Summary 1We mapped and identified all trees ≥ 10 mm in diameter in 25 ha of lowland wet forest in Amazonian Ecuador, and found 1104 morphospecies among 152 353 individuals. The largest number of species was mid-sized canopy trees with maximum height 10-20 m and understorey treelets with maximum height of 5-10 m. 2 Several species of understorey treelets in the genera Matisia and Rinorea dominated the forest numerically, while important canopy species were Iriartea deltoidea and Eschweilera coriacea . 3 We examined how species partition local topographic variation into niches, and how much this partitioning contributes to forest diversity. Evidence in favour of topographic niche-partitioning was found: similarity in species composition between ridge and valley quadrats was lower than similarity between two valley (or two ridge) quadrats, and 25% of the species had large abundance differences between valley and ridge-top. On the other hand, 25% of the species were generalists, with similar abundance on both valley and ridges, and half the species had only moderate abundance differences between valley and ridge. 4 Topographic niche-partitioning was not finely grained. There were no more than three distinct vegetation zones: valley, mid-slope, and upper-ridge, and the latter two differed only slightly in species composition. 5 Similarity in species composition declined with distance even within a topographic habitat, to about the same degree as it declined between habitats. This suggests patchiness not related to topographic variation, and possibly due to dispersal limitation. 6 We conclude that partitioning of topographic niches does make a contribution to the α -diversity of Amazonian trees, but only a minor one. It provides no explanation for the co-occurrence of hundreds of topographic generalists, nor for the hundreds of species with similar life-form appearing on a single ridge-top.
Large-scale geographical patterns of biotic specialization and the underlying drivers are poorly understood, but it is widely believed that climate plays an important role in determining specialization. As climate-driven range dynamics should diminish local adaptations and favor generalization, one hypothesis is that contemporary biotic specialization is determined by the degree of past climatic instability, primarily Quaternary climate-change velocity. Other prominent hypotheses predict that either contemporary climate or species richness affect biotic specialization. To gain insight into geographical patterns of contemporary biotic specialization and its drivers, we use network analysis to determine the degree of specialization in plant-hummingbird mutualistic networks sampled at 31 localities, spanning a wide range of climate regimes across the Americas. We found greater biotic specialization at lower latitudes, with latitude explaining 20–22% of the spatial variation in plant-hummingbird specialization. Potential drivers of specialization - contemporary climate, Quaternary climate-change velocity, and species richness - had superior explanatory power, together explaining 53–64% of the variation in specialization. Notably, our data provides empirical evidence for the hypothesized roles of species richness, contemporary precipitation and Quaternary climate-change velocity as key predictors of biotic specialization, whereas contemporary temperature and seasonality seem unimportant in determining specialization. These results suggest that both ecological and evolutionary processes at Quaternary time scales can be important in driving large-scale geographical patterns of contemporary biotic specialization, at least for co-evolved systems such as plant-hummingbird networks.
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