Posted on Authorea 11 Sep 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159986366.66136129 | This a preprint and has not been peer reviewed. Data may be preliminary. 09 nsai 00.Atitrcin ihncaiso ihple-edn net a affect can phloem-feeding with or nectaries with Lach interactions 2004; Ant (Beardsley Bronstein 2020). partners & Anastasio (Ness 2019; plants heterospecific Tarnita host from & Risch of nectar honeydew forests, extrafloral collect subalpine Milligan on and in feed 2004; can ants Bronstein ants, wood invasive native Second, & (e.g., expected First, carbon (Ness be al. of reasons. ants might several et distribution invasive ants for spatial invasive by dynamics the of displaced carbon modulate effects in often particular, shifts are In via invasions which ecosystems ant understood. of throughout consequences poorly reverberate the to are diverse, cycles and documented biogeochemical well for are impacts community-level their Although (Vanbergen pce,adotnla osit ntebhvo,fntoa oe n bnac fterntv counterparts native their of abundance Cabal and role, functional (Vanbergen behavior, networks the in shifts to (Holway lead (Bradshaw often and costs McGeoch species, economic species, foundational large native imposing suppressing 2010; (Pimenteland Herms biodiversity & Gandhi threatening cycles, Hill communities, biogeochemical 2008; ecological Englund affect profoundly 2001; can insects Non-native the highlighting scales. up, ecosystem scale at largely likely cycles Introduction resulted trees biogeochemical individual other fixation from and carbon results fixation reduced Our carbon that alter areas. to invaded demonstrate native in species we costly invasive herbivores energetically herbivores, of large of potential large by displacement trees and the on despite ants with season, browsing coinciding excluding growing from the rates experimentally during P. photosynthetic fixation By by on carbon ant- post-invasion effects whole-tree ants. lower and positive the 69% pre- year) africana) in of transpiration 1 resulted (Loxodonta is invasion stands (< and ant elephants photosynthesis shorter-term years, processes monodominant to 5 drepanolobium invades ecological vulnerable A ca. trees megacephala) fundamental After quantified We rendering megacephala. (Pheidole on and ungulates. ant effect defenders browsing big-headed ant their other native the and but displacing Africa, species, drepanolobium, East Acacia native In tree of interactions understood. and poorly assemblages shape ants Invasive Abstract 2020 11, September 4 3 2 1 Carpenter arc Milligan Patrick East foundational a ant-plant for African fixation carbon reduce ants Invasive aeUniversity Yale Wyoming Univ Conservancy Nature The Florida of University 03,o naei n-ln uulssta nunehs ln abndnmc Pige2016). (Pringle dynamics carbon plant host influence that mutualisms ant-plant in engage or 2013), tal. et tal. et 4 02 n olnto eg,Fuster (e.g., pollination and 2012) tal. et n odPalmer Todd and , 02 Bertelsmeier 2002; 2018). 1 ioh Martin Timothy , tal. et tal. et 08,itrutse ipra eg,Hriz&Shmk 96 Rodriguez- 1986; Schemske & Horvitz (e.g., dispersal seed interrupt 2018), 03,dsutn motn clgclpoess(oetcnp structure, canopy (forest processes ecological important disrupting 2013), 1 tal. et 1 rc John Grace , 07.I odig o-aieat a etutr pollination restructure can ants non-native doing, so In 2017). tal. et tal. et 1 06 Paini 2016; 00,adsra iessi olntrcommunities pollinator in diseases spread and 2020), 1 oin Riginos Corinna , tal. et tal. et 06.Ivsv nscmrs >240 comprise ants Invasive 2016). 92 Zhou 1982; 2 ao Goheen Jacob , tal. et tal. et 06,cndirectly can 2016), tal. et 07 Demian 2017; tal. et 05 Fin´er2005; tal. et 3 Scott , 2015), 2009) tal. et Posted on Authorea 11 Sep 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159986366.66136129 | This a preprint and has not been peer reviewed. Data may be preliminary. 0 do How trees? Site (3) invaded Study interactions? for plant photosynthesis vertebrate canopy and in changes ant-plant Methods to by contribute influenced ants rate invasive and that herbivores is vertebrate how and sites, invasion rmhmnhbtto ra nOCit lc-otnsvna o h past the for savannas black-cotton into OPC on al. areas habitation human on from damage heavy imposes disproportionately hot where and records) dry OPC typically availability, are forage periods intervening on and December, of to rainfall October monthly May; with to (March seasons wet in rainfall substantially regarding can questions photosynthesis research of by three of rates rate addressed plant photosynthetic We host leaf which 2010). the during Caylor ( seasons & long-term dry (King and and differ wet year) in invasion (<1 of short- impacts the how both evaluated over we 2011), (Simberloff how invader investigate to observations in and fixation experiments in canopy invasive field investigated (via fixation which conducted not We carbon by were canopy ant modes and invasive dynamics) potential this source-sink herbivores). are (via of by interactions rate damage role photosynthetic vertebrate-plant rate the leaf and photosynthetic impact and ant-plant may influences interactions ants both ants ant-plant and Thus, native direct King study. by but increase. herbivory their tree, herbivory) of host intense prevention of the the risk of that (through demonstrated costs (2010) longer-term Caylor as even invasion, Palmer after & immediately (Stanton habitats, mammals invaded large In mutualist, because by 2010). herbivory common by Palmer reduces most extirpated & and Huntzinger pletely The (Goheen 2018) (e.g., elephants 2004). Palmer domatia including (Palmer & spine 2011) (Prior ants hollow honeydew of and and thousands nectar nectar house extrafloral and producing time, feed a at species ( species sjostedti ant native four with mutualisms Laikipia, of vannas bnac,dsrbto,addvriyo aieiscs(es&Bosen20;Hffan&Pr 2008; Parr & Hoffmann 2004; Bronstein & (Ness insects native of diversity Riginos and Beardsley distribution, (e.g., abundance, Riginos insects 2004; phloem-feeding Bronstein ( & with tree (Ness thorn mutualists ant whistling native the tirpating species, foundation megacephala mono-dominant Pheidole cycles. and carbon widespread local a on in effects by disproportionate invasion have how investigated producers would We invasion primary that dominant such are could 2010), Frederickson that ants Palmer gardens, invasive ant-plants & Finally, devil’s invading fixation. (e.g., by communities carbon cycling some whole-plant and in affect carbon growth to ecosystem plant rates influence for exchange consequences especially carbon with leaf plants to host changes Nebauer on with Savage 1992; herbivory (e.g., Huber facilitate size & or canopy (Goldschmidt overall deter rates can exchange ants carbon invasive leaf Third, decrease or increase can which h abnsuc-ikrtoo otpat (Albani plants host of ratio source-sink carbon the ° ’26”,36 0’52.62”N, 05,weete cuytesadsi.Drn hsstudy, this During soil. and trees occupy they where 2015), .megacephala P. .megacephala P. tal. et .megacephala P. .drepanolobium A. ilwr a odce rmJl 07t etme 08a lPjt osrac (“OPC”; Conservancy Pejeta