Biologia 64/2: 343—349, 2009 Section Zoology DOI: 10.2478/s11756-009-0045-3

Farming system and habitat structure effects on rove (Coleoptera: Staphylinidae) assembly in Central European apple and pear orchards

Adalbert Balog1,2,ViktorMarkó2 & Attila Imre1

1Sapientia University, Faculty of Technical Science, Department of Horticulture, 1/C Sighisoarei st. Tg. Mures, RO-540485, Romania; e-mail: [email protected] 2Corvinus University Budapest, Faculty of Horticultural Science, Department of Entomology, 29–43 Villányi st., A/II., H-1118 Budapest, Hungary

Abstract: In field experiments over a period of five years the effects of farming systems and habitat structure were in- vestigated on staphylinid assembly in Central European apple and pear orchards. The investigated farms were placed in three different geographical regions with different environmental conditions (agricultural lowland environment, regularly flooded area and woodland area of medium height mountains). During the survey, a total number of 6,706 individuals belonging to 247 species were collected with pitfall traps. The most common species were: angustula, caesum, canaliculata, Oxypoda abdominale, Philonthus nitidulus, Dexiogya corticina, Xantholinus linearis, X. lon- giventris, Aleochara bipustulata, Mocyta orbata, Oligota pumilio, Platydracus stercorarius, Olophrum assimile, Tachyporus hypnorum, T. nitidulus and Ocypus olens. The most characteristic species in conventionally treated orchards with sandy soil were: Philonthuss nitidulus, Tachyporus hypnorum, and Mocyta orbata, while species to be found in the same regions, but frequent in abandoned orchards as well were: , Oxypoda abdominale, Xantholinus linearis and Drusilla canaliculata.ThespeciesDinaraea angustula, Oligota pumilio, Dexiogya corticina, Xantholinus longiventris, Tachyporus nitidulus and Ocypus olens have a different level of preferences towards the conventionally treated orchards in clay soil. The species composition of the staphylinid fauna in apple and pear orchards could not be considered uniform. The environmental conditions and the soil together have a significant influence upon the richness of species, and the cumulative effects of these factors can modify even the dominance structures of the communities. Key words: environment; seasonal dynamics; soil; treatment

Introduction horst, 1802) and Philonthus cognatus (Stephens, 1832). The response to prey spatial heterogeneity, the ag- Staphylinidae is one of the largest families of Coleoptera gregation and aphid and mildew preferences were stud- with species that are mostly predacious. More than ied by many authors (Bryen & Wratten 1985; Sunder- 45,000 species are known worldwide and probably over land et al. 1987; Dennis et al. 1991; Good & Giller 1991; 75% of tropical species are still undescribed (Howard et Birken & Cloyd 2007). Several species aggregated in al. 1998). Although 1,500–1,700 species were recorded patches of aphids and presented a positive numerical re- from Central Europe, there are some ecosystem types, sponse to high aphid densities (Bryen & Wratten 1985). like pine forests and sub-alpine regions (Zerche 1994), Under laboratory conditions the average aphid con- in which the staphylinid fauna is still little known Al- sumption was 1 mg/day for a lot of the species, which is though many authors studied the staphylinid fauna more than 34% of their body weight. In gut-dissection (Dennis et al. 1990; Majzlan & Holecová 1993; War- work carried out by Sunderland et al. (1987) three dle et al. 1993; Heyer 1994; Krooss & Schaefer 1998; categories of food, other than aphids, were identified Andersen 1991, 2000; Perner & Malt 2002) there is lit- in the diet of the Tachyporus spp.: non-aphid arthro- tle information concerning the effects of farming sys- pods, rusts and non rust fungi. Dennis et al. (1991) tems in Central Europe. Andersen (1991) presented a and Birken & Cloyd (2007) showed that these species list of staphylinid beetles in Norway occurring in spring presented a positive numerical response to high density barley, cabbage, carrot, potato, strawberries and grass- of rusts and non rust fungi, while aphid predation de- land fields. The author caught 103,000 individuals be- creased significantly in the same time. Other species, longing to 226 species. The most frequently found were like Philonthus spp., fed on a wide range of arthro- Aloconota gregaria (Erichson, 1839), rugosus pod prey, aphid predation averaged 200 aphid individ- (F., 1775), Atheta fungi (Gravenhorst, 1806), Amischa uals/day and there is no record of mycophagy (Good analis (Gravenhorst, 1802), Tachinus signatus (Graven- & Giller 1991). Staphylinidae were used as bioindica-

c 2009 Institute of Zoology, Slovak Academy of Sciences 344 A. Balog et al.

