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.     

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.     

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.

     

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 Onitis belial and Onitis ion (Coleoptera, Scarabaeidae)

Abstract. 1. Female beetles working alone or in cooperation with a male buried dung to make brood masses. O.belial brood masses were packed close together in clusters; each mass was constructed as a horizontal thick-walled tube of dung which was filled with a dung sausage containing two to six eggs. O.ion made vertical sausage-shaped brood masses with one to four eggs.
2. The larvae of both species were able to survive in artificial brood balls as well as in multi-egg brood masses because of their ability to repair larval chambers with their own excrement.
3. The multi-egg brood mass of Onitis has probably evolved from a simple one-egg brood mass. It does not resemble the underground dung mass from which brood balls are made by certain Coprini.     

Nesting behaviour of a subsocial African ball-roller Kheper platynotus (Coleoptera, Scarabaeidae)

ABSTRACT. 1. The nesting behaviour of Kheper platynotus (Bates) was observed in the Tsavo West National Park, east Kenya.
2. The construction of the brood ball seemed to be initiated by the female; thereafter, the male joined in. After its construction, the ball was rolled and buried by the male, whilst the female rode on the ball and occasionally helped the male.
3. After burying the ball, the male came out on the ground, leaving the female under-ground. The female coated the ball with soil and her excrement, and let it ferment for about 2 weeks. Thereafter, the female converted the ball into one to four free-standing brood pears, and laid one egg in each pear. The number of pears produced depended on the size of ball. Even after oviposition, the female remained in the nest to care for the brood.
4. Oocytes matured a little before being laid. Excess oocytes were gradually assimilated during brood care.
5. Females seemed most likely to breed in two or more successive rainy seasons.
6. Although male-male aggression was observed, size assortative mating was not observed.     

Payoffs of the two alternative nesting tactics in the African dung beetle, Scarabaeus catenatus

1. Scarabaeus catenatus is a ball-rolling scarab in the subfamily Scarabaeinae. This species, however, makes use of two tactics for nest building: rolling and tunnelling. The tunnelling tactic differs substantially from the rolling tactic in that (1) it always involves repeated movements to and from the dung source and the nest, whereas rolling does not, and (2) it involves a shorter distance between the two sites.
2. Brood-nest founders were usually males and less often females, with about 25% adopting the rolling tactic and 75% adopting the tunnelling tactic. During nest building, the founder paired off with a scarab of the opposite sex, and they co-operated in the work. The female made one to four brood balls from the dung in the nest, each of which contained one egg.
3. Each scarab seemed to be able to employ both tactics. The tactic employed was independent of an individual's status, e.g. body size and timing of nest founding.
4. The rolling tactic offered only male founders a greater nest-defence success than the tunnelling tactic due to a lower intrusion into the rolled nest and a higher intensity of male–male fighting. The tunnelling tactic offered both male and female founders a larger number of brood balls than the rolling tactic because it enabled scarabs to take a larger amount of dung into the nest.
5. The reproductive success for the two tactics was estimated from the product of nest-defence success and the number of brood balls. As a result, the two tactics had equal fitness payoffs for males, but unequal payoffs for females.
6. The results suggest that male alternation of tactics is controlled by a mixed strategy. Female alternation, however, cannot be explained by mixed strategy, alternative strategies or conditional strategy.     

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.

     

Development of Copris tripartitus (Coleoptera: Scarabaeidae) in Two Different Rearing Media

ABSTRACT Like many other sub-social insects, Copris tripartitus Waterhouse requires parental care during the development period in the brood balls. Mass-rearing is achieved by using indoor rearing systems and adding more dung after taking away the brood balls containing eggs. Without parental care, larvae are often affected by desiccation, as well as by fungal or sciarid fly intrusion of the brood balls. This study developed a good substitute for brood ball rearing medium. A significantly greater number of progeny survived to adulthood from broods produced by beetles reared on vermiculite medium (90.69±10.98%) than on peat moss medium (44.82±13.92%). Results of the study suggest that to protect the brood balls from desiccation and to produce healthy adults, 15-20% moisture content vermiculite must be used as brood ball rearing medium for most species reared in the laboratory.     

Do weather conditions correlate with findings in failed, provision-filled nest cells of Megachile rotundata (Hymenoptera: Megachilidae) in western North America?

