Ground beetle fauna (Coleoptera: Carabidae) of protected areas in the Łódź Province. Part I. Nature reserves

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1 FRAGMENTA FAUNISTICA 55 (2): , 2012 PL ISSN MUSEUM AND INSTITUTE OF ZOOLOGY PAS Ground beetle fauna (Coleoptera: Carabidae) of protected areas in the Łódź Province. Part I. Nature reserves Radomir JASKUŁA and Anna STĘPIEŃ Department of Invertebrate Zoology and Hydrobiology, University of Łódź, Banacha 12/16, Łódź, Poland; Abstract: Ground beetle fauna of 35 nature reserves in the Łódź Province is summarized based on the literature data and unpublished records. For all studied reserves new faunistic data are presented, including first records of Carabidae for eight of them. In total, 140 ground beetle species (27.22% of Polish fauna and 54% of species recorded in the area of the Łódź Province) are noted from 35 (39%) nature reserves located in the Łodź Province. Moreover, it has been found, that the reserves Rawka and Spała encompass some of the highest Carabidae species richness noted among the all Polish nature reserves. Among the beetles recorded in the studied nature reserves, there are 16 species protected by Polish law, including 13 species of Carabus L. and 3 of Calosoma Web., what means 50% of Polish fauna and 76.19% of protected carabid species occurring in the Łódź Province. Twelve species are included in the Polish Red Data List: three species as critically endangered (CR) (Badister dorsiges, Patrobus australis, Trechus austriacus), one as endangered (EN) (Carabus clatratus), two as vulnerable (VU) (Odacantha melanura and Oodes helopioides), four as near threatened (NT) (Calosoma sycophanta, Carabus convexus, Diachromus germanus, and Omophron limbatum), one as least concern (LC) (Carabus intricatus), and one in data deficient category (DD) (Broscus cephalotes). High number of Carabidae species recorded on a relatively very small area of investigated nature reserves (in total only 0.29% of the Łódź Province) proves their importance in the conservation of species diversity of Carabidae in Poland. Key words: Carabidae, Central Poland, nature reserve, biodiversity, protected species, Red List, Red Book INTRODUCTION Ground beetles are the coleopteran family with worldwide distribution, including more than species (Lorenz 1998, 2005). Beetles belonging to this group can be found in all major terrestrial habitat types, both in lowlands and mountain areas, including eg. forests, meadows, banks of water reservoirs (freshwater and marine), deserts, caves. Some species are typically epigeic, other can be found in top of trees and bushes, or under the soil surface (eg. in mammal burrows) (eg. Lövei & Sunderland 1996). Beetles belonging to this family are both predators and plant eaters, some species are known as insect parasites too (eg. Weber et al. 2008). All this makes Carabidae an important group from the economic point of view (both as natural pest regulators and agricultural/forest pests) (Kotze et al. 2011). Most ground beetle species are active during vegetation season but at least some of them can stay active also during winter (Jaskuła & Soszyńska-Maj 2011). Even if many species can be characterized by wide distribution, great number of carabid beetles have relatively narrow habitat specialization (Eversham et al. 1996) what makes them an important bioindicator group in different studies focused eg. on habitat fragmentation, forestry, agriculture, or urban ecology. The family is often used as flagship group for estimation of local animal species richness and diversity, for environmental monitoring, classification of habitat type and assessment of site quality (for review see: Kotze et al. 2011). Polish carabid fauna includes 518 species (Aleksandrowicz 2004). Among them, 261 species have been noted in the Łódź Province till now (Jaskuła et al. 2002b, Jaskuła 2003, 2006, Schwerk & Szyszko 2006). According to the surface area of the province ( ha,

2 102 R. Jaskuła and A. Stępień 5.82% of the country) and not much diversified landscape (lack of some types of habitats like mountains, large lakes and rivers, and sea coast) this number is relatively high (50,2% of Polish fauna). However, for many species noted in this region there are only scarce published data (even if species are generally common in Poland), and their distribution within the area is still poorly recognized (Jaskuła et al. 2002b) Among the ground beetles of Poland there are 115 species (over 22%) that have the special protection status in Poland. Thirty-two species belonging to genera Carabus Linnaeus, 1758 and Calosoma Weber, 1801 (including Callisthenes Fischer von Waldheim, 1821) are legally protected (Dz. U. 2011), 100 taxa are placed in the Red Data List (Pawłowski et al. 2002), five species are included in the Red Data Book (Pawłowski 2004a e), and two are indexed in the Natura 2000 appendix II of the EU Habitat Directive (Kubisz 2004a b). Present knowledge on the ground beetles diversity and distribution in protected areas of the Łódź Province is based upon some 40 papers. Only 19 publications concern nature reserves but more than a half of these papers provide only single records for species which are known as rare or endangered in Poland or in the region, as well as for ground beetles protected by Polish law (Kowalczyk & Watała 1986, Kowalczyk et al. 1998, Soszyński et al. 1998, 2008, Kurzawa 1999, Jaskuła & Grabowski 2001, 2003, Jaskuła et al. 2002a b, Jaskuła & Ruta 2003a b). More detailed data (in total with Carabidae checklists for 18 nature reserves) are given by Jaskuła (2003, 2006), Jaskuła et al. (2005, 2009, 2010), Jaskuła & Socha (2007), and Stępień & Jaskuła (2010). As a result of all these faunistic studies, data from 27, i.e. 1/3 of all nature reserves of the Łódź Province are known until now. However, from six of these areas less than 10 Carabidae species are reported. The aim of this paper is to present the actual state of knowledge on ground beetle species richness and distribution in the nature reserves of the Łódź Province. It is provided on the basis of literature data and new faunistic records. This paper is the first part of a series presenting the ground beetle fauna of protected areas of the Łódź Province. The next two will present data on Carabidae occurring in the Natura 2000 areas as well as in the landscape parks of this region. STUDY AREA The Łódź Province (ŁP) is located in Central Poland and covers an area of km 2 what makes 5.82% of the country (GUS 2010). Geographically, the Łódź Province belongs to the three subprovinces: Central Poland Lowlands (Niziny Środkowopolskie), Małopolska Upland (Wyżyna Małopolska), and Silesian- Kraków Upland (Wyżyna Śląsko-Krakowska) (Kondracki 2000). The first one covers the northern part of area (three macroregions are distinguished: Nizina Południowowielkopolska north-western and western part of ŁP, Central Mazovian Lowland (Nizina Środkowomazowiecka) north-eastern part of ŁP, and Wzniesienia Południowomazowieckie eastern part of ŁP), the second is located in the south-eastern part (as macroregion Przedbórz Upland (Wyżyna Przedborska)), and the third one is placed in the south-western part (as macroregion Wyżyna Woźnicko-Wieluńska). Biogeographically, the Łódź Province can be classified in three different ways (Starkel 1999). Floristic classification includes the entire area in the region named Pas Wielkich Dolin. Phytosociological division classifies the Łódź Province to the North European Plain (Niż Środkowoeuropejski) and its two main regions: Central Poland Lowlands (Niziny Środkowopolskie) (divided into Niziny Środkowo i Wschodniopolskie in eastern part, and Wielkopolska Lowlands (Niziny Wielkopolskie) in western part; both of them covering ca. 45% of ŁP) and Wyżyny Południowopolskie (only a small south-eastern part of ŁP as Silesian-

