Habitat heterogeneity affects species distribution, and a better understanding of this relationship can inform biodiversity conservation. To understand how habitat heterogeneity affects multiple dimensions of biodiversity, we sampled fruit-feeding butterflies within savannah woodlands and gallery forests from July 2012 to June 2013 in the Cerrado. A total of 3459 individuals representing 56 species were collected, with 32 found in savannah and 51 in gallery forest. In addition to taxonomic diversity, gallery forests host higher functional (Gower = 0.72) and phylogenetic diversity (Faith's PD = 641.5) compared to savannahs (Gower = 0.61; Faith = 416.1, respectively). Within gallery forests, butterfly assemblages exhibited higher functional diversity in the understory (Gower = 0.8) than in the canopy (Gower = 0.6). Similarly, phylogenetic diversity was higher in understory (PD = 573.9) compared to canopy (PD = 391.9) within the savannah. Species with smaller bodies (CWMsize = 27.8), broader diet breadth (CWMdiet = 2642), and defense-related traits (CWMeyespots = 65% of the assemblage), tended to be found in gallery forests compared to savannah woodland (CWMsize = 29.4; CWMdiet = 1527; CWMeyespots = 35% of the assemblage), respectively. Gallery forests are marked by more stable and less severe microclimate, reinforcing their pivotal role as a climatic refuge for most species, with a particular emphasis on smaller, desiccation-prone species. Furthermore, habitat heterogeneity may provide new ecological niches, which contribute to the high butterfly diversity in the Cerrado biome. Gallery forests offer favorable conditions for a higher proportion of butterfly assemblages, whereas savannah woodlands host distinct functional groups, including larger-bodied, camouflaged, and dietary specialist species. In light of this, we strongly advocate for the preservation of the gallery forest-savannah ecotone, in order to promote the coexistence of diverse butterfly lineages and functional groups.
Body-size covaries with many species’ traits, with implications for population and ecosystem-level patterns. Body size and seasonality of a species may covary if, for certain body sizes and optimal resource availability, meet the larger energy requirements of large-bodied species, are restricted to a narrow temporal window. Here, we examine the relationship between body size and seasonality of fruit-feeding butterflies in the Cerrado, as well as its association with larval diet breadth and synchrony with fruit phenology. Relative to smaller-bodied clades, the adults within larger clades were less abundant, more generalized in their larval diet-breadth, more seasonal, and synchronized with fruit phenology. Body-size covaries with species traits that are sensitive to anthropogenic drivers. In the Cerrado realm, Brazil, larger-bodied butterflies tend to be more temporally specialized and more synchronized with food plants—therefore, more vulnerable to climate-driven changes in phenology (i.e., fruiting season). To account for climate driven changes in synchrony with resources in the Cerrado, conservation should continue to focus on preserving habitat, especially corridors between savannah and gallery forests, so that resources are diverse and sufficiently abundant to sustain populations. Results from our study suggest that to conserve larger-bodied nymphalids, habitat restoration projects should prioritize seed sources that will maximize interaction diversity. This may not only be achieved by planting a diversity of native host plants but also cultivars that offset seasonal changes in the timing of flushing or fruiting.
Phytochemical diversity is an effective plant defensive attribute, but much more research has focused on genetic and environmental controls of specific defensive compounds than phytochemical diversity per se . Documenting plasticity in phytochemical richness and plant chemical composition as opposed to individual compounds is important for understanding plant defense. This study outlines a multi-site transplant experiment in Cerrado gallery forests in central Brazil, utilizing Piper arboreum (Piperaceae), a prevalent and widespread neotropical shrub. Clones from four distinct populations were planted either at their origin site or in a different forest. Secondary metabolite composition varied between populations initially and then changed after transplanting. Interestingly, clones with chemical profiles that were distinct from the populations where they were introduced experienced reduced specialist chrysomelid herbivory compared to clones that were more chemically similar to the existing P. arboreum populations where they were planted. Specialist Lepidoptera herbivory also declined in clones transplanted to a new forest, but this change could not be ascribed to chemical profiles. In contrast, generalist herbivory was unaffected by chemical dissimilarity and transplanting. This research adds to the expanding body of evidence suggesting that phytochemical diversity is a dynamic trait exerting unique effects on different herbivore guilds.
