CVR - Coronavirus Vaccines R&D Roadmap

Bat

Alphacoronaviruses

AmalacovirusAlphacoronavirus almalfi

bat alphacoronavirus isolate AMA_L_F (BtCoV-AMA-L-F)

RI: AMA_L_F

RefSeq: MT663548.1

UnknownBatBergner 2020
ColacovirusAlphacoronavirus myotis

bat coronavirus CDPHE15 (BtCoV CDPHE15)

RI: bat/USA/CDPHE15/2006

RefSeq: NC_022103.1

UnknownBatDominguez 2007
DecacovirusAlphacoronavirus australiense

alphacoronavirus sp. WA3607 (ACoV-WA3607)

RI: WA3607

RefSeq: MK472070.1

UnknownBatPrada 2019
DecacovirusAlphacoronavirus ferrumequini

BtRf-AlphaCoV/HuB2013 (BtRf-AlphaCoV)

RI: BtRf-HuB2013

RefSeq: KJ473807.1

UnknownBatWu 2015
DecacovirusAlphacoronavirus hipposideri

Hipposideros pomona bat coronavirus CHB25 (HipPBCoV-CHB25)

RI: CHB0025

RefSeq: MN611525.1

UnknownBatLi 2020
DecacovirusAlphacoronavirus rousetti

Rousettus bat coronavirus HKU10 (Ro-BtCoV HKU10)

RI: 183A

RefSeq: JQ989270.1

UnknownBatLau 2012
DuvinacovirusAlphacoronavirus chicagoense

human coronavirus 229E (HCoV_229E)

RI: 229E

RefSeq: AF304460.1

APNHuman

Letko 2020

Thiel 2001

LuchacovirusAlphacoronavirus ratti

Lucheng Rn rat coronavirus (LRNV)

RI: Lucheng-19

RefSeq: KF294380.2

UnknownRodent

Wang 2020

Wang 2015

MinacovirusAlphacoronavirus neovisontis

mink coronavirus (MCoV)

RI: WD1127

RefSeq: HM245925.1

UnknownMinkVlasova 2011
MinunacovirusAlphacoronavirus miniopteri

Miniopterus bat coronavirus HKU8 (Mi-BatCoV_HKU8)

RI: AFCD77

RefSeq: EU420139.1

UnknownBatChu 2008
MinunacovirusAlphacoronavirus pusilli

Miniopterus bat coronavirus 1 (Mi-BatCoV_1A)

RI: AFCD62

RefSeq: EU420138.1

UnknownBatChu 2008
MyotacovirusAlphacoronavirus ricketti

BtMr-AlphaCoV/SAX2011 (BtMr-AlphaCoV)

RI: BtMr-SAX2011

RefSeq: KJ473806.1

UnknownBatWu 2015
NyctacovirusAlphacoronavirus chalinolobi

alphacoronavirus sp. WA2028 (ACoV-WA2028)

RI: WA2028

RefSeq: MK472068.1

UnknownBatPrada 2019
NyctacovirusAlphacoronavirus nyctali

BtNv-AlphaCoV/SC2013 (BtNv-AlphaCoV)

RI: BtNv-SC2013

RefSeq: KJ473809.1

UnknownBatWu 2015
NyctacovirusAlphacoronavirus pipistrelli

alphacoronavirus bat coronavirus (PK-BatCoV)

RI: P.kuhlii/Italy/3398/2015

RefSeq: MH938449.1

UnknownBatDe Sabato 2019
NyctacovirusAlphacoronavirus tylonycteridis

Tylonycteris bat coronavirus HKU33 (TyBCoV-HKU33)

RI: GZ151867

RefSeq: MK720944.1

UnknownBatLau 2019
PedacovirusAlphacoronavirus finnoniense

bat alphacoronavirus BtCoV/020_16/M.dau/FIN/2016 (BtCoV/020_16/M.dau/FIN/2016)

RI: 020_16

RefSeq: MG923574.2

UnknownBatKivistö 2020
PedacovirusAlphacoronavirus gouldii

alphacoronavirus sp. WA1087 (ACoV-WA1087)

