910 resultados para Litter


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A multi-disciplinary study was conducted to compare stands of ancient and secondary origin within a single wood, the Gearagh woodland, County Cork. These sites were compared with adjacent areas of grassland, which provided a reference for the former land-use (pasture) of the secondary woodland. A historical study confirmed that while the core of the Gearagh has been subject to minimal human interference, other sections have been cleared in the past for agricultural purposes. Investigations into soil structure and composition showed that soil properties in these secondary woodland areas were significantly altered by this past woodland clearance and conversion to agriculture, while the soil of the ancient woodland showed little signs of disturbance. The vegetation community also differed between the two woodland areas, partly due to altered environmental conditions. Many of the ancient woodland plant species were unable to form a persistent seed bank, while there was increased representation of species associated with more open-habitat conditions in the seed bank of the secondary woodland. While germination of woodland species was low in all sites, overall, seeds tended to germinate more successfully in the ancient woodland. The ancient woodland also provided a suitable habitat for many soil and ground detritivores, most notably enchytraeids, although earthworms were not abundant. Past agricultural use, however, changed the decomposer community considerably, with increased representation of earthworm species and a decline in the abundance of enchytraeids in the secondary stands. In conclusion, the legacies of historical agricultural activities can continue to significantly affect the structure and composition of present-day woodlands so that they may differ considerably from undisturbed ancient woodland stands, even within the same woodland. A greater understanding of the origin, development and ecological functioning of ancient woodlands should aid in determining future conservation and management requirements.

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As a prominent form of land use across much of upland Europe, extensive livestock grazing may hold the key to the sustainable management of these landscapes. Recent agricultural policy reform, however, has resulted in a decline in upland sheep numbers, prompting concern for the biodiversity value of these areas. This study quantifies the effects of varying levels of grazing management on plant, ground beetle and breeding bird diversity and assemblage in the uplands and lowlands of hill sheep farms in County Kerry, Ireland. Farms represent a continuum of light to heavy grazing, measured using a series of field indicators across several habitats, such as the internationally important blanket bog, home to the ground beetle, Carabus clatratus. Linear mixed effects modelling and non-metric multidimensional scaling are employed to disentangle the most influential management and environmental factors. Grazing state may be determined by the presence of Molinia caerulea or Nardus stricta, and variables such as % traditional ewes, % vegetation litter and % scrub prove valuable indicators of diversity. Measures of ecosystem functioning, e.g. plant biomass (nutrient cycling) and % vegetation cover (erosion rates) are influenced by plant diversity, which is influenced by grazing management. Levels of the ecosystem service, soil organic carbon, vary with ground beetle abundance and diversity, potentially influencing carbon sequestration and thereby climate change. The majority of species from all three taxa are found in the lowlands, with the exception of birds such as meadow pipit and skylark. The scale of measurement should be determined by the size and mobility of the species in question. The challenge is to manage these high nature value landscapes using agri-environment schemes which enhance biodiversity by maintaining structural heterogeneity across a range of scales, altitudes and habitats whilst integrating the decisions of people living and working in these marginal areas.

