Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. fisppltos opoiea nih notevgtto epne ops ocncdsubne,a disturbances, volcanic past to responses vegetation the fragmentation into the insight increased have an might provide volcanism To that suggested was populations. the it its as Yet active such of disturbances. been ecosystems have to unique Andes adapted variability. southern support Hotspot. well of in Rainforest and Volcanoes Temperate range far ka, Valdivian occurred process. historical 10 the vegetation ecological its today last of in understanding the and imprint our during dynamics significant limiting with vegetation past, eruptions the in and in infrequent role are events important large an However, play eruptions Volcanic Aims Abstract forest Araucaria in disturbance Volcanic title Running the at Patagonia. status. disturbances ern conservation volcanic current past its to current affected responses of the have Vegetation at would stability distribution eruptions relative geographical volcanic past The that its Conclusion evidence in change no no found eruption. suggests I vegetation large tephrafall thus that several the showed ecotone, after analysis after area forest-steppe study The fast the landscape. relatively in the Nothofagus pollen recovering in Araucaria and changes, distributed Araucaria permanent legacies from Nothofagus percentage biological facilitating without sparse pollen perhaps resisted of of stage, increase indicative successional Slight be early years. could dominated ˜500 obliqua-type Ephedra forest after the record. condition changes. severely original significant pollen Lake affecting recovering caused conditions the around regeneration environmental seldom by vegetation the but suggested the change tephrafall as buried might small ka) tephrafall grassland, (˜3 to 39 and Sollipulli-Alpehue sensitive kyr, of was 9 eruption vegetation current large last The the Pollen the the ka. However, if Methods During 4-2 and Results Patagonia. between dynamics frequently northern record. vegetation more in sedimentary the Relem, ecotone a on forest-steppe from have disturbance Araucaria analysis volcanic Location might tephra of disturbances, volcanism volcanism. and volcanic role by that past the caused to suggested assess been responses unique an was has vegetation to is support the fragmentation conducted it into Araucaria and was Yet insight un- an study ka, Hotspot. our provide disturbances. paleoecological To 10 Rainforest limiting a to populations. last Temperate its past, adapted Valdivian the of well the the fragmentation during the and of in active increased fragmented part far been strongly as have occurred forest species, large Andes vegetation However, Araucaria-Nothofagus endangered southern variability. the today in of as in Volcanoes range such imprint historical ecosystems process. significant its ecological with and of dynamics eruptions derstanding vegetation and in infrequent role are important events an play eruptions Volcanic Aims Abstract 2021 22, July 1 Moreno-Gonzalez Ricardo Patagonia northern Araucaria in the ecotone at forest-steppe disturbances araucana volcanic past to responses Vegetation nvriyo G¨ottingen of University 1 Araucaria 1 ruai araucana Araucaria sa nagrdseis togyfamne and fragmented strongly species, endangered an is Araucaria-Nothofagus oetsep ctn nnorth- in ecotone forest-steppe oeta atof part as forest Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. nadsatae,btteautctx hne u oercmn fntins nietoeaes after areas, those Unlike nutrients. an of taxa America, enrichment pollen North to terrestrial due western in al., changed changes Mazana, et taxa significant (Egan aquatic vegetation trigger Mt. the regional not and the but did local area, in eruptions on distant impacts Also that a the concluded infer in authors 2013). to lake The al., veg- small to climate the a 2016). et contribute past disturbed in (Paine affect conducted volcanoes to was that could Europe aimed experiment evidences that western and weak volcanoes in or eruption Icelandic contradictory etation of of showed large-eruptions impact studies for palynological past conditions, example, illustrate 2018). For to Crisafulli, ecology. responses. & attempted disturbance vegetation (Swanson have and past disturbances studies sedimentary the volcanic in in Paleoecological past sub-fossil far active pollen understand occurred and to last since layers had help tephra responses challenging or of could the is study span, records, biogeographical resources, the human-life dynamic many diverse through in requires studies, vegetation highly monitored monitoring paleoecological the Well-designed in long-term be of the to especially areas, long study 2002) remote designing too the in al., in However, located help et of are may Franklin patterns volcanism. volcanoes patterns and 2005; diversity by the process of al., influenced of develop example, et areas effects the development In (Dale lasting or vegetation the recovery management of Understanding resistance, rate nature 1998). the the 1999). arrange Franklin, like and Grishin, and responses patterns & & type vegetation regeneration the Moral Knight, drove while (del 2018), areas Foster, disturbance Antos, disturbed & after 2018; in (Zobel legacies tephrafall Dale, by biological mechanism buried & of disturbance being Volcanism Crisafulli the after 1998). on