RI: WA1087

RefSeq: MK472067.1

APN

APN+TMPRSS2

Bat

Gallo 2024

Prada 2019

PedacovirusAlphacoronavirus porci

porcine epidemic diarrhea virus (PEDV)

RI: CV777

RefSeq: AF353511.1

APNSwine

Bridgen 1993

Li 2007

Liu 2015

PedacovirusAlphacoronavirus scotophili

Scotophilus bat coronavirus 512 (Sc-BatCoV_512)

RI: 512/2005

RefSeq: DQ648858.1

APNBat

Chen 2019

Tang 2006

RhinacovirusAlphacoronavirus rhinolophi

Rhinolophus bat coronavirus HKU2 (Rh-BatCoV_HKU2)

RI: GD/430/2006

RefSeq: EF203064.1

UnknownBatLau 2007
RhinacovirusUnclassified Rhinacovirus

Swine acute diarrhoea syndrome coronavirus (SADS-CoV)

RI: HKU2-related isolate 160660

RefSeq: MN611522.1

UnknownSwineZhou 2018
SetracovirusAlphacoronavirus amsterdamense

human coronavirus NL63 (HCoV_NL63)

RI: Amsterdam I

RefSeq: AY567487.2

ACE2

ACE2+TMPRSS2

Human

Hofmann 2005

Gallo 2024

van der Hoek 2004

SetracovirusAlphacoronavirus triaenopis        

NL63-related bat coronavirus (BtKYNL63)

RI: BtKYNL63-9b

RefSeq: KY073745.1

UnknownBatTao 2017
SoracovirusAlphacoronavirus soricis

common shrew coronavirus Tibet-2014 (Sa-CoV_T14)

RI: N/A

RefSeq: KY370053.1

UnknownShrewWu 2018
SunacovirusAlphacoronavirus sunci

Suncus murinus coronavirus Xingguo-74 (Sm-CoV_X74)

RI: N/A

RefSeq: KY967715.1

UnknownShrewWang 2017
TegacovirusAlphacoronavirus suis

transmissible gastroenteritis virus (TGEV)

RI: PUR46-MAD

RefSeq: AJ271965.2

SA

APN

Swine

Almazán 2000

Delmas 1992

Schwegmann-Weßels 2011

TegacovirusAlphacoronavirus 1

canine coronavirus (SD-F3)

RI: N/A

RefSeq: OM451122.1

APN

APN+TMPRSS2

CanineGallo 2024
TegacovirusAlphacoronavirus 1

feline coronavirus (FCoV)

RI: FCoV

RefSeq: NC_002306.3

APNFelineTresnan 1996
Pangolin

Betacoronaviruses

EmbecovirusBetacoronavirus gravedinis

human coronavirus OC43 (HCoV_OC43)

RI: ATCC VR-759

RefSeq: AY585228.1

N-acetyl-9-O-acetylneuraminic acid

SA

Human

Krempl 1995

St-Jean 2004

Vlasak 1988

EmbecovirusBetacoronavirus hongkongense

human coronavirus HKU1 (HCoV_HKU1)

RI: HKU1

RefSeq: AY597011.2

TMPRSS2Human

Gao 2024

Saunders 2023

Woo 2005

EmbecovirusBetacoronavirus muris

murine hepatitis virus (MHV)

RI: A59

RefSeq: AY700211.1

CEACAM1Rodent

Coley 2005

Tsai 2003

Williams 1991

EmbecovirusBetacoronavirus myodae

Myodes rufocanus vole coronavirus 2/JL2014 (MrufCoV_2JL14)

RI: N/A

RefSeq: KY370046.1

UnknownRodentWu 2018
EmbecovirusUnclassified Embecovirus

Grimso virus

RI: GRP215-15

RefSeq: OM373090.1

UnknownRodentWasberg 2022
EmbecovirusBetacoronavirus ratti

betacoronavirus HKU24 (ChRCoV_HKU24)

RI: HKU24-R05005I

RefSeq: KM349742.1

UnknownRodentLau 2015
EmbecovirusBetacoronavirus 1

bovine coronavirus (BCoV-ENT)

