Browse characteristics and impact of browsing on trees and shrubs in African savannas

Abstract. Studies on browse and browsing impact on woody plants in African savannas show that browse can be a significant part of the total food for large herbivores. Utilization of browse is little studied and patterns are therefore difficult to examine. The quality of browse has been described mainly in terms of nutrients, fibre and digestibility. Studies on chemical and physical defences of plants against herbivory can add considerably to the understanding of the browse-browser system. The various ways woody plants respond to browsing are little known. Interactions between browsers and woody plants are dynamic, since responses of woody plants to browsing result in changes in feeding behaviour of the browsing animal, both in savannas and in temperate ecosystems.     

Predispersal seed predation in native leguminous shrubs and trees in savannas of southern Botswana

Seed predation by insects in the seed-crop of nine Acacia species, Dichrostachys cinerea , and Peltophorum africanum was examined in various tree savannas in Botswana. The degree of infestation varied strongly between and within species from 0% ( P. africanum ) to more than 80% ( A. hebeclada, D. cinerea ). Whereas in all Acacia species only one phytophagous hymenopteran ( Oedaule sp.) was present, the number of bruchid species varied between one ( Bruchidius sahlbergi on A. hebecluda, B. uberatus on A. nilotica ) and eight species ( Bruchidius div. spec., Caryedon sp., Spermophagus rufonotutus on A. tortilis ).
The life history of B. sahlbergi was studied in detail and lasted at least 100 days from egg to adult beetle. The life-span of adult beetles may extend to a further 57 days.

Résumé

Des observations ont été réalisées sur les dégâts d'insects dans les graines de neuf espèces d' Acacia , de Dichrostachys cinerea et de Peltophorum africanum dans les savanes arborées du Botswana. Le taux d'infestation varie fortement aussi bien entre les espèces que parmi les espèces de 0% ( P. africanum) à plus de 80% ( A. hebeclada, C. cinerea ). Alors que toutes les especes d' Acacia n'héberge qu'un seul Hyménoptère phytophage ( Oedaule sp.), le nombre d'espèces de Bruchidae observées varie de une ( Bruchidius sahlbergi sur A. hebeclada, B. uberatus sur A. nilotica) à huit ( Bruchidius div. spec., Caryedon sp., Spermophagus rufonotatus sur A. tortilis ).
Le cycle de développement de B. sahlbergi a étéétudié et dure au moins une certaine de jours. La longévité des adultes de cette espèce peut atteindre 57 jours.     

Microclimatic parameters influencing nitrogen fixation in the phyllosphere in a Costa Rican premontane rain forest

The acetylene reduction method was used to measure nitrogen fixation in the phyllosphere of attached leaves of different phorophytes under natural conditions in a premontane rain forest in Costa Rica. Maximum rates of nitrogen fixation (26 ng N · cm⁻² leaf area · h⁻¹) – mainly due to the activity of two species of Scytonema (Cyanobacteria) – were measured in the rainy season in bright sunlight. Rates of nitrogen fixation were correlated with the leaf area covered by Scytonema. In periods without precipitation the fixation activity decreased to zero within 2–3 days. As long as the epiphylls were sufficiently supplied with water, other microclimatic factors like temperature and light intensity also influenced nitrogen fixation rates, but to a lesser extent. Relative humidity and species of phorophyte showed no direct influence. It was concluded that the most important factor for nitrogen fixation in the phyllosphere was the availability of liquid water. Linking these results to meteorological data, the input of nitrogen by biological nitrogen fixation in the phyllosphere in the investigation area was estimated to be as much as 1.6 ± 0.8 kg N · ha⁻¹ · year⁻¹ per unit of leaf area index (LAI). For an LAI of 2 for the understory the nitrogen input would vary between 2 and 5 kg N · ha⁻¹ · year⁻¹. This work also demonstrates the importance of detailed knowledge of variation in microclimate throughout the year as a basis for extrapolation of the annual nitrogen input. Received: 21 December 1997 / Accepted: 14 June 1998     

