PMC full text:
Published online 2015 Mar 25. doi: 10.1128/CMR.00102-14
TABLE 5
Evidence of zoonotic sources of MERS-CoV and closely related CoVsa
| Animal species and region (virus) | Country (area)/specimen collection date | Main findings | Reference(s) |
|---|---|---|---|
| Bats | |||
| Superfamily Vespertilionoidea, family Vespertilionidae | |||
| Asia | |||
| Tylonycteris pachypus (Ty-BatCoV HKU4) | China (Hong Kong)/April 2005 to August 2012 | Detected in 29/99 (29.3%) alimentary samples; shared 90.0% (RdRp), 67.4% (S), and 72.3% (N) aa identities with MERS-CoV (HCoV-EMC/2012). | 13, 99 |
| Pipistrellus abramus (Pi-BatCoV HKU5) | China (Hong Kong)/April 2005 to August 2012 | Detected in 55/216 (25.5%) alimentary samples; shared 92.3% (RdRp), 64.5% (S), and 70.5% (N) aa identities with MERS-CoV (HCoV-EMC/2012). | 13, 99 |
| Vespertilio superans (Bat CoV-BetaCoV/SC2013) | China (southwestern part)/June 2013 | Detected in 5/32 (15.6%) anal swabs; shared 75.7% (complete genome of 1 strain) nt identity; and 96.7% (816-nt RdRp fragment) and 69.0% (S) aa identities with MERS-CoV (HCoV-EMC/2012). | 312 |
| Africa | |||
| Neoromicia capensis (NeoCoV) | South Africa (KwaZulu-Natal and Western Cape provinces)/2011 | Detected in 1/62 (1.6%) fecal samples; shared 85.6% (complete genome) nt identity; and 64.6% (S), 89.0% (E), 94.5% (M), and 91.7% (N) aa identities with MERS-CoV from humans and camels, placing them in the same viral species based on taxonomic criteria. | 110, 111 |
| Europe | |||
| Pipistrellus, Pipistrellus nathusii, and Pipistrellus pygmaeus (Pipistrellus bat βCoVs) | Romania (Tulcea county) and Ukraine (Kiev region)/2009–2011 | Detected in 40/272 (14.7%) fecal samples; shared 98.2% (816-nt RdRp fragment) aa identity with MERS-CoV (HCoV-EMC/2012). | 313 |
| Pipistrellus kuhlii, Hypsugo savii, Nyctalus noctula, and an unknown Pipistrellus sp. (βCoVs) | Italy (Lombardia and Emilia regions)/2010–2012 | 10 βCoVs detected in fecal specimens of Pipistrellus kuhlii (7), Hypsugo savii (1), Nyctalus noctula (1), and an unknown Pipistrellus sp. (1) bats; shared 85.2% to 87% nt identity and 95.3% to 96.1% (329-nt RdRp fragment) aa identity with MERS-CoV (HCoV-EMC/2012). | 314 |
| Superfamily Emballonuroidea, family Emballonuridae | |||
| Taphozous perforatus (βCoV) | KSA (Bisha)/October 2012 | A βCoV detected in 1/29 (3.4%) fecal samples; shared 100% nt identity (182-nt RdRp fragment) with MERS-CoV (HCoV-EMC/2012). | 109 |
| Superfamily Molossoidea, family Molossidae | |||
| Nyctinomops laticaudatus (Mex_CoV-9) | Mexico (Campeche)/2012 | Detected in 1/5 (20.0%) rectal swabs; shared 96.5% (329-nt RdRp fragment) aa identity with MERS-CoV (HCoV-EMC/2012). | 315 |
| Superfamily Noctilionoidea, family Mormoopidae | |||
| Pteronotus davyi (BatCoV-P.davyi49/Mexico/2012) | Mexico (La Huerta)/2007–2010 | Detected in 1/4 (25.0%) intestinal samples; shared 71.0% (439-nt RdRp fragment) nt identity with MERS-CoV (HCoV-EMC/2012). | 316 |
| Superfamily Rhinolophoidea, family Nycteridae | |||
| Nycteris gambiensis (Nycteris bat CoV) | Ghana (Bouyem, Forikrom, and Kwamang)/2009–2011 | Detected in 46/185 (24.9%) fecal samples; shared 92.5% aa identity (816-nt RdRp fragment) with MERS-CoV (HCoV-EMC/2012). | 313 |
| Hedgehogs | |||
| Europe | |||
| Erinaceus europaeus (Erinaceous CoV) | Northern Germany/unknown date | Two clades detected in 146/248 (58.9%) fecal samples; shared 89.4% (816-nt RdRp fragment), 58.2% (S), 72.0% (E), 79.4% (M), and 72.1% (N) aa identities with MERS-CoV (HCoV-EMC/2012); RNA concn was higher in the intestine and fecal samples than in other solid organs, blood, or urine, suggestive of viral replication in the lower intestine and fecal-oral transmission; 13/27 (48.2%) sera contained nonneutralizing antibodies. | 113 |
| Camelids | |||
| Middle East | |||
| Camelus dromedarius | KSA (countrywide)/1992 to 2010 and November to December 2013 | Serum Ab: 150/203 (73.9%) (2013) and 72%–100% (1992 to 2010); adults (95%) > juveniles (55%). Viral RNA: nasal > rectal swabs; juveniles (36/104; 34.6%) > adults (15/98; 15.3%). Virus isolation: two nasal swabs cultured in Vero E6 cells. | 123, 137 |
| KSA (Riyadh and Al Ahsa)/2012 to 2013 | Serum NAb: 280/310 (90.3% with titer ≥ 1:20); adults (233/245; 95.1%) > juveniles (47/65; 72.3%). | 127 | |
