Notes on the mating system of Hylesinus varius (F.)(Col., Scolytidae), a putatively bigynous bark beetle

Abstract: Previous research has left confusion about central features of the mating system of the common ash bark beetle Hylesinus varius; in particular, reports conflict as to whether this species is monogynous or bigynous. It was found that females initiated egg tunnel construction; copulation takes place either on the bark in the initial phase of egg tunnel construction, or inside the tunnel system. Observations of one male copulating with several females and of solitary females in galleries with eggs suggested that the males sometimes leave after copulation. In contrast to findings for populations in Slovakia, Norwegian male H. varius were most commonly paired with only one female. Bigynous galleries contained significantly more eggs than monogynous galleries, but bigynous males did not double their reproductive success as found in some other bigynous bark beetles. This suggests that bigynous galleries at first are the work of one female but some males are later joined by a second female.     

Breeding season influxes and the behaviour of adult male samango monkeys (Cercopithecus mitis albogularis)

Troops comprising a high density population of samango monkeys (Cercopithecus mitis) in Natal province, South Africa, experienced an influx of adult males during the breeding season. Observation of one troop revealed that these males competed with one another and with two resident males for access to receptive females. Although both sexes initiated copulation, attempts to do so were more often successful if female-initiated. Males did not interact with non-receptive females and there were no recorded attempts at infanticide. Male-male interactions were agonistic in the presence of receptive females and neutral at other times. No ritualized displays of dominance and subordinance were seen. The significance of these observations for male reproductive strategies is discussed.     

Male egg-brooding in the millipede Yamasinaium noduligerum (Diplopoda: Andrognathidae)

Yamasinaium noduligerum is a Japanese fungivorous millipede that is usually found on decaying logs fallen on the forest floor. Here we present observations on paternal egg-brooding behavior in this species. At the end of May to early June, we found males with eggs on the bark of the underside of fallen decaying logs and in old burrows and galleries excavated by wood-dwelling insects under the bark of the logs. The males showed a typical brooding posture where they curl the anterior part of the body around the egg mass. The egg masses contained, on average, 30.2 ± 13.7 (SD) eggs. The brooding males showed no aggressive responses when disturbed.     

Effects of density on the reproductive success of pine engravers: is aggregation in dead trees beneficial?

1. The pine engraver bark beetle Ips pini (Say) (Coleoptera: Scolytidae), aggregates primarily on dead or dying pine trees. In this study pine engravers were laboratory-reared on logs at a range of low densities to determine whether there was a fecundity advantage of breeding aggregations.
2. Mean reproductive success for both males and females declined exponentially with increasing density.
3. Female pine engravers had shorter egg galleries at higher densities, suggesting that they left high-density breeding sites earlier. This would reduce the number of eggs that failed to survive due to larval competition.
4. Some pine engravers colonized the logs voluntarily during the experiment. These volunteers settled independently of the original density.
5. The fungus Ophiostoma sp. was present on the logs and may be competing with the pine engravers for limited bark area.
6. Aggregation resulted in a considerable cost to pine engraver reproductive success even at low densities. Thus, it remains perplexing why pine engravers aggregate actively in nature.     

Tree wetas (Hemideina crassicruris) (Orthoptera: Stenopelmatidae) of Stephens Island,Cook Strait

Abstract

Tree wetas (Hemideina crassicruris) were studied in kohekohe (Dysoxylum spectabile) forest on Stephens Island, Cook Strait, between 5 and 17 January 1978. Mark and recapture observations suggested a population density of at least 5300 wetas ha^?1 and a biomass of 42 kg ha^?1. Activity of wetas increased with temperature and humidity. Tuataras are important predators on Stephens Island and probably deter wetas from moving onto the forest floor. The enlarged head and jaws of male wetas are associated with head-on fights for occupancy of galleries. Single, adult males occupy galleries with a harem of up to 9 females. Stridulation was associated with agonistic encounters and was most frequent shortly before dawn. Seventeen percent of males and 42% of females were damaged. Moulting, copulation, and oviposition were observed, and females carried up to 96 mature eggs. Stephens Island tree wetas reach larger sizes overall, and males have relatively larger heads, than in other populations of the Cook Strait region. The number and size of galleries and the presence or absence of predators may be important determinants of weta population density, sex ratio, harem formation, and activity on the ground.     

