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Species distribution and assemblages of centipedes
(Chilopoda) in open xeric sites of Saxony-Anhalt
(Germany).
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| Abstract: | As part of a large-scale research program investigating endangered biotopes in the state of SaxonyAnhalt in eastern Germany, the centipede fauna of 50 sites, belonging to five types of xeric biotopes, was studied by pitfall trapping from 1995 to 1998 (each site for one year). The most abundant species was Lithobius calcaratus. By applying a combination of the criteria dominance, constancy and representativity, 'characteristic species' and 'companion species' were defined. For seven of the 20 recorded centipede species, it was possible to delineate biotope-characteristic assemblages. Mesoxeric meadows are characterised by L. calcaratus and L. forficatus, mesoxeric meadows with shrub and pioneer forest (advanced succession sites) by L. crassipes, L. mutabilis and L. forficatus and Schendyla nemorensis. In grassland contaminated by heavy metals, a typical group consisting of L. melanops and L. forficatus with Cryptops parisi was found. Dwarf-shrub heaths (L. calcaratus) and sandy xeric-meadows (Lamyctes emarginatus and Lithobius e. erythrocephalus) were only characterised by companion species. The results show that some centipede species and characteristic species assemblages are very good indicators for minor habitat differences. |
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| Subject: | Grasslands |
| Author: | Voigtlander, Karin |
| Pub Date: | 08/01/2003 |
| Publication: | Name: African Invertebrates Publisher: The Council of Natal Museum Audience: Academic Format: Magazine/Journal Subject: Zoology and wildlife conservation Copyright: COPYRIGHT 2003 The Council of Natal Museum ISSN: 1681-5556 |
| Issue: | Date: August, 2003 Source Volume: 44 Source Issue: 1 |
| Accession Number: | 204859791 |
| Full Text: |
INTRODUCTION Centipedes are known to be useful indicators for specific abiotic site conditions in Middle Europe (Frund 1996; VoigtlAnder & Dunger 1998; Spelda 1999). The value of certain species as bioindicators or 'characteristic species', respectively, depends on knowledge of their ecofaunistical behaviour, bionomical strategy and phenology. Investigations of centipedes in open xeric habitats provided an opportunity to increase our knowledge of their habitat preferences and fidelity to special site-types. This study of centipedes is part of a large-scale research program investigating endangered biotopes (defined by the biotopes 'Red List') in the state of SaxonyAnhalt, eastern Germany. Compared with investigations that only utilise a few arthropod groups, e.g. Carabidae or Saltatoria to check the necessity of landscape protection, including centipedes offers the possibility of a more comprehensive site characterisation. The present study aims to answer the following questions: Which centipede species inhabit open xeric habitats in Saxony-Anhalt? Which centipede species are characteristic of specific xeric site-types? Do centipedes form characteristic species assemblages on different xeric sites? Characteristic species are usually recognised on the basis of various criteria such as presence, dominance, constancy, frequency, preference or representativity. However, different criteria can give different results. A combination of dominance, constancy and representativity is used here to define characteristic species or species groups. SITES, METHODS AND DEFINITIONS The investigations were concentrated on sites at the Altmark with the Elbe Valley in the northern part of Saxony-Anhalt, the Nordharzvorland and Sudharz (south-west of the country) and the Unstrut-Trias-Land in the southern part (Fig. 1). The 50 sites investigated comprised: 