The influence of habitat provisioning: use of earthworm burrows by the terrestrial salamander, Plethodon cinereus

Studies examining the influence of habitat provisioning by one species on the behavior of other species can provide key insights regarding impacts of ecosystem engineers on the availability of resources to other species. More specifically, an organism’s use of additional habitat provided by ecosystem engineers may affect the interpretation of observational or demographic data. We chose to examine the possible influence of earthworms, as ecosystem engineers, on the behavior of terrestrial salamanders, common forest vertebrates in North America. We conducted two experiments in microcosms to examine whether the earthworm Lumbricus terrestris acted as an ecosystem engineer by providing additional habitat (burrows) for Plethodon cinereus, a common woodland salamander. We also examined whether the behavioral changes of the non-burrowing, fossorial P. cinereus differed across age classes. Adults and juveniles responded similarly, with both adults and juveniles found under cover objects more often when earthworms were absent and using earthworm burrows when available. Our field data supported our experimental data in that greater numbers of earthworms on field plots were associated with a lower proportion of cover objects being occupied by salamanders (i.e., salamanders were likely underground). We found no differences in the response of adults and juveniles to the presence of earthworms. By providing underground habitat for a common terrestrial amphibian, the presence of earthworms may impact demographic studies on salamanders and conclusions drawn from those studies regarding aboveground faunal communities in eastern North America.     

Home-Range Dynamics in a Solitary Subterranean Rodent

Despite an important role of subterranean rodents as ecosystem engineers, their belowground mobility is poorly documented. It is supposed that their underground burrow systems, once established, are relatively stable because of high-energy costs of digging. We chose the silvery mole-rat, Heliophobius argenteocinereus (Bathyergidae, Rodentia) from mesic Afrotropics as a representative of solitary subterranean rodents to investigate how, and how fast these rodents process their established burrow systems. We combined radio-tracking of individual animals with subsequent mapping of their burrow systems, and we developed a new method for assessing the rate of burrowing. Mole-rats continuously rebuilt their burrow systems; they excavated approx. 0.7 m of new tunnels per day and backfilled on average 64% of all tunnels. On average, every 32 d they established a new nest. They often completely backfilled newly excavated peripheral burrows, while other parts of their burrow systems were more permanent. Their home-ranges were dynamic and continuously shifted in space. Burrow system processing continued even in the advanced dry season, when soil is difficult to work.     

Nonlinearity of effects of invasive ecosystem engineers on abiotic soil properties and soil biota

Invasions of non‐indigenous species into natural communities are currently rated as one of the most important threats to biodiversity. Particularly exotic ecosystem engineers such as earthworms potentially have profound impacts on community assembly and functioning. We investigated the impact of invasion by the lumbricid earthworms into an aspen forest of the Canadian Rocky Mountains on soil organic matter, microorganisms and microarthropod communities. Building on the results of previous studies in this forest, we expected positive effects of Lumbricus terrestris middens and negative effects of Octolasion tyrtaeum on soil biota (increase and decrease in soil nutrient concentrations, microbial parameters and soil microarthropod density and diversity, respectively). Further, we expected that earthworm effects change with time. Combined results of previous and the present study suggest a wavelike colonization pattern for Dendrobaena octaedra and O. tyrtaeum and that indeed the impact of earthworms on soil biota changed with time, likely due to changes in earthworm density. Unexpectedly, L. terrestris middens neither affected soil abiotic nor soil biotic properties. By contrast and in contrast to our hypothesis, carbon and nitrogen concentration and C‐to‐N ratio in deeper soil layers increased in presence of O. tyrtaeum, thereby likely enhancing nutrient availability for soil microorganisms and microarthropods. Even though the density of this endogeic species was rather low, presence of O. tyrtaeum resulted in increased densities of a number of microarthropod taxa and increased microarthropod diversity. The results suggest that at low density, invasive ecosystem engineers, such as O. tyrtaeum, cause disturbances of intermediate strength thereby beneficially affecting soil microorganisms and most microarthropods. This contrasts earlier effects during the wavelike invasion of O. tyrtaeum into the aspen forest when densities of O. tyrtaeum were high resulting in generally detrimental effects on soil biota. The results emphasize the nonlinearity of earthworm effects on abiotic and biotic soil properties and call for further long‐term investigations.     