rapidly (e.g., al., growth depending eruptions vegetation et vegetation magnitude (Turner volcanic the ecosystem on Large the an processes of and dynamic 2018). history and the Dale, patterns large in complex and & events creates sudden stochastic (Crisafulli trigger and vegetation can episodic impacts the infrequent, the are on and particularly Earth, on changes, agents environmental disturbance important are eruptions Volcanic Introduction volcanic hotspot that Rainforest evidence Temperate Valdivian no found status. I conservation Keywords thus current ecotone, its forest-steppe affected current have would the eruptions at distribution geographical past changes, of stability permanent relative without The resisted vegetation that showed eruption. Conclusion large analysis the The after fast relatively landscape. recovering the in distributed the However, from the changes. percentage record. severely significant pollen affecting caused the 4-2 conditions by environmental between seldom facilitating suggested the frequently but as change more grassland, might tephrafall Relem, ka) and small Lake forest (˜3 around Sollipulli-Alpehue to vegetation of sensitive eruption the was large buried vegetation tephrafall The 39 kyr, ka. 9 last the During record. sedimentary Results a from analysis tephra and Pollen Methods Patagonia. northern in ecotone dynamics forest-steppe vegetation Araucaria the volcanism. on by disturbance caused volcanic been Location of has role fragmentation the current assess the if to and conducted was study paleoecological Nothofagus itrac eie eeainrslec,ln-emvgtto yais ocncecology, volcanic dynamics, vegetation long-term resilience, vegetation regime, disturbance : Araucaria eeeainrcvrn rgnlcniinatr˜0 er.Sih nraeo pollen of increase Slight years. ˜500 after condition original recovering regeneration Araucaria and ohfgsobliqua Nothofagus olni h td raatrsvrltprfl ugssn hnei its in change no suggests tephrafall several after area study the in pollen tp ol eidctv fsas ilgcllegacies biological sparse of indicative be could -type 2 Ephedra oiae al ucsinlsae perhaps stage, successional early dominated Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. Fgr ) npriua,vgtto itr a eosrce rmasdmn oeotie rmLake from obtained core sediment a from reconstructed was history vegetation particular, In current the 1). in (Figure located is area study The increase should Methods fragmentation resistant forest more Then, tephrafall. be thick would to vegetation than of where main tephrafall. tephrafall magnitude thickness the thick the thin tephra is after that to the therefore predict Tephrafall faster with I recover vegetation. related assumption, and/or the be this buried Following should area. that responses study tephrafall the vegetation the by on around mostly influence record type volcanic affected disturbance pollen the is volcanic published into it a insight Andes, separate of the a use of have made to I re-analysed end, and Araucaria this 2021) To volcanic al., status. et the conservation (Moreno-Gonzalez reconstruct its to affecting if 1) ecotone evaluate aims to forest-steppe 3) work and this disturbance regime, the volcanic disturbance in volcanic history the disturbance Despite and is 2001). dynamics the Wagstaff, hypothesis in vegetation & 1991) far the of (Kershaw so (Burns, Yet conducted ecosystems been disturbance 2016). dominated have angiosperm volcanic structions al., an resist et in Veblen to persist Araucaria’ 2002; adaptation to al., helped morphological might et recognized that Bekessy the (e.g., with resist dynamics contradictory to regeneration of capability variability the and di- or volcanic genetic bution recovery of the of role rate influenced the the might about Patagonia, the known area. northern in is the in particularly little in dynamics disturbance, vegetation, remaining vegetation unique legacies the the that biological on of and of and sturbance blast-zone characteristics types gradient, direct volcanic elevation the the the and with Despite in strategy associated that life-traits patches is on different out establishment depend created happened pointed vegetation regeneration those that (2019) early the to areas the al. similar specific gradient et as impacted disturbance Moreno-Gonzalez disturbance type the Furthermore, of disturbance types landscape. Swanson each Chait´en different volcano. the of where the in 2008-eruption of Helens, the responses St. early is Mount the case after described Another by buried (2013) 2015). was al. vegetation al., et steppe et as the such (Ghermandi 2011, correspond species would disappear in geophytes cohorts PCC phytes several from rhizomatous 2011 Eastward and and in tephra, 2016). tephra cm (PCC) al., cm ˜50 et Complex by (Montiel buried Volcanic eruptions as zones past Caulle in such to del resprouting species species Puyehue-Cordon principal pioneer the landscape the keep was the of in to eruption forest the responsible age uneven after be location sustain would tree-line or the and 1977), volcanism eruptions, al., species Moreover, the et Veblen of Andean 2004; result 2012). this Veblen, As en- in (Kitzberger, & high 2016). functions (Daniels a al., and depressed by et structure be Veblen featured the affecting can et (e.g., ecosystems, volcanoes (Fontijn underneath vegetation forest these region surrounding Nazca-plate by unique Andes and influenced the