Effect of population density on reproduction of two sympatric dung beetle species,Aphodius haroldianus andA. Elegans (Coleoptera: Scarabaeidae)

The effect of population density on reproduction as well as the oviposition modes of two sympatric dung beetle species, Aphodius haroldianus (a low fecundity/high parental effort species) and A. elegans (a high fecundity/low parental effort species) was studied at a pasture in central Japan from 1982 to 1986.

1. The adult population density of A. haroldianus was high (>40 per dung pat). The density of A. elegans was low (<5 per dung pat). Oviposition of A. haroldianus was suppressed above the density of about 10 adults of conspecifics but not affected by the density of other species. The intra- and interspecific density effects on oviposition were not detected in A. elegans.
2. The number of A. haroldianus adults per pat decreased with dung age, while that of A. elegans increased until sixth day after deposition and then decreased. The number of eggs laid per pat was not different between 1-day-and 3-day-old pats for A. haroldianus. However, more eggs of A. elegans were found in 3-day-old pats than in 1-day-old ones.
3. In the both species, the amplitude of population fluctuations was not remarkable; the maximum/minimum ratio for five years being 2.3 for A. haroldianus and 2.9 for A. elegans. Different density dependent processes were suggested to function for the small fluctuation of population density between the two species, i.e. intraspecific density effect on oviposition for A. haroldianus and contest type competition for food among larvae of A. elegans.

     

Reproductive properties of two sympatric dung beetles,Aphodius haroldianus andA. Elegans (Coleoptera: Scarabaeidae)

1. Reproductive properties of two sympatric dung beetles, Aphodius haroldianus and A. elegans, were studied at a pasture in the central Japan.
2. Overwintered adults of A. haroldianus came flying to dung pats from May to early August and bred from June to July. A. elegans overwintered as larvae, new adults came to dung pats from mid May to mid June. After aestivation, they reproduced from October to November.
3. A. haroldianus was a species with low fecundity and large food reserve for larvae. Female of this species had short duration of residence and laid fewer number of eggs in one dung pat. On the other hand, A. elegans was a species with high fecundity and small food reserve for larvae. Female of this species had long duration of residence and laid larger number of eggs in one dung pat.
4. The daily egg production in A. haroldianus reached a peak (0.40 eggs/female/day) at middle of oviposition period, but, that in A. elegans reached a peak (6.49 eggs/female/ day) at the beginning of oviposition period. The mortality of A. haroldianus female occurred after the daily egg production reached a peak and then rapidly increased, whereas that of A. elegans began soon after the beginning of oviposition and then gradually increased.

     

Duration and benefits of biparental brood care in the dung beetle Onthophagus vacca (Coleoptera: Scarabaeidae)

Abstract.

• 1 Single males, single females or pairs of dung beetles, Onthophagus vacca, were released on artificial small (100 g) or large (1000 g) dung pats in the laboratory. Emigrating beetles were trapped at 12 h intervals, and the number and size of the brood chambers were recorded after each replicate.
• 2 Emigration of males was delayed if females were present in the same dung pats, whereas emigration times of females were independent of the presence or absence of males.
• 3 A residency of 60 h proved to be a threshold value. Females emigrating before this time did not breed, whereas those emigrating later had built at least two brood chambers.
• 4 Females paired with males built more brood chambers than single females.
• 5 The reproductive success of pairs was not influenced by the size of the dung pats.

     

Spatial distribution, resource utilisation and intraspecific competition in the dung beetle Aphodius ater

Competitive interactions in northern temperate dung beetles are poorly understood. This investigation therefore comprises a series of field and experimental work on a dung beetle species common in northern Europe, Aphodius ater, with special focus on intraspecific competitive interactions. The between-pat distribution of adult A. ater in relation to the age of sheep dung pats was studied in the field. The distribution of both sexes was contagious in the fresh pats but became more regular with increasing pat age. The successional occurrence of males and females did not differ, but immature females tended to occur in fresh pats while mature females were mainly found in older pats. With increasing age of pats, the egg load of females also increased. Egg-laying behaviour of the beetles was studied in laboratory experiments. The mean number of eggs laid per female per dung pat decreased with increasing beetle density. Thus, density-dependent processes seem to regulate resource utilisation with regard to breeding behaviour, resulting in equal exploitation of the available pats. Survival and weight of recently hatched beetles decreased with increasing initial density of eggs. Hence, in A. ater, competition between larvae for food within pats does occur. Received: 4 February 1998 / Accepted: 20 April 1998     

Dung burial strategies in some South African coprine and onitine dung beetles (Scarabaeidae: Scarabaeinae)

ABSTRACT.

