Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. eaehLazaro Xenabeth upland malouinus in endozoochory of Evidence geese mosses? disperse Do eree rmterfcsaeval n aal frgnrtn e ln ise lvnfcsds ape otiigunde- containing samples disc feces Eleven tissue. gametophytes plant bryophyte new same of regenerating these fragments of in that from capable endozoochory hypothesized fragments and diaspores We of moss viable of role termined are viability Chile. the feces the sub-Antarctic evaluated their for We in evidence from species ambiguous. of retrieved the remains regenerate, totipotency sympatric tract to the intestinal known and Given bird herbivorous been have the shown feces through diaspores. has bird passage bryophyte Research from following of fragments dispersal. ( few bryophyte dispersal goose Although endozoochorous upland exogenous to fragments. the the contribute of also for samples could vectors fecal birds that potential bryophytes, many as of act tissue vegetative to known are Birds Abstract 2021 2, March 3 2 1 obrsdses oss vdneo noocoyi padgeese upland L´azaro in A. Xenabeth endozoochory of Evidence mosses? white-bellied disperse birds relationship Do birds, This Keywords: change. dispersal. climate bryophyte facing habitats in can colonize role sub-Antarctic and and mosses, critical viable disperse endozoochory, bryophytes a remained dispersal, way play bryophyte diaspores the altitudinal may in and moss birds implications fliers that robust important sub-Antarctic fact have these that may of The tracts suggests intestinal growth. migrants, the new through latitudinal passage produced and from the mosses following fragments conditions wild of experimental from under 20% propagules regenerate days, of 91 67% After and feces, medium. agar-Gamborg a using ( mosses mi drs:[email protected] USA address: 33145, Email Florida, Miami, ST, 23rd SW 3190 L´azaroXenabeth Author: Corresponding USA Texas, Denton, Texas, 3 2 1 nvriyo ot Texas North of University Magallanes de Universidad Florida of University u-naci iclua osrainPorm nvria eMglae,PnaAea,Chile Arenas, Punta Magallanes, USA de Florida, Universidad Gainesville, Program, Florida, Conservation of Biocultural University Sub-Antarctic Conservation, and Ecology Wildlife of Department eateto ilgclSine n dacdEvrnetlRsac nttt,Uiest fNorth of University Institute, Research Environmental Advanced and Sciences Biological of Department ootmmtetragonum Conostomum hopaapicta Chloephaga 1 nsbAtrtcChile sub-Antarctic in 1 1 o Mackenzie Roy , o Mackenzie Roy , .picta C. tai malouinus Attagis , ytiharobusta Syntrichia hopaapicta Chloephaga ( N 6 and =6) 2 am .Jim´enez E. Jaime , n ht-ele seedsnipes white-bellied and 2 n am Jim`enez Jaime and , .malouinus A. and , n ht-ele ednp ( white-bellied and ) nsbAtrtcChile sub-Antarctic in oyrcu strictum Polytrichum 1 ( 3 N 5 n i osfamn ape rmwl collected wild from samples fragment moss six and =5) .malouinus A. 3 eegrown were ) ee,5%o rget from fragments of 50% feces, tai malouinus Attagis xsitu ex hopaapicta Chloephaga Attagis npa oland soil peat in oti bryophyte contain ) .picta C. nvitro in and Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. etad,tnr,adapn ein Gffie ta. 02.Wt lmt hnea rwn ocr for concern growing a forests, as temperate change with climate associated With typically 2012). anywhere al., found are et are they (Goffinet and However, Earth regions rainforest. on alpine tropical plants and of the tundra, forms to peatlands, earliest tundra the the of descendants from the considered are Bryophytes interest the would with that line and in processes is natural research Introduction of this readers. understanding Hence, world. of our the furthering audience the of dispersal in broad and moss regions Evolution a in other change and shift many Ecology paradigm climate in a of to occurs to aims out likely context points and that prevalent the evidence is In convincing that presents cannot. paper this factors through important. that abiotic been consider especially disperse We may is has the therefore this dispersal in that and communities, moss defecate endozoochory places biotic for through that into of vectors paradigm birds “reshuffling” biotic and the as migratory ways far, serve flying in birds Thus strong that found. mosses report are are We these rain. mosses mosses and that the viable wind considering where defecate as especially birds substrates finding, Further,like that novel mosses. indicated a the is species fact of This The bird totipotency Chile. individuals. both of the new by south indicated produce far mosses can fragments the bird-ingested that in moss sites, of evaluated wild same concurrently regeneration from the We the from protonemas substrates. collected different obtained two mosses we on from that feces fragments moss the of content of in the viability found birds. examined the mosses wild We of birds. these by fragments birds, by endozoochory the herbivorous ground ( through cultivated the by seedsnipes dispersed on be white-belied ingested can feces of being mosses feces the that after in idea We individual deposited the mosses. tested new being of We and a dispersal system the establish digestive for to their mechanism novel mosses through potential moving of a ability addresses the that upland research investigated in experimental an endozoochory Evolution. is and of This Ecology Evidence in