Maturation of optics and resolution in adult dung beetle superposition eyes

Summary The maturation of the morphology and refractive-index gradients of the crystalline cones and cornea of the superposition compound eye of the nocturnal dung beetle Onitis aygulus was traced as a function of age following adult ecdysis. Intracellular recordings from retinula cells were also made to trace the maturation of angular sensitivity, thereby determining the resolution of the eye at each age. Maturation proceeded quickly during the first 3 days following ecdysis, and then more slowly, with full maturation attained by the end of the second week. The main experimental results obtained during the first 2 weeks after ecdysis were: (1) the mean length of the crystalline cones increases from 67 m to 79 m; (2) the mean thickness of the cornea increases from 36 m to 50 m; (3) the mean refractive index along the axis of the crystalline cones increases from 1.459 to 1.511 in the distal cone region, from 1.434 to 1.501 in the waist region and from 1.425 to 1.486 in the proximal region; (4) the shape of the refractive-index gradient becomes more parabolic; (5) the mean rhabdom cross-sectional area increases from 190 m² to 230 m²; (6) the angular-sensitivity function narrows, with the acceptance angle decreasing from 22 ° to 4 °. An optical ray tracing model predicts an image quality limited only by the immaturity of the optics, a prediction confirmed by the electrophysiological results. The results are compared to optical maturation in day-active moths and skipper butterflies (which mature very quickly) and discussed in relation to ecological efficiency.     

Screening pigment,aperture and sensitivity in the dung beetle superposition eye

Summary Sensitivity to light was investigated in the refracting superposition eye of the dung beetle Onitis alexis using electrophysiological measurements and optical modelling. Intracellular recordings were made from single retinula cells over 24-h periods, with cells light and dark adapted, in order to measure the response/intensity (V-LogI) functions. The combined effects of a circadian rhythm and light adaptation allow the determination of the relative contributions of screening-pigment migration and transduction gain to changes in sensitivity in the eye. Between the extremes of dark adaptation at night and light adaptation during the day, the maximum sensitivity change possible is at least 4 log units, of which approximately 2 log units can be accounted for by changes in the transduction gain and at least 2 log units by screening-pigment migration. The role of the superposition aperture (the number of facets that contribute light to one rhabdom) in 3 species of dung beetle was investigated with an optical ray-tracing model of the eye. The facets of the superposition aperture do not contribute light equally to the target rhabdom; except in one species, the greatest contribution comes from facets located away from both the centre and periphery of the aperture. Light adaptation increases the optical density of the superposition aperture and decreases its size.     

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.     

Geometrical optics of a galatheid compound eye

The eyes of galatheid squat lobsters (Munida rugosa) are shown to be of the reflecting superposition type. In the dark-adapted state corneal lenses focus light at the level of the rhabdoms and light from more than 1000 facets is redirected to the superposition focus by the reflecting surfaces of the crystalline cones. When the eye is light adapted, apposition optics are used. In this state paraxial light is focused by the corneal lens and the parabolic proximal end of the cone onto the distal end of a rhabdomeric lightguide. The latter transmits light across the clear zone to the rhabdom layer. In the dorsal part of the eye the individual ommatidia become progressively shorter until the cones and rhabdoms are no longer separated by a clear zone. Although formerly considered to be developing ommatidia, they are shown to be retained specifically for scanning the downwelling irradiance.Abbreviations RI refractive index - SEM scanning electron microscope     

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.

     

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.

     

Ocellar optics in nocturnal and diurnal bees and wasps

Nocturnal bees, wasps and ants have considerably larger ocelli than their diurnal relatives, suggesting an active role in vision at night. In a first step to understanding what this role might be, the morphology and physiological optics of ocelli were investigated in three tropical rainforest species – the nocturnal sweat bee Megalopta genalis, the nocturnal paper wasp Apoica pallens and the diurnal paper wasp Polistes occidentalis – using hanging-drop techniques and standard histological methods. Ocellar image quality, in addition to lens focal length and back focal distance, was determined in all three species. During flight, the ocellar receptive fields of both nocturnal species are centred very dorsally, possibly in order to maximise sensitivity to the narrow dorsal field of light that enters through gaps in the rainforest canopy. Since all ocelli investigated had a slightly oval shape, images were found to be astigmatic: images formed by the major axis of the ocellus were located further from the proximal surface of the lens than images formed by the minor axis. Despite being astigmatic, images formed at either focal plane were reasonably sharp in all ocelli investigated. When compared to the position of the retina below the lens, measurements of back focal distance reveal that the ocelli of Megalopta are highly underfocused and unable to resolve spatial detail. This together with their very large and tightly packed rhabdoms suggests a role in making sensitive measurements of ambient light intensity. In contrast, the ocelli of the two wasps form images near the proximal boundary of the retina, suggesting the potential for modest resolving power. In light of these results, possible roles for ocelli in nocturnal bees and wasps are discussed, including the hypothesis that they might be involved in nocturnal homing and navigation, using two main cues: the spatial pattern of bright patches of daylight visible through the rainforest canopy, and compass information obtained from polarised skylight (from the setting sun or the moon) that penetrates these patches.     

