Seed predators limit plant recruitment in Neotropical savannas

Despite the well‐documented impacts of consumers on seed abundance the link between seed predation and plant population dynamics remains poorly understood because experimental studies linking patterns of predation with seedling establishment are rare. We used experimental manipulations with six woody plant species to elucidate the effects of seed predator type, habitat, and plant species identity on rates of seed predation and seedling recruitment in the Neotropical savannas known as the Cerrado. We found that seed predation rates are consistently high across a diversity of local habitat types, with important inter‐habitat variation in seed predation for three of the six species used in our experiments. We also found that seed predation has a clear demographic signal – experimentally excluding predators resulted in higher rates of seedling establishment over the course of two seasons. Because the intensity of seed predation varied between species and habitats, it may play a role in structuring local patterns of plant abundance and community composition. Finally, our results lend support to the recent hypothesis that herbivores have major and underappreciated impacts in Neotropical savannas, and that top–down factors can influence the demography of plants in this extensive and biodiversity‐rich biome in previously unexplored ways.     

木本植物多度在草原和稀树干草原中增加的研究进展

木本植物多度在草原和稀树干草原中增加已经成为全球范围普遍发生的现象。为揭示这一现象发生的原因,从放牧和气候变化与木本植物多度增加的关系、木本植物多度增加过程中的正反馈作用以及木本植物侵入的关键阶段——幼苗的补充和定居,这三个方面综述了目前的研究结果。强调放牧和气候变化之间的相互共同作用,可能引发了木本植物向草原和稀树干草原中的入侵;而生物引起的正反馈作用则进一步促进了木本植物的扩展。从生态系统干扰的角度,讨论了木本植物多度增加机制的复杂性,并指出木本植物幼苗补充和定居的连续性和间断性两种方式,对于草原和稀树干草原木本植物多度增加的贡献。     

Secondary metabolites and nutrients of woody plants in relation to browsing intensity in African savannas

Carbon-based secondary metabolites (CBSMs) are assumed to function as defences that contribute to herbivore-avoidance strategies of woody plants. Severe browsing has been reported to reduce concentrations of CBSMs and increase N concentrations in individual plants, causing heavily browsed plants to be characterised by N-rich/C-poor tissues. We hypothesised that concentrations of condensed tannins (CT) and total polyphenols (TP) should decrease, or N increase, in relation to increasing intensity of browsing, rendering severely browsed plants potentially more palatable (increased N:CT) and less N-limited (increased N:P) than lightly browsed ones. We sampled naturally browsed trees (taller than 2 m) of four abundant species in southern Kruger National Park, South Africa. Species-specific relationships between N:CT, CT, TP and P concentrations and increasing browsing intensity were detected, but N and N:P were consistently invariable. We developed a conceptual post-hoc model to explain diverse species-specific CBSM responses on the basis of relative allocation of C to total C-based defence traits (e.g. spines/thorns, tough/evergreen leaves, phenolic compounds). The model suggests that species with low allocation of C to C-based defence traits become C-limited (potentially more palatable) at higher browsing intensity than species with high allocation of C to C-based defences. The model also suggests that when N availability is high, plants become C-limited at higher browsing intensity than when N availability is low.     

Neotropical savannas: Their flora and vegetation

Covering approximately three million km(2), the savannas, the second largest major plant formation in the American tropics, are gaining increasing importance for land planning and occupational strategies. There has been much recent research on the floristic and ecological characteristics of these herbaceous ecosystems, resulting in considerable progress in the understanding of their complex ecological interrelationships. Hitherto little-known savanna "islands" within the Amazon region are being intensively studied, offering interesting information on their origin and dynamic relationships with the surrounding forests. Important gaps remain to be filled, however, before a critical evaluation of the great variety of neotropical savanna ecosystems is achieved and the preservation of at least some of these unique plant formations is assured.     

