Patch size and colonisation patterns: an experimental analysis using north temperate coprophagous dung beetles

The relationship between dung pad size and both adult colonisation and larval development was investigated in an assemblage of north temperate dung beetles ( Geotrupes. Aphodius and Sphaeridium ) using both dung pads and baited pitfall traps. Wet weight of 22-day-old natural dung pads was found to vary widely in the field (< 100 g – >1000 g). Across all sampling dates in field experiments, dung pad size had a significant influence on dung beetle biomass sampled from pads. Closer examination of experimental dung pads on the second day after deposition, when beetle biomass was at a maximum, revealed not only a general positive relationship between pad size and dung beetle biomass but, more importantly, a positive relationship between dung pad size and dung beetle density (dung beetle biomass per unit dung volume). There was a strong trend for Aphodius species richness to increase, and maintain higher values for longer periods of time, in larger pads. Although dung pad and pitfall trap samples could differ in the actual numbers of beetles captured, the relationship between different dung sizes and dung beetle biomass was similar, indicating that the phenomenon is largely related to immigration processes. Pat residence times oi A. rufipes in the laboratory were significantly positively correlated with dung pad size. In two field experiments, positive correlations were found between dung pad size and numbers of larvae in pads of different sizes and in one of these experiments, larval densities (numbers per unit dung volume) were significantly and positively correlated with dung pad size. In one experiment, Aphodius larvae in the early stages of development were found to preferentially occupy the basal area of dung pads. We discuss the implications of our findings in the context of resource utilisation by north temperate dung beetles.     

Resource quality and the colonisation and succession of coprophagous dung beetles

In order to examine the degree of resource selectivity in a north temperate dung beetle assemblage und to identify major parameters that influence such selectivity, the occurrence of adult dung beetles ( Aphodius. Geotrupes and Sphaeridium ) in up to five different types of dung over a period of up to 25 d was examined in a series of field experiments using standardised dung pats. There were significant physical and chemical differences in dung quality between dung types and over time during succession. Dung beetle species showed distinct preferences for particular types of dung which were generally similar in data sets from both pitfall traps and dung pat samples. Species also showed distinct patterns of successional occurrence. Ordinations produced by Canonical Correspondence Analysis, based on species occurrences in dung types and over time, usually selected dung pat age as the most important environmental variable influencing dung beetle assemblages. Dung quality parameters contributed a significant element of structure to the species ordinations but ordinations using dung types or dung quality parameter values as the environmental variables were very similar in terms of sample and species placement within the ordination for each data set. Most importantly the CCA ordinations clearly grouped species according to their breeding behaviour. Early-successional species laid eggs in the soil, or in silken egg cocoons, which allowed them to exploit wet dung. Mid and late-successional species laid eggs in the dung pat: late-successional species could exploit normally wetter dung types than mid-successional species, probably due to increased crust formation and drying as the dung pat ages. Thus, species appear to be differentially adapted to exploit varying types of dung microhabitats. Therefore, where two or more species of large herbivores are present, dung quality preferences probably constitute an important niche dimension.     

Spatial and temporal variation in species composition of dung beetle assemblages in southern Ireland

Abstract. 1. This study attempts to identify the main community characteristics that contribute to variability in dung beetle assemblage composition and structure across a range of spatial and temporal scales.
2. Dung beetle assemblages ( Aphodius , Sphaeridium, and Geotrupes species) were monitored by dung-baited pitfall trapping at 10-day intervals during the seasonally active period at eleven sites in southern Ireland. Three of the sites were monitored over at least 2 years between 1991 and 1996.
3. Although the species composition of the above taxonomic groups was comparable among sites and years, relative abundances of component species varied considerably. Detrended correspondence analysis ordinations indicated a similar level of variability in dung beetle assemblage structure among years, and among sites ≈1–180 km apart.
4. Processes that may contribute to spatio-temporal variability in dung beetle assemblages are discussed, and strategies for future research are suggested.     

Testing for seasonal displacement in a dung beetle guild

The temporal overlap between seasonal distribution of adults of 14 congeneric dung beetle species ( Aphodius spp) was determined during one year, based on 11122 captures The mean seasonal distribution of adults (mean of the sampling dates weighted by the abundance value) was analyzed to elucidate deviations from ram-domness In addition, the effect of the shape of seasonal distribution of adults was investigated In both analyses randomization methods were used A clear trend exists maximizing the temporal displacement between middle points of chronologically consecutive species newly emerging species appear every 26 ± 16 days This trend minimizes the number of species pairs with no or very little temporal overlap In contrast, the mean temporal overlap of the whole guild seems unaffected, with some species pairs showing a large temporal overlap Whether the observed pattern can be attributed to interspecific competition must be addresed experimentally     