supports the forest been Andes of in have located the kyr also subduction is of 10 which The section areas last demism, active southernmost the most The 2004). in the (Stern, 2014). large-eruptions of several al., Pacific one triggered the but explored. has Earth, of further American-plate the south be part in to south-eastern of dispersed needs the are role patterns eruption Holocene in diversity expected volcanic the the the that during and disturbance in Despite mechanism volcanoes conditions, influence the of Active impacted. original their about degree strongly its known and variable were is dynamics to layer a little vegetation recover tephra suffered hotspots past not biodiversity thin drove vegetation could the global by the forest several covered destroyed vent in the crater eruption volcanoes the areas the this many of from that In far east indicate sites km 1996). evidences 170 McGlone, pollen & to Zealand, (Wilmshurst up New and nearby, BP), forest yrs (˜1850 eruption Taupo oetdnmc.Det h itnet h ocncsuc ftefrs-tpeeooeeastward ecotone forest-steppe the of source volcanic the to distance the to Due dynamics. forest oi-clgcliprac n t urn osrainsau,fwln-emvgtto recon- vegetation long-term few status, conservation current its and importance socio-ecological s ruai araucana Araucaria Araucaria ruai araucana Araucaria sdmnn Vbe n stn17;Brs 91.Teeiecsshow evidences The 1991). Burns, 1978; Ashton and (Veblen dominant as oetsep ctn,2 oass h eeainrsosst past to responses vegetation the assess to 2) ecotone, forest-steppe Araucaria Araucaria Araucaria Araucaria oetsep ctn rud39 around ecotone forest-steppe oet twshpteie htps ocnceruptions volcanic past that hypothesized was It forest. itiuinwsiflecdb ocns ttecurrent the at volcanism by influenced was distribution 3 Poa oetsep ctn.T otiuet h knowledge the to contribute To ecotone. forest-steppe u loaetdngtvl h ouaindistri- population the negatively affected also but , p.and spp. Rumexacetosella ° ,i otenPatagonia northern in S, nrae hl thero- while increased ohfgspumilio Nothofagus Nothofagus < 5 Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. vr10 er,adte a oeldwt mohsln ehd(pr07.Bt aibe,frequency events layer variables, of tephra Both sum the a (spar=0.7). as below method calculated smooth-spline value was events with the volcanic modelled from of therefore was records frequency sedimentary then normal, The describe in and quiet cases. to (1993), years, by some is aim Birks 1000 tephra in we and over tephra each only so, Lotter multiple of 0 of doing event to of record In the corresponded the occurrence of centimetres. Unlike magnitude in the eruptions. the thickness Patagonian the describe tephra of to the magnitude preferred to the the we corresponding manuscript, ash-value a of ( this the equation magnitude as In giving robust the modelled but 0. of and simple of a measure variable deposition is a and explanatory model as -0.5, quantitative This tephrafall considered to a 1993). -since is Birks as event and where thickness coded (Lotter disturbance ), tephra process was independent the decay tephra and exponential and layer each single simple time- tephra Furthermore, a short every as a of layers impact. to use tephra made limited these & we is of (Bennett regime, each coefficient disturbance considered dissimilarity 50 we volcanic a 5. of the interval as of reconstruct time complexity distances To regular with Euclidean based a method the was at smooth-spline with point samples a then starting pollen 1995). through and Humphry, The the fitted interpolated smooth-spline, was we data. curve with (RoC), the compositional years Rate-of-Change and the the samples, to estimate centralized. the To (PC) and of transformed Curve age root Principal the square on a Using was fitted percentage The we 1992). (PCA). Furthermore, analysis 400 Line, component of & principal count through pollen (Birks assess terrestrial E(T To tool minimum ratio the powerful the 1996). at (E(T a obtain (Bennett richness grains to palynological is distances the summed pollen Euclidean analysis estimated were in rarefaction we as based analysis, constrained individual expressed samples rarefaction stratigraphically changes, a and pollen conducted diversity sample the we per pollen for Later, taxa (CONISS) percentage. terrestrial analysis pollen as from cluster expressed data abundance, pollen relative the only of used isopach the we within Here cal located 2951 is to Relem back Lake dated analysis eruption, the is Data So-A it (hereafter the Patagonia, eruption to northern 2016). Sollipulli-Alpehue at relation in al., The In erupted past et (Fontijn recent have 1993). 