Post-Fire Effect of Savannah Vegetation on the Establishment of New Colonies of Atta sexdens rubropilosa (Hymenoptera: Formicidae)

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1 Post-Fire Effect of Svnnh Vegettion on the Estblishment of New Colonies of Att sexdens rubropilos (Hymenopter: Formicide) Author(s): Denise Alves d Silv, Mrcelo Felix Luiz, Flávio Gonçlves de Jesus, Ednldo Cândido Roch, Mrco Antônio de Oliveir nd Márcio d Silv Arújo Source: Florid Entomologist, 99(4): Published By: Florid Entomologicl Society URL: BioOne ( is nonprofit, online ggregtion of core reserch in the biologicl, ecologicl, nd environmentl sciences. BioOne provides sustinble online pltform for over 170 journls nd books published by nonprofit societies, ssocitions, museums, institutions, nd presses. Your use of this PDF, the BioOne Web site, nd ll posted nd ssocited content indictes your cceptnce of BioOne s Terms of Use, vilble t Usge of BioOne content is strictly limited to personl, eductionl, nd non-commercil use. Commercil inquiries or rights nd permissions requests should be directed to the individul publisher s copyright holder. BioOne sees sustinble scholrly publishing s n inherently collbortive enterprise connecting uthors, nonprofit publishers, cdemic institutions, reserch librries, nd reserch funders in the common gol of mximizing ccess to criticl reserch.

2 Post-fire effect of svnnh vegettion on the estblishment of new colonies of Att sexdens rubropilos (Hymenopter: Formicide) Denise Alves d Silv 1, Mrcelo Felix Luiz 1, Flávio Gonçlves de Jesus 2, Ednldo Cândido Roch 1, Mrco Antônio de Oliveir 3, nd Márcio d Silv Arújo 1, * Abstrct Estblishing their initil colony is probbly the most criticl moment in the life of lef-cutting nts. The non-estblishment is connected to biotic nd biotic fctors, nd the high mortlity rtes of initil colonies re possibly ssocited with entomopthogenic or ntgonistic microorgnisms to the symbiotic fungus present in the soil, hosted by these nts. Fire in the vegettion, depending on the intensity, is known to cuse significnt chnges to the soil physicl, chemicl, nd microbiologicl properties. The impct of fire in svnnh vegettion (Cerrdo) on the estblishment of erly colonies of Att sexdens rubropilos Forel (Hymenopter: Formicide) ws evluted. For this end, two res were selected, one where there hd been n ccidentl fire, nd contiguous one with the sme size nd vegettion chrcteristics without burning. In these res nd in soil collected in the sme res nd stored in the lbortory, femles recently fertilized in the nuptil flight were plced to excvte the soil nd estblish their colonies. Post-fire chnges in the soil chemicl nd microbiologicl properties were quntified nd correlted successfully in the estblishment of new colonies of this lef-cutting nt. Under field conditions, the femles of A. sexdens rubropilos did not show preference for which res to excvte: the ones tht hd been burned or the ones tht were unburned; under this condition, no colony survived ccording to the evlution performed 120 d fter the nuptil flight. Under lbortory conditions, the mjority of the femles excvted the soil, whether it hd been burned or not. However, the estblishment of initil colonies ws significntly higher in soils collected fr from the surfce nd in res tht hd not directly been ffected by the fire, showing negtive effect of fire on colony estblishment under lbortory conditions. Key Words: femle; initil nest; nuptil flight; lef-cutting nt; fire Resumo Estbelecer su colôni inicil provvelmente sej o momento mis crítico d vid ds formigs-cortdeirs. O não-estbelecimento está ligdo ftores bióticos e bióticos e, possivelmente, s lts txs de mortlidde de colônis iniciis estejm ssocids os microrgnismos entomoptogênicos ou ntgônicos o fungo simbionte