Ol at 2018 September to 2017 July from conducted was Fieldwork - 05 Milligan 2015; ° 15.4E 80maoesalvl.Ti 6 km 360 This level). sea above m 1800 51’58.64”E, .drepanolobium A. 2 ostela htsnhtcrt of rate photosynthetic leaf the Does (2) ? .megacephala P. a nae rpcladsbrpcleoytm rudtewrd(etrr21) ex- 2012), (Wetterer world the around ecosystems subtropical and tropical invaded has a nae (Riginos invaded has Mayr ca osntcnueetaflrlnca,hs re a xeineeegtcsavings energetic experience may trees host nectar, extra-floral consume not does 05 m(P eod) h P lpatpplto ( population elephant OPC The records). (OPC mm 30-50 . tal. et , eas h ffcso nainfeunl a eidteiiilarvlo the of arrival initial the behind lag frequently invasion of effects the Because . and .drepanolobium A. tal. et hioemegacephala Pheidole 06.I aansudranb lyrc etsl ie,‘lc-otn)sa- ‘black-cotton’) (i.e., vertisols clay-rich by underlain savannas In 2016). erpnr penzigi opie 9%o od oe (Young cover woody of >95% comprises hc osntdtrhrioe (Riginos herbivores deter not does which , 09 ah&Hffan21;Klkwk I22) hc a combine may which 2020), II Kulikowski 2011; Hoffmann & Lach 2009; rmtgse mimosae Crematogaster tal. et .megacephala P. tal. et 05.Ground-dwelling 2015). hnesotyatrteetraino otyatmutualists ant costly of extirpation the after shortly change ca tal. et er) eivsiae hs hr-adlong-term and short- these investigated We years). 5 . 92 Gaigher 1982; 2 arcu te“i-eddat)aet abncycling carbon affects ant”) “big-headed (the Fabricius ar.Hs lnsecuieyhueoentv ant native one house exclusively plants Host Mayr). tal. et 00 Del-Claro 2010; 2005; .drepanolobium A. nainiflecshs ln abnfixation carbon plant host influences invasion tal. et .megacephala P. 2 Santchi ccadrepanolobium Acacia tal. et osrac receives conservancy .megacephala P. 05,frigfclaiepartnerships facultative forming 2015), .megacephala P. tal. et rmtgse nigriceps Crematogaster , 03,btohriesuppressing otherwise but 2013), .mimosae C. tal. et .drepanolobium A. ca 06 ro amr2018) Palmer & Prior 2016; ute hnei ogterm long in change further tal. et .drepanolobium A. xeddec monitored each extended w eae (Riginos decades two . 96 n om obligate forms and 1996) .mimosae C. ccadrepanolobium Acacia nainaet carbon affects invasion ca 05.Hwvr and However, 2015). uulssaecom- are mutualists 3-0 depending 130-300 . nshv expanded have ants ca. aans Goheen savannas, 5-0 mof mm 250-300 tal. et tal. et consumes , 1 Does (1) : 04 to 2004) nareas in Emery 2011). et ). , Posted on Authorea 11 Sep 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.159986366.66136129 | This a preprint and has not been peer reviewed. Data may be preliminary. otoldcvte n ieyaehge hnntpooytei n rnprto fate nnaturally in tree a of transpiration and photosynthesis net than higher (McGarvey are conditions likely variable and cuvette, controlled idealized estimate A to our rates in transpiration treatment capacity” photosynthetic and per “canopy photosynthetic acacias [henceforth ( leaf 7-17 transpiration Uninvaded by and of and area photosynthesis subset whole-canopy Invaded leaf light-saturated random estimated long-term a multiplied ( In for and 2018 07:30-11:30. area S1), in leaf from canopy days experiment estimated cloudy factorial visually partly also or we light-saturated sunny sites, Leaf-level during canopy. 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(e.g., as trees both 20 reapplied and of ants, trunks specifications the health (Tanglefoot to public barriers Canada) Organization sticky m Health applied World 50 and sunlight; of x insecticides) full for total 50 in half-life a a day with from (2–3 density alpha-cypermethrin kg) plots, tree (>20 two and We herbivores comprised area sites). large site similar (“Uninvaded” exclude density of to tree plot site for comparable a invaded each S3). with containing at been (Fig. sites (0.25-ha) exclosure uninvaded treatments had away) plot fence four were acacias km electric in factors 2 where an Treatment resulting (< sites constructed 2018). neighboring absent), three May 3 vs. (July in in and (present dry and experiment two 2017 ants in our (November and rates seasons conducted absent) exchange We wet gas vs. and two (present potential and herbivores water 2018) large leaf measured September We and exchange. gas 2017 canopy invasive season. and of dry invasion leaf effects on of 2018 indirect impact September and years) direct and (>5 tested season from long-term wet excluded comparing and experiment 2018 2018 Factorial May May and the 2017 dry in December 2017 between Transition site elephants) July each by each analyses. at (evidently the at destroyed tall) In were surveys meters trees herbivores. repeated (1.5-2 Five large trees we to adult and accessible 20-24 2017, surveyed were sites. we by sites Control season, unaffected and All wet (pre-invasion) remained study. 2017 that the November sites) and of (“Control” season course non-manipulated site the surveyed Transition also over each and concurrent invasion, expansion from to after and km those before compared <1 sites) and trees (“Transition” invasion, front after invasion native and the by near before protected plots were trees that same trees the uninvaded on on measurements plant. potential water the leaf by and management rates water in (Gebrehiwot changes experiment acacias indicate African BACI to East rates other exchange of gas studies leaf to with relative that appropriate assumptions were confirm each 1) seasons For al. to “dry” seasons. dawn each and within before dry “wet” status and water and of mid-day soil rainy at similar potential had during sites water S3) all leaf measured (Fig. we experiments tree, surveyed factorial and (BACI) After-Control-Impact Regime Survey by front invasion sa daie siaeo rnprto;tesuffix the suffix transpiration; the of while area, estimate leaf idealized unit an is ” A max-leaf max-canopy 05 Gebrekirstos 2005; and E leaf E n cnp rnprto aaiy ( capacity” transpiration “canopy and ) emaue efgsecag pooytei n rnprto)i ocretBefore- concurrent in transpiration) and (photosynthesis exchange gas leaf measured We - leaf ]wr esrduigaL-40TPral htsnhssSse L-o Biosciences, (Li-Cor System Photosynthesis Portable LI-6400XT a using measured were )] ca oass hr-emipcsof impacts short-term assess To – r xrpltdfo a xhnertsmaue nteielevrnetwti a within environment ideal the in measured rates exchange gas from extrapolated are 0my Perke l nrevision). in al. et (Pietrek m/yr 50 . tal. et ca 0auttes(.- eestl) emre 0auttes(.- al in tall) m (1.5-2 trees adult 40 marked We tall). meters (1.5-2 trees adult 40 . 06,ad3 ocluaela ae oeta ag,wihcnb compared be can which range, potential water leaf calculate to 3) and 2006), canopy tal. et = N hioemegacephala Pheidole - ecnutdalpatpyilg esrmnso ul-xaddleaves fully-expanded on measurements physiology plant all conducted We 0 nwtseason, wet in 102 04.“ 2004). eest siae fteielzdcnp a xhne etherefore We exchange. gas canopy idealized the of estimates to refers ca. ca 0 rmec ecdpo osrea necdcnrl.Each controls. unfenced as serve to plot fenced each from m 200 0mre re tec ie efge aoiswt 0.6% with canopies fogged We site. each at trees marked 80 . .megacephala P. 