© Entomologica Fennica. 31 August 2016

Evaluation of ecological niches of abundant species of and Pterostichus (Coleoptera: Carabidae) in forests of steppe zone of

Viktor V. Brygadyrenko

Brygadyrenko V. V. 2016: Evaluation of ecological niches of abundant species of Poecilus and Pterostichus (Coleoptera: Carabidae)in forests of steppe zone of Ukraine. — Entomol. Fennica 27: 81–100. This article analyses abundances of seven Poecilus and Pterostichus species sampled from 836 forest sites located in five geographical regions of the steppe zone of Ukraine. The abundances are analysed in relation to eight forest condi- tions. New information about associations of the following five species was found. Poecilus versicolor was most abundant in sparse forest sites and on moist sandy soil. Pterostichus melanarius reached its maximum abundance on sites with a thick litter layer, mesohygrophilous condition, clay soils and a sparse her- baceous layer. Pterostichus niger reached maximum abundance in forests with mesohygrophilous and hygrophilous conditions and on sandy loam soils. Abun- dance of Pterostichus oblongopunctatus was at the highest on sites with a closed tree canopy, a thin litter layer and loamy soils of average salinity. Pterostichus ovoideus was most abundant in forests with low to average litter depth, hygrophilous conditions and loamy soils of low to average salinity. For Poecilus sericeus and Poecilus cupreus the results of this study agreed with the published information. V. V. Brygadyrenko, Department of Zoology and Ecology, Oles’ Honchar Dni- propetrovsk National University, pr. Gagarina, 72, Dnipropetrovsk, 49010, Ukraine; E-mail: [email protected] Received 21 November 2015, accepted 22 March 2016

1. Introduction fying limiting ecological factors can only be ad- dressed through extensive and carefully planned Many species of ground-dwelling invertebrates field research. This requires a very large number are very sensitive to changes in environmental of sample plots, preferably in the order of magni- conditions, while others are less demanding. For tude of several hundreds (Brygadyrenko 2006). every species in every ecosystem, there is a spe- This gives the best chance of finding sites which cific set of limiting factors (Thiele 1977). How- collectively represent the full range of values for a ever, because of the sheer variety of elements in a studied ecological factor in a given climatic zone given ecosystem, it is exceedingly difficult to or geographical region. identify precisely which of the many factors is a The range of tolerance of certain ground bee- probable, as opposed to merely a potential, limit- tle species for a particular ecological factor can ing factor (Hutchinson 1957). The task of identi- vary considerably. Thus, simple, unambiguous 82 Brygadyrenko • ENTOMOL. FENNICA Vol. 27 assessments of a species’ preferences (for exam- al. 2014, 2015). Accordingly, moisture is the fun- ple, moisture conditions)do not often provide an damental limiting factor. A dense tree crown accurate description of the situation (Thiele 1977, layer and a thick litter layer can act as consider- Kryzhanovskij 1983). Rather, each ecological able compensating factors for the lack of moisture factor should be defined more precisely through in the surrounding environment. The influences application of a range of gradations and this of moisture, tree crown density and litter depth should be incorporated into the research. This is are modified by the herbaceous layer, soil texture inseparably linked with the number of studied and salinity (Brygadyrenko 2015a). All these fac- sample sites and the species’ frequency of occur- tors may vary spatially and temporally and cause rence (i.e.% of the sampled plots where a given their influence on the ground-dwelling inverte- species is found). Of course, it is easier to numeri- brates, including ground . In addition, the cally assess the relationship of abundant ground presence of ants shows a strong correlation with species to a certain ecological factor than the composition of populations the relationship of rare ones. This is why 7 com- (Hawes et al. 2002, Reznikova & Dorosheva mon species were chosen for this study (Sharova 2004). The aim of this article is to find associa- et al. 1998, Brygadyrenko 2003). The analyzed tions of the 8 above-mentioned ecological factors species were Poecilus cupreus Linnaeus, 1758, with the 7 ground beetle species of the Ptero- P. versicolor (Sturm, 1824), P. sericeus (Fischer stichini tribe, which are dominant in forest eco- von Waldheim, 1823), Pterostichus melanarius systems of the steppe zone of Ukraine. (Illiger, 1798), P. niger (Schaller, 1783), P. ob- longopunctatus (Fabricius, 1787)and P. ovoi- deus (Sturm, 1824). 2. Materials and methods For studying occurrences of the 7 abundant ground beetle species, eight ecological factors The material was collected from 836 sample plots important in forest ecosystems were examined: (Fig. 1), located in five geographical regions type of forest, tree crown density, herbaceous (Dnipropetrovsk, Zaporizhia, Mykolaiv, Do- layer density, litter depth, moisture conditions, netsk and Kharkiv)of Ukraine’s steppe zone be- soil texture and salinity, and abundance of ants. In tween 2001 and 2014 (Brygadyrenko 2014, conditions of insufficient moisture in the steppe 2015a, b, c). These plots varied considerably in zone of Ukraine, populations of invertebrate spe- size, according to the extent of the forest or tree cies are subjected to severe stress (Kul’bachko et line plantation where they were located. In most

Fig. 1. Map of sample plots indicated by a star, located in five geo- graphical regions of Ukraine: Dnipro- petrovsk (568 plots), Zaporizhia (70), Mykolaiv (21), Donetsk (83) and Kharkiv (94). ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 83

Table 1. Gradients of ecological factors with numbers of study plots (total 836) in forest ecosystems of the steppe zone of Ukraine. For explanations of gradients, see Fig. 2.