cultivdo por esss formigs presentes no solo. Sbe-se que o fogo n vegetção, dependendo de su intensidde, provoc lterções significtivs ns proprieddes físics, químics e microbiológics do solo. Assim, neste trblho vliou-se o impcto de um incêndio em vegetção de svn (cerrdão) no estbelecimento de colônis iniciis de Att sexdens rubropilos Forel (Hymenopter: Formicide). Pr isso, form selecionds áres onde ocorreu um queim cidentl e outr, de mesmo tmnho, com mesms crcterístics vegetcionis e contígu sem queim. Nesss áres e em solo coletdos nesss mesms áres e condiciondos em lbortório form colocds fêmes recém copulds no di d revod pr perfurrem o solo e fundrem sus colônis. Alterções ns proprieddes químics e microbiológics do solo pós-fogo form quntificds e correlcionds com sucesso no estbelecimento de novs colônis dess formig-cortdeir. Em condições de cmpo, s fêmes de A. sexdens rubropilos não presentrm preferênci pr perfurr o solo de áres queimds ou não e, ness condição, nenhum colôni sobreviveu, em vlição relizd 120 dis depois d revod. Em condições de lbortório, miori ds fêmes perfurou o solo, independentemente de o solo onde els form condicionds ter sido queimdo ou não. Entretnto, o estbelecimento inicil de colônis foi significtivmente mior em solos coletdos distntes d superfície e em res que não sofrerm ção diret do fogo, o que mostrou efeito negtivo do fogo no estbelecimento de colônis no lbortório. Plvrs Chve: fêmes; ninhos iniciis; voo nupcil; formigs-cortdeirs; fogo Unlike other disturbing physicl gents, fire is influenced by the type of plnt community, nd its occurrence plys key role in the ecosystem dynmics of mny world biomes (Wheln 1995). The direct effects nd subsequent implictions depend on the fire severity, which in turn depends on the vilbility nd rrngement of the combustible mteril, soil moisture nd temperture, topogrphy, wind, nd frequency nd durtion of het (Frizzo et l. 2011). The negtive effects of fire re in generl less evident in nts thn in other rthropods becuse most of them build nests in plces tht protect them ginst the intense het. Furthermore, due to their socil orgniztion, nts re dpted for fst recovery (Nves 1996). The direct impct of fire on Att nd Acromyrmex (Hymenopter: Formicide) nests in the field hs lredy been studied by some u- 1 Universidde Estdul de Goiás, GO 330, km 241, Anel Viário, s/n, , Ipmeri, GO, Brsil; E-mil: denise_florestl@hotmil.com (D. A. S.), mrcelofelixluiz@hotmil.com (M. F. L.), ednldoroch@yhoo.com.br (E. C. R.), mrcio.rujo@ueg.br (M. S. A.) 2 Instituto Federl Goino, , Urutí, GO, Brsil; E-mil: fgjgronomi@zipmil.com.br (F. G. J.) 3 Universidde Federl de Viços, , Florestl, MG, Brsil; E-mil: moliveirc@yhoo.com.br (M. A. O.) *Corresponding uthor; E-mil: mrcio.rujo@ueg.br (M. S. A.) Florid Entomologist Volume 99, No. 4

3 Silv et l.: Post-fire effect on the estblishment of Att sexdens rubropilos colonies 745 thors (Anjos et l. 1998; Arújo et l. 2004,b; Oliveir et l. 2011). High rtes of colony mortlity hve been reported in reforesttion res in Brzil during the fires (Anjos et l. 1998). In this sense, the impct on the colonies could be direct, s mentioned erlier, nd indirect becuse fter the fires there re scrce resources to be forged by worker nts. The mortlity of initil lef-cutting nt colonies is ssocited with the physicl, chemicl, nd microbiologicl conditions of the soil where the nts try to estblish them fter the nuptil flight (Bento et l. 1991). Accidentl fires nd controlled burns not only keep the soil bre until the beginning of rinfll but lso cuse chnges to the soil chemicl nd microbiologicl properties. They prevent the estblishment of colonies possibly due to microorgnisms tht re entomopthogenic or ntgonistic to the nt s symbiotic fungus (Arújo et l. 2003). The lef-cutting nts (Att) hve