Coronavirus disease 2019 (COVID-19), caused by severe acute respiratory syndrome coronavirus 2 43 (SARS-CoV-2) rapidly spread worldwide, and recent studies suggest that pets and other animals 44 could also be infected by SARS-CoV-2 through natural contact [1, 2] . Captive rhesus macaques (M. 45 mulatta), inoculated with SARS-CoV-2 in pathological studies, exhibited a moderate infection as 46 observed in the majority of human cases [3, 4] . Besides captive animals and pets, wild animals are 47 also susceptible to the infection of coronaviruses transmitted from humans. For instance, in 2016, 48 wild chimpanzees in Côte d´Ivoire were infected by the human coronavirus OC43 [5] . 49 The close evolutionary relationship between humans and NHPs is thought to support pathogen 50 transmission [6] and many viruses have been described that circulate between humans and NHPs. 51 In captive and wild NHPs, various viruses including coronaviruses, enteroviruses, enteric 52 adenoviruses, rotaviruses, and picobirnaviruses have been detected, which are also found in 53 humans [7] [8] [9] . The most prominent cases of virus transmission from wild NHPs to human are 54 simian foamy virus (SFV), yellow fever virus (YFV), Zika virus (ZIKV), and human 55 immunodeficiency virus (HIV) [10] [11] [12] [13] . Conversely, viruses such as poliovirus and measles have 56 been reported in NHPs and likely derived from local human populations [14] . To block the 57 potential circular transmission route of different viruses between human and NHPs, precautions and 58 regulations are needed. 59
Here we performed a survey on documented virus infections in NHPs (VI-NHPs) based on 60 published data. First, we generated a summary statistics of worldwide reported VI-NHPs. We then 61 identified and predicted NHP species with a high risk of virus transmission from humans and 62 predicted geographic locations where disease outbreaks are likely to occur. 63 64 4 / 13
Global information of VI-NHPs was extracted from the Global Mammal Parasite Database (GMPD, 67 http://www.mammalparasites.org/). We also used literature searches for publications describing 68 VI-NHPs, which were not included in GMPD. Only the natural virus infections in captive and wild 69
NHPs have been recorded, while the virus inoculations for pathological studies are not included. 70 We then built host-virus ecological networks in which nodes represent NHPs that are linked 71 through shared viruses. Since centrality in primate-virus networks could assess the potential for the 72 circulation of viruses among NHPs and humans, we estimated the centrality using four metrics: 73 strength degree centrality, eigenvector centrality, betweenness centrality, and closeness centrality 74 implemented in the R package "igraph" and UCINET 6. Table S1 ), indicating that they detected similar NHP species as most central. 108 A single factor found in PCA explained 85.2% of the variance of the indices, which was used as the 109 composite index to assess the centrality of each node ( Cercopithecus nictitans (Figure 2a and 2b) . 114 After controlling for phylogeny, virus number in each NHP species and the number of viruses 115 shared with humans in each NHP species were significantly and positively associated to the 116 centrality of each NHP species (strength degree centrality, eigenvector centrality, betweenness 117 centrality, closeness centrality, and the composite centrality; Figure 2c and 2d, Table S3 Results showed the some trend with the analysis without controlling the sampling efforts, and for 151 the sake of brevity we provide results in the supplementary metarials (Table S5-S9 and Figure  152 S1). In the future, more efforts ought to be made for the collection, documentation and analysis 153 of VI-NHP, especially for NHP species with higher potential of virus transmission. Since 154 coronaviruses have been reported in macaques and other primates [7, 8] , viral surveys should 155 first target such species, not only to find known coronaviruses in such populations, but also to 156 find new strains with high zoonotic potential. 157 Experts in animal health and conservation are starting to urge for the protection of great 158 apes during human COVID-19 pandemics, since the transmission of the human virus to apes 159 could result in severe outbreaks and local extinctions [25] . We suggest to expand such efforts to 160 various Old World monkeys, as many of them, for instance, baboons or macaques, are widely 161 distributed and often in close proximity to humans (Figure 3a, 3b and 3c) . 