11 XM: The xeric meadows were mostly concentrated in the northern part of SaxonyAnhalt. They were divided into sites with two different plant communities: * pioneer sandy xerix meadows--Corynephorion canescentis Klika 1934 with the association Spergulo morisonii-Corynephoretum canescentris (R.Tx. 1928) Libb. 1933 on dry and warm, nutrient-poor inland dunes with loose Pleistocene sand. The herb cover was very small (5-60 %) with a more or less dense cover of lichens and mosses (50-85 %). Without exposition. Altitude: 30-170 m. * persistent sandy xerix meadows--Armerion elongatae Krausch, 1961 on sandy, dry soils with a more or less dense sod. The herb cover was 80 % and the moss layer 20 %. Aspect: S and SW. Altitude: 50-70 m. 22 SMM: submediterranean mesoxeric sites with Mesobromion erecti (Br.Bl. et Moor 1938) R. Knapp 1942 ex Oberd. 1957 on limestone ('Muschelkalk') and loess sites with continental mesoxeric meadows (Cirsio-Brachypodion Mahn 1959 emend.). Associations: Festuco rupicolae-Brachypodietum pinnati Mahn 1959 emend.; Gentiano-Koelerietum pyramidatae Knapp 1942 ex. Bornk. 1960. The sites had a relatively closed vegetation cover (herb layer 20-95 %) with a more or less balanced water and heat budget. They were concentrated in the higher south-western part of Saxony-Anhalt between altitudes 160 to 480 m, and with S, SW, SSW and SSO aspects. 4 HMG: grasslands contaminated by heavy metals on copper-slate seams with low proportions of fine soil and spoil dumps originating from copper-slate mining at the surroundings of Eisleben and Ilsenburg. The high content of copper, zinc and lead compounds only allowed a Armerietum halleri Libb. 1930 plant association to exist there. The sites were subjected to extremely high solar radiation (aspect: SW) and drying up. The herb cover differed widely between the sites (5-80 %). Altitude: 210-265 m. 8 DSH: subatlantic dwarf-shrub heaths (Genistion Boch. 1943) of: * the association Genisto pilosae-Callunetum R.Tx. 1937 on level sites (45-90 m) without exposition on nutrient-poor, sandy soils (podsol) in the northern part of Saxony-Anhalt. * the association Euphorbio-Callunetum on shallow, acid soils caused by weathering of sandstone, porphyry or gipsum at higher sites (altitude160-340 m) with a N and WSW aspect in the Harz region. 5 ASS: xeric shrub societies (Berberidion Br.Bl. 1950 with the association LigustroPrunetum spinosae) and pioneer forests tending towards the association Potentillo albae-Quercetum petraeae Jakucs 1967 on loamy-mary soils (advanced succession sites). Altitude 160-210m with a S aspect. A detailed description of the investigated sites will be given in: Landesamt fur Umweltschutz Sachsen-Anhalt (ed.) (in prep.). The results are based on pitfall trap catches during the years 1995 to 1998. Per site, six traps (diameter 6.5 cm) were arranged in a single row, with a distance of 8 to 10 m between traps. They were filled with 3 % formalin and were cleared monthly. Each site was investigated for one uninterrupted year. For standardisation purposes, the results were based on individuals per trap and week. Dominance and constancy give little evidence of the main occurrence (e.g. the site with the highest density) of a species along a factor gradient (e.g. temperature, moisture, exposure) of different biotope-types. For preferences within the study sites, the 'representativity' (based on biomass, abundance or activity abundance), as defined by Dunger (1978) and Muller et al. (1978) is a suitable measure. The representativity of a species is the percentage of individuals trapped in a particular site-type, relative to the number trapped in all site-types under study, where numbers are per week and trap. A 'characteristic species' for a specific site-type, as here defined, fulfils the following three criteria: