Beyond the tropics: forest structure in a temperate forest mapped plot

Question: How do the diversity, size structure, and spatial pattern of woody species in a temperate (Mediterranean climate) forest compare to temperate and tropical forests? Location: Mixed evergreen coastal forest in the Santa Cruz Mountains, California, USA. Methods: We mapped, tagged, identified, and measured all woody stems (≥1 cm diameter) in a 6‐ha forest plot, following Center for Tropical Forest Science protocols. We compared patterns to those found in 14 tropical and 12 temperate forest plots. Results: The forest is dominated by Douglas‐fir (Pseudotsuga menziesii) and three species of Fagaceae (Quercus agrifolia, Q. parvula var. shrevei, and Lithocarpus densiflorus), and includes 31 woody species and 8180 individuals. Much of the diversity was in small‐diameter shrubs, treelets, and vines that have not been included in most other temperate forest plots because stems <5‐cm diameter had been excluded from study. The density of woody stems (1363 stems ha^?1) was lower than that in all but one tropical plot. The density of large trees (diameter ≥30 cm) and basal area were higher than in any tropical plot. Stem density and basal area were similar to most other temperate plots, but were less than in low‐diversity conifer forests. Rare species were strongly aggregated, with the degree of aggregation decreasing with abundance so that the most common species were significantly more regular than random. Conclusions: The patterns raise questions about differences in structure and dynamics between tropical and temperate forests; these need to be confirmed with additional temperate zone mapped plots that include small‐diameter individuals.     

Implications of floristic and environmental variation for carbon cycle dynamics in boreal forest ecosystems of central Canada

Abstract. Species composition, detritus, and soil data from 97 boreal forest stands along a transect in central Canada were analysed using Correspondence Analysis to determine the dominant environmental/site variables that differentiate these forest stands. Picea mariana stands were densely clustered together on the understorey DCA plot, suggesting a consistent understorey species composition (feather mosses and Ericaceae), whereas Populus tremuloides stands had the most diverse understorey species composition (ca. 30 species, mostly shrubs and herbs). Pinus banksiana stands had several characteristic species of reindeer lichens (Cladina spp.), but saplings and Pinus seedlings were rare. Although climatic variables showed large variation along the transect, the CCA results indicated that site conditions are more important in determining species composition and differentiating the stand types. Forest floor characteristics (litter and humus layer, woody debris, and drainage) appear to be among the most important site variables. Stands of Picea had significantly higher average carbon (C) densities in the combined litter and humus layer (43530 kg‐C.ha^‐1) than either Populus (25 500 kg‐C.ha^‐1) or Pinus (19 400 kg‐C.ha^‐1). The thick surface organic layer in lowland Picea stands plays an important role in regulating soil temperature and moisture, and organic‐matter decomposition, which in turn affect the ecosystem C‐dynamics. During forest succession after a stand‐replacing disturbance (e.g. fires), tree biomass and surface organic layer thickness increase in all stand types as forests recover; however, woody biomass detritus first decreases and then increases after ca. 80 yr. Soil C densities show slight decrease with ages in Populus stands, but increase in other stand types. These results indicate the complex C‐transfer processes among different components (tree biomass, detritus, forest floor, and soil) of boreal ecosystems at various stages of succession.     

Home Range Size and Micro‐habitat Density Requirements of Egernia napoleonis: Implications for Restored Jarrah Forest of South Western Australia

