In sti tu te of Ge o gra p hy and Spa tial Ma na ge ment ul. Gro no sta jo wa 7, 30-387 Kra ków
w.ma cie jo wski@geo.uj.edu.pl; a.ka c przak@geo.uj.edu.pl
In flu en ce of se le c ted soil pro per ties on the stru c tu res of be e t le as sem b la ges
in the so u t hern part
of the Kraków-Czê sto cho wa Up land
Wp³yw wy bra nych cech gle by na stru ktu rê zgru po wañ chrz¹szczy w po³ud nio wej czê œci
Wy ¿y ny Krako wsko- Czêsto cho wskiej
Abs trakt: W ni nie j szej pra cy ana li zo wa no zwi¹zki po miê dzy wy bra ny mi ce cha mi gleb, jak typ i pod typ gle by oraz ro dzaj ma te rii w po zio mach ekto hu mu so wych a zgru po wa nia mi chrz¹szczy (ro dzi ny Cur cu lio nidae i Ca ra bi dae). Trzy le t nie ba da nia te re no we i pó Ÿ nie j sza ana li za sta ty sty cz na wy ka za³a ist nie nie wp³ywu tych cech po kry wy gle bo wej na pa ra me try zgru po wañ oby d wu grup chrz¹szczy. Nie jest to wp³yw si l ny, gdy¿ za sto so wa na ana li za re gre sji kro ko wej wste cz nej wy ka za³a istotn¹ sta ty sty cz nie ko re la cjê przy po zio mie isto t no œci oko³o 20%. Na roz mie sz cze nie ryj kow co wa tych wp³yw maj¹ typ i pod typ gle by, któ ry de cy du je o bo ga c twie ga tun ko wym zgru po wañ, a ta k ¿e ro dzaj ekto hu mu su (okre œla wska Ÿ ni ki ró¿ no rod no œci Si m p so na, Ber gera -Par ke ra i McIn to s ha).
U bie ga czo wa tych do mi nuj¹cy wp³yw po sia da typ i pod typ gle by, któ ry wp³ywa na rów no mie r noœæ rozk³adu po szcze gó l nych ga tun ków w zgru po wa niach i de cy du je o wa r to œciach in de ksów ró¿ no rod no œci (wska Ÿ ni ki Si m p so na, Ber gera -Par ke ra i McIn to s ha). Sta no wi to wy nik pra wdo podo b nie ki l ku nak³adaj¹cych siê na sie bie przy czyn. Po pie r wsze, pro fil gle bo wy mo¿e de cy do waæ u wie lu ga tun ków o su kce sie roz ro d czym. W bar dziej mi¹¿szym pro fi lu ist nie je wiê ksza do stê p noœæ po ka r mu wyp³ywaj¹ca z wiê kszej „po je mno œci œro do wi ska”, czy li eg zy sten cji li cz nie j szej gru py or ga ni z mów bêd¹cych po ¿y wie niem dla dra pie ¿ nych bie ga czo wa tych. Ponad to bar dziej roz wi niê ty pro fil gle bo wy zwiê ksza za kres re a kcji za mie sz kuj¹cej j¹ fa u ny na nie ko rzy st ne zmia ny fizy ko che micz ne w jej ob rê bie, np. na nie sprzy jaj¹ce dla larw owa dów prze su sze nie czy prze ma rz niê cie gle by. Drug¹ przy czyn¹ jest re a go wa nie przez ga tun ki chrz¹szczy na w³aœci wo œci fizy ko che micz ne wy ¿szych po zio mów gle bo wych, a trze ci¹ typ u¿y t ko wa nia zie mi.
