Butterflies (Papilionoidea), with over 18,000 described species [1], have captivated naturalists and scientists for centuries. They play a central role in the study of speciation, community ecology, biogeography, climate change, and plant-insect interactions and include many model organisms and pest species [2, 3]. However, a robust higher-level phylogenetic framework is lacking. To fill this gap, we inferred a dated phylogeny by analyzing the first phylogenomic dataset, including 352 loci (> 150,000 bp) from 207 species representing 98% of tribes, a 35-fold increase in gene sampling and 3-fold increase in taxon sampling over previous studies [4]. Most data were generated with a new anchored hybrid enrichment (AHE) [5] gene kit (BUTTERFLY1.0) that includes both new and frequently used (e.g., [6]) informative loci, enabling direct comparison and future dataset merging with previous studies. Butterflies originated around 119 million years ago (mya) in the late Cretaceous, but most extant lineages diverged after the Cretaceous-Paleogene (K-Pg) mass-extinction 65 mya. Our analyses support swallowtails (Papilionidae) as sister to all other butterflies, followed by skippers (Hesperiidae) + the nocturnal butterflies (Hedylidae) as sister to the remainder, indicating a secondary reversal from diurnality to nocturnality. The whites (Pieridae) were strongly supported as sister to brush-footed butterflies (Nymphalidae) and blues + metalmarks (Lycaenidae and Riodinidae). Ant association independently evolved once in Lycaenidae and twice in Riodinidae. This study overturns prior notions of the taxon's evolutionary history, as many long-recognized subfamilies and tribes are para- or polyphyletic. It also provides a much-needed backbone for a revised classification of butterflies and for future comparative studies including genome evolution and ecology.
We introduce a novel framework for conceptualising, quantifying and unifying discordant patterns of species richness along geographical gradients. While not itself explicitly mechanistic, this approach offers a path towards understanding mechanisms. In this study, we focused on the diverse patterns of species richness on mountainsides. We conjectured that elevational range midpoints of species may be drawn towards a single midpoint attractor - a unimodal gradient of environmental favourability. The midpoint attractor interacts with geometric constraints imposed by sea level and the mountaintop to produce taxon-specific patterns of species richness. We developed a Bayesian simulation model to estimate the location and strength of the midpoint attractor from species occurrence data sampled along mountainsides. We also constructed midpoint predictor models to test whether environmental variables could directly account for the observed patterns of species range midpoints. We challenged these models with 16 elevational data sets, comprising 4500 species of insects, vertebrates and plants. The midpoint predictor models generally failed to predict the pattern of species midpoints. In contrast, the midpoint attractor model closely reproduced empirical spatial patterns of species richness and range midpoints. Gradients of environmental favourability, subject to geometric constraints, may parsimoniously account for elevational and other patterns of species richness.
Aims We aim to document elevational richness patterns of geometrid moths in a globally replicated, multi‐gradient setting, and to test general hypotheses on environmental and spatial effects (i.e. productivity, temperature, precipitation, area, mid‐domain effect and human habitat disturbance) on these richness patterns. Location Twenty‐six elevational gradients world‐wide (latitudes 28° S to 51° N). Methods We compiled field datasets on elevational gradients for geometrid moths, a lepidopteran family, and documented richness patterns across each gradient while accounting for local undersampling of richness. Environmental and spatial predictor variables as well as habitat disturbance were used to test various hypotheses. Our analyses comprised two pathways: univariate correlations within gradients, and multivariate modelling on pooled data after correcting for overall variation in richness among different gradients. Results The majority of gradients showed midpeak patterns of richness, irrespective of climate and geographical location. The exclusion of human‐affected sampling plots did not change these patterns. Support for univariate main drivers of richness was generally low, although there was idiosyncratic support for particular predictors on single gradients. Multivariate models, in agreement with univariate results, provided the strongest support for an effect of area‐integrated productivity, or alternatively for an elevational area effect. Temperature and the mid‐domain effect received support as weaker, modulating covariates, while precipitation‐related variables had no explanatory potential. Main conclusions Despite the predicted decreasing diversity–temperature relationship in ectotherms, geometrid moths are similar to ants and salamanders as well as small mammals and ferns in having predominantly their highest diversity at mid‐elevations. As in those comparative analyses, single or clear sets of drivers are elusive, but both productivity and area appear to be influential. More comparative elevational studies for various insect taxa are necessary for a more comprehensive understanding of elevational diversity and productivity.
The structure of ecological networks reflects the evolutionary history of their biotic components, and their dynamics are strongly driven by ecoevolutionary processes. Here we present an appraisal of recent relevant research, in which the pervasive role of evolution within ecological networks is manifest. Although evolutionary processes are most evident at macroevolutionary scales, they are also important drivers of local network structure and dynamics. We propose components of a blueprint for further research, emphasizing processbased models, experimental evolution and phenotypic variation, across a range of distinct spatial and temporal scales. Evolutionary dimensions are required to advance our understanding of foundational properties of community assembly and to enhance our capability of predicting how networks will respond to impending changes.
Summary1. Gradients in elevation are increasingly used to investigate how species respond to changes in local climatic conditions. Whilst many studies have shown elevational patterns in species richness and turnover, little is known about how food web structure is affected by elevation. 2. Contrasting responses of predator and prey species to elevation may lead to changes in food web structure. We investigated how the quantitative structure of a herbivore-parasitoid food web changes with elevation in an Australian subtropical rain forest. 3. On four occasions, spread over 1 year, we hand-collected leaf miners at twelve sites, along three elevational gradients (between 493 m and 1159 m a.s.l). A total of 5030 insects, including 603 parasitoids, were reared, and summary food webs were created for each site. We also carried out a replicated manipulative experiment by translocating an abundant leaf-mining weevil Platynotocis sp., which largely escaped parasitism at high elevations (≥900 m a.s.l.), to lower, warmer elevations, to test if it would experience higher parasitism pressure. 4. We found strong evidence that the environmental change that occurs with increasing elevation affects food web structure. Quantitative measures of generality, vulnerability and interaction evenness decreased significantly with increasing elevation (and decreasing temperature), whilst elevation did not have a significant effect on connectance. Mined plant composition also had a significant effect on generality and vulnerability, but not on interaction evenness. Several relatively abundant species of leaf miner appeared to escape parasitism at higher elevations, but contrary to our prediction, Platynotocis sp. did not experience greater levels of parasitism when translocated to lower elevations. 5. Our study indicates that leaf-mining herbivores and their parasitoids respond differently to environmental conditions imposed by elevation, thus producing structural changes in their food webs. Increasing temperatures and changes in vegetation communities that are likely to result from climate change may have a restructuring effect on host-parasitoid food webs. Our translocation experiment, however, indicated that leaf miners currently escaping parasitism at high elevations may not automatically experience higher parasitism under warmer conditions and future changes in food web structure may depend on the ability of parasitoids to adapt to novel hosts.
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