Dynamics of pheromone production and communication in the mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae)

Summary. The mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say), often co-exist in lodgepole pine, Pinus contorta var. latifolia Engelmann. Intra- and interspecific semiochemical communication occurs in both species and their complete semiochemical repertoire and precise dynamics of pheromone production have not been elucidated. Porapak-Q extracts of captured volatiles from beetles of each species aerated at different attack phases (freshly emerged, pioneer sex alone in the log and both sexes paired in new galleries), followed by gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectroscopic analyses identified 17 compounds (seven compounds common to both species, six present in D. ponderosae and four present in I. pini) that excited the antennae of either or both species. Seven compounds for D. ponderosae and nine for I. pini had not been assessed for behavioural activity. In field trapping experiments, 2-phenylethanol produced by both species inhibited the response of D. ponderosae to its aggregation pheromones. exo- and endo-Brevicomin produced by D. ponderosae significantly decreased the response of I. pini to its aggregation pheromone ipsdienol. Nonanal, a ubiquitous compound found in the volatiles of lodgepole pine, various nonhosts and in both beetle species deterred the response of I. pini to ipsdienol. The occurrence of cis-verbenol, trans-verbenol and verbenone in emergent I. pini, and verbenone and 2-phenylethanol in emergent D. ponderosae suggests that these compounds may inhibit aggregation and induce dispersal following emergence. Termination of aggregation in D. ponderosae appears to depend on the production of frontalin in combination with changes in the relative ratios of verbenone, exo-brevicomin, trans-verbenol and 2-phenylethanol. In I. pini, the cessation of ipsdienol production by males is probably the main factor in terminating aggregation. Received 16 November 1999; accepted 7 August 2000     

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.     

Trace metals in mussels (Mytilus edulis) from southwest Iceland

The temperate population of the tropical anemonefish Amphiprion clarkii has a seasonality of reproduction, larval settlement and growth. There was a considerable difference in size (15–47 mm) among 0-year olds in December, after the season of larval settlement and growth. This difference could be attributed mainly to a difference in the duration of the first growing season spent after settlement and a social inhibition of growth of the subordinate late settlers by frequent attacks by the dominant early settlers. The growth equation which was used to describe the growth of tagged individuals estimated that an anemonefish would get an opportunity to spawn within four of five years after settlement. Breeding adult pairs occupied almost all sea anemones essential to breeding within a 50×50 m study area, and all 17 new adults that matured from non-breeding juveniles to breeding adults between June 1983 and August 1985 were found in the territories from which either or both of the mated adults had disappeared. The new adults were large juveniles who had resided near those vacant territories before. These results suggest that the onset of breeding by an individual is not only determined by his age, but also by his ranking in the dominance hierarchy. Therefore, the difference in size among 0-year olds in December might give rise to the difference in age of the earliest breeding and might be a factor in making a female apply more reproductive effort at the beginning of the season.     

Spatial variability of biotic and abiotic tree establishment constraints across a treeline ecotone in the Alaska range

Throughout interior Alaska (U.S.A.), a gradual warming trend in mean monthly temperatures occurred over the last few decades (approximatlely 2-4 degrees C). The accompanying increases in woody vegetation at many alpine treeline (hereafter treeline) locations provided an opportunity to examine how biotic and abiotic local site conditions interact to control tree establishment patterns during warming. We devised a landscape ecological approach to investigate these relationships at an undisturbed treeline in the Alaska Range. We identified treeline changes between 1953 (aerial photography) and 2005 (satellite imagery) in a geographic information system (GIS) and linked them with corresponding local site conditions derived from digital terrain data, ancillary climate data, and distance to 1953 trees. Logistic regressions enabled us to rank the importance of local site conditions in controlling tree establishment. We discovered a spatial transition in the importance of tree establishment controls. The biotic variable (proximity to 1953 trees) was the most important tree establishment predictor below the upper tree limit, providing evidence of response lags with the abiotic setting and suggesting that tree establishment is rarely in equilibrium with the physical environment or responding directly to warming. Elevation and winter sun exposure were important predictors of tree establishment at the upper tree limit, but proximity to trees persisted as an important tertiary predictor, indicating that tree establishment may achieve equilibrium with the physical environment. However, even here, influences from the biotic variable may obscure unequivocal correlations with the abiotic setting (including temperature). Future treeline expansion will likely be patchy and challenging to predict without considering the spatial variability of influences from biotic and abiotic local site conditions.     

