Chilopoda) Diversity in the Wielkopolska-Kujawy Lowlands of Western Poland

Biologia 63/5: 711—719, 2008 Section Zoology DOI: 10.2478/s11756-008-0121-0

Eﬀect of urbanization on centipede (Chilopoda) diversity in the Wielkopolska-Kujawy Lowlands of western Poland

Malgorzata Le´sniewska1,PiotrLe´sniewski2 &KrystynaSzybiak1

1Department of General Zoology, Adam Mickiewicz University, Umultowska 89,PL-61614 Pozna´n, Poland; e-mail: [email protected] 2Department of Logic and Methodology of Science, Institute of Philosophy, Adam Mickiewicz University, Szamarzewskiego 89 C, PL-60569 Pozna´n, Poland

Abstract: Over fifty years of research data from various sources were compiled and analyzed in order to determine the effect of urbanization on centipede diversity in the Wielkopolska-Kujawy Lowlands of western Poland. Urbanization has had a significant effect on species composition and community structures. However, it has not reduced overall species richness and diversity in the centipede fauna. The centipede fauna from built-up areas was found to be rich and varied. The habitats with the highest levels of species richness were the city of Poznań and the beech forests in the surrounding region. Key words: Chilopoda; diversity; species richness; forests; urbanization; Poland

Introduction of Wielkopolska and Kujawsko-Pomorskie in west-central Poland. This is a lowland area with maximum elevations Centipedes (Chilopoda) are a group of invertebrates not exceeding 200 m above sea level. Total annual rainfall that originally evolved in forest soils (Bornebush 1930). is lower here than in other parts of the Polish lowlands, In Europe, forest ecosystems are among the ecosys- and averages between 450 and 500 mm. Most of the area is under cultivation, and there are relatively few forests. The tems least altered by human activity. Therefore, certain area has one national park, six landscape parks, and more groups of forest centipedes are good reference systems than a dozen nature reserves (Kondracki 2002). for studying the degree to which natural communities The data used in this study were collected at 313 sites, of centipedes are affected by anthropogenic stress, es- comprising various types of forest, including alder, beech, pecially urbanization. oak, oak-hornbeam, oak-pine and marshy forests. Other There have been numerous publications on species data were obtained from various built-up areas, including diversity in centipede communities in synanthropic and Poznań (no. of inhabitants 565,000), Zielona Góra (120,000), ´ urban environments of Europe (Enghoff 1973; Tischler Leszno (64,000), Srem (30,000), Turek (30,000), and Pyzdry 1980; Andersson 1983; Lewis 1985; Fründ 1989; Zap- (3,000). The classification of environments and habitats used paroli 1992; Christian 1996; Le´sniewska 1996; Wytwer in this study largely corresponds to the classification used 1996). There have also been many publications on by the original researchers. Detailed descriptions of the re- centipede communities in forest ecosystems (Wignara- search areas can be found in the relevant works. jah & Phillipson 1977; Geoffroy 1979; Albert 1982; Wytwer 1990; Fründ 1991; Országh & Országhová 1997; Tajovský 1999; Tuf 2000). However, only one paper has Material and methods been published on the relationships between the urban The following published materials were used: Kaczmarek centipede fauna and the fauna of the natural habitats (1952, 1964, 1977, 1979, 1980, 1989), Zych (1989), Le´sniew- of the surrounding countryside (Wytwer 1995). ska & Wojciechowski (1992), Le´sniewska (1996, 1997, 2004), The aim of this study was to compile and analyze Le´sniewska et al. (2005). Unpublished data collected for over fifty years of research data from various sources master theses were also provided by K. Baron, M. Chi- in order to determine the effect of urbanization on cen- ciak, K. Cieplicka, M. Ciszak, A. Drewniak, I. Gonerka, tipede diversity in the Wielkopolska-Kujawy Lowlands M. Górecka, T. Gruszczyńska, M. Kaliszak, H. Polańska, of western Poland. M. Rzepka, U. Smolińska, M. Walerowicz, M. Wilkosz and J. Wojciechowski (Table 1). These data were collected between 1949 and 2005, using both qualitative and quantita- Study area tive methods. Basic data on the duration of the studies and on the methods used are presented in Tables 1 and 2. The data used in this study were collected from the Direct capture from under branches, stones, pieces Wielkopolska-Kujawy Lowlands, which lie in the provinces of bark, and other accessible micro-habitats in which

