© Entomologica Fennica. 31 August 2016
Evaluation of ecological niches of abundant species of Poecilus and Pterostichus (Coleoptera: Carabidae) in forests of steppe zone of Ukraine
Viktor V. Brygadyrenko
Brygadyrenko V. V. 2016: Evaluation of ecological niches of abundant species ofPoecilus 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 ofPterostichus 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 oflow to average salinity. For Poecilus sericeus and Poecilus cupreus the results ofthis study agreed with the published information.
’ V. V. Brygadyrenko, Department ofZoology 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 ofmagni- 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 ofvalues for a ever, because ofthe sheer variety ofelements 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 oftolerance 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- a]. 2014, 2015). Accordingly, moisture is the fun- ple, moisture conditions) do not often provide an damental limiting factor. A dense tree crown accurate description ofthe situation (Thiele 1977, layer and a thick litter layer can act as consider- Kryzhanovskij 1983). Rather, each ecological able compensating factors for the lack ofmoisture 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). Ofcourse, it is easier to numeri- brates, including ground beetles. In addition, the cally assess the relationship of abundant ground presence of ants shows a strong correlation with beetle species to a certain ecological factor than the composition of ground beetle populations the relationship of rare ones. This is why 7 com- (Hawes et a]. 2002, Reznikova & Dorosheva mon species were chosen for this study (Sharova 2004). The aim of this article is to find associa- et a1. 1998, Brygadyrenko 2003). The analyzed tions ofthe 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. 0b- 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) ofUkraine’s steppe zone be- soil texture and salinity, and abundance ofants. In tween 2001 and 2014 (Brygadyrenko 2014, conditions of insufficient moisture in the steppe 2015a, b, c). These plots varied considerably in zone ofUkraine, 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
Kharkiv
Fig. 1. Map of sample plots indicated by a star, located in five geo- graphical regions of Ukraine: Dnipro— petrovsk (568 plots), Zaporizhia (70), 100 km Mykolaiv (21), Donetsk (83) and Kharkiv (94). ENTOMOL. FENNICA Vol. 27 ° Forestpreferences ofcarabids 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 ofthe 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 (chemozem, 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: Do, Dac; average salinity or (composition of the tree, shrub and herbaceous slightly below, 3: Dn; above average salinity, layers) and type offorest ecosystem. All the plots 4: De, E; high to extremely high salinity) studied in this paper were classified using the fol- — Abundance ofants (according to the results of lowing eight ecological factors: counts from pit-fall traps on a logarithmic scale: 1: S 4, 2: 5—16, 3: 17—64, 4: 65—256, 5: — Tree crown density in percentage (average of 2 256 individuals/ 10 trap-days) five random measurements for each collec- tion plot: 1: S 20%, 2: 21—40%, 3: 41—60%, 4: The number of study plots according to each eco- 61—80%, 5: 2 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 ofthe herbaceous layer in percentage pit-fall traps in each plot, made each made of0.5 1 (average of five measurements for each col- glass jar (with 20% NaCl solution). They were lection plot: 1: S 20%, 2: 21—40%, 3: 41— placed in a line at a distance of 2—10 m from one 60%, 4: 61—80%, 5: 2 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 (1 period close to the middle of June, the ofdepth: 1: S 10 mm, 2: 11—20, 3: 21—30, 4: exact dates varying from year to year. This partic- 31—40, 5: 2 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. versico/or 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 2 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 ofthe 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 ofANOVAs (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 ofthe steppe zone ofUkraine, a and its average abundance in the total number of cluster analysis with linkage distance was per- plots was 0.027 d: 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 ofindividuals 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 d: 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 ofthe 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 ofthe different ground beetle species conditions (Fig. 4e). Its frequency of occurrence ofthe tribe Pterostichini in the studied plots in the was 2.1%. Its average abundance was 0.052 d: forests ofthe steppe zone in Ukraine. Soil texture 0.551 individuals/ 10 trap-days.
