Two nesting behaviours and life history of a subsocial African dungrolling beetle,Scarabaeus catenatus (Coleoptera: Scarabaeidae)

Nesting behaviour of a large-sized dung beetle, Scarabaeus catenatus (Gerstaecker), was observed at the Tsavo West National Park, south-eastern Kenya. Although this species is taxonomically a member of a dung-roller group (that is, subfamily Scarabaeinae), it had not only a rolling behaviour but also a tunneller behaviour for nesting. In the former case, the scarab rolled a chunk or a ball of dung some distance (0·5–15·5m) away from the dung pat and buried it under the ground. In the latter case, it dug a tunnel near the dung pat (0–1 m) and transported several pieces of dung into the burrow. In both cases, brood nests were completed by a female alone or by male-female co-operation. Four days after dung burial, the female made one to four brood balls out of buried dung, in each of which she deposited an egg. On the other hand, the male left the nest soon after the female completed oviposition. Even after oviposition, the female stayed in the nest and cared for her progeny until they emerged. This indicates that S. catenatus is subsocial. A major source of offspring mortality was likely to be predation by driver ants (Dorylus sp.). Most females seemed to breed one time in each of two or more successive rainy seasons.     

“Riding” behaviour by males of Conops quadrifasciata (Diptera: Conopidae): Do females set up “riders” as targets for takeovers by larger males?

Males of Conops quadrifasciata “claim” females in late morning and then spend the rest of the day exposed as “riders” to attacks by other males. Male riders can respond to an attack by performing an abdomen‐lift which denies the attacker access to the female's genitalia. Attacking males can tailor their behaviour to suit the prevailing conditions. If a rider is engaged in a “repeat” copulation (as he does at regular intervals), the attacker “waits” beside the rider. If the rider is not in genitalic contact with the female, the attacker tries to mate with her immediately. It is proposed that females deliberately expose their riders to takeovers in order to gain the fittest mate.     

The role of the calceoli in precopulatory behaviour and mate recognition of Gammarus pseudolimnaeus Bousfield (Crustacea,Amphipoda)

The importance of calceoli (male antennular organs) in mate recognition of Gammarus pseudolimnaeus Bousfield was examined in the laboratory. The relationship between female receptivity and their moult stage was examined by comparing the moult stage of amplexed and non-amplexed females. The former were clearly in later moult stages and thus closer to ovulation. Ablation of the antennular 2 flagella, which uniquely possess the calceoli, had no effect on the ability of males to find, evaluate and amplex with receptive females. Morphological examination verified the complex nature and apparent lack of innervation of calceoli. We propose that the calceolus functions as a general environmental vibrioreceptor, necessary in the male because of its greater level of activity.     

Antennae and the role of olfaction and contact stimulation in mate recognition by males of the pollinating fig wasp Ceratosolen gravelyi (Hymenoptera: Agaonidae)

Finding a mate is a crucial step in sexual reproduction. The mutualism between figs and their pollinators is exploited by many figwasps that develop within their galls and attain sexual maturity simultaneously. Male pollinators carry the specific burden of detecting and opening mating holes in galls containing conspecific females before mating with them. The mating behaviourand antennal sensilla of male pollinators associated with Ficus semicordata was investigated to understand mate recognition. Male Ceratosolen gravelyi located female-containing galls from a distance, but only attempted to chew a mating hole after antennal contact with the gall surface was made. They showed similar responses to females and body extracts but failed to respond to washed female bodies. This behaviour indicates that unidentified chemicals present on the body surfaces of females are sufficient to elicit olfactory attraction and tactile confirmation. Multiporous plate sensilla are candidates for olfactory reception from a distance while basiconic sensilla may be involved in contact chemoreceptor. All of the sensilla are highly localised, on the distal part of the terminal flagellomere, suggesting a response to selection for strong directionality in the complex odour environment of the fig in which messages produced by hundreds of females may confuse precise mate localisation.     

