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Rangifer Tarandus Platyrhynchus) Females Subject/Study Track Ecology and Evolutionary Biology Level Month and Year Number of Pages Master’S Thesis 12.10.2020 43P

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Rangifer Tarandus Platyrhynchus) Females Subject/Study Track Ecology and Evolutionary Biology Level Month and Year Number of Pages Master’S Thesis 12.10.2020 43P Life History Effects in the Behavior of Svalbard Reindeer (Rangifer tarandus platyrhynchus) Females Master’s thesis Laura Puikkonen Faculty of Biological and Environmental Sciences Ecology and Evolutionary Biology University of Helsinki October 2020 Faculty Degree Programme Faculty of Biological and Environmental Sciences Master’s programme in ecology and evolutionary biology Author Laura Puikkonen Title Life history effects in the behavior of Svalbard reindeer (Rangifer tarandus platyrhynchus) females Subject/Study track Ecology and evolutionary biology Level Month and year Number of pages Master’s thesis 12.10.2020 43p. + references and supplements Abstract Individuals of long-lived animal species can improve their reproductive success through experience. While individual’s resources available for survival and reproduction decrease toward the end of its lifespan through senescence, terminal investment hypothesis predicts the less likely old individuals reproduce again the more they invest in their current offspring. Experience gained through a long lifespan might have an important role in changing behavior to optimize the use of resources and compensate the effects of senescence. In addition, behavioral plasticity allows animals to respond changes in their habitat within much shorter timespan than on an evolutionary timescale. Svalbard reindeer (Rangifer tarandus platyrhynchus) is a wild subspecies of reindeer. It is only found in Svalbard, a remote archipelago in the Arctic with extreme weather conditions rapidly changing due to climate change. It has been isolated at least 5000 years and adapted to a barren habitat with nearly no hunting, predation or harassment of flying insects. The objective of this study is to investigate the effect of age and a calf at heel in Svalbard reindeer females’ maternal, vigilant, and social behavior and time budget in the light of life history theory and its senescence and terminal investment hypotheses. I carried out the field work for the study in two periods in summer in Semmeldalen valley and the south-western part of Reindalen valley on the island of Spitsbergen, Svalbard. I collected behavioral data on marked individuals by instant scan sampling and focal watch methods, wrote observations down manually and later fed them into computer. In addition, I have used birth year data collected by the long-term monitoring program by the Norwegian Polar Institute. I used generalized linear mixed models to analyze the effects of age and calf at heel to the behavior of females. The main results include that young dams maintained shorter distance to their calf in July than in August, and old females were less vigilant. Younger dams and older females without calves were in smaller groups than older dams and younger females without calves. In addition, females with calves spend proportionately less time lying down than females without calves. Dams maintained a longer distance to the nearest neighbor than females without calves. Older dams spend proportionately more time feeding and in groups in August than younger dams. These results show that the age and calf at heel do play a role in the behavior of Svalbard reindeer females and the effect varies over the course of the short Arctic summer. Experience may make older females more effective mothers by optimize the use of resources for example from vigilance to feeding in a predator-free environment. On the other hand, senescence may affect the amount of energy females can spend on their calves, potentially influencing their survival. Keywords life history theory, reindeer, age, senescence, terminal investment, maternal behavior, vigilance, time budget Supervisor or supervisors Emma Vitikainen Where deposited HELDA - Digital Repository of the University of Helsinki Additional information Field work was conducted in cooperation with University Center in Svalbard (UNIS). Tiedekunta Koulutusohjelma Bio- ja ympäristötieteellinen tiedekunta Ekologian ja evoluutiobiologian maisteriohjelma Tekijä Laura Puikkonen Työn nimi Elämänhistorian vaikutus Huippuvuorten peuranaaraiden (Rangifer tarandus platyrhynchus) käyttäytymiseen Oppiaine/Opintosuunta Ekologia ja evoluutiobiologia Työn laji Aika Sivumäärä Pro gradu -tutkielma 12.10.2020 43 s. + lähdeluettelo ja liitteet Tiivistelmä Pitkäikäisten eläinlajien yksilöt voivat parantaa lisääntymismenestystään kokemuksen avulla. Samalla, kun yksilön resurssit selviytymiseen ja lisääntymiseen laskevat senesenssin vuoksi, terminaalisen panostuksen hypoteesi ennustaa vanhan yksilön käyttävän sitä enemmän resursseja jälkikasvunsa hyväksi, mitä epätodennäköisemmin kyseinen