Role of soil mesofauna in dispersal of Coniothyrium minitans: mechanisms of transmission

Soil mesofauna are thought to play a role in the dispersal of Coniothyrium minitans Campbell but the mechanisms of transmission are unclear. The mite Acarus siro L. and the collembolan Folsomia candida Willem are known to be able to transmit the mycoparasite to uninfected sclerotia of Sclerotinia sclerotiorum (Lib.) de Bary in soil, and consequently the mechanisms of transmission of the mycoparasite by these animals have been investigated in detail. To facilitate this work, a novel method for surface sterilising mites was developed. In an initial investigation, A. siro was found to carry inoculum externally and in the gut. Indeed, almost all faecal pellets collected from either A. siro or F. candida, following feeding on C. minitans, contained germinable inoculum of the mycoparasite. Microscopic examination showed that faeces from these animals consisted mainly of conidia or fragments of conidium, but that only intact conidia gave rise to growth of C. minitans. Passage through the gut of either animal reduced percentage germination of conidia to 11 and 60% in mite and collembolan faeces, respectively, compared to 99% in controls not subjected to gut passage. In addition, germination in mite faeces was delayed by approximately 12 h compared to controls. However, subsequent germ tube extension from surviving conidia appeared unaffected by gut passage. Individual faecal pellets of A. siro were found to contain sufficient inoculum of the mycoparasite to initiate infection of sclerotia in moist sterile sand and non-sterile soil. The relevance of these results to the glasshouse and field situation is discussed.     

Survival of Coniothyrium minitans associated with sclerotia of Sclerotinia sclerotiorum in soil

The development and survival of the mycoparasite Coniothyrium minitans associated with sclerotia of the plant pathogen Sclerotinia sclerotiorum was studied in pasteurised and non-sterile (untreated) soil. Using scanning electron microscopy, developing pycnidia were first seen within the sclerotial medulla at 7 days post-inoculation with the mycoparasite in pasteurised soil. However, by 14 days post-inoculation, pycnidia had developed fully in both pasteurised and non-pasteurised treatments, and conidial droplets were exuded onto the outer surface of the infected sclerotia. Thirty days post-inoculation, irrespective of soil treatment, the majority of the sclerotial medulla had been converted to pycnidia, with the sclerotial rind remaining largely intact. The pycnidia and dried intact droplets were still observed 6 months post-inoculation with C. minitans, although the conidia on the outer surface of the dried droplets had largely collapsed by this stage. Germinability studies at 10 months post-inoculation showed that approximately 13% of the conidia in dried droplets were still viable. This work shows the potential for infected sclerotia of S. sclerotiorum to provide a unique reservoir for the survival of C. minitans.     

Quantitative real-time PCR effectively detects and quantifies colonization of sclerotia of Sclerotinia sclerotiorum by Trichoderma spp.

Some members of the fungal genus Trichoderma are able to colonize and destroy sclerotia, the thick-walled resting structures of the soilborne plant pathogenic fungus Sclerotinia sclerotiorum, thus providing a potential means of biological disease control. However, current methods to detect and quantify colonization of sclerotia are labor-intensive, and generally qualitative rather than quantitative in nature. Our objective was to develop quantitative real-time PCR (polymerase chain reaction) methods to detect and measure colonization of sclerotia by Trichoderma spp. Specific PCR primer/probe sets were developed for Trichoderma spp. and for S. sclerotiorum. A total of 180 ITS1 (internal transcribed spacer) and ITS2 sequences from different species in the genus Trichoderma were aligned, and consensus sequences were determined. Six candidate primer sets were based on conserved regions of the consensus sequence, and the specificity of each nucleotide sequence was examined using BLAST (Basic Local Alignment Search Tool; NCBI) software. Each candidate primer set was tested on genomic DNA of T. harzianum strain ThzID1-M3, as well as six different Trichoderma isolates from soil, and on genomic DNA of S. sclerotiorum. The optimum primer/probe set selected, TGP4, successfully amplified genomic DNA of all Trichoderma isolates tested, and showed high precision and reproducibility over a linear range of eight orders of magnitude, from 85 ng to 8.5 fg of T. harzianum genomic DNA. TGP4 did not amplify S. sclerotiorum DNA. A specific PCR primer/probe set (TMSCL2) was developed for S. sclerotiorum, based on the calmodulin gene sequence. TMSCL2 did not amplify Trichoderma DNA. Quantitative real-time PCR with the primers then was evaluated in experiments to test differential effects of two soil moisture levels (−50 kPa, −1000 kPa matric potential) on biocontrol of S. sclerotiorum by indigenous Trichoderma spp. Periodically over 40 days, Trichoderma and S. sclerotiorum DNA in sclerotia were quantified by PCR with appropriate primers. Over 90% of the sclerotia were colonized by indigenous Trichoderma spp. at −1000 kPa, over the 40-day period, compared to only 60% at −50 kPa. In addition to determining incidence of colonization, the PCR method allowed measurement of the extent of sclerotial colonization, which also was significantly greater in the drier soil. Quantitative real-time PCR with the TGP4 primer/probe set provides a sensitive detection and measurement tool to evaluate colonization of sclerotia by Trichoderma spp.     