2014). m the or al., al., 2 in volcano) et et largest (Naranjo Villarrica (Fontijn the BP years (e.g., of yr are k years magnitude one which 10 was 10 considerable SVZ, active eruption) last of every the been So-A eruptions the erupted to have regular have during far that more so once volcanoes volcanoes had recorded least 7 other has eruptions are while volcano all there Lonquimay [?]3) list 33 Relem The (2014) between (VEI Lake al. 1. extended around Table subduction et km is Fontijn the in 100 1). and of summarized of result (Figure plate, radius a Holocene American is a the SVZ South In during The continental 2014). (SVZ). the al., Andes et beneath the (Gilbert of plate Zone Nazca Volcanic Southern the the of in lies area study The history eruptive and analysis, 1). setting sediment (Figure Volcanic Giesecke, the fragmentation Moreno-Gonzalez, of and strong description (2020) a Moreno-Gonzalez full in a showing found (2021). as 2006) be Fontana well can al., and chronology as et the description, and (Gonzalez vegetation analysis and a.s.l. pollen climate m on 1700 details at Further when plants tree-line conditions and Climatic patterns oceanic the 2013). vegetation with reaching 1998) al., complex discharging Roig, et generate temperate effect 2012; (Mundo disturbances is rain-shadow Andes (Kitzberger, natural the sharp associations region and of a topography the section create local western in with winds the interacting climate westerly on precipitation with The the arriving outflow. of masses most or air inflow Pacific river The without influence. respectively), ha, ˜1 (38 Relem x ° 83”S 71 S; 58’39” stevlefrteah(rirrl e o10b h authors), the by 100 to set (arbitrarily ash the for value the is t 10 stesml et =ie.As,teatosabtaiyasge au eoetephra before value a assigned arbitrarily authors the Also, (=time). depth sample the is /E(T )/ 400 ) h olndvriyetmtda E(T at estimated diversity pollen the )), 400 ° ’1 ;16 ...,Lk ee sasalwadsallk ˜. depth, m (˜2.5 lake small and shallow a is Relem Lake a.s.l.), m 1268 W; 4’51” Mtha ta. 05.Cmoiinltedo h ersra aawsexplored was taxa terrestrial the of trend Compositional 2015). al., et (Matthias ) Araucaria . 4 oetocr anyaoe10 lvto,normally elevation, m 1000 above mainly occurs forest 10 ,adple vneswscluae as calculated was evenness pollen and ), α stedcycecetequal coefficient decay the is > . mtik Here thick. cm 0.5 ° o46 to S ξ ° -ατ S Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. owtsadn nyfwsgicn hne Fgr c.Tems eeatcagsocre ttetime the at occurred changes relevant most The disturbances, volcanic 4c). to in (Figure composition variation changes vegetation The significant the iterations. of few 21 sensitivity only after some cm fitted notwithstanding indicates Curve grains units cm Principal gradient 100-500 The grains distance between the 2000 disturbances. PAR other record, than vegetation in or the more of Along processes decrease indicator decayed sedimentary an PAR taxa. small as pollen eruption interpreted dominant caused both likely be So-A both of normally could was after PAR any of events vegetation where affected abundance Volcanic 4b), have when (Figure to the ka, variable seem biomass. is depressed 4.5 not together eruption do taxa before So-A all So-A notorious of after Only PAR events more few is grasses. the it while other taxa, but and other thickness, Poaceae and by tephra line) ( dominated to red Poaceae (vertical proportional in eruption not So-A (increase) is the decrease with a vegetation caused the of indicators tephra several unknown shows 4 figure The that suggesting eruptions. magnitude the volcanic following influ- analysis, with significant multivariate abundance related no analysis the somewhat had further In tephra-layers while However, frequency, small significant composition. volcanic other too. a pollen the vegetation vol- had that the the variables, indicate in eruptions explanatory on eruption ence volcanic So-A The influence The the significant data. after second a 3a). the samples had of The excluding (Figure magnitude, 20% So-A and explaining diagram. of frequency 3b) ordination response (Figure canic the the composition vegetation to of on related side influence where mostly left trend is overall the axis the on second split located and variance 1, of the (e.g., zone dance dominance of the taxa 11% in steppe explains of from abundant component dominance shift more long-term the the are explains with as component diagram others, forest interpreted first ordination be a The would unconstrained to change. and the vegetation Poaceae) variance, in gradual total Likewise, the the represent of 2021). trends 78% al., compositional forest the et the to (Poaceae) 3a) (Moreno-Gonzalez encompasses grassland (Figure fire It steppe than from (2020). transition rather vegetation Moreno-Gonzalez regime overall in an by published characterized was is ( it history and vegetation kyr lake. the 9 the last of into tephra description up of full ka was cm A 1.5 1.5 frequency events last than volcanic volcanic the more For ka to deposited ka. 4 responses none 5 Vegetation Before but and that recorded, 7 ka. are indicates around layers, 4-1.8 tephrafall disturbance frequency few without between tephra disturbance present, periods years the the with The 1000 all to years, every events/1000 Considering 2c). 6 occurred eruption than (Figure 2b). the had lower frequent decay, (Figure to eruptions exponential relatively regarding threshold 7 an effect is the around as lowest above regime modelled a tephra-value, peaks disturbance indicates The some but volcanic cm. thickness, missing 216 tephra eruption while So-A recorded the the magnitude core regards are to the 18 corresponding patterns layer in only well-defined similar tephra the which layers but The shows in 2a) one, tephra 2a). (Figure biggest content 39 Figure thick the in layer registers cm (crosses is core tephra 0.5-7.5 peaks sediment significant between the as the are considered from them total, be of reconstructed In Most were 2). thick. regime (Figure cm Relem disturbance Lake its of and sediment history volcanic The regime volcanic the of Reconstruction Results al., et (Oksanen 2015). 