Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. t Gugerli Felix Coupland George Germany Heidelberg, University, Koch A. Marcus Albani C. Italy. Maria Rome, (CNR-IAS), Council Research National ment, Andrello Germany (Main), Marco Frankfurt Centre, Research Climate and Biodiversity W¨otzelStefan alpina distantly Arabis more or species REVIEW Brassicaceae TECHNICAL other INVITED in adaptation. developed environmental and be history might life research We perennial progress future its recent model. which families. plant to discuss plant a and related from related as system, are questions alpina mating that open A. its traits of of indicate developmental emergence environments. diversification We of rapid the harsh dissection the has including in molecular facilitated alpina, alpina genomics the that A. Arabis ecological in properties of and stimulate perennial the history genome evolution of arctic-alpine to evolutionary overview life-history complete the the reference an study summarize a family, provide the to we Once Brassicaceae as thaliana review, the A. thaliana this level. Within to In A. molecular complementary biology. established the model different further at a with and become processes species and accelerated biological other model markedly numerous renowned in has most elucidate studies the research to is available, system thaliana was study Arabidopsis sequence plants, as non-crop used characteristics, In life-history been their considerations. of has practical combination advantageous also an and to due properties status genetic prominent a obtained have organisms model Many Abstract 2021 1, March 6 5 4 3 2 1 Gugerli Felix W¨otzelStefan genomics ecological for evolution plant life-history model and perennial a alpina: Arabis ws eea nttt o oetSo n adcp Research Landscape and Snow Forest for Research Institute Breeding Federal Plant Swiss for Institute Heidelberg Planck (COS) Max Studies Organismal of Centre K¨olnUniversit¨at zu Council Research National Italian Main am Goethe-Universit¨at Frankfurt hs uhr aecnrbtdeulyt h aucitadsol ecniee sfis authors first as considered be should and manuscript the to equally contributed have authors these eena oe ln o clgclgnmc n iehsoyevolution life-history and genomics ecological for plant model perennial a : 6 S ws eea eerhIsiue imndr,Switzerland Birmensdorf, Institute, Research Federal Swiss WSL t 1 ac Andrello Marco , nttt fEooy vlto n iest,Geh nvriyFakutadSenckenberg and Frankfurt University Goethe Diversity, and Evolution Ecology, of Institute t nttt o ln cecs nvriyo oon,Clge Germany Cologne, Cologne, of University Sciences, Plant for Institute idvriyadPatSseais etefrOgnsa tde CS,Heidelberg (COS), Studies Organismal for Centre Systematics, Plant and Biodiversity nttt o h td fAtrpcIpcsadSsanblt ntemrn environ- marine the in Sustainability and Impacts Anthropic of study the for Institute et fPatDv ilg,MIfrPatBedn eerh oon,Germany Cologne, Research, Breeding Plant for MPI Biology, Dev. Plant of Dept. 2 ai Albani Maria , 1 3 acsKoch Marcus , 4 ereCoupland George , 5 and , Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. n drs eefnto npyoeeial lsl eae pce htvr o ri fitrs.The interest. of trait a for vary species. related that distantly species even in related on particularly studies closely and phylogenetically with Here, in gained variation. function knowledge environmental gene species, as to address related such relation and closely questions in systemic of change more biology study a address indicate developmental where the can might and or studies underpinnings, today available, mechanistic evolution broadly on are trait more knowledge methods sufficient studied analytical with are powerful species above For increasingly mentioned sciences. plant species in the paradigm of of 2014). most al., that et phylogeography fact (Hay The 2005), dispersal seed Witham, and & development leaf Berardo, compound Dart, 2017), related (Mable, distantly Bomblies, shifts more & (H¨am¨al¨a Similarly, Yant system 2019). 2019; mating Savolainen, J al., on & (Schmickl, et studies (Kol´aˇr (Monnahan 2016). for genomics al., for hybridization system ecological et attention with evolutionary model for associated perennial recently a polyploidy more the into 2019). and for Kudoh, within developed model & been from Honjo a in has as species (reviewed and adaptation sister 2008), and ecology using al., population et addressed (Hanikenne initially tolerance Koenig were heavy-metal 2013; topics of al., (Pyh¨aj¨arvi 2021). specific number et fields Mattila, comparability, genus the Kane many & Consequently, of in 2020; research interest. reasons al., future For biological et benefit (Cesarino of will continuously areas which increased 2015), all Weigel, has variation cover & species distribution, can ease model wide and model plant transformation of cycle, dedicated facile no life tractability, understanding However, In short genetic the cultivation. size, 2010). model: genome in plant of Grossniklaus, small self-compatibility, successful advances M¨uller & traits, advantages, a tremendous in life-history of practical in to e.g. characteristics with key (reviewed led many organism biology has of combines single of This establishment a disciplines generated. the sciences, on many plant be on focusing in can depends principles By knowledge forms fundamental platforms. of life study wealth of common a diversity as overwhelming species the model understanding and Describing INTRODUCTION | 1 adap- families. local plant associations, perennial related gene–environment distantly tation, experiments, research more functional future Brassicaceae, or its which environment, species to from arctic-alpine Brassicaceae related questions other are open that in indicate traits We developed developmental adaptation. be of environmental might dissection and molecular history the of life in emergence perennial progress rapid recent the of discuss facilitated history and that evolutionary system, properties the the summarize of We overview model. an plant provide markedly we review, has this research In available, was perennial to arctic-alpine sequence complementary the family, genome Brassicaceae the complete Within biology. a different established Once further level. and plants, molecular accelerated life- non-crop the their In of at considerations. combination processes advantageous practical an also to and due thaliana properties status prominent genetic a characteristics, obtained history have organisms model Many Abstract head [email protected] Email: Switzerland. Birmensdorf, CH–8903 Z¨urcherstrasse 111, stitute, Correspondence: Arabidopsis stems eondmdladhsbe sda td ytmt lcdt ueosbiological numerous elucidate to system study as used been has and model renowned most the is ø gne,Bytn,&Kc,21) dpain(eie l,21)adpplto genomics population and 2013) al., et (Kemi adaptation 2010), Koch, & Brysting, rgensen, rbdpi thaliana Arabidopsis clgclgnmc of genomics