Cerambycid Beetle Communities in Caatinga Dry Forests Are Structured by Seasonal Species Turnover.

Dry forests in the Caatinga biome of Brazil are seasonal ecosystems where diversity is driven by water availability. Understanding how the distribution of communities is driven by temporal climate changes has intrigued researchers for decades. However, temporal diversity patterns should be more evident in dry environments, since seasonality is characterized by being highly limiting to insect activities. Cerambycid beetles are considered good ecological indicators because they respond well to impacts of environmental changes. Thus, we asked two questions: (i) How do climatic changes affect the diversity of these insects across seasons? (ii) Are diversity components correlated with increasing air humidity, rainfall, and temperature? Our results showed a marked seasonality of cerambycid beetles, with higher abundance and richness in the wet season. The mean temperature and relative humidity were predictors of the composition of beetle assemblages. However, the variation of cerambycid assemblages between seasons is related mainly to species turnover. Our study demonstrates that the combined effect of temperature and humidity drives the temporal distribution of the cerambycids in dry forests. Although thermal sensitivity was low, the decrease in air moisture during the dry season was the limiting factor for these insects. Species turnover increased continuously with air moisture and temperature rise, creating temporal segregation among cerambycid species and maintaining the stability of the assemblage. Thus, our results are consistent with mechanisms invoking activity patterns, desiccation resistance, and physiologic constraints that predict a decrease in richness and abundance of the cerambycids from warmer and moister to colder and drier conditions.

Microbial Communities of the Gut and Nest of the Humus- and Litter-Feeding Termite Procornitermes araujoi (Syntermitinae).

The evolution of the symbiotic association with microbes allowed termites to decompose ingested lignocellulose from plant-derived substrates, including herbivore dung and soil humus. Representatives of the Syntermitinae (Termitidae) range in their feeding habits from wood and litter-feeding to humus-feeding species. However, only limited information is available about their feeding ecology and associated microbial communities. Here we conducted a study of the microbial communities associated to the termite Procornitermes araujoi using Illumina sequencing of the 16S and ITS rRNA genes. This species has been previously included in different feeding guilds. However, most aspects of its feeding ecology are unknown, especially those associated to its symbiotic microbiota. Our results showed that the microbial communities of termite guts and nest substrates of P. araujoi differed significantly for bacteria and fungi. Firmicutes dominated the bacterial gut community of both workers and soldiers, whereas Actinobacteria was found in higher prevalence in the nest walls. Sordariomycetes was the most abundant fungal class in both gut and nest samples and distinguish P. araujoi from the grass/litter feeding Cornitermes cumulans. Our results also showed that diversity of gut bacteria were higher in P. araujoi and Silvestritermes euamignathus than in the grass/litter feeders (C. cumulans and Syntermes dirus), that could indicate an adaptation of the microbial community of polyphagous termites to the higher complexity of their diets.

Food Storage by the Savanna Termite Cornitermes cumulans (Syntermitinae): a Strategy to Improve Hemicellulose Digestibility?

It has been suggested that food storage inside the nest may offer termites with a nutritional provision during low resource availability. Additionally, feces employed as construction material provide an excellent environment for colonization by microorganisms and, together with the storage of plant material inside the nest, could thus provide some advantage to the termites in terms of lignocellulose decomposition. Here, we conducted for the first time a comprehensive study of the microbial communities associated to a termite exhibiting food storage behavior using Illumina sequencing of the 16S and (ITS2) regions of rRNA genes, together with enzymatic assays and data collected in the field. Cornitermes cumulans (Syntermitinae) stored grass litter in nodules made from feces and saliva located in the nest core. The amount of nodules increased with nest size and isolation, and interestingly, the soluble fraction of extracts from nodules showed a higher activity against hemicellulosic substrates compared to termite guts. Actinobacteria and Sordariales dominated microbial communities of food nodules and nest walls, whereas Spirochetes and Pleosporales dominated gut samples of C. cumulans. Within Syntermitinae, however, gut bacterial assemblages were dissimilar. On the other hand, there is a remarkable convergence of the bacterial community structure of Termitidae nests. Our results suggest that the role of nodules could be related to food storage; however, the higher xylanolytic activity in the nodules and their associated microbiota could also provide C. cumulans with an external source of predigested polysaccharides, which might be advantageous in comparison with litter-feeding termites that do not display food storage behavior.

