Mário M. Espírito-Santo

Changes in tree and liana communities along a successional gradient in a tropical dry forest in south-eastern Brazil

We investigated changes in species composition and structure of tree and liana communities along a successional gradient in a seasonally dry tropical forest. There was a progressive increase in tree richness and all tree structural traits from early to late stages, as well as marked changes in tree species composition and dominance. This pattern is probably related to pasture management practices such as ploughing, which remove tree roots and preclude regeneration by resprouting. On the other hand, liana density decreased from intermediate to late stages, showing a negative correlation with tree density. The higher liana abundance in intermediate stage is probably due to a balanced availability of support and light availability, since these variables may show opposite trends during forest growth. Predicted succession models may represent extremes in a continuum of possible successional pathways strongly influenced by land use history, climate, soil type, and by the outcomes of tree–liana interactions.

How Many Species of Gall-Inducing Insects Are There on Earth, and Where Are They?

Abstract During the last two decades, there was considerable debate on the actual number of insect species on Earth, and current estimates range from 5 to 30 million species. For gall-inducing insects, a cosmopolitan group of specialist herbivores, the last, 40-yr-old estimate of global richness indicated 13,000 species, mostly from temperate regions. To update these estimates, we used 10 studies that provided the average number of gall-inducing insect species per plant species (galled and nongalled) in different vegetation types around the world, and a recent calculation of 422,000 seed plant species on Earth. Considering that most of the gall-inducing insects are monophagous, these data were extrapolated to the global flora, assuming a positive relationship between gall-inducing insect richness and plant richness. Estimates of the global richness of gall-inducing insects ranged from 21,000 to 211,000 species, with an average of 132,930 species. The global distribution pattern described for gall-inducing insects pointed to a higher richness in warm temperate regions with sclerophyllous vegetation. However, it is likely that the tropical rain forests would harbor a higher absolute number of gall-inducing insects, due to their outstanding plant richness. Most of this fauna will be found on the poorly explored canopies of rain forests, especially in the Neotropical region. An increased sampling effort in tropical regions should fill the taxonomy gap represented by gall-inducing insects and possibly change the biogeographic patterns described so far.

Plant architecture and meristem dynamics as the mechanisms determining the diversity of gall-inducing insects

Plant architecture is considered to affect herbivory intensity, but it is one of the least studied factors in plant–insect interactions, especially for gall-inducing insects. This study aimed to investigate the influence of plant architecture on the speciose fauna of gall-inducing insects associated with 17 species of Baccharis. Five architectural variables were evaluated: plant height, number of fourth-level shoots, biomass, average level and number of ramifications. The number of galling species associated with each host plant species was also determined. To test the effects of plant architecture on gall richness at the individual level, we used another data set where the number of fourth-level shoots and gall richness were determined for B. concinna, B. dracunculifolia, and B. ramosissima every 3 weeks during 1 year. The average similarity between host species based on gall fauna was low (9%), but plants with the same architectural pattern tended to support similar gall communities. The most important architectural trait influencing gall richness at the species level was the number of fourth-level shoots, which is indicative of the availability of plant meristems, a fundamental tissue for gall induction and development. This variable also showed a positive correlation with gall richness at the individual level. We propose that variations in gall richness among host species are driven by interspecific differences in plant architecture via availability of young, undifferentiated tissue, which is genetically controlled by the strength of the apical dominance. Plant architecture should have evolutionary consequences for gall communities, promoting insect radiation among architecturally similar plants through host shift and sympatric speciation. We also discuss the role of plant architecture in the global biogeography of gall-inducing insects.

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

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.

