This study aimed to compare canopy herbivore diversity and resultant insect damage to vegetation in two distinct and adjacent ecosystems, specifically a dry forest ecosystem and a cerrado (savanna) ecosystem that occur together in an abrupt transition zone in southeastern Brazil. In the dry forest, the canopy was reached using a single rope climbing technique, whereas the shorter canopy of the cerrado was assessed using a 7m ladder. Insect specimens were collected by beating the foliage, and 20 representative leaves were collected to calculate the specific leaf mass (SLM) and leaf area loss through herbivory. Also, we collected ten soil samples from each habitat to determine soil nutrient content. We sampled 118 herbivorous insects from ten families, mostly in dry forest trees (96 individuals belonging to 31 species). A higher abundance of chewing and sap-sucking insects were observed in dry forest trees than in cerrado trees. The same pattern was observed for the richness of chewers, with a higher degree of diversity of chewers found in dry forest trees than in cerrado trees. Herbivorous insects were not affected by SLM regardless of guild and habitat. However, we observed a negative correlation between the herbivory rate and the specific leaf mass (SLM). The cerrado trees showed a higher SLM and lower herbivory rates than trees occurring in the dry forest. These results suggest that herbivory rates in the transition dry forest‐cerrado may be driven by soil nutrient content, which is thought to influence leaf sclerophylly.

Canopy Herbivory and Insect Herbivore Diversity in a Dry Forest–Savanna Transition in Brazil

Plants have traits against herbivory that may occur together and increase defense efficiency. We tested whether there are defense syndromes in a cerrado community and, if so, whether there is a phylogenetic signal in them. We measured nine defense traits from a woodland cerrado community in southeastern Brazil. We tested the correlation between all pairs of traits and grouped the species into defense syndromes according to their traits. Most pairwise correlations of traits were complementary. Plants with lower specific leaf area also presented tougher leaves, with low nitrogen, more trichomes, and tannins. We found five syndromes: two with low defenses and high nutritional quality, two with high defenses and low nutritional quality, and one with traits compensating each other. There were two predominant strategies against herbivory in cerrado: “tolerance” and “low nutritional quality” syndromes. Phylogeny did not determine the suite of traits species presented. We argue that herbivory exerts significant selection pressure on these plant defense traits.

Defense syndromes against herbivory in a cerrado plant community

This study determined the temporal patterns of herbivory on Handroanthus ochraceus (Cham.) Mattos (Bignoniaceae) along a successional gradient in a seasonally dry tropical forest (SDTF) in southeastern Brazil. We assessed the diversity of free-feeding herbivore insects (sap-suckers and leaf-chewers), leaf herbivory rates, leaf nitrogen content, phenolic compounds, and spider abundance through the rainy season in three different successional stages (early, intermediate, and late). Sampling was conducted in December, at the beginning of the rainy season (with fully expanded young leaves), February (mid-aged leaves), and April, at the end of rainy season (old leaves). Fifteen reproductive trees of H. ochraceus were sampled per successional stage in each month of sampling. Herbivore diversity was highest in the early stage of succession, but herbivory rates were highest in the intermediate and late stages. This result was probably related to differences in herbivore community composition and leaf quality across successional stages. The highest herbivore abundance was found in April in the early successional stage. In addition, we found low levels of herbivory in the intermediate and late successional stages in the second half of the rainy season. For each successional stage, leaf nitrogen content decreased through the rainy season, whereas the concentration of phenolic compounds increased. For the intermediate and late successional stages, the temporal changes that took place as the rainy season progressed corroborated the following hypotheses postulated for SDTFs: (1) both the abundance of chewing insects and herbivory rates decreased, (2) the abundance of natural enemies (i.e., spiders) increased, and (3) leaf quality decreased. These results suggest that the described herbivory patterns are robust for advanced successional stages (intermediate and late) of the SDTFs, but may not apply to early successional stages of these forests.