3 Coleoptera of the nature reserves (Łódź Province, Poland) 103 Kraków Upland (Wyżyna Śląsko-Krakowska). Zoogeographical division classifies the entire area of the Łódź Province to Podokręg Wielkopolsko-Podolski (as a part of Okręg Środkowopolski). There are 89 nature reserves within the area of the Łódź Province, covering in total ha (0.41% of ŁP) (Fig. 1). According to the protection status distinguished by Rąkowski (2006) and GUS (2010) they can be divided into 8 main types (Table 1). Fig. 1. Localization of nature reserves in the Łódź Province (for number explanation see text with description of reserves). Including the new data presented in the current paper, ground beetle fauna was studied in the 35 nature reserves of the Łódź Province (Fig. 1). This makes 39% of such objects located in the area of the Łódź Province. Most of studied nature reserves are typically forest (22, 62.86% of studied NR), five are floristic (14.29%), three peat-bog (8.57%), two landscape (5.71%), while each geological, water, and faunistic NR are represented only by one (each 2.86%) (Rąkowski 2006, Główny Urząd Statystyczny 2010) (characterization given after Rąkowski 2006):

4 104 R. Jaskuła and A. Stępień 1. Ciosny (UTM: CC95) 2.42 ha; floristic type; protects a part of dune overgrown by 150 years old individuals of Juniperus communis, as well as some Pinus sylvestris and Betula spp. trees. None carabid species recorded so far. 2. Dąbrowa Grotnicka (UTM: CC85) ha; forest type; protects an area overgrown mainly by Potentillo albae-quercetum (with 150 years old Quercus trees) and Tilio- Carpinetum; part of the Dąbrowa Grotnicka Natura 2000 area (PLH100001). Twentynine species of Carabidae noted so far (Jaskuła et al. 2002b, Jaskuła 2003). 3. Dąbrowa w Niżankowicach (UTM: CB49) ha; forest type; protects mainly Potentillo albae-quercetum with 100 year old Quercus tress; part of the Załęcze Landscape Park and the Załęczański Łuk Warty Natura 2000 area (PLH100007). Four species of Carabidae noted so far (Jaskuła et al. 2002a). 4. Gać Spalska (UTM: DC41) ha; forest type; protects valley of Gać River overgrown mainly by Fraxino-Alnetum and Carici elongatae-alnetum; part of the Spała Landscape Park and the Lasy Spalskie Natura 2000 area (PLH100003). Thirty-four Carabidae species noted so far (Jaskuła & Socha 2007, Jaskuła et al. 2009). 5. Gaik (UTM: DB29) 32.86; forest type; protects mainly Tilio-Carpinetum; part of the Sulejów Landscape Park. One species of Carabidae noted so far (Soszyński et al. 1998). 6. Gałków (UTM: DC13) ha; forest type; protects mainly Tilio-Carpinetum with Fagus sylvatica and Abies alba occurring on border of their species ranges; part of the Buczyna Gałkowska Natura 2000 area (PLH100016). Seventeen species of Carabidae noted so far (Jaskuła et al. 2002a b, Jaskuła 2003). 7. Grądy nad Lindą (UTM: CC85) ha; landscape type; protects part of Linda River and its surroundings overgrown mainly by Tilio-Carpinetum and Fraxino-Alnetum. Part of the Grądy nad Lindą Natura 2000 area (PLH100022). None species of Carabidae noted so far. 8. Grądy nad Moszczenicą (UTM: CC95) ha; forest type; protects natural part of Moszczenica River and its valley overgrown mainly by Tilio-Carpinetum. None species of Carabidae noted so far. 9. Hołda (UTM: CB49) ha; forest type; protects mainly Fraxino-Alnetum and Tilio- Carpinetum floristic formations located near Warta River; part of the Warta and Widawka Landscape Park. None species of Carabidae noted so far. 10. Jaksonek (UTM: DB29) ha; floristic type; protects part of Pilica River valley overgrown mainly by Potentillo albae-quercetum, Tilio-Carpinetum, and Querco roboris- Pinetum; part of the Sulejów Landscape Park. One species of Carabidae noted so far (Soszyński et al. 1998). 11. Jeleń (UTM: DC30) ha; forest type; protects natural part of forest with 200 years old Quercus spp. and Abies alba; part of the Spała Landscape Park and the Lasy Smardzewickie Natura 2000 area (PLH100024). Thirty-nine Carabidae species noted so far (Jaskuła & Socha 2007, Jaskuła et al. 2009). 12. Jeziorsko (UTM: CC33) ha, faunistic (bird) type; protects Jeziorsko water reservoir created on the Warta River; part of the Zbiornik Jeziorsko Natura 2000 area (PLB100002). None species of Carabidae noted so far 13. Konewka (UTM: DC41) ha; forest type; protects natural parts of forest overgrown mainly by Potentillo albae-quercetum and Tilio-Carpinetum; part of the Spała Landscape Park and the Lasy Spalskie Natura 2000 area (PLH100003). Twentyfive species of Carabidae noted so far (Kurzawa 1999, Jaskuła et al. 2002a, 2009, Jaskuła & Socha 2007). 14. Kopanicha (UTM: DC46) ha; forest type; protects part of Rawka River valley, overgrown mainly by Fraxino-Alnetum and Carici elongatae-alnetum; part of the

5 Coleoptera of the nature reserves (Łódź Province, Poland) 105 Bolimów Landscape Park and the Dolina Rawki Natura 2000 area (PLH100015). Twenty-eight species of Carabidae noted so far (Stępień & Jaskuła 2010). 15. Korzeń (UTM: CC50) ha; peat-bog type; protects peat-bog overgrown by Erico- Sphagnetum magellanici and Caricetum limose; part of the Warta and Widawka Landscape Park. None species of Carabidae noted so far. 16. Las Łagiewnicki (UTM: CC95) ha; forest type; protects a forest complex overgrown mainly by Potentillo albae-quercetum, Fraxino-Alnetum, and Quercetea robori-petraeae; part of the Łódź Hills Landscape Park. Forty-eight species of Carabidae noted so far (Kowalczyk et al. 1998, Jaskuła et al. 2002a b, 2003, 2005, 2010, Jaskuła & Grabowski 2003). 17. Lubiaszów (UTM: DC20) ha; forest type; protects forest areas overgrown mainly by Tilio-Carpinetum with Abies alba occurring on border of its species range; part of the Sulejów Landscape Park and the Lubiaszów w Puszczy Pilickiej Natura 2000 area (PLH100026). Twenty-seven species of Carabidae noted so far (Soszyński et al. 1998, Jaskuła 2003). 18. Molenda (UTM: CC92) ha; forest type; protects mainly Tilio-Carpinetum and Abietetum polonicum with Abies alba and Fagus sylvatica occurring on border of their species ranges. Twenty-one species of Carabidae noted so far (Jaskuła et al. 2002b, Jaskuła 2003). 19. Niebieskie Źródła (UTM: DC30) ha; landscape type; protects natural springs and their surroundings overgrown mainly by Fraxino-Alnetum and Carici elongatae-alnetum; located in buffer zone of the Sulejów Landscape Park; part of the Niebieskie Źrodła Natura 2000 area (PLH100005). None species of Carabidae noted so far. 20. Parowy Janinowskie (UTM: DC14) ha; forest type; protects arroyos overgrown mostly by Luzulo luzuloidis-fagetum; part of the Łódź Hills Landscape Park and the Buczyna Janinowska Natura 2000 area (PLH100017). Thirty-four species of Carabidae noted so far (Kowalczyk & Watała 1986, Jaskuła et al. 2002a, 2010, Jaskuła 2003, Jaskuła & Ruta 2003a b). 21. Piskorzeniec (UTM: DB35) ha; peat-bog type; protects mainly peat-bog with surroundings overgrown by Carici elongatae-alnetum and Vaccinio uliginosi-pinetum; part of the Przedbórz Landscape Park and the Ostoja Przedborska Natura 2000 area (PLH260004). One species of Carabidae noted so far (Soszyński et al. 2008). 22. Polana Siwica (UTM: DC36) ha; floristic type; protects an area with meadows and peat-bog; part of the Bolimów Landscape Park and the Polany Puszczy Bolimowskiej Natura 2000 area (PLH100028). Seventeen species of Carabidae noted so far (Stępień & Jaskuła 2010). 23. Polesie Konstantynowskie (UTM: CC83) 9.80 ha; forest type; protects a natural part of forest (mainly Tilio-Carpinetum and Fraxino-Alnetum); it is located in Łódź city. Twentyone species of Carabidae noted so far (Jaskuła et al. 2000b, Jaskuła 2003). 24. Rawka (UTM: DC46) ha; water type; protects Rawka River and its valley with oxbows; part of the Bolimów Landscape Park and the Dolina Rawki Natura 2000 area (PLH100015). Seventy-eight species of Carabidae noted so far (Jaskuła 2006, Stępień & Jaskuła 2010). 25. Ruda-Chlebacz (UTM: DC45) ha; forest type; protects an area overgrown mainly by Fraxino-Alnetum; part of the Bolimów Landscape Park and the Dolina Rawki Natura 2000 area (PLH100015). Twenty-three species of Carabidae noted so far (Stępień & Jaskuła 2010).