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Coastal lagoons are defined as shallow coastal water bodies partially separated from the adjacent sea by a restrictive barrier. Coastal lagoons are protected under Annex I of the European Habitats Directive (92/43/EEC). Lagoons are also considered to be “transitional water bodies” and are therefore included in the “register of protected areas” under the Water Framework Directive (2000/60/EC). Consequently, EU member states are required to establish monitoring plans and to regularly report on lagoon condition and conservation status. Irish lagoons are considered relatively rare and unusual because of their North Atlantic, macrotidal location on high energy coastlines and have received little attention. This work aimed to assess the physicochemical and ecological status of three lagoons, Cuskinny, Farranamanagh and Toormore, on the southwest coast of Ireland. Baseline salinity, nutrient and biological conditions were determined in order to provide reference conditions to detect perturbations, and to inform future maintenance of ecosystem health. Accumulation of organic matter is an increasing pressure in coastal lagoon habitats worldwide, often compounding existing eutrophication problems. This research also aimed to investigate the in situ decomposition process in a lagoon habitat together with exploring the associated invertebrate assemblages. Re-classification of the lagoons, under the guidelines of the Venice system for the classifications of marine waters according to salinity, was completed by taking spatial and temporal changes in salinity regimes into consideration. Based on the results of this study, Cuskinny, Farranamanagh and Toormore lagoons are now classified as mesohaline (5 ppt – 18 ppt), oligohaline (0.5 ppt – 5 ppt) and polyhaline (18 ppt – 30 ppt), respectively. Varying vertical, longitudinal and transverse salinity patterns were observed in the three lagoons. Strong correlations between salinity and cumulative rainfall highlighted the important role of precipitation in controlling the lagoon environment. Maximum effect of precipitation on the salinity of the lagoon was observed between four and fourteen days later depending on catchment area geology, indicating the uniqueness of each lagoon system. Seasonal nutrient patterns were evident in the lagoons. Nutrient concentrations were found to be reflective of the catchment area and the magnitude of the freshwater inflow. Assessment based on the Redfield molar ratio indicated a trend towards phosphorus, rather than nitrogen, limitation in Irish lagoons. Investigation of the decomposition process in Cuskinny Lagoon revealed that greatest biomass loss occurred in the winter season. Lowest biomass loss occurred in spring, possibly due to the high density of invertebrates feeding on the thick microbial layer rather than the decomposing litter. It has been reported that the decomposition of plant biomass is highest in the preferential distribution area of the plant species; however, no similar trend was observed in this study with the most active zones of decomposition varying spatially throughout the seasons. Macroinvertebrate analysis revealed low species diversity but high abundance, indicating the dominance of a small number of species. Invertebrate assemblages within the lagoon varied significantly from communities in the adjacent freshwater or marine environments. Although carried out in coastal lagoons on the southwest coast of Ireland, it is envisaged that the overall findings of this study have relevance throughout the entire island of Ireland and possibly to many North Atlantic coastal lagoon ecosystems elsewhere.

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After becoming competent for resuming meiosis, fully developed mammalian oocytes are maintained arrested in prophase I until ovulation is triggered by the luteotropin surge. Meiotic pause has been shown to depend critically on maintenance of cAMP level in the oocyte and was recently attributed to the constitutive Gs (the heterotrimeric GTP-binding protein that activates adenylyl cyclase) signaling activity of the G protein-coupled receptor GPR3. Here we show that mice deficient for Gpr3 are unexpectedly fertile but display progressive reduction in litter size despite stable age-independent alteration of meiotic pause. Detailed analysis of the phenotype confirms premature resumption of meiosis, in vivo, in about one-third of antral follicles from Gpr3-/- females, independently of their age. In contrast, in aging mice, absence of GPR3 leads to severe reduction of fertility, which manifests by production of an increasing number of nondeveloping early embryos upon spontaneous ovulation and massive amounts of fragmented oocytes after superovulation. Severe worsening of the phenotype in older animals points to an additional role of GPR3 related to protection (or rescue) of oocytes from aging. Gpr3-defective mice may constitute a relevant model of premature ovarian failure due to early oocyte aging.

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Intersex in largemouth bass (Micropterus salmoides) has been correlated with regional anthropogenic activity, but has not been causally linked to environmental factors. Four groups of hatchery-reared largemouth bass (LMB) and fathead minnows (FHM) of varying ages and sex were exposed to aqueous poultry litter mixtures, 17β- estradiol (E2), and controls. Water samples were analyzed for estrogens through liquid chromatography tandem mass spectrometry and estrogenicity through the bioluminescent yeast estrogen screen assay. Fish plasma was analyzed for the egg yolk protein vitellogenin (Vtg) using enzyme–linked immunosorbent assay and gonad tissue was examined histologically for enumeration of testicular oocytes (TO). Water chemistry revealed typical E2 conversion to Estrone with subsequent decay over the exposure periods. A modest prevalence of TO (9.4%) was detected with no apparent treatment effect. While significant Vtg induction was found in E2 exposed FHM, minimal Vtg induction was found in male LMB. Despite field findings of intersex in male LMB, this species may be poorly suited for laboratory investigations into endocrine disruption.

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Microplastic litter is a pervasive pollutant present in aquatic systems across the globe. A range of marine organisms have the capacity to ingest microplastics, resulting in adverse health effects. Developing methods to accurately quantify microplastics in productive marine waters, and those internalized by marine organisms, is of growing importance. Here we investigate the efficacy of using acid, alkaline and enzymatic digestion techniques in mineralizing biological material from marine surface trawls to reveal any microplastics present. Our optimized enzymatic protocol can digest >97% (by weight) of the material present in plankton-rich seawater samples without destroying any microplastic debris present. In applying the method to replicate marine samples from the western English Channel, we identified 0.27 microplastics m−3. The protocol was further used to extract microplastics ingested by marine zooplankton under laboratory conditions. Our findings illustrate that enzymatic digestion can aid the detection of microplastic debris within seawater samples and marine biota.

