Spectral information as an orientation cue in dung beetles

During the day, a non-uniform distribution of long and short wavelength light generates a colour gradient across the sky. This gradient could be used as a compass cue, particularly by animals such as dung beetles that rely primarily on celestial cues for orientation. Here, we tested if dung beetles can use spectral cues for orientation by presenting them with monochromatic (green and UV) light spots in an indoor arena. Beetles kept their original bearing when presented with a single light cue, green or UV, or when presented with both light cues set 180° apart. When either the UV or the green light was turned off after the beetles had set their bearing in the presence of both cues, they were still able to maintain their original bearing to the remaining light. However, if the beetles were presented with two identical green light spots set 180° apart, their ability to maintain their original bearing was impaired. In summary, our data show that ball-rolling beetles could potentially use the celestial chromatic gradient as a reference for orientation.     

The role of the sun in the celestial compass of dung beetles

Recent research has focused on the different types of compass cues available to ball-rolling beetles for orientation, but little is known about the relative precision of each of these cues and how they interact. In this study, we find that the absolute orientation error of the celestial compass of the day-active dung beetle Scarabaeus lamarcki doubles from 16° at solar elevations below 60° to an error of 29° at solar elevations above 75°. As ball-rolling dung beetles rely solely on celestial compass cues for their orientation, these insects experience a large decrease in orientation precision towards the middle of the day. We also find that in the compass system of dung beetles, the solar cues and the skylight cues are used together and share the control of orientation behaviour. Finally, we demonstrate that the relative influence of the azimuthal position of the sun for straight-line orientation decreases as the sun draws closer to the horizon. In conclusion, ball-rolling dung beetles possess a dynamic celestial compass system in which the orientation precision and the relative influence of the solar compass cues change over the course of the day.     

The Dung Beetle Dance: An Orientation Behaviour?

An interesting feature of dung beetle behaviour is that once they have formed a piece of dung into a ball, they roll it along a straight path away from the dung pile. This straight-line orientation ensures that the beetles depart along the most direct route, guaranteeing that they will not return to the intense competition (from other beetles) that occurs near the dung pile. Before rolling a new ball away from the dung pile, dung beetles perform a characteristic "dance," in which they climb on top of the ball and rotate about their vertical axis. This dance behaviour can also be observed during the beetles' straight-line departure from the dung pile. The aim of the present study is to investigate the purpose of the dung beetle dance. To do this, we explored the circumstances that elicit dance behaviour in the diurnal ball-rolling dung beetle, Scarabaeus (Kheper) nigroaeneus. Our results reveal that dances are elicited when the beetles lose control of their ball or lose contact with it altogether. We also find that dances can be elicited by both active and passive deviations of course and by changes in visual cues alone. In light of these results, we hypothesise that the dung beetle dance is a visually mediated mechanism that facilitates straight-line orientation in ball-rolling dung beetles by allowing them to 1) establish a roll bearing and 2) return to this chosen bearing after experiencing a disturbance to the roll path.     

Dung beetles ignore landmarks for straight-line orientation

Upon locating a suitable dung pile, ball-rolling dung beetles shape a piece of dung into a ball and roll it away in a straight line. This guarantees that they will not return to the dung pile, where they risk having their ball stolen by other beetles. Dung beetles are known to use celestial compass cues such as the sun, the moon and the pattern of polarised light formed around these light sources to roll their balls of dung along straight paths. Here, we investigate whether terrestrial landmarks have any influence on straight-line orientation in dung beetles. We find that the removal or re-arrangement of landmarks has no effect on the beetle’s orientation precision. Celestial compass cues dominate straight-line orientation in dung beetles so strongly that, under heavily overcast conditions or when prevented from seeing the sky, the beetles can no longer orient along straight paths. To our knowledge, this is the only animal with a visual compass system that ignores the extra orientation precision that landmarks can offer.     

How dim is dim? Precision of the celestial compass in moonlight and sunlight

Prominent in the sky, but not visible to humans, is a pattern of polarized skylight formed around both the Sun and the Moon. Dung beetles are, at present, the only animal group known to use the much dimmer polarization pattern formed around the Moon as a compass cue for maintaining travel direction. However, the Moon is not visible every night and the intensity of the celestial polarization pattern gradually declines as the Moon wanes. Therefore, for nocturnal orientation on all moonlit nights, the absolute sensitivity of the dung beetle's polarization detector may limit the precision of this behaviour. To test this, we studied the straight-line foraging behaviour of the nocturnal ball-rolling dung beetle Scarabaeus satyrus to establish when the Moon is too dim--and the polarization pattern too weak--to provide a reliable cue for orientation. Our results show that celestial orientation is as accurate during crescent Moon as it is during full Moon. Moreover, this orientation accuracy is equal to that measured for diurnal species that orient under the 100 million times brighter polarization pattern formed around the Sun. This indicates that, in nocturnal species, the sensitivity of the optical polarization compass can be greatly increased without any loss of precision.     

