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Abstract Global warming is causing rapid changes to the cryosphere. Predicting the future trajectory of the cryosphere requires quantitative reconstruction of its past variations. A recently identified sea-ice-associated haptophyte, known as Group 2i Isochrysidales, has given rise to a new sea-ice proxy with its characteristic alkenone distributions. However, apart from the occurrence of Group 2i Isochrysidales in regions with sea ice, and the empirical relationship between C_37:4alkenone abundance and sea-ice concentration, little is known about the ecology of these haptophyte species. Here, we systematically mapped the spatial and temporal occurrence of known Group 2i Isochrysidales based on environmental DNA in both marine and lacustrine environments. Our results indicate Group 2i is widely distributed in icy marine and lacustrine environments in both Northern and Southern Hemisphere, but is absent in warm environments. Temporally, Group 2i is part of the sea-ice algae bloom during the cold seasons, in contrast to other Isochrysidales that bloom in open waters during warm seasons. Our results indicate that ice is a prerequisite for the occurrence of the psychrophilic Group 2i haptophytes in marine and lacustrine ecosystems and further affirms its value for past ice reconstructions. 

Alkenones are long-chain ketones produced by phytoplankton of the order Isochrysidales. They are widely used in reconstructing past sea surface temperatures, benefiting from their ubiquitous occurrence in the Cenozoic ocean. Carbon isotope fractionation (εp) between alkenones and dissolved inorganic carbon may also be used as a proxy for past atmospheric pCO2 and has provided continuous pCO2 estimates back to ca. 45 Ma. Here, an extended occurrence of alkenones from ca. 130 Ma is reported. We characterize the molecular structure and distribution of these Mesozoic alkenones and evaluate their potential phylogenetic relationship with Cenozoic alkenones. Using δ13C values of the C37 methyl alkenone (C37:2Me), the first alkenone-based pCO2 estimates for the Mesozoic are derived. These estimates suggest elevated pCO2 with a range of 548−4090 ppm (908 ppm median) during the super-greenhouse climate of the Early Cretaceous, in agreement with phytane-based pCO2 reconstructions. Finally, insights into the identity of the Cretaceous coccolithophores that possibly synthesized alkenones are also offered. 

Abstract Alkenones are biomarkers produced solely by algae in the order Isochrysidales that have been used to reconstruct sea surface temperature (SST) since the 1980s. However, alkenone-based SST reconstructions in the northern high latitude oceans show significant bias towards warmer temperatures in core-tops, diverge from other SST proxies in down core records, and are often accompanied by anomalously high relative abundance of the C₃₇tetra-unsaturated methyl alkenone (%C_37:4). Elevated %C_37:4is widely interpreted as an indicator of low sea surface salinity from polar water masses, but its biological source has thus far remained elusive. Here we identify a lineage of Isochrysidales that is responsible for elevated C_37:4methyl alkenone in the northern high latitude oceans through next-generation sequencing and lab-culture experiments. This Isochrysidales lineage co-occurs widely with sea ice in marine environments and is distinct from other known marine alkenone-producers, namelyEmiliania huxleyiandGephyrocapsa oceanica. More importantly, the %C_37:4in seawater filtered particulate organic matter and surface sediments is significantly correlated with annual mean sea ice concentrations. In sediment cores from the Svalbard region, the %C_37:4concentration aligns with the Greenland temperature record and other qualitative regional sea ice records spanning the past 14 kyrs, reflecting sea ice concentrations quantitatively. Our findings imply that %C_37:4is a powerful proxy for reconstructing sea ice conditions in the high latitude oceans on thousand- and, potentially, on million-year timescales. 

Abstract Members of the order Isochrysidales are unique among haptophyte lineages in being the exclusive producers of alkenones, long‐chain ketones that are commonly used for paleotemperature reconstructions. Alkenone‐producing haptophytes are divided into three major groups based largely on molecular ecological data: Group I is found in freshwater lakes, GroupIIcommonly occurs in brackish and coastal marine environments, and GroupIIIconsists of open ocean species. Each group has distinct alkenone distributions; however, only GroupsIIandIIIIsochrysidales currently have cultured representatives. The uncultured Group I Isochrysidales are distinguished geochemically by the presence of tri‐unsaturated alkenone isomers (C_37:3bMe, C_38:3bEt, C_38:3bMe, C_39:3bEt) present in water column and sediment samples, yet their genetic diversity, morphology, and environmental controls are largely unknown. Using small‐subunit (SSU) ribosomalRNA(rRNA) marker gene amplicon high‐throughput sequencing of environmental water column and sediment samples, we show that Group I is monophyletic with high phylogenetic diversity and contains a well‐supported clade separating the previously described “EV” clade from the “Greenland” clade. We infer the first partial large‐subunit (LSU)rRNAgene Group I sequence phylogeny, which uncovered additional well‐supported clades embedded within Group I. Relative to GroupII, Group I revealed higher levels of genetic diversity despite conservation of alkenone signatures and a closer evolutionary relationship with GroupIII. In Group I, the presence of the tri‐unsaturated alkenone isomers appears to be conserved, which is not the case for GroupII. This suggests differing environmental influences on Group I andIIand perhaps uncovers evolutionary constraints on alkenone biosynthesis.