The influence of tree species on canopy soil nutrient status in a tropical lowland wet forest in Costa Rica

The canopy is host to a large percentage of the flora and fauna in tropical wet forests and is distinct from the forest floor in plant richness, soil type and microclimate. In this study, we examined the influence of tree species and season on soil nutrient cycling processes in canopy soils of four tree species common to Costa Rican wet forests. We also compared the canopy soils to the associated forest floor mineral soils. Both tree species and season had strong effects on canopy soil nutrients and processes. Canopy soils from trees with high litter lignin concentrations had higher net N-mineralization rates and higher dissolved inorganic N concentrations than those with low lignin concentrations. During the dry season, net N-immobilization occurred and dissolved organic and inorganic N and available P concentrations were significantly higher than during the wet season. Overall, canopy soils had higher N levels and higher fungi + bacteria richness than forest floor mineral soils. The differences in canopy soil properties observed among tree species indicates that these species have distinct N cycles that reflect differences in both soil origin and biological controls.     

Factors associated with plant species richness in a coastal tall‐grass prairie

Abstract. In this study we examine the factors associated with variations in species richness within a remnant tall‐grass prairie in order to gain insight into the relative importance of controlling variables. The study area was a small, isolated prairie surrounded by wetlands and located within the coastal prairie region, which occurs along the northwestern Gulf of Mexico coastal plain. Samples were taken along three transects that spanned the prairie. Parameters measured included micro‐elevation, soil characteristics, indications of recent disturbance, above‐ground biomass (including litter), light penetration through the plant canopy, and species richness. Species richness was found to correlate with micro‐elevation, certain soil parameters, and light penetration through the canopy, but not with above‐ground biomass. Structural equation analysis was used to assess the direct and indirect effects of micro‐elevation, soil properties, disturbance, and indicators of plant abundance on species richness. The results of this analysis showed that observed variations in species richness were primarily associated with variations in environmental effects (from soil and microtopography) and were largely unrelated to variations in measures of plant abundance (biomass and light penetration). These findings suggest that observed variations in species richness in this system primarily resulted from environmental effects on the species pool. These results fit with a growing body of information that suggests that environmental effects on species richness are of widespread importance.     

宁夏黄土丘陵区不同生态恢复生境地表甲虫多样性

为了解宁夏黄土丘陵区不同生境地表甲虫群落多样性变化规律及与环境因子的关系, 并探讨不同生态恢复措施对维持地表甲虫群落多样性的影响, 2013年7-8月, 作者利用陷阱法调查了该区6种生境内的地表甲虫群落多样性。结果表明: 灌草混交林地、乔灌混交林地、生态薪炭林地中地表甲虫物种丰富度和个体数量均较高, 天然封育草地、生态经济林地和水平农田中物种丰富度较低, 生态经济林地和水平农田中甲虫个体数量显著高于天然封育草地。不同生境间, 灌草混交林地、乔灌混交林地与生态薪炭林地之间、生态经济林地与水平农田之间甲虫群落组成相似性较高。多元回归分析表明, 草本层生物量、灌木层盖度及土壤含水量是影响甲虫物种丰富度的决定因素, 林冠层盖度和枯落物厚度是决定地表甲虫个体数量的重要因素。CCA分析表明, 枯落物盖度、枯落物厚度、林冠层盖度及草本层盖度是影响地表甲虫群落组成的重要环境因子。研究表明, 灌草混交林地为地表甲虫群落多样性维持较好的生境类型, 是宁夏黄土丘陵区典型生态恢复的最优模式。     

Pattern and dynamics of the ground vegetation in south Swedish Carpinus betulus forests: importance of soil chemistry and management