Survival of sclerotia of sclerotinia sclerotiorum in soil

Factors affecting longevity of sclerotia in sandy clay loam (s.c.l.) and sandy loam (s.l.) were examined, using sclerotia from a laboratory culture of S. sclerotiorum and from natural infestations on beans and lettuce.Survival of sclerotia from culture and lettuce was compared in s.l. Recovery and viability were less, and incidence of Fusarium, Mucor and Trichoderma spp. greater, in sclerotia from lettuce than from culture. Rinds of sclerotia from lettuce were more perforated than those from culture.Burial of sclerotia at 4 cm for 35 weeks reduced recovery of sclerotia to zero in s.c.l. and by 50% in s.l. At the soil surface recovery was reduced by 55% in s.c.l. and by 10% in s.l. Less than 50% of sclerotia recovered were viable. Neither a chloropicrin-methyl bromide fumigant nor a tomato compost treatment affected recovery or viability. Fumigation increased incidence of Trichoderma spp. and decreased incidence of Fusurium and Mucor spp. isolated from sclerotia.Apothecia were produced over 6 weeks in s.c.l. and over 20 weeks in s.l. Production was increased by the low rate of fumigant in s.c.l. and by tomato compost in s.l.     

Evaluation of different Coniothyrium minitans inoculum sources and application rates on apothecial production and infection of Sclerotinia sclerotiorum sclerotia

The effects of three Coniothyrium minitans isolates (Conio, IVT1 and Contans^®), applied to soil as conidial suspensions or as maizemeal-perlite (MP) inocula (Conio), on apothecial production and infection of Sclerotinia sclerotiorum sclerotia were assessed in two soil pot bioassays and two novel box bioassays in the glasshouse at different times of the year. C. minitans isolate Conio applied as either MP or ground MP at full rate (10⁶-10⁷ cfu cm⁻³ soil) consistently decreased the carpogenic germination, recovery and viability of sclerotia and increased C. minitans infection of the sclerotia of S. sclerotiorum by in comparison with either MP or conidial suspension treatments applied at lower rates (10³-10⁴ cfu cm⁻³ soil). Additionally, when applied at the same rate, MP inoculum of C. minitans was consistently more effective at reducing carpogenic germination than a conidial suspension. The effect of MP and ground MP at full rate on carpogenic germination was expressed relatively early as those sclerotia recovered before apothecia appeared on the soil surface already had reduced numbers of apothecial initials. In general, there were few differences between the isolates of C. minitans applied as conidial suspensions. Box bioassays carried out at different times of the year indicated that temperature and soil moisture influenced both apothecial production and mycoparasitism. Inoculum concentration of C. minitans and time of application appear to be important factors in reducting apothecial production by S. sclerotiorum.     

杀虫剂和除草剂对生防菌盾壳霉生长的影响

本研究采用FAO推荐的菌落直径法测定了8种杀虫剂、4种除草剂、3种植物生长调节剂和1种农用抗菌素对盾壳霉菌丝的毒力。研究结果表明,杀虫剂中氯氰菊酯、阿维菌素、毒死蜱和敌敌畏对盾壳霉生长的抑制作用较大,但除敌敌畏外,其他杀虫剂EC50值均高于相应的田间最大推荐使用浓度(MRAR);辛硫磷和氧乐果对盾壳霉的抑制作用次之;而吡虫啉和灭多威则对其生长几乎没有抑制作用。4种除草剂的EC50值均明显低于相应的MRAR值,对盾壳霉生长有强的抑制作用。在正常使用浓度下植物生长调节剂萘乙酸、赤霉素、云大120对盾壳霉的生长几乎没有抑制作用。链霉素对其生长有一定影响,但EC50明显高于MRAR值。因此,在使用盾壳霉防治菌核病的田中,可以合理使用较安全的植物生长调节剂及吡虫啉和灭多威等杀虫剂,而应尽量减少和避免使用吡氟乙草灵、草甘膦、百草枯等除草剂。     