c 2008 Institute of Zoology, Slovak Academy of Sciences 712 M. Le´sniewska et al. 9 2 14 10 26 33 34 96 39 99 149 325 4 109 65 235 16 287 550 553 1 43 88 2 15 71 7 257 75 1 538 1 17 107 1 13 12 79 256 1116 2 16 12 23 1 1351 21222324Total 8 60 8 41 5 115 95 4 10 + 3 + 4 + 28 350 171 + 210 149 414 16 284 7642 5 4 + 4 1 16 51 429 + 2+ 3 29 + 46 8 26 20 + 68 5 10 12 617 7 22 + 6 23 + 71 43 65 3 64 6 + 440 15 70 7 179 1170 4 23 7 1 2 35 1 1 3 + 104 7 3 11 1 11 252 15 7 + 17 and sample type. 1 11 13 16 100 32 54 + 339 22 1 5 6 61 1 5 2 3 1 6 3 1 6 2 46 8 2 24 41 + 19 5 15 1 13 1 4 6 1011121314151617181920 2 + + 3 + 3 + 1 1 + 2 3 4 4 2 2 3 + 6 9 3 85 + 9 14 8 11 24 13 33 667 + 965 343 405 73 113 994 18 89 1 1 + 2 4 2 4 2 3 3 6 + 2 2 + 1 1 5 + 1 3 3 1 2 1 1 + 54 2 + 18 1 + 4 6 1 1 1 1 23 1 21 2 4 1 37 7 60 + 4 10 + 9 3 ede species detected in Wielkopolska, with study period 5 9 3 + 8 + 1 2 3 4 5 6 7 8 9 49 3 31 + 1 1 + 9 4 6 67 5 35 7 9 + 36 4 4 15 20 4 55 + 15 36 + 325 1 2 54 34 17 15 1 + 1 3 1 15 1 14 1 12 2 + 2 3 1 25 25 11 + 6 15 + 30 7 2 15 44 278 299 148 + 39 53 + 291 25 18 o&Meinert, (Leach, 1817) (C.L. Koch, 1835) C.L. Koch, 1847 Leach, 1814 [e, E, (Newport, 1844) [e, E, (C.L. Koch, 1837) [e, L. Koch, 1862 [f, E] 10 2 2 (Bergs¨ (Leach, 1814) [f, E, L. Koch, 1862 [f, E] 2 Attems, 1895 [f, E] 1 C.L. Koch, 1844 [f, E] 26 49 + (L., 1758) [e, s, E] 5 1 2 Kaczmarek, 1962 [f, E] 46 6 5 + Meinert, 1872 [f, E] C.L. Koch, 1847 [f, E (C.L. Koch, 1835) [e, E, C.L. Koch, 1847 [f, EA] (L., 1758) [e, WP] 268 1045 716 + 352 L. Koch, 1862 [e, E, Naf, Leach, 1814 [e, E] Latzel, 1880 [f, E] C.L. Koch, 1844 [f, E] C.L. Koch, 1847 [f, P] 29 3 107 + C.L. Koch, 1835 [s, E] Meinert, 1868 [f, E] (De Geer, 1783) [e, s, ES] 72 77 17 23 C.L. Koch, 1847 [f, E] Reference \ Stigmatogaster subterranea [e, E, iNA] Strigamia acuminata Lithobius aeruginosus iNA] NAm, Af, Au] Lithobiomorpha Lamyctes emarginatus Naf, CA] Strigamia crassipes Lithobius calcaratus Lithobius cyrtopus Lithobius forficatus Lithobius lapidicola Lithobius macilentus Lithobius curtipes s, E, Naf, iNA] Schendyla furcidens Schendyla nemorensis Lithobius agilis Lithobius borealis Lithobius erythrocephalus [e, E] Lithobius crassipes CA] Lithobius dentatus Pachymerium ferrugineum [e, H] Geophilus truncorum 1866) [e, E] iNA] Geophilus electricus Geophilus flavus Geophilus insculptus Geophilus linearis Geophilus proximus Table 1. Number of specimens of each ofSpecies the centip Scolopendromorpha Cryptops hortensis Geophilomorpha Clinopodes flavidus Geophilus carpophagus Naf] Effect of urbanization on centipede diversity 713 8 23 24 158 25 154 ssa; qual; 16 – Chiciak 1 19 ––– 6 155 636 45 3756 thropic, WP – West ek (1979, 1980); 8 – dc; ssa ls; 2005 2004 2005 sniewska et al. (2005); 24 sfl ls ssa dc;pf 21 22 23 24 Total 313 705 930 2010 21167 1997 2003 2003 1988 qual; qual; quan; quan; 0 12 12 15 21 37 4 14 35 104 2 ssa; qual; ); 23 – Le´ opean, e – eurytopic, E – European, ltraps. 