The vegetation and environmental conditions of south Swedish hornbeam Carpinus betulus forests are described with data from 35 permanent sample plots The main floristic gradient of the ground vegetation is closely related to acid-base properties of the top soil base saturation, pH and organic matter content Other floristic differences are related to tree canopy cover and the distance of the sample plots to the Baltic coast Species richness of herbaceous plants typical of forests increases with soil pH, The number of other herbaceous species, occurring in both forests and open habitats, and of woody species is not related to pH Comparisons of vegetation data from 1983 and 1993 show relatively small compositional differences of the herbaceous forest flora The number of other herbaceous species increased considerably m those plots where canopy trees had been cut after 1983 The number of new species in managed plots increases with soil pH Species losses and gains of the herbaceous forest flora between 1983 and 1993 are generally lower as compared with other herbaceous species and woody species However, the ground cover of herbaceous forest species, especially of Oxalis acetosella and Lamium galeobdolon , was considerably lower in 1993 as compared to 1983 in both unmanaged and managed plots Possible explanations for this decrease are current soil acidification and drought during the growing season     

Disturbance of the herbaceous layer after invasion of an eutrophic temperate forest by wild boar

Disturbances of the soil and the tree canopy are crucial factors determining the diversity, composition and biomass of the herbaceous layer in forests. This study presents a detailed account of ground vegetation in permanent plots surveyed before and after invasion of wild boar (Sus scrofa) to a temperate deciduous broadleaf forest. Specifically, we aimed to quantify the effect of wild boar rooting on cover, richness and composition of spring ephemerals, summer green herbs and saplings of woody species in relation to tree canopy cover. Rooting frequency in sample plots increased from 0% in 2010 to 61% in 2013. In heavily rooted plots, the mean cover of spring ephemeral geophytes (mainly Anemone nemorosa, A. ranunculoides and Ranunculus ficaria) decreased from 75% to 39% between 2010 and 2013. Species richness of summer green herbs generally increased between 2010 and 2013 and was additionally positively affected by heavy rooting and low canopy cover. Rooting also caused heterogenization of the herbaceous layer and amplified ongoing compositional changes induced by changing light conditions. Frequency and richness of spring ephemeral and woody species remained unchanged. We conclude that overall species richness of the herbaceous layer may increase in the short‐term as a result of increased plant recruitment and seed dispersal. However, wild boar rooting can greatly reduce the ground cover of spring ephemerals in eutrophic broadleaf forests, thereby threatening their important ecological function. To avoid long‐term losses of characteristic spring flora elements, local population control of wild boar is necessary to reduce abundance and frequency of soil rooting.     

The effect of canopy disturbance on species richness in a central Himalayan oak forest

Non-epiphytic species richness was studied in different disturbance classes within a Quercus semecarpifolia forest. Nine disturbance classes were defined according to the degree of biomass removal (lopping) and their spatial mixture. Six of these were observed in the study area. The species were divided into three functional groups: climbers, phanerophytes, and field-layer plants. The primary aim was to test if there is an elevated species richness under an intermediate disturbed canopy for (i) all vascular plants, (ii) lianas, (iii) phanerophytes and (iv) field-layer species. The richness of the different plant groups and all species were fitted against the disturbance gradient by means of Generalized Linear Models (GLM). Other environmental variables such as altitude, potential solar radiation, light intensity, canopy cover and soil parameters were also evaluated as predictors. Disturbance classes, canopy cover and light intensity were combined into a new variable, disturbance-complex, using Principal Component Analyses.Phanerophytes did not respond to any variable. Climbers were mostly related to pH and canopy cover, and were the only group related to altitude, nitrogen and loss-on-ignition. Herbaceous plants and total species richness showed a unimodal response to disturbance classes and the complex disturbance gradient, which supports the intermediate disturbance hypothesis. Relative radiation and slope also supported a unimodal response in herbaceous plants, but disturbance had a significant additional contribution to this pattern. The most significant predictor for these two groups was pH. The responses to organic carbon and phosphorus were not significant for any of the subsets.The results indicate that a small-scale lopping regime will enhance species richness of vascular plants; only a few species in the intermediate disturbed forest are weedy ruderals. In such a situation, the conservation policy may accept small-scale human impact as part of the forest landscape.     

