Normal and atypical nesting behaviour of Copris lunaris (L.): comparison with related species (Coleoptera, Scarabaeidae)

Abstract. 1. The lifecycle, mating and larval behaviour of Clunaris are described. Adults appeared in the autumn and nested in the following spring. The female beetle remained in the nest with the brood and could nest again the following year.
2. Nesting was initiated when virgin females were mated in the spring. Brood balls were formed by techniques sirnilar to those used by Scarabaeini. The female beetle left the nest soon after the first imagos broke out of the brood balls.
3. Nesting behaviour was readily modified by external conditions. Many parts of the sequence could be repeated or omitted. The female beetle left the nest if the brood was removed, but she remained for longer than usual if younger brood was substituted near the end of the normal nesting period.
4. Certain experimental conditions released behaviour patterns typical of other species. These were formation of superficial nests or of two-chambered nests, oviposition before completing the brood ball, and coating of the brood balls with soil (all found in other Coprina), as well as ball rolling and ball burial (found in Scarabaeini). The results are discussed in relation to the evolution of Copris nesting behaviour.     

Brood ball construction by the non-brooding Coprini Sulcophanaeus carnifex and Dichotomius torulosus (Coleoptera, Scarabaeidae)

ABSTRACT.

• 1 Feeding burrows made by S.carnifex and D.torulosus adults consisted of tunnels filled with sausage-shaped masses of dung. Brood balls were made by taking fragments of dung from an adjacent feeding burrow and aggregating them into a small sphere which was gradually enlarged and then coated with soil.
• 2 S.carnifex adult females showed no response to their own eggs (unlike Copris lunaris: Klemperer, H.G. (1982) Ecological Erltonzology, 7, 155–167). S.carnifex larvae made and maintained an air channel to the upper pole of the ball and they are in this respect preadapted to receive parental care.
• 3 Compared with a brood mass, a brood ball was less likely to be attacked by Meptoparasitic larvae. The soil coat retarded drying of the brood ball by increasing the total mass of moisturecontaining material.

     

Parental behaviour in Copris lunaris (Coleoptera, Scarabaeidae): care and defence of brood balls and nest

Abstract. 1. Nesting female beetles righted brood balls (so as to replace the egg or larva in the uppermost position) and repaired damaged balls. This behaviour required the presence of an egg or larva in the ball, or of a short-lasting material found just after oviposition. The shape of the ball was also a righting stimulus since artificial ellipsoids were stood on end.
2. Balls containing dichloromethane extracts of C.lunaris brood were righted and repaired. Eggs and larvae of several other Scarabaeidae did not release these responses but were destroyed.
3. Righting behaviour was released when brood was absent from the top of the ball. The beetle then crawled vertically downwards and, if it encountered the displaced apex, a novel rolling action followed which automatically turned the ball towards the correct position.
4. An opening made in the nest was repaired with soil excavated from the chamber floor. Clunaris adults and Aphodius fossor larvae were attacked if they were encountered in the nest.     

The evolution of parental behaviour in Scarabaeinae (Coleoptera, Scarabaeidae): an experimental approach

ABSTRACT.

• 1 Dung beetle lifestyles are reviewed. Most Scarabaeinae lay their eggs in dung masses that are packed into underground chambers, but Coprini and Scarabaeini typically lay their eggs in free-standing brood balls and it is in these tribes that parental care of the brood has evolved.
• 2 Brood balls are constructed by aggregating fragments of dung. This technique is derived from the method of gathering dung at the surface. Larvae developing in brood balls are better protected against dehydration and parasite attack. The repair technique of Scarabaeine larvae preadapts them to life in brood balls.
• 3 Parental care by Copris lunaris depends on appropriate responses by the female to the brood, and it has the selective advantage of protecting the brood against parasites. Preadaptations for parental care in‘non-brooding’Coprini are discussed.
• 4 Variations in the basic Copris nesting behaviour are summarized. Similar variations can occur spontaneously in C.lunaris and can also be released by unusual circumstances.
• 5 C.lunaris females could in principle cooperate but certain factors have prevented this social evolution. The significance of the transient cooperation with the male beetle is discussed.