Gradient Tree Type of Density of Thickness Moisture Soil Soil Abun- of factor crown forest herbaceous of conditions texture salinity dance density ecosystem layer litter of ants

1 197 27 249 334 266 125 176 428 2 46 128 143 234 305 51 125 229 3 118 681 70 173 87 125 464 110 4 277 n. a. 134 53 107 535 71 45 5 198 n. a. 240 42 71 n. a. n. a. 24 n. a. – not analysed. cases, usually in forests, the plot was about (1950)for the steppe zone of the Soviet Un - 10,000 m2, whereas in linear windbreak planta- ion: 1: xeromesophilous, 2: mesophilous, 3: tions the plot sizes varied from 1,000 to 15,000 hygromesophilous, 4: mesohygrophilous, 5: m2. This paper is a part of a wider study which hygrophilous) took into account the origin of the forests – Texture of the soil horizon (from samples of (whether they are natural forests, usually not pri- 20 cm in depth, obtained from the soil dug for meval but at least three generations old, or planta- each of the 10 pit-fall traps per plot: 1: sandy, tions, the oldest of which dates from the 1890s), 2: sandy loam, 3: loam, 4: clay) geomorphological factors (whether they are – Soil salinity (using the same herbaceous layer floodplain forests, ravine forests, or plateau fo- indicator species that were used for defining rests), nature of the soil (chernozem, floodplain the moisture conditions, according to soils of different types, with different depths of Belgard’s (1950)scale: 1: trophotopes AB, B, humus horizon), phytocenotic characteristics C; low salinity, 2: Dc, Dac; average salinity or (composition of the tree, shrub and herbaceous slightly below, 3: Dn; above average salinity, layers)and type of forest ecosystem. All the plots 4: De, E; high to extremely high salinity) studied in this paper were classified using the fol- – Abundance of ants (according to the results of lowing eight ecological factors: counts from pit-fall traps on a logarithmic scale: 1: £ 4, 2: 5–16, 3: 17–64, 4: 65–256, 5: – Tree crown density in percentage (average of ³ 256 individuals/10 trap-days) five random measurements for each collec- tion plot: 1: £ 20%, 2: 21–40%, 3: 41–60%, 4: The number of study plots according to each eco- 61–80%, 5: ³ 81%) logical factor and its gradient in forest ecosystems – Type of forest ecosystem (1: pine, 2: mixed, is shown in Table 1. 3: broad-leaved forest) The ground beetles were collected using 10 – Density of the herbaceous layer in percentage pit-fall traps in each plot, made each made of 0.5 l (average of five measurements for each col- glass jar (with 20% NaCl solution). They were lection plot: 1: £ 20%, 2: 21–40%, 3: 41– placed in a line at a distance of 2–10 m from one 60%, 4: 61–80%, 5: ³ 81%) to another. Depending on the amount of precipi- – Depth of the litter in millimeters (average of tation, the traps were checked every 5 d. This arti- ten random measurements for each collection cle analyses the results from each collection site plot; analysis made using five 10 mm classes for a 20 d period close to the middle of June, the of depth: 1: £ 10 mm, 2: 11–20, 3: 21–30, 4: exact dates varying from year to year. This partic- 31–40, 5: ³ 41 mm) ular period was chosen because in steppe forests – Moisture conditions (using indicator species there is still a high number of species with a of the herbaceous layer, the same as used in spring period of activity (Brygadyrenko 2015a, assessment of the density of this layer, ac- b), and at the same time, a sufficiently high num- cording to the scale devised by Belgard ber of species with a summer-autumn period of 84 Brygadyrenko • ENTOMOL. FENNICA Vol. 27

Table 2. Statistical significances of associations of Pterostichini beetles with eight ecological factors in forest eco- systems of the steppe zone of Ukraine based on the analyses of 836 samples from pitfall traps. For exact ANOVA results, see Figs. 2–8.

Species Tree Type of Density of Thickness Moisture Soil Soil Abun- crown forest herbaceous of conditions texture salinity dance density ecosystem layer litter of ants

Poecilus cupreus * n.s. n.s. n.s. n.s. n.s. n.s. n.s. P. versicolor * n.s. n.s. n.s. * * n.s. n.s. P. sericeus n.s. n.s. n.s. n.s. n.s. n.s. n.s. n.s. Pterostichus melanarius n.s. * * * * * * n.s. P. niger n.s. n.s. n.s. n.s. * * n.s. n.s. P. oblongopunctatus * n.s. n.s. * n.s. * * n.s. P. ovoideus n.s. n.s. n.s. * * * * n.s.

* and n.s.: significant (P < 0.00625) and insignificant (P ³ 0.00625) association of the ecological factors after Bonferroni correc- tion for eight ANOVAs for each species. activity. For the analyses, the numbers of each was associated with the abundance of 5 of the 7 species of ground beetle at each plot were calcu- studied species, moisture conditions were associ- lated for uniform time periods (individuals per 10 ated with the abundance of 4, soil salinity, tree trap-days), i.e. the total numbers from the 10 traps crown density and depth of the litter were associ- of the 20 d collection periods were divided by ated with the abundance of 3, and density of the twenty. herbaceous layer and type of forest ecosystem The number of individuals for the different were correlated with the abundance of 1 species. factor gradations was compared using ANOVA The abundance of the analyzed species varied in Statistica 8.0. A large number of ANOVAs (8 significantly in response to 0–6 out of the 8 ana- for each ground beetle species, Figs. 2–8)was lyzed ecological factors. Pterostichus melanarius performed, so Bonferroni tests were applied to was the species with most associations (6)with adjust the P values. Therefore, in Figs. 2–8 only P the studied ecological factors. values less than 0.00625 (=0.05/8)were consid- The abundance of Poecilus cupreus was sig- ered statistically significant. nificantly higher in conditions of low tree crown In order to find out relative similarity of the density (up to 20%, Fig. 2a). Its frequency of oc- distributions of the ground beetle species in the currence in the total number of plots was 2.6% forest ecosystems of the steppe zone of Ukraine, a and its average abundance in the total number of cluster analysis with linkage distance was per- plots was 0.027 ± 0.228 individuals/10 trap-days. formed (single linkage, Euclidean distances, Poecilus versicolor reached its maximum Statistica 8.0). For this analysis, the raw data of abundance on sites with 20–40% tree crown den- the abundances were normalized: (xi–x)/SD, sity (Fig. 3a), hygrophilous moisture conditions where xi is the number of individuals for the sepa- (Fig. 3e)and on sandy soils (Fig. 3f).Its fre - rate ecosystems, x is the average number of indi- quency of occurrence was 7.6% and the average viduals for all ecosystems, and SD is the standard abundance was 0.081 ± 0.792 individuals/10 deviation of the number of individuals for all fo- trap-days. rest ecosystems. Poecilus sericeus showed no significant cor- relation in abundance with any of the studied eco- logical conditions. It was slightly more numerous 3. Results than average in areas with low tree crown density (Fig. 4a), with average numbers of ants (Fig. 4h), Table 2 provides an overall comparison of the and in xerophilous and mesophilous moisture distribution of the different ground beetle species conditions (Fig. 4e). Its frequency of occurrence of the tribe Pterostichini in the studied plots in the was 2.1%. Its average abundance was 0.052 ± forests of the steppe zone in Ukraine. Soil texture 0.551 individuals/10 trap-days. ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 85