Key words: soil, Ca ra bid be e t les, we evils, Kraków-Czê sto cho wa Up land
S³owa klu czo we: gle ba, bie ga czo wa te, ryj kow co wa te, Wy ¿y na Krako wsko- Czêsto cho wska
In tro du c tion
The an i mal world is an in te gral part of the nat u ral en vi ron ment sys tem, i.e. ac cord ing to A. Richling (1992) a sys tem of mu tu ally con nected and mu tu - ally in flu enc ing el e ments of na ture. Yet in the struc ture of the en vi ron ment the
Wa r sza wa 2006
role of par tic u lar con stit u ents is not the same. When one takes a given pair of com po nents, one of them usu ally plays a lead ing role, while the other is led.
A. Richling (1992) emphasises, how ever, that the com po nent be ing led also influences the lead ing one. The is sue is im por tant in so far as vir tu ally all ex ist - ing con cep tions of or der ing the en vi ron ment el e ments place the an i mal world in the last sec tion of the hi er ar chi cal lad der, as the one be ing the most sus cep ti - ble to the in flu ence of the oth ers (So³ncew 1965, Armand 1980, Milkow 1981). Hence a con clu sion can be drawn, that an i mals are in flu enced by vir tu - ally all el e ments of the nat u ral en vi ron ment, both through di rect in ter ac tions (fig. 1A) as well as in di rect ones (fig. 1B).
Soil con sti tutes the en vi ron ment of de vel op ment and life of an enor mous quan tity of or gan isms, in clud ing in sects, and the or gan isms them selves also play an im por tant role in the course of soil-form ing pro cesses. They in flu ence the for ma tion of a spe cific soil struc ture, shape its phys i cal prop er ties, en hance the cir cu la tion of el e ments, as well as con trol the course of many bio chem i cal pro cesses, which lead to the de com po si tion of or ganic mat ter and the or i gin of hu mus (Brady, Weil 1999, Singer 2002). In the case of bee tles, most of them make use of only the top soil ho ri zons, where they ac tively move, seek ref uge (dig ging small bur rows), get food or lay eggs.
Over re cent years, a num ber of pa pers con cern ing mu tual con nec tions be - tween the en vi ron ment fea tures and the dis tri bu tion of bee tles have been pub - lished. How ever, most of them deal, first of all, with the in flu ence of var i ous ag ri cul tural op er a tions within soil (e.g. plough ing) on the abun dance and diversity of in sect as sem blages. Such stud ies con cern ing the group of Carabid
270 Wo j ciech Ma cie jo wski, An drzej Ka c przak
Fig. 1. Hie ra r chi cal mo dels of con ne c tions be twe en the na tu ral environment co m po nents (A, B) (af ter: Ma cie jo wski 2004)
Ryc. 1. Hie rar chi cz ne mo de le powi¹zañ po miê dzy ele men ta mi œro do wi ska przy rod ni cze go (A, B) (Zród³o: Ma cie jo wski 2004)´
bee tles were car ried out by, among oth ers: B.R. Stassart et al. (1983), T. Hance, C. Grégoire-Wibo (1987), T. Hance et al. (1990) or M. Ba guette, T. Hance (1997). An in ter est ing ex per i ment was car ried out by a team of re search ers (Fadl et al. 1996), who in ves ti gated changes in the pop u la tion of a Carabid bee tles Pterostichus melanarius un der the in flu ence of ag ri cul tural op er a tions over the veg e ta tion sea son.
There are, in fact, only a few pa pers that de scribe re la tion ships be tween the oc cur rence of in sects and soil prop er ties. One of the first was the work by W.W. Dowdy (1944), who proved the in flu ence of tem per a ture on ver ti cal mi gra tions of soil ver te brates in var i ous types of soil. The re la tion ships be - tween the prop er ties of ectohumus ho ri zons and the oc cur rence of var i ous groups of soil fauna (in clud ing in sects) in south ern Po land have re cently been in ves ti gated by M. Drewnik et al. (1999), W. Maciejowski (2000) and T. Skalski et al. (2003). Those stud ies did not show any close re la tions, though. What was proven was a clear de pend ence of in sects’ abun dance on the kind of ectohumus and its thick ness, as well as a sig nif i cant in flu ence of soil re ac tion in up per soil ho ri zons on the abun dance of some spe cies.