Movement patterns and gallery use by the sexually dimorphic Wellington tree weta

The Wellington tree weta, Hemideina crassidens, is a harem-polygynous nocturnal insect whose males defend and mate groups of females residing in cavities in trees. In this study I examined sexual differences in gallery use (number of galleries occupied per unit time), distance travelled per night and activity patterns after sunset. In addition, I investigated how gallery size affected each of these variables. On average, males and females did not differ in the number of galleries visited per night, or in the distance they travelled per night; however, adult males were more likely to be seen earlier in the night than later. Although males and females did not differ in their gallery use or distance travelled, adult males occupied a significantly greater number of galleries and travelled greater distances per night at sites with small galleries than did males at sites with large galleries. These results suggest that habitat structure of a forest patch influences intrasexual tree weta behaviour.     

Effects of soil cations on the foraging behavior of Reticulitermes flavipes (Isoptera: Rhinotermitidae)

This study examined soil cation concentration gradients around decomposing logs and determined whether Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae) builds galleries in soil substrates containing those cations versus soils without, presumably in an effort to forage efficiently. Soil samples were collected at different depths and lateral distances from decomposing logs containing termites, freshly fallen logs without termites, and bare soil locations at 10 sites to determine whether cation gradients existed in a forest floor. Analyses of cation gradients were significant for potassium, calcium, and magnesium; however, only potassium showed a significant gradient where concentrations decreased with depth and lateral distance from a decomposing log. R. flavipes was then tested in laboratory bioassays to determine if they were attracted to salt cation solutions. Linear foraging distance was significantly greater in artificial K2CO3 gradients than in controls. The results from this study suggest that potassium is found in gradients around decomposing logs and that R. flavipes is attracted to this cation.     

The Relationship Between Resource Control, Association with Females and Male Weapon Size in a Male Dominance Insect

In species with a resource‐defence (male dominance) mating system, males are expected to maximize fitness by controlling resources deemed more valuable by sexually receptive females because these sites attract more mates. Furthermore, males, which control more valuable resources should themselves be of high quality. I experimentally tested these predictions in the laboratory using the sexually dimorphic Wellington tree weta, Hemideina crassidens (Blanchard) (Orthoptera: Tettigonioidea: Anostostomatidae). Male H. crassidens use their mandibular weaponry to fight for control of harems (groups of adult females) that seek shelter in trees cavities (galleries). As predicted, larger galleries housed significantly larger groups of females and males with larger weaponry controlled large galleries significantly more often. Therefore, galleries with a larger volume are likely considered more valuable by males because they house larger harems. However, contrary to prediction, males with larger weaponry did not reside with significantly more females overall because females did not always form the largest possible groups in galleries and males with smaller weaponry were able to reside with single females in small galleries. The latter observation suggests a possible alternative mating strategy by disadvantaged males.     

Formation and maintenance of mating aggregations in a stink bug,Megacopta punctissimum (Montandon) (Heteroptera: Plataspidae)

Adults of a stink bug,Megacopta punctissimum, formed mating aggregations on their host plants: a few pairs in copula and a few bachelor males (males not in copula) stayed in the aggregation and the bachelor males waited for arriving females to mate with. The processes of formation and maintenance of the aggregations were observed using the individual marking technique. Aggregations initiated by 2 males or by 1 male and 1 female were usually joined by 1 or more individuals and lasted for several days, but aggregations initiated by 2 females broke up within 1 h. Aggregations were not maintained by the same members. The residence time (time from joining an aggregation to leaving the aggregation without copulating) of males was longer than that of females. There was a negative correlation between the residence time of males and the number of bachelor males in an aggregation when bugs joined it, while the residence time of females was positively correlated with the number of bachelor males in the aggregation. When bugs copulated after joining an aggregation, the postcopulatory residence time (time from completing copulation to leaving the aggregation) was longer in males than in females. Thus, males had a stronger tendency to initiate and maintain aggregations than females.     