dominance: eudominant (>32 %) or dominant (10.0-31.9 %) (according to Engelmann 1978); constancy: present in more than 50 % of the sites of this type; representativity: maximum representativity on sites of this type. A 'companion species' for a specific site-type fits only two of these criteria. Following Braun-Blanquet (1951), a 'characteristic species assemblage' is composed of species with more than 50 % common occurrence on sites of a specific site-type. RESULTS General results From the 50 sites under study, approximately 1200 centipedes from 20 species were recorded (Table 1). The most abundant species was Lithobius calcaratus (n = 423), followed by L. forficatus (n = 237) and L. melanops (n = 130). Most of the other species were more or less rare and were sometimes concentrated in only a single site-type. Table 1 presents the results with dominance, constancy and representativity highlighted. Characteristic and companion species Characteristic species are typical for only three site-types: L. forficatus and L. melanops for the contaminated grasslands (HMG), L. mutabilis and L. crassipes for the advanced successional sites (ASS), and L. calcaratus for the mesoxeric meadows (SMM) (Table 2). Only L. calcaratus was found as a companion species in the dwarf shrub heath (DSH) and Cryptops parisi for the HMG site. Lamyctes emarginatus and Lithobius e. erythrocephalus were recorded as companion species in the xeric meadows on sand (XM). Thus, the ASS sites were highly characterised by three companion species (L. forficatus, Schendyla nemorensis, Strigamia crassipes), and the SMM sites by two companion species (L. forficatus, L. microps). Characteristic species assemblages Characteristic species assemblages were established for the advanced successional sites (ASS): L. mutabilis and L. crassipes, as characteristic species, were assembled with L. forficatus, St. crassipes and Sch. nemorensis in 60 % of the ASS sites. The common occurrence of L. melanops and L. forficatus (as characteristic species) in 75 % of the contaminated grassland (HMG) was noticeable. Additionally, an assemblage could be characterised in 50 % of the HMG sites by the occurrence of L. forficatus and L. melanops together with C. parisi (as a companion species). Similarly less differentiated was the assemblage found in the mesoxeric meadows (SMM): L. calcaratus as a characteristic species together with L. forficatus and L. microps as companion species (54 %). For the other studied site-types, such assemblages were not found. DISCUSSION The establishment of characteristic species for the different types of open and xeric habitats is supported by knowledge of their autecology. The most frequent species L. calcaratus is generally described as an open-land species, which prefers warm sites independent of their moisture content; i.e. thermophilous behaviour is the deciding factor in habitat choice (VoigtlAnder 1995; VoigtlAnder & Dunger 1998). In the present investigation, L. calcaratus was found eudominantly in the sandy xeric and mesoxeric meadows (here as a characteristic species) and dwarf-shrub heaths. The shadier shrubs (ASS) and the copper-contaminated sites (HMG) were avoided. Lamyctes emarginatus is a typical species for poorly vegetated sites. This member of Henicopidae is known to be a successful pioneer species in spoil dumps, freshly created wine-growing areas, as well as flooded areas (Dunger & VoigtlAnder 1990 2003; VoigtlAnder et al. 2001; Zulka 1991; Armbruster 1992; Zerm 1997). Its parthenogenetic life cycle predestines this species for a quick (re-)settlement of (disturbed) open lands. Correspondingly, in the present investigation this species was found, with the exception of only a few records in contaminated grassland, exclusively in xeric meadows. For L. erythrocephalus, the subspecies