Reduction of sclerotial inoculum of Sclerotinia sclerotiorum with Coniothyrium minitans

Aspects of the biology of C. minitans and its potential for control of S. sclerotiorum were investigated.Temperatures below 7°C resulted in comparatively slow rates of germination and infection of sclerotia by C. minitans. The optimum temperature for germination, growth, infection of sclerotia, and destructive parasitism by C. minitans was 20°C. The optimum relative humidity for germination, growth and infection by C. minitans was above 95%.Autumn inoculations with suspensions of conidia, pycnidia and mycelium of C. minitans in the field resulted in negligible numbers of sclerotia remaining viable after 1 month. With culture-grown sclerotia 2 months were required for a similar reduction of sclerotial viability. In the absence of C. minitans mulching had no significant effect on sclerotial viability. In the presence of C. minitans mulching did, however, influence the viability and infection by C. minitans of culture-grown sclerotia. Populations of field sclerotia also differed from culture-grown sclerotia in that they harboured an internal population of microorganisms, which included C. minitans, and had a lower level of viability at the commencement of the treatments.A winter application of C. minitans did not result in significant infection of sclerotia nor in a reduction in viability of sclerotia. This failure is believed to have resulted from low temperatures and dry conditions.     

Time and burial depth influencing the viability and bacterial colonization of sclerotia of Sclerotinia sclerotiorum

Sclerotia are the primary over wintering inoculum of Sclerotinia sclerotiorum (Lib.) de Bary. The effects of tillage on the primary inoculum are not well understood. The purpose of this research was to study sclerotial viability over time and between burial depths in soil, to identify bacteria colonizing and degrading the sclerotia, and determine whether these bacteria may be utilized as biological control agents. Correlation analysis indicated that a significant negative relationship existed between sclerotial viability and elapsed temporal factors (R²=−0.68, P<0.0001), and depth of burial (R²=−0.58, P<0.0001). After twelve months, sclerotia on the soil surface had the highest viability (57.5%), followed by those at the 5 cm depth (12.5%), and only 2.5% of those placed at the 10 cm depth remained viable. A significant negative relationship between sclerotial viability and bacterial populations also existed (R²=−0.60, P<0.0001). Two hundred and sixty-eight bacteria were isolated from sclerotia, 29 of which showed strong in vitro antagonism to the mycelial growth of S. sclerotiorum. Biodiversity of the inhibitory bacterial isolates was minimal on sclerotia from the soil surface and within all depths sampled at three months (i.e. in January). All burial depths within the April and July sampling dates produced bacterial diversities that were distinct from each other.     

Diallyl-disulphide to reduce the numbers of sclerotia of Sclerotium cepivorum in soil

A single injection of 0.2 ml diallyl disulphide (DADS) at 0.156% (v/v) into soil containing naturally-produced sclerotia of Sclerotium cepivorum and maintained in the laboratory at 15°C stimulated sclerotial germination and reduced sclerotial numbers by 67%; ungerminated sclerotia remained viable. Higher concentrations of DADS had no additional effect except that at 20% (v/v), germination was slightly inhibited. A similar reduction in sclerotial numbers was obtained when the mixture of soil and sclerotia was exposed to DADS vapour. Four, monthly applications of DADS at 0.2 ml 0.15% (v/v) per application did not give a further reduction.The effect of DADS was temperature dependant, with a reduction in sclerotial numbers of 65 and 9% at 15 and 5°C respectively.     