Soil acidification and vegetation changes in deciduous forest in southern Sweden

Summary Thirtyfour deciduous forest sites in southern Sweden, originally studied in 1949–1970, were resampled in 1984/85. The average pH change in the humus layer was-0.78 and-0.23 in soils originally studied 30–35 and 15–20 years ago, respectively. Cover changes in the field layer were measured and related to pH changes. The number of species had increased in spite of pH decreases, reaching a maximum at pH 4.0–5.0, while the total cover of the field layer was unchanged.Two groups of species showed no correlation with pH decreases in the humus layer. A large number of species had increased in cover on a majority of sites over the entire species specific pH interval, including nitrophilic species (Rubus idaeus, Chamaenerium angustifolium, Aegopodium podagraria, Stellaria nemorum). A few species had decreased in cover on a majority of sites (Polygonatum multiflorum, Pulmonaria officinalis, Dentaria bulbifera). Some species showed a covariation with pH changes, decreasing in cover in sites in the acidic part of the pH interval (Mercurialis perennis, Lamium galeobdolon, Galium odoratum, Oxalis acetosella, Luzula pilosa). Without excluding other factors, this paper suggests that the increased nitrogen deposition and the increased acidity in the humus layer might cause some of the cover changes in the presented species.     

Long-term effects of metals in sewage sludge on soils,microorganisms and plants

Summary This paper reviews the evidence for impacts of metals on the growth of selected plants and on the effects of metals on soil microbial activity and soil fertility in the long-term. Less is known about adverse long-term effects of metals on soil microorganisms than on crop yields and metal uptake. This is not surprising, since the effects of metals added to soils in sewage sludge are difficult to assess, and few long-term experiments exist. Controlled field experiments with sewage sludges exist in the UK, Sweden, Germany and the USA and the data presented here are from these long-term field experiments only. Microbial activity and populations of cyanobacteria,Rhizobium leguminosarum bv.trifolii, mycorrhizae and the total microbial biomass have been adversely affected by metal concentrations which, in some cases, are below the European Community's maximum allowable concentration limits for metals in sludge-treated soils. For example, N₂-fixation by free living heterotrophic bacteria was found to be inhibited at soil metal concentrations of (mg kg^–1): 127 Zn, 37 Cu, 21 Ni, 3.4 Cd, 52 Cr and 71 Pb. N₂-fixation by free-living cyanobacteria was reduced by 50% at metal concentrations of (mg kg^–1): 114 Zn, 33 Cu, 17 Ni, 2.9 Cd, 80 Cr and 40 Pb.Rhizobium leguminosarum bv.trifolii numbers decreased by several orders of magnitude at soil metal concentrations of (mg kg^–1): 130–200 Zn, 27–48 Cu, 11–15 Ni, and 0.8–1.0 Cd. Soil texture and pH were found to influence the concentrations at which toxicity occurred to both microorganisms and plants. Higher pH, and increased contents of clay and organic carbon reduced metal toxicity considerably. The evidence suggests that adverse effects on soil microbial parameters were generally found at surpringly modest concentrations of metals in soils. It is concluded that prevention of adverse effects on soil microbial processes and ultimately soil fertility, should be a factor which influences soil protection legislation.     

Invasive grass litter facilitates native shrubs through abiotic effects

Questions: Plant invasions are considered one of the top threats to the biodiversity of native taxa, but clearly documenting the causal links between invasions and the decline of native species remains a major challenge of invasion biology. Most studies have focused on impacts of invaders' living biomass, rather than on mechanisms mediated by litter. However, invasive plant litter, which is often of a very different type and quantity than a system's native plant litter, can have multiple important effects on ecosystem processes – such as nitrogen cycling and soil microclimate – that may influence native plants. Location: We studied effects of litter of invasive grass species that are widespread throughout western North America on native shrubs in southern California's semi‐arid habitat of coastal sage scrub. Methods: We combined a 3‐year field manipulation of non‐native litter with structural equation modeling to understand interacting effects on non‐native grasses, native shrubs, soil nitrogen (available and total), and soil moisture. Results: Litter addition facilitated non‐native grass growth, revealing a positive feedback likely to enhance invasion success. Contrary to a major paradigm of invasion biology – that competition with invasive plant species causes declines of native plants – we found that litter also facilitated growth of the native dominant shrub, a result supported by observational trends. Structural equation models indicated that enhanced soil moisture mediated the positive effects of litter on shrub growth. Conclusions: We demonstrate that invasive plants, via their litter, can facilitate dominant native plants by altering soil moisture. Our results highlight that understanding the impacts and mechanisms of plant invasions may be enhanced by considering the role of invasive plant litter on native plants and ecosystem properties.     

