Prescribed burning effects on soil physical properties and soil water repellency in a steep chaparral watershed, southern California, USA

Chaparral watersheds associated with Mediterranean-type climate are distributed over five regions of the world. Because brushland soils are often shallow with low water holding capacities, and are on slopes prone to erosion, disturbances such as fire can adversely affect their physical properties. Fire can also increase the spatial coverage of soil water repellency, reducing infiltration, and, in turn, increasing overland flow and subsequent erosion. We studied the impacts of fire on soil properties by collecting data before and after a prescribed burn conducted during Spring 2001 on the San Dimas Experimental Forest, southern California. The fire removed the litter layer and destroyed the weak surface soil structure; leaving a thin band of ash and char on top of, and mixed in with, an unstable, granular soil of loose consistency. Median litter thickness and clay content were significantly decreased after fire while soil bulk density increased. At 7 d post-fire, soil surface repellency in the watershed was significantly higher than prior to the burn. At 76 d post-fire, surface soil water repellency was returning to near pre-fire values. At the 2 and 4 cm depths, 7 d post-fire soil repellency was also significantly higher than pre-fire, however, conditions at 76 d post-fire were similar to pre-fire values. Variability in soil water repellency between replicates within a given 15 × 15 cm site was as large as the variability seen between sites over the 1.28 ha watershed. The increase in post-fire persistence of water repellency was largest beneath ceanothus (Ceanothus crassifolius) as compared to a small increase beneath chamise (Adenostoma fasciculatum). However, pre-fire persistence was higher under chamise than for ceanothus. Post-fire changes to soil properties may increase the watershed hydrologic response, however the mosaic distribution of water repellency may lead to a less severe increase in hydrologic response than might be expected for a spatially more homogenous increase in repellency.     

Effects of chaparral conversion on small vertebrates in Southern California

Lizard and small mammal faunas were sampled on eight sites of chaparral and artificial chaparral-grass conversions in southern California. The abundance and diversity of these faunas were characteristically greater in chaparral than in areas of grass. The deer mouse (Peromyscus maniculatus) was the only animal studied that was abundant on both chaparral and grass sites. In early stages of chaparral-grass conversion, mechanical brush clearing (Bulldozing, discing)_appeared to cause greater initial reductions in small vertebrate populations than did the use of selective herbicides (primarily 2,4-D). Both lizards and small mammals were virtually absent from grass that was heavily grazed. Lizards reached peak abundance in recently burned (relatively open) chaparral.     

Mirex incorporation in the environment: Toxicity in two soil macroarthropods and effects on soil community respiration

The KD (knockdown dosage) and LD (lethal dosage) values of mirex were determined for the land isopod Armadillidium vulgare and the soil millipede Oxidus gracilis by feeding with a diet containing different concentrations of technical mirex powder. KD₅₀ and LD₅₀ values for A. vulgare at 10 days exposure were 11.6 ppm and 35.2 ppm, respectively; and for O. gracilis, 5.4 ppm and 198.7 ppm, respectively. The isopod A. vulgare was less susceptible to mirex poisoning as symptomized by knockdown behavior but was killed sooner than the millipede O. gracilis. Poisoned animals showed increased O₂ uptake after two to three days of feeding on mirex diet presumably due to increased muscular activity as induced by elevated oxidative metabolism. Technical mirex showed a significant effect on soil community respiration at 100 ppm concentration after 24 h but no change was observed at the higher concentrations of 500 and 1000 ppm. The addition of mirex as fire ant baits at the rate of 1:100, 1 : 1000, and I :10 000 bait to soil ratio also did not affect soil community respiration, but 02 uptake was nevertheless elevated due to the microbial population (bacteria and fungi) that immediately invaded the bait material.     