Spatial distribution of perfluorooctane sulfonate (PFOS) in major rivers in southwest Nigeria

Concentrations of perfluorooctane sulfonate (PFOS) were measured for the first time in major rivers within southwest, Nigeria. PFOS was concentrated from water and sediment using solid-phase extraction, identified and quantified with high-performance liquid chromatography coupled with tandem mass spectrometry. Concentrations of PFOS ranged from 1.71 to 16.19 ng L^?1 in water, and from 1.64 to 10.29 ng g^?1 in sediments across all locations. Comparatively, the concentrations of PFOS observed in this study were within the range ever measured in the environment. Field-based sediment water distribution coefficients (log K_d, L kg^?1) ranged from 2.08 to 3.56. While no correlation was observed between K_d and organic carbon contents, there was significant positive correlation between K_d and salinity (r² = 0.7867), which suggested that activities capable of increasing salinity can enhance PFOS removal from the environment.     

Demonstration of sex pheromones in the predaceous diving beetle Rhantus suturalis (MacLeay 1825) (Dytiscidae)

Sex pheromones are used by a wide variety of species in terrestrial ecosystems. Much less is known about these pheromones in aquatic systems e.g., for diving beetles. To test the use of pheromones and visual signals for mate finding by the diving beetle Rhantus suturalis (MacLeay 1825), behavioral experiments were performed using three different types of vessels containing conspecific. In experiments with non-permeable glass flasks, which did not allow the diffusion of chemical substances, males and females did not stimulate any reaction by conspecific of either sex. In experiments with opaque but water permeable vessels, and in experiments with vessels made of finely woven steel, which allowed the perception of chemical as well as optical cues, male diving beetles were significantly attracted to females. Females were attracted to other females only in the last experiment, when simultaneously perceiving chemical and optical cues. These results indicate that R. suturalis females release sex pheromones to attract mating partners, which is the first experimental demonstration of pheromones in the Dytiscidae.     

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     

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.     

Demography and interannual variability of salp swarms (Thalia democratica)

Swarms of the pelagic tunicate, Thalia democratica, form during spring, but the causes of the large interannual variability in the magnitude of salp swarms are unclear. Changes in asexual reproduction (buds per chain) of T. democratica populations in the coastal waters of south-east Australia (32–35°S) were observed in three austral springs (October 2008–2010). T. democratica abundance was significantly higher in 2008 (1,312 individuals m^?3) than 2009 and 2010 (210 and 92 individuals m^?3, respectively). There was a significant negative relationship (linear regression, r ² = 0.61, F _1,22 = 33.83, P < 0.001) between abundance and asexual reproduction. Similarly, relative growth rates declined with decreasing abundance. Generalised additive mixed modelling showed that T. democratica abundance was significantly positively related to preferred food >2 μm in size (P < 0.05) and negatively related to the proportion of non-salp zooplankton (P < 0.001). Salp swarm magnitude, growth, and asexual reproduction may depend on the abundance of larger phytoplankton (prymnesiophytes and diatoms) and competition with other zooplankton.     

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.     