Habitat heterogeneity affect species distribution, and a better understanding of this relationship can inform biodiversity conservation. To understand how habitat heterogeneity affects multiple dimensions of biodiversity, we sampled fruit-feeding butterflies within savannah woodlands and gallery forests in the Cerrado. Understory traps within gallery forests had greater functional diversity than canopy traps, and understory traps within savannah woodlands had greater phylogenetic diversity than canopy traps. Species with smaller bodies, broader diet breadth, and defense-related traits, tended to inhabit gallery forests. Our results reinforce the positive association between habitat heterogeneity and multiple dimensions of biodiversity. These results shed light on the mechanisms contributing to the maintenance of biodiversity in seasonal tropical savannahs, which might help to understand the impacts of intense anthropogenic pressures and climate change on the Cerrado biota.
Abstract Introduction: Diet and temporal specialists, especially those in tropical seasonal environments, require synchronous phenology with hosts. Species with life-history-traits that covary with temporal specialization may be particularly vulnerable to climate-driven changes in phenology. In the Cerrado, larger fruit-feeding butterflies species tend to be less abundant and more seasonal than their smaller-bodied relatives. It is important to consider how ecological process and phylogenetic history are associated with these attributes. Aim/Methods: Here we evaluated the effects of climate and body size on the seasonality of abundance of fruit-feeding butterflies as well as their association with natural history traits including diet breadth and synchrony with fruits.Results: The distribution of abundances of these butterflies species was not uniform throughout the year, was associated with body size and shaped by habitat type and climate. Overall, hot and wet months were associated with higher abundance and species richness of these nymphalids in the Cerrado. Lineages of large-bodied species were less abundant, more seasonal and more generalized in larval diet breadth. In addition, these large-bodied clades were more tightly synchronized with fruit phenology.Discussion: Our findings support the hypothesis that large and less abundant species to occur in a narrow temporal window and synchronized with fruit phenology. Implications for insect conservation: Particular attention should be given for the conservation of larger-bodied temporal specialist species as they are vulnerable to phenology mismatching as a result of rapid global warming, but they will also face additional pressures with the advancing deforestation in the Cerrado.
The Cerrado biome is a biodiversity hotspot with a rich biota, and intense anthropogenic pressures. Despite its importance, it is still poorly understood how insects occupy different spatial dimensions of the landscape. We investigated how the beta diversity of plants influences the beta diversity of nymphalids at the regional‐scale (gallery forest vs. savannah) and local‐scale (transects). We expected greater beta‐diversity in nymphalid assemblages in gallery forests given the heterogeneity of plants and canopy structure. We sampled nymphalids in each habitat monthly for 1 year and observed 3459 individuals from 62 species. Nymphalidae abundance was significantly higher in the understory than in the canopy. Specifically, only 18% of the individuals were captured in the canopy of gallery forests compared to 11% in savannah canopies. Species richness was higher in gallery forests (55 species) compared to savannahs (34 species). Savannah transects were very similar in plant species composition and butterfly assemblages within the understory and canopy, whereas the species composition of plants and butterflies in gallery forests was highly heterogeneous which is consistent with the habitat heterogeneity hypothesis. Gallery forests are spatially more complex, which probably results in higher niche availability that may contribute to reduce the community saturation and help to maintain greater beta diversity in these habitats.