Plant Phenology and Absence of Sex-Biased Gall Attack on Three Species of Baccharis

Background Dioecy represents a source of variation in plant quality to herbivores due to sexual differences in intensity and timing of resource allocation to growth, defense and reproduction. Male plants have higher growth rates and should be more susceptible to herbivores than females, due to a lower investment in defense and reproduction. Methodology/Principal Findings We compared resource investment to growth and reproduction and its consequences to herbivore attack on three Baccharis species along one year (B. dracunculifolia, B. ramosissima, and B. concinna). Phenological patterns presented by the three species of Baccharis were quite different over time, but the number of fourth-level shoots and plant growth rate did not differ between sexes in any studied species. Intersexual difference in reproductive investment was only observed for B. concinna, with female individuals supporting higher inflorescence density than male individuals throughout the year. Gall abundance on the three Baccharis species was not influenced by plant sex. However, all plant traits evaluated here positively influenced the gall abundance on B. concinna, whereas only the number of fourth-level shoots positively influenced gall abundance on B. ramosissima and B. dracunculifolia. Conclusions/Significance The absence of differential reproductive allocation may have contributed to similar growth and shoot production between the sexes, with bottom-up effects resulting in gender similarities in gall abundance patterns. The number of fourth-level shoots, an indicator of meristem availability to herbivores, was the most important driver of the abundance of the galling insects regardless of host plant gender or species. Albeit the absence of intersexual variation in insect gall abundance is uncommon in the literature, the detailed study of the exceptions may bring more light to understand the mechanisms and processes behind such trend.

Efeitos de remanescentes de Mata Atlântica no controle biológico de Euselasia apisaon (Dahman) (Lepidoptera: Riodinidae) por Trichogramma maxacalii (Voegelé e Pointel) (Hymenoptera: Trichogrammatidae)

This study evaluated the effects of Atlantic Rain Forest remnants on the natural biological control of Euselasia apisaon (Dahman) by the parasitoid Trichogramma maxacalii (Voegelé e Pointel) in Eucalyptus plantations. The number of E. apisaon eggs/leaf was higher in the center than in the edge of the plantations (23.5 +/- 7.61 vs. 14.8 +/- 3.14), but parasitism showed the reversed pattern (72.4% in the center and 80.5% in the edge). The results indicated that natural regulation exerted by T. maxacalii on populations of E. apisaon may be enhanced by the preservation of fragments of native vegetation surrounding Eucalyptus plantations.

Ontogenetic and Temporal Variations in Herbivory and Defense of Handroanthus spongiosus (Bignoniaceae) in a Brazilian Tropical Dry Forest

ABSTRACT We compared the richness and abundance of free-feeding herbivore insects (sap-sucking and leaf-chewing), leaf herbivory damage, leaf toughness and total phenolic content between two ontogenetic stages (juvenile and reproductive) of Handroanthus spongiosus (Rizzini) S. O. Grose (Bignoniaceae) throughout the rainy season in a Brazilian seasonally dry tropical forest. Twenty marked individuals of H. spongiosus were sampled per ontogenetic stage in each period of the rainy season (beginning, middle, and end). Herbivore richness and abundance did not differ between ontogenetic stages, but higher percentage of leaf damage, higher concentration of phenolic compounds, and lower leaf toughness were observed for juvenile individuals. The greatest morphospecies abundance was found at the beginning of the rainy season, but folivory increment was higher at the end, despite the fact that leaf toughness and total phenolic content increased in the same period. No significant relationships between leaf damage and both total phenolic content and leaf toughness were observed. These results suggest that insect richness and abundance do not track changes in foliage quality throughout plant ontogeny, but their decrease along rainy season confirms what was predicted for tropical dry forests. The general trends described in the current study corroborate those described in the literature about herbivores and plant ontogeny. However, the lack of relationship between herbivore damage and the two plant attributes considered here indicates that the analyses of multiple defensive traits (the defense syndrome) must be more enlightening to determine the mechanisms driving temporal and spatial patterns of herbivore attack.