Birds nesting survival in disturbed and protected Neotropical savannas

Fragmentation and other habitat disturbances are long known to negatively affect birds, in large part by decreasing nest success due to high nest predation rates. The factors, however, that cause this decrease in nest success are still poorly understood and may vary among regions or species. Here, we show that nest survival is also lower in a disturbed landscape versus a protected cerrado (savanna-like) Neotropical landscape. Also, we tested the importance of garbage in the nest, brood parasitism, microhabitat and bird family in nest survival, controlling for temporal effects. We monitored 144 birds’ nests in a disturbed landscape and 150 nests in a natural reserve of cerrado vegetation in central Brazil, between September and December 2006. We used Program MARK to estimate nest survival probabilities and evaluate the effect of covariates in nest success in the disturbed area. Nest daily survival rate (DSR) was higher in the reserve (survival probability = 29.4%) than in the disturbed landscape (survival probability = 16.6%). Nest daily survival rate (DSR) was smaller in nests with garbage (survival probability = 9.3%) than in nests without garbage (survival probability = 19.5%) in the disturbed landscape. Effects of habitat disturbance on nest survival differed among bird families, with finches and tanagers being more affected mostly due to high nest predation rates. Conservation and management of birds in disturbed landscapes should include actions to decrease nest predation. In poor rural or suburban areas in developing countries, such as Brazil, actions like better garbage treatment may help conserve birds in disturbed landscapes.     

Determinants of woody encroachment and cover in African savannas

Savanna ecosystems are an integral part of the African landscape and sustain the livelihoods of millions of people. Woody encroachment in savannas is a widespread phenomenon but its causes are widely debated. We review the extensive literature on woody encroachment to help improve understanding of the possible causes and to highlight where and how future scientific efforts to fully understand these causes should be focused. Rainfall is the most important determinant of maximum woody cover across Africa, but fire and herbivory interact to reduce woody cover below the maximum at many locations. We postulate that woody encroachment is most likely driven by CO₂ enrichment and propose a two-system conceptual framework, whereby mechanisms of woody encroachment differ depending on whether the savanna is a wet or dry system. In dry savannas, the increased water-use efficiency in plants relaxes precipitation-driven constraints and increases woody growth. In wet savannas, the increase of carbon allocation to tree roots results in faster recovery rates after disturbance and a greater likelihood of reaching sexual maturity. Our proposed framework can be tested using a mixture of experimental and earth observational techniques. At a local level, changes in precipitation, burning regimes or herbivory could be driving woody encroachment, but are unlikely to be the explanation of this continent-wide phenomenon.     

The role of fire in structuring trait variability in Neotropical savannas

Intraspecific trait variability plays a fundamental role in community structure and dynamics; however, few studies have evaluated its relative importance to the overall response of communities to environmental pressures. Since fire is considered a key factor in Neotropical savannas, we investigated to what extent the functional effects of fire in a Brazilian savanna occurs via intra- or interspecific trait variability. We sampled 12 traits in communities subjected to three fire regimes in the last 12 years: annual, biennial, and protected. To evaluate fire’s relative effects, we fitted a general linear mixed models with species as random and fire as fixed factors, using: (1) all species in the communities (i.e., considering intra- and interspecific variabilities); (2) 18 species common to all fire regimes (i.e., intraspecific variability only); and (3) all species with their overall average trait values (i.e., interspecific variability only). We assessed the relative role of intra- or interspecific variability by comparing the significance of each trait in the three analyses. We also compared the within and between fire variabilities with a variance component analysis. Five traits presented larger intraspecific than interspecific variability, and the main effect of fire occurred at the intraspecific level. These results confirm that it is important to consider intraspecific variability to fully understand fire-prone communities. Moreover, trait variability was larger within than among fire regimes. Thus, fire may act more as an external filter, preventing some of the species from the regional pool from colonizing the cerrado, than as an internal factor structuring the already filtered cerrado communities.     