Micro-site performance of evergreen and deciduous dwarf shrubs in a subarctic heath in relation to nitrogen status

The relationship between micro-site nitrogen status (total soil N conc., total vegetation N, and leaf N conc.) and performance (biomass) for two evergreen and two deciduous dwarf shrubs was studied in a subarctic heath. One deciduous species, Vaccinium uliginosum , dominated on sites having the highest vegetation nitrogen content and highest leaf nitrogen concentrations. The evergreen Empetrum hermaphroditum dominated on sites with a low total vegetation nitrogen content and low leaf nitrogen concentrations. Two other species, V. myrtillus (deciduous) and V. vitis-idaea (evergreen), showed intermediate patterns that were not clearly separated from either V. uliginosum or Empetrum . Soil nitrogen content showed no significant differences between micro-sites. Possible physiological reasons for the observed patterns are discussed.     

Culm recruitment,dry matter dynamics and carbon flux in recently harvested and mature bamboo savannas in the Indian dry tropics

Culm recruitment, standing crop biomass, net production and carbon flux were estimated in mature (5 years after last harvest) and recently harvested bamboo (Dendrocalamus strictus (Roxb.) Nees) savanna sites in the dry tropics. During the 2 study years bamboo shoot recruitment was 1711–3182 and 1432–1510 shoots ha⁻¹ in harvested and mature sites, respectively. Corresponding shoot mortality was 66–93% and 62–69%, respectively. Total biomass was 34.9 t ha⁻¹ at the harvested site and 47.4 t ha⁻¹ at the mature site. Harvesting increased the relative contribution of belowground bamboo biomass. Annual litter input to soil was 2.7 and 5.9 t ha⁻¹ year⁻¹ at the harvested and mature sites, respectively. The bulk of the annual litterfall (78–88%) occurred in the cool dry season (November to February). The mean litter mass on the savanna floor ranged from 3.1 to 3.3 t ha⁻¹; at the harvested site wood litter contributed 70% of the litter mass and at the mature site leaves formed 77% of the litter mass. The mean total net production (TNP) for the two annual cycles was 15.8 t ha⁻¹ year⁻¹ at the harvested site and 19.3 t ha⁻¹ year⁻¹ at the mature site. Nearly half (46–57%) of the TNP was allocated to the belowground parts. Short lived components (leaves and fine roots) contributed about four-fifths of the net production of bamboo. Total carbon storage in the system was 64.4 t ha⁻¹ at the harvested site and 75.4 t ha⁻¹ at the mature site, of which 23–28% was distributed in vegetation, 2% in litter and 70–75% in soil. Annual net carbon deposition was 6.3 and 8.7 t ha⁻¹ year⁻¹ at harvested and mature sites, respectively.     

Changes in dry matter and nutrient yields during the restoration of species-rich grasslands

Abstract. In two moderately productive grasslands that had been withdrawn from agricultural use, one on humic sandy soil and one on heavy clay, the decrease in dry matter, N-, P- and K-yields were measured for 14 and 11 years respectively, after fertilization had been stopped. The grasslands were mown twice a year. The yields and concentrations of nutrients were compared with those of already unproductive grasslands under the same growing conditions. On sandy soil, the annual dry matter yield decreased from 10.2 to 6.5 ton / ha in four years. After nine years, the dry matter (4.1 ton / ha) and K (20.5 kg / ha) yields were as low as those of the unproductive grassland. At that time yields and concentrations of N and P were still greater. The low K- concentration of 0.50 % and the lack of response upon N fertilizing indicate that the supply of K limits biomass production. On clay, during the first three years, the dry matter yield decreased from 10.2 to 5.0 ton / ha, after 6 yr the production increased again. The low N-concentration of 1.52 % after 10 yr indicates that N is likely to limit biomass production, but the low P-concentration of 0.14% indicates that P shortage can also be expected. In the lower productive grassland on sand the number of species that invaded was higher.     

Nitrogen cycling in tropical and temperate savannas

Savannas are the most common vegetation type in the tropics and subtropics, ranging in physiognomy from grasslands with scattered woody plants to woodlands with heterogeneous grass cover. Productivity and organic matter turnover in savannas are controlled by interactions between water and nutrient availability, and this basic environmental structure is modified by fire frequency and land management practices. We compared temperate and tropical savannas in order to understand the strength of nitrogen (N) limitation of productivity. American tropical and temperate savannas are N limited systems, and the N cycle differs according to the woody plant density, fire frequency, land use change, N deposition and N fixation. Grazing and conversion to pasture have been the predominant land-use changes in most savannas. In the Cerrado and the Llanos tropical savannas, intensified use of fire for pasture management is leading to decreased woody plant density. Oppositely, in the Chaco and North American temperate savannas, fire suppression and grazing are leading to increases in woody density. In addition, the higher soil P availability in the Gran Chaco and the higher N deposition in North American savannas may be contributing to increases of N cycling and net productivity rates. Some aspects of the N budget for savannas of the American continent are still unclear and require further analysis to determine rates of N fixation, and to understand how spatial and temporal soil heterogeneity control N fluxes through soil solution and into streams.     