4 1433 55 11 1 160 13 + 84 116 15 383 1513 2 75 2 + 23 2 19 4 238 – ––––– – –– ssa; qual; nska (unp.); 14 – Kaczmarek (1989); 15 – Zych (1989); 4 – Kaliszak (unp.); 6 – Górecka (unp.); 7 – Kaczmar an, CA – Central Asian, CE – Central Eur rican, NAm – North American, P – Palearctic, s – synan apture; ls – litter-soil samples; pf – pitfal 5 1 1 8 sfl sfl sfl pf sfl sfl sfl sfl sfl dc dc; dc; dc; sfl; dc; dc; dc; dc; dc; ls; dc; 1982 1982 1984 1984 1989 1992 1992 1996 1992 1994 1998 1980 1980 1983 1981 1987 1990 1990 1995 1988 1987 10 11 12 13 14 15 16 17 18 19 20 19 18 20 20 10 1 16 17 13 18 2 ual; qual; qual; qual; quan; qual; qual; qual; qual; qual; quan; 503 310 225 1295 1662 120 501 881 390 – 3105 sniewska (1997); 21 – Drewniak (unp.); 22 – Rzepka (unp. 3 + + qual; 8 + 32 12 19 1 + 12 32 16 1 1 50 1228 63 + 52 5 166 33 37 120 231 nska (unp.); 12 – (unp.); Polańska 13 – Smoli´ 3 + 3 1 + 1 4 3 2 55 sniewska (1996); 20 – Le´ + qual; 50 + ens not given; unp. – unpublished. 11 – Gruszczy´ ); 19 – Le´ 5 wicz (unp.); 3 – Baron (unp.); 4 – Kaczmarek (1977); 5 Holarctic, iNA – introduced to N. Am., Naf – North Af fication of Chilopoda species: Af – African, Au – Australi 1 15 7 4 1 + 1 2 3 4 5 6 7 8 9 ––– – –– – –––––– 27 25 24 16 27 24 31 24 20 dc sfl sfl sfl sfl sfl sfl sfl sfl sfl sfl; dc; dc; dc; dc; dc; dc; dc; dc; dc; 1064 1708 1566 – 549 994 – 1850 486 1963 1965 1975 1971 1976 1978 1980 1979 1981 1981 1949 1964 1974 1968 1975 1977 1949 1978 1980 1980 qual; qual; qual; quan; qual; qual; quan; qual; qual; q itative; ssa – soil samples; sfl – sifting litter; dc – direct c Lohman- Latzel, 1880 [f, E] Newport, 1845 [e, s, E] 3 9 8 16 L. Koch, 1862 [f, E] 33 64 199 + 1 Meinert, 1872 [f, E] 1 1 1 1 Haase, 1880 [f, E] C.L. Koch, 1847 [f, E] 5 8 Meinert, 1868 [e, s, E] 4 12 5 + 14 L. Koch, 1862 [f, CE] 7 33 5 + Reference \ sniewska (unp.); + species present, number of specim EA – Eurasian, ES – Euro-Siberian, f – forest, H – Table 1. (continued) Species Lithobius melanops Number of specimens Sample type Explanations. Ecological and zoogeographical classi Lithobius microps Lithobius muticus Lithobius pelidnus Number of species Palearctic. References: 1 – Kaczmarek (1952, 1964); 2 – Walero Sample type: qual – qualitative; quan – quant Lithobius nodulipes Research period Lithobius mutabilis Lithobius piceus Lithobius tenebrosus fennoscandius der, 1948 [f, E] Lithobius tricuspis Ciszak (unp.); 9 – Cieplicka (unp.); 10 – Wilkosz (unp.); (unp.); 17 – Wojcie–Le´ chowski (unp.); 18 – Gonerka (unp. 714 M. Le´sniewska et al.

