Spatial and temporal patterns in the activity of European moles

 Using ratio-tracking data obtained at three sites, we assessed the effects of season and of neighbour avoidance on the activity and patterns of home range use by European moles (Talpa europaea). The home ranges of non-breeding male and female moles did not differ significantly in size, and averaged 2324 m² (minimum convex polygon). Although overlap between ranges was small (an average of 12.8% of each range being shared with neighbours and an average of only 3.3% of 2×2 m grid cells were shared with an individual neighbour, ranges were not oriented to avoid neighbours. Non-breeding male/female neighbours tended to share more of their grid cells (3.9±5.7%) mean ±SD than did neighbours of the same sex (male:male 1.2±0.95%; female:female 1.1±1.3%), but there was no significant difference in overlap between any combination of sex pairings. On average, each mole spent only 0.9% of its time within 6 m of another mole, and only 3 out of 46 dyads showed evidence of being attracted to each other; there was no evidence from the simultaneous movement patterns of neighbouring moles that they avoided each other. Although moles tended to return to the same part of their range at the same time on successive days, there was also some indication of gradual changes in the spatial pattern of daily home range use. Moles had a triphasic pattern of activity, but this became tetraphasic under drought conditions. There were significant differences between sites, but not between sexes, in sleeping behaviour and activity patterns. These differences could be related to seasonal differences in soil moisture and thus probably to prey renewal rates. We conclude that in our sites, the activity patterns and movements of moles depend on the temporal and spatial dispersion of food, rather than on short-term interactions between the movements of neighbours. Received: 13 January 1996 / Accepted: 26 June 1996     

The soil fauna as a food source for moles

Moles feed mainly on insects and earthworms and may eat 18 to 36 k (40 to 80 lb) per annum, collected in an area of about 1/25 hectare (1/10 acre). This largely restricts them to deciduous woodland and old grassland on "mull" soils since field populations of soilinsects cannot provide them with enough food but earthworm populations can. Slugs and millipedes may be relatively unpalatable.
Caches of earthworms stored by moles consist almost entirely of Lumbricus terrestris immobilized by removal or multilation of the anterior 3–5 segments. It is still uncertainhow and when the caches are made. Earthworm cocoons are prominent in the diet of moles in pastures but how they are located is not yet known.     

Comparison of ectomycorrhizas of Quercus garryana (Fagaceae) on serpentine and non-serpentine soils in southwestern Oregon

The diversity of ectomycorrhizal communities associated with Quercus garryana on and off serpentine soils was compared and related to landscape-level diversity. Serpentine soils are high in magnesium, iron, and heavy metals and low in fertility. In plant communities on serpentine soils, a high proportion of flowering plant species are endemic. At three sites with paired serpentine and nonserpentine soils in southwestern Oregon, we sampled Q. garryana roots and categorized ectomycorrhizas by morphotyping and by restriction fragment length patterns. Ectomycorrhizas were abundant at all sites; no single fungal species dominated in the ectomycorrhizas. Of 74 fungal species characterized by morphotype and pattern of restriction fragment length polymorphisms, 46 occurred on serpentine soils, and 32 were unique to serpentine soil. These species are potentially endemic to serpentine soil. Similarities in species composition between paired serpentine and nonserpentine soils were not significantly lower than among three serpentine sites or among three nonserpentine sites. We conclude that mycorrhizal communities associated with oaks on serpentine soil do not differ in species richness or species evenness from those on neighboring nonserpentine soil.     

The edaphic factor and patterns of variation in Lasthenia californica (Asteraceae)