Ecological : .thaliana A. . rbdpi halleri Arabidopsis ei uel,Boiest n osrainBooy S ws eea eerhIn- Research Federal Swiss WSL Biology, Conservation and Biodiversity Gugerli, Felix .thaliana A. osuylf-itr vlto n clgclgnmc nhrhenvironments. harsh in genomics ecological and evolution life-history study to n t eaie sas xrml aubewe efrigcomparative performing when valuable extremely also is relatives its and a evda ieyue oe o eea eae Wie,21) It 2012). (Weigel, decades several for model used widely a as served has .thaliana A. .thaliana A. o xml,cnrbtdt nesadn h oeua ai of basis molecular the understanding to contributed example, for , rbsalpina Arabis evsa eeec oietf,freape eeorthologs gene example, for identify, to reference as serves sterfrnet tmlt tde nohrseiswith species other in studies stimulate to reference the as 2 .alpina A. admn hirsuta Cardamine nldn h iesfiaino t mating its of diversification the including , rbdpi arenosa Arabidopsis rbsalpina Arabis rbdpi lyrata Arabidopsis a enue ostudy to used been has a eoeamodel a become has rgnlyserved originally .alpina A. a attracted has Arabidopsis sa as Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. vnaogtetx urnl osdrda lsl eae,including revision. taxonomic related, further closely undergo as likely will considered and currently group taxa monophyletic a which the form for among not Even 2013), does 18 present Koch, at and & genus mya (Karl the 20 at species 2020). estimated 100 al., roughly approximately Despite et be A-A). German, can Figure German, Walden, tribe Walden, 1, 2020; Genus (Box the Koch, 2019; II of & al., lineage ages German, et evolutionary group (Huang, Nikolov Brassicaceae crown respectively 2020; to and mya, close al., group except tribe et stem lineages the uncertainty, Mabry placed all this 2019; that contradicts to emerging al., 2020) finding basal early al., et likely this the et Kiefer is However, of (e.g. that 2020). one et analyses Brassicaceae, Schranz, considered German, phylogenomic Mand´akov´a, in (Walden, & been Nguyen, described lineages has Lysak, (Walden, main lineages Arabideae Aethionemeae evolutionary all evidence, and in the cytogenetic genera, found among recent accepted is tribes 18 on complexes among Based distributed trait 2020). species and al., 545 traits roughly morphological with of assemblage convergence monophyletic a is It family. includes which Arabideae, Tribe PHYLOGEOGRAPHY AND SYSTEMATICS EVOLUTION, | 2 perspectives provide HERE finally complex 1 and of FIGURE it, dissection of the underpinnings research. and physiological future under evolution history and potential life system molecular on and perennial mating the its history adaptation, understanding and in of conditions systematic progress latest studies natural evolutionary the for include and we model Moreover, phylogeographic traits. a developmental as species’ use the its summarize discuss on We knowledge current developed. of in niche, recently overview both an ecological occurrence, provide broad plant we of Here, and extremes exemplifies distribution which the into plants, wide elevation. expanding individual its by and of Given species latitude lifespan flexibility. model the (Figure aforementioned developmental restrict birds, ravines the can or or a complements and creeks ungulates for transient along wild more need are communities cattle, in habitats moss-dominated the of they thrive in these deposition conditions of where also dung moist Many individuals debris, very to 1). However, rocky tolerate due substrate. can and accumulate unstable they often slopes of and the nutrients scree habitats between where range, calcareous shoots areas broad sheltered Ansell on elongating (cf. its by occur America to persist plants North Typically, can Corresponding of elevation. 1). parts and Figure northern amplitude cf. and Greenland 2011; southern al., Scandinavia, reference et into first extends and a Africa, of release to arctic-alpine The the similar and With possible, 2009). become have genus al., analyses Arabis the genomic et addressing in comparative Wang for development 2015), system (R. al., the model et a perenniality (Willing into assembly of developed genome has mechanisms species molecular the Consequently, biology. the molecular of toolkit by its the to manipulated contrast genetically in be is which equally Brassicaceae, the of self- state of ancestral breakdown differential 2015). alpina the al., Arabis that et et Russell, Tedder Grundmann, highlighted (Ehrich (Ansell, 2008; self-compatibility studies history Vogel, of experimental postglacial degrees & varying Schneider, and and with populations 2006), Pleistocene to About led al., species’ has Scandinavia. incompatibility et the and Koch Africa inferred from 2007; studies populations on al., phylogeographic compared Research later, who years. years (1962), fifteen Hedberg 50 by past studies the taxonomic during with evolution life-history and alpina Arabis Arabis eg ifre l,2017). al., et Kiefer (e.g. .(rsiaee,teApn okrs,hseegda oe pce o clgclgenetics ecological for species model a as emerged has Rockcress, Alpine the (Brassicaceae), L. sdpodwt aecrmsm ubrof number chromosome base a with diploid is hc om h oeo rb rbda,i elsuidpr-adplpyei e of set polyphyletic and para- well-studied a is Arabideae, tribe of core the forms which , rbsalpina Arabis Arabidopsis a iegorpi itiuini h uoenAp,San rba East Arabia, Spain, Alps, European the in distribution geographic wide a has .alpina A. Agrobacterium- tde aewdndt nld te pce rmtegenus the from species other include to widened have studies , soeo h otpoiettie ihnteBrassicaceae the within tribes prominent most the of one is , 3 .alpina A. eitdtasomto n,hne saeal to amenable is hence, and, transformation mediated n Arabidopsis .alpina A. 8 n t aytp eebe h putative the resembles karyotype its and =8, n nrdc h aiu ye fresources of types various the introduce and .alpina A. evsa h yeseis However, species. type the as serves eaie with relatives .caucasica A. .alpina A. tre ery6 er ago years 60 nearly started pnawd ecological wide a span n hti fe sdas used often is that 5 h pce can species The =5. .alpina A. Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. n uhoai ein ftegnm,eceigta fohrBascca pce Wlige l,2015). al., within et accumulate (Willing In of often 2000). burst but species Initiative, genome, transposition Genome hetero- Brassicaceae the Arabidopsis both other The across in (e.g. of distributed regions retrotransposons randomly that pericentromeric of not exceeding are accumulation genome, elements the transposable the to Usually, of relates largely regions size euchromatic a genome and of in difference establishment of This genome the 2015). the was Mbp, direction 475 APPLICATIONS With this AND in Corresponding GENOME step REFERENCE loci. | neutral major 3 at a processes and for selective resources, genome past mimic reference of genomic may high-quality imprints solid drift genomic the valuable because require genetic of a selection, as analyses knowledge of is such signatures Likewise, history inferring processes genetics. demographic when demographic essential ecological underlying is in the structure questions and genetic neutral hypothesis-driven structure selective Al- investigating genetic a glaciers. for spatial to European foundation the led northern on have the 2007). knowledge might to al., ability Detailed vast et Alps pro- colonization the (Ehrich the given authors for diversity of unlikely genomic selection The edge appears decreasing strong 2007). sweep scenario northern with al., this the migrations et However, from multiple (Ehrich Europe. ternatively, expanded diversity in that genetic refugium area of single periglacial a levels Mediterranean, from low the colonization very and pose show Pyrenees contrast, the by as such populations, climate, & milder Holderegger, persisted with Parisod, have the regions Graf, remote Alps, Rogivue, pattern 2007; more the differen- This al., In within of et 2007). possibly Ehrich mosaic al., and 2008; a et al., around Ehrich form 2018). et refugia 2009; Gugerli, (Ansell and different Tatras al., diversity from the et genetic and recolonizations (Alvarez overall Carpathians of multiple structure of populations from spatial levels lineage, result East–West habitats third high might an alpine this show with when Within Carpathians groups periods, 2011). the tiated glacial probably al., colder and Europe et to during Alps also (Ansell Migration Marmara elevations and the 2006). of lower events, al., Sea at et immigration the (Koch located multiple around Africa were Northwest lineage through region in third the populations populations a through the European Anatolia, occurred for Western northern source From Eh- and a 2006). (Assefa, central as al., lineage served all et African to Koch from East resulted 2007; the rise likely with Brochmann, gave 2011). which contact & al., Ethiopia, secondary Nemomissa, et into in Taberlet, the (Ansell came groups rich, lineage, clade that phylogeographic this independent two Within populations Anatolian formed an 2006). isolated lineage form al., the previously et southern ranges through Koch more Lebanon 2011; Plateau Mount second, al., A and Iranian et Anti-Taurus (Ansell the the mountains of and high for populations African Caucasus habitats East stepping-stone the the provided reach likely to tually system migrated mountain high-elevation lineage This first diagonal. a (Ansell Anatolia, local periods From undergoing glacial colder possibly and Anatolia, interglacial the in During warmer persisted plants. between al., centres alpine fluctuations marked for survival suitable was during local habitats migrations period of of elevational This expansion network transition. 2006) the fragmented and al., Pliocene–Pleistocene a 1992) et the Bartlein, Pleistocene, Koch at & 2011; (Webb ago, al., cooling years et rapid million that by (Ansell 2-2.7 lineages indicate about ancestral DNA diverged three that with nuclear established and was distribution chloroplast Present-day on studies knowledgeand Phylogeographic of as state serve taxonomic may Nevertheless, HERE and that 1 resolved. systematic characteristics BOX the be life-history 1. of to of Box account variety requires in detailed great given still uncertaintyis A a that systems. with uncertainty study species comparative encloses taxonomic group ample taxonomic is this there plant, ornamental 2011). nsitu in uiggailpros(hihe l,20) otenErpa n North-American and European Northern 2007). al., et (Ehrich periods glacial during Gypsy lmnshsldt h xaso fprcnrmrcrgosad consequently, and, regions pericentromeric of expansion the to led has elements .alpina A. .alpina A. sruhy35tmstesz fta of that of size the times 3.5 roughly is sdsrbdbelow. described as , 4 .alpina A. .alpina A. .thaliana A. neouinrl recent evolutionarily an , rgntdi Anatolia. in originated .alpina A. .alpina A. Wlige al., et (Willing .alpina A. oeven- to might in et Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. l 21)soe httecpct o efpliaincnee tesavnaei h pollinator-poor the in advantage fitness a conveyed self-pollination as Tor¨ang for such Scandinavia, et capacity selfing, in the experiment of field that consequences a showed negative when In However, (2017) reproduction advantage. migration. assure al. this might during confirmed counterbalance self-incompatibility might subsequently e.g. of depression, was restricted, loss inbreeding which are a Such Alps, mating 2011). the for al., reached chances et species (Tedder trials the km. pollination when from by 1 lost originating to was seeds up self-incompatibility distances collected refugia, over of events 84% found mating on (2012) effective with Based Gugerli indicated predominant & analyses Spain was Holderegger, paternity Graf, inbreeding and these Buehler, France but in- Nevertheless, Alps, Scandinavia, mating, selfing. translated Swiss (Tedder the from mixed orientation in production populations anther for scale pollen selfing and evidence local lower a largely distance On and between stigma–anther 2017). flowers success in al., (Tor¨ang et smaller po- differences pollination markedly selfing differential subtle e.g. and Moreover, to having outcrossing 2015). between ones al., e.g. morphology latter et ( flower the species in with within differences pulations, variation shown mating-system have approaches targeted have Comparative few e.g. also ( only level but interspecific but the selfing, 2019), at halleri done fully al., been outcrossing, et have studies Bachmann fully such spp.; now, in Until lineage. resulting evolutionary 2011) an within Mable, Hence, & Vogel, populations. mixed-mating Lao, is Ansell, which (Tedder, 2002), function (Barrett, selfing to increased 2008). Similar and (Mable, self-compatibility Brassicaceae adaptive to in for angio- transition also opportunities among observed repeated thus, pattern commonly the ubiquitous and, in- One is 1996) system Gl´emin, 2017). plants Godt, & mating Bataillon, sperm & the Hartfield, (Hamrick because 2006; diversity (Charlesworth, pivotal, genetic evolution is of evolution distribution the mating-system governing fluences processes the Understanding EVOLUTION 2019). SYSTEM genomic al., MATING et studying | (Dittberner by 4 efforts expanded conservation been assist eventually has and coverage comparisons phylogenetic recently, of Only evolution. properties of genome from sister study benefit annual to might the studies is Future and 2020). which of of individual tetraploidy al., species using example, et sister For assembly (Rellstab perennial resources. genome population structured reference Swiss phylogenetically fragmented a further, more of a population sequencing in-depth of and pooled-sample for efforts Lobr´eaux, 2013) opportunities sequencing & outstanding ample for represented offer (Melodelima resources 2019) (Laenen data