2.4-2 multivariate vegan-package (Juggins, statistical 2016), a 0.9-15 Team, All in Rioja (RStudio samples regime. and 3.3.1 pollen disturbance 2017) RStudio volcanic the through to constrain conducted to responses were variable vegetation analyses explanatory assessing an Analysis) as (Redundant used analysis later were magnitude, and ohfgsdombeyi Nothofagus Ephedra < nteple eodrs fe h oAeuto Fgr ,Apni ) hrfr the therefore 1), Appendix 3, (Figure eruption So-A the after rose record pollen the in 0c hc vria rylns.Tprfl htmre itrac vn,normally event, disturbance a marked that Tephrafall lines). grey (vertical thick cm 10 tp)ta curd˜.k Apni ) ieya euto hnei precipitation in change a of result as likely 1), (Appendix ˜4.5ka occurred that -type) .dombeyi N. .dombeyi N. 5 tp)ple ecnae(iue4) h decrease The 4a). (Figure percentage pollen -type) tp.Poaceae, -type. Araucaria -2 yr -1 te rp nPRcnb eae to related be can PAR in drops Other . hw ekngtv eainwith relation negative weak a shows Mulinum Ephedra > . mtik htcould what thick, cm 1.5 n yeaee among Cyperaceae, and sdmnn.Teabun- The dominant. is Araucaria nrae its increased -2 yr -1 while , > 0.5 Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. namlena-cl trtre ls oisoiia tt fe prxmtl 0 er Fgrs4ad5). and 4 (Figures years 500 approximately after state original its conditions climatic to stable close relatively returned to analysis it due rate-of-change but millennial-scale the composition, with in domi- a vegetation peak vegetation short-after in in a the imprint started register strong to revegetation not evidenced impact to So-A, after eruption some enough the revegetation After fast left 2001) recovering of 4c). tephrafall al., but et rate (Figure Small 4b), Lamy (Figure 4). the 2003; compo- grasses al., (Figure and by et vegetation grasses (Jenny nated species the by low colonizer was dominated determining precipitation potential was by when ka vegetation influence responses 4.5 turn environmental before example, in the For which disturbance. in structure, role northern and a in sition play records may palynological condition the in Climatic increase of an none show knowledge, of on not my increase increase do to slight significant eruptions. sites and, a a other 2017) depict show Unlike, Giesecke, So-A, 1988), Patagonia bottom. & the that al., core Fontana of et eruption the 2020; zone (Heusser So-A at al., isopach layer Arco the m tephra del of –perhaps Mall´ın thick Paso 2 result presence instance, a local as the For very increase within conditions. a following record certain of the a indicative under and be expansion species- only its other of should facilitate PAR with abundance and co-existing is pollen percentage Relem or Lake The pollen suppressed to in close increase 1964). but such 2001). (Maher, from 32.4%, but southward al., of area km et maxima source ˜200 with (Paez around the 2.7%, 20% studied averages been to it has up pollen-samples it founded and modern disturbances In after Relem. species Lake abundant Patagonia. most the in not significance is palaeoecological its and on ecology its around about occurring patterns those since contextualize to eruptive attempt one Ephedra I and from extension impact paragraph, spatial (e.g., the following and legacies of time the biological in Relem. type In and vary Lake conditions, can 1986), vegetation factors climatic another. Zobel, operating the as & to those such (Antos that Moreover, event factors traits point 2005). plants of al., 2015) 1999), result et Grishin, al., Dale a and et is Moral recovery (Crisafulli (del of ecology topography rate volcanic the in and advances resistance likely Recent presented succession, a findings ecological the the factors. indicate new in model, interacting dynamic not a expected complex this initiate do more to a that Contrary PAR, showed volcanism here species. as overall by shadow-intolerant an and well caused suggests pioneer diversity as model by and dominated successional evenness, biomass traditional The plant and in biomass. richness and decrease biodiversity pollen decreasing the of pattern episodes, unique disturbance volcanic After was Discussion taxa small dominant other the after As occurred normally areas. that study. pattern sheltered this a in in 4e), eruptions remained (Figure volcanic with increased individual destroyed, evenness survival completely palynological not and depressed, was vegetation legacies disturbance, Perhaps biological volcanic of increased. some richness lasting expansion to richness are palynological the palynological long responses eruption richness eruption, with have In So-A So-A palynological coinciding after the not values in 4e). before did smaller particular, Changes (Figure In the deposition disturbance events. reached decrease. volcanic tephra or the increase the to small after can On richness related rapidly (Figure after palynological always So-A recover changes time. for not could the small except and and events, along the variable vegetation volcanic stable that to the relatively sensitive indicates on was not effects result were composition change The pollen of that rate 4d). the suggests by also depicted curve peaks contrary, the but So-A, of Nothofagus Nothofagus a h nxetddmnn aadrn h al ucsinlsaeatrteS- eruption So-A the after stage successional early the during taxa dominant unexpected the was and pce and species