Grazing and neighborhood interactions limit woody encroachment in wet subtropical savannas

Woody encroachment in savannas is a worldwide concern, and there is growing consensus that anthropogenic activities play a central role in changing tree – grass interactions. We evaluated the influence of livestock grazing and neighborhood interactions on seedling emergence and survival of the native tree Acacia caven in wet savannas of northeastern Argentina. We hypothesized that grazing and grass competition act as biotic barriers limiting tree recruitment, but the relative magnitude of such barriers differs according to grass patch type. In two consecutive years (cohort 1 and 2) we sowed seeds and transplanted seedlings of Acacia in two grass patch types (prostrate/palatable and tussock/unpalatable grasses) in both, grazed and ungrazed plots. Each grass patch type was further manipulated to create three levels of grass competition (unclipped control, above-ground biomass removal and total biomass removal).Cattle grazing diminished seedling emergence of both cohorts and seedling survival of cohort 1. The effect of grass competition changed according to grass patch type. Prostrate grass cover enhanced emergence but lowered early survival, while tussock grass cover and also its total biomass removal facilitated early survival. During the second year, a severe drought drastically reduced Acacia recruitment, and it was strong enough to eliminate any grazing effects although the effect of grass competition on seedling establishment remained significant.Our results suggest that grazing and grass competition additively diminished the risk of woody establishment in this wet savanna. However, the stocking rate should be carefully balanced, thus contributing to the maintenance of a competitive grass cover to limit tree recruitment.     

Phytoalexins of woody plants

Summary Phytoalexins accumulated in selected woody plants in response to microbial attack or stress are reviewed and listed with respect to their chemical structure and probable biogenetic origin. The host-pathogen systems from which they have been isolated are described. The review also considers the antimicrobial activity of the phytoalexins to the causal pathogens and other microorganisms.     

Drought resistance in woody plants

The Botanical Review -     

Rainfall and topo-edaphic influences on woody community phenology in South African savannas

Woody community phenology was studied in the central lowveld, South Africa, over a twelve month period at three sites along a rainfall gradient, with both toplands and bottomlands sampled at each site. Each month, individual plants, in replicated samples, were scored into a number of categories describing their phenological state. Position on the rainfall gradient influenced: (1) onset and magnitude of leaf emergence, (2) onset and duration of mature leaves, and (3) the proportion of leafless trees. Generally, the moist site demonstrated earlier leaf growth than the intermediate or arid sites. Emergent and mature leaves were recorded earlier, and in the case of mature leaves, retained longer. Overall, there was a lower proportion of leafless trees during the dry season at the moist site, followed by the semi-arid site, followed by the arid site. Differences with respect to catenal position were evident for the proportion of trees in winter with mature leaves, and the proportion of trees with senescent leaves. Bottomlands had a greater proportion of trees with leaves during winter, but a lower proportion of trees recorded with senescent leaves. Both of these findings were a result of the greater proportion of evergreen species in bottomlands, as well as increased leaf retention by the deciduous species. Phenological activity of leaves was related to plant stem size. In particular, there was greater leaf retention during the dry period by small stems, relative to large stems.     

Classifying the Neotropical savannas of Belize using remote sensing and ground survey