Fig. 2. Association of forest ecosystem condi- tions with Poecilus cupreus as means ±1.96·SD of numbers of ground beetles / 10 trap-days, and results of ANOVAs. – a. Tree crown density (1: £ 20%, 2: 21–40%, 3: 41–60%, 4: 61–80%, 5: ³ 81%).–b.Typeoffo- rest ecosystem (1: pine, 2: mixed, 3: broad- leaved forest). – c. Density of herbaceous layer cover (1: £ 20%, 2: 21–40%, 3: 41–60%, 4: 61–80%, 5: ³ 81%). – d. Thickness of litter (mm) (1: £ 10, 2: 11– 20, 3: 21–30, 4: 31–40, 5: ³ 41). – e. Moisture conditions (1: xero- mesophilous, 2: meso- philous, 3: hygomeso- philous, 4: mesohygro- philous, 5: hygro- philous). – f. Soil tex- ture (1: sandy, 2: sandy loam,3:loam,4:clay). – g. soil salinity (1: trophotopes, low salin- ity, 2: average salinity or slightly below, 3: above average salinity, 4: high to extremely high salin- ity ). – h. Abundance of ants as individuals / 10 trap-days (1: £ 4, 2: 5– 16, 3: 17–64, 4: 65– 256, 5: ³ 256). ANOVA results shown in bold are considered signifi- cant, after Bonferroni correction for eight ANOVAs; in these cases, letters a and b mark differences be- tween the values (Tukey’s test).

Pterostichus melanarius reached its maxi- ous forests (Fig. 5b), on soils with a raised level of mum abundance in deep litter (30–40 mm, Fig. salinity (Fig. 5g)and in sites with low levels of 4d), in mesohygrophilous moisture conditions herbaceous cover (<20%, Fig. 5c). Its frequency (Fig. 5e)and on clay soils (Fig. 5f).The species of occurrence was 20.0% and its average abun- was also significantly more abundant in decidu- dance was 0.49 ± 2.17 individuals/10 trap-days. 86 Brygadyrenko • ENTOMOL. FENNICA Vol. 27

Fig. 3. Association of forest ecosystem condi- tions with Poecilus versicolor, and results of ANOVAs. For expla- nations, see Fig. 2.

Pterostichus niger was significantly more 0.455 individuals/10 trap-days. abundant in mesohygrophilous and hygrophilous Pterostichus oblongopunctatus varied signif- moisture conditions (Fig. 6e)and on sandy loam icantly in abundance in relation to 5 out of the 8 soils (Fig. 6f). Its frequency of occurrence was analysed ecological factors. Its abundance was 6.9% and its average abundance was 0.050 ± higher in conditions of 40–100% tree crown den- ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 87

Fig. 4. Association of forest ecosystem condi- tions with Poecilus sericeus,andresultsof ANOVAs. For explana- tions, see Fig. 2. sity (Fig. 7a), loamy soils (Fig. 7f), average soil In P. ovoideus, four out of the eight analyzed salinity (Fig. 7g)and thin litter layer (10–20 mm, factors showed a significant association with the Fig. 7d). Its frequency of occurrence was 19.5% abundance of the species. It reached maximum in and its average abundance was 0.29 ± 1.33 indi- sites with low to average litter depth (10–20 mm, viduals/10 trap-days. Fig. 8d), in hygrophilous moisture conditions 88 Brygadyrenko • ENTOMOL. FENNICA Vol. 27

Fig. 5. Association of forest ecosystem condi- tions with Pterostichus melanarius, and results of ANOVAs. For expla- nations, see Fig. 2.

(Fig. 8e), on loamy soils (Fig. 8f) of low to aver- similarity in distribution occured between P. age salinity (Fig. 8g). Its frequency of occurrence cupreus and P. niger (Fig. 9). The least similarity was 12.4% and its average abundance was 0.33 ± with other species of ground beetles was ob- 2.23 individuals/10 trap-days. served among P. melanarius, P. ovoideus and P. According to the cluster analysis, the greatest oblongopunctatus. ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 89

Fig. 6. Association of forest ecosystem condi- tions with Pterostichus niger, and results of ANOVAs. For explana- tions, see Fig. 2.

4. Discussion and review zhanovskij et al. 1995, Hurka 1996, Freude et al. 2004). It forms the basis of the ground beetle 4.1. Poecilus cupreus fauna in the fields of and Asia (Kry- Poecilus cupreus is a West Palaearctic species, zhanovskij 1983, Kromp 1990, Lys & Netwig reaching Central Siberia and Central Asia (Kry- 1991, Karpova & Matalin 1993, Soboleva- 90 Brygadyrenko • ENTOMOL. FENNICA Vol. 27

Fig. 7. Association of forest ecosystem condi- tions with Pterostichus oblongopunctatus, and results of ANOVAs. For explanations, see Fig. 2.