Creating robust datasets of plant–insect interactions is important for understanding ecosystem dynamics, and data on species interactions can be used to evaluate conservation interventions. In the present work, we collected plant–herbivore–parasitoid data on an understudied but critical ecosystem—gallery forests in the Brazilian cerrado. We collected caterpillars on shrubs of Piper (Piperaceae) over the course of a year in seven gallery forests of varying sizes in order to compare seasonal changes in α- and β- diversity and tritrophic interaction networks as well as the role of fragment size in determining species and interaction diversity. Caterpillars were more abundant and diverse in the wet season and also increased with resource availability—the more Piper individuals present, the greater the abundance and richness of herbivores. The number of unique interactions between (i) plants and herbivores and (ii) herbivores and parasitoids did not change across seasons, but there was a high degree of turnover in the herbivore fauna between sites and seasons. Specialization was greatest in the dry-rainy season transition, when new leaves typically flush. Consistent with records of parasitism rates in the cerrado sensu stricto, parasitism in the gallery forests was greatest in the dry-rainy seasons. Forest size was not related to caterpillar richness. Overall, this work demonstrates the conservation value of gallery forests in supporting plant species that span the Amazon and the Atlantic Rainforest as well as diverse and highly seasonal trophic interactions.
Introduction: In animals, body size is correlated with many aspects of natural history, such as life span, abundance, dispersal capacity and diet breadth. However, contrasting trends have been reported for the relationship between body size and these ecological traits. Methods: Fruit-feeding butterflies were used to investigate whether body size is correlated with species abundance, dispersal, permanence, and larval diet breadth in a Neotropical savanna in Brazil (Cerrado). We used Blomberg’s K and Phylogenetic Generalized Least Squares models (PGLS) to measure phylogenetic signal strength in species traits and to estimate size–dispersal–diet breadth associations, while also taking shared ancestry into account. Results: 539 individuals from 27 species were captured, and 190 individuals were recaptured, representing a 35% recapture rate. We found body size to be negatively associated with butterfly abundance, and positively associated with dispersal level, distance traveled, number of traps visited, individual permanence, and diet breadth. These results indicate that larger butterflies are more likely to disperse over longer distances. Moreover, larger butterflies have more generalized larval diets, based on the number of host plant families, genera, and phylogenetic diversity of the host plants they consume as larvae. Smaller butterflies rely on fewer resources, which is reflected in their higher survival in small patches and may explain their lower dispersal ability and higher diet specialization. Nevertheless, lower dispersal ability may, if not compensated by large population sizes, threaten small-bodied species inhabiting environments, such as the Cerrado, which have intense deforestation rates. Conclusions: Body size is positively associated with dispersal and diet breadth for the fruit-feeding butterflies collected in this study.
Fire is one of the main disturbance agents globally and one of the main threats on the Brazilian cerrado (Neotropical savanna), acting as environmental filter for species selection. Individuals of Morpho helenor achillides (C. Felder and R. Felder, 1867) and M. menelaus coeruleus (Perry, 1810) were captured weekly using entomological nets and fruit-baited traps over a 36-mo period (from May 2005 to April 2008). The present study analyzed the impact of a fire event in both the above species, showing that they present different responses to this disturbance. Morpho helenor was persistent as adults during the dry season, which could fly away from the flames to neighbor unburned areas and return after dry-season bushfires. Conversely, Morpho menelaus persists only as caterpillars during the dry season, which are unable to escape from the flames resulting in high impacts on local population (the species was not captured up to the end of this study). In addition, based on host plant species reported in the literature, we assigned a broader host plant range to M. helenor, suggesting that a generalist diet could help in the maintenance of individuals during the dry season, as they have more options to breed all year round. A better understanding of the temporal dynamics of adult and immature stages could help predict the amplitude of the impacts of dry season fire events on insects, especially when preventive fires are strategically used inside protected areas.