Aim and me t hods
The aim of the pre sented study was an at tempt to in ves ti gate the in flu ence of se lected soil fea tures on the struc tures of bee tle as sem blages. Tak ing into con - sid er ation the ex cep tional di ver sity of that group of an i mals, the study was fo - cused on wee vils (Nemonychidae, Rhynchitidae, Attelabidae, Apionidae, Dryophthoridae, Erirhinidae, Curculionidae) and Carabid bee tles (Carabidae), as those fam i lies oc cur widely and are well stud ied in Po land. The re search was based on the as sump tion that the thicker the ectohumus ho ri zons, the greater the num ber of bee tles, and that there might ex ist de pend en cies be tween bee tle as sem blages char ac ter is tics and var i ous soil types. Those the ses were put for ward on the ba sis of ear lier stud ies by the au thors ei ther deal ing with bee tle as sem blages but con ducted in the moun tains (Skalski et al. 2003), or con cern ing the com po si tion of mesofauna in hu mus ho ri zons of moun tain soils in the Bieszczady Moun tains (Drewnik et al. 1999, Maciejowski 2000) as well as the ex am ple of the May bee tle (Melolontha melolontha), which needs a soil pro file at least 40-50cm thick to overwinter and pu pate (Janssens 1960, Stebnicka 1978).
The field work, con sist ing of col lect ing in di vid u als be long ing to the two bee tle groups at sites with dif fer ent soil cover fea tures, was car ried out at 30 sites in the south ern part of the Kraków-Czêstochowa Up land from 2001 to 2003. Dur ing the field work a set of meth ods of phys i cal ge og ra phy was used, in clud ing the method of physicogeographical map ping (Czeppe, Ger - man 1978). Soil map ping was car ried out us ing pro ce dures com monly used in soil sur vey (Skiba et al. 2000, Singer 2002); soil units (types and sub types)
mapped and used for sta tis ti cal anal y ses were dis tin guished ac cord ing to the current Pol ish clas si fi ca tion (Systematyka gleb Polski 1989) and in ter na - tional names of soil units were given us ing the World Ref er ence Base sys tem (WRB 1998). En to mo log i cal meth ods used to col lect bee tles com prised pit fall traps (Greenslade 1964, Thiele 1977) and sweepnet col lec tion (Karpiñski 1958, Duelli i in. 1999).
The ob tained ma te rial was iden ti fied us ing the key by K. Hurka (1996) and was sub ject to sta tis ti cal anal y sis. For the de scrip tion of de pend en cies be tween soil fea tures (soil type and sub type, kind of ectohumus) and bee tle as sem - blages’ struc tures the method of mul ti ple re verse stepwise re gres sion was used (Jongman et al. 1995), and the sta tis ti cal sig nif i cance of re sults was cal cu lated us ing vari ance anal y sis (ANOVA). Bee tle as sem blages’ struc tures were de - scribed with the fol low ing pa ram e ters: abun dance, spe cies rich ness, spe cies di ver sity de scribed with the Shan non-Wiener (Magurran 1988), Simpson (Lud wig and Reynolds 1988), Berger-Parker (Berger and Parker 1970) and McIntosh (Magurran 1988) in dexes, and also with spe cies dis tri bu tion even - ness (Whittaker 1975).
Area of stu dy
The re search area was lo cated in the south ern part of the Kraków - -Czêstochowa Up land, within a 10–15 km dis tance north-west of Cra cow.
