Increasing deforestation worldwide has expanded the interfaces between fragmented forests and non-forest habitats. Human-made edges are very different from the original forest cover, with different ...microclimatic conditions. Conversely, the natural transitions (i.e., ecotones) are distinct from human-made forest edges. The human-made forest edges are usually sharp associated with disturbances, with abrupt changes in temperature, humidity, luminosity and wind incidence towards the forest interior. However, the natural forest-lake ecotones, even when abrupt, are composed of a complex vegetal physiognomy, with canopy structures close to the ground level and a composition of herbaceous and arboreal species well adapted to this transition range. In the present study, fruit-feeding butterflies were used as models to investigate whether faunal assemblages in natural ecotones are more similar to the forest interior than to the anthropic edges. Butterflies were sampled monthly over one year in the Rio Doce State Park, Southeastern Brazil, following a standardized design using a total of 90 bait traps, in three different forest habitats (forest interior, forest ecotone and anthropic edges), in both canopy and understory. A total of 11,594 individuals from 98 butterfly species were collected (3,151 individuals from 79 species in the forest interior, 4,321 individuals from 87 species in the ecotone and 4,122 individuals from 83 species in the edge). The results indicated that the butterfly richness and diversity were higher in transition areas (ecotones and edges). The ecotone included a combination of butterfly species from the forest interior and from anthropic edges. However, species composition and dominance in the ecotone were similar to the forest interior in both vertical strata. These results suggest that human made forest edges are quite distinct from ecotones. Moreover, ecotones represent unique habitats accommodating species adapted to distinct ecological conditions, while anthropic edges accommodate only opportunistic species from open areas or upper canopies.
IMPORTANCE: Acute respiratory distress syndrome (ARDS) due to coronavirus disease 2019 (COVID-19) is associated with substantial mortality and use of health care resources. Dexamethasone use might ...attenuate lung injury in these patients. OBJECTIVE: To determine whether intravenous dexamethasone increases the number of ventilator-free days among patients with COVID-19–associated ARDS. DESIGN, SETTING, AND PARTICIPANTS: Multicenter, randomized, open-label, clinical trial conducted in 41 intensive care units (ICUs) in Brazil. Patients with COVID-19 and moderate to severe ARDS, according to the Berlin definition, were enrolled from April 17 to June 23, 2020. Final follow-up was completed on July 21, 2020. The trial was stopped early following publication of a related study before reaching the planned sample size of 350 patients. INTERVENTIONS: Twenty mg of dexamethasone intravenously daily for 5 days, 10 mg of dexamethasone daily for 5 days or until ICU discharge, plus standard care (n =151) or standard care alone (n = 148). MAIN OUTCOMES AND MEASURES: The primary outcome was ventilator-free days during the first 28 days, defined as being alive and free from mechanical ventilation. Secondary outcomes were all-cause mortality at 28 days, clinical status of patients at day 15 using a 6-point ordinal scale (ranging from 1, not hospitalized to 6, death), ICU-free days during the first 28 days, mechanical ventilation duration at 28 days, and Sequential Organ Failure Assessment (SOFA) scores (range, 0-24, with higher scores indicating greater organ dysfunction) at 48 hours, 72 hours, and 7 days. RESULTS: A total of 299 patients (mean SD age, 61 14 years; 37% women) were enrolled and all completed follow-up. Patients randomized to the dexamethasone group had a mean 6.6 ventilator-free days (95% CI, 5.0-8.2) during the first 28 days vs 4.0 ventilator-free days (95% CI, 2.9-5.4) in the standard care group (difference, 2.26; 95% CI, 0.2-4.38; P = .04). At 7 days, patients in the dexamethasone group had a mean SOFA score of 6.1 (95% CI, 5.5-6.7) vs 7.5 (95% CI, 6.9-8.1) in the standard care group (difference, −1.16; 95% CI, −1.94 to −0.38; P = .004). There was no significant difference in the prespecified secondary outcomes of all-cause mortality at 28 days, ICU-free days during the first 28 days, mechanical ventilation duration at 28 days, or the 6-point ordinal scale at 15 days. Thirty-three patients (21.9%) in the dexamethasone group vs 43 (29.1%) in the standard care group experienced secondary infections, 47 (31.1%) vs 42 (28.3%) needed insulin for glucose control, and 5 (3.3%) vs 9 (6.1%) experienced other serious adverse events. CONCLUSIONS AND RELEVANCE: Among patients with COVID-19 and moderate or severe ARDS, use of intravenous dexamethasone plus standard care compared with standard care alone resulted in a statistically significant increase in the number of ventilator-free days (days alive and free of mechanical ventilation) over 28 days. TRIAL REGISTRATION: ClinicalTrials.gov Identifier: NCT04327401