L. e. erythrocephalus and L. e. schuleri have been described, and the elevation of these subspecies to species status is under discussion. Besides different morphological characters, it seems that fundamental ecological differences between these forms exist as well. Lithobius e. erythrocephalus mostly occurs in open, warm habitats at lower altitudes, whereas L. e. schuleri is found in humid mountain woods to alpine meadows and stony open fields (Attems 1954; Minelli & Iovane 1987; Moser 1999; Spelda 1999). All individuals from Saxony-Anhalt belong to L. e. erythrocephalus and were found to be especially frequent on the driest sites (xeric meadows). Lithobius forficatus and L. melanops were established as characteristic species for the copper-contaminated grasslands (HMG). Both species evidently occur there in stable populations, as there are records of juveniles in different stages. Among all Lithobiomorpha, L. forficatus is probably the species with the broadest ecological tolerance, and was found at all investigated sites. Lithobius melanops has been described as a woodland species (Eason 1964) and as a thermophilous species with high affinities for synanthropic sites (e.g. Jeekel 1964; Dunger & VoigtlAnder 1990; Zapparoli 1990; Spelda 1999; VoigtlAnder 1999). Its occurrence at the HMG sites supports the opinion of the majority of authors. Cryptops parisi is mostly classified as a thermophilous species and an inhabitant of mesophilic woods, meadows and suburban areas (e.g. Eason 1964; Peter 1984; Minelli & Iovane 1987). This is also the case in the present investigation. Additionally, C. parisi preferred the mesoxeric meadows along a moisture gradient on a south-exposed limestone-slope ('Muschelkalk') in the Leutra Valley/Thuringia (VoigtlAnder & Dunger 1998). Other authors claim a high cold-resistance for C. parisi (Koren 1986; Spelda 1999). Lithobius forficatus, L. melanops and C. parisi occur in areas in Thuringia strongly effected by emissions (dust, Na, F and P), on sites with the highest levels of contamination (Peter 1984). This finding is in agreement with the present investigation, in which these species were not only shown to tolerate warm-dry microclimatic conditions, but also high contamination with copper on HMG sites. According to the literature, L. crassipes settles in various habitats and is even able to tolerate dryness and high solar radiation (Matic et al. 1979), but the species actually mostly prefers woodlands or sparse woods and shrublands (e.g. VoigtlAnder 1983; Wytwer 1992; Vossel & Assmann 1995; Spelda 1999; VoigtlAnder et al. 2001), as is also shown by its occurrence in the present study on ASS sites. Lithobius mutabilis is a woodland inhabitant and one of the most common species in Central European deciduous (beech) woods (e.g. Albert 1976 1982; Wytwer 1996; Frund 1991; Frund et al. 1997; VoigtlAnder & Dunger 1998; Spelda 1999). This species prefers cooler and moister habitats with a well-developed vegetation layer, which was reflected in its high abundance in the advanced successional sites (ASS) investigated here. In summary, it may be said that the distributions of the species investigated within the different site-types of the present study are closely in accordance with published data on their ecological behaviour. This can be clearly seen for the woodland species L. mutabilis and L. crassipes, for the euryoecious open land species L. emarginatus and L. calcaratus, as well as for the euryoecious species L. forficatus, L. melanops and C. parisi. The study shows that centipedes can be well utilised for site classification. They clearly indicate the different biotope types and are in accordance with the overall results for the studied sites (Landesamt fur Umweltschutz Sachsen-Anhalt, ed., in prep.). Pitfall trapping proved to be a useful and