Rock outcrop habitats in the Venezuelan Guayana lowlands: their main vegetation types and floristic components

The Guayana Shield, located in north-eastern South America, consists of a highly complex and composite mosaic of landscape elements. Amongst these, inselbergs are very conspicuous, because of their peculiar shape and their unusual associated vegetation. Geologically, these rock outcrops are part of the underlying ancient igneous-metamorphic basement and occur mainly in the lowlands of the periphery of the shield. As azonal habitats, inselbergs harbour a highly specialized flora. The characteristic vegetation is composed of lithophytic and savanna-like plant communities, as well as low dry forests. As a whole, the vegetation of an inselberg may be interpreted as a mosaic of marginal habitats. Therefore a large number of taxa find suitable niches in a quite condensed space. Gradients of soil depth and water availability are the main factors determining the floristic composition. A preliminary floristic inventory of the Venezuelan inselberg flora comprises 614 vascular plant species. 24% of them are endemic to the Guayana region, 15% are endemic to inselbergs of the Guayana region. The distribution patterns of these latter, eco-endemic species allows to distinguish a northern and a southern inselberg district. The two districts overlap in the Átures area, in the surroundings of Puerto Ayacucho, where a true centre of endemism is located. The distinction into two districts is emphasised by different phytogeographic relations. The southern inselberg district shows connections to the "tepui" flora, whereas the northern district reveals phytogeographic relations to the Caribbean region as well as to the Brazilian Shield. Possible explanations for the floristic interchange across the equator are discussed.

Floristics, structure and soil of insular vegetation in four quartzite-sandstone outcrops of "Chapada Diamantina", Northeast Brazil

Soil islands on rocky surfaces often harbor aggregated vegetation that consists of insular plant communities. These islands are typical of the rocky outcrops and in various parts of Brazil form the so-called "campos rupestres" vegetation. Four of such sites have been selected in the state of Bahia, Northeast Brazil, for this comparative study on floristics and vegetation structure: three areas situated inside the "Parque Nacional da Chapada Diamantina" (Guiné, Fumaça and "Gerais da Fumaça") and one is at the border of the Environmental Protection Area of "Marimbus-Iraquara" ("Mãe Inácia"). All occurring vegetation islands were studied in four random plots of 10 × 10 m per site. Soil was often shallow, sandy and acidic. Vascular plant species were determined, with respective life forms and canopy coverage areas. The total number of species when all four sites were added was 135, and the number of species per island varied from 2 to 32. The areas of the 214 soil islands varied from 0.015 to 91.9 m², totaling 568 m² in the four sites. Monocotyledon families were dominant, essentially Velloziaceae, as well as Orchidaceae, Bromeliaceae, Amaryllidaceae and Cyperaceae. Among the eudicotyledons, dominant families were mainly Clusiaceae, Asteraceae and Melastomataceae. The biological spectra revealed that phanerophytes and hemicryptophytes predominated among the life forms, while chamaephytes had the largest coverage area. Epilithic and desiccant chamaephytes composed the most conspicuous interspecific associations, and were probably related to early successional processes. Sites closest to one another were not the most similar in structure, indicating that other factors more relevant than distance might be involved in the abundance of species in space.

Structural analysis of the vegetation on a highland granitic rock outcrop in Southeast Brazil

Granitic rock outcrops of the Brazilian southeast are either coastal or inland. The latter can often have high altitudes, such as in the summits of "Serra do Mar" and "Serra da Mantiqueira", where they are known as "Campos de Altitude". The landscape on these high altitude plateaux is often a mosaic of shrubs and treelets within a bunchgrass matrix, with sparse pteridophytes and other herbs, interspersed with variable extensions of rock outcrops. Despite the pervasiveness of rock outcrops in the Brazilian landscape, studies on the structural analysis of the vegetation on such formations are scarce. This study aimed to analyze the structure of the vegetation on a highland granitic rock outcrop in "Parque Estadual da Serra do Brigadeiro", state of Minas Gerais, Southeast Brazil (42°20' - 42°40' S and 20°20' - 21°00' W, 1,722 m of elevation ). Quantitative parameters of absolute and relative frequency and dominance (cover) were estimated. The group analysis used the Jaccard similarity index. Trilepis lhotzkiana, Panicum sp. 1, and Vellozia variegata presented the highest relative frequencies, relative dominances and importance values. These three species, along with Dyckia bracteata, Rhynchospora emaciate, and Tibouchina cf. manicata, represented 98.3% of the relative dominance. The remaining 1.7% referred to 22 remaining species. The distinction among quadrats within formed groups by cluster analysis was due to the great number of low frequency species.

