Female mimicry and resource defense polygyny by males of a tropical rove beetle,Leistotrophus versicolor (Coleoptera : Staphylinidae)

Summary Adults of the staphylinid beetle Leistotrophus versicolor Grav. aggregate at vertebrate dung and carrion where males and females forage for adult Diptera. Some males aggressively exclude others from dung and carrion. Winners in male combat gain access to many females, which are often receptive at these foraging sites. The mating system can be categorized as resource defense polygyny. Males vary greatly in size, are larger than females on average, and have allometrically enlarged mandibles that they use in fighting. Large males consistently defeat smaller ones. Some males employ female mimicry in order to avoid aggression, remaining at dung where they forage and even obtain copulations while being courted by rival males. Female mimicry is most often practiced by males that are smaller than their rivals or by males that are unable to use their jaws aggressively because they are feeding or courting females when encountered by an opponent. Female mimicry is a conditional tactic of mature males; some individuals behave like females toward larger males but attack smaller rivals. Offprint requests to: J. Alcock     

Isopropyl (Z9)-hexadecenoate as a male attractant pheromone from the sternal gland of the rove beetle Aleochara curtala (Coleoptera: Staphylinidae)

Summary. In the rove beetle Aleochara curtula, a male specific sternal gland is described. Isopropyl (Z9)-hexadecenoate has been identified by GC/MS of surface and sternal gland extracts as a male specific compound. Its emission in the air was demonstrated by closed-loopstripping-analyses. In field experiments, conspecific males and females were attracted by the odor of carrion, caged males from laboratory cultures, and by the synthetic ester. Isopropyl (Z9)-hexadecenoate or live males, combined with fresh carrion attract more beetles than a fresh carcass alone. Isopropyl hexadecanoate as a minor compound was not attractive. The term “aggregation” pheromone and the ecological significance of attracting females to a fresh carcass allowing early copulation and egg-deposition are discussed. Received 20 August 1998; accepted 15 January 1999.     

Chemical camouflage by myrmecophilous beetles Zyras comes (Coleoptera: Staphylinidae) and Diaritiger fossulatus (Coleoptera: Pselaphidae) to be integrated into the nest of Lasius fuliginosus (Hymenoptera: Formicidae)

Summary. The myrmecophilous beetles, Zyras comes (Staphylinidae) and Diaritiger fossulatus (Pselaphidae) are guests of the black shining ant Lasius fuliginosus. Host worker ants never attacked these beetles, and often gave regurgitant to Z. comes following tactile communication with the beetle. By contrast, the workers from colonies without the myrmecophiles showed hostile responses towards Z. comes before tactile contact, but were not aware of D. fossulatus until contact. In L. fuliginosus, workers within a colony shared profiles, but the profiles differed among colonies. GC analyses showed that both Z. comes and D. fossulatus beetles had the same hydrocarbons as L. fuliginosus, and the profiles were more similar to those of the host colony workers than the foreign workers. Both Z. comes and D. fossulatus appear to imitate the hydrocarbon profile of their host workers, allowing integrating into the host nest. A Y-maze bioassay indicated that Z. comes can follow the trail pheromone of L. fuliginosus. This suggests that Z. comes may detect other chemical signals of L. fuliginosus to keep closer interactions with the workers. Received 22 June 2001; accepted 12 November 2001.     

The cause of size-assortative mating in the leaf beetle Trirhabda canadensis (Coleoptera : Chrysomelidae)

Summary Examples of positive assortative mating by body size are abundant but its causes remain controversial. I show that size-assortative mating occurs in the chrysomelid beetle Trirhabda canadensis and I test a series of alternative hypotheses to explain how this mating pattern comes about. Results suggest that assortative mating in this beetle is due to the greater ease with which size-matched pairs can achieve intromission, and not due to size-biased skews in the availability of mates or mate choice favoring large individuals. There was no correlation between male and female elytron length (a measure of body size) at the initiation of courtship, but pairs assorted positively by size at the onset of intromission. Moreover, in the laboratory, there was a negative correlation between male and female size for pairs engaged in courtship that terminated without mating. Assortative mating was not associated with a large-male mating advantage and there was no evidence of female choice of large males. Nor was there unequivocal evidence for male choice of large females; although mating females were slightly larger and considerably heavier than solitary females, males did not differ in the frequency with which they rejected large and small females. Assortative mating in T. canadensis appeared to be caused by the lower ability of mismatched pairs to achieve intromission after an encounter, both when males were larger and when they were smaller than the female.     