RI: N/A

RefSeq: AF391541.1

SA

N-acetyl-9-O- acetylneuraminic acid

Bovine

Chouljenko 2001

Schultze 1994

Vlasak 1988

EmbecovirusBetacoronavirus 1

equine coronavirus (NC99)

RI: N/A

RefSeq: EF446615.1

UnknownEquineZhang 2007
EmbecovirusBetacoronavirus 1

porcine hemagglutinating encephalomyelitis virus (PHEV)

RI: PHEV/67N/US/1970

RefSeq: MW165134.1

DPEP1

Swine

Rodent

Dufloo 2025

Hirano 2004

HibecovirusBetacoronavirus hipposideri

bat Hp-betacoronavirus/Zhejiang2013 (Bat_Hp-BetaCoV)

RI: Zhejiang2013

RefSeq: KF636752.1

UnknownBatWu 2015
MerbecovirusBetacoronavirus cameli

Middle East respiratory syndrome-related coronavirus (MERS-CoV)

RI: HCoV-EMC

RefSeq: JX869059.2

DPP4

GRP78

Camel

Human

Chu 2018

Raj 2013

Soriano-Tordera 2025

MerbecovirusBetacoronavirus erinacei

hedgehog coronavirus 1 (EriCoV)

RI: 2012-174/GER/2012

RefSeq: KC545383.1

UnknownHedgehog

Corman 2014

Cruz 2024

MerbecovirusBetacoronavirus pipistrelli

Pipistrellus bat coronavirus HKU5 (Pi-BatCoV_HKU5)

RI: HKU5

RefSeq: EF065509.1

ACE2Bat

Catanzaro 2024

Park 2025

Wang 2025

Woo 2007

MerbecovirusBetacoronavirus tylonycteridis

Tylonycteris bat coronavirus HKU4 (Ty-BatCoV-HKU4)

RI: HKU4

RefSeq: EF065505.1

DPP4Bat

Wang 2014

Woo 2007

Yang 2014

MerbecovirusUnclassified Merbecovirus

Middle East respiratory syndrome-related coronavirus (NeoCoV)

RI: Neoromicia/PML-PHE1/RSA/2011

RefSeq: KC869678.4

ACE2BatXiong 2022
NobecovirusBetacoronavirus cororeum

Rousettus bat coronavirus (Ro-BatCoV_GCCDC1)

RI: GCCDC1 356

RefSeq: KU762338.1

UnknownBatHuang 2016
NobecovirusBetacoronavirus eidoli

Eidolon helvum bat coronavirus CMR704-P12 (Ei-BatCoV_C704)

RI: N/A

RefSeq: MG693168.1

UnknownBatYinda 2018
NobecovirusBetacoronavirus rousetti

Rousettus bat coronavirus HKU9 (Ro-BatCoV_HKU9)

RI: HKU9

RefSeq: EF065513.1

GRP78Bat

Chu 2018

Woo 2007

SarbecovirusBetacoronavirus pandemicum

severe acute respiratory syndrome coronavirus (SARS-CoV)

RI: Tor2

RefSeq: AY274119.3

ACE2Human

Li 2005

Li 2003

Prabakaran 2004

SarbecovirusBetacoronavirus pandemicum

SARS coronavirus PC4-227 (SARS-CoV)

RI: PC4-227

RefSeq: AY613950.1

UnknownPalm civetSong 2005
SarbecovirusBetacoronavirus pandemicum

severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2)

RI: Wuhan-Hu-1

RefSeq: MN908947.3

ACE2

GRP78

Human

Feline

Canine

Cervid

Rodent

Mink

Monkey

Rhino

Raccoon

and more

Brielle 2020

Carlos 2021

Han 2023

Wang 2020

Wu 2020

SarbecovirusBetacoronavirus pandemicum

severe acute respiratory syndrome-related coronavirus (SARSr-CoV)

RI: BtKY72

RefSeq: KY352407.1

ACE2Bat

Su 2024

Tao 2019

SarbecovirusBetacoronavirus pandemicum

Bat SARS-like coronavirus WIV1 (WIV1)

RI: WIV1

RefSeq: KF367457.1

ACE2BatGe 2013
SarbecovirusBetacoronavirus pandemicum

Bat SARS-like coronavirus (Rs4084)

