Humus characteristics and seasonal changes of soil arthropod communities in a natural sessile oak (Quercus petraea L.) stand and adjacent Austrian pine (Pinus nigra Arnold) plantation

In order to assess the effects of conversion of natural stands into plantations, soil invertebrate micro- and macroarthropod communities were evaluated for their abundance and richness in a sessile oak (SO; Quercus petraea L.) stand and adjacent Austrian pine (AP; Pinus nigra Arnold) plantation. Sites were sampled four times a year in 3-month intervals from May 2009 to February 2010. Humus characteristics such as total mass; carbon, lignin, and cellulose contents; and C/N ratio were significantly different between SO and AP. Statistically significant differences were detected on soil pH, carbon and nitrogen contents, and electrical conductivity between the two sites. The number of microarthropods was higher in AP than in the SO site. The annual mean abundance values of microarthropods in a square meter were 67,763 in AP and 50,542 in SO, and the annual mean abundance values of macroarthropods were 921 m^?2 in AP and 427 m^?2 in SO. Among the soil microarthropods, Acari and Collembola were the dominant groups. Shannon’s diversity index was more affected by evenness than species number despite the species diversity (H′) of soil arthropods being generally higher in the SO stand. The abundance of microarthropods showed clear seasonal trends depending upon the humidity of the soil.     

Impacts of logging and wildfire on an upland black spruce community in northwestern Ontario

Plant species composition and community structure were compared among four sites in an upland black spruce community in northwestern Ontario. One site had remained undisturbed since the 1930s and three had been disturbed by either logging, fire, or both logging and fire. Canonical correspondence ordination analyses indicated that herbaceous species composition and abundance differed among the disturbance types while differences in the shrub and tree strata were less pronounced. In the herb stratum Pleurozium schreberi, Ptilium crista-castrensis and Dicranum polysetum were in greatest abundance on the undisturbed forest site, while the wildfire and burned cutover sites were dominated by Epilobium angustifolium and Polytrichum juniperinum. The unburned harvested site was dominated by Epilobium angustifolium, Cornus canadensis and Pleurozium schreberi. Species richness was lower on the undisturbed site than on any of the disturbed sites while species diversity (H) and evenness (Hill's E5) were higher on the unburned harvested site than on the other sites. Results suggest that herb re-establishment is different among harvested and burned sites in upland black spruce communities and we hypothesize that differences in the characteristics of the disturbance were responsible, in particular, the impact of burning on nutrient availability. These differences need to be taken into account in determining the effects of these disturbances on biodiversity and long-term ecosystem management.     

The new flora of northeastern USA: quantifying introduced plant species occupancy in forest ecosystems

Introduced plant species have significant negative impacts in many ecosystems and are found in many forests around the world. Some factors linked to the distribution of introduced species include fragmentation and disturbance, native species richness, and climatic and physical conditions of the landscape. However, there are few data sources that enable the assessment of introduced species occupancy in native plant communities over broad regions. Vegetation data from 1,302 forest inventory plots across 24 states in northeastern and mid-western USA were used to examine and compare the distribution of introduced species in relation to forest fragmentation across ecological provinces and forest types, and to examine correlations between native and introduced species richness. There were 305 introduced species recorded, and 66 % of all forested plots had at least one introduced species. Forest edge plots had higher constancy and occupancy of introduced species than intact forest plots, but the differences varied significantly among ecological provinces and, to a lesser degree, forest types. Weak but significant positive correlations between native and introduced species richness were observed most often in intact forests. Rosa multiflora was the most common introduced species recorded across the region, but Hieracium aurantiacum and Epipactus helleborine were dominant in some ecological provinces. Identifying regions and forest types with high and low constancies and occupation by introduced species can help target forest stands where management actions will be the most effective. Identifying seemingly benign introduced species that are more prevalent than realized will help focus attention on newly emerging invasives.     

