MUSEUM AND I N S T I T U T E OF Z O O L O G Y P O L I S H A C A D E M Y OF S C I E N C ES FRAGMENTA Fragm. faun. FAUNISTICA W arszawa, 30.12.2000 43 Supplem ent 333-342 11 th International Congress of M yriapodology, 20-24 July 1999, Białowieża, Poland PROGRESS IN ST UDIE S ON M Y R IA P O D A A N D O N Y C H O P H O R A Jolanta W ytwer C entipede (C hilopoda) com m unities of som e forest habitats of Puszcza B iałow iesk a in Poland Abstract: Species com position and structure of C hilopoda com m unities w ere studied in five n atural forest habitats of Puszcza Białowieska, eastern Poland: fresh pine forest, pine-spruce m ixed forest, linden-oakhornbeam forest, ash-alder flood plain forest, and bog-alder forest. Species num ber ranged from 7 to 9 and was the highest in the linden-oak-hornbeam forest. The centipede com m unities w ere very sim ilar in species composition. The abundance of centipedes increased along the fertility gradient (pine forest —> linden-oakhornbeam forest). The edaphic com ponent of each com m unity w as dom inated by Lithobius curtipes, while Lithobius mutabilis w as the dom inant species in the epigeic com ponent, w ith the exception of the bog-alder forest w here the com m unity w as also dom inated by L. curtipes. In this type of forest, com m unity structure w as the least sim ilar to those of the other com m unities, particularly w ith respect to the epigeic com ponent. C entipede com m unities in habitats characterised by extreme m oisture conditions (fresh pine forest, ashalder flood plain forest and bog-alder forest) show ed either the highest or the low est hom ogeneity of dom inance structure. Key w ords: Chilopoda, Puszcza Białowieska, forest type, diversity, com m unity structure A uthor's address: M useum and Institute of Zoology, PAS, Wilcza 64, 00-679 W arszaw a, POLAND; e-mail: jolawyt@ robal.miiz.waw.pl INTRODUCTION Puszcza Białowieska is a unique com plex of natural prim eval low land forests of the boreo-nemoral zone, w here the primary phytosociological and ecological differentiation corresponds to the habitat variety specific for denudation plains of the eastern part of the postglacial North European Lowland (F a l i n s k i 1994). J. 2000. C entipede (Chilopoda) com m unities of som e forest habitats of Puszcza Białowieska in Poland. In: W y t w e r J. & GOLOVATCH S. (eds), Progress in Studies on M yriapoda and O nychophora, W arszawa, XIV+ 396 pp. Fragm. faun. 43 (Suppl.): 333-342. WYTWER http://rcin.org.pl | 334 J. W ytw er O w ing to this, a com prehensive analysis of centipede com m unities in different forest habitats provides an opportunity to learn more both about the fauna of the region and the factors governing the population and biotic structure characteristic of a given com m unity (WYTWER 1999). STUDY AREA C entipedes w ere collected in forests representing the five basic units of vegetation landscape, described by KWIATKOWSKI (1994) as forest biogeocenoses of Puszcza Białowieska, on the basis of a classification of forest associations by SOKOŁOWSKI (1980), namely: 1. Fresh pine forest (Vaccinio vitis-idaeae-Pinetum) - a thinned coniferous forest with pine as dom inant and admixtures of spruce and birch; ground cover is com posed chiefly of m osses and undershrubs. There is lack of species typical of m oist habitats w h ile xerothermal species are num erous. This type of forest is found on plains of aeolian accumulation; ground water is present at m edium depths (2-5 m). 2. Pine-spruce m ixed forest (Calamagrostio arundinaceae-Piceetum) - typical of ablation m oraine plains, w ith spruce as dom inant and a high contribution of pine, and an adm ixture of birch, sporadically, of oak and hornbeam; a lush ground flora is com p osed of species characteristic of spruce forests and, generally, of forest habitats, reflecting a relatively eutrophic biocenosis; ground water table lies deep, at 5-15 m; 3. Linden oak-hornbeam forest (TUio-Carpinetum typicum) - m ulti-species and m ulti layer forest stands found on ground moraine flat plains, com posed of hornbeam, maple, sm all-leaved linden, oak and spruce; fertile-habitat species abound in the ground flora; ground water table beyond the soil profile. 