6 106 R. Jaskuła and A. Stępień 26. Sługocice (UTM: DC30) 8.57 ha; floristic type; protects mainly an area overgrown by Tilio-Carpinetum; located in buffer zone of the Spała Landscape Park. Eighteen species of Carabidae noted so far (Jaskuła & Socha 2007, Jaskuła et al. 2010). 27. Spała (UTM: DC40) ha; forest type; located on both side of Pilica River, protects one of the oldest forest in Łódź region, mainly Tilio-Carpinetum and Fraxino-Alnetum; part of the Spała Landscape Park and the Lasy Spalskie Natura 2000 area (PLH100003). Eighty species of Carabidae noted so far (Jaskuła et al. 2002a, 2009, Jaskuła & Socha 2007). 28. Starodrzew Lubochniański (UTM: DC41) ha; forest type; protects an area overgrown mainly by Tilio-Carpinetum with 150 years old Quercus spp. and Pinus sylvestris trees. One species of Carabidae noted so far (Jaskuła et al. 2002a). 29. Struga Dobieszkowska (UTM: DC04) ha; forest type; protects part of Struga Dobieszkowska River and its valley overgrown by Fraxino-Alnetum and Tilio- Carpinetum; part of the Łódź Hills Landscape Park. Forty species of Carabidae noted so far (Jaskuła et al. 2002a b, 2003, 2010, Jaskuła 2003). 30. Torfowisko Rąbień (UTM: CC84) ha; peat-bog type; protects an area of peat-bog with its surroundings. Twenty species of Carabidae noted so far (Jaskuła et al. 2002a b, Jaskuła 2003). 31. Węże (UTM: CB36) ha; geological type; protects the Hill of Zelce with underground cave system and with xerothermic flora and fauna; part of the Załęcze Landscape Park and the Załęczański Łuk Warty Natura 2000 area (PLH100007). One species of Carabidae noted so far (Jaskuła & Grabowski 2001). 32. Wiączyń (UTM: DC03) 8.80 ha; forest type; protects natural forest with some 300 years old Fagus sylvatica trees. Thirteen species of Carabidae noted so far (Jaskuła et al. 2002a b, 2003). 33. Winnica (UTM: CC40) 1.54 ha; floristic type; protects part of Warta River valley, with xerothermic meadows; part of the Warta and Widawka Landscape Park. None species of Carabidae noted so far. 34. Wolbórka (UTM: CC92) ha; forest type; protects mainly springs of Wolbórka River and their surroundings overgrown by Fraxino-Alnetum and Carici elongatae- Alnetum. Nineteen species of Carabidae noted so far (Jaskuła et al. 2002a, Jaskuła 2003) 35. Żądłowice (UTM: DC51) ha; forest type; protects mosaic area overgrown mainly by Carici elongatae-alnetum and Peucedano-Pinetum; part of the Spała Landscape Park and two Natura 2000 areas: the Dolina Pilicy (PLB140003) and the Dolina Dolnej Pilicy (PLH140016). Twenty-eight species of Carabidae noted so far (Jaskuła et al. 2002b, 2009, Jaskuła & Socha 2007). MATERIAL AND METHODS All available literature data on diversity and distribution of Carabidae species occurring in nature reserves of Łódź Province were analyzed. In total, faunistic records published in 19 papers were compiled (Kowalczyk & Watała 1986, Kowalczyk et al. 1998, Kurzawa 1998, Soszyński et al. 1998, 2008, Jaskuła 2003, 2006, Jaskuła & Grabowski 2001, 2003, Jaskuła et al. 2002a b, 2003, 2005, 2009, 2010, Jaskuła & Ruta 2003a b, Jaskuła & Socha 2007, Stepień & Jaskuła 2010). Moreover, new unpublished records from all 35 investigated areas were added to the checklists of ground beetles of each nature reserve (Table 1). All the new data are based on material collected during fieldwork in years The main collection method used in all NR was active searching for beetles under stones and pieces of decaying wood,

7 Coleoptera of the nature reserves (Łódź Province, Poland) 107 under bark of trees, and among litter. Moreover, light traps were used in Wiączyń NR, Lubiaszów NR, and Rawka NR as well as pitfall traps in Lubiaszów NR and Jeziorsko NR. Table 1. General characteristics of the nature reserves of the Łódź Province (Rąkowski 2006, GUS 2010). Nature reserves of the Łódź Province Number Area Type of nature Share in the region Percentage in the reserve Total Total area [ha] [%] region [%] Forest Floristic Peat-bog Landscape , Faunistic Aquatic Geological Halophyte Nature reserves included in present study Number Area Type of nature Share in the region Percentage in the reserve Total Total area [ha] [%] region [%] Forest 22 24,7 1328, Floristic 5 5, Peat-bog 3 3, Landscape 2 2,3 231, Faunistic Aquatic Geological Halophyte RESULTS AND DISCUSSION Carabid species richness in nature reserves In total 140 species belonging to 61 genera were noted (Table 2). This makes 27.22% of Polish Carabidae fauna (Aleksandrowicz 2004) and 54.02% of ground beetle taxa recorded in the Łódź Province (Jaskuła et al. 2002b, Jaskuła 2003, 2006, Schwerk & Szyszko 2006). For eight NR data presented in this paper are the first known records of Carabidae from these objects. These nature reserves are: Ciosny NR, Grądy nad Lindą NR, Grądy nad Moszczenicą NR, Hołda NR, Jeziorsko NR, Korzeń NR, Niebieskie Źródła NR, and Winnica NR. Moreover, according to literature data, for all the other 27 nature reserves number of noted ground beetle species increased from 8.8% to even 97% comparing to published data (Kowalczyk & Watała 1986, Kowalczyk et al. 1998, Kurzawa 1998, Soszyński et al. 1998, 2008, Jaskuła 2003, 2006, Jaskuła & Grabowski 2001, 2003, Jaskuła et al. 2002a b, 2003, 2005, 2009, 2010, Jaskuła & Ruta 2003a b, Jaskuła & Socha 2007, Stepień & Jaskuła 2010). The highest numbers of Carabidae species were recorded in Rawka NR and Spała NR 97 and 87 taxa respectively (Table 2, Fig. 2). These two nature reserves are characterized by the highest species richness not only among all NR of the Łódź Province but also among all such areas in Poland. As far as we know, only in two other Polish nature reserves higher or similar numbers of Carabidae species have been noted. These are Cisy Staropolskie im. Leona Wyczółkowskiego NR located in the Kujawy-Pomeranian Province (Błażejewski 1964, Stachowiak & Wilcz 2001) and Zwierzyniec NR placed in the Lower Silesia Province (Drozd-Nurek & Kornalewicz 1986, Twardowski & Thoma 2011), where respectively 139 and 84 species were found.

