Effects of large herbivore browsing on the functional groups of woody plants in a southern African savanna

This study examined the effect of different large herbivore species and stocking rates in savanna ecosystems of Zimbabwe on the richness and abundances of woody plant functional groups and woody plant functional attributes. Seven fence-lines with different herbivore species and stocking rates on either side of the fence were sampled. Plots were placed on both sides of each fence at each of 18 randomly selected positions. The size and species of each woody plant was recorded for each plot. It was found that the number of species with different functional attributes of spinescence, leaf longevity, fruit type and dispersal mechanism and in the functional groups of palatability were not different on the different sides of the fence. However, there were differences in plant abundances for 26 out of the 35 tests carried out on plant abundances with different functional attributes and functional groups. It was hypothesised that the time needed to change woody plant species richness is hundreds of years in these systems, whereas the time needed to change woody plant abundances is decades.     

Plant community response to loss of large herbivores: comparing consequences in a South African and a North American grassland

Loss of biodiversity poses one of the greatest threats to natural ecosystems throughout the world. However, a comprehensive understanding of the impacts of species losses from upper trophic levels is still emerging. Here we compare the impacts of large mammalian herbivore species loss on grassland plant community structure and composition in a South African and North American grassland. Herbaceous plant communities were surveyed at sites without large mammalian herbivores present and at sites with a single species of herbivore present in both locations, and additionally at one site in South Africa with multiple herbivore species. At both the North American and South African locations, plant communities on sites with a single herbivore species were more diverse and species rich than on sites with no herbivores. At the multi-herbivore site in South Africa, plant diversity and richness were comparable to that of the single herbivore site early in the growing season and to the no herbivore site late in the growing season. Analyses of plant community composition, however, indicated strong differences between the multi-herbivore site and the single and no herbivore sites, which were more similar to each other. In moderate to high-productivity ecosystems with one or a few species of large herbivores, loss of herbivores can cause a significant decrease in plant diversity and richness, and can have pronounced impacts on grassland plant community composition. In ecosystems with higher herbivore richness, species loss may also significantly alter plant community structure and composition, although standard metrics of community structure may obscure these differences.     

Reproductive biology in a tropical shrubland of Venezuelan Guayana

Plant sexual systems were evaluated in a tropical shrubland community of Venezuelan Guayana. Of 89 plant species, 73.0% were hermaphrodite, 12.4% monoecious, 10.1 % submonoecious, and 4.5 % dioecious. Temporal variation of sexual expression showed that hermaphrodite species were mostly adichogamous and monoecious, submonoecious species dichogamous (protandrous andprotogynous). Herkogamy was an important mechanism promoting cross-pollination. Of 38 hermaphrodite species, 39.5 % were herkogamous. Plant sexual systems were not associated with life form. Most species were pollinated mostly by large and medium-size bees. Pollinator specificity and pollen load transportation, were higher for monoecious species than for hermaphrodite, submonoecious and dioecious ones. Both pollinator specificity and pollen transportation were not significantly different among representatives of the latter three sexual systems. Plant sexual systems were significantly associated with dispersal syndromes; dioecious species had bird-dispersal syndromes, the other sexual systems mostly anemochory, epizoochory and endozoochory; but many hermaphrodites were anemochorous.     

Habitat dependent pollination success in a Cape orchid

Summary We examined the pollination success of Disa uniflora (Orchidaceae) in two habitats. Plants occurring in a rocky gorge were far more successful than plants occurring in an adjacent open valley. More than 55% of flowers in the gorge were pollinated and set fruit compared with less than 25% of flowers in the valley. These differences are explained by the preference of Meneris tulbaghia (Satyridae), the exclusive pollinator of the orchid, for rocky, sheltered habitats. Fruit set of hand-pollinated flowers did not differ significantly between the two habitats, indicating that resources did not account for the variation in fruiting success.     