86 Brygadyr'enko *9 ENTOOL. FE
H4. 833) m 8.01, p < 0.001 H2, 835) = 0.20, p 2 0.819 1.0 ti 0.0 ‘
0.8 '
0.7 '
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0.3 '
0.2 '
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Tree croum density Type of forest ecosystem
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' 0.03 " 0.04 / " trap 0.00 i 0.00 10 1 2 3 4 5 / Density of herbaceous layer Thickness of litter beetles H4, 833) a 5.54, p < 0.001 H3, 834) a 4.00, p m 0.002 0.7 0.50 of ~-a '
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M 0i sture conditions Soil texture
3 1:73.834): 2.07, p 0.031 H4, 833) a 1.25, p m 0.288 0.40
0.38 ‘
0.32 '
0.28 '
0.24 ‘-
020 ' 0.18 ' Fig. 3. Association of 0.12 ' forest condim " ecosystem 0.08
* tions with Poeciius 0.04 0.00 versicoior, and results 5 of ANOVAS. For expla— Soil salinity Abundance at 0013 nations, see Fig. 2.
Prerosrichus Niger“ was significantly more 0.455 individuals/l 0 trap—days. abundant in mesohygmphilous and hygrophilous .Pz‘emsz‘ichus Oblongopuncmrm varied signifi— moisture conditions (Fig. 6e) and on sandy loam icantly in abundance in relation to 5 out of the 8 soils (Fig. 6t). its frequency of occurrence was analysed ecological factors. its abundance was 6.9% and its average abundance was 0.050 i higher in conditions of40ml 00% tree crown den- Vol. 27 a Fcrestpreferences afcambids in Ukraine 87
3 H4, 333) 2 3.20, p == 0.011 b H2. 335) 2 0.35. p = 0.422
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.\ » 0.02 ~ 0.02 \ g \\ \ . ~ -~ ~ - t ‘3 t . - m 0-00 0.00 2- 1 2 3 4 5 1 2 3 4 5 “*2... 8 Density of herbaceous Bayer Thicmess of litter ‘5 2 g 6 H4, 333) 2 0.33. p 2 0.443 f F(3.334):1.42, p 0.234 . . . 4.... 0.13 . = = . e 0.13 O
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g as, 334) 2 2.03, p 2 0.103 h H4, 333) 2 2.39., p 2 0.013
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,. - . . / \1 - :, sericeus, and results of 0.00 0,00 . , 1 2 3 4 1 2 3 ‘4' 5 ANOVAs. For explanam Soil Abundance of ants tions, see Fig. 2. salinity sity (Fig. 7a)) loamy soils (Fig. 73, average soil In P. maidens, four out of the eight analyzed salinity (Fig. 7g) and thin litter layer (102-20 mm.) factors showed a significant association with the Fig. 7d). its frequency of occurrence was l9.5% abundance ofthe species. it reached maximum in and its abundance was 0.29 i 1.33 indi- sites with low to litter l 0—20 average average depth ( ’9 viduals/l0 trapndays. Fig. 8d), in hygrophflous moisture conditions 88 Brygadyrenko *9 ENTO
a H4. 033) m 3.10. p m 0.013 b H2. 035) m 0.23. p a 0.002
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Tree crown density Type of forest ecosystem
a 0 H4, 033) m 3.00. p m 0.003 d H4. 033) 17.42, p < 0.001
~ 1.0 2.4 - '
- a b
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-
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m u : Densityof herbaceous layer Thacmess of latter (0 d) g 8 H4, 033) a mo. )3 «a: 0.001 f H3. 034) a 5.37. p m 0.001 1.0 0.0 b b _ f - 1 o 1.4 0.7
- . a b
- 1.2 - 0.0 g n ab “
~ - 1.0 - 0.5 /
~ — ~48 - 0.8 4 0.4 0 /
-
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/i\\ - 0.1 0.2- ______
~
- . . . . ._ 0.0 0.0 i; . {/i 1 2 3 4 5 1 2 3 4
Moisture conditions Soil texture
9 H3. 034) m 7.21, p «a: 0.001 h H4, 033) m 2.00, p m 0.005
0.9 , = = . 0.0 - = ~ -
-
