Geographical range is considered a good predictor of the levels of isozyme variation in plants. Widespread species, often consisting of historically larger and more continuous populations, maintain higher polymorphism and are less aected by drift, which tends to erode genetic variation in more geographically restricted species. However, widespread species occurring in small and disjunct populations may not ®t this pattern. In this study we examined genetic variation in Pilgerodendron uviferum, a conifer endemic to temperate forests of southern South America, and is such a widespread and habitat-restricted species. Twenty populations along the whole range of Pilgerodendron were analysed by isozyme electrophoresis to resolve 14 putative genetic loci. Eleven were polymorphic in at least one population although only six of them were polymorphic in more than one population. We found reduced within-population levels of isozyme variation, with only 11% polymorphic loci (0.95 criterion), 1.2 mean number of alleles per locus, and mean observed and expected heterozygosities of 0.024 and 0.033, respectively. Most genetic diversity was found within populations (H T 0.039, H S 0.033, F ST 15%). Greater polymorphism and lower divergence was estimated in the more geographically restricted and closely related Fitzroya. Thus, total range, in combination with information on the degree of among-population isolation, may be a better predictor of the levels of polymorphism than range size alone.
Prezygotic barriers may represent effective mechanisms to avoid the deleterious effects of inbreeding. This study reports the existence of distance-dependent prezygotic barriers in self-compatible Alstroemeria aurea, a clonal herb native to temperate forests of the southern Andes. We analyzed pollen germination and tube growth as indicators of donor-recipient affinity using crossing distances of 1, 10, and 100 m. We used allozyme electrophoresis to determine the actual genetic relatedness between donor and recipient ramets. Pollen germination was not affected by distance between mates, but the number of pollen tubes reaching the base of the style increased strongly with distance between donor and recipient. This pattern was related to an increase in genetic dissimilarity with distance between mates. In contrast, pollen tube-style interactions did not change with distance when we restricted analysis to individuals at different distances that appeared to be genetically identical. This test implied genetic dissimilarity as the critical factor affecting pollen performance. We propose that the existence of prezygotic barriers might contribute to the high degree of genetic mixing exhibited by some clonal species.
The effects of Pleistocene glaciations on the genetic characteristics of the most austral conifer in the world, Pilgerodendron uviferum , were analysed with specific reference to the hypothesis that the species persisted locally in ice-free areas in temperate South America. It was expected that genetic variation would decrease with latitude, given that ice fields were larger in southern Patagonia and thus refugia were probably located towards the northern distributional limit of the species as suggested by the fossil record. In addition, an increase in among-population genetic divergence was expected with increasing distance to putative glacial refugia. We examined the relationship between location and within-population variability indices of 20 Pilgerodendron populations derived from isozyme analyses. We analysed possible refugia hypotheses by the distribution of allele frequencies using multivariate discriminant analysis. The degree of genetic differentiation with geographical distance between all population pairs was investigated by Mantel tests. Results indicated that Pilgerodendron populations are highly monomorphic, probably reflecting past population bottlenecks and reduced gene flow. Southernmost populations tend to be the least genetically variable and were therefore probably more affected by glacial activity than northern ones. Populations located outside ice limits seem to have been isolated during the glacial period. The presence of centres of genetic diversity, together with the lack of a significant correlation between genetic and geographical distances and the absence of geographical patterns of allelic frequencies at most analysed alleles, may indicate that Pilgerodendron did not advance southward after the last glaciation from a unique northern refugium, but spread from several surviving populations in ice-free areas in Patagonia instead.
Patterns of trait variation may be adaptive when vary in relation to an environmental gradient. In particular, leaf traits can affect productivity and competitive ability. We identify patterns of leaf size and shape variation with environmental heterogeneity in one of the most widespread tree species within temperate South America: Embothrium coccineum (Proteaceae). We collected leaf specimens and composite soil samples from 35 populations between 38º and 55º S latitude in Patagonia, covering a wide range of mean annual precipitation (MAP) and mean annual temperature (MAT). At each location, we measured nine morphological traits, some of which were cross correlated hence we focus on a smaller number of representative traits. We hypothesized that leaf area (LA), dry mass (DM), and specific leaf area (SLA) would increase, and that leaf shape (SF) would be more elongated, with increasing temperature, precipitation, and soil nutrient availability. We also expected growing season climate to be more closely associated with leaf traits than mean annual metrics. We used bivariate and backward stepwise multiple regressions to analyse the dependence of morphological traits with climatic and edaphic metrics. LA and DM increased with increasing summer rainfall or winter temperature, as hypothesized. Opposite to our hypothesis, LA and DM decreased with increasing summer temperature suggesting that in terms of leaf size, E. coccineum may sense summer conditions largely as an increasing aridity stressful gradient. Surprisingly, SLA increased with increasingly warm or dry summers. SF was related positively to MAT and negatively to MAP, suggesting that under more benign western climate regimes E. coccineum leaves tend to be elongated. Across sites, LA and DM increased with soil organic carbon and available phosphorus, and decreased with soil nitrogen and exchangeable cations. The opposite pattern was observed for SLA. Biologically meaningful climate metrics and soil nutrient conditions are useful predictors for leaf size and structure in the widespread E. coccineum. The SLA patterns probably resulted from lower values in long lasting leaves, in addition to increasing soil nitrogen, so leaves in the south are thicker. Alternatively, it could be consequence from non-isometrical scaling of LA and DM, so larger leaves such as those under oceanic western climates have lower SLA. Patterns of multiple leaf trait variation along complex environmental gradients may become uncoupled from each other, differing from what is suggested in the literature for traits that vary along simple environmental gradients.
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