The area com prises the south ern mar gin of the Olkusz Up land be ing a wavy planation sur face with mogotes, cut by deep karst river val leys of the Rac³awka, Szklarka and Bêdkówka, as well as frag ments of the Krze - szowice Graben and the Tenczyn Horst. The bed rock of most of the area is built of lime stones and dolomites (De vo nian-Car bon if er ous, Ju ras sic-Cre ta - ceous), strongly sub ject to karst pro cesses, de formed by fault tec ton ics (Gradziñski 1972), and in the Krzeszowice Graben oc cur im per me able Mio - cene sed i ments. A large frag ment of the area is cov ered with loess. The veg e ta - tion cover is char ac ter ised by the oc cur rence of 25 com mu ni ties, with the larg - est ar eas oc cu pied by ar a ble fields (Secali-Violetalia arvensis), con ti nen tal mixed for est (Querco roboris-Pinetum), sub con ti nen tal de cid u ous for est (Tilio -Carpinetum) and the Carpathian beech for est (Dentario glandulosae - -Fagetum). Places ex posed to sun are oc cu pied by rocky out crops com mu ni - ties (Festucetum pallentis) and warm bush (Peucedano cervariae -Coryletum) (Michalik 1980). The var ied prop er ties of bed rock, which is the sub strate for soil for ma tion, as well as the va ri ety of land forms, strongly con di tion ing microclimatic con di tions and veg e ta tion, are re spon si ble for the fact that the soil cover of the area is a mo saic (Greszta, Bitka 1977, Adamczyk, Kobylecka 1980). Data from Soil-Farm ing Maps 1:5000 (1978), pa pers by Komornicki (1980), Skiba and Trafas (1987) and Zalewa (2001), as well as the au thors’ ob - ser va tions, made it pos si ble to dis tin guish four ba sic soil as so ci a tions
272 Wo j ciech Ma cie jo wski, An drzej Ka c przak
(Leptosols, Luvisols and Cambisols, Fluvisols, anthropo genic soils and soil ma te ri als), each con sist ing of sev eral dif fer ent soil tax o nom i cal units.
Leptosols in the stud ied area are de vel oped from hard, mas sive car bon ate rocks (mainly dolomites and lime stones). The as so ci a tion oc cu pies the west - ern slopes of the Rac³awka, Szklarka and Bêdkówka karst val leys, as well as small patches on the out crops of car bon ate rocks around mogotes and on the steep slopes of the Krzeszowice Graben. The dom i nant units in that as so ci - a tion are Calcari-Lithic Leptosols and Humi-Rendzic Leptosols.
The as so ci a tion of Luvisols and Cambisols de vel oped mainly on loess, and – much less fre quently – on flu vial and glacifluvial sandy ma te ri als. That com - posed soil as so ci a tion oc cu pies the larg est area in the re search area (nearly 70%), form ing ex ten sive patches on the sur faces of the Olkusz Up land and the Tenczyn Horst, as well as the el e vated part of the Krzeszowice Graben.
Soils of that as so ci a tion de vel oped un der mixed for ests, the Carpathian beech for est and the drier va ri et ies of de cid u ous for ests, but pres ently they are used for farm ing. The dom i nant tax o nom i cal units are Haplic Luvisols and Stagnic Luvisols, ac com pa nied by Eutric and Epidystric Cambisols. Deep en ing of soil pro files, es pe cially hu mus ho ri zons, due to slope pro cesses is very of ten ob - served, some times re sult ing in in clu sions of Phaeozems.
Fluvisols cover all al lu vial ma te ri als along river chan nels, most of them de - vel oped in the fine gravel and silt frac tions. The larg est ar eas of Fluvisols are sit u ated in the bot tom of the Krzeszowice Graben, where they are used for ag - ri cul tural pur poses.
The anthropogenic soils and soil ma te ri als, strongly in flu enced by hu man ac tiv ity, have de vel oped in two ways. Firstly, there are Hortic Anthrosols oc - cur ring in built-up ar eas, most of ten un der or chards. Sec ondly, there are places of in ten sive ex ploi ta tion of rock ma te ri als – lime stone, do lo mite and sinter quar ries, re sult ing in the oc cur rence of anthropogeomorphic soil ma te ri als, with out sig nif i cant ex pres sion of pedogenetic pro cesses and mainly of the Spolic char ac ter (WRB 1998).