Dokuchaeva 1995, Sharova et al. 1998). Accord- radically distributed in the Balkans (Guéorguiev ing to Lindroth (1986), it is distributed through- & Guéorguiev 1995, Hristovski & Guéorguiev out Europe, south to the Mediterranean; Asia Mi- 2015)at altitudes of 0 to 1,300 m. It occurs nor; Siberia east to River Lena. In P. throughout Ukraine (Putchkov 2011, 2012): P. c. cupreus is ubiquitous (Serrano 2013). It is spo- cupreus Linnaeus, 1758 in the Transcarpathian ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 91

Fig. 8. Association of forest ecosystem condi- tions with Pterostichus ovoideus, and results of ANOVAs. For explana- tions, see Fig. 2. lowland, Carpathian mountains, forest and forest- and fields, preferably on clayey soil with rather steppe zone, and P. c. dinniki Lutchnik, 1912 in dense vegetation of grasses and sedges” (Lind- the steppe zone and Crimean peninsula. roth 1986). According to Hellqvist (2014), the In Fennoscandia and P. cupreus is species inhabits clayey, not too dry soils in Swe- common but local “on open, not too dry meadows den. 92 Brygadyrenko • ENTOMOL. FENNICA Vol. 27

Fig. 9. Results of cluster analysis (single linkage, Euclidean distances) of the distribution of the ground beetle species of the tribe Pterostichini studied in forest ecosystems of the steppe zone of Ukraine.

In Great Britain P. cupreus is found “in open, mesophile, inhabiting steppe areas and pastures, not too dry meadows and fields, sometimes near ravine forests and floodplain forests, agricultural water; common, but local” (Lindroth 1974). Luff and urban landscapes. (1992)states that P. cupreus “is common in In Ukraine P. cupreus is an eurybiontic spe- southern Britain, but much more local in the north cies, which reaches its greatest abundance within and in Ireland. It is found in open, moderately dry the steppe zone in fields and meadows (Bryga- and warm habitats such as short grass and agricul- dyrenko 2000, 2003). It is considered a predator, tural fields”. which sometimes damages sprouted seeds of Hurka (1996)writes that in the Czech and corn, peas, haricot, beet, garden radish, clover, lu- Slovak Republics P. cupreus is a “eurytopic spe- cerne, maple, hornbeam, ash, young sprouts of cies, very common in unshaded habitats: fields, potatoes, fruit pulp of strawberries, tomatoes, and steppe, water edges; lowlands to mountains”. It fallen fruit in gardens (Petrusenko & Petrusenko accounted for 2.6% of the total number of ground 1973). In the southern part of its range, in beetles collected from Acer campestre, Carpinus Ukraine’s steppe zone, the species occurs in all betulus and Fraxinus angustifolia forest in the types of ecosystems: from shores of reservoirs, Ranšpurk National Nature Reserve (Southern including salines (Brygadyrenko 2000), to Moravia)(Šejnohová 2006).In the Bialowieza xeromesophilous and mesoxerophilous steppe Primeval Forest (Northern )the species is sites and sparse, dry plantations (Brygadyrenko rare, just 0.1% of the total count of ground beetles 2003). Here its range of moisture preferences is (Sklodowski 2006). In P. cupreus was dom- very wide. The thickness of litter in different sites inant (over 17% of the total count of ground bee- inhabited by P. cupreus also differs: from com- tles)in all restored habitats in an agricultural area plete absence of litter on agricultural fields and of the Po Plain (Pilon et al. 2013). areas of sand steppe, to thick shoreline drifts of In , in the Lower Volga Region, P. cu- decaying Phragmites australis (Cav.)Trin. ex preus is seen “everywhere, often, in agro- Steud. Only a small proportion of the P. cupreus coenoses, an abundant species” (Kaljuzhnaja et populations in the steppe zone of Ukraine inhab- al. 2000). In the Republic of Adygheya, it is its forests; more individuals are recorded annu- “abundant. A polytopic mesophile which inhabits ally in fields and near water. Because of the spe- a wide range of zonal communities, from cies’ well-developed flight capacity, one cannot bottomland areas near floodplain rivers to the consider the forest individuals of this species to subalpine zone. Is often found in agrocoenosis. belong to separate populations or subpopulations. Flies towards light. Is considered a pest” (Za- Our data contained positive correlation of the motajlov & Nikitsky 2010). Sigida (1993) states species’ abundance with sparse tree crown cover. that in Ciscaucasia, the species is a polytopic This correlation was found to be very strong. ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 93