The influence of earthworms on the mobility of microelements in soil and their availability for plants

The influence of earthworms (Aporrectodea caliginosa, Lumbricus rubellus, L. terrestris, and Eisenia fetida) on the mobility of microelements and their availability for plants was studied. The contents of water-soluble Fe and Mn compounds extracted from the coprolites were 5–10 times higher than that in the soil (enriched in calcium carbonate and dried) consumed by the earthworms. This digestion-induced effect became higher with the age of the coprolites (up to 9 days) and took place under their alkalization. In the excreta (surface + enteric) of earthworms, the Fe concentration exceeded those of Mn and Zn by many times. Iron and manganese were mostly concentrated (>80% and >60%, respectively) in the organic part of the excrements. In the tests with hydroponics, the excreta were found to be a source of iron compounds available for plants that were similar to Fe₂(SO₄)₃ or Fe-citrate by their physiological effect in the case when the Fe concentration in the excretions was above 0.7 μM. However, the single application of excreta of different earthworm species into the CaCO₃ enriched soil did not significantly affect the plant (cucumber) nutrition. The analysis of the transport of microelements with xylem sap showed that this fact appeared to be due to the absence of an Fe deficit in the cucumber plants because of their high capability for the absorption of weakly soluble iron compounds.     

The influence of soil compaction and the removal of organic matter on two native earthworms and soil properties in an oak-hickory forest

 Earthworms may alter the physical, chemical, and biological properties of a forest soil ecosystem. Any physical manipulation of the soil ecosystem may, in turn, affect the activities and ecology of earthworms. The effects of removing organic matter (logs and forest litter) and severely compacting the soil on native earthworm species were measured in a central USA hardwood region (oak-hickory) forest in the Missouri Ozarks (USA). Soils in this region are characterized by a cherty residuum that is primarily of the Clarksville series (Loamy-skeletal, mixed, mesic Typic Paledults). Earthworms were collected from 0–15 cm depth each spring and fall for 2 years by handsorting, and densities were determined on a per meter square basis. Two native earthworm species, Diplocardia ornata and Diplocardia smithii, were dominant on this site. Organic matter removal decreased the average individual biomass of both species. However, both species responded differently to soil compaction. Soil compaction affected D. ornata adversely and D. smithii favorably. This suggested that the degree of soil compaction was not as restrictive with respect to D. smithii (2 mm diameter) as to D. ornata (5 mm diameter). Moreover, the apparently improved soil environmental conditions resulting from the remaining organic matter in compacted soil enhanced the population and growth of D. smithii. Sampling position on the landscape affected D. ornata but not D. smithii. Soil microbial biomass C and soil microbial biomass N were decreased under soil compaction when the organic matter was removed. Other factors influencing the ecology and activity of these two species will require further study. Received: 6 January 1999     

Leaf litter decomposition in a chaparral ecosystem, Southern California

Decomposition losses from leaves of three evergreen chaparral species, scrub oak (Quercus dumosa), ceanothus (Ceanothus crassifolius), and manzanita (Arctostaphylos glauca), were quantified over a 2-y field exposure using litterbags. Changes in ash-free dry mass, C, and N were monitored at 2- to 6-month intervals at four replicate sites composed of patches of these three chaparral species. Three proximate C fractions were extracted from fresh and decomposing litter samples: polar and non-polar extractives (EXT), acid-solubles (ACID), and acid-insolubles (KLIG). The chemical structure of fresh and decomposed litter was additionally characterized using high-resolution solid-state ¹³C NMR spectroscopy, while morphological properties were examined by scanning electron microscopy (SEM). After 2 y, the litters had lost between 20.7%±1.2 (Ceanothus) and 35.2%±6.8 (Quercus) of their original ash-free dry mass. The manzanita decomposed at a significantly faster rate than the other two litter types during the first few months of field exposure. Yet, after 2 y, mass loss was greater for the oak. Differences in decomposition rates could not be accounted for based on a single litter quality index. Fresh manzanita exhibited a significantly higher N content, which could explain its initially faster decay rate. Fresh oak litter, on the other hand, had a relatively high ACID and O-alkyl C (O-ALK) content, which may have been responsible for its decay pattern. Fresh ceanothus contained a relatively low KLIG content, yet it decomposed more slowly than the two other species. The solid-state ¹³C NMR spectra of the ceanothus litter had two peaks characteristic of proanthocyanidins, which likely contributed to the recalcitrance of this litter type. SEM revealed that ceanothus leaf surfaces were left nearly unchanged after field exposure. In comparison, the oak and manzanita leaf surfaces were pitted and covered by microbial growth to the point of being unrecognizable. Taken together, our results indicate that a combination of biological, physical and chemical factors need to be examined to clarify the different decomposition rates and patterns of these three chaparral species.     