| KSA (Jeddah)/3 November 2013 | Direct camel-to-human transmission: phylogenetic (identical full genome sequences of patient strain and an epidemiologically linked camel strain) and serological (the virus was circulating in the epidemiologically linked camels but not in the patient before the human infection occurred) evidence. | 138 | |
| KSA (Jeddah)/14 November to 9 December 2013 | Serum Ab: 4-fold rise in paired sera in 2/9 (22.2%). Viral RNA: detected in nasal swabs of both camels (upE and ORF1a). | 128 | |
| KSA (Al-Hasa)/November 2013 to February 2014 (peak calving season) | Serum NAb: 280/310 (90.3%). Viral RNA: nasal > fecal specimens. Viral genome: highly stable with an estimated mutation rate of 0 nt substitutions per site per day. Clinical: both calves and adults could be infected; symptoms included mild respiratory symptoms (cough, sneezing, respiratory discharge), increased body temp, and decreased appetite; acute infection was not associated with prolonged viremia or viral shedding. | 129 | |
| UAE (Dubai)/2003 and 2013 | Serum Ab: 151/151 (100%) (2003) and 481/500 (96.2%) (2013); high titers of NAb > 1:640 in 509/651 (78.2%). | 124 | |
| UAE (Dubai)/February to October 2005 | Serum NAb: 9/11 (81.8%). | 125 | |
| Oman/March 2013 and Spain (Canary Islands)/April 2012 to May 2013 | Serum Ab: 50/50 (100%) of Omani and 15/105 (14.3%) of Spanish camels; all 50/50 (100%) of Omani (titers 1/320 to 1/2560) and 9/105 (9%) of Spanish camels had NAb (titers 1/20 to 1/320). | 121 | |
| Oman (countrywide)/December 2013 | Viral RNA: high concentrations in nasal and conjunctival swabs of 5/76 (6.6%) camels (≥2 gene targets). | 317 | |
| Jordan (al Zarqa governorate)/June to September 2013 | Serum NAb: 11/11 (100%). | 126 | |
| Qatar/17 October 2013 | Serum NAb: 14/14 (100%); titers 1/160 to 1/5120. Viral RNA: 5/14 (35.7%) nasal swabs by 3 gene targets (upE, N, and ORF1a), 1/14 (7.1%) by 2 gene targets, and 5/14 (35.7%) by 1 gene target. Viral genome: 3/5 samples shared 100% identity (357-nt S fragment) with sequences from 2 epidemiologically linked patients; further sequencing of 4.2-kb concatenated fragments of a camel strain and 2 epidemiologically linked patient strains; only 1 nt difference in ORF1a and 1 nt difference in ORF4b. | 133 | |
| Qatar (Doha)/February 2014 | Viral RNA: 1/53 (1.9%) nasal swabs from an 8-month-old camel (1/53, 1.9%) (upE and N). Viral genome: complete genome (MERS-CoV camel/Qatar_2_2014) shared 99.5% to 99.9% nt identities with other camel and patient strains. | 131 | |
| Qatar (Al Shahaniya and Dukhan)/April 2014 | Serum and milk Ab: all 33 camels had IgG in serum and milk. Viral RNA: detected in the nose swabs and/or feces of 7/12 camels and the milk of 5/7 of these camels in Al Shahaniya. | 144 | |
| KSA (Al Hasa, As Sulayyil, Hafar Al-Batin, Medina)/1993, Egypt/2014, and Australia (central Australia and Queensland)/2014 | Serum NAb: 118/131 (90.1%) of KSA camels and 0/25 (0%) of Australian camels. | 135 | |
| Africa | |||
| Camelus dromedarius | Somalia/1983 to 1984, Sudan/June and July 1983, Egypt/June and July 1997 | Serum NAb: Somalia (70/86, 81.4%), Sudan (49/60, 81.0%), and Egypt (34/43, 79.1%) by MNT. | 132 |
| Kenya/1992 to 2013 | Serum Ab: 213/774 (27.5%), including 119/774 (15.4%) with NAb; seropositive camels were found in all sampling sites throughout the study period; increased seroprevalence was significantly correlated with increased camel population density. | 130 | |
| Nigeria/2010 to 2011, Tunisia/2009 and 2013, and Ethiopia/2011 to 2013 | Serum Ab: Nigeria (94.0% of adults) and Ethiopia (93.0% of juveniles and 97.0% of adults); lower rates in Tunisia (54.0% of adults and 30.0% of juveniles). | 318 | |
| Egypt (Cairo and Qalyubia governorate)/June 2013 | Serum NAb: 103/110 (93.6% with titer ≥ 1:20) by MNT and 108/110 (98% with titer ≥ 1:20) by spike ppNT. | 122 | |
| Egypt (Alexandra, Cairo, and Nile Delta region)/June to December 2013 | Serum NAb: 48/52 (92.3% with titers between 1:20 and ≥1:640); 0/179 abattoir workers. Viral RNA: 4/110 (3.6%) nasal swabs (upE and ORF1a). | 134 |
aAbbreviations: aa, amino acid; concn, concentration; KSA, Kingdom of Saudi Arabia; N, nucleocapsid; NAb, neutralizing antibody; nt, nucleotide; ORF, open reading frame; MNT, microneutralization test; RT-PCR, reverse transcription-PCR; ppNT, pseudoparticle neutralization test; RBD, receptor-binding domain; RdRp; RNA-dependent RNA polymerase; S, spike; UAE, United Arab Emirates.