Dietary shifts of sympatric buteos during a prey decline

Summary Diets of nesting Red-tailed Hawks (Buteo jamaicensis) and Ferruginous Hawks (Buteo regalis) were sampled before and after a decline in the hawks' principal prey species. Diets of pairs that shared their foraging ranges with interspecifics were contrasted with those of pairs whose home ranges did not overlap with interspecifics. Current theory predicts that diets should diverge during prey shortages and that overlap should be especially reduced in ranges shared with interspecifics. Species composition of the two hawks' diets diverged during the prey shortage, but the divergence was most pronounced in hawks that did not share foraging ranges with interspecifics. In contrast to predictions, the two species converged on similar-sized prey during the prey shortage. Available data on differences in habitat composition and prey availability at the sample sites did not explain the deviations from the predicted response. Although our findings on diet shifts differ from those of most other studies, the implications are consistent with observations of others who have studied assemblages of mammal-feeding raptors. We conclude that diet composition of the hawks we studied was not directly affected by interspecific competition and that shifts in diet overlap during prey shortages do not necessarily imply that interspecific competition is occurring.     

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     

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.     

Distinguishing food from fodder through the study of charred plant remains: an experimental approach to dung-derived chaff

Distinguishing human food from fodder in the archaeobotanical record is a difficult task. These categories are culturally defined and, therefore, not obvious from the plant species represented in archaeological samples, while context, such as storage area or container is not necessarily distinctive for each category. While grain can be consumed as either human food or fodder, depending on the particular needs of a given society, chaff is less suitable for human consumption and more appropriate for other uses, one of which is fodder. Archaeobotanical assemblages from two Late Neolithic sites in northern Greece (Makriyalos and Makri) are dominated by glume-wheat chaff, often in association with fig seeds. This material may well represent the remains of dung, but it was not known if the cereal component was eaten by livestock or mixed with the dung to form dung cakes. Our contribution presents the results of an experiment involving the feeding of einkorn grain and chaff (whole and pounded spikelets) and figs to goats to investigate the effect of livestock digestion on plant material. The implications of this work for the interpretation of archaeobotanical assemblages that may derive from animal dung are considered using the Greek Neolithic assemblages. Electronic Supplementary Material Supplementary material is available for this article at     

Temporal coexistence of dung-dweller and soil-digger dung beetles (Coleoptera, Scarabaeoidea) in contrasting Mediterranean habitats

The western part of the Mediterranean basin is a transitional biogeographical region for the distribution of the representatives of the main guilds of dung beetles; towards the south, Aphodiinae (dung-dwellers) become scarce, whereas northwards Scarabaeinae (soil-diggers) progressively disappear. The number of species in local dung beetle assemblages is enhanced by this double faunistic contribution. Annual dung beetle assemblages were sampled in two sub-Mediterranean sites, which differed by 600 m in elevation, in order to determine the phenological dynamics related to the way of using dung (dung-dwellers/Aphodiinae vs. soil-diggers/Scarabaeinae and Geotrupinae). Aphodiids were active all year round, although they were affected by summer drought and, at high elevation, by the length of the cold season. This reduced activity was related to an impoverishment of Aphodiinae and to reduced temporal segregation between species. In contrast, soil-diggers were not active all year round and showed different species assemblages in the two sites. An extension of the activity period of these beetles was observed due to the occurrence of cold resistant species at high elevation. Our results suggested that the occurrence of soil-diggers seemingly did not affect the seasonality of dung-dwellers; their local abundance showed no negative correlation and, most importantly, phenological differences between dung-dwellers were always significantly higher than the seasonal differences between dwellers and diggers.     

Use of habitat resources by scarab dung beetles in an savanna

In the Queen Elizabeth National Park, Uganda, we compared the scarab beetle assemblages in the dung of three wild ungulates (African buffalo, a ruminant foregut fermenter; hippopotamus, nonruminant foregut fermenter; and warthog, nonruminant hindgut fermenter). Dung was collected from two sandy-clay soils with different percentage of coarse sand. We aimed at investigating habitat resource selection by dung beetle species within a savanna natural contest with abundant and diverse food availability. Analyses were performed to detect differences for dung beetle assemblages in abundance, diversity, functional groups. Species richness in the three dung types and in the two soil types was similar. However, warthog dung and sandy-rich soil appeared the preferred habitat resources, in terms of abundance and biomass, while hippopotamus dung hosted the lowest values for these parameters. The analysis of functional groups revealed that slow-burying tunnellers held the major role, both in terms of abundance and biomass, and were mainly found in warthog dung.     