genomic available range al., Additional publicly et species analyses. the the (Rogivue genomic across and Alps from Swiss populations, individuals western plant at 304 the natural accessed and and be 35 2018) can of resources al., re-sequencing selection. other et Subsequent and of 2017). genome signatures al., reference et underlie the (Jiao that of release polymorphisms depend recent causal which most identify experiments, The to mapping genetic groups & linkage future Gugerli, Rogivue, up complicate (Choudhury, might breaking that selection natural implies features on natural which genomic in in traits, Such role adaptive chromosomes 2019). key underlying Parisod, all a are play that along density dynamics genes for genome disequilibrium Transposon enriched transposon-mediated linkage were 1992). regions of those al., pe- sweeps; patterns et selective of with (Tanksley expansion correlated landscape hence, be alpina rates; recombination to recombination the shown meiotic alter was low of can by pericentromeres regions characterized short ricentromeric the be in can located regions are Pericentromeric genes few a only contrast, of By space gene the of half about ouain ihnteSisAp,adalrepooto ftelne lcssoe intrsof signatures showed blocks linked the of proportion large a and Alps, Swiss the within populations rffi il,2014). Willi, & Griffin ; F Arabidopsis IS .sagittata A. aus nele l 20)sgetdta,atrpsgailrclnzto rmsuhr Italian southern from recolonization postglacial after that, suggested (2008) al. et Ansell values, , .alpina A. and .alpina, A. .alpina A. .alpina A. .nemorensis, A. .alpina A. a prpyi eficmaiiiysse htprilyhsls its lost has partially that system self-incompatibility sporophytic a has n h uhsalrgnm of genome smaller much the and scnandi h eeohoai oprmn ftechromosome. the of compartment heterochromatic the in contained is ugssta te pce rmtieAaieecudb used be could Arabideae tribe from species other that suggests , ffr utbepafr o tdigmtn-ytmvariation mating-system studying for platform suitable a offers lutaighwgnmcrsucscnalwfrinterspecific for allow can resources genomic how illustrating .alpina A. 5 nld h hl-hools eoesequence genome whole-chloroplast the include .montbretiana A. .thaliana A. .nordmanniana A. Wlige l,2015). al., et (Willing www.arabis-alpina.org Kee ta. 2017), al., et (Kiefer hc sthe is which , Capsella A. A. Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. rgn ihpat rmhge lvtossoigls lsiiyfrtemaue ris ugsiganother suggesting traits, measured of the population for the on plasticity depended less plasticity showing phenotypic elevations of degree higher the from (2018), plants al. with et Villemereuil origin, seasons. de growing may of which longer study key the populations, and In high-elevation the temperatures from average was diffe- those grew higher phenotypic than temperature sites respective output revealed low-elevation Again, to reproductive from sites relate higher performance. originating had natural Plants populations reproductive and the patterns: vigorously six and observed to more vegetative from the close (de explaining to plants gardens Alps parameter related of French environmental common the traits comparison in within in ranges A scale in rentiation mountain 2018). regional also different a Till-Bottraud, and two on & 2012) detected in al., Gaggiotti, also et were Mouterde, (Manel adaptation Villemereuil, local species of plant signatures alpine Significant several in were 2017), precipitation animals al., and and plants et temperature in alpina (Siepielski Both summer differentiation North. globe adaptive of the intensity shaping the parameters in the across environmental than and key counterpart, Spain as minimum southern in identified for winter its pronounced previously than as in more such than was local Scandinavia differences, which for in drought, habitat higher lower large-scale was consistently towards which adaptation were temperature, suggested parameters detected This of fitness negligible. range was sites, were European adaptation transplantation the ( both of local provenances At edges foreign strong Scandinavian 2015). experiment, and al., Spanish transplantation et the reciprocal from populations long-distance gardens between analyses. classical common genomic a and landscape transplantations for In showcase reciprocal a approaches: become Experimental has | and 5.1 Consequently, adaptation adaptation. local on broad-scale local for studies to test for history to used exploited biogeographic short be of a can of contribution clines aspects and the These continentality. share temperature Mountains of in Carpathian degree mean variation to the the natural and annual located in Alps Alps Habitats differ low Central the differentiated. but the strongly a habitats, from Pyrenees, is alpine intensity, as the gradient light mountains, such latitudinal Cantabrian also the but features, a in length, typical on photoperiod share while located Pa- season, to Habitats growing Leempoel, scales. expected 2013; of spatial are Donohue, distribution over different Scandinavia & broad diversity at along The Kim habitat plants processes 2018). 2007; large alpine adaptive Joost, Hoffmann, in to & found & due Geiser, were Papst, adaptation differentiation risod, (Byars, of local gradients patterns on narrow and K¨orner, studies geographically 2011), & for (Scherrer settings distances ideal short involving has are studies ENVIRONMENTS latter environments ALPINE early Alpine The AND of adaptation. ARCTIC series TO local ADAPTATION a | for including 5 basis research, section. the evolutionary following form the in in may topic newly outlined variation Here, prime combination seed. genetic and a in pollen standing growth, become by alleles or asexual how beneficial mutations evaluate potentially including spreading arising help reproduction through will or pollinator-independent plasticity, eventually phenotypic by systems with persistence mating varying its reduced of increase a consequences suggesting alpina and Spanish populations, causes A. 2018). al., Greek the and et French on and (Laenen with Information Italian mixed-mating populations associated outcrossing mixed-mating of likely in obligate load bottleneck, self-compatibility the strong and of to cost a diversity similar and genome genetic were selfing using contrast, populations increased by By diversity from this: genetic recolonization. resulting substantiate reduced strongly post-glacial load not had genetic could populations increased (2018) Scandinavian that and al. purged demonstrated et been they This had Laenen data, distance. depression, However, resequencing anther–stigma inbreeding reduced populations. inducing and potentially Scandinavian anthers alleles, in introrse detrimental more that with imply flowers may for observation selection to leading tundra, ihnteErpa ls(..Mnl oct eede uel,&Hleegr 2010). Holderegger, & Gugerli, Legendre, Poncet, Manel, (e.g. Alps European the within a oewt niomna hne,eg u ociaewrig oa ouainmay population local A warming. climate to due e.g. changes, environmental with cope may .alpina A. sensu aek br,20) hl ieecsbtenrgoa elct populations replicate regional between differences while 2004), Ebert, & Kawecki oevr oa uvy a erpiae ots o ovreteouinor evolution convergent for test to replicated be can surveys local Moreover, . 