Ecological and morphological characteristics of the sclerotia of Rhizoctonia solani Kühn produced in soil

Ecological and morphological characteristics of the sclerotia of R. solani produced in soil were compared among Japanese isolates belonging to five anastomosis groups (AG). Twenty-three out of 24 isolates produced sclerotia in soil and the number and size of sclerotia varied considerably among isolates within the same AG. Sclerotia from each group ranged in size from 0.25 mm to more than 2.5mm dia, with most between 0.5–1.0mm dia. Sclerotia formed in soil were smaller than those formed in pure culture. Sclerotia of each AG were tested for viability after 30–270 days incubation in artificially-inoculated soil. Sclerotia produced in soil were brown to dark-brown and these were divided into two distinct groups on external morphology: firm and irregularly globose to subglobose with either a pitted or smooth surface in one group, and rather soft with an indefinite shape in the other. On the basis of internal morphology, the sclerotia fell into three groups. Sclerotia of AG-1 were composed of three well-defined layers; the central cells had dense contents and were surrounded by an outer layer of empty cells which were bordered by a darkly-pigmented mucilaginous surface-layer. Sclerotia of AG-2, Type-1 had only two well-defined layers; The darkly-pigmented mucilaginous layer was absent. Sclerotia of the other AG were constructed of very loosely-arranged brown cells without any well-defined zones.     

Polyethylene mulching of soil to reduce viability of sclerotia of Sclerotium oryzae

Mulching of Sclerotium oryzae infested soil (moist or dry) with polyethylene sheets during hot summer days of May and June increased the soil temperature at 5 cm from 36°C (unmulched) to 48°C (wet) and from 44 to 52°C (dry) and at 20cm from 32 to 38°C (wet) and from 35 to 39°C (dry). In artificially-infested soil, the sclerotia were not eradicated but 95–100% loss in viability was observed at 5 cm by a mulch treatment for 1 week and at 20 cm by mulching for 8 weeks. Mulching effects were not influenced by moisture content of soil or by amendments with lucerne or wheat straw. Mulching of naturally-infested soil at a second site did not eradicate S. oryzae but reduced sclerotial viability by 93%.     

Survival of sclerotia of Sclerotium cepivorum berk. In muck soil as influenced by drying and the location of sclerotia in soil

The proportion of viable sclerotia of Sclerotium cepivorum placed in field plots in Burnaby, British Columbia, decreased with time (P = 0.05). Sclerotia that had been air-dried for 48–72 hr had a lower percentage survival than those that had not been dried. Sclerotia placed on the soil surface decayed more rapidly than those buried at 15 cm (P = 0.05). Loss of viability was due to decay of sclerotia rather than to a reduction in the ability of the sclerotia to germinate which did not decline with time (P = 0.05). After 16 months in the field 23.6; 2.1; 11.7 and 8.9% of the sclerotia remained viable in the not-dried buried, not-dried surface, dried buried and dried surface treatments respectively.     

Some factors affecting survival of sclerotia of Macrophomina phaseolina in soil

At least 75% of the sclerotia of Macrophomina phaseolina survived for 1 yr in most natural soils kept at 26°C and at 50–55% of the soil moisture holding capacity (m.h.c.). Although survivability was reduced in a very acid soil (pH 4.5) collected under a pine stand, 33% of the sclerotia survived for 1 yr. Soil pH had very little or no effect on sclerotial survivability. Of three organic amendments tested (alfalfa hay, chitin, pine needles) only ground alfalfa hay at 0.8% (w/w) reduced survivability of sclerotia in soil by about 75% in a year. Alfalfa hay at 0.4% reduced survivability by 36%. Various N sources added at 200 μg Ng^?1 soil had no effect on survival. Of 13 fungicides tested, only benomyl and captan at 20 μg a.i. g^?1 soil appreciably reduced populations of sclerotia in soil.Soil temperature and moisture content were the two most important factors affecting survivability of sclerotia. At ?5 or 5°C the biggest drop in sclerotial survivability occurred when the soil was incubated moist (at 50% m.h.c. or more). At 26°C the biggest drop occurred in air-dried soil (2–3% m.h.c.) and survivability was decreased to some extent at 15 and 30% m.h.c. Survivability also dropped rapidly in moist soil (50–55% m.h.c.) exposed to four cycles each having 3-week freezing (?5°C) and 1 week thawing (26°C). Sclerotia in air-dried soil (2–3% m.h.c.) continuously kept at ?5°C maintained nearly complete survivability after 16 weeks. Sclerotia survived almost 80–90% in moist soil (50–55% m.h.c.) kept for 16 weeks at 26°C or in moist soil exposed to four cycles each having 3-week thawing (26°C) and 1-week freezing (?5°C).     

