Folia faunistica Slovaca 21 (1) 2016: –98

Winter observation of weevils
(Coleoptera, Curculionoidea) on Scots pines in Borská nížina lowland (SW Slovakia)

Milada Holecová1, Adrianna Králiková2, Katarína Hollá1
& Miroslava Šebestová1

1 Department of Zoology, Faculty of Natural Sciences, Comenius University,
Mlynská dolina, Ilkovičova 6, SK – 842 15 Bratislava, Slovakia
[holecova@fns.uniba.sk, holla@fns.uniba.sk, kupkova@fns.uniba.sk]

2 Department of Environmentalistics and Zoology, Faculty of Agrobiology and Food Resources, Slovak Agricultural University, A. Hlinku 2, SK – 949 76 Nitra, Slovakia [adrianna.kralikova@uniag.sk]

Abstract: During two winter seasons (2013–2014 and 2014–2015), we collected insects from branches of Scots pine trees in the Borská nížina lowland (SW Slovakia) using beating method. A total of 21 weevil species were collected at five localities and ten study plots. Pine specialists were represented by six species (Anthonomus phyllocola, Brachonyx pineti, Brachyderes incanus, Magdalis memnonia, Magdalis phlegmatica, and Pissodes castaneus). The rest of the winter fauna were tourist species. Three eudominant pine specialists (Anthonomus phyllocola, Brachonyx pineti, Brachyderes incanus) exhibited different trends in abundance during the examined period. The abundance of Brachyderes incanus gradually rose from November until to March. On the other hand, the higher abundance of Brachonyx pineti was observed in late autumnal and prevernal months (November and the beginning of March). Adults of Anthonomus phyllocola hatch in early March and therefore they were not observed during late autumnal and winter months. The highest species diversity and abundance of the hibernating weevil fauna was observed in younger and middle-aged, dense-canopied forest stands. They probably provide a better opportunity to shelter in comparison with forest patches and non-canopied, free-growing trees. It is likely that increased activity of leaf-eating insects in forest canopies in the non-growing season may be a result of global warming and climate change.

Key words: winter activity, weevils, Scots pine, Pinus sylvestris, Borská nížina lowland, SW Slovakia, global warming, climatic change.

Introduction

Scots pine (Pinus sylvestris L.) is one of the most widely distributed conifer species in the world, with an extensive natural range stretching from Spain to Norway and from Scotland to Siberia (Nikolov & Helmisaari 1992). Among numerous other organisms, it hosts many insects, often beetles living on needles or occupying a subcortical niche. While the faunistics of several conifer species have been investigated to some extent, the associated fauna of pine canopies, particularly in Europe, is rather poorly known.

The first complex studies on arboricolous arthropods including phyllophagous and xylophagous beetles on Pinus sylvestris were published namely from the western and northern Europe (Ozanne et al. 2000, Thunes et al. 2003, 2004). Weevil assemblages associated with Scots pine canopies in northern Poland were studied by Cholewicka-Wiśniewska (1994a, 1994b).

Currently, the most extensive pine stands in Slovakia are situated on the blown sands of the Borská nížina lowland as a part of the original mixed oak-pine forests. Pine stands of the Záhorie region, replacing the original ecosystems, for the most part, play an important role in soil protection in Borská nížina lowland. Our knowledge on Scots pine entomofauna has been poorly published in Slovakia. Only in the last years, from this territory there were published several faunistic and ecological papers concerning phyllophagous and xylophagous insects living on Scots pines, namely lepidopteran and sawfly larvae, weevils and bark beetles (e.g. Holecová & Kulfan 2011, Holecová et al. 2016, Kulfan & Holecová 2010, Kulfan et al. 2011, Kupková et al. 2014, Olšovský 2007, Olšovský et al. 2013).

The winter activity of phyllophagous insects associated with pine and other coniferous canopies in Slovakia is poorly known. There are only a few data about overwintering lepidopteran and sawfly larvae dwelling on branches of coniferous trees (e.g. Dvořáčková & Kulfan 2009, Parák et al. 2015, Šebestová et al. 2015, J. Kulfan et al. 2016, M. Kulfan et al. 2016). The winter activity of weevil beetles associated with coniferous canopies has not yet been studied.