Identification of volatile emissions from Platypus mutatus (=sulcatus) (Coleoptera: Platypodidae) and their behavioral activity

We report here the identification and behavioral activity of volatile compounds emitted by male Platypus mutatus (=sulcatus) Chapuis while boring galleries in living poplar, Populus deltoides Marshall, trees. Headspace analysis using solid phase microextraction techniques showed the presence of 6-methyl-5-hepten-2-ol (sulcatol) and 6-methyl-5-hepten-2-one (sulcatone). Only one enantiomer of sulcatol, retusol, was found to be part of the volatile emission. Behavioral assays showed that females are more attracted than males to galleries with boring males inside. Both sulcatol and sulcatone elicited electroantennographic responses by female P. mutatus. Furthermore, behavioral bioassays showed that both sulcatol and sulcatone elicit behaviorally attractive responses by females. These results suggest that male P. mutatus releases a sex pheromone composed mainly of retusol and sulcatone.     

Observations on sex ratio and behavior of males in Xyleborinus saxesenii Ratzeburg (Scolytinae, Coleoptera)

Strongly female-biased sex ratios are typical for the fungalfeeding haplodiploid Xyleborini (Scolytinae, Coleoptera), and are a result of inbreeding and local mate competition (LMC). These ambrosia beetles are hardly ever found outside of trees, and thus male frequency and behavior have not been addressed in any empirical studies to date. In fact, for most species the males remain undescribed. Data on sex ratios and male behavior could, however, provide important insights into the Xyleborini's mating system and the evolution of inbreeding and LMC in general.In this study, I used in vitro rearing methods to obtain the first observational data on sex ratio, male production, male and female dispersal, and mating behavior in a xyleborine ambrosia beetle. Females of Xyleborinus saxesenii Ratzeburg produced between 0 and 3 sons per brood, and the absence of males was relatively independent of the number of daughters to be fertilized and the maternal brood sex ratio. Both conformed to a strict LMC strategy with a relatively precise and constant number of males. If males were present they eclosed just before the first females dispersed, and stayed in the gallery until all female offspring had matured. They constantly wandered through the gallery system, presumably in search of unfertilized females, and attempted to mate with larvae, other males, and females of all ages. Copulations, however, only occurred with immature females. From galleries with males, nearly all females dispersed fertilized. Only a few left the natal gallery without being fertilized, and subsequently went on to produce large and solely male broods. If broods were male-less, dispersing females always failed to found new galleries.     

Spinning behaviour and morphology of the spinning glands in male and female Aposthonia ceylonica (Enderlein, 1912) (Embioptera: Oligotomidae)

Embioptera (webspinners) are unique among insects in that juvenile and adults of both sexes spin silk. They possess spinning apparatuses in the basitarsomeres of their prothoracic legs, which they use to build galleries as habitat and protection. Embioptera are primitively social and cooperate in building the galleries. They also show sexual dimorphism that comprises modifications of the mandibles in males, the winglessness of the females and differences in the morphology of the forelegs. In the present investigation we address the correlation of spinning behaviour and sexual dimorphism in the spinning apparatus of Aposthonia ceylonica (Enderlein, 1912). To analyse spinning behaviour we conducted video observations of Ap. ceylonica in artificial habitats. We observed females and males alone as well as female-male pairs to cover possible effects of interactions between sexes. The morphology of the spinning apparatus was analysed and reconstructed using high resolution X-ray computed tomography (SRμCT). The observations show that during trials of 24 h adult males and females produce similar amounts of silk per body weight, despite the fact that adult males do not feed, perhaps due to modifications of their mandibles related to courtship that interfere with feeding. Spinning glands in males are distinctly smaller than in females in absolute values, which reflect the general size difference in females and males. Despite their smaller body size, the volumes of reservoirs of spinning glands are larger in males in relative as well as in absolute values. Together with spinning behaviour and the amount of silk production, this indicates that males produce and store gland secretions in the large reservoirs prior to their final moult for later use.     

Host preference and sex allocation of three hymenopteran parasitoid species (Ichneumonidae and Braconidae) of a longicorn pest, Oemona hirta (Fabr.) (Col., Cerambycidae)

The host preference and sex allocation of two ichneumond parasitoids, Xanthocryptus novozealandicus (Dalla Torre) and Campoplex sp., and one braconid parasitoid, Apsicolpus hudsoni Turner of the longicorn borer, Oemona hirta , were studied in New Zealand. The size of the borer larvae attacked by the parasitoids can be determined by measuring the gallery width of the borers. These parasitoids are idiobionts and their secondary sex ratio is female-biased. Females of these species are significantly larger than males, and the females allocate the sex of offspring depending on the size of host larvae, by laying female-producing eggs on significantly larger host larvae. Xanthocryptus novozealandicus is larger in body size and shorter in ovipositor length than Campoplex sp. and A. hudsoni , and consumes late instar larvae or prepupae in shallow galleries whereas the latter two utilize early to middle instar larvae in deep galleries. Campoplex sp. and A. hudsoni preferred host larvae in significantly larger tree twigs than X. novozealandicus .     

