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196  |  Kibenge et al.

          enteritis with occasional high mortality in turkey poults (Ger-  from young geese with lesions of splenitis, hepatitis with miliary
          showitz and Wooley, 1973; McFerran et al., 1976; Saif et al., 1985),   necrotic foci, epicarditis, arthritis, and tenosynovitis at necropsy
          decreased weight gain (Spackman et al., 2005a), age dependent   (Palya et al., 2003).
          moderate to severe atrophy of the bursa of Fabricius causing
          transient or possibly permanent immunosuppression (Spackman   Avian orthoreovirus disease in Bobwhite quail
          et al., 2005a; Day et al., 2008), and myocarditis (Shivaprasad et   (Colinus virginianus) and other captive exotic avian
          al., 2009; França et al., 2010). Nersessian et al. (1985b) noted   species
          that turkeys inoculated with a turkey enteric orthoreovirus had   Avian  orthoreovirus  as  a  pathogen  in  young  Bobwhite  quail
          a viraemia 7 days PI with virus distributed in most organs by   (Colinus virginianus) was first reported in a commercial operation
          3–7 days PI and virus recovered from tendons at 3–7 days and   (Magee et al., 1993). Affected quail were lethargic, had respira-
          28 days PI. Turkey reovirus isolated by Sharafeldin et al. (2014)   tory distress, and had hepatic necrosis, air sac lesions and sinusitis
          fulfilled Koch’s postulates which established the causal rela-  at necropsy with 95% flock mortality. Guy et al. (1987) and Ritter
          tionship between reoviruses and tenosynovitis in turkeys, and   et al. (1986) reported concomitant orthoreovirus and  Crypto-
          recently, seven more orthoreoviruses were isolated and geneti-  sporidium in bobwhite quail with enteritis; however, neither
          cally characterized from clinical cases of arthritis in turkeys from   claimed that orthoreovirus was the primary causative agent. Guy
          Pennsylvania, USA (Tang et al., 2015).                et al. (1987) suggested that Cryptosporidium promoted systemic
            Avian orthoreoviruses were implicated in a number of multi-  spread of orthoreovirus and orthoreovirus intensified  Crypto-
          factorial enteric syndromes in young turkeys, and the milder form   sporidium infection in Bobwhite quail.
          was characterized by diarrhoea and poor weight gain. However, in   Avian orthoreoviruses were isolated from other commercial
          some cases, high mortality was observed in the affected flocks and   poultry species. Tanyi et al. (1994) reported pancreatitis caused
          in these instances the disease was called poult enteritis complex   by  an  orthoreovirus  in  commercially  reared,  1-  to  3-week-old
          (PEC) (Spackman et al., 2005a), and an even more severe form   guinea-fowl. An orthoreovirus was isolated from fresh intestinal
          of the disease has been documented as poult enteritis mortality   content of a healthy, 1-year-old ostrich (Struthio camelus) from a
          syndrome (PEMS) (Day et al., 2008). PEMS is highly infectious   breeder farm in Japan (Sakai et al., 2009).
          in young turkeys and is characterized by diarrhoea, increased
          feed consumption, increased time-to market runting, stunting,   Orthoreoviruses in wild avian species
          decreased weight gain, immune dysfunction, and mortality (Day   Avian orthoreoviruses have been associated with a variety of
          et al., 2008). In addition to orthoreoviruses, numerous viruses,   disease syndromes in captive and free living (wild) birds. The
          including turkey coronavirus and turkey astrovirus type 2, have   first recoded isolation of orthoreovirus from pigeons (Columba
          been associated with PEC and PEMS and have been the cause of   spp.) was by McFerran et al. (1976). The virus was from a pigeon
          substantial economic losses to the turkey industry (Spackman et   with diarrhoea and was isolated on chicken kidney and chicken
          al., 2005b). However, (Heggen-Peay et al., 2002) demonstrated   embryo liver cell cultures. This pigeon reovirus isolate shared a
          that orthoreovirus alone was capable of inducing some of the   common antigen with known chicken orthoreoviruses which
          clinical signs associated with PEMS, including the intestinal   provided the evidence for the ability of avian orthoreoviruses to
          lesions and suppression of bursal and hepatic growth and devel-  cross species boundaries. Although diarrhoea and hepatitis were
          opment.                                               commonly  associated  with  orthoreovirus  infections  in  pigeons
                                                                (McFerran et al., 1976; Vindevogel et al., 1982), the attempts at
          Avian orthoreovirus disease in Muscovy ducks          experimental reproduction of the lesions failed, suggesting that
          (Cairina moschata), ducks and geese                   other factors might have been involved with the naturally occur-
          Orthoreoviruses were isolated from Muscovy Ducks (Cairina   ring disease (Vindevogel et al., 1982).
          moschata) in South Africa, France, Israel, and Hungary (Malkin-  Avian orthoreoviruses were regularly isolated from imported
          son et al., 1981; Heffels-Redmann et al., 1992; Palya et al., 2003),   psittacine birds (Meulemans et al., 1983), and as the commercial
          and the disease caused by orthoreoviruses usually affects young   market for imported psittacine birds has grown, outbreaks in these
          Muscovy ducklings between 2 and 4 weeks of age. The clinical   bird species have increased in frequency over the years (van den
          signs  included  general  malaise,  diarrhoea,  respiratory  signs,   Brand et al., 2007). Rigby et al. (1981) tested 269 consignments
          stunted growth, (Malkinson et al., 1981; Heffels-Redmann et   of imported birds (mainly psittacine and passerine avian species)
          al., 1992), and microscopic, multifocal hepatic, splenic and renal   to Canada, between January 1977 to August 1980 and isolated
          necrosis (Malkinson et al., 1981). Liu et al. (2011) and Chen   orthoreovirus from 22 consignments, and 17 of the isolates were
          et al. (2012) reported the isolation of a highly virulent duck   associated with enteritis in the shipped birds. Meulemans et al.
          orthoreovirus from Pekin ducklings (Anas platyrhynchos) suffer-  (1983) tested 28 batches of dead imported psittacine birds and
          ing from severe mortality due to hepatic and splenic necrosis, and   isolated orthoreoviruses from 15 batches. The main lesions pre-
          further studies revealed that the isolates were closely related to   sent were enteritis and hepatic congestion with focal necrosis, and
          Muscovy duck orthoreoviruses. McFerran et al. (1976) isolated   splenomegaly was present in some cases. An outbreak of reovirus
          an  orthoreovirus  from  faeces  of  normal  farmed  mallard  ducks   disease with high mortality was reported in Italy among imported
          in chick kidney and chicken embryo liver cell cultures. A goose   African grey parrots (Psittacus erithacus erithacus) and Australian
          reovirus related to Muscovy duck orthoreovirus was isolated   king parrots (Alisterus scapularis) (Conzo et al., 2001). Senne et
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