The risk of harmful algal blooms (HABs) in the oyster-growing estuaries of New South Wales, Australia

The spatial and temporal variability of potentially harmful phytoplankton was examined in the oyster-growing estuaries of New South Wales. Forty-five taxa from 31 estuaries were identified from 2005 to 2009. Harmful species richness was latitudinally graded for rivers, with increasing number of taxa southward. There were significant differences (within an estuary) in harmful species abundance and richness for 11 of 21 estuaries tested. Where differences were observed, these were predominately due to species belonging to the Pseudo-nitzschia delicatissima group, Dinophysis acuminata, Dictyocha octonaria and Prorocentrum cordatum with a consistent upstream versus downstream pattern emerging. Temporal (seasonal or interannual) patterns in harmful phytoplankton within and among estuaries were highly variable. Examination of harmful phytoplankton in relation to recognised estuary disturbance measures revealed species abundance correlated to estuary modification levels and flushing time, with modified, slow flushing estuaries having higher abundance. Harmful species richness correlated with bioregion, estuary modification levels and estuary class, with southern, unmodified lakes demonstrating greater species density. Predicting how these risk taxa and risk zones may change with further estuary disturbance and projected climate warming will require more focused, smaller scale studies aimed at a deeper understanding of species-specific ecology and bloom mechanisms. Coupled with this consideration, there is an imperative for further taxonomic, ecological and toxicological investigations into poorly understood taxa (e.g. Pseudo-nitzschia).     

Ants as Indicators of Exposure to Environmental Stressors in North American Desert Grasslands

The relative abundance of ant species was measured by pit-fall trapping at 44 sites in southern New Mexico and southeastern Arizona, U.S.A.. Sites were selected for study based on documentation of a history of disturbance or protection from disturbance, exposure to varying intensities of livestock grazing, dominance by an exotic species of plant and vegetation change resulting from disturbance or restoration efforts. Ant community composition, relative abundances of species, and species richness were the same on disturbed and undisturbed sites. None of the metrics based on hypothesized responses of ants to disturbance clearly distinguished between disturbed and undisturbed sites. Ant communities on sites where restoration efforts have resulted in distinct differences in vegetative cover and composition were similar to the ant communities on degraded unrehabilitated sites on the same soil type. Ant communities in riparian cottonwood gallery forests in Arizona and New Mexico were similar but differed from the assemblages in exotic salt cedar and native ash riparian woodlands. Ant species exhibited remarkable resistance to human-induced disturbances in these rangeland areas. In grasslands dominated by the South African grass, Eragrostis lehmanniana Nees, large seed harvesting ants, Pogonomyrmex spp., were greatly reduced in abundance compared to native grasslands. Other ant metrics were not different in E. lehmanniana grasslands and native grasslands. We conclude that ants cannot be used as indicators of exposure to stress, ecosystem health or of rehabilitation success on rangeland ecosystems. Ants are also not useful indicators of faunal biodiversity in rangeland ecosystems.     

Quantifying the consequences of conifer succession in aspen stands: decline in a biodiversity-supporting community

Quaking aspen (Populus tremuloides Michaux) stands are important for biodiversity in conifer-dominated forest landscapes. Our goal was to quantify the consequences of conifer succession on understory diversity and litter quality, as well as associated changes in aspen stand condition. We studied aspen stands on national park land in the transition zone between the northern Sierra Nevada and southern Cascade mountain ranges. We field-measured ten metrics of aspen stand condition in 29 aspen stands. Along a gradient of increasing current conifer cover, we observed decreases in herbaceous species diversity and richness and an increase in forest floor O horizon depth. We interpreted aerial photos from 1952 and 1998 to determine whether directional changes in conifer cover had occurred in the stands over the past half century, and used regression modeling to associate succession with the observed range of aspen stand condition. From the period 1952 to 1998, we found that conifer encroachment occurred in half the sampled stands, with an average increase in conifer cover of 1 % a year. Aspen were persistent in the remaining stands. Stand cover dynamics and percent total canopy cover interacted to influence species richness, diversity, aspen sprouting, and litter quality. In stands with conifer encroachment, both understory species richness and diversity declined. Although aspen sprouting increased, aspen establishment declined and the relative mass of woody to fine soil litter increased.     