Expulsion and consumption of male ejaculates by promiscous female Euxesta eluta and Euxesta mazorca (Diptera: Ulidiidae)

While expulsion of male ejaculates by the females after copulation has been reported for various animal groups, expulsion followed by consumption of the expelled ejaculate is a rare behaviour outside spermatophylax-producing orthopterans. Among Diptera, this behaviour has been reported for a few species of Piophilidae, Empididae and Ulidiidae. Here we report on its occurrence among Euxesta eluta and Euxesta mazorca (Diptera: Ulidiidae). We also attempt to characterize the mating system of E. eluta in order to facilitate future hypothesis testing to understand the behavioural factors leading to the evolution of this peculiar behaviour. For this, courtship sequences, copulation duration, frequency of ejaculate expulsion and subsequent consumption, and latency to ejaculate expulsion for both E. eluta and E. mazorca were recorded. The time of sexual maturation, the time window of sexual receptivity during the day, and the mating frequency and variance in mating success for males and females (degree of polygamy) were determined for E. eluta. Both E. eluta and E. mazorca males engaged in elaborate courtship sequences involving visual and tactile displays before copulation. Females of both species almost invariably expelled and consumed ejaculates after copulation. Female E. eluta, required a 6- to 9-day period feeding on protein and sugar before becoming sexually receptive. Reproductive activity occurred continuously over the day with an early morning and late afternoon peak. Both males and females could mate multiply, with multiple partners over a 2-h observational period. Sexual network analysis revealed that some males and females had greater mating success than others. Ejaculate consumption appears to be widespread in the genus Euxesta. It is possible that females obtain nutrients from this behaviour while exerting some control over egg fertilization.     

XXX.—A new Cicada from Arabia

Summary

Carebara nests occur in between 2% and 10% of Macrotermes mounds, but can also occur well away from mounds. A brood of alate ants is regularly produced ready to fly in the short rains (November) but flights can also take place in the long rains (April). Male and female alates are usually produced in separate nests. After the flight, male alates seek out the females, and several males appear to mate with one female.

Dealate female ants exposed to Macrotermes workers remained passive and elicited no aggression. Fecundated female ants reared broods of about 1000 workers after 39 days in the laboratory. These workers were able to overpower, kill and eat the much larger workers of Macrotermes.     

Antennation and sexual performance of male digger wasps Sphex ingens Smith (Hymenoptera: Sphecidae)

Antennation can play several roles in hymenopteran copulation, mainly in wasps. The digger wasp Sphex ingens Smith has a very peculiar sexual behaviour, in which forced copulation is a striking element. However, communication through antennation during the pre-copulation and copulation phases and the relationship between the sexual performance of males and the final result of copulation require further clarification. The sexual behaviour of wild populations of S. ingens was filmed during the breeding season in a site between the beaches Meros and Aventureiro, Ilha Grande, southeastern Brazil. We assessed antennation behaviour, courtship duration, sexual performance, and genital clasping. Only successful males had the genital clasping assessed. Evidence found so far points out that the successful males showed better sexual performance – that is, they antennated more in a shorter courtship duration. Also, genital clasping in successful males did not depend on antennation, courtship duration, or sexual performance. Our results shed light on some aspects of the sexual behaviour of wasps, as they explain part of the sexual selection mechanisms adopted by the species.     

Spatial and temporal patterns of territorial mate locating behaviour in Hypolimnas bolina (L.) (Lepidoptera: Nymphalidae)

