BIOLOGY LETTERS royalsocietypublishing.org/journal/rsbl Research a 2 Cite this ate: Krabi Mine Wines 6, Soies MA, Waters GM, 2021 Coxsal regions ofthe orthem datactic Penis ae hey foe getoo popations. Bok ter. WP 0700708, hms. door. oats 2020.0708 Fei § Ceber 2020, Accepted: 4 January 2021 Subject dreas: cxclogy, erroameial sence rps popoa, dinate ange, tonge expansion, satelite telemetry “huthor for corespondence: Malgorata Korzat-Abshie malt mlagibh wow pl ecrenc supplementary mate avalable nie at hp co/O004 isha. cae, THE ROYAL SOCIETY PUBLISHING. ene Coastal regions of the northern Antarctic Peninsula are key for gentoo populations ‘Malgorzata Korczak-Abshire’, Jefferson T. Hinke?, Gennadi Milinevsky?*, Marina A. Juares*® and George M. Watters? “intite of Bachem) and Bhi, Posh Madey of Scr, 02-206 Warm, Peland atic Ease. Beech Cte, Sauies Feris Shee Cine, Rana Marne Fetes See, Nitcnl Gauci and Asoxphsc Hina, Ls Jl, CK S300, USA “ppm ef aarp Pipes Grp aol Arak See Cte sw, 0, inane ‘phys aca, aes Shewherto sonal Uniersty fyi, Kye OBOE, Urine “aparently, Ha, Bees As AC, Anger "ron Scena el earth Cees (COW, Ca toms ue Ae, CHOSE Agent © WA, ooo 765M, obo 0002 aan 1g Southern Ocean ecosystems are rapidly changing. dow to-climate variability. ‘An apparent beneficiary of such change in the western Antarctic Peninsula (WAP) i the gctitoo penguin Pygusclisprpir, whieh has increased its popti- lation size and expanded its range southward in the last 20 years. To bether lundersiand how this species has respondast to large-scale changes, we tracked individuals during the nos-breeding winter period from five colo- nics across the latitudinal range af breeding stos in the WAP including from a recenily establishes! colony. Results hightight latitudinal grastients in movement; strong associations with shallow, coastal habilats along, the ‘enti Antarctic Peninsula; and movements that are inddepensent of, yet con= strained by, sea ioe, It is clear that coastal habitats essential to. gentoo penguins during the breeding season are similarly critical during winter. Langer movements of birds from northern colonies in the WAP further suggest that Ieap-frog migration may influence colonization events by facl- ating nestarea prospecting and use of new haul-out sites. Chur results support efforts to develop a marine protected area around the WAP. Winter habitats used by gentoo penguins eutline high priority areas for Improving the management of the spatio-temporally concentrated. kril (Explaasia superba) fishery that operates in this region during, winter 1. Introduction ‘Climate change fundamentally alters the structure and funetion of marine eco- systems by medlifying ocean productivity. altering food-web dysiamics and shifting species distributions [1], Polar ecosystems are expecially sonitive to elimate perturbations because they ate largely structured by the seasonal dynamics of sea ice [21 which have exhibited trends in extent and duration in both hemispheres [3]. An area of particular concern is the western Antarctic Peninsula (WAP), where increases in air andl sex-surface temperatures over the Inst 40 years have reduced mgional seat ioe extent and duration [4J. Such phys ‘cal perturbations are associated with changes in the distribution, abundance and survival of several species in the WAPecosystery [5]. Further compounding: risk to this ecosystem is the expanding fishery for Antarctic krill (Euphesir superto} {671 the largest, by mass, in the Southern Ocean [8 (© A The At Pale by he Ral Sd der he em le Cr Canis tin Lerse hp:hmatiearmens pers, which poms uated ee, pad he cig eran sec am enDownloaded from htips://royalsocietypublishing.org/ on 08 February 2022 ‘able Stir (CAS, Cera Cove (CIR) and the Aagentine ands (AS), All tags wre released Between 4 Febrary and 29 March 2017 er a lente tattee adit up Su OM oy 5k a as 080 6 oa 10.98 ou BS. 125 58 ene as 30.80 rs on 0.98 a6 