Aggregation pheromone compounds of the black larder beetle Dermestes haemorrhoidalis Kuster (Coleoptera: Dermestidae)

Gas chromatography with simultaneous flame ionisation and electroantennographic detection (GC–EAD) and gas chromatography with mass spectrometry analysis (GC–MS) of abdominal extracts of adult male Dermestes haemorrhoidalis Kuster (Coleoptera: Dermestidae) revealed the presence of electrophysiologically and behaviourally active compounds to its conspecific males and females. Isopropyl dodecanoate (3), isopropyl (Z)-9-tetradecenoate (5), isopropyl tetradecanoate (6), isopropyl (Z)-9-hexadecenoate (7) and isopropyl hexadecanoate (8) were detected in male abdominal extracts only. Analysis of collected male headspace volatiles revealed the presence of six EAD-active compounds (3), (5), (6) and isopropyl tridecanoate (4) plus two unidentified compounds (1) and (9). Synthetic compounds (3), (4), (5), (6) and (7) showed EAD activity with antennae of both sexes in contrast to synthetic (8) which showed EAD activity with female antennae only. Male and female antennae of D. haemorrhoidalis reacted with high receptor potentials to isopropyl (Z)-9-dodecenoate (2), although this compound itself was detected in neither male nor female abdominal extracts or headspace volatiles. Petri dish bioassays indicated that male abdominal extracts and compounds (2), (3), (5) and (6) aroused and attracted conspecific male and female beetles significantly (P < 0.05) compared to female extracts. These results suggested the presence of a male-produced aggregation pheromone in D. haemorrhoidalis. Field assays with any of the described compounds, however, did not result in attraction of this beetle in significant numbers.     

Metabolism of neolignans from P. regnellii (Piperaceae) in the beetle Naupactus bipes (Coleoptera: Curculionidae)

Piper regnellii (Piperaceae) contains high levels neolignans with diverse biological activities, including insecticidal. Despite the insecticidal activity of these neolignans, Naupactus bipes (Coleoptera: Curculionidae) larvae and adults were found feeding on the leaves (adults) and roots (larvae) of P. regnellii. The present study investigated the metabolic pathway of neolignans from P. regnellii leaves in the beetle N. bipes, focusing on possible biotransformation or sequestration of these compounds by the beetle. Two of the four plant neolignans could be recovered in the feces of adults feeding on P. regnellii leaves. In addition, four degradation products were detected, including the neolignan 3-[(2R,3R)-2,3-dihydro-3-(methyl)-2-(4-hydroxyphenyl)-5-benzofuranyl]-(2E)-propenal (7), an oxidation product of conocarpan, the major plant neolignan. The adult beetle sequesters the neolignan conocarpan selectively from leaves as well as accumulates neolignan 7. Neolignan 7 was also detected in the larvae of lab-reared beetles.     

Rove beetles (Coleoptera: Staphylinidae) collected during the long term ecological research in a Hungarian oak forest

Along term ecological research was carried out in a Hungarian oak forest, in "Bükk" National Park starting with 1972. During the faunistical studies 3,602 insect species and more than 200,000 individuals were collected. The dominant orders were Coleoptera (1,051 species), Lepidoptera (803 species), Hymenoptera (470 species) and Diptera (400 species). The relative species abundance (RSA) for all insects collected in all years of sampling period suggests a rather J shape curve than a not clear scaling property. This means that we were able to identify almost three quarters of the insect species from one ha European oak forest during the survey (from 1987 to 2003), and two third of the staphylinides expected. Considering the staphylinid fauna a total number of 160 species and 4,022 individuals were collected. The most widely occurring species in dominance order were: Ocypus biharicus, Pseudocypus mus, Atheta gagatina, Philonthus quisquiliarius, Oxypoda acuminate, Platydracus chalcocephalus, Atheta crassicomis, Latrimaeum atrocephalum, Haploglossa puncticollis, Philonthus succicola and Anotylus mutator. The pooled value of alpha diversity was 1.51. The Shannon-Weiner Index (H') was relatively high (3.29) in comparison with other studies.     