Male Hypolimnas bolina (L.) (Lepidoptera: Nymphalidae) defend perch sites as a method of visually locating receptive females. In this study we charted the spatio-temporal activity of perching males in tropical Australia to investigate (a) the extent to which male activity is thermally constrained, and (b) the physical attributes of selected perching sites. Butterflies were surveyed along two 5-15m wide open corridors through dense vegetation, and this allowed the prediction that males should favour narrower corridor sections in order to maximize their visual search capability. This prediction was supported. Beyond corridor width, the distribution of favoured perches was not explained any further by patterns of shading, larval food plant distribution, or the presence of nectar resources. Males were active from 0800 to 1700 h, but the number of perching individuals varied throughout the day, and this pattern varied between the two transects. Most individuals perched along one transect in the morning (0900-1100 h), whereas activity along the other peaked around midday (1100-1300 h). This between-transect difference in male activity followed changes in shading between the transects, however this variable did not predict male distribution at the territory scale. Although ambient and black body temperatures were significantly related to population-level activity, these variables only predicted 15-55% of the variance in male counts in individual transects. This result, viewed in conjunction with the limited available information on female receptivity, suggests that the timing of mate location in H. bolina may be primarily influenced by the daily pattern of female availability.     

Copulatory courtship and morphology of genitalic coupling in seven Phyllophaga species (Coleoptera: Melolonthidae)

Males in six of seven species of Phyllophaga perform apparent courtship behaviour during copulation. As predicted by the hypothesis that such copulatory courtship is under sexual selection by female choice, behavioural details differ between species. The modified male abdominal sternites, which rub against the female's pygidium during copulation and whose morphology also differs between species, and the flattened, setose tarsi of males of one species probably function as contact courtship devices during copulatory courtship. Observations of events inside copulating beetles were made through the semi-transparent abdominal sternites of females and male genitalia in three species. They revealed cryptic genitalic thrusting behaviour of male genitalia within the female which occurred even when there was little or no external movement of the male genitalia. Dissections of pairs frozen in copula demonstrated that males transfer large quantities of material to females during copulation that is not physically related to sperm transfer. The type of material varied between species. In at least two species most of the transfer apparently occurred after sperm transfer. The possibility that some male genitalic structures function as holdfasts and/or stimulatory devices was suggested by their positions relative to female structures, and/or their movements within the female. Portions of male genitalia with strong bristles tended to lie against setose or more heavily sclerotized portions of the female reproductive duct. Some portions of female genitalia which vary between species seem ill-designed to exclude the genitalia of cross-specific males. The female reproductive tract is partially everted during copulation in one species.     

Palpal loss,single palp copulation and obligatory mate consumption in Tidarren cuneolatum (Tullgren, 1910) (Araneae,Theridiidae)

Males of Tidarren cuneolatum (Tullgren, 1910) amputate one of their palps a few hours after the penultimate moult, like T. sisyphoides (Walckenaer, 1841) from the USA (Branch, 1942). Hence adult males, which are minute, have only one palp, either left or right randomly. This palpal organ is not oversized, when compared with other small spiders. During courtship females are unusually active, signalling receptivity by continuous twanging with legs II. Males construct a mating thread. Copulation involves one insertion, which lasts ca 4 min. Thus, only one receptaculum is inseminated during copulation. With the advance of insertion the male's prosoma becomes shrunken. Copulation regularly ends in mate consumption. In copulation with a virgin female the palp was inserted contralaterally. Females taken in the field had both receptacula filled with sperm and therefore were polyandrous. Re-mating was also observed in the laboratory. Remarkably the second male performed an ipsilateral insertion, if it possessed the same palp as the previous male. Probably the virgin receptaculum was recognized. Postembryonic development is rapid in males, which moult three times and mature ca 41 days after hatching from the cocoon. Females need four or five moults and ca 69 days to reach maturity and then survive ca 2-4 months.     

Population dynamics,age structure and sex ratio of the bromeliad-dwelling jumping spider,Psecas chapoda (Salticidae)

In this study, the population fluctuations, phenology and sex ratio of the bromeliad-dwelling jumping spider Psecas chapoda were investigated in an area of semi-deciduous forest in south-eastern Brazil. Psecas chapoda occurred and reproduced on Bromelia balansae (Bromeliaceae) throughout the year. The number of egg sacs increased at the beginning of the rainy season (September to December), but the population size and phenology were stable over time and correlated weakly with rainfall and temperature. The principal factor affecting the dynamics of P. chapoda was the blooming of B. balansae, which expelled spiders because of the drastic change in plant architecture. Male spiders were more frequent than females during the juvenile phase, whereas females were more frequent than males during the subadult and adult phases, suggesting a sex ratio that favoured males. The biotic and abiotic factors that influence the population dynamics and sex-dependent vulnerability determination are discussed.     