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No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. .7 i.SB o ogtr sites long-term for S1B) Fig. 0.87; [?] (Henry (Lambers loss conductivity water (Gebrehiwot stomatal experience by will created leaf fundamentally ([?] a ([?] potentials that range conditions water potential water water leaf leaf mid-day diurnal calculated and we tree each For season dry Corvallis, and recorded Instruments, MPa region (PMS -1.5 the Chamber to Pressure -1.0 Plant 610 . Model ( a means fixation. using Treatment and carbon site OR). BACI study on invasion each both of for in measurements effect trees absolute ( for the pre-dawn exchange estimate measured We gas not in do they differences but relative experiments, estimate factorial to approximations these compare eddntfiddffrne in differences find decreased not slightly did also we but S1B), Fig. higher 0.0001; had areas Invaded in however, S1A); S1A), Fig. [?] oeta ags efgsecag ae,adcnp a xhnecpcte.Drn e esn,tesin trees seasons, wet During capacities. exchange gas canopy reduced and had rates, areas exchange Invaded gas physiology leaf drepanolobium ranges, Acacia potential on herbivores and mimosae occupants C. ant consistent of exhibited effects years(F sites Long-term across consistent both remained at trees trees Transition significantly while 1D), a 2017, Fig. exhibited in trees than Control 2018 while in trees, trees (F newly-invaded decline for 2017 interannual in smaller than 2018 1C; in Fig. lower 0.0027, substantially ( = traits P those 8.99, of = any in physiology- differences drepanolobium significant A. no on herbivores lower and had occupants trees ant Carolina, of North effects Cary, Short-term Institute, (SAS S2. 14.1.0 from Note Pro term in this Results but JMP removed are effect, we using GLMs random so conducted on a scores, were details AICc as Further higher site Analyses in USA). included resulted that analyses. and sampling models final models from tested all our data also for and we non-significant identity, was experiments, occupant term ant both this For within nested that pooled. effects were trees fixed periods were Control occupants ant paired constructed and ( the we herbivores identity invaded experiment, ant for for were GLMs, factorial (wet/dry) differences that experiment the season factorial interannual For trees each invasion. for to for terms no traits GLMs relative but interaction physiological separate surveys, conditions significant leaf environmental 2018 in report similar differences and specifically experienced (wet/dry) identify We 2017 to season the effects. results, each between fixed analysis for were BACI GLMs the term individual for interaction constructed their we and Control) experiment, BACI the for For experiments. rial - Analysis Statistical Hochberg 2017; ψ ψ A leaf leaf max-leaf sarsl fomtco tmtlajsmns(Inoue adjustments stomatal or osmotic of result a as eg,Gebrekirstos (e.g., A vs. max-canopy , E [?] A .megacephala P. leaf tal. et max-leaf leaf n [?] and , ψ i o infiatyvr (F vary significantly not did PD eue eeaie iermdl GM)t nlz aai h AIadfacto- and BACI the in data analyze to (GLMs) models linear generalized used We 08 Zhang 2018; tal. et ψ (F ± of lower , PD 1,94 A E)of SEM) tal. et ψ ca A max-leaf leaf [?] n i-a efwtrptnil( potential water leaf mid-day and ) A 08 Scoffoni 2008; max-leaf 1,127 20Mai r odtos(Gebrekirstos conditions dry in MPa -2.0 . 23,P 42.33, = nln-emUivddadIvddstsdffrdsbtnilyi efwater leaf in substantially differed sites Invaded and Uninvaded long-term in max-canopy E leaf 06,wiepooytei n rnprto a aywtotaffecting without vary can transpiration and photosynthesis while 2006), nteBC Ls apigya 21,21)adst ye(Transition, type site and 2018) (2017, year sampling GLMs, BACI the In . leaf . ψ tal. et 77,P 17.78, = F : PD ψ (F PD n higher and , 1,122 (F 1,205 ψ agdfrom ranged 09.Nt 1frhrdsrbste hsooymethods. physiology tree describes further S1 Note 2019). λεαφ .mimosae C. 1,212 and < (F .8 .48 i.1) uigdyseasons, dry During 1E). Fig. 0.0488; = P 3.88, = 15,P 11.55, = tal. et .01 i.2) and 2A), Fig. 0.0001; ψ [?] = 1,53 A .5 P 4.55, = MD A max-canopy < leaf max-leaf [?] ψ 1,193 .6 .0 i.2)or 2B) Fig. 0.80; = P 0.06, = [?] .1 i.1B). Fig. 0.01; ΠΔ r nTbe 2adS.Wtseason Wet S3. and S2 Tables in are 07.Pat ilotnrmi ihnaspecies-specific a within remain often will Plants 2017). leaf (F ca leaf tal. et 4 or 13,P=02;Fg 1) uigdysaos trees seasons, dry During S1C). Fig. 0.24; = P =1.39, 1,208 nbt er (F years both in 19t 21Ma tde frltdte pce in species tree related of studies MPa; -2.1 to -1.9 . < – n21 hni 07 u rniintesexhibited trees Transition but 2017, in than 2018 in .megacephala P. = tal. et ψ F : .0;Fg 2A), Fig. 0.001; ΜΔ 09 n nrae yls fvsua hydraulic vascular of loss by increased and 2019) .2 .34 i.SD by S1D) Fig. 0.0334; = P 4.52, = , E .39 i.2)and 2B) Fig. 0.0329; 1,131 canopy tal. et ). 05 Gebrekirstos 2005; ψ .6 P 5.16, = E ψ MD E [?] leaf leaf 07 atıe-iat Garcia-Forner Mart´ınez-Vilalta & 2017; canopy , ψ A ) leaf stedffrnebtenpre-dawn between difference the as ) max-leaf 1,127 ntesm a stegsexchange gas the as day same the on a infiatyhge o Control for higher significantly was a xdeet h xlso of exclusion the effect, fixed a was ) eosrtsterneo viable of range the demonstrates E tal. et (F = leaf .0 .9 i.1F). Fig. 0.99; = P 0.00, = 1,94 .21 i.1A; Fig. 0.0231, , (F E E 2006). E 27,P 42.70, = canopy 1,205 leaf tal. et leaf 1,127 nwtsaos Control seasons, wet In n [?] and , ψ 56,P 15.68, = (F 06:ta ag is range that 2006): re cuidby occupied Trees - .3 0.0332; = P 4.53, = PD (F 1,212 1,53 agdfrom ranged < A ca E ψ .6 P 5.96, = max-leaf .9 = P 0.49, = .01 Fig. 0.0001; leaf leaf . MPa; 0.1 . < nthe In . F : 0.0001; 1,131 was ca < Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. hioemegacephala Pheidole S3). (Note interaction their or exclusion Discussion herbivore removal, ant to due traits by occupied e ora oprbeuivddte uigtewtsao,apiaygoigpro o ayAfrican many for period growing primary a only season, of for wet invaded the capacity during been photosynthetic tree have canopy uninvaded that comparable a savannas a has In as tree areas. hour Uninvaded 2-meter-tall per vs. a areas years, Invaded 5 long-term in ( that capacity herbivory found of intensity thetic we maintenance low results, that short-term under suggest with growth Contrasting results support These dry to the trees. strategy in during invaded sustainable negated species target a apparently woody Gadd to be were on began only invasion forage 1999) may that of photosynthesis O’Connor feeders benefits mixed & These because (Illius invasion). perhaps rates season (after photosynthetic period, 2018 consistent drought had subsequent and trees invasion) the Transition (before while 2017 rates, photosynthetic in in (Caylor decline measurements interannual season large wet 2018 and 2017 Muraoka Wolf (e.g., & species that (Mathur a tree mechanisms Glanz-Idan apparatuses removes biochemical photosynthetic 2012; (28-31 likely and to Helliker molecular temperatures areas costly damage & invaded support heat-related recently (Wiley may mitigate availability in processes carbohydrate mutualists