Table 1. The characteristics of the investigated orchards.

Farm 1 2 345 6 7 8 9

Plantation apple apple apple apple apple apple, apple apple, pear apple, pear apple, pear Year of planting 1988 1977 1992 1960 1963 1992, 1995 1950 1990 1977, 1980 Tree distance 4 × 1.6 m 1.2 × 3.2 m 4 × 2m 7× 7m 5× 4m 5× 2m 5× 4m 7× 4m 6× 4m Soil clay clay clay sand sand sand, sand clay sand sand, sand Weed management M M M M NV M, NV M NV M Treatment CON CON CON CON ABA CON, ABA CON CON CON, CON Environment WAM WAM ALE WAM ALE ALE, ALE ALE ALE RFA, RFA Landscape fields fields fields fields fields fields fields fields fields

Explanation: WAM – woodland area of medium height mountains; ALE – agricultural lowland environment; RFA – regularly flooded area; CON – conventionally treated orchard; ABA – abandoned orchard; M – mowed; NV – natural vegetation. tors of environmental changes in natural and modified to species level under a stereomicroscope. Species identifi- ecosystems because of their relative ease of capture by cation was based on the works of Freude et al. (1964), Tóth pitfall trapping, responsiveness to environmental condi- (1982, 1984) and Zerche (1994). tions, mobility and widespread distributions (Wardle et We carried out analyses of variance (two-way ANOVA) al. 1993; Heyer 1994; Krooss & Schaefer 1998; Andersen using NUCOSA statistical software to determine whether there were any differences in Staphylinidae species rich- 1991, 2000; Boháč 1999; Perner & Malt 2002; Shah et al. ness and abundance between the environmental conditions 2003). In Central Europe, reports on inhab- and soil (Tóthmérész 1996). Two-way ANOVA may be used iting orchard ecosystems concentrated on other groups to examine the effects of two categorical variables (fac- than Staphylinidae (Mészáros et al. 1984; Markó et al. tors), both individually and together, on an experimental 1995; Bogya et al. 1999; Kutasi et al. 2001; Balog et al. response. In our case these two categorical variables were 2003, 2007a, b, 2008). In this work, we are presenting two environmental conditions (WAM and ALE) and soil some aspects of species richness and activity-density of structure (sand and clay) of the investigated orchards. The Staphylinidae in apple and pear orchards under differ- following standardizations were used for the test: for envi- ent management system, environmental conditions and ronmental conditions WAM, apple orchards from conven- tional farms 1 and 2 were considered as replicates and com- soil structures. pared with apple orchards from conventional farms 3 and 7 in ALE. Orchards from farm 3 and 7 (apple, clay), and 6 (CON), 8 (apple, sand) were compared for soil studies (Ta- Material and methods ble 1). We used ANOVA to determine whether a significant difference between the interactions exists. In this respect Studies were performed from 1998 to 2002 on nine farms it is a preliminary test that informs us whether we should and 13 plantations. The latitude and longitude ranges of continue the investigation of the data. If the null hypothesis ◦ the investigated farms are the followings: Farm 1: 46 56 N, (no difference among interactions) is accepted, there is an ◦ ◦ ◦ ◦ ◦ 16 58 E; Farm 2: 48 N, 18 52 E; Farm 3: 46 3 N, 17 59 implication that no relation exists between the factor levels ◦ ◦ ◦ ◦ E; Farm 4: 47 25 N, 17 48 E; Farm 5: 46 54 N, 19 42 and the response. If a significant F -value is found for one ◦ ◦ ◦ ◦ E; Farm 6: 47 49 N, 21 30 E; Farm 7: 48 12 N, 21 40 independent variable, then this is referred to as a significant ◦ ◦ ◦ ◦ E; Farm 8: 47 36 N, 19 36 E; Farm 9: 47 16 N, 18 59 E. main effect. However, when two or more independent vari- Five farms consisted of one apple orchard each, one of two ables are considered simultaneously, there is also an interac- apple orchards, while three consisted of one apple and one tion between the independent variables which may or may pear orchard (Table 1). not be significant (Pielou 1984; Tóthmérész 1993, 1995). The investigated farms were placed in three dif- Another method is the Cluster Analysis which was used ferent geographical regions with different environmental to study the forming similarity of staphylinid communities conditions. These were agricultural lowland environment under the different geographical regions. Horn index was (ALE), regularly flooded area (RFA – overflooded basin of used to compare the dominance structure of the commu- the Danube River near Budapest) and woodland area of nities, considering the relative abundance of species. Horn medium height mountains (WAM). Five farms were located index can be derived from: on sand, whereas four on clay (Table 1). Eleven planta-  tions were treated with mainly organophosphate insecticides Ro = (Xij +Xik)log(Xij +Xik) − (methidation, fosalon, fosfamidon) during the study period.    These were applied on average 10 times during the growing − Xij log Xij − Xiklog Xik season. Two apple plantations were untreated and neither pesticides nor fertilizers had been used for five years before [(Nj +N k)log(N j +N k) − Nj log N j − Nklog N k], we started our investigation. Ten covered pitfall traps (300 3 cm in size, 8 cm in diameter, half-filled with 30% ethylene where: Ro = Horn similarity index in samples j and k; Xij , glycol) were placed in transect from a field margin towards Xik = number of individuals of i species in samples j and k; the field centre at 10 m intervals within each plantation. Nj= Xij = total number of individuals in sample j ; Nk Five traps were placed in the middle of the plantation and = Xik = total number of individuals in sample k. five in the inner edges. Pitfall traps are considered a useful We calculated Horn and Horn log10 indices (Krebs method to study assemblages of epigeic arthropods (Luff & 1989). Horn log10 was used to reduce the importance of Eyre 1988). Samples were collected fortnightly from April the dominant species and emphasizing the importance of to October. All Staphylinidae were sorted and identified up subdominant species. Rove beetles in Central European orchards 345

