Table 2.
Cryptosporidium species and genotypes identified by molecular tools in wild terrestrial mammals and their zoonotic importance.
| Cryptosporidium species/genotypes | Wildlife hosts | Zoonotic importance | gp60 subtypes reported in wildlife | References |
|---|---|---|---|---|
| C. hominis | Fallow deer (Dama dama), Dugong (Dugong dugon), Chinchillas (Chinchilla lanigera), Baboons (Pabio anubis), Chimpanzees (Pan troglodytes schweinfurthii), Red colobus (Procolobus rufomitratus), Black-and-white colobus (Colobus guereza), Rhesus macaque (Macaca mulatta), Cynomolgus monkey (Macaca fascicularis), Francois' leaf monkey (Trachypithecus francoisi), Lemurs (Lemur sp.), Bandicoots (Isoodon obesulus), Bushtail possums (Trichosurus vulpecula), Estern grey kangaroos (Macropus giganteus), Brush-tailed rock-wallabies (Petrogale penicillata), Wild dingo (Canis lupus dingo), Squirrel monkey (Saimiri sciureus) | Main Cryptosporidium species infecting humans | IbA12G3, IbA10G2R2, IiA17, IfA12G2, IaA13R7, IaA13R8, IaA14R7, IdA20, IeA11G3T3, IfA16G2, IkA7G4 (novel subtype) | Morgan Ryan et al., 2002, Salyer et al., 2012, Ye et al., 2012, Ng et al., 2011, Dowle et al., 2013, Nolan et al., 2013, Karim et al., 2014, Ryan et al., 2014, Liu et al., 2015b, Parsons et al., 2015, Zahedi et al., 2015 |
| C. parvum | Alpaca (Lama pacos), Swamp deer (Cervus duvauceli), Red deer (Cervus elaphus), Roe deer (Capreolus capreolus), Fallow deer (Dama dama), Addaxes (Addax nasomaculatus), Arabian oryx (Oryx leucoryx), Gemsboks (Oryx gazella), Sable antelopes (sable antelopes), White-tailed deer (Odocoileus Virginianus), Game grey wolves (Canis lupus), Racoon dog (Nyctereutes procyonoides viverrinus), Rabbit (Oryctolagus cuniculus), Nutria (Myocastor coypus), Prezewalski's wild horse (Equus przewalskii), Alpaca (Lama quanico pacos), Eastern grey squirrel (Sciurus carolinensis), Ground Squirrels (Spermophilus beecheyi), Siberian chipmunk (Tamias sibiricus), Hamsters (Cricetinae), Wood mice (Apodemus sylvaticus), White-footed mouse (Peromyscus leucopus), Yellow-bellied marmot (Marmota flaviventris), Bamboo rats (Rhizomys sinensis), Small brown bat (Myotis lucifugus), Campbell hamster (Phodopus campbelli), Golden hamster (Mesocricetus auratus), Capybara (Hydrochoerus hydrochaeris), Racoon dog (Nictereutes procyonoides viverrinus), Red fox (Vulpes vulpes), Rhesus macaques (Macaca mulatta), Toque macaques (Macaca sinica sinica), Grey langurs (Semnopithecus priam thersites), Purple-faced langurs (Trachypithecus vetulus philbricki), Common dolphins (Delphinus delphis), Golden takins (Budorcas taxicolor bedfordi), Eastern grey kangaroos (Macropus giganteus), Asian house rat (Rattus tanezumi), Brown rat (Rattus norvegicus), Bamboo rats (Rhizomys sinensis) | Major | IIdA15G1, IIdA18G1, IIdA19G1, IIaA15G2R1, IIaA19G2R1, IIaA19G3R1, IIaA19G4R1, IIaA20G3R1, IIaA20G3R2, IIaA20G4R1, IIaA21G3R1, IIaA16G2R1, IIaA14G1R1, IIaA14G2R1, IIaA16G3R1, IIcA5G3, IIcA5G3a, IIoA13G1, IIpA9 (novel subtype) | Morgan et al., 1999a, Atwill et al., 2001, Perez and Le Blancq, 2001, Matsui et al., 2000, Matsubayashi et al., 2004, Ryan et al., 2003, Ryan et al., 2004, Ekanayake et al., 2007, Feng et al., 2007, Meireles et al., 2007, Paziewska et al., 2007, Starkey et al., 2007, Ziegler et al., 2007, Cinque et al., 2008, Lv et al., 2009, Feng, 2010, Gómez-Cousoa et al., 2012, Ravaszova et al., 2012, Ye et al., 2012, Dowle et al., 2013, Nolan et al., 2013, Liu et al., 2014, Lv et al., 2009, Reboredo-Fernández et al., 2014, Bodager et al., 2015, Liu et al., 2015a, Montecino-Latorre et al., 2015, Qi et al., 2015a, Wang et al., 2015, Wells et al., 2015; Zahedi et al., 2015; Zhao et al., 2015a, Zhao et al., 2015b |
