Unveiling community patterns and trophic · 2018-08-22 · Unveiling community patterns and trophic niches of tropical and temperate ants using an integrative framework of ﬁeld

Unveiling community patterns and trophicniches of tropical and temperate antsusing an integrative framework of fielddata stable isotopes and fatty acidsFelix B Rosumek12 Nico Bluumlthgen1 Adrian Bruumlckner13Florian Menzel4 Gerhard Gebauer5 and Michael Heethoff1

1 Ecological Networks Technische Universitaumlt Darmstadt Darmstadt Germany2Department of Ecology and Zoology Federal University of Santa Catarina Florianoacutepolis Brazil3 Division of Biology and Biological Engineering California Institute of TechnologyPasadena CA USA

4 Institute of Organismic and Molecular Evolution Johannes-Gutenberg UniversitaumltMainz Mainz Germany

5 BayCEER ndash IBG Universitaumlt Bayreuth Bayreuth Germany

ABSTRACTBackground The use and partitioning of trophic resources is a central aspect ofcommunity function On the ground of tropical forests dozens of ant species may befound together and ecological mechanisms should act to allow such coexistence Onehypothesis states that niche specialization is higher in the tropics compared totemperate regions However trophic niches of most species are virtually unknownSeveral techniques might be combined to study trophic niche such as fieldobservations fatty acid analysis (FAA) and stable isotope analysis (SIA) In this workwe combine these three techniques to unveil partitioning of trophic resources in atropical and a temperate community We describe patterns of resource use comparethem between communities and test correlation and complementarity of methodsto unveil both community patterns and speciesrsquo nichesMethods Resource use was assessed with seven kinds of bait representing naturalresources available to ants Neutral lipid fatty acid (NLFA) profiles and d15N andd13C isotope signatures of the species were also obtained Community patternsand comparisons were analyzed with clustering correlations multivariate analysesand interaction networksResults Resource use structure was similar in both communities Niche breadths(Hprime) and network metrics (Q and H2prime) indicated similar levels of generalizationbetween communities A few species presented more specialized niches such asWasmannia auropunctata and Lasius fuliginosus Stable isotope signatures andNLFA profiles also indicated high generalization although the latter differedbetween communities with temperate species having higher amounts of fat andproportions of C181n9 Bait use and NLFA profile similarities were correlated aswell as speciesrsquo specialization indices (dprime) for the two methods Similarities in d15Nand bait use and in d13C and NLFA profiles were also correlatedDiscussion Our results agree with the recent view that specialization levels do notchange with latitude or species richness Partition of trophic resources alone does notexplain species coexistence in these communities and might act together with

How to cite this article Rosumek et al (2018) Unveiling community patterns and trophic niches of tropical and temperate ants using anintegrative framework of field data stable isotopes and fatty acids PeerJ 6e5467 DOI 107717peerj5467

Submitted 11 June 2018Accepted 27 July 2018Published 22 August 2018

Corresponding authorsFelix B RosumekrosumekhotmailcomMichael Heethoffheethoffbiotu-darmstadtde

Academic editorMarcio Pie

Additional Information andDeclarations can be found onpage 25

DOI 107717peerj5467

Distributed underCreative Commons CC-BY 40

behavioral and environmental mechanisms Temperate species presented NLFApatterns distinct from tropical ones which may be related to environmental factorsAll methods corresponded in their characterization of speciesrsquo niches to some extentand were robust enough to detect differences even in highly generalizedcommunities However their combination provides a more comprehensive picture ofresource use and it is particularly important to understand individual niches ofspecies FAA was applied here for the first time in ant ecology and proved to be avaluable tool due to its combination of specificity and temporal representativenessWe propose that a framework combining field observations with chemical analysis isvaluable to understand resource use in ant communities

Subjects Biodiversity Ecology EntomologyKeywords Formicidae Trophic niche Baits Fatty acids Stable isotopes Atlantic forestTemperate forest Trophic ecology Methodology Food resources

INTRODUCTIONThe use and partitioning of trophic resources is a central aspect of community functioningTrophic interactions govern the flux of matter and energy in food webs and lead toother fundamental interactions such as competition and mutualism (Polis amp Strong 1996Reitz amp Trumble 2002) Trophic niche partitioning is one of the most importantmechanisms allowing species coexistence and may ultimately link to evolutionaryprocesses of adaptation and character displacement (Schluter 2000)

Ants (Hymenoptera Formicidae) are among the most abundant groups ofinvertebrates in terrestrial ecosystems presenting a wide range of feeding habitsnesting sites and interactions with organisms from all trophic levels In general theyare regarded as omnivorous feeding on a combination of living prey dead arthropodsseeds and plant exudates (Bluumlthgen amp Feldhaar 2010 Lanan 2014) On the ground oftropical forests dozens of species may coexist at the same spot which raises thequestion how ecologically different are these species Although the role of interspecificcompetition in ant communities has recently been hotly debated (Cerdaacute Arnan amp Retana2013) the combination of high species richness with high biomass may lead toevolutionary pressure for more diversified niches MacArthur (1972) suggested thatspecialization increases in tropical communities and as a result more species cancoexist However this idea was put in question by recent studies (Schleuning et al 2012Morris et al 2014 Frank et al 2018) For ants behavioral and environmentalmechanisms of coexistence have been proposed (Cerdaacute Retana amp Cros 1997Andersen 2000 Parr amp Gibb 2012) The use of food resources itself is surprisinglyunderstudied and trophic niches of most species remain poorly known This isparticularly evident in rich tropical communities (Rosumek 2017) but also true for sometemperate species (Lanan 2014)

Field observations are the most straightforward way of gathering information butthere are trade-offs between the number of species studied (eg single species naturalhistory vs community patterns Medeiros amp Oliveira 2009 Houadria et al 2015)

Rosumek et al (2018) PeerJ DOI 107717peerj5467 231

the number of resources assessed (eg proteinsugar comparisons vs all resourcescollected by workers Kaspari amp Yanoviak 2001 Lopes 2007) and the sampling intensity(eg seasonal studies vs temporal ldquosnapshotsrdquo Albrecht amp Gotelli 2001 Rosumek 2017)Moreover many species present cryptic habits and the sheer complexity of interactionsmakes the assessment of trophic niches a laborious task Baiting is a method widelyused in ant ecology to assess communities and infer resource use (Bestelmeyer et al 2000)but it is affected by the aforementioned drawbacks

Several techniques have been applied in ecology to deal with these issues amongthem stable isotope analysis (SIA) and fatty acid analysis (FAA) Indirect techniquescould be faster and reduce fieldwork effort but also rely on several assumptions tointerpret their results Since every method has its assets and caveats the choice dependson the nature of the questions being asked (Birkhofer et al 2017) However this alsoworks the other way around complementary methods can be combined to provide adetailed and integrative perspective on the community being studied

Stable isotopes have been widely applied to address several questions in ant biology(Feldhaar Gebauer amp Bluumlthgen 2010) Most commonly used are the relative abundanceof heavy nitrogen (d15N) and carbon (d13C) (Hyodo 2015) d15N increases predictablywhen one organism consumes another thus indicating whether species are at the top orbottom of the food web (Heethoff amp Scheu 2016) d13C could be used to distinguishbetween main carbon sources at the bottom of the food web because C3 C4 and CAMplants have different signatures (OrsquoLeary 1988 Gannes Del Rio amp Koch 1998) SIAprovides time-representative clues about trophic position but limited information onspecific food sources or feeding behaviors For instance if two species feed exclusively onprimary consumers they will have similar d15N regardless of what prey items they actuallyconsume or whether the food is obtained through predation or scavenging As such stableisotope signatures are not suitable to calculate niche breadth or overlap or to be analyzedas species-resources interaction networks

Fatty acids obtained from the diet are mainly stored as neutral lipid fatty acids (NLFAs)in the fat body of insects Some fatty acids can be synthesized de novo by organismsfrom carbohydrates or other fatty acids Synthesis of C160 C180 and C181n9 (palmiticstearic and oleic acids) is widespread and they are the most abundant NLFAs in insects(Stanley-Samuelson et al 1988 but see Thompson 1973) Ability to synthesize otherNLFAs is highly variable among taxonomic groups such as C182n6 (linoleic acidMalcicka Visser amp Ellers 2018) When fatty acids are reliably assigned to specific foodsources they may act as biomarkers (Ruess amp Chamberlain 2010) Even when suchbiomarkers are not identified all fatty acids assimilated without modification (ie throughdirect trophic transfer) influence the composition of the fat reserves including the relativeamounts of de novo-synthesized NLFAs Hence the stored fat preserves information oningested carbon sources and NLFA profiles can be compared to infer niche differences(Budge Iverson amp Koopman 2006) However the application of FAA in field studies ofterrestrial organisms still is limited Most studies focused on soil detritivores such ascollembolans and nematodes (Ruess et al 2007 Haubert et al 2009 Ngosong et al 2009)So far FAA was not used to study trophic ecology of ants

In this work we combine field observations with SIA and FAA to unveil the use andpartitioning of trophic resources in a tropical and a temperate epigeic ant communityOur main goal is to describe patterns for each community and test differences betweencommunities and methods Specifically we aim to (1) assess use of multiple resourcesstable isotope signatures and NLFA profiles of the most abundant species in bothcommunities (2) compare patterns between communities using descriptive and statisticalapproaches (3) test whether different methods provide convergent or complementaryinformation on patterns of resource use

MATERIALS AND METHODS

BaitingFieldwork in Brazil was carried out in Florianoacutepolis (Desterro Conservation Unit2731prime38PrimeS 4830prime15PrimeW altitude ca 250 m) in December 2015 and January 2016 undersampling permit SISBIO 51173-1 (ICMBio) and export permits 15BR019038DF and17BR025207DF (IBAMA) The vegetation consists of a secondary Atlantic forest with atleast 60 years of regeneration High rainfall rate along the coast results in high productivityant species richness and a tropical aspect for the Atlantic forest even at higher latitudessuch as in our work (Silva amp Brandatildeo 2014) In Germany it was carried out in Darmstadt(Prinzenberg 4950prime14PrimeN 0840prime01PrimeE altitude ca 250 m) in July 2015 (no permitsneeded there) The vegetation consists of patches of mixed forest beech forest andorchards which were all covered by the sample grids