Table 2. Numbers of rove species and individuals in indi- Table 3. The mean species number (log10) and coef- vidual study sites. ficient of variations in orchards using two-way ANOVA.

Farms Sites Species Individuals Two-way ANOVA by species df FP

1 Vámosmikola 57 537 Environment (ALE, WAM) 1 19.70 0.057 2 Pókaszepetk 58 971 Soil (clay, sand) 1 22.63 0.102 3 Szentl˝orinc 73 547 Interaction 1 33.09 < 0.01 4 Bakonygyirót 97 992 5 Kecskemét 59 961 6 Újfehértó 69 500 7Gy¨orgytarló 58 438 8 Tura 86 1040 Table 4. The mean rove beetle individual number (log10)and 9 Szigetcsép 92 720 coefficient of variations in orchards using two-way ANOVA. Total 247 6706 Two-way ANOVA by individuals df FP

Environment (ALE, WAM) 1 8.72 0.104 Soil (clay, sand) 1 5.80 0.253 Interaction 1 72.51 < 0.001 The habitat preferences of the species were compared with DECORANA, which is able to detect relationships be- tween species and external (environmental) variables. These relationships can be “unimodal” rising and falling again as the optimum environmental conditions for a species are ap- proached and passed. DECORANA avoids the arch or horse- shoe problem where the second axis is frequently a quadratic distortion of the first axis (Oksanen & Minchin 1997). For this method all species from all investigated orchards were considered and the catches of the species with significant preferences were graphed. The activity-density of the dom- inant species was calculated for different environment, soil and treatment methods and we considered the number of the individuals caught in each pitfall trap. Dominant species with relative abundance higher than 10% in at least one of the investigated orchards were taken into account.

