Marcus J. Byrne

Dung Beetles Use the Milky Way for Orientation

When the moon is absent from the night sky, stars remain as celestial visual cues. Nonetheless, only birds, seals, and humans are known to use stars for orientation. African ball-rolling dung beetles exploit the sun, the moon, and the celestial polarization pattern to move along straight paths, away from the intense competition at the dung pile. Even on clear moonless nights, many beetles still manage to orientate along straight paths. This led us to hypothesize that dung beetles exploit the starry sky for orientation, a feat that has, to our knowledge, never been demonstrated in an insect. Here, we show that dung beetles transport their dung balls along straight paths under a starlit sky but lose this ability under overcast conditions. In a planetarium, the beetles orientate equally well when rolling under a full starlit sky as when only the Milky Way is present. The use of this bidirectional celestial cue for orientation has been proposed for vertebrates, spiders, and insects, but never proven. This finding represents the first convincing demonstration for the use of the starry sky for orientation in insects and provides the first documented use of the Milky Way for orientation in the animal kingdom.

Lunar orientation in a beetle

Many animals use the suns polarization pattern to orientate, but the dung beetle Scarabaeus zambesianus is the only animal so far known to orientate using the million times dimmer polarization pattern of the moonlit sky. We demonstrate the relative roles of the moon and the nocturnal polarized–light pattern for orientation. We find that artificially changing the position of the moon, or hiding the moons disc from the beetles field of view, generally did not influence its orientation performance. We thus conclude that the moon does not serve as the primary cue for orientation. The effective cue is the polarization pattern formed around the moon, which is more reliable for orientation. Polarization sensitivity ratios in two photoreceptors in the dorsal eye were found to be 7.7 and 12.9, similar to values recorded in diurnal navigators. These results agree with earlier results suggesting that the detection and analysis of polarized skylight is similar in diurnal and nocturnal insects.

A Review of the Biological Control Programmes on Eichhornia crassipes (C.Mart.) Solms (Pontederiaceae), Salvinia molesta D.S.Mitch. (Salviniaceae), Pistia stratiotes L. (Araceae), Myriophyllum aquaticum (Vell.) Verdc. (Haloragaceae) and Azolla filiculoides Lam. (Azollaceae) in South Africa

Biological control against water hyacinth, Eichhornia crassipes (C.Mart.) Solms (Pontederiaceae), salvinia, Salvinia molesta D.S.Mitch. (Salviniaceae), water lettuce, Pistia stratiotes L. (Araceae), parrots feather, Myriophyllum aquaticum (Vell.) Verdc. (Haloragaceae), and red water fern, Azolla filiculoides Lam. (Azollaceae) has been ongoing in South Africa since the release of the first biological control agent on water hyacinth in 1974. This review provides an account of progress for the period from 1999. Post-release evaluations over the last three years have shown that, with the exception of water hyacinth, all of these problematic aquatic plants have been suppressed effectively using classical biological control. In eutrophic water bodies at high elevations that experience cold winters, an integrated approach, that includes herbicide application and augmentive biological control, is required against water hyacinth. The grasshopper Cornops aquaticum (Brüner) (Orthoptera: Acrididae: Leptysminae) has recently been released as a new agent for water hyacinth, and Megamelus scutellaris Berg (Hemiptera: Delphacidae) and Taosa longula Remes Lenicov (Hemiptera: Dictyopharidae) are being considered for release on water hyacinth. The longterm management of alien aquatic plants in South Africa relies on the prevention of new introductions of aquatic plant species that could replace those that have been controlled, and, more importantly, on a reduction in nutrient levels in South Africas aquatic ecosystems.

Neural coding underlying the cue preference for celestial orientation

Significance Many animals use the sun or moon and the polarization pattern for navigation. We combined behavioral experiments with physiological measurements of brain activity to reveal which celestial cue dominates the orientation compass of diurnal and nocturnal dung beetles. The preference found behaviorally precisely matches the preference encoded neurally and shows how the brain dynamically controls the cue preference for orientation at different levels: The sun or moon always dominates the orientation behavior and neural tuning of diurnal beetles, whereas in nocturnal beetles, celestial bodies dominate tuning only in bright light, with a switch to polarized light at night. This flexible neural tuning in the nocturnal species provides a simple mechanism that allows it to use the most reliable available orientation cue. Diurnal and nocturnal African dung beetles use celestial cues, such as the sun, the moon, and the polarization pattern, to roll dung balls along straight paths across the savanna. Although nocturnal beetles move in the same manner through the same environment as their diurnal relatives, they do so when light conditions are at least 1 million-fold dimmer. Here, we show, for the first time to our knowledge, that the celestial cue preference differs between nocturnal and diurnal beetles in a manner that reflects their contrasting visual ecologies. We also demonstrate how these cue preferences are reflected in the activity of compass neurons in the brain. At night, polarized skylight is the dominant orientation cue for nocturnal beetles. However, if we coerce them to roll during the day, they instead use a celestial body (the sun) as their primary orientation cue. Diurnal beetles, however, persist in using a celestial body for their compass, day or night. Compass neurons in the central complex of diurnal beetles are tuned only to the sun, whereas the same neurons in the nocturnal species switch exclusively to polarized light at lunar light intensities. Thus, these neurons encode the preferences for particular celestial cues and alter their weighting according to ambient light conditions. This flexible encoding of celestial cue preferences relative to the prevailing visual scenery provides a simple, yet effective, mechanism for enabling visual orientation at any light intensity.

