Our study is the first to establish surveillance for zoonotic viruses in wild animal products illegally imported into the United States in an effort to prevent the transmission of infectious agents from these shipments. The restricted number of samples included in this study were tested for a limited range of pathogens only and thus presence of additional pathogens not included in this study cannot be ruled out. We identified four SFV strains and two different herpesviruses (in some cases in the same tissues) in smuggled NHP bushmeat. Using phylogenetic analysis and gross examination, we were able to determine that bushmeat from nine NHP species and at least two rodent species were attempted to be smuggled into the United States. These results are consistent with the origin of the shipments from West Africa and included species of conservation importance (P. papio
, Cercocebus atys
, and P. t. ellioti
are classified as “near threatened”, “vulnerable”, and “endangered”, respectively by the International Union for Conservation of Nature), suggesting more education efforts or harsher penalties are needed regarding the handling, consumption, and illegal transportation of products from wildlife of conservation concern. In addition, the finding of mangabey, guenon, and cane rat bushmeat in our study is consistent with that reported by Chaber et al
who found these and bushmeat from nine other species entering Paris-Charles de Gaulle Airport 
Our finding of SFV DNA in smuggled NHP specimens comprising of four species (baboon, chimp, mangabey, and AGM) is significant because SFV is a known zoonotic infection of humans exposed to NHPs. However, the mode of transmission to humans is poorly understood and while most infected people reported sustaining a NHP exposure (mostly bites) others did not, suggesting a less invasive mode of infection is possible 
. These viruses are probably not easily spread from human-to-human, although persistent infection has been documented 
. Several SFV-positive people reported donating blood while infected and because blood banks do not screen for SFV, secondary transmission via contaminated blood donations may be possible 
. Further research into the possibility of secondary transmission of SFV is required. The finding of SFV DNA in the bushmeat samples highlights a potential public health risk of exposure to these tissues along the hunting, transportation, and consumption continuum with multiple opportunities for primary transmissions. Unlike most retroviruses whose RNA genome is packaged in the viral particles, foamy viruses are unusual in that DNA and/or RNA can be present in the infectious virus particles. Thus, finding of only DNA does not exclude that SFV in these tissues is not infectious, especially in the more recently CDC confiscated items which contained fresher tissue compared to the USFWS items confiscated in 2006 that were partially degraded at the time of analysis in 2010. Human infection with SFV is of further concern because increases in the pathogenicity of simian retroviruses following cross-species transmission have been documented (e.g., HIV-1 and HIV-2) 
. However, the limited number of cases, short follow-up duration, and selection biases in the enrolling of healthy workers or hunters to identify cases all limit the identification of potential disease associations 
Although we did not find SIV or STLV in the limited number of specimens in this study, these viruses have been found in high prevalences in NHP specimens at bushmeat markets and in hunted NHPs 
. HIV-1 and HIV-2 emerged as a result of several spillover events of SIV from chimpanzees and mangabeys, respectively, that were likely hunted for bushmeat in central and western Africa 
. Serosurveillance studies have shown thirty-five different species of African NHPs harbor lentivirus infections, with a prevalence of SIV in up to 35% of free-ranging chimpanzees, and 30–60% of free-ranging sooty mangabeys and green monkeys 
To date, four groups of HTLV viruses found in humans are believed to have originated from corresponding STLV strains in NHP species (including mangabeys, baboons, and chimpanzees) via multiple transmission events 
. HTLV-1, closely related to STLV-1 group viruses, infects 15 to 20 million people worldwide and is spread from person to person via bodily fluids 
. These viruses are capable of causing leukemia, lymphoma and neurologic disease in humans 
. Discoveries of HTLV-3 and HTLV-4, and a novel STLV-1 strain were recently made in NHP hunters in Cameroon 
, and 89% of hunted bushmeat in Cameroon has been shown to be infected with STLV strains 
. Although imported wildlife products are often not in a freshly-killed state, many are not smoked or processed in any manner, thus screening of larger sample collections of smuggled bushmeat may reveal evidence of these viruses.
Like retroviruses, herpesviruses can cause long-term latent infections in their host. Most herpesviruses are host-specific, yet particular strains are capable of causing severe disease in the non-host, examples of which include agents of malignant catarrhal fever and Herpes B virus 
. CMVs are in the betaherpesvirus subfamily. Human CMV is typically asymptomatic in humans, with the exception of immunocompromised persons. Similarly, many NHPs are asymptomatic hosts of CMV that do not typically infect other species, including humans. However, baboon CMV (bCMV), like that identified in our study, has been shown to replicate in human tissues in vitro as well as infect and replicate in humans following a bCMV-positive liver xenotransplant 
Lymphocryptoviruses (LCV) are in the gammaherpesvirus subfamily, and include human LCV, and Epstein-Barr virus (EBV), the agent of infectious mononucleosis. Nearly 90% of adults in the United States have antibodies indicating exposure at some point to EBV. LCVs are typically asymptomatic in their host, with the exception of immunocompromised individuals who may develop B-cell tumors. Although much less efficient, baboon LCV can infect human B cells in immunocompromised persons or in persons co-infected with EBV and replicate in EBV-immortalized B cells with the theoretical potential for viral recombination 
. However, it is unknown if the novel herpesviruses found in bushmeat specimens in our study can easily infect humans handling these tissues. Systematic studies examining herpesvirus transmission risks associated with handling or consumption of infected animal tissues have not been reported. In addition, virus isolation was not performed in our study to determine the infectiousness of the specimens at the time of confiscation.
In summary, our study establishes initial surveillance methodology to detect and identify zoonotic pathogens and species of origin of wildlife products entering the United States. While we were successful in demonstrating the presence of SFV and herpesviruses in bushmeat specimens, our pilot study was limited by the range, number, and variable condition of products available to us and was not intended to be a comprehensive review of presence or to measure prevalence of all pathogens imported in wildlife products. Because our study only included a small number of CDC-regulated species and excluded products of ungulate, carnivore, reptile, avian and other origin, as well as any live animal imports, all of which may carry zoonotic pathogens or diseases that threaten domestic livestock or native wildlife, in addition to the fact that virus isolation was not performed in our study to determine the infectiousness of the specimens at the time of confiscation, there is a large component of zoonotic disease risk assessment not included in this study. A further understanding of pathogen movements through the trade will only be recognized through broader surveillance efforts and pathogen identification and discovery techniques in wildlife and wildlife products arriving at US ports of entry so that appropriate measures can be taken to further mitigate potential risks.