The potential of natural regeneration of rocky outcrop vegetation on rupestrian field soils in "Serra do Cipó", Brazil

An area's innate potential to regenerate represents a crucial factor for its conservation and management. The seed rain and seed bank are important agents in the regeneration process. Seed banks are particularly important in communities where there is a high proportion of obligate seeders. Rocky outcrops are habitats where most part of the plant species depends on their seeds to reproduce and maintain viable populations. Therefore, seed banks ought to be important in this vegetation physiognomy. We test the hypotheses that the seed bank of the rocky outcrops found in the rupestrian fields of "Serra do Cipó", Brazil, is richer in species and denser than those formed on different vegetation physiognomies neighboring the outcrops. We then compared species abundance, species richness and composition in the rocky outcrops' seed banks with those of sandy and peaty bogs, forests, gallery forests, and "cerrados". Furthermore, we report on the natural regeneration potential of these soils by assessing a greenhouse study on seedling emergence. Soil samples were collected from 0 to 5 and 5 to 10 cm of depth. Rocky outcrops had the poorest in species and less dense seed bank and showed segregation in species composition. Emergence was greater in the most superficial layer. However, soils on rocky outcrops showed the greatest proportion of endemic threatened species in their seed banks, demonstrating their importance for biodiversity conservation of the "Serra do Cipó" rupestrian fields.

Reproductive phenology of Melocactus (Cactaceae) species from Chapada Diamantina, Bahia, Brazil

This paper discusses the phenological strategies of Melocactus glaucescens Buining & Brederoo, M. paucispinus G. Heimen & R. Paul, M. ernestii Vaupel and M. ×albicephalus Buining & Brederoo, species from Chapada Diamantina, northeastern Brazil. Melocactus glaucescens, M. ernestii and M. ×albicephalus occur sympatrically in an area of "caatinga"/"cerrado" vegetation, and M. paucispinus in an area of "cerrado"/"campo rupestre". The superposition of flowering in these sympatric taxa was compared and analyzed. The phenology of M. paucispinus was correlated with both abiotic and biotic factors. Flowering of M. glaucescens and M. ×albicephalus were observed to be continuous (though with moderate peaks of activity), while fruiting was sub-annual. Melocactus ernestii exhibited an annual pattern of both flowering and fruiting; while in M. paucispinus the same patterns were sub-annual. These sympatric taxa showed 40% overlap of flowering periods, reaching to more than 50% in paired combinations of taxa, considering both the number of specimens flowering, as well as the quantity of resources being offered. Available information indicates that these taxa share pollinators, but phenological data rejects the hypothesis of shared pollinators and supports the hypothesis of hybridization in the study area. Rainfall was negatively correlated with flowering in M. paucispinus, but positively correlated with fruiting. Flowering of M. paucispinus in dry periods of the year avoids that erect flowers positioned in terminal cephalium, exposed in open areas of the vegetation, be damaged for the rains, while fruiting in rainy periods can be favorable to the dispersion and germination of this species.

Influence of soil properties on the abundance of plant species in ferruginous rocky soils vegetation, southeastern Brazil

Ferruginous "campos rupestres" are a particular type of vegetation growing on iron-rich primary soils. We investigated the influence of soil properties on plant species abundance at two sites of ferruginous "campos rupestres" and one site of quartzitic "campo rupestre", all of them in "Quadrilátero Ferrífero", in Minas Gerais State, southeastern Brazil. In each site, 30 quadrats were sampled to assess plant species composition and abundance, and soil samples were taken to perform chemical and physical analyses. The analyzed soils are strongly acidic and presented low fertility and high levels of metallic cations; a principal component analysis of soil data showed a clear segregation among sites due mainly to fertility and heavy metals content, especially Cu, Zn, and Pb. The canonical correspondence analysis indicated a strong correlation between plant species abundance and soil properties, also segregating the sites.