Alternative reproductive tactics and male-dimorphism in the horned beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae)

Adult dung beetles (Onthophagus acuminatus) exhibit continuous variation in body size resulting from differential nutritional conditions experienced during larval development. Males of this species have a pair of horns that protrude from the base of the head, and the lengths of these horns are bimodally distributed in natural populations. Males growing larger than a threshold body size develop long horns, and males that do not achieve this size grow only rudimentary horns or no horns at all. Previous studies of other horned beetle species have shown that horned and hornless males often have different types of reproductive behavior. Here I describe the mating behaviors of the two male morphs of O. acuminatus during encounters with females. Females excavate tunnels beneath dung, where they feed, mate and provision eggs. Large, horned males were found to guard entrances to tunnels containing females. These males fought with all other males that attempted to enter these tunnels. In contrast, small, hornless males encountered females by sneaking into tunnels guarded by other males. In many instances, this was accomplished by digging new tunnels that intercepted the guarded tunnels below ground. Side-tunneling behavior allowed sneaking males to enter tunnels beneath the guarding male, and mate with females undetected. Both overall body size and relative horn length significantly affected the outcome of fights over tunnel ownership. These results suggest that alternative reproductive tactics may favor divergence in male horn morphology, with long horns favored in males large enough to guard tunnels, and hornlessness favored in smaller males that adopt the “sneaking” behavioral alternative. Received: 12 October 1996 / Accepted after revision: 8 August 1997     

Quantitative composition of the defensive secretion ofBledius species (Coleoptera: Staphylinidae: Oxytelinae) is adapted to naturally occurring predators

Summary The adaptation of defensive secretions to their target organisms was examined for the abdominal gland secretions ofBledius furcatus, B. spectabilis andB. arenarius. Therefore the target organisms of the secretion of theseBledius species (i.e. their predators) had to be identified. At the collection sites examined these were the earwigLabidura riparia, the antCataglyphis bicolor, the flyLispe candicans, different carabids of the generaPogonus, Dichirotrichus, Dyschirius, Bembidion andCalathus and the wading birdsHaematopus ostralegus andCalidris alba. The secretion of the abdominal glands contains the toxin ptoluquinone dissolved in either-dodecalactone and 1-undecene (B. furcatus andB. spectabilis) or in octanoic acid and octyloctanoate (B. arenarius). The ratio of these solvents is species-specific. Application experiments using some of the natural insect predators (L. riparia, C. bicolor, Pogonus, Di. gustavii, Dyschirius) revealed that these solvent ratios provided a more effective deterrent than other possible ratios. Thus by combining the solvents in certain ratios, the capability of cuticular penetration and therefore the effectiveness of the defensive secretions are adapted to their natural targets.     

Offensive and defensive sperm competition roles in the dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae)

Sperm competition is predicted to generate opposing selection pressures on males. On one hand, selection should favour ‘defensive’ adaptations that protect a male’s ejaculate from displacement, while, on the other hand, selection should favour ‘offensive’ adaptations that overcome paternity assurance mechanisms of rivals. Here, we use the sterile male technique to assess sperm precedence when a male dung beetle Onthophagus taurus mates in both a defensive (first male) and an offensive (second male) role. Significant variation in a male’s sperm precedence (both P ₁ and P ₂) was detected, and an individual’s defensive (P ₁) and offensive (P ₂) abilities were positively correlated. Thus, it appears that sexual selection simultaneously selects for ‘defensive’ and ‘offensive’ adaptations in O. taurus. We discuss a variety of male traits in O. taurus that potentially contribute to a male’s ability to be successful when mating in an ‘offensive’ and a ‘defensive’ role.     