The soil cover of the study area re flects the di ver sity of bed rock and the va ri ety of land forms (Kobylecka 1981, Skiba and Trafas 1987), show ing cer - tain, spe cific reg u lar i ties (fig. 2). An in ter est ing phe nom ena can be ob served on the slopes of karst val leys. Their west ern slopes are oc cu pied by as so ci a - tions of Calcari-Lithic Leptosols and Rendzic Leptosols, while the east ern slopes are oc cu pied by Luvisols, whose ho ri zons are of ten deep ened due to slope trans port of silt. The rea son for that pat tern is the asym me try of karst val leys re sult ing from the monoclinal ar range ment of bed rock strata and dif - fer ences in loess ac cu mu la tion, caused by the ir reg u lar dis tri bu tion of rain - fall and wind di rec tions.
The de vel op ment of the sur face, ectohumus ho ri zons is in flu enced by the characteristics of veg e ta tion com mu ni ties and the re lief of ter rain. On flat
hill tops and on slopes not steeper than 35o, un der de cid u ous Tilio-Carpinetum, Luzulo-Fagetum and Dentario glandulosae-Fagetum, char ac ter ised by a rather weakly de vel oped storey of ground cover, ectohumus ho ri zons of the Ol type de velop. They are built of rel a tively freshly fallen leaves and their thick ness does not ex ceed a cou ple of centi metres. Morphogenetic pro cesses act ing with a greater force on steeper slopes cause a poorer de vel op ment or even a lack of Ol ho ri zons. In for est com mu ni ties with a well-de vel oped storey of ground cover (Querco roboris-Pinetum, Carici-Fagetum, Alno-Ulmion) ectohumus ho ri zons are usu ally thicker and en riched with undecomposed frag - ments of grasses, so they are of the Olf type. Then, un der grasses and herbs of nat u ral and sec ond ary xerothermic swards thick ectohumus ho ri zons (Of), built of weakly de com posed grass frag ments, de velop.
Stru c tu res of be e t le as sem b la ges in re la tion to soil ty pes and kinds of ec to hu mus ho ri zons
The anal y sis of the in flu ence of soil fea tures on the struc ture of the as sem - blages of wee vils (Curculionidae) and Carabid bee tles (Carabidae) showed the ex is tence of cer tain de pend en cies. Sta tis ti cal tests of cor re la tions (tab. 1 and 2) and the anal y sis of re verse stepwise re gres sion (tab. 3) point out that the as sem blages’ pa ram e ters are in flu enced by cer tain fea tures of the soil cover i.e. soil type and sub type and the kind of sub stance form ing ectohumus ho ri zons.
274 Wo j ciech Ma cie jo wski, An drzej Ka c przak
Fig. 2. A sche me of the di stri bu tion of so ils in re la tion to be drock in the Ol kusz Up land (A) and in the Krze szo wi ce Gra ben (B) (af ter: Ski ba, Tra fas 1987, with chan ges) Ryc. 2. Sche mat roz mie sz cze nia gleb w za le ¿ no œci od pod³o¿a na Wy ¿y nie Ol ku skiej (A) oraz w Ro wie Krze szo wi c kim (B) (•ród³o: Ski ba, Tra fas 1987, nieco zmienione)
Tab. 1. Pearson’s cor re la tion co ef fi cients and sta ti sti cal ly si g ni fi cant re la tion s hips be twe en soil pro per ties and pa ra me ters of the as sem b la ges of we evils (Cur cu lio nidae) and gro und be e t les (Ca ra bi dae) (sta ti sti cal sig ni fi can ce le vel p < 0,05)
Tab. 1. Wspó³czyn ni ki ko re la cji Pe a r so na i isto t ne sta ty sty cz nie za le ¿ no œci po miê dzy ce cha mi po kry wy gle bo wej a pa ra me tra mi zgru po wañ ryj kow co wa tych i bie ga czo wa - tych (sta ti sti cal sig ni fi can ce le vel p < 0,05)
Soil pro per ty Be e t le as sem b la ges pa ra me ters N RP p
Cur cu lio nidae (sno ut be e t les)
Soil type and sub ty pe Spe cies ri ch ness (S) 30 0,397 0,033