Sampling design followed similar protocols in both sites (Table 1) Seven bait typeswere offered as proxies for resources that are widely used by ants in general (Kaspari 2000Bluumlthgen amp Feldhaar 2010 Lanan 2014 for a full description of baits see SupplementalDocument S1) Sample points were distributed in grids and separated by 10 m In eachsampling session only a single bait was offered per point and bait types were randomizedamong points Baits were set up in transparent plastic boxes and retrieved after 90 minThis procedure was repeated in different days until all bait types had been offered ateach point (twice in Brazil) The design was based on Houadria et al (2015) and evaluatesuse of multiple resources differing from a typical cafeteria experiment which is designedto assess preferred resources (Krebs 1999)

Pitfall samplingWe performed a concomitant pitfall assessment to verify whether bait records representedwell the epigeic community (Table 1) One vial per sample point was previously buried toavoid the digging-in effect (Greenslade 1973) and replaced after collection for the nextround Pitfall and bait sampling were not performed simultaneously at the same pointVials were buried at ground level had six cm diameter and 150 ml volume and contained40 ml propylene glycol 50

Fatty acid analysisIn Brazil samples were obtained from baits and complemented by colony sampling inNovember 2017 We only used ants from melezitose sucrose and seed baits to avoid

interference of bait lipids In Germany they were obtained by colony samplingbetween July and August 2017 Samples were frozen at -18 C directly from the fieldTotal lipids were extracted from the ants whole body using one ml chloroformmethanolsolution 21 (vv) The solution was applied to SiOH-columns and the neutral parcel(= mono- di- and triglycerides) eluted with four ml chloroform Samples were analyzedwith gas chromatographyndashmass spectrometry following the same procedures describedin Rosumek et al (2017) NLFA amounts were obtained comparing their proportions toan internal standard (C190 in methanol ρi = 220 ngml) Ants were subsequently driedto obtain their lean dry weight (= without lipids)

Stable isotope analysisAnts collected in baits and conserved in ethanol 70 were used to analyze d15N and d13CC and N isotope abundances were measured in a dual element analysis mode with anelemental analyzer coupled to a continuous flow isotope ratio mass spectrometer asdescribed in Bidartondo et al (2004) Relative abundances were calculated following theequation dx = (RsampleRstandard - 1) 1000 [permil] where R denotes the ratio betweenheavy and light isotopes of samples and international standards (N2 in the air and CO2 inPeeDee belemnite) Gasters were removed prior to analysis to avoid interference of gutcontent (Bluumlthgen Gebauer amp Fiedler 2003)

Taxonomic identificationAnts were identified with taxonomic revisions and comparison to identified specimens incollections and AntWeb images (AntWeb 2016) Updated names were checked with

Table 1 Details of the sampling design applied in this study

Brazil Germany

Sampling effort

Sampling points 64 80

Period of the day Day and night Day

Baits 64 per resource per period (= 896 baits) 80 per resource (= 560 baits)

Pitfall sampling Three 10-h rounds per period (= 60 h) Three 12-h rounds (= 36 h)

Resource represented

Larger faster and harder prey Living crickets(Achaeta domesticus Linnaeus 1758)

Smaller slower and softer prey Living termites(Nasutitermitinae)

Living maggots(Lucilia sericata Meigen 1826)

Dead arthropods Crushed crickets and maggotsmealworms(Tenebrio molitor Linnaeus 1758)

Bird droppings Chicken feces from organicbreeding

Seeds Seed mixture of diverse sizes andshapes without elaiosomes

Seeds of Chelidonium majus(L) with elaiosomes

Oligosaccharides in honeydew Melezitose 20

Disaccharides in nectar and fruits Sucrose 20

Antcat (Bolton 2018) Identifications were partially confirmed by taxonomists (seeAcknowledgements)

In Brazil ants were identified to genus level with Baccaro et al (2015) and to specieslevel with AcanthognathusmdashGalvis amp Fernaacutendez (2009) AcromyrmexmdashGonccedilalves (1961)CephalotesmdashDe Andrade amp Baroni Urbani (1999) CrematogastermdashLongino (2003)CyphomyrmexmdashKempf (1965) and Snealling amp Longino (1992) GnamptogenysmdashLattke(1995) HylomyrmamdashKempf (1973) LinepithemamdashWild (2007) OctostrumamdashLongino(2013) Odontomachus and PachycondylamdashFernaacutendez (2008) PheidolemdashWilson (2003)WasmanniamdashLongino amp Fernaacutendez (2007) Camponotus and Strumigenys were identifiedsolely by comparison with collections

In Germany ants were identified to genus and species with Seifert (2007) Seifert ampSchultz (2009) and Radchenko amp Elmes (2010)

All material is stored in the collections of the Ecological Networks research groupTechnische Universitaumlt Darmstadt Darmstadt Germany and Department of Ecology andZoology Federal University of Santa Catarina Florianoacutepolis Brazil

Data analysisAs a first step we compared speciesrsquo incidences in baits and pitfalls (ie number ofsampling points where it was recorded with each method) We assumed incidence inpitfalls to represent abundance in the community and qualitatively compared it toincidence in baits to check whether common species were underrepresented in baitsTo account for different efficiencies between methods expected incidences were indicatedby a line of slope m = IbaitsIpitfalls where I is the sum of all incidences for each method(Houadria et al 2015)

Number of species replicates and individuals per sample differed between methodsbased on sample availability and ant size In Brazil we analyzed 24 species for baits41 for FAA and 31 for SIA Method comparisons were performed only with 22 speciesconsidered in all three datasets In Germany seven species were analyzed with all methodsFor a full list of recorded species and respective labels used in plots see Table S1

Unless noted otherwise similarity matrices were based on unweighted BrayndashCurtisdissimilarities andMantel tests and correlations used Spearmanrsquos coefficient (rho) Analyseswere run in R 343 (R Core Team 2017) and PAST 314 (Hammer Harper amp Ryan 2001)

For all bait analyses we used proportion of occurrence on each bait type relative tototal records for each species Only species with at least 10 records from five or moresample points were considered In Brazil day and night records were considered asindependent to calculate proportions For FAA we calculated proportions of eachNLFA relative to total composition and used average proportions for each species AllNLFAs with average proportion gt001 were considered For SIA we also used speciesrsquoaverages and analyzed d15N and d13C separately using Euclidean distances to buildsimilarity matrices A special case was Lasius fuliginosus in Germany which wasrepresented by a single colony that foraged over a large area Bait records from differentsample points were considered independent and chemical results represent the average ofdifferent samples from that colony

To analyze resource use we used clustering and network analysis UPGMA clusteringwas used for species to show functional groups based on similar use of resources andfor baits to show the structure of resource use in each community Statistical significanceof clusters was tested with SIMPROF (Clarke Somerfield amp Gorley 2008) using the packageldquoclustsigrdquo (Whitaker amp Christman 2015) A Mantel test was used to compare similaritymatrices of Brazil and Germany

For network analysis we used quantitative modularity (Q) (Dormann amp Strauss 2014)and specialization indices for speciesresources (dprime) and whole networks (H2prime) (BluumlthgenMenzel amp Bluumlthgen 2006) using the package ldquobipartiterdquo (Dormann Fruend amp Gruber2017) Modularity shows how compartmentalized is a community that is if there aregroups of species that strongly interact with groups of resources In turn dprime indicateswhether individual species are specialized in certain resources or resources that are usedby a specialized group of species H2prime is an extension of dprime and shows how specialized thenetwork is overall H2prime = 0 would mean that all species used resources in the sameproportions and H2prime = 1 that each species has its exclusive pattern of resource use

Specialization indices were also used to analyze species fatty acids contingencytables In this case they indicate how exclusively NLFAs are distributed across species(Bruumlckner amp Heethoff 2017) H2prime = 0 would mean that all compounds occur in the sameproportion in all species and H2prime = 1 that each species has its exclusive compoundsCorrespondingly relatively high dprime represents NLFAs that occur more exclusively incertain species or species with more exclusive proportions of certain NLFAs Low dprimemeansa compound that is widespread among species or species with similarly generalizedprofiles Additionally we tested whether the two communities differed in their overallNLFA composition with PERMANOVA using site as a fixed factor (Anderson 2001)Homogeneity of multivariate dispersion was tested a priori with PERMDISP (Anderson ampWalsh 2013) To detect which NLFAs contributed to differences we used SIMPER(Clarke 1993) These tests were performed using package ldquoveganrdquo (Oksanen et al 2017)

To test whether niche breadths and NLFA profile diversity were different betweencommunities at species level we calculated Shannon diversity indices for each ant speciesas Hprime = Spilnpi where pi is the proportion of each resource i used by the species orNLFA found in its profile and compared them with MannndashWhitney tests

To test whether particular NLFAs were related to use of certain resources weperformed principal component analyses (PCA) using baits species contingency tablesreplacing zeros by small values (0000001) and using centered log-ratio transformation todeal with the constant-sum constraint (Bruumlckner amp Heethoff 2017) PC axes werecorrelated with NLFAs using function ldquoenvfitrdquo from package ldquoveganrdquo We also comparedproportions amounts (in mgmg the amount of fat divided by lean dry weight) andunsaturation indices (UI the sum of percentages of each unsaturated NLFA multipliedby its number of double bounds) between Brazil and Germany with MannndashWhitney testsWe did this for total fat and the three most abundant NLFAs (C160 C180 and C181n9)To test whether there was a direct relationship between total fat amount and C181n9or total amount and UI we correlated values for all individual samples of each community(166 in Brazil 32 in Germany)

Finally to test whether the results yielded by the three methods were correlatedwe performed Mantel tests between similarity matrices of species for each methodWe also correlated speciesrsquo dprime values for baits and NLFAs to test whether specializationlevels were related

RESULTSUse of resourcesMost common species were recorded in baits in proportions similar to the expected giventheir frequency in the community (Fig S1 and Table S1) A few species wereunderrepresented in baits (eg Pachycondyla harpax Myrmica scabrinodis Stenammadebile) but in general species with few bait records were also rare in pitfalls Thus weconsider that a representative part of the epigeic communities was properly sampledDespite strong variation in total number of records the number of species recorded in eachbait was similar with exception of large prey (Table 2)

Similarities in resource use were correlated between Brazil and Germany (Fig 1Mantel test rho = 063 p = 003) In both communities large prey was set apart from theother resources being used less frequently and by fewer species Seeds and melezitosechanged positions between communities In Germany all ants used both sugarsindiscriminately while in Brazil several species used more sucrose (eg Camponotuszenon Gnamptogenys striatula Pachycondyla striata Odontomachus chelifer Solenopsissp6) and others used more melezitose (eg Pheidole aper Solenopsis sp8 Wasmanniaaffinis) (Table 2) Both modularity (QBR = 016 QGE = 014) and network specialization(H2primeBR = 013 H2primeGE = 012) were relatively low and similar between sites Species usedresources in different ways and a few were more specialized (see below) but there were noclear links between particular resources and species or groups of species