Volatile emissions from the flea beetle Altica litigata (Coleoptera: Chrysomelidae) associated with invasive Ludwigia hexapetala

The flea beetle Altica litigata (Chrysomelidae) is an insect herbivore to plants within the families Lythraceae and Onagraceae, including ornamentals such as crape myrtle, Lagerstroemia spp. This insect is important both as a pest species and as a naturally occurring biological control agent due to its aggregate feeding behavior, which typically results in severe defoliation of the host plant. Despite the negative economic impact to ornamentals and contrary benefits as a biological control agent, there are few reports on the semiochemical communication of this family of insects. Uruguayan primrose-willow (Ludwigia hexapetala) is an invasive aquatic weed in California and serves as a host to A. litigata. To better characterize this association, the volatile emissions of A. litigata were collected while the flea beetles were: in containers by themselves, in containers with L. hexapetala leaves, in situ on L. hexapetala leaves in a growth chamber, and in situ on L. hexapetala leaves in the field. For comparison, the volatile emissions of A. litigata associated with two subspecies of creeping water primrose (L. peploides) were also evaluated. Two himachalene-type sesquiterpenes, showing the same carbon skeleton as compounds previously reported from Aphthona flava and Epitrix fuscula, were detected as volatiles from A. litigata.     

Patterns of parental provisioning covary with male morphology in a horned beetle (Onthophagus taurus) (Coleoptera: Scarabaeidae)

Male dung beetles, Onthophagus taurus, are dimorphic for a secondary sexual trait, head horns. Horned males participate in the production of brood masses while hornless male do not. Here we examine the reproductive performance of females mated with males exhibiting alternative horn morphologies. We found that exposure to males may be costly for females in that it reduced the total number of brood masses produced. However, females paired with horned males produced significantly larger brood masses than females paired with hornless males or females producing broods alone. We discuss the possible selection pressures that may underly horn evolution in this genus. Received: 22 August 1997 / Accepted after revision: 19 January 1998     

Proximal traits and mechanisms for biasing paternity in the red flour beetle Tribolium castaneum (Coleoptera: Tenebrionidae)

A comprehensive understanding of sexual selection requires knowledge of the traits and mechanisms responsible for increasing a male’s paternity share (proportion of progeny sired) relative to that of other males mating with the same female. In this study we manipulated by starvation the expression of traits that might influence male paternity share in Tribolium castaneum. We then conducted experiments to examine how male starvation affects male performance during sequential episodes of sexual selection from mating to progeny production, and investigated female control over specific stages by using live vs dead females. Comparison of starved vs fed males revealed that T. castaneum females have control over spermatophore transfer during mating, as live females rejected inseminations by starved (“low quality”) males. None of the measured male copulatory behaviors (leg-rubbing frequency, asymmetry, and percent of time spent rubbing) affected the probability of successful insemination, but the last two were positively associated with male paternity share. Spermatophore positioning within the female reproductive tract was not affected by male treatment (starved/fed), by female treatment (live/dead), or by male copulatory behaviors. Starvation, however, had a dramatic effect on male reproductive physiology, decreasing both accessory gland size and total number of sperms transferred (but not sperm viability in seminal vesicles). In addition, females who mated to starved males stored fewer sperms in their spermathecae, which, together with decreased ejaculate size, may explain the reduced paternity share of starved males compared to fed males. This study elucidates some cryptic mechanisms influencing male reproductive success and aids our understanding of trait evolution through sexual selection.     

