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Fattorini et al.

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                  lnS

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                       0

                          -8 -6         -4 -2                        0  2  4  6  8

                                                                  lnA

Fig. 2 – Regression of number of species (lnS) against island area (lnA, in km2). Light grey lines: 95% confidence intervals. Dark grey
lines: 95% prediction intervals.

the best fit models for species richness and environmental              ure 1. All coefficients reported in the graphs were signif-
heterogeneity or homogeneity, we found that, in general,                icant at P < 0.05, except b2 in the pathway obtained for
a power function explained a larger fraction of variance                number of land cover categories, which indicates that, in
(R2 comprised between 0.617 and 0.821) than linear mod-                 this particular case, the contribution given by environmen-
els (R2 comprised between 0.221 and 0.824), except in the               tal heterogeneity virtually obliterated the effect of area.
case of Margalef richness and number of land cover cat-                 	 When maximum elevation was used as a measure of
egories, where the two models explained virtually identi-               environmental heterogeneity, the magnitude of the indirect
cal percentages of variance (R2 = 0.792 for the linear model            effect of area (a1b1) on species per island was much lower
and R2 = 0.787 for the power model of Margalef index; R2                than its direct effect (b2). The direct effect of area (b2) on
= 0.824 for the linear model and R2 = 0.821 for the pow-                species richness greatly exceeded the direct environment
er model of number of land cover categories), and Berg-                 effect on species (b1), but the total effect of area, when di-
er-Parker dominance, where the linear model explained a                 rect and indirect effects are summed (a1b1+b2), was much
slightly larger percentage of variance (R2 = 0.720) than the            greater (2.9 times larger) than the effect of environment.
power model (R2 = 0.708).                                               	 For Shannon index, Pielou equitability and Margalef
	 Results of structural equation models are shown in Fig-               richness, the magnitude of the indirect effect of area (a1b1)
                                                                        on species per island was roughly similar to (Shannon) or
Table 2 – Spearman rank correlation coefficients (rs) of species        lower to (Pielou and Margalef) its direct effect (b2). Us-
richness and area with environmental variables. N = number of           ing these indices, the direct effect of environment (b1) on
land cover categories, H = Shannon index; J = Pielou equitabi-          species richness exceeded the direct area effect on species
lity (evenness); DMg = Margalef richness index; C = Simpson             (b2), but the total effect of area, when direct and indirect
dominance index; d = Berger-Parker dominance. P = probability.          effects are summed (a1b1+b2), was greater (1.7 times larg-
                                                                        er) than the effect of environment.
                             Species         Area                       	 For the two indices of dominance (Simpson and Berg-
                         rP             rP                              er-Parker) the indirect effect of area on species per island
                                                                        was substantially lower than its direct effect. The direct
                                 s        s                             effect of environmental heterogeneity on species richness
                                                                        was lower than the direct area effect on species, but the
Area                   0.854 <0.00001   0.693    0.0001                 total effect of area, when direct and indirect effects were
Elevation              0.588 <0.002     0.883   <0.00001                summed, was about twice the effect of environmental het-
N                      0.871 <0.00001   0.816   <0.00001                erogeneity.
H                      0.831 <0.00001   0.602   <0.00001                	 Finally, when the number of habitats was considered,
J                      0.734 <0.00001   0.802   <0.00001                the indirect effect of area (a1b1 = 0.517) on species per is-
DMg                    0.826 <0.00001   -0.799  <0.00001                land was substantially higher than its direct effect (b2 =
C                      -0.841 <0.00001  -0.777  <0.00001                0.372). The direct effect of environmental heterogeneity
d                      -0.873 <0.00001                                  (b1 = 0.565) on species richness exceeds the direct area ef-
                                                                        fect on species (b2 = 0.372), but the total effect of area,

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