Results

In total, we collected 6,706 rove beetle individuals Fig. 1. Forming dominance (Horn log10) of rove beetle communi- belonging to 247 species and 11 subfamilies. More- ties under different environmental conditions. 1–10: species com- over, we identified the subfamily and position in each trap in ALE; 11–20: species composition in each found 80 species belonging to it. After cumulative as- trap in RFA; 21–30: species composition in each trap in WAM. say studies of the communities, we identified the most widespread species. Altogether, 16 species presented relative abundances from 2% to 10%, comprising 56% The highest activity-density occurred in orchards of all rove beetles recorded. They were: Dinaraea angus- with sandy soil and in WAM orchard. Furthermore, we tula (Gyllenhal, 1810), Omalium caesum Gravenhorst, observed that the environment and soil together has a 1806, Drusilla canaliculata (F., 1787), Oxypoda abdomi- significant effect upon the activity-density of rove bee- nale Mannerheim, 1831, Philonthus nitidulus (F., 1781), tles in orchards (Table 4). Dexiogya corticina (Erichson, 1837), Xantholinus lin- The forming of dominance structure of rove bee- earis (Olivier, 1795), X. longiventris Heer, 1839, Aleo- tle communities did not differ significantly in ALE and chara bipustulata (L., 1761), Mocyta orbata (Erichson, WAM under the three environmental conditions stud- 1837), Oligota pumilio Kiessenwetter, 1858, Platydracus ied, and some of the species found in RFA were com- stercorarius (Olivier, 1795), Olophrum assimile (Pay- mon with ALE and WAM (Fig. 1). On the other hand, cul, 1800), Tachyporus hypnorum (F., 1775), T. nitidu- the dominance structure differed significantly, which lus (F., 1781) and Ocypus olens (O.F. M¨uller, 1764). leads us to the belief that the most common rove beetle The cumulative species and individuals’ number for species in the assemblage have preferences for different each study site are presented in Table 2. Two-way environmental conditions (F = 2.05, P < 0.01) (Fig. 2). ANOVA showed that the effect of environment (WAM These preferences can be observed by studying the and ALE) on species richness was statistically insignif- similarity of the abundance for particular factors like icant (Table 3). The effect of soil (sandy and clay) on treatment and soil structure (DECORANA). In Fig. 3, species richness was insignificant, too. However, the in- along axis 1, species on the right were frequently found teraction of environment and soil has a significant effect in conventionally treated orchards, while species lo- upon species richness (Table 3). cated on the left were captured both in conventional 346 A. Balog et al.

ferent level of preferences towards the conventionally treated orchards in clay soil, while Olophrum assimile and Platydracus stercorarius were also frequent in con- ventional and abandoned orchards (Fig. 3). Seasonal dynamics of rove beetles were calculated only for dominant species with relative abundance higher than 10% in at least one of the investigated orchards and all individuals from the whole investi- gation period were considered. These species were Di- naraea angustula, Omalium caesum, Drusilla canalicu- lata, Philonthus nitidulus, Xantholinus linearis, Oxy- poda abdominale, Mocyta orbata, Aleochara bipus- tulata, Dexiogia corticina, and Platydracus sterco- rarius. Fig. 2. Dominance structure (Horn) of rove beetle communities Xantholinus linearis, Oxypoda abdominale and under different environmental conditions. For explanations see Mocyta orbata occurred in orchards situated on sand Fig. 1. in ALE, where their abundance was the highest in May, June, October and November (Fig. 4). Philonthus nitidulus and Aleochara bipustulata were found in or- and abandoned orchards. Along axis 2, species captured chards on sand in WAM. The greatest abundance was in orchards with sandy soil are located underneath, observed in the middle of the growing season and the while species frequent in clay orchards are located lowest one in spring and autumn (Fig. 5). above (Fig. 3). Consequently, the most characteristic Frequently occurring species in apple orchards on staphylinid species in conventionally treated orchards clay in ALE were Aleochara bipustulata and Dexiogia with sandy soil were Philonthus nitidulus, Tachyporus corticina with high occurrence in the first half of the hypnorum, and Mocyta orbata, while species found in growing season (Fig. 6). and Platy- the same regions, but also frequent in abandoned or- dracus stercorarius were present in the same orchards chards were Omalium caesum, Oxypoda abdominale, but in WAM, with high activity-density in the middle Xantholinus linearis and Drusilla canaliculata. Aleo- of the growing season (Fig. 7). chara bipustulata was also present in relatively high Widely occurring species in abandoned orchards numbers in clay orchards. Dinaraea angustula, Oligo- were Omalium caesum, Oxypoda abdominale and Dru- ta pumilio, Dexiogya corticina, Xantholinus longiven- silla canaliculata. The first two were numerous in May, tris, Tachyporus nitidulus and Ocypus olens had a dif- whereas Drusilla canaliculata in June and July (Fig. 8).

Fig. 3. Habitat preferences of the dominant rove beetle species in conventionally treated and abandoned apple orchards with different soil (DECORANA). Rove beetles in Central European orchards 347

90 350 Dinaraea angustula 80 Xantholinus linearis 300 Oxypoda abdominale Platydracus stercorarius 70 Mocyta orb ata 250 60 200 50

40 150

30 100 No. of individuals / trap / individuals of No. No. of individuals / trap / individuals of No. 20 50 10

0 0 MAY JUN JUL AUG SEP OCT NOV MAY JUN JUL AUG SEP OCT NOV ALE, sand WAM, clay

Fig. 4. Seasonal dynamics of dominant rove beetle species in or- Fig. 7. Seasonal dynamics of dominant rove beetle species in or- chards situated on sand and surrounded by ALE (farm 6 and 8 chards situated on clay and surrounded by WAM (farm 1 and CON). 2).