RI: Rs4084

RefSeq: KY417144.1

ACE2BatLetko 2020
SarbecovirusBetacoronavirus pandemicum

Bat SARS-like coronavirus (Rs4231)

RI: Rs4231

RefSeq: KY417146.1

ACE2BatLetko 2020
SarbecovirusBetacoronavirus pandemicum

Bat SARS-like coronavirus (Rs7327)

RI: Rs7327

RefSeq: KY417151.1

ACE2BatLetko 2020
SarbecovirusBetacoronavirus pandemicum

SARS coronavirus Urbani (RsSHC014)

RI: SHC014

RefSeq: KC881005.1

ACE2Bat

Ge 2013

Letko 2020

SarbecovirusBetacoronavirus pandemicum

Bat coronavirus RaTG13n (RaTG13)

RI: RaTG13

RefSeq: MN996532.2

ACE2Bat

Li 2021

Liu 2021

SarbecovirusUnclassified Sarbecovirus

Bat coronavirus BM48-31/BGR/2008 (BM48-31)

RI: BM48-31/BGR/2008

RefSeq: GU190215.1

ACE2BatKosugi 2025
SarbecovirusUnclassified Sarbecovirus

Sarbecovirus RhGB01 (RhGB01)

RI: RhGB01

RefSeq: MW719567.1

ACE2BatKosugi 2025
SarbecovirusUnclassified Sarbecovirus

Sarbecovirus sp (YN2020H)

RI: YN2020H

RefSeq: OK017858.1

ACE2BatKosugi 2025
SarbecovirusBetacoronavirus pandemicum

Civet SARS CoV 007/2004 (Civet007)

RI: 007/2004

RefSeq: AY572034.1

ACE2CivetKosugi 2025
SarbecovirusBetacoronavirus pandemicum

Bat SARS-like coronavirus (Rs7327)

RI: Rs7327

RefSeq: KY417151.1

ACE2BatKosugi 2025
SarbecovirusUnclassified Sarbecovirus

Sarbecovirus sp (YN2016A)

RI: RsYN2016A

RefSeq: OK017847.1

ACE2BatKosugi 2025
SarbecovirusBetacoronavirus pandemicum

Rhinolophus affinis coronavirus (LYRa11)

RI: LYRa11

RefSeq: KF569996.1

ACE2BatKosugi 2025
SarbecovirusBetacoronavirus pandemicum

SARS-like coronavirus WIV16 (WIV16)

RI: WIV16

RefSeq: KT444582.1

ACE2BatKosugi 2025
SarbecovirusUnclassified Sarbecovirus

Sarbecovirus sp (Rc-os20)

RI: Rc-os20

RefSeq: LC663958.1

ACE2BatKosugi 2025
SarbecovirusBetacoronavirus pandemicum

severe acute respiratory syndrome-related coronavirus (

RI: Rc-o319

RefSeq: LC556375.1

ACE2BatKosugi 2025
SarbecovirusUnclassified Sarbecovirus

Sarbecovirus sp (Rc-kw8)

RI: Rc-kw8

RefSeq: LC663793.1

ACE2BatKosugi 2025
SarbecovirusUnclassified Sarbecovirus

Sarbecovirus sp (Rc-mk2)

RI: Rc-mk2

RefSeq: LC663959.1

ACE2BatKosugi 2025
SarbecovirusUnclassified Sarbecovirus

Pangolin coronavirus (GX-P1E)

RI: Guangzi strain

RefSeq: MT040334.1

ACE2PangolinKosugi 2025
SarbecovirusBetacoronavirus pandemicum

Horseshoe bat sarbecovirus (Ra22QT77)

RI: Ra22QT77

RefSeq: OR233324.1

ACE2BatKosugi 2025
SarbecovirusBetacoronavirus pandemicum

Pangolin coronavirus (MP789)

RI: Guangdong strain

RefSeq: MT121216.1

ACE2PangolinKosugi 2025
SarbecovirusUnclassified Sarbecovirus

Horseshoe bat sarbecovirus (Rp22DB159)

RI: Rp22DB159

RefSeq: OR233302.1

ACE2BatKosugi 2025
Canada goose

Gammacoronaviruses

BrangacovirusGammacoronavirus brantae

Canada goose-Branta canadensis-coronavirus-Cambridge Bay 2017 (BcanCoV_CB17)