Sperm viability in the reproductive tract of females in a population of Sceloporus mucronatus exhibiting asynchronous reproduction

Asynchronous reproduction is a common phenomenon in high-elevation populations of lizards from Central México. Sperm storage in the reproductive tract of females is the mechanism for making oocyte fertilization possible. Our study addresses questions related to functional oviductal sperm storage of females mating on different dates throughout the reproductive season. A population of Sceloporus mucronatus with copulation in the summer and ovulation in the fall was chosen for this experiment. Eleven females that copulated in the field during June and 13 females that copulated in captivity during August were maintained in the laboratory until parturition. The number of pregnant females and the litter sizes produced in each experimental group were indicative of the viability of the stored sperm. Sperm stored in the reproductive tract of females were able to fertilize eggs after 4 months. No significant differences were found in the number of pregnant females between the 2 experimental groups nor in the litter sizes that they produced. We found that the amount of time sperm were held in the female reproductive tract (ca. 3 months) had no effect on the capacity of sperm to fertilize eggs. Histological examination of 8 oviducts collected before the mating season eliminated the possibility of sperm storage from one year to the next. In this system, sperm retention could have evolved as a response mechanism to deal with the asynchrony between sexes in the reproductive cycles. However, we cannot rule out alternative hypotheses.     

Mating behaviour and spermatophore transfer in the shrimp Heptacarpus pictus (Stimpson) (Decapoda: Caridea: Hippolytidae)

Summary

Heptacarpus pictus, a small caridean shrimp inhabiting the low intertidal of southern and Baja California, breeds during the winter, spring, and summer months. Fall is a period of growth. Life span of an individual does not exceed 18 months, with fish predation as the most likely source of mortality.

Females are multiple brooders, carrying developing embryos concomitant with increase in ovary size. Hatching of larvae is followed by a moult, after which the female is attractive to males and receptive to copulation. A distance pheromone does not appear to be involved in attraction of males to females. Males apparently respond to a non-diffusible substance on the exoskeleton of newly moulted females.

Precopulatory behaviour is absent. Copulation can be divided into a series of relatively stereotyped events. Female rejection of the male or his spermatophore is the chief cause of unsuccessful matings.

Males deposit the spermatophore on the underside of the female's first abdominal segment. Sperm packets are formed upon extrusion from the male's genital openings, and are composed of a mucoid material in which sperm are mixed. The glutinous spermatophores adhere to the female's smooth abdominal sternite.

The endopods of pleopods 1 and 2 of the male are different in shape, size and setation from homologous rami of females and juveniles. Endopods of pleopod 1 possess a distally located appendix interna, absent in juveniles and females. An anteriorly projecting process, the appendix masculina, is located on the endopod of pleopod 2 in males. Experiments were performed which showed that these modifications insure proper deposition of spermatophores. Males which had the copulatory rami removed did not transfer spermatophores as successfully as normal males.

Transfer of the spermatophore from the male to the female is a result of the interplay of male pleopods 1 and 2 during copulation. The large expanded abdominal pleurae of females prevent the male's genital opening from contacting her abdominal sternite. Thus, the male's anterior pleopods have become modified to lift the emitted spermatophore from his genital orifices to the first abdominal sternite of the female.     