Diversity patterns in savanna grassland communities: implications for conservation strategies in a biodiversity hotspot

For any conservation strategy to be effective, it must be preceded by knowledge of how diversity is configured within the area of interest. Here, data from 40 savanna-grassland sites were used to examine how plant biodiversity and species composition varied across spatial scales at the Kalakad-Mundanthurai Tiger Reserve (KMTR), south India. Grasslands surveyed contained 278 plant species, and were characterized by high spatial variability in species association patterns. Fourteen distinct community assemblages were identified, organized primarily along an elevation gradient in the reserve. Overall, grasslands at KMTR were characterized by the dominance of a few, widespread species. The bulk of species richness, however, resulted from subordinate species with fairly restricted distributions. At low elevations, grasslands had high species richness and species composition differed greatly between sites. Mid-elevation grasslands contained about half the number of species present at low elevations, but sites were more similar in species composition. Richness of high-elevation grasslands was a third of that found at low elevations, but different sites harbored unique sets of species. Herbivore use of grasslands varied between communities and showed patterns that coincided with elevation. Herbivore use of low-elevation communities was high albeit variable, of mid-elevation sites was consistently low, and increased at higher elevations. Tall grass communities were the least utilized by herbivores at all elevations. Most species surveyed were rare and restricted in their distribution suggesting that conservation efforts must, perforce, encompass the entire reserve. However, differences in the structuring of diversity across elevations, and in herbivore use of grasslands, suggest that conservation efforts can be partitioned differentially across locations, specifically targeting low and high elevation grasslands in the reserve.     

Elevated CO2 enhances resprouting of a tropical savanna tree

The savannas (cerrado) of south-central Brazil are currently subjected to frequent anthropogenic burning, causing widespread reduction in tree density. Increasing concentrations of atmospheric CO₂ could reduce the impact of such frequent burning by increasing the availability of nonstructural carbohydrate, which is necessary for resprouting. We tested the hypotheses that elevated CO₂ stimulates resprouting and accelerates replenishment of carbohydrate reserves. Using a factorial experiment, seedlings of a common Brazilian savanna tree, Keilmeyera coriacea, were grown at 350 ppm and 700 ppm CO₂ and at two nutrient levels. To simulate burning, the plants were either clipped at 15 weeks or were left unclipped. Among unclipped plants, CO₂ and nutrients both stimulated growth, with no significant interaction between nutrient and CO₂ effects. Among clipped plants, both CO₂ and nutrients stimulated resprouting. However, there was a strong interaction between CO₂ and nutrient effects, with CO₂ having a significant effect only in the presence of high nutrient availability. Under elevated CO₂, carbohydrate reserves remained at higher levels following clipping. Root total nonstructural carbohydrate remained above 36% in all treatments, so carbohydrate reserves did not limit regrowth. These results indicate that under elevated CO₂ this species may be better able to endure the high frequency of anthropogenic burning in the Brazilian savannas. Received: 19 May 1999 / Accepted: 8 November 1999     

Community dynamics of carrion-attendant arthropods in tropical african woodland

Summary Carcasses are temporary resources which are unbredictable and inconsistent in their availability and locality. A recognisable community of interacting user arthrocods comprising sarcophages, coprophages, dermatophages, keratophages, detritivores, predators and parasites has evolved to exploit the carcass habitat. The large number of arthropods, close confinement, and limited duration of resources necessitates aggressive utilisation. The trophic relations, competition and successionary pattern of these arthropods is discussed. Several pathways to reduce competitive conflict are described. Succession at carcasses is viewed as being inherently different from the traditional concept as the habitat is non-replenishing and does not lead to a climax community.     

Bat diversity of a Cerrado habitat in central Brazil

The Cerrado is considered one of the 25 biodiversity hotspots in the world by conservation organizations. There are few studies on bat taxonomic groups for this biome. Herein we present a bat survey employing mist nets in the protected area of a private natural heritage reserve Reserva Particular do Patrimônio Natural Pousada das Araras, located in the west-central Brazil. We investigate the hypothesis that the Cerrado habitat complexity plays a role on the different structuring forces in bat ecological communities. Bats represent a diversity of trophic levels, and they occupy a wide range of available Cerrado habitats and microhabitats. The patterns and processes we discuss represent the factors influencing coexisting species of bats in different habitats and their implications for conservation. We captured 758 individuals of 25 species belonging to four families. Phyllostomidae was the dominant family, represented by 20 species (80%). The average recapture rate was 6.2%, and the species with most proportional recapture was A. caudifer. Greater species richness was observed among bats with predominantly insectivore habits, followed by frugivores. Glossophaga soricina was a dominant species, with about 30% of the captures. There was seasonal variation relating to the number of bats captured, with greater bat frequency occurring during the wet season, although some species occurred at higher rates during the dry season, such as C. perspicillata and S. lilium. Sanguivore bats were abundant, reflecting the availability of shelters and food supply (livestock) in the surrounding area. Compared to other surveyed areas, Pousada das Araras may be considered of high species diversity, supplanting the majority of known Cerrado studied areas. Apparently Cerrado favours the occurrence of insectivore species, with emphasis on the foliage-gleaning insectivores belong to the subfamily Phyllostominae. This study indicates that apparently the conservation of the Cerrado savanna fragments can support a considerable diversity of bat species.     