yksilö lisääntyy tulevaisuudessa. Pitkän elämänkaaren varrella kerätty kokemus saattaa olla tärkeä elementti käyttäytymisen muutoksessa, jonka avulla yksilö pystyy optimoimaan resurssiensa hyödyntämisen ja kompensoimaan senesenssin vaikutuksia. Sen lisäksi, käyttäytymisen muovautuvuus mahdollistaa eläimen sopeutumisen elinympäristön muutoksiin paljon lyhyemmällä aikavälillä kuin evolutiivinen kehitys. Huippuvuorten peura (Rangifer tarandus platyrhynchus) on peuran eli poron villi alalaji. Sitä tavataan ainoastaan Huippuvuorilla, joka on syrjäinen saaristo arktisella alueella, ja jonka äärimmäiset sääolot muuttuvat nopeasti ilmastonmuutoksen seurauksena. Alalaji on ollut eristyksissä vähintään 5000 vuoden ajan ja sopeutunut karuun elinympäristöön, sekä metsästyksen, saalistajien ja lentävien hyönteisten aiheuttaman häirinnän puuttumiseen. Tämän tutkielman tavoite on tutkia iän ja vasan vaikutusta Huippuvuorten peuranaaraiden maternaaliseen ja sosiaaliseen käyttäytymiseen, vigilanssiin ja aikabudjettiin elämänhistoriateorian sekä senesenssi ja terminaalinen panostus -hypoteesien puitteissa. Tein tutkielmaan kuuluvat kenttätyöt kahtena ajanjaksona kesällä Semmeldalenin laakossa ja Reindalenin laakson lounaisosissa Spitsbergenin saarella Huippuvuorilla. Havainnoin merkittyjen yksilöiden käyttäytymistä instant scan sampling ja focal watch - metodeilla, kirjasin havaintoni ylös käsin, ja siirsin ne myöhemmin tietokoneelle. Lisäksi hyödynsin Norwegian Polar Institute - instituutin pitkäaikaisen seurantaohjelman keräämiä tietoja peurojen syntymävuosista. Analysoin iän ja alle vieroitusikäisen vasan vaikutusta peuranaaraiden käyttäytymiseen yleistettyjen lineaaristen sekamallien avulla. Tuloksien perusteella nuoret emät ovat lähempänä vasojaan heinä- kuin elokuussa, ja vanhemmat peuranaaraat ovat vähemmän valppaita. Nuoremmat emät ja vanhemmat naaraat ilman vasoja olivat pienemmissä ryhmissä kuin vanhemmat emät ja nuoremmat naaraat. Emien etäisyys lähimpään naapuriin oli keskimäärin pidempi, kuin vasattomien naaraiden. Lisäksi naaraat ilman vasaa viettivät suhteessa enemmän aikaa maaten kuin emät. Vanhemmat emät viettivät suhteessa enemmän aikaa syöden sekä ryhmissä elokuussa, kuin nuoremmat emät. Tulokset näyttävät, että ikä ja alle vieroitusikäinen vasa vaikuttavat Huippuvuorten peuranaaraiden käyttäytymiseen, ja vaikutus vaihtelee kesän edetessä. Kokemus saattaa tehdä vanhemmista naaraista tehokkaampia emiä auttamalla niitä optimoimaan resurssien käyttöä esimerkiksi valppaudesta syömiseen saalistajista vapaassa ympäristössä. Toisaalta senesenssi saattaa vähentää vasojen hoitoon käytettävissä olevan energian määrään, potentiaalisesti vaikuttaen niiden selviytymiseen. Avainsanat elämänkaariteoria, peura, ikä, senesenssi, terminaalinen panostus, maternaalinen käyttäytyminen, valppaus, aikabudjetti Ohjaaja tai ohjaajat Emma Vitikainen Säilytyspaikka HELDA - Helsingin yliopiston digitaalinen arkisto / HELDA Muita tietoja Kenttätyö tehtiin University Center in Svalbard (UNIS) -yliopistokeskuksen avustuksella. “The differential allocation of energy to either reproduction or survival represents a major conflict with important implications for patterns of life history.” - Loison & Strand (2005) 4 Table of contents 1 Introduction ............................................................................................................................................................ 6 1.1 Life history theory: traits and trade-offs .............................................................................................. 6 1.2 Senescence and terminal investment in ungulates ............................................................................... 7 1.3 Objectives, research questions and hypotheses...................................................................................10 2 Materials and methods ..........................................................................................................................................12 2.1 Svalbard reindeer as a study species ..................................................................................................12 2.2 Study area .........................................................................................................................................14 2.3 Studied individuals ............................................................................................................................15 2.4 Observation period ............................................................................................................................16 2.5 Behavioral observations ....................................................................................................................17 2.5.1 Instantaneous scan sampling ......................................................................................................18 2.5.2 Focal watches ............................................................................................................................21
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