Strategies for retinal design in arthropod eyes of low F-number

Summary The superposition eyes and simple eyes of many arthropods have apertures (A) with a diameter bigger than, or about the same size as, the focal length of the eye (f). That is, these eyes have low F-numbers (f/A). Many of the light rays focussed onto a photoreceptor will not be trapped by total internal reflection in the photoreceptor and will therefore pass through and be absorbed in photoreceptors other than that for which the light was intended. This spread of light in the retina leads to a broadening of the angular-sensitivity function and a consequent degrading of the image at the retinal level. A number of solutions to this problem are found in nature, with the most effective that of isolating the photoreceptors with a sheath of either light-absorbing pigment or reflecting tapetum. A ray-tracing model was used to assess the relative merits of the tapetal and pigment sheath designs in low F-number superposition eyes, and also to investigate the effect of changing the refractive index and absorption coefficient of the rhabdom. Which sheathing solution is best depends on the quality of the image on the retina, on the spacing of the rhabdoms in the retina and on the intensity of light normally experienced by the eye. In a retina with closely packed rhabdoms, the model predicts full sheathing to be the optimal solution if the image is well-focussed, partial sheathing if poorly focussed and no sheathing if moderately well-focussed. In a retina with rhabdoms spaced apart and a well-focussed image, the model predicts partial sheathing to be optimal. A pigment sheath is predicted to be useful in eyes which experience bright light and have no need for high sensitivity. A tapetal sheath is predicted to be useful at any intensity. A survey of arthropod eyes with low F-number supports the predictions of the model.     

Movement patterns in dung beetles and in the dung fly

Movements in coprophagous beetles between cow pats are discussed in relation to two hypotheses: (i) that the movement patterns are results of selection to maximize the net rate of energy intake; (ii) that the rates of immigration to and emigration from droppings are governed by simple stochastic rules of behaviour. Observations support the latter hypothesis, and contrast with predictions from the simplest optimization model. Reasons for this discrepancy are discussed. Spatio-temporal occurrence of the dung fly, Scatophaga stercoraria, around cow pats is re-analysed from Parker's data. A simple Markov chain model predicts the observations well, but it does not solve the question of the type of selection responsible for the values of the behavioural parameters. It is argued that difficulties in coping with a stochastic environment, and the consequences of movements being manifested on many spatial and temporal levels, favour evolution consisting of few key behavioural parameters being singled out and subsequently adjusted by selection.     

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.

     

Intensity and polarization of the eyeshine in butterflies

The compound eyes of most diurnal butterflies have a reflecting tapetum below the retina. Light that enters the eye is guided down the rhabdom, reflected by the tapetum, and then guided back up the rhabdom. The light that is not absorbed by the rhabdom is reemitted and gives rise to an eyeshine. We have measured the fraction of the incident light that is re-emitted, and also the degree to which this light retains its original polarization. The following conclusions are drawn:

Superposition of modes in a caricature of a model for morphogenesis

In a model proposed for cell pattern formation by Nagorcka et al. (J. Theor. Biol. 1987) linear analysis revealed the possibility of an initially spatially uniform cell density going unstable to perturbations of two distinct spatial modes. Here we examine a simple one-dimensional caricature of their model which exhibits similar linear behaviour and present a nonlinear analysis which shows the possibility of superposition of modes subject to appropriate parameter values and initial conditions.     

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...     