Transectional studies of Lasthenia californica in the Jasper Ridge Biological Preserve (Stanford University) have documented the existence of two races (A and C) based upon flavonoid chemistry, achene morphology, allozymes, and flowering time differences. The two races coexist on a serpentine outcrop and have maintained a sharply defined pattern of distribution for a period of at least 15 yr. The present study has revealed significant differences in the physical and chemical features of the soils harboring the two races. Soils at the lower ends of the transects, where race A plants grow, have higher pH, cation exchange capacity, relative water content, total ionic strength, percentage clay, and sodium and magnesium concentrations than do soils harboring race C plants at the upper ends of the transects. Soils supporting race C plants have higher calcium, potassium, and nickel concentrations and higher calcium:magnesium ratios. Plant tissue concentrations of ions were also significantly different in the two races. Race A plants accumulated sodium to concentrations three times those observed with race C plants. Plants from an additional 22 sites gave very similar results. Greenhouse studies indicated that the two races from Jasper Ridge show differential responses to ridge-top and ridge-bottom soils. Race A achenes germinated, grew to maturity, and set seed about equally in the two soils. Race C achenes germinated in both types of soils but showed significantly poorer growth and absolutely no flowering when found in the soils of race A plants. Differential responses to edaphic conditions on the ridge may contribute to the pattern of distribution observed over the years. It is suggested that race A plants are more tolerant of edaphic stress than race C plants and that physiological specialization may contribute to the present distribution of the two races throughout the species' range. It is not yet possible to state which is the more significant factor in driving this specialization, the chemistry of the soil or its physical characteristics, or whether there is interaction between the two. This is the first study to present evidence for soil/plant variation within a serpentine site. The linking of sodium levels to racial differentiation within the serpentine habitat is also a new discovery.     

Mycorrhizal colonisation, root browning and soil properties associated with a growth disorder of cotton in Australia

The symptoms of a growth disorder of cotton and associated properties of the soil were quantified. Data were collected from 35 sites across two irrigated fields that showed gradients in the severity of early season stunting of cotton. Ordination analysis of soil characteristics distinguished three groups of sites (A, B, and C) which corresponded to patterns of yield and early season growth. Group A and B soils had lower pH, finer texture and higher P, Zn, Mn and exchangeable Mg, K and Na than group C soils. Early season growth and arbuscular mycorrhizal colonisation of cotton at group A and B sites was much slower than at group C sites. Group B sites showed a recovery of yield later in the season while group A sites did not. Nutrient deficiencies, waterlogging, soil compaction, soil sodicity and Mn toxicity were unlikely causes of early season stunting, although soil Mn was a good predictor for the disorder. Thielaviopsis basicola, Verticillium dahliae and unidentified Chytridiomycetes were not associated with stunting and reduced yield, although other fungal pathogens may have been present. Root browning was a symptom of the disorder and suggests that pathogens, perhaps bacteria, play a causal role. This study showed that the growth disorder involved an interaction between cotton and the soil flora which was associated with heavy soil texture.     

Australia's first fossil marsupial mole (Notoryctemorphia) resolves controversies about their evolution and palaeoenvironmental origins

Fossils of a marsupial mole (Marsupialia, Notoryctemorphia, Notoryctidae) are described from early Miocene deposits in the Riversleigh World Heritage Area, northwestern Queensland, Australia. These represent the first unequivocal fossil record of the order Notoryctemorphia, the two living species of which are among the world's most specialized and bizarre mammals, but which are also convergent on certain fossorial placental mammals (most notably chrysochlorid golden moles). The fossil remains are genuinely 'transitional', documenting an intermediate stage in the acquisition of a number of specializations and showing that one of these-the dental morphology known as zalambdodonty-was acquired via a different evolutionary pathway than in placentals. They, thus, document a clear case of evolutionary convergence (rather than parallelism) between only distantly related and geographically isolated mammalian lineages-marsupial moles on the island continent of Australia and placental moles on most other, at least intermittently connected continents. In contrast to earlier presumptions about a relationship between the highly specialized body form of the blind, earless, burrowing marsupial moles and desert habitats, it is now clear that archaic burrowing marsupial moles were adapted to and probably originated in wet forest palaeoenvironments, preadapting them to movement through drier soils in the xeric environments of Australia that developed during the Neogene.     

Habitat selection by threatened burrowing spiders (Araneae: Atypidae,Eresidae) of central Europe: evidence base for conservation management

Habitat selection in sedentary, long-lived burrowing spiders is a key life-history event that directly affects their reproductive success. In this study, we analyzed the role of the chemical and physical properties of soil in habitat selection by threatened temperate burrowing spiders. We examined 296 burrows of three Atypus spp. and three Eresus spp. at 68 sites in Czechia. We found that the study species were associated with soils that have high or very high cation exchange capacity, which allows the presence of a stable and humid microclimate in their burrows. We found that specific bedrock types can be used as predictors of the presence of particular study species. All Eresus spp. avoided compacted soils. However, when present in soils with very low penetration resistance, they were limited to sites with high soil cohesion. The burrows of all study species were located at well-drained sites. The study species seemed to be selective for a steppe-like character of their microhabitat but not necessarily for its southward orientation. We found the study species even at sites where steppes were present in the early 1950s and that underwent later afforestation but not extensive landscaping or plowing. What types of disturbances can be sustained by the study species and how long it takes the decreased populations to recover after, e.g., trampling associated with grazing, remain to be investigated.     