metabolic Increased nectivorous trees other before 2020). of control and rates neighboring loss upregulation, photosynthetic their The photosynthetic in season rates wet for period. photosynthetic sink similar time of carbohydrate maintain decline same to a that despite trees ecosystem in invasion, invaded on after newly immediately species enable and invasive may of that which suggest effects sites trees, Transition the from examining results Our studies long-term relevant 2011). of Strayer Simberloff ecologically (sensu 2005; value some of (Crooks processes that the effect habitat a net highlight illuminates in the rates, appearance of scales fixation initial sign their carbon in behind change leaf lag The on can invasion species invasive of of effects effect (Hooper positive processes initial ecosystem an fundamental alter can species invasive al. that demonstrating studies other with or 3D), in treatments these (F by occupied or 3A) by 4C) Fig. nwtrptnilrneadgsecag nbt e n r esn.Drn e esn,Ivddtrees Invaded seasons, wet During seasons. dry and wet and both of 3C) in cleared exchange experimentally gas were and that of range occurrence potential linked we water experiment, in factorial our in removal ant Through higher slightly or 3D), in Fig. differ 0.19; not did treatment [?] (F uigteriysao,tesi nae ra htwr xoe ovrert ebvrshdlower had herbivores vertebrate to exposed were that areas Invaded in trees season, A rainy the During max-leaf 2,53 2,94 leaf 05 Morales 2005; tal. et .8 .08 i.4)than 4B) Fig. 0.0038; = P 8.38, = 03,P 50.30, = E i o ie (F differ not did E [?] leaf canopy (F leaf 01 nfvrbeaitcconditions. abiotic favorable in 2001) ca 2,205 (F .mimosae C. .megacephala P. E (F 2,205 .5Ma eddntfidsgicn ieecsin differences significant find not did we MPa; 0.15 . leaf 2,208 (F .3 P 5.03, = [?] tal. et < 2,94 A .7 .9 i.SA ntescerdof cleared trees on S2A) Fig. 0.79; = P 0.07, = leaf ° .01 i.4) n o-infiatylower non-significantly and 4A), Fig. 0.0001; (F ) hc a as usata neana hne npeooyo te deciduous other of phenology in changes interannual substantial cause can which C), max-leaf .7 .1 i.4) ncnrs otesfo ogtr nae ie,trees sites, Invaded long-term from trees to contrast In 4D). Fig. 0.11; = P 2.57, = .drepanolobium A. nainsrnl nune ae fcro xto in fixation carbon of rates influenced strongly invasion 2,212 07.Teeln-emngtv ffcsof effects negative long-term These 2017). 2,193 07,P 10.77, = P=09;Fg D,btte nIvddaesta eeepsdt etbae had vertebrates to exposed were that areas Invaded in tree but 4D), Fig. 0.93; = (P tal. et tUivddstsddntehbtdffrn a xhneo efwtrpotential water leaf or exchange gas different exhibit not did sites Uninvaded at tal. et A = .9 P 3.99, = max-leaf .0 .6.Drn h r esn nae re nec herbivory each in trees invaded season, dry the During 0.66). = P 0.20, = .2;Fg 3A), Fig. 0.025; a lgtyhigher slightly had (F 2006). 2,212 00 eercre nteJnayMrhdysao htocre between occurred that season dry January-March the in recorded were 2010) < .megacephala P. .1 .2 i.3B), Fig. 0.92; = P 0.01, = = (F .0;Fg3) n lgtysmaller slightly and 3A), Fig 0.001; .megacephala P. .49 i 2)and S2B) Fig 0.0459; Satn&Ple 01,feigrsucst upr efgrowth, leaf support to resources freeing 2011), Palmer & (Stanton 2,212 .megacephala P. .4 .6 i.3B), Fig. 0.16; = P 1.94, = A .megacephala P. max-canopy E tal. et leaf .drepanolobium A. a higher had 5 nhs recro xto cosdffrn temporal different across fixation carbon tree host on (F 00.Ti a xli h oto re hwda showed trees Control why explain may This 2020). eoa re,btw on odffrne between differences no found we but trees, removal 2,212 (F 2,94 rgessottr eet o el invaded newly for benefits short-term triggers E .6 .17 i 2)and S2B) Fig 0.0117; = P 6.36, = canopy A E 30,P 53.07, = A .megacephala P. max-canopy leaf max-canopy A .megacephala P. a akdylwrcnp photosyn- canopy lower markedly had max-leaf (F (F tal. et .megacephala P. 2,53 2,205 [?] ca A < leaf (F .0 .19 i.4B). Fig. 0.0189; = P 5.70, = max-canopy .drepanolobium A. 04.Hg aiu daily maximum High 2014). (F .01 i.3) and 3C), Fig. 0.0001; .9 P 2.99, = (F 1 ftecro fixation carbon the of 31% . 2,94 (F 2,205 2,53 ncro xto,despite fixation, carbon on 2,193 05,P 10.57, = .1 .8 Fig. 0.08; = P 3.11, = ndysaos trees seasons, dry In . .9 .7 Fig. 0.67; = P 0.19, = nswt differences with ants .2 0.0378; = P 4.32, = = (F .8;Fg S2A); Fig. 0.084; 2,53 < .1 = P 1.71, = consistent , .0;Fig. 0.001; E E canopy canopy sensu et > Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. eetciaecag oespeitta h atArcnrgo ilsitt etrciae ihless with climates wetter to shift reduced. will is region pressure herbivore African if (Shongwe East have reversible century can the by next be that that the may tissues over so predict in droughts and severe models occur photosynthetic 2010) mechanisms change net Mohapatra proposed create climate & may These (Jha Recent thus trees. acacias and uninvaded in fixation, for turnover carbon level high in canopy increases change the large a at allow inducing benefits also without would capacities but and costly Additionally, rates (Inoue lically 1982). exchanges range gas Sharkey potential water high & leaf allow (Farquhar in can photosynthesis which to leaves, cost in a concentrations at po- management. ranges water water drepanolobium potential leaf A. leaf water in but leaf differences trees to similar uninvaded attributable in be and may invaded altogether long-term which drepanolobium between not, systems. differed did other exchange ranges and tential gas this canopy in and physiology & plant Leaf (Szarek host acacias affect to Nesting African ants other aquaporins). invasive (Moutinho of (e.g., enable content management membranes water Kebbas solute soil cell 1978; and and in Woodhouse status transpiration, channels water Lambers photosynthesis, water plant soils, influences of dominated affect status clay activation can in near and uptake ants water production invasive for by the factor key reduce (a or hairs ) root fine fewer produce also rudte ot nti edeprmn,btteecvto fns aiisaon ot a enobserved been has roots around cavities nest of excavation (Cockfield the taxa but experiment, plant field in many this can in of roots insects tree efficiency phloem-feeding around use of Haavik Infestations water canopy. 