peculir behvior: After the nuptil flight, the newly fertilized femles fll to the ground, lose their wings, quickly strt moving round looking for plce to excvte their new nest, nd then cloister to form colony (Arújo et l. 2011). Femle preference for plces clered of vegettion, such s rod sides, open pths, or even res with little vegettion creted by fires, hs been observed in Att cephlotes (L.) nd in Acromyrmex octospinosus (Reich) (Cherrett 1968; Vsconcelos 1990), in Att sexdens (L.) (Diehl- Fleig 1995), in Acromyrmex stritus (Roger) (Diehl-Fleig & Roch 1998), nd in Att bispheric Forel (Arújo et l. 2003). The nuptil flights usully occur fter rins tht moisten the soil, which not only fcilittes excvtion but lso provides fvorble microenvironment for initil cultivtion of the nts symbiotic fungus (Bento et l. 1991; Arújo et l. 2003). The nest building process is the most crucil period in the life of n nt colony (Wilson 1971), nd filure of Att nts to estblish new colonies is directly ssocited with high femle mortlity due to predtion (Autuori 1950; Mintzer & Vinson 1985; Diehl-Fleig 1995), to rinfll (Mriconi 1970), nd to microorgnisms nturlly existing in soils (Bento et l. 1991; Arújo et l. 2003). Other fctors my lso ffect the estblishment of new lef-cutting nt nests, though they re not well known, nd one of these fctors my be fire. The long-term effects of fires re known to bring bout chnges to environmentl conditions nd shelters, s well s to food qulity (Mjer 1984; Collett 1998; Arújo et l. 2004,b). Given the pprent economic unimportnce nd complex procedures required to study the impct cused by fire on smll soil fun, little hs been reserched in this re when compred with plnts nd lrge vertebrtes. Consequently, the post-fire ecology of such orgnisms is not well understood (McCullough et l. 1998). The knowledge generlly ccepted bout nts is tht they re less ffected by fires thn mny other insects. However, the effect of climte conditions nd disorders cused by humn interference, such s controlled burns or fires, on the estblishment of new colonies of these eusocil insects is poorly studied (Arújo et l. 2003). Thus, this study ws imed t ssessing the impct of fire in svnnh vegettion re (Cerrdo) in the Ipmeri Region, Goiás Stte, Brzil, on the erly estblishment of Att sexdens rubropilos Forel colonies. Mterils nd Methods STUDY AREA This study ws conducted t the Experimentl Frm of Goiás Stte University, Ipmeri Câmpus, in the municiplity of Ipmeri, GO, Brzil. This site is locted t S, W, nd 764 m ltitude. In this re, the svnnh vegettion (in regenertion) type is dominnt nd is chrcterized by trees, bushes, nd subshrubs, with herbceous plnts composed predominntly of exotic grsses Urochlo decumbens Stpf nd Hyprrheni ruf (Nees) Stpf (Pocee), with no history of burns in the lst 6 yr. According to Köppen, the climte type is Aw. The verge temperture of the region is 21.9 C, with verge ir humidity rnging from 58 to 81% nd nnul rinfll of 1,448 mm, with round 80% of rin in Dec, Jn, nd Mr, nd the rest distributed minly throughout Oct, Nov, nd Feb (Rodrigues et l. 2009). The predominnt soil type is dystrophic red-yellow ltosol (Embrp 2006). STUDY SITES AND CHARACTERIZATIONS The re of the svnnh frgment studied ws 13 h. Four hectres of this re were burned. The rest of the frgment, which hd not directly been ffected by the fire, hd vegettion nd soil chrcteristics similr to the burned res. In order to infer the burning intensity, combustible mteril ws collected in five 1.0 m 2 smples rndomly selected hed of the firing line. This shllow combustible mteril, like dried leves nd shrub brnches nd grss, ws weighed. On the nuptil flight dy of A. sexdens rubropilos in the studied region, which hppened 67 d fter vegettion burning, homogenized soil from burned (B) nd