8 108 R. Jaskuła and A. Stępień Table 2. Occurrence of the ground beetle species in nature reserves of the Łódź Province: published data, new records. Protection status No. Species Abax carinatus (Duftschmidt, 1812) 2. Acupalpus parvulus (Sturm, 1825) 3. Agonum dolens (C.R. Sahlberg, 1827) 4. Agonum fuliginosus (Panzer, 1809) 5. Agonum gracile Sturm, Agonum micans (Nicolai, 1822) 7. Agonum sexpunctatum (Linnaeus, 1758) 8. Amara aenea (De Geer, 1774) 9. Amara apricaria (Paykull, 1790) 10. Amara aulica (Panzer, 1796) 11. Amara bifrons (Gyllenhal, 1810) 12. Amara brunnea (Gyllenhal, 1810) 13. Amara communis (Panzer, 1797) 14. Amara fulva (O.F. Müller, 1776) 15. Anchomenus dorsalis (Pontoppidan, 1763) 16. Anisodactylus binotatus (Fabricius, 1787) 17. Anisodactylus nemorivagus (Duftschmid, 1912) 18. Anisodactylus signatus (Panzer, 1796) 19. Anthracus consputus (Duftschmidt, 1812) 20. Asaphidion flavipes (Linnaeus, 1761) 21. Asaphidion pallipes (Duftschmidt, 1812) 22. Badister bullatus (Schrank, 1798) 23. Badister dorsiger (Duftschmid, 1812) CR 24. Badister sodalis (Duftchmidt, 1812) 25. Badister unipustatulus Bonelli, Bembidion quadrimaculatum (Linnaeus, 1761) 27. Blemus discus (Fabricius, 1792) 28. Bradycellus csikii Laczó, Bradycellus harpalinus (Audinet-Serville, 1821) 30. Bradycellus verbascii (Duftschmidt, 1812) 31. Broscus cephalotes (Linnaeus, 1758) DD 32. Calathus cinctus Motschulsky, Calathus erratus (C. R. Sahlberg, 1827) 34. Calathus fuscipes Goeze, Calathus melanocephalus (Linnaeus, 1758) 36. Calathus micropterus (Duftschmid, 1812) 37. Calosoma auropunctatum (Herbst, 1784) P 38. Calosoma inquisitor (Linnaeus, 1758) P 39. Calosoma sycophanta (Linnaeus, 1758) P, NT 40. Carabus arvensis Herbst, 1784 P 41. Carabus auronitens Fabricius, 1792 P 42. Carabus cancellatus Illiger, 1798 P 43. Carabus clathratus Linnaeus, 1761 P 44. Carabus convexus Fabricius, 1775 P 45. Carabus coriaceus Linnaeus, 1758 P, NT 46. Carabus glabratus Paykull, 1790 P 47. Carabus granulatus Linnaeus, 1758 P Ciosny Dąbrowa Grotnicka Dąbrowa w Niżankowicach Gać Spalska Gaik Gałków Grądy nad Lindą Grądy nad Moszczenicą Hołda Jaskonek Jeleń

9 Coleoptera of the nature reserves (Łódź Province, Poland) 109 Table 2 continuation (2) Jeziorsko Konewka Kopanicha Korzeń Las Łagiewnicki Lubiaszów Molenda Niebieskie Źródła Parowy Janinowskie Piskorzeniec Polana Siwica Polesie Konstantynowskie Rawka Ruda Chlebacz Sługocice Spała Starodrzew Lubochniański Struga Dobieszkowska Torfowisko Rąbień Węże Wiączyń Winnica Wolbórka Żądłowice No

10 110 R. Jaskuła and A. Stępień Table 2 continuation (3) Protection status No. Species Carabus hortensis Linnaeus, 1758 P 49. Carabus intricatus Linnaeus, 1758 P 50. Carabus linnaei Panzer, 1810 P 51. Carabus nemoralis O. F. Müller, 1764 P, NT 52. Carabus violaceus Linnaeus, 1758 P 53. Chlaenius nigricornis (Fabricius, 1787) 54. Chlaenius nitudulus (Schrank, 1781) 55. Cicindela campestris Linnaeus, Cicindela hybrida Linnaeus, Clivina collaris (Herbst, 1784) 58. Clivina fossor Linnaeus, Cychrus caraboides Linnaeus, Demetrias atricapillus (Linnaeus, 1758) 61. Diachromus germanus (Linnaeus, 1758) NT 62. Dicheirotrichus placidus (Gyllenhal, 1827) 63. Dromius agilis (Fabricius, 1787) 64. Dromius fenestratus (Fabricius, 1794) 65. Dromius quadraticollis (A. Morawitz, 1862) 66. Dromius quadrimaculatus A. Morawitz, Dromius schneideri (Crotch, 1871) 68. Dyschirius aeneus (Dejean, 1825) 69. Dyschirius globosus (Herbst, 1784) 70. Dyschirius nitidus (Dejean, 1825) 71. Dyschirius obscurus (Gyllenhal, 1827) 72. Elaphrus cupreus Duftschmid, Elaphrus riparius Linnaeus, Emphanes azurescens (Dalla Torre, 1877) 75. Emphanes tenellus (Erichson, 1837) 76. Epaphius secalis (Paykull, 1790) 77. Harpalus affinis (Schrank, 1781) 78. Harpalus griseus (Panzer, 1796) 79. Harpalus latus (Linnaeus, 1758) 80. Harpalus rufipes (De Geer, 1774) 81. Harpalus tardus (Panzer, 1796) 82. Lebia cruxminor (Linnaeus, 1758) 83. Leistus ferrugineus (Linnaeus, 1758) 84. Leistus rufomarginatus (Duftschmid, 1812) 85. Leistus terminatus (Panzer, 1793) 86. Limodromus assimilis (Paykull, 1790) 87. Loricera pilicornis (Fabricius, 1775) 88. Metallina lampros (Herbst, 1784) 89. Metallina properans (Stephens, 1828) 90. Microlestes maurus (Sturm, 1827) 91. Microlestes minutulus (Goeze, 1777) 92. Nebria brevicollis (Fabricius, 1792) 93. Notaphus dentellus (Thunberg, 1787) 94. Notaphus obliquus (Sturm, 1825) Ciosny Dąbrowa Grotnicka Dąbrowa w Niżankowicach Gać Spalska Gaik Gałków Grądy nad Lindą Grądy nad Moszczenicą Hołda Jaskonek Jeleń

