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EVOLUTION OF HYBRID STICK-INSECTS                                                                                       65

                                                                 Bacillus rossius: GAMETE DUPLICATION                     Bacillus lynceorum:APOMIXIS WITH DNA EXTRASYNTHESIS
                                                                                                                                                very low recombination
                                                       with recombination        without recombination

                                                                 "y* 1         primary oocyte                             primary oocyte            TETRA-PACHYTENE

                                                                                1st division
                                                                              secondary oocyte

                                                       /\              / \ 2nd division                                   1st division

                                                                       / \ Igamete duplication]

                                                                                                                          secondary oocyte

                                                                   \ / cleavage nuclei \ /                             d  genotype          A1A2A3

                                                                  ffjK zygoid /f^                                      Fig. 3 - Maturation processes of parthenogenetic Bacillus, a:
                                                                                                                       automictic facultative parthenogenesis of B. rossius;b, c: automictic
                                                                 AA or aa genotype AAor                                obligate parthenogenesis of B. atticus and of B. whitei, respectively;
                                                                                                                       d: apomictic obligate parthenogenesis of B. lynceorum. Note that:
Downloaded by [31.185.101.124] at 08:47 23 March 2016  Bacillus atticus: AUTOMIXIS WITH FUSION OF THE PRODUCTS         type a makes all loci homozygous at once; type b can produce
                                                                              OF THE 1st MEIOTIC DIVISION              homozygous genotypes from a heterozygous one following a
                                                                                                                       crossover; type c, although reductional and automictic, produces
                                                       without recombination                     with recombination    clonal offspring; type d is apomictic, butallows rare recombinations
                                                                                                                       to occur in the early synaptic phase before tetra-pachitene.
                                                                              primary oocyte

                                                                                 1st division                          1992); rare recombination events appear possible,
                                                                                                                       however, very likely originating during the short, partially
                                                                              secondary oocyte                         synaptic pachytenic phase (unpublished). As a consequen-
                                                                                2nd division                           ce, B. lynceorum unisexuals do not remain completely in-
                                                                                                                       variant over long periods.

                                                                                                                         The comparative picture stemming from both facultative
                                                                                                                       and obligatory parthenogenesis of Bacillus supports the
                                                                                                                       idea that parthenogenetic mechanisms follow species-
                                                                                                                       specific patterns (Fig. 3).

                                                       Aa genotype                                                     INTERACTING HYBRIDS AND PARENTAL TAXA

                                                       Bacillus  whitei:      CENTRAL FUSION WITH UNASSORTED GENOME      The data discussed above, besides demonstrating the oc-
                                                                              SEGREGATION ANDEXTRASYNTHESIS OF DNA     currence of interspecific interactions that ultimately
                                                                                                                       defined the composition of the genus, also showed thatall-
                                                                                no recombination                       female taxa oî Bacillus didnot become completely isolated
                                                                                                                       from their bisexual ancestors. A first type of evidence is
                                                                 primary oocyte  TETRA-PACHYTENE                       represented by the temporary Sardinian complex of
                                                                                                                       atticus/rossius hybrids: since it kept crossing to B. rossius
                                                       grandii genome                                  rossius genome  and its meiotic system apparently allowed recombination,
                                                                                                                       it didnotpersist asa defined hybrid andonly produced the
                                                                                 2nd division                          already reported paleo-introgression of atticus genes into
                                                                              secondary oocyte/                        the rossius pool.

                                                                              central fusion                             As far as stabilized hybrids are concerned, allozyme
                                                                                     zygoid                            markers and parallel karyological analyses consistently
                                                                                                                       evidenced two main distinct mechanisms of natural
                                                                                   genotype                            reproductive interactions occurring between allodiploid
                                                                                         Aa                            hybrids - either hemiclonal or clonal - andtheir ancestors:
                                                                                                                       «androgenesis» and «genome incorporation». Fertilized
                                                                                                                       hybridogenetic and parthenogenetic Bacillus allodiploids
                                                                                                                       are able to produce about 1 non-hybrid progeny, namely
                                                                                                                       all-paternal fertile bisexuals (androgenetics) from the mixis
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