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Historical dispersal of white sharks C. Gubili et al.  1681


                                 GW44

                                                                                  GW10
                                 GW43
                                                                                GW8        GW9
                  GW40    GW39
                                GW37    7   GW35
                                                             GW15
                                                                          GW12
                   2         2                                        4
                                            2  GW42  43    GW14      GW11  GW6
                                                                                GW7
                     GW41      GW36                              GW17  GW2
                                      GW38
                                                          GW13                  GW1
                                                                        GW4   GW3
                                   missing haplotype
                   17                                      GW16
                                                                           GW18
                                   NW Atlantic                                           GW32   GW33

                                   Mediterranean                                GW31
                   GW45
                                                                                            GW30
                                   South Africa                       3     GW29                       GW27
                                                                                                   3
                                   New Zealand                                     2
                                                                                                   GW28
                                   Australia                  GW34
                                                                                              GW24
                                   NE Pacific                                                         GW25
                                                                               GW23
                                                                                                  2
                                                                                       GW22
                                                                          GW20
                                                                                               GW26
                                                                                  GW19      2
                                                                                    GW21
             Figure 1. A median-joining network of 45 mtDNA control region haplotypes from white shark sampling areas, illustrating the
             close affinity of Mediterranean samples with the Indo-Pacific clade. Circle size is proportional to the frequency of the haplo-
             type. Unless specified, single mutational steps are assumed between haplotypes; line breaks with a number indicate the
             mutational steps.


             Mediterranean white sharks from European museum col-  4. DISCUSSION
             lections [14] are presently inaccessible for destructive  Analysis of the mitochondrial control region of several
             analysis. However, it is unlikely that full sequences  rare Mediterranean white sharks suggests that they have
             would be obtained from this material. The South African  little genetic variability and a close affinity with the
             haplotypes form a distinct group separated from the NWA  Indo-Pacific clade. Current understanding of white
             by only seven mutational steps. However, the SA/NWA  shark biology suggests recent Lessepsian migrations
             group differs from the Indo-Pacific/MED group by    [37], characteristic of other Indo-Pacific taxa inhabiting
             43 mutations. Both Bayesian and ML analyses corrobo-  the eastern Mediterranean, are an unlikely explanation.
             rate the close genetic similarity of the Mediterranean  Here, we argue that this highly vulnerable stock may
             and western Indo-Pacific Ocean haplotypes (figure 2).  owe its origins to a historical anomalous migratory
             Further, SH tests of an alternative topology to the ML  event, rather than persistent historical or contemporary
             inferred tree (difference in ln L ¼ 32.36, p , 0.003)  dispersal via the Atlantic.
             firmly rejected the NWA/SA as sister clade to the     The close affinity of the MED and Indo-Pacific
             Mediterranean. Evolutionary rates for the white shark  (figure 2) dismisses founding of the Mediterranean popu-
             control region were calculated as either 1.19 or 0.74 per  lation solely by a simple dispersal scenario from the
             cent divergence between lineages per million years,  geographically closest populations of the NWA (separated
             calibrated with the rise of the Panama Isthmus or the  by more than 51 mutational steps; figure 1). Conse-
             Sunda-Sahul shelves, respectively. This corresponds to a  quently, the question arises as to whether the five or six
             Mediterranean–Indo-Pacific population divergence of  mutations differentiating MED and Indo-Pacific haplo-
             348–565 Ka. Significantly, this dates divergence of the  types indicate an isolated Mediterranean subpopulation,
             Mediterranean stock to the Late Pleistocene, a period of  or a stock with contemporary and persistent demographic
             extreme dynamic climatic and eustatic events, suggested  connections with the Indo-Pacific. The most divergent
             to have dramatic impacts on the distribution and size of  haplotypes within either the south-western Pacific or the
             fish populations.                                   NEP differ by nine (GW21 to either GW27 or GW33)
             Proc. R. Soc. B (2011)
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