In Brazil W auropunctata occupied a highly specialized niche using only feces baitswhich lead to the highest dprime values for any species and resource Linepithema iniquum alsoshowed a relatively higher specialization level due to its preference for dead arthropods andlow use of sugars P striata and O chelifer used more large prey dead arthropods andsucrose C zenon grouped with them based on use of dead arthropods and sucrosebut avoided large prey P aper was the only species to have melezitose as its preferredresource Other species showed higher redundancy and clustered together including allSolenopsis and most Pheidole (Fig 1 Table 2)

In Germany only L fuliginosus showed a relatively high specialization level andclustered separately due to its almost exclusive use of animal resources (living prey anddead arthropods) Other species showed low specialization and dissimilarity (Fig 1Table 2)

Niche breadths were similar between communities (MannndashWhitney p = 044) AverageShannon index was 16 plusmn 04 SD in Brazil and 17 plusmn 01 SD in Germany (Table 2)

Fatty acidsTemperate species contained much higher amounts of total fat than tropical ones (Fig 2)Fatty acid compositions changed between communities (PERMANOVA r2 = 035

Table 2 Resource use of ant species in Brazil and Germany Values for the seven baits are given in of the total records for each species Onlyspecies with at least 10 records from five sample points are listed

Species Large prey Small preyDeadarthropods

Feces Seeds Melezitose Sucrose dprimeShannonindex

Records

Brazil

Camponotus zenon ndash 14 36 7 7 7 29 006 16 14

Gnamptogenys striatula 2 23 21 19 11 6 17 004 18 47

Linepithema iniquum 10 10 50 20 ndash 10 ndash 016 14 10

Linepithema micans ndash 6 31 6 19 19 19 003 17 16

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Odontomachus chelifer 26 5 19 5 5 10 31 012 17 42

Pachycondyla striata 14 3 42 ndash 1 6 34 016 13 88

Pheidole aper 4 ndash 15 19 7 37 19 008 16 27

Pheidole lucretii 4 4 26 8 14 20 24 002 18 50

Pheidole nesiota 4 9 20 4 16 25 21 002 18 89

Pheidole sarcina 4 8 16 14 20 18 22 001 19 51

Pheidole sigillata 4 10 24 10 16 13 22 000 18 91

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp3 ndash 4 16 8 32 20 20 005 16 25

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

Wasmannia affinis ndash 25 10 10 30 20 5 009 16 20

Wasmannia auropunctata ndash ndash ndash 100 ndash ndash ndash 062 0 19

dprime 017 009 009 024 014 007 009 H2prime = 013

Total richnessdagger 26 31 33 32 32 34 34

Total recordsdagger 107 203 422 215 344 327 366

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius fuliginosus 20 27 33 13 ndash ndash 7 031 15 15

Lasius niger 7 14 19 11 9 19 20 001 19 118

Lasius platythorax ndash ndash 17 17 4 30 30 011 15 23

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

Myrmica ruginodis ndash 10 27 14 6 18 24 003 17 49

Temnothorax nylanderi ndash 4 21 14 18 21 22 006 17 165

dprime 029 014 002 005 013 011 005 H2prime = 012

Notes Species not considered in comparisons between methodsdagger Including species with less than 10 recordsndash Species not recorded in this bait

p lt 001) Multivariate dispersion was heterogeneous being higher in Brazil than inGermany (PERMDISP F = 1132 p lt 001) This does not change the previousresult because in this case PERMANOVA becomes overly conservative (Anderson ampWalsh 2013)

The main reason for this difference was the predominant role of C181n9 in temperatespecies (SIMPER dissimilarity contribution = 47 p lt 001 Figs 2 and 3 Table 3) InBrazil composition was more balanced which led to higher proportions of C180(contribution = 21 p lt 001) although amounts were similar C160 was proportionallythe most abundant NLFA in Brazil and the difference from Germany was marginallysignificant (contribution = 20 p = 006) although amounts again were higher intemperate species A few other NLFAs had statistically significant but very smallcontributions to the difference (Table S2)

Fatty acid compositions were generalized overall but more homogeneous in Germanybecause of the predominance of C181n9 (H2primeBR = 009 H2primeGE = 003) Accordingly NLFAprofile diversity was higher in tropical species (average Shannon index = 15 plusmn 02 SD) thantemperate ones (09 plusmn 02 SD) (MannndashWhitney p lt 001) (Fig 3 Table 3)

In samples from Germany there was no correlation between total amount of fatand percentage of C181n9 (rho = 019 p = 030) or unsaturation index (rho = 024p = 089) In samples from Brazil there was weak negative correlation between total fatand both C181n9 (rho = -016 p = 004) and unsaturation index (rho = -022 p gt 001)

In Brazil several fatty acids were related to resource use (Fig 4 see Table S3 for PCAeigenvalues and full Envfit results) Species with higher C181n9 also used more dead

Figure 1 UPGMA clustering of resources and species in Brazil (A C) and Germany (B D) based onBrayndashCurtis dissimilarities Red lines link elements from the same statistically significant cluster(SIMPROF p lt 005) Full-size DOI 107717peerj5467fig-1

arthropods (Envfit r2 of the NLFA with PC axes = 032 p = 002) while C182unk1(an unidentified NLFA) was related to use of sucrose and large prey (r2 = 031 p = 003)C140 (mystric acid) tended to be higher in species that used more seeds feces andsmall prey (r2 = 039 p = 001) Notice that the first two Principal Components explainedonly 60 of the variance and linear regressions were not strong In Germany mostvariation was along the sugar-protein axis C180 and C170 (margaric acid) werestrongly correlated with PC axes (r2 = 084 p = 005 and r2 = 078 p = 004 respectively)Both were higher in species that used more sugars and C170 also was related to use offeces although its relative abundance was very low in all species (lt05 Table 3)

Stable isotopesIn Brazil W auropunctata presented distinctive signatures for both isotopes It was thespecies with highest d15N while most species ranged from 58 to 82 and six showedconspicuously lower signatures Besides W auropunctata d13C varied less ranging from-241 to -27 (Fig 5 Table 4)

In Germany d15N was lower overall ranging from 36 (Lasius niger) to -11 (Lasiusfuliginosus) Species varied little in d13C (from -254 to -263) with values within the rangeof Brazilian species (Fig 5 Table 4)

p lt 001 p = 029

p lt 001

C160 C180 C181n9 All NLFAs

(microg

p lt 001

p lt 001 p lt 001

C160 C180 C181n9 Unsaturation index

Figure 2 Comparison of the three most abundant NLFAs total amounts and unsaturation indicesbetween tropical and temperate species (a) Amounts (b) percentages Green = tropical species red= temperate species Significant differences are in bold (MannndashWhitney test)

Full-size DOI 107717peerj5467fig-2

Isotope signatures were correlated for tropical species (rho = 043 p = 002)For temperate species the correlation lacked statistical significance (rho = -046p = 03) but their inclusion slightly strengthened the correlation for all species together(rho = 047 p lt 001)

largeprey smallprey deadarth feces seeds melezitose sucrose

formfu lasifu lasini lasipl myrmrb myrmrg temnny

largepreysmallprey deadarth feces seeds melezitose sucrose

Figure 3 Networks of resource use and fatty acid composition in Brazil (A) and Germany (B) Specieslabels are standardized to represent 100 of resource useNLFA composition The width of connectinglines represents proportion of bait useNLFA abundance for each species BaitNLFA labels show the sumof proportions for the whole community Only species analyzed with all three methods are included

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

Dataset comparisonsIn Brazil similarities in bait use and NLFA profiles were correlated (Table 5) While d15Nsimilarities were correlated with similarities in bait use but not with NLFAs theopposite was found for d13C That is similar use of resources among species was reflectedin similar body fat composition and both were related to their long-term trophic positionalbeit in different ways In Germany no such correlations were found between datasets(although it was marginally significant for NLFAs and d15N)

minus15 minus10 minus05 00 05 10 15

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus15 minus10 minus05 00 05 10

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

Figure 4 Principal component analysis of resource use in Brazil (A) and Germany (B) Bluesetae = bait direction vectors Red setae = NLFAs correlated to the two main PC axes (Envfit onlystatistically significant relationships are plotted) Setae sizes indicate relative strength of the relationshipsbut are scaled independently for each plot Only species analyzed with all three methods are included

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

minus275 minus250 minus225 minus200 minus175

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Figure 5 Stable isotope signatures of species in Brazil (A) and Germany (B) Gray bars displaystandard deviations for species averages d15N and d13C are displayed in permil following the equationdescribed in the methods Full-size DOI 107717peerj5467fig-5

Table 4 Stable isotope signatures of ant species in Brazil and Germany Average d15N and d13C aregiven in permil following the equation described in the methods

Species d15N d13C Samples

Brazil

Acromyrmex aspersus 342 -2576 1

Camponotus lespesii 244 -2699 3

Camponotus zenon 350 -2506 4

Crematogaster nigropilosa 363 -2697 2

Gnamptogenys striatula 818 -2500 5

Linepithema iniquum 450 -2671 5

Linepithema micans 771 -2462 4

Linepithema pulex 763 -2648 4

Nylanderia sp1 594 -2512 5

Odontomachus chelifer 769 -2528 5

Pachycondyla striata 769 -2552 5

Pheidole aper 671 -2526 5

Pheidole avia 584 -2546 3

Pheidole lucretii 789 -2579 3

Pheidole nesiota 613 -2555 5

Pheidole sarcina 777 -2502 4

Pheidole sigillata 735 -2467 5

Pheidole sp1 640 -2518 5

Solenopsis sp1 614 -2512 5

Trachymyrmex sp1 229 -2677 1

Wasmannia affinis 628 -2628 4

Wasmannia auropunctata 1120 -1779 4

Germany

Formica fusca 315 -2572 5

Lasius fuliginosus -109 -2581 5

Lasius niger 363 -2631 5

Lasius platythorax 080 -2540 5

Myrmica rubra 178 -2603 5

Myrmica ruginodis 166 -2556 5

Temnothorax nylanderi 053 -2565 5

Notes Species not considered in comparisons between methods

Table 5 Correlations between methods in Brazil and Germany Results are for Mantel tests usingSpearmanrsquos rho based on similarities matrices (BrayndashCurtis for baits and NLFAs Euclidean distances forisotopes) Asterisks indicate significant correlations