Sex ratio and female sexual status of the coconut pest, Oryctes monoceros (Coleoptera: Dynastidae), differ in feeding galleries and pheromone-baited traps

Oryctes monoceros is a serious coconut pest, causing up to 40% damage in tropical Africa. Synthetic aggregation pheromone, ethyl 4-methyloctanoate, has been used to lure adults to traps. Traps with pheromone plus decaying palm material captured a high proportion of males. This raises the question whether individuals, which damage palms are receptive to the pheromone. We studied the sex ratio of the insects feeding on coconuts and those attracted to pheromone traps. Sixty two percent of adults from feeding galleries on living coconut palms were females. Pheromone with rotting palm material lured 43% females. To investigate the reasons for this difference, we compared the reproductive system of females lured to the odour traps or feeding in coconut galleries, or present in old rotting stems. Ninety six percent of the females trapped by pheromone had mated, and were sexually mature. In the galleries on living palms, 46% of females were immature, and 24% had not mated. In old rotting stems where eggs are laid and larvae develop, a mixture of 52% mated and 48% virgin females was found. Therefore, the pheromone together with the odour of rotting coconut stems signals a reproduction site to beetles, particularly mature females. In practice, the pheromone-baited traps will help in reducing the dissemination of gravid females, but will not affect directly the numbers of immature ones attacking palms. Our results show that when using pheromones for monitoring or controlling insects, the physiological status of the insects may have unexpected effects on the outcome.     

Sexual behavior in brown capuchins (Cebus apella)

We observed sexual behavior patterns in two captive groups of Cebus apella.We obtained data on intersexual solicitations and intersexual competition. The length of proceptivity cycles by females averaged 18.8 ± 1.2 days; this corresponds with published reports of physiological measures of ovulatory cycles. Females initiated sexual interactions with males significantly more than males initiated interactions with females, and female solicitations increased in frequency over the duration of displayed periods of proceptivity. In contrast, male solicitations of females were infrequent and did not change significantly throughout female proceptive phases. Each female solicited the same male throughout a given proceptive phase. We observed no evidence of male-male or female-female competition. Males seem to respond to female sexual behavior directed toward them, rather than initiating sexual interactions themselves. Even so, male sexual interactions with females followed only a small proportion of solicitations.     

Sexual behavior in brown capuchins ( Cebus apella )

We observed sexual behavior patterns in two captive groups of Cebus apella.We obtained data on intersexual solicitations and intersexual competition. The length of proceptivity cycles by females averaged 18.8 ± 1.2 days; this corresponds with published reports of physiological measures of ovulatory cycles. Females initiated sexual interactions with males significantly more than males initiated interactions with females, and female solicitations increased in frequency over the duration of displayed periods of proceptivity. In contrast, male solicitations of females were infrequent and did not change significantly throughout female proceptive phases. Each female solicited the same male throughout a given proceptive phase. We observed no evidence of male-male or female-female competition. Males seem to respond to female sexual behavior directed toward them, rather than initiating sexual interactions themselves. Even so, male sexual interactions with females followed only a small proportion of solicitations.     

Ips pini (Curculionidae: Scolytinae) is a vector of the fungal pathogen, Sphaeropsis sapinea (Coelomycetes), to Austrian pines, Pinus nigra (Pinaceae)