| C. cuniculus | European rabbit (Oryctolagus cuniculus), Eastern grey kangaroo (Macropus giganteus) (single report) | Responsible for several waterborne outbreaks and sporadic cases of cryptosporidiosis in the UK and has been identified in a human in Australia | VaA18,VbA18, VbA19, VbA21, VaA22, VbA24, VbA26, VbA29, VbA32, VbA22R4, VbA23R3, VbA24R3, VbA25R4,VbA26R4 | Xiao et al., 2002a, Ryan et al., 2003, Chalmers, 2012, Nolan et al., 2010, Robinson et al., 2010, Elwin et al., 2012, Zhang et al., 2012, Nolan et al., 2013, Kaupke et al., 2014, Koehler et al., 2014, Liu et al., 2014, Puleston et al., 2014 |
| C. ubiquitum | Swamp deer (Cervus duvauceli), Deer mouse (Peromyscus), Eastern grey squirrels (Sciurus carolinensis), Red squirrel (Sciurus vulgaris), Eastern chipmunk (Tamias striatus), Lemur (Lemuroidea), North American beaver (Castor canadensis), Woodchuck (Marmota monax), Raccoon (Procyon lotor), White-tailed deer (Odocoileus virginianus), Sika deer (Cervus Nippon), Roe deer (Capreolus capreolus), Blesbok (Damaliscus pygargus phillipsi), Ibex (Capara sibirica), Nyala (Niyala anagasii), Coquerel's sifaca (Propithecus coquereli), Large Japanese field mouse (Apodemus speciosus), Foxes | Emerging human pathogen | XIIa, XIIb, XIIc, XIId, XIIe, XIIf | Perez and Le Blancq, 2001, da Silva et al., 2003, Ryan et al., 2003, Feng et al., 2007, Karanis et al., 2007, Ziegler et al., 2007, Wang et al., 2008, Fayer et al., 2010, Cinque et al., 2008, Feng, 2010, Robinson et al., 2010, Robinson et al., 2011, Feng et al., 2012, Abu Samraa et al., 2013, Nolan et al., 2013, Murakoshi et al., 2013, Li et al., 2014, Ma et al., 2014, Perec-Matysiak et al., 2015, Qi et al., 2015a, Qi et al., 2015b, Stenger et al., 2015b |
| C. muris | Wild rats (Rattus sp.), Mice (Mus sp.), Greater bilblies (Macroties lagotis), Girrafes house mice (Mus musculus), Eastern grey squirrel (Sciurus carolinensis), Golden hamster (Mesocricetus auratus), Rock hyrax (Procavia capensis), Large footed mouse-eared bat (Myotis adversus), Japanese field mouse (Apodemus argenteus), Bilbies (Macrotis lagotis), Bank voles (Clethrionomys glareolus), Campbell hamster (Phodopus campbelli), Siberian hamster (Phodopus sungorus), Golden hamster (Mesocricetus auratus), Mountain goats (Oreamnos americanus), Cynomolgus monkeys (Macaca fascicularis), East African mole rat (Tachyoryctes splendens), Ringed seal (Pusa hispida), Donkey (Giraffa camelopardalis), Ringed seal (Phoca hispida), Large Japanese field mouse (Apodemus speciosus), Cynomolgus monkey (Macaca fascicularis), Slow loris (Nycticebus coucang), Ostriches (Struthio camelus), Mountain gorillas (Gorilla beringei beringei), Asian house rat (Rattus tanezumi), Brown rat (Rattus norvegicus), House mouse (Mus musculus) | Numerous reports in humans | Chalmers et al., 