EnglishNatural biological control is one of the major causes responsible for reduction of pest population in agricultural ecosystem. However, natural biological control importance is usually minimized by not being estimated. This study reports the occurrence of Campoletis sonorensis (Cameron, 1886) (Hymenoptera: Ichneumonidae), Archytas marmoratus (Townsend, 1915) and Archytas incertus (Macquart, 1851) (Diptera: Tachinidae) associated with Helicoverpa armigera (Hubner, 1809) (Lepidoptera: Noctuidae). The rate of larval parasitism was up to 41% in structured refuge areas of cotton, without chemical control with insecticides. This study strengthens our knowledge relating to parasitoids associated with natural control of H. armigera in the American continent. In addition, it documents the rule that structured refuge areas could play as a source of natural enemies, besides their contribution to production of non-selective populations to Bt technology. Key words: biological control; integrated pest management; natural enemies portuguesO controle biologico natural e um dos principais componentes responsaveis pela reducao populacional de pragas em agroecossistemas. Entretanto, a importância do controle biologico natural, usualmente, e minimizada devido nao ser estimada. Este estudo documenta a ocorrencia de Campoletis sonorensis (Cameron, 1886) (Hymenoptera: Ichneumonidae), Archytas marmoratus (Townsend, 1915) e Archytas incertus (Macquart, 1851) (Diptera: Tachinidae) associados a Helicoverpa armigera (Hubner, 1809) (Lepidoptera: Noctuidae). A taxa de parasitismo larval foi superior a 41% em areas de refugio estruturado de algodao, sem uso de inseticidas quimicos. Este estudo fortalece o conhecimento relacionado com os parasitoides associados ao controle natural de H. armigera no Continente Americano. Adicionalmente, documenta o efeito que areas de refugio estruturado podem desempenhar como fonte de inimigos naturais, alem da sua contribuicao para produzir insetos nao expostos a tecnologia Bt. Palavras-chave: controle biologico; manejo integrado de pragas; inimigos naturais
The use of GMO expressing Bt toxin in soybean production has increased significantly in the last years in Brazil in order to manage the damage caused by lepidopteran pests. In this study, we compared the richness and abundance of owlet moths (Noctuoidea) associated with Bt and non-Bt soybean. We determined the temporal variations as a function of phenology, and correlated the population variations of the most common species with meteorological variables. The research was conducted at the experimental area of Embrapa Cerrados. The collection method used was differentiated being suppressive and absolute. A total of 13 species were collected, of which eight occurred on Bt soybeans. The most representative taxa were Chrysodeixis includens (72.87%), Anticarsia gemmatalis (18.17%) and Spodoptera spp (5.22%). The number of larvae belonging to species targeted by the Bt technology was 10 times lower on Bt than on non-Bt soybeans. Utetheisa ornatrix and Elaphria deltoides were recorded on soybean for the first time, observing larvae of both species in non-Bt soybean and those of U. ornatrix also in Bt soybean. Only A. gemmatalis larvae correlated (p <0.05) negatively with precipitation. This study provided field information on the abundance and species richness of owlet moths on non-Bt soybeans, associated with the effects of Bt soybean. When considering the different levels of infestation between cultivars as a criterion, larvae monitoring is of substantial importance in order to develop the lost control program.
Some moths species are considered good indicators of habitat quality because they are very responsive to human disturbance, vegetation type, and successional processes. However, Saturniidae moths have not yet been considered as indicators of environmental quality. Little is known on the distribution of moth species in different vegetation types and the moths’ seasonal variations in the Brazilian savannah. Therefore, this study aims to describe the spatial distribution and temporal patterns of moths belonging to the Saturniidae family in two vegetation types—Cerrado sensu stricto on rocky outcrops and semi-deciduous forest—in both the rainy and dry seasons. It addresses the influence of the climatic seasons and vegetation types on abundance, richness, and species composition to describe the temporal and spatial distribution patterns and the relationship between the ecological aspects and the life history of these moths. This study was conducted in the Cerrado phytogeographical domain, in Pireneus State Park, Goiás, Brazil. The results revealed that most Saturniidae species sampled are present during the rainy season and typically found in forest habitats. Furthermore, a clear positive connection was found between the abundance pattern and rainfall seasonality and humid habitats; this is apparently related to the physiological tolerance of these moths, due to rudimentary mouthparts during their adulthood. Thus, rainfall and a forest habitat are important to and fundamental requirements for the persistence of the Saturniidae species in the Cerrado domain. Based on the results of this study, we suggest the use of saturniid species as indicators of changes in vegetation and climatic conditions.