Fifteen-spined stickleback (Spinachia spinachia) females prefer males with more secretional threads in their nests: an honest-condition display by males

In the fifteen-spined stickleback (Spinachia spinachia), the male alone builds a nest and provides care for the eggs until they hatch. The nest is made of filamentous algae held together with shiny secretional threads of a glycoprotein, here called tangspiggin. Tangspiggin is produced by transformed kidney cells. I investigated how tangspiggin production was related to food intake by giving males low or high food rations. Males in the high-food group produced significantly more tangspiggin than those in the low-food group. Since low food rations did not lead to a significant drop in resting metabolic rate (oxygen consumption), tangspiggin production appears to be one of the first energy-demanding processes to be dropped during food deprivation. Among males given equal food rations, females preferred to mate with those with elevated levels of tangspiggin in their nests. Tangspiggin threads may signal safety for the offspring by keeping the eggs within the nest and protecting them from egg predators. Moreover, the threads become less shiny with age, due to particle deposition, and older nests that may house older (more valuable) eggs are thereby well camouflaged. Tangspiggin provides an honest signal of male condition, acting as an extra-bodily ornament, signaling direct benefits to females or possibly also indirect benefits of "good genes" from a male who is a successful forager.     

Pheromone of the elm bark beetle Scolytus laevis (Coleoptera: Scolytidae): stereoisomers of 4-methyl-3-heptanol reduce interspecific competition

Stereoisomers of 4-methyl-3-heptanol (MH) are pheromone components of several Scolytus bark beetles. The elm bark beetle Scolytus laevis (Coleoptera: Scolytidae) has in previous studies been caught in traps baited with commercial MH containing all four stereoisomers, but the lure has been considered a weak attractant. In this study, we addressed the question whether stereospecific responses by S. laevis to stereoisomers of MH might contribute to its niche separation from other sympatric Scolytus species. Using GC–MS, we analyzed extracts of hindguts and abdomens from male and female S. laevis and the sympatric S. triarmatus. We also tested all four MH-stereoisomers individually and in combinations in the field to determine their role for S. laevis. All four stereoisomers were synthesized via a boronic ester method with 1,2-dicyclohexylethanediol as chiral director. In addition, the (3S,4R)-stereoisomer of MH was prepared through enantioselective, lipase-mediated transesterification of a mixture of the four stereoisomers of MH. Females of both species contained small amounts of syn-MH, and males contained trace amounts of anti-MH. The anti stereoisomer (3R,4S)-MH was attractive to male and female S. laevis, whereas the syn stereoisomer (3S,4S)-MH acted as an inhibitor or deterrent and reduced the catch when added to the attractive isomer. The syn isomer is the main aggregation pheromone component of the larger and sympatric S. scolytus and possibly also of S. triarmatus. The avoidance response of S. laevis to the (3S,4S)-stereoisomer may reduce interspecific competition for host trees.     

Prolonged mating in the milkweed leaf beetle Labidomera clivicollis clivicollis (Coleoptera: Chrysomelidae): a test of the “sperm-loading” hypothesis

Summary Female milkweed leaf beetles (Labidomera clivicollis clivicollis) frequently mate with more than one male, and pairs form mating associations which last for up to 42 h in the field. I tested the hypothesis that males remaining with females for long periods of time benefit by numerically overwhelming the sperm of their competitors. Male L.c.clivicollis copulated intermittently with females throughout an 11 hour period in the laboratory. When virgin females were allowed a single copulation, 94.3% of the sperm they received were located in the spermatheca immediately afterward. Males were not sperm-depleted, for they had large numbers of sperm available after one copulation (mean=230,000±43,200); the maximal number of sperm a male transferred to a female in 24 h was 30,500. There was a positive linear relationship between the number of sperm transferred and time up to 24 h after mounting (r ²=0.178, P<0.003). These data suggest that males transfer increasing numbers of sperm throughout a 24-h-period. Mating duration was the most important determinant of paternity when females were placed with one male for 24 h and another male for 6 hours. Females whose first matings were longer showed first male sperm predominance (as determined by starch-gel electrophoresis), while females whose second matings were longer showed last male sperm predominance. In view of these data, it is puzzling that males do not inseminate with large numbers of sperm immediately after mounting the female. It is possible that female refractory behaviors make insemination difficult and favor prolonged mating by male milkweed leaf beetles.     

