1. Sedimentary sea-ice proxies in the Arctic: Seasonal production, vertical export and taxonomic insights
- Author
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Luostarinen, Tiia, University of Helsinki, Faculty of Biological and Environmental Sciences, Doctoral Programme in Interdisciplinary Environmental Sciences, Helsingin yliopisto, bio- ja ympäristötieteellinen tiedekunta, Ympäristöalan tieteidenvälinen tohtoriohjelma, Helsingfors universitet, bio- och miljövetenskapliga fakulteten, Doktorandprogrammet i tvärvetenskaplig miljöforskning, Caissie, Beth, Heikkilä, Maija, and Ribeiro, Sofia
- Subjects
ympäristötieteet - Abstract
Arctic sea-ice cover is decreasing at an alarming rate with severe consequences for ecosystems across the Arctic and the climate system globally. The changes observed in the ice cover amplify global climate warming by decreasing the ice-albedo effect and may alter ocean circulation and weather patterns. The loss of sea ice is likely to increase primary production in the Arctic, leading to drastic changes in marine food webs and biodiversity at all trophic levels. For over four decades, satellite imagery has provided high-resolution spatial and temporal understanding of changing Arctic sea-ice cover. However, the paucity of any human observations of Arctic sea ice beyond a few centuries limits our understanding of the current trajectory and its implications for ecosystems and the global climate. Marine sediments provide a natural archive of past sea-ice variability over timescales from centuries to millions of years. Commonly used sedimentary sea-ice proxies are biogenic: (sub)fossil remains and biogeochemical tracers originating from micro-organisms inhabiting sea ice (sympagic), or ice-associated under-ice and ice-edge ecosystems. State-of-the-art sea-ice reconstructions typically use diatoms, resting stages (cysts) of dinoflagellates and the highly branched isoprenoid (HBI) lipids IP25, IPSO25, HBI III and HBI IV, which are synthetized by certain diatom species. Current understanding of the associations of these proxies with sea ice is largely based on their relative abundances in surface sediment assemblages with respect to sea-ice cover, as well as limited ecological field and laboratory studies. Each of these proxies has some advantages over others, but there are important limitations to consider, largely stemming from the lack of adequate understanding of their behaviour in modern environments seasonally with respect to habitat source (sea-ice, under-ice, ice-edge or open-water) and factors controlling proxy production, vertical transport and deposition on the sediment. While certain diatom species are known to inhabit the sea-ice matrix, these species are rarely observed in sediment archives. In contrast, dinoflagellate cysts are preserved exceptionally well in the sediment, but observations of key sea-ice proxy species from modern habitats are rare. Furthermore, the motile stage forms for the dinoflagellate cyst species routinely used as sea-ice proxies are in many cases unknown or only recently discovered, which is why ecological knowledge of these species is very limited. The sea-ice source specificity for IP25 and IPSO25 is well established, but the diatom species producing specific HBIs need further work, particularly for HBI III and HBI IV, which are generally used as proxies for ice-edge or open-water production. Furthermore, the environmental regulators of HBI production are still incompletely understood. Very little is also known of the seasonal dynamics of proxy production with respect to sea-ice cover, especially at Arctic coastal sites. This thesis aims to fill some of the knowledge gaps in sea-ice proxy understanding by examining the seasonality of proxy production, their habitat (ice, under-ice, ice-edge, open-water) sources and vertical export to the sediment, as well as taxonomic relationships of Arctic dinoflagellate cysts. This was achieved by investigating 1) diatom species assemblages and vertical fluxes over a full annual production cycle using automatically sequencing sediment traps at two sites in Greenland with differing ice regimes (high-Arctic Young Sound and sub-Arctic Godthåbsfjord), 2) seasonal changes, habitat sources and vertical transport of diatom, dinoflagellate cyst and HBI assemblages during spring and summer from sea-ice, water-column, sediment-trap and surface-sediment samples from a Hudson Bay Archipelago and 3) taxonomic relationships (single-cell LSU rDNA) of spiny brown dinoflagellate cyst sea-ice proxies with unknown motile affinities from Disko Bay area, Greenland. The results provide novel evidence of the seasonality and habitat sources of the three proxies and offer insights into their association with sea ice in different geographical settings. The results support state-of-the-art interpretations of diatoms associated with under-ice or ice-edge blooms, namely Fragilariopsis oceanica, Fragilariopsis reginae-jahniae and Fossulaphycus arcticus, as reliable indicators for seasonal sea-ice conditions when sympagic diatom species are not detected in the sediment archives and provide important new insights into their seasonal and source dynamics. Fragilariopsis oceanica, Fragilariopsis reginae-jahniae and Fossulaphycus arcticus were highly abundant in both Young Sound and the Hudson Bay Archipelago, the two sites experiencing seasonal sea ice. The species showed clearly seasonal bloom windows associated with spring ice break-up both in Young Sound and the Hudson Bay archipelago. Although sympagic diatoms were abundant in some of the sediment-trap assemblages, their signal in the surface-sediment assemblages was diluted by the significant summer open-water bloom in the Hudson Bay archipelago. Despite this, the under-ice and ice-edge species F. oceanica, F. reginae-jahniae and F. arcticus nevertheless formed a significant portion of the surface sediment assemblage. While these species were also present in minute abundances in ice-free Godthåbsfjord, their presence was linked with drifting ice from a nearby area. Importantly, grazing and dissolution appear to have a marked effect on diatom fluxes to the sediment, likely controlled by differences in the water-column depth between the sites and possibly varying grazing rates. Thus, comparing species fluxes from sites with different water depths should be treated with caution, since it may significantly affect the species composition. Here, the sympagic diatom communities were highly diverse, but species composition was largely decoupled from that of the water column, in particularly in sediment traps and surface sediment. Unlike diatoms that were abundant in sea-ice, water-column and sediment-trap samples throughout the spring melting season, dinoflagellate cysts were absent. One cyst of Islandinium minutum subsp. minutum and one of Echinidinium karaense, both routinely used for sea-ice reconstruction, were detected in the sediment trap after the ice break-up, while the surface sediment assemblage displayed a typical sub-Arctic composition (11% contribution by the two species). This indicates that key dinoflagellate cyst proxies for sea ice are not unequivocally associated with the sea-ice environment, but are likely produced later in the season, and their use as sea-ice proxies should be carefully considered for each research site and question. The most common HBI in the spring sea-ice samples was surprisingly HBI IV, which also exhibited trends in concentrations similar of those of the diatom Pleurosigma rhomboides, a known IP25 producer. IP25 was the second most abundant HBI in the sea ice and predominated in the sediment-trap samples towards the ice break-up. It was also the most abundant HBI in the surface sediment. While two IP25-producing diatom species were detected (P. rhomboides and Haslea kjelmannii), all HBIs were found in all sample types. This suggests that further research is needed to identify not only the potential diatom source species, but also to understand the environmental factors affecting the physiological responses to lipid synthetization, since this study implies seasonal trends in the production of HBI IV and IP25, in particular. Furthermore, the use of HBI IV as a proxy for ice-edge or open-water conditions also needs re-evaluation. This study revealed a previously undiscovered complexity in taxonomic relationships with the spiny brown dinoflagellate cysts routinely used as sea-ice proxies. Single-cell LSU rDNA and phylogenetic analyses revealed that the dinoflagellate cyst species Protoperidinium tricingulatum, previously reported only from temperate areas and bearing conspicuous morphological similarity to common sea-ice proxy species Islandinium? cezare (morphotype 1, Head et al. 2001), was present in Disko Bay sediment. Morphological variation of P. tricingulatum and I.? cezare (morphotype 1) specimens was also described from standard microscopic slides from the same material. This