β diversity of herbivorous insects in the tropics is usually very high, and there is often strong dissimilarity in herbivore species composition across different spatial scales and different abiotic gradients. Similarly, turnover is high for trophic interactions between herbivorous insects and their host plants. Two factors have been proposed to explain temporal or spatial differences in trophic interactions: changes in species composition and temporal changes in the behavior of shared species. The goal of this study was to evaluate determinants of high β diversity of trophic interactions between lepidopteran caterpillars and their host plants across dry and rainy seasons and their transitions. Over the course of a year, interaction diversity data were collected from 275 temporary plots in Cerrado vegetation, comprising 257 species of caterpillars, 137 species of host plants and 503 different trophic interactions. All these diversity parameters varied across seasons. Species assemblages of caterpillars and plants were different among the four seasons, and there was a high turnover of interactions between the seasons. The high temporal β diversity of trophic interactions was mostly due to interaction rewiring between co-occurring species, as opposed to changes in species composition over time.
This study examined temporal variation in the abundance of immature stages of Elbella luteizona (Hesperiidae) and describes the morphology and behavior of the larvae on their host plants, Byrsonima coccolobifolia and Myrsine guianensis. Five hundred sixty-eight 10 m diameter plots were searched for caterpillars in the Brazilian Cerrado over a period of one year. We inspected 5968 host plants, and found 31 eggs and 262 larvae on 244 plants. Similar numbers of immatures were found in both species of host plants. The abundance of immature stages varied monthly and was significantly higher in the dry season on both host plants, which may be due to the low density of natural enemies during that time. E. luteizona is univoltine, and larvae present relatively little morphological variation. However, during development, substantial changes occur in the architecture of leaf shelters that caterpillars construct. In addition, E. luteizona larvae develop very slowly, taking more than 300 days to complete metamorphosis.
Documented patterns of specialization and species interactions often omit plasticity in resource use across space and time, yet such variation is an important part of species interactions. To examine temporal variation in resource use, we compared species‐ and community‐level patterns of host plant use by folivorous caterpillars between the dry and rainy seasons in four preserved areas of cerrado vegetation in the Distrito Federal, Brazil. We sampled plants and caterpillars in 10 m circular plots monthly from March 2010 to March 2011. At the community level, we found a significant increase in the mean diet breadth of dry season caterpillars relative to those collected in the rainy season. Families and species of moths varied in diet breadth, but most exhibited a seasonal expansion. In particular, intraspecific comparisons showed a 30% increase in the number of host plants used in the dry season compared to the rainy season. These results provide a clear example of how temporal variation in resource use is heterogeneous, and more generalized patterns of resource use can emerge from studies at large temporal scales. Thus, seasonal or annual heterogeneity may obscure ecologically relevant specialized interactions that occur at smaller scales.
Specialist insects are more sensitive to spatial variations than generalists, which are able to exploit diverse hosts in various habitats. This study investigated whether specialist lepidopteran larvae feeding on a single host, Roupala montana Aubl. (Proteales: Proteaceae), maintain consistent abundance rates across spatial scales. We compared the abundance of specialist and generalist larvae at local and regional scales during the same period of collection, with equal sample efforts, and in the same type of vegetation, within a Brazilian savanna biome. Particularly, we focused on the following questions: Does spatial scale matter to the abundance of specialist larvae on a single host plant species? What is the relationship between the abundance of specialists and generalists among spatial scales? As predicted, in general, specialist larvae were present at higher densities on their specific host plants than generalists. However, we sought to learn how this abundance changed or did not change with spatial scale, as well as whether community similarity increased with spatial proximity. In this study, most larvae of specialist species on R. montana occurred at both local and regional scales, but they differed in abundance at different spatial scales. Moreover, although specialist larvae exhibited higher densities on R. montana than generalists, this pattern was not always consistent. The assemblage of larvae in neighboring areas showed greater mutual similarity, and there was a negative relationship between distance and similarity.
