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.     

Establishment of exotic dung beetles in Queensland: the role of habitat specificity

Between 1968 and 1983, 22 species of dung beetles from southern Africa were released into pastoral regions of Queensland in northern Australia. Seven species have become widespread and abundant and the others either occur in isolated patches, are rare or have failed to establish self perpetuating populations. These species all occur in the Hluhluwe district in southern Africa which has a climate similar to that of coastal central and southern Queensland. The numberical abundance and habitat associations of the 22 speices were examined at Hluhluwe to seek an explanation for their performance in Australia. It was concluded that habitat specificity and the adequacy with which habitat was matched in Australia were critical factors. Species which established in Australia were relatively common in cattle dung in Africa where they showed a preference for grassland on clay-loam soils.      

Honeybee navigation: critically examining the role of the polarization compass

Although it is widely accepted that honeybees use the polarized-light pattern of the sky as a compass for navigation, there is little direct evidence that this information is actually sensed during flight. Here, we ask whether flying bees can obtain compass cues derived purely from polarized light, and communicate this information to their nest-mates through the ‘waggle dance’. Bees, from an observation hive with vertically oriented honeycombs, were trained to fly to a food source at the end of a tunnel, which provided overhead illumination that was polarized either parallel to the axis of the tunnel, or perpendicular to it. When the illumination was transversely polarized, bees danced in a predominantly vertical direction with waggles occurring equally frequently in the upward or the downward direction. They were thus using the polarized-light information to signal the two possible directions in which they could have flown in natural outdoor flight: either directly towards the sun, or directly away from it. When the illumination was axially polarized, the bees danced in a predominantly horizontal direction with waggles directed either to the left or the right, indicating that they could have flown in an azimuthal direction that was 90° to the right or to the left of the sun, respectively. When the first half of the tunnel provided axial illumination and the second half transverse illumination, bees danced along all of the four principal diagonal directions, which represent four equally likely locations of the food source based on the polarized-light information that they had acquired during their journey. We conclude that flying bees are capable of obtaining and signalling compass information that is derived purely from polarized light. Furthermore, they deal with the directional ambiguity that is inherent in polarized light by signalling all of the possible locations of the food source in their dances, thus maximizing the chances of recruitment to it.     

蜣螂的生态价值和保护意义

蜣螂通常以粪便或腐殖质为食,在生态系统平衡中具有重要的作用。文章系统介绍蜣螂的生态价值,包括转将运粪便到地下、对种子二次传播的作用、传粉作用、对有害生物的控制作用和寄生虫的中间宿主,以及广阔的开发应用前景、重要仿生学意义和重大理论意义;并进一步提出蜣螂保护的迫切性和重要意义。     

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.

     

Behavioural and physiological mechanisms of polarized light sensitivity in birds

Polarized light (PL) sensitivity is relatively well studied in a large number of invertebrates and some fish species, but in most other vertebrate classes, including birds, the behavioural and physiological mechanism of PL sensitivity remains one of the big mysteries in sensory biology. Many organisms use the skylight polarization pattern as part of a sun compass for orientation, navigation and in spatial orientation tasks. In birds, the available evidence for an involvement of the skylight polarization pattern in sun-compass orientation is very weak. Instead, cue-conflict and cue-calibration experiments have shown that the skylight polarization pattern near the horizon at sunrise and sunset provides birds with a seasonally and latitudinally independent compass calibration reference. Despite convincing evidence that birds use PL cues for orientation, direct experimental evidence for PL sensitivity is still lacking. Avian double cones have been proposed as putative PL receptors, but detailed anatomical and physiological evidence will be needed to conclusively describe the avian PL receptor. Intriguing parallels between the functional and physiological properties of PL reception and light-dependent magnetoreception could point to a common receptor system.     

Artificial light and sun compass orientation in the sandhopper Talitrus saltator (Crustacea, Amphipoda)

Experiments on compass orientation under artificial light were conducted with adult individuals of Talitrus saltator. The aim was to reproduce in the laboratory an orientation based on the sun compass corresponding to that recorded in conditions of the true sun and sky. This was obtained by the creation within an opaque Plexiglas dome of a scenario that permitted variation of the brightness of the artificial sky and sun. The results show that it is possible to obtain sun compass orientation corresponding to the natural situation even in an artificial environment. It can be concluded that sandhoppers identify an artificial light source as the sun if the artificial sky is also illuminated and if the intensities of the artificial sun and sky exceed certain threshold values (1.13 and 10 μW cm⁻², respectively). The results of other experiments under the natural blue sky with an artificial sun and with the real sun attenuated are discussed. Accepted: 23 May 1997     

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.     