efficient method for bioindication with centipedes. No doubt, the exclusive application of pitfall trapping restricts the species inventory to active epigeic forms. However, the presented results demonstrate that 'pitfall species' are sufficient for site characterisations of high quality. This can also be seen in other investigations (Frund 1996; VoigtlAnder & Dunger 1998). The method used here for the recognition of characteristic species and species assemblages has been shown to be useful for centipedes as well as for millipedes (VoigtlAnder & Duker 2001). ACKNOWLEDGEMENTS I would like to thank Dr P. Schnitter (Landesamt fur Umweltschutz Sachsen-Anhalt) for his confidence in the study of the myriapods sampled in the project 'Tierokologische Untersuchungen in gefAhrdeten Biotoptypen des Landes Sachsen-Anhalt. I. Zwergstrauchheiden, Trocken- und Halbtrockenrasen'. I am especially indebted to Prof. W. Dunger for reading, commenting on and providing valuable discussions on the manuscript. REFERENCES ALBERT, A. 1976. Biomasse von Chilopoden in einem Buchenaltbestand des Solling. Verhandlungen der Gesellschaft fur Okologie (Gottingen) 1976: 93-101. --1982. Species spectrum and dispersion patterns of Chilopods in Solling habitats. Pedobiologia 23: 337-347. ARMBRUSTER, C. 1992. Wiederbesiedlung und Sukzession bei Chilopoden im flurbereinigten RebgelAnde des Kaiserstuhls. Diplomarbeit Univ. Freiburg im Breichsgau. ATTEMS, O. 1954. Myriopoda. In: Franz, H., ed., Die Nordostalpen im Spiegel ihrer Landtierwelt. Bd. 1: 289-328. Innsbruck: UniversitAtsverlag Wagner. BRAUN-BLANQUET, J. 1951. Pflanzensoziologie. Grundzuge der Vegetationskunde. Wien: Springer. DUNGER, W. 1978. Parameter der Bodenfauna in einer Catena von Rasen-Okosystemen. Pedobiologia 18: 310-340. DUNGER, W. & VOIGTLANDER, K. 1990. Succession of Myriapoda in primary colonisation of reclaimed land. In: Minelli, A., ed., Proceedings of the 7th International Congress of Myriapodology 1987: 219-227. Leiden: E. J. Brill. --2003. Soil-biological development parameters in post-mining landscapes. Journal of Soils and Sediments (in press). EASON, E. H. 1964. Centipedes of the British Isles. London, New York: Frederick Warne & Co Ltd. ENGELMANN H.-D. 1978. Zur Dominanzklassifizierung von Bodenarthropoden. Pedobiologia 18: 378-380. FRUND , H.-C., 1991. Zur Biologie eines Buchenwaldbodens. 14. Die Hundertfusser (Chilopoda). Carolinea 49: 83-94. --1996. Chilopoda. In: Rombke, J., Beck, L., Forster, B., Frund, H.-C., Horak, F. Ruf, A., Rosciczweski, C., Scheurig, M. & Woas, S., Fortfuhrung der Literaturstudie: Bodenfauna und Umwelt. Bd.1. Karlsruhe: Landesanstalt fur Umweltschutz Baden-Wurttemberg pp.113-121. FRUND , H.-C., BALKENHOL, B. & RUSZKOWSKI, B. 1997. Chilopoda in forest habitat-islands in north-west Westphalia, Germany. Entomologica Scandinavica Supplement 51: 107-114. JEEKEL, C. A. W. 1964. Beitrag zur Kenntnis der Systematik und Okologie der Hundertfusser (Chilopoda) Nordwestdeutschlands. Abhandlungen und Verhandlungen des Naturwissenschaftlichen Vereins Hamburg, N.F. 8: 111-153. KOREN, A. 1986. Die Chilopoden-Fauna von KArnten und Osttirol. Teil 1 Geophilomorpha, Scolopendromorpha. CarinthiaII Sonderhefte 43. Landesamt fur Umweltschutz Sachsen-Anhalt (ed.) (in prep.). Tierokologische Untersuchungen in gefAhrdeten Biotoptypen des Landes Sachsen-Anhalt. I. Zwergstrauchheiden, Trocken- und Halbtrockenrasen. Berichte des Landesamtes fur Umweltschutz Sachsen-Anhalt, Sonderheft. MATIC, Z. E., SCHNEIDER, A. & WEISS, I. 1979. Untersuchungen uber die Arthropodenfauna xerothermer Standorte im siebenburgischen Hugelland. VIII. Die Chilopoden eines Sudhanges im Hugelland Siebenburgens. Studii si Comunicari / Stiinte naturale, Muzeul Brukenthal 23: 263-274. MOSER, K. 1999. Vertikalverteilung und Habitatwahl der Steinkriecher im Exkursionsgebiet um Innsbruck (Nordtirol, Osterreich) (Chilopoda, Lithobiomorpha). Berichte des NaturwissenschaftlichMedizinischen Vereins in Innsbruck 86: 213-228. MINELLI, A. & IOVANE, E. 1987. Habitat preferences and taxocoenoses of Italian centipedes (Chilopoda). Bolletino del Museo Civico di Storia Naturale di Venezia 37: 7-43. MULLER H.