11 Coleoptera of the nature reserves (Łódź Province, Poland) 111 Table 2 continuation (4) Jeziorsko Konewka Kopanicha Korzeń Las Łagiewnicki Lubiaszów Molenda Niebieskie Źródła Parowy Janinowskie Piskorzeniec Polana Siwica Polesie Konstantynowskie Rawka Ruda Chlebacz Sługocice Spała Starodrzew Lubochniański Struga Dobieszkowska Torfowisko Rąbień Węże Wiączyń Winnica Wolbórka Żądłowice No

12 112 R. Jaskuła and A. Stępień Table 2 continuation (5) Protection status No. Species Notaphus varius (Olivier, 1795) 96. Notiophilus aestuans Dejean, Notiophilus aquaticus (Linnaeus, 1758) 98. Notiophilus biguttatus (Fabricius, 1779) 99. Notiophilus palustris (Duftchmid, 1812) 100. Ocydromus bruxellensis (Wesmael, 1835) 101. Ocydromus cruciatus polonicus J. Müller, Ocydromus modestus (Fabricius, 1801) 103. Ocydromus tetracolus (Say, 1823) 104. Odacantha melanura (Linnaeus, 1767) VU 105. Odontium argenteolum (Arhens, 1812) 106. Odontium littorale (Olivier, 1790) 107. Omophron limbatum (Fabricius, 1777) NT 108. Oodes helopioides (Fabricius, 1792) VU 109. Ophonus rupicola (Sturm, 1818) 110. Oxypselaphus obscurus (Herbst, 1784) 111. Panagaeus bipustulatus (Fabricius, 1775) 112. Paradromius linearis (Olivier, 1795) 113. Paranchus albipes (Fabricius, 1796) 114. Patrobus atrorufus (Strom, 1768) 115. Patrobus australis J. Sahlberg, 1875 CR 116. Philochthus bigutatus (Fabricius, 1779) 117. Philochthus gutulla (Fabricius, 1792) 118. Philochthus mannercheimii (C. R. Sahlberg, 1827) 119. Philorhizus sigma (Rossi, 1790) 120. Poecilus cupreus (Linnaeus, 1758) 121. Poecilus lepidus (Leske, 1785) 122. Poecilus versicolor (Sturm, 1824) 123. Princidium punctulatum (Drapiez, 1821) 124. Princidium ruficolle (Panzer, 1796) 125. Pterostichus burmeisteri Heer, Pterostichus diligens (Sturm, 1824) 127. Pterostichus melanarius (Illiger, 1798) 128. Pterostichus minor (Gyllenhal, 1827) 129. Pterostichus niger (Schaller, 1783) 130. Pterostichus nigrita (Paykull, 1790) 131. Pterostichus oblongopunctatus (Fabricius, 1787) 132. Pterostichus strenuus (Panzer, 1796) 133. Stenolophus mixtus (Herbst, 1874) 134. Stenolophus teutonus (Schrank, 1781) 135. Synuchus vivalis (Illiger, 1798) 136. Tachyta nana (Gyllenhal, 1810) 137. Trechus austriacus Dejean, 1831 CR 138. Trepanes articulatus (Panzer, 1796) 139. Trepanes assimilis (Gyllenhal, 1810) 140. Trepanes doris (Panzer, 1796) Ciosny Dąbrowa Grotnicka Dąbrowa w Niżankowicach Gać Spalska Gaik Gałków Grądy nad Lindą Grądy nad Moszczenicą Hołda Jaskonek Jeleń

13 Coleoptera of the nature reserves (Łódź Province, Poland) 113 Table 2 continuation (6) Jeziorsko Konewka Kopanicha Korzeń Las Łagiewnicki Lubiaszów Molenda Niebieskie Źródła Parowy Janinowskie Piskorzeniec Polana Siwica Polesie Konstantynowskie Rawka Ruda Chlebacz Sługocice Spała Starodrzew Lubochniański Struga Dobieszkowska Torfowisko Rąbień Węże Wiączyń Winnica Wolbórka Żądłowice No

14 114 R. Jaskuła and A. Stępień Fig. 2. Species richness of Carabidae in nature reserves of the Łódź Province. 114

15 Fig. 3. Number of protected and endangered Carabidae noted in nature reserves of the Łódź Province. Coleoptera of the nature reserves (Łódź Province, Poland) 115

16 116 R. Jaskuła and A. Stępień In the area of the Łódź Province data on ground beetles are known from 39% of nature reserves. Comparing with other provinces of Poland this value is very high. In most Polish provinces regular faunistic studies on Carabidae were undertaken in total only in several nature reserves while from great number of such areas only single, if any, species are known. Regular investigations (both on ground beetles or on Coleoptera in general with ground beetles as a part) were made in NR of the Świętokrzyskie Province ( Białe Ługi NR Huruk 2001, Leśniak 2001b, 2004; Cisów NR Huruk 2000, Leśniak & Górecka 2000; Świnia Góra Leśniak 2001a, 2004), in the Kujawy-Pomerania Province ( Cisy Staropolskie im. Leona Wyczółkowskiego NR Błażejewski 1964, Stachowiak & Wilcz 2001); in the West- Pomerania Province ( Bielinek NR Wojas 1997; Wzgórze widokowe nad Miedzyodrzem Wolender 2002; Jezioro Szare NR Aleksandrowicz & Dąbrowski 2007; Leśne Źródła and Wielki Bytyń Gutowski & Ruta 2004), in the Małopolska Province ( Madohora NR Kaźmierczak 1997, Dolina Szklarki NR and Dolina Racławki NR Maciejowski 2007), in the Wielkopolska Province ( Kuźnik NR Ruta & Melke 2002, 2011; Krajkowo NR Wojas 1992, Sienkiewicz 2000, 2003, Sienkiewicz & Konwerski 2004, 2005), in the Warmia- Masuria Province ( Redykajny NR Nietupski et al. 2006, 2008a, Mszar NR Nietupski et al. 2008b, and Las Warmiński NR Czachorowski et al. 1998), in the Podlasie Province ( Karczmisko NR and Krzemionka NR Jaskuła & Rachalewski 2011), in the Lower Silesia Province ( Zwierzyniec NR Drozd-Nurek & Kornalewicz 1986, Twardowski & Kornalewicz 2004, Twardowski & Thoma 2011), in the Pomerania Province ( Łebskie Bagno NR and Czarne Bagno NR Jaskuła & Rewicz 2011), in the West Pomerania Province ( Gołębia Góra RN Aleksandrowicz & Genzel 2011), and in the Masovian Lowland ( Las Bielański NR Czechowski 1982, Dębina NR, Las im. Jana III Sobieskiego NR, Modrzewina NR, Cyganka NR, Bolesław Hryniewiecki NR Czechowski 1989). In case of a few Carabidae species new faunistic data are needed to confirm their occurrence in investigated areas. Among them are: Emphanes tenellus recorded from Żądłowice NR (Jaskuła et al. 2002b), and Carabus clathratus noted in Lubiaszów NR (Soszyński et al. 1998). Moreover, all literature data on Calathus mollis occurrence in NR areas of the Łódź Province relate to C. cinctus (Jaskuła et al. 2002b, 2003, 2005), as well as record of Patrobus assimilis from Torfowisko Rąbień (Jaskuła et al. 2002b, Jaskuła 2003) applies to P. atrorufus. Rare, endangered and protected ground beetle species Among 140 Carabidae species recorded from the investigated nature reserves of the Łódź Province, 24 have special protection status (Table 2, Fig.3.). Sixteen taxa are protected by Polish law, including 13 species of Carabus L. and 3 of Calosoma Web., one is included in the Polish Red Data Book (Carabus clathratus in EN category; Pawłowski 2004a) and 12 species (including three protected and one from the Red Book) are placed in the Polish Red Data List (Pawłowski et al. 2002). Among them are three taxa classified as critically endandered (CR), one as endangered (EN), two as vulnerable (VU), four as near threatened (NT), one as least concern (LC), and one in data deficient category (DD). In total 50% of protected Polish ground beetle fauna (Aleksandrowicz 2004, Dz.U. 2011) and 79.19% of such species occurring in the Łódź Province (Jaskuła 2002a) have been noted in the investigated nature reserves. According to the area of ŁP, this number can be even higher (84.2%) as occurrence of two Carabidae species previously noted from this region should be confirmed by new data (Carabus auratus known only as a single individual collected near Rogów (Kinelski & Szujecki 1959) and C. ulrichii a mountain species noted from Meszcze by Borusiewicz & Kapuściński (1950)). In all NR species protected by Polish law were recorded (Table 1, Fig. 3). The richest faunas of such ground beetles were found in Las