The aims of our study are to:

■ characterize the structure of hibernating weevil assemblages associated with Scots pine canopies on the territory of the Borská nížina lowland;

■ analyze the winter weevil fauna in forest stands of different age, canopy and fragmentation as well;

■ find out whether there are differences in species diversity of weevils between individual forest stands and in individual months of the winter aspect;

■ compare qualitative and quantitative representation of pine species and tourists during the winter aspect.

Material and Methods

Area description

The study of weevils was carried out in Scots pine forests of different age and structure, growing on the sandy soils in the Borská nížina lowland, southwestern Slovakia (Tab. 1, Figs 1 and 2). The ten investigated study plots belong to the biotope of managed pine forests and semi-native pine-oak forests. The area is warm with moderately dry climate and mild winters, whereby the average temperature in January is usually above -3 °C. The average annual temperature is 9 °C and annual rainfall about 550 mm (Lapin et al. 2002). Study plots were visited during December 2013–March 2014 and November 2014–March 2015. Both of these periods of our field study were warmer comparing with the long-term average (1971–2000). The average air temperature went up by about 3.18 °C in the first research period (from 2.5 °C in December 2013 to 4.5 °C in March 2014) and about 2.8 °C (from 3.5 °C in November 2014 to 2.2 °C in March 2015) in the second research period. The number of days with the average temperatures below freezing point was the highest in January 2014 and February 2015 (9 days). Meteorological data (according to measurements of the station Moravský Svätý Ján) were provided by the Slovak Hydrometeorological Institute in Bratislava (Figs 3 and 4).

Sampling weevils

Weevil adults were sampled monthly by beating from Scots pine branches at heights of 1–3 m above the ground, using a beating tray with a diameter of 1 m. One sample consisted of weevils that had dropped into the beating tray from a total of 20 branches which were 1 m long each. In total, ten samples (200 branches) were taken every month from each study plot. The weevil beetles were preserved in 70% ethanol and examined in the laboratory, using a stereomicroscope Stemi 2000. The voucher specimens of all weevil species detected in the present study are deposited in the collection of the first author.

Data analysis

The species dominance is characterized by the scale proposed by Tischler (1949) and completed by Heydemann (1955): ED = eudominant, D = dominant, SD = subdominant, R = recedent, SR = subrecedent. The indices of Shannon-Wiener (H’, using natural logarithm of a number), Pielou (e) and Simpson (c) were used as the alpha diversity indices (Odum 1977, Spellerberg & Fedor 2003). All the couples of Shannon-Wiener’s diversity indices were compared with a t-test, to see if they are significantly different (Poole 1974). The weevil assemblages of individual study plots and months of a field study were compared by non-metric multidimensional scaling (nMDS) using computer programs NCLAS (Podani 1993) and STATISTICA (StatSoft Inc 2001). The analyses were based on Bray-Curtis dissimilarity index (Bray & Curtis 1957, Faith et al. 1987). The diversity of weevil assemblages was computed using the program PAST (Hammer et al. 2001).

Results and discussion

Hibernating assemblages

A total of 441 weevils belonging to 21 species and three families were collected at ten study plots (in the further text SP 1 – SP 10; Tab. 2). Despite the comparatively low average temperature on several sampling days, i.e. the temperature did not rise above 0 °C, all weevil adults were active during most of the time (Fig. 4, Tab. 2). Brachyderes incanus and Brachonyx pineti, two eudominant pine species, occurred on Scots pine branches during all months of the non-growing period. Adults of Brachyderes incanus are feeding on pine needles; their larvae are root-eating. Both the adults and larvae of Brachonyx pineti live and feed on pine needles (Scherf 1964). According to Dieckmann (1980, 1988), adults of two aforementioned weevils overwinter in the forest soil and leaf litter. Anthonomus phyllocola, the third eudominant pine specialist, was recorded only in early March (at study plots in surroundings of Lakšárska Nová Ves village; SP 1 – SP 4). Its larvae develop in the male flower of Scots pines, and adults can frequently be found feeding on current-year foliage in stands 10 years old and older (Lindelöw & Björkman 2001). According to our observations, this weevil species strongly preferred middle-aged canopies (SP 3).