Herbivory on Handroanthus ochraceus (Bignoniaceae) along a successional gradient in a tropical dry forest

Plant development is the main factor that determines the insect-ontogeny interaction, since it leads to variations in resource quality and availability. The aim of this study was to test the hypothesis that plant development and varying tannin concentration leads to changes in species richness, abundance and composition of ants, free-feeding herbivores and galling insects associated with Copaifera langsdorffii (Fabaceae). The plant ontogeny and tannin concentration effects on insects were tested on 60 individuals with height varying from 0.9 to 11.0 m. A positive correlation was observed for tree height and species richness and abundance of ants, free-feeding and galling insects. In contrast, we did not find a significant relation between leaf tannin concentration and plant height, or richness and abundance of the different insect guilds. The assemblage of ants (composition of species) did not change between saplings and adults of C. langsdorffii. However, the assemblage of free-feeding herbivores and galling insects varied between the two development stages studied. The present study reveals an ontogenetic succession pattern for herbivore insects along the C. langsdorffii growth, probably due to both indirect and direct benefits from the host plant architecture and quality. Those plants with more complex architectures should support a wider diversity of insects, since they present higher number of sites for egg laying, housing, feeding and better environmental conditions. This is the first work to investigate the host plant ontogeny effect on insects in Cerrado “Savanna” vegetation. The pattern described, along with other previous studies, suggests a vast occurrence of ontogenetic succession in tropical areas.

Relationship between plant development, tannin concentration and insects associated with Copaifera langsdorffii (Fabaceae)

Herbivores are attracted to young shoots and leaves because of their tender tissues. However, in extrafloral nectaried plants, young leaves also attract patrolling ants, which may chase or prey on herbivores. We examined this scenario in extrafloral nectaried shrubs of Banisteriopsis malifolia resprouting after fire, which promoted both the aseasonal production of leaves and the activity of extrafloral nectaries (EFNs). Results were compared between resprouting (burned) and unburned control plants. The aggressive ant species Camponotus crassus and the herbivorous thrips Pseudophilothrips obscuricornis were respectively rapidly attracted to resprouting plants because of the active EFNs and their less sclerophyllous leaves. The abundance of these insects was almost negligible in the control (unburned) shrubs. Ants failed to protect B. malifolia, as no thrips were preyed upon or injured by ants in resprouting plants. Consequently, on average, 37 % of leaves from resprouting shrubs had necrosis marks. Upon contact with ants, thrips released small liquid droplets from their abdomen, which rapidly displaced ants from the surroundings. This study shows that P. obscuricornis disrupted the facultative mutualism between C. crassus and B. malifolia, since ants received extrafloral nectar from plants, but were unable to deter herbivore thrips.

Fire triggers the activity of extrafloral nectaries, but ants fail to protect the plant against herbivores in a neotropical savanna

Susceptibility of Didymopanax vinosum (Apiaceae) to insect herbivores was investigated in three sites of a cerrado mosaic - composed of campo cerrado (a grassland with scattered trees and shrubs), cerradao (a tall woodland) and cerrado sensu stricto (intermediate between the two) - situated in Cerrado Pe-de-Gigante, Santa Rita do Passa Quatro, SP, Brazil. We also examined the relationship of folivory with the composition and abundance of the insect herbivore fauna, and with several nutritional and defensive plant characteristics (water, nitrogen, cellulose, lignin, tannin leaf contents, and leaf toughness). We collected insects associated with D. vinosum every month, and we measured leaf damage every three months. In general, the annual folivory differed among sites. It reached the highest rates in site 1 and site 3: 7.33 and 8.5 percent, respectively. Only 1.32 percent of annual folivory was observed in site 2. These levels resulted from the higher abundance, in sites 1 and 3, of the thrips Liothrips didymopanacis (Phlaeothripidae), the most abundant herbivore sampled, responsible for more than 90 percent of the observed damage. However, no significant relationship was found between insect activity and the chemical and physical composition of the leaves. Our findings suggest that, at least in this species, other chemical compounds or variables related to plant apparency and resource availability to herbivores (e.g. plant architecture) might play a more decisive role in the spatial variation of folivory than the nutritional and defensive traits that were analyzed.

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M. P. Pais

Papers

Insect folivory in Didymopanax vinosum (Apiaceae) in a vegetation mosaic of Brazilian cerrado.