Compared to natural forests, coarse woody debris (CWD) is typically scarce in restored forests due to the long time it takes to develop naturally. In post‐mining restored forests in the Jarrah forest of south western Australia, CWD is returned at densities of one log pile per hectare. We tested the adequacy of these densities for meeting the micro‐habitat requirements of Napoleon's skink (Egernia napoleonis), a species rarely found within restored sites. Home range size and overlap, and micro‐habitat densities used by skinks, were measured by radio‐tracking 12 individuals in natural, unmined forest. Napoleon's skinks had small home ranges (0.08 ± 0.02 ha), based on 8 individuals with sufficient fixes. All skinks overlapped in home ranges, with average overlaps of 43.5 ± 8.6%. Ten of the 12 skinks shared micro‐habitats and 4 shared them simultaneously, which indicates some social tolerance. This will influence as to how many micro‐habitats are required. Micro‐habitats were used at high densities: logs at 49.2 ± 8.8 ha^?1 and woody debris piles at 12.4 ± 4.8 ha^?1. Based on these densities, it is recommended that CWD is returned to restored forests at densities of 60 ha^?1, which should provide sufficient micro‐habitats for multiple skinks. Due to the infeasibility of returning such CWD densities across large areas of restored forest, CWD could be preferentially returned as patches, large enough for numerous home ranges, adjacent to unmined forest, or as corridors between unmined forest. These recommendations for returning micro‐habitats should be tested for effectiveness in encouraging recolonization of restored forest by Napoleon's skink and other species.     

Influence of soils and topography on Amazonian tree diversity: a landscape‐scale study

Question: How do soils and topography influence Amazonian tree diversity, a region with generally nutrient‐starved soils but some of the biologically richest tree communities on Earth? Location: Central Amazonia, near Manaus, Brazil. Methods: We evaluated the influence of 14 soil and topographic features on species diversity of rain forest trees (≥10 cm diameter at breast height), using data from 63 1‐ha plots scattered over an area of ～400 km². Results: An ordination analysis identified three major edaphic gradients: (1) flatter areas had generally higher nutrient soils (higher clay content, carbon, nitrogen, phosphorus, pH and exchangeable bases, and lower aluminium saturation) than did slopes and gullies; (2) sandier soils had lower water storage (plant available water capacity), phosphorus and nitrogen; and (3) soil pH varied among sites. Gradient 2 was the strongest predictor of tree diversity (species richness and Fisher's α values), with diversity increasing with higher soil fertility and water availability. Gradient 2 was also the best predictor of the number of rare (singleton) species, which accounted on average for over half (56%) of all species in each plot. Conclusions: Although our plots invariably supported diverse tree communities (≥225 species ha^?1), the most species‐rich sites (up to 310 species ha^?1) were least constrained by soil water and phosphorus availability. Intriguingly, the numbers of rare and common species were not significantly correlated in our plots, and they responded differently to major soil and topographic gradients. For unknown reasons rare species were significantly more frequent in plots with many large trees.     

Microhabitat Preference of Egernia napoleonis in Undisturbed Jarrah Forest,and Availability and Introduction of Microhabitats to Encourage Colonization of Restored Forest

An animal's microhabitat requirements can impact its ability to colonize restored areas, particularly species requiring slow developing microhabitats, such as logs and woody debris piles. Introduction of these microhabitats may be required to facilitate colonization by some species. Restored bauxite mine‐pits in the Jarrah (Eucalyptus marginata) forest of south‐western Australia contain introduced log piles at densities of 1 ha^?1. However, these have not facilitated colonization by Napoleon's skink (Egernia napoleonis), which rely on logs for habitat and are largely absent from restored sites. We radio‐tracked 12 skinks in unmined forest to determine their microhabitat preferences and examined differences in vegetation structure, and microhabitat and food availability, between restored and unmined forests to identify reasons for their absence. Restored and unmined forests differed in canopy, mid‐ and understory cover and ground substrates, which were all potential barriers to colonization. Food availability was similar between restored and unmined forest, thus not a barrier to colonization. Skinks primarily utilized long logs, large woody debris piles, and large trees; microhabitats that were scarce or absent in restored sites and, therefore, potential barriers to colonization. Using this information, we introduced small woody debris piles into restored sites in close proximity to unmined areas containing skinks to facilitate skink colonization. This showed early signs of success and suggested that the lack of logs and woody debris were barriers to colonization. However, further monitoring is required to accurately determine the long‐term value of woody debris piles in facilitating skink colonization.     