Seedling establishment after endozoochory in disturbed and undisturbed grasslands

Local plant community composition and structure may be largely influenced by germination and seedling establishment from seeds dispersed in animal dung, through seed input, gap creation and nutrient enrichment. With an experimental approach we assessed (1) what the effect is of dung deposition on the number of seedlings in the plant community 3 months and 1 year after dung deposition, (2) what the effect is of this seedling establishment on the local plant community characteristics such as species richness and (3) if this effect interacts with large-scale soil disturbance which removes the close canopy, such as sod-cutting. Viable seeds of monocotylous species were abundantly present in the dung, and dung deposition led to a higher number of monocotylous seedlings after 3 months. However, this effect was no longer significant after 1 year. Moreover, the proportion of viable monocotylous seeds that effectively established in the field after 3 months was less than 5%. A lower number of viable seeds of the less-dominant dicotylous species was dispersed in the dung but they had a higher cover and species richness after 1 year. This resulted in an increased total small-scale species richness and diversity after dung deposition through a decreasing dominance of monocotylous species. Sod-cutting had a pronounced effect on seedling emergence: viable seeds dispersed by dung had a higher probability of successful establishment when the dung was deposited in large gaps. This indicates that an increase of safe sites associated with disturbance strengthens the effects of seed dispersal and gap creation by dung deposition.     

No consistent legacy effects of invasion by giant goldenrod (Solidago gigantea) via soil biota on native plant growth

Aims Changes in soil microbial communities after occupation by invasive alien plants can represent legacy effects of invasion that may limit recolonization and establishment of native plant species in soils previously occupied by the invader. In this study, for three sites in southern Germany, we investigated whether invasion by giant goldenrod (Solidago gigantea) leads to changes in soil biota that result in reduced growth of native plants compared with neighbouring uninvaded soils.Methods We grew four native plant species as a community and treated those plants with soil solutions from invaded or uninvaded soils that were sterilized, or live, with live solutions containing different fractions of the soil biota using a decreasing sieve mesh-size approach. We measured aboveground biomass of the plants in the communities after a 10-week growth period.Main Findings Across all three sites and regardless of invasion, communities treated with <20 μm soil biota or sterilized soil solutions had significantly greater biomass than communities treated with the complete soil biota solution. This indicates that soil biota>20 μm are more pathogenic to the native plants than smaller organisms in these soils. Across all three sites, there was only a non-significant tendency for the native community biomass to differ among soil solution types, depending on whether or not the soil was invaded. Only one site showed significant differences in community biomass among soil solution types, depending on whether or not the soil was invaded; community biomass was significantly lower when treated with the complete soil biota solution than with soil biota <20 μm or sterilized soil solutions, but only for the invaded soil. Our findings suggest that efforts to restore native communities on soils previously invaded by Solidago gigantea are unlikely to be hindered by changes in soil microbial community composition as a result of previous invasion.     

Dung beetle activity affects rain forest seed bank dynamics and seedling establishment

Dung beetles relocate vertebrate feces under the soil surface, and this behavior has many ecological consequences. In tropical forests, for example, seeds defecated by mammals that are subsequently buried by dung beetles are less likely to suffer predation. While the effects of dung beetles on the fate of defecated seeds have been relatively well studied, their effect on seeds already buried in the soil has not. To contribute to fill this gap, we designed a study with three objectives: (a) Describe the vertical re‐distribution of soil seeds that occurs due to dung beetle activity; (b) Determine if beetle activity favors establishment of seedlings from the soil seed bank; and (c) Determine if the effect of dung beetles is stronger in sites of recurrent mammal defecation. We carried out three complementary field experiments, one with artificial seeds (plastic beads) of three sizes buried at known depths, one with two species of seeds buried at those same depths, and one with the natural soil seed bank in sites of single vs. recurrent defecation. Buried beads were moved by dung beetles along the vertical axis, both upwards (9.5%) and downwards (11.5%); smaller beads were more frequently moved downwards while the contrary occurred for larger beads. Dung beetle activity caused an increase in seedling establishment, both from experimentally buried seeds and from the natural seed bank. Defecation recurrence had no effect on seedling establishment. We conclude that dung beetle activity affects seed bank dynamics with important consequences for seedling establishment in tropical forests.Abstract in Spanish is available with online material.     