mosses? publication disperse its birds for “Do dered manuscript our submit geese to excited are We Letter Cover bryophytes way change. the Keywords climate in implications facing important habitats have a colonize may play relationship and may This disperse can birds robust dispersal. sub-Antarctic and bryophyte these in that viable of role suggests remained tracts critical migrants, intestinal diaspores latitudinal the moss and through that passage altitudinal fact the and The following fliers growth. conditions experimental new under produced regenerate mosses from wild fragments from of propagules 20% days, ( 91 mosses collected feces wild strictum Eleven from passage tissue. Polytrichum plant samples following new from fragment regenerating fragments same moss diaspores of moss these six capable of bryophyte undetermined in and of containing viability endozoochory viable fragments samples that are of the disc hypothesized feces role We their for Chile. the from sub-Antarctic evaluated evidence retrieved in We gametophytes species the ambiguous. bird remains sympatric regenerate, Given and tract endozoochorous ( to herbivorous intestinal diaspores. goose to known bird bryophyte upland contribute the been the through of also have of dispersal could feces samples exogenous birds bird fecal ( bryophytes, the that seedsnipe many shown for white-bellied of has vectors Research tissue potential dispersal. vegetative as bryophyte of act totipotency to the known are Birds Abstract hopaapicta Chloephaga id,bypyedsesl noocoy oss sub-Antarctic mosses, endozoochory, dispersal, bryophyte birds, : eegrown were ) n ht-ele seedsnipes white-bellied and tai malouinus Attagis tai malouinus Attagis .malouinus A. xsitu ex npa oland soil peat in oti roht rget.Atog e rget from fragments few Although fragments. bryophyte contain ) ee,5%o rget from fragments of 50% feces, n fsmarculn es ( geese upland sympatric of and ) tai malouinus Attagis 2 ootmmtetragonum Conostomum .picta C. nvitro in ( N sn grGmogmdu.After medium. agar-Gamborg a using nsbAtrtcCie ob consi- be to Chile” sub-Antarctic in 6 and =6) .picta C. .malouinus A. , ytiharobusta Syntrichia hopaapicta Chloephaga hopaapicta Chloephaga ee,ad6%of 67% and feces, ( N 5 and =5) and ) and ) and , Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. ta.(00 bevdta ea ape olce nadyn nwetbdwr opie faot50- about of comprised were bed snowmelt drying a by on bryophytes collected of samples consumption fecal the that of observed reported observations (2020) been first al. et the has knowledge, endozoochory malouinus 2017; our through To al., A. dispersal 2019). et bryophyte ( al., (Wilkinson of foxes et endozoochory evidence flying through Additional spectacled bryophytes 2020). dispersing in al., of et capable ( up ducks Russo km be level mallard might 400 sea as shorebirds, some such from birds, and observed migration that 1000-1500 altitudinal been comm.). suggests daily between have research pers. Jim´enez, perform and Previous 2020, Island (J. geese ground Navarino there some wintering on masl and the 700 tagged Argentina over on in geese to masl Gallegos Upland Rio 100 ground. study near than same breeding Santa north lower the to the as to grounds) altitudinally, on wintering moved migrates male (the masl individual also Aires a between species the Buenos of migrating This from that route grounds). America, km found breeding migratory 1986). 1485 South (the of the al., through Patagonia distance tracked province, movements minimum et (2015) Cruz a larger (Hayman al. migrated have Islands he et to that Falkland Pedrana found known grounds. the is wintering to Seedsnipe that and Patagonia White-bellied breeding goose be leave del “e-Bird: migratory could to 1990; a 2003; (Fjelds˚a and known Krabbe, is al., winters & America been harsh et the occasionally South Jaramillo during has sub-Antarctic 1996; flats al., lowland inhabit to et that habitats Hoyo upland species its dispersal. bird from bryophyte downwards herbivorous for two vectors are potential 1996)) al., al., also Herv´ıas-Parejo ( et et has 2019; seedsnipe as al., white-bellied ingestion, the et internal and ( Blanco through 2018; goose endozoochory, upland al., via The et this (Lovas-Kiss transportation 2020). do herbs al., external to other et by able Silva and ectozoochory, 2019; ferns be via & with may Am´elio (Chmielewski distances shown and 2012; movements long been 2014), al., migratory bryophytes et al., through Parnikoza disperse transcontinentally et 2006; and (Lewis can al., 2018), et act Birds may al., (Calvelo them 2019). they reach nesting et Eppley, as help and Parnikoza bryophytes or foraging 2017; of movement otherwise, through al., long-distance reach al., sites et and not local et directed would in in Zhang they aid dispersers as that 2007; can serve as areas birds may al., reach of birds to et behavior these plants The gametophyte habitats, Proctor quicker. small same entire these 2002; the the allow share Cleavitt, bryophytes that regrow and 1997; vectors to birds Longton, some facilitated able because also 1963; mechanism However, is (Anderson, 2003). colonization reproduction fragments and asexual dispersal tissue an Bryophyte attract from 2011). totipotency, to al., matter, of cell et organic bryophyte number (Jofre decomposing by vectors mimicking large odor, dispersal strong a potential a reproduction as