4. Ash-alder flood plain forest (Circaeo-Alnetum) - a sw am py alder forest grow ing along stream s on ground moraine plains, w ith a considerable contribution of ash and a sm all admixture of either spruce or hornbeam and linden; it has a poorly d eveloped understory and a w ell developed herb layer with num erous bogassociated species and a tufted, m osaic structure; ground water table reaches the soil profile but periods of stagnation and flooding are not long. 5. Bog-alder forest (Carici elongntae-Alnetum) - a marshy alder forest of m osaic and tussock structured ground vegetation; spruce and birch are constant elem ents and, together w ith scrub layer species, form and occupy bog tussocks; prolonged periods of ground water stagnation; the ground flora of tussocks is com posed of species characteristic of m eso- and oligotrophic habitats, w hile the w aterlogged areas are occupied by marshy vegetation, with a high proportion of tall perennials; plants typical of water habitats are found locally in hollow s in the ground. The first three forest types represent lithogenic habitats and have been grouped in the order of increasing fertility from an oligotrophic fresh pine forest via a m esotrophic pine-spruce mixed forest to an eutrophic linden-oak-hornbeam forest. In contrast, forest associations 4 and 5 are hydrogenic habitats; the ash-alder flood plain http://rcin.org.pl C entipede com m unities of Puszxza Białowieska 335 forest resem bles the linden-oak-hornbeam forest in terms of fertility w hile the bogalder forest provides a m esotrophic environment. Sam pling w as only carried out in mature natural stands, i.e. approxim ately 100year old in the pine forest and linden-oak-hornbeam forest, and at least 60-year old in both alder forests. METHODS OF SAMPLING AND FAUNISTIC ANALYSES Two m ethods w ere used for quantitatively sam pling the Chilopoda: . litter sifting w herever it w as found practicable, i.e. in both types of coniferous forest and the linden-oak-hornbeam forest; in both types of inundated alder forest, w here the upper soil layer, litter and ground cover were difficult to separate, soil sam ples w ere taken to a depth of about 15 cm; sam ple size was the sam e for both (10 x 0 . 1 m 2); the specim ens w ere then collected by hand. 2. Barber's pitfall trapping, used from April to October; 10 open traps filled w ith a 1 0 % glycol solution w ith som e detergent were set up in each forest section. Data on the periods and m ethods of quantitative sam pling at individual sites are presented in Table I. 1 Table I. Study sites and m ethods of sam pling of Chilopoda. Sites (forest section) 538, 667, 668 Soil sa m p lin g /litte r sifting N um ber of samples** Time of study Habitat Fresh pine forest (Peucedano1986-1987 Pinetum) 2 Pine spruce m ixed forest 1994-1995 (Calamagrostio arundinaceae- 288/318, 521, 699 Piceetum) 3 Linden oak-hornbeam forest 334, 399, 633 1994-1995 (Tilio-Carpinetum typicum) 4 A sh-alder flood plain forest 315, 599, 600 1991 (Circaeo-Alnetimi) forest (Carici 5 Bog-alder 514/539 1997, 2000 elongatae-Alnetum) bog tussocks Sections m arked in bold w ere sam pled also by pitfall trapping *A sam ple m eans 10 x 0.1 m 2 1 Pitfall trapping Frequency of Time of study trap em pting 24 1986-1987 2 weeks 8 1995 4 weeks 8 1995 4 weeks 7 1991 2 weeks 2 1997 4 weeks The material obtained from siftings and from soil sam ples w as regarded as representative of the edaphic com ponent of com m unities, whereas those collected by m eans of pitfall trapping and reflecting the intensity of centipede penetration into the surface layer of the forest soil served as representive of the epigeic com ponent. Qualitative sam pling, i.e. looking for centipedes specifically in dead w ood , m ould, stum ps etc., w as carried out several times during the season. The analyses of the Chilopoda com m unities used standard indices em ployed in faunistic studies and soil zoology (M A GURRA N 1988, GÓRNY & G RU M 1993): http://rcin.org.pl J. W ytwer 336 □ index of density = number of individuals per m 2 of soil sam ple or sifting; □ trapability index = number of individuals caught by 10 Barber's pitfall traps over 14 days; □ the M arczewski-Steinhaus index of similarity in species com position: w here Sx and Sy represent the number of species in habitats X and Y, respectively, and Sc is the number of species com m on to both habitats; □ index of dominance: x, = N 100% w here n, represents the density (trapability) of a given species, N represents the density (trapability) of the community; □ Morisita's index of similarity of dominance structures, m odified by HORN (1966) s w here x, and y, are dom inance structure indices of z-th species in habitats X and Y, repectively, and S is the total number of species; □ Shannon's index of species diversity: w ith the same sym bols as above; □ Pielou's index of eveness: where H max = log2 S* and is equal to potential diversity at the m axim um possible num ber of species S* in a community; ABUNDANCE The Chilopoda com m unities studied show ed variation in abundance relative to the habitat (Fig. 1). As the lithogenic habitats increased in fertility, both density and trapability rose. The density, estim ated from soil sam ples, w as found to range from 3.25 in d ./m 2 in the pine forest to 19.75 in d ./m 2 in the linden-oak-hornbeam forest. The trapability, estim ated by pitfall trapping, w as revealed as ranging from 0.75 to 2.33 in d ./1 0 traps x 14 days. In the inundated, though fertile, ash-alder flood plain forest, the values of both indices were lower, with the trapability index reaching its m inim um http://rcin.org.pl Centipede com m unities of Puszcza Białowieska 337 there (0.20 i n d ./ 10 traps x 14 days). Conversely, tussocks in the bog alder forest show ed the highest density of centipedes (39.50 in d ./m 2) and, also, a very low trapability index (0.69 in d ./1 0 traps x 14 days). Depressions in this type of forest were either excluded from sam pling on account of being inundated or they were too sw am py to yield any centipedes. D E N S IT Y 1 2 3 T R A P A B IL IT Y 4 5 1 2 3 4 5 Fig. 1. Values of the abundance indices (density and trapability) in different forest habitats of Puszcza Białowieska. SPECIES COMPOSITION A total of 10 species of Chilopoda w ere identified in the study habitats (Table II). All study habitats w ere characterized by relatively similar species richness estimates. Interestingly, different habitats of similarly high fertility were show to contain both the highest (9 - linden-oak-hornbeam forest) and the low est ( 6 - ash-alder flood plain forest) number of species. Four (sub)species were com m on to all study habitats: Lithobius forficatus, L. tenebrosus fennoscandius, L. lapidicola, and L. curtipes. Table II. O ccurrence of C hilopoda species in five different habitats of the Białowieża Forest. H a b ita t No 1 2 3 4 5 6 7 8 9 10 S p e c ie s 1 Geopliilus proximus C. L. KOCH, 1847 Pachymerium ferrugineum (C . L. K o c h , 1847) Lithobius eri/throcephalus C . L. KOCH, 1847 Lithobius forficatus (LINNAEUS, 1758) Lithobius lapidicola M e i n e r t , 1872 Lithobius mutabilis L. K o c h , 1862 Lithobius pelidnus H a a s e , 1880 Lithobius tenebrosus fennoscandius L o h m a n d e r , 1948 Lithobius curtipes C . L. K o c h , 1847 Lithobius crassipes L. KOCH, 1862 Total + + + + + + + + 8 http://rcin.org.pl 2 3 4 5 + + + + + + + + + + + + + + + + + + 9 7 + + + + + + + + + + + 6 7 J. W ytwer 338 5 All the com m unities of centipedes included in this study were characterized by relatively high similarities in species com position, w ith values of the M arczewski-Steinhaus index ranging from 0.56 for the fresh pine forest and ash-alder forest centipede com m unities to 0.78 for the linden-oakhornbeam forest and bog alder forest centipede associations. The similarity in species com position of the five study com m unities is presented graphically by a dendrogram in Fig. 2, obtained using M oUNTFORD's (1962) method. 4 1.0 09 08 0 7 06 05 0 4 03 02 Fig. 2. D endrogram of sim ilarities in species com position of centipede com m unities in the stu d y forest habitats of Puszcza Białowieska. 0 1 0.0 DOMINANCE STRUCTURE The edaphic com ponent of all Chilopoda communities studied in the different habitats of Puszcza Białowieska (estimated from soil sam ples or by sifting) w as dominated by Lithobius curtipes, w hose estimated percentage contribution ranged from nearly 45% in the bog alder forest to more than 90% in the mixed pine-spruce forest and the ash-alder flood plain forest (Table III). The epigeic com ponent (estimated by Barber's pitfall trapping) w as dominated by Lithobius mutabilis, with the exception of the bog alder forest where L. curtipes was the dominant species in both com munity components. In this com m unity, Geophilus proximus was subdominant, this holding true for both components. It w as also subdominant in the ash-alder