• 1 The dung colonization and dung burial behaviour of twelve crepuscular/nocturnal tunnelling (paracoprid) species of beetles were examined in order to identify mechanisms which might facilitate resource (dung) partitioning. The species were selected from a diverse assemblage of dung beetles, the members of which coexist in the sandy-soil regions of Natal, South Africa.
• 2 The pattern of dung colonization in relation to dung age was examined in the field using baited pitfall traps. Some species, e.g. Onitis deceptor Peringuey, Catharsius tricornutus De Geer and Copris elphenor Klug, showed a marked preference for fresh dung (1–2 days old) whereas other species, e.g. O. viridulus Boheman and Copris fallaciosus Gillet, preferred older dung (3–7 days old).
• 3 Two distinct patterns of dung burial were recognized. In the Coprini, dung burial was complete within 24–48h of pad colonization, and the level of dung burial was similar in the laboratory and in the field. In the Onitini, dung burial occurred progressively over a 12-day period, although the timing of initiation of dung burial varied between species: in O. deceptor nearly all individuals had begun burial within 2 days of pad colonization, whereas only 20% of O. viridulus had commenced dung burial by that time. However, nearly all O. viridulus had buried substantial quantities of dung by day 12.
• 4 The mass of dung buried per pair by the larger coprine beetles (100–300 g) and onitine beetles (400–1000 g) suggests that there is potential for inter- and intraspecific competition, even in pads colonized by relatively few beetles. The colonization and use of dung of different ages are discussed as means of resource partitioning in relation to the relative abilities of species to compete for dung.

     

Asymmetric competition in cattle dung between two species of Onthophagus dung beetle and the bush fly, Musca vetustissima

Abstract.

• 1 Competition in cattle dung pads between two dung beetles, Onthophagus ferox Harold and Onthophagus binodis Thunberg, and the bush fly, Musca vetustissima Walker, was investigated in laboratory experiments, to determine why spring fly abundance in the field did not fall following the introduction of O. binodis.
• 2 At low beetle densities, the number of eggs laid by each species was reduced by the second species. A similar amount of dung was buried by each species alone or by both together.
• 3 At high beetle densities O. binodis egg production was substantially affected by each additional O.ferox, but O.ferox egg production was not affected by each additional O.binodis. Asymmetric competition occurred because O.ferox buried more dung than O.binodis, and a greater proportion in day 1 (pre-emptive dung burial).
• 4 O.ferox caused greater M. vetustissima egg-puparia mortality than O. binodis. Mortality mostly occurred in young M. vetustissima larvae less than 1 day old. Total egg-puparia fly mortality was correlated better with the dung buried on day 1 than dung buried on day 8 (pre-emptive dung burial). O.binodis did not add to fly mortality by O.ferox at high densities because of asymmetric competition between the beetles.

     

Evidence that sex‐specific signals may support mate finding and limit aggregation in the dung beetle Aphodius fossor

1. In temperate climates, dung is often colonised by several species of endocoprid (dwelling) dung beetles which use pats for feeding, shelter, and reproduction. 2. Endocoprid beetles aggregate even when offered patches (dung pats) of consistent age, size, and origin, suggesting that beetles themselves might influence the attractiveness of patches to members of their own species. Both pheromones, and physical changes to the structure of dung pats caused by colonising beetles have been suggested as mechanisms facilitating intraspecific aggregation, but neither of these hypotheses have been empirically tested. 3. Using a common European dung beetle (Aphodius fossor L.), we conducted a simple choice experiment designed to test whether (i) earlier colonisation by conspecifics could alter dung attractiveness and (ii) whether attraction was influenced by sex‐specific signals. 4. We found that female beetles are repelled by dung colonised by conspecific females and are attracted to dung colonised by conspecific males. Male beetles show no evidence of attraction or repellence for dung colonised by either sex. Neither in females nor males was uncolonised dung found to be significantly more or less attractive than predicted by non‐preference. 5. Our results suggest that for A. fossor male‐produced signals may support mate finding in patchy environments, and that female‐produced signals may serve to discourage subsequent colonisation by additional females.     