17 Coleoptera of the nature reserves (Łódź Province, Poland) 117 Łagiewnicki NR (12 species), Lubiaszów NR (11), Parowy Janinowskie NR (11), Rawka NR (11), Wiączyń NR (11), Dąbrowa w Niżankowicach (10), and Konewka NR (10). In all these areas more than 50% of all protected Carabidae species known from the Łódź Province were noted (Jaskuła et al. 2002a, Jaskuła 2006). Only Carabus marginalis, C. nitens, C. problematicus, C. ulrichii, and C. auratus were not found in any of investigated NR, but at least occurrence of the last two of them should be confirmed by new data in the area of the entire Łódź Province (Jaskuła et al. 2002a). In the next 21 NR five to nine protected species were recorded (23 40% of ŁP fauna), while less than 20% of all protected Carabidae were found in 7 NR, with the smallest number (only 1 species) noted in Cisy NR and Polana Siwica NR. Most of the legally protected ground beetle species recorded in the investigated nature reserves are widely distributed in Poland and in the Łódź Province, and usually they are not rare or even common (Burakowski et al. 1973, Jaskuła et al. 2002a). To this group belong: Calosoma inquisitor, Carabus arvensis, C. auronitens, C. cancellatus, C. convexus, C. glabratus, C. granulatus, C. hortensis, C. nemoralis, and C. violaceus. All these taxa were noted in most of the investigated areas (15 35 NR; Table 2, Fig. 3). To the group of species rarely collected in the area of ŁP belong: (1) species rare and very rare in Polish fauna C. clathratus and Calosoma sycophanta (Burakowski et al. 1973) both known from the Łódź Province from only 2 3 localities (Jaskuła et al. 2002a), (2) species with the northern border of species distribution in Poland located in the Łódź Province (Kowalczyk & Watala 1986) Carabus linnaei, recorded in three nature reserves ( Las Łagiewnicki NR, Parowy Janinowskie NR and Wiączyń NR), 3/ other rare species in the Łódź Province: Calosoma auropunctatum (among NR known from Rawka NR only; records from Gaik NR and Lubiaszów NR given by Jaskuła et al. (2002a) were wrongly cited as in reality they refer to C. inquisitor (Soszyński et al. 1998)), Carabus intricatus (noted only from Parowy Janinowskie NR), and C. coriaceus (known only from two nature reserves placed in the Bolimów Landscape Park Rawka NR and Ruda Chlebacz NR (Stępień & Jaskuła 2010). According to the Red List, in total 12 Carabidae species were noted in 17 of investigated NR (Table 2, Fig. 3), what makes 12% of Polish fauna (Pawłowski et al. 2002). Among investigated areas Rawka NR and Spała NR are characterized by the highest number of such taxa (3 species). In other 15 NR only 1 2 species from the List were noted. The only nature reserve where Carabidae species included in the Polish Red Data Book was Lubiaszów NR (Table 2, Fig. 3). In this area one species Carabus clathratus was found, what makes 20% of the Polish Red Book ground beetle fauna (Pawłowski 2004a e), and 50% of such beetle species occurring in the Łódź Province (Jaskuła et al. 2002b, Jaskuła 2003, 2006, Schwerk & Szyszko 2006). The Red List critically endangered ground beetles are: 1/ Badister dorsiger noted in Polesie Konstantynowskie NR (Jaskuła 2003), 2/ Trechus austriacus recorded in Węże NR (Jaskuła & Grabowski 2003; new material: 3 ex., Stalagmitowa Cave, VI , leg. R. Jaskuła et J. Szymański; 2 ex., Za Kratą Cave, IV.2004., leg. J. Szymański; 1 ex., Niespodzianka Cave, IV.2004., leg. J. Szymański), 3/ Patrobus australis collected in Wiączyń NR (new material: 1 ex., I.VIII.2002., light trap, leg. R. Jaskuła et J. Kalisiak). The endangered beetle is Carabus clathratus recorded in Lubiaszów NR (Soszyński et al. 1998). The vulnerable carabids are: Odacantha melanura found in Rawka NR (Jaskuła 2006) and Oodes helopioides noted in Gać Spalska NR, Jeleń NR, Spała NR (Jaskuła & Socha 2007, Jaskuła et al. 2009), and in Struga Dobieszkowska NR (new material: 1 ex., , leg. R. Jaskuła). The threatened species are: 1/ Calosoma sycophanta Konewka NR (Jaskuła et al. 2009) and Las Łagiewnicki NR (Jaskuła et al. 2010), 2/ Carabs convexus Dąbrowa