Endogeic earthworms differentially influence bacterial communities associated with different soil aggregate size fractions

Endogeic earthworm activities can strongly influence soil structure. Although soil microorganisms are thought to be central to earthworm-facilitated aggregate formation, how and where within the soil matrix earthworm-facilitated influences on soil microbial communities are manifested is poorly defined. In this study we used 16S rRNA gene-based terminal restriction fragment polymorphism (T-RFLP) analyses to examine bacterial communities associated with different aggregate size fractions (macroaggregates, microaggregates-within-macroaggregates and inner-microaggregates-within-macroaggregates) of soils incubated for 28 d with and without earthworms. We hypothesized that bacterial communities in different soil aggregate size fractions are differentially influenced by earthworm activities. Our results indicate significantly enhanced aggregate formation (both macroaggregates and microaggregates within macroaggregates) in earthworm-worked soils relative to soils receiving only plant litter. Although significant differences were found between bacterial communities of earthworm and litter-only treatments for all soil fractions, communities associated with earthworm-worked macroaggregate fractions exhibited the least similarity to all other soil fractions regardless of treatment. In addition to differences in terminal restriction fragment (T-RF) size distributions, T-RFLP profiles of earthworm-worked soil macroaggregates had significantly fewer T-RF sizes, further suggesting less species evenness and more extensive alteration of bacterial communities within this fraction. These findings suggest that, due to rapid occlusion of organic materials, microbial communities associated with microaggregates-within-macroaggregates formed during or shortly after passage through the earthworm gut are relatively inactive, and therefore change relatively little over time compared to macroaggregate populations as a whole.     

Optimum temperatures for soil respiration along a semi-arid elevation gradient in southern California

Soil respiration (R) has not been adequately studied at temperatures above 35 °C, which are common temperatures for soils in the southwestern United States and may be important for C dynamics in semi-arid regions. While frequently excluded from ecosystem models or set to 35 °C, the optimum temperature for soil R is poorly understood. Optimum temperatures are likely controlled by substrate availability, soil moisture content, and previous climate. To quantify the optimal temperature for soil R and hypothesized relationships, we collected soils from beneath and between plant canopies at three sites along a semi-arid elevation gradient. Processed soil samples were incubated at three soil moisture contents and soil R was measured at 6 temperatures, successively (25–55 °C). From these data, an activation energy for reaction kinetics and deactivation energy for enzyme functionality model was used to generate soil R curves from which two parameters were derived: R_max, the maximum rate of soil R and T_opt, the optimum temperature for soil R. R_max was significantly greater for soils at the highest elevation and at medium soil moisture content. T_opt was greater than 35 °C at all locations. In addition, T_opt was significantly greater for soils with greater amounts of SOM but not significantly different along the elevation gradient or at different moisture contents. These results support inclusion of much higher optimum temperatures than currently used in many ecosystem and land-surface models and provide support for explaining variation in T_opt as regulated by substrate quantity within a site and general insensitivity across climate differences.     

Effects of endogeic earthworms on soil processes and plant growth in coniferous forest soil

Summary The effects of the endogeic earthworm, Aporrectodea caliginosa tuberculata (Eisen) on decomposition processes in moist coniferous forest soil were studied in the laboratory. The pH preference of this species and its effects on microbial activity, N and P mineralization, and the growth of birch seedlings were determined in separate pot experiments. Homogenized humus from a spruce stand was shown to be too acid for A. c. tuberculata. After liming, the earthworms thrived in the humus and their biomass increased (at pH above 4.8). In later experiments in which the humus was limed, the earthworms positively influenced the biological activity in humus and also increased the rate of N mineralization. A. c. tuberculata increased the growth of birch seedlings, with increases observed in stems, leaves, and roots. Neither NH ₄ ⁺ -N fertilizer nor mechanical mixing with artificial worms affected seedling growth. No plant-growth-affecting compounds (e.g., hormone-like compounds) due to the earthworms were present in the humus. The shoot: root ratio in the birch seedlings was not affected by either the earthworms or the fertilizer. The experiments revealed the impact of earthworm activity on soil processes and plant growth.     