Exotic Ecosystem Engineers Change the Emergence of Plants from the Seed Bank of a Deciduous Forest

The anthropogenic spread of exotic ecosystem engineers profoundly impacts native ecosystems. Exotic earthworms were shown to alter plant community composition of the understory of deciduous forests previously devoid of earthworms. We investigated the effect of two exotic earthworm species (Lumbricus terrestris L. and Octolasion tyrtaeum Savigny) belonging to different ecological groups (anecic and endogeic) on the emergence of plants from the seed bank of a northern North American deciduous forest using the seedling emergence method. We hypothesized that (1) exotic earthworms change the seedling emergence from the plant seed bank, (2) L. terrestris increases the emergence of plant seedlings of the deeper soil layer but decreases that of the upper soil layer due to plant seed burial, and (3) O. tyrtaeum decreases plant seedling emergence due the damage of plant seeds. Indeed, exotic earthworms altered the emergence of plant seedlings from the seed bank and the functional composition of the established plant seedlings. Surprisingly, although L. terrestris only marginally affected seedling emergence, O. tyrtaeum changed the emergence of native plant species from the seed bank considerably. In particular, the number of emerging grass and herb seedlings were increased in the presence of O. tyrtaeum in both soil layers. Moreover, the impacts of earthworms depended on the identity of plant functional groups; herb species benefited, whereas legumes suffered from the presence of exotic earthworms. The results highlight the strong effect of invasive belowground ecosystem engineers on aboveground ecosystem characteristics and suggest fundamental changes of ecosystems by human-spread earthworm species.     

Worm grunting, fiddling, and charming--humans unknowingly mimic a predator to harvest bait

Background

For generations many families in and around Florida''s Apalachicola National Forest have supported themselves by collecting the large endemic earthworms (Diplocardia mississippiensis). This is accomplished by vibrating a wooden stake driven into the soil, a practice called “worm grunting”. In response to the vibrations, worms emerge to the surface where thousands can be gathered in a few hours. Why do these earthworms suddenly exit their burrows in response to vibrations, exposing themselves to predation?

Principal Findings

Here it is shown that a population of eastern American moles (Scalopus aquaticus) inhabits the area where worms are collected and that earthworms have a pronounced escape response from moles consisting of rapidly exiting their burrows to flee across the soil surface. Recordings of vibrations generated by bait collectors and moles suggest that “worm grunters” unknowingly mimic digging moles. An alternative possibility, that worms interpret vibrations as rain and surface to avoid drowning is not supported.

Conclusions

Previous investigations have revealed that both wood turtles and herring gulls vibrate the ground to elicit earthworm escapes, indicating that a range of predators may exploit the predator-prey relationship between earthworms and moles. In addition to revealing a novel escape response that may be widespread among soil fauna, the results show that humans have played the role of “rare predators” in exploiting the consequences of a sensory arms race.     

Root Foraging Influences Plant Growth Responses to Earthworm Foraging

Interactions among the foraging behaviours of co-occurring animal species can impact population and community dynamics; the consequences of interactions between plant and animal foraging behaviours have received less attention. In North American forests, invasions by European earthworms have led to substantial changes in plant community composition. Changes in leaf litter have been identified as a critical indirect mechanism driving earthworm impacts on plants. However, there has been limited examination of the direct effects of earthworm burrowing on plant growth. Here we show a novel second pathway exists, whereby earthworms (Lumbricus terrestris L.) impact plant root foraging. In a mini-rhizotron experiment, roots occurred more frequently in burrows and soil cracks than in the soil matrix. The roots of Achillea millefolium L. preferentially occupied earthworm burrows, where nutrient availability was presumably higher than in cracks due to earthworm excreta. In contrast, the roots of Campanula rotundifolia L. were less likely to occur in burrows. This shift in root behaviour was associated with a 30% decline in the overall biomass of C. rotundifolia when earthworms were present. Our results indicate earthworm impacts on plant foraging can occur indirectly via physical and chemical changes to the soil and directly via root consumption or abrasion and thus may be one factor influencing plant growth and community change following earthworm invasion. More generally, this work demonstrates the potential for interactions to occur between the foraging behaviours of plants and soil animals and emphasizes the importance of integrating behavioural understanding in foraging studies involving plants.     