The butterfly family Nymphalidae contains some of the most important non-drosophilid insect model systems for evolutionary and ecological studies, yet the evolutionary history of the group has ...remained shrouded in mystery. We have inferred a robust phylogenetic hypothesis based on sequences of 10 genes and 235 morphological characters for exemplars of 400 of the 540 valid nymphalid genera representing all major lineages of the family. By dating the branching events, we infer that Nymphalidae originated in the Cretaceous at 90 Ma, but that the ancestors of 10-12 lineages survived the end-Cretaceous catastrophe in the Neotropical and Oriental regions. Patterns of diversification suggest extinction of lineages at the Cretaceous/Tertiary boundary (65 Ma) and subsequent elevated speciation rates in the Tertiary.
The morphological species delimitations (i.e. morphospecies) have long been the best way to avoid the taxonomic impediment and compare insect taxa biodiversity in highly diverse tropical and ...subtropical regions. The development of DNA barcoding, however, has shown great potential to replace (or at least complement) the morphospecies approach, with the advantage of relying on automated methods implemented in computer programs or even online rather than in often subjective morphological features. We sampled moths extensively for two years using light traps in a patch of the highly endangered Atlantic Forest of Brazil to produce a nearly complete census of arctiines (Noctuoidea: Erebidae), whose species richness was compared using different morphological and molecular approaches (DNA barcoding). A total of 1,075 barcode sequences of 286 morphospecies were analyzed. Based on the clustering method Barcode Index Number (BIN) we found a taxonomic bias of approximately 30% in our initial morphological assessment. However, a morphological reassessment revealed that the correspondence between morphospecies and molecular operational taxonomic units (MOTUs) can be up to 94% if differences in genitalia morphology are evaluated in individuals of different MOTUs originated from the same morphospecies (putative cases of cryptic species), and by recording if individuals of different genders in different morphospecies merge together in the same MOTU (putative cases of sexual dimorphism). The results of two other clustering methods (i.e. Automatic Barcode Gap Discovery and 2% threshold) were very similar to those of the BIN approach. Using empirical data we have shown that DNA barcoding performed substantially better than the morphospecies approach, based on superficial morphology, to delimit species of a highly diverse moth taxon, and thus should be used in species inventories.
Although mutualisms have been intensively investigated, demonstration of indirect effects between co‐occurring mutualistic systems is rare. For instance, the ecological consequences of co‐occurrence ...of ant‐tended insects on a plant have never been examined for survival effects on either trophobiont species. Here, we assess the selective pressures mediating co‐occurrence of a facultative ant‐tended butterfly (Parrhasius polibetes) with ant‐tended treehoppers (Guayaquila xiphias) onSchefflera vinosashrubs. We evaluated host plant selection and caterpillar survival inP. polibetesin the presence and absence of ant‐treehopper associations. Paired trials revealed that butterflies preferably oviposit on branches hosting ant‐tended treehoppers when they had a choice between those and branches without this interaction. Presence of ant‐tended treehoppers on a branch reduced the abundance ofP. polibetes’ natural enemies and improved caterpillar survival in both premyrmecophylic and ant‐tended phases. Thus ant‐tended treehoppers create an enemy‐free space on foliage that butterflies exploit to protect larval offspring. These findings connect two widely documented ant‐trophobiont mutualisms and highlight the importance of considering multiple interactions for a proper understanding of ant‐plant‐herbivore systems. Detection of other ant‐based mutualisms on oviposition to improve offspring survival may have represented an important evolutionary step in the process of host plant selection in facultative myrmecophilous butterflies.