unburned (NB) res, collected rndomly t multiple loctions (n = 15) nd 2 depths (0.0 to 10.0 cm nd 10.1 to 20.0 cm), ws plced in 250 ml plstic cups. From this mteril, only 1 smple of the soil ws tken, nd routine chemicl nlysis ws performed. Also, polyvinyl chloride (PVC) tubes (10 cm in dimeter) were used to collect whole section of soil (B nd NB) in the field, without disruption. For this purpose, 25 cm pieces of PVC tubes were pressure inserted into the soil with the use of mllet. After tht, PVC tubes contining the soil were pulled up nd tken to the lbortory. COLONY FOUNDATION IN THE FIELD In ech tretment frgment (B nd NB), 10 A. sexdens rubropilos femles fertilized on the nuptil flight dy were plced long 60 m trnsect seprted by 6.0 m. These queens were surrounded by plstic brrier to prevent them from escping their site nd prevent them from excvting the ground. All femles when cptured where without wings nd in serch of site to excvte their nest outside the experimentl re. Excvting behvior in regrd to soil to build the nest ws evluted for period of 24 h, nd the number of surviving colonies ws counted 120 d fter the femles hd excvted the sme soil. Under field conditions, the durtion of survivl of lef-cutting nt colonies on verge is between 71 nd 118 d when the workers mke the first scouting opening (Autuori 1941). Surviving colonies were considered those tht hd living queen with workers nd symbiotic fungus forms. In the surviving colonies, the presence of living queen ws sought, the numbers of workers, pupe, nd lrve counted, nd the dry weight of the symbiotic fungus colony mesured. COLONY FOUNDATION IN THE LABORATORY A newly fertilized femle ws plced into ech cup or PVC tube. Twenty-five femles were plced in plstic cups with B nd NB substrte tretments, nd 45 femles were plced in PVC tubes to excvte the soil. The method to observe lef-cutting nt colony estblishment under lbortory conditions ws the sme s tht dopted by Arújo et l. (2003). The continers were kept t room temperture, with dily wter supply in cotton pds ttched to their covers, which ensured fvorble environment for the development of the entire colony, s recommended by Bento et l. (1991). The prts of the soil (0.0 to 10.0 nd 10.1 to 20.0 cm deep) collected from the B nd NB res to fill the 250 ml plstic cups were subjected to chemicl nd microbiologicl nlyses. The chemicl nlysis ws the stndrd usully recommended for minerl fertiliztion of gricul-

4 Florid Entomologist Volume 99, No. 4 turl crops. The microbiologicl nlysis ws the sme s tht dopted by Arújo et l. (2003), where the mount of microorgnisms present in the soil ws inferred by mesuring the CO 2 production. STATISTICAL ANALYSES A Chi-squred (χ 2 ) test nd Mnn Whitney U tests were used to compre the femles choices for nest plce nd their successful estblishment of initil colonies. CO 2 production in the soils from B nd NB res collected t 2 depths ws compred using the Mnn Whitney U test. An nlysis of vrince (ANOVA) nd n HSD Tukey test were performed to compre the morphometric chrcteristics nd composition of surviving colonies between nests excvted in B nd NB res. The Levene test ws used to check for homogeneity of vrince between tretments, nd the Shpiro Wilk test ws used to verify whether the dt were normlly distributed. These nlyses were processed with the softwre SISVAR 5.3 (Ferreir 2008). Results SITE CHARACTERIZATIONS The dry weight (verge ± stndrd devition) of the combustible plnt mteril by 1.0 m 2 portion ws ± kg. The burning of this mteril cused chemicl chnges in the soil, especilly n increse in ph nd in K, P, C 2+, nd Mg 2+ (Tble 1). The orgnic mtter content of the soil ws 2.1% in the B res nd 1.8% in the NB res. The cly contents in the nlyzed smples were g/ kg in B nd g/kg in NB, mening tht the occurrence