Conservation strategies for poorly surveyed taxa: a dung beetle (Coleoptera, Scarabaeidae) case study from southern Africa

Despite being the focus of an international research effort spanning decades, the spatial distribution of southern African scarab beetles remains poorly documented. As well as reinforcing the magnitude of the challenge facing biodiversity scientists, this raises real concerns about best practice conservation strategies in the absence of detailed distribution information. However, dung beetles appear to be well represented in established conservation areas. This apparent contradiction could be ascribed to anthropogenic transformation, successful conservation efforts, the presence of dung generalists and reserve-biased or mesic-biased dung beetle collection efforts. It is suggested that all of the above contribute to the observed pattern to varying degrees. The implications of selecting areas that are either rich in species, contain rare species or contain taxonomically distinct species from a group whose taxonomy is well known but for which inadequate distribution data exist are explored. Best practice, in the face of inadequate data, appears to revolve around a subtle interplay between advantages and disadvantages associated with data interpolation techniques, reserve selection algorithms that use criteria more robust than database rarity (such as taxonomic distinctiveness) and the long-term economic costs of proceeding with the data at hand versus investing in biological surveys.     

The status of the European Mole Talpa europaea L. as an agricultural pest and its management

Although the pest status of the European Mole Talpa europaea L. in Britain has never been adequately assessed, it is perceived as being a pest of agricultural and amenity land and is widely persecuted. We review the Mole's pest status, and evaluate current and proposed control methods.
While there has been a recent decline in the demand for strychnine, abuse of this poison leads annually to the death of many wild and domestic animals. The need remains for a humane, practical and safe alternative. If strychnine were banned without such an alternative being available, the use of mole-traps would probably increase. Through post-mortem examination of trapped Moles we evaluate the humaneness of these traps.
Our returned questionnaires showed that, while most farmers perceive Moles as pests, the damage which is attributed to them is slight on the great majority of farms. Control of Moles was none the less common and widespread, and was undertaken by 49.5% of respondents in 1992. Perceived pest status and the favoured method of control varied regionally; these trends may be related to soil quality and cultivation type.
Silage pollution was the most widely cited agricultural problem attributed to Moles. Practical measures to reduce silage pollution by Moles are discussed, particularly with regard to silage collection and treatment using bacteriophages. The efficiency of these additives merits detailed study because they have the potential, regardless of whether pollution has occurred or not, to improve the quality of the silage to an extent that killing Moles may be unnecessary; less than 1% of respondents reported that 10% or more of their silage was seriously affected by Mole activity.     

The Old and Young Amazon: Dung Beetle Biomass, Abundance, and Species Diversity

The Amazon Basin can be divided into two geomorphological regions based on the age of its soils: young (< 30 mya) and old (> 300 mya). We tested the effects of soil age on dung beetle communities by comparing biomass, abundance, and species between reserves in Ecuador on young soils and reserves in Brazil on old soils. Beetle biomass in the old Amazon was one-third that in the young Amazon, and beetle abundance in the old Amazon was one-fourth that in the young Amazon. Species richness, rarefied to equal sample sizes, was not significantly different between old and young soils. These data suggest young soils of the Amazon support a significantly greater biomass and abundance of dung beetles than old soils, but that species richness across the Basin is similar. As dung beetles are bio-indicators of mammals, our data support previous studies indicating a greater biomass of mammals on young versus old Amazon soils.     