Aim
The landscape of the Neotropical region has undergone dynamic evolution throughout the Miocene, with the extensive Pebas wetland occupying western Amazonia between 23 and c. 10 Ma and the ...continuous uplift of the Andes mountains. The complex interaction between the Andes and Amazonia probably influenced the trajectory of Neotropical biodiversity, but evidence from time‐calibrated phylogenies of groups that diversified during this period is lacking. We investigate the role of these landscape transformations in the dynamics of diversification in the Neotropical region using a 26‐Myr‐old endemic butterfly radiation.
Location
Neotropics.
Time period
Oligocene to present.
Major taxa studied
Ithomiini butterflies.
Methods
We generated one of the most comprehensive time‐calibrated molecular phylogenies of a large clade of Neotropical insects, the butterfly tribe Ithomiini, comprising 340 species (87% of extant species) and spanning 26 Myr of diversification. We applied a large array of birth–death models and historical biogeography estimations to assess the dynamics of diversification and biotic interchanges, especially at the Amazonia–Andes interface.
Results
Our results suggest that the Amazonian Pebas wetland system played a major role in the timing and geography of diversification of Ithomiini, by constraining dispersal and diversification in the Amazon basin until c. 10 Ma. During the Pebas wetland period, Ithomiini diversification mostly took place in the Andes, where terrestrial habitats were not affected. An explosion of interchanges with Amazonia and with the Northern Andes accompanied the demise of the Pebas system (11–8 Ma) and was followed by local diversification in those areas, which led to a substantial renewal of diversification.
Main conclusions
Many studies on Neotropical diversity have focused only on the Andes, whereas we show that it is the waxing and waning of the Pebas mega‐wetland, interacting with Andean uplift, that determined the timing and patterns of regional interchanges and diversification in Ithomiini.
How evolution creates and maintains trait patterns in species-rich communities is still an unsolved topic in evolutionary ecology. One classical example of community-level pattern is the unexpected ...coexistence of different mimicry rings, each of which is a group of mimetic species with the same warning signal. The coexistence of different mimicry rings in a community seems paradoxical because selection among unpalatable species should favor convergence to a single warning pattern. We combined mathematical modeling based on network theory and numerical simulations to explore how different types of selection, such as mimetic and environmental selections, and habitat use by mimetic species influence the formation of coexisting rings. We show that when habitat and mimicry are strong sources of selection, the formation of multiple rings takes longer due to conflicting selective pressures. Moreover, habitat generalist species decrease the distinctiveness of different mimicry rings' patterns and a few habitat generalist species can generate a "small-world effect", preventing the formation of multiple mimicry rings. These results may explain why the coexistence of mimicry rings is more common in groups of animals that tend towards habitat specialism, such as butterflies.
Understanding why species richness peaks along the Andes is a fundamental question in the study of Neotropical biodiversity. Several biogeographic and diversification scenarios have been proposed in ...the literature, but there is confusion about the processes underlying each scenario, and assessing their relative contribution is not straightforward. Here, we propose to refine these scenarios into a framework which evaluates four evolutionary mechanisms: higher speciation rate in the Andes, lower extinction rates in the Andes, older colonization times and higher colonization rates of the Andes from adjacent areas. We apply this framework to a species‐rich subtribe of Neotropical butterflies whose diversity peaks in the Andes, the Godyridina (Nymphalidae: Ithomiini). We generated a time‐calibrated phylogeny of the Godyridina and fitted time‐dependent diversification models. Using trait‐dependent diversification models and ancestral state reconstruction methods we then compared different biogeographic scenarios. We found strong evidence that the rates of colonization into the Andes were higher than the other way round. Those colonizations and the subsequent local diversification at equal rates in the Andes and in non‐Andean regions mechanically increased the species richness of Andean regions compared to that of non‐Andean regions (‘species‐attractor’ hypothesis). We also found support for increasing speciation rates associated with Andean lineages. Our work highlights the importance of the Andean slopes in repeatedly attracting non‐Andean lineages, most likely as a result of the diversity of habitats and/or host plants. Applying this analytical framework to other clades will bring important insights into the evolutionary mechanisms underlying the most species‐rich biodiversity hotspot on the planet.
