Respiration of wood ant nest material affected by material and forest stand characteristics

Biologia 68/6: 1193—1197, 2013 Sec ion Zoology DOI: 10.2478/s11756-013-0261-8 Respi a ion o wood an nes ma e ial affec ed by ma e ial and o es s and cha ac e is ics Ve onika Jilko a1,2,TimoDomisch3,4,ZuzanaHo icka1,2 &JanF ouz1,2 1Ins i u e o Soil Biology, AS CR, Na Sadkach 7,CZ-37005 Ceske Budejo ice, Czech Republic; e-mail: jilko a. e [email protected] 2Ins i u e o En i onmen al S udies, Cha les Uni e si y in P ague, Bena ska 2,CZ-12800 P aha, Czech Republic 3Facul y o Fo es y, Uni e si y o Joensuu, P.O. Box 111,FI-80101 Joensuu, Finland 4Finnish Fo es Resea ch Ins i u e, Joensuu Resea ch Uni , P.O. Box 68,FI-80101 Joensuu, Finland Abs ac : We s udied diffe ences in espi a ion o ma e ials om diffe en pa s o wood an nes ( op, bo om, and im) and om he nes su oundings (humus laye and mine al soil). Samples we e aken om 8 wood an (Fo mica aquilonia) nes s in each o he wo ypes o o es (bi ch and pine) in eas e n Finland. The diffe ences we e ela ed o ma e ial and o es s and cha ac e is ics (i.e., mois u e, pH, ca bon con en , and C:N a io). As a esul , he highes espi a ion pe g DW was measu ed a he op o an nes s in he bi ch o es . Howe e , espi a ion did no significan ly diffe be ween he pa s o an nes s in he pine o es . Respi a ion o he humus laye s in bo h o es s ands was on a e age highe , whe eas espi a ion o he mine al soils in bo h o es s ands was lowe in compa ison wi h espi a ion o he nes ma e ials. The espi a ion pe g C did no show any significan diffe ences be ween diffe en pa s o nes s and su ounding soil. The mos impo an ac o s influencing espi a ion o he ma e ials appea ed o be mois u e, ca bon con en , and pH. In conclusion, espi a ion o wood an nes ma e ial is affec ed by he specific ma e ial and o es s and cha ac e is ics. Key wo ds: Fo mica aquilonia; bi ch o es ; pine o es ; CO2; mois u e; ca bon con en ; pH In oduc ion Wood an s (Fo mica s. s ., Hymenop e a: Fo micidae) include species o an s ha a e dominan in empe a e and bo eal o es ecosys ems (Dlusskij 1967). They a e widesp ead om he empe a e zone o Eu ope and Asia o he No h-Palaea c ic a ea, e en no h o he A c ic Ci cle (Czechowski e al. 2002; Pun ila & Kilpel¨ainen 2009). Wood an species a e mainly associa ed wi h coni e ous and mixed o es s, al hough hey occu also in deciduous o es s (Czechowski e al. 2002). Due o hei o aging and building ac i i ies hey affec especially nu ien cycling (F ouz e al. 2005; F ouz & Jílko á 2008). Recen ly, hei nes s we e ound o be “ho spo s” o CO2p oduc ion (Ohashi e al. 2005; Risch e al. 2005; Domisch e al. 2006). This CO2o igina es om se e al sou ces: an espi a ion, decompose communi y espi a ion, and espi a ion o oo s pene a ing in o he belowg ound pa s o he nes (Lenoi e al. 2001; Risch e al. 2005). In ou s udy, we ocused on he decomposi ion ac i i y o o ganisms (i.e., CO2efflux), which is an impo an pa o he ca bon cycle (S eubing 1970). The knowledge o CO2sou ces and hei dependence on en i onmen al cha ac e is ics is impo an o he unde s anding o ecosys em ca bon balance in o es ecosys ems. Respi a ion o he decompose communi y can be influenced by many ma e ial p ope ies, such as mois- u e, acidi y, ca bon con en , C:N a io, o he ela- ion be ween cellulose and lignin (Edwa ds e al. 1970; S eubing 1970; Paul & Cla k 1996). Al hough he an nes ma e ial is epo ed o be d ie in compa ison o he nes su oundings, decomposi ion p ocesses a e a he a ou ed he e (F ouz e al. 1997; Laakso & Se ¨al¨a 1997; Lafleu e al. 2002). Since he wa e con- en diffe s be ween he pa s o he nes , he mois u e is highes in he uppe pa o he nes whe e he e was also measu ed he highes empe a u e (Coenen-S aß e al. 1980). Thus, mic obial ac i i y and decomposi ion p ocesses a e encou aged especially in mois su ace lay- e s o an nes s (Laakso & Se ¨al¨a 1998; Daube e al. 2001; F ouz e al. 2003). Wood an nes s and hei close icini y a e usually epo ed o be less acidic in compa - ison