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S. Renaud and J. R. Michaux

              Table 5 Characteristics of the islands, physical (area, maximum altitude, distance to continent) and ecological (number of competitors of
              the wood mice and qualitative estimation of the competition pressure, including density effects). The amount of morphological differen-
              tiation is estimated as the difference in size (estimated by A 0 for the M1 and area H1 for the mandible) or as Euclidean distances on FCs for
              estimating shape divergence, between the island value and the one of the closest mainland population of the same genetic clade. Data from
              Michaux (1996), Michaux et al. (1996b, 2002a,b) and Vigne (1999)

                       Island                                 Age of
                       area  No. of  Competition Dist. to  Altitude colonization Differential  M1 size Md size  M1 shape Md shape
                          2
              Island   (km ) competitors (low/high)  mainland (km) (m)  (years)  island/mainland  (A 0 )  (area H1) [d(RFT9)] [d(EFT7)]
              Ole ´ron  17.5  4     0        Bridge     Low   > 5000  OL-FRCH     )0.057 )0.063  0.018  0.024
              Re ´     9.3  3       0        4          Low   > 5000  RE-FRCH     )0.024  0.003  0.013  0.040
              Yeu      2.2  3       0        20         Low   > 5000  YEU-FRCH     0.671  0.259  0.024  0.063
              Noirmoutiers 4.9  4   0        Pathway    Low   > 5000  NM-FRCH     )0.289 )0.122  0.015  0.031
              Porquerolles 12.5  2  1        3          142   > 5000  PORQ-FRS     0.228 )0.073  0.020  0.032
              Port Cros  6.4  2     1        9.2        196   > 5000  PTCR-FRS    )0.261 )0.103  0.015  0.033
              Corsica  8722  4      0        82         2710   5000   CO-ITTARQ    0.098  0.113  0.007  0.037
              Sardinia  23,833 4    0        220        1834   5000   SARD-ITTARQ  0.062 )0.286  0.018  0.021
              Elba     224  4       1        8          1019  > 5000  E-ITTARQ     0.655  0.361  0.010  0.024
              Sicily   25,460 4     0        2          1950  > 5000  SICFIC-ITCAL  0.079 )0.063  0.021  0.038
                                                                      SICGRAT-ITCAL  0.163 )0.093  0.020  0.032
              Ibiza      541 3      0        180         475   5000   IB-SPMUR     0.043  0.427  0.006  0.044


              Table 6 Relationship between size and shape divergence and insular characteristics (cf. Table 5). The dependent variable was size difference
              or shape distance for the M1 and the mandible; grouping variable was derived from the insular characteristics in Table 5. The differences
              between the categories were tested using a Kruskall–Wallis test, a nonparametric analogue of a one-way analysis of variance. Significant
              effects (5%) are in bold. Some insular characteristics were discontinuous (number of competitors, competition level, age of colonization).
                                                                             2
                                                                                               2
                                                                                                         2
              Others were continuous and had to be categorised: (1) island area, three categories (< 20 km , 20 < area < 20,000 km , > 20,000 km ), (2)
              distance to mainland, three categories (< 10 km, 10 km < D < 200 km, > 200 km), (3) altitude, three categories (< 200 m,
              200 < A < 1500 m, > 1500 m)
                                                           Size                        Shape
                                                           M1 size      Md size        M1 shape        Md shape
              Type of parameter   Parameter                (A 0 )       (area H1)      [d(RFT9)]       [d(EFT7)]
              Physical            Island area              0.520        0.048          0.027           0.784
                                  Distance to continent    0.596        0.048          0.677           0.048
                                  Altitude                 0.527        0.093          0.149           0.938
              Historical          Age of colonization      0.683        0.683          0.102           1.000
              Ecological          Number of competitors    0.952        0.127          0.853           0.040
                                  Competition level        0.518        0.926          0.782           0.405



              both mandibles and molars as opposed to various individual  to Bergmann’s rule (Bergmann, 1847) within the considered
              patterns of differentiation on islands.           latitudinal range.
               The possible causes of the clinal gradients are discussed first,  The northward decrease in body size has been interpreted in
              as they provide a background to consider the factors influen-  many ways. Firstly, A. sylvaticus displays a distribution area
              cing insular differentiation.                     shifted towards warmer environments compared with the
                                                                related Apodemus flavicollis. Accordingly, its relative fitness is
                                                                thought to decrease towards colder environments, leading to a
              North–south size gradient of mandibles and molars on
                                                                progressive decrease in size (Niethammer, 1969). Another, not
              mainland
                                                                mutually exclusive, interpretation relates to the size variations
              Geographical variations in body size of the wood mouse,  of A. sylvaticus resulting from competition pressure exerted by
              Apodemus sylvaticus, have been reported for over 50 years  A. flavicollis (Alca `ntara, 1991). The negative effect of
              (Ursin, 1956; Saint Girons, 1966; Niethammer, 1969; Kah-  A. flavicollis on A. sylvaticus has been documented in Sweden
              mann & Niethammer, 1971; Alca `ntara, 1991). Our results  (Hoffmeyer & Hansson, 1974). In such environments, resource
              using mandible and molar size as proxies for body size support  partitioning occurs that could favour a small size for
              the increase in body size from the north to the south contrary  A. sylvaticus (Angelstam et al., 1987). As the competition

              348                                                                  Journal of Biogeography 34, 339–355
                                                          ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
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