Heterotrophic succession within dung-inhabiting beetle communities in northern Spain

Successional patterns of beetles inhabiting dung pats were examined during May and July 1993 in a mountain area in northern Spain (Picos de Europa). Beetles belonging to six families were caught during the course of succession (30 d). Coprophagous beetles were more abundant in dung pats than predatory beetles (89 and 11 %, respectively). A trophic sequence was observed in relation to age of the dung, coprophagous beetles occurring earlier in the dung than predatory beetles. The pattern was observed on two occasions during the season, though succession proceeded somewhat faster in July than in May. These results suggest that food availability and microclimatic conditions in dung pats appear to determine the successional occurrence of beetle taxa. On the other hand, coprophagous species (Aphodius) were poorly segregated along the successional axis. Null models failed to support the hypothesis that successional overlap and differences in successional mean occurrence between species could be the result of competition. Successional patterns at the specific level probably reflect differences in behaviour, such as pat location, feeding, mating, egg-laying and larva requirements, rather than competitive replacement.     

Brood care in the dung beetle Onthophagus vacca (Coleoptera: Scarabaeidae): the effect of soil moisture on time budget, nest structure, and reproductive success

Under laboratory conditions brood care behaviour, nest structure and weight of dung supply in brood chambers of the dung beetle Onthophagus vcca proved to depend on water content of the soil beneath the dung. The substrate in a bucket beneath the dung pat was dry sand (4% water content) or moist sand (8% water content). Emigrating beetles were trapped and counted at 12 h intervals. In a total of 109 replicates one pair was released on an artificial 1000 g dung pat. From 95 replicates in which brood chambers were built the following results were derived: 1) Breeding females and resident males which helped the female stayed longer in dung pats on dry sand than in those on moist sand. 2) Nest architecture was influenced by substrate moisture: length of main tunnels did not differ between nests in dry and moist sand, but total length of side tunnels was shorter in dry sand. 3) Numbers of brood chambers were equal in both substrate types, weight of the dung supplies was larger in dry sand. 4) Offspring size was not only influenced by dung provision in the brood chambers. Beetles emerging from chambers in dry sand were smaller than those emerging from moist sand even if the amount of dung supply was equal.     

Experimental warming disrupts reproduction and dung burial by a ball‐rolling dung beetle

1. Insects are sensitive to climate change. Consequently, insect‐mediated ecosystem functions and services may be altered by changing climates. 2. Dung beetles provide multiple services by burying manure. Using climate‐controlled chambers, the effects of warming on dung burial and reproduction by the dung beetle Sisyphus rubrus Paschalidis, 1974 were investigated. Sisyphus rubrus break up dung by forming and rolling away balls of manure for burial and egg deposition. 3. To simulate warming in the chambers, 0, 2 or 4 °C offsets were added to field‐recorded, diurnally fluctuating temperatures. We measured dung ball production and burial, egg laying, survival and residence times of beetles. 4. Temperature did not affect the size or number of dung balls produced; however warming reduced dung ball burial by S. rubrus. Because buried balls were more likely to contain eggs, warming could reduce egg laying via a reduction in ball burial. Warming reduced the humidity inside the chambers, and a positive relationship was found between the number of dung balls produced and humidity in two temperature treatments. Temperature did not affect survival, or whether or not a beetle left a chamber. Beetles that did leave the chambers took longer to do so in the warmest treatment. 5. This study demonstrates that climate warming could reduce reproduction and dung burial by S. rubrus, and is an important first step to understanding warming effects on burial services. Future studies should assess warming effects in field situations, both on individual dung beetle species and on aggregate dung burial services.     