Patterns of litter production in a secondary alluvial Atlantic Rain Forest in southern Brazil

Above-ground litter production is one of the most accessible ways to estimate ecosystem productivity, nutrient fluxes and carbon transfers. Phenological patterns and climatic conditions are still not fully explained well for tropical and subtropical forests under less pronounced dry season and non-seasonal climates, as well as the interaction of these patterns with successional dynamics. Monthly litterfall was estimated for two years in a 9 to 10 year old secondary alluvial Atlantic Rain forest. Total litterfall was higher in the site with more developed vegetation (6.4 ± 1.2 ton ha-1 year-1; 95% confidence interval) as compared to the site with less developed vegetation (3.0 ± 1.0 ton ha-1 year-1). The monthly production of 11 litter fractions (eight fractions comprising the leaf litter of the seven main species of the community and other species; reproductive parts, twigs £ 2 cm diameter, and miscellaneous material) were correlated with meteorological variables making possible to identify three patterns of deposition. The main pattern, dominated by leaf-exchanging species, consisted of a cycle with the highest litterfall at the beginning of the rainy season, preceding by basically three months the peaks of the annual cycles of rainfall and temperatures. Other two patterns, dominated by brevi-deciduous species, peaked at the end of the rainy season and at the end of the non-rainy season. Tropical and subtropical dry forests that present the highest leaf fall gradually earlier than rain forests (as the studied sites) are possibly related to the start of senescence process. It seems that such process is triggered earlier by a more severe hydric stress, besides other factors linked to a minor physiological activity of plants that result in abscission.

Floristic and vegetation structure of a granitic grassland in Southern Brazil

A floristic and structural survey of a natural grassland community was conducted on Morro do Osso, a granitic hill in Porto Alegre, RS, Brazil. Structural data were surveyed in 39 one square meter plots placed over two major grassland areas. An accidental fire has occurred in one of the areas approximately one year prior to our survey, leading to further analysis of parameters differences between sites. The floristic list contains 282 species, whereas the structural survey has found 161 species. Families with highest accumulated importance values were Poaceae, Asteraceae and Fabaceae. The diversity and evenness indexes were 4.51 nats ind-1 and 0.86, respectively. Cluster analysis denoted two groups coinciding with the areas distinguished by the fire disturbance. A similarity analysis between our data and two other data sets from nearby granitic hills resulted in 28% to 35% similarity, with equivalent species-family distribution and many common dominant species, corroborating the concept of a continuous flora along the South Brazilian granitic hills.

Plant-plant associations and population structure of four woody plant species in a patchy coastal vegetation of Southeastern Brazil

We examined plant population structure and interspecific associations for juveniles and adults of four woody species (Andira legalis (Vell.) Toledo, Clusia hilariana Schltdl., Protium icicariba (DC.) Marchand and Vernonia crotonoides Sch. Bip. ex Baker) in a patchy vegetation on a sandy coastal plain (restinga) in SE - Brazil. We found 101 vegetation patches in a 0.5 ha grid and these were divided into two distinct size classes, with large patches (> 20 m²) containing the majority of adult individuals of the species studied. The most abundant species, P. icicariba (465 individuals) and C. hilariana (312), had actively regenerating populations, whereas A. legalis (20) and V. crotonoides (338) showed evidence of intermittent regeneration. The regeneration niches of the four species differed as did their investment in vegetative reproduction: for instance, 81% of C. hilariana seedlings were found growing inside tank-bromeliads contrasting with only 3% of P. icicariba in this habitat. Additionally, 28% of regenerants of C. hilariana originated vegetatively, contrasting with only 6% for P. icicariba. All significant associations between species found in the study were positive. There was a positive association between adults of C. hilariana and P. icicariba, as well as between adults of C. hilariana and juveniles of both. This suggests that P. icicariba is successfully establishing under the canopy of C. hilariana and highlights the role of C. hilariana in generating vegetation cover that will be later dominated by other woody plant species, as an important process for maintenance of plant diversity in this restinga vegetation.