Distribution and abundance of dung beetles in fragmented landscapes

Related species utilising similar resources are often assumed to show similar spatial population structures and dynamics. This paper reports substantial ecological variation within a set of Aphodius dung beetles occurring in the same patchily distributed resource, livestock dung in pastures. We show how variation in habitat and resource selectivity, in the rate of movements between pastures, and in the distribution of local population sizes all contribute to interspecific differences in spatial population structures. Local dung beetle assemblages are compared between two landscapes with different densities of pastures. In one of the landscapes, we contrast the abundances and regional distributions of Aphodius before and after 15 years of rapid habitat loss. Different species show very dissimilar responses to changes in the structure of the landscape. Our results suggest that generalist Aphodius species, and specialist species with high dispersal powers, occur as large "patchy" populations in the landscape. In contrast, a strict pasture specialist species with limited dispersal powers (A. pusillus) forms classical metapopulations. At the community level, interspecific differences in spatial population structures make the local community composition a function of the structure of the surrounding landscape.     

Molecular evolution and radiation of dung beetles in Madagascar

Madagascar is the world's fourth largest island and has a wide range of climates and ecosystems. Environmental diversity combined with long history of isolation (160 Myr) has generated a high level of endemism at different taxonomic levels, making Madagascar one of the hotspots of global biodiversity. Dung beetles, represented by the two tribes of Canthonini and Helictopleurini, exemplify a large insect taxon. Helictopleurini are completely endemic to Madagascar while Canthonini are endemic at generic level. Using data from mitochondrial and nuclear genes, phylogenetic relationships were investigated in a sample of 44 species. The phylogeny for Canthonini consists of several distinct clades, possibly reflecting multiple colonization of Madagascar. The phylogeny does not support the current taxonomy for all genera. The phylogeny for Helictopleurini lacks statistical support at supra‐specific level, and genetic divergence among the Helictopleurini species is comparable with that among species within genera in Canthonini. These results suggest that Helictopleurini has undergone rapid speciation and most likely more recently than Canthonini, consistent with the estimated radiation time based on mtDNA mutation rates in insects and with knowledge about the systematics and geographic distribution of dung beetles worldwide. A detailed analysis of sequence composition identified common patterns in Malagasy dung beetles and other insects. © The Willi Hennig Society 2007.     

Thermoregulation in endothermic dung beetles (Coleoptera: Scarabaeidae): effect of body size and ecophysiological constraints in flight

We explore the physiological constraints of body temperature as related to body mass and ambient temperature during flight in endothermic dung beetles showing a mass-related breakpoint where species show strong vs. weak endothermy. We found two different strategies in the dung beetles prior to flight; larger beetles (>1.9 g) elevate and maintain their body temperature (T(b)) at levels well above ambient temperature (T(a)) whereas smaller beetles' (<1.9 g) T(b) tends to conform with T(a). Physiological constraints analysis revealed a constant maximum tolerated temperature (in flight) of 42 degrees C and a minimum temperature for flight of around 25 degrees C. These, with body mass, may play a role in thermal niche partitioning and geographical distribution patterns.     

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。     

甲虫表皮碳氢化合物研究进展

昆虫表皮碳氢化合物(cuticular hydrocarbons)是正烷烃、不饱和烃以及甲基支链烃的混合物.除了保持水分的基本功能外,还具有多种生物功能.鞘翅目昆虫(俗称甲虫)是多样性最为丰富的昆虫类群,目前对其表皮碳氢化合物研究开展颇多,主要见于化学通讯领域,内容涉及到性信息素、标记信息素、聚集信息素、化学拟态、寄主...     

Habitat fragmentation and the functional efficiency of temperate dung beetles

To understand how current patterns of habitat loss and fragmentation will ultimately affect ecosystem functioning, we need to match experimental manipulations of community structure with real changes occurring in the landscapes of today. In this study, we examine the consequences of habitat fragmentation on a key function: the decomposition of dung by invertebrates. In a microcosm experiment, we use previous observations of dung beetle assemblage structure in fragmented and intact landscapes to create realistic differences in assemblages of small, dung‐dwelling species in the genus Aphodius. We ask whether such differences will affect ecosystem functioning, and how their effects compare to those of removing full functional groups: dung‐dwelling Aphodius, tunnelling Geotrupes stercorarius, and/or earthworms. As measured by changes in dung fresh weight, we observe an overriding impact of removing G. stercorarius, with the amount of dung remaining at any one time doubling if the species is excluded. Compared to this major effect, there seem to be less effects of removing Aphodius, ambiguous effects of excluding earthworms, and no detectable effects of relatively minor changes in Aphodius assemblages as induced by current levels of fragmentation. Overall, our results support the general notion that different species contribute highly unevenly to overall ecosystem functioning. Most importantly though, our findings suggest that the functional consequences of habitat loss will depend on taxon‐specific responses to landscape modification. Only by addressing these responses may we predict the actual consequences of habitat loss.     

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.