-J., BAHRMANN, R., HEINRICH, W., MARSTALLER, R., SCHALLER, G. & WITSACK, W. 1978. Zur Strukturanalyse der epigAischen Arthropodenfauna einer Rasen-Katena durch KescherfAnge. Zoologische Jahrbucher, Abteilung fur Systematik, Okologie und Geographie der Tiere 105: 131-184. PETER, H.-U. 1984. Uber den Einfluss von Luftverunreinigungen auf Okosysteme. IV. Isopoda, Diplopoda, Chilopoda und Auchenorrhyncha aus BodenfallenfAngen in der Umgebung eines Dungemittelwerkes. Wissenschaftliche Zeitschrift der Friedrich-Schiller-UniversitAt Jena, naturwissenschaftliche Reihe 33 (3): 291-307. SPELDA, J. 1999. Verbreitungsmuster und Taxonomie der Chilopoda und Diplopoda Sudwestdeutschlands. Diskriminanzanalytische Verfahren zur Trennung von Arten und Unterarten am Beispiel der Gattung Rhymogona Cook, 1896 (Diplopoda: Chordeumatida: Craspedosomatidae). Dissertation, UniversitAt Ulm (Teil II. Abhandlung der einzelnen Arten). VOIGTLANDER, K. 1983. Chilopoden aus FallenfAngen im Waldgebiet Hakel, nordostliches Harzvorland der DDR. Herzynia N. F. (Leipzig) 20: 117-123. --1995. Diplopoden und Chilopoden aus FallenfAngen im Naturschutzgebiet 'Dubringer Moor' (Ostdeutschland/Oberlausitz). Abhandlungen und Berichte des Naturkundemuseums Gorlitz 68 (8): 39-42. --1999. Untersuchungen zur Diplopoden- und Chilopodenfauna des Brockengebietes (Myriapoda: Diplopoda et Chilopoda). Abhandlungen und Berichte fur Naturkunde.Museum fur Naturkunde, Magdeburg 22: 27-38. VOIGTLANDER, K. & DUKER, C. 2001. Distribution and species grouping of millipedes (Myriapoda, Diplopoda) in dry biotopes in Saxony-Anhalt/Eastern Germany. European Journal of Soil Biology 37: 325- 328. VOIGTLANDER, K. & DUNGER, W. 1998. Centipedes of the nature reserve "Leutratal" near Jena (Thuringia, East Germany). In: Pizl, V. & Tajovsky', K, eds, Soil zoological problems in Central Europe. Ceske Budejovice. Ceske Budejovice: Institute of Soil Biology, Academy of Science of the Czech Republik pp. 255-265. VOIGTLANDER, K., KOBEL-LAMPARSKI, A. & LAMPARSKI, F. 2001. Die Chilopodenfauna (Myriapoda) im RebgelAnde des Kaiserstuhls--Einfluss verschiedener Bodenbearbeitungsverfahren. Carolinea 59: 73-80. VOSSEL, E. & ASSMANN, T. 1995. Die Chilopoden, Diplopoden und Carabiden unterschiedlich genutzter WaldflAchen bei Bentheim (Sudwest-Niedersachsen): Vergleich eines Wirtschaftshochwaldes mit zwei ehemaligen HudeflAchen. Drosera 95 (2): 127-143. WYTWER, J. 1992. Chilopoda communities of the fresh pine forest of Poland. Berichte des naturwissenschaftlich-medizinischen Vereins in Innsbruck, Supplement 10: 205-211. --1996. Chilopoda of urban greens in Warsaw. In: Geoffroy, J.-J., Mauries, J.-P. & Nguyen DuyJacquemin, M., eds, Acta Myriapodologica. Memoires du Museum National d' Histoire Naturelle 169: 213-220. ZAPPAROLI, M. 1990. Chilopodi di ambiente urbani e suburbani della citta di Roma. Bolletino, Associazone Romana di Entomologia 44 (1989): 1-12. ZERM, M. 1997. Distribution and phenology of Lamyctes emarginatus and other lithobiomorph centipedes in the floodplain of the lower Oder Valley, Germany (Chilopoda, Henicopidae: Lithobiidae). Entomologica Scandinavica 51: 125-132. ZULKA, K. P. 1991. Uberflutung als okologischer Faktor: Verteilung, PhAnologie und Anpassungen der Diplopoda, Lithobiomorpha und Isopoda in den Flussauen der March. Dissertation, UniversitAt Wien. Karin VoigtlAnder (Staatliches Museum fur Naturkunde Gorlitz, PF 30 01 54, 02806 Gorlitz, Germany; Karin.Voigtlaender@smng.smwk.sachsen.de) TABLE 1.