Reproductive biology of Protium spruceanum (Burseraceae), a dominant dioecious tree in vegetation corridors in Southeastern Brazil

We investigated the reproductive biology of Protium spruceanum (Benth.) Engler in vegetation corridors of secondary Atlantic forest in Lavras, southern Minas Gerais State, Brazil. The reproductive phenology was investigated fortnightly over a one year period. Floral biology studies involved pollen viability analysis, nectar production, stigmatic receptivity, pollen tube growth, visiting insect species and visit rates. The small, pale yellowish flowers (0.3-0.4 cm diameter) are functionally unisexual and organized in dense inflorescences (ca. 45 flowers). P. spruceanum presented annual flowering between September and November. Staminate flowers supplied a high percentage of viable pollen (90.6%) and relatively abundant nectar (x = 4.5 μL). Pistillate flowers produced only nectar to flower visitors (x = 4.0 μL). The effective pollinators were Apis mellifera and Trigona sp. (Hymenoptera, Apidae). Pollen tubes of cross-pollinated flowers were observed entering the ovaries 48 h after pollination. The fruiting season is from October to March, with a peak in November, coinciding with the rainfall peak. Ecological implications of these findings, and alternative arguments to explain the high genetic diversity at regional landscape are discussed.

Test of hypotheses about herbivory and chemical defences of Qualea parviflora (Vochysiaceae) in Brazilian Cerrado

(Test of hypotheses about herbivory and chemical defences of Qualea parviflora (Vochysiaceae) in Brazilian Cerrado). Qualea parviflora Mart. (Vochysiaceae), a widely distributed tree found in different habitats in Brazilian Cerrado (savanna), provides resources for a great variety of insects. In this study, we tested two hypotheses about plant investment in anti-herbivore defences along a fertility gradient in Cerrado: the carbon/nitrogen balance (CNBH) and resource availability (RAH). We also investigated how the pattern of herbivory varies through the year and among three types of vegetation in Brazilian Cerrado - campo sujo, cerrado sensu strictu and cerradão. Sampling was conducted in three types of vegetation and in rainy (January and November) and dry months (April and July). Damage on 20 completely expanded leaves, leaf nutrients, sclerophlylly, total phenols and tannins were recorded for each plant (n = 30). When leaves were young, less sclerophyllous, and with higher concentration of nutrients and tannins, damage by herbivores was about 7% in cerrado sensu stricto and 3% in campo sujo. Mature leaves did not show any significant difference on herbivory among habitats, that varied from 6 to 9%. Nutrient availability to plants is an important factor determining production of secondary metabolites in Q. parviflora, corroborating the CNBH. The absence of correlation between damaged leaf area and tannin concentration did not corroborate the RAH, suggesting that tannin production is not strongly influenced by herbivores on Q. parviflora.

Impacts of the Belo Monte hydroelectric dam construction on pioneer vegetation formations along the Xingu River, Pará State, Brazil

There is an important pioneer vegetation formation along the Xingu River in the area where the Belo Monte hydroelectric dam is being constructed that is highly adapted to a seasonally fluctuating water levels. The aim of this study was to examine the habitat and flora of the pioneer formations in the Belo Monte area. The area was divided in three sections for study purposes (Reservoir, Low Flow, and Control) that were expected to experience different degrees of impact from the dam project. The calculations of habitat losses were based on satellite imagery classifications, and a total of 111 plots were established in the three areas for vegetation sampling. Habitat losses of the pioneer formations will total 89.7% when the project is fully functional. Forty-five of the 72 recorded species are restricted to single areas. Species richness and diversity were significantly lower in the control area. The completion of the Belo Monte reservoir will result in habitat reductions and will consequently reduce the richness and diversity of pioneer formations. Studies suggest monitoring the populations located in the reduced flow area to determine possible impacts resulting from changes in the regional hydrological cycle caused by the Xingu River dam.