Mortality pattern and age structure in two carabid populations with different seasonal life cycles

1. Adult populations of two carabid species, a spring breeder, Carabus yaconinus, and an autumn breeder, Leptocarabus kumagaii, were studied in a lowland habitat (area: 16,500 m²) by pitfall sampling and mark-recapture method. The seasonal change in population number, age structure and mortality pattern were clarified and related to their seasonal life cycles.
2. The survival rate of new adults from emergence to the first reproductive season was on the average 48% in C. yaconinus and 55% in L. kumagaii. In C. yaconinus, beetles which emerged later in the season survived more to the first reproductive season than those emerged earlier. C. yaconinus which had survived the pre-reproductive period mostly died out during the first reproductive season of 5 months, and about 8% survived until the second season. In L. kumagaii which had a short reproductive period in autumn, about 20% survived to the second reproductive season, and a small proportion even to the third reproductive season. Accordingly, the proportion of old beetles in the reproductive population was higher in L. kumagaii than in C. yaconinus.
3. The reproductive population of C. yaconinus contained on the average 1600 beetles and produced 3300 new adults. the L. kumagaii population contained on the average about 530 reproductives in autumn, and about 820 beetles emerged in the following year. The recruitment rate of new adults of C. yaconinus was higher than that of L. kumagaii, and this resulted in its higher population density. In L. kumagaii, however, the high adult survivorship and iteroparous reproduction were important for its population growth.
4. Relationship between seasonal adaptation and demographic strategies in the carabid populations were discussed.

     

Dung beetle community composition affects dung turnover in subtropical US grasslands

1. An important service in many ecosystems is the turnover and degradation of dung deposited by cattle. Dung beetles are the primary group of insects responsible for dung turnover, and factors affecting their abundance and distribution thus impact dung degradation. Lands lost to grazing due to dung buildup and pasture contamination total millions of acres per year in US pastures.
2. We evaluated the structural differences in dung beetle assemblages in natural grasslands versus a managed agroecosystem in subtropical southeastern Florida (USA). We measured the direct effect of dung longevity when dung beetle fauna normally inhabiting dung pats were excluded.
3. Our results indicate dung beetle abundance, functional diversity, and species richness have a substantial impact on the rate of dung turnover in subtropical pastoral lands with ~70% of dung removed from the soil surface after three months. Functional diversity and evenness did not have a significant positive effect on dung removal in managed, versus natural grasslands demonstrating a strong relationship between dung beetle assemblage composition and delivery of a key ecological process, dung degradation.
4. We suggest the importance of trees, which provide a thermal refuge for beetles, should be dispersed within matrixes of open pasture areas and within proximity to adjacent closed‐canopy hammocks to facilitate the exchange of dung beetles between habitats and therefore maintain the provisioning of dung degradation services by dung beetle assemblages.

     

Patterns of daily flight activity in onitine dung beetles (Scarabaeinae: Onitini)

Different species of African dung beetles emerge from the soil at characteristic times of the day to fly and colonize the freshly-deposited dung of mammalian herbivores. Onitine dung beetles in their natural habitat displayed one of five distinctive daily flight behaviours: dusk crepuscular (Onitis alexis Klug, O. caffer Boheman, O. fulgidus Klug, O. tortuosus Houston, O. vanderkelleni Lansberge, O. westermanni Lansberge); dusk/dawn crepuscular (O. pecuarius Lansberge and O. viridulus Boheman); dusk/dawn crepuscular and nocturnal (O. aygulus (Fabricius), O. mendax Gillet, O. uncinatus Klug); late afternoon-dusk and dawn-early morning [Heteronitis castelnaui (Harold)]; or diurnal flight activity [O. belial (Fabricius), O. ion (Olivier)]. These diagnostic daily flight behaviours span a light intensity range of over 6 orders of magnitude and have been retained in selected species introduced into Australia. Ambient light intensity appears to be the primary determinant of the daily flight period in onitine dung beetles. Because the dung of mobile herbivores is rapidly exploited by onitine species for feeding and breeding purposes, different flight behaviours result in a spatial and temporal partitioning of species in the local dung beetle community. The timing of flight may contribute to, or lead to avoidance of, competition between species which may ultimately affect colonization success. Many onitines show a strong preference for dung of specific herbivores, which may further reduce interspecific competition. All crepuscular-nocturnal species examined raised their thoracic temperatures endothermically to between 35°C and 40°C before the onset of flight. In O. aygulus the thoracic temperature excess was as large as 19.3°C. The thermal threshold below which the frequency of flight onsets drops off rapidly is about 12°C for O. aygulus and 17°C for O. alexis and O. pecuarius. Radiant loss of body heat during cool nights and dawns may explain why smaller species (<0.4 g body weight), in particular, are adapted behaviourally so that they fly only during the day or early dusk.     