of emit flies like sexual that production sporophytes, species, by colored the Some brightly 2018). facilitated and or al., dispersal sporophytes, et Zanatta tall wind 2017b; spores, or changing a anemophily, al. exposed Barb´e in (Mu˜noz, et survival 2004; as through spores their such to possible fundamental features is made areas morphological challenge new is in a establish This represent and may disperse planet. to which bryophytes regime 2005), of Faced climate ability Graham, The ancestral species. mosses. & or a as niches (Wiens by such their tolerances organisms habitat, “following” conser- climatic sessile by a niche for respond their as by to adapting such driven likely montane characteristics, than is more in that ecological rather are followed shift 2018) ancestral species (23%), upslope al., change, of highest et climate an retention Freeman be by causing the 2015; will is Tingley, is species change & which 3 of climate (Elsen vatism, by bry- risks Additionally, communities increases extinction for animal (14%). temperature where potential and Zealand Earth’s regions plant and New the three mechanisms and if of dispersal Australia (2015), one the by Urban be understand to latitude will to According high America important habitats. in South is new endemism colonize bryophyte it to of 2008), ophytes proportion al., large et the (Rozzi and regions habitats, high-latitude and high-elevation nsbAtrtcSuhAeiawr aeb eln ta.(06.Mr eety Russo recently, More (2016). al. et Behling by made were America South sub-Antarctic in hopaapicta Chloephaga trpsconspicillatus Pteropus tai malouinus Attagis , 06,o i ae,sc sgmao pahcp Gie 07;Glime, 2017a; (Glime, cups splash or gemma as such water, via or 2016), re neioms aiyAaia Croea iwn 2020)) Kirwan, & (Carboneras Anatidae family Anseriformes, order , .malouinus A. alradubyi Tayloria Prose l,20)adfehae s (Boedeltje fish freshwater and 2007) al., et (Parsons ) re hrdifre,fml hncrde(e Hoyo (del Thinocoridae family , order , 3 sa liuia irtr hrbr htmoves that shorebird migratory altitudinal an is nsplatyrhynchos Anas aeee enkont aesik spores sticky have to known been even have , tai malouinus Attagis n u-naci geese sub-Antarctic and ) .picta C. .picta C. ,nd)and n.d.) ”, .picta C. and and ° C, Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. rmec ea ape(prxmtl mi hcns) ahds a isle eaaeyi la dishes clear in eliminate separately to dissolved sliced was feces then disc were the Each discs of thickness). thin layer in Two mm fragments. outer 1 and the (approximately spores remove sample soil-borne fecal to or each wind- knife from by precision contamination disinfected 2010). of possibility Goffinet, a the & were used (Buck We field Navarino the processing. Isla in before of collected mosses laboratory, Bryophytes compared to the conditions. capabilities key In growing preliminary regeneration same the for the using test RM under to by samples sites identified fecal sample from fecal fragments near to sampled were Bartramiaceae), family two ( robusta were processing there for ( site samples, families each six three from those selected Of was one species. picta only bird through later each field and from the site three each in from identified 586779.97E collected were 3908430.45S,19F were Za˜nartu (19F birds 587490.15E birds fragments both these Lake (19F observing of summit line, level, experience samples Bandera tree personal sea Fecal our the Cerro 3907270.36S). above near the 586790.75E sites on sites 19F four ponds and 3907237.18S, two large 3911044.12S), opportunistically and 585969.41E streams (19F sampled Bay near We 13 Robalo additional sites. and 3911888.71S) an lowland 587064.87E 2019. over and April monitored to upland was fresh 2018 on conducted fragments December field, were recovered from work the of Island, laboratory Navarino and growth In in Field the line. chamber and tree growth the weeks a above five in m weeks of 100 some course meadows tundra the open Magellanic ( over in a in masl, species thrive 700 beech mosses, ca. and southern at lichens of graminoids, some mix (M´endez al. plants, seasons. environment a et cushion moderate including by still plants, covered but small marked, is of with forest topography the rugged pumilio a Nothofagus ranges, has lower island the The Americas. At the of end (54 Island southern Navarino on conducted was research The 2018). al., the vegetation et the like altering Bjorkman Methods Chile, is 2008; & change sub-Antarctic al., climate Materials and et beyond 2005) (Klein regions Bergersen, relations & to (Fox plant-herbivore widespread mosses and a applicable on be communities feed bryophyte be to also likely the could birds is Our conditions, where Endozoochory findings endozoochory. Arctic, proper individual. through our the new a sub-Antarctic under Therefore, regenerate cultivated the to phenomenon. and in have totipotence defecated the bryophytes species have ingested, avian for will being fragments both vectors after that dispersal that propose is species, as We prediction serve bird feces. to to herbivorous lab their necessary potential under two from was grown retrieved the of when gametophytes it role bryophyte regenerating feces, potential fragmented of bird of capable the in were of feces fragments observations malouinus digestive bird viable A. make bird’s in of we found the search Here, fragments through bryophyte