of fire did not ffect this soil chrcteristic. The mount of CO 2 relesed in the soil (vlue ccumulted during 10 d) did not differ between the B nd NB res for the 2 depths studied (Mnn Whitney U test, P > 0.05) (Fig. 1). COLONY FOUNDATION IN THE FIELD AND THE LABORATORY Under field conditions, A. sexdens rubropilos femles excvted B nd NB soils similrly (χ 2 = 1.82; P = 0.178), nd none of these excvting queens mnged to estblish their own colony (Tble 2). However, in the lbortory, the collection of soil smples with PVC tubes ws imed t obtining undisturbed smples to observe femles nturlly excvte them s under field conditions. The soils from B nd NB res were eqully excvted by the A. sexdens rubropilos femles (χ 2 = 1.91; P = 0.167) (Tble 3). No depth difference in initil chmbers ws observed in the soils from B nd NB res (ANOVA, F (1, 28) = 4.196). The verge (± stndrd error) depths of constructed chmbers were 8.0 ± 0.12 cm nd 8.5 ± 0.14 cm in the B nd NB res, respectively. Likewise, no difference ws observed in the dimensions of the initil chmber built between B nd NB soils (ANOVA, F (1, 28) = 4.194). Chmber dimensions were ± 6.02 cm 3 in B nd ± 4.64 cm 3 in NB. In PVC pipes, successful estblishment of initil nt colonies (11.1%) occurred in NB soils but not in B soils (Tble 3). The number of A. sexdens rubropilos femles tht excvted the soil from B nd NB res collected t 2 depths is shown in Tble 4. Soils from B or NB surfce res were eqully excvted by the femles (χ 2 = 0.80; P = 0.371), with similr rtes of initil colony survivl (χ 2 = 0.60; P = 0.440). Non-surfce soils collected t 10.1 to 20.0 cm depth from B or NB res were lso eqully excvted by the femles (χ 2 = 7.71; P = 0.057). However, in this sitution, the fire negtively ffected the initil colony estblishment: The survivl rte of initil nt colonies ws significntly greter in colonies tht were estblished in NB soils compred with B soils (χ 2 = 8.01; P = 0.004). The chrcteristics of lef-cutting nt colonies tht survived in the 250 ml plstic cups re shown in Figs. 2 nd 3. There ws no difference in the numbers of lrve, pupe, workers, nd symbiont fungl dry weight of initil colonies estblished in B or NB res (ANOVA, P > 0.05). Discussion The study site ws predominntly composed of U. decumbens nd redily enbled fire spreding. According to Mirnd et l. (2002), fire in Brzilin svnnhs is usully superficil nd mostly consumes herbceous plnts. The fire s unpredictbility mde it impossible to relibly mesure the speed of the firing line, nd therefore the burning intensity could not be estimted in this study. In previous study during controlled burning of dry sugr cne strw, which ws monitored with higher mteril volume thn tht used in this study, the fire intensity (615 KJ/m/s) ws low (Arújo et l. 2003). Bsed on this previous ssessment, the fire in our study could be considered of low intensity. The increse in nutrient content ws probbly due to the shes resulting from the fire nd incorported into the soil by the rins, s lso reported by Redin et l. (2011). According to Nuernberg et l. (1984) nd Ctteln & Vidor (1990), these nutrients directly foster soil microorgnisms. In our study, 67 d fter direct fire ction, the mount of CO 2 relesed in the soil (vlue ccumulted during 10 d) did not differ significntly between B nd NB res t the 2 depths studied. In the present work, the microorgnism community in the soil ws not identified, but it probbly interfered with the new colony setting of A. sexdens rubropilos (Rodrigues 2007). Up to the dte when we observed no survivl of the initil colonies (120 d fter nuptil flight) in the field, the microorgnism community ws most likely influenced by climte fluctution such s temperture nd rinfll. Under field conditions, none of the colonies survived in B or NB res (Tble 2). It