New palaeoecological evidence for the potential role of fire in the Gran Sabana, Venezuelan Guayana, and implications for early human occupation

The neotropical Gran Sabana region of Venezuela is dominated by apparently anomalous vegetation types, treeless savannas and savanna-forest mosaics, considering the present-day warm and wet bioclimatic conditions. Past climatic changes and fire have been proposed as the more probable causes. Recent palynological studies show that savanna vegetation has been present since the beginning of the Holocene, but the earliest fires recorded so far only go back to 3,800 cal years b.p. This paper uses pollen and charcoal analyses to show the existence of early Holocene regional fires in the Gran Sabana, and to show the intimate connection between the proxies for fire (charcoal) and savanna vegetation (pollen) throughout the Holocene. Although the cause of such fires is not yet known, the possibility of early Holocene human occupation of the Gran Sabana is suggested.     

Pollination and breeding systems of woody plant species in tropical dry evergreen forests,southern India

Tropical dry evergreen forests (TDEF) are a unique forest type found along the east coast of India. They mostly occur as small, isolated fragments of varying sizes (0.5 to ≈10 ha) and are considered as endangered forests types in peninsular India. Although plant diversity is well documented in these forests, there is a paucity of ecological studies vital for conservation and for planning restoration activities. We studied reproductive biology of 13 woody species: four trees, six shrubs, and three lianas in fragments of TDEF in southern India. The phenology of reproduction, floral biology, anthesis and sexual system of each species were recorded. The pollination mode was assessed through observations of the visitation frequency of pollinators and from the floral characters. The breeding system was determined by hand-augmented self- and cross-pollination experiments. The plants flowered during the dry season from January to July. Plants of nine species had both flowers and fruits at the same time. Twelve species were hermaphrodites and one was polygamo-dioecious. Flowers of 11 species opened at dawn and two at dusk. Four species were self-incompatible and six were self-compatible. Natural fruit set ranged from 10% to 56%, self-incompatible species having low fruit set. Cross-pollen augmentation increased fruit set, suggesting presence of outcrossing in all species. The majority of plants species (85%) had a generalized pollination system, receiving visits from diverse insects, such as social bees, solitary bees, wasps, moths and flies. However, only few of them were functionally important for the species. Two species namely: Capparis brevispina and C. zeylanica had butterflies and birds, respectively, as their main pollinators. Our data reveal that there is a predominance of outcrossing in plant species and a generalized pollination system in these forests. We suggest that restoration of TDEFs is crucial as habitats, not only for wild plants but also for pollinating insects.     

Variation in pollinator assemblages in a fragmented landscape and its effects on reproductive stages of a self-incompatible treelet, <Emphasis Type="Italic">Psychotria suterella</Emphasis> (Rubiaceae)

Few studies of plant–pollinator interactions in fragmented landscapes evaluate the consequences of floral visitor variation on multiple stages of plant reproduction. Given that fragmentation potentially has positive or negative effects on different organisms, and that self-incompatible plant species depend on pollinators for sexual reproduction, differences in floral visitor assemblages may affect certain plant reproductive stages. We evaluated how pollinator assemblage, availability of floral resources, pollination, reproductive output, and seed and seedling performance of Psychotria suterella Muell. Arg. varied among three fragmentation categories: non-fragmented habitats, fragments connected by corridors, and isolated fragments. Richness and frequency of floral visitors were greater in fragments than in non-fragmented sites, resulting mainly from the addition of species typically found in disturbed areas. Although 24 species visited Psychotria suterella flowers, bumblebees were considered the most important pollinators, because they showed the highest frequency of visits and were present in eight out of ten sites. Additionally, the number of pollen tubes per flower per visit was lower in areas without bumblebees. The increased visitation in fragments seemed to enhance pollination slightly. However, fruit and seed output, germination, and seed and seedling mass were similar in non-fragmented sites, connected sites, and isolated fragments. Our results suggested that, even for a self-incompatible species, responses to habitat fragmentation at different stages of plant reproduction might be decoupled from the responses observed in floral visitors, if fruit set is not pollen limited. If all reproductive stages were considered, variation on the small scale was more important than the variation explained by fragmentation category. In spite of its self-incompatible breeding system, this plant–pollinator system showed resilience to habitat fragmentation, mainly as a result of high availability of potential mates to P. suterella individuals, absence of pollen limitation, and the presence of bumblebees (Bombus spp.) throughout this highly connected landscape.     