Austrocedrus Araucaria fe r itrac eg,Rffee&Vbe,19) nta study, that In 1998). Veblen, & Raffaele (e.g., disturbance fire after r osdrdpoerte ntergo Fgr ) itei known is Little 5). (Figure region the in tree pioneer considered are Ephedra’ Ephedra Araucaria olngan ol ipresvrlklmte a from far kilometres several disperse could grains pollen s fe neuto o aebe eotddrn recent during reported been have nor eruption an after 6 oetsep ctn,wa a ecnrle ymany by controlled be may what ecotone, forest-steppe Ephedra .dombeyi N. Ephedra eoeS- rpinwsawy low, always was eruption So-A before Ephedra hltfrs eeoe ae.This later. developed forest whilst tp nteae;btinterestingly but area; the in -type Ephedra ih aeanreyeffect nursery a have might Ephedra rud3k,above ka, 3 around Dcsnet (Dickson Ephedra Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. ra erytecreteooehv ensihl nraigfrtels 4k ept fteiflec of influence the of in despite ka abundance ˜4 pollen last that the showed Dickson for (2020) 2020) pattern, increasing al., al., generalised slightly et a been et Nanavati have not (Dickson and after ecotone Although Cilantro (2021) abundance current eruption. relative al. Lake the So-A pollen nearby et from after in areas Moreno-Gonzalez PAR record changes (2020), affected in neither pollen volcanism al. changes Relem, recent that et Lake nor as unlikely in A layers source is observed tephra volcanic (2002). it those small from ka), al. as distant 9 patterns et fragmentation last similar Bekessy the the shows increasing by for from populations area suggested abundance the the it pollen of in in distribution eruption change the soil largest significant negatively to (the found for change eruption to last So-A chemical unlikely not the the therefore If might in growth 2012), population. deposited Since tree-ring moderate al., whole records. tephrafall decreasing et resist sedimentary the as (Tognetti small to such to time that harms species long damage possible physiological a physical the is deposition, significant allow tephra it cause after leafs Hence, occurred not 1995). smooth in did al., and years crown et branches trees 500 Veblen flexible 1991; after bark, (Burns, area thick disturbance the the into that expanded documented that and of indicated pioneer abundance analyses of as Multivariate pollen decay behave the the not following So-A, stages did of successional species large-eruption advanced the Nothofagus after that However, shows cm. 20 than dombeyi- of smaller abundance the tephrafall records by palynological In 2019). al., none events. Chait´en et eruption Volcano from (Moreno-Gonzalez recent dombeyi eruption eruption the the last N. after the After year of 1982). one blast-zone found (Veblen, the particularly disturbances in tephrafall, However, volcanic 2017). small after by species Puyehue, affected pioneer of less considered be Chait´en are volcano, the to establishment that vegetation of expected early eruption the are in last Trees fleshy- role the some important in an of 2019). playing and al., pollen found 1996); et the also (Moreno-Gonzalez were McGlone where fleshy-fruits eruption, and with Taupo (Wilmshurst species after the found suggested were was taxa Probably dispersal 2015). fruits plant al., of et mechanism (Loera rodents similar and birds through principally Ephedra mechanism, dispersal temperate and of in fleshy-fruit the expansion eruptions its following of past the elevation repeated Furthermore, after 2015). low Chait´en Remarkably, Volcano,al., 2019). to the al., close in et areas canopy (Moreno-Gonzalez the zones forest mid-elevation the and the low suggesting blast-zone under in thus the grows occurred to tephrafall, normally same Close by The that forest. 1996). buried system, McGlone area & root te- (Wilmshurst wide by rhizome eruption a the disturbance fern of in buried the impacts of records to with the resisting spore several response after Pliscoff, in the in spread & plants BP), increasing species (Luebert of (˜1850 found conditions mechanism were eruption weather common cold Taupo system, a and the be arid After to resist the phrafall. to seem while canopy. capacity growing cm, undisturbed the of ˜15 Roots the and by establishment 2006). under shoots, buried early shrubs sprouting being the and soil, resist eruption, trees poor not So-A tolerant could the shadow performan- herbs the to species that by the Regarding found determine dominated may (1986) was and Zobel establishment deposition & early tephra Antos resist to example, For role important ce. an Patagonia, also play in traits significantly. augmented Plant disturb precipitation not after did when condition or original Later, of deposition its 1996). development tephra to McGlone, returned the vegetation & promoted the (Wilmshurst it Zealand, years New 120-250 in eruption about Taupo after comparison, In lae-ar ta.