Aim This paper evaluates a method of combining data from GPS ground survey with classifications of medium spatial resolution LANDSAT imagery to distinguish variations within Neotropical savannas and to characterize the boundaries between savanna areas and the associated gallery forests, seasonally dry forests and wetland communities. Location Rio Bravo Conservation Area, Orange Walk District, Belize, Central America. Methods Dry season LANDSAT data for 10 April 1993 and 9 March 2001 covering a conservation area of 240,000 acres (97,459 ha), were rectified to sub‐pixel accuracy using ground control points positioned by GPS ground survey. The 1993 image was used to assess the accuracy with which the boundaries between the savanna matrix and gallery forests, high forests, wetlands and water bodies could be discriminated. The image was classified by a maximum likelihood (ML) classifier and the shapes and areas of forest and wetland classes were compared with an interpretation of these land cover types from 1 : 24,000 aerial photography, mapped at 1 : 50,000 scale in 1993. The 2001 image was used to assess whether different subtypes of savanna could be distinguished from LANDSAT data. This required the creation of a reference (‘ground truth’) data set for testing classifications of the image. One hundred and sixty sample patches (650 ha, distributed over an area of 7000 ha) of ten sub‐types of savanna vegetation and associates identified using a physiognomic classification scheme, were delineated on the ground by GPS and divided into two subsets for training and testing. Continuous classifications of LANDSAT data covering the savannas were developed that estimated potential contributions from up to five sub‐types of land cover (grassland, wetland, pine woodland, gallery forest and palmetto). The accuracy of each classification was assessed by comparison against ground data. An ML classification was also produced for the 2001 image using the same areas for training. This allowed a comparison of the relative accuracy of both continuous and Boolean ML methods for classifying savanna areas. Results The boundary between savannas and evergreen forests, gallery forests and open water in the study region could be delineated by the ML classifier to within 2 pixels (60 m) using LANDSAT imagery. However, the constituent sub‐types within the savanna were poorly discriminated. Whilst the shape and extent of closed canopy forest, gallery forest, wetlands and water bodies agreed closely with the distributions interpreted from aerial photography, classes such as ‘open pine savanna’ or ‘grassland’ were only 45–65% accurate when tested against ground data. A continuous classification, estimating the proportions of three savanna vegetation subtypes (grassland, marshland and woodland) present in each pixel, correctly classified more of the ground data for these cover types than the comparable ML result. Proportional mixtures of the land cover estimated by the continuous classifier also compared realistically with the vegetation formations observed along ground transects. Main conclusions By using GPS, a ground survey of vegetation cover was accurately matched to remotely sensed imagery and the accuracy of delineating boundaries and classifying areas of savanna was assessed directly. This showed that ML classification techniques can reliably delineate the boundaries of savannas, but continuous classifiers more accurately and realistically represent the distribution of the subtypes comprising savanna land cover. By combining these ground survey and image classification methods, medium spatial resolution satellite sensor data can provide an affordable means for land managers to assess the nature, extent and distribution of savanna formations. Over time, using the archives of LANDSAT (and SPOT) data together with marker sites surveyed in the field, quantitative changes in the extents and boundaries of savannas in response to both natural (e.g. fire, hurricane and drought) and anthropogenic (e.g. cutting and disturbance) factors can be assessed.     

Rates of woody encroachment in African savannas reflect water constraints and fire disturbance

Aim

The aims of this study were to (1) estimate current rates of woody encroachment across African savannas; (2) identify relationships between change in woody cover and potential drivers, including water constraints, fire frequency and livestock density. The found relationships led us to pursue a third goal: (3) use temporal dynamics in woody cover to estimate potential woody cover.

Location

Sub‐Saharan African savannas.

Methods

The study used very high spatial resolution satellite imagery at sites with overlapping older (2002–2006) and newer (2011–2016) imagery to estimate change in woody cover. We sampled 596 sites in 38 separate areas across African savannas. Areas with high anthropogenic impact were avoided in order to more clearly identify the influence of environmental factors. Relationships between woody cover change and potential drivers were identified using linear regression and simultaneous autoregression, where the latter accounts for spatial autocorrelation.

Results

The mean annual change in woody cover across our study areas was 0.25% per year. Although we cannot explain the general trend of encroachment based on our data, we found that change rates were positively correlated with the difference between potential woody cover and actual woody cover (a proxy for water availability; p < .001), and negatively correlated with fire frequency (p < .01). Using the relationship between rates of encroachment and initial cover, we estimated potential woody cover at different rainfall levels.

Main conclusions

The results indicate that woody encroachment is ongoing and widespread across African savannas. The fact that the difference between potential and actual cover was the most significant predictor highlights the central role of water availability and tree–tree competition in controlling change in woody populations, both in water‐limited and mesic savannas. Our approach to derive potential woody cover from the woody cover change trajectories demonstrates that temporal dynamics in woody populations can be used to infer resource limitations.