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

Figure 6 Relationship between specialization indices (dprime) for bait use and NLFAs Green = tropicalspecies the dashed line indicates significant correlation red = temperate species no correlationobserved Only species analyzed with all three methods are included

Exclusiveness (dprime) of bait choices and NLFAs profiles were also correlated in Brazil(rho = 051 p = 002) suggesting that specialization in resources was reflected in morespecific compositions of fatty acids No correlation was observed in Germany (rho = -007p = 086) (Fig 6)

DISCUSSIONOur main findings in this work are (1) patterns of resource use are similar in bothcommunities although the role of oligosaccharides is distinct (2) both communitiesare similarly generalized in resource use regardless of species richness (3) temperateants present higher amounts of fat and more homogeneous NLFA compositions(4) composition and specialization in resource use and NLFAs are correlated and are alsorelated to speciesrsquo trophic position (5) some species show specialized behaviors that can bebetter understood by method complementarity

The hypothesis proposed by MacArthur (1972) suggested that specialization is higherin tropical communities because the environmental stability allows species to adapt tomore specialized niches without increasing extinction risk thus allowing more speciesto coexist However this idea was put in question by recent studies where thelatitude-richness-specialization link was not confirmed or an inverse trend was found(Schleuning et al 2012 Morris et al 2014 Frank et al 2018) Our work is not anexplicit test of this hypothesis but several results agree with the view that specializationdoes not necessarily increase with higher richness toward the tropics despite the differentnumber of species network metrics of resource use and niche breadths were similarlygeneralized in both communities fatty acid compositions were also highly generalizedalthough in this case in different level possibly due to other factors (see discussion on fattyacids below) cluster analysis of resource use showed similar patterns betweencommunities and both species clusters and stable isotopes indicated strong overlap insideeach community

The bait protocol we applied is efficient to assess niches of generalists andspecialized species were seldom recorded Nevertheless these generalists represent themajority of the communities (as highlighted by our pitfall data) and one might expect morediversified niches to allow coexistence but that was not the case The differenceswe observed might still play a role in coexistence of some species particularly when theyshare other traits such as O chelifer and Pachycondyla striata Both are large solitaryforaging Ponerinae species very common on the ground of the Atlantic forest butO chelifer is more predatory and Pachycondyla striata is more scavenging (Rosumek 2017)Coexistence is result of a complex interplay of habitat structure interspecific interactionsand species traits and no single factor governs ant community organization (CerdaacuteArnan amp Retana 2013) Trophic niche alone does not explain coexistence of the commonspecies in these two communities but likely is one of the many factors structuring them

Use of resourcesResource use in Brazil was discussed in detail in Rosumek (2017) as well as the literaturereview on trophic niche of our identified tropical species Large prey was the less used

resource because size and mobility of the prey limits which species are able toovercome them Small prey and feces were also relatively less used the first because also isrelatively challenging to acquire and the second probably due to smaller nutritional valueOn the other hand the other resources are nutritive and relatively easy to gatherparticularly dead arthropods which was by far the most used resource Consideringthe similarity in resource use patterns most general remarks in that work apply toGermany as well The two main differences we found are discussed below

The role of insect-synthesized oligosaccharides seems to be distinct between temperateand tropical communities In Brazil the aversion to melezitose showed by some speciescould represent a physiological constraint since tolerance to oligosaccharides differsamong ant species (Rosumek 2017) For ants without physiological constraints melezitoseuse might be opportunistic and does not necessarily mean that they interact withsap-sucking insects However honeydew is the only reliable source of oligosaccharides innature so the few species that preferred this sugar may engage in such interactions(particularly Pheidole aper) In Germany on the other hand all species used both sugarssimilarly The two Myrmica Lasius and Formica fusca are known to interact withsap-sucking insects and Temnothorax nylanderi uses honeydew opportunistically whendroplets fall on the ground (Seifert 2007)

Seeds were other resource used differently but this probably is consequence of ourmethodological choice of seeds with elaiosomes in Germany Elaiosomes are thought tomimic animal prey and attract predators and scavengers (Hughes Westoby amp Jurado1994) not only granivores Effectively elaiosomes of Chelidonium majus are attractive to awide range of ants (Reifenrath Becker amp Poethke 2012) However seeds were moreextensively used in Brazil A higher diversity of shape and sizes of seeds was offered therewhich allowed more ants to use them

Fatty acidsFatty acid compositions were generalized but differed between communities In GermanyC181n9 plays a prominent role making up for more than 70 of the NLFAs stored byants The amounts of fat also differed remarkably in average temperate ants storedover five times more fat Similarly high amounts of total fat and percentages of C181n9were observed in laboratory colonies of F fusca and M rubra (Rosumek et al 2017)which suggest that it might be a general trend for temperate species In Brazil NLFAabundance at community level was more balanced between C160 C180 and C181n9Both amounts and proportions of C181n9 were variable among species

Organisms can actively change their fatty acid composition in response toenvironmental factors and physiological needs (Stanley-Samuelson et al 1988)Temperature and balance between saturated and unsaturated NLFAs are importantbecause the fat body should present a certain fluidity that allows enzymes to access storednutrients (Ruess amp Chamberlain 2010) C181n9 seems to be the only unsaturated fattyacid that ants are able to synthesize by themselves in large amounts (Rosumek et al 2017)However there was no positive correlation between amount of fat and C181n9 orunsaturation index of samples which would be expected if C181n9 synthesis was a direct

mechanism of individuals to balance saturationunsaturation ratios (the weak negativecorrelation in Brazil also does not fit this hypothesis)

Therefore we suggest that differences in C181n9 percentages and total amounts couldbe consequence of two distinct environmental factors Under lower temperatures higherproportion of unsaturated fatty acids is needed to maintain lipid fluidity (Jagdale ampGordon 1997) Thus temperate species might be adapted to synthesize and store moreC181n9 to withstand the cold seasons If this hypothesis were correct the ants wouldmaintain this high proportion throughout the year since we collected in summer In turnhigh amount of fat could be a direct consequence of the marked seasonality intemperate regions These species might be adapted to quickly acquire and accumulateenergy reserves during the short warm season while there is less pressure for this inregions where resources are available throughout the year

We observed relationships between certain NLFAs and resources although overallthey were not strong and not necessarily result of direct trophic transfer C181n9was related to use of dead arthropods in Brazil This NLFA is considered aldquonecromonerdquo a chemical clue for recognition of corpses by ants and other insects(Sun amp Zhou 2013) so it presumably increases in dead arthropods However onlypolyunsaturated fatty acids can be degraded to form C181n9 during decompositionand C181n9 itself turns into C180 (Dent Forbes amp Stuart 2004) Thus for high C181n9to be a direct result of scavenging prey items should previously possess high levels ofunsaturation This might be an indirect effect as well scavenger ants might be betterat tracking and retrieving food items that are naturally rich in C181n9 No correlationwas found in Germany which could also be related to the special role of this NLFAin temperate species its predominance due to environmental factors may override itsdietary signal

C182n6 occurs independently of diet only in very small amounts and it is a potentialbiomarker (Rosumek et al 2017) The differences we observed among species are directresult of diet Its occurrence was more widespread in Brazil but we observed no clearcorrelation with specific resources C182n6 is found in elaiosomes seeds and otherarthropods in different amounts (Thompson 1973 Hughes Westoby amp Jurado 1994)Since it can come from different sources C182n6 cannot be straightforwardly used as abiomarker for specific diets but depends on a deeper analysis of the resources actuallyavailable in the habitat

The biological significance of the correlations of NLFAs and resources in Germany isdifficult to grasp C180 does not appear to be preferably synthesized from carbohydratescompared to C160 and C181n9 (Rosumek et al 2017) Adding to the fact that suchcorrelation was not found in Brazil this might not represent a physiological link betweensugar consumption and C180 synthesis With low number of species in Germany evenstrong correlations might be result of species-specific factors other than diet The samemight be said for C170 a fatty acid that occurs in very low amounts in several vegetableoils (Beare-Rogers Dieffenbacher amp Holm 2001)

Interestingly we did not observe any 183n3 or 183n6 (a- and -linolenic acids)Ants are not able to synthesize them and they are assimilated through direct trophic

transfer (Rosumek et al 2017) In the studied communities these fatty acids seem to becompletely absent from food sources used by ants This is an unexpected result sincetheir occurrence is well documented in elaiosomes and a wide range of insect groups thatmight serve as prey (Thompson 1973 Hughes Westoby amp Jurado 1994)

The use of fatty acids as biomarkers to track food sources is one of the greatest potentialsof this method However it might be more suitable to detritivore systems where thebiomarkers are distinctive membrane phospholipids from microorganisms thatdecompose specific resources and that end up stored in the fat reserves of the consumers(Ruess amp Chamberlain 2010) The NLFA profiles we observed are generalized and themost relevant fatty acids could represent distinct sources andor be synthesized de novo inlarge amounts The biomarker approach might not be suitable at community level forground ants contrary to NLFA profiles (see Method Comparison below) However itstill might be useful to unveil species-specific interactions or in contexts with less potentialsources that can be better tracked (eg leaf-litter or subterranean species)

Stable isotopesTrophic shift (ie the degree of change in isotopic ratios from one trophic level toanother) varies among taxonomic groups and according to other physiological factors(McCutchan et al 2003) ldquoTypicalrdquo values of ca 3permil for d15N and 1permil for d13C wereexperimentally observed in one ant species (Feldhaar Gebauer amp Bluumlthgen 2010)Establishing discrete trophic levels is unrealistic in most food webs particularly foromnivores such as ants (Polis amp Strong 1996) but species within the range of onetrophic shift are more likely to use resources in a similar way d15N ranges of ca 9permilwere observed for ant communities in other tropical forests representing three trophicshifts (Davidson et al 2003 Bihn Gebauer amp Brandl 2010) This is similar to ourrange of 89permil but discounting W auropunctata the remaining range of 61permil ismore similar to what was observed in an Australian forest (71permil Bluumlthgen Gebauer ampFiedler 2003) In Germany only Lasius fuliginosus presented a distinct signature In bothcommunities most species fell within the range of one trophic shift

d13C showed smaller but meaningful variations that were correlated to d15N d13Cis less applied to infer trophic levels as it is more sensitive to sample preservationmethod and diet composition (Tillberg et al 2006 Heethoff amp Scheu 2016) An averagechange of 061permil was observed in samples stored in ethanol by Tillberg et al (2006)However we observed correlations (including with NLFAsmdashsee below) despite thiseventual change and it would not affect the similarity among species and betweencommunities Primary consumers using distinct plant sources may present differencesof up to 20permil and this will influence the signature of secondary consumers (OrsquoLeary 1988Gannes Del Rio amp Koch 1998) However in our case only W auropunctata presentedsuch distinct value