Dynamics of viable nitrifier community, N-mineralization and nitrification in seasonally dry tropical forests and savanna

The study was conducted in Vindhyan region, to assess the N-mineralization, nitrification and size of viable community of ammonium- and nitrite-oxidizing bacteria as affected by different sites and seasons. Six different ecosystems (four forests and two savannas), which differ in terms of topography, vegetation and moisture status, were selected for the present study. The soils of the study sites differ significantly in its physico-chemical properties. The savanna site had significantly higher pH (7.2), bulk density (1.37 g cm(-3)) and silt content (67.80%) but lower water holding capacity (1.37%), total-C (16,356 microg g(-1) dry soil), N (1090 microg g(-1) dry soil) and P (213 microg g(-1) dry soil) than forest sites. The soil moisture content, N-mineralization, nitrification rates and numbers of ammonium- and nitrite-oxidizing bacteria were highest in the wet season and lowest in dry season, while the size of mineral-N (NH4(+)-N and NO3(-)-N) showed a reverse trend at the sites. The N-mineralization, nitrification and nitrifier population size differ significantly across the site and season. The numbers of free-living cells of ammonium- and nitrite-oxidizing bacteria were significantly related to each other and to N-mineralization, nitrification, soil moisture and mineral-N components. The N-mineralization, nitrification and the viable number of nitrifying cells were consistently higher for forest soils compared to savanna sites. It was concluded that soil microbial process (N-mineralization and nitrification) and nitrifier population size were dependent on site topography, vegetation cover and soil moisture status.     

Composition of the pollinator community,pollination and the mating system for a shrub in fragments of species rich kwongan in south-west Western Australia

In this study we investigate the composition of the potential honeyeater pollinator community, patterns of honeyeater visitation, pollination and the mating system in a range of population fragments for the bird-pollinated mixed mating system shrub Calothamnus quadrifidus R.Br. Specifically, we aimed to answer the following questions. For smaller and more isolated population fragments are honeyeater species lost from the pollinator community, patterns of visitation different, levels of pollination lower and rates of selfing, biparental inbreeding and correlated paternity higher. The composition of the honeyeater community was similar across population fragments and there was no relationship between the abundance of birds and population fragment size. Honeyeaters were most commonly observed visiting numerous inflorescences within single plants in all populations, but as population fragments became larger movements between plants were more commonly observed. Our observations of honeyeater visitation were generally consistent with our measurements of pollination and patterns in the mating system across population fragments. We found no significant relationship between population fragment size and levels of pollination. Mating system studies showed outcrossing rates (t _m) comparable to those found in other bird-pollinated Myrtaceae, and ranged from 0.54 to 0.90 across populations. Outcrossing rates were not significantly correlated with log population size, but correlations of outcrossed paternity indicate a clear trend from low correlated paternity in larger populations to significantly higher correlated paternities in smaller populations. As a consequence mating in small populations will occur between much smaller groups of plants, and this may affect population fitness in subsequent generations.     

Liana habitat associations and community structure in a Borneanlowland tropical forest

Lianas (woody vines) contribute substantially to the diversity and structure of most tropical forests, yet little is known about the importance of habitat specialization in maintaining tropical liana diversity and the causes of variation among forests in liana abundance and species composition. We examined habitat associations, species diversity, species composition, and community structure of lianas at Sepilok Forest Reserve, Sabah, Malaysia in northeastern Borneo among three soil types that give rise to three distinct forest types of lowland tropical rain forest: alluvial, sandstone hill, and kerangas (heath) forest. Alluvial soils are more nutrient rich and have higher soil moisture than sandstone soils, whereas kerangas soils are the most nutrient poor and drought prone. Lianas ≥0.5-cm in diameter were measured, tagged, and identified to species in three square 0.25-ha plots in each forest type. The number of lianas ≥0.5 cm did not differ significantly among forest types and averaged 1348 lianas ha⁻¹, but mean liana stem diameter, basal area, estimated biomass, species richness, and Fisher’s diversity index were all greater for plots in alluvial than sandstone or kerangas forests. Liana species composition also differed greatly among the three habitats, with 71% of species showing significant positive or negative habitat associations. Sandstone forests were intermediate to alluvial and kerangas forests in most aspects of liana community structure and composition, and fewer species showed significant habitat associations with this forest type. Ranking of forest types with respect to liana density, biomass, and diversity matches the ranking in soil fertility and water availability (alluvial > sandstone hill > kerangas). These results suggest that edaphic factors play an important role in maintaining liana species diversity and structuring liana communities.     