is worth mentioning tht under such field conditions, the beginning of soil excvtion by the femles took plce under intense solr rdition, with the femles being completely exposed to direct rdition, fct tht probbly hindered their ctivity. Under lbortory conditions, the behvior of excvting the soil to estblish colony ws similr to tht observed by Arújo et l. (2003) with nother lef-cutting nt species, A. bispheric, which lso excvted the soil offered regrdless of its being from burned or unburned res. The higher excvtion rtes in the lbortory (91.1 nd 97.8% Tble 1. Chemicl nlysis of soils from burned res (B) nd surrounding unburned res (NB). Ipmeri, GO, Brzil, Nov Are Depth (cm) ph P K C 2+ Mg 2+ Al 3+ H + + Al 3+ OM b (mg/dm 3 ) (cmol c /dm 3 ) (%) B NB ph determined in wter; b OM = Orgnic mtter. Texture (g/kg)

5 Silv et l.: Post-fire effect on the estblishment of Att sexdens rubropilos colonies 747 Fig. 1. Microbil ctivity (men nd stndrd error) in soils collected t 2 depths (0.0 to 10.0 cm nd 10.1 to 20.0 cm) in res with burned nd unburned vegettion in Ipmeri, GO, Brzil. Mens followed by the sme letter (ccumulted CO 2 production t 10 d), for ech depth where the soil ws collected, did not differ from ech other (Mnn Whitney U test, P > 0.05). Tble 2. Number of femle lef-cutting nts (Att sexdens rubropilos) tht excvted the soil fter the nuptil flight in burned (B) nd unburned (NB) res, nd success in estblishing colonies fter 100 d in the field. Ipmeri, GO, Brzil, Nov Prmeter B NB χ 2 vlue P vlue Number of excvting femles n.s Successful estblishment Success in initil colony estblishment ws mesured through the presence of the queen, workers, lrve, nd symbiotic fungi 120 d fter femle reclusion; n.s. = not significnt. for B nd NB, respectively), when compred with those observed in the field (70 nd 40% for B nd NB, respectively), were probbly due to fvorble lbortory conditions such s temperture, reltive ir humidity, nd soil moisture. Under these fvorble lbortory conditions, no significnt difference ws observed in the depth of initil chmber construction or in chmber size in soils from B nd NB res. The colony depths were similr to those observed by Dell Luci & Arújo (1993) for the sme lef-cutting species in soil unffected by fire. In PVC tubes, femles were ble to estblish colonies in NB soils (success rte 11.1%) but not in B soils. Arújo et l. (2003) reported tht immeditely fter the fire, there ws reduction in the soil microbiologicl ctivity, nd tht fter few months, like in this study, nutrients originting from the burning my hve benefited microorgnisms, Tble 3. Estblishment nd survivl of lef-cutting nt (Att sexdens rubropilos) colonies in soil collected with PVC tubes nd stored in the lbortory (n = 45 femles). B: Soils from burned re; NB: soils from unburned re. Ipmeri, GO, Brzil. Prmeter B NB χ 2 vlue P vlue Number of excvting femles (%) n.s. Successful estblishment (%) * 0.02 Success in initil colony estblishment ws mesured through the presence of the queen, workers, lrve, nd symbiotic fungi 120 d fter femle reclusion. * = Significnt; n.s. = not significnt. including those not beneficil to the colony, which my hve occurred in our study becuse no colony survived in B soils. When the femles were llowed to excvte soil tht hd been collected t 2 soil depths (0.0 to 10.0 cm nd 10.1 to 20.0 cm) from B nd NB res, we observed tht, lthough femles eqully excvted soils from both res, the highest survivl rte of initil colonies ws in non-surfce soil from the NB re (Tble 4). Even without fire effect, there ws lower colony survivl in topsoils thn non-surfce soils, reinforcing the hypothesis of the ction of entomopthogenic microorgnisms, microprsites, or symbiotic fungi competitors of lef-cutting nts. Bento et l. (1991), without considering the fire