Visitor impact on rocky shore communities of Qeshm Island, the Persian Gulf, Iran

The influence of visitors on macroinvertebrates of rocky intertidal shores was investigated in southern coasts of the Qeshm Island, the Persian Gulf, Iran. Qeshm Island located at the Strait of Hormuz, with an area of 1,491 km², is the largest island in the region. This island consists of a number of important natural habitat types including creeks, mangroves, corals, and sandy, muddy, and rocky shores that accommodate diverse marine flora and fauna communities. Two rocky shores were selected at the touristic beaches being visited regularly, and further two control locations selected at pristine shores. Intertidal macroinvertebrates were collected from six microhabitats including rock platforms, cobbles, boulders, crevices, sea walls, and rock pools during two different periods representing high and low tourist seasons. Species richness, density, and assemblage structure in heavily visited shores were compared with that of control locations. Striped barnacles (Balanus amphitrite) were present on platforms of all locations, thus the changes in their size were used as the obvious contrast associated with visitor’s impact. A total of 70 macroinvertebrate species from 11 phyla were recorded. Significant differences were detected in taxonomic richness, density, and assemblage structure of macroinvertebrates between heavily visited and pristine shores, suggesting that macroinvertebrates were adversely affected by visitors’ impact at heavily visited shores. The test of changes in species richness, density, and assemblage structure from high to low seasons yielded mixed results. The significant changes in density and assemblage structure from high to low seasons were only observed in one heavily visited shore. A significant reduction in size of striped barnacles was observed only in one heavily visited shore. The opportunistic or fugitive species (e.g., small macroalgae and barnacles) were dominant macroinvertebrates on heavily visited shores indicating early succession stage. The results presented here showed that macroinvertebrates were adversely affected by human activities in subtropical rocky shore.     

Types of Ectomycorrhiza as Pollution Stress Indicators: Case Studies in Slovenia

Mycorrhiza is the main spatial and temporal linkage between different constituents in a forest ecosystem. The functional compatibility and stress tolerance of ectomycorrhizal types is species specific, and therefore the information on the ectomycorrhizal community structure can add to the understanding of processes in forest ecosystems and can also be applied as tools for bioindication of pollution stress in forest soils. We have studied the effects of pollution (N and S) on trees and forest soils by: (1) quantification of ECM types diversity as in situ indicators in forest stands, (2) determination and quantification of pollution-sensitive or -insensitive ECM types as passive monitors, (3) root growth and development of ECM on nonmycorrhizal spruce seedlings, planted at the studied sites (active monitors), and (4) ECM infection (a bioassay based on mycorrhizal inoculum potential) of seedlings in an experimental set-up as ex situ testers. ECM species richness for Norway spruce trees (Picea abies) showed higher values in unpolluted sites than in polluted ones, while the differences were not significant for European beech trees (Fagus sylvatica). As pollution-sensitive or -insensitive ECM species in spruce forests, we suggest Hydnum rufescens (sensitive) and Paxillus involutus (unsensitive). Mycorrhizal potential in Norway spruce seedlings as a bioassay for soil N and S pollution was effective, and is suggested as an additional, standardized and widely comparable system in bioindication of soil pollution.     

Spatiotemporal variation of macroinvertebrates in relation to canopy cover and other environmental factors in Eriora River, Niger Delta, Nigeria

Canopy cover is well known to influence the distribution of macroinvertebrates in temperate streams. Very little is known about how this factor influences stream communities in Afrotropical streams. The effects and possible interactions of environmental factors and canopy cover on macroinvertebrate community structure (abundance, richness, and diversity) were examined in four stations in Eriora River, southern Nigeria bimonthly from May to November 2010. The river supported diverse macroinvertebrates in which the upstream sampling stations with dense canopy cover were dominated by Decapoda, Ephemeroptera, Odonata, Gastropoda, Trichoptera, and Coleoptera while Diptera and Coleoptera were the benthic organisms found predominant at downstream stations with less canopy cover. Some caddisfly species such as Agapetus agilis, Trichosetodes species and the stonefly Neoperla species were present upstream and were found to be potential bioindicators for a clean ecosystem. The blood worm Chironomus species and Tabanus sp. were abundant at the downstream of the river and are considered potential bioindicators for an organically degrading ecosystem. Some environmental factors varied temporally with significantly higher macroinvertebrate abundance and richness in May. We found out that canopy cover and environmental factors affected macroinvertebrates abundance, diversity, and richness and that the individual taxon had varying responses to these factors. These results help identify the mechanisms underlying the effects of canopy cover and other environmental factors on Afrotropical stream invertebrate communities.     