18 118 R. Jaskuła and A. Stępień Grotnicka NR (Jaskuła 2003), Dąbrowa w Niżankowicach (new material: 1 ex., V.2004., leg. R. Jaskuła), Konewka NR and Spała NR (Jaskuła & Socha 2007, Jaskuła et al. 2009), Las Łagiewnicki NR (Jaskuła et al. 2005, 2010), Lubiaszów NR (new material: 3 ex., VI.2005., leg. R. Jaskuła, J. Hejduk et M. Olczyk), Molenda NR (Jaskuła 2003), Parowy Janinowskie (Jaskuła 2003, Jaskuła et al. 2010), Ruda Chlebacz (Stępień & Jaskuła 2010), Starodrzew Lubochniański NR (new material: 2 ex., VIII.2008., leg. R. Jaskuła), Węże (new material: 1ex., VI.2004., leg. R. Jaskuła), Wiączyń NR (new material: 2 ex., VIII.2003., 3 ex., IX.2005., leg. R. Jaskuła), 3/ Diachromus germanus Parowy Janinowskie NR (Jaskuła & Ruta 2003a), 4/ Omophron limbatum Gać Spalska NR and Spała NR (Jaskuła & Socha 2007, Jaskuła et al. 2009), Rawka NR (Jaskuła 2006, Stępień & Jaskuła 2010), and Struga Dobieszkowska NR (new material: 1 ex., , leg. R. Jaskuła). In least concern category and data deficient category only single species are included, Carabus intricatus and Broscus cephalotes respectively. The first species was regularly found in Parowy Janinowskie (Jaskuła et al. 2002a, Jaskuła 2003; new material: several specimens noted in years by the first author), while the second was noted in Rawka NR (Jaskuła 2006, Stępień & Jaskuła 2010). CONCLUSIONS Present knowledge allows to provide faunistic data from 35 nature reserves of the Łódź Province (140 species, including 16 protected taxa and 12 included in the Polish Red Data List and Book). According to the very small area of the investigated NR (in total only 0.29% of ŁP) and relatively high number of recorded Carabidae species (27.22% of Polish fauna or 54% of species recorded in the area of ŁP), we can assume that nature reserves play an important role in nature conservation of ground beetle fauna in the Łódź Province. Moreover, as for now Carabidae were studied in only 39% of all NR located in this region, it can be expected that further studies allow to find additional species in such areas as well as to confirm their significant role in protection of ground beetles in the studied area. ACKNOWLEDGEMENTS We would like to thank to Karolina Banasik, Anna Drozd, Kamila Dunajska, Anna Fijarczyk, Janusz Hejduk, Jacek Hikisz, Paweł Hikisz, Piotr Jóźwiak, Jacek Kalisiak, Łukasz Kępczyński, Andżelika Kucharska, Dorota Laskowska, Damian Lewiński, Marek Michalski, Magdalena Nowakowska-Wójcik, Marcin Olczyk, Piotr Pawlikiewicz, Michał Rachalewski, Tomasz Rewicz, Grzegorz Socha, Agnieszka Soszyńska-Maj, Jakub Szymański, Ewelina Włodarczyk, and Karol Zemko for their help during field works in nature reserves of the Łódź Province. Additionally, special thanks to Piotr Jóźwiak for preparing map of the Łódź Province. REFERENCES ALEKSANDROWICZ O. R Biegaczowate (Carabidae). Pp In: BOGDANOWICZ W., CHUDZICKA E., PILIPIUK I. & SKIBIŃSKA E. (eds). Fauna Polski Charakterystyka i wykaz gatunków. Vol. 1. Muzeum i Instytut Zoologii PAN, Warszawa., 509 pp. ALEKSANDROWICZ O. & DĄBROWSKI P Preliminary studies on epigeic beetle fauna (Coleoptera) in surroundings of "Szare Lake" nature reserve. Parki Narodowe i Rezerwaty Przyrody 26 (4): [In Polish with English summary] ALEKSANDROWICZ O., GENZEL A Species composition and carabid s assembly structure (Coleoptera: Carabidae) of old pine mixed forest in protected terrain Gołębia Góra. Słupskie Prace Biologiczne 8: [In Polish with English abstract]

19 Coleoptera of the nature reserves (Łódź Province, Poland) 119 BŁAŻEJEWSKI F From the invertigation of ground-beetles (Carabidae) in the yew-tree reserve Wierzchlas. Przyroda Polski Zachodniej 7 (1 4): [In Polish with English summary] BORUSIEWICZ A. & KAPUŚCIŃSKI S Materials to the knowledge about the dissemination of the genus Carabus Lin. (Carabidae, Coleoptera) in the Polish forests. Prace Rolniczo-Leśne PAU 54: [In Polish with English summary] BURAKOWSKI B., MROCZKOWSKI M. & STEFAŃSKA J Chrząszcze Coleoptera, Biegaczowate Carabidae. Katalog Fauny Polski. Warszawa, XXIII. 2: CZACHOROWSKI S., BUCZYŃSKI P., ALEXANDROVITCH O., STRYJECKI R., KURZĄTKOWSKA A Material required for knowledge of insects and arachnids of the Warmiński Forest nature reserve (the Olsztyn Lake District). 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Informacje i opracowania statystyczne], Warszawa, 609 pp. GUTOWSKI J. M. & RUTA R Estimating natural values of the Tuczno commune on the basis of the beetle fauna survey (Insecta: Coleoptera). Nowy Pamiętnik Fizjograficzny 3 (1 2): [In Polish with English summary] HURUK S Carabid (Carabidae, Col.) of the Cisów reserve and adjoining cultivated fields. Rocznik Świętokrzyski, Ser. B. Nauki Przyrodnicze 27: [In Polish with English summary] HURUK S Biegaczowate (Col., Carabidae) wybranych zespołów roślinnych Białych Ługów. Pp In: ŻUREK S. (ed.), Rezerwat torfowiskowy Białe Ługi. Wydawnictwo Homini, Kraków, 268 pp. JASKUŁA R Carabid beetles (Coleoptera: Carabidae) of some nature reserves near Łódź. Parki Narodowe i Rezerwaty Przyrody 22 (4): JASKUŁA R Dalsze dane do poznania biegaczowatych (Coleoptera: Carabidae) Bolimowskiego Parku Krajobrazowego ze szczególnym uwzględnieniem rezerwatu Rawka. Przyroda Polski Środkowej 8: JASKUŁA R. & GRABOWSKI M New records of some interesting species of Carabidae (Coleoptera) from Poland. Wiadomości Entomologiczne 22 (1): [In Polish] JASKUŁA R. & GRABOWSKI M New localities of interesting eight species of ground beetles (Coleoptera: Carabidae) in Poland. Wiadomości entomologiczne 22: [in Polish] JASKUŁA R., KALISIAK J. & SZCZEPKO K New localities of some rare species of ground beetles (Coleoptera: Carabidae) in Poland. Parki Narodowe i Rezerwaty Przyrody 22 (2): [In Polish with English abstract] JASKUŁA R., KOWALCZYK J. K., LOGA B., SOSZYŃSKA MAJ A. & WATAŁA C Ground beetles (Coleoptera: Carabidae) of Łagiewniki Forest near Łódź with special regard to Las Łagiewnicki Nature Reserve. Parki Narodowe i Rezerwaty Przyrody 24 (1 4): [In Polish with English abstract] JASKUŁA R., KOWALCZYK J. K. & WATAŁA C. 2002a. Protected ground beetles (Carabidae) occurring on Łódź Upland. Parki Narodowe i Rezerwaty Przyrody 21 (2): [In Polish with English abstract] JASKUŁA R., KOWALCZYK J. K. & WATAŁA C. 2002b. Ground beetles (Coleoptera: Carabide) of Lodz Upland, Central Poland. Baltic Journal of Coleopterology 2 (2): JASKUŁA R., PRZEWOŹNY M. & MELKE A Chrząszcze (Coleoptera). Pp In: JASKUŁA R. & TOŃCZYK G. (eds), Owady Spalskiego Parku Krajobrazowego. Część I. Mazowiecko-Świetokrzyskie Towarzystwo Ornitologiczne, Spała, 119 pp. JASKUŁA R., PRZEWOŹNY M., MELKE A. & SOSZYŃSKA-MAJ A Chrząszcze Coleoptera. Pp In: JASKUŁA R. & TOŃCZYK G. (eds), Owady (Insecta) Parku Krajobrazowego Wzniesień Łódzkich. Dyrekcja Parku Krajobrazowego Wzniesień Łódzkich/Mazowiecko-Świętokrzyskie Towarzystwo Ornitologiczne, Łódź, 213 pp. JASKUŁA R. & RACHALEWSKI M Materiały do znajomości biegaczowatych (Coleoptera: Carabidae) dwóch rezerwatów przyrody zachodniej części Parku Krajobrazowego Puszcza Knyszyńska, p. 30. In: 12 Sympozjum Polskich Karabidologów. Biegaczowate i inne bezkręgowce w badaniach środowiskowych. Olsztyn, VI Materiały konferencyjne, Olsztyn, 36 pp. JASKUŁA R. & REWICZ T Chrząszcze biegaczowate. Pp In: HERBICHOWA M. & HERBICH J. (eds), Przyroda rezerwatów Łebskie Bagno i Czarne Bagno. Fundacja Rozwoju Uniwersytetu Gdańskiego, Gdańsk, 330 pp. JASKUŁA R. & RUTA R. 2003a. New localities of some interesting species of ground beetles (Coleoptera: Carabidae) in Poland. Wiadomości entomologiczne 22 (1): [In Polish] JASKUŁA R. & RUTA R. 2003b. New localities of seven interesting species of ground beetles (Coleoptera: Carabidae) from Poland. Wiadomości entomologiczne 21 (4): [In Polish]