Effects of dung beetles on seedling emergence from herbaceous seeds in the dung of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) in a temperate Japanese grassland ecosystem

The effects of dung form and condition and of dung beetles on the emergence of seedlings from herbaceous seeds in sika deer dung were examined in a temperate grassland ecosystem dominated by Zoysia japonica and Hydrocotyle maritima. I conducted field experiments to compare seedling emergence between dung exposed to dung beetles and intact dung using both dung pellets and pats during a typical rainy month (June) and the hottest, drier month (August), when large numbers of seeds of the dominant species were present in the dung. The exposed dung was immediately attacked and broken up by dung beetles, whereas dung protected from the beetles remained intact. In June, at least 12 herbaceous species, including Z. japonica, H. maritima, Mazus pumilus, and Plantago asiatica, emerged from the dung, versus at least six species in August. Decomposition rates of the pellets in June and decomposition scores of the pats in June and August were positively correlated with the number of emerging seedlings, suggesting that the acceleration of decomposition by dung beetles can positively affect seed germination. In this system of interactions among sika deer, herbaceous plants, and dung beetles, sika deer dung prevented seeds from germinating, and beetles had an indirect positive effect on seedling emergence by accelerating decomposition of the dung, although the extent of the effect may depend on the dung type, plant species, and environmental factors.     

Herbivore dung as food for dung beetles: elementary coprology for entomologists

1. How do dung beetles and their larvae manage to subsist on herbivore dung consisting of plant remains that are at least partly indigestible, mixed with various metabolic waste products? To clarify what is known and not known about this basic aspect of dung beetle biology, the present review summarises information on dung composition and discusses the feeding of beetles (food: fresh dung) and larvae (food: older dung) in relation to this information. 2. There is 70–85% water in typical fresh dung, and undigested lignocellulose or ‘fibre’ constitutes about 70% of the organic matter which also contains 1.5–3% N. About 75% of this is ‘metabolic faecal nitrogen’, mostly associated with dead and alive microbial biomass. As all essential amino acids and cholesterol are probably present, additional synthesis by microbial symbionts may not be needed by the beetles. 3. Beetles minimise the intake of lignocellulose by filtering fibre particles out of their food which is probably microbial biomass/debris with much smaller particle size. Excess fluid may be squeezed out of this material by the mandibles before ingestion. 4. All larvae are bulk feeders and unable to filtrate, but little is known about the composition of their food, i.e. older dung in pats or underground brood masses. Larvae in dung pats may depend on easily digestible dung components, probably microbial biomass, whereas the nutritional ecology of larvae in brood masses is still not understood. Unravelling the composition of their food might answer some of the so far unanswered questions.     

Ecosystem functioning in relation to species identity,density, and biomass in two tunneller dung beetles

1. Species abundance, biomass, and identity are the main factors that influence ecosystem functioning. Previous studies have shown that community attributes and species identity help to maintain natural ecosystem functioning. 2. This study examined how species identity, biomass, and abundance in dung pats (i.e. density) of dung beetles affect multiple ecological functions: dung removal, seed dispersal, and germination. Specifically, two species of tunnellers were targeted: Onthophagus illyricus (Scopoli, 1763) and Copris lunaris (Linnaeus, 1758). In accordance with their natural abundance, densities ranging from 10 to 80 individuals were considered for O. illyricus, and those from two to eight were considered for C. lunaris, spanning the total biomass per treatment from 0.22 to 1.76 g. 3. Results showed that, even at higher abundance, O. illyricus is not as efficient as C. lunaris. These results show that species identity, biomass, and density are crucial factors for maintaining ecosystem functioning. The combined effect of species identity and density/biomass facilitated dung removal and seed dispersal. Conversely, species identity is the only relevant factor for germination. Moreover, relationships among functions depend on the species investigated: C. lunaris showed a positive correlation between dung removal and seed dispersal, whereas O. illyricus showed a positive correlation between germination and dung removal. 4. In conclusion, optimal ecosystem functioning depends on multiple factors, such as density and species identity, and thus also on body size, nesting strategies and ecological functions investigated. Moreover, the loss of larger and efficient species cannot be compensated by higher abundances of small species.     