Phyllostomid Bat Occurrence in Successional Stages of Neotropical Dry Forests

Tropical dry forests (TDFs) are highly endangered tropical ecosystems being replaced by a complex mosaic of patches of different successional stages, agricultural fields and pasturelands. In this context, it is urgent to understand how taxa playing critical ecosystem roles respond to habitat modification. Because Phyllostomid bats provide important ecosystem services (e.g. facilitate gene flow among plant populations and promote forest regeneration), in this study we aimed to identify potential patterns on their response to TDF transformation in sites representing four different successional stages (initial, early, intermediate and late) in three Neotropical regions: México, Venezuela and Brazil. We evaluated bat occurrence at the species, ensemble (abundance) and assemblage level (species richness and composition, guild composition). We also evaluated how bat occurrence was modulated by the marked seasonality of TDFs. In general, we found high seasonal and regional specificities in phyllostomid occurrence, driven by specificities at species and guild levels. For example, highest frugivore abundance occurred in the early stage of the moistest TDF, while highest nectarivore abundance occurred in the same stage of the driest TDF. The high regional specificity of phyllostomid responses could arise from: (1) the distinctive environmental conditions of each region, (2) the specific behavior and ecological requirements of the regional bat species, (3) the composition, structure and phenological patterns of plant assemblages in the different stages, and (4) the regional landscape composition and configuration. We conclude that, in tropical seasonal environments, it is imperative to perform long-term studies considering seasonal variations in environmental conditions and plant phenology, as well as the role of landscape attributes. This approach will allow us to identify potential patterns in bat responses to habitat modification, which constitute an invaluable tool for not only bat biodiversity conservation but also for the conservation of the key ecological processes they provide.

TANNINS IN BACCHARIS DRACUNCULIFOLIA (ASTERACEAE): EFFECTS OF SEASONALITY, WATER AVAILABILITY AND PLANT SEX

Several ecological, genetic, and environmental factors are known to influence tannin concentration in plant tissues. In this study, the effects of seasonality, water availability, and sex of the plant on tannin concentration in the dioecious shrub Baccharis dracunculifolia were assessed. The effects of water availability on plant shoot growth and its relationship to tannin concentration were also experimentally evaluated. Tannins occurred in B. dracunculifolia from November to May, peaking during the summer (December to March). However, no relationship between tannin concentration, rainfall and temperature was found. No difference in tannin concentration and shoot growth was found between plants in the irrigated and control treatments, this suggesting that tannin production was affected by factors other than water availability, such as light intensity or plant phenology. There was a negative relationship between tannin concentration and shoot growth, possibly due to a trade-off between these metabolic activities. Plant gender did not influence tannin concentration and shoot growth, indicating that B. dracunculifolia did not show a differential resource allocation between male and female individuals.

Understanding patterns of land-cover change in the Brazilian Cerrado from 2000 to 2015.

Clearing tropical vegetation impacts biodiversity, the provision of ecosystem services, and thus ultimately human welfare. We quantified changes in land cover from 2000 to 2015 across the Cerrado biome of northern Minas Gerais state, Brazil. We assessed the potential biophysical and socio-economic drivers of the loss of Cerrado, natural regeneration and net cover change at the municipality level. Further, we evaluated correlations between these land change variables and indicators of human welfare. We detected extensive land-cover changes in the study area, with the conversion of 23 446 km2 and the natural regeneration of 13 926 km2, resulting in a net loss of 9520 km2. The annual net loss (−1.2% per year) of the cover of Cerrado is higher than that reported for the whole biome in similar periods. We argue that environmental and economic variables interact to underpin rates of conversion of Cerrado, most severely affecting more humid Cerrado lowlands. While rates of Cerrado regeneration are important for conservation strategies of the remaining biome, their integrity must be investigated given the likelihood of encroachment. Given the high frequency of land abandonment in tropical regions, secondary vegetation is fundamental to maintain biodiversity and ecosystem services. Finally, the impacts of Cerrado conversion on human welfare likely vary from local to regional scales, making it difficult to elaborate land-use policies based solely on socio-economic indicators. This article is part of the themed issue ‘Tropical grassy biomes: linking ecology, human use and conservation’.