At the study sites, from 3 to 9 weevil species were recorded. The highest diversity of overwintering species was observed in 8-years old, strong canopied Scots pine forest without contact with an open landscape (SP 9) (Table 3). On the other hand, the highest number of weevil individuals were observed in middle-aged, strongly canopied forests with or without contact with the non-forest environment (SP 2 – SP 4).

In individual months of the non-growing season from 4 (January) to 14 species (March) were found. The weevil assemblages were the most diversified in early spring (March). There is a highly significant difference (P < 0.001, t-test) between the diversity of this assemblage (H’ = 1.703, 14 spp.) and the assemblage diversities in the late autumnal and winter months (Tab. 4).

The weevil assemblages of individual study plots and months of our field study were compared by non-metric multidimensional scaling (nMDS). The scatter of the study site assemblages based on Bray-Curtis dissimilarity showed two groups of communities. The first one is formed by assemblages of five study sites (SP 1 – SP 5) with a predominance of pine specialists Anthonomus phyllocola and Brachonyx pineti. The second, different group on the diagram consists of assemblages of the rest study plots (SP 6 – SP 10) with higher abundance of pine monophage Brachyderes incanus (Tab. 2, Fig. 5).

Non-metric multidimensional scaling also confirmed the strong difference between weevil assemblages of the late autumn (November), early spring (March), and winter months (December, January, February) (Fig. 6).

The winter weevil fauna is represented by two groups of species – pine specialists and tourists which find shelter in dense pine canopies. With the exception of early spring (March) the pine specialists predominated qualitatively and quantitatively as well (Tab. 2, Fig. 7).

Three eudominant pine specialists (Anthonomus phyllocola, Brachonyx pineti, Brachyderes incanus) exhibited different trends in abundance during the examined period (Fig. 8). The abundance of Brachyderes incanus gradually rose from November until to March. On the other hand, the higher abundance of Brachonyx pineti was observed in late autumnal and prevernal months (November and early March). Anthonomus phyllocola emerges in early March and therefore its adults were not observed during late autumnal and winter months.

Faunistic data

Note: Weevil species living and feeding on Scots pines are highlighted in bold.

Apionidae

Catapion pubescens (Kirby, 1811)

SP 2: 5.12.2014, 1 ♂.

Catapion seniculus (Kirby, 1808)

SP 2: 21.11.2014, 1 ♂, 5.12.2014, 2 ♂.

Ceratapion gibbirostre (Gyllenhal, 1813)

SP 10: 6.3.2014, 1 ♀.

Oxystoma craccae (Linnaeus, 1767)

SP 6: 20.11.2014, 1 ♀.

Perapion curtirostre (Germar, 1817)

SP 6: 20.11.2014, 2 ♀.

Nanophyidae

Nanophyes marmoratus (Goeze, 1777)

SP 1: 17.1.2014, 1 ♀.

Curculionidae

Anthonomus phyllocola (Herbst, 1795)

SP 1: 15.3.2014, 2 ♂ 4 ♀, SP 2: 15.3.2014, 13 ♂, 4 ♀, 19.3.2015, 1 ♂, SP 3: 15.3.2014, 33 ♂, 27 ♀, 19.3.2015, 1 ♀, SP 4: 15.3.2014, 4 ♂, 7 ♀.