1992; and Whitlow the cerambycid rates in tolerate to not Milligan) photosynthetic appeared did P. and affect we observation 1999) Fraser While (pers. & canopy. infestations Eastwood the by Australia, larvae in in damage a association root pests to widespread by insect a driven though here, Palmer; other be physiology, relationships may with leaf these This interactions examine areas. impacted facultative invaded explicitly directly in through herbivores trees or large workers invaded of nesting long-term occurrence the in rates than photosynthetic ants degree in invasive lesser invasion. changes of ant similar after occurrence see herbivory to The intense expect experience may well that we is plants so rates and other photosynthetic species, for and plant concentrations trees invaded across phenolic on therein) leaf occurring references between be tradeoff Ishida could The (e.g., which well. 2002), as conserved Young photosynthesis example, sites For & canopy study (Ward areas. our herbivory of invaded at to in driver response function strongest in and branches the structure 1-m-tall leaf is to trees changes drepanolobium invaded induce A. in also can area herbivores leaf but decline, Reduced herbivores. large large that with showed (2010) Africa Caylor megacephala and Sub-Saharan P. King in for and plants rates 2010), photosynthetic Palmer woody ( leaf & many Goheen suppress Large of primary 2002; can habitats. Invaded Jacobs herbivores size the in & canopy is trees Biggs for herbivory the 1983; transpiration vertebrate (Pellew suppress and that photosynthesis elephants, canopy suggest particularly and experiment leaf herbivores, the exclusion in ant changes and of driver herbivore our from (Kurz Results insects non-native by are defoliation that to areas subjected leaf forests canopy by The magnified areas. greatly Invaded are vaded is that long-term but rates in photosynthetic trees, trees invaded leaf for for by season driven wet partially the is during difference This 1995). (Gourlay acacias rmtgse sjostedti Crematogaster .drepanolobium A. .drepanolobium A. .drepanolobium A. nln-emivddhbtt a ls tmt omnmz ae os hc ol result could which loss, water minimize to stomata close may habitats invaded long-term in naintigr ag hne nla n aoypooytei aaiywe co-occurring when capacity photosynthetic canopy and leaf in changes large triggers invasion nUivddhbtt a aemr abhdae ospottemngmn fsolute of management the support to carbohydrates more have may habitats Uninvaded in cuidb esdfnientv uulssices efpeo ocnrtosin concentrations phenol leaf increase mutualists native defensive less by occupied alns(Milligan saplings oteeciai hne a ie nivddaduivddsavannas. uninvaded and invaded in differ may changes climatic these to tal. et tal. et htieetvl eeshrioe Mris21) u eut iial hwthat show similarly results Our 2010). (Martins herbivores repels ineffectively that ) ssmlri antd odcie npooytei nNrhAeia hardwood American North in photosynthesis in declines to magnitude in similar is tal. et 08 Golan 2008; 08 Sumbele 2008; 05.Tu,tesml rcs fns xaainaon ln ot may roots plant around excavation nest of process simple the Thus, 2015). tal. et tal. et 07 Zhang 2017; tal. et .megacephala P. .megacephala P. tal. et npeaain.Mroe,rsuc-iie nae re may trees invaded resource-limited Moreover, preparation). in 05.W i o uniytesz of size the quantify not did We 2015). tal. et 02 n lsi eg,Kea inmt 06and 2016 Niinemets & Keenan (e.g., plastic and 2012) tal. et .drepanolobium A. 6 01 Haile 2011; 09.Teeomtcajsmnscnb metabo- be can adjustments osmotic These 2019). die eln npooytei aaiyfrin- for capacity photosynthetic in decline -driven tal. et ed yandctrilr pr.osrainT. observation (pers. caterpillars lycaenid tends 08 Albani 2008; tal. et 00,adpyilgcladjustments physiological and 2020), re hnocpe yantv ant native a by occupied when trees tal. et 00 Clark 2010; tal. et tal. et .megacephala P. 03,adwater and 2003), 97 ee & Meyer 1987; ca. ca. tal. et tal et 5 lower 65% 3 lower 13% Acacia 2010). 2008 . nests ca . Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. ybotatvt n olrsiain iia oeet eni ag-cl idigeprmn naboreal a in N-fixing experiment both girdling large-scale reduce a turn in in seen could effects this to habitats, similar invaded respiration, and in savannas, soil roots and cotton activity to black symbiont allocation in contributes. photosynthate productivity tree plant their foundational for reduce this resource which limiting to a nolobium processes is reduces ecosystem nitrogen habitat other invaded example, affect For in may ant-plants which for longer pool, capacity photosynthetic across carbohydrate photosynthesis of loss on long-term effects protection. The net herbivore positive effective plants yielding to other while owing plants, against scales host competitiveness time to increasing costs even metabolic or significant 2009) Holland (Fiala & (Chamberlain damage herbivore Villamil other consuming al. including for 2006; ways, decisions of variety (Ness allocation a Heil in visitation these 1980; plants (O’Dowd of host bodies cost to costs food the argument plant-provisioned impose despite the may symbionts mutualists support Ant ant sites processes. Transition metabolism, for biological invaded resources recently as our the prioritize well from ant-plants of results that (as understanding The broadly. our rates more to mutualisms photosynthetic plant contribute 2012). in Helliker also & results decline Wiley Our a by reviewed to defense; contribute chemical and and regeneration, budget carbohydrate 2017; Zvereva tree’s & Kozlov (e.g., systems Sharpe other production, many in plants for and Wilson 2002) Young & Ward metabolites, like (Rubanza undefended this defense plants minerals both in and for for both costs protein intense forage respiration crude quality and of high frequent amounts for Roques becomes high selective (e.g., paralleled thus more savannas clearly herbivory often are other are and others and herbivores but 2008) conditions, context-specific, (Veblen are dry system fixation In carbon herbivores systems. in vertebrate other hypotheses. decline starvation with observed in our carbon interactions invasive of study, broad system-specific aspects this for Some via generalize in fixation to demonstrated difficult As carbon resources. be affect may its strongly that exhausted can even also carbon insects invasive has in can how tree describes declines insects (2011) the large McDowell while before to Helliker shifts, mortality due climatic and increase to rises Wiley mortality Allen mortality plant and costs. in plant increases metabolic (2011) where initiate global in McDowell starvation, to increases shown both carbon strong been or of example, have fixation For deficits process Such systems. the plant. fixation, many discuss foundational a carbon (2012) in in limit declines “deficit” to herbivores carbon performance vertebrate a plant with to interact contribute that can turn suggests ant neighbors. in invasive study and conspecific an our how their demonstrates and though study trees neighbors, Our ant-defended plant to understory applies their other principle and between same conservancy spines) the Laikipia on (i.e. another both defense at seen inducible levels defenses deploy comparable associational to weak for plants evidence with removal found on plants potential browsing where reduced one have 2009), mimosae may but al. which clear, ants, et defensive not of (Barbosa ( are adjacent defense” Our typically neighbors. pattern were “associational well-defended areas to this of proximity by idea underlying protection gain the mechanisms defenses to The relates performance. of explanation because effect tree negative expected, a reduce be or would exclusion can herbivore effects of These effect habitats. positive uninvaded a in find not did we but habitats, and exclusion herbivore Both 05.Hwvr n-ln atesistpclyyedln-emntbnfisfrtepatb reducing by plant the for benefits net long-term yield typically partnerships ant-plant However, 2005). tal. et tal. et mot irgnit hs ytm hog -xto (Fox-Dobbs N-fixation through systems these into nitrogen imports ocpe lnsi pnaes n otpat ihnhrioeecoue.Coverdale exclosures. herbivore within plants host and areas, open in plants -occupied 08.Idcbersosst ebvr omtcajsmn,Freeland adjustment, (osmotic herbivory to responses Inducible 2018). 