Local distribution of native and invasive earthworms and effects on a native salamander

North America is home to both native and invasive earthworms acting as ecosystem engineers as they build burrows that can serve as habitat for other species or otherwise alter soil structure, affecting nutrient cycling and other ecosystem processes. Here I determine where and what earthworm species commonly occur in my study area, and compare effects of native and invasive earthworms on the common woodland salamander, Plethodon cinereus, in field surveys and laboratory experiments. The native earthworm Eisenoides carolinensis was the most common earthworm, followed by two invasive species Dendrobaena octaedra and Octolasion tyrtaeum. The presence of O. tyrtaeum was associated with a narrower O-horizon (i.e., organic layer in the soil). Using structural equation modeling to explore direct and indirect pathways of these three most common earthworm species on salamanders, I found O. tyrtaeum occurrence was negatively correlated with nighttime salamander counts, a proxy for total salamander numbers, mediated by negative effects on O-horizon depth and microinvertebrate numbers. In the laboratory, O. tyrtaeum and D. octaedra consumed more leaf litter per gram of earthworm per day than the native E. carolinensis. However, salamanders consumed earthworms and used burrows of all native and invasive species of earthworms similarly. The potential for negative indirect effects of the invasive earthworm O. tyrtaeum on P. cinereus was demonstrated both in the field and laboratory, highlighting that seemingly small differences between native and invasive ecosystem engineers have the potential to significantly alter the effects of these closely related native and invasive organisms.     

Colonization and Recovery of Invertebrate Ecosystem Engineers during Prairie Restoration

Ants (Hymenoptera: Formicidae) and earthworms (Oligochaeta) are considered ecosystem engineers because they form biogenic structures in the soil that influence resource supply. The objectives of this study were to quantify recovery dynamics of these invertebrate groups across a chronosequence of restored prairies and elucidate whether changes in the abundance and biomass of ants and earthworms were related to key plant and ecosystem properties. We sampled ants and earthworms from cultivated fields, grasslands restored from 1 to 21 years, and native prairie. Ant abundance and biomass peaked between 5 and 8 years of restoration and abundance was 198 times greater than cultivated fields. Earthworm abundance increased linearly across the chronosequence and became representative of native prairie, but all earthworm populations were dominated by European species. Ant abundance and biomass were positively correlated with plant diversity and plant richness, whereas earthworm abundance biomass was only related to surface litter. These results demonstrate that earthworm abundance increases with time since cessation of cultivation and concomitant with prairie establishment, whereas the abundance and biomass of ants are more related to the structure of restored plant communities than time. The dominance of exotic earthworms in these restorations, coupled with their capacity to alter soil properties and processes may represent novel conditions for grassland development.     

Earthworms, as ecosystem engineers, influence multiple aspects of a salamander��s ecology

Ecosystem engineers create habitat that can be used by other species in multiple ways, such as refuges from predators, places to breed, or areas with increased prey resources. I conducted a series of enclosure experiments to: (1) determine if salamanders use earthworm burrows, and (2) examine the potential influence of earthworm burrow use and indirect effects on salamander intra- and interspecific competition, predator avoidance, and seasonal performance. I found that one species of woodland salamander, Plethodon cinereus, used earthworm burrows 50% of the time when burrows were present. Neither adults nor juveniles of the congeneric P. glutinosus used earthworm burrows. Intraspecific, but not interspecific, competition by P. cinereus affected salamander behavior when earthworms were absent, with P. cinereus found under cover objects >70% of the time when alone or with a P. glutinosus, but only 40% of the time when with another P. cinereus. When earthworms were present, the behavior of P. cinereus was similar across salamander treatments. Earthworms decreased the amount of leaf litter and microinvertebrates, although this did not affect salamander mass. In subsequent experiments using only P. cinereus, the refuge provided by earthworm burrows increased the survival of P. cinereus over the winter and allowed P. cinereus to avoid being consumed by the common garter snake (Thamnophis sirtalis). Because earthworm burrows provide a refuge for P. cinereus during intraspecific encounters, in the presence of a predator and over the winter, they may serve as an important belowground?Caboveground linkage in eastern forests where salamanders are common.     