The species richness–productivity relationship in the herb layer of European deciduous forests

Aim In contrast to non‐forest vegetation, the species richness–productivity (SR‐P) relationship in forests still remains insufficiently explored. Several studies have focused on the diversity of the tree layer, but the species richness of temperate deciduous forests is mainly determined by their species‐rich herb layer. The factors controlling herb‐layer productivity may differ from those affecting tree layers or open herbaceous vegetation, and thus the SR‐P relationship and its underlying processes may differ. However, the few relevant studies have reported controversial results. Here we explore the SR‐P relationship in the forest herb layer across different areas from oceanic to continental Europe, and put the effect of habitat productivity on species richness into context with other key factors, namely soil pH and light availability. Location North‐western Germany, Czech Republic, Slovakia and southern Urals (Russia). Methods We measured herb‐layer species richness and biomass, soil pH and tree‐layer cover in 156 vegetation plots of 100 m² in deciduous forests. We analysed the SR‐P relationship and the relative importance of environmental variables using regression models for particular areas and separate forest types. Results We found a consistent monotonic increase in the herb‐layer species richness with productivity across all study areas and all forest types. Soil pH and light availability also affected species richness, but their relative importance differed among areas. Main conclusions We suggest that the monotonically increasing SR‐P relationship in the forest herb layer results from the fact that herb‐layer productivity is limited by canopy shading; competition within the herb layer is therefore not strong enough to exclude many species. This differs fundamentally from open herbaceous vegetation, which is not subject to such productivity limits and consequently exhibits a unimodal SR‐P relationship. We present a conceptual model that might explain the differences in the SR‐P relationship between the forest herb layer and open herbaceous vegetation.     

Effects on soil,microclimate and vegetation of the native-invasive <Emphasis Type="Italic">Retama monosperma</Emphasis> (L.) in coastal dunes

The aim of this study was to analyse and quantify the effects of the canopy of the native-invasive N-fixer woody shrub Retama monosperma in the dune ecosystem, affecting the structure and function of the dune environment as well as plant community, in the context of the facilitation mechanism. Air temperature and relative humidity; soil pH, electric conductivity, organic matter (OM) and nutrient content; above and below-ground vegetation biomass, litter mass, species richness and Shannon diversity were determined and compared from sampling plots below the R. monosperma canopy and in canopy gaps within a coastal dune system in SW Spain. The relationships between soil OM and nutrient contents and above and below-ground vegetation biomass, litter mass, species richness and Shannon diversity were also assessed. A predominance of positive interactions was confirmed. The canopy of R. monosperma ameliorated temperature extremes beneath, and soil OM and nutrient concentrations were increased by 188–466%, compared to those found in gaps. Plant biomass increased by 442% beneath the canopy and was composed almost exclusively of herbaceous annuals. Plant diversity was not affected. Plant communities were clearly structured as fertility islands, distributed in an environmentally stressful dune matrix characterized by scarce vegetation cover and low biomass.     

Interactions between Canopy Structure and Herbaceous Biomass along Environmental Gradients in Moist Forest and Dry Miombo Woodland of Tanzania

We have limited understanding of how tropical canopy foliage varies along environmental gradients, and how this may in turn affect forest processes and functions. Here, we analyse the relationships between canopy leaf area index (LAI) and above ground herbaceous biomass (AGB_H) along environmental gradients in a moist forest and miombo woodland in Tanzania. We recorded canopy structure and herbaceous biomass in 100 permanent vegetation plots (20 m × 40 m), stratified by elevation. We quantified tree species richness, evenness, Shannon diversity and predominant height as measures of structural variability, and disturbance (tree stumps), soil nutrients and elevation as indicators of environmental variability. Moist forest and miombo woodland differed substantially with respect to nearly all variables tested. Both structural and environmental variables were found to affect LAI and AGB_H, the latter being additionally dependent on LAI in moist forest but not in miombo, where other factors are limiting. Combining structural and environmental predictors yielded the most powerful models. In moist forest, they explained 76% and 25% of deviance in LAI and AGB_H, respectively. In miombo woodland, they explained 82% and 45% of deviance in LAI and AGB_H. In moist forest, LAI increased non-linearly with predominant height and linearly with tree richness, and decreased with soil nitrogen except under high disturbance. Miombo woodland LAI increased linearly with stem density, soil phosphorous and nitrogen, and decreased linearly with tree species evenness. AGB_H in moist forest decreased with LAI at lower elevations whilst increasing slightly at higher elevations. AGB_H in miombo woodland increased linearly with soil nitrogen and soil pH. Overall, moist forest plots had denser canopies and lower AGB_H compared with miombo plots. Further field studies are encouraged, to disentangle the direct influence of LAI on AGB_H from complex interrelationships between stand structure, environmental gradients and disturbance in African forests and woodlands.     