Table 2. Characteristics of the quantitative samples used in the study.

Reference Kaczmarek (1977) Kaczmarek (1989) Le´sniewska (1997) Le´sniewska et al. Le´sniewska (unp.) Le´sniewska (2005) (unp.)

Site W˛agrowiec Jakubowo Buczyna Lutomska DusznikiLopuchówko Pozna´n

Habitats oak-wood, alder- oak-hornbeam I, II marshy forest, oak-hornbeam III, oak-pine forest park wood beech forest IV

Sample type litter-soil sifting litter 1988–1992 sifting litter-soil litter-soil litter-soil litter-soil 1993–1994 litter- soil

Surface area of 1/20 m2 1/20 m2 1/16 m2 1/ 16 m2 1/16 m2 1/16 m2 square frame

Collection date 1968: May, July, 1981: each month 1988: May (only each month from 2000: November, 1996: October October from May to Octo- beech forest), July, May 2003 December ber August, September 1969: May, July, 1982: each month 1989: May (2x), to April 2004 2001: each month 2003: May October from May to Octo- June, September from March ber 1971: May, June, 1983: each month 1990: May, July, to October 2004: May September, Octo- from May to Octo- September ber ber 1984: each month 1991: May, July 2005: each from April to Oc- month tober 1992: June, Au- from January gust, October to December each month from April 1993 to June 1994

Surface area per 1m2 at each 1m2 at each 1m2 at each 1/2 m2 at each 1m2 1m2 sample habitat habitat habitat habitat

Total surface area 20 m2 49 m2 61 m2 12 m2 10 m2 15 m2 of samples centipedes live was the most frequently used qualitative species and subspecies of centipedes have been con- method. Other qualitative methods included pitfall traps, firmed. This represents 100% of the species and sub- sifting of litter and soil, and sampling of litter and soil. The species found in lowland areas in Poland (Kaczmarek most frequently used quantitative methods were sifting lit- 1979, 1980; Wytwer 1990; Le´sniewska & Wojciechowski ter and sampling of soil using square frames. However, not 1992) and 61% of the species found in the whole country all researchers used these techniques. It was therefore pos- (Table 1). sible to compare quantitative data only from published reports or reports ready for publication. Not all data could be Among the identified species there were 22 species included in the database. The only built-up area in which from the order Lithobiomorpha, 14 from the order quantitative studies were carried out was Poznań. Detailed Geophilomorpha, and one from the order Scolopendro- information on the quantitative methods used in this study morpha (Table 1). is presented in Table 2. The specimens collected during the course of the The data include information on 21,167 specimens study most often included the following species: which had been identified to species level. The data also in- – eurytropic: Lithobius forficatus, L. erythrocepha- clude information gathered from publications which did not lus, Geophilus flavus, Schendyla nemorensis; specify the number of specimens of each species (Table 1). –forest:Lithobius mutabilis; Data were ranked and analyzed using standard bio- –synanthropic:Lithobius microps. cenotic indexes, including: dominance (D), frequency (F ), Shannon index of diversity (H’), Pielou index of evenness These species were probably the most common and (J ), Morisita-Horn index of similarity, and cluster analy- most numerous species in the investigated area. On the sis. These indices and their significance have been described other hand, only a few specimens of Lithobius dentatus, elsewhere (Balogh 1958; Horn 1966; Trojan 1994; Magurran L. nodulipes and Clinopodes flavidus were collected. 1996; Everitt et al. 2001). In the investigated forests thirty-three species of centipede were found. The number of species found per Results individual forest type ranged from seven to nineteen (Table 3). Eleven species were found only in forests. In the Wielkopolska-Kujawy Lowlands, thirty-seven Three species were found in all of the different for- Effect of urbanization on centipede diversity 715

Table 3. Occurrence of centipedes in forests and built-up areas in Wielkopolska and in Warsaw (data for Warsaw are taken from Wytwer 1995).