Discontinuous gas exchange in dung beetles: patterns and ecological implications

Abstract This study correlates a distinctive pattern of external gas exchange, referred to as the discontinuous gas exchange cycle (DGC), observed in the laboratory, with habitat associations of five species of telecoprid dung beetles. The beetles were chosen from a variety of habitats that would be expected to present different amounts of water stress. All five species exhibited DGC. Sisyphus fasciculatus has been recorded only in woodland areas, and does not have strict spiracular control during its DGC. Anachalcos convexus and Scarabaeus rusticus are associated with open mesic habitats. Both species exhibit a distinct DGC, previously found in some other insect species, but intermediate within this study group. Sc. flavicornis and Circellium bacchus are typically found in arid regions, and have the most unusual form of DGC, with spiracular fluttering during the burst phase. These results support the hypothesis that spiracular fluttering reduces respiratory water loss. From this study we conclude that the DGC is an ancestral adaptation, most probably as a result of anoxic environments in underground burrows, but that spiracular control is enhanced to reduce respiratory water loss in beetle species that live in arid habitats.

The influence of climate on the establishment and success of the biocontrol agent Gratiana spadicea, released on Solanum sisymbriifolium in South Africa

Abstract The tortoise beetle Gratiana spadicea (Klug) (Coleoptera: Chrysomelidae) was released as a biological control agent for the South American weed, Solanum sisymbriifolium Lamarck (Solanaceae), in South Africa. Many of the releases were carried out in the high altitude regions (Highveld). However, the insect failed to establish at some sites, and where establishment was confirmed, its impact has been variable. Cold winters were blamed for the insects nonestablishment or for the variable success at some sites. This hypothesis was tested by comparing the climate in the native range of beetles with that of the release sites in South Africa and by exposing different life stages of the beetle to the extremes of temperature and humidity, typical of winters at these sites. Climate comparison indicated that moisture stress would be expected at the high elevation release sites. The lower lethal humidity for the eggs was calculated to be 56.6% RH. Humidity at the high elevation release sites dropped below this value every month, creating potential moisture stress for the eggs. Humidity had no significant effect on the survival of G. spadicea pupae. The critical minimum temperature (CT min ) of the adult beetles was 4.9±1.3 ° C (n=20) and 6.8±1.3 ° C (n=20) for the larvae. The calculated LT 50 of the adult beetles was −7.1 ° C (n=120;−9.1 to −5.3 ° C ) , while eggs and larvae survived 2-h exposure to −10 ° C . Although the beetle is tolerant of temperature extremes experienced at the Highveld sites, this area was shown to be a marginal habitat for the beetles because of humidity stress.

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 beetles 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.

Visual cues used by ball-rolling dung beetles for orientation

Ball rolling by dung beetles is considered to be a derived behaviour that evolved under pressure for space, and from competitors at the dung pat. Straight-line orientation away from the pat using a celestial cue should be the most successful rolling strategy to move dung to an unknown burial site. We tested this hypothesis in the field and the laboratory by presenting five species of ball-rolling beetles with different orientation tasks, involving reaction to obstacles as well as to reflected sunlight and artificial light sources. Beetles were found to consistently orientate along a chosen route, usually in the direction of the sun. Beetles rolling dung balls successfully negotiated barriers and returned to the original path as did beetles falling from ramps, or rotated about a fixed point while rolling a ball. The sun was found to be the main orientation cue, which could be substituted by reflected or artificial light. However, beetles reoriented themselves less accurately in response to lights in the laboratory, than they did to the reflected sun in the field. It is probable that phototactic orientation using the sun, which is widespread amongst arthropods, has been incorporated in the straight-line foraging behaviour that has evolved in ball-rolling dung beetles.

Predicting the distribution of Eccritotarsus catarinensis, a natural enemy released on water hyacinth in South Africa

Water hyacinth [Eichhornia crassipes (Mart.) Solms (Pontederiaceae)] is the most damaging aquatic weed in South Africa, where five arthropod biological control agents have been released against it. The most recent introduction of Eccritotarsus catarinensis (Carvalho) (Heteroptera: Miridae) has failed to establish permanent populations at a number of sites in South Africa where water hyacinth is a problem. Cold winter temperatures at these sites are assumed to be the reason for these establishment failures. This assumption was tested by investigating the thermal physiology of the mirid, then incorporating these data into various predictive distribution models. Degree‐day models predict 3–14 generations per year at different localities in South Africa, and five generations at a Johannesburg site where the mirid failed to overwinter. The inability to develop sufficiently rapidly during winter months may hinder overwintering of this insect, which was predicted to develop through only one generation during the winter months of April to August in Johannesburg. A CLIMEX model also showed that cold stress limits the mirids ability to overwinter in the interior of the country, while determination of the lower lethal limit (–3.5 °C) and critical thermal minimum (1.2 ± 1.17 °C) also indicated that extreme temperatures will limit establishment at certain sites. It is concluded that E. catarinensis is limited in its distribution in South Africa by low winter temperatures.

Larval performance of the pyralid borer Eldana saccharina Walker and stalk damage in sugarcane: Influence of plant silicon, cultivar and feeding site

Abstract Applied silicon (Si) can improve resistance of plants to insect attack. We investigated the interactions between soil applied Si, sugarcane cultivar, and three documented feeding/penetration sites (internode, leaf bud, and root primordia), each at three heights (top, middle, and base) on the stalk and their effects on stem borer, Eldana saccharina Walker (Lepidoptera: Pyralidae) performance and the stalk damage it inflicts. Four sugarcane cultivars, treated (Si+) and untreated (Si−) with calcium silicate, were artificially infested with E. saccharina and the larvae were examined 21 days later. Silicon treatment significantly increased stalk Si content in all sugarcane cultivars and, overall, significantly reduced borer growth rate and also reduced stalk damage, particularly at the internode. Borer survival and growth rate and stalk damage were lowest at the internode, particularly in Si+ resistant cultivars. Although the internode had the hardest rind, hardness at each site was not significantly increased by Si+.