Root, mycorrhiza and earthworm interactions: their effects on soil structuring processes, plant and soil nutrient concentration and plant biomass

Earthworms, arbuscular mycorrhiza fungi (AMF) and roots are important components of the belowground part of terrestrial ecosystem. However, their interacting effects on soil properties and plant growth are still poorly understood. A compartmental experimental design was used in a climate chamber in order to investigate, without phosphorus (P) addition, the single and combined effects of earthworms (Allolobophora chlorotica), AMF (Glomus intraradices) and roots (Allium porrum) on soil structure, nutrient concentration and plant growth. In our experimental conditions, plant roots improved soil structure stability (at the level of macroaggregates) whereas earthworms decreased it. AMF had no effect on soil structure stability but increased P transfer from the soil to the plant and significantly increased plant biomass. Earthworms had no direct influence on P uptake or plant biomass, and the N/P ratio measured in the shoots indicated that P was limiting. Interactions between AMF and earthworms were also observed on total C and N content in the soil and on total root biomass. Their effects varied temporally and between the different soil compartments (bulk soil, rhizosphere and drilosphere). After comparison with other similar studies, we suggest that effects of earthworms and AMF on plant production may depend on the limiting factors in the soil, mainly N or P. Our experiment highlights the importance of measuring physical and chemical soil parameters when studying soil organism interactions and their influence on plant performance.     

Earthworm effect on root morphology in a split root system

Plants respond to their environment through adaptations such as root proliferation in nutrient-rich patches. Through their burrows and casts production in soil, earthworms create heterogeneity which could lead to local root adaptations or systemic effects. To investigate the effect of earthworms on root system morphology and determine whether earthworm effect is local or systemic, we set up two independent split root experiments with rice or barley, (i) without earthworm (CC), (ii) with earthworms in both compartments (EE), and (iii) with earthworms in one single compartment (CE). Earthworms had an effect on belowground plant biomass. The relative length of thick roots decreased with an increasing abundance of earthworms. Some root diameter classes responded to earthworm number in a linear or curvilinear way, making simple conclusions difficult. We found no difference in root biomass or morphology between the two compartments of the split root system in the CE treatment, but a positive effect of earthworm biomass on root biomass, volume, surface area, and length at the whole plant level. Results supported a systemic effect dependent on earthworm abundance. Modification of nutrient mineralization, soil physical structure, and/or the concentration of signal molecules could all be responsible for this systemic effect.     

两种代表性蚯蚓对设施菜地土壤微生物群落结构及理化性质的影响

不同生态型蚯蚓的取食偏好和生境有所差异,因此蚯蚓的生态型差异可能关乎其对土壤性质的不同影响;有关不同生态型蚯蚓对土壤性质尤其是微生物学性质影响的研究有助于了解蚯蚓生态功能的作用机制。在野外调控试验的第4年采集土壤,研究了牛粪混施和表施处理下内层种威廉腔环蚓(Metaphire guillelmi)和表层种赤子爱胜蚓(Eisenia foetida)对设施菜地土壤微生物群落结构和主要理化性质的影响。结果表明,土壤微生物群落结构同时受到蚯蚓种类和牛粪施用方式的影响。牛粪表施时,两种蚯蚓均显著降低了菌根真菌、真菌生物量和原生动物生物量(P0.05);牛粪混施时,不同蚯蚓的影响有所差异,威廉腔环蚓明显增加了菌根真菌、真菌生物量和放线菌生物量,而赤子爱胜蚓的作用不明显。此外,两种蚯蚓均提高了土壤孔隙度、团聚体稳定性和土壤p H、矿质氮以及微生物生物量碳氮水平,但提高幅度取决于蚯蚓种类和牛粪施用方式。冗余分析表明蚯蚓影响下土壤微生物群落结构的变化与团聚体稳定性、p H、速效磷、矿质氮呈正相关,而与土壤容重呈负相关。     

Wildlife associates of nine‐banded armadillo (Dasypus novemcinctus) burrows in Arkansas