Forests Built-up areas of Wielkopolska Environment Alder Beech Marshy Oak- Oak Oak- Unspe- Total in Lesz- Poz- Pyz- Srem´ Tu- Zielona Total in War- No. Species horn- pine ciﬁed forests no na´n dry rek Góra towns saw beam

1 C. hortensis ++ + + + + + + 2 C. flavidus ++ 3 G. carpophagus ++ 4 G. electricus +++++++ 5 G. flavus +++ + +++++++++ 6 G. insculptus ++++ 7 G. linearis +++ 8 G. proximus ++ + + 9 G. truncorum ++ + + + + ++ + + 10 P. ferrugineum ++ +++ + 11 S. furcidens ++ 12 S. nemorensis +++ +++ +++++++ 13 S. subterranea ++ 14 S. acuminata ++ + ++++ + + + + 15 S. crassipes ++++++++ 16 L. emarginatus ++ + + +++ 17 L. aeruginosus ++ 18 L. agilis ++ + ++ + + + + + 19 L. borealis +++ +++ 20 L. calcaratus +++++ 21 L. crassipes ++ + + + +++ + + + 22 L. curtipes ++ + ++ + ++ + 23 L. cyrtopus ++ 24 L. dentatus ++ 25 L. erythrocephalus ++ + ++ + + ++++++ + 26 L. forficatus ++ + ++++ + ++++++ + + 27 L. lapidicola ++ + ++ + + + + 28 L. macilentus +++ + 29 L. melanops ++ +++++ + ++ 30 L. microps ++ + ++++++ + + 31 L. mutabilis ++ + ++++ + + + + 32 L. muticus ++ + 33 L. nodulipes ++ 34 L. pelidnus ++ + + 35 L. piceus ++++ + 36 L. t. fennoscandius ++ + + + + + 37 L. tricuspis +++ +

Number of species 11 19 18 15 12 7 19 33 12 21 9 12 12 11 26 12

est types investigated: Strigamia acuminata, L. muta- were urban parks (21 species), beech forests (19 bilis and L. forficatus. Ten of the species found in the species), marshy forests (18 species), cemeteries (16 forests belonged to the Geophilomorpha, twenty-two to species), and gardens (12 species) (Tables 3, 5). The the Lithobiomorpha and one to the Scolopendromor- habitats with the smallest number of species were pha. houses, rubbish heaps, flowerbeds and marginal waste- In the investigated built-up areas 26 species of cen- lands. These have few microhabitats suitable for centipede were found (Table 3). In each of the locations tipedes. between nine and 21 different species were found. Four Twenty-two species were found in both forests and species were found in all of the six investigated built- built-up areas. However, the centipede communities in up areas: L. forficatus, G. flavus, L. erythrocephalus, forests and built-up areas were distinctly different from and L. microps. Four species of centipedes were found each other (Fig. 1). exclusively in built-up areas: C. flavidus, Geophilus Species diversity as estimated on the basis of the linearis, G. proximus and Stigmatogaster subterranea. Shannon index was the highest in beech forests and Three species were found only in Poznań: S. subter- marshy forests, which both had a Shannon index of ranea, C. flavidus and G. linearis. 2.4 (Table 4). In the rest of the examined habitats the Twelve of the species found in cities belonged to Shannon index ranged from 1.4 to 1.6. The Pielou index the Geophilomorpha, thirteen to the Lithobiomorpha, ranged from 56.5 to 77.8%. However, these data apply and one to the Scolopendromorpha (Table 1, Fig. 3). only to those locations in which quantitative studies The habitats with the highest number of species were carried out (Table 4). 716 M. Le´sniewska et al.

(1) Zielona Góra

(2) Śrem (3) Pyzdry

(4) Poznań (5) Leszno

(6) Turek (A) marshy

(B) beech (C) oak-hornbeam

(D) oak-pine (E) alder (F) oak

1 0

Fig. 1. Species composition similarity (distance measure: Sokal and Mitchener; cluster method: Lance and Williams). (1) – (6): built-up areas; (A) – (F): forests.

beech forest

marshy forest

oak-hornbeam III

oak-hornbeam II

oak-hornbeam I

oak-hornbeam IV

alder-wood

oak-wood

oak-pine

park

1.1 0.0

Fig. 2. Dendrogram of the mutual similarities of dominance structures based on Morisita-Horn index (cluster method: nearest neighbor).