influence, lso showed tht femles of lef-cutting nts prefer to estblish their colonies under lbortory conditions in soils from non-surfce res (A horizon) (Bento et l. 1991). Despite observing higher colony survivl in soils from NB res nd in non-surfce Tble 4. Percentge of femle lef-cutting nts (Att sexdens rubropilos) tht excvted the soil under lbortory conditions, nd their success in estblishing initil colonies in homogenized soils coming from res with burned (B) nd unburned (NB) vegettion collected t depths of 0.0 to 10.0 cm nd 10.1 to 20.0 cm (n = 25 femles). Ipmeri, GO, Brzil. Depth (cm) Excvting femles (%) B NB χ 2 vlue P vlue A 40 A A 88 Ab χ 2 vlue P vlue Rte of successful estblishment (%) B A 20 A A 68 Bb χ 2 vlue P vlue Success in initil colony estblishment ws mesured through the presence of the queen, workers, lrve, nd symbiotic fungi 120 d fter femle reclusion. Percentge followed by different uppercse letters within rows or different lowercse letters within columns indicte significnt differences between tretments (Chi-squred test, P < 0.05). NB

6 Florid Entomologist Volume 99, No. 4 Acknowledgments We cknowledge Universidde Estdul de Goiás for the finncil support through reserch llownce (PBIC/UEG), nd the Coordintion for the Improvement of Higher Eduction Personnel (CAPES) t AUXPE 2370/2014 Project. We thnk the editor nd the nonymous reviewers for their vluble comments nd suggestions to improve the mnuscript. We lso thnk the English nd Compny Ltd. for revising the English in the submitted version of the mnuscript. References Cited Fig. 2. Chrcteriztion of Att sexdens rubropilos surviving colonies in burned nd unburned res, fter 120 d of queen reclusion (femle). Mens (nd stndrd errors) followed by the sme letter did not differ from ech other (ANOVA, P > 0.05). soils, no significnt difference ws seen in the composition of surviving colonies (fungl dry weight, number of lrve, pupe, nd dults) (Figs. 2 nd 3). In summry, A. sexdens rubropilos femles did not show preference in excvting soils from res with burned or unburned vegettion, nd initil colony survivl ws higher in non-surfce soils, from res tht did not hve vegettion burning. However, the composition of initil colonies creted in the lbortory (numbers of lrve, pupe, nd dults, nd dry weight of symbiont fungus) ws similr in soils collected from burned nd unburned res. Fig. 3. Chrcteriztion of Att sexdens rubropilos surviving colonies in burned nd unburned res, fter 120 d of queen reclusion (femle), in soils from 2 depths ( cm nd cm). Mens (nd stndrd errors) followed by the sme letter did not differ from ech other (ANOVA, P > 0.05) Anjos N, Dell Luci TMC, Myhé-Nunes AJ Gui prático sobre formigs cortdeirs em reflorestmentos. Grff Cor Ltd., Ponte Nov, Brzil. Arújo MS, Dell Luci TMC, Ribeiro GA, Ksuy MCM Impcto d queim controld d cn-de-çúcr n nidificção e estbelecimento de colônis de Att bispheric Forel (Hymenopter: Formicide). Neotropicl Entomology 32: Arújo MS, Dell Luci TMC, Nscimento IC, Veig CE O fogo como gente de distúrbio em comunidde de formigs. Ecologi Austrl 14: Arújo MS, Dell Luci TMC, Picnço MC. 2004b. Impcto d queim d plhd d cn-de-çúcr no ritmo de forrgemento de Att bispheric Forel (Hymenopter, Formicide). Revist Brsileir de Zoologi 21: Arújo MS, Mrinho CGS, Oliveir MA, Ribeiro MMR, Dell Luci TMC Voo nupcil ou revod de formigs-cortdeirs, pp In Dell Luci TMC [ed.], Formigs-cortdeirs: d bioecologi o mnejo. UFV, Viços, Brzil. Autuori M Contribuição pr o conhecimento d súv (Att spp. Hymenopter: Formicidde). I: Evolução do Suveiro (Att sexdens rubropilos Forel, 1908). Arquivos do Instituto Biológico 12: Autuori M Contribuição pr o conhecimento d súv (Att spp. Hymenopter: Formicidde). V: Número de forms lds e redução dos suveiros iniciis. Arquivos do Instituto Biológico 