6 .alpina A. ffr ayopruiist investigate to opportunities many offers .alpina A. a enextensively been has .alpina A. .alpina, A. (Tor¨ang A. as Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. nig eeucvrdwti h rnhadSisprso h uoenAp,i ean ob tested of be range to distribution remains broad it the Alps, of European advantage the full local of taking and parts regional elsewhere, Swiss discussed exist the and are patterns of French precipitation, most similar the While and if within preferences. habitat temperature uncovered broad were as but findings specific, such has drivers, that in plant environmental differentiation pioneer common adaptive that with appears associated it summary, In 2020; Nowak, & 2020). environment Brochmann, al., Arctic Brysting, et Gustafsson, the (Birkeland, Rellstab to species-specific adaptation remained in may genes involved adaptive assumption genes phylogeneticallymost general the including in this pathways compared species though or studies different even Two genes in above). 2013), same (see (Stern, (K¨orner, the 2003; true niches targets species hold ecological selection plant not similar that many occupy possible for that hence relevant species is are distant It 2012). factors al., environmental populations et similar Manel among environment, alpine shared defence an largely and In terms. elements stress response to transposable temperature adaptation identified to local also in for corresponding involved SNPs in and study candidate genes flowering this identified of Interestingly, of (2019) orthologs al. response. regulation included et the Rogivue these resolution, and and (Lobr´eaux,spatial cover 2020), response snow Miquel, stress and Lobr´eaux & in precipitation repre- abiotic 2014; in performance temperature, reduced Melodelima, with vegetative performance on & associated in growth based regions Manel, patterns with genomic scan observed 19 associated genome the Additional gene A 2018). explain candidate data. could sequencing a which identified whole-genome , of of data sets use sequencing independent that sentation made different suggesting studies study, in not recent original genes More could different the involve 2017). in 2014) may al., as Gugerli, et cues types (Rellstab environmental & habitat populations similar Schnyder, contrasting to with were Holderegger, adaptation associate loci Brodbeck, local to outlier (Buehler, variants such SIT4 populations confirm cases, in of two protein set family In family phosphatase-associated independent 2013). SIT4 nucleotidyltransferase the et Gugerli, a matched one (Manel & to regions: adaptation from Schnyder, gene within local known Zulliger, located others, of protein 2010; be than drivers to al., differentiation found environmental and genetic et main sequenced higher Poncet the significantly In as 2010; to (AFLPs). due precipitation al., polymorphisms adaptive and length be temperature fragment to landscape indicated amplified as considered to using loci, referred scans AFLP commonly adaptation, outlier genome local on of signals relied genomic genomics, identify to attempting studies elevation. Earlier same the genomics signals at Moreover, Landscape pressure, located phenotypes. | herbivore populations growth–defence 5.2 between among to link even associated a site-specific they found were which authors adaptation compounds, the of defence adaptation, using and of experiments drivers traits chamber biotic leaf including climate study and single observations yet (2019) field Moraes combined De of & but Mescher, populations clines, Widmer, Buckley, elevational contrasts. elevational considered in and also adaptation latitudinal to local relation in regional on variation with studies might phenology fulfilled mentioned flowering it partially likely The synchronize Scandinavia, only is to and 2009). are it Spain photoperiod Heide, that Hence, between & with cues site. distance (King interacts specific Spanish latitudinal conditions temperature includes the the that scales Given at expected regional site. much be transplantation and so foreign large not the the but between in Scandinavian, both markedly the on differed flowering adaptation at of that populations of onset comparison foreign the the and 2015): in al., detected (Tor¨anglocal also et accessions was Spanish plasticity and Phenotypic environments. Scandinavian contrasting to adaptation of layer .alpina A. .thaliana A. oivsiaevraini risrltdt rwhadhrioedfne nthis In defence. herbivore and growth to related traits in variation investigate to .alpina A. and .lyrata A. n on htaatto novdsmlrfntoa ahas but pathways, functional similar involved adaptation that found and .alpina A. .thaliana A. Belre l,21) n h te n a homologous was one other the and 2013), al., et (Buehler .alpina A. 7 hsmgtntb nxetdfra arctic-alpine an for unexpected be not might This . oue naitcdiesadtat htshow that traits and drivers abiotic on focused Zlie ta. 03.Hwvr eoyigan genotyping However, 2013). al., et (Zulliger .thaliana A. yicesn ohgnmcand genomic both increasing By . .alpina A. d ilmrule al., et Villemereuil (de .alpina A. .thaliana A. .alpina, A. and Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. itr a curdmn ie Kee ta. 07.I snwpsil odsettemcaim that mechanisms the HERE dissect 2 to FIGURE between possible studies now comparative is using It level in life 2017). molecular monocarpic relatives the al., to annual at et polycarpic its evolution from (Kiefer life-history transition times the to many life Brassicaceae, contribute two the occurred these whereas Within between plants, has plants. difference polycarpic monocarpic in history major branches in One reproductive 2017). global to Hughes, restricted is is see it flowering parity after of senescence modes that is on strategies details (for model die plant they classical before the like plants monocarpic as EVOLUTION such TRAIT plants section. AND Polycarpic next HABIT the GROWTH that in PERENNIAL traits conditions exemplified | environmental of as prevalent biology 7 life pivotal, developmental the its is the to of understanding according trajectories Hence, stage vary demographic individual. particular can with the life-cycle stage a interact of that enters plant composition within individual individual genetic spends any the of it which and contribution time at the differential seen time and be a The cycle also persistence. infer can population can which to we 2018), components observations, al., demographic et occurrences. such Philipp local From 1993; former Prock, of & legacy (Diemer genetic of soils a seeds arctic-alpine as viable several Conversely, of populations selfing. bank through studied seed inbreeding six with the combined between likely differentiation genetic strong Ac- However, extinct. 