the conditions. passing the with after if together test propagate Thus, first reasons, successfully dispersal. successful these fragments with For these correlated inappropriate directly system. that is is of conclude it feces establishment bird conditions, and cannot in field success dispersal we fragments under the bryophyte little successful that finding with Given be that 2020). to fragments assume al., to steps these et on interdependent Russo regenerate and feed 2017; sequential or that al., involves cultivate et plants species Wilkinson to avian 2007; attempted al., from et has recovered (Parsons research been have previous fragments plants, gametophyte these bryophyte viable though Even as eecmrsdo bu 010 roht prpye.O l ea ape h uhr noee,91% uncovered, authors meadows the flooded samples on fecal collected all Of those sporophytes. and bryophyte Polytrichaceae, 80-100% family of about moss of the comprised from were fragments sporophyte 80% .picta C. Polytrichum ee rma padst,adthree and site, upland an from feces Ho.&Ge. ..Zne,fml otaee and Pottiaceae; family Zander, R.H. Grev.) & (Hook. n 5 of 85% and sedzohru roht ipresb etn h iblt n eeeaiecapabilities regenerative and viability the testing by dispersers bryophyte endozoochorous as , oyrcu strictum Polytrichum p,wt tlatoegnrtn e growth. new generating one least at with sp., .picta C. .picta C. rwn nkumozfraina h reln.Aoetete ie ihcommunity rich a line, tree the Above line. tree the at formation krummholz in growing ) .malouinus A. , and 03.W sampled We 2013). and .malouinus A. .malouinus A. rd,fml oyrcaee lokona pgo wheat”; “pigeon as known also Polytrichaceae, family Brid., , ape otie roht rget,icuigfamnsidentified fragments including fragments, bryophyte contained samples .malouinus A. nsitu in ea ape eesoe npprbg tro temperature room at bags paper in stored were samples fecal .picta C. ea ape eecletdfo i oain,focusing locations, six from collected were samples fecal ° ,67 S, utpefclsmlsta iil otie bryophyte contained visibly that samples fecal Multiple . 4 ee tsalvland level sea at feces ° ) aelncrgo nsbAtrtcCie tthe at Chile, sub-Antarctic in region Magellanic W), ee rmuln ie.Adtoal,mse from mosses Additionally, sites. upland from feces ootmmtetragonum Conostomum .picta C. ee rmlwadsts one sites, lowland from feces .picta C. ohfgsbetuloides Nothofagus and .malouinus A. Hd. Lindb., (Hedw.) .picta C. Syntrichia n feces =6), and and C. Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. ml rcino osdaprsrmisval olwn h asg hog h netnltato both of tract intestinal the through that reveal passage results the These following 1). Table viable (see remains treatment diaspores soil moss peat bryophyte the recovery of of in fragment evidence fraction bryophyte clear species small observed allow bird we a would both small, any, was of size if feces sample which, from the test regeneration Although to conditions. lab was under treatments growth chosen and the for growth. reasoning of Our days would 42 from samples until and fecal cultures 1961) and al (Proctor, our et moss days from (Wilkinson wild 60 days after both 11 spores from after bryophyte fragments fragments from bryophyte the growth that described to research predicted feces We Previous regenerate. the field. the from from recovered taken fragments samples the an an of for and consisted conditions soil, These peat growing growth. bryophyte two of set probability the to increase decided we feces However, in methods. found fragments tested cultivate inoculations al to six et attempted the of has any research species. for Previous bird agar-Gamborg both the of in vegetative observed showed feces medium was the agar-Gamborg growth from solid no the contrary, in the mosses On wild culture treated fragmented growth. five of inoculations of fragment inoculations occurrence of from fecal (100%) the (56%) (three three One on nine treatment all the based tested. soil of 1) Five (Table treatments peat 1). growth the (Figure the new to beds produced in moss (67%) and growth samples shoots fragment green produced moss light-colored wild (44%) with six 18 of of four mediums culture with 8 mediums of total 11 days. A 91 January of on total agar-Gamborg a Discussion sample for the and moss chamber 8 in Results wild 9 growth January January The inoculated the on on h. in fragments containers chamber 16:8 left soil sample growth of were peat the moss cycle samples the day:night in wild in a placed inoculated and at fragments were ratio, sample sample containers 4:1 fecal soil A a the between peat parafilm. by in fluctuating followed the lights with temperature LED in sealed the blue inoculated and with and fragments covered chamber, red then growth on a and grown into were plate 5 placed agar-Gamborg agar of were humidity the minimum 28-well mediums inner occasio- For a a culture experiment. High with in 18 the humid. experiment of placed throughout containers the total were containers culture of sealed samples the remainder the the keep the in treatment, to for observed in thoroughly was period sealed only soil, observation condensation and placed the one and the covered on therefore of placed throughout then was were misting exception and and fragments nal fragment, water the the bryophyte treatment, filtered with soil one from with propagated, peat contained and misted the be that sample For disc disc treatment. to fecal