The Nine‐banded Armadillo (Dasypus novemcinctus) is a widespread burrowing species with an expanding geographic range across the southeastern and midwestern United States. Armadillos dig numerous, large burrows within their home ranges and these burrows are likely used by a diverse suite of wildlife species as has been reported for other burrowing ecosystem engineers such as Gopher Tortoises (Gopherus polyphemus), Desert Tortoises (Gopherus agassizi), and Black‐tailed Prairie Dogs (Cynomys ludovicianus). We used motion‐triggered game cameras at 35 armadillo burrows in 4 ecoregions of Arkansas and documented 19 species of mammals, 4 species of reptile, 1 species of amphibian, and 40 species of bird interacting with burrows. Bobcat (Lynx rufus), Coyote (Canis latrans), Eastern Cottontail (Sylvilagus floridanus), Gray Fox (Urocyon cinereoargenteus), Gray Squirrel (Sciurus carolinensis), Northern Raccoon (Procyon lotor), Virginia Opossum (Didelphis virginiana), and unidentified rodents (mice and rats) were documented using burrows in all four ecoregions. We documented wildlife hunting, seeking shelter, rearing young in, and taking over and modifying armadillo burrows. The rate of use was highest in the Mississippi Alluvial Valley, a landscape dominated by agriculture, where natural refugia may be limited and rodents are abundant. Armadillo burrows are clearly visited and used by numerous wildlife species to fulfill various life stage requirements, and this list will likely expand if more attention is devoted to understanding the role of armadillos burrows. Armadillos are important ecosystem engineers, and their ecological role warrants more investigation and attention as opposed to only being viewed and managed as agricultural and garden pests.     

Earthworm-mycorrhiza interactions can affect the diversity, structure and functioning of establishing model grassland communities

Both earthworms and arbuscular mycorrhizal fungi (AMF) are important ecosystem engineers co-occurring in temperate grasslands. However, their combined impacts during grassland establishment are poorly understood and have never been studied. We used large mesocosms to study the effects of different functional groups of earthworms (i.e., vertically burrowing anecics vs. horizontally burrowing endogeics) and a mix of four AMF taxa on the establishment, diversity and productivity of plant communities after a simulated seed rain of 18 grassland species comprising grasses, non-leguminous forbs and legumes. Moreover, effects of earthworms and/or AMF on water infiltration and leaching of ammonium, nitrate and phosphate were determined after a simulated extreme rainfall event (40 l m(-2)). AMF colonisation of all three plant functional groups was altered by earthworms. Seedling emergence and diversity was reduced by anecic earthworms, however only when AMF were present. Plant density was decreased in AMF-free mesocosms when both anecic and endogeic earthworms were active; with AMF also anecics reduced plant density. Plant shoot and root biomass was only affected by earthworms in AMF-free mesocosms: shoot biomass increased due to the activity of either anecics or endogeics; root biomass increased only when anecics were active. Water infiltration increased when earthworms were present in the mesocosms but remained unaffected by AMF. Ammonium leaching was increased only when anecics or a mixed earthworm community was active but was unaffected by AMF; nitrate and phosphate leaching was neither affected by earthworms nor AMF. Ammonium leaching decreased with increasing plant density, nitrate leaching decreased with increasing plant diversity and density. In order to understand the underlying processes of these interactions further investigations possibly under field conditions using more diverse belowground communities are required. Nevertheless, this study demonstrates that belowground-aboveground linkages involving earthworms and AMF are important mediators of the diversity, structure and functioning of plant communities.     

Differential effects of ecosystem engineering by the superb lyrebird Menura novaehollandiae and herbivory by large mammals on floristic regeneration and structure in wet eucalypt forests

Ecosystem engineers that modify the soil and ground‐layer properties exert a strong influence on vegetation communities in ecosystems worldwide. Understanding the interactions between animal engineers and vegetation is challenging when in the presence of large herbivores, as many vegetation communities are simultaneously affected by both engineering and herbivory. The superb lyrebird Menura novaehollandiae, an ecosystem engineer in wet forests of south‐eastern Australia, extensively modifies litter and soil on the forest floor. The aim of this study was to disentangle the impacts of engineering by lyrebirds and herbivory by large mammals on the composition and structure of ground‐layer vegetation. We carried out a 2‐year, manipulative exclusion experiment in the Central Highlands of Victoria, Australia. We compared three treatments: fenced plots with simulated lyrebird foraging; fenced plots excluding herbivores and lyrebirds; and open controls. This design allowed assessment of the relative impacts of engineering and herbivory on germination rates, seedling density, vegetation cover and structure, and community composition. Engineering by lyrebirds enhanced the germination of seeds in the litter layer. After 2 years, more than double the number of germinants were present in “engineered” than “non‐engineered” plots. Engineering did not affect the density of seedlings, but herbivory had strong detrimental effects. Herbivory also reduced the floristic richness and structural complexity (<0.5 m) of forest vegetation, including the cover of herbs. Neither process altered the floristic composition of the vegetation within the 2‐year study period. Ecosystem engineering by lyrebirds and herbivory by large mammals both influence the structure of forest‐floor vegetation. The twofold increase in seeds stimulated to germinate by engineering may contribute to the evolutionary adaptation of plants by allowing greater phenotypic expression and selection than would otherwise occur. Over long timescales, engineering and herbivory likely combine to maintain a more‐open forest floor conducive to ongoing ecosystem engineering by lyrebirds.     