1997, Hurkova et al., 2003 | Dubey et al., 2002, Morgan et al., 1999a, Xiao et al., 2002a, Xiao et al., 2004b, Warren et al., 2003, Nakai et al., 2004, Hikosaka and Nakai, 2005, Santín et al., 2005, Azami et al., 2007, Brikan et al., 2008, Kváč et al., 2008, Lupo et al., 2008, Lv et al., 2009, Kodádková et al., 2010, Feng, 2010, Murakoshi et al., 2013, Yang et al., 2011, Yang et al., 2013; Ng-Hublin et al., 2013, Karim et al., 2014, Qi et al., 2014, Sak et al., 2014, Du et al., 2015, Laatamna et al., 2015, Petrincová et al., 2015, Zhao et al., 2015b |
| C. andersoni | Bacterian camel (Camelus bactrianus), European wisent (Bison bonasus), Marmots Campbell hamster (Phodopus campbelli), Golden hamster (Mesocricetus auratus), Golden takins (Budorcas taxicolor bedfordi), Giant panda (Ailuropoda melanoleuca), Macaca mulatta (Rhesus macaque), American mink (Mustela vison) | Minor | – | Matsubayashi et al., 2005, Wang et al., 2008, Lv et al., 2009, Stuart et al., 2013, Du et al., 2015, Wang et al., 2015, Zhao et al., 2015a |
| C. felis | Rhesus macaques (Macaca mulatta); Pallas's cat (Felis Manul) | Numerous reports in humans | – | Lucio-Forster et al., 2010, Ye et al., 2012, Beser et al., 2015, Ebner et al., 2015, Li et al., 2015c |
| C. canis dog genotype | Unidentified fox, Coyote (Canis latrans) | Numerous reports in humans | – | Xiao et al., 2002a, Ryan et al., 2004, Zhou et al., 2004, Trout et al., 2006, Ziegler et al., 2007, Elwin et al., 2012; Koompapong et al., 2014 |
| C. canis fox genotype | Fox | No reports in humans to date | – | Zhou et al., 2004, Swaffer et al., 2014 |
| C. canis coyote genotype | Coyotes | No reports in humans to date | – | Xiao et al., 2002a, Zhou et al., 2004 |
| C. erinacei | European hedgehog (Erinaceus europaeus), Horses | One report in humans | XIIIaA21R11, XIIIaA22R9, XIIIaA21R10, XIIIa20R10, XIIIaA19R12, XIIIaA22R11 | Dyachenko et al., 2010, Laatamna et al., 2013, Nolan et al., 2013, Kváč et al., 2014a, Kváč et al., 2014b, Meredith and Milne, 2009 |
| C. fayeri | Southern brown bandicoot (Isodon obesulus), Western-barred bandicoot (Permeles bougainville), Koala (Phascolarctos cincerus), Red kangaroo (Macropus rufus), Eastern grey kangaroo (Macropus giganteus), Yellow footed rock wallaby (Petrogale xanthopus), Western grey kangaroo (Macropus fuliginosus), Koalas (Phascolarctos cinereus) | Minor | IVaA9G4T1R1, IVaA10, IVaA7, IVbA9G1T1, IVcA8G1T1, IVdA7G1T1, IVfA12G1T1 | Power et al., 2005, Ryan et al., 2008; Yang et al., 2008, Yang et al., 2011, Power, 2010, Waldron et al., 2010, Feng et al., 2011b, Nolan et al., 2013, Swaffer et al., 2014, Vermeulen et al., 2015 |
| Opossum genotype I (C. fayeri) | Opossum (Didelphimorphia) | No reports in humans to date | XIaA4G1T1 | Feng et al., 2011b |
| Opossum genotype II | Virginia opossum (Didelphis virginiae) | No reports in humans to date | – | Xiao et al., 2002b, Oates et al., 2012 |
| C. meleagridis | Mountain gorillas (Gorilla beringei beringei), Brush-tailed rock wallabies (Petrogale penicillata), deermouse (Peromyscus sp.) | Major | IIIbA, IIIgA (closest match to IIIeA19G2R1) | Morgan et al., 2000, Cama et al., 2003, Gatei et al., 2006, Leoni et al., 2006, Muthusamy et al., 2006, Feng et al., 2007, Elwin et al., 2012, Silverlås et al., 2012, Kurniawan et al., 2013, Adamu et al., 2014, Ryan and Xiao, 2014, Ghaffari and Kalantari, 2014, Sak et al., 2014, Rahmouni et al., 2014, Wang et al., 2014, Stensvold et al., 2015, Vermeulen et al., 2015 |