RI: CGCoV

RefSeq: MK359255.1

UnknownAvianPapineau 2019
CegacovirusGammacoronavirus delphinapteri

beluga whale coronavirus (BWCoV)

RI: SW1

RefSeq: EU111742.1

UnknownBeluga whaleMihindukulasuriya 2008
CegacovirusGammacoronavirus delphinapteri

bottlenose dolphin coronavirus HKU22 (BdCoV_HKU22)

RI: HKU22

RefSeq: KF793824.1

UnknownDolphinWang 2020
IgacovirusGammacoronavirus anatis

duck coronavirus DK/GD/27/2014 (DuCoV)

RI: DuCoV_2714

RefSeq: KM454473.1

UnknownAvianZhuang 2015
IgacovirusGammacoronavirus galli

infectious bronchitis virus (IBV)

RI: IBV

RefSeq: M95169.1

SAAvian

Boursnell 1987

Winter 2006

IgacovirusGammacoronavirus pulli

infectious bronchitis virus (ACoV_9203)

RI: ACoV_9203

RefSeq: KR902510.1

UnknownAvianShilpa 2022
Moorhen

Deltacoronaviruses

AndecovirusDeltacoronavirus marecae

wigeon coronavirus HKU20 (WiCoV_HKU20)

RI: HKU20-9243

RefSeq: JQ065048.1

UnknownAvianWoo 2012
BuldecovirusDeltacoronavirus gallinulae

common moorhen coronavirus HKU21 (CMCoV_HKU21)

RI: HKU21-8295

RefSeq: JQ065049.1

UnknownAvianWoo 2012
BuldecovirusDeltacoronavirus lonchurae

munia coronavirus HKU13 (MunCV_HKU13)

RI: HKU13-3514

RefSeq: FJ376622.1

APNAvian

Liang 2023

Woo 2009

BuldecovirusDeltacoronavirus pycnonoti

Bulbul coronavirus HKU11 (BulCV_HKU11)

RI: HKU11

RefSeq: FJ376619.2

APNAvian

Liang 2023

Woo 2009

BuldecovirusDeltacoronavirus suis

porcine coronavirus HKU15 (PoCoV_HKU15)

RI: HKU15-155

RefSeq: JQ065043.2

APNSwine; Human

Lednicky 2021

Li 2018

Woo 2012

BuldecovirusDeltacoronavirus zosteropis

white-eye coronavirus HKU16 (WECoV_HKU16)

RI: HKU16

RefSeq: JQ065044.1

UnknownAvianWoo 2012
BuldecovirusUnclassified Buldecovirus

sparrow coronavirus HKU17 (HKU17)

RI: HKU17

RefSeq: JQ065045.1

APNAvianLiang 2023
HerdecovirusDeltacoronavirus nycticoracis

night heron coronavirus HKU19 (NHCoV_HKU19)

RI: HKU19

RefSeq: JQ065047.1

UnknownAvianWoo 2012
Bat

Unclassified Coronaviruses

UnclassifiedUnclassified

Bat coronavirus (BANAL-20-52)

RI: BANAL-20-52

RefSeq: MZ937000.1

ACE2Bat

Kosugi 2025

Temmam 2022

UnclassifiedUnclassified

Bat coronavirus (BANAL-20-236)

RI: BANAL-20-236

RefSeq: MZ937003.2

ACE2Bat

Kosugi 2025

Temmam 2022

Coronaviruses

Human Tissue Distribution

ACE2Intestinal enterocytes, renal proximal tubules, testis, alveolar type II cells (low)

Hikmet 2020

López-Cortés 2023

DPP4Lung alveolar epithelium, kidney proximal tubules, liver hepatocytes, T cells

Hikmet 2020

Koyanagi 2023

Luu 2023

Nassar 2021

Seys 2017

ANPEPIntestinal epithelium, kidney, airway epithelial cells, neutrophils

Hikmet 2020

Lendeckel 2023

9-O-Acetylated Sialic AcidsRespiratory mucins, colon epithelial cells, CNS

Barnard 2019

Hikmet 2020

Wasik 2017

TMPRSS2Stomach, pancreas, lungs, small intestine, salivary glands, kidneys, prostate, parathyroid glandCao 2021
Marmoset