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Human activities within the marine environment give rise to a number of pressures on seabed habitats. Improved understanding of the sensitivity of subtidal sedimentary habitats is required to underpin the management advice provided for Marine Protected Areas, as well as supporting other UK marine monitoring and assessment work. The sensitivity of marine sedimentary habitats to a range of pressures induced by human activities has previously been systematically assessed using approaches based on expert judgement for Defra Project MB0102 (Tillin et al. 2010). This previous work assessed sensitivity at the level of the broadscale habitat and therefore the scores were typically expressed as a range due to underlying variation in the sensitivity of the constituent biotopes. The objective of this project was to reduce the uncertainty around identifying the sensitivity of selected subtidal sedimentary habitats by assessing sensitivity, at a finer scale and incorporating information on the biological assemblage, for 33 Level 5 circalittoral and offshore biotopes taken from the Marine Habitat Classification of Britain and Ireland (Connor et al. 2004). Two Level 6 sub-biotopes were also included in this project as these contain distinctive characterising species that differentiate them from the Level 5 parent biotope. Littoral, infralittoral, reduced and variable salinity sedimentary habitats were excluded from this project as the scope was set for assessment of circalittoral and offshore sedimentary communities. This project consisted of three Phases. • Phase 1 - define ecological groups based on similarities in the sensitivity of characterising species from the Level 5 and two Level 6 biotopes described above. • Phase 2 - produce a literature review of information on the resilience and resistance of characterising species of the ecological groups to pressures associated with activities in the marine environment. • Phase 3 - to produce sensitivity assessment ‘proformas’ based on the findings of Phase 2 for each ecological group. This report outlines results of Phase 2. The Tillin et al., (2010) sensitivity assessment methodology was modified to use the best available scientific evidence that could be collated within the project timescale. An extensive literature review was compiled, for peer reviewed and grey literature, to examine current understanding about the effects of pressures from human activities on circalittoral and offshore sedimentary communities in UK continental shelf waters, together with information on factors that contribute to resilience (recovery) of marine species. This review formed the basis of an assessment of the sensitivity of the 16 ecological groups identified in Phase 1 of the project (Tillin & Tyler-Walters 2014). As a result: • the state of knowledge on the effects of each pressure on circalittoral and offshore benthos was reviewed; • the resistance, resilience and, hence, sensitivity of sixteen ecological groups, representing 96 characteristic species, were assessed for eight separate pressures; • each assessment was accompanied by a detailed review of the relevant evidence; Assessing the sensitivity of subtidal sedimentary habitats to pressures associated with human activities • knowledge gaps and sources of uncertainty were identified for each group; • each assessment was accompanied by an assessment of the quality of the evidence, its applicability to the assessment and the degree of concordance (agreement) between the evidence, to highlight sources of uncertainty as an assessment of the overall confidence in the sensitivity assessment, and finally • limitations in the methodology and the application of sensitivity assessments were outlined. This process demonstrated that the ecological groups identified in Phase 1 (Tillin & Tyler-Walters 2014) were viable groups for sensitivity assessment, and could be used to represent the 33 circalittoral and offshore sediments biotopes identified at the beginning of the project. The results of the sensitivity assessments show: • the majority of species and hence ecological groups in sedimentary habitats are sensitive to physical change, especially loss of habitat and sediment extraction, and change in sediment type; • most sedimentary species are sensitive to physical damage, e.g. abrasion and penetration, although deep burrowing species (e.g. the Dublin Bay prawn - Nephrops norvegicus and the sea cucumber - Neopentadactyla mixta) are able to avoid damaging effects to varying degrees, depending on the depth of penetration and time of year; • changes in hydrography (wave climate, tidal streams and currents) can significantly affect sedimentary communities, depending on whether they are dominated by deposit, infaunal feeders or suspension feeders, and dependant on the nature of the sediment, which is itself modified by hydrography and depth; • sedentary species and ecological groups that dominate the top-layer of the sediment (either shallow burrowing or epifaunal) remain the most sensitive to physical damage; • mobile species (e.g. interstitial and burrowing amphipods, and perhaps cumaceans) are the least sensitive to physical change or damage, and hydrological change as they are already adapted to unstable, mobile substrata; • sensitivity to changes in organic enrichment and hence oxygen levels, is variable between species and ecological groups, depending on the exact habitat preferences of the species in question, although most species have at least a medium sensitivity to acute deoxygenation; • there is considerable evidence on the effects of bottom-contact fishing practices and aggregate dredging on sedimentary communities, although not all evidence is directly applicable to every ecological group; • there is lack of detailed information on the physiological tolerances (e.g. to oxygenation, salinity, and temperature), habitat preferences, life history and population dynamics of many species, so that inferences has been made from related species, families, or even the same phylum; • there was inadequate evidence to assess the effects of non-indigenous species on most ecological groups, and Assessing the sensitivity of subtidal sedimentary habitats to pressures associated with human activities • there was inadequate evidence to assess the effects of electromagnetic fields and litter on any ecological group. The resultant report provides an up-to-date review of current knowledge about the effects of pressures resulting from human activities of circalittoral and offshore sedimentary communities. It provides an evidence base to facilitate and support the provision of management advice for Marine Protected Areas, development of UK marine monitoring and assessment, and conservation advice to offshore marine industries. However, such a review will require at least annual updates to take advantage of new evidence and new research as it becomes available. Also further work is required to test how ecological group assessments are best combined in practice to advise on the sensitivity of a range of sedimentary biotopes, including the 33 that were originally examined.