Seobinds are important indicators of exnsystem status ancl ane among the species impacted by climate change and fish~ ries (729-111, Changes in population sizes and phenologics due to envimonmental variation in the Southern Hemisphere ane evident [9.12-14], For example, in the WAR, populations of ioo-dependent Adélie penguins (Pygosclis alin) and iee-olerant chinstrap penguins (P. mfarcticus have declined [1215] while the abundance and range of ioe-avoiding _gentoo penguins (P. pypua) have increased [121 Notably, range expansion and rapid population growth af gentoo pen- uins is occurring, at the southern margin of theit brvesting range, where at least seven nevily established colnnies have been identified in last 20 years (electronic supplementary material, figure 51) [16 Despite divergent population trends among the pygoscelid penguins, all three species have been affected by rooent krill fishing during, the non= Dressing period (7 hereafter winter. Thus. althensgh _gencrally considered to be climate ‘winners’ [17M the risks te gentoo. penguin populations should be farther assessed te better inform conservation and management actions ‘Acrans the WAR. genino penguins typically forage within 20 km of breceling sites during the austral summer (18-22) During winter, gentoos ane not constrained by’ the need to provision chicks on can undertake longer range movements. Prior tracking stuclos from the South Shetland! Istands (1821) stiggested Winkor mavements wp to I times farther than ‘during summer. Such dispersal to distant foraging. areas ix the primary mechanism by which range expansion could jccur 23}, but a lack of tracking data fram colonies through cvot the WAP Timits understanding of howe this seabird distributes during the winter and whether there is variation im movement among colonies. Given rapidly changing cavironmental and anthropogenic drivers in the WAT, identi fying winter movements and patterns of habitat use by genio penguins aw also useful for assessing. population satus and risks to the species We therfore trackd the Winter movements of gentvo penguins from five colonics -acrors the latitulinal range ofthis species in the WA inciul- ing from a recently established colony [24] near the southern limit of the species’ range. 2. Material and methods We wacked 10 Asdgling and 65 post moult adult gentoo pomguins from five colonies across the tnitucinal range of breeding colonics in the WAP from Febuary 2017 through January 2018 Mable 1; electronic. supplemeniary’ material, figure Sto). We awd Sirtrack Kiwisat MOK2G72A, satelite duration (6 asim =) 6 105 (rb-148) 109 (32-229), a 19 s-157) 28 (20-186) 9 100 45-194) 151 (ar-2st) 10 92 (16-220) 63 (28-141), u 196 (57-308) 2 (2-190) $ 30 (12-78) 8-13) $ 85) 680-112) trasaithers 64) = 27 = 17 man, 34 gb te track fledgling.s amd Wikd- lle Compters Spt275 satelite transmis (17 I mm, 58) fo tack adults All binds were captured on beaches and the traneaiters were afixed ta hack feathers using phe and cle tes [25] Tronsmiter were cheat to tons aly from 12:80 to 18D UTC, coresponding. to daylight howe shen bir should be foraging, We prcesed raw location estimates by removing four adult epayments tat wore tracks fess than 7 ays, ant al ero rs hatin estinstes indicated by quality codes oF unspent slipse cram Nes, we applied sped ter [26 Assiming ® conservative swim spe of 25 mvs. Remaining tracks wore smoothed with 3 atespace mod [27] sing, the 28 package ‘crow [29 Male Rts Were used fe eneate 100 alleative tock foreach deployment, vith leations ext crated every 2h. Allerative racks were pawl and mapped to hexagonal polygons with cents spaced 15 km apart (area = 87 kr’) to estimate habitat utilization distributions CHUDs) ming the R package ‘cre? OL TA spi scale ppeonimates daly foraging ranges by gentoo penguins dig the breeding sone (25). Weewed several ysl wars to compare the habitats tom by binds fom ferent