Air pollen dispersal in a tropical area

Volumetric data on airborne pollen have been gathered for two consecutive years at a neotropical location (Caracas). Among the 65 taxa which were identified, pollen from aCupressus species (introduced) and from aCecropia species (indigenous) were dominant. Less numerous but also abundant (daily averages ≥5 grains/m³ air) were pollen from Gramineae, Urticaceae,Alcalypha, Pinus, Piperaceae andMimosa. Pollen grains were recorded daily throughout the year. They increased in numbers during April–May and again during November–December. The first peak was contributed mainly by indigenous species, the second peak mainly by introduced species.     

Fire responses of three co-occurring Asteraceae shrubs in a temperate savanna in South America

South American temperate savannas have undergone significant woody plant encroachment through changes in their disturbance regimes. We studied fire strategies and the spatial variation in individual fire responses of Eupatorium buniifolium, Baccharis medullosa and B. dracunculifolia, the dominant shrub species at El Palmar National Park, in areas with different fire history. In recently burnt sites, all Baccharis dracunculifolia individuals died, whereas all E. buniifolium and B. medullosa individuals sprouted. The relative growth rate was higher in E. buniifolium, which invested more biomass in shoots, and in B. medullosa, which invested more biomass in leaves. At the unburnt site, B. dracunculifolia showed the highest growth rate and leaf biomass. B. medullosa produced capitula immediately after fire, whereas E. buniifolium reproduction was delayed for 1 year. In both species, production of capitula and reproductive effort were higher in recently burnt sites. For B. dracunculifolia, production of capitula was lower and the reproductive effort was higher in burnt than in unburnt sites. Seedling establishment was extremely low for E. buniifolium, low for B. medullosa and high for B. dracunculifolia in both recently burnt sites. E. buniifolium behaved as a sprouter, B. medullosa as facultative sprouter and B. dracunculifolia as obligate seeder species. In order to control the increase and/or attain desirable population levels of these species, fire management practices in these savannas should consider individual species’ responses, particularly to the time since the last fire.     

Thornbush invasion in a savanna ecosystem in eastern Botswana

The paper describes the thornbush invasion (bush encroachment) found in the savanna at Olifants Drift in eastern Botswana. Overgrazing by cattle is responsible for the opening up of the grass sward and has enabled woody species to establish. A quantification of this process between 1950 and 1975 in an area of ca. 100 km² is given in terms of plant density as well as aboveground biomass. The loss of herbaceous biomass which generally accompanies bush encroachment and the possibilities for pasture restoration are mentioned.Species nomenclature follows the Flora of Southern Africa and the system in use at the herbarium of the Botanical Research Institute at Pretoria, South Africa.     

Possibilities for dispersal in annual and perennial grasses in a savanna in Botswana

Dispersal of 11 dominant grass species in the savanna of southern Botswana was investigated. The dispersal is autochorous, anemochorous and epizoochorous independent of the life-cycle (annuals, perennials). Dispersal distances were estimated experimentally. Anemochorous species with a plume-like spikelet such as Chloris virgata and Enneapogon cenchroides have a low rate of descent (<1 m s^-1) and a low Reynold number (100–110). Nevertheless they can only be transported up to 13 m from the originating infructescence at a wind velocity of 10 m s^-1. Therefore, the majority of the disseminules remained near the parent plant. By analysing the seed pool under the canopy of trees of Dichrostachys cinerea epizoochorous species such as Tragus berteronianus were dominant at the cattle resting sites under trees. The results are discussed in relation to the three seed dispersal hypotheses and the model of selective interaction of dispersal, dormancy, and seed site as adaptions to variable environments.