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Plastic debris is now ubiquitous in the marine environment affecting a wide range of taxa, from microscopic zooplankton to large vertebrates. Its persistence and dispersal throughout marine ecosystems has meant that sensitivity toward the scale of threat is growing, particularly for species of conservation concern, such as marine turtles. Their use of a variety of habitats, migratory behaviour, and complex life histories leave them subject to a host of anthropogenic stressors, including exposure to marine plastic pollution. Here, we review the evidence for the effects of plastic debris on turtles and their habitats, highlight knowledge gaps, and make recommendations for future research. We found that, of the seven species, all are known to ingest or become entangled in marine debris. Ingestion can cause intestinal blockage and internal injury, dietary dilution, malnutrition, and increased buoyancy which in turn can result in poor health, reduced growth rates and reproductive output, or death. Entanglement in plastic debris (including ghost fishing gear) is known to cause lacerations, increased drag—which reduces the ability to forage effectively or escape threats—and may lead to drowning or death by starvation. In addition, plastic pollution may impact key turtle habitats. In particular, its presence on nesting beaches may alter nest properties by affecting temperature and sediment permeability. This could influence hatchling sex ratios and reproductive success, resulting in population level implications. Additionally, beach litter may entangle nesting females or emerging hatchlings. Lastly, as an omnipresent and widespread pollutant, plastic debris may cause wider ecosystem effects which result in loss of productivity and implications for trophic interactions. By compiling and presenting this evidence, we demonstrate that urgent action is required to better understand this issue and its effects on marine turtles, so that appropriate and effective mitigation policies can be developed.

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During twao years soil and litter pH of 31 permanent plots grown by several rock-rose scrubs (jarales) of Cistion laurifolii at different dynamic stages have been measured. Acidily records show the existence of important seasonal variations and according to a certain rythm. The different syntaxa are characterized as for this ecological factor and the results for these matorrals are compared with other data of heath-scrubs. Adult jaral-phases tend to decrease soil pH when compared with younger dwarf-scrubs-phases, nevertheless upper soil levels generally remain less acidic. Multivariable analysis do not show preference of t he different syntaxa for a certain soil-pH range.