wi h he wide su oundings (Jílko á e al. 2011, 2012), which could also a ou he ac i i y o bac e ia and ac inomyce es, no only ungi ypical o acidic soils (Paul & Cla k 1996). Because o o aging and building ac i i ies o an s, amoun o o ganic ma e (i.e. ca bon con en ) is enhanced in an nes s (F ouz e al. 1997), bu he quali y o o ganic ma e (C:N a io and he ela ion be ween cellulose and lignin) depends on he sou ce o o ganic ma e and hus on he o es s and (Edwa ds e al. 1970). In his s udy we examined ma e ial espi a ion in diffe en pa s o an nes s in wo o es s ands and in he humus laye and mine al soil om he su oundings c 2013 Ins i u e o Zoology, Slo ak Academy o Sciences B ough o you by | Me la Finnish Fo es Resea ch Ins i u e Au hen ica ed Download Da e | 12/21/16 11:43 AM 1194 V. Jilko a e al. Table 1. P ope ies o ma e ials om he espec i e sampling loca ions a he bi ch and he pine o es . Sampling Mois u e (%) pH Ca bon con en (%) C:N a io loca ion F(4,56) =5.04P<0.05 F(4,56) =6.16P<0.001 F(4,56) =6.79P<0.001 F(4,56) = 15.76 P<0.001 Bi ch Pine Bi ch Pine Bi ch Pine Bi ch Pine TOP 41.15 ±1.85a 19.55 ±1.40w 4.92 ±0.14a 4.50 ±0.05w 43.08 ±0.71a 52.30 ±1.11w 20.97 ±0.13a 67.70 ±6.08w BOTTOM 22.50 ±1.43b 16.80 ±1.33w 5.23 ±0.08a 5.19 ±0.05x 32.58 ±0.88b 25.40 ±5.44x 19.22 ±0.78a 30.27 ±2.81x RIM 21.80 ±5.00b 11.46 ±1.22w 5.24 ±0.07a 4.86 ±0.06y 12.07 ±4.14c 9.14 ±0.26y 19.05 ±0.42a 30.12 ±0.15x HUMUS 59.01 ±2.18c 51.19 ±4.97x 3.97 ±0.07b 3.73 ±0.07z 45.44 ±1.01a 29.74 ±2.55x 22.50 ±0.80a 31.23 ±0.35x SOIL 16.15 ±1.82b 20.00 ±0.75w 4.81 ±0.12a 5.18 ±0.03x 0.63 ±0.03d 2.23 ±0.16y 30.78 ±3.08b 36.07 ±1.46x Explana ions: TOP e e s o he op o he an nes s, BOTTOM o he base o he nes s, RIM o he pe iphe y o he nes s, and HUMUS and SOIL o he humus laye and mine al soil, espec i ely, as con ols. Mean ±SEM and esul s o h ee-way ANOVAs a e shown. S a is ically homogeneous g oups o alues o he same ma e ial p ope y a e ma ked by he same le e (one-way ANOVA, Tukey HSD pos hoc es ). as con ols. We ela ed he diffe ences o mic osi e and o es s and cha ac e is ics. Ou hypo heses we e ha (1) ma e ial espi a ion is he highes in he op o he nes s, (2) he diffe ences in ma e ial espi a ion a e ex- plained by ma e ial p ope ies (e.g., mois u e, pH, ca - bon con en ), and (3) he o es s and cha ac e is ics a e esponsible o he diffe ences in ma e ial espi a- ion, especially due o ca bon con en o he li e . Ma e ial and me hods S udy si es The s udy was conduc ed in Augus 2009 in wo o - es s ands, bi ch (Be ula pendula Ro h) and pine (Pinus syl es is L.), in eas e n Finland nea Joensuu (62◦3928 N, 29◦3938 E, bi ch s and; 62◦4120 N, 29◦4433 E, pine s and). The bi ch o es was a 47-yea old s and wi h a ee densi y o 2,376 ees pe ha and an an nes densi y o 9.9 nes s pe ha. An nes s in he bi ch o es we e cons uc ed mainly om bi ch wigs, ba k, and lea es. Howe e , a small po ion o he pine needle ma e ial was also included since he bi ch o es s and was si ua ed ca. 100 m om he pine o es s and. The pine o es was a 37-yea old s and wi h a ee densi y o 2,235 ees pe ha and an an nes den- si y o 6.3 nes s pe ha. An nes s in he pine o es we e cons uc ed mainly om pine wigs and needles, wi h a con- side able amoun o coni e esin. Sampling design Samples o ma e ials we e aken wi h a sho el om 8 mounds o Fo mica aquilonia Ya ow, 1955 an s in bo h o es s ands. Sampling loca ions we e he op (20 cm be- low he su ace), he bo om ( he cen e o he mounds on he le el o he o es floo ) and he soil im o he mounds ( he pe iphe y o he mounds), and he humus laye and mine al soil app oxima ely 10 m om an adjacen mound as con ols. In o al, 80 ma e ial samples we e aken. F esh samples we e immedia ely anspo ed o he labo a o y o analyses. Ma e ial analyses Respi a ion o he ma e ials, mois u e, pH, and con en o ca bon and ni ogen we e measu ed. Respi a ion was es- ablishedusing5go he eshma e ialsa e a wo-day incuba ion a 20◦ C by i a ion o NaOH acco ding o Page (1982). Mois u e was de e mined as a diffe ence be ween