Sphaeropsis sapinea (Fr.:Fr.) Dyko and Sutton, is among the most common and widely distributed pathogens of conifers worldwide. S. sapinea is disseminated over short distances by rain splash and moist wind, but significant knowledge gaps regarding long-range dispersal remain. Our objective was to determine whether or not the pine engraver beetle, Ips pini Say, is a vector of the pathogen onto Austrian pines (Pinus nigra Arnold). In 2004 and 2005, individuals of I. pini were collected with pheromone traps at two locations in central Ohio (197 and 1,017 individuals for 2004 and 2005, respectively) and screened for the presence of S. sapinea. In the field, fresh logs of Austrian pine were baited with pheromone lures, mechanically wounded, or left undisturbed. After 2 mo, logs were evaluated for insect feeding and the presence of S. sapinea along beetle galleries. Fresh logs were also inoculated in the greenhouse with adult I. pini that were either artificially infested or uninfested with S. sapinea spores to determine vectoring potential. Phoresy rates for individual collections ranged from 0 to 4.1%; average rates were 1.5 and 2.0% for 2004 and 2005, respectively. Isolation frequencies of S. sapinea from baited (15 +/- 5%) and unbaited logs (3 +/- 1%) differed significantly (P=0.009). I. pini was also capable of transmitting the pathogen under controlled conditions. Based on phoresy rates, association, and artificial inoculation studies, we conclude that I. pini is able to transmit S. sapinea to Austrian pine stems.     

Flight movement of Scapanes australis australis (Boisduval) (Coleoptera: Scarabaeidae: Dynastinae) in Papua New Guinea: a radiotelemetry study

Abstract We used radiotelemetry and/or chemical light-tags to track the flight of 15 individuals of Scapanes australis in Madang Province, Papua New Guinea. This species causes severe economic impacts on coconut palms in young plantations. Flights to feeding, mating, resting, and possibly oviposition sites covered distances of 52 to 835 m in males, and from 245 m to>1000 m in females. Upon release, females flew in a tight upward spiral above canopy level (>20 m), then usually flew along a single bearing out of radio reception within 1 min of initiating flight. Dispersing females probably follow scent trails to pheromone-releasing males that occupy feeding galleries excavated most frequently in coconut palms, or search for oviposition sites. Most tagged females were not found again, because they dispersed beyond the tracking capabilities of our radio-receivers, but one female was followed for 245 m to a feeding gallery excavated by an adult male. Males typically flew within 5 m of the ground, took erratic flight paths with numerous turns, and frequently circled coconuts and other host plants. We followed males from the release point until they ceased flight for a night. Males passed daylight hours either in a feeding gallery within a host plant or under soil litter.     

Identifying a causal agent of sexual selection on weaponry in an insect

In many animal species, males do not seek females directly butinstead locate and defend sites that contain spatially or temporallylimited resources essential to female survival and reproduction.Resident males that successfully repel conspecific rivals canmate with females attracted to these resources. In theory, increasingresource value increases harem size and thus increases the opportunity(I_mates) for and strength of sexual selection on traits crucialto male resource-holding potential and mating success. I experimentallytested this hypothesis in the field using the Wellington treeweta, Hemideina crassidens (Orthoptera: Tettigonioidea: Anostostomatidae),a sexually dimorphic insect in which males use their enlargedmandibles as weapons in male–male contests over accessto females sheltering in tree cavities (galleries). By manipulatinggallery size, I showed that, compared with smaller galleries,larger galleries housed larger harems. Variation in gallerysize was an important determinant of I_mates, but contrary toexpectation, greater opportunity existed in small galleriescompared with large galleries. As predicted, male weapon sizewas under stronger directional selection in large galleriesbecause the fitness benefits were greater under these conditionscompared with small galleries. My results help explain the positiveassociation between average weapon size and average gallerysize observed within and among tree weta populations in NewZealand.     

Selection in the absence of males does not affect male-female conflict in the parasitoid wasp Leptopilina clavipes (Hymenoptera: Figitidae)

Divergent reproductive interests of males and females can lead to sexually antagonistic coevolution (SAC). In the absence of males, adaptations evolved under SAC are released from selection and expected to deteriorate. In this study, we investigated this prediction using two populations of the parasitoid wasp Leptopilina clavipes, one arrhenotokous and one thelytokous. Thelytokous females were induced to produce sons by curing them of their Wolbachia-infection. We examined whether thelytokous males were less able to inhibit female remating than arrhenotokous males and whether thelytokous females were more susceptible to male-induced longevity reduction than arrhenotokous females. The results showed that females were monandrous, regardless of whether mated with an arrhenotokous or thelytokous male. While ongoing courtship of males reduced female life span, there was no longevity cost of mating for either arrhenotokous or thelytokous females. Our results therefore do not support the idea that adaptations evolved under SAC deteriorate under prolonged female-only selection.