Interactions between earthworms and their soil environment

Laboratory experiments were used to study the effect of food quantity and quality on the biomass of earthworms, and the influence of earthworms on plant growth and infiltration of water into soil. Earthworms with the most food gained weight faster than those with little or no supplementary food. The latter also failed to become reproductively mature. Earthworms lost weight on a nitrogenpoor diet, but this was not rectified by supplementing such food with inorganic nitrogen added to the soil 2 weeks before the worms. Ryegrass grown in soil in which earthworms (Allolobophora trapezoides) had been kept grew more slowly than in soil which had no previous worm activity, perhaps indicating that earthworms had converted relatively-available organic N into less available forms.Microscolex dubius gave the fastest infiltration rates of water into soil when clover mulch was present. With Eisenia foetida there was little effect of worm density on infiltration rates; the highest density significantly increased infiltration but only when clover hay had been mixed in the soil. The surface casting behaviour of the two earthworm species varied with the placing of the food offered.     

Soil moisture affects survival of microorganisms in heated chaparral soil

Prescribed burning, the planned application of fire to reduce the hazards of wildland fuels, is coming into wider use in southern California chaparral. Soil was heated to various temperatures in factorial combination with various soil moistures to determine the direct effects of fire and soil moisture on the survival of microorganisms in chaparral soil. For fungi, mild heating increased germination of dormant forms yielding significantly higher counts than those in unheated soil. With increasing temperatures, microbe populations showed an exponential decrease. For heterotrophic soil bacteria, this decrease was summarized as a function of soil moisture and temperature. Microbial groups differed significantly in sensitivity to temperature: fungi > nitrite oxidizers > heterotrophic bacteria. Physiologically-active populations in moist soil were significantly more sensitive than were dormant populations in dry soil. The mathematical model presented shows qualitatively that more of the microbial biomass will be killed when the soil is moderately moist—as during prescribed burning—than when it is dry. Mineralization of killed microbial biomass in soil and release of plant nutrients may partially explain the increased plant growth and reduced response to fertilizer at burned sites.     

Distribution of earthworms and influence of soil properties across a successional sand dune ecosystem in NW England

An investigation of earthworms across a sand dune system in NW England examined species distribution and abundance with respect to soil physico-chemical conditions and management. Replicated 0.1 m² quadrats were examined every 50 m along 700 m transects across areas of vegetation succession and samples were then taken every 10 m where earthworms were first encountered. A translocation of Aporrectodea longa assessed the ability of this species to live in soils with a high sand content and laboratory work examined growth, maturation and survival of this species in soils with increasing proportions of sand (0–100%). Nine earthworm species were found on the dunes, but none in yellow dunes where organic matter (OM) content was <1%. Dendrobaena octaedra and Lumbricus rubellus were located 300 m from the strand line in grey dunes with an OM content of 3.9%. Allolobophora chlorotica and Lumbricus castaneus occurred within a wet dune slack at 340 m (OM content 11%). In areas of human disturbance (dune car park), A. longa, Aporrectodea caliginosa and Lumbricus terrestris were present, and in soil below pine trees L. castaneus and L. rubellus were present in litter. Laboratory growth of A. longa demonstrated significant (p < 0.05) increases in growth (2–2.75 g) with 25 and 50% sand compared with 0, 75 and 100% over 24 weeks and a greater rate of maturation. Initial results show earthworm colonisation to be influenced by dune successional stage. Laboratory findings suggested A. longa could exist in higher sand content areas but experimental design needs development. The translocation was unsuccessful. Future investigations could examine soil properties more closely and undertake monitoring on site throughout the year.     

