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Hanna-Kaisa Lakka JYU DISSERTATIONS 295 Hanna-Kaisa Lakka Environmental Changes in Arctic Freshwaters The Response of Indicator Species to Global Warming and Acidification in the Arctic JYU DISSERTATIONS 295 Hanna-Kaisa Lakka Environmental Changes in Arctic Freshwaters The Response of Indicator Species to Global Warming and Acidification in the Arctic Esitetään Jyväskylän yliopiston matemaattis-luonnontieteellisen tiedekunnan suostumuksella julkisesti tarkastettavaksi lokakuun 23. päivänä 2020 kello 12. Academic dissertation to be publicly discussed, by permission of the Faculty of Mathematics and Science of the University of Jyväskylä, on October 23, at 12 o’clock noon. JYVÄSKYLÄ 2020 Editors Jari Haimi Department of Biological and Environmental Science, University of Jyväskylä Päivi Vuorio Open Science Centre, University of Jyväskylä “The Arctic region is one of the most vulnerable systems on our planet, if and when global warming continues.” “I have said this before: if we lose the Arctic, we lose the globe.” – Sauli Niinistö, President of the Republic of Finland, The Arctic Biodiversity Congress, Rovaniemi, Finland, 9th October 2018. Copyright © 2020, by University of Jyväskylä Permanent link to this publication: http://urn.fi/URN:ISBN:978-951-39-8326-0 ISBN 978-951-39-8326-0 (PDF) URN:ISBN:978-951-39-8326-0 ISSN 2489-9003 ABSTRACT Lakka, Hanna-Kaisa Environmental changes in Arctic freshwaters: The response of indicator species to global warming and acidification in the Arctic Jyväskylä: University of Jyväskylä, 2020, 51 p. (JYU Dissertations ISSN 2489-9003; 295) ISBN 978-951-39-8326-0 (PDF) Diss. In the Arctic region, global warming progresses quickly. Arctic freshwater indicator species were used as model organisms to study environmental change. The objective of the thesis was to evaluate the severity of warming and acidification for fauna in Arctic freshwaters. Freshwater acidification was confirmed in northern Finland. Precipitation was acidic and water pH was an alarmingly low 4.9 in small ponds. This measured low pH was also the mean precipitation pH in northern Finland. Increasing acidic precipitation together with warming are major threats to acid sensitive Arctic species especially in ponds in northern Finland, due to their low water volume and their limited buffering capacity. The new method to calculate climate risk for selected species predicts physiologically harmful temperatures in Lapland. The Arctic indicator species have not experienced these predicted high temperatures during the last 60-years. The findings highlight that the lethal effect of heat stress caused by global changes need to be studied in a species- perspective to allow predictions for Arctic populations. Behavioural patterns, such as sexual reproduction, cannibalism and colonisation can help the species to survive in the severe and fast changing Arctic. The result shows that these beneficial behavioural patterns were present in the Arctic tadpole shrimp populations in the High Arctic Svalbard. Colonisation after the Last Glacial Maximum (LGM) happened from refugia in the South-West resulting in diverse invertebrate fauna in Arctic archipelagos Svalbard, Franz Josef Land and Novaya Zemlya. Together, the results highlight populations where the possible management actions should be carried out. Keywords: Acidification; Biogeography; Branchiopoda; Climate change; Climate risk calculation; pH. Hanna-Kaisa Lakka, University of Jyväskylä, Department of Biological and Environmental Science, P.O. Box 35, FI-40014 University of Jyväskylä, Finland TIIVISTELMÄ Lakka, Hanna-Kaisa Ympäristönmuutos arktisissa vesissä: Indikaattorilajien vaste ilmaston lämpenemiseen ja happamoitumiseen arktisella alueella Jyväskylä: Jyväskylän yliopisto, 2020, 51 S. (JYU Dissertations ISSN 2489-9003; 295) ISBN 978-951-39-8326-0 (PDF) Yhteenveto: Ympäristönmuutos arktisissa järvissä ja lammissa Arktinen alue lämpenee kiihtyvällä vauhdilla. Arktisia makeanveden indikaattorilajeja käytettiin tässä tutkimuksessa malliorganismeina ympäristönmuutosten tutkimuksessa. Työn tavoitteena oli arvioida lämpenemisen ja happamoitumisen vaikutuksia arktisten sisävesien eliöstöön Pohjois-Suomen lammissa mitattiin hälyttävän hapanta vettä (pH 4,9), mikä johtui haitallisen happamasta sadevedestä. Sademäärän voimakas lisääntyminen yhdessä lämpenemisen kanssa on merkittävä uhka happamoitumista huonosti sietäville arktisille lajeille. Erityisen huono tilanne on lammissa johtuen niiden pienestä vesimäärästä ja rajallisesta puskurikapasiteetista. Uusi menetelmä ilmastoriskin laskemiseksi arktisille lajeille ennustaa fysiologisesti haitallisia lämpötiloja Lapissa. Arktisen alueen indikaattorilajit eivät ole kokeneet näitä ennustettuja korkeita lämpötiloja viimeisten 60 vuoden aikana. Tulokset