有机肥施用对农田中小型土壤动物群落的影响

2016年玉米生长季6～9月,以农田土壤为研究对象,设置了有机肥4个不同施用量处理,即0 kg·hm~(-2)(CK)、15 000 kg·hm~(-2)(OF1)、30 000 kg·hm~(-2)(OF2)、45 000 kg·hm~(-2)(OF3),并对其中小型土壤动物群落及其多样性的变化进行调查研究。研究结果表明,1)从试验样地共捕获中小型土壤动物2 821只,其中:OF处理显著提高了中小型土壤动物个体密度(P0.05),OF1、OF2和OF3处理分别较CK提高2.28%、31.93%和60.70%;有机肥施用有降低类群数的趋势,但无显著差异(P0.05);2)各处理对中小型土壤动物多样性指数无显著影响(P0.05);3)农田中小型土壤动物具有明显表聚性,各处理0～10 cm土层的个体密度和类群数显著高于10～20和20～30 cm土层的个体密度(P0.05)。有机肥施用对农田中小型土壤动物具有一定的影响,有机肥对土壤的影响是深远的,目前的结果着重表现在对个体密度的影响较大,对多样性的变化,需要长期观测。     

Reduction in viability of sclerotia of Macrophomina phaseolina with polyethylene mulching of soil

Mulching of Macrophomina phaseolina-inksted soil (moist or dry) with transparent polyethylene sheets during the hot days of May increased temperature of wet soil at 5 cm from 37°C (unmulched) to 52°C (mulched) and of dry soil from 52°C (unmulched) to 65°C (mulched). At 20 cm mulching increased temperature from 30°C to 41°C (wet) and from 38°C to 42°C (dry). In artificially-infested soil. the sclerotia of M. phaseolina were eradicated at 5 cm by a mulch treatment for 1 week and at 20 cm depth 50% sclerotia lost viability in wet soil but were not affected in dry soil. In a naturally infested soil (5–7 sclerotia g^?1), which gave 20% infection on Vigna, the sclerotia were reduced to such an extent that after 1 week mulching no disease was observed on Vigna.     

Short-term mesofauna responses to soil additions of corn stover biochar and the role of microbial biomass

Biochar additions have been suggested to influence soil microbial communities that, through a cascade effect, may also impact soil fauna. In turn, any direct biochar effects on fauna can influence microbial communities through grazing, physical fragmentation of organic debris (and biochar) and modifying soil structure. If biochar creates a favorable environment for soil microorganisms, it is also plausible for fauna to be attracted to such microbially enriched habitats. However, how soil fauna respond to biochar addition to soil and what are the main factors that drive their behavior has rarely been experimentally addressed. Therefore, the behavior of two mesofauna species was assessed as a result of corn stover biochar (slow pyrolysis at 600 °C) additions to a loamy temperate soil, after preincubation for 2, 17, 31 and 61 d, and related to variations in microbial biomass and activity. Microbial biomass increased by 5–56% and activity by 6–156% with increasing biochar rates for the different preincubation times. Over the incubation time, microbial biomass did not change or increased at most 15% with the different biochar rates, while in turn microbial activity decreased steadily (around 70–80% at day 61). Enchytraeids generally did not show avoidance or preference to biochar when provided with an alternative unamended soil, while collembolans often showed avoidance responses. However, collembolan avoidance to biochar decreased or disappeared in biochar mixtures with higher microbial biomass and water extractable NH₄-N content, agreeing with the plausible role of microorganisms to potentially attract soil fauna after biochar applications. Avoidance response was mainly explained by environmental preferences of the test species and not by any toxic effect of the biochar in this study. However, avoidance after the application of biochar may still need to be considered due to the potential negative impacts of individuals’ migration on soil ecosystem functioning.     