Results of trapped centipedes from 50 sites (5 site-types)
in Saxony-Anhalt. All figures are percentages; see text for
definitions. d = dominance ((a) eudominant (b) dominant),
c = constancy ((c) >50%), r = representativity ((d) highest r)
XM
sandy xeric
meadows
Species d c r
Lithobius forficatus (Linne, 1758) 8.4 54 (c) 1.9
Lithobius melanops Newport, 1845 0 0 0
Lithobius calcaratus C. L. Koch, 1844 47.0 (a) 45 20.9
Lithobius mutabilis L. Koch, 1862 2.5 18 3.8
Lithobius crassipes L. Koch, 1862 1.7 9 6.2
Schendyla nemorensis
(C. L. Koch, 1837) 5 36 16.7
Strigamia crassipes
(C. L. Koch, 1835) 0 0 0
Lithobius microps Meinert, 1868 0 0 0
Lamyctes emarginatus (Newport, 1844) 19.3 (a) 18 51.5 (d)
Lithobius e. erythrocephalus
C. L. Koch, 1847 14.2 (a) 18 62.5 (d)
Lithobius piceus L. Koch, 1862 0 0 0
Cryptops parisBr6lemann, 1920 0 0 0
Cryptops hortensis Leach, 1814 0 0 0
Lithobius nodulipes Latzel, 1880 0 0 0
Necrophloeophagus flavus
(De Geer, 1778) 0 0 0
Geophilus electricus (Linne, 1758) 0 0 0
Lithobius macilentus L. Koch, 1862 1.7 9 27.3
Lithobius muticus C. L. Koch, 1847 0 0 0
Strigamia acuminata (Leach, 1814) 0 0 0
Lithobius dentatus C. L. Koch, 1844 0 0 0
DSH
dwarf-shrub
heaths
Species d c r
Lithobius forficatus (Linne, 1758) 1.6 12 0.2
Lithobius melanops Newport, 1845 0 0 0.8
Lithobius calcaratus C. L. Koch, 1844 39.1 (a) 62 (c) 10.7
Lithobius mutabilis L. Koch, 1862 3.1 25 3.8
Lithobius crassipes L. Koch, 1862 3.2 12 6.2
Schendyla nemorensis
(C. L. Koch, 1837) 12.5 (a) 50 250
Strigamia crassipes
(C. L. Koch, 1835) 0 0 0
Lithobius microps Meinert, 1868 17.2 (a) 25 20.8
Lamyctes emarginatus (Newport, 1844) 0 0 0
Lithobius e. erythrocephalus
C. L. Koch, 1847 6.3 12 250
Lithobius piceus L. Koch, 1862 0 0 0
Cryptops parisBr6lemann, 1920 0 0 0
Cryptops hortensis Leach, 1814 1.6 0 3.1
Lithobius nodulipes Latzel, 1880 4.7 37 91.7 (d)
Necrophloeophagus flavus
(De Geer, 1778) 0 0 0
Geophilus electricus (Linne, 1758) 0 0 0
Lithobius macilentus L. Koch, 1862 0 0 0
Lithobius muticus C. L. Koch, 1847 0 0 0
Strigamia acuminata (Leach, 1814) 0 0 0
Lithobius dentatus C. L. Koch, 1844 0 0 0
SMM
mesoxeric
meadows
Species d c r
Lithobius forficatus (Linne, 1758) 10.3 (a) 64 (c) 50
Lithobius melanops Newport, 1845 1.1 9 0.8
Lithobius calcaratus C. L. Koch, 1844 62.0 (a) 86 (c) 57.0 (d)
Lithobius mutabilis L. Koch, 1862 1.9 27 3.8
Lithobius crassipes L. Koch, 1862 1.9 27 6.2
Schendyla nemorensis
(C. L. Koch, 1837) 2.1 18 16.7
Strigamia crassipes
(C. L. Koch, 1835) 2.1 27 12.7
Lithobius microps Meinert, 1868 11.8 (a) 32 46.5 (d)
Lamyctes emarginatus (Newport, 1844) 0 0 0
Lithobius e. erythrocephalus
C. L. Koch, 1847 1.3 9 12.5
Lithobius piceus L. Koch, 1862 0.4 9 3.5
Cryptops parisBr6lemann, 1920 0.2 4 50
Cryptops hortensis Leach, 1814 0.4 9 3.1
Lithobius nodulipes Latzel, 1880 0.2 4 8.3
Necrophloeophagus flavus
(De Geer, 1778) 1.3 18 31.3
Geophilus electricus (Linne, 1758) 0.9 14 24.1
Lithobius macilentus L. Koch, 1862 0 0 0
Lithobius muticus C. L. Koch, 1847 0.9 9 100 (d)
Strigamia acuminata (Leach, 1814) 0.7 14 100 (d)
Lithobius dentatus C. L. Koch, 1844 0.2 4 100 (d)
ASS
advanced
successional sites
Species d c r