13C and 15N in microarthropods reveal little response of Douglas-fir ecosystems to climate change

Understanding ecosystem carbon (C) and nitrogen (N) cycling under global change requires experiments maintaining natural interactions among soil structure, soil communities, nutrient availability, and plant growth. In model Douglas-fir ecosystems maintained for five growing seasons, elevated temperature and carbon dioxide (CO₂) increased photosynthesis and increased C storage belowground but not aboveground. We hypothesized that interactions between N cycling and C fluxes through two main groups of microbes, mycorrhizal fungi (symbiotic with plants) and saprotrophic fungi (free-living), mediated ecosystem C storage. To quantify proportions of mycorrhizal and saprotrophic fungi, we measured stable isotopes in fungivorous microarthropods that efficiently censused the fungal community. Fungivorous microarthropods consumed on average 35% mycorrhizal fungi and 65% saprotrophic fungi. Elevated temperature decreased C flux through mycorrhizal fungi by 7%, whereas elevated CO₂ increased it by 4%. The dietary proportion of mycorrhizal fungi correlated across treatments with total plant biomass (n= 4, r²= 0.96, P= 0.021), but not with root biomass. This suggests that belowground allocation increased with increasing plant biomass, but that mycorrhizal fungi were stronger sinks for recent photosynthate than roots. Low N content of needles (0.8–1.1%) and A horizon soil (0.11%) coupled with high C : N ratios of A horizon soil (25–26) and litter (36–48) indicated severe N limitation. Elevated temperature treatments increased the saprotrophic decomposition of litter and lowered litter C : N ratios. Because of low N availability of this litter, its decomposition presumably increased N immobilization belowground, thereby restricting soil N availability for both mycorrhizal fungi and plant growth. Although increased photosynthesis with elevated CO₂ increased allocation of C to ectomycorrhizal fungi, it did not benefit plant N status. Most N for plants and soil storage was derived from litter decomposition. N sequestration by mycorrhizal fungi and limited N release during litter decomposition by saprotrophic fungi restricted N supply to plants, thereby constraining plant growth response to the different treatments.     

Earthworm and belowground competition effects on plant productivity in a plant diversity gradient

Diversity is one major factor driving plant productivity in temperate grasslands. Although decomposers like earthworms are known to affect plant productivity, interacting effects of plant diversity and earthworms on plant productivity have been neglected in field studies. We investigated in the field the effects of earthworms on plant productivity, their interaction with plant species and functional group richness, and their effects on belowground plant competition. In the framework of the Jena Experiment we determined plant community productivity (in 2004 and 2007) and performance of two phytometer plant species [Centaurea jacea (herb) and Lolium perenne (grass); in 2007 and 2008] in a plant species (from one to 16) and functional group richness gradient (from one to four). We sampled earthworm subplots and subplots with decreased earthworm density and reduced aboveground competition of phytometer plants by removing the shoot biomass of the resident plant community. Earthworms increased total plant community productivity (+11%), legume shoot biomass (+35%) and shoot biomass of the phytometer C. jacea (+21%). Further, phytometer performance decreased, i.e. belowground competition increased, with increasing plant species and functional group richness. Although single plant functional groups benefited from higher earthworm numbers, the effects did not vary with plant species and functional group richness. The present study indicates that earthworms indeed affect the productivity of semi-natural grasslands irrespective of the diversity of the plant community. Belowground competition increased with increasing plant species diversity. However, belowground competition was modified by earthworms as reflected by increased productivity of the phytometer C. jacea. Moreover, particularly legumes benefited from earthworm presence. Considering also previous studies, we suggest that earthworms and legumes form a loose mutualistic relationship affecting essential ecosystem functions in temperate grasslands, in particular decomposition and plant productivity. Further, earthworms likely alter competitive interactions among plants and the structure of plant communities by beneficially affecting certain plant functional groups.     