Seedling regeneration,environment and management in a semi‐deciduous African tropical rain forest

Questions: How is seedling regeneration of woody species of semi‐deciduous rain forests affected by (a) historical management for combinations of logging, arboricide treatment or no treatment, (b) forest community type and (c) environmental gradients of topography, light and soil nutrients? Location: Budongo Forest Reserve, Uganda. Methods: Seedling regeneration patterns of trees and shrubs in relation to environmental factors and historical management types were studied using 32 0.5‐ha plots laid out in transects along a topographic gradient. We compared seedling species diversity, composition and distribution patterns along topographic gradients and within types of historical management regimes and forest communities to test whether environmental factors contributed to differences in species composition of seedlings. Results: A total of 85 624 woody seedlings representing 237 species and 46 families were recorded in this rain forest. Cynometra alexandri C.H. Wright and Lasiodiscus mildbraedii Engl. had high seedling densities and were widely distributed throughout the plots. The most species‐rich families were Euphorbiaceae, Fabaceae, Rubiaceae, Meliaceae, Moraceae and Rutaceae. Only total seedling density was significantly different between sites with different historical management, with densities highest in logged, intermediate in logged/arboricided and lowest in the nature reserve. Forest communities differed significantly in terms of seedling diversity and density. Seedling composition differed significantly between transects and forest communities, but not between topographic positions or historical management types. Both Chao‐Jaccard and Chao‐Sørensen abundance‐based similarity estimators were relatively high in the plot, forest community and in terms of historical management levels, corroborating the lack of significant differences in species richness within these groups. The measured environmental variables explained 59.4% of variance in seedling species distributions, with the three most important being soil organic matter, total soil titanium and leaf area index (LAI). Total seedling density was positively correlated with LAI. Differences in diversity of >2.0 cm dbh plants (juveniles and adults) also explained variations in seedling species diversity. Conclusions: The seedling bank is the major route for regeneration in this semi‐deciduous tropical rain forest, with the wide distribution of many species suggesting that these species regenerate continuously. Seedling diversity, density and distribution are largely a function of adult diversity, historical management type and environmental gradients in factors such as soil nutrient content and LAI. The species richness of seedlings was higher in soils both rich in titanium and with low exchangeable cations, as well as in logged areas that were more open and had a low LAI.     

Nitrogen deposition and forest nitrogen cycling along an urban–rural transect in southern China

There is increasing concern over the impact of atmospheric nitrogen (N) deposition on forest ecosystems in the tropical and subtropical areas. In this study, we quantified atmospheric N deposition and revealed current plant and soil N status in 14 forests along a 150 km urban to rural transect in southern China, with an emphasis on examining whether foliar δ¹⁵N can be used as an indicator of N saturation. Bulk deposition ranged from 16.2 to 38.2 kg N ha^?1 yr^?1, while the throughfall covered a larger range of 11.7–65.1 kg N ha^?1 yr^?1. Foliar N concentration, NO₃^? leaching to stream, and soil NO₃^? concentration were low and NO₃^? production was negligible in some rural forests, indicating that primary production in these forests may be limited by N supply. But all these N variables were enhanced in suburban and urban forests. Across the study transect, throughfall N input was correlated positively with soil nitrification and NO₃^? leaching to stream, and negatively with pH values in soil and stream water. Foliar δ¹⁵N was between ?6.6‰ and 0.7‰, and was negatively correlated with soil NO₃^? concentration and NO₃^? leaching to stream across the entire transect, demonstrating that an increased N supply does not necessarily increase forest δ¹⁵N values. We proposed several potential mechanism that could contribute to the δ¹⁵N pattern, including (1) increased plant uptake of ¹⁵N‐depleted soil NO₃^?, (2) foliage uptake of ¹⁵N‐depleted NH₄⁺, (3) increased utilization of soil inorganic N relative to dissolved organic N, and (4) increased fractionation during plant N uptake under higher soil N availability.     