Recent trends in beech tree health in southern Britain and the influence of soil type

The health of beech (Fagus sylvatica L.) at sixteen sites of high nature conservation value in the U.K. has been assessed annually since 1987. Survey results show that, from 1987–1992, beech health (as measured by crown thinness) deteriorated markedly at many sites, but a substantial recovery was seen in 1993 and 1994. However, results from a crown architecture assessment indicate that beech health has continued to decline steadily throughout the study period. Tree health is related to soil type, with health being worse on poorly drained, acidic soils. An involvement of climate and perhaps also ozone is suggested given the deterioration in health after the dry summers and high ozone concentrations of 1989 and 1990. These results support earlier findings that climate, in particular the severe drought of 1975/76, had long term effects on canopy development of trees at some of the sites.     

蚯蚓与黑麦草相互作用对土壤中荧蒽去除的影响

Earthworms can promote the bioremediation of contaminated soils through enhancing plant growth and microorganism development. The individual and combined effects of earthworms and ryegrass (Lolium multiflorum Lam.) on the removal of fluoranthene from a sandy-loam alluvial soil were investigated in a 70-d microcosm experiment. The experiment was set up in a complete factorial design with treatments in four replicates: without earthworms or ryegrass (control, CK), with earthworms only (E), with ryegrass only (P), and with both earthworms and ryegrass (EP). The residual fluoranthene, microbial biomass C, and polyphenol oxidase activity in the soil changed significantly (P<0.01) with time. In general, the residual concentration of fluoranthene in the soil decreased sharply from 71.8-88.7 to 31.7-37.4 mg kg 1 in 14 d, and then decreased gradually to 19.7-30.5 mg kg 1 on the 70th d. The fluoranthene concentration left in the soil was the least with both earthworms and ryegrass, compared to the other treatments at the end of the experiment. Half-life times of fluoranthene in the E, P, and EP treatments were 17.8%-36.3% smaller than that of CK. More fluoranthene was absorbed by earthworms than ryegrass. However, the total amounts of fluoranthene accumulated in both the ryegrass and earthworms were small, only accounting for 0.01%-1.20% of the lost fluoranthene. Therefore, we assumed that microbial degradation would play a dominant functional role in fluoranthene removal from soil. We found that earthworms significantly increased microbial biomass C and polyphenol oxidase activity (P<0.01) in the presence of ryegrass at the end of the experiment. Furthermore, microbial biomass C and polyphenol oxidase activity were significantly (P<0.05) and negatively related to the residual fluoranthene concentration. This implied that earthworms might promote the removal of fluoranthene from soil via stimulating microbial biomass C and polyphenol oxidase activity.     

The effect of earthworms and liming on soil microbial communities

The effect of liming and earthworms on the composition and function of soil microbial communities was investigated in an upland soil from the UK in order to understand interactions between the biotic and abiotic components of soil systems. A factorial experiment was established using soils from the Sourhope Farm, near Kelso, with lime or no lime added, with or without earthworms added and a combined treatment of both lime and earthworm additions. The soils were incubated and destructively sampled after 180 days. Measurements of soil microbial biomass, dehydrogenase activity, phenotypic structure (by phospholipid fatty acid analysis (PLFA) and responses to four carbon substrates (d-glucose, l-arginine, α-ketoglutaric acid, α-cyclodextrin) were determined. Statistically significant results were limited to the litter layers, with no significant observations in either the H or Ah horizons. There were significant decreases in the soil microbial biomass and microbial activity in the litter layers caused by the addition of earthworms; liming reduced microbial biomass only. The addition of earthworms caused a significant difference in the PLFA principle component analysis (PCA) profile, as did liming. For the PLFA PCA profile, earthworm plus lime treatment was indistinguishable from the liming result. Addition of earthworms significantly suppressed the response to glucose; this effect was removed by liming. This indicates that liming may significantly alter the ecological interactions between earthworms and the microbial community.     