esh mass and d y mass a e d ying a 105◦ C o 12 h. pH was measu ed in a 1:10 ma e ial:wa e suspension wi h a glass elec ode. The o ganic ma e con en was assessed based on loss-on-igni ion a e 5 h in 600◦ C. The con en o ca bon and ni ogen was analyzed in samples o d y c ushed soil using elemen al analyze EA 1108 (Ca lo E ba Ins u- men s). S a is ical analyses Th ee-way ANOVA wi h o es s and, sampling loca ion and nes as ca ego ical a iables, wi h nes as a andom effec nes ed in o es s and, we e used o analyze diffe ences in ma e ial p ope ies. I he in e ac ion be ween o es s and and sampling loca ion was significan , one-way ANOVA and pos hoc es (Tukey HSD) was pe o med o assess di - e ences be ween he sampling loca ions. All compu a ions we e made using he p og am S a is ica 10. Resul s Ma e ial espi a ion pe g DW soil diffe ed signifi- can ly be ween he sampling loca ions and o es s ands (Fig. 1). Diffe ences we e mo e p onounced in he bi ch o es han in he pine o es . The highes espi a ion was measu ed a he op o an nes s in he bi ch o - es , whe e i was se en imes highe han espi a ion a he op o an nes s in he pine o es . The bo om and im in bo h o es s ands, as well as he op o an nes s in he pine o es we e no ma kedly diffe en in ma e ial espi a ion. Respi a ion o he humus lay- e s in bo h o es s ands was significan ly highe han espi a ion o mine al soils; i was also highe han es- pi a ion a he bo om and im o an nes s in bo h o es s ands, whe eas espi a ion o he mine al soils was lowe . When we calcula ed espi a ion pe g C, he diffe ences be ween indi idual pa s o nes s and nes su oundings we e no significan . The o he ma e ial p ope ies also conside ably di - e ed be ween he wo o es s ands (Table 1). An nes s in he bi ch o es we e on a e age mois e han an nes s in he pine o es wi h he highes mois u e a he op o an nes s. Ma e ial pH was simila in bo h o es s ands and was lowe a he ops o an nes s in compa ison wi h he bo oms and ims. The ca bon con en was he highes a he ops o an nes s in bo h o es s ands wi h g ea e diffe ences be ween sampling loca ions in he pine o es . C:N a io was on a e age highe in he pine o es wi h he highes alue a he B ough o you by | Me la Finnish Fo es Resea ch Ins i u e Au hen ica ed Download Da e | 12/21/16 11:43 AM Respi a ion o wood an nes ma e ial 1195 Fig. 1. Respi a ion o ma e ials pe g DW soil om sampling lo- ca ions in he bi ch and pine o es s ( h ee-way ANOVA, F4,56 = 22.35, P<0.001). TOP e e s o he op o he an nes s, BOTTOM o he base o he nes s, RIM o he pe iphe y o he nes s, and HUMUS and SOIL o he humus laye and mine al soil, espec i ely, as con ols. Columns wi h diffe en le e s a e s a- is ically diffe en (one-way ANOVA, Tukey HSD pos hoc es ). Means ±SEM a e shown. Fig. 2. Respi a ion o ma e ials pe g ca bon om sampling lo- ca ions in he bi ch and pine o es s ( h ee-way ANOVA, F4,56 = 1.42, P= 0.24). TOP e e s o he op o he an nes s, BOT- TOM o he base o he nes s, RIM o he pe iphe y o he nes s, and HUMUS and SOIL o he humus laye and mine al soil, e- spec i ely, as con ols. Means ±SEM a e shown. op o an nes s. In he bi ch o es , C:N a io did no diffe be ween he sampling loca ions. Discussion In ag eemen wi h ou hypo heses, ma e ial espi a ion pe g DW diffe ed be ween he sampling loca ions and, also, be ween he wo o es s ands. This confi ms ea - lie findings ha he same species can affec soil p op- e ies diffe en ly in diffe en ecosys ems (F ouz e al. 2003; Holec & F ouz 2006). Respi a ion o he ma e- ial om he op o an nes s in he bi ch o es was significan ly highe han espi a ion in he o he pa s o an nes s. Since li e decomposi ion is a complex p ocess dependen on mois u e, empe a u e, o li e quali y (Edwa ds e al. 1970; S eubing 1970; Paul & Cla k 1996), his significan diffe ence could be mos p obably explained by highe mois u e o he ma e ial om he op o an nes s oge he wi h qui e low C:N a io in he bi ch o es in compa ison wi h he pine o - es . Howe e , in he pine o es he espi a ion o he nes ma e ials