Sexual dimorphism is a pronounced pattern of intraspecific variation in Lepidoptera. However, moths of the family Sphingidae (Lepidoptera: Bombycoidea) are considered exceptions to this rule. We used geometric morphometric techniques to detect shape and size sexual dimorphism in the fore and hindwings of seven hawkmoth species. The shape variables produced were then subjected to a discriminant analysis. The allometric effects were measured with a simple regression between the canonical variables and the centroid size. We also used the normalized residuals to assess the nonallometric component of shape variation with a t-test. The deformations in wing shape between sexes per species were assessed with a regression between the nonreduced shape variables and the residuals. We found sexual dimorphism in both wings in all analyzed species, and that the allometric effects were responsible for much of the wing shape variation between the sexes. However, when we removed the size effects, we observed shape sexual dimorphism. It is very common for females to be larger than males in Lepidoptera, so it is expected that the shape of structures such as wings suffers deformations in order to preserve their function. However, sources of variation other than allometry could be a reflection of different reproductive flight behavior (long flights in search for sexual mates in males, and flight in search for host plants in females).
Moths exhibit different levels of fidelity to habitat, and some taxa are considered as bioindicators for conservation because they respond to habitat quality, environmental change, and vegetation types. In this study, we verified the effect of two phytophysiognomies of the Cerrado, savanna and forest, on the diversity distribution of moths of Erebidae (Arctiinae), Saturniidae, and Sphingidae families by using a hierarchical additive partitioning analysis. This analysis was based on two metrics: species richness and Shannon diversity index. The following questions were addressed: 1) Does the beta diversity of moths between phytophysiognomies add more species to the regional diversity than the beta diversity between sampling units and between sites? 2) Does the distribution of moth diversity differ among taxa? Alpha and beta diversities were compared with null models. The additive partitioning of species richness for the set of three Lepidoptera families identified beta diversity between phytophysiognomies as the component that contributed most to regional diversity, whereas the Shannon index identified alpha diversity as the major contributor. According to both species richness and the Shannon index, beta diversity between phytophysiognomies was significantly higher than expected by chance. Therefore, phytophysiognomies are the most important component in determining the richness and composition of the community. Additive partitioning also indicated that individual families of moths respond differently to the effect of habitat heterogeneity. The integrity of the Cerrado mosaic of phytophysiognomies plays a crucial role in maintaining moth biodiversity in the region.
Understanding variation in resource specialization is important for progress on issues that include coevolution, community assembly, ecosystem processes, and the latitudinal gradient of species richness. Herbivorous insects are useful models for studying resource specialization, and the interaction between plants and herbivorous insects is one of the most common and consequential ecological associations on the planet. However, uncertainty persists regarding fundamental features of herbivore diet breadth, including its relationship to latitude and plant species richness. Here, we use a global dataset to investigate host range for over 7,500 insect herbivore species covering a wide taxonomic breadth and interacting with more than 2,000 species of plants in 165 families. We ask whether relatively specialized and generalized herbivores represent a dichotomy rather than a continuum from few to many host families and species attacked and whether diet breadth changes with increasing plant species richness toward the tropics. Across geographic regions and taxonomic subsets of the data, we find that the distribution of diet breadth is fit well by a discrete, truncated Pareto power law characterized by the predominance of specialized herbivores and a long, thin tail of more generalized species. Both the taxonomic and phylogenetic distributions of diet breadth shift globally with latitude, consistent with a higher frequency of specialized insects in tropical regions. We also find that more diverse lineages of plants support assemblages of relatively more specialized herbivores and that the global distribution of plant diversity contributes to but does not fully explain the latitudinal gradient in insect herbivore specialization.