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In were (Knops, 2015). responses rates to processes Miller, contributed opposite life-cycle & Brys, different further from Ochocki, between Jacquemyn, and result trade-offs elevation can from for low components or compensated at drivers growth life-cycle elevation than between population higher reduces higher correlations at which at ve probabilities, probabilities smaller survival survival was in increased rates, components variation variation life-cycle particular, interannual spatial different Moreover, of In among fecundity. lack compensation 2020). lower This demographic 2020). al., to al., ascribed et et partly (Andrello (Andrello be range could elevational rates full growth in its elevation, across Swenson, along rates Moorhead, growth rates because demographic population (Read, temperature, in temperature of clines effect marked with 2015). leaf Despite an Poschlod, increase R¨omermann, specific or & Rosbakh, commonly large plants 2014; area other with Sanders, with leaf species & competition Bailey, specific tall-growing to and of response height composed a vegetative was indicate vegetation could 2013). surrounding This individual Schweingruber, areas. the & that Nobis when observed 2017; fruits also Obeso, elevation. & more (2020) with (Laiolo al. decrease vice-versa and et to elevations, tended Andrello high fecundity inhabit to and tend growth found species while authors in increase, These findings to Alps. 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All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. age l,20) eebigteeryflwrn hntp of phenotype flowering early addition, the In resembling 2009). 2009), al., al., et et Wang Wang R. of 2018; alleles al., non-functional et with (Lazaro buds flower type), form (wild plants accession reference the the of of plants orthologue the is which to 2009), al., et C Wang optimal LOCUS R. 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Data may be preliminary. n ucinltat eg de ta. 04 oi cwigue,21;Ra ta. 04 a eobserved be can 2014) al., et Read 2013; Schweingruber, & Nobis 2014; the al., et at Adler 2017), (e.g. Bomblies traits & functional (Yant and adaptation of endorses study populations the evolution outcrossing for of fields drawback availability a very the the as in but resources considered research been biological future available has guide selfing of in may Predominant wealth genetics that the avenues molecular using some and formulate evolutionary addressed we ecological, be Here, of to However, knowledge. life-history. await current perennial that expand of unknowns evolution to many the are and environments still arctic-alpine there harsh to that plants demonstrate evolved. of have adaptation we the histories overview, extent life this which contrasting and to In the annual see which to of in interesting divergence environments PERSPECTIVES be the the will | It to understand 8 2017). back to al., related et tool be Kiefer a can 2019; as al., traits relevant et Arabideae. developed (Hyun the been histories within have life histories perennial background life genetic perennial com- common from and step a result annual the might between often of synergies function Further lines is gene adaptation. 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Bosch, (Chopra growth formation monocarpic trichome the leaf resembles closely that in alpina strategy Arabis reduced life probably rapid-cycling be is a This to towards 2019). shift in found Albani, potential observed & been Soppe, perseverance have Hughes, vegetative 2012; reduced dormancy al., the et (Albani auxin to survival PEP1 of related common plant of In supply for role 2020). relevant continuous Vayssi`eres al., is This 2021; the et non-functional function 2009). al., ensuring with al., et accessions by (Soppe et trials, flowering Wang buds after garden maintenance R. dormant even the 2018; branches maintain ensure axillary al., to vegetative to et from contribute vernalization (Lazaro pleiotropically year during flowering following also the to may growth habit commit polycarpic growth not vegetative the did of to that contribute suggests meristems also This in may 2018). upregulated meristems, al., vegetative et of and (Lazaro habit reproductive vegetative vernalization-mediated between remain in observed which differences as age-pathway, meristems, the juvenile besides in that not but exposure, cold silences stably Vernalization 1999). intra .alpina A. within pcfi ee.Oecud o xml,seiclyadesaatto oeteeenvironmental extreme to adaptation address specifically example, for could, One level. specific a lobe sdt td ute eeomna ris(xmlfidi i.2,wihinclude which 2), Fig. in (exemplified traits developmental further study to used been also has igeseis iial,i a etse whether tested be can it Similarly, species. single a Tbe1 eiwdi op,V˜er el or,&Abn,2021). 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RAlvl are levels mRNA in Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. rvosyngetdacsin rmArc Drauae l,21) ti lopsil htacsin from accessions that possible also is It the 2017). of al., understanding of et the example (Durvasula the change habitats Africa for might seasonal from shown accessions extreme lineages been has neglected the distinct what previously from such to differs similarly of history, radically analyses evolutionary exposed, the species’ Moreover, are daytime latitudes. and equatorial populations Northern the Alps African Particularly variantsof habitat. the genetic European divergent which unexpected strongly the contain to a might to climate, from islands adaptation provide sky populations that African traits on and underlying mountains centred Anatolian the is from using Accessions research tested aforementioned accession and reference 2006) the Spanish al., of et Whitham most 2013; While al., to et and developed (Gugerli pathways, be concept could physiological phenotype hypotheses alpina common involved Here, extended A. plant. how the the host in the unclear to of relation co-adaptations remains trajectory in evolutionary to it the lead under influence Nevertheless, alternative they they evolve uptake degree symbiosis. can which whether phosphorus mycorrhizal soils, are, for nutrient-poor common associations often support otherwise alpine, these microbial on the growing indicate to taxa results that solutions suggest Their and content. conditions phosphorus limiting only on date, low To microbiota with symbioses. fungal mutualistic soils characterized close (2017) form many In to al. organisms. roots et how mutualistic plant on with available with associate is to associate information 2019). ability microbiota limited their al., other is et or