High retention of 15N‐labeled nitrogen deposition in a nitrogen saturated old‐growth tropical forest

The effects of increased reactive nitrogen (N) deposition in forests depend largely on its fate in the ecosystems. However, our knowledge on the fates of deposited N in tropical forest ecosystems and its retention mechanisms is limited. Here, we report the results from the first whole ecosystem ¹⁵N labeling experiment performed in a N‐rich old‐growth tropical forest in southern China. We added ¹⁵N tracer monthly as ¹⁵NH₄¹⁵NO₃ for 1 year to control plots and to N‐fertilized plots (N‐plots, receiving additions of 50 kg N ha^?1 yr^?1 for 10 years). Tracer recoveries in major ecosystem compartments were quantified 4 months after the last addition. Tracer recoveries in soil solution were monitored monthly to quantify leaching losses. Total tracer recovery in plant and soil (N retention) in the control plots was 72% and similar to those observed in temperate forests. The retention decreased to 52% in the N‐plots. Soil was the dominant sink, retaining 37% and 28% of the labeled N input in the control and N‐plots, respectively. Leaching below 20 cm was 50 kg N ha^?1 yr^?1 in the control plots and was close to the N input (51 kg N ha^?1 yr^?1), indicating N saturation of the top soil. Nitrogen addition increased N leaching to 73 kg N ha^?1 yr^?1. However, of these only 7 and 23 kg N ha^?1 yr^?1 in the control and N‐plots, respectively, originated from the labeled N input. Our findings indicate that deposited N, like in temperate forests, is largely incorporated into plant and soil pools in the short term, although the forest is N‐saturated, but high cycling rates may later release the N for leaching and/or gaseous loss. Thus, N cycling rates rather than short‐term N retention represent the main difference between temperate forests and the studied tropical forest.     

Low stocks of coarse woody debris in a southwest Amazonian forest

The stocks and dynamics of coarse woody debris (CWD) are significant components of the carbon cycle within tropical forests. However, to date, there have been no reports of CWD stocks and fluxes from the approximately 1.3 million km² of lowland western Amazonian forests. Here, we present estimates of CWD stocks and annual CWD inputs from forests in southern Peru. Total stocks were low compared to other tropical forest sites, whether estimated by line-intercept sampling (24.4 ± 5.3 Mg ha⁻¹) or by complete inventories within 11 permanent plots (17.7 ± 2.4 Mg ha⁻¹). However, annual inputs, estimated from long-term data on tree mortality rates in the same plots, were similar to other studies (3.8 ± 0.2 or 2.9 ± 0.2 Mg ha⁻¹ year⁻¹, depending on the equation used to estimate biomass). Assuming the CWD pool is at steady state, the turnover time of coarse woody debris is low (4.7 ± 2.6 or 6.1 ± 2.6 years). These results indicate that these sites have not experienced a recent, large-scale disturbance event and emphasise the distinctive, rapid nature of carbon cycling in these western Amazonian forests.     

Woody Species Alpha‐diversity and Species Abundance Distributions in an African Semi‐deciduous Tropical Rain Forest

Understanding plant species diversity patterns and distributions is critical for conserving and sustainably managing tropical rain forests of high conservation value. We analyzed the alpha‐diversity, species abundance distributions, and relative ecological importance of woody species in the Budongo Forest, a remnant forest of the Albertine Rift in Uganda. In 32 0.5‐ha plots, we recorded 269 species in 171 genera and 51 families with stems of ≥2.0 cm in diameter at breast height (dbh). There were 53 more species with stems of ≥2.0 cm dbh than with stems of ≥10 cm dbh, of which 33 were treelets and 20 were multi‐stemmed shrubs. For both minimum stem diameter cut‐offs (i.e., ≥2 cm dbh vs. ≥10 cm dbh), the Fabaceae, Euphorbiaceae, Ulmaceae, and Meliaceae families and the species Cynometra alexandri, Lasiodiscus mildbraedii, and Celtis mildbraedii had the highest relative ecological importance. The relative ecological importance of some species and families changed greatly with the minimum stem diameter measured. Alpha‐diversity, species richness, and species abundance distributions varied across historical management practice types, forest community types, and as a function of minimum stem diameter. Species richness and Shannon–Weiner diversity index were greater for species with stems of ≥2.0 cm dbh than of ≥10 cm dbh. The decrease in species evenness with an increasing number of plots was accompanied by an increase in species richness for trees of both minimum diameters. This forest is characterized by a small number of abundant species and a relatively large proportion of infrequent species, many of which are sparsely distributed and with restricted habitats. We recommend lowering the minimum stem diameter measured for woody species diversity studies in tropical forests from 10 cm dbh to 2 cm dbh to include a larger proportion of the species pool.     