| C. tyzerri | Mice (Mus musculus), Brown rats (Rattus norvegicus), Large-footed bat (Myotus adversus), Yellow-necked mouse (Apodemus flavicollis), Bank vole (Myodes glareolus), Common vole (Microtus arvalis), Red panda (Ailurus fulgens), Leopard (Panthera pardus), Takin (Budorcas taxicolor), Prairie bison (Bison bison), Lesser panda (Ailurus fulgens), Black leopards (Pantera pardus), Bobcats (Lynx rufus) | Occasionally reported in humans | IXaA5R2, IXaA6R1, IXaA6R2, IXaA6R3, IXbA6, IXbA6R2 | Morgan et al., 1999a, Xiao et al., 2002a; Bajer et al., 2003; Alves et al., 2005, Foo et al., 2007, Karanis et al., 2007, Ziegler et al., 2007, Lv et al., 2009, Feng et al., 2011b, Carver et al., 2012, Kváč et al., 2012, Ren et al., 2012, Rasková et al., 2013, Silva et al., 2013, Swaffer et al., 2014 |
| C. macropodum | Red kangaroo (Macropus rufus), Eastern grey kangaroo (Macropus giganteus), Swamp wallaby (Wallabia bicolor), Western grey kangaroos (Macropus fuliginosus) | No reports in humans to date | – | Power et al., 2004, Power et al., 2005, Power and Ryan, 2008, Power, 2010, Yang et al., 2011, Nolan et al., 2013 |
| C. bovis | Yaks, foxes, Gorillas (single report), Roe deer (Capreolus capreolus) | Occasionally reported in humans | – | Robinson et al., 2011, Helmy et al., 2013, García-Presedo et al., 2013b, Sak et al., 2013, Qi et al., 2015b, Qin et al., 2014 |
| C. ryanae | Roe deer (Capreolus capreolus), Water buffaloes (Bubalus bubalis) | No reports in humans to date | – | Feng et al., 2012, García-Presedo et al., 2013b |
| C. wrairi | Guinea pig (Cavia porcellus), California ground squirrels (Spermophilus beecheyi) | No reports in humans to date | VIIaA13T1, VIIaA17T1, VIIaA16T1 | Atwill et al., 2004, Feng et al., 2007, Feng et al., 2011b, Lv et al., 2009 |
| C. scrofarum | Asian house rat (Rattus tanezumi), Brown rat (Rattus norvegicus), Eurasian wild boars (Sus scrofa) | Occasionally reported in humans | – | García-Presedo et al., 2013a, Ng-Hublin et al., 2013, Němejc et al., 2013, Bodager et al., 2015, Parsons et al., 2015 |
| C. suis | Chimpanzees (Pan troglodytes schweinfurthii), Eurasian wild boars (Sus scrofa), Rodents | Occasionally reported in humans | – | Němejc et al., 2012, Němejc et al., 2013, Bodager et al., 2015, Parsons et al., 2015 |
| C. suis-like | Asian house rat (Rattus tanezumi) | No reports in humans to date | – | Ng-Hublin et al., 2013 |
| C. rubeyi | California ground squirrel (S. beecheyi), Belding's ground squirrel (S. beldingi), Golden Mantled ground squirrel (S. lateralis) |
No reports in humans to date | – | Pereira et al., 2010, Li et al., 2015a |
| Bear genotype | Black bear (Ursus americanus) | No reports in humans to date | – | Xiao et al., 2000 |
| Bat genotype I | Chinese rufous horseshoe bat (Rhinolophus sinicus), Stoliczka's trident bat (Aselliscus stoliczkanus) | No reports in humans to date | – | Wang et al., 2013b |
| Bat genotype II | Chinese rufous horseshoe bat (Rhinolophus sinicus), Fulvus roundleaf bat (Hipposideros fulvus), Leschenault's rousette (Rousettus leschenaultii) | No reports in humans to date | – | Wang et al., 2013b |