Soil fertility and herb facilitation mediated by Retama sphaerocarpa

Abstract. Woody legumes growing in dry climates can increase soil nutrient content and facilitate plant growth in their understorey. We investigated differences in soil fertility and herbaceous community in relation to the presence and absence of the legume Retama sphaerocarpa in a shrubland in a mediterranean type climate. The results showed a higher content of limiting nutrients for plant growth, such as phosphorus and nitrogen, under the R. sphaerocarpa canopy. Herbaceous biomass, cover and nitrogen content increased below the canopy. However, species richness and diversity were diminished in the presence of a dense canopy of R. sphaerocarpa. Nitrogen isotopic analysis did not indicate a clear relationship between symbiotic fixation in R. sphaerocarpa and nitrogen content of soil and plants under its canopy. Nevertheless, herbs growing in the understorey showed a linear correlation between foliar N content and ¹⁵N values. The existence of a dense shrub canopy induced a smaller monthly variation in herb richness, diversity and biomass, suggesting that it provides a stable microhabitat facilitating herbaceous vegetation establishment and growth. The ‘shelter’ effect was more important when environmental conditions became stressful for herbaceous plants.     

Environmental correlates of plant diversity in Korean temperate forests

Mountainous areas of the Korean Peninsula are among the biodiversity hotspots of the world's temperate forests. Understanding patterns in spatial distribution of their species richness requires explicit consideration of different environmental drivers and their effects on functionally differing components. In this study, we assess the impact of both geographical and soil variables on the fine-scale (400 m²) pattern of plant diversity using field data from six national parks, spanning a 1300 m altitudinal gradient. Species richness and the slopes of species–area curves were calculated separately for the tree, shrub and herb layer and used as response variables in regression tree analyses. A cluster analysis distinguished three dominant forest communities with specific patterns in the diversity–environment relationship. The most widespread middle-altitude oak forests had the highest tree richness but the lowest richness of herbaceous plants due to a dense bamboo understory. Total richness was positively associated with soil reaction and negatively associated with soluble phosphorus and solar radiation (site dryness). Tree richness was associated mainly with soil factors, although trees are frequently assumed to be controlled mainly by factors with large-scale impact. A U-shaped relationship was found between herbaceous plant richness and altitude, caused by a distribution pattern of dwarf bamboo in understory. No correlation between the degree of canopy openness and herb layer richness was detected. Slopes of the species–area curves indicated the various origins of forest communities. Variable diversity–environment responses in different layers and communities reinforce the necessity of context-dependent differentiation for the assessment of impacts of climate and land-use changes in these diverse but intensively exploited regions.     

荒漠草原人工柠条林对冠下草本植物群落的保育作用

以宁夏盐池县荒漠草原人工柠条（Caragana intermedia）林为研究对象,分别选取柠条林冠下东侧（SE）,冠下西侧（SW）及带间（Gap）为研究样地,从群落水平探讨柠条对冠下草本植物群落结构、物种多样性及功能群分布的影响。结果表明：（1）在3种微生境中均鉴定出12种植物,SW以蒙古冰草为优势种,SE以蒙古冰草和中亚白草为优势种,Gap则以蒙古冰草和牛枝子为优势种。（2）与Gap相比,SW和SE植物群落的平均高度分别增加了41.06%和81.75%,地上生物量分别增加了40.88%和38.73%。SW和SE中,禾本科植物地上生物量分别占地上总生物量的67.10%和58.40%,显著高于Gap （P<0.05）。（3）柠条冠层效应使得草本植物的物种丰富度指数增加,但Shannon-Winner指数、Simpson指数和Pielou指数显著差异（P>0.05）,变化范围分别为1.620-1.756、0.701-0.730和0.775-0.878。（4）冗余分析表明：土壤温度、空气相对湿度及土壤有机碳是影响草本植物物种多样性及生物量的主要因子,解释量分别为42.70%,11.70%和8.80%。研究表明,柠条对冠下草本植物群落尤其是禾本科植物具有一定的保育作用,该效应的产生主要是由于柠条冠下微气候及土壤环境因子的改善为草本植物的生长发育提供了有利条件。柠条对草本植物的保育作用对荒漠草原生态环境的保护与恢复具有重要意义。     