Cavity-nesting alfalfa leafcutting bees, Megachile rotundata (F.) (Hymenoptera: Megachilidae), are excellent pollinators of alfalfa, Medicago savita L., for seed production. In commercial settings, artificial cavities are placed in field domiciles for nesting and, thereby, bee populations are sustained for future use. For this study, cells from leafcutting bee nests were collected in late summer from commercial seed fields. Over 3 yr (2003-2005), 39 samples in total of approximately equal to 1,000 cells each were taken from several northwestern U.S. states and from Manitoba, Canada. X-radiography of 500 cells from each sample was used to identify "pollen balls" (i.e., cells in which the pollen-nectar provision remained, but the egg or larva, if present, was not detectable on an x-radiograph). Most U.S. samples seemed to have higher proportions of pollen ball cells than Manitoba samples. Pollen ball cells were dissected to determine the moisture condition of the mass provision and true contents of each cell. Most pollen ball cells from Manitoba samples contained fungus, the frequency of which was positively correlated with cool, wet weather. In the United States, most pollen ball cells had moist provisions, and many of them lacked young brood. Correlation analysis revealed that pollen ball cells occurred in greater proportions in fields with more hot days (above 38 degrees C). Broodless pollen ball cells occurred in greater proportions under cool conditions, but dead small larvae (second-third instars) seemed to occur in greater proportions under hot conditions. Pollen ball cells with unhatched eggs and first instars (in the chorion) occurred in lesser proportions under hot conditions.     

Egg removal by cuckoos forces hosts to accept parasite eggs

Many avian brood parasites remove one or more host eggs before laying their own eggs in the host nest. Various hypotheses have been proposed to explain the adaptive significance of this behaviour, but none of them provides an adequate explanation. Here we provide a new hypothesis for explaining why a parasite removes host eggs before laying its own. In this study, we attempted to answer this question by constructing a mathematical model that focused on the changes in host decision making according to reduced clutch size as a consequence of egg removal by parasites. We assume that a host selects one of the following two options to maximise the number of its own chicks: trying to eject a suspicious egg from the nest (trying‐to‐eject) or acceptance without trying to eject the egg (acceptance). The option selected depends on the number of eggs in the nest. Our model provides a new explanation for egg removal behaviour by showing that the host should select trying‐to‐eject if there is a large number of eggs in the nest but acceptance with a small number of eggs. This is because the relative payoff for a host that selects trying‐to‐eject decreases with the number of eggs in the nest. Therefore, parasites benefit by removing the host egg because this behaviour reduces the number of eggs in the nest, thereby increasing the probability of their own eggs being accepted. Thus, hosts have evolved egg ejection to combat brood parasites, but it may also have facilitated the evolution of egg removal by parasites. This hypothesis may also apply to brood parasitic species that do not eject host chicks. In addition, this hypothesis may explain other parasitic behaviours, such as egg damaging and egg puncturing, which lead to reductions in the host clutch size.     

Brood ball size but not egg size correlates with maternal size in a dung beetle,Onthophagus atripennis

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The earliest burst of necrophagous dung beetles in South America revealed by the Cenozoic record of Coprinisphaera

New ichnological, sedimentological and palaeobotanical information from a Miocene palaeosol succession from Patagonia bearing abundant fossil brood balls of dung beetles (Coprinisphaera) allow inferring novel aspects of the evolutionary history, biology and feeding habits of Scarabaeinae, along with the palaeoenvironmental and palaeoclimatic conditions in which they nested. Coprinisphaera tonnii and Coprinisphaera akatanka, both attributed to necrophagous dung beetles, represent 40.2% of the specimens. Considering their scarce or null record in older Cenozoic units from South America, these high values reveal that the first burst of necrophagous Scarabaeinae took place during the Miocene. Some Coprinisphaera preserve characters that indicate the developmental stage of the ball occupant and the adult emergence success. Both ichnospecies show the egg chamber isolated from the provision chamber, which was associated with a higher preference of cleptoparasites for necrophagous balls. The comparison among the abundance of traces of nest intruders in Coprinisphaera attributed both to necrophagous and to coprophagous beetles from different units of South America indicates that the presence of such intruders would be independent of the nature of the organic matter contained within the balls. Phytolith analyses performed in Coprinisphaera and extant necrophagous and coprophagous balls indicate that the comparison between the relative abundance of phytoliths in the wall of the brood ball, their infilling and the bearing palaeosol is a useful tool for inferring the feeding behaviour of the trace makers and support the attribution of Coprinisphaera tonnii and Coprinisphaera akatanka to necrophagous Scarabaeinae. Sedimentary and palaeosol analyses indicate that the beetles would have nested in well-drained soils showing mollic-like features, in grass-dominated habitats, under a seasonal sub-humid, temperate–warm climate.     

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.

     

Mothers vigilantly guard nests after partial brood loss: a cue of nest predation risk in a paper wasp

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Sex-specific defence behaviour against brood parasitism in a host with female-only incubation

Nest protection against intruders is an indispensable component of avian parental care. In species with biparental care, both mates should evolve nest defence behaviour to increase their reproductive success. In most host-parasite systems, host females are predicted to have more important roles in nest defence against brood parasites, because they typically are primarily responsible for clutch incubation. Male antiparasitic behaviour, on the other hand, is often underestimated or even not considered at all. Here we investigated sex-specific roles in four aspects of great reed warbler (Acrocephalus arundinaceus) nest defence against a brood parasite—the cuckoo (Cuculus canorus), namely (1) mobbing, (2) nest attendance/guarding, (3) nest checking and (4) egg ejection. Using dummy experiments, simulating brood parasitism and by video-monitoring of host nests we found that males took the key roles in cuckoo mobbing and nest guarding, while females were responsible for nest checking and egg ejection behaviours. Such partitioning of parental roles may provide a comprehensive clutch protection against brood parasitism.     