Managing open habitats by wild ungulate browsing and grazing: A case‐study in North‐Eastern Germany

Question: Can wild ungulates efficiently maintain and restore open habitats? Location: Brandenburg, NE Germany. Methods: The effect of wild ungulate grazing and browsing was studied in three successional stages: (1) Corynephorus canescens‐dominated grassland; (2) ruderal tall forb vegetation dominated by Tanacetum vulgare; and (3) Pinus sylvestris‐pioneer forest. The study was conducted over 3 yr. In each successional stage, six paired 4 m²‐monitoring plots of permanently grazed versus ungrazed plots were arranged in three random blocks. Removal of grazing was introduced de novo for the study. In each plot, percentage cover of each plant and lichen species and total cover of woody plants was recorded. Results: Wild ungulates considerably affected successional pathways and species composition in open habitats but this influence became evident in alteration of abundances of only a few species. Grazing effects differed considerably between successional stages: species richness was higher in grazed versus ungrazed ruderal and pioneer forest plots, but not in the Corynephorus sites. Herbivory affected woody plant cover only in the Pioneer forest sites. Although the study period was too short to observe drastic changes in species richness and woody plant cover, notable changes in species composition were still detected in all successional stages. Conclusion: Wild ungulate browsing is a useful tool to inhibit encroachment of woody vegetation and to conserve a species‐rich, open landscape.     

Tree diversity and distribution in undisturbed and human-impacted sites of tropical wet evergreen forest in southern Western Ghats, India

The composition, abundance, population structure and distribution patterns of all woody species ( 30 cm gbh) were investigated in an undisturbed and two adjacent human-impacted sites of a tropical wet evergreen forest in Kalakad National Park, Western Ghats, south India. Three 1-ha plots were established, one each in (i) an undisturbed site (named site UD), (ii) in a site selectively felled 35 years ago (site SF – small stems felled leaving the large trees (as shade) for developing it into a cardamom estate, on the failure of which the site was abandoned) and (iii) a frequently disturbed site (site FD – round woods logged for use in ovens for curing cardamom). These sites are 1 to 3 km apart in the same wet evergreen forest. In the three study plots a total of 2150 stems (mean density 716 ha^–1) covering 122 species in 89 genera and 41 families were enumerated. Species richness was greatest (85 species ha^–1) in the undisturbed site UD, intermediate (83) in SF and lowest (80) in FD. Tree density was greatest (855 stems ha^–1) in site SF, intermediate (720) in UD and lowest (575) in FD. The forest stand was exceptionally voluminous in site UD (basal area 94.64 m² ha^–1), intermediate (66.9 m²) in SF and least (61.7 m²) in FD, due to tree removal for fuel in the latter sites. Species composition and abundance patterns markedly varied between the three sites. In UD and SF, primary forest species (Cryptocarya bourdillonii , Cullenia exarillata Myristica dactyloides etc.) occurred in greater density. In FD heliophilic secondary forest species (Elaeocarpus venustus, Litsea wightiana, Viburnum punctatum and Vitex altissima) were abundant, while these were absent in UD and SF. The species–area curve did not reach an asymptote in any of the sites on the 1-ha scale. The stand population structure was clearly reverse J shaped in UD and SF, while small stems were 2- to 3-fold fewer in FD. Most trees exhibited clumped distribution of individuals on the 1-ha scale. Variation in the kind and richness of species and their abundance is related to human interference and the need for forest conservation is emphasized.     