Effects of herd management and the use of ivermectin on dung arthropod communities in grasslands

Agroecosystems represent a large geographical footprint in most terrestrial landscapes, and management decisions within these systems affect their function in species conservation. We evaluated the effects that rangeland management systems (based on stocking density, rotation frequency, and the number of avermectin applications) have on conserving the dung arthropod community in the Northern Great Plains of North America. Comprehensive bioinventories of arthropods were collected from 16 rangelands using core samples of dung pats. Ivermectin was quantified in pats from each ranch using enzyme-linked immunosorbant assay (ELISA). Arthropods in dung were abundant (116,244 specimens) and diverse in eastern South Dakota (172 morphospecies). Rangelands managed with more regenerative practices (frequent rotation at high stocking densities and lack of ivermectin applications) had greater species richness, diversity, predator species abundance, and dung beetle abundance than more conventionally managed rangelands. Ivermectin quantity in cattle pats was negatively correlated with dung beetle abundance and diversity. This work shows that herd management (specifically high-intensity, frequent rotational grazing and eliminating prophylactic ivermectin use) that aims to mimic intensive grazing of large migrating herds of ruminants can foster dung arthropod community structure, a key trait correlated with nutrient cycling, pest suppression, and productivity of cattle-grazed rangelands.     

Some effects of rearing the yellow dung fly Scatophaga stercoraria in cattle dung containing invermectin

Invermectin was added to cattle dung in controlled concentrations like those found in the pats of injected cattle, and the medium was used to rear larvae of the dung fly Scatophaga stercoraria. Ivermectin at 0.036 ppm (wt/wet weight) debilitates 50% of the larvae within 48 h. At 0.015 ppm, 50% of the larvae are unable to pupariate, while at 0.001 ppm, 50% of the larvae fail to reach the adult stage. Adults produced from larvae reared in pats containing 0.0005 ppm invermectin show high levels of fluctuating asymmetry in wing characteristics as well as deformities in the wing veins themselves. The data are discussed in relation to the effects of excreted ivermectin on pastureland biology.     

Experimental warming alters the relative survival and emigration of two dung beetle species from an Australian dung pat community

To preserve insect‐mediated ecosystem services under ongoing climate change, it is necessary to first understand the impact that warming will have on the insects that provide or mediate these services. Furthermore, because responses of a species may be modified by interactions with competitors, it is informative to examine warming effects on organisms and service provision under competition. Dung beetles provide numerous services to agriculture by burying the manure of other animals. To understand the potential impacts of climate warming on ecosystem service provision, we exposed two dung beetle species (Sisyphus rubrus and Euoniticellus fulvus (Coleoptera: Scarabaeidae)), occurring together in the same experimental pats, to warming and measured reproduction (dung ball production and burial, brood production, and egg laying), pat departure behaviour and survival of both species. These two species are likely competitors in pastures in northern New South Wales. To simulate climate warming, we used custom‐built chambers to add offsets (+0, +2 or +4°C) to field recorded, diurnally fluctuating baseline temperatures. There was no direct effect of increased temperature on any measured trait in either species. We did find however that the relative survival of the two species depended on temperature; S. rubrus had a higher probability (resulting in greater odds) of surviving than E. fulvus in the +0 and +4°C offset chambers, but not in the +2°C offset chambers. Likewise, the relative likelihood of the different species leaving a dung pat was temperature dependent; in the +2°C offset chambers, E. fulvus were more likely to leave than S. rubrus, but not in the +0 and +4°C offsets chambers. Our results highlight that it may be important for future studies to consider warming effects on relative survival and emigration because such effects could potentially lead to changes in dung beetle species composition.     