Host plant invests in growth rather than chemical defense when attacked by a specialist herbivore.

Plant defensive compounds may be a cost rather than a benefit when plants are attacked by specialist insects that may overcome chemical barriers by strategies such as sequestering plant compounds. Plants may respond to specialist herbivores by compensatory growth rather than chemical defense. To explore the use of defensive chemistry vs. compensatory growth we studied Brugmansia suaveolens (Solanaceae) and the specialist larvae of the ithomiine butterfly Placidina euryanassa, which sequester defensive tropane alkaloids (TAs) from this host plant. We investigated whether the concentration of TAs in B. suaveolens was changed by P. euryanassa damage, and whether plants invest in growth, when damaged by the specialist. Larvae feeding during 24 hr significantly decreased TAs in damaged plants, but they returned to control levels after 15 days without damage. Damaged and undamaged plants did not differ significantly in leaf area after 15 days, indicating compensatory growth. Our results suggest that B. suaveolens responds to herbivory by the specialist P. euryanassa by investing in growth rather than chemical defense.

Soldiers initiate foraging activities in the subterranean termite, Heterotermes tenuis.

Caste polyethism has been recorded in some termite species, however the foraging behavior of subterranean termites remains poorly known. Heterotermes tenuis Hagen (Isoptera: Rhinotermitidae) is a subterranean termite that is native to Brazil and is an agricultural and urban pest. The aim of this study was to investigate which caste acts as scouts when searching for food sources and determinate the percentages of each caste present in the foraging territories of field colonies of H. tenuis. Our results showed no significant differences among the caste proportions present in the foraging territories of the three colonies studied in the field. Laboratory experiments showed that minor soldiers were the most frequent initiators of foraging activities. This result suggests that the exploratory phase of the foraging behavior may be regulated by the number of soldiers present in the foraging territories of each colony.

Attraction of Bucephalogonia xanthophis (Hemiptera: Cicadellidae) to volatiles of its natural host Vernonia condensata (Asteraceae)

Plant volatiles are important cues for the orientation of herbivorous insects. It is possible that these compounds indicate whether the plant is suitable for feeding and larval development, or for mating aggregation. Vernonia condensata (Asteraceae) is known to attract species of leafhoppers, most of them important vectors of the citrus variegated chlorosis (CVC). In this study, we evaluated the role of volatiles of V. condensata on the orientation of Bucephalogonia xanthophis (Hemiptera: Cicadellidae). Four-arm olfactometer bioassays showed that only males were attracted to the volatiles of the host-plants Citrus sp. and V. condensata. Furthermore, fresh leaves of V. condensata induced a stronger response than volatiles from hexane-extracted leaves. This study opens the possibility to utilize V. condensata volatiles for pest management programs of B. xanthopis.

Os voláteis de plantas são sinais importantes para a orientação de insetos herbívoros. É possível que esses compostos indiquem que a planta é apropriada para alimentação e desenvolvimento larval, ou para agrupamento reprodutivo. Vernonia condensata (Asteraceae) atrai espécies de cigarrinhas, muitas delas vetores da clorose variegada dos citros (CVC). Neste trabalho, foi avaliado o papel dos voláteis de V. condensata na orientação de Bucephalogonia xanthophis (Hemiptera: Cicadellidae). Bioensaios em olfatômetro de quatro braços mostraram que somente os machos foram atraídos pelos voláteis das plantas hospedeiras Citrus sp. e V. condensata, sendo que os voláteis das folhas frescas de V. condensata induziram uma maior resposta quando comparada aos extratos hexânicos da planta. A possibilidade de usar os voláteis de V. condensata abre novas perspectivas para programas e alternativas para o manejo de pragas.

Attraction of Bucephalogonia xanthophis (Hemiptera: Cicadellidae) to volatiles of its natural host Vernonia condensata (Asteraceae)

Plant volatiles are important cues for the orientation of herbivorous insects. It is possible that these compounds indicate whether the plant is suitable for feeding and larval development, or for mating aggregation. Vernonia condensata (Asteraceae) is known to attract species of leafhoppers, most of them important vectors of the citrus variegated chlorosis (CVC). In this study, we evaluated the role of volatiles of V. condensata on the orientation of Bucephalogonia xanthophis (Hemiptera: Cicadellidae). Four-arm olfactometer bioassays showed that only males were attracted to the volatiles of the host-plants Citrus sp. and V. condensata. Furthermore, fresh leaves of V. condensata induced a stronger response than volatiles from hexane-extracted leaves. This study opens the possibility to utilize V. condensata volatiles for pest management programs of B. xanthopis.