Reproductive strategies of two sympatric species of Hyalella Smith, 1874 (Amphipoda,Dogielinotidae) in laboratory conditions

This study was performed in order to test theoretical predictions that sympatric species might show different reproductive strategies to facilitate their coexistence. The duration of precopulatory mate‐guarding behaviour, body size of ovigerous females, embryonic and postembryonic development time, fecundity, and viability of the sympatric amphipod species Hyalella pleoacuta and H. castroi were investigated under laboratory conditions (19±1°C and 12 h light/dark photoperiod). Specimens were collected with nets in autumn and spring 2004, and summer and winter 2005 in the Vale das Trutas, São José dos Ausentes County, southern Brazil, and transported to the laboratory. Both species of Hyalella showed similar reproductive strategies, especially in respect to the duration of the precopula, duration of the embryonic period, and fecundity. However, these species differed in body size of ovigerous females, duration of the postembryonic period, viability, and body size of juveniles that remained in the marsupium and those released by females. These differences in the reproductive traits of H. pleoacuta and H. castroi may facilitate their coexistence in nature.     

Spatial patterns of the semi-aquatic rodent Nectomys squamipes in Atlantic forest streams

Nectomys squamipes is a semi-aquatic rodent with a wide geographic range in Brazilian forests and savannas. This species is a host for several parasites associated with aquatic environments, especially Schistosoma mansoni, and thus its spatial pattern may influence some diseases’ patterns. Here we present important aspects of water rat spatial behaviour, including technical and ecological aspects. Water rats were studied by live trapping in rivers and in gallery forests between January 2005 and August 2008, with a total sampling effort of 9511 trap-nights. We also studied some animals through radio tracking and the spool-and-line technique. Water rat movement was distributed evenly between the terrestrial and aquatic environments. Nectomys squamipes occurrence in small tributaries was higher than in larger and lotic rivers. Both home range, the conventional two-dimensional approach, and home length, a one-dimensional approach, proved to be useful to study N. squamipes, depending on the shape of the water body used by the individual. Although Nectomys squamipes’ home length was positively correlated with body weight, sex was also important to explain home length variation; at similar body weights, males have larger home lengths than females. Males explored larger areas during the rainy season, likely in search of females, as this is the water rat reproductive season in Atlantic forest. In contrast, females explored larger areas during the dry season, likely looking for additional food supplies as a result of deficient water rat food supplies during the Atlantic forest dry season. Females overlapped home lengths with more than one male, and males with more than one female, suggesting a promiscuous mating system for this species.     

The green June beetle (Cotinis nitida) (Coleoptera: Scarabaeidae): local variation in the beetle’s major avian predators and in the competition for mates

The green June beetle (Cotinis nitida) was observed in northern Virginia for a second and third flight season. The study yielded some results that were consistent with earlier findings, namely that mate-locating beetles were vulnerable to attack by avian predators and matings occurred primarily early in the flight season. Novel findings included the following: (1) blue jays (Cyanocitta cristata) were the major predators, not common grackles (Quiscalus quiscala), (2) some jays appeared to selectively prey on female beetles, (3) competition for mates among male beetles could be so intense that some males attempted to copulate with already mated (unreceptive) females, and (4) the sex ratio of the beetle population feeding on wild blackberries remained near equality late in the flight season despite the fact that the sex ratio of the population of beetles at the emergence site became highly female-biased over time.     

Sexual dimorphism in chelicerae,forelegs and palpal traits in two burrowing wolf spiders (Araneae: Lycosidae) with sex-role reversal

Allocosa alticeps and Allocosa brasiliensis are two sex-role-reversed spiders. Females locate males and initiate courtship. Copulation occurs inside male burrows and after copulation the males deliver their burrows to the females. We explored the occurrence of differences in morphological traits related to sex roles in both Allocosa species and compared the results with two non-burrowing wolf spiders with typical sex roles. We measured the length of the foreleg's tibia-patella and the chelicerae. Scanning electron micrographs of palpal tarsus distal sections were taken. Males showed higher values than females in all the traits considered. Adult and penultimate males lacked true claws compared with juveniles and females, but showed modified spines. The palpal organ was more proximal to the tibia compared with the non-burrowing lycosids considered in this study. Spines and palpal organ location could be associated with more effective digging. We discuss how natural and sexual selection could interact on morphology in this scenario.     