colon. We extracted bution eps along each track from the ETOPO! dataet (31) We xt mmitad ditince to the nestest point of find using the ‘Wri simp database im the [281 package ‘maptosl [2 Fo amine near nal-time experience of sey surface temperatures {S5TP and sew ine concentrations (SIC, exprowed an per cet soned, we matched ta pstion eskimutes with daily STs from the mailtscalewlta-high olan SST data eave om 4 Tham grid 35] and iy SIC fen the EUMETSAT Ossi GiadSen lee Settle Appian Face oft rion a ative 10m grt on gi We ase! the Talay honest sgniicantditence (HSD) ta Vealily ‘oderplesd difreion. by rveniet sod lint mnbsad-ticts models ting the K [2] package “met” [3] to identify colony trvel illrences in physical tabi waiables We ted spams cts fe cach physien! vanible and inckated meth an a ford oflec inal ert to secon for potential seasonal trends. Inivaduals were tated rancor tis, 3, Results Positions (N 29119) of fledging and adult penguins wer, opectively, ported for an averge of 29 days (range: 9-86 days; table 1) and 100 days Gange: 16-306 days), Maxirmam dlistanocs fom tagging sites varied by colony Mable 1D. Gentoo penguins originating from the northern exge of their range in the WAP dispersed farther and with sipnifi- cantly greater shifts to the south (Tukey 15D Fy 128, Tagging locations and mean deployment dations (range in parentheses) Tagging latins were 2 ons. ump (URE), Stanger Foot SS), Cape BOLOOOE SAL HT ye RUA EaoremsrMALroOSeing.org/ isocietypubl ‘royal: n https: 04 4 * 100, 104 6 lautade (89 03 ote W) Figure 1. ol Mean and 95% ronfdenceitenals fr psec shits for bids om the Arends ark lel, Gena Coe (pul ape Shi rf igh ue, Sanger Poet (ange) ass Rump (yelio). 2) HLDs for al tated bids. lon cons ae ica with ccs cour to atc pane (oh 36 p<0.01) than adults tagged at colonies further southwest (figure Io). Longitudinal movements were not different across colonics (figure la, Tukey HSD Fag 19, p= th10) Given high overlap of fledglings and adults (clectranic supplementary material, figure 82), all tracks were pooled for further analyses. The physical habitats encountered during winter wore largely similar acrocs colonice (electronic supplementary material, figure $3) The modol-predicted distance from shore averaged 6.4 km with little variation among, colonies (table 29. Near shore areas corresponded to shallow habitats that averages! 42 m dewp across all colonies (able 2), nating, that birds from Capo Shirwlf and the Argentine Idlands had the shallowest habitats. Binds dom Cape Shirreff, with ‘more northern distribution for much of the wintes, typically encountered warmer water than birds from other colonies able 2). Responses tor the distribution of SIC were not colony-speciic when ice was encountered (able 2}, and all, bores usually occupied foe-fe waters (Fig The aggregate HUD for all tracks during winter (figure I demonstrates an affinity for coastal arco. Winter HUDSs were concentrated near tagging sites, araund islands inthe Bransficlel Sait, and along the enti margin of the WAP from the “Angentine lands to the tip of the Peninsula (Fggure 18). Over- ES lap of colonytevel HUDS (figure 2) was common, This was particularly evident for bids from Stranger Point and. Lions Rump, swhich overlapped extensively around King Goorgo/25 de Mayo Band and along the northern tip of the Peninsula from 59°WY to 57° W. Similarly, birds: from Cape Shireff and Cierva Cove owelapped extensively along, the contal WAP from 63° to S8°W. The HUDS far gentoo penguias from the Argentine Islands, the mest southern colony, were largely Isola from other eokmies. Differences in calony-level movements from February te Apeil occurred prine to the presence of sea ice near any colony (figure 2), demonstrating that sea ise was not the main driver of colony-sperfic, ever-winter dispersal patterns. Nonetheless, the evolution of dense sen ice (SIC » 50) did affect the distribution