Characterization of a Novel Betacoronavirus Related to Middle East Respiratory Syndrome Coronavirus in European Hedgehogs

Bats are known to host viruses closely related to important human coronaviruses (HCoVs), such as HCoV-229E, severe-acute respiratory syndrome coronavirus (SARS-CoV), andMiddle East respiratory syndrome CoV (MERS-CoV). As RNA viruses may coevolve with their hosts, we sought to investigate the closest sister taxon to bats, the Eulipotyphla, and screened European hedgehogs (Erinaceus europaeus) from Germany for CoV by nested reverse transcriptase PCR. A novel betacoronavirus species in a phylogenetic sister relationship toMERS-CoV and clade c bat CoVs was detected and characterized on the whole-genome level. A total of 58.9 % of hedgehog fecal specimens were positive for the novel CoV (EriCoV) at 7.9 log10 mean RNA copies per ml. EriCoV RNA concentrations were higher in the intestine than in other solid organs, blood, or urine. Detailed analyses of the full hedgehog intestine showed the highest EriCoV concentrations in lower gastrointestinal tract specimens, compatible with viral replication in the lower intestine and fecal-oral transmission. Thirteen of 27 (48.2%) hedgehog sera contained non-neutral-izing antibodies against MERS-CoV. The animal origins of this betacoronavirus clade that includes MERS-CoVmay thus include both bat and nonbat hosts. The Coronaviridae subfamily Coronavirinae contains the fourgenera Alpha-, Beta-, Gamma- and Deltacoronavirus (1, 2). Betacoronaviruses are further discriminated into clades a to d. Until recently, five human coronaviruses (HCoVs) were known, namely, the alphacoronaviruses HCoV-229E and HCoV-NL6