agar-Gamborg one treatment Once each the had 2006). either from only al., fragments (Gonz´alez to which et growth the sample sample, moss assigned species for randomly medium we moss specific processed, each a were 5.8, samples of fecal pH parafilm, all a with with openings Laboratories) the technology at together (Kekkil sealed vitro soil cups bryophyte in 138 plastic peat two and sterile container, analyzed, commercial culture samples with a disc either 11 to were in belonging tested resulted 4 (106 ments species) at inoculated bird days each and fragments. few recovered from into a probe fragments (three cut for samples a also stored fecal were filtered using were six with sampled samples The dishes filled mosses These partially the collected protocol. and wild from sample (2020) the 100x out disc al.’s Additionally, and each picked et inoculation. 40x to Russo were at corresponding following microscope possible tubes water, compound Eppendorf river as into HD fragments placed ICC50 bryophyte forceps, Leica and many a under As observed (0.22- were magnification. River contents Robalo disc the The from Ireland). collected water filtered using ., 07 us ta. 00.I hssuy eprilyfloe us ta.s(00 fragment-processing (2020) al.’s et Russo followed partially we study, this In 2020). al., et Russo 2017; ramn a aeuiga grGmogMxueBslSl B at)mdu G6,Phyto- (G768, medium salts) (B5 Salt Basal Mixture agar-Gamborg an using made was treatment .malouinus A. nvitro in º n aiu f25 of maximum and C odto sn grGmogmdu o ohfamnsfo ea n idmoss wild and fecal from fragments both for medium agar-Gamborg using condition rgetsmls(0) he fsxwith six of three (20%), samples fragment .picta C. rfsinlSbtae ramcolt for microplate a or Substrate) Professional ¨ a º hogotteetrt fteeprmn.Bypyecultures Bryophyte experiment. the of entirety the throughout C n from one , ., 07.Hwvr eddntosresgso regeneration of signs observe not did we However, 2017). 5 .picta C. .malouinus A. μ VFseiesrnefitr,Mlioe Cork, Millipore, filters, syringe sterile PVDF m n 2to 32 and xsitu ex nvitro in .picta C. n n from one and , rwhcniinuigcommercial using condition growth .malouinus A. ihsm ucs (Wilkinson success some with rgetsmls(0) and (50%), samples fragment Polytrichum nvitro in ape) h treat- The samples). th th 09 n fecal and 2019, , 09 l the All 2019. , .malouinus A. rwh The growth. sp ° th until C 2019, , . and ) Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. eln,E,Cvns,T . ei,L . iee,J . ont . oz,R 21) Dispersal (2016). Savannah, R. Rozzi, 2016. & Botany B., 224. Goffinet, Abstract E., Conference J. Jimenez, birds. R., by ingestion L. following Lewis, diaspores E., materi- bryophyte T. of Caviness, a E., as events Behling, dispersal Bryophytes Long-distance away: far (2017). so yet and 2745.12637. F. reassembly. close So metacommunity (2016). D. bryophyte Y. govern Bergeron, Peralta, & J., N. & Fenton, Barb´e, M., Mosses. S., (1963). Brazil. E. J. L. in Anderson, Lima, nests M., birds’ D. build doi:10.31055/1851.2372.v52.n2.17434. to Carmo, al landscapes. D., high-latitude can these we L. at Am´elio, so species dispersal conditions, bryophyte bryophyte field the in under identify results zoochory to these of References reproduce more type samples to this replication, fecal need of and the relevance the size in is the sample there understand found Finally, conducting larger However, fragments and in them. a soil, bryophyte birds. birds culturing of suggest of the before to types we role distinct of accessible to research, the variably sequencing are and of respond DNA but might endozoochory line fragments through rain, bryophyte in this comm.). as dispersal birds or along treatments, and bryophyte pers. substrate wind as progress test habitat Jim´enez, 2019, by To to bryophytes, suitable (J. established done process. of generally be graze this be to a dispersal they to needs within unlikely the where research propagate are further in conditions effectively they habitat role to where like critical bryophytes locations in likely enabling defecate but potentially to Thus, passive, likely holds a more and play are 2012) may axes al., Navarino horizontal birds et in and Sub-Antarctic ecosystems. bryophytes (Goffinet vertical their of the importance” to viability in changes ecological the by scales al. highest maintaining affected (M´endez time et of in species short dispersal “ecosystem critical endemic in as an especially many process serving contains is mosses this By which as process populations. in such This Island, viable aid organisms maintain landscape. to conditions, to the able warming latitudes of with be or as would altitudes short but important higher birds for locally, occur especially migrants to vectors, only move is altitudinal to not to fliers, This might capability have robust dispersers scale. their would as are broader increasing role species a and their bird distances, at rain, long possibly also both or for for sub-Antarctic, As migrants wind plausible latitudinal 2). the beyond and is (Figure in disperse distances types it birds to habitat herbivorous that region specific these shows this reach by in research bryophytes endozoochory our aiding through Consequently, further dispersed conditions. be laboratory herbivorous to of in fragments feces the regenerate from retrieved could gametophytes bryophyte birds fragmented that hypothesis the soil. confirmed