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Chapter 3: Transmission / Natural cycle


          Argasidae  with  genera  (soft  ticks),  and  the   well beyond the basis capituli. In Europe, TBEV
          Nuttalliellidae,  represented  by  only  one   has  been  recovered  from  Haemaphysalis
          species,  Nuttalliella  namaqua. 6,7,8   All  the  tick   punctata (‘the red sheep tick’), Haemaphysalis
          species  involved  in  the  eco-epidemiological   concinna   in   Europe   and   Asia,   and
          cycle of TBEV  belong to the Ixodidae. Details   Haemaphysalis  longicornis  in  Asia,  Australia,
          of  tick  biology  generally  can  be  found  in  a   New  Zealand,  and  recently  also  in  North
          variety  of  publications,  for  example  in   America. 7,14,15
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          Nicholson  et  al.,   Petney  et  al.,   and
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          Sonenshine  and  Roe,   and  a  list  of  valid   The  genus  Hyalomma  is  relatively  small  with
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          species names in Guglielmone and Nava.  The   27 species of small- to large-sized ticks.  They
          following  genera  of  ticks  contain  species   are  characterized  by  their  elongated  palps,
          known to transmit TBEV.                     which are at least twice as long as wide. The
                                                      distinct eyes are located in sockets adjacent to
          Ixodes  is  the  largest  tick  genus,  with  244   the postero-lateral edges of the scutum that is
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          described  species  worldwide .  Ixodes  species   unornamented. The distribution of Hyalomma
          are  characterized  by  a  distinct  groove  that   species is limited to the Old World, primarily
          encircles  the  anus  anteriorly  and  a  lack  of   to  arid  or  semiarid  habitats.  Hyalomma
          eyes.  Males  have  7  sclerotized  ventral  plates   marginatum (‘the Mediterranean Hyalomma’)
          that are absent in the males of other genera.   is the only member of this genus from which
          The  genus  Ixodes  has  been  subdivided  in   TBEV has been recovered.
          roughly   15   subgenera   (e.g.   Ixodes,
          Pholeoixodes) on the basis of morphology. 12,13
          The  genus  has  a  worldwide  distribution,
          including parts of Antarctica. 7,14  Some species   The biology of hard ticks
          are particularly important as vectors of TBEV:
          Ixodes ricinus the ‘castor bean tick’ or ‘sheep   All the species known to transmit TBEV have a
          tick’  in  Europe  and  middle  Asia,  Ixodes   3-host  life-cycle  (Figure  1).  Each  active  life
          persulcatus  ‘the  taiga  tick’  in  northeastern   stage  requires  a  blood  meal  from  a  suitable
          Europe and northern Asia, and Ixodes ovatus   host, after which the tick detaches and molts
          in middle Asia and Japan.                   on the ground. The arrows with broken lines
                                                      in the figure show the potential transmission
          The  genus  Dermacentor  has  35  species   paths  to  humans.  The  line  from  larvae  to
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          worldwide.   The  basis  capituli  appears   humans  indicates  that  transovarial  trans-
          rectangular  when  viewed  dorsally.  A  pair  of   mission  from  an  infected  female  can  happen
          medially directed spurs occurs on the first pair   which  results  in  infective  larvae.  Infection  of
          of  coxae.  The  palps  are  short  and  thick.  The   the  tick  can  occur  when  larvae,  nymphs,  or
          scutum  is  almost  always  ornamented.     females feed on an infective host (see below).
          Dermacentor  species  are  found  mostly  in
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          Europe, Asia, and North America.  In Europe,   The larva, nymph, and adult (female or male –
          TBEV  has  been  recovered  from  2  species,   Figures  2a,  2b,  2c,  and  2d)  are  active  stages
          Dermacentor  reticulatus  (‘the  ornate  dog   that  require  a  host  (this  is  not  the  case  for
          tick’),  Dermacentor  marginatus  (‘the  ornate   males  of  the  genus  Ixodes,  which  can  mate
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          sheep tick’), and in Asia, Dermacentor nuttalli.   off-host  without  feeding).   Larvae  are  easily
          Haemaphysalis  is  the  second  largest  tick   recognizable by the presence of only 3 pairs of
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          genus.  This eyeless genus can, in most cases,   legs,  and  absent  spiracular  and  genital
          be  identified  by  a  pronounced  lateral   apertures (Figures 3a and 3b). Nymphs have 4
          projection of palpal segment 2, which extends   pairs of legs and spiracles (Figures 4a and 4b).



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