Are New Zealand's Nothofagus species in equilibrium with their environment?

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Abstract
Past explanations of the large disjunctions in the distribution of New Zealand's four Nothofagus species have emphasized displacement during glacial cycles followed by slow re‐occupation of suitable sites, or the effects of plate tectonics coupled with ecological and/or environmental limitations to further spread. In this study the degree of equilibrium between Nothofagus distribution and environment was compared with that of other widespread tree species by statistical analysis. Generalized additive regression models were used to relate species distribution data to estimates of temperature, solar radiation, soil water deficit, atmospheric humidity, lithology and drainage. For each species, the amount of spatial patterning remaining unexplained by environment was assessed by adding a variable describing species presence/absence on adjacent plots. Results indicate that Nothofagus species occur more frequently in environments suboptimal for tree growth, i.e. having various combinations of cool temperatures, low winter solar radiation, high root‐zone water deficit, low humidity, and infertile granitic substrates. Despite these demonstrated preferences, they exhibit substantially more spatial clustering which is unexplained by environment, than most other widespread tree species. Predictions formed from regressions using environment alone confirm that several major Nothofagus disjunctions are not explicable in terms of the environmental factors used in this analysis, but more likely reflect the effects of historic displacement coupled with slowness to invade forest dominated by more rapidly dispersing endomycorrhizal species. The technique used in this study for detecting residual spatial autocorrelation after fitting explanatory variables has potentially wide application in other studies where either regression or ordination techniques are used for analysis of compositional data.