We moss peat wild and hand, medium sub-Antarctic the other agar-Gamborg in Conclusions bryophytes the in these On of both endozoochory substrate. and bryophyte bird inoculated of soil of substrate longer hypothesis environment. evidence samples the peat take agar-Gamborg was support in the may results the there these observed in However, diaspores to Therefore, 2014). feces was moss acclimation al., from growth as necessary et recovered gametophyte 1), (Sabovljevic the fragments Table medium to in (see the due regeneration in species conditions conditions bird pH sterile both neutral of in tested germinate fragments to sample bryophytes mechanisms. the the fecal endozoochory that that the effective confirmed for plausible agar-Gamborg condition is the necessary Additionally,It in by a number. multiplied mosses absolute is were large wild which results a three our totipotent, into if the Given translate are bird, would in significant. sampled each results observed are by our evidence time, daily cases growth and defecated our area feces the small, per of birds Although number of the conditions. number and suitable the feces in the regeneration of volume of the capable is and birds nvitro in aa-abr)cniin ol aeyeddflengtv eut from results negative false yielded have could conditions (agar-Gamborg) h rooit 66 Bryologist, The , oe´ nd aScea retn eBtaia 52 Bot´anica, de Argentina Sociedad la de Bolet´ın 03,adjs sciia lblyweebypye r lobeing also are bryophytes where globally critical as just and 2013), ora fEooy 104 Ecology, of Journal 3,1719 doi:10.2307/3240710. 107-119. (3), 6 6,10-79 doi:10.1111/1365- 1707-1719. (6), 2,199-208. (2), Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. lm,JM 21a.Aatv taeis pr ipra etr.Cat -.I .M. J. In at available 4-9. and 2020 June Chapt. 3 updated vectors. Last dispersal Bryologists. Spore of Association International strategies: < the Adaptive and versity causes change (2017a). Climate community.Glime, J.M. bird (2018). tropical W. Glime, J. a Fitzpatrick, in 115 & extirpations Sciences, V., mountaintop of Academy Ruiz-Gutierrez, and N., shifts M. upslope Scholer, G., B. Freeman, geese barnacle between competition of 36 Lack (2005). geese E. footed Bergersen, & D., A. Fox, 876. under pp. species (1990). Copenhagen. montane N. of of Krabbe, University fate the & and J., topography Fjeldsa, mountain Global (2015). W. change. M. Tingley, climate & 2021). R., P. 12, February Elsen, (Accessed: In https://ebird.org/species/whbsee2/AR-V. Available: Seedsnipe (Seedsnipes). York. White-bellied Thinocoridae New Family eBird: occupied (n.d.). (1996). their J. eBird. to Sargatal, relation & world A., the in of Elliott, species birds D., moss J. Hoyo, common viable del and of rare vector of potential. dispersal tolerance dispersal 2745.2002.00713.x. novel Stress asexual a and (2002). as environments L. passerines N. Forest Cleavitt, (2019). M. S. 10.1098/rspb.2018.2253. Eppley, propagules. & bryophyte W., M. Chmielewski, 10.2173/bow.uplgoo1.01. doi: ( USA. Goose NY, Upland Ithaca, Ornithology, (2020). M. in G. nests Kirwan, World hummingbird & of C., structure Carboneras, and composition Botanical Antartica (2006). (Argentina). doi:10.1080/00222930500371000. (Prov. V. Patagonia Chile northwestern Ojeda, Navarino, from & habitats Isla different of A., Trejo, mosses the S., to Calvelo, key Preliminary of (2010). role B. P. The Goffinet, Chilena). E. & habits: C. R. feeding W. W. contrasting Verberk, Buck, with & availability. species M., propagule fish P. and by J. season, stream Lenssen, doi:10.3389/fevo.2019.00054. diet, temperate M., fish Vos, a traits, P., in plant Sollman, endozoochory dispersal Herb M., Plant Schaap, (2019). (2019). B., L. J. Klutman, Tella, G., & Boedeltje, fruit’? F., Weiher, the Hiraldo, biome. ‘foliage . A, Lovas-Kiss, Is tundra . D., cockatoos: warming Chamorro, . by C., S., a Bravo, P. across G., Beck, Blanco, N., change Ruger, trait S., Normand, functional C., Plant S. doi:10.1038/s41586-018-0563-7. Elmendorf, H., (2018). I. E. Myers-Smith, D., A. Bjorkman, 3. 30-August July Georgia. http://digitalcommons.mtu.edu/bryophyte-ecology/ 3,1318 doi:10.1111/j.0908-8857.2005.03540.x. 173–178. (3), J e oo .Elot .Sraa,D .Crsi,adE eJaa dtr) onl a of Lab Cornell Editors). Juana, de E. and Christie, A. D. Sargatal, J. Elliott, A. Hoyo, del (J. roht ecology Bryophyte oaHdii,Biet 138 Beiheft, Hedwigia, Nova ne brachyrhynchus Anser aueCiaeCag,5 Change, Climate Nature Vl ,p.5855.Breoa pi:Ln Ediciones. Lynx Spain: Barcelona, 538-545). pp. 3, (Vol. rceig fteRylSceyB ilgclSine,286 Sciences, Biological B: Society Royal the of Proceedings 4) 18-18.di 10.1073/pnas.1804224115. doi: 11982-11987. (47), Vl ,p.14) bo pnoe yMcia ehooia Uni- Technological Michigan by sponsored Ebook 1-43). pp. 1, (Vol. id ftehg Andes high the of Birds uta clg,45 Ecology, Austral uigtepebedn eidi Svalbard. in period pre-breeding the during 215-229. , 8,7276 o:10.1038/nclimate2656. doi: 772-776. (8), tai malouinus. Attagis ora fEooy 90 Ecology, of Journal 7 > Acse:Jl 3 2020). 23, July (Accessed: . 1,1216 doi:10.1111/aec.12835. 