of the bitas during, winter, For example, the HLUDs of birds from Lions Rump contracted between June and July when high SIC blankie! the eastern Bransficld Strait, Likewise, birds from the Argentine Islands shifted northeast into the Gerlache Strait coincident with the expan- sion of dense sca ice in waters south of Anvers Island from, ‘August to October (Figure 2), However, SIC<50% did not preclude gentoo occupation of those areas (figure 2), . Discussion We seport clone latitinal gradi in winter movements of gentoo penguine from five enlonies of varying, population sizes ans trends along the WAP, Dispersal distances were lager for bieds from northern colonies than. fen southern colonics. The HUDs of birds fom different colonics over: lapped in the relatively shallows, 4eefoe coastal margin of the WAP. Of particular importance were the coastal ragions fof the Antarctic Peninsula from the Argentine Inds to the Lip f the Antarctic Poninsuls. Our muilt-colony tracking study demonstrates that, in the WAP. the coastal habita sential to brecling. gentoo penguins during summer ie similarly critical during winter, The observed graicnt in rmavement patterns suggests that birds from noxtheen eo figs are immigrant sources forthe current range expansion ‘of this species Marine top predators are often expected to. change foraging behaviours, movement pattcms and atsoa dist butions 135) in response to elimate-driven changes in prey distribution (9 rather than to direct changes in theie physical ‘vironment Inthe WAP, however, sca ioe dynamics can fu damentally aller the availability of foraging, habitats, As an Yointolerant species [36], we expectest the movement pat= tems of gentoo penguins to be driven ‘by avoidance of developing sa ice. However, all Jongedistance dispersal observed! here wats initiated before the presence of sea ie at study colonics. Indevd, wintcr movements of geno pew gins fn the Faldand/Matvinas Klan, where se foe doce fot exist, wens even farther than observed in the WAP [371 Thus, sea ice is not the main driver of ditferences in. colony speviic dispersal patterns acnong genioo penguins. As sex ice extent and duration in the WAP are expect to dectine under most climate-change sccnaries [81 the observed lati- tudinal gradients in movement and the afilation of gosto ppenuguins with coastal regions along the WAP may be ‘expostel in the Future Dispersal of seabirds from bresing sites to winter fort ging amas must be driven by niiable availablity of pry. INOUE {ZA U2] IB paKNRWINe| Eun foxysee0snwy o sine sic S04 sic a prumal in fos) aouoncoe <2 Bat yo ‘on 08 February 2022 cietypublish Figure 2. Monthy etents of colom-pectic MUD ovetad on menthiy SC. Calon locations ae denied by fled cles. Cony locations and their HDS ze coloured as in figure 1 Table 2. Mixeeefea mode pedicons foe the fuedrefects (95% Cl) of colony agin on distance fom shor, depth, sex-surface temperatures (SSN and sea ice conentars (Cfo al tracked penguins, colony depth (m) 0 EC we 52 82-4 103 7e-13 1 05-51) rs 40 01-98) oan) 19 07-125, BBN as 43 24-100) 14 149) 142 (135-150) 4 an48) on 43 05-80 BB 03 54-1. Fat) Ns 79 (2-118) 8 (01-2) 106 77-135) sas Gentoo penguins typically echibit benthic and pelagic fora- invertebrates and fishes that can vary by location [36] sing dives (les than 150 m depths) im coastal rons [20] Within the WAR, shorter dispersal ranges of gomtoos from with dicts of crustaceans (mainly Antarctic krill), other southern colonies relative to northern colonies suggestDownloaded from htips://royalsocietypublishing.org/ on 08 February 2022 greater food availablity along the margins of the Antarctic Peninsula molative to the south Shetland Islands during winter: The continental shelves of the WAP. over which the HUDs were concentrated, are known to harbour high krill densities during winter (eg, (9240), consistent with expes- tations of a southwand contraction of krill