Earthworms as paratenic hosts of toxoplasmosis in eastern barred bandicoots in Tasmania

An experimental feeding study was designed to assess the role of earthworms in the transmission of Toxoplasma gondii infection to eastern barred bandicoots (Perameles gunnii). Six animals with no agglutinating antibodies to T. gondii were fed artificially cultured earthworms that had been maintained in autoclaved nutrient-enriched soil. Two animals were given earthworms that had been maintained in soil contaminated with T. gondii oocysts (P89/VEG strain); two animals were fed on earthworms, which initially had been exposed to soil containing T. gondii oocysts then transferred through three changes of sterile soil; two control bandicoots were fed earthworms maintained in sterile soil. Both bandicoots fed earthworms maintained in T. gondii contaminated soil died 11 and 14 days after feeding. The necropsy findings were consistent with acute toxoplasmosis. Bandicoots fed earthworms exposed to oocysts but then transferred through changes of sterilized soil remained healthy as did control animals. All surviving animals remained seronegative over the 6 wk observation period after feeding. These findings confirm that earthworms, a major component of the natural diet of P. gunnii, can transmit T. gondii infection. It appears that oocysts present in the alimentary tracts of the worms, rather than infective stages of T. gondii in worm somatic tissues, are responsible for these infections.     

Mole activity in woodlands, fens and other habitats

Monks Wood, a deciduous wood of 155 hectares on clay soil, has an estimated population of 400 moles. There is an anastomosing system of permanent burrows, and when a mole is trapped its area of burrow is soon occupied by another animal. Little new digging occurs, except in winter, after frost has driven the soil fauna deeper into the ground. The burrows usually act as pit-fall traps for the food. Young animals migrating in summer may live more superficially, in tunnels in moss and grass, but survivers usually move into a permanent tunnel system by autumn.
Woodwalton Fen has an easily worked peat soil, and few earthworms. Here moles burrow at all seasons, presumably needing an extensive burrow system to catch sufficient food. In times of flood, the moles leave submerging ground but return very soon after the water subsides. They probably swim across flooded areas but iis yet we do not know if the same animals return after floods to the same burrows.
Moles are not good indicators of soil fertility, particularly as many mole heaps are made by few moles in poor soil, and fewmew heaps may appear in good soil with permanent burrows. We do not understand why there are so few moles in some apparently suitable pasture, with high worm populations, or how others manage to obtain sutficient food in infertile areas.     

Response of bacteria to earthworm surface excreta

Response of bacteria to the surface excreta of the Aporrectodea caliginosa earthworm was studied. The excreta were obtained by a 1 h incubation of the earthworms in petri dishes with subsequent collection of the slime. Both inhibition and stimulation of growth were revealed, as well as suppression of the respiratory activity of some bacterial species treated with A. caliginosa surface excreta. The organisms studied included various taxa of soil bacteria (19 strains), bacteria isolated from A. caliginosa intestine and excrements (82 strain), and 48 Bacillus thuringiensis strains. For the cultures of soil bacteria, the respiratory activity was determined using the formazan color reaction due to the activity of the respiratory cycle enzymes. Earthworm excreta caused a consistent 30–50% decrease of dehydrogenase activity in 13 out of the 19 cultures. Determination of the growth rates (derived from OD₆₂₀ of cell suspensions) after 10 h of incubation revealed growth stimulation in 48 out of the 82 strains isolated from intestines and excrement. Other strains exhibited no reaction to the excreta. For 29 out of 45 B. thuringiensis strains, growth stimulation was observed, while growth of two strains was suppressed; other strains exhibited no reaction to the excreta. No relation was found between bacterial reaction to the excreta and their taxonomic position. These results correlate with the research, demonstrating antibacterial and antifungal activity of the extracts from the earthworm body and digestive tract. Thus, earthworms, apart from their medium-forming function, affect the formation of soil microbial communities by direct stimulation or suppression of specific microbial populations.