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We present descriptions of a new order (Ranunculo cortusifolii-Geranietalia reuteri and of a new alliance (Stachyo lusitanicae-Cheirolophion sempervirentis) for the herbaceous fringe communities of Macaronesia and of the southwestern Iberian Peninsula, respectively. A new alliance, the Polygalo mediterraneae-Bromion erecti (mesophilous post-cultural grasslands), was introduced for the Peninsular Italy. We further validate and typify the Armerietalia rumelicae (perennial grasslands supported by nutrient-poor on siliceous bedrocks at altitudes characterized by the submediterranean climate of central-southern Balkan Peninsula), the Securigero-Dasypyrion villosae (lawn and fallow-land tall-grass annual vegetation of Italy), and the Cirsio vallis-demoni-Nardion (acidophilous grasslands on siliceous substrates of the Southern Italy). Nomenclatural issues (validity, legitimacy, synonymy, formal corrections) have been discussed and clarified for the following names: Brachypodio-Brometalia, Bromo pannonici-Festucion csikhegyensis, Corynephoro-Plantaginion radicatae, Heleochloion, Hieracio-Plantaginion radicatae, Nardetea strictae, Nardetalia strictae, Nardo-Callunetea, Nardo-Galion saxatilis, Oligo-Bromion, Paspalo-Heleochloetalia, Plantagini-Corynephorion and Scorzoneret alia villosae. 

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Climate change is occurring most rapidly in the Arctic where warming has been twice as fast as the rest of the globe over the last few decades. Arctic soils contain a vast store of carbon and warmer arctic soils may mediate current atmospheric CO2 concentrations and global warming trends. Warmer soils could increase nutrient availability to plants, leading to increased primary production and sequestration of CO2. Presumably because of these effects of warming on shrub ecosystems, shrubs have been expanding across the arctic over the last 50 years, Arctic shrub expansion may track or cause changes in nutrient cycling and availability that favour growth of larger, denser shrubs. This study aimed at measuring gross and net nitrogen cycling rates, major soil nitrogen and carbon pool sizes, and elucidating controls on nutrient cycling and availability between a mesic birch (Betula nana) hummock tundra ecosystem and an ecosystem of dense, tall, birch (B. nana) shrubs. Nitrogen cycling and availability was enhanced at the tall shrub ecosystem compared to the birch hummock ecosystem. Net nitrogen immobilization by microbes was approximately threefold greater at the tall shrub ecosystem. This was in part because of larger microbial biomass nitrogen and carbon (interpreted as a larger microbial community) at the tall shrub ecosystem. Nitrogen inputs via litter were significantly larger at the tall shrub ecosystem and were hypothesized to be the major contributor to the higher dissolved organic and inorganic nitrogen pools in the soil at the tall shrub ecosystem. The results from this study suggest a positive feedback mechanism between litter nitrogen inputs and the enhancement of nitrogen cycling and availability as a driver of shrub expansion across the Arctic.

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The present study was undertaken to test whether inhibition of the proangiogenic inflammatory cytokine tumor necrosis factor (TNF)-alpha can modulate retinal hypoxia and preretinal neovascularization in a murine model of oxygen-induced retinopathy (OIR). OIR was produced in TNF-alpha-/- and wild-type (WT) control C57B6 neonatal mice by exposure to 75% oxygen between postnatal days 7 and 12 (P7 to P12). Half of each WT litter was treated with the cytokine inhibitor semapimod (formerly known as CNI-1493) (5 mg/kg) by daily intraperitoneal injection from the time of reintroduction to room air at P12 until P17. The extent of preretinal neovascularization and intraretinal revascularization was quantified by image analysis of retinal flat-mounts and retinal hypoxia correlated with vascularization by immunofluorescent localization of the hypoxia-sensitive drug pimonidazole (hypoxyprobe, HP). HP adducts were also characterized by Western analysis and quantified by competitive enzyme-linked immunosorbent assay. TNF-alpha-/- and WT mice showed a similar sensitivity to hyperoxia-induced retinal ischemia at P12. At P13 some delay in early reperfusion was evident in TNFalpha-/- and WT mice treated with semapimod. However, at P17 both these groups had significantly better vascular recovery with less ischemic/hypoxic retina and preretinal neovascularization compared to untreated retinopathy in WT mice. Immunohistochemistry showed deposition of HP in the avascular inner retina but not in areas underlying preretinal neovascularization, indicating that such aberrant vasculature can reduce retinal hypoxia. Inhibition of TNF-alpha significantly, improves vascular recovery within ischemic tissue and reduces pathological neovascularization in OIR. HP provides a useful tool for mapping and quantifying tissue hypoxia in experimental ischemic retinopathy.