Ec to hu mus type Si m p son in dex (D) 30 0,376 0,044
Ber ger - Par ker in dex (d) 30 0,378 0,043
McIn tosh in dex (Q) 30 0,373 0,046
Ca ra bi dae (gro und be e t les)
Soil type and sub ty pe Si m p son in dex (D) 30 0,454 0,013
Ber ger - Par ker in dex (d) 30 0,451 0,014
McIn tosh in dex (Q) 30 0,450 0,014
Di stri bu tion even ness (J) 30 –0,617 0,000
N – ma xim um num ber of sam ples, RP – Pe ars on’s cor rel ati on co eff ici ent, p – sta tis tical si gnif ica nce level After: Ma ciej owski 2004
Tab. 2. Spearman’s ran k -or der cor re la tion co ef fi cients, and sta ti sti cal ly si g ni fi cant re la tion s hips be twe en soil pro per ties and pa ra me ters of the as sem b la ges of we evils (Cur cu lio nidae) and gro und be e t les (Ca ra bi dae) (sta ti sti cal sig ni fi can ce le vel p < 0,05) Tab. 2. Wspó³czyn ni ki ko re la cji porz¹dku rang Spe a r ma na i isto t ne sta ty sty cz nie za le -
¿ no œci po miê dzy ce cha mi po kry wy gle bo wej a pa ra me tra mi zgru po wañ ryj kow co wa - tych i bie ga czo wa tych (po ziom isto t no œci sta ty sty cz nej p < 0,05)
Spearman’s ran k -or der cor re la tion be twe en soil pro per ties
and be e t le as sem b la ges pa ra me ters N RS t(N-2) p
Cur cu lio nidae (sno ut be e t les)
Soil type and sub ty pe & abun dan ce (A) 29 0,372 2,085 0,047
Ca ra bi dae (gro und be e t les)
Soil type and sub ty pe & Shan non - Wie ner in dex (H’) 29 –0,387 –2,183 0,038 Soil type and sub ty pe & Si m p son in dex (D) 29 0,422 2,418 0,023 Soil type and sub ty pe & Ber ger - Par ker in dex (d) 29 0,460 2,695 0,012 Soil type and sub ty pe & McIn tosh in dex (Q) 29 0,422 2,418 0,023 Soil type and sub ty pe & di stri bu tion even ness (J) 29 –0,618 –4,082 0,000 N – ma xim um num ber of sam ples, RS – Spe arm an’s rank– or der cor rel ati on co eff ici ent, t – num ber of sam - ples anal yzed in a se ries, p – sta tis tical si gnif ica nce level. After: Ma ciej owski 2004
Soil type and sub ty pe
There is a sta tis ti cally sig nif i cant re la tion ship be tween the soil type and sub - type and the struc tures of both stud ied bee tle groups’ as sem blages. In wee vil as sem blages it shows ex clu sively pos i tive cor re la tions with abun dance (RS= 0,372) and spe cies rich ness (RP = 0,397). It means that the soil type and sub type, de ter mined by its phys i cal and chem i cal prop er ties, and its pro file mor phol ogy and thick ness, in flu ences the growth of the val ues of the stud ied wee vil as sem blages’ pa ram e ters. In Carabid bee tle as sem blages that soil fea - ture cor re lates with all in dexes of spe cies di ver sity (H’, D, d, Q) and dis tri bu - tion even ness (J). For the Shan non-Wiener di ver sity in dex (RS = –0,387) and dis tri bu tion even ness (RP = – 0,617, RS = –0,618) it shows neg a tive cor re la - tions, whereas for the Simpson (RP = 0,454, RS = 0,422), Berger-Parker (RP = 0,451, RS = 0,460) and McIntosh (RP = 0,373, RS = 0,422) in dexes the cor re la tions are pos i tive (ta ble 1 and 2). So, changes of soil tax o nom i cal units, in the study area usu ally con nected with changes in pro file thick ness, from the shal low est Calcari-Lithic Leptosols to the deep est Cambisols, in flu ence an in - crease in the con tri bu tion and abun dance of the dom i nant spe cies and a si mul - ta neous de crease in the con tri bu tion of rare, oc ca sion ally found spe cies. It is prob a bly a re sult of sev eral over lap ping causes. First, a thicker pro file for many spe cies de ter mines a larger pro cre ation suc cess. It is an ef fect of a higher avail abil ity of food re sult ing from an in creased “en vi ron ment ca pac ity”, i.e. there can ex ist a more nu mer ous group of or gan isms be ing Carabid bee tles’
