Anselmo Nogueira

Vertical distance from drainage drives floristic composition changes in an Amazonian rainforest

Background: Plant composition changes with topography and edaphic gradients that correlate with soil-water and nutrient availability. Data on soil water for the Amazon Basin are scarce, limiting the possibility of distinguishing between soil and soil-water influences on plant composition. Aim: We tested a new proxy for water table depth, the terrain height above nearest drainage (HAND), as a predictor of composition in trees, lianas, palms, shrubs, and herbs and compared HAND to conventional measures of height above sea level (HASL) and horizontal distances from nearest drainage (HDND). Methods: Plant-species composition in 72 plots distributed across 64 km2 of lowland evergreen terra firme forest was summarised using non-metric multidimensional scaling (NMDS). NMDS scores were regressed against estimates of HAND, HASL and HDND. Results: Plant composition was highly correlated with the vertical distance from water table, capturing up to 82% of variation. All life forms showed highest turnover rates in the zone with seasonally water-saturated soils, which can extend 350 m from stream margins. Conclusions: Floristic composition is closely related to water table depth, and HAND appears to be the most robust available topographical metric of soil-water gradients. Brazilian conservation laws protecting 30-m-wide riparian buffers are likely to be too narrow to encompass the full zone of highest floristic turnover and may be ineffective in safeguarding riparian plant diversity.

Do extrafloral nectaries present a defensive role against herbivores in two species of the family Bignoniaceae in a Neotropical savannas

Despite the general belief that the interaction between extrafloral nectaries (EFNs) and ants is mutualistic, the defensive function of EFNs has been poorly documented in South American savannas. In this article, we evaluate the potential impact of EFNs (benefits and costs) on two species of plants from the dry areas of Central Brazil, Anemopaegma album and Anemopaegma scabriusculum (Bignoniaceae). In particular, we characterize the composition of substances secreted by the EFNs, test whether EFNs attract ants, and whether ants actually present a defensive role, leading to reduced herbivory and increased plant fitness. Histochemical analyses indicated that EFNs from both species of Anemopaegma secrete an exudate that is composed of sugars, and potentially lipids and proteins. Furthermore, EFNs from both species were shown to present a significant role in ant attraction. However, contrary to common expectations, ants were not found to protect plants against herbivore attack. No effect was found between ant visitation and flower or fruit production in A. album, while the presence of ants led to a significant decrease in flower production in A. scabriusculum. These results suggest that EFNs might present a similar cost and benefit in A. album, and a higher cost than benefit in A. scabriusculum. Since the ancestor of Anemopaegma occupied humid forests and already presented EFNs that were maintained in subsequent lineages that occupied savannas, we suggest that phylogenetic inertia might explain the presence of EFNs in the species of Anemopaegma in which EFNs lack a defensive function.

Trichome structure and evolution in Neotropical lianas

BACKGROUND AND AIMS Trichomes are epidermal outgrowths generally associated with protection against herbivores and/or desiccation that are widely distributed from ferns to angiosperms. Patterns of topological variation and morphological evolution of trichomes are still scarce in the literature, preventing valid comparisons across taxa. This study integrates detailed morphoanatomical data and the evolutionary history of the tribe Bignonieae (Bignoniaceae) in order to gain a better understanding of current diversity and evolution of trichome types. METHODS Two sampling schemes were used to characterize trichome types: (1) macromorphological characterization of all 105 species currently included in Bignonieae; and (2) micromorphological characterization of 16 selected species. Individual trichome morphotypes were coded as binary in each vegetative plant part, and trichome density and size were coded as multistate. Ancestral character state reconstructions were conducted using maximum likelihood (ML) assumptions. KEY RESULTS Two main functional trichome categories were found: non-glandular and glandular. In glandular trichomes, three morphotypes were recognized: peltate (Pg), stipitate (Sg) and patelliform/cupular (P/Cg) trichomes. Non-glandular trichomes were uniseriate, uni- or multicellular and simple or branched. Pg and P/Cg trichomes were multicellular and non-vascularized with three clearly distinct cell layers. Sg trichomes were multicellular, uniseriate and long-stalked. ML ancestral character state reconstructions suggested that the most recent common ancestor (MRCA) of Bignonieae probably had non-glandular, Pg and P/Cg trichomes, with each trichome type presenting alternative histories of appearance on the different plant parts. For example, the MRCA of Bignonieae probably had non-glandular trichomes on the stems, prophylls, petiole, petiolule and leaflet veins while P/Cg trichomes were restricted to leaflet blades. Sg trichomes were not present in the MRCA of Bignonieae independently of the position of these trichomes. These trichomes had at least eight independent origins in tribe. CONCLUSIONS The patterns of trichome evolution indicate that most morphotypes are probably homologous in Bignonieae and could be treated under the same name based on its morphological similarity and common evolutionary history, in spite of the plethora of names that have been previously designated in the literature. The trichome descriptions presented here will facilitate comparisons across taxa, allowing inferences on the relationsthips between trichome variants and future studies about their functional properties.