Savanna tree influence on understory vegetation and soil nutrients in northwestern Kenya

Abstract. Contrary to observations and models in which trees and herbaceous plants are viewed as competitors, we found that trees in an African savanna have positive impacts on herbaceous biomass production and composition, and on soil nutrient status. In the Turkana District of northwestern Kenya, we investigated vegetation and soil gradients along equi-angular transects radiating from the boles of individual Acacia tortilis trees. Total herbaceous biomass averaged 260 ± 17(se) g/m² at the bole and declined to 95 ± 8 g/m² in the tree interspaces. Soil organic carbon and total nitrogen concentrations were greatest (0.72 % and 0.083 %, respectively) in shallow soils near the bole and declined rapidly toward the interspaces and with increasing depth. Transects were also established between tree pairs to assess effects of differential canopy proximities. Grass production averaged 220 ± 21 g / m² below overlapping canopies, 150 ± 15 g / m² under individual canopies, and 95 ± 8 g / m² in interstitial areas. Detrended correspondence analysis revealed that shifts in species composition were correlated with distance from tree bole out to the edge of the canopy. Species response, in terms of relative cover, to increasing distance from the bole, seemed to fall into five general classes: 1) greatest at the bole, 2) increasing with distance from the bole, 3) greatest in the mid canopy zone, 4) least at the bole and 5) no response. Trees did not influence herbaceous compositionbeyondtree canopies. It is assumed that shade cast by the tree canopy with subsequent reductions of understory water stress and temperature and increased nutrient concentrations may be the most important factors affecting understory soil and vegetation.     

Above-ground biomass and species richness in a Mediterranean salt marsh

Abstract. Species composition, above-ground biomass and diversity of herbaceous communities were recorded in 50 sites on the upper salt marsh of the Guadalquivir delta (SW Spain). Through Canonical Correspondence Analysis relationships were shown between plant-species characteristics abundance and environmental variables. Seven community types were distinguished and related to hydro logical regime and physical and chemical soil parameters. The higher species richness was associated with moderate values of above-ground biomass (Biomass at maximum diversity, BMD = 302 g/m²). Salinity was negatively correlated with species richness. There was no significant correlation with total above-ground biomass. Water regime and soil nutrient (Ca, Mg) content were significant predictors of the community above-ground biomass.     

Effects of Quercus emoryi on herbaceous vegetation in a semi-arid savanna

Ten trees (5–70 m² canopy area) were selected to determine effects of tree size (crown area) on herbaceous species composition and biomass in a Quercus emoryi savanna in southeastern Arizona. Consistent with most studies in temperate savannas, herbaceous biomass was reduced beneath the canopy relative to grassland areas. However, tree size appeared to exert no influence over herbaceous biomass. In contrast to most temperate savannas, Q. emoryi trees did not affect distribution of herbaceous species.     