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     

Abundance and reproductive strategy of the dung beetle Aphodius rufipes (L.) (Scarabaeidae)

Abstract. 1. Adult A.rufipes invade fresh dung pats (mainly in August) and stay for a few days in each pat to feed and oviposit; maximum abundances: 0–100 beetles per cattle dung pat. Larval growth is completed within 40–55 days.
2. The dispersion of beetles is contagious or seemingly random. In sunny weather, with rapid crust formation on the dung, pats deposited in the afternoon are invaded by more beetles than those deposited in the morning.
3. In the field, the total number of eggs per pat is significantly less variable than the number of adult beetles; this might indicate a density-dependent, regulatory, rate of oviposition. The subsequent larval mortality is probably density-independent.
4. Experiments confirmed that the number of eggs laid per female-day in the pats was density-dependent.
5. The non-opportunistic reproductive strategy of A.rufipes may have evolved as an adaptation to life in dung pats: at excessive population densities pats might disappear before completion of larval growth.
6. Finally, A.rufipes in dung pats is compared with blowflies breeding in carrion.     

An analysis of the nesting behaviour of Geotrupes spiniger Marsham (Coleoptera, Scarabaeidae)

Abstract. 1. Female G.spiniger adults working alone made nests each consisting of a vertical shaft leading to a series of horizontal brood chambers filled with dung brood masses. Oviposition near the tip of the brood mass occurred while the egg cell was being completed over the expanded ovipositor. The shaft above each brood mass was filled with soil excavated from the next brood chamber. A similar response also filled artificial diverticula. An avoidance reaction towards buried dung prevented damage to pre-existing brood masses. 2. Virgin females did not make nests and did not avoid buried dung, but after mating (at about 4 weeks after eclosion) both types of behaviour were released within a few hours. 3. The presence of dung was required to initiate but not to maintain nesting behaviour. If dung was removed after oviposition the chamber was filled with soil produced by renewed excavation. Cellulose pulp could substitute for dung in brood mass formation. 4. Beetles interchanged between burrows at different stages before oviposition readily repeated all pre-oviposition behaviour. They appeared to respond to the length of the shaft and of the brood chamber since they extended short shafts and short brood chambers considerably more than those of normal length. After oviposition the beetles continued to make brood masses even under abnormal conditions. 5. Tilting the cage through 90° caused beetles before oviposition to re-orientate their burrowing direction, but tilting just after oviposition caused them to make vertical brood masses. Placing the shaft in a horizontal position towards the end of brood mass formation postponed the termination of this phase. 6. Beetles repeatedly excavated shafts and chambers when transferred to new cages. Conversely they repeatedly made brood masses when maintained in preformed plaster-of-Paris burrows. 7. This nesting behaviour can be described as a reaction chain in which each action generates its own terminating stimulus and initiates the subsequent response. The behaviour before oviposition could be omitted or repeated as required by the environment, but after oviposition there was little response to external interference. These characteristics have direct relevance to the survival of the larvae.     

The action of post-dispersal beetles (Coleoptera: Scarabaeidae) and ants (Hymenoptera: Formicidae) on scats of Didelphis spp. (Mammalia: Didelphidae)

A two year study of dung beetles and ants acting on scats of two species of opossum (Didelphis spp.) was carried out. Scats were left in the field in order to detect post-dispersal agents. A portion of each scat (30 %) was examined for seeds in the laboratory. Beetles were recovered from burrows (51% of 84 faecal samples left in the field) where they either buried scats of opossums or were attracted, together with ants, to pitfalls (N = 10) baited with opossum scats. Dung beetles were the main post-dispersal agents of seeds found in scats of opossums, rolling the scats away or burying then on the site of deposition. They buried faeces at 4 to 15 cm in depth (N = 22 tunnels). The main dung beetles identified (medium to large size) were Eurysternus (28.7 % in pitfalls) and Dichotomius (13.7 %), Coprophanaeus (seen only directly on faeces), besides small-bodied beetles (< 10 mm; 57.6 %). The ant Acromirmex sp. transported some seeds from scats. This species was present in 25.5 % of all Formicidae samples (pitfall). These post-dispersal agents contribute to avert scat seed predators such as rodents, and to accelerate seed bank formation.     

Bearing selection in ball-rolling dung beetles: is it constant?