Contradictory results on the role of polarized light in compass calibration in migratory songbirds

Experiments with migrating birds on the interaction between magnetic and celestial cues have produced heterogeneous results. A recent study claimed that the magnetic compass in passerine migrants is calibrated by the pattern of polarized light at sunset and sunrise and that the area just above the horizon is crucial for this calibration. To test the latter hypothesis, we performed a similar experiment with Australian Silvereyes. It produced contrary results, however, the birds, in spite of observing the natural polarization pattern at sunrise and sunset down to the horizon in an altered magnetic field, continued in their normal southerly magnetic direction when subsequently tested in the local geomagnetic field—the conflict between magnetic and polarized light cues had not caused them to recalibrate their magnetic compass. This contradicts the assumption that skylight polarization patterns generally serve as a primary calibration reference for migratory songbirds.

Development of lateralization of the magnetic compass in a migratory bird

The magnetic compass of a migratory bird, the European robin (Erithacus rubecula), was shown to be lateralized in favour of the right eye/left brain hemisphere. However, this seems to be a property of the avian magnetic compass that is not present from the beginning, but develops only as the birds grow older. During first migration in autumn, juvenile robins can orient by their magnetic compass with their right as well as with their left eye. In the following spring, however, the magnetic compass is already lateralized, but this lateralization is still flexible: it could be removed by covering the right eye for 6 h. During the following autumn migration, the lateralization becomes more strongly fixed, with a 6 h occlusion of the right eye no longer having an effect. This change from a bilateral to a lateralized magnetic compass appears to be a maturation process, the first such case known so far in birds. Because both eyes mediate identical information about the geomagnetic field, brain asymmetry for the magnetic compass could increase efficiency by setting the other hemisphere free for other processes.     

Field evidence of compass orientation in migrating juvenile grunts (Haemulidae)

Juvenile French and white grunts, Haemulon flavolineatum (Desmarest) and H. plumieri (Lacé-pède), were captured during their daily migrations between diurnal resting sites on coral patch reefs and nocturnal feeding grounds in seagrass beds. Grunts captured during morning and evening migrations were released on the route and after displacement up to 100 m or 5 km away. Grunts generally moved in the direction which would have taken them back towards their home reef or to their accustomed feeding sites, indicating that familiar landmarks are not essential for orientation. The spatial precision of migration may serve to partition the feeding area most efficiently. The timing of migrations is also very precise, and appears to be adapted to reduce the vulnerability of grunts to predation near their home reef.     

Dung pad residence time covaries with male morphology in the dung beetle Onthophagus taurus

1. The immigration and emigration behaviours of Onthophagus taurus were investigated in a combination of field and laboratory experiments to examine patterns of dispersal in this species. 2. On average, the mean proportion of major males immigrating to baited pitfall traps decreased significantly with time. In contrast, the mean proportion of minor males and females did not change with time. 3. The time taken to emigrate from a dung pad differed significantly among major males, minor males, and females. This difference arose because major males spent a significantly shorter period of time residing within the dung pad. On average, more than 50% of major males had emigrated from the dung pads after 30 h, some 4 h earlier than minor males and females. 4. When the effects of body size were controlled, major males with longer horns were shown to have longer wings. Because major males spend more time in flight, longer wings may be an adaptation to more frequent dispersal. Alternatively, longer wings in major males may compensate for the production of aerodynamically costly horns.     

Orientation to shorelines by the supratidal amphipod Talorchestia longicornis: Wavelength specific behavior during sun compass orientation

If released in water or on sand the supratidal amphipod Talorchestia longicornis Say amphipods moves in the onshore direction. The present study was designed to determine whether this species uses the sun as a cue for orientation and if so, which visual pigment in the compound eyes is involved. When tested in an apparatus with a view of only the sun and sky amphipods were disoriented when the sun was obscured by clouds. However, when the sun was visible, they oriented in the onshore direction of their home beach in both water and air during both the morning and afternoon. Resetting the time of their circadian rhythm in activity with either an altered light:dark or diel temperature cycle also reset the chronometric mechanism associated with sun compass. orientation. T. longicornis has two visual pigments with absorption maxima near 420 nm and 520 nm. Only the 420 nm pigment is used for sun compass orientation, which may be an adaptation for increasing the contrast between the sun and background scattered skylight or for detecting the radiance distribution of skylight. Irradiating the 520 nm absorbing pigment alone induced positive phototaxis to the sun but not onshore orientation. Thus, T. longicornis shows wavelength specific behavior by using only one of its visual pigments for sun compass orientation.     

Comparative phylogeography and speciation of dung beetles from the Australian Wet Tropics rainforest