.7 _
_ 0.0
_. a? ab
“ 30 0.5 » _ \ 0' - 0.4
_ 0': ' - a 0.3 -- 0‘3 _ Fig. 5. Association of / *
~ _ 0.2 forest condim 02 \3 \ : ecosystem
~ ~ ~ 0.1 0.1 _ tions with Pierostichus
- ~ 0.0 . 0.0 . . \\ . (“rm—s meianarius, and results 1 2 3 4 ”l 2 3 4 5 01 ANOVAs. For explam SO“ Abundance 0f ants Salinity matiofis5 see Fig“ 2
(Fig. 80), on loamy soils (Fig. SD of iOW to aver- similarity in distribution occured between P. age salinity (Fig. 8g). its frequency ofoccurrence cupreus and P. Niger (Fig. 9). The ieast similarity was 12.4% and its average abundance was 0.33 :l: with other species of ground beetles was obn- 2.23 individuals/10 trapudays. served among P. meianarius. P. ovoideus and. P. According to the cluster analysis, the greatest oblongopmcmms. V01. 27 a F074657 preferences afcambids in Ukraine 89
3 H4, 833) 3 0.86, 1:) 3 0.488 b H2. 335) z 0.50, p 2 0.307 0.21 0.20
0.18 ' 0.18 " 0.16 ‘ 0.15 ‘ 0.14-
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Tree crown density Type of forest ecosystem
H4, 333)=1.77, p = 0.133 d H4, 333)=1.77, p = 0.133 0.24 0.27
- 0.21 " 0.24
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-
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r ' ' ' 0.00 1 0.00 1/1 1 trap 1 2 3 4 5 1 2 3 4 5 10 Density of herbaceous Sayer Thickness of litter
H4, 333) a 4.00, p a 0.003 f H3. 334) a 5.02, p < 0.001 0.30 0.50
- " -—b beeties/ 0.27 0.45 of i.) 13 — 0.24 " 0.40
— No. 0.21 " 0.35 0.18 “ 0.30 -
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Moisture conditions Soil texture
1:13.334): 0.51, p 2 0.507 h H4, 333) = 0.50, p = 0.555 0.20 0.16 ' 0.18 '
- 0.14 0.16-
- 0.12 0.14 *
' 0.10 ' 0.12
* 0.10' 0.08 0.08 ‘ 3x Fig. 6. Association of 0.06 ‘ 0.06“ forest condim ' ecosystem 0.04 ‘
}/ ' 0.04
" tions with Prerosz‘ichus - 0.02 0.02
' niger, and resuits of 0.00 T 0.00 - 2 3 4 1 2 3 4 5 ANOVAS. For explanau Soil Abundance of ants time, see Fig. 2. salinity
zhanovskij 61‘ 0d. 1995, Hurka 19969 Freude er ed. 2004). it toms the basis of the ground beetle 4.1. Paeeiius cupreus fauna in the fields of Europe and Asia Poecflus capreus is a West Paiaearctic species, zhanovskij 19839 1990, Lys & Netwig reaching Central Siberia and Central Asia (_ s 1991. Karpova & Matahn 1.993, Soboieva~ 90 BWgadyrenkg “ 0L.
H4, 333) a 4.52, p a 3.3m F(2,335):1.73, p 2 0.133 0.8 0.0
'
0.7 ' 0.5 0.0 * ab 0.4 * ' ah 0.5 ah (/0
" 0.4 ' 0.3
'
0.3 ' /.-...... _... 0.2 0.2 ' 0.1 ' 0.1 '
0.0 0.0 1 2 3 4 5 1fi/J2 3 Tree crown density Type of forest ecosystem
= H4, 333)=1.25, p 0.237 H4, 333) m 3.34. p < 3.331 0.7 0.8 '
'
0.0 " 0.7 ab
" 0.0 0.0 "
( " 0.4 " 0.5
0.4 ' * 0.3 /MW ‘ days ' 0.1 0.1 ' trap 0.0 0.0 1 2 3 4 5 10 / Densityof herbaceous layer Thicimess of litter H4, 333) z 2.33, p a 0.043 H3. 334) m 22.14, p < 3.331 beeties 1.0 1.4 of * b 0.0 ' 1.2 0.8 ' " No. ' 0.7 1.0 ' 0.0 0.8 " * " 0.0 0.4 ' 0.3 ' a gov/l 0.4 / 0.2 " 3- ' \a " 0.2 0.1 " 2/ ' 0.0 0.0 i 1 2 3 4 5 i 2 3 4 Moisture conditions Soil tenure H3. 