Observations on zygopine weevil behaviour (Coleoptera: Curculionidae: Zygopinae)

Observations on the behaviour and chosen environment are reported of the zygopine weevils Mecopus audinetii, M. bispinosus, M. fausti, M. pulvereus, Phylaitis v-alba and Osphilia sp. Mecopus audinetii and M. bispinosus were not distinguished in the field, but can be separated by details of the male and female genitalia and by the presence in the male of M. bispinosus of long black hairs on the pronotum. All the species studied were found on the bark of fallen or dying trees, the Mecopus species and Phylaitis v-alba being associated with whitish-silvery bark and the Osphilia species with greenish-brown substrates. The small Phylaitis species was found on branches of smaller diameter than those frequented by the larger Mecopus species. All the species are fairly cryptic in coloration, but when moving the larger species are more visible than the smaller, and the ones on pale bark are more visible than those on greenish-brown bark. Potential predators may be confused by jerky movement patterns observed in all species and the distant shadows cast by the long-legged larger Mecopus. The weevils are not believed to be involved in any mimicry, although this has been suggested for other members of the same subfamily. A single contest between males of Mecopus audinetii/bispinosus was observed, in the absence of a female. Each male attempted to dislodge its opponent from the substrate using its rostrum; the thoracic spines were not seen to be used. Contests between males of Osphilia sp. and Phylaitis v-alba were observed in the presence of females of the appropriate species. The males of Osphilia species attempted to dislodge one another from the back of the female in contention, and also engaged in a spiralling flight together. The males of Phylaitis v-alba did not come into physical contact, but responded to the threat of an approach. During copulation in Mecopus audinetii/bispinosus and Osphilia sp. the male repeatedly brushed the head and thorax of the female with his front legs. In the first species the male stood on the substrate and ‘caged’ the female with his legs and rostrum but in the second the male, ‘rode’ the female. Both species oviposit into the bark of their trees, the females using the rostrum to drill a hole for the ovipositor. The length of the rostrum probably determines the thickness of the bark through which eggs are laid, and hence the area of the region of the tree inhabited by the beetles.     

The function of the sphragis in Cressida cressida (Fab.) (Lepidoptera,Papilionidae): a visual deterrent to copulation attempts

During mating, males of Cressida cressida (Lepidoptera, Papilionidae) form a large external mating plug or sphragis that covers the female's copulatory opening and physically prevents remating. The sphragis has lateral and distal projections that make it an obvious structure. We tested experimentally the hypothesis that the sphragis acts as a signal of female mating status. Males pursue mated females with an intact sphragis only briefly and rarely make physical contact with them. When the sphragis of a mated female is removed or reduced in size, males are significantly more likely to physically contact the female and initiate the aerial takedown that preceeds a copulation attempt. These results suggest that the sphragis deters male sexual interest at a distance and thus functions as a signal of female mating status. The discussion focuses on the fitness consequences for females and their mates of the signal function of the sphragis.     

Assortative mating in soldier beetles (Cantharidae: Chaulignathus ): test of the mate-choice hypothesis

Soldier beetles of 2 species, Chauliognathus basalis and C. deceptus , were examined to test the Crespi hypothesis that positive assortative mating by size is caused by mate choices. Specifically, we tested the prediction that if mate choice involves choosing the largest mate available, then mating individuals will be larger than nonmating individuals. Four samples were taken, at different times during the mating season, from each of 2 sites. Each sample consisted of mating pairs, nonmanting males, and nonmating females. Some of the samples contained beetles of both species; others contained beetles of a single species. For each gender elytron lengths of mating individuals were compared with elytron lengths of nonmating individuals. We found no effect of mating status (mating vs. nonmating) on elytron lengths in samples that exhibited assertive mating (which occurs where 2 species coexist). Surprisingly, we found a consistent effect of mating status on elytron lengths in samples that did not exhibit assortative mating (which occurs where only 1 species exists). Our results do not support the mate-choice hypothesis. Instead, mate choice and assortative mating appear to be alternative mating patterns in which mate choice occurs where a single species exists and assortative mating occurs where 2 species coexist.