Influence of edaphic variables on the floristic composition and structure of the tree-shrub vegetation in typical and rocky outcrop cerrado areas in Serra Negra, Goiás State, Brazil

The present study analyzed the influence of edaphic variables on the floristic compositions and structures of the arboreal and shrub vegetation of typical cerrado (TC) and rocky outcrop cerrado (RC) communities in the Serra Negra mountain range in Piranhas Municipality, Goiás State, Brazil. Ten 20×50m plots were established in each community, and all individuals with minimum diameters ³5cm measured at 30cm above soil level were sampled. Composite soil samples were collected at 0-20cm depths in each plot for physical and chemical analyses. The proportions of above-ground rock cover work also estimated in each RC plot. A total of 2,009 individuals (83 species, 69 genera, and 34 families) were recorded. Qualea parviflora was the only species consistently among the 10 most structurally important taxa in both communities, and was considered a generalist species. The observed and estimated species richnesses were greater in RC than in TC, although plant basal areas and heights did not differ between them. There were positive correlations between rock cover×plant density and rock cover×basal areas. TWINSPAN and PCA analysis separated the TC and RC plots, and three RC habitat specialist species (Wunderlichia mirabilis, Norantea guianensis, and Tibouchina papyrus) were identified. Soil variables were found to have greater effects on the species compositions of the TC and RC sites than the geographic distances between sampling plots. According to CCA analysis, the exclusive (or more abundant species) of each community were correlated with soil variables, and these variables therefore determined the selection of some species and influenced the differentiation of the vegetation structures of the communities studied.

The tree-shrub vegetation in rocky outcrop cerrado areas in Goiás State, Brazil

We describe the floristic composition of the tree-shrub vegetation in 10 areas of rocky outcrop cerrado in Goiás State, Brazil. Ten 20×50m plots (totaling 1ha) were established and all of the individuals with diameters at 30cm above soil level (DB30) ³5cm were included in the sampling. Comparative analyses of the flora were realized using similarity indices (Sørensen and Czekanowski), classification analysis (TWINSPAN), and the Mantel test. A total of 13,041 tree-shrub individuals were sampled, distributed among 219 species, 129 genera and 55 families. Fabaceae was the most well-represented family, followed by Myrtaceae, Melastomataceae, Vochysiaceae, Malphigiaceae, and Rubiaceae. Fully 42.3% of the comparisons evaluated by the Sørensen index were >0.50, while all the values were <0.50 for the Czekanowski index, with the exception of Jaraguá and Mara Rosa areas. The TWINSPAN classification generated four divisions and, in general, only the differences in the size of the population were responsible for the groupings. The Mantel test indicated that there was no relationship between floristic similarity and the distances between the areas (r=0.32, P=0.05). It therefore appears that the areas of rocky outcrop cerrado in Goiás State are relatively floristically homogeneous and that they are principally distinguished by the differences in the sizes of the populations of their dominant species, and the presence of exclusive species in certain areas.

NADPH-diaphorase activity in area 17 of the squirrel monkey visual cortex: neuropil pattern, cell morphology and laminar distribution

We studied the distribution of NADPH-diaphorase activity in the visual cortex of normal adult New World monkeys (Saimiri sciureus) using the malic enzyme "indirect" method. NADPH-diaphorase neuropil activity had a heterogeneous distribution. In coronal sections, it had a clear laminar pattern that was coincident with Nissl-stained layers. In tangential sections, we observed blobs in supragranular layers of V1 and stripes throughout the entire V2. We quantified and compared the tangential distribution of NADPH-diaphorase and cytochrome oxidase blobs in adjacent sections of the supragranular layers of V1. Although their spatial distributions were rather similar, the two enzymes did not always overlap. The histochemical reaction also revealed two different types of stained cells: a slightly stained subpopulation and a subgroup of deeply stained neurons resembling a Golgi impregnation. These neurons were sparsely spined non-pyramidal cells. Their dendritic arbors were very well stained but their axons were not always evident. In the gray matter, heavily stained neurons showed different dendritic arbor morphologies. However, most of the strongly reactive cells lay in the subjacent white matter, where they presented a more homogenous morphology. Our results demonstrate that the pattern of NADPH-diaphorase activity is similar to that previously described in Old World monkeys