122-126. (1), rnir nEooyadEouin 7 Evolution, and Ecology in Frontiers hopaapicta Chloephaga ora fNtrlHsoy 40 History, Natural of Journal oehgn emr:Zooia Museum, Zoological Denmark: Copenhagen, . Br,CrelLbo rihlg,Ithaca, Ornithology, of Lab Cornell eBird, 5,7575 o:10.1046/j.1365- doi: 785-795. (5), ,vrin10 In 1.0. version ), aue 562 Nature, rceig fteNational the of Proceedings rnaluossand leucopsis Branta ora fAinBiology, Avian of Journal 19) -.doi: 1-8. (1897), adoko the of Handbook 91) 589-603. (9-10), 72) 57-62. (7725), id fthe of Birds 5) 1-15. (54), pink- Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. asn,J . ars . ono,C . osn .K . hlo,L . eto,D .(2007). A. D. Westcot, & A., L. Shilton, A., K. S. discovery. Robson, novel N., A C. foxes: gull: 006-0639-1. flying Johnson, kelp by the A., dispersal by Cairns, Bryophyte vegetation G., Antarctic J. of Parsons, Spread Peter, C., (2018). Braun, I. O., Kozeretska, regions. Mustafa, & 2274-9. Antarctic R., maritime V., Ochyra, two J., Kunakh, of Esefeld, Comparison J., Convey, M., Smykla, Veselski, & U., P., Convey, R., H. A., Ochyra, Rozhok, dispersal. I., A., plant Parnikoza, Rozhok, in role V., possible Kunakh, its 10.1007/s00300-012-1212-5. and I., Antarctica) of Kozeretska, (maritime Hemisphere. Use V., area Southern Ivanets, the (2012). I., in P. Dykyy, vehicle dispersal I., long-distance Parnikoza, a 10.1126/science.1095210. as doi: Wind 1144-1147. (5674), (2004). J. Navarino Isla Munoz, la de Chile. altoandina ferns zona Hornos, la aquatic de en musgos of Cabo 66432013000200011. vascular Endozoochory Biosfera Flora de (2013). (2018). Reserva L. J. Cavieres, (55oS), & A. R., Green, Rozzi, M., & Mendez, A., Europe. central V. in Molnar, mallards by O., angiosperms Vincze, and B., strategies. Vizi, life-history migrant and A., transequatorial Lovas-Kiss, biology of Reproductive Rozzi, plumage J., (1997). the Liebezeit, E. in J., R. Bety, diaspores Longton, F., bryophyte J. of Lamarre, species evidence S., First on C. Elphick, influences (2014). E., birds. B. change Qian, Goffinet, H., Climate & Goussel, E., R., (2008). Behling, R., M. L. Philipp, Lewis, & R., 10.1016/S0065-2504(07)00004-9. Lundgren, Greenland. high-Arctic doi: H., in H. distributions and Bruun, evi- interrelationships R., First D. (2011). R. Klein, Rozzi, & of F., case Massardo, The Chile. S., Splachnaceae: 0317. Horn, S. Hemisphere Cape Nihei, Southern Biosphere a A., by Reserve R. attraction Raguso, insect P., of Marino, dence spores B., fern Goffinet, (2003). of J., D. Jofre, Dispersal Beadle, (2019). & P., R. Burke, Heleno, A., & Jaramillo, A., Traveset, M., finches. Nogales, Galapagos M., by J. Olesen, S., 412. Hervias-Parejo, pp. Mifflin. (1986). Houghton T. and London: Prater, & micropropagation . J., vitro Marchant, In P., Hayman, (2006). J. moss 10.1007/s10535-006-0047-8. Rodriguez-Oubina, endangered 445. doi: & pp. the 339-345. J., Magallanes. of lupa. at de Reinoso, conservation con Universidad Ecoturismo R., available long-term Ediciones Hornos: Mallon, Chile: and M. de M.L., Arenas, Cabo Punta 2020 J. Gonzalez, (2012). del & miniatura F. May In Press en Massardo, Texas 14 Bosques North & of W., lens. updated University 4-11. Buck, hand a L., Last with Lewis, tourism R., Chapt. Rozzi, Bryologists. B., Goffinet, of vectors. Association dispersal International Vegetative < the strategies: and Adaptive sity (2017b). Glime, J.M. Glime, http://digitalcommons.mtu.edu/bryophyte-ecology/ er,2 PeerJ, roht ecology Bryophyte 44 o:10.7717/peerj.424. doi: e424. , ecapi antarctica Deschampsia ora fOntooy 160 Ornithology, of Journal Vl ,p.14) bo pnoe yMcia ehooia Univer- Technological Michigan by sponsored Ebook 1-42). pp. 1, (Vol. oaHedwigia,92 Nova o etbidn ytekl uli h retn Islands Argentine the in gull kelp the by building nest for id fChile of Birds oa ilg,41 Biology, Polar ora fEooy 00, Ecology, of Journal hrbrs nietfiaingiet aeso h world the of to guide identification An Shorebirds: pahu ampullaceum Splachnum 3,8183 o:10.1007/s10336-019-01656-9. doi: 831-833. (3), aaaBtnc,70 Botanica, Gayana 8 > Acse:Jl 3 2020). 23, July (Accessed: . eooi,152 Oecologia, 3,3736 o:10.1127/0029-5035/2011/0092- doi: 317-326. (3), odn hitpe em p 288. pp. Helm. Christopher London: . dacsi clgclRsac,40 Research, Ecological in Advances 6,14-15 o:10.1007/s00300-018- doi: 1143-1155. (6), oa ilg,35 Biology, Polar iitr oet fCp on Eco- Horn: Cape of forests Miniature -0 o:10.1111/1365-2745.12913. doi: 1-10. dacsi rooy 6 Bryology, in Advances 1,1214 o:10.1007/s00442- doi: 112-114. (1), 2,3733 doi:10.4067/s0717- 337-343. (2), . ilgaPatrm 50 Plantarum, Biologia alradubyi Tayloria 1) 7315.doi: 1753-1758. (11), etn Texas: Denton, rt.i the in Broth. cec,304 Science, 65-101. , 81-100. , (3), Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. hn,L,M,P,Cu -. u,&P-.(03.Tremdso sxa erdcini h moss the in reproduction asexual of modes Three (2003). PP-H. & But, L-M., Chu, albidum and P., Octoblepharum perfect Ma, with species L., between Zhang, comparison A released? spores peristomes. moss are specialized conditions humidity which bryophyte Under Heden large A., of con- Vanderpoorten, Endozoochory F., (2017). and J. Zanatta, A. ecology, Green, & evolution, A., waterbirds. D. Integrating by Callaghan, fragments A., conservatism: Lovas-Kiss, M., Niche D. Wilkinson, (2005). H. C. 10.1146/annurev.ecolsys.36.102803.095431. Graham, biology. & servation J., change. J. climate Wiens, from risk extinction Accelerating (2015). 