31 51 52 Coronaviruses are a diverse family of positive-sense, single-stranded RNA viruses, found widely 53 in mammals and birds 1 . They have a broad host range, a high mutation rate, and the largest 54 genomes of any RNA viruses, but they have also evolved mechanisms for RNA proofreading and 55repair, which help to mitigate the deleterious effects of a high recombination rate acting over a 56 large genome 2 . Consequently, coronaviruses fit the profile of viruses with high zoonotic potential. 57There are seven human coronaviruses (two in the genus Alphacoronavirus and five in 58Betacoronavirus), of which three are highly pathogenic in humans: SARS-CoV, SARS-CoV-2, and 59MERS-CoV. These three are zoonotic and widely agreed to have evolutionary origins in bats 3-6 . 60 61Our collective experience with both SARS-CoV and MERS-CoV illustrate the difficulty of tracing 62 specific animal hosts of emerging coronaviruses. During the 2002-2003 SARS epidemic, SARS-63 CoV was traced to the masked palm civet (Paguma larvata) 7 , but the ultimate origin remained 64 unknown for several years. Horseshoe bats (family Rhinolophidae: Rhinolophus) were implicated 65 as reservoir hosts in 2005, but their SARS-like viruses were not identical to circulating human 66 strains 4 . Stronger evidence from 2017 placed the most likely evolutionary origin of SARS-CoV in 67 Rhinolophus ferrumequinum or potentially R. sinicus 8 . Presently, there is even less certainty in the 68 origins of MERS-CoV, although spillover to humans occurs relatively often through contact with 69 dromedary camels (Camelus dromedarius). A virus with 100% nucleotide identity in a ~200 base 70 pair region of the polymerase gene was detected in Taphozous bats (family Emballonuridae) in 71 Saudi Arabia 9 ; however, based on spike gene similarity, other sources treat HKU4 virus from 72 Tylonycteris bats (family Vespertilionidae) in China as the closest-related bat virus 10,11 . Several 73 bat coronaviruses have shown close relation to MERS-CoV, with a surprisingly broad geographic 74 distribution from Mexico to China 12,13,14,15 . 75 76 Coronavirus disease 2019 (COVID-19) is caused by severe acute respiratory syndrome 77 coronavirus-2 (SARS-CoV-2), a novel virus with presumed evolutionary origins in bats. Although 78 the earliest cases were linked to a wildlife market, contact tracing was limited, and there has been 79 no definitive identification of the wildlife contact that resulted in spillover nor a true "index case." 80 Two bat viruses are closely related to SARS-CoV-2: RaTG13 bat CoV from Rhinolophus affinis 81 (96% identical overall), and RmYN02 bat CoV from Rhinolophus malayanus (97% identical in one 82 gene but only 61% in the receptor-binding domain and with less overall similarity) 6,16 . The 83 divergence time between these bat viruses and human SARS-CoV-2 has been estimated as 40-50 84 years 17 , suggesting that the main host(s) involved in spillover remain unknown. Evidence of viral 85 recombination in pangolins has been proposed but is unresolved 17 . S...