     

Nesting of Afrotropical Oniticellus (Coleoptera, Scarabaeidae) and its evolutionary trend from soil to dung

Abstract. 1. Oniticellus egregius Klug constructs brood ovoids of dung in the soil immediately under the edge of animal droppings. Each successive brood ovoid is enveloped within a soil shell. After completion of brood construction, loose earth is cleared from around the broods to produce a brood chamber. The immatures are then abandoned as eggs or first instar larvae.
2. O.planatus Castelnau and O.formosus Chevrolat usually construct brood balls of dung within animal droppings. Each brood is progressively enlarged by the addition of further dung after egg-laying. This enlargement is slight in O.planatus and marked in O.formosus. Parental females of both species remain in the brood chambers during development of the immatures which are abandoned principally as pupae.
3. Under very moist experimental conditions, O.planatus buries dung and constructs broods shallowly in the soil. Such nests are frequently connected to the pad by a short tunnel.
4. From a consideration of behavioural patterns it is suggested that the specialized nesting habits of these species have been derived from those of dung-burying ancestors similar to the modern genus, Euoniticellus, through reduction and loss of tunnelling in the soil.     

Subsocial behaviour in Oniticellus cinctus (Coleoptera, Scarabaeidae): effect of the brood on parental care and oviposition

ABSTRACT. In Oniticellus cinctus (F.) the nest chambers each contain about twenty brood balls. Females enlarge the brood balls during the egg and larval stages and remain in the chamber for the whole period of brood development (1 month); they then make a new nest after 1 week. The presence of the brood releases parental care and ensures that the mother remains in the nest: she repairs defects in the brood balls and the nest, and expels other O. cinctus females. A new ball is formed around a naked O. cinctus larva, but larvae of other species are killed. In addition, the brood inhibits oviposition: removal (or addition) of brood balls stimulates (or inhibits) egg laying. In inhibited ovarioles, existing follicles are resorbed and production of new ones ceases. Control of clutch size by the brood is an adaptation to the nest structure and life history of O. cinctus. It may have an important role in the reproductive strategy of other insects with parental care.     

Duration and benefits of biparental brood care in the dung beetle Onthophagus vacca (Coleoptera: Scarabaeidae)

Abstract.

• 1 Single males, single females or pairs of dung beetles, Onthophagus vacca, were released on artificial small (100 g) or large (1000 g) dung pats in the laboratory. Emigrating beetles were trapped at 12 h intervals, and the number and size of the brood chambers were recorded after each replicate.
• 2 Emigration of males was delayed if females were present in the same dung pats, whereas emigration times of females were independent of the presence or absence of males.
• 3 A residency of 60 h proved to be a threshold value. Females emigrating before this time did not breed, whereas those emigrating later had built at least two brood chambers.
• 4 Females paired with males built more brood chambers than single females.
• 5 The reproductive success of pairs was not influenced by the size of the dung pats.

     

Effect of food and larval density on survival and growth of early instar greyback canegrub, Dermolepida albohirtum (Waterhouse) (Coleoptera: Scarabaeidae)

Abstract First- and early second-instar larvae of Dermolepida albohirtum (Coleoptera: Scarabaeidae) survived best and gained most weight on living roots of lawn seedlings, sugarcane or Guinea grass ( Panicum maximum var. maximum ) when compared with decaying plant material or soil alone. Survival of first-instar D. albohirtum was not density-dependent, but survival of older larvae (late first-instar and early second-instar) was reduced at high larval density. There was no evidence for larval combat between first instars. First instars kept at high density gained less weight and were slower to develop into second instars compared with first instars kept at low density. When food was limited, survival at high larval densities was reduced. Survival of early instars was high in sugarcane fields, probably reflecting the availability of suitable food. We conclude that food type and supply are critical factors affecting the survival and development of early instar D. albohirtum .     

Two Newly Recorded Coprophagid Species from Korea (Coleoptera, Scarabaeidae and Aphodiidae)

A species of Scarabaeidae, Panelus parvulus Waterhouse and a species of Aphodiidae, Aphodius fossor (Linné) are newly recorded from Korea. The diagnosis and photographs of them are provided.     

Nest construction, fighting, and larval behaviour in a geotrupine dung beetle, Ceratophyus hoffmannseggi (Coleoptera: Scarabaeidae)

A pair of beetles worked together to make a series of horizontal brood masses by packing dung into underground chambers. An egg was laid in the soil just beyond the tip of each mass. Resident beetles used their horns to resist intruders of either sex. During head-to-head pushing contests in the nest entrance-tunnel, a stable position was adopted where the horns of each opponent thrust harmlessly against the pronotum of the other. The larva prevented collapse of its chamber by building a rigid tube from accurately positioned faecal pellets. Defects in the larval chamber released no obvious repair behaviour and the larva was, therefore, not able to survive in a free-standing brood ball or in the presence of another larva in the same chamber.
It is a pleasure to thank Sr J. de Ferrer (8 Av. F. Franco, Algeciras) for his help and advice.     