forest, yet only in the epigeic component, but it w as totally absent from the coniferous forest habitats. Table III. Percentage contribution of individual species of Chilopoda in the m aterial collected from the different habitats in Puszcza Białowieska, estim ated from sifting/soil sam ples (ss) or Barber's pitfall traps (pt); + refers to species collected using other techniques. H abitat 3 ss Pt + 3.8 No Species 1 ss 1 2 3 4 5 6 7 8 Geophilus proximus Pachymerium ferrugineum) Lithobius erythrocephalus L. forficntus L. lapidicola L. mutabilis L. pelidnus L. tenebrosus fennoscandius Lithobius (L.) sp. 9 L. curtipes 10 L. crassipes 2.6 + + 5.1 10.3 12.8 + 5.1 64.1 2 Pt + + 8.0 4.3 73.0 1.6 6.6 4.7 1.8 Ss Pt + + 6.3 1.6 + + 6.5 74.7 10.3 + + + + 17.1 0.6 + + + + 75.1 4.0 + 78.5 21.0 + + 4 ss 7.1 5 Pt + + + + + + 43.2 + 14.2 92.9 42.6 8.6 92.1 + http://rcin.org.pl ss 43.0 P‘ 28.8 + 8.9 1.3 + + + 1.3 + 1.3 44.3 + + 71.2 Centipede com m unities of Puszcza Białowieska 339 The structure of dom inance of the centipede communities in the five study habitats appears relatively similar w hen the material obtained from sifting or soil sam ples is analysed, i.e. in the edaphic component. For this com ponent, the low est values of Morisita's index of similarity of pairs of com m unities were 0.66-0.69. W hen the individual com m unities are clustered using M ountford's m ethod, the bog alder forest centipede com m unity stands out as the least similar to the others, being segregated only at a similarity value of 0.69 (Fig. 3A). The structure of dom inance of the centipede com m unities show ed a greater diversity w h en material obtained by pitfall trapping w as compared. The values of Morisita's index for pairs of com m unities ranged w idely, from 0.01 to 0.98. A graph show in g the similarity for the epigeic com ponent of Chilopoda com m unities reveals that the highest similarity of dominance structure can be seen betw een com m unities from lithogenic habitats (fresh pine forest, pine-spruce m ixed forest and linden-oakhornbeam forest), the index values being at least 0.96 (Fig. 3B). The ash-alder forest com m unities displayed a similarity index of 0.74 as regards these three com m unities, w hile the bog alder forest Chilopoda com m unities were the least similar to all others, at 0.26. 10 1.0 0.9 0.9 0.8 08 . 0.7 0.7 0.6 0.6 0.5 0.5 0.4 - 0.4 0.3 - 0.3 - 0.2 02 . 0.1 0.1 00 J 0.0 , Fig. 3. D endrogram s of dom inance structure sim ilarity for the two com ponents of C hilopoda comm unities: A - edaphic, B - epigeic. SPECIES DIVERSITY The linden-oak-hornbeam forest habitat supported the highest num ber of species of centipedes recorded and, consequently, the highest m aximum diversity (Hmax) of the Chilopoda com m unity (Tab. IV). Flowever, this did not entail a high actual diversity (FI') or eveness (J) of the dominance distribution in either of the com m unity com ponents. The highest values of these two indices for the edaphic com ponent were http://rcin.org.pl T. W ytw er 340 obtained for both habitats characterized by extreme moisture conditions, i.e. the fresh pine forest and the bog alder forest. The epigeic com ponent of the com m unities is subject to more com plex relations. Here the values of the index of eveness, more suitable for com parisons, were the highest in the ash-alder flood plain forest, follow ed by the fresh pine forest, indicating m ost even (=flat) dom inance structures. Table IV. Com parison of various diversity param eters for C hilopoda com m unities in the five different habitats of Puszcza Białowieska. H abitat Hmax E daphic p art of the com m unity Epigeic p art of the com m unity H' 1 2 3.00 1.48 1 H' 0.49 1 0.43 1.28 3 2.58 0.46 0.18 0.91 0.35 4 3.17 0.89 0.28 0.97 0.31 5 2.58 0.37 0.14 1.45 0.56 2.81 1.35 0.48 0.87 0.31 DISCUSSION The study habitats ot Puszcza Białowieska are not as abundant in centipedes as Central European beech forests (ALBERT 1982, F r u n d 1987, LEŚNIEWSKA 2000), where studies have yielded several dozen to several hundred centipede individuals per sq. m and usually a dozen or so species. N onetheless, the linden-oak-hornbeam forests of Puszcza Białowieska are quite rich in centipedes (almost 20 ind./rr?) com pared to the sam e type of forest in the W ielkopolska (Great Polish) Lowland or the M azovian region, where the same collecting techniques normally reveal low