内蒙古典型草原家畜粪中的粪金龟子群落特征

2008年6月-2009年9月,在内蒙古典型草原放牧区,分析了马、牛和羊鲜粪中粪金龟子的物种多样性、群落结构、功能群和取食偏好.共捕获粪金龟子7805头,隶属于3科4属32种,优势种为符号蜉金龟(Aphodius comma)、游荡蜉金龟(Aphodius erraticus)和泥蜉金龟(aphodius sordec...     

Differences in temporal utilization patterns of dung pats among three scarabaeid dung beetles

1. Differences in the temporal utilization patterns of dung pats were investigated among three dung beetle species, Aphodius haroldianus, Onthophagus lenzii and Liatongus phanaeoides, and density effects of three species on emigration from a dung pat were compared.
2. A. haroldianus preferred fresh dung pats, immigrated to fresh pats and then emigrated from those pats quickly (mean duration of residence 48.4 h), whereas L. phanaeoides immigrated to older dung pats (over 7 days after deposition) as well as fresh ones and remained in the pats for the longest time (165.6 h).O. lenzii showed intermediate behavior in this respect (63.3 h).
3. In A. haroldianus, the presence of conspecifics induced faster emigration than presence of the other two species. L. phanaeoides showed the opposite relationship. In O. lenzii, the response to conspecifics was intermediate to those of the other two species.

     

A functional classification for analysis of the structure of dung beetle assemblages

Abstract. 1. Regional scarabaeid dung beetle assemblages in southern Africa may contain over 100 species, ranging in live weight from 10 mg to 10 g. These show a wide variety of dung-use and reproductive strategies.
2. To facilitate analysis of these diverse assemblages, a system of classification analogous to guilds is proposed. Scarabaeid dung beetle species are allocated to one of seven functional groups (FGs) according to the way they use and disrupt dung. Each group therefore contains a set of species which are functional analogues of each other. This classification provides a conceptual framework within which to analyse the structure of dung beetle assemblages and the interactions between dung beetles and other dung-breeding species such as coprophagous flies.
3. There is a clear hierarchy of functional groups in their ability to compete for dung. Competitively dominant groups such as the large ball rollers (FG I) and fast-burying tunnellers (FG III) are mostly large, aggressive beetles which rapidly remove dung from the pad. The smaller ball rollers (FG II) are also effective competitors for dung. Subordinate groups are those which bury dung slowly over many days (FG IV and V) and those which breed inside the pad (FG VII, endocoprids). Kleptocoprids (FG VI) breed in dung buried by other beetles and so are not part of the hierarchy.
4. The use of this classification is illustrated by reference to three contrasting assemblages of dung beetles in a summer rainfall region of southern Africa. The potential of these beetles for biological control of dung-breeding flies is discussed.     

Dung beetle movements at two spatial scales

To understand the dynamics of spatially structured populations, we need to know the level of movements at different spatial scales. This paper reports on Aphodius dung beetle movements at two scales: movements between dung pats within pastures, and movements between pastures. First, I test an assumption common to many recent models of spatially structured populations – that the probability of an individual moving between habitat patches decreases exponentially with distance. For dung beetles, I find sufficient evidence to reject this assumption. The distribution of dispersal distances was clearly leptokurtic, with more individuals moving short and long distances than expected on the basis of an exponential function. In contrast, the data were well described by a power function. I conclude that dung beetle movements include an element of non-randomness not captured by the simplistic exponential model. The power function offers a promising alternative, but the actual mechanisms behind the pattern need to be clarified. Second, I compare several species of Aphodius to each other. Although these species occur in the same network of habitat patches, their movement patterns are different enough to result in a mixture of different spatial population structures. Movements between pastures were more frequent the larger the species, the more specific its occurrence in relation to pat age, and the more specialized it is on cow dung and open pasture habitats. Within pastures, all species form "patchy" populations, with much movement among individual pats.     

Effects of dung-pad conditions and density on coprophagous beetle assemblages in a Mediterranean rangeland

Dung beetles highly depend on the ephemeral microhabitat dung which is food resource and larval habitat at the same time. Environmental conditions surrounding a dung pad, such as vegetation structure, have an impact on dung beetle assemblages. We investigated the influence of dung conditions and surrounding habitat characteristics on Mediterranean dung beetle assemblages in a permanently grazed landscape in northern Sardinia. We sampled the dung beetle assemblages of donkey and horse dung in three different vegetation types and assessed species richness and abundance of dung beetles. Species richness was determined by dung and surrounding habitat conditions, whereas abundance was solely affected by dung conditions. However, species richness and abundance decreased with increasing dung density. The effect of dung density on species richness varied depending on vegetation type, with dry grassland exhibiting the highest number of dung beetles species at high dung density. Species composition in dung pads was influenced by abiotic factors with dwellers being negatively affected by increasing dung-pad temperature. Our results underline the importance of diverse vegetation, particularly with respect to the complexity of vegetation which interrelates with the microclimate. Furthermore, our findings illustrate the negative effect of high dung densities on dung beetle assemblages, suggesting that the degree of the intensity of use by grazing animals is important when considering measures for the conservation of dung beetles.     

