During the past decade, our understanding of eukaryotic evolution has increased immensely. Newly recognized eukaryotic supergroups have been established1-3, and most enigmatic orphan lineages have had their relationships resolved4-6. Studies on unicellular protist eukaryotes have also been key to understanding the evolution of mitochondria, the fundamental organelles of the eukaryotic cell, which originated from an alphaproteobacterial ancestor. The retention of ancestral alphaproteobacterial pathways in some protist lineages reveals that the mitochondrion of the last eukaryotic common ancestor was more metabolically versatile than are the highly derived mitochondria that are found in most modern eukaryotes7,8. Here we report the discovery of such a unicellular eukaryote, Solarion arienae gen. et sp. nov., an inconspicuous, free-living heterotrophic protist with two morphologically distinct cell types and a novel type of predatory extrusome. We assign Solarion to the new phylum Caelestes. Together with Provora, hemimastigophoreans and Meteora, they form a new eukaryotic supergroup, Disparia. Moreover, S. arienae has some noteworthy traits associated with the mitochondrial genome; in particular, the mitochondrially encoded secA gene, a remnant of an ancestral alphaproteobacterial protein secretion pathway, which has been lost almost entirely in extant mitochondria9,10. The discovery of S. arienae broadens our understanding of early eukaryotic evolution and facilitates the study of proto-mitochondrial metabolic remnants, shedding light on the complexity of ancestral eukaryotic life.

Department of Biological Sciences Mississippi State University Starkville MS USA

Department of Biological Sciences Texas Tech University Lubbock TX USA

Department of Marine Sciences University of Puerto Rico Mayagüez Mayagüez Puerto Rico

Department of Parasitology Faculty of Science BIOCEV Charles University Vestec Czechia

Department of Zoology Faculty of Science Charles University Prague Czechia

Imaging Methods Core Facility at BIOCEV Faculty of Science Charles University Vestec Czechia

Institute for Genomics Biocomputing and Biotechnology Mississippi State University Starkville MS USA

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