Heliconius butterflies are widely distributed across the Neotropics and have evolved a stunning array of wing color patterns that mediate Müllerian mimicry and mating behavior. Their rapid radiation ...has been strongly influenced by hybridization, which has created new species and allowed sharing of color patterning alleles between mimetic species pairs. While these processes have frequently been observed in widespread species with contiguous distributions, many Heliconius species inhabit patchy or rare habitats that may strongly influence the origin and spread of species and color patterns. Here, we assess the effects of historical population fragmentation and unique biology on the origins, genetic health, and color pattern evolution of two rare and sparsely distributed Brazilian butterflies, Heliconius hermathena and Heliconius nattereri.
We assembled genomes and re-sequenced whole genomes of eight H. nattereri and 71 H. hermathena individuals. These species harbor little genetic diversity, skewed site frequency spectra, and high deleterious mutation loads consistent with recent population bottlenecks. Heliconius hermathena consists of discrete, strongly isolated populations that likely arose from a single population that dispersed after the last glacial maximum. Despite having a unique color pattern combination that suggested a hybrid origin, we found no genome-wide evidence that H. hermathena is a hybrid species. However, H. hermathena mimicry evolved via introgression, from co-mimetic Heliconius erato, of a small genomic region upstream of the color patterning gene cortex.
Heliconius hermathena and H. nattereri population fragmentation, potentially driven by historical climate change and recent deforestation, has significantly reduced the genetic health of these rare species. Our results contribute to a growing body of evidence that introgression of color patterning alleles between co-mimetic species appears to be a general feature of Heliconius evolution.
The underlying mechanisms mediating the use of multiple host plants were investigated in Parrhasius polibetes (Lycaenidae), a florivorous and facultative myrmecophilous butterfly. Plant traits such ...as presence of ant-treehopper associations as a source of enemy-free space, flower bud dimensions, toughness, thickness, trichomes, and the corresponding performance and wear of P. polibetes mandibles were examined for three natural hosts: Schefflera vinosa (Araliaceae), Pyrostegia venusta (Bignoniaceae) and Luehea grandiflora (Malvaceae). Parasitism levels of larvae found on the three hosts were also determined. Almost all Luehea had ant-treehopper associations, and all larvae found on this host were non-parasitized. Parasitism was low in larvae found on Schefflera, half of which hosted ant-treehopper associations. No ant-treehopper association was found on Pyrostegia, where parasitism was significantly higher compared to other hosts. In the laboratory, P. polibetes performed well on Schefflera, followed by Pyrostegia. No larvae survived when fed with Luehea. Flower buds of Luehea were thicker and tougher than those of Schefflera and Pyrostegia. Indeed, mandibles of larvae reared on Luehea showed substantial wear, whereas those reared either on Schefflera or Pyrostegia presented no significant damage. Additionally, we suggest that co-occurrence with ant-treehopper associations on a plant provides parasitoid-free space for P. polibetes larvae. Our results support the hypothesis that ecological trade-offs among host plants (i.e., food quality and enemy-free space) promote polyphagy in natural populations of P. polibetes. Host morphological traits seem to play a relevant role in P. polibetes performance. To our knowledge, this is the first report showing the costs of polyphagy in a myrmecophilous butterfly.