Brachonyx pineti (Paykull, 1792)

SP 1: 21.11.2014, 2 ♂, 6 ♀, 5.12.2014, 1 ♀, 15.3.2014, 2 ♂, 6 ♀, SP 2: 21.11.2014, 2 ♂, 3 ♀, 5.12.2014, 2 ♂, 10 ♀, 15.3.2014, 5 ♂, 1 ♀, SP 3: 21.11.2014, 2 ♂, 2 ♀, 15.3.2014, 23 ♂, 4 ♀, 19.3.2015, 1 ♂, SP 4: 21.11.2014, 1 ♂, 5 ♀, 15.1.2015, 3 ♂, 9 ♀, 19.2.2015, 1 ♀, 15.3.2014, 7 ♂, 3 ♀, 19.3.2015, 2 ♀, SP 5: 20.11.2014, 3 ♂, 5 ♀, 20.3.2015, 1 ♀, SP 6: 20.11.2014, 1 ♀, 11.3.2014, 2 ♀, SP 7: 20.11.2014, 1 ♀, SP 8: 10.3.2015, 1 ♂, SP 9: 7.3.2014, 1 ♂, 21.3.2015, 1 ♀, SP 10: 18.3.2015, 1 ♂.

Brachyderes incanus (Linnaeus, 1758)

SP 2: 20.2.2014, 1 ♂, 3 ♀, 21.11.2014, 1 ♂, 5.12.2014, 2 ♀, 15.3.2014, 1 ♀, 19.3.2015, 1 ♂, SP 3: 15.3.2014, 5 ♀, 19.3.2015, 2 ♀, SP 4: 21.11.2014, 1 ♂, 15.3.2014, 3 ♂, 3 ♀, 5.12.2014, 2 ♂, 15.1.2015, 2 ♂, 4 ♀, SP 5: 16.1.2014, 1 ♀, SP 6: 16.1.2014, 1 ♂, 4 ♀, 18.2.2014, 3 ♂, 6 ♀, 11.3.2014, 1 ♂, 5 ♀, 20.2.2015, 2 ♀, SP 7: 18.2.2014, 3 ♀, 11.3.2014, 2 ♀, 20.11.2014, 2 ♀, 8.12.2014, 1 ♂, 16.1.2014, 2 ♀, 20.2.2015, 1 ♀, 20.3.2015, 1 ♂, 1 ♀, SP 8, 19.1.2014, 2 ♀, 15.2.2014, 1 ♂, 5.3.2014, 3 ♀, SP 9: 28.12.2013, 5 ♂, 2 ♀, 23.1.2014, 1 ♂, 2 ♀, 7.3.2014, 6 ♂, 10 ♀, SP 10: 23.1.2014, 1 ♂, 2 ♀, 6.3.2014, 4 ♂, 6 ♀, 21.2.2014, 7 ♂, 9 ♀, 7.3.2014, 4 ♂, 5 ♀, 22.1.2015, 3 ♂, 1 ♀, 27.2.2015, 2 ♂, 1 ♀, 18.3.2015, 4 ♂, 3 ♂.

Ceutorhynchus pallidactylus (Marsham, 1802)

SP 1: 19.3.2015, 1 ♀, SP 8: 15.2.2014, 1 ♂, 5.3.2014, 1 ♀, 10.3.2015, 6 ♀, SP 9: 7.3.2014, 6 ♂, 10 ♀, SP 10: 21.2.2014, 1 ♂.

Dorytomus ictor (Herbst, 1795)

SP 8: 10.3.2015, 1 ♀, SP 9: 7.3.2014, 2 ♂.

Dorytomus melanophthalmus (Paykull, 1792)

SP 2: 5.12.2014, 1 ♂.

Gymnetron rostellum (Herbst, 1795)

SP 5: 11.3.2014, 1 ♂, SP 9: 7.3.2014, 1 ♂.

Gymnetron stimulosum (Germar, 1821)

SP 8: 5.3.2014, 1 ♂, SP 9: 7.3.2014, 1 ♂.

Magdalis memnonia (Gyllenhal, 1837)

SP 9: 19.1.2015, 1 ♀.

Magdalis phlegmatica (Herbst, 1797)

SP 1: 15.3.2014, 2 ♂, 1 ♀, SP 4: 20.2.2014, 1 ♀, SP 5: 11.3.2014, 2 ♂, SP 9: 21.3.2015, 1 ♂.