Again both isotopes suggest that the core of these communities is composed bygeneralists that broadly use the same resources Since we did not establish baselines lowervalues in Germany do not necessarily mean lower trophic levels in this communityIsotope signatures for the same species are highly variable among sites in Europe

(Fiedler et al 2007) and this variation can be the result of either different isotope baselinesor actual changes in speciesrsquo ecological roles

Low d15N suggest that a species obtain most of their nitrogen from basal trophiclevels mainly plant sources (Bluumlthgen Gebauer amp Fiedler 2003 Davidson et al 2003)This fits the six species with lowest d15N in Brazil Two were fungus-growing ants(Acromyrmex aspersus Trachymyrmex sp1) which use mostly plant material to grow itsfungus The others were species that forage frequently on vegetation besides the ground(Camponotus lespesii Camponotus zenon Crematogaster nigropilosa Linepithemainiquum) Arboreal species that heavily rely on nectar or honeydew usually present lowd15N which may be the case for these species Linepithema represents well this trendthe two mainly ground-nesting species Linepithema micans and Linepithema pulexpresented higher signatures than the plant-nesting Linepithema iniquum (Wild 2007)

Community patterns and method comparisonThe correlations we observed between methods are interesting from both themethodological and the biological perspective From a methodological viewpoint forterrestrial animals this is the first time an empirical relationship is shown betweenpatterns of resource use and composition of stored fat in natural conditions and that bothrelate to their long-term trophic position Although differences between species weresmall these relationships were robust enough to be detected by different methods From abiological viewpoint it highlights several physiological mechanisms involved in suchrelationships We will discuss in the following some of these mechanisms as well as caveatsthat are often cited for these methods They probably still influence our results andcorrelations but did not completely override the patterns

A commonly cited caveat for using baits is that ants could be attracted to the mostlimited resources instead of the ones they use more often Evidence for this comes mainlyfrom nitrogen-deprived arboreal ants (Kaspari amp Yanoviak 2001) and some cases arediscussed below (see method complementarity) However this effect might be lesspronounced in epigeic species and our results suggest that there is convergence betweenbait attendance and medium- and long-term use of resources

Diet may significantly change NLFA composition in a few weeks (Rosumek et al 2017)and persist for a similar time (Haubert Pollierer amp Scheu 2011) Therefore theldquosnapshotrdquo of resource use we observed with baits should represent at least the seasonalpreferences of the species A seasonal study on NLFA compositions can bring valuableinformation on resource use changes or if they are stable throughout the year

Adult ants are thought to feed mostly on liquid foods due to the morphology of theproventriculus which prevents solid particles to pass from the crop to the midgut(Eisner amp Happ 1962) Larvae are able to process solids and possess a more diversified suitof enzymes and are sometimes called the ldquodigestive casterdquo of the colony (Houmllldobler ampWilson 1990 Erthal Peres Silva amp Ian Samuels 2007) Trophallaxis is an importantmechanism of food sharing between workers and larvae Our results suggest that thetrophic signal of NLFAs is not lost in this processes and that might be true even for solid

items such as arthropods or seeds However the similarities could be as well the result ofdirect digestion and assimilation of liquid sources (sugars hemolymph)

We also found correlations with stable isotopes They were weaker than betweenbaits and NLFAs and different for each isotope For d15N it shows that patterns ofresource use are more correlated with trophic level Protein amino acids must be obtainedfrom diet or synthesized from other nitrogenated compounds so the signal relative tonitrogen sources should be more preserved This also fits to the idea that d15N reflectslarval diet because it is in this stage that ants grow and build most of their biomass(Bluumlthgen amp Feldhaar 2010) On the other hand it makes sense that the signal relative tocarbon sources is related to NLFA composition We should note that we removed thegaster of the ants used in SIA so we observed only the signal of carbon incorporated inthe other body parts This is related to dietary carbon but a stronger signal could beexpected if the fat body is included

The low source-specificity of stable isotope signatures might also lead to relativelyweak correlations Pachycondyla striata and O chelifer had the same d15N despite theirdifferent preferences Other species that appear to be mostly scavengers had similar orhigher d15N than those two ldquopredatorsrdquo such as Linepithema micans Pheidole sarcinaPheidole lucretii and Pheidole sp4

In Germany no correlation was observed between methods This is probably aconsequence of the low number of species available in the community The relationshipsfound in Brazil might be valid for other communities although ecological context andphysiology might change their significance or strength

Species niches and method complementarityNiche differences were correlated at community level but the use of different techniquesallows better understanding of speciesrsquo niches Method complementarity is particularlyimportant if one is interested in the functional role of individual species not only in overallpatterns Some cases are described below

In Brazil W auropunctata was distinct from the remaining community both inresource use and isotopic signature (unfortunately no NLFA samples were obtained forthis species) Strong preference for feces is a novel behavior for this species known toinvade and dominate disturbed habitats but less dominant inside forests (Rosumek 2017)Its isotopic signature confirms that they have a highly differentiated diet and could bedirect result of a feces-rich diet In herbivorous mammals feces are usually enrichedin d15N relative to diet (Sponheimer et al 2003 Hwang Millar amp Longstaffe 2007)The proposed mechanism of 15N enrichment along trophic levels states that this happensdue to preferential excretion of 14N and it is assumed that most nitrogen is excreted in theurine which is depleted in 15N (Peterson amp Fry 1987 Gannes Del Rio amp Koch 1998 butsee Sponheimer et al 2003) However 15N-enriched feces were also observed in uricotelicorganisms such as birds and locusts (Webb Hedges amp Simpson 1998 Bird et al 2008)Thus high d15N is consistent with a diet based on 15N-enriched feces from otherconsumers The relationship with the d13C signature is less clear but it also suggestshigh specialization

This behavior might be a local adaptation but also could indicate thatW auropunctatashifts to less disputed resources inside native forests (although the exact resourcesused may be context-dependent) In Davidson (2005) Wasmannia species (includingW auropunctata) presented relatively high d15N and were considered highly carnivorousHowever our result shows that high d15N should not be taken from granted torepresent high-level consumers It might be a solid generalization for communities butother trophic pathways may lead to such signatures Due to their lack of specificityisotope signatures should be combined with field observations to provide reliableinformation at species level As another example the second highest d15N in our workwas observed in Pheidole sp7 a species that used mainly seeds and was seldom recorded inanimal (or feces) baits

Another example where results seem to be contradictory is L fuliginosus which showedstrong preference for animal baits but low d15N In this case the natural history of thespecies is well known and it strongly interacts with aphids particularly the giant oak aphidStomaphis quercus (Seifert 2007) This suggests that this aphidrsquos honeydew is not enrichedin 15N and has a composition similar to the plants on which they feed The honeydewsupply should be abundant since ants basically ignored sugar baits but also relatively poorin nitrogen which makes L fuliginosus use animal sources whenever possible A similarpattern may apply to Linepithema iniquum in Brazil which also combined low d15N withpreference for animal baits This species is also known to use extra-floral nectaries andhoneydew (Rosumek 2017) but does not have such strong and specific interactions as itstemperate counterpart

CONCLUSIONSIn this work we investigated two communities with three distinct methods and providedinformation on community patterns of resource use and speciesrsquo trophic niches Ourresults agree with the view that ant communities are mostly composed by generalistspecies that share similar resources and suggest that such patterns do not differbetween tropical and temperate communities Although high richness may lead to morespecialists in the tropics the generalist core of the community should be maintained by acombination of several factors

Overall we observed that the three methods corresponded in their characterization ofthe communities but their combination provided a more comprehensive picture ofresource use However the time and costs demanded should limit the broad application ofthis framework and some techniques are more suitable to answer particular questionsWe gave special focus on FAA in this work because it was the first time this method wasapplied to study ant ecology in the field Considering that NLFA profiles provide a moretime-representative snapshot than baits and are more specific than stable isotopes wesuggest FAA as a powerful tool to study trophic niche relationships in species-rich antcommunities It allows the researcher to obtain quantitative data related to diet withrelatively short fieldwork time or from systems where direct observation is limited andthen use it to infer niche breadths similarities and overlap However their use asbiomarkers has yet to be developed and seems to be limited for epigeic ant communities

Combining NLFA compositions with field observations is strongly recommended if theresearcher is interested in source-specificity Finally stable isotopes (particularly d15N)might be added as a long-term representation of trophic position which can corroborateor complement other results

ACKNOWLEDGEMENTSWe thank Cristian Klunk Frederico Rottgers Marcineiro Larissa Zanette da Silva LukasKauling and Felix Schilcher for assistance in fieldwork and sample sorting AnnaRuppenthal for assistance in fieldwork and FAA of German ants Christine Tiroch fortechnical assistance in EA-IRMS and Rodrigo Machado Feitosa Alexandre CasadeiFerreira Thiago Sanches Ranzani da Silva and Francisco Hita Garcia for taxonomicassistance We also thank the two anonymous reviewers for their valuable comments andsuggestions

ADDITIONAL INFORMATION AND DECLARATIONS

FundingFelix B Rosumek and Adrian Bruumlckner were supported by PhD scholarships from theBrazilian National Council of Technological and Scientific Development (CNPq) and theGerman National Academic Foundation (Studienstiftung des deutschen Volkes)respectively This study was funded by CNPq (2900752014-9) the German ResearchFoundation (DFG HE 45935-1) and the Open Access Publishing Fund of DFGTechnische Universitaumlt Darmstadt The funders had no role in study design datacollection and analysis decision to publish or preparation of the manuscript

Grant DisclosuresThe following grant information was disclosed by the authorsBrazilian National Council of Technological and Scientific Development (CNPq) 2900752014-9German National Academic Foundation (Studienstiftung des deutschen VolkesGerman Research Foundation (DFG) HE 45935-1Open Access Publishing Fund of DFGTechnische Universitaumlt Darmstadt

Competing InterestsThe authors declare that they have no competing interests

Author Contributions Felix B Rosumek conceived and designed the experiments performed the experimentsanalyzed the data contributed reagentsmaterialsanalysis tools prepared figures andortables authored or reviewed drafts of the paper approved the final draft