Lithobius forficatus (Linne, 1758) 30.2 100 (c) 19.5
Lithobius melanops Newport, 1845 0.7 20 0.6
Lithobius calcaratus C. L. Koch, 1844 2.9 20 3.5
Lithobius mutabilis L. Koch, 1862 21.6 100 (c) 84.6 (d)
Lithobius crassipes L. Koch, 1862 12.9 60 (c) 81.3 (d)
Schendyla nemorensis
(C. L. Koch, 1837) 4.3 60 (c) 33 (d)
Strigamia crassipes
(C. L. Koch, 1835) 10.8 40 87.3 (d)
Lithobius microps Meinert, 1868 6.5 80 (c) 32.7
Lamyctes emarginatus (Newport, 1844) 0 0 0
Lithobius e. erythrocephalus
C. L. Koch, 1847 0 0 0
Lithobius piceus L. Koch, 1862 7.20 20 85.9 (d)
Cryptops parisBr6lemann, 1920 0 0 0
Cryptops hortensis Leach, 1814 0 0 0
Lithobius nodulipes Latzel, 1880 0 0 0
Necrophloeophagus flavus
(De Geer, 1778) 2.2 20 68.7 (d)
Geophilus electricus (Linne, 1758) 2.2 40 75.9 (d)
Lithobius macilentus L. Koch, 1862 0.7 20 31.8
Lithobius muticus C. L. Koch, 1847 0 0 0
Strigamia acuminata (Leach, 1814) 0 0 0
Lithobius dentatus C. L. Koch, 1844 0 10 0
HMG
contaminated
grasslands (a)
Species d c r
Lithobius forficatus (Linne, 1758) 42.9 (a) 75 (c) 73.4 (d)
Lithobius melanops Newport, 1845 40.9 (a) 75 (c) 98.6 (d)
Lithobius calcaratus C. L. Koch, 1844 2.7 50 8.1
Lithobius mutabilis L. Koch, 1862 0.3 25 3.8
Lithobius crassipes L. Koch, 1862 0 0 0
Schendyla nemorensis
(C. L. Koch, 1837) 0.3 25 8.3
Strigamia crassipes
(C. L. Koch, 1835) 0 0 0
Lithobius microps Meinert, 1868 0 0 0
Lamyctes emarginatus (Newport, 1844) 2.7 25 48.5
Lithobius e. erythrocephalus
C. L. Koch, 1847 0 0 25,0
Lithobius piceus L. Koch, 1862 0.3 25 10.6
Cryptops parisBr6lemann, 1920 6.6 75 (c) 95.2 (d)
Cryptops hortensis Leach, 1814 3.3 25 93.8 (d)
Lithobius nodulipes Latzel, 1880 0 0 0
Necrophloeophagus flavus
(De Geer, 1778) 0 0 0
Geophilus electricus (Linne, 1758) 0 0 0
Lithobius macilentus L. Koch, 1862 0.3 25 40.9 (d)
Lithobius muticus C. L. Koch, 1847 0 0 0
Strigamia acuminata (Leach, 1814) 0 0 0
Lithobius dentatus C. L. Koch, 1844 0 0 0
TABLE 2.
Characteristic and companion species of the different biotope-types.
species XM DSH SMM
sandy xeric dwarf- mesoxeric
meadows shrub meadows
heaths
Lithobius forficatus
(Linnaeus, 1758) companion
Lithobius melanops
Newport, 1845
Lithobius calcaratus
C. L. Koch, 1844 companion characteristic
Lithobius mutabilis
L. Koch, 1862
Lithobius crassipes
L. Koch, 1862
Schendyla nemorensis
(C. L. Koch, 1837)
Strigamia crassipes
(C. L. Koch, 1835)
Lithobius microps
Meinert, 1868 companion
Lamyctes emarginatus
(Newport, 1844) companion
Lithobius e.
erythrocephalus
C. L. Koch, 1847 companion
Cryptops parisi
Brolemann, 1920
species ASS HMG
advanced contaminated
successional grasslands
sites
Lithobius forficatus
(Linnaeus, 1758) companion characteristic
Lithobius melanops
Newport, 1845 characteristic
Lithobius calcaratus
C. L. Koch, 1844
Lithobius mutabilis
L. Koch, 1862 characteristic
Lithobius crassipes
L. Koch, 1862 characteristic
Schendyla nemorensis
(C. L. Koch, 1837) companion
Strigamia crassipes
(C. L. Koch, 1835) companion
Lithobius microps
Meinert, 1868
Lamyctes emarginatus
(Newport, 1844)
Lithobius e.
erythrocephalus
C. L. Koch, 1847
Cryptops parisi
Brolemann, 1920 companion |
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