Ecological specialization enables the partitioning of resources and thus can facilitate the coexistence of species and promote higher species richness. Specialization and niche partitioning are expected to exert a decisive influence on local spatial scales, while species richness at regional scales should be shaped mostly by historical factors and abiotic conditions. Moreover, specialization is expected to be particularly important in communities that are exceptionally species rich for their environmental conditions. Concurrently, niche overlap in these communities should be minimized to enable species coexistence. We tested these hypotheses by studying specialization–richness relationship and niche overlap in assemblages of 298 species of songbirds (Passeriformes) across Australia. We used local (2–6 ha) to regional (bioregions) spatial scales and detailed data on habitat, diet and foraging behaviour (method, substrate and stratum). We expected the richness–specialization relationship to be particularly strong (a) on local spatial scales and (b) in communities exceptionally species rich for given environmental conditions (approximated by moisture and vegetation complexity). We also expected (c) low niche overlap in assemblages with specialized species. Only the third prediction was partly supported. First, while the specialization and species richness were often positively related, the strength and the direction of the relationship changed between traits and across spatial scales. The strength of the specialization–richness relationship was consistently positive only in foraging stratum, and it increased towards smaller spatial scales only in case of habitat and diet. Simultaneously, species in local communities demonstrated high overlap in habitat and diet. Second, we did not find particularly strong specialization–richness relationships in exceptionally species‐rich communities. Third, we found the expected negative relationship between specialization and overlap in foraging stratum and substrate (in local communities), suggesting that species partition ecological space locally in terms of where they find food. Our expectations were only weakly supported. Specialization on foraging stratum was probably important in facilitating species coexistence. Conversely, although species were often specialized on habitat and diet, high overlap in these traits did not preclude their local coexistence. Overall, specialization and overlap in foraging traits were more important for species coexistence than habitat or diet.
Vegetation complexity is an important predictor of animal species diversity. Specifically, taller vegetation should provide more potential ecological niches and thus harbor communities with higher species richness and functional diversity (FD). Resource use behavior is an especially important functional trait because it links species to their resource base with direct relevance to niche partitioning. However, it is unclear how exactly the diversity of resource use behavior changes with vegetation complexity. To address this question, we studied avian FD in relation to vegetation complexity along a continental‐scale vegetation gradient. We quantified foraging behavior of passerine birds in terms of foraging method and substrate use at 21 sites (63 transects) spanning 3,000 km of woodlands and forests in Australia. We also quantified vegetation structure on 630 sampling points at the same sites. Additionally, we measured morphological traits for all 111 observed species in museum collections. We calculated individual‐based, abundance‐weighted FD in morphology and foraging behavior and related it to species richness and vegetation complexity (indexed by canopy height) using structural equation modeling, rarefaction analyses, and distance‐based metrics. FD of morphology and foraging methods was best predicted by species richness. However, FD of substrate use was best predicted by canopy height (ranging 10–30 m), but only when substrates were categorized with fine resolution (17 categories), not when categorized coarsely (8 categories). These results suggest that, first, FD might increase with vegetation complexity independently of species richness, but whether it does so depends on the studied functional trait. Second, patterns found might be shaped by how finely we categorize functional traits. More complex vegetation provided larger "ecological space" with more resources, allowing the coexistence of more species with disproportionately more diverse foraging substrate use. We suggest that the latter pattern was driven by nonrandom accumulation of functionally distinct species with increasing canopy height.
Disentangling regional and local drivers of species richness in communities is a long-term focus of ecology. Regional species pools affect local communities by providing their constituent species. Additionally, the amount and variety of resources enhance diversity locally. Here, we investigated whether the same ecological factor (vegetation complexity) shapes both regional and local species richness and thus drives local diversity both indirectly (via pool size) and directly (via facilitating the coexistence of species). We studied passerine birds of woodlands and forests in eastern Australia. We quantified regional species pool size and sampled local bird communities at 63 transects spanning 3,000 km. We estimated canopy height both regionally using satellite imagery and locally using vegetation sampling in the field. We studied how species pool size changed with regional canopy height and water availability, and how local species richness changed with pool size and local canopy height. Local species richness increased with both local canopy height and the size of the regional species pool. Pool size, in turn, increased with regional canopy height, which itself increased with water availability. Moreover, local species richness expressed as a proportion of the regional pool also increased with local canopy height. In sum, vegetation complexity indexed by canopy height had a doubly positive effect on local species richness: indirectly by promoting a large regional species pool and directly by facilitating the coexistence of disproportionately many species locally. Regional pools were larger in tall forests probably due to the legacy of extensive moist forests that once covered most of Australia, thus providing a sizeable potential for speciation, diversification, and species persistence. Local species richness was greater in tall, more productive forests with more vegetation layers likely due to more and varied resources (i.e., more potential niches), allowing the coexistence of more individuals and species of consumers.
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