Evolution of extrafloral nectaries: adaptive process and selective regime changes from forest to savanna

Much effort has been devoted to understanding the function of extrafloral nectaries (EFNs) for ant–plant–herbivore interactions. However, the pattern of evolution of such structures throughout the history of plant lineages remains unexplored. In this study, we used empirical knowledge on plant defences mediated by ants as a theoretical framework to test specific hypotheses about the adaptive role of EFNs during plant evolution. Emphasis was given to different processes (neutral or adaptive) and factors (habitat change and trade‐offs with new trichomes) that may have affected the evolution of ant–plant associations. We measured seven EFN quantitative traits in all 105 species included in a well‐supported phylogeny of the tribe Bignonieae (Bignoniaceae) and collected field data on ant–EFN interactions in 32 species. We identified a positive association between ant visitation (a surrogate of ant guarding) and the abundance of EFNs in vegetative plant parts and rejected the hypothesis of phylogenetic conservatism of EFNs, with most traits presenting K‐values < 1. Modelling the evolution of EFN traits using maximum likelihood approaches further suggested adaptive evolution, with static‐optimum models showing a better fit than purely drift models. In addition, the abundance of EFNs was associated with habitat shifts (with a decrease in the abundance of EFNs from forest to savannas), and a potential trade‐off was detected between the abundance of EFNs and estipitate glandular trichomes (i.e. trichomes with sticky secretion). These evolutionary associations suggest divergent selection between species as well as explains K‐values < 1. Experimental studies with multiple lineages of forest and savanna taxa may improve our understanding of the role of nectaries in plants. Overall, our results suggest that the evolution of EFNs was likely associated with the adaptive process which probably played an important role in the diversification of this plant group.

Floral nectar production and nectary structure of a bee‐pollinated shrub from Neotropical savanna

Biotic pollination is critical for tropical ecosystem functioning, and nectar plays an essential role as it represents the main trophic resource for pollinators. Nevertheless, little is known about the mechanisms that underlie its production, which is essential for understanding the basis of nectar-mediated interactions in ecological and evolutionary approaches. Therefore, this study explores the relationship between the nectar secretion pattern and nectary functional changes in Anemopaegma album, a bee-pollinated species. We analysed the pattern of nectar production under field conditions and investigated floral nectary structural changes in two different developmental stages using light, transmission and scanning electron microscopy. We measured 30.95 ± 23.02 μl (mean ± SD, n = 30) of nectar accumulated inside the nectar chamber (29.26 ± 3.48% sucrose equivalents) at the moment of flower opening. Nectar removal did not influence the pattern of floral nectar production in terms of volume or total sugar but reduced the concentration of the nectar produced during the first 24 h of anthesis. The nectary consisted of an epidermis, a nectary parenchyma and a subnectary parenchyma supplied only by phloem. Starch grains decreased in size and abundance from the subnectary parenchyma toward the epidermis. We observed the degradation of starch grains and incorporation of amyloplasts into vacuoles at the pre-anthesis stage as well as the transformation of amyloplasts into elaioplasts during anthesis. Nectar secretion was continuous during the A. album flower life span, which was related to the functional features of its floral nectary, especially the presence of starch stored in the parenchyma.

Geographic Mosaic of Plant Evolution: Extrafloral Nectary Variation Mediated by Ant and Herbivore Assemblages

Herbivory is an ecological process that is known to generate different patterns of selection on defensive plant traits across populations. Studies on this topic could greatly benefit from the general framework of the Geographic Mosaic Theory of Coevolution (GMT). Here, we hypothesize that herbivory represents a strong pressure for extrafloral nectary (EFN) bearing plants, with differences in herbivore and ant visitor assemblages leading to different evolutionary pressures among localities and ultimately to differences in EFN abundance and function. In this study, we investigate this hypothesis by analyzing 10 populations of Anemopaegma album (30 individuals per population) distributed through ca. 600 km of Neotropical savanna and covering most of the geographic range of this plant species. A common garden experiment revealed a phenotypic differentiation in EFN abundance, in which field and experimental plants showed a similar pattern of EFN variation among populations. We also did not find significant correlations between EFN traits and ant abundance, herbivory and plant performance across localities. Instead, a more complex pattern of ant–EFN variation, a geographic mosaic, emerged throughout the geographical range of A. album. We modeled the functional relationship between EFNs and ant traits across ant species and extended this phenotypic interface to characterize local situations of phenotypic matching and mismatching at the population level. Two distinct types of phenotypic matching emerged throughout populations: (1) a population with smaller ants (Crematogaster crinosa) matched with low abundance of EFNs; and (2) seven populations with bigger ants (Camponotus species) matched with higher EFN abundances. Three matched populations showed the highest plant performance and narrower variance of EFN abundance, representing potential plant evolutionary hotspots. Cases of mismatched and matched populations with the lowest performance were associated with abundant and highly detrimental herbivores. Our findings provide insights on the ecology and evolution of plant–ant guarding systems, and suggest new directions to research on facultative mutualistic interactions at wide geographic scales.

Morphology, secretion composition, and ecological aspects of stipular colleters in Rubiaceae species from tropical forest and savanna.

Colleters are secretory structures that produce and release mucilage or a mucilage-resin mixture protecting meristems and young structures against desiccation, herbivores, and pathogens. The secretions may vary in colleters of same or different types, indicating that the functionality of colleters may be more specific than previously thought. In this study, we compared 17 Rubiaceae species from savanna and forest environment focusing on colleter secretions and its ecological role. First, we evaluated the morphology, distribution, and histochemistry of stipular colleters using light and scanning electron microscopy. Additionally, we investigated the phenology, microclimate, and the proportion of damaged apices in the savanna and forest species. We recorded standard-type colleters, variable in distribution and size, in 14 of the 17 studied species. The secretion varied from predominantly hydrophilic, mixed to predominantly lipophilic. During the budding period, secretion covered the vegetative apices. Savanna species had a prevalence of lipid secretion in habitats with higher luminosity, which had a lower proportion of damaged apices. In contrast, forest species occurred in habitats with lower luminosity and had a higher proportion of damaged apices, in general with the absence of lipids in the colleters. These results highlight that colleters with similar morphology clearly differed in secretions among species, especially between species from savanna and forest, in which the colleters appear potentially associated with protection against irradiation in savanna, but not in the forest environment.