Soil pH influences embryonic survival in Pseudophryne bibronii (Anura: Myobatrachidae)

Abstract: There has been limited research examining the role that terrestrial habitat characteristics play in influencing persistence of amphibian populations. In this study we investigated the influence of a terrestrial habitat attribute on the distribution of the terrestrial egg‐laying toadlet, Pseudophryne bibronii. Eggs of this species are deposited in depressions, or under leaf litter, and develop to a stage where they can hatch to free swimming tadpoles when water covers breeding sites or embryos are washed into water. Because rainfall can be intermittent, eggs may sit for extended periods in terrestrial nests before sufficient rainfall initiates hatching. Appropriate egg‐laying sites must be chosen by both males and females to ensure embryos survive these periods. A study of 20 sites found that soil pH at sites where P. bibronii were recorded as present was significantly lower than pH at sites where P. bibronii were recorded as absent. To determine whether soil pH influenced the hatching success in P. bibronii, experiments were conducted in which batches of eggs were translocated to local areas with different soil pH. Survival rate of eggs was significantly influenced by relatively small changes in pH, with hatching success higher at lower pH. In a laboratory trial survival of eggs raised on sterilized soil was not affected by soil pH. Fungal infection of eggs was observed in the field and laboratory suggesting that hatching success might be affected via a complex interaction between soil pH, fungi and other soil biota. Choice trials using P. bibronii metamorphs indicated that individuals were capable of distinguishing between small differences in pH. Therefore, distribution of P. bibronii may be influenced by either differential mortality of embryos, or habitat choice by post metamorphic individuals, and further experiments are required to determine the relative importance of these factors.     

Regional and local influence of grazing activity on the diversity of a semi-arid dung beetle community

This study analyses the effect of resource availability (i.e. sheep dung) on dung beetle communities in an arid region of Central Spain, both at regional and at local scales. A total of 18 sites within 600 km² were sampled for the regional analysis and 16 sites within the 30 km² of an Iberian municipality were sampled for the local analysis. Spatial and environmental characteristics of sampling sites were also compiled at both scales, including measures of grazing activity (livestock density at regional scale, and two counts of rabbit and sheep dung at local scale). At a regional scale, any environmental or spatial variable can help to explain the variation in abundance. However, species richness was related to summer precipitation and composition was related to elevation. At local scale, abundance is not significantly related to any of the environmental variables, but species richness was related to the local amount of sheep dung (27% of variance). The amount of dung in a 2‐km buffer around the site accounts for 27–32% of variance in abundance and 60–65% of variance in species richness. The presence of the flock with the highest sheep density explains 53% of abundance variability and 73% of species richness variance. A cluster analysis of localities identified two main groups, one characterized by a lower abundance and species richness that can be considered a nested subsample of the species‐rich group. The mean and maximum amount of sheep dung in the sites separated by less than 2 km are the only significant explanatory variables able to discriminate both groups. These results suggest that grazing intensity (and the associated increase in the amount of trophic resources) is a key factor in determining local variation in the diversity and composition of dung beetle assemblages. However, dung beetle assemblages are not spatially independent at the analysed resolution, and the amount of dung in the surroundings seems to be more important for locally collected species than the dung effectively found in the site. Although differences in the availability and quantity of trophic resources among nearby sites could be affecting the population dynamics and dispersion of dung beetles within a locality, sites with larger populations, and greater species numbers would not be able to exercise enough influence as to bring about a complete local faunistic homogenization.     

Soil properties and the distribution of the endangered Juliana's golden mole

The endangered Juliana's golden mole Neamblysomus julianae occurs in three small, geographically isolated populations within South Africa. As a sand-swimmer, it is a habitat specialist and has a highly fragmented distribution within each population. There is a need for a conservation action plan for this species, but this is difficult because of the dearth of knowledge of its biology, including its habitat characteristics. To better understand its patchy distribution, we selected 48 plots that were occupied or unoccupied by golden moles and measured the soil properties in each plot. The size distribution of sand particles influenced the density and compactability of the soil, and both were positively correlated with the presence of golden moles. These soil characteristics are thus vital in understanding and describing the distribution of this cryptic species.     

Baseline Levels of Metals in Volcanic Soils of the Azores (Portugal)

Data on metal concentrations present in the soils of the Azores (Portugal) are scarce. The goal of this study was to measure the current levels of several metals in the top horizon of soils of two areas, distinguishable by their volcanic activity and physical characteristics, in order to establish some baseline concentrations of these elements. Soil samples were taken in similar ways from five sites in a volcanically active area and another five sites in an area without volcanic activity. Particle-size fractions, % organic matter, moisture content, pH, and major and trace elements compositions were measured. In general, the concentrations of trace metals in the soils from Santa Maria (inactive volcanism) were higher than those from Furnas (active volcanism), with the exception of Zn. The soils from Furnas, which have slightly lower pH and less % clay-silt than Santa Maria, will probably make such trace metals as Zn become more readily bioaccessible, and therefore pose a larger threat to living organisms inhabiting these soils.     