Ball rolling in dung beetles is thought to have evolved as a means to escape intense inter- and intra-specific competition at the dung pile. Accordingly, dung beetles typically roll along a straight-line path away from the pile, this being the most effective escape strategy for transporting dung to a suitable burial site. In this study, we investigate how individual diurnal dung beetles, Scarabaeus (Kheper) nigroaeneus, select the compass bearing of their straight-line rolls. In particular, we examine whether roll bearings are constant with respect to geographic cues, celestial cues, or other environmental cues (such as wind direction). Our results reveal that the roll bearings taken by individual beetles are not constant with respect to geographic or celestial references. Environmental cues appear to have some influence over bearing selection, although the relationship is not strong. Furthermore, the variance in roll bearing that we observe is not affected by the presence or absence of other beetles. Thus, rather than being constant for individual beetles, bearing selection varies each time a beetle makes a ball and rolls it away from the dung pile. This strategy allows beetles to make an efficient escape from the dung pile while minimizing the chance of encountering competition.     

Generalist dung attraction response in a New Zealand dung beetle that evolved with an absence of mammalian herbivores

1. Dung beetles (Scarabaeidae: Scarabaeinae) are integral parts of many ecosystems because of their role in decomposition of dung; particularly mammal dung, which forms the diet of both larvae and adults. 2. New Zealand dung beetles are unusual as they are flightless and evolved on islands with a highly depauperate mammal fauna and thus without the usual dung resource used by dung beetles elsewhere. The diet of New Zealand dung beetles is unknown. 3. We hypothesised (1) that the endemic dung beetle Saphobius edwardsi would be attracted to a broad range of food types, and (2) that S. edwardsi would be able to survive and reproduce on a range of dung types and puriri (Vitex lucens) humus. 4. Laboratory choice tests identified that S. edwardsi was attracted to a range of mammal, bird, invertebrate, and reptile dung types, but not to non‐dung food sources. Five‐month no‐choice tests found that beetle survival rates were lower for beetles fed with humus compared with those fed on mammal, bird, or invertebrate dung. None of the beetles reproduced. 5. This study suggests S. edwardsi have a strong preference for dung, and are likely to be broad dung generalists in their feeding behaviour.     

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.     

Interspecific and intraspecific variation in diet preference in five Atlantic forest dung beetle species

1. Dung beetles are commonly assumed to be generalist feeders, but there has been limited work in identifying whether there is interspecific variation in feeding preference. Equally, there has been no work exploring whether generalist feeding behaviour in a species is a result of within‐species specialisation. 2. This study identified the individual and species‐level feeding preferences of five dung beetle species towards human, jaguar and pig dung using a choice experiment. 3. It was found that species varied in their preference for the dung types, but there was no evidence that within a species, individual beetles varied in their dung choice. These findings were similar to results from field experiments that are more typically used to assess feeding preferences in dung beetles. 4. The results suggest that individual specialism in feeding may not be common in dung beetles. However, there is variation in feeding preferences among species, which is often overlooked and can have implications for the ecosystem functions they provide. It is suggested that choice arenas can be used to assess feeding preferences in dung beetle species that are not abundant enough to reliably estimate dietary choice from field studies.     

Roles of endothermy in niche differentiation for ball-rolling dung beetles (Coleoptera: Scarabaeidae) along an altitudinal gradient

Abstract.  1. An analysis of whether niche differentiation in ball-rolling dung beetles can be explained by the way in which they regulate their body temperature was conducted.
2.  A priori assumptions were: (i) if thermoregulation affects niche partitioning, sympatric species must have different endothermic strategies that minimise encounters; or, alternatively (ii) if two co-occurring species show the same thermoregulation pattern and their flight periods overlap, they might be avoiding competition by exhibiting different resource preferences or different food relocation behaviour.
3. The ball-rolling dung beetles studied showed a hierarchical structure based on the species' endothermic capacity, measured as temperature excess [ T _ex= difference between body temperature ( T _b) and ambient temperature ( T _a)]. Those with a high T _ex (10–15 °C) were located exclusively at altitudes >1000 m a.s.l. On the coastal plains, species with a high T _ex were restricted to flying at night when the T _a was lower. Species with a lower T _ex (less than 10 °C higher than T _a) were found in the coastal plains zone.
4. Where there was sympatry with similar trophic habits, the species involved showed very different thermal niches, and where there was significant overlap of thermal niches between sympatric species, trophic habits of species were very different.
5. The results suggest that it is possible to use the concept of the thermal niche as a tool to explain interspecific interactions and the spatial distribution of species.     

Are ball-rolling (Scarabaeini,Gymnopleurini, Sisyphini) and tunnelling scarabaeine dung beetles equally choosy about the size of ingested dung particles?