城镇化对西苕溪大型底栖动物群落的影响

于2004—2010年5月对西苕溪流域20个参照点,5个中度干扰点以及2个重度干扰点的底栖动物进行了调查,共鉴定出74科190属226个种;受干扰后,底栖动物群落物种丰富度、EPT物种丰富度、Shannon-wiener多样性指数和B-IBI指数显著下降,BI指数显著升高。CCA分析结果表明,参照、中度和重点干扰样点的底栖动物群落差异明显。城镇化引起的溪流水温上升、营养盐升高、泥沙输入量增多和堤岸固化是导致溪流底栖动物群落退化的主要原因。     

Effect of diel period and season on seining effort required to detect changes in Lake Erie beach fish assemblages

Using beach seining data from Lake Erie beaches, we examined the effect of diel period and season on two common study objectives considered sensitive to sampling effort: (1) has fish assemblage composition changed between two sampling time periods?, and (2) how many samples are necessary to provide a representative sample of the fish assemblage? Across a range of effect sizes and power levels, sample sizes were estimated for species richness, total fish abundance, round goby (Neogobius melanostomus) abundance and emerald shiner (Notropis atherinoides) abundance. For most sampling periods and effect sizes, several hundred to several thousand seine hauls were estimated to be required for detecting large-scale (e.g. basin-wide) changes in round goby and emerald shiner abundance. Depending on the indicator of interest, night seining (either spring or fall) was interpreted as the most efficient approach. For both seasons, sampling effort beyond 40 to 50 night seine hauls results in few additional species being detected. Both species accumulation curves and estimated sample sizes indicate that less effort is required to detect species richness changes than fish abundance. For most effect sizes and power levels, estimates of the number of seine hauls required to detect changes in fish abundance are unrealistically large for most monitoring programs. More practical monitoring strategies could be achieved by adopting alternative indicators (e.g. guilds), a more liberal significance level (p=0.1), a paired-site sampling design, or including reference sites.     

Eight nonnative plants in western Oregon forests: Associations with environment and management

Nonnative plants have tremendous ecological and economic impacts on plant communities globally, but comprehensive data on the distribution and ecological relationships of individual species is often scarce or nonexistent. The objective of this study was to assess the influence of vegetation type, climate, topography, and management history on the distribution and abundance of eight selected nonnative plant taxa in forests in western Oregon. These eight taxa were selected as being reliably detected by a multi-resource inventory of 1127 systematically-placed plots on nonfederal forest lands from 1995 to 1997 by the USFS Forest Inventory and Analysis (FIA) program. One or more of the eight nonnative taxa studied were found on 20% of the sampled subplots in the study area, but relatively few stands were dominated by them. Overall abundance of nonnative taxa was likely much greater, because few composites and graminoids were identified to species in this general-purpose inventory. Distribution of most taxa was more closely associated with low density of overstory trees than with climate. Nonnative taxa were significantly more abundant in stands that had been recently clearcut or thinned than in stands that had not. Frequencies of several taxa decreased with elevation, which may reflect proximity to source populations and intensive land use rather than any climatic constraints. Although the greatest potential for displacement of native forest species appears to be in early-successional communities, the potential for spread of some shade-tolerant evergreen shrubs also seems high.     

The Monitoring of Insects to Maintain Biodiversity in Ogawa Fores Reserve

The results of a biodiversity monitoring program conducted in the Ogawa Forest Reserve and its vicinity, situated in a cool temperate region of Japan, identified three different patterns for species richness. Forests of the region are characterized by a mosaic of secondary deciduous stands of various ages scattered among plantations of conifers. The three different types of change in species richness observed in response to the stand age are as follows: Type I (butterflies, tube-renting bees and wasps, hoverflies, fruit flies, and longicorn beetles), the species diversity was highest in open areas, just after clear-cutting, decreasing with the stand age; Type II (mushrooms and mites associated with them), older stands showed greater diversity than younger stands; and, Type III (moths, oribatid mites, collembolas, carabid beetles, and ants), the number of species did not change greatly with the stand age, though ordination analysis revealed that there was variation in species compositions. These results indicate that combinations of stands of different ages, or heterogeneously arranged stands, can contribute to the maintenance of insect biodiversity at the landscape level.     