Coronavirus Receptors in Animal Models

MiceNot naturally permissive due to differences in ACE2 sequence. Transgenic hACE2-expressing mice or mouse-adapted SARS-CoV-2 strains are used for research. High-dose SARS-CoV-1 infection in BALB/c mice can overcome poor binding to a degree, epseially with mouse-adapted strains.Not naturally susceptible to MERS-CoV due to differences in DPP4. Introducing human-like mutations (A288L and T330R) or using transgenic hDPP4 mice allows infection.Not naturally susceptible to HCoV-229E due to differences in ANPEP. However, transgenic hANPEP-expressing mice allow infection and pathogenesis studies.Express 9-O-acetylated sialic acids, making them susceptible to some betacoronaviruses, though infection is generally limited.
HamstersSupport SARS-CoV-2 infection and exhibit mild to moderate lung pathology. However, ACE2 expression is limited in the trachea and lung, affecting viral tropism.Do not naturally support MERS-CoV infection due to receptor incompatibility.Susceptibility to HCoV-229E is unclear; no strong evidence supports infection.Express 9-O-acetylated sialic acid and can be infected by betacoronaviruses.
FerretsExpress ACE2 mainly in the upper respiratory tract, leading to mild SARS-CoV-2 infections without severe lung pathologyNot permissive to MERS-CoV due to DPP4 sequence differences.Limited data on ferret ANPEP expression and susceptibility to HCoV-229E.Likely express 9-O-acetylated sialic acids, but data is limited.
Non-Human PrimatesACE2 highly similar to humans, making them good models for SARS-CoV-2. Most show mild to moderate disease.Rhesus macaques develop mild to moderate MERS-CoV infections, whereas African green monkeys show more severe disease.NHPs likely support HCoV-229E infection due to ANPEP compatibility, but data is limited.Expression of 9-O-acetylated sialic acids in NHPs is species-dependent.
Common MarmosetsExpress ACE2 and are susceptible to SARS-CoV-2, but disease severity varies.Develop more severe MERS-CoV infections than rhesus macaques, making them a valuable model.Limited data on ANPEP distribution in marmosets.Limited data on marmoset expression of 9-O-acetylated sialic acids.

Coronavirus Receptors in Animal Models: References

Cockrell 2014: Mouse dipeptidyl peptidase 4 is not a functional receptor for Middle East respiratory syndrome coronavirus infection

Gong 2018: The battle against SARS and MERS coronaviruses: reservoirs and animal models

Gretebeck 2015: Animal models for SARS and MERS coronaviruses

Lassnig 2005: Development of a transgenic mouse model susceptible to human coronavirus 229E

Lean 2021: Differential susceptibility of SARS-CoV-2 in animals: evidence of ACE2 host receptor distribution in companion animals, livestock and wildlife by immunohistochemical characterisation

Lin 2022: Animal models for studying coronavirus infections and developing antiviral agents and vaccines

Liu 2023: Mouse models susceptible to HCoV-229E and HCoV-NL63 and cross protection from challenge with SARS-CoV-2

Pandey 2020: Animal models for SARS-CoV-2 research: a comprehensive literature review

Qin 2022: Review of selected animal models for respiratory coronavirus infection and its application in drug research

Roberts 2008: Animal models and vaccines for SARS-CoV infection

Saravanan 2022: Animal models for SARS-CoV-2 and SARS-CoV-1 pathogenesis, transmission and therapeutic evaluation

Singh 2020: A comprehensive review of animal models for coronaviruses: SARS-CoV-2, SARS-CoV, and MERS-CoV

Snouwaert 2023: Human ACE2 expression, a major tropism determinant for SARS-CoV-2, is regulated by upstream and intragenic elements

Sutton 2015: Development of animal models against emerging coronaviruses: from SARS to MERS coronavirus

van Doremalen 2015: Animal models of Middle East respiratory syndrome coronavirus infection

Yuan 2020: Animal models for emerging coronavirus: progress and new insights