4.2. Poecilus versicolor The species is common on fields of Russia, both in European (Kromp 1990, Karpova & Poecilus versicolor is a Palaearctic species, dis- Matalin 1993)and Asian side (Shilenkov 1978). tributed in most of Europe, the Caucasus, Siberia In the Lower Volga Region, P. versicolor occurs and Japan (Lindroth 1986, Kryzhanovskij et al. everywhere, being common in the north-west re- 1995, Hurka 1996). It is sporadically distributed gion but rare in the southern part (Kaljuzhnaja et in the Balkans (Guéorguiev & Guéorguiev 1995, al. 2000). In the Republic of Adygheya, P. Guéorguiev 2007), occurring at altitudes from versicolor “is quite common, inhabits forest ar- 300 to 2,000 m (Hristovski & Guéorguiev 2015) eas, most commonly found in areas of mixed fo- and it has a limited distribution also in Spain rests, seen in steppified areas, in agrocoenoses, (Serrano 2013). It has been also recorded in the considered a pest” (Zamotajlov & Nikitsky Transcarpathian lowland, Carpathian mountains, 2010). Sigida (1993) states that in Ciscaucasia the and forest and forest-steppe zone of Ukraine species is classified as a polytopic mesophile (Putchkov 2011, 2012). which inhabits natural steppe areas and pastures, Lindroth (1986)writes that in Fennoscandia ravine and floodplain forests as well as agricul- and Denmark P. versicolor occurs “on different tural and urban landscapes. Poecilus versicolor is kinds of open ground, e.g. meadows, fields and a predator but the beetles sometimes damage pulp heaths; often also on cultivated soil. It is more of mushrooms, fruits of tomatoes, sprouted seeds xerophilous than P. cupreus, preferring rather of hornbeam, and fallen fruit in gardens (Petru- dry, sandy soil with scattered vegetation. Occa- senko & Petrusenko 1973). sionally found together with P. cupreus on clay- In Ukraine, P. versicolor is an eurybiontic mixed sandy soil”. In Great Britain P. versicolor species, which is most abundant in meadows, is “more xerophilous than P. cupreus, in all kinds agrocoenoses, in steppe areas and in forests of the of open fields; common but possibly becoming steppe zone (Brygadyrenko 2000, 2003). It toler- rarer” (Lindroth 1974). According to Luff ates very strong anthropogenic transformation of (1992), in Britain P. versicolor “is especially forest ecosystems; as a forest edge species it is of- common in the wetter, western areas. It is found ten abundant on sites affected by fire, and in clear in open habitats such as grassland (including up- felled areas (Moroz et al. 2011). In Ukraine’s land moors)and arable land, especially if either steppe zone, the species is often found on shores near water or with high rainfall”. of reservoirs, including those with a high level of Hurka (1996)writes that in the Czech and salinity (Brygadyrenko 2000). In the south of Slovak Republics P. versicolor is “common in Ukraine, the species is a eurybiont. Forest eco- unshaded habitats: meadows, pastures, fields, systems are habitats where the species can sur- vegetated water margins, forest clearings; low- vive through unfavourable weather conditions lands to mountains, mostly in hills”. When study- (for example summer droughts); a considerable ing the ecotonic effect of P. versicolor distribu- number of individuals are concentrated in clear- tion in a forest clearing, Brigiæ et al. (2014)found ings and at forest edges. that the species was attracted to meadowland In the present study, the species showed unex- (8.94% of the total count of ground beetles), and pectedly a significant increase in abundance in ar- not to the forest of Fagus sylvatica L. and Abies eas of high tree crown density. This contrasts alba Mill. (0.02% of the total count of ground with our findings for P. cupreus, which it strongly beetles). In the Bialowieza Primeval Forest resembles morphologically. Besides this, P. ver- (Northern Poland), the species was abundant (on sicolor was found to be more abundant on sandy average 2.1% of the total count of ground bee- soils. tles), replacing ground beetle species typical of standard forests (Sklodowski 2006). The species 4.3. Poecilus sericeus is common (over 2.5% of the total count of ground beetles)in restored habitats in an agricul - Poecilus sericeus is a West Palaearctic species, tural area of the Po Plain (Lombardy, Italy)(Pilon reaching Central Siberia and Central Asia et al. 2013). (Kryzhanovskij et al. 1995, Hurka 1996). Freude 94 Brygadyrenko • ENTOMOL. FENNICA Vol. 27 et al. (2004)writes that P. sericeus extends from Amur River in Siberia, and introduced in North South-West Europe to Siberia, and has numerous America (Lindroth 1986, Kryzhanovskij et al. subspecies. It does not occur in Great Britain 1995, Hurka 1996, Freude et al. 2004). It forms (Lindroth 1974), Fennoscandia and Denmark the basic element of the ground beetle fauna in (Lindroth 1986), (Guéorguiev 2007) and fields (Kromp 1990, Lys & Netwig 1991, Kar- (Guéorguiev & Guéorguiev 1995). The pova & Matalin 1993, Soboleva-Dokuchaeva species is found only in 4 of the 93 geographical 1995, Sharova et al. 1998, Eyre et al. 2009). The regions of the Republic of Macedonia (Hristovski subspecies, P. m. bulgaricus Lutshnik, 1915, is & Guéorguiev 2015)and 2 of the 23 geographical distributed in and much of Bulgaria, oc- regions of Spain (Serrano 2013). Petrusenko & curring at altitudes from 0 to 2,290 m (Guéor- Petrusenko (1973)write that P. sericeus is a pre- guiev & Guéorguiev 1995). The rest of Europe is dator which sporadically damages sprouting inhabited by the nominate subspecies, P. m. seeds of peas, haricot, beet, lupins, cabbage melanarius (Illiger, 1798), being rare in southern sprouts, fruit pulps of strawberries, tomatoes, Europe (Guéorguiev 2007, Serrano 2013, Hri- fallen drops in gardens. stovski & Guéorguiev 2015). The species occurs Hurka (1996)writes that in the Czech and throughout Ukraine (Putchkov 2011, 2012). It is Slovak Republics P. sericeus is rare, inhabiting a predator, but, in conditions of insufficient mois- “unshaded, dry habitats: steppe, fields; lowlands ture, eats sprouts of corn, soy-beans, beet, lettuce, to hills”. The species is found in fields there sprouting seeds of maple, hornbeam, ash, fruits of (Karpova & Matalin 1993). blackberries, strawberries, stone berries, and In Russia, in the Republic of Adygheya, it is fallen fruits in gardens (Petrusenko & Petrusenko “common, steppe mesophile, is often found in 1973, Kryzhanovskij 1983). agrocoenoses, is considered a pest of many agri- Lindroth (1986)writes that in Fennoscandia cultural crops” (Zamotajlov & Nikitsky 2010). In and Denmark P. melanarius is “very eurytopic, the Lower Volga Region, P. sericeus is common usually occurring on open and not too dry and and ubiquitous (Kaljuzhnaja et al. 2000). Sigida sandy ground, e.g. meadows and grassland. It has (1993)states that in Ciscaucasia P. sericeus is clearly been favoured by human cultivation, be- classified as a steppe mesophile, inhabiting natu- ing a common inhabitant of arable land, parks, ral steppe, pastures, and also urban landscapes. gardens, etc. It occurs also in forest edges and The species occurs throughout Ukraine ex- light woodland, living in company with eurytopic cept the Transcarpathian lowland and the Car- forest-dwelling Carabidae, e.g. P. niger”. Lind- pathian mountains (Putchkov 2011, 2012). In roth (1974)writes that in Great Britain P. mela- Ukraine’s steppe zone, P. sericeus is most abun- narius inhabits “all kinds of open, not too dry dant on ploughed sites, and slightly less so in country; common though somewhat local”. Luff steppe and meadow ecosystems, river valleys and (1992)states that in Britain and Ireland it is wide - in agrocoenoses (Brygadyrenko 2000, 2003). spread and common, “it prefers more open habi- The species dominates among ground beetles on tats than P. madidus, and is commoner in non-ba- the driest sites. In general, this is due to the de- sic grasslands and cultivated fields, where it can crease in the number of abundant, eurybiontic often be the dominant species”. ground beetle species. It does not often penetrate Hurka (1996)writes that in the Czech and into forests and stays near forest edges or in wind- Slovak Republics P. melanarius is a “very com- breaks with open tree canopies. Overall, the data mon, very eurytopic species of fields, meadows, from the current study confirm the information gardens, as well as forests; lowland to moun- presented in the existing literature. tains”. It accounted for 5.5% of the total number of ground beetles collected in Acer campestre, 4.4. Pterostichus melanarius Carpinus betulus and Fraxinus angustifolia fo- rest in the Ranšpurk National Nature Reserve Pterostichus melanarius is a Eurosiberian spe- (Southern Moravia)(Šejnohová 2006).The spe - cies, distributed across most of Europe, except cies accounted for 7.9% of the ground beetles col- the far north and south, reaching eastward to the lected in a beech wood (Melittio-Fagetum)in ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 95

