ------2------1-12------1------111------1-241111 ------21-2-- --1------1 ------1--2121111-- ---1------1------1------1------1-----1------1------1--1------1------1------1-----1------1------1------1------1112111------1 -1111- -21112- -1------1------1------1------1-----1- 1----11--111-- -1------1--11-1---- 11------1------1-3----1--1 -3--1-111-----1 11-2------1 ------1------1 ------2------2 111 ------2------46--1-11------1 ------1------1------1---2------1------1----1------331 ------2-- - -1------55--3--1111--11 ------1--1----1---1--- 311------33---11--1----- 1--- -1-43 24------6 ------3------221-1-1--21221- 1---24--1- --1 -----1------1------1-----1 ------1-----1------12---1------1------1------1------1------121 ------12124 45-----111---11 ---14-1------2 ------1------1---1-1------224 ------22466 ------66-12-211111323 ------534 ------2 461---1--1---11 ------1------1-----2------251--1------11------1--1------2 11-1---1-- 21- 36------21-13 0000000000000 0 1111111111111 1 1111111111111 1 1111111111111 1 0111111111111 1 0000000000000 0 -1-11- 11--1--1111------1-1-- -2------1------111- -111111- 1 -2------11111111111121- -111- 3- -11 2------11------1--- -1-----1------1----1-----12 - 1----1---- 311111211 ------2------31--12 31- - 1- - 14------2--6 31-2141-1------1111-111-1111- -1 535 36- - 42111- - -1-1 5355361123-1--21 1------1------352------23------1-----2--1------12312-1 ------1----1---1- 1-1 1---211------3 1-141--2-- r continuation Groups of biotopesGroups A-II-a-1 A-II-a-2 B-I-a B-I-b lipsocus pumilis ctopsocus meridionalis pipsocus lucifugus lipsocus abdominalis nderleinella obsoleta euterella helvimaculata lipsocus hyalinus tenopsocus immaculatus stigmaticustenopsocus tenopsocus lachlani 2: Elipsocus moebiusi Elipsocus Mesopsocus laticeps S S Caecilius flavidus E Amphigerontia contaminata Caecilius gynapterus quercus E Graphopsocus cruciatus Psococerastis gibbosa Peripsocus phaeopterus parviceps Philotarsus Peripsocus subfasciatus atricornis Caecilius Peripsocus parvulus Trichadenotecnum majus Blaste quadrimaculata Kolbia quisquiliarum E corrodensLiposcelis E E Hemineura dispa R Caecilius piceus didymus Peripsocus Peripsocus alboguttatus Metylophorus nebulosus fasciata Loensia variegata Caecilius fuscopterus E S Trogium pulsatorium burmeisteriCaecilius Loensia pearmani Philotarsus picicornis Cuneopalpus cyanops Amphigerontia bifasciata Trichadenotecnum sexpunctatum Lachesilla pedicularia Mesopsocus unipunctatus Caecilius despaxi 58 O. Holuša VS4Dsm VS4Emd BE5Sbk BE6Sbk PB4Dsm BE4Ssm OR8Zjan VS3Ltrn VS5Asm VS5Bjd BE6Spod BE7Spod OR5Sjd BE8Zpod OR8Zkos OR6Bsm BE5Fjd BE5Fsm BE6Gsm BE7Fbk VS4Djd VS4Sjd BE5Sjd BE6Fsm BE6Sjd BE6Ssm BE6Zsm BE7Fsm BE7Sjiv BE7Ssm BE7Zsm BE8Zbk BE8Zkos BE8Zma BE8Zsm OR7Ssm OR8Sjr OR8Ssm OR8Zsm OR9Kkos OR9Ksm