The interactions between tree-herb layer diversity and soil properties in the oriental beech (<Emphasis Type="Italic">Fagus orientalis</Emphasis> Lipsky) stands in Hyrcanian forest

This study investigated the interactions between tree-herb layer diversity and some physico-chemical and eco-physiological characteristics of soil in natural oriental beech stand in western Guilan, Iran. The data were collected from nine research sites (50 m?×?50 m) which were described as a gradient from pure oriental beech (Fagus orientalis Lipsky) stands to mixed stands with up to nine deciduous tree species (n =?27) in Hyrcanian forest. Herbaceous plants were sampled within ten 1 m?×?1 m sub-plots in two plots of 400 m² which were installed randomly in each research site. Composite soil samples were taken at five positions in each research site. We found that the increase in tree diversity in mature oriental beech stands brought about an increase in microbial biomass carbon, soil carbon content, and the ratio of microbial biomass carbon to the organic carbon (C_mic/C_org). Increased soil organic carbon raised microbial biomass carbon through creating suitable environment for microorganisms. The findings also indicated that the ratio of microbial biomass carbon to the organic carbon (C_mic/C_org) increased as a quantitative indicator of soil carbon dynamics that finally benefits soil fertility of mixed oriental beech stands compared to pure oriental beech stands. The results showed that humus layer and litter thickness were negatively correlated with tree layer richness. Generally, it can be stated that maintaining a mixture of tree layer species in natural oriental beech stands results in an increase in richness and diversity values of herb plants as well as carbon content and microbial biomass carbon of soil.     

Derivation and Use of Sediment Quality Guidelines for Ecological Risk Assessment of Metals and Radionuclides Released to the Environment from Uranium Mining and Milling Activities in Canada

The Screening Level Concentration (SLC) approach was used to derive Lowest Effect Level (LEL) and Severe Effect Level (SEL) concentrations for nine metals (As, Cr, Cu, Pb, Mo, Ni, Se, U and V) and three radionuclides (²²⁶Ra, ²¹⁰Pb, and ²¹⁰Po) released to the aquatic environment during the mining and milling of uranium ore. This method was chosen because it allowed for the best use of the considerable historical and current data collected for diverse purposes in the uranium mining and milling regions of Canada (20,606 data points used in the analysis). Except for Cr, all the LELs derived in this study using the weighted method and published sediment quality guidelines (SQGs) were highly reliable (> 85%) in predicting sites unimpacted by uranium mining/milling defined as sites where reductions in the abundance and species richness of benthic invertebrate communities were < 20%. The derived SEL values and corresponding published SQGs (with the exception of Ni) were not reliable predictors (≤60%) of severe impacts on benthic invertebrate communities when severe impacts are defined as a reduction in abundance and species richness ≥40%. Most of the severely impacted sites had sediment contaminant concentrations well below the SEL values. It is concluded that LELs derived using the weighted method can reliably be used in ecological risk assessments as concentrations below which adverse effects on benthic invertebrate communities are not expected. In contrast, it is recommended that SELs not be used in assessments of uranium mining/milling activities as concentrations above which adverse effects are anticipated.     

An exploration of the relationships betweenmacroinvertebrate community composition and physical andchemical habitat characteristics in farm dams

Recently, Australian interest in farm dams has focused on rates of harvest of surface waters (runoff), and the impact this has on nearby natural systems. Little research has been directed towards the role of these artificial water bodies in sustaining biological reserves within the wider ecosystem. Macroinvertebrate communities in three farm dams in the Central Tablelands of New South Wales were surveyed, and water quality variables were correlated with species richness and abundance. Community responses to habitat factors including sediment depth, stock use, vegetation and debris were also examined. Communities were described at several taxonomic levels in addition to allocation to trophic groups and primary functional feeding groups.Species richness and abundance of communities were found to vary between dams and between habitat types within dams. The extent of these differences was decreased when communities were described by either trophic status or functional feeding mechanisms. Habitats were influenced by water quality and by physical features of the habitat, with the two factors interacting to define equilibrium conditions. Localised conditions resulted in different macroinvertebrate communities. Physicochemical parameters that correlated most closely with communities included light penetration, chlorophyll-a and conductivity. Habitat factors that were most frequently linked with communities were sediment depth and canopy cover, with localised disturbances related to stock use affecting feeding groups rather than specific taxa.One of the major problems associated with increasing modification of landscapes by agriculture or urbanisation is the fragmentation of undisturbed habitats. Creation of joint aquatic and woodland habitats enhances biodiversity corridors. The recognition of the potential for farm dams as reservoirs of biodiversity and development of management practices that optimise this neglected biodiversity reserve may have much wider benefits biologically, aesthetically and productively.     