99.Orsuyad oti ieaue eosrtn htntv n soitsimpose associates ant native that demonstrating literature, this to adds study Our 1989). tal. et 96 pn rdcin on klo19)cncnuealreprino the of portion large a consume can 1998) Okello & Young production, spine 1986; ca. .megacephala P. tal. et )t well-defended to m) 5 .drepanolobium A. 00,o asn oa atain(Stanton castration floral causing or 2020), eoa oiieyipce otte hsooyi invaded in physiology tree host impacted positively removal tal. et tal. et 7 97 tno amr21) irpigpollinator disrupting 2011), Palmer & Stanton 1997; .drepanolobium A. tal. et rgoigls oig,Gadd foliage, lost (regrowing tal. et .drepanolobium A. 21)piaiyatiueosre contemporary observed attribute primarily (2010) 07.Crnchrioyipsscumulative imposes herbivory Chronic 2007). 01 Kos 2001; .mimosae C. .drepanolobium A. .mimosae C. tal. et re cuidb auecolonies mature by occupied trees atmdlmtaimadant- and model -ant tal. et togyrpl ebvrsthat herbivores repels strongly 02 Abraham 2012; tal. et rmvltesi unfenced in trees -removal 00.I otpat must plants host If 2010). tal. et ihlae containing leaves with tal. et 95 tannin/saponin 1985; .mimosae C. .drepanolobium A. Acacia 99 Gaume 1999; 01 producing 2001; tal. et pce that species tal. et .drepa- A. removal (2018) 2019), C. et ’s Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. rdhw .. eo,B,Blad . oz . let . orir A. Fournier, C., insects. Albert, invasive D., of Roiz, costs C., global National Bellard, underestimated Kruger B., the Leroy, in C.J., trees Bradshaw, marula on elephant African the 8. of impact ant The global (2002). governs O. Park. Jacobs, history & human R. Recent Biggs, (2017). L. Keller, 7. & A. Liebhold, dynamics. S., invasion Hawaii. Ollier, in C., disease interrelationships wilt Bertelsmeier, the mealybug on pineapple investigations and Field mealybug, pineapple (1982). 6. gray D. the Gerling, ant, & big-headed F. the McEwen, of T.H., Su, J.W., Beardsley, University. Clark complexities 5. ants. habitat and of effects distribution edge spatial interaction, ant-hemipteran the invasive on of Impacts (2020). O.E. Anastasio, forests. for risks change 4. climate M. emerging Vennetier, reveals N., management mortality McDowell, tree and ecology heat-induced D., Forest and Bachelet, drought H., of overview Chenchouni, global A.K., Macalady, C.D., of Allen, impact the forests. Predicting States United (2010). 3. eastern D.R. of Foster, dynamics & D.A. carbon Research Orwig, on adelgid A.M., woolly Ellison, P.R., hemlock Moorcroft, M., Albani, herbivores. savanna of strategies 2. Drought-response (2019). A.C. Staver, evolution and & Ecology G.P. Hempson, J.O., Abraham, JRG. and CR to 1. TMP, grant to Explorer 1556905) Young DEB Society (NSF Cited Geographic grant National grantLiterature Foundation RADS a Science Center PDM, substantial National International to a Florida provided grant and of ForestGeo Gituku) University PDM, Smithsonian a Benard a by and PDM, supported was to Mutisya Pejeta research Ol Samuel This and the Kipkoech, support. (particularly and Isaac logistical administration Buseinei, team Centre Research Gilbert management Mpala Ekadeli, assistance. Jackson Conservancy field Maiyo, excellent Nelly provided Mizell, Lemosiany Gabriella John work. this conduct savannas.to these in processes (Riginos Acknowledgements: ecosystem mortality connected the and species, increasing tree cycling By monodominant carbon productivity. this alter understory of in to performance shown linked the been decreasing mineralization be and has also content as may total (Riginos the trees system, plants (Glaser both the understory reduces soils into of species in productivity acacia inputs N of N and removal C where reducing Africa of further East term, of areas longer other the over savannas invaded H by forest ot fia ora fWllf eerh2-ot eae pnaccess open delayed Research-24-month Wildlife of Journal African South 0 119-133. 40, , ¨ ogberg tal. et aueEooy&Evolution & Ecology Nature ,7047-7056. 9, , etakteKna oenet(AOT//847/49 o hi permission their for (NACOSTI/P/18/4376/9459) government Kenyan the thank We 20) oetal opudn hseet lpat a euete oe within cover tree reduce may elephants effect, this compounding Potentially (2001). tal. et 5,660-684. 259, , 01.Finally, 2001). tal. et auecommunications Nature 09,adtu h abnfiainpromneo invaded of performance fixation carbon the thus and 2009), ccadrepanolobium Acacia ,0184. 1, , 8 .megacephala P. a est-eedn ffcso the on effects density-dependent has ,1-8. 7, , tal. et aainJunlo Forest of Journal Canadian nainmyfundamentally may invasion 21) asv e grossly yet Massive (2016). 2 13-22. 32, , tal. et tal. et 05 and 2015) 21) A (2010). Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. il,B,Mshiz . og ..&Hli,AJ 18) tde faSuhEs sa ant-plant Asian East South a of Studies (1989). A.J. borneensis. Helbig, Crematogaster by 21. & trees T.Y. Macaranga Pong, of protection U., association: Maschwitz, B., Fiala, 20. photosynthesis. and conductance Stomatal (1982). physiology T.D. Sharkey, & G.D. Farquhar, mea- conservation and biodiversity insect In: 19. aquatic Polynesia. native French to and Australia. threats Hawai’i in in species Invasive sures ants (2008). and R.A. butterflies Englund, lycaenid between Associations 18. (1999). Ecology A.M. of Fraser, Journal in Australian & species R. non-native Eastwood, between interactions Ant-plant-hemipteran 17. (2019). C.E. In: Tarnita, Kenya. 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Chamberlain, Hydrological Centre. and Research Meteorological Mpala Centre Kenya. Research 10. Laikipia, Mpala 2018-2020, (2020). data Weather D.J. Summary Martins, & Dataset. J. Gitonga, K.K., Caylor, 9. 3 317-345. 33, , Ecoscience 6 88-101. 16, , 3 207-221. 63, , clg n vltoayBiology Evolutionary and Ecology Ecology 2 316-329. 12, , 4 503-537. 24, , 0 2384-2392. 90, , netCnevto n Islands and Conservation Insect 9 rneo University. Princeton . Oecologia Ecology pigr p 221-234. pp. Springer, . niomna entomology Environmental 9 1724-1736. 99, , 9 463-470. 79, , nulrve fplant of review Annual Global tal. et Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. ln-dn .&Wl,S 22) peuaino htsnhssi iea urto-ecettomato nutrition-deficient mineral in photosynthesis of Upregulation 33. ratio. (2020). source-to-sink reduced S. by Wolf, plants & tree N. co-occurring lands. Glanz-Idan, degraded five of of restoration Adaptation in implication 32. (2006). its R. and stress Mitlohner, water & Management to M. species Fetene, shrub D., and Teketay, A., Gebrekirstos, characterize Ethiopia. to northern relations water of plant drylands of 31. use the The in (2005). sites R. and Mitloehner, species & M. tree para- Haile, B., castration Muys, of K., Gebrehiwot, case novel a and conflicts Ant–plant 30. (2005). ecological myrmecophyte. R.M. on a Borges, in herbivores & sitism M. insect Zacharias, alien L., of Gaume, effects indirect America. and North eastern Direct 29. of forests (2010). in interactions D.A. and Herms, processes & K.J. enemy ant- Gandhi, natural destructive diverse a a from ecosystem from tropical 28. support a with Saving mutualism (2013). urbicola) S. Pulvinaria assemblage. Noort, megacephala, Van (Pheidole & scale M.J. 