food. More over, a better-de vel oped soil pro file in creases the re ac tion span of the fauna in hab it ing the soil to un fa vour able phys i cal and chem i cal changes in the soil, e.g. dry ness or oc cur rence of frost, be ing un fa vour able for in sects’
lar vae. That is be cause it en ables an es cape down the soil pro file or even sea - sonal ver ti cal mi gra tions (Demel 1967). Tarnawski (2000) re ports that Staphylinidae lar vae dur ing au tumn chills can even go down to a depth of 50 cm. An other cause is the re ac tion of bee tle spe cies to the phys i cal and chem i cal prop er ties of the up per soil ho ri zons. They may pre fer spe cific phys i - cal prop er ties, e.g. tex ture and com pact ness, which fa cil i tate dig ging a ref uge (nest), get ting food or en hance the de vel op ment of lar vae (Demel 1967), or ap - pro pri ate mois ture con di tions (Hurka 1996). Lindroth (1945, 1953) proved a conscious choice of car bon ate ground by Carabid spe cies from the Harpalus ge nus. It is also known that the Carabidae subfamily of Cicindelinae is con - fined to sandy ground. That fact might ex plain a lack of that subfamily spe cies in the study area, as sand ter rains prac ti cally do not oc cur there at all. Soil chem i cal prop er ties are not in dif fer ent to an i mals, ei ther. For ex am ple, some Carabid spe cies re spond to changes in soil re ac tion (Hurka 1996), and Lindroth (1949) pointed out such pref er ences for two Carabid spe cies from the Bembidion ge nus (Bembidion aeneum i B. min i mum), whose zoo geo graph -
In flu en ce of se le c ted soil pro per ties on the stru c tu res... 277
i cal dis tri bu tion was closely re lated to the chem is try of the ground. A sim i lar choice may be no ticed in the study area, as on the al ka line and shal low Calcari-Lithic Leptosols only four Carabid spe cies (Carabus cancellatus, C. coriaceus, C. nemoralis, Abax parallelepipedus) were re corded, while on the leached and much deeper Luvisols a dom i nance of nu mer ous other spe cies was ob served. It may be ex plained with a low avail abil ity of food and, si mul ta - neously, ex cep tion ally un fa vour able hab i tat con di tions (e.g. high tem per a ture am pli tudes, strong dry ness) in Calcari-Lithic Leptosols, none the less there is a third rea son that most prob a bly has a cru cial in flu ence on the ob served dis tri - bu tion of spe cies and as sem blages. It is the type of land use. That is sup ported by both the proven re la tion ships in di cat ing Carabid bee tles’ pref er ence of for - est com mu ni ties as most suit able for them, as well as the fact that in ar eas used by man (mead ows, pas tures, ar a ble fields) oc cur their as sem blages of a de - formed dom i nance struc ture. Prob a bly that has the larg est in flu ence on the in - crease in the Simpson and Berger-Parker in dexes, and thus a si mul ta neous de - crease in the Shan non-Wiener and dis tri bu tion even ness in dexes. That prob - lem re mains an open is sue, be cause ob serv ing such spe cies as Asaphidion pallipes, Carabus violaceus or Amara curta one can no tice that they ap pear in ar a ble lands and in for ests, but only on acid ground, lack ing cal cium car bon ate (usu ally on Luvisols and Epidystric Cambisols). The ul ti mate an swer may be found dur ing fur ther stud ies com par ing a se ries of ar eas on dif fer ent soil units and of var ied land use, with other el e ments of the en vi ron ment re main ing uni - form.