Abstract.  1. The maximum size of ingested particles was determined in 11 species of ball-rolling, adult dung beetle (Scarabaeidae: Scarabaeinae) by mixing small latex or glass balls of known diameter into their food. The tribes Scarabaeini, Gymnopleurini, and Sisyphini (four, four, and three species respectively) were represented, with mean body sizes ranging from 0.33 to 4.0 g fresh weight.
2. Only particles with maximum diameters of 4–85 µm were ingested. Hence rollers, like other known beetles feeding on fresh dung, filter out larger, indigestible plant fragments and confine ingestion to small particles of higher nutritional value.
3. The maximum diameter of ingested particles increased significantly with body weight, whereas taxon (tribe) had no additional effect. Because big rollers accept larger particles than do tunnellers (which make dung stores for feeding and breeding in the soil immediately below the pat) of similar weight, the slope of the diameter-against-weight regression for rollers was significantly higher than that found earlier for tunnellers.
4. An explanation could be that a typical food ball made by a roller is considerably smaller than the amount of dung available to a feeding tunneller of the same size. If the roller were as choosy about particle size as the tunneller, it might not get enough food. This applies to large rollers in particular because their food balls contain a higher proportion of coarse fibres than those made by small species.     

Alternate Reproductive Tactics in an African Dung Beetle, Circellium bacchus (Scarabeidae)

Male-male competition is recognized as a potent force of sexual selection. When intra-sexual competition is strong then selection theory predicts that alternative male phenotypes will evolve. Circellium bacchus is a large, hornless, ball-rolling dung beetle with extensive variation in size and subject to intense male-male contest competition. We proposed that small male C. bacchus, which are unlikely to be successful in male contest competition, would adopt the alternative reproductive tactic of sneaking copulations. This alternative tactic is likely to influence not only behavior, but morphology with an expectation that sneaking males would invest more resources in testes development. Investigation of testis allometry revealed that smaller male C. bacchus beetles had relatively larger testes than their bigger conspecifics. Furthermore, as resources may be limited during larval development, differential investment in testes development should result in adult male beetles either competing for fertilizations or for access to mates. This is seen in C. bacchus as two alternate male phenotypes; smaller beetles with a relatively low body mass invest proportionately more in testes development compared to larger, heavier form in which testes size does not scale with condition. To our knowledge this study provides the first investigation of alternative phenotypes in the reproductive tactics of a ball-rolling dung beetle species.     

半干旱草原大中型土壤动物在畜粪分解中的作用

土壤动物是陆地生态系统的重要组分, 在有机质分解过程中具有重要作用。目前有关土壤动物在生态系统分解中的作用研究主要聚焦于植物凋落物的分解, 而对动物粪便分解的研究稀少。本研究在内蒙古典型草原设置了马粪和牛粪分解原位实验, 使用不同孔径的金属隔离网排除不同体型大小的土壤动物, 通过测定大中型土壤动物对畜粪分解过程中质量损失、碳氮含量和微生物呼吸以及土壤养分动态变化的影响, 解析其在分解中的作用。设置5个处理, 即CK, 仅土壤, 无粪; T0, 粪添加+0.425 mm隔离网(排除了粪居型和掘洞型粪金龟和中型土壤动物); T1, 粪添加+1 mm隔离网(排除了粪居型和掘洞型粪金龟); T2, 粪添加+2 mm隔离网(排除了掘洞型粪金龟); T3, 仅粪添加(不排除土壤动物)。结果表明: (1)在畜粪分解60天内, 土壤动物对畜粪的干质量损失没有显著的促进作用(P > 0.05); 相反, 在畜粪分解360天, 不隔离土壤动物处理(T3)显著地提高了牛粪干质量损失(P < 0.05), 而降低了马粪干质量损失(P < 0.05)。(2)在畜粪分解的60天内, 畜粪中碳和氮含量下降速度在有土壤动物存在的情况下(T3)快于隔离土壤动物(T0和T1)。(3)两种畜粪添加增加了土壤微生物的呼吸, 且这种增加趋势在实验的第15天和第30天在土壤动物存在时(T3)最明显。(4)与对照(CK)相比, 马粪添加处理提高了土壤速效氮、有机碳的含量和土壤含水量, 且这种增加趋势在排除掘洞型粪金龟(T2)和不排除土壤动物(T3)条件下表现更显著(P < 0.05), 而牛粪添加处理没有明显改变这些指标(P > 0.05)。研究表明, 分解初期粪金龟的取食和活动会改变畜粪的理化性质, 进而影响分解后期土壤生物在畜粪分解中的作用。