| Bat genotype III | Big brow bat (Eptesicus fuscus) | No reports in humans to date | – | Kváč et al., 2015 |
| Bat genotype IV | Western barbastelle (Barbastella barbastellus) | No reports in humans to date | Kváč et al., 2015 | |
| Beaver genotype | North American beaver (Castor canadensis) | No reports in humans to date | – | Feng et al., 2007 |
| Brushtail possum I | Brushtail possum (Trichasuris vulpecula) | No reports in humans to date | – | Hill et al., 2008 |
| Chipmunk genotype I | Chipmunk sp. (Tamias sp.), Eastern grey squirrel (Sciurus carolinensis), Deer mice (Peromyscus maniculatus) | Emerging human pathogen | XIVaA18G2T1, XIVaA18G2T2 | Jiang et al., 2005, Feltus et al., 2006, Feng et al., 2007, ANOFEL Cryptosporidium National Network, 2010, Insulander et al., 2013, Lebbad et al., 2013, Guo et al., 2015 |
| Chipmunk genotype II | Eastern chipmunk (Ramias striatus) | No reports in humans to date | – | Feng et al., 2007, Stenger et al., 2015a |
| Chipmunk genotype III | Siberian chipmunk (Tamias sibiricus) | No reports in humans to date | – | Lv et al., 2009 |
| Deer mouse genotype I | Deer mouse (Peromyscus) | No reports in humans to date | – | Xiao et al., 2002b, Feng et al., 2007, Feng et al., 2011b |
| Deer mouse genotype II | Deer mouse (Peromyscus) | No reports in humans to date | – | Xiao et al., 2002b, Feng et al., 2007 |
| Deer mouse genotype III | Deer mouse (Peromyscus) | No reports in humans to date | – | Feng et al., 2007, Stenger et al., 2015b |
| Deer mouse genotype IV | Deer mouse (Peromyscus) | No reports in humans to date | – | Feng et al., 2007 |
| Ferret genotype | Ferret (Mustelidae), Siberian chipmunk (Tamias sibiricus), River otters (Lontra canadensis), Black-footed ferret (Mustela nigripes), Red squirrel (Sciurus vulgaris) | No reports in humans to date | VIIIaA5G2 | Xiao et al., 2002a; Abe and Iseki, 2003, Gaydos et al., 2007; Kváč et al., 2008, Lv et al., 2009, Feng et al., 2011b |
| Giant panda genotype | Giant panda (Ailuropoda melanoleuca) | No reports in humans to date | – | Liu et al., 2013 |
| Squirrel genotypes I–III | Golden-mantled ground squirrels (Callospermophilus lateralis), Belding's ground squirrels (Urocitellus beldingi), California ground squirrels (Otospermophilus beecheyi), Black-tailed prairie dog (Cynomys ludovicianus) | No reports in humans to date | – | Atwill et al., 2004, Pereira et al., 2010, Stenger et al., 2015b |
| Hamster genotype | Siberian hamster (Phodopus sungorus) | No reports in humans to date | – | Lv et al., 2009 |
| Horse genotype | Przewalski's wild horse (Equus przewalski), Four-toed hedgehog (Atelerix albiventris) | Identified in humans in the UK | VIaA11G3, VIbA13 | Ryan et al., 2003, Robinson et al., 2008, Abe and Matsubara, 2015 |
| Mink genotype | River otter (Lontra canadensis), American minks (Mustela vison), Ermine (Mustela ermine) | Several reports in humans | XaA5G1 | Feng et al., 2007, Wang et al., 2008, Feng et al., 2011b, Ng-Hublin et al., 2013, Stuart et al., 2013, Ebner et al., 2015 |
| Mouse genotype II Mouse genotype III |
House mouse (Mus musculus) House mouse (Mus musculus) |
No reports in humans to date No reports in humans to date |