Nest construction and larval behaviour of Bubas bison (L.) and Bubas bubalus (01.) (Coleoptera, Scarabaeidae)

Abstract.

• 1 Female beetles working alone or in cooperation with a male excavated vertical, tunnel-shaped brood chambers. Each chamber was filled with dung to form a cylindrical brood mass which contained two eggs, one near each pole.
• 2 To examine the possible relationship with other Onitini (which lay either one or several eggs per brood mass) factors that influence the two-egg programme were studied. Brood masses with only a single egg were formed if excavation was resumed prematurely. Conversely, when excavation was suppressed several oviposition programmes fused to produce a multi-egg brood mass.
• 3 The larvae repaired their chambers in the typical Scarabaeine manner by building a self-supporting wall formed from their own excrement. This behaviour also prevented direct contact and fighting between adjacent larvae in the same brood mass, and it allowed the larvae to survive inside artificial brood balls. Similar behaviour was observed in larvae of Onthophagus taunts and Ontho-phagus vacca (which develop in one-egg brood masses). The evolution of nesting habits that involve multi-egg brood masses or free-standing brood balls may depend on the pre-existence of this larval repair behaviour.

     

Relative abundance and species composition of some dung beetles (Coleoptera: Scarabaeinae) in Bangladesh

1. The occurrence and community structure of dung beetle species (Coleoptera: Scarabaeinae) in the faecal deposits of man and some domestic vertebrates varied according to dung type, seasonal conditions and soil type. 2. Of the twenty-four species belonging to six genera, Onthophagus was the prevalent genus and O. ramosellus the most abundant species. 3. Cattle dung was the dominant food type.     

Effect of agronomic factors on the spatial distribution of greyback canegrub, Dermolepida albohirtum (Waterhouse) (Coleoptera: Scarabaeidae), in sugarcane in the Burdekin River Irrigation Area

Abstract A series of studies were undertaken examining the factors that impact on the spatial distribution of greyback canegrub infestations in sugarcane in the Burdekin region of Queensland. Historic records of damage by greyback canegrub showed that sugarcane blocks planted or harvested earlier than surrounding blocks were more likely to be damaged than blocks planted or harvested later. The derived hypothesis that sugarcane height may be the primary determinant of where damage occurs was confirmed in field studies. The tallest sugarcane blocks at the time of oviposition consistently had the highest grub populations. There was no difference in the distribution of damage between the cultivars Q96 and Q117 and the age of these crops also had no impact. The finding that sugarcane height is the primary determinant of where damage occurs on Burdekin farms opens the possibility of using it as a tool to manipulate where canegrubs oviposit and the development of a range of associated cultural control strategies.     

Phylogeography of the Namib Desert dung beetles Scarabaeus (Pachysoma) MacLeay (Coleoptera: Scarabaeidae)

Aim Namib biogeography in many instances remains reliant on advanced and detailed systematic studies. This study attempts to combine molecular phylogenetic data, geology and palaeo‐climatic data to (i) resolve the relationships of the 13 morphological species of Scarabaeus (Pachysoma) and (ii) relate their evolution to past climatic and geological events. Location South Africa and Namibia. Methods Sequencing of a 1197 bp segment of the mitochondrial cytochrome oxidase I (COI) gene of the 13 species within Scarabaeus (Pachysoma) was undertaken. Analyses performed included Parsimony and Maximum Likelihood as well as imposing a molecular clock. Results The molecular phylogeny showed strong support for 11 of the 13 morphological species. The remaining two species, S. (P.) glentoni and S. (P.) hippocrates, formed a complex and could not be assigned specific status on the basis of the COI gene phylogeny. Strong support for the three species formerly classified within the genus Neopachysoma was consistently obtained. The subgenus appears to have arisen c. 2.9 Ma. Species within the subgenus arose at different times, with the common ancestor to Neopachysoma and the hippocrates complex having evolved 2.65 and 2.4 Ma, respectively. Scarabaeus (P.) denticollis, S. (P.) rotundigenus, S. (P.) rodriguesi and S. (P.) schinzi are some of the youngest species, having diverged between 2 million and 600,000 years ago. Main conclusions Scarabaeus (Pachysoma) is a derived monophyletic clade within the Scarabaeini. The subgenus appears to be young in comparison with the age of the Namib Desert, which dates back to the Miocene (c. 15 Ma). The psammophilous taxa are shown to disperse with their substratum and habitat, barchan dunes. Clear south/north evolutionary gradients can be seen within the species of this subgenus, which are consistent with the unidirectional wind regime. Species with a suite of mostly plesiomorphic characters have a southerly distribution while their derived psammophilous relatives have central to northern Namib distributions. Major rivers such as the Orange, Buffels and Holgat appear to be gene barriers to certain species as well as areas of origin of speciation events.     