Losses of biota from American aquatic communities due to acid rain

Models based on chemical survey data and geochemical assumptions were calibrated for areas where rates of acidification are known, then used to predict the declines in alkalinity and pH of lakes in the eastern and midwestern U.S.A. These results were combined with known acid tolerances of different taxonomic groups to estimate the extent of damage caused by acid rain to biological assemblages.An average of over 50% of the species in some taxonomic groups have probably been eliminated from lakes in the Adirondacks, Poconos-Catskills and southern New England. Moderate damage to biotic communities was predicted for lakes in central New England, and north-central Wisconsin. Damage predicted in Maine, upper Michigan, northeastern Minnesota and the remainder of the upper Great Lakes region was slight. Crustaceans, molluscs, leeches and insects were among the most severely affected groups. Among fishes, species of minnows (Cyprindae) were depleted in the most heavily acidified regions, with some declines in salmonid and centrarchid species.Predicted damage to individual lakes in all areas was highly variable. In areas receiving highly acidic deposition, 100% of the species in acid-sensitive taxonomic groups were eliminated in some lakes, while damage to other lakes was predicted to be slight.Estimated damage varied from lake to lake within each subregion, based on chemical characteristics. The most heavily damaged lakes in the Adirondacks and Pocono-Catskills have probably lost all species of molluscs, leeches and crustaceans. On the other hand, lakes of the Midwest showed either slight increases or decreases in the richness of predicted biotic communities.The possible ranges of original sulfate concentrations in lakes and the proportion of sulfuric acid in precipitation that liberated base cations from catchments were confined to relatively narrow limits by the model.Contribution from Fourth World Wilderness Congress-Acid Rain Symposium, Denver (Estes Park), Colorado, September 11–18, 1987.     

Geographic Analysis of Species Richness and Community Attributes of Forest Birds from Survey Data in the Mid-Atlantic Integrated Assessment Region

Species richness of local communities is a state variable commonly used in community ecology and conservation biology. Investigation of spatial and temporal variations in richness and identification of factors associated with these variations form a basis for specifying management plans, evaluating these plans, and for testing hypotheses of theoretical interest. However, estimation of species richness is not trivial: species can be missed by investigators during sampling sessions. Sampling artifacts can lead to erroneous conclusions on spatial and temporal variation in species richness. Here we use data from the North American Breeding Bird Survey to estimate parameters describing the state of bird communities in the Mid-Atlantic Assessment (MAIA) region: species richness, extinction probability, turnover and relative species richness. We use a recently developed approach to estimation of species richness and related parameters that does not require the assumption that all the species are detected during sampling efforts. The information presented here is intended to visualize the state of bird communities in the MAIA region. We provide information on 1975 and 1990. We also quantified the changes between these years. We summarized and mapped the community attributes at a scale of management interest (watershed units).     

Response of soil invertebrates to disturbance across three resource regions in North Carolina

We evaluated the potential of soil microarthropods and enchytraeid worms to be useful as bioindicators of soil condition in forest, wetland, and agricultural ecosystems over a range of ecoregions. Selected mesofauna and soil characteristics in soil and litter in relatively undisturbed and disturbed examples of each of three ecosystems within each of three land resource regions were monitored over two years. Optimal times of year to sample these organisms as indicators of disturbance were April, May, July and September. No single measure reflected disturbance across all three ecosystems. Among forest sites, Simpson's diversity index, evenness, abundance of ants, and proportion of enchytraeids in the mesofauna differed between soils of different disturbance levels. Among agricultural sites, richness, evenness, abundance of mites, and proportions of collembolans and of enchytraeids in the mesofauna differed between disturbance levels. Among wetland sites, Shannon's and Simpson's diversity indices, richness based on the total mesofauna, and abundances of mites, diplurans, ants, and isotomid and onychiurid collembolans differed between disturbance levels. Covariates most frequently associated with abundance and diversity of the measured mesofauna were soil electrical conductivity, available N, organic matter, and pH. Canonical correspondence analysis provided information somewhat different to bivariate analysis. Using both approaches to examine soil and litter taxa that have distinctive responses to disturbance may help to identify candidate groups applicable for use in large-scale environmental monitoring programs.