Kind of ec to hu mus
The kind of ectohumus does not in flu ence sig nif i cantly any pa ram e ters of Carabid as sem blages, while it sig nif i cantly de scribes the pa ram e ters of wee vil as sem blages. Those pa ram e ters are: the spe cies di ver sity in dexes of Simpson (RP = 0,376), Berger-Parker (RP = 0,378) and McIntosh (RP = 0,373). In each of the cases the cor re la tion is pos i tive. It means that with a change of lit ter type from leaf lit ter (Ol), through mixed leaf-grass lit ter (Olf) to pure grass lit ter (Of) an in crease in the num ber of pop u lar, fre quently oc cur ring, spe cies along with a si mul ta neous in crease in the con tri bu tion of the dom i nant spe cies in di - vid u als is ob served in wee vil as sem blages. The ob tained re sults con firm the theses put for ward ear lier claim ing that the di ver sity of wee vils de pends largely on the kind of land use, as it is the kind of land use that plays an im por - tant role in the de vel op ment of ectohumus ho ri zons, or rather – the pres ence or lack of them. In for est ar eas the lit ter ho ri zon is con sti tuted by fallen tree leaves (Ol) or leaves mixed with with ered grass frag ments (Olf). The kind of ectohumus, then, de pends on the de gree of de vel op ment of ground cover and its char ac ter is tics. The better the ground cover is de vel oped, the higher the contribution of undecomposed grass frag ments in lit ter will be. Dur ing
the vegetation pe riod such a ho ri zon has a very small thick ness and may even be not pres ent at all, due to a rel a tively fast de com po si tion of or ganic mat ter. It achieves its great est thick ness and best ex pres sion only in the au tumn time, when veg e ta tion dies and leaves fall down. On mowed mead ows and ar a ble fields lit ter ho ri zons do not oc cur at all. In the study area the best-de vel oped ectohumus ho ri zons of grass lit ter char ac ter (Of) oc cur un der nat u ral com mu - ni ties of xerothermic swards or for mer ar a ble lands be com ing over grown, sub - ject to renaturalisation. Thus wee vil spe cies find ing fa vour able con di tions in such hab i tats form abun dant clus ters. A change of plant spe cies, in turn, trig - gered by suc ces sion will elim i nate the con tri bu tion of sin gle bee tle spe cies, which will re treat slightly with a de crease in the di ver sity of com mu ni ties.
There fore the in flu ence of the kind of ectohumus is not sig nif i cant in that area, es pe cially as the de vel op ment of ectohumus de pends on a veg e ta tion com mu - nity un der which it forms while its pres ence is con nected with the type of land use.
Con c lu sions
An anal y sis of the in flu ence of se lected soil fea tures on the as sem blage struc tures of se lected bee tle groups showed the ex is tence of cer tain de pend en - cies. The in flu ence is not very strong as the anal y sis of re verse stepwise re gres - sion showed sta tis ti cally sig nif i cant cor re la tion with a small per cent age of ex - plained cases of the de pend ency usu ally reach ing ca. 20%. The val ues of R2 os cil lated form 0,14 to 0,36 (ta ble 3). The dis tri bu tion of wee vils is in flu enced by the type and sub type of soil, con trol ling spe cies rich ness, and the kind of ectohumus, con trol ling the in dexes of di ver sity (Simpson, Berger-Parker, McIntosh). As for Carabid bee tles, there is a sig nif i cant in flu ence of the type and sub type of soil, con trol ling the Simpson, Berger-Parker and McIntosh di - ver sity in dexes as well as de ter min ing the dis tri bu tion even ness of par tic u lar spe cies within as sem blages.
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