19: Bento JMS, Dell Luci TMC, Muchovej RMC, Vilel EF Influênci d composição químic e d populção microbin de diferentes horizontes do solo no estbelecimento de suveiros iniciis de Att levigt (Hymenopter: Formicide) em lbortório. Anis d Sociedde Entomológic do Brsil 20: Ctteln JM, Vidor C Flutução n biomss, tividde e populção microbin do solo, em função de vrições mbientis. Revist Brsileir de Ciênci do Solo 14: Cherrett JM Some spects of the distribution of pest species of lef-cutting nts in the Cribben. Cribben Proceedings of the Americn Society for Horticulturl Science Tropicl Region 12: Collett NG Effects of two short rottion prescribed fires in utumn on surfce ctive rthropods in dry sclerophyll euclypt forest of west-centrl Victori. Forest Ecology Mngement 107: Dell Luci TMC, Arújo MS Fundção e estbelecimento de formigueiros, pp In Dell Luci TMC [ed.], As formigs cortdeirs. Folh de Viços, Viços, Brzil. Diehl-Fleig E Sucesso no estbelecimento de colônis de Acromyrmex stritus (Roger) (Hymenopter: Formicide). Anis d Sociedde de Entomologi do Brsil 24: Diehl-Fleig E, Roch ES Escolh de solo por fêmes de Acromyrmex stritus (Roger) (Hymenopter: Formicide) pr construção de ninho. Anis d Sociedde de Entomologi do Brsil 27: Embrp Sistem brsileiro de clssificção de solos. 2nd Edition. Embrp, Rio de Jneiro, Brzil. Ferreir DF Sisvr: um progrm pr nálises e ensino de esttístic. Revist Symposium 6: Frizzo TLM, Bonizário C, Borges MP, Vsconcelos HL Revisão dos efeitos do fogo sobre fun de formções svânics do Brsil. Oecologi Austrlis 15: Mjer JD Short-term responses of soil nd litter invertebrtes to cool utumn burn in Jrrh (Euclyptus mrgint) forest in Western Austrli. Pedobiologi 26: Mriconi FAM As súvs. Ceres, São Pulo, Brzil. McCullough DG, Werner RA, Neumnn D Fire nd insects in northern nd borel forest ecosystems of North Americ. Annul Review of Entomology 43:

7 Silv et l.: Post-fire effect on the estblishment of Att sexdens rubropilos colonies 749 Mintzer A, Vinson B Coopertive colony foundtion by femles of the lef-cutting nt Att texn in the lbortory. Journl of the New York Entomologicl Society 93: Mirnd HS, Bustmente MMC, Mirnd AC The fire fctor, pp In Oliveir PS, Mrquis RJ [eds.], The Cerrdos of Brzil. Columbi University Press, New York, New York. Nves MA Efeito do fogo n populção de formigs (Hymenopter: Formicine) em cerrdo do Distrito Federl, pp In Anis do Simpósio impcto ds queimds sobre os ecossistems e mudnçs globis, III Congresso de Ecologi do Brsil, 6 11 Oct 1996, Brsíli, Brzil. Nuernberg NJC, Vidor C, Stmmel JC Efeito de sucessões de culturs e tipos de dubção n densidde populcionl e tividde microbin do solo. Revist Brsileir de Ciênci do Solo 8: Oliveir MA, Arújo MS, Mrinho CGS, Ribeiro MMR, Dell Luci TMC Mnejo de formigs-cortdeirs, pp In Dell Luci TMC [ed.], Formigs-cortdeirs: d bioecologi o mnejo. UFV, Viços, Brzil. Redin M, Sntos GF, Miguel P, Deneg GL, Luptini M, Donel A, Souz EL Impctos d queim sobre tributos químicos, físicos e biológicos do solo. Ciênci Florestl 21: Rodrigues OD Influênci d comunidde microbin do solo no estbelecimento de suveiros iniciis de Att sexdens rubropilos Forel, 1908 (Hymenopter: Formicide). M.Sc. Disserttion, University of São Pulo, Pircicb, Brzil. Rodrigues RA, Oliveir GA, Fri ALP, Oliveir Júnior A Crcterizção climátic do entorno d Usin Hidrelétric Serr do Fco (GO), pp In Torres FTP, Dgnino RS, Oliveir Júnior A [eds.] Contribuições geográfics. Geogrphic, Ubá, Brzil. Vsconcelos HL Atividde de forrgemento de dus espécies de formigs cortdeirs (Att) em um florest primári d Amzôni Centrl. Insectes Sociux 3: Wheln RJ The Ecology of Fire. Cmbridge University Press, Cmbridge, United Kingdom, pdf (lst ccessed 5 Aug 2016). Wilson EO The Insect Societies. Hrvrd University Press, Cmbridge, Msschusetts.