(00 eosrtdta eldvlpdfrs a o ffce ntelong-term the in affected not was forest developed well that demonstrated (2020) al. et Alvarez-Barra ´ a h anfo upyfrbrsadrdnsi ieae eattdatrS- rpin A eruption. So-A after devastated area wide a in rodents and birds for supply food main the was yewssrnl ffce ti td n iko ta.22) ned h aapeetdhere presented data the Indeed, 2020). al. et Dickson and study (this affected strongly was type tp sawy aibe n e tde aebe sesddrcl h epne ovolcanic to responses the directly assessed been have studies few and variable, always is -type ohfgspumilio Nothofagus ohsraquadripinnata Lophosoria Araucaria Lophosoria Nothofagus pce eefudrgoigi ra uidb eha(ote tal., et (Montiel tephra by buried areas in regrowing found were species Lophosoria Araucaria i o niaesgicn hnei olnpretg n A after PAR and percentage pollen in change significant indicate not did fe h 08euto fCate ocn.Teseis lowith also species, The Chait´en volcano. of 2008-eruption the after a n ftefis pce erwn n eratrteeruption the after year one regrowing species first the of one was pce nteae,tevgtto a essniiet small to sensitive less was vegetation the area, the in species epne oiieysotatrteeutos(Henr´ıquez et eruptions the after short positively responded a o ffce yayo h ml ehaal ti largely is It tephrafall. small the of any by affected not was Ephedra Ephedra 7 nteae Fgr ,PaeI)cnb eae to related be can II) Phase 5, (Figure area the in Ephedra Fgr ,paeIII). phase 5, (Figure Pteridium, a ieydet pca dpainto adaptation special to due likely was Araucaria enseiswt hzm root rhizome with species fern a Nothofagus and Nothofagus edigwere seedling species N. Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. otj,K,Lcoyz .M,Rwo,H,Pl,D . ahr .A,Nrno .A,&Mrn-o,H. Moreno-roa, & A., J. Naranjo, A., Argentina. T. and Mather, Chile M., southern D. of at , Pyle, tephrostratigraphy Holocene Quaternary H., the dy- Late Rawson, during M., (2014). millennial-scale change S. vegetation and Lachowycz, of K., Argentina. Centennial patterns Fontijn, Patagonia, and northern Processes in (2020). (2017). ecotone T. forest-steppe I. Giesecke, the P. & L., Moreno, S. Fontana, & L., T. https://doi.org/10.1111/jbi.14017 Hall, M., in (Ed.), Walker namics R. Fletcher, L. In in B., recovery. Treeline ecosystem Dickson, Altitudinal and disturbances on Volcanic Ground Climate (1999). Disturbed Y. of of S. 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Reference macro-charcoal northern and core, Pollen sediment (2020): Relem L Lake Sonia of Fontana, Thomas; Giesecke, Ricardo; the Moreno-Gonzalez, in used hotspot. past be Rainforest of Temperate can statement Valdivian role that accessibility the the and Data in forest understand species Araucaria to threatened in done patterns certain be diversity of current to conservation determining need in studies eruptions palaeoecological volcanic Further eruptions. volcanic several ´ 89 08.https://doi.org/10.1016/j.quascirev.2014.02.007 70–84. , Vegetatio 19 Araucaria , Springer. . 64 h eeeaindnmc fAacraaraucana Araucaria of dynamics regeneration The Ecology 23,1311 https://doi.org/10.2307/20037246 103–111. (2/3), – Nothofagus , 85 1te. .88.Elsevier. 868). p. ed., (1st 5,18–26 https://doi.org/10.1890/03-0092 1284–1296. (5), p.2726.Srne.https://doi.org/https://doi.org/10.1007/0-387- Springer. 277–286). (pp. oet ntesuhr Andes. southern the in forests clgclrsossa on t ees:rvstd3 er after years 35 revisited : Helens St. Mount at responses Ecological ° S). eiwo aaooayadPalynology and Palaeobotany of Review h Holocene The h nylpdao Volcanoes of Encyclopedia The 8 08:1512.doi:10.1177/0959683620913920. 30(8):1115-1128. Pages-OSM h Holocene The ora fBiogeography of Journal nvriyo Colorado. of University . 234. , utraySineReviews Science Quaternary , clgclRsosst the to Responses Ecological ac 2018 March , , 2 85 1,11.htt- 1–10. (1), 34,263–287. (3–4), Heredity 1265–1284. , jbi.14017. , , 88 , Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. ibr,D,Fent . utrl,S,&Bret .(04.Ps-lca iesre fexplosive of series central time south Post-glacial volcano, Lonquimay of (2014). system C. plumbing Burkert, magma & the in S., the changes on Kutterolf, Chile. deposition associated A., ash and volcanic Freundt, eruptions of D., Effects Gilbert, (2015). steppes. F. Patagonian Northwestern Oddi, https://doi.org/10.1016/j.jaridenv.2015.06.020 in & vegetation 154–160. J., gap Franzese, of recovery S., early example. Gonzalez, an devel- as L., structural forests and Ghermandi, Douglas-fir Disturbances using B., (2002). D. implications, J. Lindenmayer, Management Chen, silvicultural R., and & with D. Ecology K., Berg, ecosystems Forest Bible, A., forest C., D. natural D. Thornburgh, Shaw, of B., S., opment A. W. Carey, Van, Keeton, R. E., Pelt, M. A., Harmon, T. Spies, F., J. Franklin, nNwWrdEhda ihrrtso ih vlto eae odseslability. 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Rate top Sollipulli-Alpehue Accumulation the the Pollen by at caused and taxa other curve) main indicate (dotted the of abundance lines patterns vertical Successional grey 5. Figure eruption; Sollipulli-Alpehue Palynological d) the indicates time. deposited Principal the line tephra b) along Relem. vertical changes Lake changes. Red compositional biomass from significant disturbances. vegetation record showing for change sediment proxy of E(T the a rate richness in the as registered c) constrained. used and b) Pollen-Accumulation-Rate eruptions Curve and taxa volcanic unconstrained pollen to a) All smooth responses Relem, a) by Vegetation Lake obtained for 4. was composition Figure frequency pollen of of diagrams Trend Ordination years. 3. =0.7. years. 