Northern (Andorko & Kadar 2006). In fer in untreated and treated barley fields in the largest archipelago of islands in Poland, on and accounted for 18.4–30.5% of the the Mazurian Lake Mamry, P. melanarius ac- overall number of ground beetles, regardless of counted for over 30% of the total count of ground whether or not the agricultural crop was treated beetles, the species being found on 15 of the 17 is- with Chlorpyrifos-methyl (Kos et al. 2010). lands investigated (Zalewski & Ulrich 2006). The Studies of the trophic specialization of this species is dominant in forests in Northeastern Po- species in relation to invertebrates has allowed us land, accounting for 13.7% of the total number of to identify which invertebrate species it con- ground beetles (Kwiatkowski 2011). In the sumes or only consumes to a low extent (Korolev Bialowieza Primeval Forest (Northern Poland) & Brygadyrenko 2012, 2014). The abundance of the species is very numerous (about 5.5%), dis- the species in steppe zone forests, which are not placing species of ground beetle which are char- damaged by recreational pressure, is usually low. acteristic inhabitants of protected forests (Sklo- Partial destruction of vegetation and leaf litter, dowski 2006). In the species inhabits pollution of a territory by industrial and domestic sandy, not too dry soils (Hellqvist 2014). This wastes lead to an increase in the number of P. species is common (over 2% of the overall count melanarius with a corresponding reduction of the of ground beetles)in restored habitats in an agri - number of other ground beetle species (Bryga- cultural area of the Po Plain (Lombardy, Italy) dyrenko et al. 2012, Brygadyrenko & Korolev (Pilon et al. 2013). 2015). The species occurs everywhere in Elek & Lövei (2005)studied the distribution Ukraine, except in the driest habitats. Its abun- of this species in the context of urbanization and dance can remain high even when other ground found that in natural forests in Denmark it ac- beetle species disappear or significantly decrease counts for 14.8% of the total number of ground in numbers. Increased abundances of P. melana- beetles, in suburban forests 5.9%, and in city rius are found in areas which are close to high- parks 40.6%. Matalin (2006)emphasizes that one ways and in city parks (Brygadyrenko & of the reasons for the high tolerance of this spe- Chernysh 2003, Brygadyrenko et al. 2012). The cies for anthropogenic influences is the variabil- species is regularly found near water, in mead- ity of its life cycle. ows, in reeds, tolerates moderate salinization of In Russia, in Tatarstan, P. melanarius occurs the soil (Brygadyrenko 1999, 2000). Natural un- in forests and damp meadows (Zherebcov 2000). damaged forest ecosystems do not play a signifi- In the Lower Volga Region P. melanarius is cant role in supporting the generally high abun- ubiquitous, but not often seen (Kaljuzhnaja et al. dance of this species in Ukraine’s steppe zone. 2000). Zamotailov & Kryzhanovskij (1992) state The data from the current study confirm the infor- that in the Caucasus the species occurs from mation presented in the existing literature. bottomland forests to subnival belt; is common in forests in the foothills and middle altitudes of the mountain zone and in highly degraded Alpine 4.5. Pterostichus niger pastures as well as in cultivated land; at altitudes of 2,000–2,100 m; the species accounts for 15% Pterostichus niger is a Palaearctic species, dis- of the total number of ground beetles. It is com- tributed across most of Europe, except in the far mon in the Republic of Adygheya. “A eurybiont north and south, ranging eastward to Iran, West- species, distributed from bottomland forests to ern Turkestan and East Siberia (Lindroth 1986, the subnival zone at altitudes ranging from 200– Kryzhanovskij et al. 1995, Hurka 1996, Freude et 2,100 m, often in cultivated grounds, in ruderal al. 2004). In Albania (Guéorguiev 2007) and landscapes, is seen in agrocoenosis. Is considered Macedonia it is rare (Hristovski & Guéorguiev a pest of a number of agricultural crops” (Za- 2015), but recorded in 17 of the 30 geographical motajlov & Nikitsky 2010). Sigida (1993) men- regions of Bulgaria at altitudes from sea level up tions that in Ciscaucasia the species is a polytopic to 2,500 m (Guéorguiev & Guéorguiev 1995). mesophile inhabiting fields and urban land- The species is distributed in Spain, being found in scapes. The abundance of the species did not dif- 10 of the 23 geographical regions of the country 96 Brygadyrenko • ENTOMOL. FENNICA Vol. 27