Structural characteristics and aboveground biomass of old‐growth spruce–fir stands in the eastern Carpathian mountains,Ukraine

Abstract

Temperate old‐growth forests are known to have ecological characteristics distinct from younger forests, but these have been poorly described for the remaining old‐growth Picea abies–Abies alba forests in the eastern Carpathian mountains. In addition, recent studies suggest that old‐growth forests may be more significant carbon sinks than previously recognized. This has stimulated interest in quantifying aboveground carbon stocks in primary forest systems. We investigated the structural attributes and aboveground biomass in two remnant old‐growth spruce–fir stands and compared these against a primary (never logged) mature reference stand. Our sites were located in the Gorgany Nature Reserve in western Ukraine. Overstory data were collected using variable radius plots; coarse woody debris was sampled along line intercept transects. Differences among sites were assessed using non‐parametric statistical analyses. Goodness‐of‐fit tests were used to evaluate the form of diameter distributions. The results strongly supported the hypothesis that old‐growth temperate spruce–fir forests have greater structural complexity compared to mature forests, including higher densities of large trees, more complex horizontal structure, and elevated aboveground biomass. The late‐successional sites we sampled exhibited rotated sigmoid diameter distributions; these may reflect natural disturbance dynamics. Old‐growth Carpathian spruce–fir forests store on average approximately 155–165 Mg ha^?1 of carbon in aboveground tree parts alone. This is approximately 50% higher than mature stands. Given the scarcity of primary spruce–fir forests in the Carpathian region, remaining stands have high conservation value, both as habitat for late‐successional species and as carbon storage reservoirs.     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

The impact of nitrogen deposition on carbon sequestration in European forests and forest soils

An estimate of net carbon (C) pool changes and long‐term C sequestration in trees and soils was made at more than 100 intensively monitored forest plots (level II plots) and scaled up to Europe based on data for more than 6000 forested plots in a systematic 16 km × 16 km grid (level I plots). C pool changes in trees at the level II plots were based on repeated forest growth surveys At the level I plots, an estimate of the mean annual C pool changes was derived from stand age and available site quality characteristics. C sequestration, being equal to the long‐term C pool changes accounting for CO₂ emissions because of harvest and forest fires, was assumed 33% of the overall C pool changes by growth. C sequestration in the soil were based on calculated nitrogen (N) retention (N deposition minus net N uptake minus N leaching) rates in soils, multiplied by the C/N ratio of the forest soils, using measured data only (level II plots) or a combination of measurements and model calculations (level I plots). Net C sequestration by forests in Europe (both trees and soil) was estimated at 0.117 Gton yr^?1, with the C sequestration in stem wood being approximately four times as high (0.094 Gton yr^?1) as the C sequestration in the soil (0.023 Gton yr^?1). The European average impact of an additional N input on the net C sequestration was estimated at approximately 25 kg C kg^?1 N for both tree wood and soil. The contribution of an average additional N deposition on European forests of 2.8 kg ha^?1 yr^?1 in the period 1960–2000 was estimated at 0.0118 Gton yr^?1, being equal to 10% of the net C sequestration in both trees and soil in that period (0.117 Gton yr^?1). The C sequestration in trees increased from Northern to Central Europe, whereas the C sequestration in soil was high in Central Europe and low in Northern and Southern Europe. The result of this study implies that the impact of forest management on tree growth is most important in explaining the C pool changes in European forests.     

Carbon of Chaillu forests based on a phytosociological analysis in Republic of Congo,more than meets the eye?