In the tested forests, the super-dominant species aims and methods of their research differed from study were Lithobius curtipes L. mutabilis,andSchendyla to study. That is why our paper is a presentation of the nemorensis. In the Citadel Park in Poznańthesuper- results of previous studies and should not be considered dominant species were L. microps and Stigmatogaster to be strictly comparative in nature. subterranea. Other eu-dominant species included L. agi- The majority of researchers collected materials lis, L. forficatus, G. flavus,andL. pelidnus (Table 4). using qualitative methods (Table 1). The results of Based on Morisita-Horn indices, similar habitats their studies were used only to determine the species generally had similar dominance structures (Fig. 2). composition in various habitats (Tables 3, 5). On the However, geographical location also played a role. other hand, the data for quantitative comparison were The species with the highest coefficients of domi- taken from various previously published and unpub- nance were generally the species which were most fre- lished sources. The methods used in these studies are quently encountered at the studied locations (Table 4). presented in Table 2. The results may have been af- Quantitative samples taken from litter and soil do not fected by the methods used, which differed in terms of give the frequency of species associated primarily with the kinds of samples collected. For example, some sam- tree stumps, logs, stones, moss and bark. In our re- ples were sifted, whereas others were not. Nevertheless, search, this is especially true for the cosmopolitan and the data set as a whole is good enough to support a few eurytopic species L. forficatus. Although L. tenebrosus conclusions and hypotheses. fennoscandius is also very common, it is clearly associ- In the Wielopolska- Kujawy Lowlands, the great- ated with tree stumps and logs, and is thus underesti- est richness of centipede species was found in Poznań. matedinquantitativesamples(Le´sniewska unpublished This finding supports the opinion expressed by En- data). ghoff (1973) that human activity does not always re- duce species diversity in animal communities. Discussion During the post-glacial sub-Atlantic Baltic period, the primeval flora of the Poznań area was dominated The materials used in this study were collected by many by mesophilic deciduous forests. The dominant forest researchers over a period of more than fifty years. The type was the oak-hornbeam forest type (Galio silvatici- Effect of urbanization on centipede diversity 717

Table 4. Dominance (D %), Frequency (F %), Shannon index of diversity (H’, Hmax) and Pielou index of evenness (J %) for selected habitats in Wielkopolska.

Buczyna Puszcza Species\Site W˛agrowiec Jakubowo Lutomska Duszniki Zielonka Pozna´n Reference (Kaczmarek (Kaczmarek (Le´sniewska (Le´sniewska (Le´sniew- (Le´sniew- 1977) 1989) 1997) at al. 2005) ska, unp.) ska, unp.)

Type of forest Oak- Alder- Oak-horn- Oak-horn- Marshy Beech Oak-horn- Oak-horn- Oak- Citadel wood wood beam I beam II forest forest beam III beam IV pine park D% F% D% F% D% F% D% F% D% F% D% F% D% F% D% F% D% F% D% F%