(Serrano 2013). It has been recorded in all parts of an agricultural area of the Po Plain (Italy)(Pilon Ukraine except for the steppe zone of the Cri- et al. 2013). mean Peninsula (Putchkov 2011, 2012). Elek & Lövei (2005)studied the distribution Pterostichus niger is a predator, but in conditions of the species in the context of urbanization and of insufficient moisture, it eats fruit pulps of found that in natural forests of Denmark the spe- strawberries (Petrusenko & Petrusenko 1973). cies accounts for 2.9% of the total number of In Fennoscandia and Denmark P. niger is a ground beetles, in suburban forests 0.6%, and in widely distributed and common “eurytopic spe- city parks 0.7%. cies, especially characteristic of woodland, oc- In Russia, in the Lower Volga Region, P. curring in almost every type of forest community, niger is ubiquitous but uncommon (Kaljuzhnaja predominantly in deciduous and mixed stands of et al. 2000). In the Republic of Adygheya, it “is a humus-rich, rather moist soil. Also in hedges, polytopic mesophile, which inhabits a wide range parks and gardens; less frequent in meadows and of plain and mountain zonal communities, is on arable land” Lindroth (1986). In Sweden the found in agrocoenoses, considered as a pest of a species inhabits humus-rich moist soils (Hell- number of agricultural crops” (Zamotajlov & qvist 2014). Lindroth (1974) writes that in Great Nikitsky 2010). Sigida (1993) states that in Britain P. niger is found “usually in parks, thin Ciscaucasia, the species is a polytopic mesophile forests, etc., on not too dry soil, often under bark; which lives in urban landscapes. common”. Luff (1992)points out that P. niger “is In Ukraine’s steppe zone, the species is con- common throughout Britain, with most Irish re- centrated mostly in the forests and on the shores cords coming from the north-east of the country. of reservoirs. The abundance of the species does It is found in woodland and damp grassland, and not considerably differ between populated areas on upland moors.” and natural ecosystems. This is one of the species In Central Europe P. niger is fairly frequent, with the highest flight migration activity, thus preferring moderately damp coniferous forests within a season its abundance can greatly change (Freude et al. 2004). Hurka (1996) states that in in different forest types. Although P. niger and P. the Czech and Slovak Republics the species is melanarius are very close morphologically and in “common in moist habitats, indifferent to shade: size, in the steppe zone they differ substantially in meadows, forests, water margins with vegetation; flight capacity. All imagines of the former can fly, lowlands to mountains, frequent in hills”. It ac- whereas only 2% of imagines of the latter can do counted for 28.7% of the overall number of so (Brygadyrenko, unpublished data). ground beetles collected in Acer campestre, An unexpected finding of the current research Carpinus betulus and Fraxinus angustifolia fo- is that P. melanarius was significantly more rest in the Ranšpurk National Nature Reserve abundant on clay soils, whereas P. niger was sig- (Southern Moravia)(Šejnohová 2006).It is a nificantly more abundant on sandy loam soils. A dominant species in the forests of Northeastern second surprise, with no indication in the litera- Poland, accounting for 27.8% of the total number ture or in our previous research, is that P. mela- of ground beetles (Kwiatkowski 2011). In the narius was significantly more abundant in areas Bialowieza Primeval Forest (Northern Poland) of very sparse (<20%)herbaceous layer, while P. the species accounted for 14.8% of the overall niger, seemed to be more abundance on sites with number of ground beetles (Sklodowski 2006). a dense herbaceous layer (60–80%), although the The species is not often found on fields (Lys & latter was not statistically significant. Netwig 1991, Karpova & Matalin 1993, Sobo- leva-Dokuchaeva 1995). On untreated and treated barley fields in Croatia the abundance of 4.6. Pterosctichus oblongopunctatus the species fell from 4.6% and 18.0% of the total number of ground beetles to 0% when the crops Pterostichus oblongopunctatus is a Transpalae- were treated with Chlorpyrifos-methyl (Kos et al. arctic species, distributed from North and Central 2010). The species is common (3.7% of the total Europe to the Caucasus and Siberia, and to Japan number of ground beetles)in restored habitats in in the East (Kryzhanovskij et al. 1995, Hurka ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 97