334) m 22.42, p < 3.331 F(4,333):1.34, p a 0.254 " 1.4 b 0.0 ‘ " 1.2 ' 0.0 1.0 ' 0.4 " ' 0.8 -_ 7. Association of 0.3 ' Fig. 0.0 ' forest ecosystem condi- ' " 0.2 M\ 0.4 a \ tions with Prerostichus " 0.2 " 0.1 “met>42“ oblongopuncz‘aws, and ' \r 0.0 85 f 0.0 i resuits of ANOVAS. For i 2 3 4 1 2 3 4 5 explanations, see Sci! salinity Abundance of ants Fig. 2. Dokuchae‘va 19953 Sharova er al. 1998). Accord radicafly distributed in the Balkans (Guécrguie‘v ing to Lindroth (1986), it is distributed through—:- & Guéorguie‘v 1995, Hristovski & Guéorguiev out Europe. south to the Mediterranean; Asia Min 2015) at aitimdes of O to 1.300 m. it occurs Siberia east to River Lena. In P. Ukraine 201 1 P. c. nor; Spain throughout (Putchko'v , 2012): cupmus is ubiquitous (Serrano 2013). it is spou cupreus Li 1758 in the Transcarpathian VOL 27 a Fare“preferences incarabids in Ukraine 91 a F(4,833):1.83, p 3 0.112 b H2, 835) 3 2.03, p 3 0.133 ‘ ''' 0‘7 ' ' ‘ 0.0 - ' 0.8 ~ 0.6 0.5 - a 0’4 0.5 /<>\ - 0.4 0.3 _ _ \ 0'3 \C’ - /V 0.2 0.2 ' - 0.1 ' 0.1 " 0.0 . ' * E 0.0 ’i 2 3 4 5 Tree crown density Type of forest ecosystem a m 0 H4, 553) a 1.57. p a 0.155 d H4. 833) 4.05. p 0.003 4 . . . 0.7 . . - . 0‘9 _ "b - 0'8 0.5 ~ __ 0.7 - 0.5 ' ‘ ab 43 06 ab “ “ ' 0.4 / 0‘5 - __ a 0.3 - a - ' - 8: 0.2 “\ 0.2 ' a ' cs 04 0.1 - “U __ ‘ ' ' * . Q. 00 V/41 0.0 1 fl E 1 2 3 4 5 1 2 3 4 5 2 Densityof herbaceous layer Thickness Of litter E 5 H4, 533) m 3.54, p a 0.004 f H3. 534) a 17.05. p < 0.001 '5 1.4 - - = = 1.5 = - - : _ b " b 48 — 1.2 1 4 45 ' - ~ 1'2 - of“: 1.0 3 -» 2 1.0 / - - 0.8 a - ~ 5 0.8 / _ 0 ' 0 0.5 - a a ac _ a 0.4- — / 04 _ - ' C 0.2 0.2 , i ' i E ' 0.0 l 0.0 1 1 1 2 3 4 5 1 2 3 4 Moisture conditions Soil texture g H3, 334131000, p < 0.001 h H4, 8.33) a 0.04, p a 0.442 1.5 , . ~ . - . b 0.7 - 1.4 " 0.5 12 _ " - 0‘5 1.0 _ ' " - - 0‘4 0.5 // a — 8. Association of 0.5 — / 0.3 Fig. a : ~ - - forest ecosystem condi~ 0,4 - 0.2 ' — — tions with Pterostichus 0‘2 0.1 /\. and results of . . . . \i ovoideus, 0,0 \EK'EWM‘WE 0.0 . ANOVAS. For explana- 1 2 3 4 1 2 3 4 5 tions. see Fig. 2. Soil salinity Abundance of ants lowianda Carpathian mountains, forest and foresin and. fields? preferably on ciayey sofl 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 Heilqvist (2014), the ° in Fennoscandia and. Denmark P. cuprcus is abits clayeya not too dry sofls in Swen 001 on but local “on open, not too dry meadows 92 Brygadyrenko ° ENTOMOL. FENNICA V01. 27 Poecilus cupreus Pterostichus niger Poecilus sericeus Poeci/us versicolor Pterostichus oblongopunctatus Pterostich us ovoideus Pterostichus melanan'us 1O 15 20 25 30 35 40 45 50 55 Linkage Distance 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, fiuit pulp ofstrawberries, tomatoes, and steppe, water edges; lowlands to mountains”. It fallen fiuit in gardens (Petrusenko & Petrusenko accounted for 2.6% ofthe total number ofground 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 meinus angustifolia forest in the types of ecosystems: from shores of reservoirs, Ranspurk National Nature Reserve (Southern including salines (Brygadyrenko 2000), to Moravia) (Sejnohova 2006). In the Bialowieza xeromesophilous and mesoxerophilous steppe Primeval Forest (Northern Poland) the species is sites and sparse, dry plantations (Brygadyrenko rare, just 0.1% ofthe total count ofground beetles 2003). Here its range of moisture preferences is (Sklodowski 2006). In Italy P. cupreus was dom- very wide. The thickness oflitter 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 a]. 2013). areas of sand steppe, to thick shoreline drifts of In Russia, 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 ofUkraine 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.