raises questions whether I.? cezare morphotype 1 is indeed P. tricingulatum or whether these species co-exist in the Arctic. Past sea-ice records are of high relevance to climate science and society. This work addresses the critical need for better understanding of the sea-ice proxies routinely used to produce these records. While the results provide enhanced understanding for the use of biogenic sea-ice proxies generally, they also highlight the importance of understanding the site-specific differences affecting the proxies. Importantly, this work also raises questions that need to be resolved by future studies. Arktisen merijääpeitteen nopea väheneminen vaikuttaa voimakkaasti arktisen alueen ekosysteemeihin, ravintoketjuihin ja monimuotoisuuteen, mutta samanaikaisesti myös maailmanlaajuisiin ilmasto-olosuhteisiin. Satelliittihavaintoihin perustuvat aikasarjat merijään muutoksista kattavat vain muutaman vuosikymmenen, vaikkakin muutoksia on tärkeää tutkia erityisesti pidemmällä aikavälillä. Tutkimuksessa voidaan hyödyntää luonnonarkistoja, kuten merenpohjan sedimenttikerrostumia, joihin tallentuu tietoa menneistä olosuhteista jopa miljoonien vuosien aikajänteellä. Sedimenttiin tallentuneista eloperäisistä (sub)fossiilisista ja biogeokemiallisista mikro-organismien jäänteistä käytetään nimitystä paleoekologinen merijääproksi. Näihin lukeutuvat muun muassa piilevät, panssarisiimalevien lepoitiöt ja korkeasti haarautuneet isoprenoidilipidit (HBI), joita tietyt piilevälajit tuottavat. Nämä proksityypit ovat yleisiä sedimentin pinnalla alueilla, joilla merijäätä esiintyy. Sen sijaan proksien esiintymisestä merijäässä ja merijääympäristöissä eri vuodenaikoina, niiden kulkeutumisesta merenpohjaan sekä yhteydestä ekologisiin tekijöihin ja tunnettuihin nykyorganismeihin, on rajallinen määrä tietoa. Tämän työn tarkoitus on lisätä tietämystä arktisista merijääprokseista tutkimalla lajiston vuodenaikaista tuotantoa, elinympäristöä ja kulkeutumista merijäästä merenpohjan sedimenttiin, sekä tutkimalla keskeisten arktisten panssarisiimalevien lepoitiöiden välisiä taksonomisia suhteita. Näihin asioihin vastataan tarkastelemalla 1) piileväyhteisöjen lajikoostumusta ja niiden vertikaalista kulkeutumista vesipatsaassa yhden vuoden kierron ajalta 2) piilevien, panssarisiimalevien lepoitiöiden ja HBI-lipidien vuodenaikaista vaihtelua eri elinympäristöissä sekä niiden vertikaalista kulkeutumista merenpohjaan ja 3) panssarisiimalevien lepoitiöiden välisiä taksonomisia suhteita käyttäen LSU rDNA -menetelmää. Tutkimuskohteet sijaitsivat Kanadan ja Grönlannin rannikkoalueilla. Tämän työn tulokset antavat uutta tietoa lajien tuotannon vuodenaikaisesta vaihtelusta ja lähteistä, sekä niiden suhteesta merijäähän erilaisissa maantieteellisissä kohteissa. Tulokset tukevat aiempia havaintoja tiettyjen merijääympäristöissä esiintyvien piilevälajien soveltuvuudesta menneiden merijääolosuhteiden rekonstruointiin. Tulokset osoittavat, että monet jäässä elävät piilevälajit eivät joko säily sedimenteissä tai niiden tuotannon vähäisen määrän vuoksi niiden esiintyvyys on verrattain heikko kevätkukkijoihin nähden. Jääpeiteajan pituus ja jään sulamisen ajoitus vaikuttavat piileväkukintoihin vahvasti, ja kukinto voi olosuhteista riippuen olla erittäin lyhytaikainen, mutta voimakas. Sen sijaan panssarisiimalevien itiöitä ei löytynyt merijäästä lainkaan, vaikka niitä esiintyy runsaasti pintasedimentissä. Tämän vuoksi niiden käyttöä merijääproksina tulee harkita tutkimuskysymys- ja aluekohtaisesti. Vaikka kullekin HBI-lipidille on todistetusti osoitettu useampia tuottajalajeja, tulokset osoittavat, että tuottajia saattaa olla tiedettyä enemmän. Tulokset painottavat, että olisi tärkeää tarkentaa eri HBI-lipidien käyttöä merijää- tai avovesi-indikaattoreina. Lisäksi DNA-tutkimukset osoittivat, että panssarisiimalevien lepoitiöiden väliset taksonomiset suhteet ovat monimutkaisempia kuin on aiemmin ymmärretty, ja lisää tietoa tarvitaan, paitsi lajien välisistä suhteista, mutta myös niiden yhteydestä merijäähän. Menneiden merijääolosuhteiden rekonstruktiot tuottavat sekä ilmastotieteelle että yhteiskunnalle merkittävää tietoa. Tässä työssä vastattiin akuuttiin tarpeeseen ymmärtää tavanomaisissa merijäärekonstruktioissa käytettyjä paleoekologisia menetelmiä paremmin. Tutkimustulokset helpottavat eloperäisten merijääproksien sedimentissä esiintymisen tulkintaa arktisella alueella, mutta myös korostavat tarvetta ymmärtää tutkimusalueiden välisiä eroja.
- Published
- 2023