Great Spotted Cuckoos Frequently Lay Their Eggs While Their Magpie Host is Incubating

Species that suffer from brood parasitism face a considerable reduction in their fitness which selects for the evolution of host defences. To prevent parasitism, hosts can mob or attack brood parasites when they approach the host nest and block the access to the nest by sitting on the clutch. In turn, as a counter‐adaptation, brood parasites evolved secretive behaviours near their host nests. Here, we have studied great spotted cuckoo (Clamator glandarius) egg‐laying behaviour and defence by their magpie (Pica pica) hosts inside the nest using continuous video recordings. We have found several surprising results that contradict some general assumptions. The most important is that most (71%) of the parasitic events by cuckoo females are completed while the magpie females are incubating. By staying in the nest, magpies force cuckoo females to lay their egg facing the high risk of being attacked by the incubating magpie (attack occurred in all but one of the events, n = 15). During these attacks, magpies pecked the cuckoo violently, but could never effectively avoid parasitism. These novel observations expand the sequence of adaptations and counter‐adaptations in the arms race between brood parasites and their hosts during the pre‐laying and laying periods.     

Biology and social behaviour of three species ofAnischnogaster (Vespidae,Stenogastrinae) in Papua New Guinea

Summary Level of social organization, adult behaviour, size and development of the brood in three species ofAnischnogaster are described. The normal colony size in all the three species was one female per nest, and colony size never exceeded two females per nest. The social interactions, which were only observed in one species, did not include any marked dominance behaviour, but there was clear caste differentiation, with the older female guarding the nest while the younger female foraged for food. InAnischnogaster sp. A only some eggs and larvae have abdominal secretion, while no secretion at all was found on the eggs and larvae ofA. laticeps. In spite of this, the Dufour's gland was found to be well developed. The significance of this is discussed. Females ofA. laticeps were found to fall into two groups distinguished by the length of the sting. The larvae seem quite similar to those of other Stenogastrinae and have, apparently, only four instars.     

Gut microbiota of dung beetles correspond to dietary specializations of adults and larvae

Vertebrate dung is central to the dung beetle life cycle, constituting food for adults and a protective and nutritive refuge for their offspring. Adult dung beetles have soft mandibles and feed primarily on nutritionally rich dung particles, while larvae have sclerotized mandibles and consume coarser dung particles with a higher C/N ratio. Here, using the dung beetles Euoniticellus intermedius and E. triangulatus, we show that these morphological adaptations in mandibular structure are also correlated with differences in basic gut structure and gut bacterial communities between dung beetle life stages. Metagenome functional predictions based on 16S rDNA characterization further indicated that larval gut communities are enriched in genes involved in cellulose degradation and nitrogen fixation compared to adult guts. Larval gut communities are more similar to female gut communities than they are to those of males, and bacteria present in maternally provisioned brood balls and maternal ‘gifts’ (secretions deposited in the brood ball along with the egg) are also more similar to larval gut communities than to those of males. Maternal secretions and maternally provisioned brood balls, as well as dung, were important factors shaping the larval gut community. Differences between gut microbiota in the adults and larvae are likely to contribute to differences in nutrient assimilation from ingested dung at different life history stages.     

Costs and consequences of superparasitism in the polyembryonic parasitoid Copidosoma koehleri (Hymenoptera: Encyrtidae)

Abstract.  1. Polyembryonic wasps provide dramatic examples of intra-specific developmental conflict. In these parasitoids, each egg proliferates into a clonal lineage of genetically identical larvae. If more than one egg is laid in a host (superparasitism), individuals of different clones may compete for food resources.
2. In the polyembryonic encyrtid Copidosoma koehleri , one larva per clone can differentiate into a sterile soldier. It is shown that soldiers are always females, and that they attack intra-specific competitors.
3. Research hypotheses were that (a) clones that develop in superparasitised hosts suffer heavier mortality than clones that develop in singly parasitised hosts, and (b) female clones cause higher mortality to their competitors than male clones, hence larval survival is lower in superparasitised hosts that contain females than in male-only broods.
4. The potential frequency of superparasitism in C. koehleri was manipulated by varying parasitoid–host ratios and exposure durations.
5. As parasitoid densities and exposure durations increased, the frequency of superparasitism rose, brood sizes increased, but the number of hosts that completed development was reduced. The number of offspring per parasitoid female decreased with increasing parasitoid–host ratios. Offspring size and longevity were inversely correlated with brood size. As superparasitism rates increased, fewer all-male broods were produced. Male–female broods were female-biased, suggesting selective killing of males by female soldiers. All-female broods were significantly smaller than all-male broods at high parasitoid densities only, possibly reflecting aggression among soldiers of competing clones.
6. The results support the working hypotheses, and suggest that female larvae outcompete males in superparasitised hosts.