Pissodes castaneus (De Geer, 1775)

SP 2: 21.11.2014, 1 ♂, 5.12.2014, 2 ♀, 15.3.2014, 1 ♀, SP 4: 20.2.2014, 1 ♂, 15.3.2014, 1 ♀.

Rhinusa antirrhini (Paykull, 1800)

SP 8: 10.3.2015, 2 ♀, SP 9: 7.3.2014, 2 ♀.

Rhinusa hispida (Brullé, 1832)

SP 5: 18.2.2014, 1 ♂.

Sitona macularius (Marsham, 1802)

SP 7: 20.3.2015, 1 ♀, SP 10: 18.3.2015, 1 ♀.

Sitona suturalis (Stephens, 1831)

SP 4: 15.3.2014, 1 ♂.

The results of our preliminary study pose several questions. Is the present stage of knowledge concerning the leaf-eating insect activity in the non-growing season sufficient in Central Europe? Is the increasing winter activity of insects a normal phenomenon or it is a result of global warming and climatic change? Responses to winter conditions are not isolated from growing-season responses to climate. Therefore it is important to investigate the impacts of winter on performance, fitness and biotic interactions in the context of growing-season biology. We suggest that an opportunity exists to extend existing long-term studies of growing-season biology to incorporate the effects of winter. At the population and community levels, inter- and intra-specific interactions strongly influence responses to winter climate change. Impacts on individual species will propagate through ecosystems, and the role of winter in modifying these interactions must be considered when predicting the ecological impacts of climate change (Wiliams et al. 2015).

Acknowledgement

This study was published with the financial support of VEGA (Scientific Grant Agency of the Ministry of Education and the Slovak Academy of Sciences), grant number 1/0066/13 and 2/0035/13. We greatly acknowledge Juraj Holec for his help with the map design in ArcGIS application software.

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© Faunima, Bratislava, 2016

e–ISSN 1336–4529 ISSN 1335–7522

Holecová M et al.: Winter observation of Curculionoidea in Borská nížina lowland

Figure 1. Study area and position of ten study plots (design: Juraj Holec).

Figure 2. Studienka (SP 5) in the winter period 2015 (photo: Katarína Hollá).

Folia faunistica Slovaca 21 (1) 2016: 91–98

Table 1. Study plot characteristics.

Site	Study plot	GPS coordinates [WGS 84]	Altitude [m]	Study plot characteristics
Lakšárska Nová Ves	1	N 48° 34' 56,85'' E 17°10' 33.41''	222	About 25-year old pines free growing on sand dunes that gradually reach the adjacent stand about 100-year old.
Lakšárska Nová Ves	2	N 48° 34' 54,46'' E 17° 10' 34,56''	218	About 10-year old pines forming a dense forest stand close to a canopied stand.
Lakšárska Nová Ves	3	N 48° 34' 51,43'' E 17° 10' 22,78''	218	About 25-year old pines forming a forest stand wall adjacent to a meadow.
Lakšárska Nová Ves	4	N 48° 34' 55,81' E 17° 9' 52,23''	218	A dense forest stand of 15-year old pines, strongly canopied and without contact with open landscape.
Studienka	5	N 48° 32' 25,65'' E 17° 8' 29,88''	218	About 100-year old pines forming a stand with grassy undergrowth and surrounded by a meadow.
Studienka	6	N 48° 32' 16,49'' E 17° 8' 15,03''	218	About 15-year old pines growing in irregular clusters with grassy undergrowth.
Studienka	7	N 48° 32' 30,73'' E 17° 8' 13,49''	219	The youngest study plot with 5-year old pine trees, canopied and strongly insolated.
Lozorno	8	N 48° 20' 3'' E 17° 0,00' 56,1''	166	About 60-year old forest with grassy undergrowth and surrounded by agricultural land.
Pernek	9	N 48° 23' 16,5'' E 17° 6' 10,7''	203	About 8-year old Scotch pines forming a strongly canopied forest stand without grassy undergrowth.
Moravský Svätý Ján	10	N 48° 33' 52,4'' E 16° 59' 54,1''	159	About 10-year old pines forming a canopied forest stand wall, on one side surrounded by an forest path and on the other side by 80-100-year old forests.