Os voláteis de plantas são sinais importantes para a orientação de insetos herbívoros. É possível que esses compostos indiquem que a planta é apropriada para alimentação e desenvolvimento larval, ou para agrupamento reprodutivo. Vernonia condensata (Asteraceae) atrai espécies de cigarrinhas, muitas delas vetores da clorose variegada dos citros (CVC). Neste trabalho, foi avaliado o papel dos voláteis de V. condensata na orientação de Bucephalogonia xanthophis (Hemiptera: Cicadellidae). Bioensaios em olfatômetro de quatro braços mostraram que somente os machos foram atraídos pelos voláteis das plantas hospedeiras Citrus sp. e V. condensata, sendo que os voláteis das folhas frescas de V. condensata induziram uma maior resposta quando comparada aos extratos hexânicos da planta. A possibilidade de usar os voláteis de V. condensata abre novas perspectivas para programas e alternativas para o manejo de pragas.

Differential attractiveness of potato tuber volatiles to Phthorimaea operculella (Gelechiidae) and the predator Orius insidiosus (Anthocoridae).

The behavioral responses of the potato tuberworm moth Phthorimaea operculella and the polyphagous predator Orius insidiosus to volatiles emanating from exposed tubers were studied by four-arm olfactometer bioassays. Mated females of P. operculella distinguished volatiles released by intact potato tubers from volatiles damaged mechanically or by conspecific larvae. Volatiles from intact potato tubers were attractive to them. On the other hand, unmated females of P. operculella did not respond to tuber volatiles. Adults of O. insidiosus were attracted to volatiles from tubers damaged by P. operculella larvae, but did not respond to intact or mechanically damaged tubers. Methyl jasmonate (MeJA) was the only compound identified from the headspace of potato tubers (GC-MS of direct headspace sampling). The amount varied with the type of induction, being 0.001 +/- 0.0003 ng g(-1) in tissues of intact fresh tubers, 0.002 +/- 0.0007 ng g(-1) in mechanically damaged tubers, and showing a six- to tenfold increase in P. operculella damaged tubers (0.090 +/- 0.006 ng g(-1)). Behavioral bioassays with synthetic MeJA confirmed that the response of the insects is dependent on MeJA concentration. Mated females of P. operculella showed the highest response at 0.001 ng g(-1) (concentration released by intact tubers), whereas O. insidiosus showed the highest response, between 0.01 and 0.05 ng g(-1), which is close to the concentration released by P. operculella damaged tubers. Based on these results, we postulate that P. operculella and O. insidiosus have adapted their responses to plant volatiles differently, enabling them to locate suitable hosts or prey.

Plant volatiles: new perspectives for research in Brazil

Os agroecossistemas consistem em complexas relações tróficas entre plantas hospedeiras, herbívoros e seus inimigos naturais. Este trabalho revisa as pesquisas com voláteis de plantas no Brasil, apresenta os múltiplos mecanismos de resistência em culturas de importância econômica e contribui para o conhecimento das defesas induzidas em plantas. Uma grande parte dos programas de manejo de pragas, incluindo controle químico e biológico, não considera o impacto dessas substâncias sobre herbívoros e seus inimigos naturais. Estratégias alternativas de controle estão sendo desenvolvidas para o entendimento dos mecanismos endógenos de defesas induzidas em plantas contra artrópodes fitófagos. A utilização de voláteis de plantas no manejo integrado de pragas é uma estratégia adicional e ecologicamente sustentável no controle de pragas. Essa técnica envolve a possibilidade de utilização de iscas como atraentes de organismos benéficos, e a manipulação dos processos bioquímicos que induzem e regulam as defesas em plantas. A determinação dos mecanismos responsáves pela defesa indireta de plantas resultará em avanços significativos no controle biológico de pragas.

Plant volatiles: new perspectives for research in Brazil.