The objectives were to quantify aboveground, belowground and dead wood carbon pools near Mayoko in the Chaillu massif of Republic of Congo and explore relationships between carbon storage and plant diversity of all growth forms. A total of 190 plots (25 m by 25 m) were sampled (5072 stems, 211 species) and data analysed using recommended central-African forest allometric equations. Mean stem diameter at breast height was 33.6 cm, mean basal area 47.7 m² ha⁻¹ and mean density of individuals 407 ha⁻¹. Mean aboveground carbon (AGC) ranged from 13.93–412.66 Mg C ha⁻¹, belowground carbon from 2.86–96.97 Mg C ha⁻¹ and dead wood from 0.00–7.59 Mg C ha⁻¹. The maximum AGC value recorded in a plot was 916 Mg C ha⁻¹. The analysis performed using phytosociological association as basis rather than broad vegetation type is unique. AGC values for undisturbed terra firme forest sites featured among the highest recorded for African tropical forests. Considering only tree diversity, a weak, yet significant, relationship existed between AGC and species richness, Shannon-Wiener index of diversity and Fisher's alpha. However, if diversity of all plant growth forms is considered, no relationship between carbon and plant diversity existed.     

Linking woody species diversity with plant available water at a landscape scale in a Brazilian savanna

Question: How is the diversity of woody species in a seasonally dry savanna related to plant available water (PAW)? Location: Savannas in central Brazil. Methods: Two‐dimensional soil resistivity profiles to 10‐m depth previously measured along three 10 m × 275 m replicate transects revealed differences in belowground water resources among and within transects: (1) driest/most heterogeneous; (2) wettest/least heterogeneous; and (3) PAW‐intermediate. All woody plants along these transects were identified to species, and height and basal circumference measured. Species diversity was evaluated for the whole transect (total diversity), 100‐m² plots (alpha‐diversity) and dissimilarity among 100‐m² plots within transects (beta‐diversity). Correlation analyses were conducted between PAW and vegetation variables at the 100‐m² scale. Results: The driest/most heterogeneous transect had the lowest total species diversity, while the wettest/least heterogeneous transect showed the lowest beta‐diversity. Floristic variation was correlated with PAW in all transects. In the most heterogeneous transect, species density was positively correlated with PAW in the 0‐400 cm soil layer. Evenness and Simpson's diversity were negatively correlated with PAW in the 700‐1000 cm soil layer. Conclusion: Woody species diversity was related to PAW at a fine spatial scale. Abundant PAW in the top 4 m of soil may favour many species and increase species total diversity. Conversely, abundant PAW at depth may result in lower evenness and total diversity, probably because the few species adapted to obtaining deep soil water can become dominant. Environmental changes altering soil water availability and partitioning in soil layers could affect the diversity of woody plants in this savanna.     

The Imprint of Land-use History: Patterns of Carbon and Nitrogen in Downed Woody Debris at the Harvard Forest

Few data sets have characterized carbon (C) and nitrogen (N) pools in woody debris at sites where other aspects of C and N cycling are studied and histories of land use and disturbance are well documented. We quantified pools of mass, C, and N in fine and coarse woody debris (CWD) in two contrasting stands: a 73-year-old red pine plantation on abandoned agricultural land and a naturally regenerated deciduous forest that has experienced several disturbances in the past 150 years. Masses of downed woody debris amounted to 40.0 Mg ha⁻¹ in the coniferous stand and 26.9 Mg ha⁻¹ in the deciduous forest (20.4 and 13.8 Mg C ha⁻¹, respectively). Concentrations of N were higher and C:N ratios were lower in the deciduous forest compared to the coniferous. Pools of N amounted to 146 kg N ha⁻¹ in the coniferous stand and 155 kg N ha⁻¹ in the deciduous forest; both are larger than previously published pools of N in woody debris of temperate forests. Woody detritus buried in O horizons was minimal in these forests, contrary to previous findings in forests of New England. Differences in the patterns of mass, C, and N in size and decay classes of woody debris were related to stand histories. In the naturally regenerated deciduous forest, detritus was distributed across all size categories, and most CWD mass and N was present in the most advanced decay stages. In the coniferous plantation, nearly all of the CWD mass was present in the smallest size class (less than 25 cm diameter), and a recognizable cohort of decayed stems was evident from the stem-exclusion phase of this even-aged stand. These results indicate that heterogeneities in site histories should be explicitly included when biogeochemical process models are used to scale C and N stocks in woody debris to landscapes and regions. Received 27 April 2001; accepted 4 January 2002.