Nico Bluumlthgen conceived and designed the experiments contributed reagentsmaterialsanalysis tools authored or reviewed drafts of the paper approved the final draft

Adrian Bruumlckner conceived and designed the experiments analyzed the data authoredor reviewed drafts of the paper approved the final draft

Florian Menzel conceived and designed the experiments authored or reviewed drafts ofthe paper contributed reagentsmaterialsanalysis tools approved the final draft

Gerhard Gebauer performed the experiments contributed reagentsmaterialsanalysistools approved the final draft

Michael Heethoff conceived and designed the experiments contributed reagentsmaterialsanalysis tools authored or reviewed drafts of the paper approved the finaldraft

Field Study PermissionsThe following information was supplied relating to field study approvals (ie approvingbody and any reference numbers)

Fieldwork in Brazil was carried out under sampling permit SISBIO 51173-1 (ICMBio)and export permits 15BR019038DF and 17BR025207DF (IBAMA) No permits wereneeded in Germany

Data AvailabilityThe following information was supplied regarding data availability

All raw data used for the analysis of this article are provided in Tables 2ndash4 and Table S1

Supplemental InformationSupplemental information for this article can be found online at httpdxdoiorg107717peerj5467supplemental-information

REFERENCESAlbrecht M Gotelli NJ 2001 Spatial and temporal niche partitioning in grassland ants Oecologia

126(1)134ndash141 DOI 101007s004420000494

Andersen AN 2000 A global ecology of rainforest ants functional groups in relation toenvironmental stress and disturbance In Agosti D Majer JD Alonso LE Schultz TR edsAnts Standard Methods for Measuring and Monitoring Biodiversity Washington DCSmithsonian Institution Press 25ndash34

Anderson MJ 2001 A new method for non-parametric multivariate analysis of varianceAustral Ecology 26(1)32ndash46 DOI 101111j1442-9993200101070ppx

Anderson MJ Walsh DCI 2013 PERMANOVA ANOSIM and the Mantel test in the face ofheterogeneous dispersions what null hypothesis are you testing Ecological Monographs83(4)557ndash574 DOI 10189012-20101

AntWeb 2016 AntWeb Available at httpwwwantweborg (accessed 15 March 2016)

Baccaro FB Feitosa RM Fernaacutendez F Fernandes IO Izzo TJ de Souza JLP Solar RRC 2015Guia para os gecircneros de formigas do Brasil Manaus Editora INPA

Beare-Rogers JL Dieffenbacher A Holm JV 2001 Lexicon of lipid nutrition (IUPAC TechnicalReport) Pure and Applied Chemistry 73(4)685ndash744 DOI 101351pac200173040685

Bestelmeyer BT Agosti D Alonso LE Brandatildeo CRF Brown WL Delabie JHC Silvestre R2000 Field techniques for the study of ground-dwelling ants In Agosti D Majer JD Alonso LESchultz TR eds Ants Standard Methods for Measuring and Monitoring BiodiversityWashington DC Smithsonian Institution Press 122ndash144

Bidartondo MI Burghardt B Gebauer G Bruns TD Read DJ 2004 Changing partners in thedark isotopic and molecular evidence of ectomycorrhizal liaisons between forest orchids andtrees Proceedings of the Royal Society B Biological Sciences 271(1550)1799ndash1806DOI 101098rspb20042807

Bihn JH Gebauer G Brandl R 2010 Loss of functional diversity of ant assemblages in secondarytropical forests Ecology 91(3)782ndash792 DOI 10189008-12761

Bird MI Tait E Wurster CM Furness RW 2008 Stable carbon and nitrogen isotope analysisof avian uric acid Rapid Communications in Mass Spectrometry 22(21)3393ndash3400DOI 101002rcm3739

Birkhofer K Bylund H Dalin P Ferlian O Gagic V Hambaumlck PA Klapwijk M Mestre LRoubinet E Schroeder M Stenberg JA Porcel M Bjoumlrkman C Jonsson M 2017Methods toidentify the prey of invertebrate predators in terrestrial field studies Ecology and Evolution7(6)1942ndash1953 DOI 101002ece32791

Bluumlthgen N Feldhaar H 2010 Food and shelter how resources influence ant ecology In Lach LParr CL Abbott KL eds Ant Ecology Oxford Oxford University Press 115ndash136

Bluumlthgen N Gebauer G Fiedler K 2003 Disentangling a rainforest food web using stableisotopes dietary diversity in a species-rich ant community Oecologia 137(3)426ndash435DOI 101007s00442-003-1347-8

Bluumlthgen N Menzel F Bluumlthgen N 2006 Measuring specialization in species interactionnetworks BMC Ecology 69 DOI 1011861472-6785-6-9

Bolton B 2018 An online catalog of the ants of the world Available at httpantcatorg(accessed 4 June 2018)

Bruumlckner A Heethoff M 2017A chemo-ecologistsrsquo practical guide to compositional data analysisChemoecology 27(1)33ndash46 DOI 101007s00049-016-0227-8

Budge SM Iverson SJ Koopman HN 2006 Studying trophic ecology in marine ecosystems usingfatty acids a primer on analysis and interpretation Marine Mammal Science 22(4)759ndash801DOI 101111j1748-7692200600079x

Cerdaacute X Arnan X Retana J 2013 Is competition a significant hallmark of ant(Hymenoptera Formicidae) ecology Myrmecological News 18131ndash147

Cerdaacute X Retana J Cros S 1997 Thermal disruption of transitive hierarquies in Mediterraneanant communities Journal of Animal Ecology 66(3)363ndash374 DOI 1023075982

Clarke KR 1993 Non-parametric multivariate analyses of changes in community structureAustralian Journal of Ecology 18(1)117ndash143 DOI 101111j1442-99931993tb00438x

Clarke KR Somerfield PJ Gorley RN 2008 Testing of null hypotheses in exploratory communityanalyses similarity profiles and biota-environment linkage Journal of Experimental MarineBiology and Ecology 366(1ndash2)56ndash69 DOI 101016JJEMBE200807009

Davidson DW 2005 Ecological stoichiometry of ants in a New World rain forest Oecologia142(2)221ndash231 DOI 101007s00442-004-1722-0

Davidson DW Cook SC Snelling RR Chua TH 2003 Explaining the abundance of ants inlowland tropical rainforest canopies Science 300(5621)969ndash972 DOI 101126science1082074

De Andrade ML Baroni Urbani C 1999 Diversity and adaptation in the ant genus Cephalotespast and present Stuttgarter Beitraumlge zur Naturkunde Serie B (Geologie und Palaumlontologie)2711ndash889

Dent BB Forbes SL Stuart BH 2004 Review of human decomposition processes in soilEnvironmental Geology 45(4)576ndash585 DOI 101007s00254-003-0913-z

Dormann CF Fruend J Gruber B 2017 bipartite visualising bipartite networks andcalculating some (ecological) indices Version 208 Available at httpscranr-projectorgpackage=bipartite

Dormann CF Strauss R 2014 Amethod for detecting modules in quantitative bipartite networksMethods in Ecology and Evolution 5(1)90ndash98 DOI 1011112041-210X12139

Eisner T Happ GM 1962 The Infrabuccal pocket of a Formicine ant a social filtration devicePsyche A Journal of Entomology 69(3)107ndash116 DOI 101155196225068

Erthal M Peres Silva C Ian Samuels R 2007 Digestive enzymes in larvae of the leaf cutting antAcromyrmex subterraneus (Hymenoptera Formicidae Attini) Journal of Insect Physiology53(11)1101ndash1111 DOI 101016JJINSPHYS200706014

Feldhaar H Gebauer G Bluumlthgen N 2010 Stable isotopes past and future in exposing secrets ofant nutrition (Hymenoptera Formicidae) Myrmecological News 13(3)13

Fernaacutendez F 2008 Subfamilia Ponerinae s str In Jimeacutenez E Fernaacutendez F Arias TMLozano-Zambrano FH eds Sistemaacutetica Biogeografiacutea y Conservacioacuten de las Hormigas Cazadorasde Colombia Bogotaacute Instituto de Investigacioacuten de Recursos Bioloacutegicos Alexander vonHumboldt 123ndash218

Fiedler K Kuhlmann F Schlick-Steiner BC Steiner FM Gebauer G 2007 Stable N-isotopesignatures of central European antsndashassessing positions in a trophic gradient Insectes Sociaux54(4)393ndash402 DOI 101007s00040-007-0959-0

Frank K Krell F-T Slade EM Raine EH Chiew LY Schmitt T Vairappan CS Walter PBluumlthgen N 2018 Global dung webs high trophic generalism of dung beetles along thelatitudinal diversity gradient Ecology Letters 21(8)1229ndash1236 DOI 101111ele13095

Galvis JP Fernaacutendez F 2009 Ants of Colombia X Acanthognathus with the description of a newspecies (Hymenoptera Formicidae) Revista Colombiana de Entomologia 35245ndash249

Gannes LZ Del Rio CM Koch P 1998 Natural abundance variations in stable isotopes and theirpotential uses in animal physiological ecology Comparative Biochemistry and Physiology119(3)725ndash737 DOI 101016S1095-6433(98)01016-2

Gonccedilalves CR 1961 O gecircnero Acromyrmex no Brasil (Hym Formicidae) Studia Entomologica4113ndash180

Greenslade PJM 1973 Sampling ants with pitfall traps diggin-in effects Insectes Sociaux20(4)343ndash353 DOI 101007BF02226087

Hammer Oslash Harper DAT Ryan PD 2001 PAST paleontological statistics software package foreducation and data analysis Palaeontologia Electronica 41ndash9

Haubert D Birkhofer K Flieszligbach A Gehre M Scheu S Ruess L 2009 Trophic structureand major trophic links in conventional versus organic farming systems as indicatedby carbon stable isotope ratios of fatty acids Oikos 118(10)1579ndash1589DOI 101111j1600-0706200917587x

Haubert D Pollierer MM Scheu S 2011 Fatty acid patterns as biomarker for trophic interactionschanges after dietary switch and starvation Soil Biology and Biochemistry 43(3)490ndash494DOI 101016jsoilbio201010008

Heethoff M Scheu S 2016 Reliability of isotopic fractionation (D15N D13C) for the delimitationof trophic levels of oribatid mites diet strongly affects D13C but not D15N Soil Biology andBiochemistry 101124ndash129 DOI 101016jsoilbio201607013