Agroecosystems consist on complex trophic relationships among host plants, herbivores and their natural enemies. This article reviews the research of plant volatiles in Brazil, in order to determine multiple resistance mechanisms of economically important crops and to contribute to the understanding of insect-plant interactions. Most pest management programs, including chemical and biological control, do not consider the impact of these chemicals on herbivores and their natural enemies. Alternative control methods are being developed in order to improve our understanding on the endogenous mechanisms of plant induced defenses against phytophagous arthropods. The use of plant volatiles technology as an additional tool in integrated pest management programs would offer a new and environmentally sound approach to crop protection. This technique involves the development of baits that attract beneficial organisms and the manipulation of biochemical processes that induce and regulate plant defenses, key factors in the improvement of control programs against economically important pests. The elucidation of the mechanisms involved in the indirect defenses of plants will result in useful tools for biological control of crop pests.

Effect of biotic and abiotic factors on the tunneling behavior of Coptotermes gestroi and Heterotermes tenuis (Isoptera: Rhinotermitidae).

Using bidimensional arenas, the construction and spatial dispersion of tunnels constructed by Coptotermes gestroi and Heterotermes tenuis (Rhinotermitidae) was determined under different laboratory conditions. Workers of both species showed an increase of the tunneled area with the rise in temperature. The activity of workers of C. gestroi also increased with the rise in soil moisture. Primary tunnels showed a high degree of dispersion in the arenas for all treatments. The presence of food did not influence the tunneling pattern of C. gestroi; but for H. tenuis there was a significant decrease in the number of tunnels. In addition, the type of substrate affected the tunneling pattern in both species. The understanding of food searching strategies of subterranean termites could contribute in the improvement of pest management programs based on the use of toxic baits.

Cuticular hydrocarbons of Heterotermes tenuis (Isoptera: Rhinotermitidae): analyses and electrophysiological studies.

Termites have become an important pest of Eucalyptus and Pinus reforestations, sugarcane and other cultures. An alternative for the control of this pest would be the use of attractive traps that take in account the social behavior of these insects. Diverse factors are important for the insects in the localization of the habitat and the choice of the food and specific odors can facilitate this. Studies referring to Heterotermes tenuis (Isoptera: Rhinotermitidae) are scarce. The objective of this work was to analyze the tergal cuticular extract of H. tenuis and determine the selectivity and sensitivity of its antennae to the components of this extract by electroantennography (EAG). The composition of the cuticular extract was determined by GC-MS analysis. The hydrocarbons found were restricted to linear alkanes, being most abundant C24 to C27 that comprises ca. 65% of the total. Olefins were not detected. EAG and behavioral test responses to the cuticular hydrocarbons were greater and significantly different from the control and the high selectivity of the antennae to the extract indicates its potential as chemical messenger. Cuticular hydrocarbons mixture is species-specific and can be used to identify a given taxon without the diagnostic castes, soldiers or imagoes Difference in the composition appears to relate with the type of habitat of specie.

Neotenic formation in laboratory colonies of the termite Coptotermes gestroi after orphaning.

The termite Coptotermes gestroi (Wasmann 1896) (Rhinotermitidae: Coptotermitinae) is an exotic species in Brazil and information concerning its reproductive developmental biology is scarce. We induced the formation of neotenics in laboratory colonies through orphaning experiments. Orphaning experiments were conducted in three-year old colonies of C. gestroi kept under laboratory conditions. After three months, eight nymphoid neotenics were observed in one colony after queen removal. Histological analysis showed that these neotenics were non-functional. The results suggest that these individuals may have arisen from the first nymphal instar (N1) or from an early N1 instar after one or two larval moults. Neotenics also were recorded on two incipient colonies of C. gestroi that lost the queen naturally.

Segmental specializations in the Malpighian tubules of the fire ant Solenopsis saevissima Forel 1904 (Myrmicinae): an electron microscopical study.

The Malpighian tubules of workers of the fire ant Solenopsis saevissima (Myrmicinae) were analyzed by scanning and transmission electron microscopy in order to determine their functional organization and association with the hindgut epithelium. The ants showed six Malpighian tubules with three segments morphologically and structurally different. The proximal segment was long and its cells showed abundant smooth endoplasmic reticulum and lipid droplets, which suggest their role in lipid secretion. The mid segment was long and undulated and it was composed by the cells that showed the typical features of ion transporting epithelia. The distal segment, short and flattened, adheres to the rectum wall. The cells of this segment showed the basal lamina fused to that of the rectum, it is probable that this part of the tubule may play a role in ion and water uptake from the feces.