Groups of biotopes B-II-a-1 B-II-a-2 B-II-b-1

Elipsocus moebiusi ---1------1------0001 Mesopsocus laticeps ------0001 Stenopsocus immaculatus ------0000 Stenopsocus stigmaticus ------0000 Caecilius flavidus --431----1------1-----11---1--11111------0010 ------0010 Amphigerontia contaminata ------0011 Caecilius gynapterus ------01000 Lachesilla quercus ------01000 Ectopsocus meridionalis ------01001 Graphopsocus cruciatus ------2------01010 Psococerastis gibbosa ------1------01010 Peripsocus phaeopterus -1------01011 Philotarsus parviceps ------1--- 01011 Peripsocus subfasciatus 1------1---11-----1--1------01100 Caecilius atricornis ------01101 Peripsocus parvulus ------01101 Trichadenotecnum majus --1------1-- --11----1---1------0111

Blaste quadrimaculata ------11000 Kolbia quisquiliarum ------11000 Epipsocus lucifugus ------110010 Liposcelis corrodens ------110010 ------110011 Enderleinella obsoleta ------110011 Hemineura dispar ------110011 Reuterella helvimaculata ------2---1------110011 Caecilius piceus ------11------1------1--- 1101 Peripsocus didymus ------1----1------1--- 1101 Peripsocus alboguttatus ------1110 Metylophorus nebulosus ------111------111100 Loensia fasciata 1-1------111101 Loensia variegata ------111101 Caecilius fuscopterus 1------11111 ------11111 Stenopsocus lachlani 1--1------11-3 ---1-11111513-451145226525 10100 Trogium pulsatorium ------1------10100 Caecilius burmeisteri 112332122211111 334- - - - 322513-43-114- -45-6 10101 Loensia pearmani ------1------10110 Philotarsus picicornis ------11--- 2233----1-3-3-22---42-11-1 10110 Cuneopalpus cyanops 1------11------10111 Amphigerontia bifasciata 11--1------1-51--11-141-1 1000 Trichadenotecnum sexpunctatum ------1------100100 Lachesilla pedicularia ------1------1-3-1111-3---1-2 100101 Mesopsocus unipunctatus 2-11--1--1------231234515145--151-12-3 100101 Caecilius despaxi - - 11- -1------11 134- - - - 3226253661366226214 10011

111111111111111 11111111111111111111111111 111111111111111 11111111111111111111111111 000000000000000 11111111111111111111111111 000000000011111 00000000000000000000000000 001111111100011 00001111111111111111111111 11100000001 01111110000000000000000000 2: continuation

CONCLUSION The results of this work with previous results for the 5th Abieto–fageta s. lat. (HOLUŠA, 2009), 6th Piceeti– fageta s. lat. (HOLUŠA, 2011), 7th Fageti–piceeta s. lat and 8th Piceeta s. lat. VTs (HOLUŠA, 2007c) create a survey of compositions of psocid taxocenoses in the majority of VTs in the Western Carpathian Mts. The 4th VT is a community of Fageta abietis s.lat. with large richness of tree edifi cators, therefore there is also larger diversity types of forest stands. With regard to richness and abundance of psocid species, 4th VT is the richest. Compared to psocid taxocenoses, there is a noticeable diff erence to the higher VTs, which also corresponds phytocenological consequently geobiocenological characteristics of VTs. This work verifi es the hypothesis that units of geobiocenological or forest-typological system i.e. at fi rst VTs, have proved to be a suitable frame for zoocenological studies. These units, as characteristics of a potential state of ecosystems, together with the description of the present tree species composition and the level of naturalness form a perfect base for studies focused on the taxocenoses structure. Therefore VTs are an ideal frame for animal (entomological) studies. And on the other side, the order of psocids can be used for geobiocenological classifi cation of ecosystems and also for evaluation of potential ecosystem changes. Structure and diversity of Psocid taxocenoses (Insecta: Psocoptera) in the forest ecosystems of the Fageta abietis s. 59

Acknowledgement This paper was worked up in the frame of the research project „Strategy of the management of territories with a special protection status“ MSM 6215648902-04 of Faculty Forestry and Wood Technology of Mendel University in Brno.

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Address doc. Ing. Otakar Holuša, Ph.D., Ústav ochrany lesa a myslivosti, Mendelova univerzita v Brně, Zemědělská 1, 613 00 Brno, Česká republika, e-mail: [email protected]