Gas exchange characteristics and dry matter accumulation of soybean treated with Nod factors

The effect of Nod factors on gas exchange characteristics and growth of soybean plants was studied. Soybean responded positively to some concentrations of Nod factors. Photosynthesis was increased up to 13% over the control, and this was accompanied by increases in stomatal conductance; plant dry weight was also increased. Near-fully expanded leaves responded more strongly to Nod factors than fully expanded leaves. Only Nod factor NodBj-V (C18:1, MeFuc) had significant effects on soybean. The finding suggests that Nod factors can affect photosynthesis, possibly indirectly, by stimulating sink strength.     

Effects of habitat and growing season fires on resprouting of shrubs in longleaf pine savannas

The effects of habitat and timing of growing season fires on resprouting of shrubs were studied in second-growth longleaf pine savannas of the west Gulf coastal plain in the southeastern United States. Within the headwaters of three different drainages of the Calcasieu River in the Kisatchie National Forest in western Louisiana, replicated permanent transects were established that extended from xeric upland longleaf pine savannas into downslope hydric seepage savannas. All shrubs were mapped and tagged, and numbers of stems were counted prior to any fires. Replicated prescribed fires were set early (June) and late (August) during the 1990 growing season; maximum fire temperatures were measured within both upland and seepage habitats within each transect. Shrubs were relocated; stems were recensused two and twelve months after the fires. At least some shrubs of all species resprouted from underground organs; none regenerated solely from seed banks in the soil. There was no reduction in total numbers of stems one year after fires compared to before fires, either in the upland or in seepage savannas. In addition, there was no reduction in total numbers of stems one year after early or late growing season fires. Fire-related mortality was restricted to small shrubs (< 18 stems) and was not associated with high fire temperatures. The rate of resprouting varied among species and between habitats. Resprouting occurred more rapidly in seepage than upland savannas, but more resprouts were produced in upland than seepage savannas one year after fires. In contrast to other upland species, Vaccinium arboreum and V. elliottii delayed resprouting more than two months following fire. Stems of Rhus copallina and Pyrus arbutifolia, species with long rhizomes, increased more after fires in June than fires in August. We suggest that growing season fires may block further recruitment of shrubs into longleaf pine savannas, but reduction in numbers of large shrubs may require additional management.     

Estimates of nitrogen fixation by trees on an aridity gradient in Namibia

Summary Nitrogen (N₂) fixation was estimated along an aridity gradient in Namibia from the natural abundance of ¹⁵N (¹⁵N value) in 11 woody species of the Mimosacease which were compared with the ¹⁵N values in 11 woody non-Mimosaceae. Averaging all species and habitats the calculated contribution of N₂ fixation (N _f ) to leaf nitrogen (N) concentration of Mimosaceae averaged about 30%, with large variation between and within species. While in Acacia albida N _f was only 2%, it was 49% in Acacia hereroensis and Dichrostachys cinerea, and reached 71% in Acacia melifera. In the majority of species N _f was 10–30%. There was a marked variation in background ¹⁵N values along the aridity gradient, with the highest ¹⁵N values in the lowland savanna. The difference between ¹⁵N values of Mimosaceae and non-Mimosaceae, which is assumed to result mainly from N₂ fixation, was also largest in the lowland savanna. Variations in ¹⁵N of Mimosaceae did not affect N concentrations, but higher ¹⁵N-values of Mimosaeae are associated with lower carbon isotope ratios (¹³C value). N₂ fixation was associated with reduced intrinsic water use efficiency. The opposite trends were found in non-Mimosaceae, in which N-concentration increased with ¹⁵N, but ¹³C was unaffected. The large variation among species and sites is discussed.This paper is prepared in memory of J. Visser, who took part in the collection of species, but died in 1990     

我国豆科树种结瘤调查(续)

据国内资料报道,我国豆科树种约94属763种,已调查过结瘤状况的有含羞草亚科的123种(包括近几年来从国外引种的54种)、蝶形花亚科的211种和苏木亚科的70种,其中可以结瘤的有含羞草亚科的116种、蝶形花亚科的207种和苏木亚科的18种。含羞草亚科中的2种、蝶形花亚科中的3种均有结瘤和不结瘤的报道。     