Following a scented beetle: larval faeces as a key olfactory cue in host location of Stegobium paniceum (Coleoptera: Anobiidae) by Lariophagus distinguendus (Hymenoptera: Pteromalidae)

Lariophagus distinguendus (Hymenoptera: Pteromalidae) is a generalist solitary ectoparasitoid parasitising immature stages of at least 11 foodstuff beetles from five families, including species developing in seeds of Poaceae and Fabaceae. In this study, we tested the role of visual and olfactory stimuli affecting L. distinguendus host location in the trophic context of Stegobium paniceum (Coleoptera: Anobiidae) infesting chickpeas, Cicer arietinum (Fabales: Fabaceae). When either visual plus olfactory cues or olfactory cues alone were provided, S. paniceum-infested chickpeas were more attractive than uninfested chickpeas to naïve L. distinguendus females. Larval faeces of S. paniceum also evoked strong attraction to female wasps. Uninfested chickpeas covered with host faeces were preferred over uninfested ones without faeces, while infested chickpeas were as attractive as healthy chickpeas plus host faeces. Overall, results demonstrated that L. distinguendus females orient their host-seeking decisions mainly on the basis of faeces-borne olfactory stimuli. Further research is on-going to characterise the volatiles from S. paniceum larval faeces to evaluate their attractiveness towards L. distinguendus females.     

Volatiles from the bark of trembling aspen, Populus tremuloides Michx. (Salicaceae) disrupt secondary attraction by the mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae)

Summary. Coupled gas chromatographic-electroantennographic detection (GC-EAD) analysis of the Porapak Q-captured volatiles from the bark of trembling aspen, Populus tremuloides Michx., revealed four compounds that consistently elicited antennal responses by mountain pine beetles (MPBs), Dendroctonus ponderosae Hopkins. One of these, 1-hexanol, disrupted the capture of MPBs in multiple-funnel traps baited with the aggregation pheromones trans-verbenol and exo-brevicomin and the host kairomone myrcene, a blend of semiochemicals that mediates the secondary attraction response in which beetles mass attack and kill living pines. The other three EAD-active aspen bark volatiles, benzyl alcohol, benzaldehyde and nonanal, were inactive alone, but in binary and ternary combinations contributed to a disruptive effect in an additive and redundant manner when all four aspen bark volatiles were tested in all possible binary and ternary blends. The best ternary blend and the quarternary blend achieved ≥ 80% disruption. The quarternary blend enhanced the disruptive effect of the antiaggregation pheromone verbenone in traps, raising the disruptive effect to 98%, and also enhanced the inhibition of attack on attractant-baited lodgepole pines. This is the first demonstration of specific compounds from the bark of angiosperm trees that disrupt the secondary attraction response of sympatric coniferophagous bark beetles. The results support the hypothesis that such bark beetles are adapted to recognize and avoid non-host angiosperm trees by responding to a broad spectrum of volatiles that can act in various blends with equal effect. Received 27 October 1997; accepted 20 February 1998.     

Solar and landscape cues as orientation mechanisms in the beach-dwelling beetle Eurynebria complanata (Coleoptera,Carabidae)

The present paper reports on the first orientation experiments conducted on the strand-living beetle Eurynebria complanata (Linnaeus, 1767) (Coleoptera, Carabidae). The experiments were carried out from June to October 1988. Two different populations were used: one inhabiting a Tyrrhenian beach in Italy and the other a beach on the Atlantic coast of France. Response to sun compass cues was demonstrated in each population at the collection site and for the French population after transportation to Italy, where experiments were carried out on a differently oriented beach. Landscape cues were shown to improve the beetle's orientation capacity when these were tested together with a visible sun. Artificial landscapes were also tested under laboratory conditions. Different-sized silhouettes were placed in the four cardinal directions, and these envoked different responses depending on their height. The beetles oriented towards a black silhouette when this subtended an angle of 25°. The results obtained for the two populations are compared and discussed from an eco-ethological point of view.