distributions over lagger scales [41]. These shallow, costal areas also: provide gentoo penguins with sevess to benthic and demersal resoutoes, which are suspected to be impartant components of thet winter dots [42] The tracking sata reported here suggest source popu lations for the ongoing ange expansion af gentoo penguins inthe WAP. Genotic analysis indicates that basin-scake disper- ‘sal and colonization events are rave for this species, and the Polar Front isan effective boundary between sub-AAntarctic and Antarctic populations [23,43]. At smafler spatial scales, gentoo penguins are well-kaown colonizers of new brevsting territory and quickly take advantage of ice-free breeing space [36]. Recent colonization events have been attributes, te emigzation from colonies at the southwestern edge of this species’ mange [1244], Hawewes, longer distance mave- ments of Birds from northern colonies. suggest that rare dispersal events to the south eoukl be instigated by birds from northem colonics. Such a leap-érog. migration strategy has been reported for other scabinds {45}. Differential move- ment patterns of birds from different breeding colonies may be driven by variation in prey availability at breeding, colo- nies but shared foraging habitat preferences that favour gradients in directed movement 145,461. While we cannot test this hypothesis dirty, the observed latitudinal gradient in the movement of gentoo penguins is consistent with a leap-frog strategy. In particular, larger scale movements ‘during the winter provide an opportunity to prospect new aul-out sites anc nesting areas that would support colonization events, The eects of ongoing climate change in the WAP region are dificil to predict 471, Hamrever,continaod reaction in ‘SIC curing winter may be advantageous for coastal predators ‘and an increase in the availabilty of feefive foraging habitats muy facilitate southwaret expansion in the beoesing range of gentoo penguins. Nonetheless, such expectations may be tempered by increases tv the biomass of salps [48] with com ‘comitant declines in krill biomass due to recruitment falas FAIL local increases in the abundances of cetaceans (potential ‘competitors with gentoo penguins for food) [49] and contin= usd growth of the krill fishing industry [6,8 Our study suggests that, im the WAR.a latitudinal gradient im the move- ment of gentoo penguins during winter might be a key to the dynamic of how gentoo populations cope with large-scale changes in the ecosystem. thio. The msenrch was permite! under US. Antarctic Conservation ‘Act Permits (permit no. 2017-012), the Posh Permit Acthity FB: PAS (permit no. /20%6; permit no. 15/2016) ate the Agente Dimccion Nacional det Antirtion Environmental Office (permit mn. 207-10 arnt 2017-489, Feld protons were appre by the Un sity of Cabloenia San Diggs fnttutioral Animal Cae aed Use (Comment: 05480, Da acesity. Data for this article are availble from the Dryad Digie tal Reponitory at: gs dion! VO fdryind SpS020 150] ates onbutons. MCA, EH, ans GMLW. concealed the seady. }THL, Ma}, MICA an GML curated the data, NECA and [TTH. confuxted formal analysis JH, MAD, MIKA. “and GMM, ocquire funing. EH. SUA MLR. rotor GM. nd GM, b= proget MIKA. ancl [TH wrote the orginal drt A, LTH, MAJ, GM, and GMAW. reviewed and cited the Final dra, Comper ees We declare we Rave ms competing intents Fone. This study ws fence by CCAMLR CEMP Sypevil Fut, ‘Connision for the Conservation of Antarstic Marin, Heneyk Ar toneski Potish Antarctie Statiy and eatitut: Amtitioo Angee — Dinvoriin Nackoul el Antirtio and NASA MESSUREs programm, Fciommiegemenr. Data te ew ware collected with cappart of Henry: Arctowaki Polish Antirtic Station aru the Akadernik Ver= radiky Ukrainian Antarctic Station We unk the Institute ‘Aststien Argentino—Dinweis Nacional el Antirtica for supp ‘a.Gierva Cone art Stranger Point. 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