Seasonal differentiation of soil mesofauna in a littoral dune of the Egyptian Mediterranean coast

Summary Soil mesofauna (sensu Ghilarov, i.e., larger than 1 mm) was sampled from under four shrub species (Lygos raetam, Thymelaea hirsuta, Lycium shawii, and Josonia candicans) growing on the littoral oolitic sand dunes at Gharbaniat, 53 km W of Alexandria, Egypt, for a period of more than 30 months, spanning 3 years. Population density (PD) was calculated in relation to area of shrub canopy. Seasons were considered, according to the prevailing climatic conditions as well as to earlier physiological studies on the animals concerned, as: four winter months, four summer months and two months for each of the transitional seasons spring and autumn. The PD values obtained for each season thus defined were combined for the 41 taxa that were sampled. The table of these data was treated by correspondence analysis (CA) and ascending hierarchic classification (AHC), by the Roux DATA-VISION programme (suitable for Apple). Results from such treatment show that fauna as related to shrub species are differentiated along the first factorial axis, while as related to seasons they are differentiated along the second axis. Taxa common to all shrubs and to all seasons are centrally placed and cause some confusion in the resolution. Summer taxa are more distantly placed than those related to the other three seasons. Fauna under Lygos is richer, more diverse, and more complementary as regards trophic levels (with predators), and could probably be the source for replenishing the fauna of the other shrubs at all seasons. The AHC method did not show a similarly clear differentiation.Dedicated to the late Prof. Dr. M.S. Ghilarov     

Effects of soil mesofauna and farming management on decomposition of clover litter: a microcosm experiment

The effects of soil mesofauna and different farming systems on decomposition of clover (Trifolium repens) litter were investigated in a laboratory experiment. Microcosms were incubated for 16 weeks with fine and coarse litterbags in soils from three types of management systems: fallow, integrated farming and organic farming, the latter two cropped with wheat. The effects were studied by analysing litter mass loss, C and N content, DOC, nitrate and pH in soil leachate, and CO₂ production, as well as mesofauna. Mesofauna significantly accelerated mass loss and C and N release from clover litter in all three soils. With mesofauna access, at the end of the experiment average clover mass loss was almost twice as high and clover C and N content were 60% lower than without mesofauna. Farming systems influenced the decomposition through affecting both element turnover and mesofauna. Although in the first weeks less N was leached from organic farming than from integrated farming soil, cumulative N leaching did not differ between these soils. However, more than 20% less N was leached from the fallow soil than from the field soils. CO₂ production was highest in fallow soil. Here, mesofauna had no effect on this variable. In soil with integrated farming, mesofauna reduced cumulative CO₂ production by 10% whereas in soil from organic farming it increased CO₂ production by 20%. Our data suggest that differences in C and N turnover in different management systems are strongly mediated by soil mesofauna.     

Diversity of potential microbial parasites colonizing sclerotia of Macrophomina phaseolina in soil

The colonization of Macrophomina phaseolina sclerotia by microbial parasites was evaluated in unsterilized field soil at different levels of soil moisture (0,-5, and-10 kPa) and temperature (20, 30, and 40°C). The maximum colonization of sclerotia was recorded in soil held at-5 or-10 kPa at 30–40°C. Trichoderma harzianum isolate 25–92 and Pseudomonas fluorescens isolate 4–92 were recorded as potential sclerotial parasites, and they significantly (P=0.05) reduced the germination of sclerotia by 60–63%. Cells of P. fluorescens and buffer-washed conidia of T. harzianum were completely agglutinated at 28°C with crude agglutinin of M. phaseolina. The ability of different antagonists to parasitize the sclerotia were correlated with the agglutination ability of the antagonists.     

Structural-functional organization of the soil mesofauna population in bogs and waterlogged forests of northwestern Karelia

The soil mesofauna was studied in two ecological series (oligotrophic and mesotrophic) of bogs in northwestern Karelia. The earlier found differences in the structural-functional organization of the soil mesofauna in the bogs investigated were confirmed. The high indices of the taxonomic diversity and abundance of mesopedobionts were characteristic of the mesotrophic biogeocenoses. Among the factors limiting the activity of the mesopedobionts, the hydrothermal regime and soil acidity played the leading part. In the series of the bog biogeocenoses studied, the abundance of mesopedobionts increased with the increasing soil acidity, lowering the groundwater table and decreasing the content of sodium extracted by the ammonium acetate solution. Zoophages predominated in the trophic structure of the oligotrophic bogs due to the unfavorable conditions for large invertebrates there; in the mesotrophic bogs, myxophages were the dominants. The abundance of myxophages increased in the mesotrophic series simultaneously with the increasing nitrogen content in the soil mesofauna-inhabited soil horizons. In the mesotrophic biogeocenoses, saprophages were not numerous (one species of litter earthworms). Probably, in the bog soils, they are substituted for myxophages.