er com m unity densities (KACZM AREK 1977, 1989, W YTWER 1990). The sam e holds for Chilopoda com m unities of pine forests in Puszcza Białowieska, as com pared to their counterparts in other regions of Poland (W YTWER 1992). The results of the study, w hich com prised five different forest habitats located in the sam e region, can be taken to show that the abundance of Chilopoda depends not only on moisture conditions, but also on soil fertility of the forest habitat. For it is unlikely that the thin mull-type litter of the linden-oak-hornbeam forest w ou ld provide better m oisture conditions for Chilopoda than the thicker rnor litter in the m ixed forest. In each of the forest habitats studied, during periods w hen the litter is drier, centipedes are able to find adequately m oist microhabitats. H owever, com m unity abundance increases markedly along w ith rising fertility in lithogenic habitats: from the pine forest to the mixed pine-spruce forest to the linden-oak-hornbeam forest. Thus, it appears that a higher abundance may sim ply be due to the presence of a better nutritive base in a more fertile habitat, for example one with a more abundant m esofauna. On the other hand, the abundance of centipedes in habitats where the soil is inundated by ground water and is subject to periodic floods, appears to be independent on soil fertility. In the ash-alder flood plain forest, where flood water can cover the entire area, even though the flooding is but short in duration, the abundance of Chilopoda is dramatically lower. Centipedes are also virtually absent from the swam py hollow s of the bog alder forest, where water is usually present during most of the year, w hile in the http://rcin.org.pl Centipede com m unities of Puszcza Białowieska 341 tussocks of the alder forest, usually situated above water level during flooding, Chilopoda are twice as abundant as on the ground surface in the linden-oak-hornbeam forest, a habitat supporting the highest number of Chilopoda species. The above changes in abundance are also associated with reshaping the dom inance structure of the Chilopoda com m unities, less so w ith change in species com position. In the study com m unities representing lithogenic forest habitats of Puszcza Białowieska, the lithobiom orphs Lithobius curtipes and L. mutabilis were absolute dominants. The latter species is typically epigeic, hence the m ost frequently captured by pitfall trapping. It is much less abundant, how ever, in the ash-alder flood plain forest and is absent from the bog alder forest, where Chilopoda are only found on sm all-sized tussocks separated from one another by stagnant water. L. curtipes remains abundant in this habitat, but equally abundant is the edaphic Geophilus proximus, a co-dom inant in the bog alder forest com m unity. This explains w hy, despite a high density of Chilopoda in the bog alder forest, not many centipedes were caught in pitfall traps. The structural changes in the com m unities as revealed by diversity indices do not lend them selves to an unequivocal interpretation. The m ost even (flat) pattern of dom inance structure observed in this study in com m unities from habitats characterised by extreme m oisture conditions can be taken as a sign of intensified com petition betw een species or m ay indicate a com m unity w ell adapted to changing habitat conditions. H ow ever, w hile the even structure of dominance was observed for both edaphic (J=0.49) and epigeic (J=0.43) com ponents in the fresh pine forest com m unity of Chilopoda, in the ash-alder flood plain forest this w as only seen in the epigeic com ponent (J=0.56) w hile in the bog alder forest this was seen in the edaphic com ponent of the Chilopoda com m unity (J=0.48). The edaphic com ponent of the ashalder forest Chilopoda com m unity and the epigeic com ponent of the bog alder forest com m unity have the same dom inant species, L. curtipes, a w idespread form (ZALESSKAJA 1978) w hich is am ong the few centipedes know n to inhabit inundated areas (ZULKA 1992, Z e r m 1997). REFERENCES M. 1982. Species spectrum and dispersion patterns of Chilopoda in Soiling habitats. Pedobiologia 23: 337-347. F a liŃ S K I J. B. 1994. Concise Geobotanical Atlas of Białowieża Forest. Phytocoenosis, W arszaw a-B iałow ieża, 6 (N. S.): 3-34. F r u n d H.-Ch. 1987. 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