Interactions between dung beetles (Coleoptera: Scarabaeidae) and the arbovirus vector Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae)

Abstract  The potential for dung beetles to reduce populations of the biting midge and arbovirus vector Culicoides brevitarsis in bovine dung was studied in the Hunter Valley, New South Wales (NSW) between 1999 and 2003 using natural populations of insects. Preliminary work to develop experimental procedures showed that: a few C. brevitarsis could survive in buried dung; dung beetles and C. brevitarsis coming to dung were unaffected by a background of shade-cloth used experimentally to prevent dung burial; the most abundant dung beetle, Onthophagus gazella L. and C. brevitarsis oviposition occurred concurrently in the first 2 d after dung deposition, and the potential for interaction between dung beetles and C. brevitarsis was greatest in open pasture adjacent to trees where cattle congregate at night. Laboratory experiments on dung burial showed that C. brevitarsis numbers decreased as numbers of dung beetles increased or as the dry weight of dung decreased due to burial. This was seldom reflected in the field where, although significant burial occurred experimentally in 9 of 20 trials over 3 years, a significant decline in C. brevitarsis numbers attributable to burial only occurred once. C. brevitarsis numbers in the field were lower in unburied dung in 70% of trials. Differences were significant twice and were considered the result of dung disturbance. In the laboratory, decreasing numbers of C. brevitarsis were related to three characteristics of disturbance: the flattening, spreading and reduction in wet weight of the dung. Evidence of C. brevitarsis activity throughout coastal NSW suggests that, while C. brevitarsis numbers may be modified by dung beetles, the interaction is insufficient to prevent their increase, spread and ability to transmit viruses to livestock.     

Description of the larvae of Dicronocephalus wallichi bourgoini Coleoptera: Scarabaeidae: Cetoniinae) with observations on nesting behavior and life cycle of two Dicronocephalus species under laboratory conditions

Larva of the fruit beetle Dicronocephalus wallichi bourgoini Pouillaude 1914 is described and illustrated. The following unique morphological characters within Cetoniinae are discussed: shape of claw, spindle-shaped last antennomera, general body shape. The presence of metathoracic egg-bursters in the first instar larva has been found. Its significance for distinguishing the first instar larvae of Cetoniinae is discussed. The nesting behaviours of D. wallichi bourgoini and D. adamsi(Pascoe 1863) are described. The course of the life cycle of both species under laboratory condition is presented and discussed.     

Site‐specific profiles of biochemical properties in the larval digestive tract of Japanese rhinoceros beetle,Trypoxylus dichotomus (Coleoptera: Scarabaeidae)

The larvae of Japanese rhinoceros beetle, Trypoxylus dichotomus, feed on dead plant material in forest soils that are derived from fallen leaves broken down by basidiomycete fungi. Our previous work provided an understanding of the degradation of polysaccharides in dead plant material by T. dichotomus larvae and reported the complexity of the physicochemical and biochemical environment of the larval gut. Here, we examined ten divisions of the digestive tract of T. dichotomus larvae for physicochemical and biochemical conditions to elucidate site‐specifically functional properties along the tract. The distribution of potassium ions, pH, and acetic acid differed markedly along the length of the digestive tract with the potassium ion concentration profile closely reflecting that of pH along the length of the digestive tract. Distinct physicochemical environments were maintained in the digestive tract along with site‐specific polysaccharide degradation. Based on these findings, we suggest that there are metabolic relationships between the activities of the enzymes involved in polysaccharide degradation, the presence of intermediate metabolites and location along the digestive tract. Furthermore, we revealed that the anterior region of the gut plays an important role in the degradation of polysaccharides in the digestive tract of T. dichotomus larvae.