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Avian Adenovirus |   305

          days and can reach 10% and occasionally be as high as 30% (Hess,   et al., 1988; Shivaprasad et al., 2001; Hess, 2013; Moura-Alvarez
          2013). Clinical signs include poor growth, apathy, prostration,   et al., 2013). Limited studies have been conducted to understand
          ruffled feathers and huddling behaviour. Concurrent infections   the epizootiology and association of aviadenoviruses and diseases
          often occur in flocks affected by IBH, which might explain the   in turkeys. The prevalence of aviadenovirus in turkey flocks asso-
          difficulties in reproducing the disease experimentally (Schachner   ciated with poult enteritis complex has been recently described in
          et al., 2018).                                        Brazil (Moura-Alvarez et al., 2013), UK and Hungary (Kajan et
            GE is mainly caused by FAdV-1 and -8 (Ono et al., 2003;   al., 2010; Marek et al., 2014a). However, the serotypes affecting
          Okuda et al., 2004) and is commonly found in slaughtered broiler   these turkey flocks are not described (Kleine et al., 2017). Studies
          chickens (Okuda et al., 2004).                        conducted in Germany in 2012 demonstrated the prevalence of
            HPS is caused by FAdV-4 and occurs in broiler birds of 3 to   TAdV-2 and -4 in turkey flocks clinically presenting with enteritis
          5 weeks of age of either sex and occasionally in 10- to 20-week-  and poor growth, respectively. TAdV-5 and members of species
          old layers and breeder pullets. HPS outbreaks are rare in older   FAdV-B and FAdV-E were identified in flocks with clinical signs
          birds and in other species of poultry. HPS outbreaks have been   of hepatitis, splenitis, enteritis and polyserositis. In addition, a
          reported in quails, pigeons, black kites, ducks and recently in   member of species  TAdV-D was isolated from flocks showing
          ostriches (Asthana et al., 2013; Changjing et al., 2016; Pan et al.,   clinical signs of HPS and increased mortality (Kleine et al., 2017).
          2017b). Some studies suggest that broilers, regardless of breed,   The negative impacts  of EDS  in egg  production  have been
          are susceptible under field conditions. It has also been shown   recognized since 1976. The natural hosts for EDSV are ducks
          that the course of disease under natural conditions or after oral   and geese, though it extends its host range to other avian spe-
          inoculation range from 7 to 15 days (Asthana et al., 2013). The   cies such as coots, grebes, herring gulls, owls, storks, swans, and
          mortality rate can range between 20–80% depending on location.   quail. EDSV in chicken is thought to have originated from ducks
          For example, outbreaks in broiler farms in Pakistan ranged from   (Smyth, 2013), probably through contaminated vaccines grown
          20% to 75% and in India from 30% to 80% (Asthana et al., 2013).  in duck cells or embryonated eggs or during blood collection
            Emerging cases of severe IBH/HPS have been recently   (McFerran, 1979). Some studies showed that EDSV isolated
          reported in China, and IBH cases in Spain (2011–2013) and   from ducks failed to cause any drop in egg production from laying
          Korea (2007–2010) (Zhao et al., 2015; Oliver-Ferrando et al.,   hens, though egg weight decreased (Brugh et al., 1984). EDSV
          2017). In particular IBH/HPS outbreaks that are associated with   infection compromises eggshell quality and causes loss of egg
          virulent FAdV-4 are reported in high numbers from China and   production. Chickens of all ages and breeds are susceptible to
          since 2015 increases in economic losses due to high mortality   infection, though disease tends to be most severe in heavy broiler-
          have also been reported (Li et al., 2016; Zhang et al., 2016; Ruan   breeders and hens producing brown eggs. There are rare reports
          et al., 2017). Although FAdV-4 seems to be the predominant   that the egg drop syndrome virus (EDSV) has caused either a
          serotype in China, other serotypes – such as FAdV-1, FAdV-2,   drop in egg production or respiratory tract disease in other spe-
          FAdV-8, FAdV-10 and FAdV-11 have also been isolated from   cies, e.g. turkeys, ducks, geese, and quail (McFerran, 1979; Ivanics
          flocks with clinical signs of IBH (Zhao et al., 2015; Changjing et   et al., 2001). EDSV can be transmitted vertically through the egg,
          al., 2016).                                           after which the virus remains latent until progeny reaches sexual
            HE outbreaks were first reported in some states of the US   maturity. Virus is then shed through the eggs and faeces infecting
          during 1960s. This disease affects confined as well as free-range   susceptible contacts. Virus can be also transmitted horizontally
          turkey  flocks.  Currently,  HE  is  distributed  worldwide  where   during lay. Horizontal transmission between flocks can occur by
          turkey  production  takes place (Pierson and  Fitzgerald, 2013).   displacement of contaminated egg trays. Indeed, outbreaks often
          HEV varies in pathogenicity from no clinical signs and death to   occur in common egg-packing stations. Transmission and out-
          severe clinical outcomes with mortality ranging between < 1% to   breaks can occur when domestic birds have direct contact with
          over 60% (Dhama et al., 2017). HEV can infect and cause disease   wild birds or through water supply contaminated with wildfowl
          in chickens, peafowl, bobwhite quail and chukars (McFerran   droppings (Smyth, 2013).
          and Smyth, 2000). Infection of chickens and pheasants with   EDS 76 was successfully eradicated from Northern Ireland by
          pathogenic virus causes MSD and hence its alternative name,   constant monitoring (through HI test) and removal of contami-
          MSD virus (MSDV). MSD has been reported in all geographic   nated and adjoining pens until all birds (40 weeks of age) were HI
          locations with confinement pheasant operation. Infections in   negative. Eggs from grandparent and parent flocks were found to
          chickens are common based on serology (Dhama et al., 2017).  be EDSV-free (McFerran, 1979).
            Disease outbreaks caused by aviadenoviruses have been   EDSV is distributed worldwide in ducks and geese, but clinical
          reported in other poultry species such as quail, turkey, pigeon,   cases of EDS have been reported mainly in Europe, Asia, Africa
          goose and ostrich (McFerran and Smyth, 2000; Changjing et al.,   and Latin America (Bishop and Cardozo, 1996; Ivanics  et  al.,
          2016). The pathogenicity of aviadenoviruses in turkeys remains   2001; Biđin et al., 2007; Ezema et al., 2010; Cha et al., 2013; Kang
          unclear to date. These viruses have been isolated in clinical cases of   et al., 2017). EDS outbreaks have also been reported in flocks of
          respiratory and enteric diseases. IBH caused by aviadenoviruses   ducks in Canada and domestic birds (chickens and ducks) in the
          in turkey seems to be responsible for lower hatchability rates (Guy   USA (Schloer et al., 1978; Villegas et al., 1979; Brash et al., 2009).
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