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124  |  Brown and Eterradossi

          et al., 2007; Turpin et al., 2008; Wei et al., 2013). Thus, it would   South America and reciprocally that of South American AMPV-A
          appear that AMPV-C have the broadest avian host range.  and B in North America, especially as both continents share
            Most of the information pertaining to the host range of AMPVs   migratory flight paths.
          has been drawn from different studies at different times in differ-
          ent locations. However, between 2014 and 2018 the susceptibility
          of SPF turkeys, chickens and Muscovy ducks to infection by   Clinical features
          each of the four subgroups was assessed in a series of controlled   Field AMPV infection of meat turkeys is most often observed
          experimental conditions (Brown et al. submitted). Globally these   between three and 12 weeks of age with peak clinical respiratory
          extensive  experimental  trials  confirmed  previous  literature  in   signs occurring 2–3 days after infection. At the onset in galliformes,
          that AMPV A, B, Turkey C and D were indeed well adapted to   watery eyes and sneezing accompanied by a clear nasal exudate
          turkeys  and chickens,  albeit  that differences in seroconversion   are observed. Thereafter, the nasal exudate becomes turbid, liquid
          and transmission of viruses in these species could be observed.   around the eyes develops a foamy appearance and often a dry
          The Muscovy duck AMPV C was well adapted to Muscovy ducks   tracheal cough develops (Eterradossi et al., 2015). Birds can also
          however, both chickens and turkeys seroconverted and were   show signs of depression. In galliformes, clinical signs culminate
          positive by virus isolation despite a lack of detection of viral   in swelling of the infraorbital sinuses. In the absence of secondary
          RNA by RT-qPCR. Details of this study should bring a deeper   bacterial infections these clinical signs will disappear within seven
          understanding of the host range of AMPVs and give insight as to   to ten days. Morbidity is usually 100% with mortality varying
          how different subgroups become adapted to their preferred host   from as little as 1% to as high as 60% in cases of poor management
          (Brown et al., 2018).                                 conditions and resulting secondary infections (Stuart, 1989; Van
            The close relationships between AMPV C viruses and HMPV   de Zande et al., 1998; Jones and Rautenschlein, 2013; Eterradossi
          and the suggestion that HMPV and AMPV C originated from a   et al., 2015). Field AMPV infection of breeding turkeys or chick-
          common ancestor about 200 years ago (de Graaf et al., 2008b)   ens is often much less pronounced and even benign in its early
          means that cross species transmission has to be considered   respiratory phase, which is suggested to be a result of better man-
          however, to date there are no reports demonstrating respiratory   agement conditions (Eterradossi et al., 2015). However, infection
          disease in humans caused by AMPV. On the other hand, one   during lay can result in a drop in egg production (10–30%) and
          study has demonstrated clinical signs; viral antigen and genomic   shell quality for up to 10–21 days.
          RNA of HMPV in HMPV infected turkeys under experimental   Similar clinical signs are observed in chickens and guinea fowl
          conditions (Velayudhan  et  al., 2006). More investigations into   following AMPV infections in the field however, in these species
          this issue are need.                                  early signs induced by virus-infection are less pronounced and can
                                                                sometimes be inapparent (Cook, 2000; Jones and Rautenschlein,
          Transmission and spread                               2013). That said, when complicated by secondary infections it
          The transmission of AMPV is thought to be mostly due to direct   can lead to swollen head syndrome (SHS) which is characterized
          contact  with excreting  individuals  or  contaminated  materials   by severe oedema of the infraorbital and periorbital sinuses that
          (Cook et al., 1991; Alkhalaf et al., 2002) and the virus is considered   progresses subcutaneously over the head hence the ‘swollen head’
          highly infectious. As a result, the density of the infected popula-  appearance, a loss of equilibrium and torticollis due to inflamma-
          tion plays an important role in its spread. Airborne transmission is   tion of the middle ear. In SHS, mortality can reach as high as 20%
          also suspected due to the replication of the virus in the respiratory   (Morley and Thomson, 1984; O’Brien, 1985; Picault et al., 1987;
          tract and observations in the field of spontaneous contamination   Pattison et al., 1989; Buys et al., 1989b).
          of different poultry houses during vaccination programs (France   Egg-drop in breeders and SHS in chickens may often develop
          anon). However, still to date only two studies focusing on this   several weeks after the triggering AMPV infection. As a result,
          issue have been reported and the results are conflicting (Giraud et   seroconversion against AMPV may already be well established
          al., 1986; Cook et al., 1991).                        and readily detected in supposedly ‘early’ serum samples collected
            The potential for AMPV to be transmitted vertically has been   in the first days when the egg-drop or swollen head signs become
          shown through detection of viral RNA in the eggs of infected   apparent. Under such a clinical course, a careful investigation of
          flocks though no infectious virus was detected. Although no   unnoticed mild respiratory episodes occurring in the preceding
          viruses have yet been isolated from the reproductive tract the fact   weeks may help unravel the original virus infection.
          that several studies detected viral RNA in the reproductive tract   The clinical signs observed in ducks following AMPV infec-
          of laying turkeys and breeder chickens (Jones et al., 1988; Khehra   tion are slightly different than those observed in turkeys, chicken
          and Jones, 1999; Cook et al., 2000; Villarreal et al., 2007) means   and guinea fowl and are somewhat more pronounced under
          that this route of infection merits further investigation.  experimental conditions. At the onset (2–3 days after initial
            The mechanism of spread of AMPV over long distances is not   infection)  watery  eyes  accompanied  by  a  clear  nasal  exudate
          known. Wild birds have been implicated though the data are con-  are observed and  the ducks have  a more  nervous  disposition.
          flicting and based on different subgroups (Turpin, 2003; Delogu   Thereafter heavy congestion in the respiratory tract develops and
          et al., 2004; Turpin et al., 2008). If wild birds are involved in the   rales not dissimilar to those of chickens following infection with
          transmission of AMPV over large distances then it is surprising   infectious bronchitis virus (IBV) can be heard. At this time the
          to note the lack of report so far for North American AMPV-C in   nasal exudate takes on a thick, cloudy appearance. Ducks tend
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