20 120 R. Jaskuła and A. Stępień JASKUŁA R. & SOCHA G Materials to the knowledge of carabid fauna (Coleoptera: Carabidae) of the Spała Landscape Park with special regard to nature reserves. Parki Narodowe i Rezerwaty Przyrody 26 (1): [In Polish with English abstract] JASKUŁA R. & SOSZYŃSKA-MAJ A What do we know about winter active ground beetles (Coleoptera, Carabidae) in Central and Northern Europe? In: KOTZE D. J., ASSMANN T., NOORDIJK J., TURIN H. & VERMEULEN R. (eds), Carabid Beetles as Bioindicators: Biogeographical, Ecological and Environmental Studies. ZooKeys 100: KAŹMIERCZAK T Entomofagi Ips typographus (Coleoptera, Scolytidae) w rezerwacie Madohora (Beskid Mały) w latach Pp In: MAZUR S., SKŁODOWSKI J. & WOJCIECHOWSKA A. (eds), VI Sympozjum Ochrony Ekosystemów Leśnych Waloryzacja ekosystemów leśnych metodami zooindykacyjnymi Jedlnia 2 3 grudnia 1996r. Fundacja Rozwój SGGW, 242 pp. KINELSKI S. & SZUJECKI A Contribution to the knowledge of Polish Coleoptera. Polskie Pismo Entomologiczne 29: [In Polish with English summary] KONDRACKI J Geografia fizyczna Polski. PWN, Warszawa, 441 pp. KOTZE D. J., BRANDMAYR P., CASALE A., DAUFFY-RICHARD E., DEKONINCK W., KOIVULA M. J., LÖVEI G. L., MOSSAKOWSKI D., NOORDIJK J., PAARMANN W., PIZZOLOTTO R., SASKA P., SCHWERK A., SERRANO J., SZYSZKO J., TABOADA A., TURIN H., VENN S., VERMEULEN R. & ZETTO T Forty years of carabid beetle research in Europe from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation. In: KOTZE D. J., ASSMANN T., NOORDIJK J., TURIN H. & VERMEULEN R. (eds), Carabid Beetles as Bioindicators: Biogeographical, Ecological and Environmental Studies. ZooKeys 100: KOWALCZYK J. K., MAJECKI J., SOSZYŃSKI B. & ŚLIWIŃSKI Z Fauna bezkręgowców In: KUROWSKI J. K. (ed.), Park Krajobrazowy Wzniesień Łódzkich. EKO-WYNIK, 183 pp. KOWALCZYK J. K. & WATAŁA C Interesting species of ground-beetles (Coleoptera, Carabidae) in Łódź Upland. Przegląd Zoologiczny 30: [In Polish with English summary] KUBISZ D. 2004a. Carabus variolosus Fabricius, Pp In: ADAMSKI P., BARTEL R., BERESZYŃSKI A., KEPEL A. & WITKOWSKI Z. (eds), Poradniki ochrony siedlisk i gatunków Natura 2000 podręcznik metodyczny. T.6. Gatunki zwierząt (z wyłączeniem ptaków). Ministerstwo Środowiska, Warszawa. KUBISZ D. 2004b. Carabus zawadzkii Kraatz, Pp In: ADAMSKI P., BARTEL R., BERESZYŃSKI A., KEPEL A. & WITKOWSKI Z. (eds) Poradniki ochrony siedlisk i gatunków Natura 2000 podręcznik metodyczny. T.6. Gatunki zwierząt (z wyłączeniem ptaków). Ministerstwo Środowiska, Warszawa. KURZAWA J Diagnoza, założenia koncepcji generalnej Planu Ochrony w części dotyczącej entomofauny (Chrząszcze) Spalskiego Parku Krajobrazowego. Plan Ochrony SpPK. Etap II. Spalski Park Krajobrazowy, Spała (manuscript) LEŚNIAK A. 2001a. Ground beetles (Carabidae, Coleoptera) of Świnia Góra Reserve in the Świętokrzyskie Mountains. Fragmenta Faunistica 44: LEŚNIAK A. 2001b. Entomofauna naziemna Białych Ługów. Pp In: ŻUREK S. (ed.), Rezerwat torfowiskowy Białe Ługi. Wydawnictwo Domini, Kraków 208 pp. LEŚNIAK A Communities of terrestrial entomofauna with special emphasis on Carabidae (Coleoptera) in the protected and manager forests in Świętokrzyski Region. Leśne Prace Badawcze 3: [In Polish with English abstract] LEŚNIAK A. & GÓRECKA J Carabidae, Coleoptera of Cisów reserve. Rocznik Świętokrzyski, Ser. B. Nauki Przyrodnicze 27: [In Polish with English summary] LORENZ W Systematic list of the extant ground beetles of the world (Insecta. Coleoptera Geadephaga: Trachypachidae and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). Lorenz, Tutzing, 502 pp. LORENZ W Nomina Carabidarum. A directory of the scientific names of ground beetles (Insecta, Coleoptera, Geadephaga: Trachypachidae and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). Lorenz, Tutzing, 993 pp. LÖVEI G. L. & SUNDERLAND K. D Ecology and behavior of ground beetles (Coleoptera: Carabidae). Annual Review of Entomology 41: MACIEJOWSKI W Wpływ cech środowiska przyrodniczego na rozmieszczenie wybranych grup chrząszczy (Coleoptera) w południowej części Wyżyny Krakowsko-Częstochowskiej. Wydawnictwo Uniwersytetu Jagiellońskiego, 177 pp. NIETUPSKI M., CIEPIELOWSKA D. & KOSEWSKA A. 2008a. Assemblages of epigeic Carabidae (Col.) in a peatbog nature reserve situated in an urban area. Polish Journal of Natural Sciences 23 (3): NIETUPSKI M., KOSEWSKA A. & CIEPIELOWSKA D Comparison of assemblages of Carabidae (Coleoptera) in a peatbog reserve Redykajny with urban woods of Olsztyn. Wiadomości Entomologiczne 25 (Supl. 1): [In Polish with English abstract] NIETUPSKI M., KOSEWSKA A., CIEPIELOWSKA D. & SĄDEJ W. 2008b. Assemblages of Carabidae in a forest peat bog reserve situated in an urban agglomeration. Sylwan 11: [In Polish with English summary] PAWŁOWSKI J. 2004a. Carabus clatratus Linnaeus, Pp In: GŁOWACIŃSKI Z. & NOWACKI J. (eds) Polish Red Data Book of Animals. Invertebrates. Instytut Ochrony Przyrody PAN. 445 pp.