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     

Diving characteristics of southern rockhopper penguins (Eudyptes c. chrysocome) in the southwest Atlantic

The diving behaviour of southern rockhopper penguins (Eudyptes c. chrysocome) was studied at two breeding sites in the Southwest Atlantic: the Falkland Islands and Staten Island, Argentina. Incubating and brooding birds were equipped with time-depth recorders to monitor their foraging activities. Rockhopper penguins from Staten Island started their breeding season about 3 weeks earlier than their conspecifics from the Falkland Islands. The foraging area used by incubating males from the Falkland Islands comprised about 150,000 km² to the northeast of the breeding site and was characterised by shelf and slope waters, whereas the foraging area of incubating males from Staten Island comprised 350,000 km² of oceanic waters to the southeast of the breeding site. A number of dive parameters were measured and compared between the four study groups: Incubating males and brooding females from the Falkland Islands, and incubating males and females from Staten Island. In all study groups, dive depth correlated positively to light intensity, dive duration and vertical velocity. However, significant differences between various diving parameters of the study groups were noted, not only in terms of diving performance, but also as regards diving efficiency (DE). A principal component analysis (PCA) on 16 variables revealed that 75% of the variance could be explained by only two principal components: diving pattern (PC1) and diving effort (PC2). PC1 indicated that the birds from Staten Island, both males and females, dived deeper, covered a greater vertical distance per hour and had higher ascent rates, but spent less time underwater and at the bottom of a dive, and had a lower DE than conspecifics from the Falkland Islands. PC2, which included the percentage of foraging dives, the number of dives per hour, dive duration, bottom time and descent rate, differed significantly between incubating males from the Falkland Islands and the other three groups, which were all very similar. Overall, the diving behaviour was notably similar to that of conspecifics from the Indian and Pacific Oceans. The implications of the results in terms of intra-specific adaptations as well as potential threats from human activities are discussed.     

Mitochondrial DNA sequence variation in deep-sea bamboo coral (Keratoisidinae) species in the southwest and northwest Pacific Ocean

The Keratoisidinae are a poorly known, but phenotypically diverse group of deepwater corals. Recent developments in deepwater trawling in the southwest Pacific have provided many more specimens of bamboo corals. Two sub-regions of the mitochondrial genome were sequenced to test genetic relationships among specimens collected over a wide geographical range (27–50° S): a sub-region of the large-subunit rRNA (16S rRNA), characterized by a highly variable insertion/deletion (INDEL#2) region; and a non-coding region between COII and COI. Based on DNA haplotypes, 14 species of Keratoisidinae were recognized among 88 specimens from deep water in the southwest Pacific Ocean. The common haplotypes also appeared in specimens collected in the northwest Pacific Ocean and may indicate that some bamboo coral species are widespread in the Pacific, or that the mitochondrial markers are insensitive to recent speciation events. Many specimens were taken from flat bottom areas and, contrary to assumptions, the bamboo corals are not endemic to seamounts. The closure of some seamounts to trawling will protect bamboo corals from extinction, but not from local depletion.Communicated by M.S. Johnson, Crawley     

Worker size variability and foraging efficiency in Veromessor pergandei (Hymenoptera: Formicidae)

Summary As foragers of the harvester ant, Veromessor pergandei, travel further from their nest they spend significantly more time sampling seeds in experimental patches. Although accepted seeds are heavier than offered seeds, mass of accepted seed is not correlated with sampling time. Variably sized V. pergandei workers do not size-match; little, if any, variance in size of seed selected can be attributed to body size of forager. The lack of size-matching in V. pergandei suggests individual performance may be an inadequate measure of colony foraging success.     

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.     

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.     

Mammalian breath: trigger of defensive chemical response in a tenebrionid beetle (Bolitotherus cornutus)

Summary The tenebrionid beetle Bolitotherus cornutus everts a pair of quinone-producing defensive glands in response to mammalian breath. Experiments with a controlled airstream indicate that the beetle recognizes breath on the basis of temperature, humidity, and airflow dynamics. Under attack by mice the beetle everts the glands immediately upon being mouthed and may secure its release as a result. Against ants the beetle is protected by its tough exoskeleton and usually refrains from everting the glands. Other arthropods also show defensive responses when breathed upon.Paper No. 77 of the series Defense Mechanisms of Arthropods     

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.