Table 2. The cumulative number of weevil species collected at ten study plots (SP 1 – SP 10) during our field study.

% – dominance; CD – category of dominance: ED – eudominant, D – dominant, R – recedent, SR – subrecedent; BIO – bionomic groups: P – pine species, T – tourist species.

	1	2	3	4	5	6	7	8	9	10	Σ	%	CD	BIO
Apionidae
Catapion pubescens (Kirby, 1811)	0	1	0	0	0	0	0	0	0	0	1	0.23	SR	T
Catapion seniculus (Kirby, 1808)	0	3	0	0	0	0	0	0	0	0	3	0.68	SR	T
Ceratapion gibbirostre (Gyllenhal, 1813)	0	0	0	0	0	0	0	0	0	1	1	0.23	SR	T
Oxystoma craccae (Linnaeus, 1767)	0	0	0	0	0	1	0	0	0	0	1	0.23	SR	T
Perapion curtirostre (Germar, 1817)	0	0	0	0	0	3	0	0	0	0	3	0.68	SR	T
Nanophyidae
Nanophyes marmoratus (Goeze, 1777)	1	0	0	0	0	0	0	0	0	0	1	0.23	SR	T
Curculionidae
Anthonomus phyllocola (Herbst, 1795)	6	18	61	11	0	0	0	0	0	0	96	21.77	ED	P
Brachonyx pineti (Paykull, 1792)	17	23	33	31	9	2	1	1	3	1	121	27.44	ED	P
Brachyderes incanus (Linnaeus, 1758)	0	9	7	15	1	22	13	6	34	46	153	34.69	ED	P
Ceutorhynchus pallidactylus (Marsham, 1802)	1	0	0	0	0	0	0	8	16	1	26	5.90	D	T
Dorytomus ictor (Herbst, 1795)	0	0	0	0	0	0	0	1	2	0	3	0.68	SR	T
Dorytomus melanophthalmus (Paykull, 1792)	0	1	0	0	0	0	0	0	0	0	1	0.23	SR	T
Gymnetron rostellum (Herbst, 1795)	0	0	0	0	0	0	0	2	4	0	6	1.36	R	T
Gymnetron stimulosum (Germar, 1821)	0	0	0	0	1	0	0	1	2	0	4	0.91	SR	T
Magdalis memnonia (Gyllenhal, 1837)	0	0	0	0	0	0	0	1	2	0	3	0.68	SR	P
Magdalis phlegmatica (Herbst, 1797)	3	0	0	1	2	0	0	0	1	0	7	1.59	R	P
Pissodes castaneus (De Geer, 1775)	0	0	0	0	0	0	0	0	1	0	1	0.23	SR	P
Rhinusa antirrhini (Paykull, 1800)	0	4	0	2	0	0	0	0	0	0	6	1.36	R	T
Rhinusa hispida (Brullé, 1832)	0	0	0	0	1	0	0	0	0	0	1	0.23	SR	T
Sitona macularius (Marsham, 1802)	0	0	0	0	0	0	1	0	0	1	2	0.45	SR	T
Sitona suturalis (Stephens, 1831)	0	0	0	1	0	0	0	0	0	0	1	0,23	SR	T

Figure 3. Long-term average monthly temperatures (1971–2000) and average monthly temperatures in examined months and years of the research (according to the measurements of the meteorological station in Moravský Svätý Ján).

Figure 4. The number of days with an average temperature below freezing point in individual months of our field study.

Holecová M et al.: Winter observation of Curculionoidea in Borská nížina lowland

Folia faunistica Slovaca 21 (1) 2016: 91–98

Figure 5. Results of nMDS analysis based on Bray-Curtis distance showing dissimilarity of weevil assemblages in individual study plots. Numbers of study plots – see in Table 1.