Oviposition of the dung beetle Aphodius ater in relation to the abundance of yellow dungfly larvae (Scatophaga stercoraria)

1. The dung beetle Aphodius ater and the yellow dungfly Scatophaga stercoraria are temporally co-occurring species in sheep dung, which they use for reproduction and nutrition ( A. ater ) or for reproduction only ( S. stercoraria ) during the spring in northern Germany. Scatophaga stercoraria uses fresh sheep dung pellets a few hours old for oviposition, whereas A. ater lays eggs into 2–10-day-old pellets. In the present study, the egg laying behaviour of A. ater in sheep dung in relation to the presence of larvae of S. stercoraria was investigated experimentally.
2. Choice experiments, based on examining the egg laying behaviour of beetles in 2- and 4-day-old pellets with and without high and low densities of fly larvae, showed the following. In 2-day-old pellets, the beetles did not distinguish between pellets without fly larvae or with fly larvae at low larval density but avoided laying eggs into pellets with a high larval density. In 4-day-old pellets, the beetles always preferred to lay their eggs into pellets without fly larvae, regardless of larval density.
3. The influence of different densities of larvae of S. stercoraria on dung depletion was examined by measuring the dry weight, organic matter content and organic nitrogen content of the remaining dung after larval development. The presence of the larvae led to a reduction in all three parameters.
4. The beetles' behaviour of laying eggs into older pellets, and their awareness of the presence of high densities of fly larvae, enables them to avoid egg laying into pellets that will have been depleted by fly larvae before the beetle larvae have finished their development.     

MULTIDIMENSIONAL ANALYSIS OF RESOURCE UTILIZATION IN ASSEMBLAGES OF RANGELAND GRASSHOPPERS (ORTHOPTERA: ACRIDIDAE)

Abstract  A study was conducted in a typical steppe area in central Inner Mongolia to determine resource utilization in time, space, and food by grasshopper assemblages. Pattern of use for these resources was analyzed from three vegetation types by examining species-specific niche breadths, overlaps and partitions of resource utilization as well as population sizes. Popular niche indices and the principal components analysis (PCA) procedure were used to indicate the interrelations among grasshopper species. In the heterogeneous grassland environments resulting from livestock grazing, eleven grasshopper species were categorized respectively based on the 3-dimensional niche segregation and overlap degree. Population sizes were determined by the calorific values of their realized population biomass. Comparison in niche breadth displayed that species which had a broad niche along one dimension at least had a narrow niche along another one. There were sufficient differences between species for overall overlap associated with resource use to explain coexistence in the assemblages by resource segregation. Distinct rule of niche complementarity in overlap was not found. Evidence from this study implied that the species-specific use of resource may be due primarily to grasshopper adaptive differentiation and to coevolutionary interactions between grasshopper and plant rather than interspecific competition among grasshopper species.     

草原蝗虫种团资源利用的多维分析(直翅目:蝗科)(英文)

本文研究了内蒙古典型草原蝗虫在时间、空间和食物资源利用方面的特点。对于这些资源利用的格局是通过对存在于三种类型植被中蝗虫种的生态位宽度、资源利用的分化和重叠以及每种蝗虫的种群大小来分析的。常用的生态位指数和主成份分析技术用来阐明蝗虫种间的关系。在放牧形成的异质性草原环境中,11种蝗虫根据它们在三维生态位方面的分化和重叠程度被划分成数个种组。种群的大小是根据种群实际生物量所含热能值来确定的。生态位宽度比较说明,在某一维生态位宽的种,在另一维上必定变窄。生态位重叠的差异说明共存的蝗虫种在资源利用上的分化。生态位重叠的补偿规律在本项研究中并不明显。本项研究结果说明,蝗虫种资源利用的分化主要归因于蝗虫的适应性分化和蝗虫——植物协同进化的相互作用关系,而不是蝗虫种间竞争所造成的。     

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.     

Stridulation in Aphodius Dung Beetles: Behavioral Context and Intraspecific Variability of Song Patterns in Aphodius ater (Scarabaeidae)

Aphodius dung beetles produce substrate vibrations by means of an abdomino-alary stridulatory organ. Applying a method that allows the recording of Aphodius vibrations under natural conditions in a small amount of dung, the stridulatory behavior of Aphodius ater was investigated. Male A. ater are acoustically active, while females rarely stridulate. Males have a complex song, which consists of a series of different patterns that are displayed in a specific order over a considerable time when a female is encountered in the dung. Different populations show the same stridulatory patterns but individual variability is high and males display songs with differing complexity. It is hypothesized that females use the information within the song in the context of mate choice.