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S., carbon in Neuvonen, group) J., rufa Kilpelainen, (Formica ecosystem. ants T., wood Domisch, of M.F., role Jurgensen, L., Finer, Ecosystems ilgclInvasions Biological 2,259-267. 229, , 6 196-208. 16, , vltoayEooyResearch Ecology Evolutionary 5 2115-2125. 15, , 3 189-193. 13, , ln inlBehav Signal Plant 10 Oikos ,435-452. 7, , 2 515-521. 92, , 5 1712543. 15, , ora fAi Environments Arid of Journal ilgclInvasions Biological mrcnJunlo Botany of Journal American Ecology 2 389-405. 12, , tal. et oetEooyand Ecology Forest 1 1296-1307. 91, , 0 581-592. 60, , 21) The (2013). Nature , Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. omn,BD ar ..(08.A nainrvstd h fia i-eddat(hioemega- (Pheidole ant big-headed African the revisited: invasion 45. An Australia. (2008). northern C.L. in Parr, cephala) & B.D. Hoffmann, plant–environment a Iso/anisohydry: trait. 44. (2018). hydraulic H. simple Cochard, a & than N.M. rather Holbrook, interaction biodi- F.E., of Rockwell, U., drivers Hochberg, ants fire ants-are Invasive (2013). 43. A.G. Hart, & F.S. loss. Gilbert, versity R.B., Rosengaus, C. J.K., Hill, Scoffoni, L.R., Fletcher, M.K., dehydration. 42. leaf Bartlett, to R., responses constrains Pan, trade-off G.P., safety-efficiency John, C., Macaranga Henry, in production body Food partners. ant 41. (1997). symbiotic P. via Menke, defence anti-herbivore & in G. investment Ecology Zotz, plant a K.E., Linsenmair, (Euphorbiaceae): triloba B., Fiala, M., Heil, Africa. East G. over Leng, patterns T., 40. drought Gebremicael, on X., change Liu, climate S.M., of Hosseini-Moghari, impacts (Enaphalodes Q., Tang, borer G.G., oak Haile, Dendrochronological red (2008). with S. infested Billings, L.(Fagaceae)) 39. & rubra S. (Quercus Leavitt, Cerambycidae)). oak V., (Haldeman)(Coleoptera: red Salisbury, rufulus northern M., Fierke, of F., parameters Stephen, L., Haavik, 38. species. Acacia African some of 121-140. characteristics ring , Growth (1995). I.D. in Gourlay, Accumulation End-Product Sugars. by Hexose Photosynthesis and 37. of Sucrose, Regulation Starch, Storing (1992). Plants S.C. of Leaves Huber, & E.E. host Goldschmidt, two in fluorescence chlorophyll and 36. pigments photosynthetic of content species. B. the G´orska-Drabik,Lagowska, plant E., on I., infestation Kot, insect Kmie´c, K., scale K., Rubinowska, K., Golan, change landscape 35. megaherbivore-driven stabilize plant-ants savanna. Defensive African an (2010). T.M. in Palmer, & J.R. Goheen, mineralization Tanzania. nitrogen Northern and Carbon of 34. soils (2001). savanna W. Zech, natural & and D. cultivated Solomon, M., in Fuhrboter, J., Lehmann, B., Glaser, 847-861. , clgclEntomology Ecological rhoo-ln Interactions -Plant urn Biology Current ilgclinvasions Biological 8 539-539. 38, , 0 1768-1772. 20, , ,55-65. 9, , rnsi ln Science Plant in Trends oetEooyadManagement and Ecology Forest 0 1171-1181. 10, , 11 ln Physiology Plant ilg n etlt fsoils of fertility and Biology at’ Future Earth’s auecommunications Nature 3 112-120. 23, , 9 1443-1448. 99, , tal. et ,e2020EF001502. 8, , ora ftoia ecology tropical of Journal tal. et tal. et 5,1501-1509. 255, , 21) stomatal A (2019). 21) matof Impact (2015). 22) Projected (2020). 3 301-309. 33, , 0 1-9. 10, , ora of Journal Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. ig ..&Cyo,KK 21) ebvrsadmtaitcat neatt oiyte photosynthesis. tree modify to interact ants 57. mutualistic and Herbivores phytologist (2010). New K.K. The Caylor, & E.G. King, 56. Plants Niinemets, & T.F. subsp. Keenan, tortilis Acacia of photosynthesis the on 55. stress stage. drought seedling of young Effect (2015). the F. at Aid, raddiana & S. Lutts, S., Kebbas, species tree major four in turnover 54. India. and of production region root fine semi-arid litterfall, the Leaf of (2010). K.P. Mohapatra, & P. Jha, angiosperms. drought-tolerant 32 across hydraulics and 53. gain carbon leaf D.L. to Lauenstein, related N., Oecologia traits Koike, stem S., and Matsuki, leaf K., ween Yazaki, T., Dryobalanops Nakano, rain- of A., trees of Ishida, canopy Effects mature (2017). in T. forest. adjustment Nakashizuka, rain 52. osmotic tropical & and Malaysian T.o. traits a Kumagai, in exchange A., (Dipterocarpaceae) gas Yoneyama, aromatica leaf T., on Kenzo, exclusion T., fall semiarid Ichie, and Y., arid Inoue, to concepts nonequilibrium of relevance 51. the On (1999). T.G. systems. O’Connor, grazing & A.W. Illius, of defenses indirect induced of 50. Relaxation herbivores. (2004). mammalian T.M. of Palmer, exclusion & following T.P. Young, acacias R., Karban, M., Huntzinger, removal in variation myrmecochore: 49. neotropical a of distance. dispersal dispersal Seed and (1986). rates D.W. Schemske, & C.C. Horvitz, knowledge. S. current Lavorel, P., of 48. 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Data may be preliminary. amr ..(04.Wr fatiin ooysz eemnscmeiieotoe nagido African of guild a in outcomes competitive 81. determines size colony attrition: ants. of Global acacia Wars (2016). (2004). T.M. M.B. Palmer, Thomas, & S.P. Worner, P.J., 80. Barro, De species. invasive D.C., from agriculture Cook, to implications. A.W., threat ecological Sheppard, D.R., pyramidale: Paini, Ochroma tree, neotropical a 79. of bodies Botany of Pearl Journal American (1980). D.J. O’Dowd, pollinators. deter also bodyguards 78. plant aggressive most the costs: indirect mutualists. mutualism’s Oikos ant A (2006). prospective J.H. on Ness, ants invasive of effects 77. The (2004). J.L. Bronstein, Invasions & J. Ness, down-regulation Citrus. Photosynthesis in 76. (2011). limitation R.-V. sink Molina, under & accumulation J.L. carbohydrate Guardiola, precedes B., Renau-Morata, S.G., Nebauer, primary gross on index 75. T.M. area Japan. 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Sack, C., Scoffoni, richness. ant native nectary-bearing reduced extrafloral and 92. ant of invasive dominance an Elevated of abundances Distributions (2009). increased and K.D. with Whitney, associated & is plants J.A. Rudgers, A.M., Tanzania.Savage, north-western protein, to of content indigenous The species 91. Acacia (2007). selected T. Fujihara, of & leaves nutrition T. of animal Ichinohe, minerals S.S., and Bakengesa, fibre M.N., Shem, C.D., Rubanza, African an in 90. dependence. encroachment density shrub and of rainfall Dynamics herbivory, 268-280. fire, 38, (2001). of A.R. influences Watkinson, relative & savanna: T.G. O’Connor, K.G., Roques, ant- of Disruption 89. chinensis). (Pachycondyla (2012). ant N.J. needle Sanders, Asian & invasive 557-565. R.R. the 14, by Dunn, mutualisms B., dispersal Guenard, seed K.L., Stuble, M.A., forests. Rodriguez-Cabal, subalpine ants in wood emissions red CO2 of and contribution 88. The pools N (2005). D.S. and Page-Dumroese, C & soil M. to Schutz, M.F., Jurgensen, A.C., ant–plant Risch, protective trees. a savanna of to Disruption damage (2015). elephant 87. T.M. increases Palmer, ant invasive & of an D.I. by effects Rubenstein, mutualism M.A., landscape-scale Karande, versus C., Local Riginos, (2009). T.P. Young, 86. & D.J. grasses. ant-plant Augustine, on keystone trees J.B., a savanna Grace, strengthens C., partner Riginos, third scale: of Economy 85. (2018). T.M. Palmer, mutualism. & K.M. Prior, ecology. mutualism 84. with dynamics carbon plant Integrating 71-75. (2016). E.G. Pringle, C. invasions. O’connell, microbe 83. C., and animal, Simmonds, plant, J., alien Wightman, of environment J., threats environmental Janecka, and S., the nomic McNair, of woodlands D., tortilis Pimentel, Acacia the upon fire and 82. giraffe elephant, of impacts Serengeti. The (1983). R.A.P. 