环境胁迫对岷江干旱河谷豆科灌木幼苗根系生长及其结瘤的影响

通过野外调查,分析了岷江干旱河谷海拔梯度上相同年龄段的白刺花、岷谷木蓝和小雀花幼苗根系结瘤、生物量积累、生物量百分比的变化及其相互关系;人为控制土壤水分（田间持水量(WHC)的80%、60%、40%和20% 4个水平）研究了干旱对2月龄的白刺花与小雀花幼苗结瘤能力的影响.结果表明：1)干旱河谷低海拔地段(1600～1950 m)幼苗的结瘤能力都很低,白刺花、岷谷木蓝和小雀花的平均结瘤数分别为每株(0.1±0.1)、(0.9±0.5)和(5.7±1.9)个,未结瘤的个体分别占651%、123%和176%;总体上随着海拔的升高,3种幼苗的根瘤数均呈增加趋势,且与细根百分比及根长正相关.2)土壤水分低于60% WHC时,白刺花与小雀花的根瘤数及生物量均明显减小;在20% WHC条件下,白刺花根瘤数为0,小雀花根瘤数为每株(9.8±3.6)个.综合分析发现,干旱河谷地区豆科灌木的结瘤固氮能力很低,土壤水分在40% WHC以上时其生物固氮才能有效发挥,与白刺花相比,小雀花结瘤能力较高,干旱忍受性较强.     

我国豆科树种结瘤调查

据国内资料报道,我国豆科树种约94属763种,已调查过结瘤状况的有含羞草亚科的123种(包括近几年来从国外引种的54种)、蝶形花亚科的211种和苏木亚科的70种,其中可以结瘤的有含羞草亚科的116种、蝶形花亚科的207种和苏木亚科的18种。含羞草亚科中的2种、蝶形花亚科中的3种均有结瘤和不结瘤的报道。     

Nitrogen and phosphorus resorption in trees of a Mexican tropical dry forest

Resorption efficiency (RE) and proficiency, foliar nutrient concentrations, and relative soil nutrient availability were determined during 3 consecutive years in tree species growing under contrasting topographic positions (i.e., top vs. bottom and north vs. south aspect) in a tropical dry forest in Mexico. The sites differed in soil nutrient levels, soil water content, and potential radiation interception. Leaf mass per area (g m^–2) increased during the growing season in all species. Soil P availability and mean foliar P concentrations were generally higher at the bottom than at the top site during the 3 years of the study. Leaf N concentrations ranged from 45.4 to 31.4 mg g^–1. Leaf P varied from 2.3 to 1.8 mg g^–1. Mean N and P RE varied among species, occasionally between top and bottom sites, and were higher in the dry than in the wet years of study. Senesced-leaf nutrient concentrations (i.e., a measure of resorption proficiency) varied from 13.7 to 31.2 mg g^–1 (N) and 0.4 to 3.3 mg g^–1 (P) among the different species and were generally indicative of incomplete nutrient resorption. Phosphorus concentrations in senesced leaves were higher at the bottom than at the top site and decreased from the wettest to the the driest year. Soil N and P availability were significantly different in the north- and south-facing slopes, but neither nutrient concentrations of mature and senesced leaves nor RE differed between aspects. Our results suggest that water more than soil nutrient availability controls RE in the Chamela dry forest, while resorption proficiency may be interactively controlled by both nutrient and water availability.     

Relative abundance of denitrifying and dinitrogen-fixing bacteria in layers of a forest soil

The populations of N(2)-fixing and denitrifying bacteria in an acid forest soil near Cologne were characterized by gene probing. The DNA isolated from the soil for this purpose was suitable for DNA-DNA hybridization using 0.4-0.7-kb probes targeting denitrification enzymes, dinitrogenase reductase (nifH) and eubacterial 16S rRNA. The densitometrical comparison of band intensities obtained in these Southern hybridizations indicated that the highest number of total bacteria, of denitrifying and N(2)-fixing microorganisms always occurred in the upper ( approximately 5 cm) soil layer. The concentration of all these organisms decreased in parallel with the soil depth. The soil investigated was rich in nitrate in all layers, and the availability of nitrate apparently did not govern the distribution of denitrifying and N(2)-fixing bacteria in this soil. Soil cores investigated in the laboratory formed N(2)O on addition of nitrate irrespective of the presence of C(2)H(2). Hybridization intensities, with a gene probe for the 16S rRNA, and MPN numbers were generally higher in soil samples taken from the roots of plants than in the bulk soil. There was no selective enrichment of denitrifying or N(2)-fixing bacteria at the roots. The data obtained by hybridizing isolated soil DNA generally matched previous results obtained with culturable bacteria.