1 C. hortensis 1.1 1.9 1.5 3.5 6.4 9.8 2 C. flavidus 3 G. carpophagus 4 G. electricus 0.7 1.0 5 G. flavus 2.3 17.1 4.3 22.0 19.0 28.3 13.1 24.8 2.2 3.1 6.6 23.2 6 G. insculptus 1.5 1.4 6.6 22.3 7 G. linearis 8 G. proximus 9 G. truncorum 4.0 10.0 1.0 1.3 1.2 1.5 0.2 0.9 10 P. ferrugineum 11 S. furcidens 25.0 0.5 4.9 12 S. nemorensis 1.3 4.9 1.4 4.9 13.9 17.7 10.5 17.9 5.2 7.3 18.2 13.9 47.4 37.5 7.0 30.4 13 S. subterranea 35.0 60.7 14 S. acuminata 1.9 6.7 4.0 15.0 2.1 14.6 2.5 22.0 4.5 9.6 3.2 7.7 3.7 5.2 3.0 2.8 7.0 14.1 15 S. crassipes 0.1 0.4 0.2 0.9 16 L. emarginatus 17 L. aeruginosus 18 L. agilis 5.1 20.0 11.2 35.0 1.1 7.3 0.7 4.9 3.4 6.3 3.0 7.3 2.2 3.1 19 L. borealis 1.9 3.1 1.8 3.3 6.7 9.4 1.5 1.4 2.2 7.8 20 L. calcaratus 0.6 5.0 21 L. crassipes 3.0 10.0 1.0 2.3 3.2 6.5 1.5 1.4 22 L. curtipes 53.2 90.0 51.5 80.0 29.5 82.9 22.4 75.6 0.1 0.2 1.1 2.5 11.9 13.5 4.5 4.2 23 L. cyrtopus 24 L. dentatus 25 L. erythrocephalus 3.2 20.0 4.0 15.0 0.6 4.9 0.9 7.3 0.4 0.8 0.1 0.4 3.0 4.2 26 L. forficatus 3.8 20.0 1.0 5.0 1.0 7.3 0.7 7.3 1.5 3.1 2.0 4.8 2.2 3.1 12.7 12.5 2.6 12.5 27 L. lapidicola 1.9 10.0 3.0 5.0 0.4 0.6 0.1 0.2 1.8 1.6 28 L. macilentus 29 L. melanops 0.8 1.9 0.9 1.9 30 L. microps 0.6 5.0 22.7 11.1 35.2 60.7 31 L. mutabilis 17.7 50.0 5.1 25.0 43.6 82.0 44.4 87.8 49.4 39.8 55.6 48.1 60.0 72.9 45.5 34.7 18.0 21.9 32 L. muticus 0.6 5.0 0.3 0.6 33 L. nodulipes 34 L. pelidnus 1.0 5.0 0.1 0.4 11.0 15.6 35 L. piceus 4.4 20.0 1.1 7.3 0.8 1.9 1.7 3.8 1.5 2.1 1.6 1.4 36 L. t. fennoscandius 1.0 5.0 0.3 0.4 0.1 0.2 37 L. tricuspis 0.3 0.6 0.7 1.9 0.7 1.0 Lithobius sp. 7.0 11.2 30.0 18.5 48.9 21.1 56.1

H’ 1.6 1.6 1.4 1.5 2.4 2.4 1.5 1.5 1.5 1.6 H max 2.5 2.5 2.2 2.4 4.2 4.3 2.5 2.2 1.9 2.2 J % 64.1 64.1 63.5 62.3 57.7 56.5 60.0 68.2 77.8 72.0

Carpinetum). Smaller areas were covered by other de- park in the city), L. curtipes and S. acuminata. ciduous forest types (Jackowiak 1990). The pre-urban In smaller built-up areas, a greater number of for- centipede fauna of the Poznań area was therefore sim- est species were found, including Lithobius agilis, L. ilar to the fauna of present-day deciduous forests in lapidicola and L. mutabilis (Table 3). This is because Wielkopolska. As the city grew, the forest cover was the forested areas at these locations have not been de- reduced or destroyed to be replaced by urban green graded to the degree that the forested areas in more spaces. urbanized locations have been. Of the 21 centipede species found in Poznań, twelve The town and its vicinity provide for different cen- were also found in beech forests, and ten were also found tipede species a great diversity of micro-habitats such in oak-hornbeam forests in the region. Four species as forested areas, parks, cemeteries, allotment gardens, found in Poznań were also found in all the deciduous rubbish heaps, roadsides, flowerbeds, marginal waste- forests studied: Strigamia acuminata, Lithobius cur- lands, greenhouses, and buildings (Table 5). The cen- tipes, L. erythrocephalus and L. forficatus.Theonly tipede urban fauna, especially of Poznań, is character- typical forest dwellers found in PoznańwereL. tene- ized by a high number of species belonging to the or- brosus fennoscandius (which occurs only in the oldest der Geophilomorpha and small species of Lithobiomor- 718 M. Le´sniewska et al.

Lithobiomorpha Geophilomorpha 100% 2 2 4 5 2 5 6 3 3 4 5 6 12 10 14 12

80% 10 9 13 9 13 5 10 12 6 22 60% 22 6 7 13 40% 5 8

20%

0% ń rem Total Ś Turek Pyzdry Leszno Pozna oak forest forest alder forest alder Total towns Total forests beech forest Zielona Góra marshy forest oak-hornbeam oak-pine forest unspecified forest

Fig. 3. Percentage participation of geophilomorph and lithobiomorph species in the studied habitats. 2–22 – number of species.