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Data may be preliminary. i o infiatydffrfrivddaduivddtesdrn h r esn eut fpiws com- pairwise of Results season. dry the P during (* significant trees uninvaded as and indicated invaded are for parisons differ significantly not did decline interannual the ( than trees larger Transition significantly for was consistent difference was ( this ( trees and 2018 Control 2017 to in for invasion) physiological than before leaf invasion) (immediately these after yet 2017 months 2017, from ( in differ than invasion) not 2018 after did in higher trees indicating was Transition terms range A for interactions potential seasons, variables significant water wet with by In leaf protected panel, and change. lower were each interannual significantly (that from in distinct sites reported invasion Control of are paired effect tests in an Effect and surveys) study). season the dry throughout and wet initial after 2017 form) of draft range ant this native in 1. suppress figures Figure invicta below Solenopsis included (also (2017). Legends Y.J. Figure solenopsis. Xu, Phenacoccus & mealybug, invasive Y.Y. the Lu, from Zoology L., exploitation mutual Zeng, excluding G.W., by for Liang, area A.M., leaf Zhou, plant whole and the on traits conditions leaf precipitation 111. of different Adjustments under pseudoacacia (2019). Robinia L. Plateau. in Wu, Loess demand & and X. supply water Zhao, balancing M., Huang, Z., Zhang, in coexistence species acacias: swollen-thorn 110. on Ants (1996). system. L.A. simple Isbell, a & spines C.H. Stubblefield, herbivores: T.P., Young, of exclusion after defense induced 109. an of Relaxation drepanolobium. (1998). Acacia B.D. on Okello, & T.P. Young, perspective. whole-tree C.S. a Thornber, species: 108. A.F., forest foundational Sommi, a C.M., on Rigsby, 1783-1791. herbivores 99, M.L., invasive of Hickin, impacts R.N., Chronic Schaeffer, carbon for C.M., support Wilson, greater lends trees in storage 107. carbon of re-evaluation (Hy- A growth. megacephala (2012). to B. Pheidole limitation Helliker, ant, & E. big-headed Wiley, African the of 106. spread Worldwide Formicidae). menoptera: (2012). J.K. Wetterer, 105. ae re nwtcniin;()ivddteshv higher have trees invaded (B) conditions; wet in trees vaded by occupied of Trees SEM) +- (means cupied photosynthesis capacity) photosynthetic canopy 2. Figure ccadrepanolobium Acacia 49. , ccadrepanolobium Acacia efpooytei ae( rate photosynthetic Leaf ieecsi ef ( leaf- in Differences giutrladFrs Meteorology Forest and Agricultural A Oecologia .megacephala P. B , C .As,i r seasons, dry in Also, ). e Phytologist New , E ymclgclNews Myrmecological Oecologia .I r esn,tesi rniinstshdlower had sites Transition in trees seasons, dry In ). 0,98-107. 109, , dlsi e n r esn tln-emIvddadUivddsts (A) sites. Uninvaded and Invaded long-term at seasons dry and wet in adults re nTasto ie ta eeivddby invaded were (that sites Transition in trees A okr aesgicnl lower significantly have workers max-leaf D 1,508-513. 115, , A ,adla ae oeta ag a ossetfraltes( trees all for consistent was range potential water leaf and ), 9,285-289. 195, , max-leaf < .5 *P ** 0.05, .. e-ntla-ra n aoylvl( canopy-level and per-unit-leaf-area) i.e., ; E leaf 7 51-62. 17, , ,la rnprto ae( rate transpiration leaf ), nrae ewe 07ad21 o oto re but trees Control for 2018 and 2017 between increased 7,107733. 279, , 17 < .0,**P *** 0.001, A max-leaf A max-leaf .megacephala P. uigdycniin,but conditions, dry during max-leaf < .01 rntsgicn (NS). significant not or 0.0001) E and leaf n E and .megacephala P. A A ) max-leaf max-canopy n efwtrpotential water leaf and leaf vs. - nCnrlteswere trees Control in aitnJunlof Journal Pakistan A max-canopy .mimosae- C. n21 ( 2018 in tal. et ca hnd unin- do than A nDecember in .mimosae C. max-canopy months 6 . Ecology F (2018). ca i.e., ; ). 9 . oc- , Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. naieatpeec assasalbtsgicn euto nla ae oeta ag.I h dry the In ( in loss range. water Figures output): potential transpirational full increased water of for leaf combination text in ( a main reduction range by effect potential significant to of water but refer leaf results affect small right; Significant a at ( causes values seasons. season, presence P dry ant left, and invasive at ) wet (factors in ( subfigures seasons, sites in wet Invaded reported long-term are 3 tests at conducted herbivores) ( conditions for dry SEM) in +- variable response 4. either affect Figure not does presence ant invasive and dlsi x ulfcoileprmn peec/bec fat,lrehrioe)cnutda long- ( leaf- in 3 in declines the at at decline conducted estimated causes are herbivores) indices herbivory large Photosynthetic ants, seasons. dry of ( and canopy-level (presence/absence wet experiment in sites full-factorial Invaded term 2x2 a in adults 3. Figure B A ieecsi htsnhss(en -SM of SEM) +- (means photosynthesis in Differences etbaehrioyand herbivory vertebrate ) ieecsi aoytasiaincpct ( capacity transpiration canopy in Differences max-canopy ccadrepanolobium Acacia A A max-canopy etbaehrioyand herbivory vertebrate ) while ; .Rslso ffc et r eotdi aes nwtsaos ( seasons, wet In panels. in reported are tests effect of Results ). A max-leaf A max-leaf D dlsi x ulfcoileprmn peec/bec fat,large ants, of (presence/absence experiment full-factorial 2x2 a in adults .Tedunlcag nla ae oeta ( potential water leaf in change diurnal The ). n ( and .megacephala P. and C .megacephala P. A max-canopy etbaehrioyadivsv n rsnebt cause both presence ant invasive and herbivory vertebrate ) 18 rsnebt as elnsin declines cause both presence r oe uigdysaos etbaeherbivores vertebrate seasons, dry during lower are A E .megacephala P. rsnebt as elnsin declines cause both presence canopy and B n yrui resistance. hydraulic and ) n efwtrptnilrne(means range potential water leaf and ) -occupied C and B ccadrepanolobium Acacia and E D canopy slkl driven likely is ) D E A A max-leaf canopy ). vertebrate ) u onot do but n ( and and ) C Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. iue1. Figure 19 Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. iue2. Figure 20 Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. iue3. Figure 21 Posted on Authorea 11 Sep 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159986366.66136129 — This a preprint and has not been peer reviewed. Data may be preliminary. iue4. Figure 22