1996). In Central Europe, the species is common tus “is common, inhabits forest zone, often is in forests (Freude et al. 2004). It is sporadically found in more damp areas, considered as a pest of distributed in the Balkans between 300 and 1,900 a number of agricultural crops” (Zamotajlov & m above sea level (Guéorguiev & Guéorguiev Nikitsky 2010). Sigida (1993) states that in 1995, Guéorguiev 2007, Hristovski & Guéor- Ciscaucasia, the species is a forest mesophile of guiev 2015). In Spain it has been recorded only in the mountain complex, belonging to the ecologi- one region (Serrano 2013). The speceis occurs cal groups of halohygrophiles, reophiles and throughout Ukraine (Putchkov 2011, 2012). The stagnophiles. beetles sometimes eat pulp of boletus, edible In Ukraine’s steppe zone, P. oblongopuncta- mushrooms, and also the fruits of strawberries tus is one of the species most adaptable to differ- (Petrusenko & Petrusenko 1973). ent types of forest habitats, including those that Lindroth (1986)writes that in Fennoscandia have been considerably transformed by human and Denmark the species is “a eurytopic wood- actions (Brygadyrenko 1999, 2003, Moroz et al. land species, occurring in both deciduous and co- 2011). In forests of steppe zone, the species has a niferous forests, usually in light stands on moder- very wide range of diet, which can be compared ately dry, mainly sour humus soil. In the Atlantic in variability to that of P. melanarius (Komarov climate of West also in open country”. & Brygadyrenko 2011). Beyond forest ecosys- According to Koivula et al. (1999), in Central tems, the species is rarely found on river banks, in this species significantly decreases in meadows near forest edges, in city parks with abundance in forests with a thin litter layer. sparse tree cover. However, in the south Ukraine According to Lindroth (1974) P. oblongo- it prefers forest areas with no steppe plants, with punctatus is, in Great Britain, “a forest species; on typical forest microclimate conditions and a de- all kinds of soil, often under bark; common, but veloped layer of forest litter. The results of the local” while Luff (1992)writes that it is classified present study agree with this information re- as Nationally Scarce (B), it “inhabits woodland ceived in previous studies (Brygadyrenko 2003). and forests, occurring over much of Britain. It has also been recorded from Ireland. Its distribution is rather patchy, with four or five apparent centres 4.7. Pterostichus ovoideus of distribution, and it is very local in Ireland. It generally shows a preference for rather open Pterostichus ovoideus is a Eurosiberian species, woods on dry soils.” distributed from the Pyrenees through Central Hurka (1996)writes that in Czech and Slovak Europe to the Caucasus and Siberia (Kryzha- Republics P. oblongopunctatus “is very com- novskij et al. 1995, Hurka 1996, Freude et al. mon, found in all types of forests; lowlands to 2004). It has been recorded in 6 of the 30 geo- mountains, frequent in hills”. The species ac- graphical regions of Bulgaria, at altitudes ranging counted for 3.8% of the overall number of ground from 200 to 1,200 m (Guéorguiev & Guéorguiev beetles in beech woods (Melittio-Fagetum)and 1995). The species has been recorded throughout oak forest (Querco-petreae-Carpinetum)in Ukraine except for the southern steppe subzone Northern Hungary (Andorko & Kadar 2006). It is (right bank of the Dnieper, westwards)and the a dominant species in the forests of Northeastern Crimean Peninsula (Putchkov 2011, 2012). Poland, making up 6.7% of the total ground bee- Hurka (1996)states that in the Czech and Slo - tle population (Kwiatkowski 2011). The abun- vak Republics P. ovoideus is “common in rather dance of this species is high in the Bialowieza Pri- moist habitats, indifferent to shade: meadows meval Forest (Northern Poland)with 18.8% of near water, edges of water with vegetation, flood- the overall number of ground beetles (Sklodow- plain forests, groves; lowlands to mountains, fre- ski 2006). quently in hills”. In Western Croatia, P. ovoideus In Russia, in the north and north-west parts of accounts for 2.1% of the total count of ground the Lower Volga Region, the species is common, beetles in meadows, while it is absent 30–50 m but it is rare in the south (Kaljuzhnaja et al. 2000). into Fagus sylvatica L. and Abies alba Mill. fo- In the Republic of Adygheya, P. oblongopuncta- rest (Brigiæ et al. 2014). The species is also absent 98 Brygadyrenko • ENTOMOL. FENNICA Vol. 27 from forests and clearings, though often occur- The abundance of the analyzed species varied ring in grassland communities 50 m from the fo- significantly in association with from 0 to 6 out of rest in the North Hungarian Mountains (Andorko the 8 analyzed ecological factors. Pterostichus & Kadar 2006). Kaljuzhnaja et al. (2000)write melanarius varied significantly in abundance in that in Russia, in the Lower Volga, P. ovoideus is association with all the factors, except for the rare with only one population known. In the Re- abundance of ants and for the tree crown density. public of Adygheya (North-West Caucasus), it is According to the cluster analysis, the greatest common, “a polytopic mesophile, belonging to a similarity in distribution among the studied wide range of zonal communities of the plains Pterostichini beetles occurs between P. cupreus and lower mountains, found in agrocenoses, flies and P. niger, and the least similarity with other towards light” (Zamotajlov & Nikitsky 2010). species was observed among P. melanarius, P. Sigida (1993)states that in Ciscaucasia the spe - ovoideus and P. oblongopunctatus. cies is a polytopic mesophile, inhabiting ravine The patterns of habitat preferences identified and floodplain forests and agricultural land- in this study for beetles of the genera Poecilus and scapes. Pterostichus may not apply to all parts of their Outside forest ecosystems of the steppe zone range. It would be interesting to identify the zonal of Ukraine, P. ovoideus is fairly common on the differences in ecological niches of these species banks of reservoirs, in reeds and on meadows in the north, central and southern parts of their (Brygadyrenko 2000, 2003). As with the pre- range using the methods described in this paper vious species, our data from the current study do and compare those data with the ones I have ob- not contradict the information presented in pre- tained. In different parts of a species’ range, the vious studies on the ecological preferences of P. importance of each limiting factor can vary con- ovoideus. siderably because of the very different ecological characteristics in each area.

5. Conclusions Acknowledgements. I would like to thank Paul Bradbeer and two anonymous referees for valuable comments on the In forests of the steppe zone of Ukraine, P. manuscript. cupreus is more abundant in conditions of low tree crown density. Poecilus versicolor reaches References its maximum abundance in conditions of 20–40% of tree crown density, in hygrophilous moisture Andorko, R. & Kadar, F. 2006: Carabid beetle (Coleop- conditions, on sandy soils, with low soil salinity. tera: Carabidae)communities in a woodland habitat in Poecilus sericeus is more abundant in areas with Hungary. — Entomologica Fennica 17: 221–228. low tree crown density and with average numbers Belgard, A. L. 1950: (The forest vegetation of the south of ants. The abundance of P. melanarius reaches east of the Ukraine.)— Kiev University Press, Kiev, its maximum in conditions of 40–60% of tree Ukraine. 264 pp. [In Russian.] Brigiæ, A., Starèeviæ, M., Hrašovec, B. & Elek, Z. 2014: crown density, deep litter, in mesohygrophilous Old forest edges may promote the distribution of forest moisture conditions, on clay soils and with low species in carabid assemblages (Coleoptera: Carabi- densities of herbaceous cover. Pterostichus niger dae)in Croatian forests. — European Journal of Ento - is more abundant in mesohygrophilous and mology 111: 715–725. hygrophilous moisture conditions and on sandy Brygadyrenko, V. V. 1999: (Regularities of distribution of the ground beetles of the tribe Pterostichini (Coleop- loam soils. The abundance of P. oblongopuncta- tera, Carabidae)from the floodplain and arena ecosys - tus increases in conditions of 40–100% tree tems of the Samarskiy Bor.)— The Kharkov Entomo - crown density, loamy soils, average soil salinity, logical Society Gazette 7: 72–74. [In Russian.] thin litter layer and hygromesophilous moisture Brygadyrenko, V. V. 2000: (The ground beetles (Coleop- conditions. The abundance of P. ovoideus is tera, Carabidae)of the nature reserve “Bulakhovskiy Liman” (Dnepropetrovsk region).) — The Kharkov greatest at sites with low to average litter depth, in Entomological Society Gazette 8: 86–94. [In Russi- hygrophilous moisture conditions, on loamy an.] soils, with low to average soil salinity. Brygadyrenko, V. V. 2003: (Fauna of ground beetles (Co- ENTOMOL. FENNICA Vol. 27 • Forest preferences of carabids in Ukraine 99

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