Litter quality effects of beech and hornbeam on undergrowth species diversity in Luxembourg forests on limestone and decalcified marl

Question: Do beech and hornbeam differ in undergrowth species diversity, and could these differences be explained by mass of the organic layer, pH or soil moisture? Could species richness under beech and hornbeam be explained by differences in N dynamics? Location: Ancient forests dominated by beech and hornbeam in Luxembourg, on decalcified and lime‐rich parent material. Methods: Species composition and soil characteristics, i.e. mass of the organic layer, pH, moisture content, bulk density and air‐filled pore space, were determined under beech and hornbeam on decalcified marl and limestone, and analysed with cluster analysis, correspondence analysis and ANOVA, with tree species and parent material as independent factors. Net N mineralization, nitrification and N in soil and microbial biomass were determined in a laboratory incubation experiment over a pH gradient, and correlated with species richness. Results: Species richness was lower under beech than under hornbeam on both decalcified marl and on limestone. Mass of the organic layer was higher under beech in all cases, but pH and soil moisture only differed between tree species on decalcified marl. Species richness increased with soil N, microbial N and nitrification in the mineral topsoil, but was not correlated with net N mineralization, and was negatively correlated with nitrification in the organic layer and with efficiency of N mineralization per unit microbe. Conclusion: Litter quality affected species richness on both limestone and decalcified marl via the organic layer, but only on decalcified marl in terms of pH or soil moisture. Species richness did not seem to be stimulated by high overall net N mineralization or high nitrate availability in the organic layer, but only by high nitrate in the mineral topsoil.     

Seedling and Sapling Dynamics of Treefall Pits in Puerto Rico1

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N–fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N–fixing sym–bionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer–term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.     

Tree diversity and species identity effects on soil fungi,protists and animals are context dependent

Plant species richness and the presence of certain influential species (sampling effect) drive the stability and functionality of ecosystems as well as primary production and biomass of consumers. However, little is known about these floristic effects on richness and community composition of soil biota in forest habitats owing to methodological constraints. We developed a DNA metabarcoding approach to identify the major eukaryote groups directly from soil with roughly species-level resolution. Using this method, we examined the effects of tree diversity and individual tree species on soil microbial biomass and taxonomic richness of soil biota in two experimental study systems in Finland and Estonia and accounted for edaphic variables and spatial autocorrelation. Our analyses revealed that the effects of tree diversity and individual species on soil biota are largely context dependent. Multiple regression and structural equation modelling suggested that biomass, soil pH, nutrients and tree species directly affect richness of different taxonomic groups. The community composition of most soil organisms was strongly correlated due to similar response to environmental predictors rather than causal relationships. On a local scale, soil resources and tree species have stronger effect on diversity of soil biota than tree species richness per se.     

Woody species as landscape modulators: their effect on the herbaceous plants in a Mediterranean maquis

“Landscape modulators” are ecosystem engineers that have an impact on community structure by creating patches in the landscape mosaic. Our aim was to study the effect of evergreen-trees, as landscape modulators, on herbaceous plants in a Mediterranean maquis system in northern Israel. We examined the effects of canopy removal and cattle grazing on species richness, plant functional types, and rare plant species in two patch-types: (1) woody—under tree canopy (or the location of a removed canopy); (2) herbaceous—in open areas with no tree canopy. Patch-type and tree removal affected species richness and plant functional types. The extreme negative effect of the woody patch-type on species richness disappeared soon after the removal of the landscape modulator canopy. We conclude that the dominant effect of the evergreen woody landscape modulators can be regulated by canopy removal and grazing for maintaining patch-type and landscape diversities, and consequent high species richness in Mediterranean ecosystems, which is a main goal of global nature conservation policy.     

Scale‐dependence of environmental effects on species richness in oak savannas

Abstract. In order to investigate how scale (grain size) affects the relationships between species richness and environmental drivers (such as stress and disturbance), we collected 12 nested quadrats (from 0.25 m² to 1023 m²) from seven remnant oak savannas located in the floodplain of the Chippewa River in western Wisconsin, USA. Large and small‐scale richness were not significantly correlated, suggesting that small‐scale richness is not strongly controlled by sampling effects of the local species pool. Linear and curvilinear regressions between species richness and disturbance, canopy cover, biomass, and soil organic matter were dependent on sampling scale (grain size). Disturbance by fire was strongly related to richness at small scales, while tree canopy cover was strongly related to richness at larger scales. While there was some evidence suggesting the transition from disturbance to canopy effects occurs between 10 and 100 m², the transition was not particularly abrupt. The results cast doubt on the general importance of local species pools in affecting small‐scale richness as well as our ability to make generalizations that do not explicity include scale.