Phylogenetic tree of microbial rhodopsins. Archaea are: BR, halorhodopsin (HR), SRI and -II from H. salinarum (Hs), Natronomonas pharaonis (Np), Haloarcula vallismortis (Hv), H. walsbyi (Hw), archaerhodopsin-3 from H. sodomense (AR3). Bacteria are: TR from T. thermophilus JL-18, T. oshimai JL-2, and Thermus sp. CBB_US3_UF1, PR from marine proteobacteria, GR from G. violaceus, E. sibiricum rhodopsin (ESR) from E. sibiricum, actinorhodopsin (ActR) from actinobacteria, xanthorhodopsin (XR), halorhodopsin (HR), and SRI from Salinibacter ruber (Sr), and Anabaena sensory rhodopsin (ASR) from Anabaena sp. PCC7120. Eukaryotes are: channelrhodopsin I (ChRI) and II (ChRII) from Chlamydomonas reinhardtii, Leptosphaeria rhodopsin (LR) from Leptosphaeria maculans, and Acetabularia rhodopsin (AR) from Acetabularia acetabulum. Dashed-lined arrows indicate domains of life. PR-like and BR-like proton pumps are highlighted in light gray and gray, respectively, and sensory rhodopsins are highlighted in dark gray. TRs were discovered from thermophiles living at higher temperatures (∼73–77 °C, indicated by the bold black arrow).