100 200 Drusilla canaliculata 90 Philonthus nitidulus 180 Omalium caesum Aleochara bipustulata 160 80 Oxypoda abdominale 70 140

60 120

50 100

40 80

No. of individuals / trap / individuals of No. 30 60 No. of individuals / trap / individuals of No. 20 40

10 20

0 0 APR MAY JUN JUL AUG SEP OCT NOV APR MAY JUN JUL AUG SEP OCT NOV WAM, sand ALE, ABA, s and

Fig. 5. Seasonal dynamics of dominant rove beetle species in or- Fig. 8. Seasonal dynamics of dominant rove beetle species in chards situated on sand and surrounded by WAM (farm 4). abandoned orchards situated on sand and surrounded by ALE (farm 5 and 6 ABA).

25 Aleochara bipustulata Dexiogyia corticina chards throughout Europe indicates that there is a 20 large variability between the composition of Staphyli- nidae communities. Out of the 17 species abundant 15 in our study, only Omalium caesum, Drusilla canali-

10 culata, Olophrum assimile and Tachyporus hypnorum were reported as common in other European agri- No. of individuals / trap / individuals of No.

5 cultural fields. The other species were mentioned as frequent in apple and pear orchards for the first 0 time. APRMAYJUNJULAUGSEPOCTNOV The species composition of the rove beetle fauna ALE, clay in apple and pear orchards cannot be considered uni- Fig. 6. Seasonal dynamics of dominant rove beetle species in or- form; the environmental conditions and soil affect the chards situated on clay and surrounded by ALE (farm 3 and 7). rove beetle fauna in different ways. The composition of the species and their activity-density are also in- fluenced by the above-mentioned factors. The expla- Discussion nation lies in different soil texture and different tol- erance of the species to microclimate and humidity. Results on rove beetle fauna in Central European Our findings are consistent with other studies, which apple and pear orchards were recorded for the first also showed higher species number in conventional time. However, few of these species were mentioned agricultural systems. The reason for a relatively high as frequent in agricultural fields by other authors number of species in conventionally treated orchards throughout Western Europe (Dennis et al. 1990; Maj- may be the higher crop density in these fields pro- zlan & Holecová 1993; Basedow & Kollát 1997). More viding a more humid microclimate, or the presence than 15% of Central European rove beetle fauna was of more suitable food resources (Basedow & Kollát present in the investigated orchards. Comparison of 1997; Shah et al. 2003). Generalist predators can in- our data with results from other apple and pear or- terfere by intraguild predation, cannibalism, predator- 348 A. Balog et al. predator competition and avoidance behaviour (Shah Birken E.M. & Cloyd R.A. 2007. Food preference of the rove et al. 2003). Moreover, almost all species recorded in beetle, Atheta coriaria Kraatz (Coleoptera: Staphylinidae) 14: this study are highly eurytopic and can not precisely under laboratory conditions. 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The activity-density of the rove beetle fauna un- Entomol. 5: 251–259. der different environmental conditions can not be con- Dennis P., Wratten S.D. & Sotherton N.W. 1990. Feeding behav- sidered uniform in time, and probably climatic condi- ior of the staphylinid beetle Tachyporus hypnorum in relation tions, especially the lowest temperature in WAM (on to its potential for reducing aphid numbers in wheat. Ann. ◦ Appl. Biol. 117: 267–276. average 3–5 C lower than in ALE) affect the rove beetle Dennis P., Wratten S.W. & Sotherton N.W. 1991. Mycophagy as fauna in different ways. Moreover, although the domi- a factor limiting predation of aphids (Hemiptera: Aphididae) nant rove beetles in the investigated orchards belonged by staphylinid beetles (Coleoptera: Staphylinidae) in cereals. to disturbance-resistant species, the rove beetles fauna Bull. Entomol. Res. 81: 25–31. Freude H., Harde, W.K. & Lohse G.A. 1964. Die K¨afer Mitteleu- in these habitats was not uniform. 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