Abstract
Biogeographic models predict that, because of increasingly unfavorable and stressful conditions, populations become less frequent, smaller, less dense, and less reproductive toward the range edges. These models have greatly influenced the thinking on geographical range limits and have broad implications for ecology, evolution, and conservation. However, empirical tests of the models have rarely investigated comprehensive sets of population properties. We studied population size and density and a broad set of fitness-related traits in 66 populations of the alpine thistle Carduus defloratus along a latitudinal (615 km) and altitudinal (342-2300 m) gradient from the European Alps in the south to the northern range limit in the low mountain ranges of central Germany. Regression analysis indicated that population size and plant density declined with decreasing altitude from the center to the range margin, but plant size increased. In spite of the larger size of plants, the number of seeds produced strongly declined toward the range margin, mainly due to an increase in seed abortion. The number of flowering plants in a population influenced all components of reproduction. Plants in large populations initiated more seeds, aborted fewer seeds, and produced more and larger seeds per plant. The probability that seeds were attacked by insect larvae and the proportion of seeds damaged decreased strongly from the center to the margin of the distribution. However, in spite of the much lower level of parasitization, plants at the range margin produced far fewer viable seeds. Fluctuating asymmetry of leaf width, an indicator of developmental instability, was similar across the range and not related to population size.
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