Houmllldobler B Wilson EO 1990 The Ants Cambridge Harvard University Press

Houadria M Salas-Lopez A Orivel J Bluumlthgen N Menzel F 2015 Dietary and temporalniche differentiation in tropical antsmdashcan they explain local ant coexistence Biotropica47(2)208ndash217 DOI 101111btp12184

Hughes L Westoby M Jurado E 1994 Convergence of elaiosomes and insect prey evidence fromant foraging behaviour and fatty acid composition Functional Ecology 8(3)358ndash365DOI 1023072389829

Hwang YT Millar JS Longstaffe FJ 2007 Do d15N and d13C values of feces reflect the isotopiccomposition of diets in small mammals Canadian Journal of Zoology 85(3)388ndash396DOI 101139Z07-019

Hyodo F 2015Use of stable carbon and nitrogen isotopes in insect trophic ecology EntomologicalScience 18(3)295ndash312 DOI 101111ens12128

Jagdale GB Gordon R 1997 Effect of temperature on the composition of fatty acids in totallipids and phospholipids of entomopathogenic nematodes Journal of Thermal Biology22(4ndash5)245ndash251 DOI 101016S0306-4565(97)00019-3

Kaspari M 2000 A primer on ant ecology In Agosti D Majer JD Alonso LE Schultz TR edsAnts Standard Methods for Measuring and Monitoring Biodiversity Washington DCSmithsonian Institution Press 9ndash24

Kaspari M Yanoviak SP 2001 Bait use in tropical litter and canopy antsmdashevidence of differencesin nutrient limitation Biotropica 33(1)207ndash211 DOI 1016460006-3606(2001)033[0207BUITLA]20CO2

Kempf WW 1965 A revision of the neotropical fungus-growing ants of the genus CyphomyrmexMayr Part II Group of rimosus (Spinola) (Hym Formicidae) Studia Entomologica 8161ndash200

Kempf WW 1973 A revision of the neotropical myrmicine ant genus Hylomyrma Forel(Hymenoptera Formicidae) Studia Entomologica 16225ndash260

Krebs CJ 1999 Ecological Methodology Menlo Park BenjaminCummings

Lanan M 2014 Spatiotemporal resource distribution and foraging strategies of ants(Hymenoptera Formicidae) Myrmecological News 2053ndash70

Lattke JE 1995 Revision of the ant genus Gnamptogenys in the New World (HymenopteraFormicidae) Journal of Hymenoptera Research 4137ndash193

Longino JT 2003 The Crematogaster (Hymenoptera Formicidae Myrmicinae) of Costa RicaZootaxa 151(1)1ndash150 DOI 1011646zootaxa15111

Longino JT 2013 A revision of the ant genus Octostruma Forel 1912 (Hymenoptera Formicidae)Zootaxa 36991ndash61 DOI 1011646zootaxa369911

Longino JT Fernaacutendez F 2007 Taxonomic review of the genus Wasmannia Memoirs of theAmerican Entomological Institute 80271ndash289

Lopes BC 2007 Ecologia do forrageio por Cyphomyrmex morschi Emery (HymenopteraFormicidae) em vegetaccedilatildeo de restinga no Sul do Brasil Revista Brasileira de Zoologia24(1)52ndash56 DOI 101590S0101-81752007000100006

MacArthur RH 1972 Geographical Ecology Patterns in the Distribution of Species PrincetonPrinceton University Press

Malcicka M Visser B Ellers J 2018 An evolutionary perspective on linoleic acid synthesis inanimals Evolutionary Biology 45(1)15ndash26 DOI 101007s11692-017-9436-5

McCutchan JH Lewis WM Kendall C McGrath CC 2003 Variation in trophic shift for stableisotope ratios of carbon nitrogen and sulfur Oikos 102(2)378ndash390DOI 101034j1600-0706200312098x

Medeiros FNS Oliveira PS 2009 Season-dependent foraging patterns case study of a neotropicalforest-dwelling ant (Pachycondyla striata Ponerinae) In Jarau S Hrncir M eds FoodExploitation by Social Insects Ecological Behavioral and Theoretical Approaches Boca RatonTaylor amp Francis Group 81ndash95

Morris RJ Gripenberg S Lewis OT Roslin T 2014 Antagonistic interaction networks arestructured independently of latitude and host guild Ecology Letters 17(3)340ndash349DOI 101111ele12235

Ngosong C Raupp J Scheu S Ruess L 2009 Low importance for a fungal based food web inarable soils under mineral and organic fertilization indicated by Collembola grazers Soil Biologyand Biochemistry 41(11)2308ndash2317 DOI 101016jsoilbio200908015

Oksanen J Blanchet FG Friendly M Kindt R Legendre P McGlinn D Minchin PROrsquoHara RB Simpson GL Solymos P Stevens MHH Szoecs E Wagner H 2017 vegancommunity ecology package Version 25-2 Available at httpscranr-projectorgpackage=vegan

OrsquoLeary MH 1988 Carbon isotopes in photosynthesis Bioscience 38(5)328ndash336DOI 1023071310735

Parr CL Gibb H 2012 The discovery-dominance trade-off is the exception rather than the ruleJournal of Animal Ecology 81(1)233ndash241 DOI 101111j1365-2656201101899x

Peterson BJ Fry B 1987 Stable isotopes in ecosystem studies Annual Reviews of Ecology andSystematics 18292ndash320 DOI 101146annureves18110187001453

Polis GA Strong DR 1996 Food web complexity and community dynamics American Naturalist147(5)813ndash846 DOI 101086285880

R Core Team 2017 R A Language and Environment for Statistical ComputingVersion 343 Vienna the R Foundation for Statistical Computing Available athttpswwwR-projectorg

Radchenko A Elmes GW 2010 Myrmica Ants (Hymenoptera Formicidae) of the Old WorldWarszawa Natura optima dux Foundation

Reifenrath K Becker C Poethke HJ 2012 Diaspore trait preferences of dispersing antsJournal of Chemical Ecology 38(9)1093ndash1104 DOI 101007s10886-012-0174-y

Reitz SR Trumble JT 2002 Competitive displacement among insects and arachnidsAnnual Review of Entomology 47(1)435ndash465 DOI 101146annurevento47091201145227

Rosumek FB 2017 Natural history of ants what we (do not) know about trophic and temporalniches of neotropical species Sociobiology 64(3)244ndash255DOI 1013102sociobiologyv64i31623

Rosumek FB Bruumlckner A Bluumlthgen N Menzel F Heethoff M 2017 Patterns and dynamics ofneutral lipid fatty acids in ants-implications for ecological studies Frontiers in Zoology 14(1)36DOI 101186s12983-017-0221-1

Ruess L Chamberlain PM 2010 The fat that matters soil food web analysis using fatty acids andtheir carbon stable isotope signature Soil Biology and Biochemistry 42(11)1898ndash1910DOI 101016jsoilbio201007020

Ruess L Schuumltz K Migge-Kleian S Haumlggblom MM Kandeler E Scheu S 2007 Lipidcomposition of Collembola and their food resources in deciduous forest standsmdashimplicationsfor feeding strategies Soil Biology and Biochemistry 39(8)1990ndash2000DOI 101016jsoilbio200703002

Schleuning M Fruumlnd J Klein AM Abrahamczyk S Alarcoacuten R Albrecht M Andersson GKSBazarian S Boumlhning-Gaese K Bommarco R Dalsgaard B Dehling DM Gotlieb AHagen M Hickler T Holzschuh A Kaiser-Bunbury CN Kreft H Morris RJ Sandel BSutherland WJ Svenning JC Tscharntke T Watts S Weiner CN Werner M Williams NM

Winqvist C Dormann CF Bluumlthgen N 2012 Specialization of mutualistic interactionnetworks decreases toward tropical latitudes Current Biology 22(20)1925ndash1931DOI 101016jcub201208015

Schluter D 2000 Ecological character displacement in adaptive radiation American Naturalist156(S4)S4ndashS16 DOI 101086303412

Seifert B 2007 Die Ameisen Mittel-und Nordeuropas Tauer Lutra

Seifert B Schultz R 2009 A taxonomic revision of the Formica rufibarbis Fabricius 1793 group(Hymenoptera Formicidae) Myrmecological News 12255ndash272DOI 101007s12275-007-0115-6

Silva RR Brandatildeo CRF 2014 Ecosystem-wide morphological structure of leaf-litter antcommunities along a tropical latitudinal gradient PLOS ONE 9(3)e93049DOI 101371journalpone0093049

Snealling RR Longino JT 1992 Revisionary notes on the fungus-growing ants of the genusCyphomyrmex rimosus group (Hymenoptera Formicidae Attini) In Quintero D Aiello A edsInsects of Panama and Mesoamerica Selected Studies Oxford Oxford University Press 479ndash494

Sponheimer M Robinson TF Roeder BL Passey BH Ayliffe LK Cerling TE Dearing MDEhleringer JR 2003 An experimental study of nitrogen flux in llamas is 14N preferentiallyexcreted Journal of Archaeological Science 30(12)1649ndash1655DOI 101016S0305-4403(03)00066-9

Stanley-Samuelson DW Jurenka RA Cripps C Blomquist GJ de Renobales M 1988Fatty acids in insects composition metabolism and biological significance Archives of InsectBiochemistry and Physiology 9(1)1ndash33 DOI 101002arch940090102

Sun Q Zhou X 2013 Corpse management in social insects International Journal of BiologicalSciences 9(3)313ndash321 DOI 107150ijbs5781

Thompson SN 1973 A review and comparative characterization of the fatty acid compositions ofseven insect orders Comparative Biochemistry and Physiology Part B Comparative Biochemistry45(2)467ndash482 DOI 1010160305-0491(73)90078-3

Tillberg CV McCarthy DP Dolezal AG Suarez AV 2006Measuring the trophic ecology of antsusing stable isotopes Insectes Sociaux 53(1)65ndash69 DOI 101007s00040-005-0836-7

Webb SC Hedges RE Simpson SJ 1998 Diet quality influences the d13C and d15N of locusts andtheir biochemical components Journal of Experimental Biology 2012903ndash2911

Whitaker D Christman M 2015 clustsig significant cluster analysis Version 11 Available athttpsCRANR-projectorgpackage=clustsig

Wild AL 2007 Taxonomic Revision of the Ant Genus Linepithema (Hymenoptera Formicidae)Berkeley University of California Press