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Species may become obligate cooperative breeders when parents are unable to raise their offspring unassisted. We measured the daily energy expenditure of mothers, helpers and offspring during peak lactation in cooperatively breeding meerkats Suricata suricatta using the doubly labelled water technique. Lactating mothers expended more energy per day than allo-lactating subordinate females, non-lactating females or suckling offspring. Metabolizable energy intakes of lactating mothers were calculated from isotope-based estimates of offspring milk energy intake, and were not significantly different from the previously suggested maximal limit for mammals. Allo-lactating females were the only category of animals that lost weight during the period of study, probably because they spent more time babysitting than non-lactating females. Daily energy expenditure (DEE) of lactating mothers increased with litter size but decreased with the number of helpers. Calculations show that for every 10 helpers, even in the absence of allo-lactators, mothers are able to reduce their DEE during peak lactation by an amount equivalent to the energy cost of one pup. These results indicate that helpers have beneficial energetic consequences for lactating mothers in an obligate cooperatively breeding mammal.

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Biological activities greatly influence the formation of many soils, especially forest soils under cool humid climates. The objective of this study was to investigate the effects of vegetation and soil biota on the formation of selected soils. Field morphology, micromorphology, and carbon and organic matter analysis were determined on six Podzols (Spodosols) and two Cambisols (Inceptisols) from the eastern United States and north-east Scotland. Humification of plant material by soil fauna and fungi occurs in all organic horizons. Thick organic coatings are observed on soil peds and rock fragments from the E1 to the Bs horizon in a Haplic Podzol from Clingmans Dome Mt., TN. Thin sections reveal large accumulations of root material in different stages of decomposition in the spodic horizons of a Haplic Podzol from Whiteface Mt., NY. Organic carbon ranges from 5.4 to 8.5% in the spodic B horizons of the Whiteface Mt. Podzol. Earthworms and enchytraeids have a great effect on the structure of the surface and subsurface horizons in the Dystric Cambisols from Huntly and Clashindarroch Forests, Scotland and a Cambic Podzol from the Corrie Burn Basin, Scotland. Podzols from Speymouth Forest, Scotland (Gleyic Podzol), Cling-mans Dome Mt., and Whiteface Mt. have thick organic horizons. The Podzols from the Flatwoods in Georgia, the Pine Barrens in New Jersey, the Corrie Burn Basin, and the Cambisol from Huntly Forest have only A horizons at the surface. The Clashindarroch Forest soil has a very thin organic horizon. Warm and humid climates and sandy parent material are responsible for thick E horizons and lack of thick organic horizons in the Flatwoods (Carbic Podzol) and Pine Barrens (Ferric Podzol) soils. Earthworms and enchytraeids thrive in the Corrie Burn Basin and Huntly Forest soils due to the vegetation and the highly weathered basic parent material. The site at Clashindarroch once carried oak, and then birch forest, both of which produce a mild litter and also encourage earthworm and enchytraeids. This fauna is responsible for much mixing of the topsoil. The present conifer vegetation will eventually produce a deep litter and cause podzolization.

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The reduction of forest floor ground cover and litter layers by prescribed fires may alter the morphology (field and micro) and physical properties of surface horizons. This study determined long-term (35 yr) changes in surface horizon bulk density, organic matter concentration and content, and morphology in response to periodic (5 yr) and annual (1 yr) prescribed fires. Soils were fine-silty, siliceous, thermic Glossic Fragiuldults, supporting mixed oak vegetation in middle Tennessee. Upper mineral soils (0- to 2-cm and 0- to 7.6-cm depths) were sampled and detailed field descriptions made. Periodic and control plots had a thin layer of Oi, Oe, and Oa horizons 5 yr after the 1993 burn, whereas on annual burn plots a 1- to 2-cm charred layer was present. Significant reductions in organic matter concentration and mean thickness of the A horizon were found from burning (A horizons thicknesses were 6.4, 4.6, and 2.9 cm in control, periodic, and annual plots, respectively). Periodic burns did not significantly alter the organic matter and bulk density of the upper 7.6 cm of mineral soil; however, annual burns did result in significantly higher bulk densities (1.01, 1.07, and 1.29 Mg m-3 in control, periodic, and annual plots, respectively) and lower organic matter concentrations and contents. Microscopic investigations confirmed that compaction was increased from annual burning. Thin sections also revealed that the granular structure of the A horizons in control and periodic plots resulted from bioterbation of macro and mesofauna, fungi, and roots. Long-term annual burning greatly affected surface soil properties, whereas periodic burning on a 5-yr cycle had only limited effects.