om he diffe en pa s o an nes s was simila . The eason o missing significan diffe ences be ween mic osi es wi hin an nes s could be simila mois u e in all he ma e ials and C:N a io in he ma- e ial om he op wice as high as in he bo om and im o an nes s. Despi e he low pH o he humus laye s in bo h o es s ands, he humus laye espi a ion was highe han in he bo oms and ims o an nes s, which could be caused by highe mois u e and ca bon con en . Appa en ly, he mois u e, C:N a io, and ca bon con- en , sepa a ely o in combina ion, had an impo an effec on ma e ial espi a ion in ou samples. E en hough he empe a u e is conside ed o be he mos impo an ac o influencing he a e o o - ganic ma e u no e , i s o e all effec is complica ed by he in e ac ion be ween empe a u e and mois u e (Edwa ds e al. 1970). In addi ion o highe empe a- u es eco ded inside an mounds (Coenen-S aß e al. 1980), an inc ease o mois u e was ound s imula ing de- composi ion p ocesses o mound ma e ial (F ouz 1996; Lenoi e al. 2001). Bu as mounds a e usually d ie han he su ounding o es floo due o an cons uc - ing ac i i ies (Sei e 1996; Lafleu e al. 2002), decom- posi ion p ocesses can be lowe ed he e (Lenoi e al. 2001; Domisch e al. 2008). Ne e heless, decomposi ion in an nes s in he pine o es was no dec eased due o lowe mois u e con en , which sugges s ha he e a e o he p ope ies affec ing his p ocess, e.g. ca bon a ail- abili y. On he o he hand, ma e ials om an nes s in he bi ch o es had on a e age highe mois u e han in he pine o es , which means ha he li e decom- posi ion was p omo ed he e p obably due o highe mois u e con en . This esul is in acco dance wi h e- sul s o Coenen-S aß e al. (1980), who ound highe mois u e inside an nes s, and hus inc eased mic obial ac i i y. Ne e heless, he ma e ial espi a ion was affec ed no only by mois u e, bu also by he C:N a io, which was ound o be lowe in he bi ch o es and hus o be ano he ac o s imula ing decomposi ion in his o es s and. C:N a io a he op o an nes s in he pine o - es was wice as high in compa ison wi h he bo om and he im. Mound ma e ial a he op o an nes s in he pine o es consis s p edominan ly o needles, whe eas he bo om and he im consis o he mix- u e o needles and mine al soil (K is iansen & Amelung 2001). Needles om coni e ous ees con ain esis an cons i uen s such as cellulose and lignin, and o en ha e la ge C:N a io (S eubing 1970). Hence, he C:N a io a he op o an nes s in he pine o es was affec ed by he high C:N a io o needles. Howe e , since he a ailabili y o ca bon is e y impo an o he espi a ion o o ganisms (Schulze e al. 2000), we calcula ed he espi a ion pe g C and B ough o you by | Me la Finnish Fo es Resea ch Ins i u e Au hen ica ed Download Da e | 12/21/16 11:43 AM 1196 V. Jilko a e al. i showed diffe en esul s han he espi a ion pe g DW. The espi a ion inc eased in an nes s, especially in compa ison o he humus laye s. The mos impo an p ope ies ha p obably affec ed he espi a ion we e pH o he ma e ial and a ailabili y o ca bon. Highe pH a o s he ac i i y o bac e ia mo e han ha o ungi (Paul & Cla k 1996). Since bac e ia ha e highe a e o me abolism, i can hus lead o highe CO2p o- duc ion. In an nes s, an s accumula e as amoun s o ood, mainly honeydew, which consis s o simple sug- a s easily a ailable o mic oo ganisms. Mic oo ganisms ha e enough ene gy o decomposi ion p ocesses and ha also leads o highe CO2p oduc ion (i.e., p iming effec ). In conclusion, ou da a showed ha ma e ial de- composi ion, and hus CO2efflux, in ou o es s ands a e affec ed mainly by mois u e, ca bon con en and a ailabili y, C:N a io, and pH. Al hough we examined an mounds o he same species (Fo mica aquilonia)in bo h o es s ands, ma e ial p ope ies o an nes s di - e ed be ween he bi ch and he pine o es s. F om his we can conclude ha nes ma e ial p ope ies a e a - ec ed no only by he an species p esen and i s way o li e, eeding and nes cons uc ion (Pe al 1978), bu also by he o es s and cha ac e is ics. 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