Significance of earthworms in stimulating soil microbial activity

 The stimulatory effect of earthworms (Lumbricus terrestris L.) on soil microbial activity was studied under microcosm-controlled conditions. The hypothesis was tested that microbial stimulation observed in the presence of a soil invertebrate would be due to the utilization of additional nutritive substances (secretion and excretion products) that it provides. Changes in microbial activity were monitored by measuring simultaneously CO₂ release and protozoan population density. The increase in CO₂ released in the presence of earthworms was found to result from both earthworm respiration and enhanced microbial respiration. The stimulation of microbial activity was confirmed by a significant increase in protozoan population density, which was 3–19 times greater in the presence of earthworms. The respiratory rate of L. terrestris was estimated to be 53 μl O₂ g^–1 h^–1. Earthworm respiration significantly correlated with individual earthworm weight, but there was no correlation between the increase in microbial respiration and earthworm weight. This finding does not support the hypothesis given above that enhanced microbial respiration is due to utilization of earthworm excreta. A new hypothesis that relationships between microbial activity and earthworms are not based on trophic links alone but also on catalytic mechanisms is proposed and discussed. Received: 26 August 1997     

Effects of rhamnolipids on bacterial communities in a dioxin-contaminated soil and the gut of earthworms added to the soil

The biosurfactants rhamnolipids and the “soil ecosystem engineers” earthworms are often used to remediate contaminated soils. However, the effects of rhamnolipids on earthworm intestinal flora and microbial community in soil containing earthworms are not clearly understood. In our study, a 21-d microcosm experiment was carried out to reveal the effects of rhamnolipids on microbial abundance, composition, and metabolism, as well as contaminant degradation capacity. Both rhamnolipids and earthworm...     

Influence of soil ingestion by earthworms on the availability of potassium in soil: An incubation experiment

Summary An incubation experiment was conducted to study the changes that occur in the K status of soil due to earthworm activity. Samples of Tokomaru silt loam soil were inoculated with the common pasture earthworm species Aporrectodea caliginosa and incubated for 21 days. Aliquots of moist soil were analyzed for exchangeable K by leaching with neutral molar ammonium acetate at 1:50 soil solution ratio. Extraction with boiling 1 M nitric acid at 1:100 soil solution ratio for 20 min was used to determine available non-exchangeable K. The results indicated that the exchangeable K content increased significantly due to earthworm activity but nitric acid-extractable K did not change significantly. It is inferred that earthworms increase the availability of K by shifting the equilibrium among the forms of K from relatively un-available forms to more available forms in the soil chosen for the study.     

Bacteria and protozoa in soil microhabitats as affected by earthworms

The effects of incorporation of elm leaves (Ulmus glabra) into an agricultural sandy loam soil by earthworms (Lumbricus festivus) on the bacterial and protozoan populations were investigated. Three model systems consisting of soil, soil with leaves, and soil with leaves and earthworms, respectively, were compared. The total, viable, and culturable number of bacteria, the metabolic potentials of bacterial populations, and the number of protozoa and nematodes were determined in soil size fractions. Significant differences between soil fractions were shown by all assays. The highest number of microorganisms was found in microaggregates of 2–53 μm and the lowest in the <0.2μm fraction. A major part of the bacteria in the latter fraction was viable, but non-culturable, while a relatively higher number of culturable bacteria was found in the macroaggregates. The number of colony-forming units and 5-cyano-2,3-ditolyl tetrazolim chloride (CTC)-reducing bacteria explained a major part of the variation in the number of protozoa. High protozoan activity and predation thus coincided with high bacterial activity. In soil with elm leaves, fungal growth is assumed to inhibit bacterial and protozoan activity. In soil with elm leaves and earthworms, earthworm activity led to increased culturability of bacteria, activity of protozoa, number of nematodes, changed metabolic potentials of the bacteria, and decreased differences in metabolic potentials between bacterial populations in the soil fractions. The effects of earthworms can be mediated by mechanical mixing of the soil constituents and incorporation of organic matter into the soil, but as the earthworms have only consumed a minor part of the soil, priming effects are believed partly to explain the increased microbial activity. Received: 7 January 1996