– – |
Foo et al., 2007, Silva et al., 2013 Silva et al., 2013 |
| Muskrat genotype I | Muskrat (Ondatra zibethicus), Boreal red-backed vole (Myodes rutilus) | No reports in humans to date | – | Xiao et al., 2002a, Zhou et al., 2004, Feng et al., 2007 |
| Muskrat genotype II | Muskrat (Ondatra zibethicus), Red fox (Vulpus vulpus), Deer mouse (Peromyscus maniculatus), Meadow vole (Microtus pennsylvanicus) | No reports in humans to date | – | Ziegler et al., 2007, Robinson et al., 2011 |
| Naruko genotype | Large Japanese field mouse (Apodemus speciosus) | No reports in humans to date | – | Murakoshi et al., 2013 |
| Rat genotype I | Brown rat (Rattus norvegicus) | No reports in humans to date | – | Ng-Hublin et al., 2013 |
| Rat genotype II | Brown rat (Rattus tanezum), Wild black rat (Rattus rattus), Brown rat (Rattus norvegicus). | No reports in humans to date | – | Lv et al., 2009, Paparini et al., 2012, Ng-Hublin et al., 2013, Silva et al., 2013 |
| Rat genotype III | Asian house rat (Rattus tanezumi), Wild black rat (Rattus rattus). | No reports in humans to date | – | Lv et al., 2009, Paparini et al., 2012, Ng-Hublin et al., 2013, Silva et al., 2013 |
| Rat genotype IV | Tanezumi rat (Rattus tanezumi), Asian house rat (Rattus tanezumi), Brown rat (Rattus norvegicus) | No reports in humans to date | – | Ng-Hublin et al., 2013 |
| Seal genotypes I and II | Ringed seals (Phoca hispida), Harbour seals (Phoca vituline), Hooded seal (Cystophora cristata) | No reports in humans to date | – | Santín et al., 2005, Bass et al., 2012 |
| Seal genotype III | Harp seal (Pagophilus groenlandicus) | No reports in humans to date | – | Bass et al., 2012 |
| Seal genotype IV (similar to skunk genotype) | Southern elephant seal (Mirounga leonina) | No reports in humans to date | – | Rengifo-Herrera et al., 2011, Rengifo-Herrera et al., 2013 |
| Seal genotype V (Weddell seal genotype) | Weddel seal (Leptonychotes weddellii) | No reports in humans to date | – | Rengifo-Herrera et al., 2013 |
| Shrew genotype | Northern short-tailed shrew (Blarina brevicauda), Wildebeest (Connochaetes), White-toothed shrew (Crocidura russula), Common shrew (Sorex araneus), Masked shrew (Sorex scinereus), Pygmy shrew (Sorex minutus), Brewer's mole (Parascalops brewer), Ermine (Mustela ermine) | No reports in humans to date | – | Torres et al., 2000; Alves et al., 2005, Feng et al., 2007, Ziegler et al., 2007 |
| Skunk/skunk-like genotype | Striped skunk (Mephitis mephitis), Raccoon (Procyon lotor), Eastern grey squirrel (Sciurus carolinensis), River otter (Lontra canadensis), Raccoon (Procyon lotor), Southern elephant seal (Mirounga leonina), Raccoon (Procyon lotor), Shunk (Mephitis mephitis), American red (Tamiasciurus hudsonicus), Fox squirrel (Sciurus niger) | Has been reported in humans | – | Xiao et al., 2002b, Zhou et al., 2004, Feng et al., 2007, Ziegler et al., 2007, Robinson et al., 2008, Chalmers et al., 2009, Feng et al., 2011b, Rengifo-Herrera et al., 2011, Elwin et al., 2012, Stenger et al., 2015b |
| Vole genotype | Meadow vole (Microtus pennynsylvanicus) | No reports in humans to date | – | Feng et al., 2007 |
| Wildbeast genotype | Black wildbeast (Connochaetos) | No reports in humans to date | – | Alves et al., 2005 |