plants Choudhury fungi of 2017; success taxa, Parisod, adaptive on & the information to Neuhaus, (Choudhury, pertinent key Another TEs the of and distribution patterns (Richards et and methylation changes Thieme activity genome-wide epigenetic types, 2018; between how of the Bucher, evaluates links ability & that establishing the (Thieme investigation in to generation any potential contribute next of unaware 2017) the are al., in we et transmission response present, Epigenetic adaptive expression. At acquired gene 2017). of an al., regulation to the lead or in integrity then gene cold can affects turn in to in e.g. which adaptation (TEs), shown, ments in of been addressed has aspects stress further been functional Environmental yet the has are there understand metabolites natural However, 2021). better storage shaping al., to of alpina et protection led (Sergeeva needed physiology frost behaviour accumulation studies the perennial with sugar eco-physiological and Moreover, trade-off growth temperatures, 2012). secondary physiological warmer to al., elevations, relation a at low et However, to suggesting by compared (Wingler levels. senescence, high identified variation sucrose from leaf was that plants leaf in accelerated trajectory of tolerance higher to similar significance stress detected by cold A adaptive and higher mediated 2017). suggesting stress found likely who elevations, Ananiev, cold (2015), (but to high & al. adaptation herbivory Markovska, et response at of Wingler and Stanev, addressed tolerance view Ivanova, have pathogens frost in (Kolaksazov, studies as relevant increased trait physiological such are with Initial stress that variation 2019). biotic traits clinal al., regarding many flowering, et of particular still Buckley regulation are in see might the environments, There research besides Future arctic-alpine traits understood. relevant processes. well to ecologically adaptive increasingly of polymorphisms in basis involved is identified genetic have being which molecular for studies the candidates genome-wide address promising and further are approaches that (cf. genomic genes taxa landscape within other of in variety processes broad physiological A system govern 2017). model also arctic-alpine Lee, that an & specialist particularities from Kim, snow-bed genomic learnt Park, alpine the lessons the Here, pinpointing 2019)? of help gene al., ability (K¨ornermay or et inhabit the temperatures basis genes molecular that freezing as same the at such the How species is grow within physiologies, What in season: found processes? specialized challenges adaptive growing differences highly these ecological are short in of involved these exist, pathways and strategies to complementary were cover different solutions or Do networks, snow evolutionary habitats? of persistent alpine basis and temperatures, arctic genomic low the irradiance, is high similar as such factors . . Pajares uuesuissol xadaogtepyoegah of phylogeography the along expand should studies future , .alpina A. .alpina A. .thaliana A. a eetfo uhro soits o xml,Almario example, For associates. root such from benefit may .alpina A. ocp ihevrnetlsrsos ee esegreat see we Here, stressors. environmental with cope to 11 oidc nacdatvt ftasoal ele- transposable of activity enhanced induce to , ot n on eti aat edmnn on dominant be to taxa certain found and roots .thaliana A. odnlapusilla Soldanella fe including after .alpina A. A. to . Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. lai,J,Jea . udr . agn . ucr,A,Culn,G,&Bce,M (2017). M. Bucher, & G., Coupland, A., Zuccaro, G., Langen, nonmycorrhizal nutrition. J., of (2012). Wunder, phorus microbiota G. G., fungal Coupland, Jeena, Root-associated . . . J., R., Wang, Almario, J., Wunder, L., J. flowering. perennial Mateos, W¨otzel, S., L., strategies. Castaings, PEP1 history C., life America M. plant of Albani, States in United the variation of explain Sciences Franco, & of traits Academy C., Mbeau-Ache, Functional J., Ray-Mukherjee, (2014). S., J. M. Hsu, A., Salguero-G´omez, Compagnoni, B., R., P. section. Adler, Information Supporting the in online with REFERENCES found article be the may wrote information FG supporting Additional and version. final MAK the MCA, approved MA, and Information SW, read Supporting outline, authors the all conceived and FG GC, from and contributions GC MCA, MA, SW, contributions Author CR32I3_149741). (GeneScale, Foundation for Science inspiration under National the Foundation) Swiss acknowledges FG his Research Sciences 390686111. Plant of German ID: on much (DFG, Excellence 2048/1Project Forschungsgemeinschaft of StrategyEXC Cluster Deutsche Excellence the the acknowledges Germany’s MCA by article. funded the of (CEPLAS) helpful version report provided Ir`ene earlier meeting Till-Bottraud and an a Schneeberger on to Korbinian comments contributions Rellstab, and Christian discussion review. valuable on for this symposium stimulating 2018, a in of Germany) participants (Cologne, all Research thank Breeding to like environment. would biotic authors and The abiotic their responses with biology evolutionary cope plant plants of evolutionary ways ways of ACKNOWLEDGEMENTS manifold complementary field the the address about enrich to knowledge will relying, to taxa contribute still studied and intensively is Additional, research cues. of environmental array to taxa, huge ecologi- other a and of evolutionary which the suite on on a overview with this Together natural With biology. the studies. of comparative across genomics for encountered basis cal traits comprehensive life-history a as in funda- serve variation understanding marked help further The of will range adaptation. scales in environmental processes and 2020; phylogenetic mental evolution al., adaptive the the et widening understanding (Birkeland together, for Taken Brassicaceae opportunities the provide in and adaptation 2020) traits. Studies of al., life-history et feasible. principles of Rellstab address increasingly 2020; to becoming al., begun are as et just Nowak genomics such have comparative kind Brassicaceae sequenced, such fully the of being of species genera plant more well-studied other in Cardamine adapted those cold to from for expanded and advantages be adaptive can provide discoveries warming. Biological may global by that threatened variants are genetic that contain species climates southerly more osrim 20) itr reooy usrt yea ao rvro pta eei tutr in structure genetic spatial of driver major IntraBioDiv a . as . . type Sch¨onswetter, P., S., Substrate Manel, ecology? R., or Holderegger, History A., (2009). Tribsch, Consortium. C., Thiel-Egenter, N., Alvarez, 10.1073/pnas.1710455114 of .alpina A. rbsalpina Arabis and Arabis .alpina A. Brassica oehrwt h ulcyaalbegnmcrsucsetbihdi eetyas will years, recent in established resources genomic available publicly the with together , rceig fteNtoa cdm fSciences of Academy National the of Proceedings LSGenetics PLoS