松嫩草地牛粪中大型节肢动物种类组成及种群动态变化

研究并分析了松嫩草地牛粪中大型节肢动物种类组成及动态变化.结果表明,在草地牛粪中共调查到大型节肢动物7目20科43种,以鞘翅目、膜翅目和双翅目为主.依其食性不同,分为粪食性型(8科19种)、腐食性型(4科4种)、植食性型(4科6种)、捕食性型(5科11种)、杂食性型(2科5种)、尸食性型(1科1种)和寄生性型(1科1种).根据鞘翅目金龟总科中营粪食性型生活的粪甲虫对牛粪利用方式的不同,分为外生粪甲虫(6种)、内生粪甲虫(5种)、远生粪甲虫(2种)和偷窃寄生粪甲虫(1种).除了双顶嗡蜣螂,牛粪中大型节肢动物数量与鲜牛粪重量具有显著正相关关系(p＜0.01).粪食性型种类数和个体数量随鲜牛粪堆置时间的延长而迅速减少,7d后消失,其个体总数量与鲜牛粪堆积时间呈显著的负相关关系(p＜0.01),而与牛粪含水量呈显著正相关关系(p＜0.01),各种大型节肢动物因其食性不同在鲜牛粪中占据不同的生态位.鲜牛粪中大型节肢动物以粪食性型占绝对优势,而干牛粪中则以捕食性型和杂食性型为主.牛粪中大型节肢动物种群季节动态变化受温度、降水量与牛粪含水量影响,鲜牛粪中大型节肢动物优势种为直蜉金龟(69.25%)和肖秋家蝇与东方角蝇幼虫(25.12%).4、5、6月份和9月份,优势种为直蜉金龟,7月和8月份,优势种为蝇幼虫.     

Molecular analyses of methanogen diversity associated with cattle dung

Methanogen communities were characterized in cattle dung of different ages by using a culture-independent approach. Community structures were determined by the phylogenetic analyses of methyl-coenzyme M reductase A (mcrA) clones of fresh, 8-month-old, and 24-month-old-dry dung samples. The clones in the mcrA libraries of fresh and 8-month old dung samples were identified as belonging to Methanomicrobiales, Methanobacteriales, and Methanosarcinales. However, clones in the library of 24-month-old dung were not affiliated to Methanomicrobiales. Anaerobic digestion of 2-month-old dung produced only 15% less methane compared to fresh dung which indicated the possibility of using dry dung to fuel the biogas plants in areas where unavailability of fresh dung hinders their continuous functioning. Our results first time showed the presence of viable methanogens in dry cattle dung stored for prolonged periods of time.     

Effect of dung burial by the dung beetle Bubas bison on numbers and viability of Cryptosporidium oocysts in cattle dung

Cryptosporidium oocysts were inoculated into fresh dung (∼1.2 × 10⁴ oocysts per gram wet weight) and fed to dung beetles to assess the effect of dung burial by the dung beetle Bubas bison on the distribution of the oocysts in small cores of soil in the laboratory. The experiment consisted of five replicates of each of two treatments; controls (dung but no dung beetles) and the experimental treatment (inoculated dung and seven pairs of dung beetles). After 5 days, when approximately 90% of the dung was buried, the surface and buried dung was recovered and subsampled. The oocysts in the subsamples were recovered and enumerated using qPCR. Oocyst viability was evaluated using an assay based on the exclusion or inclusion of two fluorogenic vital dyes, 4′,6-diamidino-2-phenylindole (DAPI) and propidium iodide (PI). Results revealed that overall 13.7% of oocysts remained on the surface and 86.3% of oocysts were buried. The viability of oocysts in buried dung was only 10% compared to oocysts the surface dung (58%). Therefore, widespread dung burial by B. bison during the winter months could substantially reduce the numbers of Cryptosporidium oocysts available to be washed into waterways following winter rains.     

荒漠草原马粪中粪金龟子组成及分解作用

2007年4-10月,采用野外和室内实验相结合的方法,研究了荒漠草原马粪中的粪金龟子群落组成和对马粪分解的影响.共捕获粪金龟子13528头,隶属于3科4属26种,优势种为符号蜉金龟(Aphodius comma),占总捕获量的88.5%,不同季节粪金龟子群落组成明显不同;在鲜马粪中活动的粪金龟子以粪居型蜉金龟为主,在鲜马粪堆置3d时种数和个体数达到最高,6d后全部离开;马粪经过136 d分解,残留率为52.5%,残留粪样中氮和磷释放较慢,钾释放较快;结合室内分解实验的结果,认为粪金龟子主要通过在马粪中活动对其分解产生影响,并与风蚀作用成为荒漠草原马粪块破碎的重要动因,主要发生在春季和秋季.