Figure 6. Results of nMDS analysis based on Bray-Curtis distance showing dissimilarity of weevil assemblages in individual months of the non-growing season.

Table 3. Species diversity test and basic coenological characteristics of weevil assemblages at ten study plots during our field study.

Σ spp. – total number of species, Σ ind.– total number of individuals, c – Simpson’s index of dominance, e – Pielou’s index of equitability, H′ – Shannon-Wiener ’s index of diversity; Div. comp. – diversity comparison: t-test values are under the diagonal and degrees of freedom are above the diagonal. Significance levels: *** P < 0.001, ** P < 0.01, * P < 0.05, ns not significant). Numbers of the study plots see in the Table 1.

Study plot	1	2	3	4	5	6	7	8	9	10
Σ spp.	5	7	3	6	5	4	3	7	9	5
Σ ind.	28	59	101	61	14	28	15	20	65	50
c	0.69	0.765	0.778	0.695	0.701	0.531	0.442	0.8	0.659	0.242
e	0.429	0.276	0.476	0.353	0.449	0.635	0.76	0.27	0.343	0.848
H′	1.11	1.488	0.855	1.245	1.128	0.736	0.485	1.557	1.447	0.39
Div. comp.
1	0	49.757	34.532	49.729	25.427	55.567	30.644	43.149	64.34	65.495
2	1.876ns	0	94.507	119.953	18.454	46.306	21.446	31.527	118.524	92.675
3	1.41ns	5.27***	0	97.764	15.286	33.443	16.85	23.332	87.861	65.213
4	0.667ns	1.636ns	3.25**	0	18.429	46.263	21.411	31.478	119.441	93.016
5	0.053ns	1.24ns	0.983ns	0.403ns	0	27.455	28.109	27.9	21.881	23.207
6	1.471ns	3.494***	0.605ns	2.364*	1.186ns	0	33.053	45.242	59.093	61.088
7	2.15*	3.898***	1.53ns	2.953**	1.791ns	0.836ns	0	32.248	26.446	28.253
8	1.682ns	0.301ns	3.333**	1.369ns	1.269ns	2.972**	3.457**	0	40.229	42.671
9	1.529ns	0.236ns	3.958***	1.167ns	1.051ns	3.052**	3.532**	0.448ns	0	110.059
10	3.164**	6.002***	2.903**	4.676***	2.382*	1.445ns	0.343ns	4.635***	5.195***	0

Table 4. Species diversity test and basic coenological characteristics of weevil assemblages in individual months of the field study (November – March) at ten study plots together.

Σ spp. – total number of species, Σ ind. – total number of individuals, c – Simpson’s index of dominance, e – Pielou’s index of equitability, H’ – Shannon-Wiener’s index of diversity; Div. comp. – diversity comparison: t-test values are under the diagonal and degrees of freedom are above the diagonal. Significance levels: *** P < 0.001, ** P < 0.01, * P < 0.05, ns not significant.

	November	December	January	February	March
Σ spp.	6	6	4	7	14
Σ ind.	42	37	40	42	276
c	0.6304	0.3616	0.5138	0.6995	0.2378
e	0.3805	0.5734	0.5708	0.3	0.3923
H′	0.8254	1.236	0.8256	0.7419	1.703
Div. comp.
November	-	77.352	72.691	83.375	51.787
December	1.753ns	-	73.216	75.114	50.483
January	0.001ns	2.1478*	-	69.232	61.877
February	0.31ns	1.999*	0.359ns	-	50.173
March	4.5784***	2.9383**	6.456***	4.6342***	-

Holecová M et al.: Winter observation of Curculionoidea in Borská nížina lowland

Folia faunistica Slovaca 21 (1) 2016: 91–98

Figure 7. Cumulative proportion of pine and tourist species in individual months of the field study based on the number of species and individuals.

Figure 8. The occurrence of three eudominant pine species in individual months of the non-growing season.

Holecová M et al.: Winter observation of Curculionoidea in Borská nížina lowland