10.1126/science.aaa4984. C. Seed (2020). seasonality. M. L. and Urban, Maltchik, species & A., bird Lovas-Kiss, on C., Stenert, 10.1111/fwb.13615. depends V., doi: waterfowl Weber, P., neotropical Hoffmann, by In J., dispersal biology: A. Green, conservation G., Bryophyte Europe. G. Silva, from (2014). bryophytes A. of conservation Sabovljevic, targeted situ & ex of https://doi.org/10.1080/11263504.2014.949328 the J., Evidence to Pantovic, approach (2020). M., vitro Vujicic, E. America. M., J. Jim´enez, South Sabovljevic, B., sub-Antarctic Goffinet, in R., birds Mackenzie, by 10.1111/aec.12858. mosses M., Russell, of lenses: Robertson, K., conservation consumption J., T. biodiversity M. N. Changing Arroyo, America. (2008). Russo, South J., B. southern Silander, J. of Callicott, F., flora & 6 Massardo, non-vascular Environment, A., Subantarctic W., L. the Buck, Cavieres, from B., Insights B., D. Goffinet, C. B. J., Anderson, Mishler, J. S., & Armesto, L., R., N. Rozzi, Cleavitt, R., of Dispersal L. (1961). Stark, W. V. P., upland L., Proctor, Alpert, an Bernad, J., of J., A. pattern Calvo, Wood, migration review. G., 2745(2007)110[595:dibar]2.0.co;2. J., the A bryophytes: into M. Castresana, in insights Oliver, O., Desiccation-tolerance First F., (2007). P. C. Rojas, (2015). M. K. A., Proctor, Putz, Leiss, & P., O., J. N. ( Isacch, Maceira, goose S., D., P. S. J. Munoz, Pon, J., Pedrana, al :Smayo roht eeeainrslsfrec idfcladwl ossample moss wild and fecal bird each for results treatment. regeneration each goose bryophyte in upland of the Summary by 1: dispersal Table bryophyte of mechanism birds. endozoochory sub-Antarctic possible ( by the endozoochory of bryophyte illustration native Schematic of bryophyte the diagram from treatment Schematic soil 2: peat ( the Figure goose in regenerated upland that an bed from moss a extracted and fragments shoots green of fragments. image bryophyte Detailed from regeneration Moss 1: Figure Tables & Figures hopaapicta Chloephaga hopaapicta Chloephaga 3,3-3.di 10.1890/070020. doi: (3),131-137. n h ht-ele ednp ( seedsnipe white-bellied the and ) nulRve fEooy vlto,adSseais 36 Systematics, and Evolution, Ecology, of Review Annual clg n vlto,00 Evolution, and Ecology . rpoai,Bylge 38 Bryologie, Cryptogamie, ae nstlietracking. satellite on based ) ora fBylg,25 Bryology, of Journal Riella s . oaso,V,Pt˜o . L Pati˜no, J., V., Johansson, L., ¨ as, prsb waterfowl. by spores hopaapicta Chloephaga -.di 10.1002/ece3.4579. doi: 1-8. , 3,1519 o:10.1179/037366803235001751. doi: 175-179. (3), 9 2,2328 o:10.7872/cryb/v38.iss2.2017.223. doi: 223-228. (2), tai malouinus Attagis Ornitolog´ıa 26 Neotropical, h rooit 110 Bryologist, The rooit 64 Bryologist, ea sample. fecal ) uta clg,45 Ecology, Austral ln Byosistems Plant ). cec,348 Science, nel . yadr .(2018). K. Hylander, & N., ¨ onnell, 1,5-1 o:10.2307/3240925. doi: 58-61. (1), 4,5561 o:10.1639/0007- doi: 595-621. (4), rswtrBooy 00 Biology, Freshwater rnir nEooyadthe and Ecology in Frontiers 245-253. , 63) 7-7.doi: 571-573. (6234), 1,5959 doi: 519-539. (1), , 3,3943 doi: 399-403. (3), 148 4,857-868. (4), 1-11. , Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. us n ihe oeto o drsigpoetrltdiqiis n air ieafrtedetailed the for Videla Javiera Nicholas and data, of inquiries, collection project-related the addressing in assistance for discus- for pictures. Robertson Sch fruitful Gillum Elke Michael and Taylor thank advice and and Polanco for to Russo Jonilee Lewis like support, Lily would logistic and we Additionally, Goffinet project Stu- research. Bernard for bryophyte to Experience on grateful N Research sions are International CONICYT-PAI We Magallanes. - project Foundation of the University Science National 1658651), the (NSF-IRES by dents funded was data. research the manuscript. This analyzed the R.M. of editing manuscript. and the writing Acknowledgements drafted conceptualization, and the data, to contributed the authors collected All experiment, the executed X.A.L. Contributions Author interests. competing no repository. declare Dryad authors the The in available publicly is study Interests present Competing the in used data raw The Accessibility Data tetragonum C. Conostomum: nus ybl n coysfrsmlsindicate samples for acronyms and Symbols chamber. growth a in substrates of types two and regimes light controlled under grown ( ) mosses collected wild of fragments ( goose ( upland mosses from and recovered feces from fragments for results Growth wieblidseedsnipe); (white-bellied OSTTL()3 100 33 (%) TOTAL MOSS Conostomum Polytrichum Syntrichia Agar-Gamborg Attagis Soil Peat Attagis Attagis Attagis Chloephaga Chloephaga Chloephaga Chloephaga Sample hopaapicta Chloephaga ossml;+ rwh :n rwh A osml nta combination. that in sample no NA: growth; no -: growth; +: sample; moss ytiha .robusta S. Syntrichia: ee - - - 0 + NA 50 - (%) TOTAL 3 feces 2 feces 1 feces ytiharobusta Syntrichia ee - - - 0 + + 100 + (%) TOTAL 3 feces 2 feces 1 feces Chloephaga n ht-ele ednp ( seedsnipe white-bellied and ) N 10 + + + - + - 1) eeeainpeec n bec ffragments of absence and presence Regeneration =17). , ossample; moss oyrcu strictum Polytrichum : ° .picta C. 9719 h nvriyo ot ea and Texas North of University the 79170119, oyrcu:P strictum P. Polytrichum: uln goose); (upland tlradTmr otdrfor Contador Tamara and uttler ¨ and , tai malouinus Attagis ootmmtetragonum Conostomum tai:A maloui- A. Attagis: ossample; moss ee and feces ) Posted on Authorea 2 Mar 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161469488.80502225/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. 11