In tropical rainforests, insects show especially high species richness and local endemism of species relative to vertebrates. One possible cause is that insects respond to historical fluctuations of rainforests on a smaller spatial scale than do vertebrates. To evaluate this hypothesis, we combine environmental niche models and mitochondrial DNA phylogeography for two pairs of sister species of the dung beetle genus Temnoplectron (T. aeneopiceum-T. subvolitans and T. politulum-T. reyi) from the rainforests of northeastern Australia, where climate-driven rainforest fluctuations in the Quaternary have strongly influenced genetic and species diversity of vertebrates. Within both species pairs, the bioclimatic niche was conserved, but the T. aeneopiceum-T. subvolitans species pair had the narrower environmental range, and thus more restricted potential distribution. Coalescent analyses indicated Late Pliocene or Early Pleistocene divergences for both species pairs, and earlier speciation in (T. aeneopiceum-T. subvolitans) than in (T. politulum-T. reyi). Phylogeographic structure in (T. aeneopiceum-T. subvolitans) was more pronounced than in (T. politulum-T. reyi), with significant isolation-by-distance in the former species-pair only. Nested clade and coalescence analyses indicated local range expansions for the T. aeneopiceum-subvolitans species pair and range-wide expansion for both T. politulum and T. reyi. We suggest that stronger phylogeographic structure and earlier divergence in (T. aeneopiceum-T. subvolitans) than in (T. politulum-T. reyi) reflects a stronger influence of environmental barriers to gene flow under the present climate and greater sensitivity to warmer and drier periods of the Quaternary. The two species pairs evidently responded to Quaternary rainforest fluctuations at spatial scales similar to those seen within low-vagility species of vertebrate. Despite this similarity of scale, these insect lineages are reproductively isolated at parapatric boundaries, whereas analogous lineages of vertebrates often are not. We suggest that rapid evolution of genitalia may facilitate geographic speciation in rainforest beetles.     

The daily onset of flight in the crepuscular dung beetle Onitis alexis

The role of light intensity and temperature in determining the onset of flight in the crepuscular dung beetle Onitis alexis Klug (Coleoptera: Scarabaeidae) was examined. Flight under natural light was highly synchronized, with two-thirds of the beetles that flew flying over a period of 10–12 min. In six flights on different days, the mean time of onset varied by up to 13 min, but mean onset occurred at fixed light intensity. Absolute light intensity therefore appears to be a vital cue in determining flight onset. Mean onset remained at this intensity when dusk was advanced artificially by up to about 8 min. However, when dusk was brought further forward, mean flight occurred at lower intensities and onset of flight took place over a longer period. This is interpreted as an overlapping of the period of light intensities suitable for flight with the circadian rhythm that brings the beetles to the surface. No beetles flew when kept under constant bright light or in the dark during the dusk period. Under conditions of constant dusk, the beetles appeared uncoordinated and, although the median time of onset of flight was only 3 min later than in the control flight, onset of flight was spread out over a much longer period. At soil temperatures of 20–22 °C, over 90% of the beetles flew. This percentage decreased with decreasing temperature, and less than 10% flew at temperatures of 16–17 °C.

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L'incitation quotidienne au vol chez e bousier crépusculaire Onitis alexis Klug (Col. Scarabaeidae)

Résumé L'examen a porté sur le rôle de l'intensité lumineuse et de la température sur l'incitation au vol chez Onitis alexis. Le vol en lumière naturelle est fortement synchronisé, avec les deux tiers des adultes qui s'envolent en 10–12 min. Sur six vols pour différents jours, le moment moyen d'envol variait de 13 min, mais l'envol moyen avait lieu à une intensité lumineuse déterminée. La valeur absolue de l'intensité lumineuse paraît ainsi un signal crucial dans la détermination et l'incitation au vol. Le moment d'envol se maintient à cette intensité quand le crépuscule est artificiellement avancé jusqu'à 8 min environ. Cependant, quand le crépuscule est rendu encore plus précoce, l'envol moyen se produit à des intensités plus basses et est étalé. Ceci peut être interprété comme un chevauchement de la période des intensités lumineuses favorables à l'envol avec le rythme circadien qui conduit les bousiers à la surface. Aucun adulte ne vole quand il y a maintien de lumière constante ou d'obscurité à l'heure du crépuscule. En présence de crépuscule constant, les réponses sont hétérogènes, et, bien que le moment médian d'envol ne soit retardé que de 3 min, par rapport aux témoins, l'incitation à l'envol est étalée sur une période plus longue. Avec des températures au sol de 20–22 °C, plus de 90% des adultes s'envolent. Le pourcentage diminue avec la température, et moins de 10% s'envolent à 16–17 °C.

     

Forecasting insect species richness scores in poorly surveyed territories: the case of the Portuguese dung beetles (Col. Scarabaeinae)

Large-scale biodiversity assessment of faunal distribution is needed in poorly sampled areas. In this paper, Scarabaeinae dung beetle species richness in Portugal is forecasted from a model built with a data set from areas identified as well sampled. Generalized linear models are used to relate the number of Scarabaeinae species in each Portuguese UTM 50 × 50 grid square with a set of 25 predictor variables (geographic, topographic, climatic and land cover) extracted from a geographic information system (GIS). Between-squares sampling effort unevenness, spatial autocorrelation of environmental data, non-linear relationships between variables and an assessment of the models' predictive power, the main shortcomings in geographic species richness modelling, are addressed. This methodological approach has proved to be reliable and accurate enough in estimating species richness distribution, thus providing a tool to identify areas as potential targets for conservation policies in poorly inventoried countries.     

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.