Table 5. Occurrence of selected centipede species in diﬀerent urban habitats in Wielkopolska.

Habitat\Species C. flavidus G. proximus S. subterranea L. calcaratus L.lapidicola L. t. fennoscandius L. mutabilis G. linearis S. acuminata L. agilis L. curtipes L. macilentus G. insculptus G. truncorum S. crassipes P. ferrugineum L. crassipes G. electricus C. hortensis L. erythrocephalus S. nemorensis L. microps L. melanops L. emarginatus G. flavus L. forficatus Total

Balcony +1 Flat/apartment +1 Pavement/sidewalk +1 Rubbish/trush +++3 Cellar ++ + 3 Wasteland ++ ++4 Greenhouse ++++4 Flowerbed ++++4 Ditch ++ + + + 5 Railway embankment + + + + + + 6 Outbuildings + + + + + + 6 Roadside + + + + + + + 7 River bank + + + + + + + + 8 Lakeshore + + +++++++ + +11 Garden ++ +++ + ++++ + +12 Cemetery + + ++++ ++++++++ + +16 Park ++ +++ ++++++ ++++++++ + +21

Total 11111112222234444666889101113

pha, which are generally more often found in soil than and a lower percentage of Geophilomorpha (Table 3, in litter. Because leaves, old trees, logs and stumps Fig. 3). are usually removed from urban areas, larger Litho- In urban areas, only some of the indigenous species biomorpha are found only in small forested areas on become part of the fauna. Other species are introduced the outskirts of the city, or in a few untended ar- in various ways. Human activity can allow new species eas of old cemeteries and parks. On the other hand, to establish themselves. For example, some species are forests, especially protected ones, with thick leaf lit- introduced with garden and landscaping plants and ter and abundant fallen trees, stumps and branches, with soil used in gardening, landscaping and construc- provide ideal conditions for centipedes of the order tion. This is probably how Stigmatogaster subterranea Lithobiomorpha. Although the number of species may made its way to Poznań. Lamyctes emarginatus is fre- be about the same as that in the city, there is a quently encountered in commercial garden soils of- much higher percentage of species of Lithobiomorpha, fered for sale in shops with garden supply depart- Effect of urbanization on centipede diversity 719 ments. Lamyctes emarginatus is a parthenogenetic pio- Kaczmarek J. 1980. Pareczniki (Chilopoda). Katalog Fauny Pol- neer species (Dunger & Voigtländer 1990; Voženílková ski. PWN, Warszawa, 43 pp. & Tajovský 2001) which easily establishes itself at Kaczmarek J. 1989. Pareczniki (Chilopoda) wybranego lasu gr˛adowego Wielkopolski na przykladzie rezerwatu Jakubowo. marginal sites in Wielkopolska (Le´sniewska 2004). Fragm. Faun. 32 (17): 369–379. In another study (Wytwer 1995), 12 species of cen- Kondracki J. 2002. Geografia regionalna Polski. PWN, Warszawa, tipedes were identified in Warsaw whereas 17 species 441 pp. Le´sniewska M. 1996. Centipedes of Poznań town (Poland), pp. were identified in forest ecosystems in the surrounding 221–224. In: Geoffroy J.-J., Mauriès J.-P. & Nguyen Duy – region of Mazovia (Table 3). There were also consid- Jacquemin M. (eds), Acta Myriapodologica. Mém. Mus. Nat. erable differences in species composition and commu- Hist. Nat. 169. nity structure between the centipede fauna of Warsaw Le´sniewska M. 1997. Zgrupowanie pareczników (Chilopoda) w rezerwacie przyrody “Buki nad jeziorem Lutomskim”. Wyd. and the surrounding forest ecosystems. Lithobius tene- Nauk. UAM, Poznań, 83 pp. brosus tenebrosus and L. tenebrosus fennoscandius had Le´sniewska M. 2004. Lamyctes emarginatus (Newport, 1844) w both been reported to occur in Wielkopolska (Kacz- Wielkopolsce. Bad. Fizj. Pol. Zach. Seria C – Zoologia 50: marek 1952, 1977; Le´sniewska 1997). However, more 45–51. Le´sniewska M., Koralewska-Batura E. & Bloszyk J. 2005. 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