1000 Figure 1000 spar every every with frequency Relem. frequency line) of Eruption Lake (blue c) the sum function into eruption. spline the after deposited indicate decay layers points exponential tephra Grey the of as of thick modelled threshold the thickness arbitrary Tephra Indicates shows a) line horizontal regime. the Grey eruption Relem, Volcanic Lake 2. of of Figure position distribution the the America representing (2020) and South area Relem, Google southern study Lake source, in to the map Holocene closes of the volcanoes Map during of b) triangles) location (red 2013). volcanoes Program, active Volcanism showing (Global Figure a) 1 Figure tables and figures of List Tec- (2016). 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Turner, imhrt .M,&MGoe .S 19) oetdsubnei h eta North central the in disturbance eruption. and Forest Resilience, Inertia, Taupo Tephra: Volcanic by Eruption BP Buried 1980 Understory the Forest (1996). In after 1850 (2018). Redevelopment. Community A. S. of J. Pattern the Antos, the & M. B., following D. Zobel, McGlone, Zealand, & https://doi.org/10.1177/095968369600600402 New M., Island, J. Wilmshurst, https://doi.org/10.1080/0028825X.2015.1130726 400 , 9 )ade ayooia vnesE(T evenness palynological e) and )) > 1,1.https://doi.org/10.2307/2844727 11. (1), .c hc notelake. the into thick 1.5cm p.1315.Srne e ok https://doi.org/10.1007/978-1-4939-7451-1 York. New Springer 113–125). (pp. Ecosystems clgclRsossa on t ees eiie 5years 35 Revisited Helens: St. Mount at Responses Ecological > . mt eemn eeateutos(e rse) b) crosses). (red eruptions relevant determine to cm 1.5 10 11 )/E(T , e eln ora fBotany of Journal Zealand New 1 clg fteSuhr Conifers Southern the of Ecology 6,511–523. (6), 400 ora fBiogeography of Journal oprn iest hne fe volcanic after changes diversity comparing ) ruai araucana Araucaria Holocene , rdplgn) Base polygons). (red 6 , 4 , 4,399–411. (4), 54 , 3,275–294. (3), JANUARY 223–246. , ora of Journal 6 , Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. rypit niaetesmo rqec vr 00yas rn ffeunywsotie ysmooth by obtained was frequency of Trend years. =0.7. years. 1000 1000 spar every every with frequency Relem. frequency line) of Eruption Lake (blue c) the sum function into eruption. spline the after deposited indicate decay layers points exponential tephra Grey the of as of thick modelled threshold the thickness arbitrary Tephra Indicates shows a) line horizontal regime. Grey eruption Volcanic 2. Figure of distribution the (2020) and Relem, Google Lake source, to map closes volcanoes of location > . mt eemn eeateutos(e rse) b) crosses). (red eruptions relevant determine to cm 1.5 12 ruai araucana Araucaria rdplgn) Base polygons). (red Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. iue3 riaindarm fple opsto o aeRlm )ucntandadb constrained. b) and unconstrained a) Relem, Lake for composition pollen of diagrams Ordination 3. Figure 13 Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. itracs e etclln niae h olpliApheeuto;ge etcllnsidct other indicate lines vertical grey eruption; Sollipulli-Alpehue Palynological d) the indicates time. deposited Principal the line tephra b) along Relem. vertical changes Lake changes. Red compositional biomass from significant disturbances. vegetation record showing for change sediment proxy of E(T the a rate richness in the as registered c) used and Pollen-Accumulation-Rate eruptions Curve taxa volcanic pollen to All responses a) Vegetation 4. Figure 400 )ade ayooia vnesE(T evenness palynological e) and )) > .c hc notelake. the into thick 1.5cm 10 14 )/E(T 400 oprn iest hne fe volcanic after changes diversity comparing ) Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. eid ftm r o xc n r sdol ihshmtcpurpose. that schematic Note with conditions. only original used of are and recovery and green. colonization and exact in area), reorganization not (yellowish represented are II III, is Phase time Phase Pre-eruption of vegetation. area. phases. the periods disturbed successional of the “collapse” indicate into the top indicates expansion the curve) area) (continuous at (reddish Rate rela- boxes I Accumulation pollen Small Phase Pollen shows the y-axis taxa. shows Primary y-axis selected Secondary eruption. of and Sollipulli-Alpehue curve) the (dotted by abundance caused tive patterns Successional 5. Figure 15 Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. al .Lcto fptnilsuc ocne ls oLk ee,isfeuny(ute eal o each for details (further frequency its Relem, 2014) Lake al., to et Fontijn close in volcanoes found source are potential eruption of Location 1. Table ocn aeDsac n ieto rmLk ee oa rpin at1 aEutosVI[?]3 VEI Eruptions 326 ka 326 km; 10 km; 88 last 80 eruptions Total Relem Lonquimay* Lake from direction and Tolhuaca Distance name Volcano ° ° N 218 1 22 1 NNW NNW 16 Posted on Authorea 22 Jul 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.162696461.14671023/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. pedx1 hr olndarmo eetdtx.Fl olndarmi ie nMrn-ozlz(2020). Moreno-Gonzalez in years given 500 is last diagram the pollen in Full occurred taxa. them selected of of diagram Most pollen 2014) uncertain. Short al., are et 1. dates dacite), (Gilbert Appendix the (mainly [?]3 eruptions tephras VEI of the to composition of equivalent the Some tephra, but ** of (2014), km3 al. 0.01 et Fontijn about by ejected provided probably not is VEI Lonquimay *In ai 1k;205 km; 81 236 km; 223 89 km; [?]3 80 270 VEI km; Eruptions 299 Lanin 38 km; 64 ka Quetropillan 10 last eruptions Total Villarica Relem Sollipulli Lake from direction and Llaima Distance name Volcano ° ° ° ° ° S 4 1 2 10 3 5 3 150** 2 55 WSS WS WS W WWN 17