Wilson EO 2003 Pheidole in the New World A Dominant Hyperdiverse Ant Genus CambridgeHarvard University Press

Unveiling community patterns and trophic niches of tropical and temperate ants using an integrative framework of field data stable isotopes and fatty acids

Introduction

Materials and Methods

Results

Discussion

Conclusions

flink6

References

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behavioral and environmental mechanisms Temperate species presented NLFApatterns distinct from tropical ones which may be related to environmental factorsAll methods corresponded in their characterization of speciesrsquo niches to some extentand were robust enough to detect differences even in highly generalizedcommunities However their combination provides a more comprehensive picture ofresource use and it is particularly important to understand individual niches ofspecies FAA was applied here for the first time in ant ecology and proved to be avaluable tool due to its combination of specificity and temporal representativenessWe propose that a framework combining field observations with chemical analysis isvaluable to understand resource use in ant communities

Subjects Biodiversity Ecology EntomologyKeywords Formicidae Trophic niche Baits Fatty acids Stable isotopes Atlantic forestTemperate forest Trophic ecology Methodology Food resources

INTRODUCTIONThe use and partitioning of trophic resources is a central aspect of community functioningTrophic interactions govern the flux of matter and energy in food webs and lead toother fundamental interactions such as competition and mutualism (Polis amp Strong 1996Reitz amp Trumble 2002) Trophic niche partitioning is one of the most importantmechanisms allowing species coexistence and may ultimately link to evolutionaryprocesses of adaptation and character displacement (Schluter 2000)

Ants (Hymenoptera Formicidae) are among the most abundant groups ofinvertebrates in terrestrial ecosystems presenting a wide range of feeding habitsnesting sites and interactions with organisms from all trophic levels In general theyare regarded as omnivorous feeding on a combination of living prey dead arthropodsseeds and plant exudates (Bluumlthgen amp Feldhaar 2010 Lanan 2014) On the ground oftropical forests dozens of species may coexist at the same spot which raises thequestion how ecologically different are these species Although the role of interspecificcompetition in ant communities has recently been hotly debated (Cerdaacute Arnan amp Retana2013) the combination of high species richness with high biomass may lead toevolutionary pressure for more diversified niches MacArthur (1972) suggested thatspecialization increases in tropical communities and as a result more species cancoexist However this idea was put in question by recent studies (Schleuning et al 2012Morris et al 2014 Frank et al 2018) For ants behavioral and environmentalmechanisms of coexistence have been proposed (Cerdaacute Retana amp Cros 1997Andersen 2000 Parr amp Gibb 2012) The use of food resources itself is surprisinglyunderstudied and trophic niches of most species remain poorly known This isparticularly evident in rich tropical communities (Rosumek 2017) but also true for sometemperate species (Lanan 2014)

Field observations are the most straightforward way of gathering information butthere are trade-offs between the number of species studied (eg single species naturalhistory vs community patterns Medeiros amp Oliveira 2009 Houadria et al 2015)

Brazil Germany

Sampling effort

Records

Brazil

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

dprime 017 009 009 024 014 007 009 H2prime = 013

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius niger 7 14 19 11 9 19 20 001 19 118

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

dprime 029 014 002 005 013 011 005 H2prime = 012

p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Brazil Germany

Sampling effort

Records

Brazil

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

dprime 017 009 009 024 014 007 009 H2prime = 013

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius niger 7 14 19 11 9 19 20 001 19 118

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

dprime 029 014 002 005 013 011 005 H2prime = 012

p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Brazil Germany

Sampling effort

Records

Brazil

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

dprime 017 009 009 024 014 007 009 H2prime = 013

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius niger 7 14 19 11 9 19 20 001 19 118

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

dprime 029 014 002 005 013 011 005 H2prime = 012

p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Brazil Germany

Sampling effort

Records

Brazil

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

dprime 017 009 009 024 014 007 009 H2prime = 013

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius niger 7 14 19 11 9 19 20 001 19 118

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

dprime 029 014 002 005 013 011 005 H2prime = 012

p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Records

Brazil

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

dprime 017 009 009 024 014 007 009 H2prime = 013

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius niger 7 14 19 11 9 19 20 001 19 118

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

dprime 029 014 002 005 013 011 005 H2prime = 012

p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Records

Brazil

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

dprime 017 009 009 024 014 007 009 H2prime = 013

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius niger 7 14 19 11 9 19 20 001 19 118

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

dprime 029 014 002 005 013 011 005 H2prime = 012

p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Records

Brazil

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

dprime 017 009 009 024 014 007 009 H2prime = 013

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius niger 7 14 19 11 9 19 20 001 19 118

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

dprime 029 014 002 005 013 011 005 H2prime = 012

p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Records

Brazil

Nylanderia sp1 7 10 25 6 9 23 21 003 18 267

Pheidole sp1 3 13 19 10 18 17 21 001 19 101

Pheidole sp2 6 11 14 14 21 20 15 001 19 322

Pheidole sp4 5 6 21 19 14 14 21 001 19 78

Pheidole sp7 6 6 6 6 41 18 18 008 16 17

Solenopsis sp1 4 14 18 13 26 12 13 001 18 141

Solenopsis sp2 2 14 24 7 28 13 13 003 18 180

Solenopsis sp4 1 5 21 10 31 14 18 003 17 96

Solenopsis sp6 2 10 29 7 17 10 26 002 17 42

Solenopsis sp8 7 11 29 7 25 14 7 003 18 28

dprime 017 009 009 024 014 007 009 H2prime = 013

Germany

Formica fusca 4 5 21 2 12 26 30 006 16 57

Lasius niger 7 14 19 11 9 19 20 001 19 118

Myrmica rubra 3 13 15 13 3 25 30 003 17 40

dprime 029 014 002 005 013 011 005 H2prime = 012

p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


p lt 001 p = 029

p lt 001

(microg

p lt 001

p lt 001 p lt 001

ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


ttyacid

antspeciesin

Brazilan

dGerman

yAverage

individu

dances

aregivenin

thetotalcom

position

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Brazil

Acanthognathu

sbrevicornis

ndash03

ndashndash

Acanthognathu

socellatus

ndashndash

aspersus

ndashndash

laticeps

ndashndash

10ndash

ndashndash

13ndash

ndashndash

subterraneus

ndashndash

Aztecasp2

ndashndash

ponotuslespesii

ndashndash

ponotuszenon

ndashndash

Cephalotes

pallidicephalus

25ndash

ndashndash

Cephalotespu

ndashndash

Crematogaster

nigropilosa

ndashndash

Cyphomyrmex

rimosus

ndashndash

ndash34

ndashndash

mptogenys

striatula

ndashndash

yrmareitteri

ndashndash

ndash55

ndashndash

ndash005

Linepithem

ainiquu

ndashndash

Linepithem

amican

ndashndash

deriasp1

ndash49

ndashndash

Octostrum

apetiolata

ndashndash

Odontom

schelifer

ndashndash

Pachycondyla

harpax

ndashndash

Pachycondyla

striata

ndashndash

Pheidoleaper

ndashndash

ndash001

Pheidolelucretii

ndashndash

Pheidolenesiota

ndashndash

Pheidolerisii

ndashndash

Pheidolesarcina

ndashndash

Pheidolesigillata

ndashndash

Pheidolesp1

ndashndash

tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


tinued)

Species

aiC150

C161n7

C161n9

aiC170

C181n9

C181n11

C182n6

C182unk1

C182unk2

Shannon

Amount

ethlg=mgTHORNUI

Samples

Pheidolesp2

ndashndash

Pheidolesp4

ndashndash

Pheidolesp5

ndashndash

ndash27

ndashndash

ndash001

Pheidolesp6

ndashndash

34ndash

ndashndash

Pheidolesp7

ndashndash

Solenopsissp1

ndash44

ndashndash

Solenopsissp2

ndash43

ndashndash

Solenopsissp4

ndashndash

Solenopsissp6

ndashndash

Solenopsissp8

ndashndash

53ndash

ndashndash

Strumigenys

denticulata

ndashndash

38ndash

ndashndash

Wasman

affinis

ndashndash

dprimelt0

ndashndash

H2prime=003

German

icafusca

ndashndash

Lasius

fuliginosus

ndashndash

Lasius

ndashndash

ndash002

Lasius

platythorax

ndashndash

Myrmicarubra

ndashndash

ndash19

ndashndash

Myrmicaruginodis

ndashndash

ndash18

ndash16

ndashndash

nothorax

nyland

ndashndash

dprimelt0

ndashndash

H2prime=003

Speciesno

tconsidered

incomparisons

betweenmetho

ndashNLF

tdetected

inthisspeciesUIun

saturation

minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


minus1

5minus

minus0

PC1 = 368

largeprey

smallprey

deadarth

melezitose

sucrose

campze

gnamst

lineinlinemi

nyla01odonch

pachst

pheiap

pheilupheinepheisa

pheisi

phei01

phei02

phei04

phei07

sole01

sole02

sole04

sole06

sole08wasmaf

C181n9

C182unk1

minus1

0minus

PC1 = 518

largeprey

smallprey

deadarth

melezitose

sucrose

formfu

lasifu

lasini

lasipl

myrmrb

myrmrg

temnny C17

acroas

cample

campzecremni

gnamst

linein

linemilinepu

nyla01

tshcap hcnodo

phei01phei02

phei04

phei05

phei07

pheiap

pheiav

pheilu

pheine

pheisapheisi

sole01sole02

sole03

sole04sole06

sole08

trac01

wasmaf

wasmau

lasiniformfu

myrmrbmyrmrg

lasipltemnny

lasifu

minus25

Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Brazil

Germany

MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


MethodBaits NLFAs

rho p rho p

Brazil

NLFAs 043 lt001

d13C 023 007 023 002

d15N 025 004 014 008

Germany

NLFAs -023 077

d13C -024 076 029 019

d15N 037 012 046 006

formifu

lasifu

lasini

lasipl

myrmrbmyrmrg

temnny

campzegnamst

linein

linemi

nyla01

odonch

pachst

pheiap

pheilupheine

pheisapheisi

phei01

phei02

phei04

phei07

sole01sole02

sole04sole06

sole08

wasmaf

0000 0025 0050 0075

NLFAs d

126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


126(1)134ndash141 DOI 101007s004420000494

Introduction


Results

Discussion

Conclusions

flink6

References


Documents

Unveiling community patterns and trophic · 2018-08-22 · Unveiling community patterns and trophic niches of tropical and temperate ants using an integrative framework of ﬁeld