eerhb ireTbre n ofHleegr n nnilspottruhthe through support financial and Holderegger, Rolf and Taberlet Pierre by research secddb w vrapn ee htcnrbt ontrlgntcvrainin variation genetic natural to contribute that genes overlapping two by encoded is K¨ mr 05 uhot,Sn,Le icelOd,21) ihmr and more With 2011). Mitchell-Olds, & Lee, Song, Rushworth, (Kr¨amer, 2015; edmntaeta hsseisi aubeeegn oe ytmi plant in system model emerging valuable a is species this that demonstrate we , , 8 1) 1010 o:10.1371/journal.pgen.1003130 doi: e1003130. (12), .alpina A. .alpina A. 12 , 111 ocoeyrltdsse pce ihnteArabideae, the within species sister related closely to opeet h ihrooutstanding hitherto the complements rbsalpina Arabis 2) 01–01.di 10.1073/pnas.1410430111 doi: 10019–10019. (27), rbsalpina, Arabis n t otiuint ln phos- plant to contribution its and Arabidopsis , 114 rceig fteNational the of Proceedings otda h P o Plant for MPI the at hosted 4) 90–91.doi: E9403–E9412. (44), , Capsella .thaliana A. , Boechera, , Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. yr,S . as,W,&Hffan .A 20) oa dpainadcgain eeto ntealpine the in selection cogradient and adaptation Local (2007). A. 5646.2007.00248.x A. Hoffmann, & locus W., plant, outlier Papst, An G., (2013). S. replicates. F. Byars, regional Gugerli, independent & across P., plant Taberlet, alpine sys- S., an Ecology Manel, mating in R., selection and habitat-mediated Holderegger, flow in N., relevant B. gene Poncet, Contemporary D., (2012). Buehler, F. 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(4), 4–7.doi: 141–173. , aueReviews Nature , , species. 66 23 PEP1 TFL1 Plant New (4), (1), Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. eainhp nAaiee(pce ubrgvn.CaedfiiinflosKr oh(03 n Kiefer and clade including (2013) clade on Koch the knowledge & artifact of phylogenetic Karl analytical areas follows present-day an Ancestral definition of (C) than Clade (2017). Summary rather given). al. number (B) phenomenon et (species biological 2020). 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Koch, included & were Ansell, Kiefer, those caucasica Arabis (Karl, of applied all be to If have would clade. 2012). concepts unresolved subspecies phylogenetically entities, monophyletic same establish the within found monophyletic resulting The genera. new into transferred be type to the contain have (i) would will species present genus of monophyletic at set a genus for the aims work in taxonomic contained future if However, 1753). genus 664, of species 2014). type 2013, The genus Koch, of & considerations that (Karl taxonomic variation syndrome and genetic selfing lower Systematic the exhibit from and result radiations, others, subsequent undergo among annuals not might, in did e.g. ranges, linked, Generally, distribution inde- often 2013). lowland the Koch, clades are & different and from (Karl convergently the world evolved ranges detailed there, the have vs. From which a of characteristics regions 2017). (for trait A), mountain Conti, Mountains morphological alpine (Figure & Pamir numerous and Staedler, region and temperate the Manafzadeh, Irano-Turanian Shan colonized region: Tian the pendently floristic the is this to on Arabideae regions review Saharo-Arabian tribe occurred and of rates Mediterranean origin eastern speciation of increased center which between The in split a species, involved clades sister various perennial in are radiations montane/alpine frequently. mean 63% Multiple family and 2013). genus decades 2020), Koch, the annual two the & past than al., of lowland (Karl the paraphyly higher concept et of demonstrates systematic times Research that German, new lineage. three tree a this (Walden, phylogenetic than of well-resolved Arabideae dynamics more a evolutionary in is tribe the resulted highlighting rate within 2020), diversification al., species net et (Huang mean 545 its and and genera neopolyploids, 18 the Among Arabideae tribe of Evolution 1. 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Arabis .deflexa A. srpeetdb h ina ooyeof holotype Linnean the by represented is .alpina A. Arabis .alpina A. , , .inclx .caucasica A. ionocalyx, A. .auriculata A. .nova A. oehrwt prxmtl 0adtoa pce htare: that species additional 20 approximately with together r h e rpn oeigtaxa flowering pink or red the are Arabis 21 ssp. rbsalpina Arabis .alpina A. iberica ld 3seis al&Kc,21;Kee tal., et Kiefer 2013; Koch, & Karl species; (3 clade , .kennedyae A. ssp. n t itrcae,hglgtn the highlighting clades, sister its and and caucasica .alpina A. .tianchanica A. Arabis, ul pawell-defined a up build ) adtoa yoysare synonyms (additional hnms ftespecies the of most then .purpurea A. .alpina A. Arabis SeisPatrm2: Plantarum (Species .nordmanniana A. Krysa)are (Kyrgyzstan) n established and .alpina A. ssp. , .cypria A. brevifolia ( A. to Posted on Authorea 1 Mar 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161461048.89520056/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. R1ABC u omnyBddrac asirse l 2020 al. et Vayssieres 2019 al. et Chopra 2021 al. Reference et Zhou 2013; al. et Bergonzi 2014 al. et Chopra perennial-model-plant-for-ecological-genomics-and-life-history-evolution figA_Box.pdf 2021 al. et Zhou file Hosted 2019 al. 2011 et perennial-model-plant-for-ecological-genomics-and-life-history-evolution al. Hyun et Wang fig2-1.pdf file Hosted perennial-model-plant-for-ecological-genomics-and-life-history-evolution W¨otzel.fig1.pdf S. Photos: (E). lateral traits at developmental file (arrow) glasshouse in Hosted large-scale rooting variation of adventitious occurring Illustration (D). naturally B), density for (A, trichome testing low population with survey variant same occurring the naturally (C), from branches plants Photos: between vernalization,p rosettes. varies to vegetative polycarpy glasshouse response illustrate of vernalization, patterning Age-dependent photos base Earth. Trichome to 2 the row Natural response close-up at with FIGURE Bottom Age-dependent The shoot made dormancy (D). W¨otzel; mature individuals. map Bud verges S. a (G) 2019 and road from fruiting al. 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Karl of intervals; source confidence a as region” ”Irano-Turanian rbdpi thaliana Arabidopsis osre n ies oe fgnsfntoal hrceie in characterized functionally genes of roles diverse and Conserved vial at available ape ouain of populations Sampled vial at available xmlso eeomna rissuidusing studied traits developmental of Examples vial at available (At). https://authorea.com/users/398897/articles/511488-arabis-alpina-a- https://authorea.com/users/398897/articles/511488-arabis-alpina-a- https://authorea.com/users/398897/articles/511488-arabis-alpina-a- rbsalpina Arabis oe ag at fisgoa ag c.Ansell (cf. range global its of parts large cover 22 .alpina A. rbsalpina Arabis acrossre() os habitats moist (B), scree calcareous : AD:Foeigtm nthe in time Flowering (A-D): rbsalpina Arabis A)compared (Aa) tal. et ,