korostavat, että maailmanlaajuisten muutosten aiheuttamaa lämpöstressin kuolleisuutta lisäävää vaikutusta on tutkittava lajien fysiologisesta näkökulmasta, jotta arktisten populaatioiden häviämisriskin ennustaminen olisi mahdollista. Seksuaalinen lisääntyminen, kannibalismi ja kolonisaatio voivat auttaa lajeja selviytymään arktisten alueiden ankarissa ja nopeasti muuttuvissa oloissa. Tulokset osoittavat, että nämä selviytymisstrategiat olivat käytössä paljakkakilpiäisten populaatioissa Huippuvuorilla. Immigraatio, eli tulomuutto, tapahtui viimeisen jääkauden jälkeen etelä-länsi suunasta Huippuvuorten, Franz Josefin maan ja Novaja Zemljan saaristossa, mikä johti monimuotoiseen selkärangatonlajistoon. Tulokset osoittavat, että toimenpiteisiin olisi ryhdyttävä ainutlaatuisten arktisen eläimistön suojelemiseksi. Avainsanat: Biogeografia; Branchiopoda; Ilmastonmuutos; Ilmastoriskin laskenta; Happamoituminen; pH. Hanna-Kaisa Lakka, University of Jyväskylä, Department of Biological and Environmental Science, P.O. Box 35, FI-40014 University of Jyväskylä, Finland SAMMENDRAG (NORSK) Lakka, Hanna-Kaisa Miljøforandringer i Arktiske ferskvann: Indikatorarters respons på global oppvarming og forsuring i Arktis Jyväskylä: University of Jyväskylä, 2020, 51 p. (JYU Dissertations ISSN 2489-9003; 295) ISSN 978-951-39-8326-0 (PDF) Diss. I den arktiske regionen utvikler den globale oppvarmingen seg raskt. Her ble indikatorarter i ferskvann benyttet som modellorganismer for å studere miljøendringer. Formålet med undersøkelsen var å evaluere betydningen av oppvarming og forsuring for faunarn i arktisk ferskvann. Forsuring ble påvist i Nord-Finland (Lappland), og her hadde små dammer en alarmerende lav pH på 4,9. Gjennomsnittlig pH målt i nedbøren viste like lave verdier. Økende forsuring, sammen med oppvarming, er alvorlige trusler mot sensitive arktiske arter i slike lokaliteter i Nord-Finland. Dette skyldes i hovedsak at de har et lite vannvolum og en begrenset bufferkapasitet. Den nye metoden for å beregne mulige klimatiske skadevirkninger for utvalgte arter tilsier også fysiologiske skader relatert til høyere temperaturer i Lappland. De arktiske indikatorartene har ikke vært utsatt for slike forventede høye temperaturer i løpet av de siste 60 årene. Funnene viser at for arktiske arter må den dødelige effekten av varmestress forårsaket av globale klimaendringer, studeres på artsnivå. Adferdsmønstre, som seksuell reproduksjon, kannibalisme og kolonisering, kan hjelpe artene til å overleve i det ugjestmilde og raskt skiftende klimaet i disse områdene. Disse gunstige atferdsmønstrene ble også funnet hos skjoldkreps på Svalbard. Kolonisering etter siste istid skjedde fra refugier i sør-vest, og resulterte i en mangfoldig fauna av virvelløse dyr i kystnære strøk på Svalbard, Franz Josef Land og Novaya Zemlya. Resultatene viser klart hvor det bør settes inn tiltak for å bevare denne spesielle faunaen. Nøkkelord: Biogeografi; Branchiopoda; beregning av klimatiske endringer; forsuring; klimaendringer; pH. Hanna-Kaisa Lakka, University of Jyväskylä, Department of Biological and Environmental Science, P.O. Box 35, FI-40014 University of Jyväskylä, Finland Author´s address Hanna-Kaisa Lakka Department of Biological and Environmental Science P.O. Box 35 FI-40014 University of Jyväskylä, Finland [email protected] Supervisors Senior Lecturer Emily Knott Department of Biological and Environmental Science P.O. Box 35 FI-40014 University of Jyväskylä, Finland Professor Stephen Coulson ArtDatabanken Box 7007 750 07 Uppsala, Sweden Senior Lecturer Panu Halme Department of Biological and Environmental Science P.O. Box 35 FI-40014 University of Jyväskylä, Finland Reviewers Professor Milla Rautio Département des sciences fondamentales 555, boulevard de l’Université, Chicoutimi (Québec) G7H 2B1 Université du Québec à Chicoutimi, Canada Professor Timo Muotka Ecology and Genetics Research Unit PL 8000 FI-90014 University of Oulu, Finland Opponent Professor Leopold Füreder Department of Ecology Technikerstrasse 25 A-6020 Innsbruck University of Innsbruck, Austria CONTENTS ABSTRACT TIIVISTELMÄ SAMMENDRAG (Norsk) LIST OF ORGINAL PUBLICATIONS ABBREVATIONS AND TERMS PREFACE 1 INTRODUCTION .............................................................................................. 13 1.1 Arctic environmental change and cold adapted species ..................... 13 1.2 Theoretical framework ............................................................................. 14 1.2.1 Environmental change in the Arctic freshwaters ..................... 14 1.2.2 Adaptation to changing conditions ............................................ 14 1.3 Contextual background ........................................................................... 17 1.3.1 Acidification (Paper
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