Seed Dispersal of the Palm Syagrus romanzoffiana by Tapirs in the Semi-deciduous Atlantic Forest of Argentina

Vertebrates play a fundamental role in the dispersal of Neotropical trees, generating different seed shadows according to their physical and behavioral features. Tapirs are capable of consuming great quantities of large fruits, and they defecate seeds far from parent trees. For instance, intact seeds of the palm Syagrus romanzoffiana have been found in tapir dung piles in the Atlantic Forest of southeastern Brazil, suggesting that tapirs effectively disperse this species. However, recruitment was not examined therein. We studied tapir endozoochory of large and medium seeds in the semi-deciduous Atlantic Forest of Argentina by examining dung piles found within Iguazu National Park. We recorded dung-pile positions to evaluate the spatial distribution. We also counted the number of juveniles in 2 × 2 m quadrats placed on old dung piles in latrines, beneath adults and in random sites to estimate recruitment levels. Syagrus romanzoffiana seeds were present in 98 percent of dung piles, averaging >200 seeds/dung pile, indicating that this species constitutes the main fruit component in the tapir's diet. Dung piles showed a clumped deposition pattern reflecting the use of latrines. Juvenile recruitment in latrines was 21 times higher than that of under-adult sites and 500 times greater than in random sites, and correlated with the frequency of use of the areas. We concluded that the lowland tapir is a major disperser of S. romanzoffiana . Given that this palm can be considered a keystone species, a disruption of this interaction might affect the entire community of frugivores in the long term.     

Endozoochory by free-ranging, large herbivores: Ecological correlates and perspectives for restoration

Seed dispersal via ingestion and defecation by large herbivores provides a possible aid for ecological restoration of plant communities, by connecting source communities of target species with habitat restoration sites. It is also a possible threat due to invasion of weeds, grasses or exotic species. Insight into the factors determining internal seed dispersal could therefore improve the management of grazed ecosystems.

We recorded viable seed density in cattle, sheep and pony dung samples and monitored dung pat colonisation in the field. In addition, we counted the distribution of dung pats in plots spread over all habitat units in our study site.

The three herbivore species appeared to disperse large quantities of many species (61 in total) from a variety of plant families, monocots as well as dicots. The density of viable seeds in herbivore dung and the colonisation of dung pats were positively correlated with Ellenberg nitrogen indicator values and seed supply, but not with seed mass or shape.

The results imply that many seeds are dispersed from high productive to low productive parts of the grazed area. In free-ranging systems, we therefore recommend enclosure and separate management of plant communities on nutrient-poor soils with high conservation interest. For habitat restoration sites we recommend integrated grazing only with target plant communities on nutrient-poor soils and not with plant communities on nutrient-richer soils.     

Deposition and decomposition of cattle dung and its impact on soil properties and plant growth in a cool-temperate pasture

Livestock dung provides an important direct pathway by which carbon and nutrients enter soils in pasture ecosystems and affects carbon and nitrogen cycling indirectly through changes in soil and plant properties. Here, we quantify dung deposition, decomposition, and the effects of dung on soil and plants in a Zoysia japonica grassland in Japan. We determined (1) the distribution of dung, (2) the mass loss rate of dung and the amount of carbon respired as CO₂, and (3) changes in soil properties and aboveground biomass of Z. japonica. Dung deposition was 4.0–9.7 g C and 0.4–1.0 g N m^?2 year^?1 and distributed patchily (Morishita’s I _δ  > 1). Most (71 %) of the carbon in dung deposited in June was lost within a single grazing period by aerobic decomposition, more than mass loss rate of Z. japonica litter in the first year (about 50 %), suggesting that grazing and defecation can accelerate carbon cycling compared with the typical litterfall–decomposition regime. Nitrogen in dung mass entered the soil as ammonium nitrogen and was nitrified. The spatiotemporal distribution of these processes corresponded to that of stimulated Z. japonica growth. These results suggested that dung deposition significantly affected the inorganic nitrogen status of soil and, therefore, the growth of Z. japonica. However, these effects were very restricted temporally (July–August) and spatially (within 10 cm from dung edge). Thus, such spatiotemporally restricted effects combined with the patchy distribution of dung may contribute to the heterogeneous structure of pasture ecosystems.